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The CANADIAN
FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
fut, a4
January—March 2006
Volume 120, Number 1
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THE CANADIAN Ty
FIELD-NATURALIST
Volume 120
2006
Volume 122
The Ottawa Field-Naturalists’ Club Transactions
Promoting the study and conservation
of northern biodiversity since 1880
THE OTTAWA FIELD-NATURALISTS’ CLUB
OTTAWA CANADA
The Canadian Field-Naturalist
January—March 2006
Volume 120, Number |
Predicting Bird Oiling Events at Oil Sands Tailings Ponds and
Assessing the Importance of Alternate Waterbodies for Waterfow!:
a Preliminary Assessment
ROBERT A. RONCONI
Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada
Present address: Department of Biology, University of Victoria, P.O. Box 3020 Station CSC, Victoria, British Columbia
V8W 3N5 Canada; e-mail: rronconi@uvic.ca
Ronconi, Robert A. 2006. Predicting bird oiling events at oil sands tailings ponds and assessing the importance of alternate
waterbodies for waterfowl: a preliminary assessment. Canadian Field-Naturalist 120(1): 1-9.
Tailings ponds are an integral part of oil sands mining development in northeastern Alberta, but waterfow! and shorebirds
often land in these ponds during spring migration where they may become covered with oil. For decades, managers have
developed and implemented methods for deterring birds from landing in these ponds, yet no deterrent strategy is fully effec-
tive. Therefore, to enhance deterrence strategies, it will be important to understand the environmental conditions that influ-
ence bird use of tailings ponds. This study quantified waterfowl flights over, and use of, tailings ponds and compared this
use to waterfowl activity at natural waterbodies in the region over a single spring migration period. Results suggest that
waterfowl are most likely to land on tailings ponds before lakes have thawed, after which migratory ducks appeared mainly
to use natural waterbodies for migratory stopover sites. Very high numbers of waterfowl were observed on one waterbody,
Kearl Lake, suggesting that this lake may be of greater importance to spring staging waterfowl than previously thought. A
small sample of birds oiled at tailings ponds were examined in relation to spring weather conditions. Logistic regression
analysis demonstrated that the probability of birds being oiled tended to increase with precipitation levels. Results of this
study suggest that (1) preservation of natural waterbodies may play an important role in minimizing bird use of tailings
ponds, and (2) future bird deterrence efforts should especially aim to deter birds during rainy weather conditions when birds
may be more likely to become oiled. These results were from a small sample size, are preliminary in nature, and should be
interpreted with caution. A concerted and careful effort to collect and thoroughly analyze long-term records of oiled birds
may reveal important environmental effects predicting bird oiling events.
Key Words: waterfowl, oil sands, mining, tailings ponds, lakes,
In northeastern Alberta, oil sands development has
grown considerably over the past several decades. This
industry promises to grow into the 21‘' century since
recent oil reserve estimates have placed Canada sec-
ond on the world list of oil reserves with 175 billion
barrels of oil in oil sands deposits (CAPP 2003*). Al-
though most of these reserves are extractable with new
drilling technologies, an estimated 20% are recoverable
only with open pit mining practices (CAPP 2003*).
One of the primary environmental concerns stemming
from oil sands mining is the existence of tailings ponds
which have oil floating on the surface. Birds landing
on these ponds might become covered in oil and die.
This problem has been recognized since the initial
development of oil sands mining (SCL 1973; Schick
and Ambrock 1974*; Ward et al. 1976*; Hennan and
Munson 1979) and significant effort has been dedicat-
ed to researching the conditions that promote and pre-
vent birds from becoming oiled (Boag and Lewin 1980;
Gulley 1980; Yonge et al. 1981*; Ronconi and St. Clair
weather, migration, radar, conservation, Alberta.
2006; also see review by Golder Associates 2000*).
Nevertheless, birds still continue to be oiled and oil
sands companies carry on their search for better means
of reducing these incidents.
Previous oil sands research recommended the cre-
ation of uncontaminated waterbodies in close proximity
to tailings ponds, allowing birds alternate landing sites
(Gulley 1980; Golder Associates 2000*). Moreover,
research in other industrial contexts indicates that birds
can be deterred from an area more effectively when
alternate roosting sites (including ponds) are available
(Martin and Martin 1984; Gosler et al. 1995; Stevens
et al. 2000). Yet others have argued that such diversion-
ary ponds would only draw more birds to the region
(Allen 1990). Instead, the preservation of naturally
occurring waterbodies may be an optimal solution to
provide alternate landing sites for birds, particularly if
these alternative bodies are already used by migrating
birds in the region. Comprehensive surveys of water-
fowl at the major waterbodies in the vicinity of Alber-
2 THE CANADIAN FIELD-NATURALIST
ta’s oil sands were conducted prior to oil sands devel-
opment (Hennan and Munson 1979), but the current
status of these waterbodies and their use by migratory
birds are unknown. Moreover, subsequent surveys of
waterbird numbers at natural waterbodies (Shortt
1985*; Van Meer and Arner 1985*) did not compare
levels of bird activity between these natural ponds and
nearby tailings ponds. Such information would make it
possible to assess the importance of alternative water-
bodies as stopover sites during migration, and to assess
temporal changes within seasons in the attractiveness
of tailings ponds to waterfowl.
In addition to information about the alternative
stopover sites for migrating birds, researchers and man-
agers have also sought to understand the conditions
under which birds become oiled on tailings ponds. Over
a three-year period, Gulley (1980) identified three main
conditions as determinants of the numbers of birds that
were killed each year. First, more birds were oiled
when more birds were migrating (Gulley 1980), though
the synchrony of migration is not fully predictable and
tends to vary among both years and species (Bellrose
1976). Second, cold spring weather and delayed ice
break-up correlated with bird oilings. Finally, greater
than average seasonal rainfall has been correlated with
higher rates of oiled birds. In contrast, other studies
failed to observe any environmental (wind, tempera-
ture, precipitation) effects on the probability of bird
landings in tailings ponds (Ronconi and St. Clair
2006). Yet neither analysis examined the frequency of
bird oiling events in relation to daily weather patterns
and this information may be most likely to reveal the
causes of variation.
The study is situated at one of more than 10 oil sands
tailings ponds operating in northeastern Alberta (Gold-
er Associates 2000*), all of which are situated along
a major migratory flyway for waterfowl traveling to
the Peace-Athabasca Delta, an internationally signifi-
cant staging area (Bellrose 1976; Hennan and Munson
1979). The objectives in this study were three-fold.
First, | examined the seasonal abundance and direc-
tion of migratory bird movements over tailings ponds.
Second, I compared numbers of waterfowl that landed
at tailings ponds with numbers of birds at natural ponds
in a broader region and with historical data. Finally, I
assessed potential correlations between daily weather
conditions and the frequency of oiled birds found dur-
ing April and May of 2003. Because data were available
only for a single season, I present these as preliminary
results and encourage future researchers to investigate
these findings over longer time periods.
Methods
The study was undertaken at the Muskeg River Mine
(MRM), Albian Sands Energy Ltd., located approxi-
mately 75 km north of Fort McMurray, Alberta (Fig-
ure 1). During April and May 2003, the mine was
beginning initial oil production stages. The tailings
Vol. 120
pond complex consists of three ponds totaling ~ 740 ha
of surface area when full, but water coverage during
this study period was approximately 530 ha. Natural
waterbodies were selected for surveys based on acces-
sibility, size, and proximity to the MRM tailings ponds.
These included two lakes near the mine, four lakes
south of the mine near other oil sands development,
and eight lakes north of the mine that were isolated
from development (Figure 1). As these lakes were sit-
uated along a North-South axis relative to the MRM,
all lakes were potentially usable by waterfowl] migrat-
ing along this corridor.
Airport radar systems have been used to study bird
migration patterns in Alberta (Richardson and Gunn
1971), and marine radar has been used by ornitholo-
gists to study various aspects of bird activity (Cooper
et al. 1991; Burger 1997; Bertram et al. 1999; Gau-
threaux and Belser 2003). In this study, marine type
radar was used to assess spring migratory activity over
the MRM tailings ponds between 3 and 29 May 2003.
Migratory activity was measured by the groups of birds
flying over the study area (groups rather than individ-
uals are identifiable by radar) and the direction of flight.
Radar observations were made from 0.5 hours before
sunrise to 4 hours after sunrise. This period was con-
sidered a high migratory period in northeastern Alberta
(Richardson and Gunn 1971). Owing to some techni-
cal difficulties, directional data were not recorded for
the first four days of this period, and data collection
was not possible on six other days because of heavy
precipitation that obscured radar observations.
Tailings pond and lake surveys were conducted from
fixed observation points using binoculars and spot-
ting scopes. Tailings ponds were surveyed regularly
from 18 April to 29 May, and lakes were surveyed 9 to
28 May as most lakes were still completely or partially
frozen during the first week of May. Observers counted
individual ducks, gulls, coots, loons, and grebes on the
water. No more than two or three geese were observed
during any lake count; therefore, goose counts were
omitted from the results. Because of the large size of
McCleland Lake, all portions of it were not simulta-
neously visible, thus counts were only a minimum esti-
mate of bird numbers on that lake. Kearl Lake was
surveyed more often than the other natural waterbod-
ies because it was nearest to the MRM tailings ponds
and thus most likely to offer an alternate landing site
for migratory birds. During the mid-1970s Mildred
Lake was dammed, converting most of the lake into a
tailings pond and leaving only a portion of the lake in
its semi-natural state. Surveys of Mildred Lake in-
cluded only the semi-natural portion. When applicable,
results were compared to previous surveys (see Hen-
nan and Munson 1979 for detailed waterfowl surveys
in the region).
Reports of oiled birds on the tailings ponds were col-
lected from 10 April to 29 May, inclusive. Although
systematic searches for oiled birds were not conduct-
2006
ed, observers spent 4 to 8 hours observing waterfowl] at
the tailings ponds on most days thus ensuring some
consistency in effort throughout this period. Weather
data for wind speed and direction, temperature, and
rainfall were recorded from meteorological stations
on site, and snowfall events were converted into rain-
fall equivalency. Mean temperature is reported here as
the daily mean of maximum and minimum tempera-
tures. Wave conditions on the ponds and percent cloud
cover were recorded by observers. All occurrences of
oiled birds are reported (below); however, analyses
of weather effects were conducted only on oiled birds
that were still alive when found, an assumption that
those birds became oiled either the day they were
found or the previous night. For analysis by logistic
regression, oiled birds were the dependent variable
(1 = days with oiled birds; 0 = days with no oiled
birds). Days when observers were not present on site
were excluded from analyses. Weather data were the
independent variables (wind, temperature, precipitation,
wave conditions, % cloud cover). Except for wave
conditions, analyses of weather effects were assessed
for three time periods: date of oiled bird finding, one
day prior (i.e. previous 24 hour conditions), and two
days prior (i.e. previous 24-48 hour conditions). Wave
condition was only analyzed for the first two time
periods since it was considered likely that rough water
effects would not persist for as long as two days.
Logistic regression analysis (SPSS 11.5) was used to
assess weather effects and to build a model for pre-
dicting their effects on the probability of birds being
oiled. Owing to small sample sizes of data from a single
season, a liberal alpha value (0.10) was used for logis-
tic regression analysis. Tests were conducted for inter-
ance
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RONCONI: PREDICTING BIRD OILING AT OIL SANDS PONDS ;
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© Lakes Surveyed if =
Ga Muskeg River Mine ) ‘a 05
@ Fort McMurray | \3
~~ Rivers 2s ao in
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FIGURE 1. Map of study area showing the location of the
Muskeg River Mine and surrounding natural water-
bodies surveyed in 2003. Lake names are as follows:
(1) Poplar, (2) Ruth and Crane, (3) Mildred, (4) Isa-
dore’s, (5) Kearl, (6) McCleland, (7) Unknown Lakes,
(8) Go Go, (9) Herb, (10) Six Lakes, (11) Larocque,
and (12) Net.
NW &N Flights
NE & E Flights
SE & E Flights
SW & W Flights
Bird Groups per Hour
26/05/03
>
FiGurE 2. Migration patterns from radar data at the Muskeg River Mine tailings ponds, 3-29 May 2003. Bird groups were the
unit of measure because group size could not be inferred from radar images. Gaps in the data set resulted from days
of heavy precipitation which obscured radar observations.
4 THE CANADIAN FIELD-NATURALIST Vol. 120
TABLE 1. Numbers of waterfowl at the Muskeg River Mine (MRM) and surrounding natural waterbodies, 2003.
Distance (km) Other
Lake from Water
Location Date Coordinates MRM Size! Gulls Ducks _ birds
Muskeg River Mine
Tailings Ponds 18 April 57°13'47"N, 111°35'10"W 0 0 B85 3
20 April 0 8 0
23 April 0 32 0
24 April 0 13 0
25 April 4 13 0
26 April 0 22 4
27 April 5) 55 0
03 April 0 86 0
04 April 0 90 0
05 April 0 16 0
06 May 0 166 58
07-29 May 0-14 < 28 <2
Near Muskeg River Mine
Kearl Lake 09 May 57°18'08"N, 111°15'06"W 19 > 15 104 1]
15 May 300 2700 present
19 May 440 859
30 May 2 8 9
Isadore’s Lake 26 May 57°13'43"N, 111°36'29"W 1 = 66 102 2)
North of Muskeg River Mine
McCleland Lake? 23 May 57°30'35"N, 111°21'16"W 32 >> 12 645 5
Unknown Lake 23 May 57°50'25"N, 111°06'29"W V2 < 4 0 0
Unknown Lake 2 23 May 57°50'28"N, 111°06'46"W 72 < 0 4 2,
Go Go Lake 23 May 57°52'48"N, 111°02'56"W Ty « 0) 0 2
Herb Lake 23 May 57°54'13"N, 111°02'11"W 80 = 0 5 1
Six Lakes 24 May 57°58'SO"N, 111°01'45"W 88 = 0 0 3
Net Lake 24 May 58°07'23"N, 110°46'55"W 108 = 0 5 3
Larocque Lake 24 May 58°06'S1"N, 110°49'11"W 107 = 3 0 1
South of Muskeg River Mine
Mildred Lake 25 May = 57°02'45"N, 111°33'52"W 18 = 0 0 0
Crane Lake 25 May 56°59'45"N, 111933'12"W 24 0 158 present
28 May 0 106 5
Ruth Lake 25 May 56°59'22"N, 111°34'06"W Ds > 19 306 present
27 May 12 350 1
Poplar Reservoir Dil Maya y5 62553 20Ns ile29 500i 3p = 0 15 2
Relative to Muskeg River Mine tailings ponds = (approximately equal), > (larger), < (smaller).
2Tncludes coots, loons, grebes.
3 Entire lake was not observable, therefore counts are minimum numbers of birds.
actions and confounding variables. For detailed descrip-
tion of the model-building approach see Ronconi and
St. Clair (2006) and Hosmer and Lemeshow (1989).
Results
Counts of bird groups by radar provided an indica-
tion of overall bird activity relative to the numbers of
birds landing at tailings ponds. The radar system was
operational from 3 — 29 May, during which two peaks
in migratory activity were observed (Figure 2). Peaks
in migratory flights (NW & N flights) coincided with
two peaks in bird activity (bird groups per hour). How-
ever, a third peak in groups per hour on 27 May showed
no corresponding peak in NW & N flights. Notwith-
standing the three peaks, these data document high
migratory volumes over most of May when counts of
waterfowl on lakes were conducted. However, the num-
bers of ducks that also landed on the tailings ponds
(Table 1) were consistently low over this same period
with no comparable peaks.
Counts of birds at tailings ponds and natural water-
bodies showed much variability and peaks occurred
at different times for different lakes. Counts of ducks
and other waterbirds at the MRM tailings ponds (Table
1) showed peaks in activity from 26 April through to
6 May, after which activity dropped considerably and
remained low throughout May. Conversely, high counts
of birds on surrounding lakes were observed in May
(Table 1), though comparable counts from April were
not possible because the natural waterbodies were still
frozen at that time. In particular, Kearl Lake showed
a peak around mid May with an estimated 2700 ducks
2006
and 300 gulls on 15 May. As numbers were low at Kearl
Lake in early May, these data, in combination with low
May counts at tailings ponds, suggest that bird use of
tailings ponds may have shifted to natural waterbodies
for much of May. North of MRM, counts showed very
low numbers except for McCleland Lake with over
600 ducks on 23 May. This high count suggests that
McCleland was still being used as a migratory stopover
at this time, but lakes farther north were likely only
occupied by small numbers of breeding pairs. Between
25 and 27 May, counts of lakes south of MRM had
higher numbers than maximum counts at MRM tail-
ings ponds throughout April or May, but by this date
many of these birds may have been resident breeders.
Although duck use of tailings ponds appeared low
in comparison with natural waterbodies, overall duck
numbers for this region were higher than historical sur-
veys reported. Comparisons with Hennan and Mun-
son’s (1979) surveys showed several changes (Table
2). Duck numbers at McCleland Lake fell within the
range of historical counts and were higher than four
of the five years surveyed between 1973-1977, even
though McCleland was only censused once in 2003.
The highest count at Kearl Lake in 2003 was 4.6 times
the maximum recorded by once annual surveys be-
tween 1973 and 1977. South of the MRM, Mildred
Lake showed no use in 2003, in contrast to abundant
use in previous surveys, whereas numbers were high-
RONCONI: PREDICTING BIRD OILING AT OIL SANDS PONDS 5
er at Ruth Lake than were recorded during the 1970s.
Due to inconsistencies in data collection methods
and dates of surveys, we can not speculate on trends
or changes in waterfowl abundance since the 1970s.
However the dramatically high count of ducks on Kear!
Lake suggest that this lake may be more heavily used
as a migratory stopover site than might have been sus-
pected from earlier surveys.
Fifteen oiled birds were observed at the MRM tail-
ings ponds during the study period (Table 3), nine of
which were alive when found. Of these, four Lesser
Scaup (Aythya affinis) were lightly oiled and able to
fly, making it probable that these were non-fatal oiling
events. The remaining live oiled birds included one
goose, one shorebird, one gull and two ducks. Days
when live oiled birds were found were plotted in rela-
tion to weather variables (Figure 3). A logistic regres-
sion analysis was conducted on 47 days of observa-
tions, six of which had reported live oiled birds. This
analysis found no effect of temperature, sea-state, wind
direction, or cloud cover. Univariate analyses found
date (P = 0.160) to be marginally significant, and rain-
fall (P = 0.086), rainfall during the previous 24 hours
(P = 0.066), and wind speed during the previous two
days (P = 0.050) as significant variables affecting the
probability of birds being oiled. However, when com-
bined in a multivariate analysis no variables retained
significance but the two rainfall variables were only
TABLE 2. Comparisons of spring 2003 duck counts with historical counts from 1973-1977 (Hennan and Munson 1979).
Maximum Number of Ducks Recorded During Spring Migration
Location 1973 1974
MRM Tailings Ponds ~ =
Kearl Lake 529 586
McCleland Lake! 194 1154
Mildred Lake? 246 313
Ruth Lake 113 20
Poplar Reservoir - ~
1975
1976 1977 2003
- - - 166
24 37 140 2700
225 360 195 645
136 17 - 0
26 1] - 350
16 l 14 15
' For 2003 surveys the entire lake was not observable; therefore, count is a minimum estimate of ducks
ca. 1975 most of Mildred Lake was converted into a tailings pond. Surveys in 2003 checked only the portion of Mildred
Lake that remains in its natural state.
TABLE 3. Summary of oiled birds found at the Muskeg River Mine tailings ponds, April and May 2003.
Date Common Name Scientific Name
26 April goose spp. unknown
03 May Common Goldeneye Bucephala clangula
19 May Lesser Scaup Aythya affinis
20 May Bonaparte’s Gull Larus philadelphia
21 May small shorebird Calidris spp.
21 May Lesser Yellowlegs Tringa flavipes
26 May Northern Shoveler Anas clypeata
26 May Northern Shoveler Anas clypeata
26 May small Shorebird unknown
26 May medium passerine unknown
26 May large shorebird unknown
Group Size Status Oiled Status
1 Live heavy — >80%
1 Live heavy — 80%
4 Live light < 10%!
] Live heavy — 70%
1 Live heavy — 60%
1 Dead moderate — 30%
] Live heavy > 50%
1 Dead heavy — 100%
1 Dead heavy — 100%
1 Dead heavy — 90%
] Dead heavy — 100%
' Lightly oiled but able to fly.
6 THE CANADIAN FIELD-NATURALIST
marginally non-significant (P = 0.107 and 0.105). To
examine the apparent, though non-significant, effect of
rainfall on birds being oiled, the cumulative rainfall was
summed for the day of plus day before oiled birds were
found and this had a significant effect (Wald = 4.973,
d.f. = 1, P = 0.026) on the probability of birds being
oiled. The positive coefficient of this variable indicated
that the probability of a bird becoming oiled increased
with higher rainfall. With the cumulative rainfall vari-
able, the model was significant (-2LL = 30.264, model
- = 5.636, df. = 1, P =0.018), but the model fit was
weak (Nagelkerke’s r? = 0.212, Hosmer and Lemeshow
test ¥° = 1.330, df. = 1, P= 0.095). Small sample size
likely contributed to the marginal significance of vari-
ables and the poor fit of the model, yet the persistence
of rainfall effects indicate that precipitation was relat-
ed to birds being oiled this year and this may be true
more generally.
Discussion
Timing of Spring Migration
Although the radar system was not operational in
April, visual observations suggested that migration
activity in the study area was fairly low in April and
early May and that migration peaked in mid-May. The
radar was also not in operation (due to weather) for
six days in May near peak migration (Figure 2); there-
fore, migration records presented here may have missed
some critical portions of the migration. Nevertheless,
the results on migration timing agree with other stud-
ies in northeastern Alberta. Richardson (1969) reported
peak spring migration over east-central Alberta in May.
Aerial surveys of lakes in this region during 1972 and
1973 reported peak numbers on 15 May and 25 May,
respectively (SCL 1973). Likewise, Schick and Am-
brock (1974*) showed peak migration to occur in mid-
dle and end of May in 1973. During aerial surveys in
1976/1977, Hennan and Munson (1979) reported peak
spring migration for ducks in early May, with most
abundant species being scaup (Aythya sp.), Ring-
necked Ducks (Aythya collaris), Buffleheads (Buce-
phala albeola), and to a lesser extent Mallards (Anas
platyrhynchos). Annual variation in timing of spring
migration is likely to vary with timing of ice break-up
of the Athabasca River and also the Peace-Athabasca
Delta (Schick and Ambrock 1974*).
Although the timing of migration in northeastern
Alberta is relatively well known, the composition and
numbers of birds passing through the area are less well
known. During the same time period of this study in
2003, Ronconi and St. Clair (2006) reported the fol-
lowing numbers of birds flying over the tailings ponds:
1140 ducks, 2132 shorebirds, 11 339 geese, 218 swans,
965 gulls, 56 terns and 205 others (loons, grebes,
cranes, herons, cormorants, coots). These numbers give
some indication of guild composition during migration
but should be used cautiously as such because these
were visual, daytime observations and many birds in
Alberta migrate at night and at high altitudes (Richard-
Vol. 120
son 1971; Blokpoel 1973; Blokpoel and Burton 1973).
Hennan (1972) reported hundreds of thousands of
waterfowl using the Peace-Athabasca Delta as a spring
staging area, thus providing some indication of the
volume of birds that may pass over the tar sands area
each year. Nevertheless, many individuals of some
waterfowl species may also take alternate migration
routes to Alaska or nest in southern Canada (Miller et
al. 2005).
Finally, in this study area we may also be observing
resident as well as migratory waterfowl. This study
makes no distinction between the two groups; howev-
er, the high proportions of northwesterly and northerly
flights (Figure 2) suggest that most of the birds ob-
served were migrants. It is likely that the small num-
bers of birds observed on the tailings ponds in May
and on some lakes in late May were residents rather
than stopover migrants. Surprisingly, the ecology of
waterfowl at staging and stopover areas is fairly poorly
understood (Arzel et al. 2006).
Numbers of Birds on Ponds and Lakes
Despite the apparent peak in migration around mid-
May, the numbers of ducks on the tailings ponds
peaked sooner than this. Working in a nearby area,
Gulley (1980) found that duck dependence on tailings
ponds was higher when spring ice break-up was de-
layed. In 2003, ice break-up on the nearby Athabasca
River began 23 April and was finished by 1 May, yet
many of the nearby lakes remained at least partially
frozen through the first week of May. Duck surveys
at Kearl Lake showed low numbers on 9 May when
this lake was still partially frozen, but numbers grew by
23 times only one week later when the lake was ice-
free. These results are mainly qualitative, but they sup-
port the hypothesis of others (Gulley 1980; D. Mar-
tindale, personal communication) that duck use of
tailings ponds is highly dependent on the availability
of open water elsewhere. After 9 May, duck numbers
at the MRM tailings ponds were seldom above 10 indi-
viduals, while thousands of individuals were using
Kearl Lake less than 20 km away. Nevertheless, these
conclusions about seasonal change in dependence on
tailings (early spring) to natural waterbodies (mid to
late May) are tenuous due to several inconsistencies
in survey methodology: (1) Kearl Lake was the only
natural water body surveyed more than twice, (2) most
natural waterbodies were surveyed only in late May
and might have missed peak migration, and (3) lack
of personnel did not allow for simultaneous surveys of
tailings ponds and natural waterbodies. Bird move-
ments between wetlands may have been a concern;
however, the extremely low numbers of ducks at the
tailings ponds after 7 May suggest little, if any, move-
ment between artificial and natural ponds after this
date. Finally, no data are available on wetland charac-
teristics, which were beyond the scope of this project
but may have had significant influence on waterfowl
use of waterbodies.
2006
oe Mean Wind Speed
cast sesees Mean Temperature
5 20 Daily Precipitation
Ss = One or More
& Live Oiled Bird Found
E
= 15
FEMS
ae
O
wee le
Se
ur
2S aoa
OO or
- ® ns U \
ep)
12)
& 5
: f
-10
g g gf
RONCONI: PREDICTING BIRD OILING AT O11. SANDS PONDS 7
. *,
cer
A)
+ gle i 1
a hee
yo tN ae
ia Ce eee | Reece
o 7
& & se \
9 IN a> VY
FIGURE 3. Weather patterns with respect to dates when live oiled birds were found at the Muskeg River Mine tailings ponds,
April and May 2003.
Comparisons with census results from the 1970s sug-
gest some changes in lake use. However, duck activity
levels can vary widely within seasons, as was indicated
in our results and other studies (e.g. SCL 1973; Gul-
ley 1980; Van Meer and Arner 1985*; Shortt 1985*).
The maximum counts between 1973 and 1977 were
typically based on only one or two surveys during May,
making them similarly prone to high variability. Al-
though qualitative and tenuous, these comparisons rein-
force the belief (SCL 1973; Schick and Ambrock
1974*; Ward et al. 1976*; Hennan and Munson 1979)
that some lakes are important as stopover sites for
migratory ducks, though not as major staging areas.
As with our results, censuses from 1973 to 1977 ranked
McCleland and Kearl lakes among the most important
stopover sites for ducks (Hennan and Munson 1979).
Other lakes in the region with high numbers included
Little McCleland, Mildred, Horseshoe, and Saline lakes
(Hennan and Munson 1979). The lack of ducks on Mil-
dred Lake in 2003 is not surprising considering only
a small portion of the lake remains in its natural state.
The lack of ducks on remote lakes north of McCleland
Lake was not surprising given the relatively small size
of the lakes surveyed. Nonetheless, Hennan and Mun-
son (1979) identified other remote lakes with high wat-
erfowl densities, though these were not accessible for
surveys by land in 2003.
Generally the oil sands region is not recognized as
an important staging area for migratory waterfowl
(SCL 1973; Schick and Ambrock 1974*; Ward et al.
1976*; Hennan and Munson 1979), although some
staging does occur nearby. The Peace-Athabasca Delta,
north of oil sands regions, has been recognized as an
internationally significant staging area for hundreds of
thousands of waterfowl (Hennan 1972). Further south
of Fort McMurray, Gordon Lake was previously rec-
ognized as an important waterfowl staging area with
spring numbers greater than 5600 in 1973 and 1974
(Hennan and Munson 1979). The count of 2700 water-
fowl at Kearl Lake in 2003 suggest that this lake may
bear greater importance as a staging area than might
have been concluded from those earlier surveys, but
further surveys may be needed to verify annual varia-
tion in waterfowl use of this lake. Surveys of other nat-
ural waterbodies also revealed that smaller ponds sup-
port small populations of birds throughout the spring
and summer (Shortt 1985*). Because availability of
natural waterbodies may minimize duck dependence
on tailings ponds, further investigations of waterfowl
use of these waterbodies is merited. Should it continue
to appear that they are important to waterfowl, future
oil sands developers may consider conserving these
locations and avoiding the establishment of tailings
ponds in close proximity to them.
8 THE CANADIAN FIELD-NATURALIST
Probability of Oiling
Logistic regression analysis of several weather vari-
ables found wind speed and precipitation to be the only
variables that might predict the few observations of
oiled birds in 2003. Strong winds may mix waters and
spread oily films over wider areas, hence creating
greater risk of oiling, but with consistent winds oily
sheets may also be pushed to one shore of the pond.
Thus any link between winds and oiling may be com-
plicated. A two-day cumulative rainfall measure best
predicted days with oiled birds, but rainfall the day of
an oiling event and rainfall from the previous 24 hours
showed some possible predictive ability. These uncer-
tain results, along with small sample size, suggest only
a potential effect of precipitation. Results of this study
support a previous analysis across years (Gulley 1980)
which showed that precipitation levels affected num-
bers of waterfowl deaths. Although Ronconi and St.
Clair (2006) found no correlation between weather and
bird landings on tailings ponds, the more important
factor may be whether or not birds become oiled when
they land.
If precipitation increases the frequency of birds be-
ing oiled, as these results cautiously suggest, I specu-
late that it may be caused by two possible mechanisms.
First, poor weather conditions such as rain, strong
winds, headwinds, and cold temperatures can delay
migration, cause changes or a temporary reversal in
migration direction, and cause migratory birds to be
temporarily grounded (Richardson 1978; Elkins 1983).
Waterfowl in particular appear to be erratic in their mig-
ratory behaviour which may be influenced by break-up
of ice cover (P6ysd 1996), temperatures (Austin et al.
2002), weather fronts (Custer et al. 1997), and espe-
cially wind patterns (Bergman 1978; Liechti 1993;
Bruderer 1994; Alerstam and Gudmundsson 1999).
The significance of precipitation as a predictor of oil-
ing may be due to correlations between rainfall and
other weather changes, which may together be affect-
ing bird movements and halting of migrations. The
more frequently migratory waterfowl are forced to
move and stop during migration, the greater the risk of
encountering oil sands ponds and oil on the water sur-
face in these ponds.
Second, precipitation may additionally increase oil-
ing probabilities through birds experiencing reduced
visibility during landings under rainy and snowy con-
ditions. Without clear views of the water, they may be
more likely to land in patches of oil which might other-
wise be avoided. Reduced visibility as a causal mech-
anism may also explain the non-significance of wind
speed, wind direction, and temperature since these
variables should be less likely to affect the visibility
of oil patches. Snow could affect in-flight visibility
of birds, and on two of six days with oiled birds, the
precipitation that occurred was a mix of rain and snow.
Moreover, for three of the remaining days with oiling
Vol. 120
events (19, 20, and 21 May), heavy snow had fallen for
two days previously (17 and 18 May). Although fog
was not measured in this study, fog may also limit vis-
ibility and thus influence oiling events (Gulley 1980).
Careful collection and thorough statistical analyses
of long-term oiled bird records at tailings ponds may
reveal important trends and weather association. These
data could be valuable to improving the effectiveness
of bird deterrent strategies, and I encourage oil sands
companies to standardize collection protocols and pool
their data for such an investigation.
Conclusions
As the development of oil sand mines continues in
the 21*' century, tailings pond and bird oiling problems
will persist, and findings from current and previous
studies will become more relevant to managers seek-
ing a solution to this issue. Findings from this study
suggest the importance of maintaining natural water-
bodies in the region, especially those that are already
used as migratory stopover sites for waterfowl. The
significance of rainfall affecting numbers of birds being
oiled suggests that bird deterrence efforts could benefit
from enhancement on rainy days, and future deterrent
development should pursue strategies that are effective
during rainy weather more so than during clear condi-
tions. With decades of data on oiled birds, and 30 years
of research on bird deterrence, the oil sands industry in
northern Alberta is positioned to offer significant insight
into waterfowl management issues for future develop-
ment here and at other sites of industrial development.
Acknowledgments
Logistical support and funding was provided by
Albian Sands Energy Ltd., with additional logistical
support from the University of Alberta, Edmonton. I
thank Colleen Cassady St. Clair for support and guid-
ance throughout this study and for reviewing drafts of
this manuscript. I am grateful for support from Darrell
Martindale, excellent field assistance from Andrew
Forrest and Sarah Wong, and technical support from
Will Neagle and Peregrine Systems Inc. Field work
for this project was made possible with the help of
Albian Sands employees Christine Theriault, Maurice
Kay, Bob McKenzie, Thor Einarson, Brian Kean, and
others on the Tailings and Site Services Teams. This
manuscript was improved by comments from A. J.
Erskine and one anonymous reviewer.
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Received 17 February 2004
Accepted 17 May 2006
A Test of Interspecific Effects of Introduced Eastern Grey Squirrels,
Sciurus carolinensis, on Douglas’s Squirrels, Tamiasciurus douglasii,
in Vancouver, British Columbia
YEEN TEN Hwanc!* and SERGE LARIVIERE!”
‘Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada
*Delta Waterfowl Foundation, R.R. #1, Box 1, Site 1, Portage La Prairie, Manitoba RIN 3A1 Canada
>Corresponding author: Department of Biology, University of Western Ontario, 1151 Richmond Street N., London, Ontario
N6A 5B7 Canada; e-mail: yeenten@ gmail.com
Hwang, Yeen Ten, and Serge Lariviére. 2006. A test of interspecific effects of introduced Eastern Grey Squirrels, Sciurus caro-
linensis, on Douglas’s Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia. Canadian Field-Naturalist
120(1): 10-14.
We compared the effects of absence and presence introduced Eastern Grey Squirrels (Sciurus carolinensis) on the demography
of native Douglas’s Squirrels (Jamiasciurus douglasii) in two urban parks in Vancouver, British Columbia: Ecological
Reserve #74 in Pacific Spirit Regional Park (Douglas’s Squirrel only) and Stanley Park (Douglas’s and Eastern Grey squirrels).
Based on the exploitative competition hypothesis, we predicted that in the presence of introduced Eastern Grey squirrels,
Douglas’s Squirrels would occur at lower densities, have larger home ranges, lower body mass, and poorer reproduction. Using
mark-recapture methods, we found no differences in density, home range, or body mass of Douglas’s Squirrel between parks.
However, the proportion of breeding Douglas’s Squirrels was higher in Ecological Reserve #74 in the absence of Eastern Grey
Squirrel, than in Stanley Park. We found no evidence that Eastern Grey Squirrels are displacing Douglas’s Squirrels in Stanley
Park, but less conspicuous negative effects such as reduced breeding propensity may still reflect the competitive interactions
of the two squirrel species.
Key Words: Density, home range, body mass, reproduction, Eastern Grey Squirrel, Sciurus carolinensis, Douglas’s Squirrel,
Tamiasciurus douglasii, exploitative competition, British Columbia.
In western Canada, Eastern Grey Squirrels (Sciurus
carolinensis) were introduced to Stanley Park in the
city of Vancouver, British Columbia, shortly before
1914. The three to four pairs of introduced Eastern
Grey Squirrels came from eastern Canada (Robinson
and Cowan 1954). In the early 1980s, Eastern Grey
Squirrels colonized the rest of the city of Vancouver,
including many of its urban parks. Since then, Eastern
Grey Squirrels have spread to many parts of the Greater
Vancouver Regional District (Gonzales 1999, 2000).
On the mainland in southwest British Columbia, the
native tree squirrel is the Douglas’s Squirrel (Jamias-
ciurus douglasii); it is the ecological counterpart of
the Red Squirrel (7. hudsonicus) that is widespread in
temperate forests of North America (Steele 1998, 1999).
The Eastern Grey Squirrel is native in eastern North
America (Hall 1981) and its geographical range over-
laps broadly with that of the Red Squirrel. Although
Red Squirrels are associated with conifer forests and
Eastern Grey Squirrels with mast-bearing hardwood
forests, the two species often coexist in mixed forests
or urban parks (Riege 1991). Forty years after Eastern
Grey Squirrels were introduced to southwestern British
Columbia, Robinson and Cowan (1954) conducted a
live-trapping and observational study on Eastern Grey
Squirrels in Stanley Park. They noted that, in decidu-
ous habitat in Stanley Park, Eastern Grey Squirrels
lived at high densities together with lower densities
of Douglas’s Squirrels. They suggested that the East-
ern Grey Squirrels outcompete Douglas’s Squirrels for
food, and thus restrict Douglas’s Squirrels to areas of
predominately coniferous habitat (Robinson and Cowan
1954). Whether competitive interactions between the
two species still exist today (> 80 years since the intro-
duction of Eastern Grey Squirrels) remains unknown.
In Great Britain, Eastern Grey Squirrels were intro-
duced at the end of the 19" century, and have caused
the decline of the smaller native European Red Squir-
rel (Sciurus vulgaris) from its former range (Gurnell
and Pepper 1993; Wauters and Gurnell 1999; Tean-
gana et al. 2000). The mechanisms of replacement of
European Red by Eastern Grey squirrels are not yet
fully understood (Skelcher 1997). Juvenile recruitment
of European Red Squirrels is typically lower when
Eastern Grey Squirrels are present (Wauters et al. 2000,
2001). Also, diseases such as parapoxvirus have been
hypothesized to facilitate the replacement of European
Red Squirrels by Eastern Grey Squirrels in the United
Kingdom (Tompkins et al. 2003).
The Eastern Grey Squirrel averages 400-700 g in
body mass (Koprowski 1984), and is larger than Doug-
las’s Squirrel, which averages 250-300 g (Steele 1999).
Douglas’s Squirrels feed primarily on coniferous seeds,
new shoots of conifers, green vegetation, acorns, nuts,
mushrooms, fruits, and berries. Also, Douglas’s Squir-
rels actively defend food middens within their territory
10
2006
(Steele 1999). In its native range, the Eastern Grey
Squirrel feeds heavily on hickory nuts, beechnuts,
acorns, and walnuts (Koprowski 1984); however, intro-
duced Eastern Grey Squirrels in British Columbia feed
extensively on maple, oak, hazelnut, mushrooms,
berries and pine cones (Robinson and Cowan 1954).
Both species prefer seeds, and other plant foods pre-
dominate in the diet only when seed crops are small
(Moller 1983). Also, in Stanley Park, a few Eastern
Grey Squirrels depend almost entirely on “hand-outs”
from park visitors as an artificial food source. Eastern
Grey Squirrels living in close proximity to humans
often venture out of the forests and feed on artificial
food sources.
Eastern Grey Squirrels have a dominance hierarchy
and are not territorial (Flyger 1955; Thompson 1977)
whereas Douglas’s Squirrels are territorial year round
(Steele 1999). Territorial behaviour exhibited by Doug-
las’s Squirrels is mainly used against conspecifics sim-
ilar to Red Squirrels (Smith 1981). Intruding Eastern
Grey Squirrels are generally free to enter their terri-
tory (Robinson and Cowan 1954).
The interspecific effects of the introduced Eastern
Grey Squirrels on the native Douglas’s Squirrels are
unknown in southwestern British Columbia. We tested
the hypothesis that Eastern Grey Squirrels negatively
impact native Douglas’s Squirrels through exploita-
tive competition for resources. With the introduction
of Eastern Grey Squirrels, resources such as food and
nest sites in areas where both squirrels occur would
decrease, causing an increase in home range size of
Douglas’s Squirrels because larger ranges are needed
to collect more food (Schoener 1971). Larger home
ranges would decrease the density of Douglas’s Squir-
rels in the given area. Also, with increased home ranges,
Douglas’s Squirrels would expend more energy forag-
ing and defending the area and would have less ener-
gy for reproduction and maintenance. Therefore, we
predicted that in areas with Eastern Grey Squirrels,
Douglas’s Squirrels would occur at lower densities,
have larger home ranges and lower body mass, and that
they would reproduce less well.
Materials and Methods
We worked at two study sites. The species co-occur
in Stanley Park, which was the original site of Eastern
Grey Squirrel introduction to western Canada, and of
Robinson and Cowan’s (1954) study. The control study
site was Ecological Reserve #74 (Pacific Spirit Region-
al Park) where only Douglas’s Squirrels occur. Al-
though Eastern Grey Squirrels do not occur in Eco-
logical Reserve #74, they do occur in residential areas
nearby and are abundant on the University of British
Columbia campus only 1-2 km distant.
Both study areas are in the Coastal Western Hem-
lock biogeoclimatic zone. The dominant coniferous
species in Stanley Park were Western Hemlock (Tsuga
HWANG AND LARIVIERE: EFFECTS OF GREY ON DOUGLAS’S SQUIRRELS 1}
heterophylla), Western Redcedar (Thuja plicata), and
Douglas Fir (Pseudotsuga menziesii). Stanley Park is
a mosaic of second- (> 50 years old) and old-growth
(> 120 years old) conifer stands. Ecological Reserve
#74 represented a mature conifer stand (> 90 years
old) with Western Hemlock, Western Redcedar, Dou-
glas Fir, and Sitka Spruce (Picea sitchensis) with scat-
tered Red Alder (Alnus rubus) and Big Leaf Maple
(Acer macrophyllum). Understory and ground cover in
both study areas consisted mainly of Vine Maple (Acer
circinatum), Salmonberry (Rubus spectabilis), Red
Huckleberry (Vaccinium parvifolium), Salal (Gaulthe-
ria shallon), leaf litter, ferns and mosses.
In each study area, we established a 9-ha live-trap-
ping grid consisting of 96 (6 by 16) stations located
at 30 m intervals. We set Tomahawk live-traps (model
201, Tomahawk Live Trap Co., Tomahawk, Wisconsin)
at alternate trap stations, resulting in 48 traps per live-
trapping grid. We live-trapped squirrels for 2 days
every 2 weeks from May to August 1997 (9 trap ses-
sions) in both study areas.
We baited the traps with sunflower seeds and set
them in early morning and checked traps 4-6 hours
later. Captured squirrels were ear-tagged with num-
bered metal tags (Monel #1, National Band and Tag
Co., NewPort, Kentucky), sexed and weighed (+ 5 g
using a Pesola® spring balance). We noted breeding
condition by palpation of male testes and female mam-
maries (Sullivan and Moses 1986).
We estimated population size, proportion of repro-
ductive squirrels, body mass, and home range size.
Population size and trappability were estimated for
each trap session using the Jolly-Seber mark-recap-
ture model (Jolly 1965; Seber 1965). We were not able
to use radio-telemetry to estimate the territory size of
Douglas’s Squirrels in this study; therefore we esti-
mated home range size by using trapping location for
animals that were captured = 2 times using the exclu-
sive boundary strip method (Hayne 1949, 1950; Stickel
1954). Also, we only considered home ranges of fe-
males because in late spring and early summer, males
roam extensively while searching for potential mates.
We used chi-square test to examine the trappability
of squirrel populations between both sites. We used
one-tailed t-test to compare densities of Douglas’s
Squirrels between sites. Due to unequal variances for
Eastern Grey Squirrel densities in Stanley Park, we
used Mann-Whitney U test to compare differences in
densities between both species in the two study areas.
We used two-way Analysis of Variance (ANOVA) to
examine the difference in body mass and home ranges
of Douglas’s Squirrels between Ecological Reserve
#74 and Stanley Park. The Z-test was used to test for
significant differences in the proportion of squirrels
in breeding condition (Sokal and Rohlf 1995). Since
sample sizes of breeding individuals were small, the
statistical power of the Z tests is very low in detecting
12 THE CANADIAN FIELD-NATURALIST
Vol. 120
Table 1. Summary of demographic parameters for Douglas’s Squirrels (DS) and Eastern Grey Squirrels (EGS) in Ecological
Reserve #74 and Stanley Park, Vancouver, British Columbia, Canada from May to August 1997. Values shown are mean#+ SE.
Ecological Reserve #74 Stanley Park
DS DS EGS
Density (individuals/ha) 3.5+04 2.9+0.1 2.5+04
Home range (ha) 0.37 + 0.096 0.44+0.11 —
Trappability (7%)
Male 74.7 60.3 30.0
Female 80.9 74.0 30.0
Body mass (g)
Male 181.8 + 2.69 184.5+3.81 608.6 + 11.91
Female 177.4 +4.62 181.1+4.40 603.7 + 15.59
Proportion of breeding females (7%) 20 5) 77
a significant effect; however, the Z-statistic was used as
an index of differences between study areas. All val-
ues are reported as mean + SE.
Results
Density and home range
From May to August 1997, we captured 30 male and
24 female Douglas’s Squirrels in Pacific Spirit Park.
In Stanley Park, we caught 19 male and 25 female
Douglas’s Squirrels. Eastern Grey Squirrels were only
found in Stanley Park, and we caught 22 males and
17 females. See Table 1 for a summary of demograph-
ic parameters.
Trappability of Douglas’s Squirrels in Pacific Spirit
Park was similar between males and females, at 74.7%
and 80.9%, respectively. Male and female Douglas’s
Squirrels in Stanley Park also have relatively high trap-
pability values, both ranging from 60.3% to 74.0%.
On the other hand, Eastern Grey Squirrels had very
low trappability, with males and females both having
30%. Trappability of Eastern Grey Squirrels was sig-
nificantly lower in Stanley Park (y? = 14.54, df = 1,
P <0.001) than for Douglas’s Squirrels.
Densities of Douglas’s Squirrels did not differ
Gia osridig—i2ene) — OMI) libetweent = colopical
Reserve #74 (3.5 individuals/ha + 0.4) and Stanley
Park (2.9 individuals/ha + 0.1). In Stanley Park, density
also did not differ (Mann-Whitney T = 56, P = 0.71)
between Eastern Grey Squirrel (2.5 individuals/ha + 0.4)
and Douglas’s Squirrels (2.9 individuals/ha + 0.1).
No effect of sex (F, ,, = 0.18, P = 0.67) or study
area tes = 1.24, P = 0.27) on the home range of
Douglas’s Squirrels, nor were significant interactions
detected (Ban = 0.44, P=0.51). Home ranges aver-
aged 0.31 + 0.05 ha (n = 96).
Body mass and reproduction
No effect of study area (F, 93 = 0.075, P = 0.78) on
body mass of Douglas’s Squirrels was found, but there
was a gender effect (F, ,,= 5.41, P = 0.022), with males
(185.3 + 4.1 g, n = 48) being heavier than females
(177.2 + 5.3 g,n = 49) in both parks.
For comparison purposes, Eastern Grey Squirrels
in Stanley Park averaged 606.4 + 19.1 g with no gen-
der difference (t = 0.25, df = 37, P = 0.80). On average,
male Eastern Grey Squirrels weighed 608.5 + 11.9 g,
and females weighed 603.8 + 15.6 g.
From 20 May to 28 August 1997, proportions of fe-
male Douglas’s Squirrels in breeding condition differed
significantly between sites (Z = 2.56, P < 0.001), with
a higher proportion of breeding females in Ecological
Reserve #74 compared to that in Stanley Park: 20%
(15 breeding out of 77 individuals) and 5% (3/62),
respectively. In Stanley Park, the proportion of fe-
male Eastern Grey Squirrels in breeding condition
(77%: 17/22) was higher than for Douglas’s Squirrels
(5%: 3/62). However, the proportion of breeding
male Douglas’s Squirrels at the two sites was similar
and averaged 98.3% (59/60).
Discussion
We did not detect strong effects of exploitative com-
petition between Eastern Grey Squirrels and Douglas’s
Squirrels. There was no evidence of reduced density,
large ranges, and lower mass in Douglas’s Squirrels
where the two species co-occurred. However, the pro-
portion of breeding Douglas’s Squirrels was significant-
ly lower in Stanley Park compared to that in Ecologi-
cal Reserve #74, suggesting possible negative effects
of Eastern Grey Squirrels on the breeding propensity
of Douglas’s Squirrels.
Densities of Douglas’s Squirrels are limited by food
availability (Sullivan and Sullivan 1982; Ransome and
Sullivan 1997), and old-growth forests provide higher-
quality habitat for Douglas’s Squirrels than younger
forests due to greater and more reliable quantities of
conifer seed (Buchanan et al. 1990). In our study, the
densities of Douglas’s Squirrels were similar at both
sites, suggesting that habitat suitability was similar, and
further suggesting that reduced breeding propensity
was caused by the presence of Eastern Grey Squirrels,
and not simply an artifact of difference in food avail-
ability among study areas.
A possible explanation for weak effects of exploita-
tive competition between the two species of squirrels
could be due to the fact that Eastern Grey Squirrels
often can exploit broadleaved and deciduous wood-
2006
land to increase breeding success by feeding on high-
energy food (Skelcher 1997; Gurnell et al. 2001), but
since both study areas consist of mixed woods with
high conifer content, advantages in breeding success of
Eastern Grey Squirrels were perhaps less pronounced
(Teangana et al. 2000). Thus, Douglas’s and Eastern
Grey Squirrels could coexist for many years although
perhaps Douglas’s Squirrel would slowly decline due
to lower reproductive outputs (Reynolds 1985; Gur-
nell 1996).
Our density estimates for Eastern Grey Squirrels
(2.5 individuals/ha) also were slightly higher than those
reported >45 years earlier (1.7 individuals/ha; Robin-
son and Cowan 1954), suggesting that the population
of Eastern Grey Squirrels in Stanley Park is relatively
stable, even if their range elsewhere has expanded
(Gonzales 1999, 2000). If the combined presence of
both Douglas’s and Eastern Grey squirrels in Stanley
Park accelerates the depletion of the autumn seed crop,
this might force a greater dependence on alternative
plant foods in the spring and summer, which Eastern
Grey Squirrels are better able to use because of their
larger body size (Ackerman and Weigl 1970; Steele
and Weigl 1993). An investigation into the feeding
behavior and space use of both species would provide
a better understanding of how Douglas’s and Eastern
Grey squirrels coexists in anthropogenic environments
such as urban parks. This study highlights the need for
more long-term population monitoring of the interac-
tion of the two squirrel species in areas where East-
ern Grey Squirrels were introduced.
Acknowledgments
Douglas Ransome and T. P. Sullivan helped design
the study and provided valuable insight, statistical help
and field equipment. Many thanks to Mike Mackintosh
(Wildlife Manager, Vancouver Parks Board) and Greg
Paris (Wildlife Manager, Greater Vancouver Regional
District) for permission and help with study area design.
J. N. M. Smith reviewed an earlier draft of this man-
uscript.
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Received 22 July 2004
Accepted 22 September 2005
Responses of Syrphids, Elaterids and Bees to Single-tree Selection
Harvesting in Algonquin Provincial Park, Ontario
Erica Nov!, HUME DouGLas!*, and WILLIAM J. CRINS?
'Biology Department, Trent University, Peterborough, Ontario K9J 7B8 Canada
*Present address: Biology Department, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6 Canada
3Planning and Research Section, Ontario Parks, Ontario Ministry of Natural Resources, 300 Water Street, Peterborough,
Ontario K9J 8M5 Canada
Nol, Erica, Hume Douglas, and William J. Crins. 2006. The response of syrphids, elaterids and bees to single-tree selection
harvesting in Algonquin Provincial Park, Ontario. Canadian Field-Naturalist 120(1): 15-21.
The species composition of hoverflies (Syrphidae), click beetles (Elateridae), and bees (Apoidea) was studied to determine
whether there was a positive response in these flower-seeking insect groups to gaps in the canopy created through single-
tree selection harvesting of Sugar Maple (Acer saccharum) and Yellow Birch (Betula alleghaniensis) in hardwood forests of
the Great Lakes-St. Lawrence forest region of Algonquin Provincial Park, Ontario. There were significantly more hoverflies
and bees collected in forest stands harvested within the previous five years than in wilderness zone (unharvested at least for
40 years) stands or stands harvested 15-20 years previously (old logged stands). Click beetles, especially Selatosomus pul-
cher (LeConte), were collected most often in old logged stands. Bees and click beetles were collected significantly later in
the season in logged than in wilderness zone stands. Malaise traps resulted in higher capture rates for syrphids than pan
traps, and only with these higher capture rates did we detect a significant increase in species richness in recently logged
stands over that in wilderness stands. Changes in the numbers and phenology of flower-visiting insects may impact on repro-
ductive success of flowering plants of the forest understory and deserves further study.
Key Words: single-tree selection logging, syrphids, hoverflies, Syrphidae, click beetles, Elateridae, bees, Apoidea, Algonquin
Provincial Park, hardwood forests.
Single-tree selection harvesting’ in shade-tolerant
hardwood forests creates cyclic perforations in the
canopy on a 20-25 year rotation and is hypothesized
to simulate conditions found under gap-phase dynam-
ics. This form of silviculture is practised in many parts
of the Great Lakes-St. Lawrence Forest Region in
Ontario and Quebec (Hunter 1990). Very little is known
about the impact on non-target species of removing
25% of the basal area of the forest through selection
cutting (Annand and Thompson 1997).
Particular invertebrate groups are most likely to
respond to changes in light conditions of the forest
floor as a result of reductions in canopy cover. Many
plants respond positively to increased light. As a result,
insects with adult stages attracted to conspicuous flow-
ers, also are expected to increase in abundance. These
include bees (Hymenoptera: Apoidea), hoverflies (Dip- .
tera: Syrphidae), a family of flies that contains impor-
tant flower visitors and pollinators (Waldbauer 1983;
Kula 1997; Goulson and Wright 1998; Gross 2001),
and many click beetles (Coleoptera: Elateridae), a less
well-known group of flower visitors (Johnson 2002).
Changes in other components of the forest as a result
of selection cutting may also benefit these insects.
Larvae of hoverflies of the subfamily Syrphinae feed
largely on aphids (Vockeroth 1992), and may benefit
from increased herbaceous growth associated with
canopy opening. Larvae of many elaterids (Johnson
2002) and some hoverflies of the subfamily Eristali-
nae (Vockeroth 1992) inhabit decaying wood. Their
response to logging may depend on the availability of
amounts, sizes and decay states of the standing and
fallen trees remaining after each phase of the logging
cycle. Soil-inhabiting elaterid larvae and ground-nest-
ing bees may be affected by changes in vegetation,
litter composition and depth, microclimate, and soil
disturbance (Penev 1992; Sugar et al. 1998). Distur-
bance caused by construction of sand logging roads
may result in greater colonization by psammophilous
bees and elaterids in a previously unsuitable forest
stand (Cane 1991).
We studied the impact of selection cutting on these
non-target invertebrates in Algonquin Park, Ontario.
We predicted positive density responses for all three
groups, either as a result of an increase in spring flora,
or other changes associated with logging. We also pre-
dicted that the presence of insects that visit flowers
would extend later into the growing season in logged
forests than in unlogged stands because of the greater
amount of light penetrating to the forest floor through-
out the spring and summer seasons. We tested these
predictions by comparing these insect communities in
Under selection harvesting, only some of the trees are cut in order to balance the volume of wood harvested with the amount
produced through new growth and to enhance biodiversity and ecological functions. This system of silviculture differs from
selective harvesting in which only the best timber trees are cut and the forest is gradually degraded.
15
16 THE CANADIAN FIELD-NATURALIST
stands cut 6 months to 3 years before our sampling,
15-20 years before, and in stands that have not been
subjected to logging for at least 40 years.
Study Area and Methods
The study was conducted in the summers of 1997
and 1998 in Algonquin Provincial Park, Ontario
(42°32'N, 78°36'W), as part of a larger study that was
examining impacts of logging on birds (Jobes et al.
2004) and ground beetles (Carabidae: Vance and Nol
2003). Forest stands were chosen randomly (within
the constraint that they be within 500 m of road access)
in wilderness designated zones (Algonquin Park Man-
agement Plan 2000), last logged at least 40 years prior
to survey), old logged (15-20 years post cutting), and
recently logged (6 months — 3 years post cutting) areas.
Canopy cover (estimated in July) ranged from 60-
70% in the wilderness zone to 25-35% in old cut and
recently cut stands, and was significantly higher in the
wilderness zone stands than in both the old logged
and recently-cut stands (Jobes et al. 2004). Other veg-
etation differences included a significant increase in
vegetation under 2 m and percent logging residue
cover (mostly tops of trees) in the recently cut stands
over that in the other two treatments, and a signifi-
cant increase in stem densities and saplings (2-5 m)
and subcanopy (5-10 m) vegetation in the old logged
treatments as compared to the other two (Jobes et al.
2004).
In 1997, we used a single Malaise trap set on a ridge
in one stand of each of the recently logged and wilder-
ness zone treatments. It was emptied at 1-week inter-
vals every other week from May to August. Thus, we
had no replication of stands for this year. In 1998, we
set yellow pan traps (21 cm? x 5 cm depth) filled with
water to a depth of 2-3 cm, and a small amount of
detergent in a total of 18 stands. These traps were set,
within stands, at 100 m intervals at permanent mark-
ers. We sampled eight stands in the wilderness zone,
seven in the recently logged areas, and three in the
old logged areas. Stands were separated by a minimum
of 200 m, but most stands were separated by > 3 km,
in stands consisting of at least 75% upland deciduous
forest (Vance and Nol 2003) so we assumed that they
were spatially independent. Each stand contained a
minimum of three traps. The sampling was uneven in
the treatments because of accessibility of stands and
because two of the initial old logged treatments were
logged in the winter of 1997-1998. These traps were
set from early May to late August, and emptied weekly
for three weeks in early, mid, and late summer periods,
for a total of nine trapping weeks. There were 69 inci-
dents of trap disturbance by mammals.
Pitfall traps were used to obtain activity-density esti-
mates for click beetles (Baars 1979). Pitfalls were con-
structed using | litre plastic 11.5 cm diameter contain-
ers lined with 500 mL plastic containers of the same
diameter (Spence and Niemela 1994). The traps were
dug into the ground so that the top of the upper contain-
Vol. 120
er was level with or slightly below the surface of the
soil. Each trap was filled with approximately 300 ml
of water, with a few drops of dish soap added to reduce
surface tension. Two pitfall traps were set at 4 m to
each side of the permanent stand markers, at the same
intervals as the pan traps. In 1997, pitfalls were set for
one week in each of May through August for a total
of four weeks. In 1998, pitfalls were set for the same
periods as the pan traps. A small number of traps (5%)
were disturbed in all treatments
All invertebrates were sorted initially to class, and
all but insects were discarded. Insects of the three target
groups were then sorted and identified with the use of
keys by Becker (1956, 1974); Brown (1934); Coovert
and Thompson (1977); Curran (1921, 1922, 1925,
1934, 1941); Curran and Fluke (1926); Dietrich (1945);
Fluke and Hull (1945); Fluke and Weems (1956); Hull
and Fluke (1950); Michener et al. 1994; Mitchell
(1960); Shannon (1939); Telford (1970); Vockeroth
and Thompson (1981); and Vockeroth (1992). Voucher
specimens are deposited in the Canadian National
Collection of Insects and Arthropods, Agriculture and
Agri-food Canada, Ottawa.
As we began sampling in all treatments at the same
time, we used the dates (1998 only) on which the tar-
get groups were captured as independent samples, for
our comparisons of phenology. These data were high-
ly skewed so we used the Kruskal-Wallis ANOVA to
test for significance among treatments. We were unable
to use stands as sampling units for any statistical com-
parisons because capture rates were too low. Thus, we
tested for significant differences among treatments in
species and numbers of individuals trapped, using chi-
square tests, with proportion of total trapping effort
among treatments used to calculate expected num-
bers. We report standardized residuals ((Observed —
Expected)/ VExpected) and their direction to show
which treatment had greater or less than the numbers
of individuals or species than expected based on our
sampling effort (Quinn and Keough 2002). For pit-
falls and pan traps we usually caught single individu-
als of the target species and these were spread across a
range of stands within that treatment, so that the as-
sumption of independence for this statistical test was
probably not violated (1.e., the samples are not biased
by many individuals caught in one particular stand and
treatment except in cases where noted below). Given
the low capture rate for any of our target groups we did
not calculate or analyse patterns of diversity.
To assess spring ephemeral abundance, we counted,
in | m* quadrats placed 5 m to the side of pitfall and pan
trap locations, all visible stems of spring-blooming forb
species during the first sampling period in May 1998,
before canopy leaf emergence.
Results
Each of the three target groups had substantially
higher captures per unit effort (combining sample
across all stands per treatment because of low num-
2006
bers captured) in at least one logged treatment than
in the wilderness zone treatment (Table 1).
Syrphids. In both years, with both pan and malaise trap-
ping methods, more species of syrphids were caught
in logged than in unlogged landscapes (Table 2). In
1997, eight species were found in wilderness zone
malaise traps and 25 species were trapped from the
recently logged stands with three species in common
between the two treatments. As Malaise traps were
opened for an equal number of days in the two stands,
we calculated expected numbers using the binomial
distribution, based on P = 0.5. Significantly more
NoL, DOUGLAS, AND CRINS: RESPONSES OF SYRPHIDS, ELATERIDS AND BEES
TABLE |. Catch per unit effort (x100) for all Syrphidae, Elateri
dae and Apoidea caught in pan traps in recently logged, old
logged, and wilderness stands in 1998, and combined catch
for pitfalls in 1997 and 1998.
Recently Old
Logged Logged Wilderness
Total pan trap days 1894 1040 2961
Total pitfall days 4796 3088 7938
Syrphidae (pan) 0.68 2.31 0.17
Elateridae (pan) 1.37 6.82 3.7]
Elateridae (pitfall) 0.27 1.49 0.30
Apoidea (pan) 3.22 2.11 1.28
TABLE 2. Species and numbers of individuals captured of Syrphidae from pan traps set in recently logged, old logged, and
wilderness stands in Algonquin Provincial Park using Malaise traps in 1997 and pan traps in 1998. ¥° statistic provided for
test of association between number of individuals and species and logging treatment. Standardized residuals give direction
and strength of deviation (Quinn and Keough 2002)
Species
Year
Sub-Family Syrphinae
Baccha elongata (Fabricius)
Dasysyrphus pauxillus (Williston)
Epistrophe nitidicollis (Meigen)
Eupeodes perplexus (Osburn)
Eupeodes americanus (Wiedemann) or pomus (Curran)
Melanostoma mellinum (Linnaeus)
Meliscaeva cinctella (Zetterstedt)
Parasyrphus semiinterruptus (Fluke)
Parasyrphus sp.
Platycheirus confusus (Curran)
Platycheirus obscurus (Say)
Sphaerophoria novaeangliae (Johnson)
Syrphus rectus (Osten Sacken)
Toxomerus geminatus (Say)
Sub-Family Eristalinae
Brachyopa notata (Osten Sacken)
Brachypalpus oarus (Walker)
Chalcosyrphus nemorum (Fabricius)
Cheilosia rita (Curran)
Cheilosia tristis (Loew)
Eristalis dimidiatus (Wiedemann)
Helophilus fasciatus (Walker)
Lejota aerea (Loew)
Lejota cyanea (Smith)
Neoascia distincta (Williston)
Pipiza femoralis (Loew)
Rhingia nasica (Say)
Sericomyia chrysotoxoides (Macquatt)
Sphegina brachygaster (Hull)
Sphegina flavomaculata (Malloch)
Sphegina keeniana (Williston)
Sphegina campanulata (Robertson)
Temnostoma balyras (Walker)
Volucella bombylans (Linnaeus)
Xylota quadrimaculata (Loew)
Xylota confusa (Shannon)
Number of individuals ¥7= 45.9, P < 0.001
Standardized residuals, individuals (1998 data only)
Number of species ¥7= 5.5, n.s.
Standardized residuals, species (1998 data only)
1997
11
Wilderness
1998
Old logged
1998
Recently logged
1997 1998
i)
bho
vs)
— Om W—
‘=
et emt KD ND
042
18 THE CANADIAN FIELD-NATURALIST
syrphid individuals and species were captured in the
Malaise trap set in the recently logged than in the
wilderness habitat (Binomial test, species: P < 0.0001,
individuals: P < 0.0001). In 1998, using pan trap catch-
es, two species, each consisting of one individual each,
were found only in the wilderness zone, whereas nine
species were found only in the recently logged land-
scape. Only six species were found in both years using
the two capture techniques. Significantly more individ-
uals were caught in 1998, in the old logged stands than
in either the recently logged or wilderness treatments
(Table 2), but this result was entirely due to a capture of
17 individuals of the species Melanostoma mellinum
on 14 May 1998 in the old logged treatment.
In 1997, we did not begin sampling until late May,
after leaf-out in the park. In 1998, there was a strong
bias in the collection times for all syrphids, with 16 of
22 collection dates for syrphids (73%) in May, prior
to leaf-out. There was, however, no significant differ-
ence among treatments in the collection dates of syr-
phids in 1998 (median sample dates (range): wilder-
ness: 29 May (26 May-6 August), n = 5 trap sample
dates; old logged: 21 May (14 May-2 July), n = 4;
recently logged: 27 May (9 May-24 June), n = 13,
Kruskal-Wallis 77 = 4.33, P =0.11).
Bees. Twelve species of bees were identified, with
five of these from wilderness and old logged stands and
all twelve in the recently logged stands (Table 3). All
but four species (excluding Andrena) were halictids.
For both number of species and number of individu-
als the recently logged stands had significantly more
bees than expected based on catch effort (Table 3).
All but 6 of the 57 dates on which we captured bees
were in May. Bees were caught at a significantly later
date in recently logged stands than in either old logged
or wilderness stands (median collection dates (range):
wilderness: 10 May (8-20 May), n = 18 trap sample
dates; old logged: 9 May (9 May-24 June), n = 13;
recently logged: 19 May (9 May-14 July), n = 26,
Kruskal-Wallis 77 = 13.8, P < 0.001).
Elateridae. In this group of insects, two species, Cteni-
cera triundulata and C. vulnerata were found only in
the wilderness stands, both in very low numbers (Table
4). The number of species among treatments did not
deviate from expected using the pan trap data but was
marginally higher than expected in the old logged
treatment using the combined 1997/1998 pitfall data
(combined years due to small sample sizes, y7 = 5.89,
P <0.06).
Numbers of individual click beetles were signifi-
cantly higher in the old logged habitat, and significant-
ly lower in the wilderness zone than expected based on
catch effort, using pan traps and pitfall traps (Table 4).
Elaterids were collected in 1998 from pan traps at
significantly later dates in old logged and recently
logged stands than in wilderness zone stands (median
date (range); wilderness zone: 13 May (8 May-5 June),
Vol. 120
‘n = 9; old logged: 28 May (8 May-9 July), n = 23:
recently logged: 19 May (9 May-2 July), n = 23; Krus-
kal-Wallis y? = 7.6, P= 0.02). Using 1998 pitfall data,
elaterids were sampled significantly later in recently
logged stands than in the other two treatments (wilder-
ness zone: 17 May (13 May-4 August), n = 20; old
logged: 17 May (14 May-3 July, n = 22; recently
logged: 24 June (12 May-24 June), n = 9; Kruskal-
Wallis 7? = 6.34, P = 0.04). The 1997 elaterid data
were too sparse for analysis.
Spring ephemerals. We recorded 15 species of spring
ephemerals in our quadrats (1998 only). The num-
bers of ephemerals found in all of the stands were
extremely variable (cv’s range from 156-165%).
Although the mean number of stems in the recently
logged stands was more than twice as large as this
number in wilderness zones, there was no significant
difference among treatments (recently logged: 59.7 +
16.7 / m’, n = 23 plots; old logged: 46.5 + 22.1 / m?,
n= 13 plots; wilderness zone: 26.5 + 18.8 /m*,n=18
plots; IB sn = (0.88, n.s.).
Discussion
Our results suggest that all three of groups of flower-
visiting insects were more abundant in one of the logged
habitat types than in the wilderness zone. This suggests
that these insect groups were positively affected by
single-tree selection harvesting as it is practised recent-
ly and/or how it was practised in the 1970s, when the
selection cuts were heavier than is currently accept-
able (Algonquin Forestry Authority, unpublished). This
result is not explained simply by greater numbers of
spring ephemerals, as, at least with our sampling
methodology, we found no significant difference among
treatments in the availability of flowers measured prior
to leaf-out. Differences were apparent in number of
individual insects, but, in general, not in insect species
richness, except in the case of the Malaise traps set in
1997, where both numbers of individuals and species
richness were higher in collections from the trap in the
recently logged stand. Given that this result is for one
trap only in each treatment, this result should be viewed
cautiously.
As many larval syrphids prey on Homoptera in
herbaceous vegetation and many adults feed on nectar
sources in this vegetation, the presence of much high-
er percent cover of herbaceous or soft woody vegeta-
tion (37 to 43% in old logged and wilderness stands
respectively, to 78% in recently logged stands, partic-
ularly Rubus idaeus L. var. strigosus (Michx), Jobes
1999) could explain the significant increase in syr-
phid abundance that we found through Malaise trap-
ping. An apparent increase also was noted in our pan
trapping results but not established statistically because
of the generally poor success of this method for cap-
turing syrphids in any of our stands. A similar positive
numerical response in syrphids (also not tested statis-
tically) was found in Poland’s polluted spruce forests,
2006
NoL, DOUGLAS, AND CRINS: RESPONSES OF SYRPHIDS. ELATERIDS AND BEES
19
TABLE 3. Species and Genera of bees collected from pan traps and numbers of individuals and minimum number of specie
from Algonquin Park, summer 1998, 122 individuals captured. Statistics and standardized residuals as in Table 2
Genus
Wilderness
Old logged Recent logged
Andrena'
Bombus perplexus Cresson
Bombus vagans Smith
Hylaeus elliptica (Kirby)
Lasioglossum admirandus (Sandhouse)
Lasioglossum divergens (Lovell)
Lasioglossum laevissimus (Smith)
Lasioglossum nymphaearum (Robertson)
Lasioglossum quebecensis (Crawford)
Lasioglossum rohweri (Ellis)
Lasioglossum versans (Lovell)
Lasioglossum zephyrus (Smith)
Osmia atriventris Cresson
Number of individuals x? = 21.2, P < 0.001
Standardized residuals, individuals
Minimum number of species y= 20.4, P < 0.001
Standardized residuals, species
36
Wr o
20
|
21
— ~
o
Nw — — WwW
Ww
to
' Not identified to species because of poor quality of specimens.
TABLE 4. Species and number of individuals of Elateridae collected from wilderness, old logged stands, and recently logged
stands in Algonquin Park in 1997 and 1998 using pan traps (individuals: y7= 175.1, P < 0.00; species: y? = 3.2. ns.) and pitfall
traps (individuals: y= 90.7, P < 0.001, species: y?= 5.9, n.s.). Analysis for pitfalls from two years combined). Standardized
residuals as in Table 2.
Wilderness
Old logged Recent logged
Species 98 Pan 97 Pit 98 Pit 98 Pan 97Pit 98 Pit 98Pan 97Pit 98 Pit
Agriotes collaris (LeConte) 2 | 5 6 l 3
Agriotes stabilis (LeConte) l
Athous acanthus (Say) |
Ctenicera hieroglyphica (Say) ((Say)) 1 2 Z, l
Ctenicera insidiosa (LeConte) l
Ctenicera resplendens (Eschscholtz) ] l
Ctenicera spinosa (LeConte) | l
Ctenicera triundulata (Randall) 2
Ctenicera vulnerata (LeConte) 1
Dalopius cognatus Brown l l
Dalopius fuscipes Brown l | l
Dalopius spp. (females) 2, 2 | 3 2
Limonius aeger LeConte 2
Limonius confusus LeConte > l l - l 2 9 a
Melanotus castanipes (Paykull) (Paykull) l
Neohypdonus tumescens (LeConte) | l 2
Selatosomus pulcher (LeConte) 2 10 59 2 28 10
Number of individuals 11 5 20 71 6 42 27 2 9
Standardized residuals, individuals -3.4 -2.6 11.8 8.6 -1.3 -3.]
Number of species 5 3 9 5 5 8 7 2 -
Standardized residuals, species =|? -0.8 1.1 22. 0.7 0.6
as an apparent result of drastic reductions in canopy
cover and greater herbaceous and grass undergrowth
(Kula 1997). Some species of at least two genera of
hoverflies that were found in the wilderness zone
(Melanostoma and Platycheirus) specialize on grami-
noid and other anemophilous pollen, enabling them
to be active later into the spring/summer when grami-
noids typically flower (Vockeroth 1992; Gilbert 1993).
However, none of the species captured is restricted to
flying in the spring.
Bees, like syrphids, were most abundant in the re-
cently logged areas. Unlike syrphids, larval bees are
fed pollen and thus are more wholly dependent on flo-
ral resources. Some bees also may be affected by the
20 THE CANADIAN FIELD-NATURALIST
availability of open flight paths present in the open
understory of the recently logged treatments, to make
pollen-gathering trips for brood provisioning, although
the effects of forest habitat on bee movement is prob-
ably species-specific, and for many species these move-
ment patterns are unknown (Kreyer et al. 2004). Wood-
land and edge ground nesting bees (the majority of
the species caught in the recently logged stands) are
thought to benefit from patches of disturbed habitat
associated with habitat fragmentation (Cane 1991) and
thus, may benefit from the building of sand logging
roads. Halictid bees can be major components of pol-
linator communities in other forest types (Pascarella
1997). Their role in northern hardwood forests has not
been adequately assessed.
Elaterids were most abundant in the old logged habi-
tats, using two methods of sampling, indicating posi-
tive responses to the much higher cover of the sapling
(2-10 m) layers found in this treatment (Jobes et al.
2004). The most abundant species, Selatosomus pul-
cher, has been trapped in open field habitats (Boiteau
et al. 2000). This species is morphologically nearly
indistinguishable from the European species C. cru-
ciatus, a species whose larvae develop in the sandy
and moist soils in forest and forest edge (Leseigneur
1972). The higher abundance of elaterids in the old
logged treatment areas may be due largely to this
species’ positive response to the edge-like conditions
present in this treatment. The lower numbers of ela-
terids in the recently logged stands may indicate the
absence of colonization because of the short time
frame. Analagous results are suggested in a study of
herbaceous plants in recently logged White Pine (Pinus
strobus) stands in Algonquin. Plants in those stands
were most similar to plants in unlogged stands pre-
sumably because there had been insufficient time for
changes in community assemblages after the logging
(Kingsley 1998).
As little is known about the role of adult food avail-
ability in the distribution of elaterids, the near signifi-
cant positive effect on species richness may also be
due to the greater availability of soil invertebrates or
plant material suitable for larval food produced by a
four-fold increase in the number of stumps in old logged
versus either wilderness or recently logged stands
(Jobes 1999).
Of the two elaterids caught only in the wilderness
treatment, only one may be an old growth dependant
species. Ctenicera vulnerata (1 specimen caught) has
rarely been collected, and has not been reported pre-
viously from Ontario. The other, Crenicera triundulata
(2 specimens) cannot be considered old growth-restrict-
ed, as it has been collected in moderate numbers in an
apparently healthy spruce (Picea sp.) in an old field
by HD.
We found very few bees, syrphids or click beetles in
our samples from the wilderness zone stands, although
we expected to find some, at least early in the season
when the canopy had not yet leafed out or of species
Vol. 120
that utilized downed woody debris (e.g., Chalcosyr-
phus, Xylota, Brachyopa). The reason for their paucity
in the wilderness zone is unclear but deserves further
study.
The numerical increase and changes in phenology of
flower-associated insect species suggests the potential
for greater pollinator efficiency in logged forests, and
hence greater reproductive success, through greater
seed set of flowers. A more detailed examination of
species-specific patterns of habitat use by pollinators
and their functional role in hardwood forests is worthy
of future research (e.g., Sheffield et al. 2003; Kreyer
et al. 2004).
Acknowledgments
We thank, in particular, Christine Vance, who tire-
lessly emptied pan traps in 1998 for this project. We
also thank Andrew Jobes for collecting some of the
data on spring ephemeral abundance and providing the
canopy cover information. We thank E. C. Becker for
verifying elaterid identifications, and R. Tuckerman,
J. Grixti and T. Romankova for identification of bees.
D. Voigt and E. Addison provided the stimulus for this
project through their project on disturbance of ecosys-
tems in Algonquin Park. We thank them for their exten-
sive logistic support. The Algonquin Forestry Authority
(AFA) also provided details on the history of the stands.
N. Quinn (Algonquin Park) kindly provided funding
for our work on birds and beetles in the park. This proj-
ect was also funded by NSERC (Canada) grants to E.
Nol.
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Received 14 July 2004
Accepted 7 February 2006
New Nesting Records of the Le Conte’s Sparrow, Ammospiza leconteii,
from Northeastern Ontario, with Some Notes on Nesting Behaviour
MICHAEL PATRIKEEV
Ontario Parks, 199 Larch Street, Suite 404, Sudbury, Ontario P3E 5P9 Canada
Present address: 3 Helen Street, Dundas, Ontario L9H 1N2 Canada; e-mail: mpatrikeev@hotmail.com
Patrikeev, Michael. 2006. New nesting records of the Le Conte’s Sparrow, Ammospiza leconteii, from northeastern Ontario,
with some notes on nesting behaviour. Canadian Field-Naturalist 120(1): 22-26.
Le Conte’s Sparrow is sparsely distributed through northeastern Ontario with no confirmed records from Algoma District and
eastern Lake Superior. Two nests were found in open poor fen between Wawa and Hawk Junction in central Algoma District
in 1999. Notes on behaviour, nests, nestling development and feeding effort were taken during 9 hours and 38 minutes at one
nest over three days.
Key Words: Le Conte’s Sparrow, Ammospiza leconteii, nesting, northeastern Ontario.
Le Conte’s Sparrow (Ammospiza leconteii) is dis-
tributed through grasslands and wetlands of central and
southern Canada east to Ontario and Quebec, and in
the north-central United States. “It is one of the least
known North American sparrows and any new infor-
mation on this species may be of interest” (Lowther
1996). A cryptic species, it nests in thick clumps of
dead grass, and its nests are difficult to find. Only about
50 nests had been recorded range-wide by the early
1960s (Walkinshaw 1968) and few have been docu-
mented since then (Lowther 1996). Only nine nests of
this species had been found in Ontario prior to 1999
(Peck and James 1987; Sandilands and Campbell 1987;
G. Peck, personal communication). In Ontario, Le
Conte’s Sparrow has been reported in widely separated
areas in the forest zone with the greatest concentrations
in the James Bay Lowland and northwestern Ontario
(Sandilands and Campbell 1987). During the first
Ontario Breeding Bird Atlas (1981-1985) this species
was found in only 20 of 137 UTM blocks (100 x 100
km) and nesting was confirmed in only four blocks
(Sandilands and Campbell 1987). None of the first Atlas
records were from Algoma District, although Le Conte’s
Sparrow was considered a rare spring species in the
vicinity of Wawa (Ontario Ministry of Natural Re-
sources 1990). In 2001-2004, surveyors for the second
Ontario Breeding Bird Atlas reported probable nesting
in two blocks (Sault Ste. Marie and St. Joseph Island),
and possible nesting in five additional blocks in the
vicinity of Sault Ste. Marie and Thessalon at the south
end of Algoma District (Ontario Breeding Bird Atlas
website: http://www.birdsontario.org/atlas).
Here I describe two new nests from central Algoma
District, northeastern Ontario. A small group with 2-3
singing males was found in an upland wetland (5.8 ha,
48°02'42.24N", 084°34'01.30W") in Esquega Township
east of Wawa and southwest of Hawk Junction (Fig-
ure 1). The wetland (329 m above sea level) encom-
passed elements of several wetland types gradually
changing from sphagnum bog with few stunted Black
Spruce (Picea mariana) to open poor fen and then to
a meadow marsh (Harris et al. 1996).
Le Conte’s Sparrows inhabited approximately 2 ha
of open poor fen in this wetland (see Figure 2). The
wetland was bordered by alder thickets (Speckled
Alder, Alnus incana) with a few dead Balsam Poplars
(Populus balsamifera) and by mixed forest dominat-
ed by Trembling Aspen (P. tremuloides) and White
Birch (Betula papyrifera). Water regime of the wet-
land changed from late May through July 1999. The
lower parts (mostly meadow marsh section) were flood-
ed in late May (up to 0.5 m deep), but water gradually —
receded by late June. The area remained damp, but no
standing water remained (natural flow in this wetland
was likely disrupted years ago when it was transsected
by Highway 101). The area occupied by Le Conte’s
Sparrows had a ground cover of sphagnum moss and
was dominated by two species of sedge: Few-seeded
Sedge (Carex oligosperma) and Aw]-fruited Sedge (C.
stipata); Canada Bluejoint (Calamagrostis canaden-
sis) was also present.
Singing males were heard at the wetland on 21 May
1999, and a display flight observed on 27 May. A flight
song is described by Murray (1969). I also observed
a “spy flight’: a sparrow would rise from the grass and
fly in a low arch quickly dropping back into the cover.
Le Conte’s Sparrow likely uses this flight to deter-
mine precise position of an approaching intruder. No
song or audible call was uttered during such flight.
Two nests were found in the wetland, on 13 June
and 17 July 1999, respectively. The first nest contained
five eggs (Figure 3), but was empty on 17 June. The eggs
matched a description of Le Conte’s Sparrow eggs
from Walkinshaw (1968). The second nest contained
five newly-hatched young on 17 July and still contained _
five young on 23 July when the young would be seven
days old. On day seven, the young were in cinnamon-_|
coloured plumage with dark streaks on the back, dark |
De
2006
_Manitoba
omens
OS eg 44 at
PATRIKEEV: NESTING OF LE CONTE’S SPARROW IN NORTHEASTERN ONTARIO 23
Hudson Bay
( co
a
—S %
a oe Quebec
Poff Xe
| Algoma District 7 |
FIGURE |. Position of Wawa and Algoma District in Ontario.
grey crowns and cheeks, tan median crown-stripe and
black shafts. The median crown-stripe was seen in at
least some chicks (Figure 4). Bill colour changed from
fleshy on days two-three to grey-pink on day seven.
Gape was bright flesh red on day seven. According to
Gollop et al. (1966), the young leave the nest at the
age of seven days.
The two nests were approximately 150 m apart and
might have belonged to the same pair. Both nests were
built of dried stems, and lined with fine dried stems of
sedge and grass. Outer rim of both nests was loosely
built. The first nest was well hidden under and inside
a clump of dried graminoids and could not be seen from
above. The second nest was attached to dried and liv-
ing sedge stems and a small Leatherleaf (Chamaeda-
phne calyculata) under a loose cover of sedges and
grasses. Both nests were 5-6 cm above the wet ground.
Observations totaling 9 hours and 38 minutes were
made at the second nest from a blind (through a tele-
photo lens) during late afternoon-early evening hours
on 19 July (1610-2045 hours), 21 July (1800-2110) and
22 July (1812-2005). All observations were conducted
during calm fair weather. The pair at the nest under
observation consisted of a slightly larger brighter male,
and a scruffier duller female. Le Conte’s Sparrows nev-
er flew directly to and from the nest, but landed some
distance away and walked (hopped) to it, fed the young
and walked away before taking on the wing. Also both
birds were never present at the nest at the same time
during the observation period. A total of 80 feeding
visits was recorded. The birds were bringing food every
6.6 minutes on average (SD + 5.2, Standard error 0.6).
Lowther (1996) reported | feeding/minute at a nest con-
taining a cowbird. Gender of feeding birds was as-
sumed on 55 occasions. The male fed the young on
average every 14.1 minutes (SD + 13.5) and the female
every 13.5 minutes (SD + 13.5). No differences be-
tween sexes were detected in feeding efforts (P = 0.50).
Intervals between feedings became shorter as the young
were growing: every 7.9 minutes + 5.9 on 19 July,
5.7 + 4.6 on 21 July and 4.9 + 3.0 on 22 July (Kruskal-
Wallis one way ANOVA: P = 0.07).
24 THE CANADIAN FIELD-NATURALIST Vol. 120
FIGURE 2. Habitat of Le Conte’s Sparrow in Esquega Township: this wetland encompassed elements of several wetland
types gradually changing from a sphagnum bog to open poor fen and then to a meadow marsh. Le Conte’s Sparrows
inhabited about 2 ha of open poor fen. Photo by Michael Patrikeev.
FiGurE 3. Nest of Le Conte’s Sparrow with five eggs. The nest was built of dead stems, and lined with fine stems of sedge
and grass. It was well hidden under and inside a clump of dead graminoids and could not be seen from above. Photo
by Michael Patrikeev. 13 June 1999.
2006
PATRIKEEV: NESTING OF LE CONTE’S SPARROW IN NORTHEASTERN ONTARIO
Nm
Ww
Ficure 4. Nestlings of Le Conte’s Sparrow seven days old. Photo by Michael Patrikeey. 23 July 1999.
Lowther (1996) noted that hardly any information
was available on this species’ diet. Although in many
cases food brought to the young was too small to iden-
tify through a telephoto lens, I observed small green
caterpillars, a white moth, a spider, a small sphinx cater-
pillar and two grasshopper abdomens. The last three
items were brought by the female who tried to feed
them to the small young. The content of the sphinx
caterpillar was eventually squeezed into the young’s
mouths, and one grasshopper abdomen was swallowed
by the female itself after futile attempts to feed it to
the young.
I did not observe wetlands similar to the one des-
cribed above in the Wawa area (Michipicoten and Es-
quega townships, and the adjacent parts of Lake Super-
ior Provincial Park) in 1999-2000 and no Le Conte’s
Sparrows were detected in other visited wetlands. How-
ever, this species likely occurs in low densities through-
out Algoma District where suitable habitat is available.
Acknowledgments
I am grateful to Carol Dersch (Lake Superior
Provincial Park) and Kim Taylor (North-Eastern Sci-
ence and Technology, Ontario Ministry of Natural Re-
sources) for help with plant identification, to George
Peck (Royal Ontario Museum) who provided me with
information on the number of confirmed nesting rec-
ords of this species in Ontario, to Richard Knapton
(University of Alberta, Edmonton), John C. Eitniear
(San Antonio, Texas), A. J. Erskine and an anonymous
reviewer for useful comments on this manuscript.
Literature Cited
Gollop, J. B., J. A. Slimmon, and R. V. Folker. 1966. Some
1965 bird records for the Saskatoon district. Blue Jay 24:
76-78.
Harris, A. G., S. C. McMurray, P. W. C. Uhlig, J. K.
Jeglum, R. F. Foster, and G. D. Racey. 1996. Field guide
to the wetland ecosystem classification for northwestern
Ontario. Ontario Ministry of Natural Resources, Northwest
Science and Technology. Thunder Bay, Ontario. Ontario
Field Guide FG-01. 74 pages + Appendices.
Lowther, P. E. 1996. Le Conte’s Sparrow (Ammodramus
leconteii). In The Birds of North America (224). Edited by
A. Poole and F. Gill. The Academy of Natural Sciences,
Philadelphia, Pennsylvania, and The American Ornithol-
ogists’ Union, Washington, D.C.
Murray, B. G., Jr. 1969. A comparative study of Le Conte's
and Sharp-tailed sparrows. Auk 86: 199-231.
Ontario Ministry of Natural Resources. 1990. Checklist
of Birds/Liste des oiseaux. Lake Superior Provincial Park
and adjacent areas/Pare provincial du lac Supérieur et les
régions environnantes. Ministry of Natural Resources/Min-
istére des Richesses naturelles. MNR Number 3752. ISBN
0-7729-6809-8. 24 pages.
Peck, G. K., and R. D. James. 1987. Breeding Birds of On-
tario: Nidiology and Distribution. Volume 2: Passerines.
Life Sciences Miscellaneous Publication, Royal Ontario
Museum, Toronto, Ontario. xi + 387 pages.
Sandilands, A., and C. Campbell. 1987. Le Conte’s Spar-
row/Bruant de Le Conte Ammodramus leconteii. Pages
26 THE CANADIAN FIELD-NATURALIST Vol. 120
454-455 in Atlas of the Breeding Birds of Ontario. Com-
piled by M. D. Cadman, P. F. J. Eagles, and F. M. Hellein-
er. Federation of Ontario Naturalists and Long Point Bird
Observatory. University of Waterloo Press.
Walkinshaw, L. H. 1968. Passerherbulus caudacutus (Lath-
am). Le Conte’s Sparrow. Pages 765— 776 in Life histories
of North American cardinals, grosbeaks, buntings, towhees,
finches, sparrows, and allies. Edited by O. L. Austin, Jr.
United States National Museum Bulletin 237, Part 2.
Received 21 October 2004
Accepted 14 September 2005
Conditions for Sexual Interactions
RICHARD P. THIEL
Canadian Field-Naturalist 120(1): 27—30.
of the two species.
Wolves and Coyotes can hybridize in captivity
(Young and Goldman 1944; Silver and Silver 1969;
Kolenosky 1971; Schmitz and Kolenosky 1985a,
1985b). Previously it was assumed that behavioral bar-
riers prevented hybridization between Canis species
(Wolf x Coyote) in the wild. Evidence of such crosses
in the wild remained elusive until Lehman et al. (1991)
reported on the presence of Coyote genes in Wolf mito-
chondrial DNA material from the Upper Great Lakes
region of North America probably from crosses be-
tween female Coyotes and male Wolves. This stimulat-
ed further investigations (Roy et al. 1994; Pilgrim et al.
1998; Boyd et al. 2001; Wayne et al. 1995; among oth-
ers), confirming the introgression of Coyote genes in
_ Wolf mtDNA in a region extending from the western
_ Great Lakes east to Quebec.
_ Confounding these findings, recent DNA studies
_ among Canis in eastern Ontario and northeastern Unit-
ed States hypothesize the existence of a unique and
putative Eastern (Timber) Wolf (Canis lycaon (Wilson
et al. 2000, 2003) while other researchers argue the
possibility that the Eastern Wolf is the Red Wolf (Canis
rufus), or some type of a Wolf (Canis lupus) x Coyote
hybrid (Schmitz and Kolenosky 1985a; Wayne et al.
1995; Nowak 2003; Wayne and Vila 2003; Phillips et al.
2003; Sears et al. 2003; among others).
It is speculated that the tumultuous ecological up-
, heavals caused by Eurasian settlement of North Ameri-
ca disrupted Canis communities creating conditions
| facilitating hybridization (Wayne et al. 1995; among
others). Forest fragmentation and heavy exploitation of
i
|
i
| Canis populations are believed responsible for current
_ hybridization between various Canis communities in
southeastern Ontario (Sears et al. 2003; Theberge and
| Theberge 2004).
_ The question remains under what environmental —
.
behavioral circumstances did (do) unions between
Coyotes and Wolves occur, given the usual aggressive
|
1
Between Wild Grey Wolves,
Canis lupus, and Coyotes, Canis latrans
Wisconsin Department of Natural Resources, Sandhill Wildlife Area, Box 156, Babcock, Wisconsin 54413 USA
Thiel, Richard P. 2006. Conditions for sexual interactions between wild Grey Wolves, Canis lupus, and Coyotes, Canis latrans.
| Genetic evidence for the hybridization of wild Grey Wolves and Coyotes was first reported by Lehmann et al (1991). Subse-
quent genetic and landscape-environmental analyses have attempted to grasp the extent of Wolf-Coyote crosses in North
America. Since Wolves are normally territorial and thus aggressive towards Coyotes, hybridization events remain rare, not
withstanding the taxonomic debates regarding Canis in eastern Ontario. In this paper I report on amicable interactions between
Wolves and Coyotes observed in Wisconsin in recent decades and discuss circumstances that may lead to pairing between individuals
Key Words: Grey Wolf, Canis lupus, Coyote, Canis latrans, interactions, hybridization, Wisconsin, Upper Great Lakes region.
tendencies of Wolves to Coyotes? Lehman et al. (1991)
believed that initial hybridization most likely occurred
when young, male Wolves dispersed into an area with
an endemic Coyote population where female Wolves
were rare. I report encounters between Coyotes and
Wolves in Wisconsin that support this theory during a
period when Wolves were re-colonizing landscapes
occupied by Coyotes. Because of the ongoing debate
over Canis affinities in the east, I confine my comments
to the Upper Great Lakes region of North America.
Methods
It is generally accepted that two taxonomically dis-
tinct wild canid species occur in Wisconsin: Grey
Wolves (Canis lupus) and Coyotes (Canis latrans).
Natural re-colonization of Wolves in Wisconsin began
in the mid 1970s from expansion of the Minnesota
population (Mech and Nowak 1981; Thiel and Welch
1981). Wolf monitoring has been conducted since 1979
in Wisconsin involving techniques that include a com-
bination of winter track surveys, aerial surveillance of
radio-collared Wolves, summer howl surveys, and col-
lection of reports by government employees and citi-
~ zens (Wydeven et al. 1995). The following cases of
Wolf-Coyote interactions are derived from visual obser-
vations from both ground and aerial surveillance, and
from interpretation of trails in snow.
Results
Florence County
Trails, raise-leg urinations (RLUs) and radio
telemetry data indicated that three lone male Wolves
occupied home ranges near one another in western Flo-
rence County (45°50'N, 88°50'W) in northeastern Wis-
consin between 1982 and 1984. At that time 17 — 25
Wolves lived in 2-4 packs along the Minnesota-Wis-
consin border (46°15'N, 92°00'W), and in two north-
central Wisconsin packs, 150 km southeast of the border
Deh
28 THE CANADIAN FIELD-NATURALIST
area (45°30'N, 90°00'W). The Florence County Wolves
were 100 km east of the two north-central packs. One
of these Wolves, male 077, was captured and collared,
and aerially radio-monitored about once-weekly for
250 days between 6 May 1983 and 11 January 1984,
occupying a 44 km* home range.
Several citizen and US Forest Service reports in-
volved sightings of a Coyote trailing a lone Wolf in
the areas occupied by the lone male Wolves. Wolf Pro-
ject personnel also observed snow sign indicating that
lone Wolves and Coyotes interacted. On 17 February
1982, Wolf Project technician, Larry Prenn, and I fol-
lowed the trail of a Coyote and Wolf over 3 km. We
were not able to ascertain the timing of each canid’s
travel, but based on highly convoluted trail maneuvers,
it appeared they were traveling together. The Wolf was
likely a male (RLUing) and the Coyote was a female
(squat urinations, including one with blood).
On 6 December 1983, while homing in on the signal
of Wolf 077 using a Cesna 180 fixed-wing aircraft, pilot
Dan Doberstein and I saw a pair of canids lying within
15- 20 m on the ice of Halsey Lake. One appeared 2 or
3 times larger than the other. After determining that nei-
ther was Wolf 077, we began a slow descent. When
approximately 200 m above them, the smaller canid
stood up and quickly ran off into thick lowland conifer
cover. We identified it as a Coyote. As it disappeared
into cover, a third, larger canid ran from shore to where
the second canid was still lying. On our third pass we
approached within 10 m of these canids; the second one
was larger, and we agreed they were both Wolves.
Shortly thereafter we located Wolf 077 male 9.6 km to
the east.
The following morning we attempted to inspect their
trails, but high winds overnight and thin ice made inter-
pretation challenging. We found outlines of two large
tracks corresponding in size to Wolf, and one small
Coyote-sized track. No evidence of a kill-site was found
that might have attracted the two species.
Wood County
In each of seven winters between 1995 and 2002
high school students under my supervision followed the
snow trails of a non-collared male Wolf, 501 that had
escaped into the 36 km? Sandhill Wildlife Area (SWA)
in Wood County (44°17'N, 90°10'W) (Thiel 2000; Thiel
unpublished data) in May 1995. This is a deer research
facility enclosed by 3 m tall x 29.5 km long fence. At
that time, Wolves were colonizing this portion of west-
central Wisconsin. In winter, 1996-1997, a territorial
pair of Wolves colonized the area surrounding SWA. In
January 1998, a pair of Wolves believed to be yearling
siblings, entered SWA, and subsequent snowfall pre-
vented them from digging out beneath the fence and
leaving. Wolf 501 paired with one of these Wolves, a
female based on bloody urine discovered on 20 Janu-
ary 1998. The third Wolf thereafter roamed the area
as a loner.
Vol. 120
In March a 6.5 year-old radioed Coyote was killed
by Wolves within SWA. Likely, Wolf 501 male was in-
volved in its death. Wolf 501 was last observed with
another Wolf in October 1998. In four additional win-
ters of snow tracking, Wolf 501 was the only Wolf de-
tected within SWA.
On 22 January 2001, High School students reported
trailing Wolf 501 male and encountered where the Wolf
had physically interacted with a Coyote, resulting in
an injury to the Wolf’s right front foot. On 5 February
several students and I encountered Wolf 501 male’s
trail where it intersected a Coyote’s trail, heading in
the same direction. In backtracking we determined the
Coyote was a proestrus female based on bloody urine.
The mutual trail led toa 5 m x 5 m site completely
padded down with Wolf and Coyote tracks. Numerous
small droplets of blood and several tufts of canid fur
were found in the padded-down site. Genetic testing to
ascertain the species leaving the blood and fur was cost-
prohibitive. By carefully back-trailing and following the
tracks of both the Coyote and Wolf 501 male as they
left the site we established the Wolf had been injured
by the proestrus Coyote at the padded down site.
The Coyote and Wolf walked on the same trail for
approximately 1 km beyond the padded down site be-
fore separating. Bloody Coyote urine was noted three
times along this trail, and the Wolf inspected each. At
one spot along the trail the Coyote and Wolf sat down
within 1.5 m of each other. We followed the Wolf an
additional 3.2 km after their trails separated. The Wolf
RLU’d 8 times, and in each of five beds we found the
imprint of his bloody limb.
It was evident that both canids were present together
and spent considerable time at the padded down site.
Since we found no evidence of prey remains, and since
Wolf 501 male did not react aggressively when injured
by the proestrus Coyote we concluded that his injury
was caused by unwanted sexual advances.
At approximately 0830 (CDT) on 22 June 2001,
amateur wildlife photographer, Rudi Van Stedum, saw
male Wolf 501 male standing on rock talus in a forest
in SWA at a distance of approximately 50 m. The Wolf
was watching her as she idled her car along a woods
trail. Simultaneously Van Stedum saw movement ob-
liquely behind and left of the Wolf. A Coyote appro-
ached but retreated when it became aware of her car.
Meanwhile, the Wolf turned and left the rock, facing
her. At that point, the Coyote again approached the
Wolf from behind and thrust its muzzle forcefully into
the left rump of the Wolf. A yelp from the Wolf was
clearly audible while the Wolf’s gaze remained fixed
on Van Stedum. The Coyote stepped back, and both
stood still for a moment before the Wolf walked a ways
off and turned momentarily to look at her. As the Wolf
began slowly walking away, the Coyote approached it
from behind and jabbed it a second time in its left hip
with its muzzle. The Wolf yelped again. Both then
trailed off into the undergrowth.
2006 THIEL: CONDITIONS FOR SEXUAL INTERACTIONS BETWEEN GREY WOLVES AND COYOTES 29
Wildlife biologist Wayne Hall and I inspected Van
Stedum’s photographs, and while they were taken in
poor light conditions and no single frame showed both
the Wolf and Coyote, the position of trees in the
background of all photographs attested to the accuracy
of her observation. Wolf 501 male roamed SWA alone
in winter 2001-2002 and disappeared sometime after
October 2002. No evidence of hybrids materialized
subsequent to the observations in 2001.
Discussion
Coyote mtDNA in Wolves was calculated as the
result of six hybridization episodes between male
Wolves and female Coyotes (Lehman et al. 1991);
Wayne and Vila (2003) speculated that, “... female
wolves and male coyotes are more closely matched in
size, and thus may also be more likely to mate...”, but
they further noted that no Coyotes sampled had Wolf-
like haplotypes.
Several researchers have speculated that male Wolf
dispersers provide the potential pool for male Wolf x
female Coyote unions (Lehman et al. 1991; Wayne and
Vila 2003; among others). While probably true, this is
not necessarily because male Wolves disperse beyond
the edge of Wolf range, as is implied. During winter
1983-1984, Michigan and Wisconsin DNR biologists
discovered several lone male Wolves living in a several
thousand square kilometer area. Some of these Wolves,
like Wolf 077, seemed to have home ranges, as evi-
denced by RLU’ ing (R. Thiel and J. Hammill, unpub-
lished notes). They likely originated from at least 100
km away where the nearest known packs then existed.
But female Wolves also dispersed to this area. In sum-
mer-fall 1986 radio-collared female Wolf 035 dispersed
from Douglas County, Wisconsin (46°15'N, 92°00'W)
277 km to Iron County, Michigan (46°20'N, 88°59'W)
and settled into a 174 km” home range, becoming the
first known Wolf to reach upper Michigan since their
demise in the late 1950s (Thiel 1988). The first breed-
ing Wolf pack in upper Michigan would establish itself
within this same area in 1990-1991 (J. Hammill, per-
sonal communication, | March 2004), implying that
other female Wolves also dispersed to this region. The
male Wolf x female Coyote hybridization hypothesis
is therefore not explained in our region by any superior
dispersal advantage displayed by male Wolves.
Because of their size, Wolves can easily overpower
and kill Coyotes. Ballard et al. (2003: page 267, Table
10.4) summarized Wolf-Coyote interactions in Yellow-
stone National Park and noted that 11 percent of inter-
actions resulted in killed Coyotes. In fatal encounters,
the number of Wolves exceeded the number of Coy-
otes. Wisconsin biologists have also encountered aggres-
Sive interactions between Wolves and Coyotes, includ-
ing killings of Coyotes. Killings involved superior
numbers of Wolves (R. Thiel, unpublished data). In
contrast, the few observations of single Wolf and Coy-
ote encounters are not known to have resulted in
killings. In the fall of 1981 pilot Jim Dienst! and I
watched dispersing lone male Wolf 023 chase a Coyote
from a White-tailed Deer (Odocoileus virginianus)
fawn kill, but made no attempt to catch or kill it. SWA
male Wolf 501 likely killed at least one Coyote when
paired with a female Wolf. Four years later, this same
male Wolf - who remained a loner in those four years
consorted with one or more Coyotes on at least three
occasions; one case involving a proestrus female Coy-
ote.
As in the Wisconsin cases reported here, Michigan
DNR biologist, J. Hammill (personal communication,
1 March 2004), witnessed two separate incidents where
a single Wolf and Coyote apparently traveled and bed-
ded down together during the early years of Wolf re-
colonization.
Within Wolf range, owing to the aggressive stance
pack Wolves normally display towards foreign Wolves
and Coyotes (Arjo and Pletscher 1999; Ballard et al.
2003), hybridization events must be exceedingly rare.
Observations reported here indicate that single Wolves
are capable of genial behavioral interactions with Coy-
otes. Further, they suggest a potential for sexual ad-
vances of male Wolves towards female Coyotes, when
and where male Wolves lack access to female Wolves.
Unions between male Wolves and female Coyotes
probably remain localized (however, note Schmitz and
Kolenosky 1985a; Sears et al. 2003; and Theberge and
Theberge 2004), and likely occur in areas of exceed-
ingly low Wolf densities where access to female Coy-
otes by single male Wolves far exceeds access to
female Wolves. Under such circumstances, hybridiza-
tion is at least possible, and then depends on the behav-
ioral nuances of these canids as individuals.
Acknowledgments
I thank L. D. Mech, United States Geological Sur-
vey, and A. Wydeven, Wisconsin DNR, for comments
on an earlier draft of this manuscript.
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Received 22 March 2004
Accepted 7 November 2005
Multiple Scale Den Site Selection by Swift Foxes, Vulpes velox,
in Southeastern Colorado
ANN M. KiITCHEN!, Eric M. GESE?, and SARAH G. Lupts!
' Department of Forest, Range and Wildlife Sciences, Utah State University, Logan, Utah 84322-5230 USA. Present address
National Zoological Park, Smithsonian Institution, Washington DC 20013-7012 USA. Corresponding author: Henderson
An@si.edu
7 U.S. Department of Agriculture, Wildlife Services, National Wildlife Research Center, Department of Forest, Range and
Wildlife Sciences, Utah State University, Logan, Utah 84322-5230 USA.
'
Kitchen, Ann M.., Eric M. Gese, and Sarah G. Lupis. 2006. Multiple scale den site selection by Swift Foxes, Vulpes velox, in
| southeastern Colorado. Canadian Field Naturalist 120(1): 31-38.
Predation by Coyotes (Canis latrans) is a major source of mortality in Swift Fox (Vulpes velox) populations. Year-round den
use by Swift Foxes is likely to be a predator avoidance strategy. Due to the importance of denning to Swift Fox ecology, we
recorded den site selection of Swift Foxes in southeastern Colorado. Den site selection was recorded at two scales: micro-
habitat characteristics at the den and den placement within the home range. The number of den entrances, height and width
of each entrance, aspect, hill position, slope, percent rock in soil, vegetative cover, and horizontal foliar density of 42 Swift
Fox dens were examined during December 1999 — April 2000. This was compared to the same microhabitat characteristics
at 42 random sites within Swift Fox home ranges to determine if Swift Foxes were using site characteristics according to
their availability. Our results indicated that Swift Foxes were not highly selective of den sites based on the microhabitat
characteristics evaluated in this study, although Swift Foxes selected areas of intermediate rock percentages. In addition,
Swift Foxes were radio-tracked throughout the sample period and the location and frequency of use of known dens were
recorded. Within the core area of home ranges, Swift Foxes used more dens (mean = 3.51 + 1.70 (SD)), and had a higher
frequency of use of dens (mean = 8.20 + 6.01) than in the mid-range area (number of dens, mean = 0.90 + 0.94; frequency,
mean = 1.27 + 2.12) and the boundary area (number of dens, mean = 0.34 + 0.53; frequency, mean = 0.45 + 0.93) of the
home range. We discuss our results in terms of the importance of dens in facilitating escape from Coyotes. These results
illustrate the need for examining den site selection at multiple scales to determine all selection factors, and to provide infor-
‘mation useful for recovery and management efforts for this species.
Key Words: Swift Fox, Vulpes velox, denning, habitat selection, home range use, Colorado.
Introduction
The Swift Fox (Vulpes velox) is native to the short
and mid-grass prairies of North America. The current
range of the Swift Fox is much reduced from the his-
toric range, and the species was formerly a candidate
for endangered species listing (United States Fish and
Wildlife Service 1996). The Swift Fox has been local-
ly extirpated in the northern prairies in Canada, North
Dakota, and much of South Dakota (Scott-Brown et al.
1987). One of the principal causes of Swift Fox pop-
ulation declines across much of the historical range is
a reduction in suitable habitat (Hillman and Sharps
1978).
__ The combined availability and distribution of suitable
_den sites and escape holes is likely a habitat compo-
nent for Swift Foxes (Egoscue 1979; Rongstad et al.
1989; Herrero et al. 1991). Swift Foxes were referred
to as the “burrowing fox” by Lewis and Clark and are
‘ considered one of the most fossorial canids in North
| America because they use dens throughout the year
| (Kilgore 1969; Egoscue 1979). Swift Foxes spend the
/ majority of diurnal hours in dens or near den entrances,
| often concurrently with a mate (Cutter 1958; Kitchen
et al. 1999). Denning might enable Swift Foxes to
maintain homeostasis by providing a cool, damp micro-
habitat refuge. Dens offer shelter from the elements and
facilitate escape from predators, especially Coyotes
(Canis latrans), in an environment that offers little nat-
ural cover (Egoscue 1979; Rongstad et al.1989; Her-
rero et al. 1991; Pruss 1999). Kitchen et al. (1999)
found all Coyote-killed Swift Foxes near the periphery
or outside their home range boundary, a significant dis-
tance from their nearest currently used den.
Thus, knowledge of habitat requirements for den
sites might be an important component of Swift Fox
conservation efforts (Hillman and Sharps 1978; Pruss
1999): however, results of studies on Swift Fox den site
‘selection have varied. Swift Foxes may modify bur-
rows dug by other animals such as American Badgers
(Taxidea taxus), ground squirrels (Spermophilus spp.)
or Black-tailed Prairie Dogs (Cynomys ludovicianus),
or dig their own dens (Kilgore 1969; Hillman and
Sharps 1978; Cameron 1984; Pruss 1999). Dens have
been found in a variety of habitat types including short-
and mid-grass prairie, grazed prairie, cultivated fields,
fence rows, and rock outcrops (Cutter 1958; Hines
1980; Cameron 1984: Urésk and Sharps 1986; Rong-
stad et al. 1989; Pruss 1999). Occasionally, dens have
been associated with man-made structures such as cul-
verts, buildings, and cemetery gravesites with a con-
crete cap (Kilgore 1969; Hillman and Sharps 1978;
Jackson and Choate 2000). Dens are frequently locat-
Sil
a2 THE CANADIAN FIELD-NATURALIST
ed on or near tops of gently sloping hills, which are
well drained and have a clear line of sight (Hillman
and Sharps 1978; Cameron 1984; Uresk and Sharps
1986; Pruss 1999; Harrison 2003). However, dens also
have been found in level, open areas and at the base
of hills (Kilgore 1969; Hines 1980; Cameron 1984).
Den entrance characteristics vary among seasons
and individual dens. There are indications that natal
dens typically have more entrances than non-natal dens
(Kilgore 1969; Hillman and Sharps 1978). Hines
(1980) reported that Swift Foxes seem to select for
entrances with west and east exposures. Similarly,
Uresk and Sharps (1986) documented a trend toward
dens with easterly exposures. In contrast, Rongstad et
al. (1989) and Pruss (1999) found entrance exposure
to be random.
An additional factor that might influence the selec-
tion of dens is the position of the site within the home
range. Previous studies investigating den use in Swift
Foxes have not considered den placement within the
range as a factor influencing selection. We contend,
however, that due to the importance of dens for pred-
ator escape (Herrero et al. 1991; Kitchen et al. 1999;
Pruss 1999), placement relative to the home range core
and boundary areas might be a driving force in selec-
tion. It is likely den placement is correlated to overall
space use patterns and dens are positioned to facili-
tate a quick escape from an approaching predator.
The objective of this study was to determine the fac-
tors affecting den site selection for Swift Foxes on the
Pinon Canyon Maneuver Site, in southeastern Col-
orado. We examined den site placement and micro-
habitat characteristics of active Swift Fox dens and
compared these to sites not used by Swift Foxes.
Methods
Study Area
We conducted the study on the 1040-km? Pinon
Canyon Maneuver Site (PCMS) in Las Animas Coun-
ty, Colorado. The dominant cover type on the PCMS
is native grassland (60%), with Juniper (Juniperus
monosperma) and Pinyon Pine (Pinus edulis) com-
munities found in the hills and canyons (Shaw et al.
1989). Annual precipitation ranges from 26 to 38 cm
(United States Department of Army 1980). Mean
monthly temperatures range from —1°C in January to
23°C in July (Andersen and Rosenlund 1991). Preda-
tors known to kill Swift Fox are Coyotes (Covell 1992;
Kitchen et al. 1999) and raptors (Covell and Rongstad
1990).
Swift Fox Capture
Swift Foxes were captured with double-door box
traps (80 x 25 x 25 cm) baited with chicken or mack-
erel (Covell 1992). We deployed traps in the evening
and checked them the following morning. Traps were
not activated during periods when nighttime temper-
atures dropped below -10°C. We attached a radiocollar
Vol. 120
and ear tag to the Swift Fox and recorded the weight,
sex and approximate age (using tooth wear) of the ani-
mal (Rongstad et al. 1989). All Swift Foxes were re-
leased at the site of capture. We followed telemetry
procedures recommended by White and Garrott (1990).
Locations were obtained by triangulating 2-3 bearings
to the animal’s position within a 10-minute period.
Triangulation angles were maintained between 20° and
160° (Gese et al. 1988). We used aerial telemetry (Mech
1983) to locate missing animals. Telemetry error was
determined by comparing telemetry locations with
actual locations of stationary reference transmitters.
When Swift Foxes were located in a den, we record-
ed the UTM coordinates of the den. We attempted to
locate Swift Foxes approximately every 1-3 days with
locations obtained throughout the 24-h period to reduce
bias in home range estimates (used for den placement
analysis).
Den Site Characteristics
Microhabitat characteristics of den sites were meas-
ured at a randomly selected sub-sample of the known
active dens used by Swift Foxes during the breeding
season (defined on the basis of energetic demands due
to climatic changes and prey abundance, and behay-
ioral characteristics to be 15 December to 14 April).
Active dens were found by tracking radio-collared Swift
Foxes. Den site selection was not stratified by sex as
Swift Foxes form pair bonds with males and females
concurrently using the same dens.
The following physical characteristics were meas-
ured at each den site: number of entrances, height and
width of each entrance, den aspect, den position on
hills, slope of site, percent rock in soil, vegetative
cover, and horizontal foliar density. These character-
istics were selected based on field observations and
published descriptions of potentially important habi-
tat characteristics (Hines 1980; Cameron 1984; Ursek
and Sharps 1986; Rongstad et al. 1989; Covell 1992;
Pruss 1999).
Den position on a hill was classified as bottom, mid-
dle, top, or flat when there was no hill. We determined
maximum slope outside the den using a hand-held cli-
nometer (Hays et al. 1981). The percentage of rock in
the soil was visually estimated from the presence of
rock in the dirt mound runway leading to the den
entrance. Vegetation cover was determined by estimat-
ing the percent cover of grasses, forbs, shrubs, and bare
soil in eight 1-m? plots randomly located within 10 m
of the den. Percent soil and vegetation cover were esti-
mated within the nearest 5% category. We indexed
horizontal foliar density at 5 distances (10, 20, 30, 40,
and 50 m) from the den in the four cardinal directions
with a vegetation profile board and techniques for the
fixed distance approach (Hays et al. 1981). The number
of squares visible on a vertical board was counted by
an observer lying down to approximate the visibility
from the height of a Swift Fox.
2006
BRITISH
COLUMBIA
KITCHEN, GESE, AND LupPIS: DEN SITE SELECTION BY Swirr FOXES 33
Current distribution
W@ Historic distribution
Study site (PCMS)
FicurE |. Swift Fox current and historic range and study area, Pinon Canyon Maneuver Site, Colorado, 2000.
Den Site Selection
To examine den site selection, we compared micro-
habitat characteristics of dens to control sites that rep-
resented available sites (Taylor et al. 1999). For each of
the active den sites, we randomly selected one control
site (not a den) within the Swift Foxes home range. We
used the control site to compare the site characteristics
of active den sites versus random sites within the Swift
Foxes’ home range. The control site was located 500 m :
away from the active den site in a random direction.
This distance was arbitrary but allowed for selection
of a random location within the home range. Because
control sites were in the home range of the Swift Fox
inhabiting the active den site, they were considered
available as a possible denning location to the Swift
Fox. The same den site characteristics were evaluated
on the control sites, excluding variables associated with
den entrances.
Den Placement
Dens used by Swift Foxes during the breeding sea-
son were recorded using telemetry procedures to assess
the placement and use of the dens within the home
range. The Swift Foxes’ home ranges were described
using a 95 % fixed kernel home-range estimator based
on > 30 locations recorded within the breeding sea-
son (Worton 1989) with Arcview 3.0 software (Envi-
ronmental Systems Research Institute, Inc., Redlands,
California). Areas within the home range were then
defined as the core area, which was the area within the
50% isopleth, the mid-range area, between the 50%
and 75% isopleths, and the boundary area, between
the 75% and the 95% isopleths. The number of dens
and the frequency of use of those dens in the core, mid-
range, and boundary areas of the home range were
compared. In addition, we compared the number of
dens in each area to the number expected from the
proportion of each area in the home range; i.e., 50%
within the core area, 25% within the mid-range area,
and 20% within the boundary area.
Statistical Analysis
Because the habitat measurements were categorical,
chi-square tests were used to assess the difference
between habitat characteristics at active dens versus
control sites (Zar 1999). Whether den entrances were
randomly distributed around a circle (i.e., no predom-
inant direction) was tested using Rayleigh’s nonpara-
metric Z test (Zar 1999). We attempted to predict class
membership (i.e., active dens or control sites) by using
a classification tree analysis with cost complexity prun-
ing (CART: Verbyla 1987; S-Plus Version 6.0 1988-
34 THE CANADIAN FIELD-NATURALIST
2001.). Also called tree regression (Rejwan et al. 1999),
this method repeatedly partitions the study sites into
two groups (active dens and control sites) that are as
similar as possible based upon the five variables com-
mon to both groups (position, slope, horizontal foliar
density, percent rock in soil, and percent vegetative
cover). Each of the independent variables is used at
each step in the analysis regardless of whether they
were previously used in the tree. The tree that is cre-
ated is hierarchically structured with the complete data
set at the top (the root) and the binary splits, referred
to as “nodes”, below to the final undivided “leaves”
or groupings at the bottom of the tree. Analysis of
variance with Tukey-adjusted post-hoc comparisons
was used to assess the difference in the number of dens
used and the number of times these dens were used
by area (SAS Version 8.2 2001). Log transformations
were used to achieve normality where deviations
occurred. In cases where more than one den per Swift
Fox was sampled, one den was randomly chosen for
analysis to avoid pseudoreplication.
Results
Forty-two dens belonging to 42 Swift Foxes were
located from December 1999 to April 2000 and used
to evaluate den site characteristics. Most active dens
had only one or two entrances and the maximum num-
ber of den entrances was seven. The mean height of
den entrances was 19.4 + 3.1 cm and the mean entrance
width was 17.9 + 2.5 cm. The percentage of dens with
entrances oriented to the north, south, east, and west
were 29.9, 21.1, 21.9, and 27.1%, respectively. These
aspects were randomly distributed around a circle
(Z,, = 0.088; P = 0.54), i.e., dens were not oriented in
a predominant direction.
The mean maximum slope of active dens was 1.3 +
1.3 degrees. We found no difference between the
slope at den sites and control sites (y? = 2.71, df = 4,
P=0.61). Most dens (88.7%) and control sites (80.3%)
were located on gentle slopes between 0 and 2 degrees.
Four percent of control sites were located on slopes
60
50 -
40 -
Percent Use
(do)
o>)
I
Vol. 120
that exceeded 5 degrees, however, no dens were found
on slopes that steep. The majority of Swift Fox dens
were positioned either on flat ground or in the middle of
a hill, and relatively few dens were located at the top or
bottom of a hill (Figure 1). Control sites were found in
similar positions with no detected difference between
site position of used and control sites (y* = 2.04, df = 3,
P=0.59; Figure 1). We also found no difference in the
pattern of visibility by distance from the den as com-
pared to control sites (y* = 0.06, df = 4, P = 0.99).
Visibility from both active den sites and control sites
decreased uniformly in each direction as one moved
away from the site. The vegetative cover of den
sites and control dens was similar (y7 = 1.98, df = 3,
P =0.598) with both dens and control sites occurring
in areas mostly covered in grasses and soil (Figure 2).
There was a significant difference in the percentage
of rock at dens versus control sites (y7 = 8.91, df = 3,
P = 0.03; Figure 3) with dens having higher propor-
tions of intermediate percentages of rock and control
sites having higher proportions of rock below 5% or
above 20%. The classification tree used to characterize
the habitat at dens versus control sites failed to pro-
duce any biologically relevant pattern.
The average number of dens used per Swift Fox
during the breeding season was 5.1 + 2.1. The num-
ber of dens used by Swift Foxes varied by area of the
home range (Wa 26 = 37.53, P < 0.001; Figure 4a).
Tukey’s adjusted post-hoc tests indicated that more
dens were used in the core area than in the mid-range
and boundary areas (mid-range: t,, = 6.29, P < 0.001;
boundary: t= - 7.54, P < 0.001). There were also sig-
nificantly more dens in the mid-range area than the
boundary area (t,, = -2.20, P = 0.035). In addition,
we found that the number of dens in each area varied
from that expected from the proportion of the area,
and thus the frequency of use of the area (y* = 38.41,
df = 3, P < 0.0001).
Overall, there was a difference in the frequency of
use of dens in the different areas of the home range
(F,.. = 44.94, P < 0.001; Figure 4b). The number of
2,33
w Den sites
O Control sites
|
(0) 5;
Bottom Middle
Position
Top
Flat
FIGURE 2. Position of Swift Fox den sites and control sites, Pinon Canyon Maneuver Site, Colorado, 2000.
2006
100 -
ioe)
i=)
Percent Use
(o>)
Oo
NO A
Sy [>
I i]
|
i —
Shrubs Grasses
KITCHEN, GESE, AND LupIs: DEN SITE SELECTION BY SWIFT FOXES
Soil
-)
wv
g Den sites
0 Control sites
Forbs
Vegetative Cover
FIGURE 3. Vegetative cover at Swift Fox den sites and control sites, Pinon Canyon Maneuver Site, Colorado, 2000.
times that dens in the core area were used was higher
than in the mid-range or boundary areas (mid-range:
t,,= 7.52, P < 0.001; boundary: t,= 7.65, P <0.001).
There was no difference in the number of times dens in
the mid-range and boundary areas were used (f,, = 1.55,
P = 0.13). The pattern of den use was variable. For
example, one Swift Fox was located in only one den
for a period spanning 67 days, while another Swift Fox
used at least four different dens in a five-day period.
Discussion
Our results indicated Swift Foxes on the PCMS did
not select den sites based on the microhabitat charac-
teristics that we evaluated. However, Swift Foxes ap-
peared to select areas of intermediate rock percentages.
The selection of areas with intermediate rock percent-
ages may be related to den structure and strength. Foxes
had no preference for particular positions or vegeta-
tion at den sites despite other studies concluding that
Swift Foxes preferentially selected den sites on hilltops
(Uresk and Sharps 1986; Pruss 1999; Harrison 2003).
The finding that Swift Foxes in our study area select
den sites with habitat characteristics that do not differ
from control sites was not unexpected considering the
homogeneous prairie environment on the PCMS. Our
result does differ from other Swift Fox studies which
reported preferences for certain habitat characteristics,
however. The importance of selection of hill position
and vegetation characteristics might be determined by
variation in topography and habitat in the geographic
area. Habitat selection by Swift Foxes on the PCMS
appears to operate at the landscape level instead of at
the den site level as Swift Foxes were only trapped on
the open prairie on the PCMS despite efforts to capture
Swift Foxes in wooded areas (Schauster et al. 2002).
Den site selection occurred at a larger scale; the
placement of dens relative to areas within the home
range was a significant factor in site selection. Foxes
®
27680):
cob)
B08. ‘m Den sit
3 40 m Den sites
Oo 20 ‘oO Control sites
0 2) ee <
o 9Q\o 9 9 9
ao. oa oY s\ oS io s
a
Percent Rock
FicurE 4. Percentage of rock at Swift Fox den sites and control sites, Pinon Canyon Maneuver Site, Colorado, 2000.
36
THE CANADIAN FIELD-NATURALIST
Vol. 120
6 -
ane
rane
oe.
@ 2
a =
0 hy Ne le
Mid-range Boundary
Area
& 15 -
Y”
®
Ey 0]
or 7)
o 7 a4
2
E
> 0
Core Mid-range Boundary
Area
FIGURE 5. Placement of Swift Fox dens within the home range (top) and frequency of Swift Fox den use by area of the home
range (bottom), Pinon Canyon Maneuver Site, Colorado, 2000.
used more dens in the core area of the home range as
compared to the mid-range or boundary areas. In addi-
tion, dens in the core area were used more frequently
by radio-tracked Swift Foxes than dens in the mid-
range or boundary area. Den placement within the
home range may be designed to facilitate escape from
Coyotes. Previous studies have indicated that Swift
Foxes use dens as a mechanism of escape from pred-
ators (Herrero et al. 1991; Kitchen et al. 1999; Pruss
1999) and that Coyote-caused mortality might be a key
factor in limiting population growth in North America
as well as the re-establishment of Swift Foxes on the
Canadian prairies (Scott-Brown et al. 1987). Kitchen
et al. (1999) found that all Coyote-caused Swift Fox
mortalities occurred near or outside the Swift Fox’s
home range boundary, a substantial distance from their
nearest currently used den.
Thus, Swift Foxes might maintain the majority of
dens in the core area of the home range where they
spend most of their diurnal hours (Schauster et al.
2002), and use dens only occasionally in the outer areas
of their range to reduce encounters with and facilitate
escape from predators. Previous studies have suggest-
ed that other preferences for den sites also might be
due to Coyote avoidance, including den locations that
are negatively associated with water sources (Pruss
1999), and dens that are closer to roads than unoccu-
pied sites (Pruss 1994; Harrison 2003).
The importance of interspecific competition in deter-
mining den site selection has also been noted in Kit
Foxes (List and Macdonald 2003) and Arctic Foxes
(Alopex lagopus; Frafjord 2003). Frafjord (2003) doc-
umented that the distance to Red Fox (Vulpes vulpes)
dens and the elevation from the tree line were the most
important factors determining den use in Arctic Foxes.
Kit Foxes in Mexico denned more often in grasslands
and less in Prairie Dog (Cynomys ludovicianus) towns
than expected (List and Macdonald 2003). These au-
thors hypothesized that Kit Foxes reduced the amount
of time they spent in Prairie Dog towns due to the
activity of Coyotes.
Thus, Swift Fox den site selection in homogenous
environments might be unrelated to microhabitat char-
acteristics and more dependent on interspecific inter-
actions and predator avoidance. This study also illus-
trates the importance of examining den site selection
at multiple scales to determine all site selection fac-
tors. Due to the importance of denning to Swift Fox
2006
ecology, a full understanding of den site selection in
various environments is essential to recovery, reintro-
duction, and management efforts.
Acknowledgments
Funding and assistance provided by the U.S. Army,
Directorate of Environmental Compliance and Man-
agement (DECAM), Fort Carson, Colorado, through
the U.S. Fish and Wildlife Service (USFWS), Colorado
Fish and Wildlife Assistance Office, Golden, Colorado,
and the Utah Cooperative Fish and Wildlife Research
Unit at Utah State University, Logan, Utah. Addition-
al support provided by the U.S. Department of Agri-
culture, Wildlife Services, National Wildlife Research
Center, Logan Field Station at Utah State University,
Logan, Utah. We thank T. Warren, G. Belew, and R.
Bunn at DECAM, and B. Rosenlund at the USFWS for
logistical support. We thank E. Bergman, E. Joyce, S.
Karki, M. Klavetter, A. Kozlowski, H. Kratz, and L.
Schutte for field assistance. Research protocols were
approved by the Institutional Animal Care and Use
Committees (IACUC) at the National Wildlife Research
Center and Utah State University.
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Vol. 120
Worton, B. J. 1989. Kernel methods for estimating the uti-
lization distribution in home range studies. Ecology 70:
164-168.
Zar, J. H. 1999. Biostatistical analysis. Prentice-Hall, Inc.
New Jersey.
Received 8 December 2004
Accepted 15 November 2006
First Records of the Northern Long-eared Bat, Myotis septentrionalis,
in the Yukon Territory
THOMAS S. JuNG!, BRIAN G. SLOUGH”, DAVID W. NAGORSEN?, TANYA A. DEwEY?, and Topp Powe!
'Yukon Department of Environment, Fish and Wildlife Branch, Box 2703, Whitehorse, Yukon Territory Y1A 2C6 Canada;
e-mail: thomas.jung@ gov.yk.ca
235 Cronkhite Road, Whitehorse, Yukon Y1A 5S9 Canada
$Mammalia Biological Consulting, 4268 Metchosin Road, Victoria, British Columbia V9C 374 Canada
4Mammal Division, Museum of Zoology, University of Michigan, 1109 Geddes Road, Ann Arbor, Michigan 48109-1079
U.S.A.
Jung, Thomas S., Brian G. Slough, David W. Nagorsen, Tanya A. Dewey, and Todd Powell. 2006. First record of the Northern
Long-eared Bat, Myotis septentrionalis, in the Yukon Territory. Canadian Field-Naturalist 120(1): 39-42.
Three adult male Northern Long-eared Bats, Myotis septentrionalis, were captured in mist nets in July 2004 in the LaBiche
River Valley, southeastern Yukon. These are the first records of M. septentrionalis in the Yukon. Further survey work is needed
to delineate the extent of the range and population structure of this and other species of bats in northwestern North America.
Key Words: Northern Long-eared Bat, Myotis septentrionalis, distribution, Yukon.
The diversity and distribution of bats in northwestern
North America (e.g., =>55° N) is poorly documented.
Few areas have been adequately surveyed, particularly
for rare or cryptic species (sensu Nagorsen and
Brigham 1993; van Zyll de Jong and Nagorsen 1994;
Parker and Cook 1996; Parker et al. 1997). Youngman
(1975) reported the Little Brown Bat (Myotis lucifu-
gus) as the only species of bat in the Yukon. However,
Youngman (1975) noted the possibility of other species
occurring in the Yukon, such as the Big Brown Bat
(Eptesicus fuscus) and Long-legged Bat (Myotis
volans), based on single specimens collected near Fair-
banks, Alaska, and Atlin, British Columbia, respective-
ly. In the 30 years subsequent to Youngman (1975), no
species of bats other than M. /ucifugus have been con-
firmed from the Yukon. Slough (2000*), however,
recorded a bat in southeastern Yukon that appeared to
be a Big Brown Bat according to its echolocation call
characteristics (C. Corben, personal communication),
but it may have been a Silver-haired Bat (Lasionycteris
noctivagans) and remains to be verified by a capture.
We believe that species other than M. lucifugus may —
extend northward into the Yukon, and that a lack of
records of other species is largely a function of previous
search effort.
Based on a specimen collected in Nahanni National
Park, Northwest Territories (van Zyll de Jong 1985),
captures in recent surveys in adjacent British Columbia
(Wilkinson et al. 1995*: Vonhof and Wilkinson 1997*),
and an unidentified Myotis in a recording of an echolo-
cation call (Slough 2000*), we suspected that the North-
em Long-eared Bat (Myotis septentrionalis) and the
Western Long-eared Bat (Myotis evotis) could be pres-
ent in southeastern Yukon. In summer 2004, we under-
took a preliminary field survey of the mammalian
diversity in the boreal forest of southeastern Yukon,
including a bat inventory. Here, we provide the first
records of the Northern Long-eared Bat in the Yukon.
Methods
On 28 July — 2 August 2004, we used mist-nets to
capture bats in the LaBiche River Valley (60.126°N,
124.064°W) of extreme southeastern Yukon. Mist-net-
ting occurred at three sites; two sites were over small
ponds (>150 m7’) within dense boreal forest, and anoth-
er at an intersection of a roughly 3 m wide forest trail
and a roughly 30 m wide forest access road. Nets were
set across and bordering the ponds and perpendicular
to the road and forest trail. Nets were opened 0.5 hr
before sunset (about 23:00 hrs PDT) and closed 2—3 hr
thereafter. A 6 m long mist net open for | hr was
used as our definition of | mist-net hr.
Captured bats were examined for external diagnostic
characteristics according to van Zyll de Jong (1985)
and Nagorsen (2002), including presence of dark
shoulder patches in the pelage, ear length, number of
hairs on the trailing edge of the uropatagium, presence
of keeled calcar, and relative length and shape of the
tragus. We measured forearm length, ear length, and
mass, and determined age as per van Zyll de Jong
(1985). No voucher specimens were procured; rather,
we obtained morphological measurements and wing
punches for mitochondrial DNA sequencing to verify
our field identifications. Wing punches were stored in
70% ethanol until lab analysis. Genetic analyses were
conducted at the University of Michigan’s Museum of
Zoology (Ann Arbor, Michigan), and included an
unweighted maximum parsimony analysis of 745 bp
of the cytochrome b gene using 21 specimens of Myotis
(Figure 1).
39
40 THE CANADIAN FIELD-NATURALIST
Results and Discussion
We amassed 42.1 mist-net hr over six nights of sam-
pling; only six bats were captured. On 28 July 2004, we
captured two adult male M. septentrionalis and one M.
lucifugus at a small, shallow pond (ca. 108 m7). On 29
July 2004, a third adult male northern M. septentrion-
alis and two M. lucifugus were captured at the same
pond. Although three of our captures were consistent
with the morphological traits of M. septentrionalis
(Table 1), this species can be difficult to distinguish
from other species of potentially sympatric long-eared
Myotis (1.e., M. evotis and M. keenii), based on external
characteristics alone (van Zyll de Jong 1979; Nagorsen
2002). DNA sequences, however, confirmed our tenta-
tive species identifications: the three M. septentrionalis
are included in the M. septentrionalis clade (Figure 1).
The presence of M. septentrionalis in the Yukon is
not surprising, considering that captures were reported
from elsewhere in the Liard River Watershed by
Wilkinson et al. (1995*), Bradbury et al. (1997*) and
Vonhof and Wilkinson (1997*). The only voucher taken
in those surveys to confirm identification, however, was
a specimen of M. septentrionalis collected by Vonhof
and Wilkinson (1997*) from the Fort Nelson River,
British Columbia, near Highway 77 (Royal British
Columbia Museum, RBCM 19516), about 140 km
southeast of our captures. It is not known if there are
breeding populations of M. septentrionalis in the Yukon,
as no juveniles, or lactating or post-lactating females
were captured. Breeding populations of the Northern
Long-eared Bat, however, have been reported from
nearby northeastern British Columbia (Wilkinson et
al. 1995*, Vonhof and Wikinson 1997*). Similarly, it is
not known if M. septentrionalis hibernates in the Yukon.
No bat hibernacula have been found in the Yukon.
Despite low capture rates, we suspect that M. septen-
trionalis were locally common in our survey area,
based on the number of captures relative to M. lucifu-
gus. Moreover, we often detected what appeared to be
M. septentrionalis at all of the mist-netting sites, based
on low intensity echolocation calls (Faure et al. 1993)
heard on narrowband bat detectors tuned to 40-50 kHz
and observations of detection distances of passing bats.
Our data, along with that of Wilkinson et al. (1995*),
Bradbury et al. (1997*), and Vonhof and Wilkinson
Vol. 120
(1997*), suggest that M. septentrionalis may be locally
common and widely distributed throughout the Liard
River Watershed in British Columbia and the Yukon.
We suspect that M. septentrionalis may occur through-
out the Liard River Watershed in the Yukon, from the
LaBiche River west to the Little Rancheria River. Other
species of bats (e.g., M. evotis, M. volans, Eptesicus
fuscus, and Lasionycteris noctivagans) may also be
present in southern Yukon, but not yet confirmed with
a capture, voucher, or DNA sample.
Perhaps the most intriguing question is whether M.
septentrionalis is a long-established member of the
Yukon mammalian fauna that has simply gone unde-
tected, or, alternatively, is it a species that has recently
(i.e. <100 years ago) colonized southeastern Yukon?
Other mammals have recently colonized the Yukon,
from the south, including: Mule Deer and White-tailed
Deer (Odocoileus hemionus and O. virginianus; Hoefs
2001), Coyote (Canis latrans; Youngman 1975), and
Cougar (Puma concolor; Jung et al. 2005). Moreover,
Humphries et al. (2002) provided data from simulation
models that suggested that the range of M. lucifugus,
and presumably other species of bats in the boreal for-
est, would substantially move northward in response to
global warming. Regardless of the origin of M. septen-
trionalis in the Yukon, these records underscore that
our knowledge of the diversity and distribution, and
population structure of bats in the Yukon is limited.
Directed surveys are needed to address these informa-
tion gaps and provide a basis for management and con-
servation initiatives that include bats.
Acknowledgments
S. Cannings and A. Runck kindly provided com-
ments on an earlier draft of this note. Additional tissue
samples for DNA sequence comparison were kindly
provided by T. Dewey, P. Myers, D. Burles, and R.
Benedict. The staff of Devon Canada Corporation’s
LaBiche River Gas Plant kindly went out of their way
to accommodate us and provide logistical support.
Funding was provided by the Yukon Department of
Environment and NatureServe Yukon. Additional finan-
cial support was provided by a Northern Research
Endowment Grant from the Northern Research Insti-
tute, Yukon College, to B. Slough.
TABLE 1. Age, sex, and morphometrics of Myotis septentrionalis (n = 3) captured in the LaBiche River valley, southeastern
Yukon, 28-29 July 2004.
Specimen GenBank Forearm Ear Mass
Label Accession Species! Sex Age Length (mm) Length (mm) g)
LaBiche-1 AY883902 M. septentrionalis Male Adult 16 37.7 7.6
LaBiche-3 AY883904 WM. septentrionalis Male Adult 15 35.8 6.6
LaBiche-9 AY883905 M. septentrionalis Male Adult 14 37.6 6.9
' As tentatively identified in the field and subsequently confirmed with genetic analyses.
2006
100
89
95
JUNG ET AL.: RECORDS OF THE NORTHERN LONG-EARED BAT IN THE YUKON 4]
LaBiche9
M. septentrionalis
(British Columbia,
5 individuals)
Mh. septentrionalis
(Nebraska)
M. septentrionatis
(South Dakota)
LaBichel
LaBiche3
M. septentrionalis
(Michigan)
M. auriculus
LaBiche2
M. lucifugus
(South Dakota)
M. lucifugus
(South Dakota)
LaBiche10
fi. lucifugus
(Michigan)
M. evotis
(South Dakota)
M. evotis
(Colorado)
M. volans
MM. yumanensis
Eptesicus fuscus
FIGuRE |. Placement of Myotis septentrionalis (LaBiche 1,3, and 9) and M. lucifugus (LaBiche 2 and 10) samples collected
in the LaBiche River Valley, Yukon, based on an unweighted maximum parsimony analysis of cytochrome b (745 bp)
for these samples compared to other Myotis. Eptesicus fuscus is used as an outgroup. Numbers above the branches
represent bootstrap support values for those nodes.
Documents Cited (marked * in the text)
Bradbury, S. M., S. Morris, and S. McNally. 1997. Bat sur-
vey of the Liard River watershed in British Columbia. Un-
published Report. British Columbia Ministry of Environ-
ment, Lands and Parks, Victoria, British Columbia. 29 pages.
Slough, B. G. 2000. A survey of the bat fauna of the Yukon
Territory: 1999 field studies. Unpublished Report. North-
em Research Institute, Yukon College, Whitehorse, Yukon.
26 pages.
Vonhof, M. J., and L. C. Wilkinson. 1997. Roosting habitat
requirements of northern long-eared bats (Myotis septentri-
onalis) in the boreal forest of northeastern British Colum-
bia. Unpublished Report. British Columbia Ministry of
Environment, Lands and Parks, Fort St. John, British
Columbia. 88 pages.
Wilkinson, L. C., P. F. J. Garcia, and R. M. R. Barclay.
1995. Bat survey of the Liard River watershed in northern
British Columbia. Unpublished Report. British Columbia
Ministry of Environment, Lands and Parks, Victoria,
British Columbia. 39 pages.
Literature Cited
Faure, P. A., J. H. Fullard, and J. W. Dawson. 1993. The
gleaning attacks of the northern long-eared bat, Myotis
septentrionalis, are relatively inaudible to moths. Journal
of Experimental Biology 178: 173-189.
Hoefs, M. 2001. Mule, Odocoileus hemionus, and White-
tailed, O. virginianus, deer in the Yukon. Canadian Field-
Naturalist 115: 296-300.
Humphries, M. M., D. W. Thomas, and J. R. Speakman.
2002. Climate-mediated energetic constraints on the distri-
bution of hibernating mammals. Nature 418: 313-316.
Jung, T. S., and P. J. Merchant. 2005. First confirmation of
cougar, Puma concolor, in the Yukon. Canadian Field-Nat-
uralist 119(4): 580-581.
42 THE CANADIAN FIELD-NATURALIST
Nagorsen, D. W. 2002. An identification manual to the small
mammals of British Columbia. Ministry of Sustainable
Resource Management, Ministry of Water, Land and Air
Protection, and Royal British Columbia Museum, Victoria,
British Columbia. 153 pages.
Nagorsen, D. W., and R. M. Brigham. 1993. Bats of British
Columbia. Volume 1. The Mammals of British Columbia.
UBC Press, Vancouver, British Columbia. 176 pages.
Parker, D. I., and J. A. Cook. 1996. Keen’s long-eared bat,
Myotis keenii, confirmed in southeast Alaska. Canadian
Field-Naturalist 110: 611-614.
Parker, D. 1I., B. E. Lawhead, and J. A. Cook. 1997. Distrib-
utional limits of bats in Alaska. Arctic 50: 256-265.
Vol. 120
van Zyll de Jong, C. G. 1979. Distribution and systematic
relationships of the long-eared Myotis in western Canada.
Canadian Journal of Zoology 57: 987-994.
van Zyll de Jong, C. G. 1985. Handbook of Canadian
mammals. Volume 2: Bats. National Museum of Natural
Sciences, Ottawa, Ontario. 212 pages.
van Zyll de Jong, C. G., and D. W. Nagorsen. 1994. A
review of the distribution and taxonomy of Myotis keenii
and Myotis evotis in British Columbia and the adjacent
United States. Canadian Journal of Zoology 72: 1069-1078.
Youngman, P. M. 1975. Mammals of the Yukon Territory.
National Museums of Canada, Ottawa, Ontario. 192 pages.
Received 4 February 2005
Accepted 12 September 2005
Recent Declines of House Sparrows, Passer domesticus, in Canada’s
Maritime Provinces
ANTHONY J. ERSKINE
16 Richardson Street, Sackville, New Brunswick E4L 4H6 Canada
Erskine, Anthony J. 2006. Recent declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces. Canadian
Field-Naturalist 120(1): 43-49.
House Sparrows, Passer domesticus, were introduced to North America after 1850, increased and spread up to 1920, and sta-
bilized or decreased thereafter until 1960. In the Maritimes (and perhaps some other areas), a further decline set in after 1970,
continuing to the present. Now the species is rare to absent in much of the Maritimes, except around farms with livestock.
Decline here since 1970 probably approaches 90 per cent in most other areas of human settlement except south of 45°N
Similar declines are known in the U.K., but seem poorly documented, if recognized, in North America outside of our region.
Key Words: House Sparrows, Passer domesticus, declines, New Brunswick, Nova Scotia, Prince Edward Island.
Fifty years ago, House Sparrows (Passer domesti-
cus; HoSp hereafter — a standard abbreviation used in
many reports of organized birding activities) were so
nearly ubiquitous in and around human settlements in
eastern Canada that one sometimes had to “tune them
out” to hear other birds. Then, it would have been un-
imaginable that this abundant species could become
scarce in our lifetimes. HoSp, and their (urban) habi-
tats, were virtually beneath notice both of birders and
academics, so few serious studies of the species were
made (see Kendeigh 1973), and none in Canada, dur-
ing the rapid expansion of research on birds after World
War 2. The result is a continuing scarcity of hard data
on HoSp, precluding rigorous discussion of its popu-
lation ecology. This study is a first look at what hap-
_ pened in the recent past and what may still be investi-
gated usefully, in the Maritime Provinces of Canada
{New Brunswick, Nova Scotia, and Prince Edward
Island (hereafter “the Maritimes”)].
HoSp were first introduced to North America around
1850, and became established in the Maritimes by the
1880s (Christie 1979; Lowther and Cink 1992). Num-
_ bers and densities in the next century are known main-
ly from anecdotal remarks in general accounts. As was
general in eastern North America by the early 20" Cen-
tury, HoSp were nearly ubiquitous and very common
around human settlements, including cities, towns, vil-
lages, and even isolated farms. HoSp thrived on grain
Wastage, as well as using other food wastes. Nearly all
_ human settlements, through the 1940s, harboured dom-
| estic animals, especially livestock and poultry, which
| were fed on small grains at least seasonally.
Motor vehicles began replacing horses in transporta-
i tion by the 1920s, and the resulting decrease in horse
droppings and other waste grain sources led to a decline
| in HoSp, documented mainly by general statements
' (e.g., Bent 1958). Motor vehicle use in the Maritimes
| was much less during the Great Depression of 1930s,
when most people could not afford cars, and in World
War 2, when gasoline and tires were reserved for mil-
itary use. Use of horses again became widespread in
those decades, and that presumably slowed or halted
the earlier decline of HoSp. After the war, motor vehi-
cles again became available, use of horses declined rap-
idly, and probably HoSp also declined. Very few bird
counts were made in the Maritimes before 1960, when
HoSp still were considered widespread and abundant
(Squires 1952; Godfrey 1954; Tufts 1962). Winter bird
counts at Wolfville, Nova Scotia (J. S. Erskine 1968)
showed no obvious trend in HoSp numbers between
1948 and 1968. Christie (1979) summarized bird status
changes in the Maritimes over the century to 1978, not-
ing arrival, increase, and subsequent decline in HoSp,
with numbers likely stable after 1930.
Starting in 1977, winter plot counts through 1988 in
Sackville, New Brunswick (Erskine 1992b), showed
further decreases in HoSp — evidently nor related to
decline of horses and other livestock, which already
were absent in such towns (compare Erskine and
McManus 2005). Also in the 1980s, Christie (1985 and
following years) remarked on generally lower HoSp
numbers on New Brunswick Christmas Bird Counts
(CBCs hereafter). Some naturalists (personal communi-
cations) remarked on similar decreases in Nova Scotia
in the early 1990s. The cooperative Breeding Bird Sur-
vey (BBS) documented HoSp declines across eastern
Canada in summers 1967-1996 (Downes and Collins
2003), with decreases each decade (significant only in
1981-1990) in the Bird Conservation Region (BCR 14)
including the Maritimes. HoSp continued to decline,
and by 2000 were rare in Sackville, New Brunswick
(personal observations), and in nearby towns.
In the near-absence of even quasi-systematic counts
of HoSp, CBCs provided the only widespread sam-
pling of bird numbers in the Maritimes, with many more
counts in more years than the BBS. Although CBC data
are poorly standardized (e.g., Dunn et al. 2005), those
surveys are extensive enough, both here and across
43
44 THE CANADIAN FIELD-NATURALIST
Canada, to allow examination of HoSp occurrence in
various habitats, in an attempt to identify factors in-
volved in their recent declines.
Methods
Christmas Bird Counts are single-day annual counts
in the period 13 December to 5 January, each restrict-
ed to a 24-km diameter circle selected subjectively.
Bird species richness and availability of observers are
usual selection criteria.
CBC data for HoSp in the Maritimes in 1960-2003
were extracted from N. B. Naturalist (formerly Nature
News), Nova Scotia Birds (formerly N.S. Bird Society
Newsletter), and Island Naturalist. Audubon publica-
tions (American Birds, formerly Audubon Field Notes)
included many fewer counts (“circles” hereafter) in the
Maritimes, and were used only to fill gaps in regional
publication series. All data were tabulated in year vs.
location matrices.
Data were selected by excluding circles surveyed
in <=20 years and others with obvious “discontinu-
ities” in coverage or effort. The selected circles were
grouped by major non-urban habitat combinations, most
including also some human settlements, thus (names
in parentheses used in text and tables hereafter):
e (Forest) inland, predominantly forested, little or
no farming;
e (Coastal) much forest or bog, farming insignifi-
cant;
e (Farm=forest) inland or coastal, farms and other
open areas minor or unproductive but roughly
equal in area to forests;
e (Farm>forest) farming major (much exceeding
forest), relatively productive, usually around a siz-
able town;
e (Major urban) cities or larger towns, other habi-
tats minor.
To allow for varying observer effort, between years
or circles, published counts (“raw data”) were also con-
verted to birds/10 party-hours (“adjusted data”; see also
Discussion).
Statistics on area changes in major habitats of the
Maritimes over the study period (roughly 1960-2000)
were assembled from Statscan websites, Canada Year
Books, and some other directories.
Extrapolation from CBC data to Maritimes HoSp
populations was attempted. Counts from the selected
circles were expanded across the five habitat groupings
above. Further adjustment was made to cover two 1m-
portant omissions in coverage:
e not all suitable areas within active CBCs were
surveyed, and
¢ not all HoSp in the areas surveyed were detected.
Those factors could not be assessed rigorously, but
“informed guesses” from personal experience (see
Appendix) allowed tentative adjustment of extrapo-
lated estimates.
Vol. 120
Results
HoSp counts (adjusted data) for each selected circle
in the various habitat-groupings were summarized as
decade-means (1960-69, 1970-79, etc.; Table 1). Circles
lacking data for >1 decade were excluded, as were
those lacking 1 decade and >=3 years in a 2"! decade.
Mean data were estimated for circles missing one
decade, using ratios of HoSp counts in other circles in
that and the next decade(s). Some general patterns were
evident:
(i) a HoSp decline (from an earlier “plateau’),
most often starting in the 1980s, was apparent
nearly everywhere, in all habitats, and whether
raw or adjusted data were compared;
(ii) the recent decline was less pronounced in cir-
cles where farmlands were important, or at least
equal in area to forests, than elsewhere;
the recent decline began earlier in cities and
large towns than elsewhere.
Observer effort in and around cities and towns in
1960-1979 increased so much that increased detec-
tion of HoSp then sometimes balanced or exceeded
declines, which were less apparent in raw data vs.
adjusted data.
Area changes in major habitats were substantial,
as follows:
(a) Forests remained the largest land cover type in
New Brunswick and Nova Scotia throughout; forest
cover increased in Prince Edward Island, but that was
minor in the overall picture. HoSp were never found
in or near forests except in association with human
settlements, so changes in forest area are ignored in
further discussion.
(b) Among farmlands, “improved lands” (= crops,
pasture, fallow; the main farmland types used by HoSp,
aside from buildings) decreased in all provinces, by
35% in Prince Edward Island (where those remain
dominant land-uses), and by 55% in New Brunswick
and Nova Scotia, between 1961 and 1996. Published
statistics available did not distinguish “commercial”
and “marginal” farms; most decline in New Brunswick
and Nova Scotia was in marginal farms, on which
before 1960 farming was only one of several sources
of income. Land-use on commercial farms became
more intensive in 1961-1996 (personal observations),
in the Maritimes as elsewhere, but that was not obvi-
ous from the published statistics.
(c) Urban area statistics allowed few generalizations.
Larger communities mostly increased in human pop-
ulation and area, whereas many smaller ones decreased,
some dwindling to scattered houses. Cultural changes
within urban areas were many and widespread, involv-
ing sweeping changes in commerce, transportation,
heating, garbage disposal, tolerance of livestock and
poultry, recreation, etc. [see review in Erskine and
McManus 2005]. Correlations with single factors in
urban change could not be made rigorously so did
not warrant statistical tests.
(ii1)
2006 ERSKINE: RECENT DECLINES OF HOUSE SPARROWS 45
Table 1. Comparison of CBC HoSp means over decades, by habitats: (a) Raw data, (b) Adjusted data. Missing data added
from ratios (of all groups) to next decade. Circles with no HoSp ever omitted.
(a) Raw data 1960-69 1970-79 1980-89 1990-99 2000-02
Forest areas
7 circles 439 530 450 117 49
Coastal settlements, no farming
11 circles 875 1306 1023 404 138
Farming & woodlands nearly equal
11 circles 2120 2482 1823 949 735
Major farming areas, larger communities
8 circles 3306 4596 4953 2695 1296
Cities
4 circles 4017 3569 1664 374 157
Grand totals: 10757 12483 9913 4539 2375
(b) Adjusted data
1960-69 1970-79 1980-89 1990-99 2000-02
Forest areas
7 circles 599 476 331 58 25
Coastal settlements, no farming
11 circles 596 442 211 101 37
Farming & woodlands near equal
11 circles 1255 927 672 216 160
Major farming areas, larger communities
8 circles 2130 1840 1153 500 240
Cities
4 circles 1568 1001 283 39 19
Grand totals: 6148 4686 2650 914 48]
Extrapolated HoSp populations by decades, expand-
ed from surveyed circles across suitable habitats, and
adjusted for incomplete coverage and detection, are
shown in Table 2.
That suggested that since the 1970s total HoSp num-
bers in the Maritimes decreased greatly, probably by
three-quarters or more, with urban numbers — in New
Brunswick and Prince Edward Island — diminishing to
less than 10% of levels prevailing before 1980. Farm-
land HoSp numbers still may be one-half to two-thirds
of those present 30-40 years earlier — more likely less.
They now account for the vast majority of all HoSp
remaining in New Brunswick, Prince Edward Island,
_ and northern and eastern parts of Nova Scotia, where
| winter conditions and especially snow cover limit sur-
vival at that season. Numbers in western and southern
Nova Scotia declined to a lesser degree.
_ Discussion
Johnston and Klitz (pages 15-52 in Pinowski and
Kendeigh 1977) stated “The house sparrow seems
» clearly a product of interaction between sparrows and
» sedentary, agricultural man... This is a long-term rela-
» tionship, sufficient to allow fine-grain adaptation of the
birds to man’s way.” There seems no reason to dispute
that generalization, on the long-term, broad-scale stage
_ for which it was stated, but regional evidence suggest-
ed that “man’s way” in the Maritimes recently changed
more rapidly than did HoSp adaptations to it.
Generalization from CBC data. Rigorous conclu-
sions regarding the recent HoSp decline in the Mar-
itimes (or elsewhere) will not emerge from CBCs
alone, but improved understanding may result when
various kinds of information are combined. The scarci-
ty of hard data on this species, anywhere in North
America, is embarrassing. In the Maritimes and adja-
cent regions, the only multi-year study found that pro-
vided HoSp density indices directly was the one in
Sackville, New Brunswick (Erskine 1992b) that stim-
ulated this investigation. Knowledge of habitats and
diet here relied on casual observations, as no planned
studies had been made — and HoSp numbers in most
areas now are too low for economical study.
CBC data are poorly standardized, and adjusting
effort using party-hours addressed only a small part
of variation in CBC data (compare Dunn et al. 2005).
Regional data have not been examined rigorously, but
it was obvious that the few — mostly urban — circles
deploying large numbers of observers detected far fewer
HoSp relative to effort than were found on most counts
with fewer observers. Thus, declines in large urban
areas may have been less drastic than indicated by the
counts adjusted by party-hours, but they were impres-
sive: compare means of raw data for 1966-1970 vs.
46 THE CANADIAN FIELD-NATURALIST Vol. 120
Table 2. Expansion of counts in Table 1 (raw data) to total Maritimes population estimates (explanation in Appendix).
Habitats Extrapolated estimates
1960s 1970s 1980s 1990s 2000s
Forest+urban 1694 2045 Wiss 452 189
Coastal+urban 16944 25290 19809 7824 2673
(Farm=forest)+urban 55506 64980 47730 24846 19242
(Farm>forest)+urban 78520 109160 117630 64010 30780
Large urban 120510 107070 49920 11220 4710
farming 67013 87070 115752 71085 45020
urban 206161 221475 121072 37267 12574
Grand totals: 273174 308545 236824 108352 57594
1996-2000 at Fredericton (3379 vs. 53), Moncton (1046
vs. 142), Saint John (S501 vs. 74).
Also, CBC data often were less adequate for HoSp
than for most other bird species. When HoSp were
abundant and ubiquitous in urban areas, some CBCs
did not actually count them during the “census”, mere-
ly inserting round-number estimates as reported totals.
Where such obvious guesses recurred in several years,
those counts were excluded. A more frequent problem
was that parts of many urban areas, where HoSp were
the most expectable birds, were ignored as unproduc-
tive (of more interesting species!) until HoSp decline
was far advanced.
HoSp declines in urban areas. That the recent
HoSp decline in the Maritimes began earlier in cities
(except perhaps “Greater Halifax’) than less urbanized
situations suggested that influences formerly encour-
aging HoSp to associate with human settlements are
now less strong. Changes since 1960 in the regional
urban milieu that might have been expected to affect
HoSp numbers included:
Favourable influences
(i) winter bird-feeding increased greatly since
1960; however, small grains — preferred by
HoSp (Bent 1958, also personal observations)
— are now much less often presented (Ruther-
ford 1984), feeding stations now offering most-
ly sunflower and thistle seeds, rather than oats,
cracked corn, or “chicken scratch feed” mix-
tures;
(11) shelter in urban hedges and other shrubs in-
creased, presumably in response to increased
disposable income of home-owners;
Negative influences
(a) food from human garbage, livestock wastage and
manure decreased greatly, by disappearance of live-
stock, especially horses, and improved garbage dispos-
al, these factors all related to increasing “urbanization”;
(b) nesting and wintering shelter within buildings
decreased, through removal of dilapidated structures
and renovation of others — especially closing of open-
ings (personal observations);
(c) transmittal of pathogens (e.g. Salmonella) may
have been aided by increased transportation between
urban areas (Erskine 1980b; a HoSp die-off at Char-
lottetown, Prince Edward Island, had been reported
in the preceding winter);
(d) avian predators in towns (especially Merlin,
Falco columbarius, and Sharp-shinned Hawk, Accip-
iter striatus) increased, with Merlins breeding in urban
areas here since the 1980s; domestic cats, Felis catus,
also increased.
The recent major decline of urban HoSp in the Mar-
itimes indicated that urban areas no longer provide a
favourable environment for this species, in winter or
summer or both. Complete disappearance of HoSp in
winter in many smaller human settlements suggested a
“threshold effect’; below some population level, which
likely varies with winter severity and availability of
suitable habitats nearby, production in urban HoSp may
no longer balance winter losses in this resident species.
HoSp declines in farmlands. HoSp decline in farm-
ing areas seemed better correlated with decreases in
farms with livestock than with total numbers of farms
or areas of “improved farmland”. Loss of invertebrate
food through pesticide use, as in England (D. Sum-
mers-Smith, letter), seems plausible but is unstudied
here; chemical treatments in regional agriculture are
obvious especially in potato- and fruit-producing areas,
less so in grain production (mostly oats). Much Mar-
itimes farmland area is inaccessible to HoSp in win-
ter owing to snow cover. Livestock disappeared com-
pletely from most urban areas, and also from many
former farms now used as rural residences or “hobby
farms”. Horses are scarce generally, and cattle now are
mostly found in large feed-lot operations, where larger
birds (gulls, pigeons, corvids, starlings) often out-com-
pete HoSp — though a few of the latter persist (per-
sonal observations). Concentrations of HoSp persist in
productive farming areas, even when disappearing from
nearby towns, presumably because livestock operations
now are mainly in better farming areas.
From the viewpoint of agriculture (and general econ-_|
omy), the Maritimes are a “backwater”. Agriculture here
ranks far behind forestry, tourism, fisheries, and “gen-
2006
eral service” on economic and employment indices,
despite the local importance of potatoes and orchards.
Snow cover and related temperatures are major fac-
tors limiting agriculture (and HoSp) here. Soils and
geology underlie the picture, with limestone areas very
limited. Agricultural changes since World War II in-
volved abandonment of former marginal agricultural
lands rather than intensified use of better farmlands.
HoSp responses to habitat changes. All CBC
_ habitat groupings used herein involved combinations
of human settlements, of varying size, with nearby
habitats, among which only farmlands were regularly
used by HoSp. Human settlements formerly provided
_HoSp with both sheltered nesting places and nearby
_ feeding opportunities. After nesting, urban HoSp often
| withdrew from built-up areas to glean waste grain in
agricultural lands within a radius of 2-3 km, return-
ing to towns and villages with the approach of winter
| (personal observations; Summers-Smith 1963). Where
| little farmland exists near urban settlements, in forest-
| ed regions (e.g., Miramichi, New Brunswick) or with-
in larger cities, urban HoSp presumably depended
| year-round on feeding opportunities within the city.
Cultural changes altered urban areas more, as well
| as in different ways, than farmlands. Thus, HoSp in
areas that combined extensive farming with urban
settlements were (and are) less restricted for feeding
' than those relying year-round on urban areas. Recent
| trends suggest that HoSp might disappear within a few
| decades from most urban habitats in New Brunswick
and Prince Edward Island, where snow-cover limits
' their use of non-urban habitats in winter. They may
persist longer in “Greater Halifax”, and in some towns
in farming areas of western Nova Scotia, where snow
cover is less and winter temperatures more moderate.
| Such changes presuppose continuation of climates
| similar to recent decades; if “global warming” brings
\ served numbers across potential HoSp habitat to
regional populations (Table 2) involved much “in-
_ formed guesswork”, as hard data do not exist for most
| needed adjustments. Valuable perspective may emerge
from such simple “modeling”, even though it lacks
| rigour and precision.
| Modeling allowed comparison with the only other
' independent estimate of Maritimes HoSp populations,
_ derived from abundance indices in the Maritimes breed-
ing bird atlas (Erskine 1992a) in 1986-1990. The Atlas
) estimates used a general algorithm (C. Field and P.
_Payzant, Appendix D in Erskine 1992a) for extrapo-
lation to areas unsampled or lacking abundance indices;
_ that algorithm may not have been fully appropriate for
_ HoSp, of which density varied greatly between habi-
tats. The Atlas estimate of HoSp (140 000 pairs) was
ERSKINE: RECENT DECLINES OF HOUSE SPARROWS 47
of a similar order to that for the same period (1980s)
in Table 2 (ca. 240 000 individuals).
The HoSp situation elsewhere. No systematic stud-
ies of HoSp were available in other parts of Canada.
CBC data there — tabulated and adjusted for party-
hours as in the Maritimes (unpublished MSS) — re-
vealed no such drastic HoSp declines as seen here in
1960-2003.
Continuing availability of small grains in both rural
and urban areas of the Prairie Provinces south of the
boreal forest were well-correlated with the highest
HoSp CBC counts on the continent, with no suggestion
of declines unrelated to count efforts. British Colum-
bia CBC data were extremely variable, with generally
low numbers, as was to be expected from the varied
geography and relative scarcity of arable farming. No
long-run counts suggested continuing declines in HoSp
there. Very few long-run counts were available in north-
ern, deep-snow areas of western Canada, where most
showed no HoSp at any time.
Quebec experiences the largest average snowfalls
in Canada outside the western mountains. Most of the
few long-run (>20 years) CBCs in Quebec suggested
some decline in recent decades, from “adjusted counts”,
as reported also from independent surveys (1970-1991)
by Cyr et Larivée (1995); those declines were less ex-
treme than in the Maritimes.
In Ontario, where CBCs are more numerous than
elsewhere in Canada, HoSp (raw) numbers remained
generally stable over the decades, except in a few long-
run urban circles near the northern limits of human set-
tlement. Adjusted data, however, suggested declines,
starting in 1980s, on nearly half the long-run counts;
Sudbury showed the largest proportional decline, and
counts at Sault Ste. Marie and Thunder Bay, also north-
ern areas with much snow cover — and despite both
being ports involved in grain shipping from the Prairies,
also declined.
In the United Kingdom, where the HoSp situation
has been followed for decades, especially by Denis
Summers-Smith (1963, 2003, and letters), declines
evidently are of a similar order to those found in the
Maritimes. Although research on HoSp is limited there
too, much detailed information is available on habitat
relations of HoSp in the U.K. Summers-Smith’s con-
clusions (op. cit.) regarding causation agree well with
tentative inferences suggested here for the Maritimes
— despite the near-absence of snow cover as a limit-
ing factor in the U.K. The intensity of agriculture there
is much more comparable to that in southern Ontario
or the Prairie Provinces than to the Maritimes. At this
time it seems likely that many different factors, some
general and widespread, others applying only region-
ally, are involved in recent declines of HoSp in the
wider picture.
A recent discussion of HoSp decline in North
America (Holder 2003) drew on wide-ranging sur-
veys, including BBS and CBC, but reached few con-
48 THE CANADIAN FIELD-NATURALIST
clusions except that competition with recent House
Finch populations had not affected HoSp numbers.
Use by Holder of only CBCs reported to Audubon
limited his understanding of the Maritimes situation.
His expressed suggestion (page 65) that the HoSp sit-
uation in Canada might parallel that in the U.K. —
where very different climatic and economic regimes
prevail — seems unlikely.
Acknowledgments
Thanks to the many observers and compilers of
CBCs whose data were used in this study, and also to
publishers and editors of magazines (named in text) in
which those reports were presented. Denis Summers-
Smith (U.K.) and Jenny De Laet (Belgium) comment-
ed very helpfully on an earlier version of the MS.
Reviewers for this journal provided much-needed en-
couragement as well as helpful suggestions for im-
provement of the text.
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Christie, D. S. (1985 and following years). [general comments
in annual “Christmas Counts” articles] in New Brunswick
Naturalist.
Cyr, A., et J. Larivée. 1995. Atlas saisonnier des oiseaux
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La Société de Loisir Ornithologique de |’’Estrie, inc. [pages
660-661].
Downes, C. M., and B. T. Collins. 2002. The Canadian Breed-
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Dunn, E. H., C. M. Francis, P. J. Blancher, S. R. Drennan,
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the Birds of North America, number 12. Edited by A.
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Rutherford, L. A. 1984. Effects of winter feeding on bird
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um. 164 pages.
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Received 14 February 2005
Accepted 13 September 2005
2006 ERSKINE: RECENT DECLINES OF HOUSE SPARROWS 49
Appendix
(i) Factors, by habitat groupings, for expansion from long-run CBC circles to total areas in Maritimes. Note: Forest areas
support very few or no HoSp, so those areas were ignored, except where CBCs had been done.
Habitats Number long-run Potential CBC circles
CBC circles + other potential habitat
| Forest + urban 7 add 8
| Coastal + urban 11 add 60
| (Farm = forest) + urban 1] add 37
(Farm > forest) + urban 8 add 1]
| Large urban 4 add 4
(ii) Expansion factors, by habitat groupings, for incomplete coverage of suitable habitats in circles and incomplete detec-
tion of HoSp in areas covered.
| Habitats Coverage Detection
| Forest + urban 1.2 1.5
» Coastal + urban 2.0 eS
(Farm = forest) + urban 3.0 2.0
(Farm > forest) + urban 5.0 2.0
) Large urban 5.0 3.0
(iii) Segregation of HoSp numbers among habitats, where appropriate. Forest and coastal habitats omitted as not used by
HoSp; thus only 3 & 4" categories above required segregation, with proportions varying over time.
Decades 1960s 1970s 1980s 1990s 2000s
| Farm:urban ratio 50:50 50:50 70:30 80:20 90:10
First Record of the Southern Red-Backed Vole, Clethrionomys
gapperi, in Newfoundland: Implications for the Endangered
Newfoundland Marten, Martes americana atrata
BRIAN J. HEARN, ! JOHN T. NEVILLE,! WILLIAM J. CURRAN,! and DEAN P. SNow?
‘Natural Resources Canada, Canadian Forest Service — Atlantic Forestry Centre, P.O. Box 960, Corner Brook, Newfoundland
A2H 6J3 Canada
> College of the North Atlantic, Fish and Wildlife Program, P.O. Box 822, Corner Brook, Newfoundland A2H 6H6 Canada
Hearn, Brian J., John T. Neville, William J. Curran, and Dean P. Snow. 2006. First record of the Southern Red-Backed Vole,
Clethrionomys gapperi, in Newfoundland: implications for the endangered Newfoundland Marten, Martes americana
atrata. Canadian Field Naturalist 120(1): 50-56.
We report on the first capture of the Southern Red-backed Vole (Clethrionomys gapperi), the eleventh non-native terrestrial
mammal established on the island of Newfoundland over the last 150 years. Red-backed Voles may have been accidentally
introduced by unknown sources in pulpwood imports or may have been deliberately introduced in an attempt to augment the
depauperate small mammal fauna as a vigilante recovery effort for the endangered Newfoundland Marten (Martes americana
atrata). We anticipate significant utilization of the Red-backed Vole as prey by both Newfoundland Marten and Red Fox
(Vulpes vulpes) with associated demographic responses within and between these species. Red-backed Voles will likely change
habitat utilization patterns for the endemic subspecies of Meadow Vole, Microtus pennsylvanicus terraenovae.
Key Words: Southern Red-backed Vole, Clethrionomys gapperi, Marten, Martes americana atra, introduced species, New-
foundland.
The island of Newfoundland has only 13 resident
native species of terrestrial mammals (Bangs 1913;
Cameron 1958: Dodds 1983; Table 1). A fourteenth
species, the Newfoundland Wolf (Canis lupus beothu-
cus) was extinct by the early 20" century (Allen and
Barbour 1937), and one of the three native mustelids,
the Newfoundland Marten (Martes americana atrata)
is currently listed by the Committee on the Status of
Endangered Wildlife in Canada as endangered (Lemon
1996*). As well, the indigenous community of terres-
trial mammals in Newfoundland has a skewed compo-
sition, with a disproportionate number of predators and
few prey species; historically, only one microtine, the
native subspecies of Meadow Vole (Microtus pennsyl-
vanicus terraenovae) occurred on the island (Dodds
1983). Moreover, Cameron (1958) considered 10 of the
14 native species of terrestrial mammals as endemic
subspecies; recent genetic analysis has confirmed the
subspecies classification of the Newfoundland Marten
(Kyle and Strobeck 2003). However, over the last 150
years, an additional nine species of mammals have been
either intentionally or accidentally introduced to the
island (Dodds 1983; Gould and Pruitt 1969; Northcott
1974*; Northcott et al. 1974; Payne 1976; Table 1). A
tenth species, the Eastern Coyote (Canis latrans) be-
came established on the island of Newfoundland in
1985, likely after crossing on sea ice from mainland
Nova Scotia (Parker 1995). Herein, we report the first
record of the eleventh non-native mammal established
on the island of Newfoundland, the Southern Red-
backed Vole (Clethrionomys gapperi), and discuss the
circumstances and implications of this recent introduc-
tion, particularly with reference to the Newfoundland
Marten.
Field Sites and Sampling
Between 2 and 5 November 1999, as part of a larger
study investigating the demography and ecology of the
Newfoundland Marten, we indexed small mammal pop-
ulations on four sites inside the Pine Marten Study
Area (PMSA) in southwestern Newfoundland (48°37'N,
57°53'W). The PMSA is a 2078-km* wildlife reserve,
that was created in 1973 by the Newfoundland and La-
brador Wildlife Division, for the protection of the New-
foundland Marten (Snyder and Bissonette 1987). Sites
1-3 were overmature (81+ years), Balsam Fir (Abies
balsamea) stands, whereas Site 4 was a regenerating
(10-15 yrs) Balsam Fir stand that regenerated after an
outbreak of Spruce Budworm (Choristoneura fumifer-
ana) and Hemlock Looper (Lambdina fiscellaria fiscel-
laria) in 1987. Site 4 contained a mix of young Balsam
Fir and early successional species, such as White Birch
(Betula papyrifera) and Pin Cherry (Prunus pensylvani-
ca), had a significant number of dead Balsam Fir
snags, and had no canopy. Each site was trapped with
100 snap traps on 3 consecutive nights. Traps were bait-
ed with peanut butter, and placed in pairs at 50 trapping
stations, spaced 15 m apart, yielding a 735-m transect,
following the methods previously used by Thompson
and Curran (1995). All captures were recorded and col-
lected for further analysis.
Forty small mammals were collected on the four sites
over 1200 trap nights: 8 Meadow Voles, three Deer Mice
(Peromyscus maniculatus), 10 Masked Shrews (Sorex
cinereus), and 19 Southern Red-backed Voles. How-
50
2006
TABLE |. Native and non-native ‘? terrestrial mammals on the island of Newfoundland prior to the discovery of the Southern
Red-backed Vole!.
Order
Insectivora
Chiroptera
Family
Soricidae
Vespertilionidae
Lagomorpha Leporidae
Rodentia Cricetidae
Scuridae
Castoridae
Muridae
Carnivora Mustelidae
Felidae
Canidae
Ursidae
Arctiodactyla Cervidae
Scientific Name (Authority)
Sorex cinereus acadicus (Gilpin)
Myotis lucifugus lucifugus (Le Conte)
Myotis septentrionalis (van Zyll de Jong)
Lepus arcticus bangsti (Rhoads)
Lepus americanus struthopus (Bangs)
Microtus pennsylvanicus terraenovae (Bangs)
Peromyscus maniculatus (Wagner)
Ondatra zibethicus obscurus (Bangs)
Tamiasciurus hudsonicus ungavensis (Anderson)
Tamias striatus lysteri (Richardson)
Castor canadensis caecator (Bangs)
Rattus norvegicus norvegicus (Berkenhout)
Mus musculus domesticus (Rutty)
Mustela erminea richardsonii (Bonaparte)
Mustela vison (Schreber)
Martes americana atrata (Bangs)
Lontra canadensis degener (Bangs)
Lynx lynx susolanus (Bangs)
Vulpes vulpes deletrix (Bangs)
Canis lupus beothucus? (Allen and Barbour)
Canis latrans var. (Lawrence and Bossert)
Ursus americanus hamiltoni (Cameron)
Rangifer tarandus terraenovae (Gmelin)
Alces alces americana (Clinton)
HEARN, NEVILLE, CURRAN, AND SNOW: RED-BACKED VOLE IN NEWFOUNDLAND
Common Name
Masked Shrew *
Little Brown Bat
Northern Long-eared Bat
Arctic Hare
Snowshoe Hare *
Meadow Vole
Deer Mouse *
Muskrat
Red Squirrel °
Eastern Chipmunk ~
Beaver
Norway Rat *
House Mouse *
Short-tailed Weasel
Mink *
Newfoundland Marten?
Otter
Canada Lynx
Red Fox
Eastern Timber Wolf
Eastern Coyote ~
American Black Bear
Woodland Caribou
Moose
'Several other species are either seasonal visitors (e.g., Arctic Fox (Alopex lagopus), Polar Bears (Ursus maritimus)) or have
been introduced to offshore islands (e.g., Bison (Bison bison), Bank Vole (Clethrionomys glareolus suecicus)). Adapted from
Cameron (1958) and Dodds (1983).
*Subspecies designation confirmed via recent genetic analysis (Kyle and Strobeck 2003).
3Extinct
ever, Red-backed Voles were captured on only two of
the four sites (Sites 3 and 4). The two sites where Red-
backed Voles were captured were approximately 6 km
apart, but on opposite sides of Little Grand Lake. Move-
ment between these two sites would require an over-
land dispersal of at least 11 km, suggesting that the spe-
cies is well established in the area. At least one of the
collected females had obvious placental implantation
scars on the uterine horns, indicating recent breeding.
Species identification was confirmed by pelage and
dental characteristics (Banfield 1974).
Discussion
Sites 1—3 had previously been snap trapped for small
mammals for 4 years, between 1990 and 1993 (Thomp-
son and Curran 1995). Additionally, Site 4, as well as the
general area encompassing all four of our sampled sites,
was extensively live trapped from 1993 to 1997, as part
of a larger study investigating Marten, small mammal
communities, and forest structure (Adair 2003). Neither
of these two previous trapping efforts captured Red-
backed Voles, suggesting that the species had recently
become established, most likely sometime during or after
1998. Moreover, a more extensive small mammal survey,
conducted two weeks before this survey in the Red Indian
Lake area, approximately 35 km east of our four trap-
ping sites, using identical methods but involving 16
sites and >4900 trap-nights, captured 116 small mam-
mals, but no Red-backed Voles. Autumn (October)
small-mammal surveys in the Red Indian Lake area,
repeated annually since 1999, have subsequently doc-
umented the arrival (2001) and eastward dispersal of
the Red-backed Vole in southwestern and south-central
Newfoundland (B. J. Hearn, unpublished data).
Two explanations for this introduction are possible.
Red-backed Voles may have been accidentally intro-
duced in pulpwood or pulp chips imported through the
port of Stephenville, (approximately 38 km SW of our
sampling sites). Alternatively, the species may have been
deliberately introduced by unknown persons to increase
prey diversity and abundance for the Newfoundland
Marten. Because Red-backed Voles are a common prey
item in the diet of American Marten elsewhere (Mar-
tin 1994), this management option was debated during
preparation of the Newfoundland Marten Recovery
Plan (Forsey et al. 1995*), resulting in a divergence of
opinions among local stakeholder groups. Although
ultimately rejected by the Newfoundland Marten Re-
covery Team as an ethical recovery action, the limited
prey base and the ecological consequences of introduc-
ing the Red-backed Vole were publicly discussed in
1998, during a two-day symposium on issues concern-
Sy THE CANADIAN FIELD-NATURALIST
ing the Newfoundland Marten. Given the coincidence
in timing (ca. 1998) and spatial distribution of this dis-
covery (inside the PMSA), we suggest that it is likely
that Red-backed Voles were deliberately introduced.
Regardless of the source of this introduction, the
direct and indirect effects of this new species on com-
munity structure in general, and the Newfoundland
Marten in particular, warrant investigation. As an exam-
ple, Cameron (1958) reported that Newfoundland
Meadow Voles are less selective in their habitat require-
ments than mainland Meadow Voles, and atypically,
occupy forested habitats — likely due to the historical
absence of Red-backed Voles (Cameron 1958;
Clough 1964; Cameron 1965; Morris 1969; Folinsbee
et al. 1973). Consequently, the introduction of the Red-
backed Vole will probably result in a niche contraction
for the endemic Meadow Vole, and we predict that
Meadow Voles will be displaced from forested areas
and restricted to their preferred habitat (1.e., moist
meadow-like areas (Folinsbee et al. 1973)). Preliminary
analysis of our 1999-2006 small-mammal trapping
data from the RIL area is documenting an irrupting
population of Red-backed Voles (>3000 Red-backed
Voles captured since 2001), and an essential elimina-
tion of Meadow Voles from forested sites where we
had previously captured Meadow Voles.
Consequently, the positive effect of increased prey
diversity and biomass for the Newfoundland Marten
will be offset, to some degree, by a decrease in distri-
bution and densities of Meadow Voles, which have
been a historically important prey species (Bateman
1986; Drew 1995; Gosse and Hearn 2005; Tucker
1988). Further, indirect effects of the Red-backed Vole
on Newfoundland Marten could occur as a result of the
impact of this species on local Red Fox (Vulpes vulpes)
populations. Given the depauperate prey base in New-
foundland, Red Fox and Marten display considerable
dietary overlap, and Red Fox are the most important
natural predator of Newfoundland Marten (B. J. Hearn,
unpublished data). Consequently, Marten may be nega-
tively affected if the introduction of Red-backed Voles
has a positive effect on Red Fox demography, leading
to increased intraguild predation on Marten (Lindstr6m
et al. 1995) or increased Marten—Fox competition for
food (Kurki et al. 1997; Lindstrém et al. 1994; Marc-
strom et al. 1988). Pine Marten (Martes martes) popu-
lations in Scandinavia increased in the 1980s, follow-
ing a decline of Red Fox due to a sarcoptic mange
epidemic, as a direct result (hypothesized) of reduced
predation by Red Fox (Lindstrom et al. 1994; Helldin
1998) or reduced competition for Microtus prey
(Storch et al. 1990). Although counterintuitive to con-
ventional competition theory, increases in food re-
sources (prey availability) can actually increase intra-
guild predation by supporting increased populations of
the superior competitor. Parallel circumstances have
been described for other threatened or endangered car-
nivores elsewhere (e.g., Coyote predation on Swift
Vol. 120
Foxes (Vulpes macrotis) (Sovada et al. 1998), Coyote
predation on San Joaquin Kit Foxes (Vulpes macrotis)
(Ralls and White 1995; White and Garrott 1997;
Cypher and Spencer 1998), and Lion (Panthera leo)
and Spotted Hyena (Crocuta croenta) predation on
African Wild Dogs (Lycaon pictus) (Creel 2001; Creel
et al. 2001)). African Wild Dog densities are actually
lowest where density of their major prey is highest, due
to intraspecific predation (Mills and Gorman 1997). It
is noteworthy that Marten live-trapping captures on the
Northern Peninsula of Newfoundland increased in 2005
following a recent (2002-2004) rabies outbreak (Whit-
ney 2004*) that virtually eliminated Red Fox from the
local landscape (M. McGrath, personal communication).
Changes in prey availability clearly affect densities
of carnivores, improving demographic performance
(1.e., increased reproductive output and survival (Fuller
and Sievert 2001). Furthermore, decreased food re-
sources have been associated with reduced population
density or trapper success, enlarged home ranges, and
lower reproductive performance of American Marten
(Bulmer 1974; Fryxell et al. 1999; Poole and Graf
1986; Simon et al. 1999; Thompson and Colgan 1987).
The reduced prey base for Marten in Newfoundland
compared with that available on the mainland portion
of the province (Labrador) has been suggested as a
likely explanation for the difference in the demograph-
ic health of these two populations (Bissonette et al.
1988*). By comparison, Labrador has 17 small mam-
mal species, including the Red-backed Vole (Tucker
1988) and Marten populations appear capable of sup-
porting a commercial harvest; 500-800 Marten were
harvested annually between 1995 and 1998 (Simon et
al. 1999).
However, an increase in vole biomass may not lead
to an immediate increase in Marten densities if territo-
ry size in Newfoundland is adjusted to the lowest (or
historical) level of prey availability. Such appears to be
the case for European Badger (Meles meles) in Scot-
land (Kruuk and Parrish 1982), Kit Foxes in California
(White and Ralls 1993), and some Coyote populations
in the western United States (Mills and Knowlton
1991). Interestingly, Payer (1999) in the largest Ameri-
can Marten study reported to date (n > 140 marten
ranges), found no difference in mean home-range area
for American Marten in Maine between 1995-1998,
despite a three-fold decline and recovery in small
mammal populations. Thompson and Colgan (1987,
1990) suggested that, based on energetics, Marten in
Ontario cannot survive exclusively on small rodents
during late winter, and further, take small mammals
opportunistically as they hunt for larger prey (i.e.,
Snowshoe Hare). Moreover, Gosse and Hearn (2005)
reported that Snowshoe Hare provide >90% of the
caloric intake of Newfoundland Marten in winter, the
most energetically stressful period annually (Buskirk et
al. 1988). As cautioned by Fuller and Sievert (2001), |
the correlation between food and density-related demo- |
2006 HEARN, NEVILLE, CURRAN, AND SNOW: RED-BACKED VOLE IN NEWFOUNDLAND 53
59°0'0"W 58°30'0"W 58°0'0"W
49°0'0"N
48°30'0"'N
* Trap Sites
48°0'0"'N
=a kilometers
0 5 10 20 30
58°30'0"W 58°0'0"W
57°30'0"W 57°0'0"W
49°0'0"N
48°30'0"N
48°0'0"'N
57°30'0"W 57°0'0"W
Ficure |. Map of southwestern Newfoundland showing location of Southern Red-backed Vole (Clethrionomys gapperi) cap-
tures in 1999, and capture sites relative to other areas mentioned in text; A indicates location of first Southern Red-
backed Vole capture in Red Indian Lake area in 2001. Inset map shows the location of areas relative to the island of
Newfoundland.
graphic parameters is statistically broad, and many site-
specific factors (e.g., prey availability, habitat patchi-
ness, habitat selection under foraging versus predation-
risk trade-offs), influence home-range characteristics
and carnivore density.
Newfoundland Marten have been geographically
and reproductively isolated from mainland Marten
populations for the last 7000 years (South 1983),
resulting in a genetically distinct subspecies amidst a
much more genetically homogenous Canadian main-
54
land population (Kyle and Strobeck 2003). Additional-
ly, Newfoundland Marten are large (males = 1254 g,n=
122; B. J. Hearn, unpublished data), and have extreme-
ly large home-range requirements (males = 30.8 km?,
n= 43; B. J. Hearn, unpublished data) compared with
nearby mainland populations in Quebec (males = 937 g,
n= 67; 74 km’, n = 40; Potvin and Breton 1997) and
Maine (males = 776 g, n = 23, Katnik 1992; 4.04 km”,
n = 96; Payer 1999). Collectively, these population
characteristics suggest that the Newfoundland Marten
is a product of a unique ecological setting and evolu-
tionary selective factors operating on a geographically
and reproductively isolated island population. Thus,
it seems unlikely, given the historical ecological set-
ting, that Newfoundland Marten spatial requirements
evolved to access or select habitat (Hearn et al. 2005*)
based on utilizing small mammals as prey.
We predict that the establishment of the Red-backed
Vole on the island of Newfoundland will cause direct
and indirect community-level effects in general and for
Newfoundland Marten in particular (e.g., increased uti-
lization of the Red-backed Vole as prey by Newfound-
land Marten and Red Fox and associated demographic
responses and interactions, more restricted habitat uti-
lization by Newfoundland Meadow Voles, and chang-
ing utilization and dispersal of seeds by introduced
Red-backed Voles and consequently changing plant
regeneration patterns). The direct and indirect effects
of previous introductions of non-native species to the
island of Newfoundland have been previously described
for predator—prey interactions (Bergerud 1971, 1983),
plant successional patterns (Thompson et al. 1992,
Thompson and Curran 1993), and species interactions
(Benkman 1993).
Additional small mammal field surveys are being
conducted to document the changing distribution of
Red-backed Voles on the island and may offer addi-
tional information about the mechanism by which this
species was introduced. In this regard, genetic analysis
may be helpful in determining the source population
for this introduction. Future field studies, in areas where
Newfoundland Marten spatial characteristics have been
well documented (Hearn et al. 2005*) are being con-
sidered to assess the effects of this new prey species
on Newfoundland Marten population characteristics.
Acknowledgments
Funding for this work was provided by Natural
Resources Canada, Canadian Forest Service; the New-
foundland and Labrador Department of Environment
and Conservation — Inland Fish and Wildlife Division;
Newfoundland and Labrador Department of Forest
Resources and Agrifoods — Forestry Division; Corner
Brook Pulp and Paper; Abitibi Consolidated; and the
Western Newfoundland Model Forest through support
of a five-year research project concerning Newfound-
land Marten. We thank D. J. Harrison, B. Krohn, J.
Loo, and two anonymous reviewers for comments on
this manuscript.
THE CANADIAN FIELD-NATURALIST
Vol. 120
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Received 21 February 2005
Accepted 15 March 2006
Long-range Homing by an Adult Female Black Bear, Ursus americanus
L. J. LANDRIAULT!, M. N. HALL?, J. HAMR?, and F. F. MALLoryY4
‘Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada
Present address: P.O. Box 521, Dowling, Ontario POM 1RO Canada
2Ontario Ministry of Natural Resources, Sudbury, Ontario P3E 1G7 Canada
3Cambrian College, Sudbury, Ontario P3A 3V8 Canada
‘Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada
Landriault, L. J.,M.N. Hall, J. Hamr, and F. F. Mallory. 2006. Long-range homing by an adult female Black Bear, Ursus amer-
icanus. Canadian Field-Naturalist 120(1): 57—60.
An adult female Black Bear was repeatedly captured and relocated as a result of nuisance behaviour. The relocation distances
ranged from 40 km to 389 km (mean = 152 km, n = 6). She homed successfully from all relocations, even when accompanied
by young-of-the-year. Differential homing ability among bears may depend on first homing from a short relocation, facilitating
subsequent responses to longer distance relocations.
Key Words: Black Bear, Ursus americanus, homing, distance, relocation, translocation, Ontario.
The ability of animals to return home when removed
from familiar territory has been recorded for several
different species including: pigeons, Columba spp. (Papi
1992); Deer Mice, Peromyscus maniculatus (Bovet
1968; Teferi and Millar 1993); Red Squirrels, Tamias-
ciurus hudsonicus (Bovet 1995); Raccoons, Procyon
lotor (Belant 1992); Wolves, Canis lupus (Fritts et al.
1984); and Brown Bears, Ursus arctos (Miller and
Ballard 1982). The use of relocation as a management
tool for nuisance bears has provided researchers with
the opportunity to study the homing behaviour of a
large mammal. Several studies conducted on Black
Bears (Ursus americanus) have determined that a high
proportion of relocated bears return to the capture area
(Harger 1970; Alt et al. 1977; Rutherglen and Herbi-
son 1977; McArthur 1981; Rogers 1986a; Shull 1994),
with adult bears more likely to return to the capture
area than juveniles (Harger 1970; Rogers 1986a; and
Landriault 1998). During a four-year study of relocat-
ed nuisance Black Bears in Sudbury, Ontario (46°N,
81°W), one bear was successful in homing over an
exceptionally long distance (Landriault 1998). This
report describes her relocation and homing history, and
briefly discusses factors that may affect homing suc-
cess in relocated bears.
Animals were captured in large barrel traps or with
the use of immobilization agents delivered via a dart
gun or jab stick. In accordance with the study protocol,
nuisance Black Bears were immobilized using a mix-
ture of ketamine hydrochloride (Ketaset®) and xylazine
hydrochloride (Rompun®) at a dosage of 4.4 mg/kg and
ae mg/kg, respectively. Captured bears were weighed,
various body measurements obtained, and the animals
were ear-tagged (Kurl-lock metal tags, Ketchum Man-
ufacturing Inc, Brockville, Ontario). A premolar was
extracted for ageing purposes. Some animals were fitted
with a very high frequency (VHF) radio-transmitter
collar (148 to 152 MHz range, Lotek Inc., Newmarket,
Ontario). If re-captured, most bears were immobilized
again to assess changes in condition by means of growth
rates for juveniles and changes in weight for adults. The
majority of capture and release locations were estimat-
ed using topographical maps or a hand-held global
positioning system (GPS) unit; however, the only infor-
mation available for the first and third captures des-
cribed in this note was the township name. In these
cases the center of the township was used to estimate
release and homing distances. In the study area, town-
ships were 10 x 10 km such that the center of the town-
ship could only be a maximum of 7 km from the true
location. Distances presented in this report are straight-
line measurements and do not take topographical fea-
tures into consideration. Returning within 20 km of the
capture site was defined as successful homing (Lan-
driault 1998).
Adult female Black Bear LRO4 was captured as a
nuisance animal and relocated a total of 6 times (Table |
and Figure 1). In all cases she was captured because
she was foraging in improperly stored refuse at resi-
dences in suburban areas. She was initially captured
as a nuisance animal in June of 1994. At this time,
she was estimated to be 9 years of age based on the
examination of premolar cementum annuli (Johnston
et al. 1987). She was ear-tagged and relocated approx-
imately 40 km south of the capture site. LRO4 was
recaptured in June of 1995, an estimated 6 km from
her first capture location. She was lactating; however,
no cubs were found. She was fitted with a radio-collar
and relocated a second time, 105 km to the north.
LRO4 was subsequently recaptured in September of
1995, 7 km from her second capture location, and was
accompanied by three cubs. The four bears were relo-
cated as a family unit, approximately 112 km. LRO4
was captured again in June of 1996, within 17 km of
her previous capture location, accompanied by three
yearlings. LRO4 and two of the yearlings were relo-
SY)
58 THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Relocation history of nuisance Black Bear LR04, including relocation distance, bearing, and proximity of consecu-
tive capture and release sites. All capture locations are in the suburban areas of the City of Sudbury, Ontario. Body weight
information has been included as a means to assess the general condition of the animal.
Capture Body Proximity of
date weight capture site to
(kg) previous capture
site (km)
20 June 1994 65
14 June 1995 76 6
18 September 1995 147 7
8 June 1996 67 13
5 July 1996 82 10
1 October 1997 140 <1
cated 169 km. LRO4 was subsequently captured a fifth
time in July of 1996, within 11 km of her previous cap-
ture site. She was not accompanied by any offspring.
On this fifth relocation she was transported 389 km to
the northwest and was recaptured in October of 1997,
within 7 km of her previous capture location. At this
time she weighed approximately 140 kg and was re-
located 100 km.
From 1994 to 1997, adult female LRO4 was relo-
cated a total of six times, with relocation distances
ranging from 40 to 389 km. She homed successfully
in all cases and the maximum distance between any
two capture sites was 10 km. The longest homing dis-
tance observed for this animal (389 km) appears to be
a record distance for Black Bears. The true distance
travelled by bear LR04 to return to her home range is
expected to be much greater than 389 km, due to vari-
ations in topography and the probable deviations from
straight-line homing as a result of landscape features,
human habitation, foraging, and interactions with other
animals. Other observations of long-range homing in
relocated Black Bears include animals returning from
distances of 99 km (Rutherglen and Herbison 1977)
and 229 km (Harger 1970) on the Upper Peninsula in
Michigan, 103 km in the Rocky Mountains of British
Columbia (Erickson and Petrides 1964), and 182 km
in Central Ontario (Landriault 1998). Although all of
these observations are from adult animals, in some in-
stances juveniles can home from long distances. As
part of the same study, a juvenile female homed from
a relocation distance of 92 km (Landriault 1998). The
differential homing ability of adults versus juveniles
has been attributed to an established home range and to
experience gained during seasonal foraging excursions
(Anderson et al. 1977; Rogers 1986b, Rogers 1987a).
Juvenile male Black Bears generally disperse be-
tween 2 and 4 years of age, prior to establishing home
ranges. In contrast, juvenile females generally take up
residence in their maternal home range (Rogers 1987b;
Schwartz and Franzmann 1992). It has been hypothe-
sized that bears without established home ranges have
limited homing success due to a lack of effort rather
Proximity of
Relocation release site to Bearing to
distance previous release relocation
(km) site (km) site
40 166°
105 141 B25 a
i 6 825%
166 76 343°
389 222 330°
100 290 323%
than ability (Anderson et al. 1977; Rogers 1986b). As
predicted by this hypothesis, lower homing success
has been observed in juvenile males than in adults and
juvenile females (Harger 1970; Rogers 1986a; and
Landriault 1998). Adult females would benefit from
a strong desire to return to their established home range
where they have been able to meet the nutritional re-
quirements necessary for reproduction (Rogers 1976;
LeCount 1982; Elowe and Dodge 1989; Kolenosky
1990) and have successfully avoided fatal conflicts
for them and/or their young, with local conspecifics
(Jonkel and Cowan 1971; Tietje et al. 1986).
Experience would also be expected to increase an
animal’s homing success. Many Black Bears cover ex-
tensive area during yearly foraging excursions (Rogers
1987a) and these foraging excursions may allow bears
to develop orientation and homing skills, resulting in
increased homing success with age. Many young bears
may not have gained enough experience to home suc-
cessfully from significant distances.
It is unknown whether bear LRO4 would have
homed from 389 km had she been relocated that dis-
tance after her initial capture. It is possible that repeat-
ed relocations result in training individual bears to
home from long distances. Black Bears that home suc-
cessfully after the first relocation are likely to be suc-
cessful after all subsequent relocations of increasing |
distance (Blanchard and Knight 1995; Landriault'
1998), suggesting that homing ability in bears is en--
hanced with experience. Homing pigeons are common--
ly trained by gradually increasing the displacement|
distance from the home loft, and young birds displaced!
too far early in their training do not home successfully;
(Papi 1992). The successful return from a relatively)
short distance appears to enhance homing effort and/
or skill, and increases the chance of success from longer
distances.
The desire for adults to return to an established home)
range, combined with experience acquired in navigat-
ing over long distances, appears to explain why adult!
bears are more adept at homing. The inadvertent train-)
ing of bear LR04 by gradually increasing the reloca-))
2006
100 0
EE _
LANDRIAULT, HALL, HAMR, AND MALLORY: LONG-RANGE HOMING 59
© Lake Huron a1 |
100 Kilometers
SIGURE 1. Capture and release locations for nuisance Black Bear LRO4. Black circles represent the capture locations and
grey circles represent release locations. The thin black lines on the map correspond to the two-lane paved provincial
highways along the relocation routes, while the shaded squares are cities and small towns along these routes. Irregu-
lar shaded areas are lakes.
ion distances may explain how she was able to home
successfully over a longer distance than any previous-
y reported for a Black Bear.
Acknowledgments
Funding for this research was provided by The Fed-
sration of Ontario Naturalists, the Ontario Ministry of
Natural Resources, the Ontario Federation of Anglers
d Hunters, the Bear Alliance-Canada, Laurentian
Jniversity, and Cambrian College.
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Received 23 February 2005
Accepted 26 September 2005
Early Ontogenetic Diet in Gray Wolves, Canis lupus, of Coastal
British Columbia
HEATHER M. BryAN!, Curis T. DARIMONT! 4, THOMAS E. REIMCHEN!, and PauL C. PagueT?=
'Department of Biology, University of Victoria, P.O. Box 3020, Station CSC, Victoria, British Columbia V8W 3N5 Canada
*Raincoast Conservation Society, P.O. Box 26, Bella Bella, British Columbia VOT 1B0 Canada
Faculty of Environmental Design, University of Calgary, Calgary, Alberta T2N 1N4 Canada
‘Corresponding author: cdarimon@uvic.ca
Bryan, Heather M., Chris T. Darimont, Thomas E. Reimchen, and Paul C. Paquet. 2006. Early ontogenetic diet in Gray
Wolves, Canis lupus, of coastal British Columbia. Canadian Field-Naturalist 120(1): 61-66.
Within populations, different age classes often consume dissimilar resources, and provisioning of juveniles by adults is one
mechanism by which this can occur. Although the diet of Gray Wolves (Canis lupus) has been studied extensively, the diet
of pups is largely unknown. We examined faeces deposited by altricial pups and adult providers during the first two months
following birth at two den sites over two years on the central coast of British Columbia, Canada. Pups and adult wolves con-
sumed similar species, and Black-tailed Deer (Odocoileus hemionus) constituted most of the diet for both age groups. Pup
and adult diet, however, diverged. Specifically, adult deer occurred significantly less frequently in the diet of pups than in
the diet of adult wolves, which suggests that adults selectively provisioned pups. We speculate that this may relate to adaptive
strategies of adult wolves to provide their offspring with food of optimal nutritional value or reduced parasitic burden, and/or
logistic factors associated with provisioning such as prey transportability and availability.
Key Words: Gray Wolf, Canis lupus, Sitka Black-tailed Deer, Odocoileus hemionus, provisioning, pups, diet, British Columbia.
Different age classes within a species often consume
dissimilar resources, increasing population niche width
and decreasing intraspecific competition (Polis 1984;
Bolnick et al. 2003). Provisioning of young by adults
is one mechanism whereby differences in diet between
young and adults may occur (Markman et al. 2002).
Foragers that select foods to optimize growth and
development of their offspring may have a selective
advantage over those that provision their young oppor-
tunistically (Krebs and Avery 1984; Wright et al. 1998).
Providers may select resources that are easy to trans-
port (Molsher et al. 2000) or that have nutritional qual-
ities important for developing juveniles (Krebs and
Avery 1984). Furthermore, providers may alter their
behaviour while rearing young to reduce exposure of
their offspring to parasites (Christe et al. 1994; Mer-
‘ila and Allander 1995; Tripet et al. 2002).
~ Young Gray Wolves (Canis lupus) rely predomi-
nantly on food killed and delivered by closely related
adult providers for many months (Paquet and Carbyn
2003). Although diets of adult wolves have been stud-
ied extensively, the diet of juvenile wolves has not been
well documented in any system (Paquet and Carbyn
2003).
_ Wolves of coastal British Columbia (BC) have a
potential niche that includes up to 15 mammal species
(Darimont and Paquet 2000*, 2002; Darimont et al.
2004), which likely differ in transportability, nutritional
characteristics, and parasite load. Herein, we investi-
gated potential age class differences in diet by exam-
ining faeces of pups and adults deposited at den sites
‘within the first two months following birth for two
social groups over two breeding seasons. Our objectives
were to describe the diet of adult and juvenile wolves
during the denning season and examine whether the
diet of pups differed from that of adults. Given the ubig-
uity of evidence from a variety of other systems, we pre-
dicted that adult wolves would selectively provision
food resources to developing pups.
Study Area
We collected faeces from two home sites (i.e., repro-
ductive areas) on the central coast of British Colum-
bia in July of 2001 and 2002. The study area has been
described well elsewhere (Darimont and Paquet 2002;
Darimont et al. 2004). One site was located on Chat-
field Island (area: 48 km?; distance from mainland:
1 km; centre island location: 52°16'N, 128°05'W) and
the other on Yeo Island (98 km?; 0.25 km; 52°21'N,
128°08'W). Both sites were <100 m ASL (above sea
level). During the study, social groups were composed
of 3-7 pups and 3-8 adults. Potential prey species in-
clude Sitka Black-tailed Deer (Odocoileus hemionus
sitkensis), mustelids, ursids, birds, rodents, fish, and
marine mammals and invertebrates (Darimont and
Paquet 2000*, Darimont et al. 2004).
Materials and Methods
Scat collection and analysis
Both sites were on or within 50 m from transects
monitored for faeces during spring (April/May) before
arrival of wolves or birth of pups, and summer (July)
after wolves had left the areas. Thus, we are confident
that faeces collected during July represent resources
61
62
acquired during May and June, at which time pups
were approximately 2-8 weeks old. This period also
corresponds to the birth pulse of many prey in the area,
including deer (Shackleton 1999).
In all cases, wolves had moved to another site by
July, so we sampled all or most faeces that surround-
ed the abandoned sites (within ~100 m). Based on
diameter, we subjectively classified faeces (n = 479)
as those deposited by pups or adults. In July, pup fae-
ces are easy to distinguish from those of adults and
there is no overlap in diameter (Weaver and Fritts 1979).
The few faeces that were mostly liquid or suggestive
of severe weathering (i.e., amorphous shape or irreg-
ular surface; n ~ 20), which we felt could not be accu-
rately classified, were discarded.
Faeces were autoclaved, washed, and dried so that
only macroscopic components such as hair, bone, teeth,
and hooves remained (Cuicci et al. 1996). To identify
mammalian prey items, we examined the entire sam-
ple with the naked eye to detect hair clumps that dif-
fered in colour, size and/or texture from others, and
other clues such as teeth, claws and hooves. Subse-
quently, we microscopically examined five sub-sam-
ples, randomly selected via a grid system in the sample
tray, which typically contained five to 10 hairs each.
We identified species by comparing with a reference
hair collection, as well as cuticular scale imprints using
dissecting and compound microscopes and dichoto-
mous keys (Mayer 1952; Adorjan and Kolenasky 1969).
We differentiated between adult deer and fawn hair
using a combination of diameter and colour charac-
teristics (Scott and Shackleton 1979). Birds and inver-
tebrates were identified by feathers, claws, shell frag-
ments, and cuticles and were scored as present or
absent. Using an 8 x 10 grid of 1-cm squares, we esti-
mated the percentage of prey species in each faeces.
All faeces were analyzed by one observer (HB) to
eliminate inter-observer bias (Spaulding et al. 2000).
Before faecal analysis, we tested observer accuracy
with a reference collection and scored 92% (23 out of
25 correct). Following identification of prey remains,
we also assessed precision and scored 100% (25/25)
on classification of taxa. Finally, we investigated our
precision in assessing adult/fawn categorization. We
classified 77% (17/22) correctly. This bias, however,
was directional: we scored five adults incorrectly as
fawns.
Data analysis
We used two indices in analysis. Percentage of whole
scat equivalents (pWSE) is the mean percentage of
each prey species found per faeces (Angerbjorn et al.
1999; Elmhagen et al. 2002). For example, two fae-
ces containing deer and seal in percentages of 40,60
and 60.40 respectively, would have pWSE of 50 for
both prey species. Percent occurrence per item (O/T) is
the number of occurrences of each prey species divid-
ed by the total number of items identified, where an
THE CANADIAN FIELD-NATURALIST
Vol. 120
item is defined as an occurrence of a prey species in
an individual faeces (Theberge et al. 1978).
We used log-linear analysis on pWSE data to test
for differences in diet between pups and adult wolves
while simultaneously assessing influences of years and
sites. We excluded birds and invertebrates from this
analysis because conversion to pWSE was not possi-
ble on presence/absence data. These items occurred
infrequently and were never found alone in faeces. In
this log-linear analysis, we also combined non-deer
items into an “other” category due to infrequent occur-
rence (see Results).
To measure dietary breadth of both age classes, we
used Levin’s Measure, standardized on a scale from
0 to 1 (Hurlbert 1978; Krebs 1999; Elmhagen et al.
2002). Levin’s measure of dietary breadth, B, is cal-
culated using pWSE data as follows:
[1] =| stato»
where p,; is the proportion of resource 7 in a diet
consisting of n resources.
We also calculated dietary overlap between age
classes using Schoener’s measure (p;,; Shoener 1970)
and Horn’s index (R,; Horn 1966). Both calculations
indicate food-niche overlap between populations j and
k, where p;; and p;, are the proportions of resource 1
in diets, and n is the number of available resources.
Although Shoener’s measure is frequently used and —
therefore comparable to other literature, Horn’s index
is less biased under changing numbers of resources,
sample size, and resource evenness (Krebs 1999).
Calculations were performed on pWSE data.
[2] P; =| ¥ (minimum p,,, p, )/100
[3]
5 _ ©, + Pi) log (p; +p.) - Sp, logp , -Y pi. logp ,
2 2log?
Results
Pup and adult wolves consumed a breadth of
resources; however, Sitka Black-tailed Deer consti-
tuted most of the diet for both age classes (Table 1).
Deer represented 54.6 to 100% of whole scat equiva-
lents, depending on site and year, and accounted for
50.0 to 98.4% of all prey items (Table 1). After deer,
Harbour Seal (Phoca vitulina), invertebrates, and birds
were the most common prey items, whereas Beaver
(Castor canadensis), Short-tailed Weasel (Mustela
ermina), Mink (Mustela vison), River Otter (Lutra can-
adensis), Marten (Martes americana), and Fisher
pups and adult wolves (Table 1). Pups and adults had
|
|
|
|
(Martes pennati) occurred infrequently in the diets of ©
|
63
ES
| WoL\
2T IN
aw De
JET: EARL
EN, AND PAQUET
IMCHEN,
T. REIM
ONT,
ARIM
N,D
BRYA
2006
srcent dict-
4 Average ae 719%.
Tz ble «). , ylves ther
(la It we hig
/ breadths and adu iderably his
, 1e a « a cons
similar foi pl ie ia tee ia 2). liet was
ary overld Ps index of 0.96 (Ta and adult die oe
a whereas eae shbitice stween pup aia of gape ed
cl iia f erage die A ference : c age clas rtion of ¢ h
Tia pa ajor diffe ’ different ropo “d wit
Dein 1) A maj ion of di a lower p compare 1).
; Ds a ortic ned a fawn ; Table
NE a nSm < he prop consun ion of 0.001; “ars
2 a oe ¥ (oe) fon n pe a << t ms Pups 2r proporti r=? Pc); across yea
a1 £ ae BAe. inet 5s ZZ sumed. a higher p 7.77, df = 2 varied ac ze class
2 3 Za pone St | and a h Ne eee ion, age >.
Epa it eilias we i | deer (Partial x f this differe en location, P=002:
Mh Koa Liaal adults itude o a etwee =i,F= ;
= Z S i “S = Hn | a chmcnens 36 oe “= Rete field Island
= S Ame I The m ions (ass¢ artial Chatfi ord-
25 Bal ess 6 nn locations item; Part of seal on > 1). Acec
a Loleas =] and lo and prey ite rtion of se (Table alysis,
gy eu Se ine iia < > of wolf, ue high propo en g-linear ana volf,
5 ages Zz. 1). A hig 3-way as: m log- ass of w
& A] é& ea] TAN venti | Table . d the 4 seal fro e class 0) 20.
E icles es ee Spee Bim das
3 9 vt Salas in 200 we exc locati Partial x areas
er are
- 5 no) *"o. i) ingly, mee smo sigican Pan to other aller
42 3 inal dee sSOC as not : d, si ificantly sm
S a — | he as ; Was foun , can d th
RS s oon Lpelait ; rey item We then d a signifi ‘olves at bo
Q» on | dp ). sume fe)
5 8 pape rec a P=0.65 consum id adult w 1)
5 = ~ © 5 ee i oa) re) ened df= 1, z that she than Fe 1, P<0.001).
a ‘Ss = ae) ears, t =1,
5g E ~ Qn+e hone eon 7a dk propia etn
7 @ OD Va) ’ 10) ia F diet
Eh pene aia a tee Gein ing wolf die
E oq a Re lie thie loca ists regarding atterns
49 = ieee Ta) Pe . ture exists temporal E 2003)
3 2 S atl iscussion of litera atial and Cuicei 2003
2 a i Sola Re << ye large body ddresses spa terson and variability.
©) lap iu er te A lars ily a 3; Peters lation >on-
a3 y Q i oI ZZ it primar n 2003; ee SUN IESE
° = | titp Carby f intrap ists 1 es
sie 5 ere + bu and ts o ion ex 0 studi
=o} % ieee (Paquet ther aspec informatio’ clude two F aint
SES Sale l not oth dietary ions in diet o
205 =: | + and little Exceptions i the berge
Balk he ively ng. Ey aon rere
= 2 a oy Relat ° ing young differen / areas ever.
Bees a raising y nt earby how
eo ©°o xt of enifica Ean a study, e.
gO iS B= |RES lac is t found sig den sites 6). Neither s s we do he
> e va} mat nN . ~
225 alo aed ien Za aha s between hko 198 adult diet as olves from
vO E as) oq) Yori wolve 78: Meles pup and d adult w the prey
5 s = < QMlho mS et al. 19) d between juvenile an iches at h and
285 eae bie differentiate how that ju retary breadth an
3 ow Il 1 1 in. we § have s : dieta ) wolves
2 iy 5'o Ton ere, we ds imilar adult ny
ane 5 2 ras) if . astal islan have ae) show that hemselves
! So a) No) oO CO S they ults s “1, to t 5. At
iD a) a N S Slaw en (erent tw ree level, a These res Ss similar for pups. ¢
2es Ie) ae Para eCcl erlap. items s Ses ; differ
= F SS = ~ a Teel Z. sp te ove ith food y spec diets d
é Set odera with food pu EEy SES at idesice
As g fe r~ SAS Saat een nting eis a show = as owe
N55 = t hu < veo fein cation an
vee ete See a
eae 3 = Lone ner res ; nd adu 3 ontrolling ved tha a
= v a} Va) x lo | a fi nN pups a isioning. Cc sults shov deer tha
oR ron ee is S It
28 é |x tela co aa eeu lective prov log-linear re ion of adu
Rese WS [i 7 MOISES: her ea T prop ced by
LS S| 20 et sue Ss year nee pres Id be E eeae ot
= = c S & S F
ee = B |nw 08 29 CI Senaene by adults. detected eee a bias in
No ao N = : i : : .
h = § a AlRSS aia SNES AER inition ae The
. — | jj 62° N. latitude) in the Taiga Shield ecozone. Records show Coyotes have been seen since the 1960s. Prior
to 2000, evidence of Coyotes breeding in the Taiga Shield has been anecdotal. In 2000, a Coyote was repeatedly seen at the
Yellowknife airport and in 2001, a pair of Coyotes was observed with two pups. Since then, Coyote pups have been observed
annually at the airport and adult Coyotes are seen regularly within the city of Yellowknife, an urban island within the Taiga Shield
ecozone. Unlike in most regions occupied by Coyotes, medium-sized prey are rarely seen. Recently, Coyotes have become a poten-
tial hazard to aircraft at the Yellowknife airport. Although Coyotes appear to have established themselves within the city of Yel-
lowknife, maintaining a presence beyond the urbanized area remains uncertain.
Key Words: Coyote, Canis latrans, distribution, range extension, reproduction, Northwest Territories.
Coyotes (Canis latrans) have been uncommon in
the Northwest Territories (NWT) north of Great Slave
Lake (north of the 62" parallel). Banfield (1974) and
Voigt and Berg (1987) suggested Coyote distribution
was limited to the Taiga Plain ecozone and did not
extend beyond the northwest shore of Great Slave Lake
where the Taiga Shield ecozone begins. The Taiga
Shield ecozone is characterized by a patchwork of
wetlands, forests, meadows, and shrublands situated
on top of glaciated bedrock with numerous lakes and
rock outcrops. Summers are short and winters are long
and cold. Although sightings of Coyotes beyond cur-
rent range limits may be expected, occurrences of re-
production beyond the Taiga Plain and into the Taiga
Mica would be less likely. Bekoff (1977) and Bekoff
and Gese (2003) estimated the range of Coyotes to be
further northward but their delineation appears based
B the northern limit of trees (“tree-line”). This paper
en on recorded sightings and breeding of Coyotes
‘in the Taiga Shield ecozone in northwestern Canada.
lidentify current conflicts and management responses
for Coyotes as their numbers have increased in a north-
em urban area.
IMethods
| Irecorded and investigated Coyote sightings in the
ellowknife area since summer 2001. I solicited sight-
‘ing and harvest information from wildlife officers, hunt-
‘ers, and trappers in the Northwest Territories and west-
m Nunavut. I searched historical records for Coyotes
ong fur harvest records, Wolf (C. /upus) bounty
‘teceipts, and annual reports. Carcasses of Coyotes ob-
ained since 2001 were examined whenever possible.
Results and Discussion
The earliest recorded sighting of Coyotes in the
Taiga Shield ecozone occurred in the Yellowknife area
in the early 1960s. In winter 1974-1975, five Coyotes
were observed in the Long Lake area near the Yellow-
knife airport (J. Hordal, Hay River, Northwest Territo-
ries, personal communication). Prior to 2000, three other
sightings occurred outside (>40 km) of the Yellowknife
area (Figure 1) and in the late 1960s, a trapper shot a
Coyote on Stark Lake, east from Lutsel K’e (formerly
Snowdrift) (A. Boucher, Lutsel K’e, Northwest Term-
tories, personal communication). One Coyote was oc-
casionally sighted at the Yellowknife airport in 2000
and in spring 2001, two adult Coyotes were often re-
ported near the airport; two pups were observed later
that summer, the first documented occurrence of Coy-
otes breeding this far north in the Northwest Territories.
There have been two oral reports of Coyotes at Kug-
luktuk (67°49'N, 115°06'W), Nunavut, near the north-
ern coast of mainland Canada. Although exact dates
and written reports are absent, the sightings are note-
worthy as they occurred on the tundra 40-60 kilome-
tres beyond the tree-line. The first was by a hunter who
shot a Coyote 70 km west of Kugluktuk around 1988.
The second was circa 1997 when two hunters saw a
Coyote southwest of Bloody Falls on the Coppermine
River (A. Niptanatiak, Kugluktuk, Nunavut, personal
communication).
The presence of Coyotes at the Yellowknife airport
is not surprising because the area is characterized by
open grassy habitat in contrast to the surrounding boreal
forest. Rodents, birds, hares, and other wildlife on the
airport property serve as food. The availability of food
67
68 THE CANADIAN FIELD-NATURALIST
scraps near the city of Yellowknife proper and at the
city dump should contribute to a continued presence by
Coyotes within the urbanized area. It is possible that the
Coyotes observed near the Yellowknife airport in win-
ter of 1974-1975 had bred in the area as three of the five
Coyotes did not appear adult size (J. Hordal, personal
communication).
Coyotes in northern urban areas
The existence and colonization of Coyotes in and
around Yellowknife is of concern to residents and air-
port authorities. Residents are primarily concerned for
the safety of their pets. Airport authorities are concerned
as Coyotes and their pups occupy the Yellowknife air-
port property and represent an air safety risk. In 2002,
2003 and 2004, 2,7, and 4 pups were observed on the
airstrip area respectively (S. Loutitt, Yellowknife air-
port, personal communication). Coyotes have not been
limited to the airport. Seven were observed together on
Back Bay in Yellowknife on 9 December 2004 (J.
Bastedo, Yellowknife, personal communication) repre-
senting the largest number of Coyotes observed together
within Yellowknife to date. In addition, a Coyote pup
was found dead on 30 August 2004 (J. Bastedo, per-
sonal communication) and another Coyote pup was
road-killed on 29 October 2004, both within Yellow-
knife city limits. Furthermore, on 21 October 2004, a
Coyote pup was killed on the highway about 60 kilo-
metres northwest of Yellowknife and may represent a
separate litter (Figure 1).
The origin of this newest incursion of Coyotes is
unknown, although they may have followed the single
highway corridor connecting Yellowknife to the south.
Coyotes have been expanding their range elsewhere
over recent decades and human-induced habitat ch-
anges are likely contributory (Moore and Parker 1992;
Chubbs and Phillips 2002; Chubbs and Phillips 2005).
Factors that might discourage Coyotes from colonizing
northward include morphology, behaviour, and repro-
duction (Phillips 1982). Although not morphologically
equipped for travel in areas with deep snow, Coyotes
might overcome this limitation by using trails and areas
with hard snow (Murray and Boutin 1991; Murray and
Lariviere 2002). Wolves often kill coyotes where their
ranges overlap (Krefting 1969; Paquet 1989; Crabtree
1998), further discouraging northern Coyote range
expansion.
Air-strike risk
Coyotes at the Yellowknife airport present a public
safety risk because of aborted take-off and landing
attempts and potential for collisions causing property
damage and personal injury (B. Webber, Yellowknife
airport, personal communication). Although the airport
perimeter is fenced, Coyotes can access the area
through holes in the chain-link or by digging under-
neath. Continual removal efforts would be required to
be effective over the long-term.
Vol. 120
Although Coyotes have the potential to cause signif-
icant damage to small planes, those struck at southern
airports are often hit by undercarriage and do little
overall damage to the aircraft (Dolbeer et al. 2000;
Cleary et al. 2004). The greater risk is from aborted
take-off and landings (Dolbeer et al. 2000), as pilots
tend to see Coyotes on runways before collisions occur.
There is anecdotal evidence that Coyotes may dis-
perse other wildlife at airports such as Sandhill Cranes
(Grus canadensis), Red Fox (Vulpes vulpes), and deer
(Odocoileus sp.), and thus reduce the overall risk of
wildlife strikes by aircraft (R. Dolbeer, United States
Department of Agriculture, personal communication; S.
Loutitt, personal communication). Coyotes are known
to predate on Red Fox in some southern regions (Voigt
and Earle 1983; Sargeant et al. 1987; Harrison et al.
1989), although fox predation by Coyotes has not yet
been reported at the Yellowknife airport. There are few
site-specific wildlife studies that address relative abun-
dance and spatial patterns of wildlife at airports (Hoff-
man et al. 1996; Dolbeer et al. 2000). The dilemma at
the Yellowknife airport is one of risk management. Air-
port management staff believe co-existence is possible
and have chosen not to eliminate Coyotes but to moni-
tor their presence and reduce numbers when required.
Coyote persistence
Whether Coyotes remain in the Yellowknife region
is uncertain, although they occupy similar habitat else-
where (e.g., Alaska; Labrador). Coyotes expanded their
range northwest to Alaska in the late 1800s by scav-
enging along trails left by the gold rush (Gier 1975)
and arrived in the Yukon by 1910 (Parker 1995). Coy-
otes appeared in the Northwest Territories fur harvest
in the early 1930s when records were first kept. Sup-
pressants to early colonization of Coyotes in the Yel-
lowknife region include hunting, trapping, predation by
Wolves, and incidental mortality at poison baits during
Wolf control in the early 1960s (Heard 1983; Cluff and
Murray 1995). Wolves are occasionally observed at
the Yellowknife airport, and in 2004 a pack of five were
reported within city limits.
To sustain a Coyote population a reasonable prey
base must exist. Birds, Snowshoe Hare (Lepus ameri-
canus) and other small mammals would be the main
source of food for forest-dwelling Coyotes (Créte et al.
2001). White-tailed Deer (O. virginianus) are only
rarely sighted in the Taiga Shield (unpublished data;
Veitch 2001). Wood Bison (Bison bison athabascae)
have expanded their range into the area since 1999 but
this may adversely impact Coyotes if Wolf numbers
also increase. Garbage and domesticated animals in the
Yellowknife area would augment the available food
supply. The Yellowknife city dump, electrified only in
summer against Black Bears (Ursus americanus), is not
a barrier to Coyotes. Cabins and homes scattered on
the landscape outside of Yellowknife may also supply
2006
CLUFF: COYOTE BREEDING RANGE IN NORTHWEST TERRITORIES
69
Area of
Detail
\ asp
Ficure |. Taiga Plain and Taiga Shield ecozones bordering Great Slave Lake in the Northwest Territories showing records of
Coyotes since the 1960s documented in text. Circles represent sites prior to 2000, squares in 2003 and 2004.
food. Consequently, Coyote persistence in Taiga Shield
of the Northwest Territories may be strongly linked to
a suburban environment, especially if an inadequate
prey base exists elsewhere. How the public and wildlife
management authorities choose to deal with Coyotes as
urban carnivores will be a key factor for Coyote persist-
ence in the Yellowknife region.
Acknowledgments
I thank the trappers, wildlife officers, and biologists
in the Northwest Territories and Nunavut for their assis-
tance in documenting and mapping sightings of Coy-
Otes in the two jurisdictions. I also appreciate the efforts
of airport management staff to document their sight-
ings of Coyotes and to facilitate logistics of my investi-
gations. E. Campbell was especially helpful in referring
me to trappers and providing earlier personal sightings
of Coyotes. Comments from A. Gunn, G. Elliott, M.
Créte, and two anonymous referees helped improve an
earlier version of the manuscript.
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Received 7 March 2005
Accepted 2 November 2005
Lake Sturgeon, Acipenser fulvescens, Movements in Rainy Lake,
Minnesota and Ontario
W. EUGENE ADAMS, Jr.!*, LARRY W. KALLEMEYN?, and Davip W. WILLIs!4
'Department of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, South Dakota 57007 USA
*B-mail: Geno.Adams@staff.sd.us
3U.S. Geological Survey, 3131 Hwy 53, International Falls, Minnesota, 56649, USA. E-mail: larry_kallemeyn@usgs.gov
4E-mail: David.Willis@sdstate.edu, corresponding author.
Adams, W. Eugene, Jr., Larry W. Kallemeyn, and David W. Willis. 2006. Lake Sturgeon, Acipenser fulvescens, movements
in Rainy Lake, Minnesota and Ontario. Canadian Field-Naturalist 120(1): 71-82.
Rainy Lake, Minnesota-Ontario, contains a native population of Lake Sturgeon (Acipenser fulvescens) that has gone largely
unstudied. The objective of this descriptive study was to summarize generalized Lake Sturgeon movement patterns through
the use of biotelemetry. Telemetry data reinforced the high utilization of the Squirrel Falls geographic location by Lake Stur-
geon, with 37% of the re-locations occurring in that area. Other spring aggregations occurred in areas associated with Kettle
Falls, the Pipestone River, and the Rat River, which could indicate spawning activity. Movement of Lake Sturgeon between
the Seine River and the South Arm of Rainy Lake indicates the likelihood of one integrated population on the east end of the
South Arm. The lack of re-locations in the Seine River during the months of September and October may have been due to
Lake Sturgeon moving into deeper water areas of the Seine River and out of the range of radio telemetry gear or simply moy-
ing back into the South Arm. Due to the movements between Minnesota and Ontario, coordination of management efforts
among provincial, state, and federal agencies will be important.
Key Words: Lake Sturgeon, Acipenser fulvescens, biotelemetry, Rainy Lake, Ontario, Minnesota.
Movement patterns of Lake Sturgeon (Acipenser ful-
vescens) were relatively unstudied through the early
1970s (Scott and Crossman 1973). However, in the past
decade, Lake Sturgeon movements have been studied
at an increasing rate throughout much of the species’
range (Rusak and Mosindy 1997; McKinley et al. 1998;
Auer 1999; Borkholder et al. 2002; Knights et al.
2002). Lake Sturgeon have been found to move large
distances for purposes such as spawning (Threader and
Brosseau 1986; Fortin et al. 1993; Rusak and Mosindy
1997; Auer 1999) while maintaining smaller ranges in
other systems (Bassett 1982*). Such movement pat-
‘terns make Lake Sturgeon management difficult, par-
ticularly for multiple agencies dealing with border
waters of substantial size.
Rainy Lake, a border water shared by Minnesota and
Ontario, contains a Lake Sturgeon population that has
gone largely unstudied. Lake Sturgeon in this system
have been segregated from upstream Namakan Lake
and downstream Lake of the Woods/Rainy River sys-
i populations by dams constructed on the outlets
/of Rainy and Namakan lakes early in the 20" century.
|The Lake Sturgeon is a Minnesota state-listed species
| of special concern (Minnesota Department of Natural
i Resources 2003*), while The Committee on the Sta-
tus of Endangered Wildlife in Canada considers west-
em populations of Lake Sturgeon to be endangered
is the Rainy River-Lake of the Woods populations
to be of special concern (Environment Canada 2005*).
| Management of the Lake Sturgeon population in
Rainy Lake is a cooperative effort among the Ontario
Ministry of Natural Resources, the Minnesota Depart-
ment of Natural Resources, and the United States fed-
eral government. Movement of Lake Sturgeon across
international borders leads to an increase in impor-
tance of appropriate harvest regulations and an under-
standing of movement patterns and spawning locations.
Such information will allow managers to protect areas
that are important for Lake Sturgeon spawning.
The objective of this descriptive study was to deter-
mine generalized Lake Sturgeon movement patterns
in Rainy Lake through the use of biotelemetry. Move-
ment data obtained from tagged Lake Sturgeon were
organized using a geographic information system (GIS)
and used to help identify seasonal movement patterns
and potential spawning locations.
Study Site
Rainy Lake, located on the Minnesota-Ontario bor-
der, consists of three main basins: the North Arm,
Redgut Bay, and the South Arm, all of which are part
of the Winnipeg-Nelson drainage system in the Lake
Winnipeg primary watershed (Figure 1). The North
Arm and Redgut Bay are both located entirely in Cana-
da. Rainy Lake has a total surface area of 92 000 ha, a
maximum depth of 49.1 m, and a mean depth of 9.9 m.
Of the three main lake basins, the South Arm has the
greatest surface area at 49 200 ha, with 27 300 ha in
Ontario and 21 900 ha in Minnesota. The South Arm
is also the deepest basin with a mean depth of 9.9 m
and a maximum depth of 49.1 m. The South Arm ex-
tends for 56 km along the border of the United States
TA
2 THE CANADIAN FIELD-NATURALIST
and Canada. The South Arm of Rainy Lake, along
with a 40-km section of the Seine River, was the area
of focus for this study.
The watershed associated with Rainy Lake encom-
passes over 37 500 km? and can be divided into two
sub-basins, including a 19 270-km? area above the
outlet of Namakan Reservoir at Kettle Falls and Squir-
rel Falls and a 19 320-km? area below Kettle Falls
and Squirrel Falls that drains directly to Rainy Lake.
Overflows between Namakan Reservoir and Rainy
Lake also occur at Bear Portage and Gold Portage.
Below Kettle Falls and Squirrel Falls there are two
principal tributaries that enter Rainy Lake, the Turtle
River with a mean discharge of 37 m*/sec and the Seine
River with a mean discharge of 48 m?/sec. Smaller,
ungauged tributaries into the South Arm of Rainy Lake
include the Rat and Pipestone rivers, both of which
may be used by Lake Sturgeon for spawning. Changes
in flow initiated at the headwaters of the watershed
take about 21 days to reach the outlet of Rainy Lake
under normal flow conditions with the water drop-
ping about 135 m in the 338 km between these two
points (International Rainy Lake Board of Control/
International Lake of the Woods Control Board 1984*).
Long-term flow records indicate that approximately
8.3 billion m? of water move through the Rainy Lake
watershed annually (Ericson et al. 1976*).
Methods
Biotelemetry was utilized in the identification and
characterization of Lake Sturgeon seasonal movement
patterns and likely spawning locations in Rainy Lake.
Lake Sturgeon were captured for transmitter implan-
tation in October of 2002 (n = 13), May and June of
2003 (n = 20), and October of 2003 (n = 8) at the mouths
of tributaries where they were known to concentrate
(tagging locations are noted in Table 1). Large mesh
multifilament gill nets with mesh sizes of 103, 114, 127,
152, and 178 mm (bar measure; i.e., distance from
knot to adjacent knot) were used to capture fish. Net
lengths varied from 30 to 91 m, with all nets having a
height of 1.83 m. Most nets were fished for approxi-
mately 24 hours, but actual times varied among sam-
pling crews. Captured Lake Sturgeon were measured,
weighed, sex was noted if gonads were identifiable
during surgery, and a pectoral fin ray section was re-
moved for aging (Adams et al. 2006).
Lotek Wireless combined acoustic/radio transmit-
ters (CART) were surgically implanted into 41 Lake
Sturgeon (>8 kg) throughout the duration of this study.
Thus, only a single tag was implanted and used for
both radio and acoustic tracking, with the radio portion
of the tag allowing tracking of fish in shallow or tur-
bulent habitats, while the ultrasonic portion of the tag
was needed for tracking of Lake Sturgeon in deeper
habitats (Winter 1996). Surgical implantation proce-
dures followed guidelines set by Hart and Summerfelt
(1975). A 5- to 8-cm incision was made on the ventral
Vol. 120
fish surface approximately 3 cm off the midline and
approximately 3 cm from the anterior end of the pelvic
girdle. A 0.5-cm exit hole for the whip antenna was
started with a scalpel on the midline approximately
3 cm posterior to the incision. A curved, 12-gauge
catheter needle was passed through the hole and out
through the incision with care taken to not penetrate or
cut the viscera. The whip antenna projecting from the
CART tag was then threaded though the end of the
catheter needle that was protruding through the incision
with the antenna being passed out of the peritoneal
cavity through the antenna hole. The CART tag was
then inserted into the peritoneal cavity with minimal
pressure exerted on the internal organs. Fish were
placed in a holding pen following implantation, if nec-
essary. When full equilibrium was regained, fish were
placed in the water at the site of capture and monitored
until they vacated the area. All fish handling and sur-
gery followed guidelines for use of wild fishes estab-
lished by Nickum et al. (2004).
Lotek CART 16-2 series tags were implanted in 33
Lake Sturgeon with the remaining eight fish receiving
CART 16-1 series tags. Lotek CART 16-2 tags were
16 mm X 85 mm with a weight in air of 36 g, while
Lotek CART 16-1 tags were 16 mm Xx 60 mm with a
weight in air of 25.3 g. Implanted CART tags did not
exceed 2% of the total body weight of any given fish
(Gallep and Magnuson 1972; McCleave and Stred
1975; Stasko and Pincock 1977; Moser et al. 1990).
Aerial tracking of Lake Sturgeon occurred at least
once per week throughout the spring spawning season
and summer months when the airplane was opera-
tional. Transects were flown covering the eastern half
of the South Arm, with every flight including a 40-km
stretch of the Seine River that flows into Rainy Lake.
Transects were chosen to maximize area covered with
the least amount of flight time. The remaining portion
of the South Arm was flown at least once every two
flights, time and weather permitting. The North Arm
of Rainy Lake was flown once in the summer of 2003
in an attempt to locate a fish that could not be located
in the South Arm.
Radio tracking by boat took place on average two
to three times per week depending upon weather con-
ditions. Tracking was also dependent on time availabil-
ity, with partial and full days devoted to boat tracking
when possible. Locations highly used by Lake Stur-
geon were sampled by boat during May and June be-
cause recognition of individual signals was difficult
from the airplane.
Distributional patterns of Lake Sturgeon were sum-
marized by month during the spawning period (May
and June) and by two-month periods throughout the
rest of the summer (July/August and September/Oc-
tober). Movement rates were calculated as the linear
distance between successive fixes divided by the num- |
ber of elapsed days. Lake Sturgeon home ranges were |
calculated using the Animal Movement extension in |
———————
=
2 ey a
2006
ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS
73
Kilometers
:
Seine River
IRS
Pipestone River
——=
5a eo te
| Kettle Falls ;
Squirrel Falls’
Sok » 7 Canadian Channel
Ca a’
x=
—
FIGURE 1. Rainy Lake, Minnesota and Ontario. The line across the South Arm indicates the border between Canada and the
United States.
_ ArcView 3.2 (Hooge et al. 1999*), with home ranges
estimated by using minimum convex polygons (White
and Garrett 1990). Home ranges included only water
area; land masses were excluded.
Lake Sturgeon eggs were sampled during 2004 in
an attempt to confirm spawning locations. Egg col-
lectors were set weekly from 7 June through 1 July
in three areas of high Lake Sturgeon use. Collectors
consisted of an 18-m mainline of vinyl-coated wire
cable (4.8 x 6.4 mm) with cement blocks (203 x 406
x 102 mm) attached at 3-m intervals and wrapped in
furnace filter material. Floats were attached to each end
of the mainline and egg collectors were set in shallow
water areas of 0.3 to 4.6-m depth. Egg collectors were
checked once weekly during the entire time period in
which they were deployed. A sample of all egg types
present on the collectors was removed and transport-
ed back to laboratory facilities to be hatched and the
fry subsequently identified.
Results and Discussion
Lake Sturgeon tagging and survival
Forty-one Lake Sturgeon were implanted with CART
tags during 2002 and 2003 (Table 1). Individual Lake
Sturgeon were re-located between 0 and 36 times. No
mortalities were positively identified. However, fish 99,
implanted in October of 2003, was not located during
the remainder of the study. Fish 30 was not located for
12 months after it was tagged and released, but then
was subsequently located 11 times throughout the re-
mainder of the study. Fish 106 was found at the same
location, adjacent to the Seine River First Nations vil-
lage on the Seine River, weekly for 11 weeks. We later
learned that fish 106 may have perished in a gill net
used in the Seine River First Nations subsistence fish-
ery that had been left on the bottom of the Seine River
due to entanglement with the substrate. All other fish
were located at least once throughout the study.
Lake Sturgeon implanted with CART tags had a
mean fork length of 1217.4 mm (range 951-1505 mm),
girth of 502.4mm (range 372-639 mm), mass of
15.35 kg (range 7.5-30 kg), and age of 25.9 (range
15-55) (Table 1). These ranges were representative of
the Lake Sturgeon population in Rainy Lake at the
time of this study (Adams et al. 2006).
Transmitter detection
Acoustic equipment was tested during the spring
of 2003. Tags were lowered at 2-m intervals between
74
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Fish description, transmitter code, and capture location at time of tagging for Lake Sturgeon implanted with Lotek
CART transmitters in Rainy Lake, 2002-2003. Ages were determined from pectoral fin ray sections. NA = not available.
Tagging Total Fork
Code date length length Girth
number Month/day/year (mm) (mm) (mm)
1 10/2/02 NA 1100 444
yD 5/21/03 1506 1433 635
3 10/9/02 NA 1210 556
4 10/9/02 NA 1275 615
5 10/9/02 NA 994 372
6 10/9/02 NA 1050 425
i] 10/9/02 NA 951 395
8 10/8/02 NA 1212 485
9 10/8/02 NA 1313 582
10 10/8/02 NA 1386 614
11 10/8/02 NA 1447 639
12 10/8/02 NA 1388 585
13 10/8/02 NA 1200 445
14 10/8/02 NA 1125 407
15 5/28/03 1372 1260 495
16 5/20/03 1400 1270 503
17 5/28/03 1404 1300 520
18 5/28/03 1410 1285 475
19 6/18/03 1270 1140 410
20 5/28/03 1200 1060 415
21 6/18/03 1425 1280 570
22 5/28/03 1505 1365 545
23 6/18/03 1120 1000 460
24 5/21/03 1460 1315 578
25 5/20/03 1379 1260 525
ay 5/23/03 1640 1505 585
30 5/29/03 1444 1291 595
39 5/23/03 1555 1429 580
43 5/29/03 1335 1205 520
Sil 5/29/03 1180 1063 405
56 5/23/03 1575 1443 589
63 5/23/03 1135 1025 435
69 5/28/03 1280 1140 410
Ve 10/8/03 1449 1325 568
82 10/7/03 1400 1275 486
87 10/8/03 1095 997 431
93 10/8/03 1265 1134 442
99 10/8/03 1217 1094 450
106 10/8/03 1288 1186 461
111 10/8/03 1272 1138 467
119 10/8/03 1160 1045 480
Weight Tagging
(kg) Sex” Age location
11.5 FI 22 Canadian Channel
30 F 5p) Brule Narrows
19 FM 26 Stokes Bay
Des) FM Di Canadian Channel
8 MM 15 Stokes Bay
95 Unk 19 Stokes Bay
eS) MM 17 Squirrel Falls
16 Unk 28 Squirrel Falls
20.5 FM 32 Canadian Channel
DiS es) FM 28 Canadian Channel
30 FM 31 Stokes Bay
DORMS FM 26 Kettle Falls
IS). Unk 18 Squirrel Falls
11.5 Unk 19 Squirrel Falls
14.5 Unk 21 Kettle Falls
16.5 Unk NA Brule Narrows
16.75 MM 21 Kettle Falls
IS) MM 35 Kettle Falls
9.5 M 32 Kettle Falls
9 MM 20 Kettle Falls
20 FM 30 Squirrel Falls
1935 FM 27 Kettle Falls
11 Unk 21 Kettle Falls
20.75 FM 22 Brule Narrows
18 Unk 26 Brule Narrows
19.25 FM 47 Seine River
VES le 29 Squirrel Falls
22 FM 47 Stokes Bay
16.5 FI 22 Stokes Bay
8.5 Unk 19 Seine River
16 Unk 44 Seine River
8 Unk 18 Seine River
10 Ip 23 Rat River Bay Mouth
21 F NA Seine River
5) Unk Bi Brule Narrows
8 Unk 18 Seine River
10.5 M 16 Seine River
9.5 Unk 16 Brule Narrows
12 M 18 Seine River
eS M Ds) Seine River
10.5 Unk 19 Seine River
“F = female, maturity unknown; FM = female, mature; FI = female, immature; M = male, maturity unknown; MM = male,
mature; Unk = sex unknown
2 and 20 m. The highest maximum detectable dis-
tance from the boat was 0.95 km when the CART tag
was lowered to 6 m. The lowest maximum detectable
distance of 0.53 km occurred at a tag depth of 18 m.
The mean maximum detectable distance was 0.65 km
with a standard error of 0.04. Limitations in the per-
formance of the acoustic portion of the tags led to the
discontinuation of acoustic telemetry from the study
beginning during July of 2003. No fish locations were
recorded with the acoustic equipment despite attempts
on 10 dates between 15 May and 15 June 2003. The
complexity of the bottom contours of Rainy Lake may
have been the reason for the lack of reception of the
directional acoustic signal.
Radio telemetry equipment was also tested. The
highest maximum detectable distance. from the boat
was 0.47 km when the CART tag was lowered to 2 m.
The lowest maximum detectable distance of 0.017 km
occurred at a tag depth of 20 m. The mean maximum
detectable distance was 0.21 km with a standard error
of 0.05. The maximum detectable distance by airplane
with the tag lowered to 8.77 m was 0.55 km with a
standard error of 38.5.
2006
Canadian Channel
-92.964° -92.823° -92.701°
ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 75
_ 4 Kilometers
-92.580° -92.458° -92.336°W
FIGURE 2. Distribution of Lake Sturgeon locations in Rainy Lake during May and June, 2003 and 2004.
Lake Sturgeon locations
The 41 implanted Lake Sturgeon were located 587
times throughout the study, with fish being tracked
through September 2004. In 2003 Lake Sturgeon were
located 285 times and in 2004 Lake Sturgeon were
located 302 times.
Lake Sturgeon utilized Squirrel Falls/Kettle Falls
and adjacent areas (Canadian Channel, Hale Bay) ex-
tensively throughout the study (37% of total locations,
including both spawning and non-spawning seasons).
A rock shelf below the Squirrel Falls Dam is one of the
areas thought to be a spawning site for Lake Sturgeon
in Rainy Lake. Egg samplers were placed below the
Squirrel Falls Dam, in the Canadian Channel, and
below the falls in the Pipestone River in 2004. Lake
Sturgeon spawning, verified by hatching eggs and iden-
tifying larvae, was confirmed only at the Squirrel Falls
Dam and occurred between 14 June and 17 June of
2004. Thus, we cannot exclude the possibility of spawn-
ing at other locations, but were able to verify spawning
below the Squirrel Falls Dam. Other sites with Lake
Sturgeon spawning habitat characteristics as listed by
Scott and Crossman (1973) (e.g., 0.6-4.6 m depth, swift
current, below falls) include the Kettle Falls Dam,
Pipestone River, Rat River, and the Seine River (Crilly
Dam and Highway 11 Bridge). Lake Sturgeon were
located at all of these sites throughout the study, in-
cluding the spawning season.
The majority of May locations (89%) for both 2003
(85%) and 2004 (90%) were associated with the cur-
rent area directly below Kettle Falls, Squirrel Falls, and
in the Canadian Channel (Figure 2), which suggests
pre-spawning aggregations. Four of the other six May
locations were associated with the Brule Narrows, also
an area with consistent current.
Fewer Lake Sturgeon were located in the Kettle
Falls/Squirrel Falls/Canadian Channel area as June
progressed in both years (Adams 2004*). We suspect
that Lake Sturgeon gradually left the spawning areas
at Kettle Falls and Squirrel Falls, and dispersed pri-
marily to the main basin east of Brule Narrows. Dur-
ing this month, Lake Sturgeon locations in the Seine
River were spread throughout Seine Bay and 36 km
upriver. The increased number of Seine River locations
from 2003 to 2004 was due to the increased number of
tagged Lake Sturgeon. Only four fish from the Seine
River were tagged for 2003 tracking, while six addi-
tional fish were tagged for 2004 tracking.
76 THE CANADIAN FIELD-NATURALIST
Vol. 120
2 4
6 Kilometers
-93.270° -93.067° -92.963°
-92.701° -92.580° -92.458°W
FIGURE 3. Distribution of Lake Sturgeon in Rainy Lake during July and August, 2003 and 2004.
During the July/August period, the majority of Lake
Sturgeon were found from the Brule Narrows east-
ward and in the Seine Bay/Seine River system, with
only five locations found west of the Brule Narrows
(Figure 3). Some Lake Sturgeon continued to be locat-
ed in the Kettle Falls/Squirrel Falls area during this
period in both years (15 locations and 35 locations,
respectively). Locations in the main body of the South
Arm were associated with the north shore more than
the south shore during this time period (96% in 2003
and 84% in 2004). There are substantial habitat dif-
ferences between the two areas with the south shore
consisting of a sharp drop-off into depths of over 30 m.
Conversely, the north shore consists of rock reefs and
depths shallower than 15 m. Either the fish were less
likely to use the deeper water, or the lack of locations
was due to limitations of the telemetry equipment with
radio signals detected from depths no greater than 18 m.
September locations were obtained during both
2003 and 2004 with October locations obtained only
in 2003. Locations during this time period were dis-
tributed throughout the South Arm (Figure 4). Loca-
tions in the Seine River were few, with zero in 2003
and two in 2004. Fish 75 was tagged in the Seine River
but located in the South Arm during this time period.
Lake Sturgeon were located on the south shore of the
South Arm for the first time in September. One poten-
tial explanation is that Lake Sturgeon were staging at
locations in the vicinity of winter habitats. Rusak and
Mosindy (1997) found that Lake Sturgeon in the Lake
of the Woods/Rainy River system demonstrated con-
sistent preferences for specific areas in the main basin
of Lake of the Woods during the winter; these areas
were adjacent to the mouth of the Rainy River where
the Lake Sturgeon would later spawn in the spring.
Lake Sturgeon movement rates
Mean movement rates of Lake Sturgeon were cal-
culated by month for descriptive purposes (Figure 5).
Movement rates increased from May to June during
2003 and 2004, with a maximum movement rate of |
0.80 km/day in June of 2004 and a minimum move- |
ment rate of 0.17 km/day in May of 2003. Movement |
rates in five of the 10 months exceeded the mean spring
2006
4 Kilometers
|
-92.963° -92.823°
-92.701°
ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 77
-92.580° -92.458°W
Ficure 4. Distribution of Lake Sturgeon locations in Rainy Lake during September and October 2003 and September 2004.
movement rate of 0.50 km/day observed by Knights
et al. (2002) for Lake Sturgeon in the upper Missis-
sippi River system. Lake Sturgeon in Rainy Lake like-
ly were dispersing into the lake away from spawning
sites during the post-spawn period in June. However,
movement rate patterns were dissimilar for the same
months between years. Further research thus will be
needed to ascertain seasonal movement patterns and
explanations for those patterns.
| Movement of fish between the Seine River and the
main body of the South Arm did occur but was mini-
mal throughout the study. Rusak and Mosindy (1997)
found that in the Lake of the Woods/Rainy River sys-
_ tem, there was a separation in the population based
| primarily on winter habitat use. “River” fish spawned
in the Rainy River and remained there throughout the
_ winter months. “Lake” fish spawned in the Rainy River
| but moved into the main basin of Lake of the Woods
| during the winter months. If a similar dichotomy exists
/ in the Rainy Lake/Seine River population, there may
| be greater movement of Lake Sturgeon between these
a
at
two areas during the time period in which we did not
attempt to locate fish (i.e., late fall and winter). Thus,
Lake Sturgeon may move from Rainy Lake into the
Seine River during this time period, but further inves-
tigation will be needed.
Movement patterns varied for individual Lake Stur-
geon in Rainy Lake. Some fish did not exhibit dis-
cernable movement patterns, such as fish 21, which
was located throughout the east end of the South Arm
from Stokes Bay to the Brule Narrows and also in
Seine Bay and at the mouth of Seine Bay (Figure 6).
Locations in 2003 began in the east and moved west
with the last location east of the Brule Narrows. The
first locations in 2004 were in the Seine Bay/Seine
River area with the remainder of locations further east.
One factor contributing to the differential patterns
between years may be the spawning interval exhibited
by Lake Sturgeon, which ranges between | and 3 years
for males and 4 and 6 years for females (Magnin 1966).
Telemetry results from Rainy Lake suggest that
some Lake Sturgeon remained in the same area for
78
THE CANADIAN FIELD-NATURALIST Vol. 120
May June July August September
Month
FIGURE 5. Mean monthly movement rate (+ standard error) of Lake Sturgeon in Rainy Lake during 2003 and 2004.
2006
ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 79
4 Kilometers
Tagging Location |
6/18/03
FiGureE 6. Distribution of locations for Lake Sturgeon number 21 in Rainy Lake, 2003 and 2004.
extended periods. For example, fish 15 was tagged at
the Squirrel Falls Dam on 28 May 2003 and was next
located in the Brule Narrows area (Figure 7). Subse-
quent locations were all in the area of the Brule Nar-
rows until 26 August 2003. The next locations were
in the Squirrel Falls Dam area beginning on 28 Sep-
tember 2003. The fish was then located in the Squirrel
Falls Dam area on 18 May 2004 and was not located
in the Brule Narrows area until 30 June 2004.
Lake Sturgeon home ranges
_ Home ranges for Lake Sturgeon in Rainy Lake var-
ied substantially, with a maximum home range size of
} 14 844 ha, a minimum size (excluding fish 106, pos-
‘sible mortality) of 84 ha, and a mean home range size
\of 4 625 ha (standard error = 642). Lyons and Kemp-
inger (1992*) reported that most Lake Sturgeon in the
Lake Winnebago system had consistent movement pat-
terns, while others demonstrated variable movement
patterns and apparently did not establish home ranges.
They also found that Lake Sturgeon in Lake Win-
nebago did not remain in any particular location for
long periods of time.
Conclusions
This research should provide biologists from mul-
tiple agencies insight into general movement patterns
of Lake Sturgeon inhabiting Rainy Lake. Telemetry
data indicated high utilization of the Squirrel Falls area,
with 37% of total locations occurring at that area.
Squirrel Falls was the only site at which spawning
was confirmed by collection of Lake Sturgeon eggs,
although other aggregations in areas associated with
Kettle Falls, the Pipestone River, and the Rat River
also could indicate spawning activity. Movement of
Lake Sturgeon between the Seine River and the South
Arm of Rainy Lake indicates the likelihood of one
integrated population on the east end of the South Arm.
The lack of locations in the Seine River during the
months of September and October might result from
Lake Sturgeon moving into deeper water areas of the
Seine River and out of the range of telemetry gear or
simply moving back into the South Arm. Further re-
search will be needed to determine the winter range
of Lake Sturgeon in Rainy Lake. The Lake Sturgeon
has historically been a resource of both cultural and
economic importance to the Rainy Lake area. Due to
80 THE CANADIAN FIELD-NATURALIST Vol. 120
TABLE 2. Home ranges and mean movement rates (SD) by year for Lake Sturgeon in Rainy Lake. Missing values (dots) indicat
that a fish was located zero or one times that year and thus movement could not be calculated.
Fish number Total Home range Mean movement rate Mean movement rate
locations (ha) 2003 (km/d) 2004 (km/d)
1 Ds 2164 0.320 (0.22) 0.265 (0.31)
2 12 7758 0.170 (0.14) 0.564 (0.26)
3 7) 2985 0.267 (0.20) 0.372 (0.33)
4 7 14844 0.619 (0.50) 2
5 13 3779 0.285 (0.33) 0.510 (0.36)
6 15 5416 0.448 (0.70) 0.663 (0.72)
7 23 3558 0.289 (0.28) 1.065 (0.74)
8 20 VAY 0.076 (0.05) 0.554 (1.11)
9 11 8600 0.156 (0.20) i
10 29 11894 0.808 (0.67) 0.977 (1.29)
11 17 10501 0.309 (0.23) 0.382 (0.23)
12 12 578 0.233 (0.25) 0.261 (0.43)
13 31 4404 0.420 (0.54) 0.120 (0.23)
14 12 3806 1.121 (1.00) 0.195 (0.21)
15 34 7141 0.352 (0.42) 0.714 (0.92)
16 19 2434 0.324 (0.25) :
17 29 10245 0.565 (0.53) 0.552 (0.41)
18 28 Sly) 0.279 (0.28) 0.513 (0.46)
19 14 6434 0.396 (0.39) 1.091 (1.74)
20 34 1859 0.299 (0.51) 0.870 (1.30)
21 15 10017 0.316 (0.11) 0.817 (0.61)
22 26 8492 0.459 (0.36) 0.511 (0.38)
23 19 4095 0.684 (0.80) 0.187 (0.20)
24 19 9411 0.587 (0.94) 0.236 (0.31)
25 7 7677 : 1.713 (1.56)
27 10 1921 0.324 (0.19) 0.200 (0.20)
30 10 1196 : 0.527 (0.66)
>I 18 1049 0.131 (0.06) 0.262 (0.38)
56 6 1499 : 0.651 (0.81)
69 10 2448 0.267 (0.36) 0.960 (1.26)
TS 14 159 1.160 (2.77)
82 6 641 0.213 (0.26)
87 5 433 0.397 (0.15)
93 8 2355 0.995 (0.97)
106 14 71 0.099 (0.04)
ita 3 84 1.114 (0.35)
119 6 380 0.583 (0.90)
the movements between Minnesota and Ontario, coor-
dination of future management efforts among provin-
cial, state, and federal agencies will be important.
Acknowledgments
Technical review was provided by Trent Sutton,
Daniel Hubbard, and three anonymous reviewers. Par-
tial funding for this project was provided by the Na-
tional Park Service through the Great Plains Cooper-
ative Ecosystem Studies Unit Cooperative Agreement
Modification Number J6820031002 (administered
through the University of Nebraska), the U.S. Geolog-
ical Survey, the Minnesota Department of Natural Re-
sources, and the Ontario Ministry of Natural Resources.
Field assistance was provided by Voyageurs National
Park employees. This manuscript was approved for
publications by the South Dakota Agricultural Exper-
iment Station as Journal Series Number 3472.
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)06
4 Kilometers
ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 8 |
| Tagging Location|. 4
bs SORNB ya ae
2003 ry
@ 2004
Ficure 7. Distribution of locations for Lake Sturgeon number 15 in Rainy Lake, 2003 and 2004.
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uer, N. A. 1999. Population characteristics and movements
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Received 8 March 2005
Accepted 11 October 2005
lorthern Range Expansion and Invasion by the Common Carp,
'yprinus carpio, of the Churchill River System in Manitoba
.SCAL H. J. BApiou! and L. GORDON GOLDSBOROUGH!
lepartment of Botany, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada; e-mail: pbadiou@shaw.ca;
Present address: 48 DesMeurons Street, Winnipeg, Manitoba, RZH 2M1 Canada
lelta Marsh Field Station, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada
diou, Pascal H.J., and L. Gordon Goldsborough. 2006. Northern range expansion and invasion by the Common Carp,
Cyprinus carpio, of the Churchill River system in Manitoba. Canadian Field-Naturalist 120(1): 83-86
cent fisheries data from northern Manitoba indicates that the Common Carp (Cyprinus carpio) has extended the northern
lit of its range. Additionally, it also appears that carp have invaded and established viable populations in the Manitoba
rtion of the Churchill River. Habitat degradation and altered flow regimes as result of hydroelectric development in north-
1 Manitoba may have facilitated the expansion of carp in the region.
y Words: Common Carp, Cyprinus carpio, exotic species, range expansion, invasion, habitat disturbance, Manitoba.
The Common Carp (Cyprinus carpio) was intro-
ced intentionally to Manitoba, Canada, when fish
ym Minnesota were stocked in the Assiniboine River
itershed southeast of Brandon in 1886 (Stewart and
atkinson 2004). These early attempts at introduc-
n failed, and carp only became established after pop-
itions from the northern U.S. entered the province
‘ough the Red River (Atton 1959). In fact, the Com-
mn Carp was largely unknown in Manitoba until 1938,
1en it was positively identified in the Red River near
ckport (Hinks 1943). Today, the Common Carp is
und throughout most of the southern and central
Zions of the province, but particularly in the Red and
siniboine rivers, and lakes Manitoba, Winnipeg, and
innipegosis. Common Carp do not appear to have ex-
nded into the eastern portion of the province, likely
e to the fact that they cannot easily migrate upstream
this region because of the many obstructions they
juld encounter (i.e., Beaver dams). Additionally, lakes
this portion of the province are characterized by
cky substrates and clear deep waters typical of the
ecambrian Shield and do not provide suitable habi-
for viable populations of the Common Carp.
Benthivorous fish such as the Common Carp can
ve profound effects in aquatic ecosystems by increas-
2 turbidity through sediment resuspension while for-
ing and spawning (Breukelaar et al. 1994; Zambra-
yet al. 2001). Carp are known to uproot submerged
acrophytes during spawning and accidentally con-
me them while foraging for benthic invertebrates
olterman 1990; Roberts et al. 1995; Lougheed et al.
98; Zambrano and Hinojosa 1999). Sediment resus-
nsion and excretion by carp can increase water col-
nn nutrient concentrations, thereby causing phyto-
ankton to flourish (Lamarra 1975; Breukelaar et al.
94: King et al. 1997). A shading effect results from
ese blooms, further suppressing submerged macro-
phytes and creating an ecological feedback mechanism
that perpetuates the turbid state (Scheffer 1998). Carp
physically disturb the sediment-water interface and the
algae associated with sediment. These algae play a
major role in the stabilization of the bottom sediments
(Taylor et al. 1998) and the regulation of nutrient fluxes
across the sediment-water interface (Goldsborough and
Robinson 1985; Carlton and Wetzel 1988; Woodruff
et al. 1999). It is thought that many of the once clear,
shallow lakes and wetlands in southern Manitoba have
switched to the turbid state due to the loss of sub-
merged macrophyte cover which, in turn, was caused
by the proliferation of Common Carp (Badiou 2005).
Before 1940, carp were found only in the waters of
the Red River (Figure 1a). After invading the province
from the south, the Common Carp spread quickly
throughout the Red and Assiniboine River watersheds
prior to the 1950s and established themselves in the
south basins of lakes Winnipeg and Manitoba (Figure
1b). However, the greatest range expansion occurred
between 1950 and 1970 when carp were easily able to
access the northern and western portions of the province
after successfully invading Lake Winnipeg, Lake Man-
itoba, and Lake Winnipegosis, the largest lakes in the
province (Figure Ic). Swain (1979) indicated that by
1976 the Common Carp was found in most large lakes
and virtually all major rivers in Manitoba, with the
exception of the Churchill River system.
A specimen collected from Split Lake (Figure Id)
in 1963 was considered to be the most northerly record
for the distribution of carp in the world (McCrimmon
1968). These original reports stated that carp caught
from Split Lake appeared emaciated and unhealthy.
According to Atton (1959), cold summer temperatures
prevented the carp from invading the Churchill River
system. However, as shown in Figure 1d, the Common
Carp has migrated northward through the Nelson River
83
84 THE CANADIAN FIELD-NATURALIST
system and, in the last decade, has become established
in the Churchill River system according to carp pro-
duction data spanning the period from 1970 to 2004.
On the Nelson River system, carp have become
established in Split Lake where commercial catches
(3-74 kg per year) have been reported starting in 1996.
Fish monitoring studies for the Limestone Generating
Station (C. Barth, personal communication) and the pro-
posed Gull Generating Station (D. J. Pisiak, unpub-
lished data) documented Common Carp in Stephens
Lake and in the forebay of the Limestone Generating
Station (Figure 1d). Furthermore, during the course of
fish studies for the proposed Conawapa Generating
Station, a large, approximately 4 kg, Common Carp
was caught in a backwater area near the lower Lime-
stone Rapids (P. Nelson, personal communication).
This is the first report of Common Carp downstream
of the Limestone Generating Station and the most
northerly location documented for the Nelson River.
More important than the northward expansion of
carp on the Nelson River system is the invasion of the
Churchill River system in the last decade. Recent fish-
eries records for northern Manitoba provided by the
Freshwater Fish Marketing Corporation indicate that
the Common Carp is now established in and around the
Churchill River system with commercial catches report-
ed from Britton, Guthrie, Highrock, Loon, Sisipuk, and
Southern Indian lakes (Figure 1d). Commercial catches
in these lakes were much higher (range 8—1184 kg per
year) relative to those from Split Lake on the Nelson
River system. Carp were likely introduced into the
Churchill River system from the Saskatchewan River
system, either by fishers using them as bait or by birds
that may have inoculated carp eggs from one system
into the other. A thorough search of the primary liter-
ature and government documents revealed that Com-
mon Carp have not been reported in the Saskatchewan
portion of the Churchill River. The fact that carp have
not been reported in the Churchill River in Saskatche-
wan indicates that the transfer of the Common Carp
between the Saskatchewan River and Churchill River
systems must have occurred within the province of
Manitoba in the vast wetland area that occurs be-
tween these two watersheds along the Manitoba/Sas-
katchewan border.
Based on the fact that carp were found in Split Lake
as early as 1963, it is not surprising that they have con-
tinued to migrate northward through the Nelson River
system. However, given the poor condition of carp in
Split Lake in 1963, a change must have occurred,
which has allowed them to persist in this system. The
two basic habitat requirements necessary to sustain carp
populations are: (1) a shallow marsh environment with
abundant aquatic vegetation for spawning and, (2) an
area of deep water where carp can overwinter (Mc-
Crimmon 1968). We hypothesize that the construction
of the Kettle Generating Station in 1974 at the outlet of
Stephens Lake on the Nelson River provided the neces-
sary habitat requirements to allow the establishment and
Vol. 120
sustain viable populations of Common Carp in Split Lake
and Stephens Lake. Construction of this generating sta-
tion increased the size of Stephens Lake by 242 km?
and resulted in the flooding of 192 km? of land (Envi-
ronment Canada / Fisheries and Oceans 1992a). This
created an extensive area of shallow marsh-like habitat
where carp could spawn in the summer. This was the
case in Ontario where Swee and McCrimmon (1966)
reported that Lake St. Lawrence near Cornwall, formed
as the result of flooding during hydroelectric develop-
ment on the St. Lawrence River, provided ideal spawn-
ing habitat for the Common Carp. However, it was also
noted that fluctuations in lake level associated with
hydroelectric activities caused many of the eggs at-
tached to vegetation to be exposed and destroyed. More
importantly, the construction of the Kettle Generating
Station, with a forebay head of 30 meters (Environment
Canada / Fisheries and Oceans 1992a), created a deep
basin with warmer profundal zone waters to which
carp can escape during cold winter months.
The completion of the Churchill River Diversion in
1976 raised the water level in Southern Indian Lake by
approximately 3 m and flooded 187 km? of surround-
ing land (Environment Canada / Fisheries and Oceans
1992a). Like the flooding experienced on Stephens
Lake after the construction of the Kettle Generating
Station, flooding on Southern Indian Lake would have
also provided ideal spawning habitat for carp. The
extensive flooding on Southern Indian Lake increased
shoreline erosion and drastically increased suspend-
ed sediment concentrations in the lake from 5 mg-L"!,
pre-impoundment, to 30-50 mg-L"', post-impoundment
(Environment Canada / Fisheries and Oceans 1992b).
This anthropogenic increase in suspended sediments
would have conferred a competitive advantage on carp
which can feed more efficiently in highly turbid waters
due to their superior olfactory senses, relative to sight-
feeders (Panek 1987).
Water flows on the Churchill and Nelson river sys-
tems have been altered dramatically as a result of the
Churchill River Diversion, which has diverted 80% of
the flow from the Churchill River at Southern Indian
Lake into the Nelson River system. Carp are found in
large, slow-moving rivers as well as fast flowing streams
(Panek 1987). Due to the ability of carp to inhabit wa-
ters of varying flow, the increased flow on the Nelson
and decreased flow on the Churchill, as well as the in-
creased variability in flows in both systems as a result
of hydroelectric development, would also have con-
ferred an advantage on the Common Carp relative to
native fish that may be less tolerant of variations in
flow regime.
Overall, it is not surprising that Common Carp are
invading northern Manitoba in step with hydroelec-
tric development, as many authors have demonstrated
the link between anthropogenic habitat alteration and
invadability of ecosystems (Moyle and Light 1996;
Keith and James 1999; Byers 2002). Keith and James
(1999) showed a positive correlation between the num-
2006 BADIOU AND GOLDBOROUGH: CARP OF THE CHURCHILL RIVER SYSTEM 85
ex
ats pen
(Wax a Ls raz
uae RZ
ip oA Hudson Bay
Ficure |. Range expansion of the Common Carp (Cyprinus carpio) in Manitoba (A) prior to 1940, (B) prior to 1950, (C)
prior to 1970, and (D) prior to 2000. Carp distributions prior to 1970 were estimated based on date of first occur-
rences reported in Swain (1979), while the range expansion between 1970 and 2000 was estimated from carp pro-
duction data provided by Manitoba Conservation and the Freshwater Water Fish Marketing Corporation for the
period from 1970 to 2000. Numbers in D indicate locations discussed in the text: (1) Split Lake, (2) Stephens Lake.
(3) Limestone Generating Station, (4) Britton Lake, (5) Sisipuk Lake, (6) Guthrie Lake, (7) Highrock Lake, (8)
Southern Indian Lake, (9) Loon Lake, (10) Churchill River Estuary, and (11) Nelson River Estuary.
86 THE CANADIAN FIELD-NATURALIST
ber of reservoirs and the number of introduced species
in North American drainages. Given that the Common
Carp has already been reported in subarctic regions
(Lukin 1999) and is tolerant of a wide range of salin-
ity levels (Lam and Sharma 1985) it is a matter of time
before the populations of carp established on the Nel-
son and Churchill river systems reach the Churchill
River and Nelson river estuaries.
Acknowledgments
We thank Ellen Smith of the Freshwater Fish Mar-
keting Corporation for providing us with carp produc-
tion data from Manitoba lakes. We also thank Patrick
Nelson, Paul Graveline and Cam Barth of North/South
Consultants Inc., in Winnipeg for helpful comments.
This is publication number 311 from the Delta Marsh
Field Station (University of Manitoba).
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Received 21 March 2005
Accepted 30 January 2006
Diversity and Range of Amphibians of the Yukon Territory
BRIAN G. SLOUGH! and R. LEE MENNELL?
' 35 Cronkhite Road, Whitehorse, Yukon Territory Y1A 5S9 Canada
2 P.O. Box 10002, Whitehorse, Yukon Territory Y1A 7A1 Canada
Slough, Brian G., and R. Lee Mennell. 2006. Diversity and range of amphibians of the Yukon Territory. Canadian Field-Naturalist
120(1): 87-92.
Four amphibian species occur in the Yukon: Western Toad (Bufo boreas; first verified record in 1961), the Boreal Chorus Frog
(Pseudacris maculata, first record in 1995), the Columbia Spotted Frog (Rana luteiventris, first record in 1993), and the Wood
Frog (Rana sylvatica; first record in 1933). The Western Toad is restricted to the Liard Basin in five geographically separated
areas. Breeding sites have been located on the lower Coal River and vicinity. The Boreal Chorus Frog has been documented
from a small area of the lower La Biche River valley near the Yukon-British Columbia-Northwest Territories border. The Colum-
bia Spotted Frog occurs in two widely separated areas, at Bennett Lake in the southwest and in the Liard Basin in the southeast
The Wood Frog is widespread below treeline. Limited survey efforts continue to hinder our knowledge of amphibian distribution.
Key Words: Bufo boreas, Western Toad, Pseudacris maculata, Boreal Chorus Frog, Rana luteiventris, Columbia Spotted Frog,
Rana sylvatica, Wood Frog, amphibian, Yukon, distribution.
Biological surveys began in the interior of the Yukon
Territory in 1899 (Osgood and Bishop 1900); however,
these were often directed at mammals and birds, and
did not report amphibian observations. The earliest
known amphibian records for the region are from
Atlin (Slevin 1928; Western Toad and Wood Frog) and
Bennett Lake, British Columbia (Logier 1932; Colum-
bia Spotted Frog, formerly Western Spotted Frog).
The first published report of amphibians in the Yukon
was by Loomis and Jones (1953), who reported a
Wood Frog collected at Dezadeash Lake in 1948 by
the University of Kansas Museum of Natural History.
Museum collections from the Canadian Museum of
Nature (CMN) and the American Museum of Natural
History (AMNH) date to 1933 and 1936, respectively.
Logier and Toner (1961) report only the single locali-
ty record of Loomis and Jones (1953). Using an ex-
panded database, including museum specimens and
inferences from locality records in Alaska, the North-
west Territories and British Columbia, Martof (1970)
shows an approximate range for the Wood Frog;
although a large area with no data existed in the east-
ern and northern Yukon. The amphibian range maps
in Stebbins (2003) were modified with information
provided by BGS.
The Western Toad (Bufo boreas) was first observed
in 1961 in the southeast Yukon (Cook 1977). The
Columbia Spotted Frog was first observed in 1993
(Mennell 1997) in the southwest Yukon. There are no
published records of the Boreal Chorus Frog or other
species for the Yukon. All species of amphibians have
been underrepresented in records due to the remote-
ness of breeding sites and lack of directed surveys.
Several amphibian surveys and expert observations
have been made since 1993 by the authors and others,
and the data have remained unpublished. We typical-
ly used visual encounter surveys and dipnetting of
terrestrial, semi-terrestrial (wetland) and aquatic habi-
tats (Thoms et al. 1997). We review unpublished data,
unpublished reports and museum collections to pres-
ent an update on the amphibian species of the Yukon
and their ranges. Anecdotal information on phenology
and habitats is provided. Scientific and common
names follow Collins and Taggart (2002).
Species Accounts
WESTERN TOAD (Bufo boreas)
Western Toads occur from south-coastal Alaska,
through western Canada and the western United States
to Baja California. Cook (1977) reported two museum
specimens of the Western Toad from the Yukon. The
Whitehorse specimen, collected in 1948, was verified
in 1997 by F. Cook, who also verified the label in 2003
as collected by William Mason July 1-10. Mason was
an entomologist and the date notation suggests a spec-
imen taken in a pan or similar trap set for that period
and contents preserved on the last date (F. Cook, per-
sonal communication). This specimen is here consid-
ered to be either an extra-limital stray or erroneously
labelled, since the nearest confirmed breeding records
of Western Toads are 100 km to the south in British
Columbia on Bennett Lake (Slough 2004*; 17 km
from the British Columbia-Yukon border) and on
Tagish Lake (Mennell and Slough 1998*; 40 km
from the border). A single adult was found in non-
breeding habitat on Tagish Lake 9 km from the border
(Mennell and Slough 1998*). All other records are
from the Liard Basin in the southeast Yukon, where
the Western Toad is now known from at least 11 popu-
lations in five geographically separated areas (Figure
1). These are summarized from west to east.
Two adult Western Toads were observed near a geo-
thermal spring on the Meister River (60°17'N,
130°07'W) in 1988 (D. Mossop, personal communi-
87
88 THE CANADIAN FIELD-NATURALIST
cation). RLM searched this area in 2004 but did not
observe Western Toads. A single adult was observed
near Upper Liard (60°05’°N, 128°58’W) in 2004 (C.
Eckert, personal communication; verified from pho-
tographs).
At least six breeding populations of Western Toads
occur in close proximity (within 16 km) on the lower
Coal River and wetlands to the west. The most visited
population and breeding site is the Coal River Springs
(60°09'N, 127°26'W). G. Scotter collected two speci-
mens in 1977 (CMN 17798-1 and -2), and at least six
Western Toad records of adults and metamorphs were
documented between 1982 and 2004 (B. Slough and
L. Mennell, unpublished data; Slough 2005a*). Two
breeding sites exist in spring marshes below the upper
and lower springs. The temperature of the main source
pool is 11°C, likely maintaining frost-free hibernating
sites nearby. Other breeding sites occur in riparian
floodplain backwater channels and ponds above the
springs (60°11'N, 127°29'W), below the springs on an
island in the Coal River (60°07'N, 127°24'W), and fur-
ther downstream on the Coal River in a bedrock pool
[60°08'N, 127°14'W] (J. Staniforth and J. Meikle, per-
sonal communication; L. Mennell, unpublished data;
Slough 2005a*). Juvenile toads have also been ob-
served in the chain of lakes southwest of the springs
(60°14'N, 127°29'W) [B. Slough, unpublished data;
Slough 2005a*] and in a tributary of the lakes (60°06'N,
127°32'W) [Slough 2005b*], suggesting the presence
of breeding populations.
Cook (1977) reported a specimen collected in 1961
from North Toobally Lake (60°20'N, 126°15'W). An
adult Western Toad was observed 23 km to the south
near South Toobally Lake (60°08’N, 126°19’W) in
1996 (C. Eckert, H. Grunberg, P. Sinclair, personal
communication; verified from photograph). A single
adult Western Toad was observed on the lower Beaver
River [60°02’N, 124°32’W] in 1997 (B. Bennett, V.
Loewen, J. Staniforth, personal communication; veri-
fied from photograph). All of the Western Toad local-
ities are low elevation sites with early deep snow accu-
mulation. The snow cover likely prevents deep frost
penetration and permits safe hibernation, as speculated
by Cook (1977).
Western Toad breeding sites are predominantly shal-
low, silty areas on ponds, lakes and rivers. Adults are
often encountered far from water in forests or mead-
ows. Breeding dates in the Yukon are unknown; how-
ever, populations in northwestern British Columbia
breed in late May (B. Slough and L. Mennell, unpub-
lished data). A population in the Atlin Warm Springs,
20 km south of Atlin, breeds in late February-early
March. Tadpoles have been observed on the Coal River
in July, and tadpoles and metamorphs were observed
there on 10 August 2004.
BOREAL CHORUS FROG Pseudacris maculata
While Pseudacris maculata is currently considered a
distinct species following Platz (1989), it was formerly
Vol. 120
called P. triseriata maculata (and earlier Pseudacris
triseriata septentrionalis). Recent phylogeographic
studies of North American chorus frogs have shown
that range limits and taxonomy of the triseriata com-
plex may need revision (Moriarty and Cannatella 2004).
The Boreal Chorus Frog ranges across much of cen-
tral North America east of the continental divide, from
the Northwest Territories to southern James Bay and
south to Arizona and New Mexico. We present the first
records of the Boreal Chorus Frog from the Yukon.
Boreal Chorus Frog calls were identified at five natural
wetlands and roadside/airstrip ditches along a 10 km
section of road and airstrip in the lower La Biche River
valley in the southeast Yukon (60°03'N 123°58'W to
60°08'N 124°03'W) between 8 and 17 June in 1995,
1997 and 1999 (Figure 1) [B. Bennett, C. Eckert, V.
Loewen, P. Sinclair, J. Staniforth, personal communi-
cation; B. Slough, unpublished data]. BGS also iden-
tified Boreal Chorus Frog tadpoles there on 10 June
1999. Extensive surveys by the authors in southeastern
Yukon in 2004 did not yield any further observations
of the species. Matsuda et al. (2006) show the species
160 km to the southeast at Fort Nelson, British Colum-
bia, and it occurs 125 km northeast at the junction of
the South Nahanni and Liard rivers (Fournier 1998*).
COLUMBIA SPOTTED FROG (Rana luteiventris)
The Columbia Spotted Frog occurs between the
Rocky Mountains and Coast Ranges from the Yukon
to Utah and Nevada in the south. Mennell (1997)
located two populations of Columbia Spotted Frogs
adjacent to the West Arm of Bennett Lake in 1993, in
a pond (60°05'N 135°01'W) and a beaver pond on a
creek (60°O1'N 135°12'W) [Figure 1]. Slough (2002)
obtained single voucher specimens and preserved toe-
tips (for phylogeographic studies, n=10 and 11, respec-
tively) from the two ponds in 2001 (Slough et al. 2002*).
A third population was found in a beaver pond in the
same area (Slough 2002) [60°03’N 134°58’W] where
a voucher specimen and six toe-tips were obtained;
specimens are held at the Redpath Museum, McGill
University [RM 3382, 3379 and 3381 for the three
populations, respectively]. Additional Columbia Spot-
ted Frog adults (n=7 in 4 sites) were observed within
2.5 km of Site 2 in apparent non-breeding habitats (i.e.,
early life-stages not observed) along the Partridge River
to Bennett Lake (R. McClure, personal communication,
verified from photograph; B. Slough and L. Mennell,
unpublished data). These three breeding populations
are the most northerly of a series of populations that
extends into northwestern British Columbia (Matsuda
et al. 2006, Mennell 1997; Mennell and Slough 1998*;
Slough et al. 2002*).
In 2003, two adult Columbia Spotted Frogs were ob-
served in Irons Creek Lake in the southeastern Yukon
(60°00’N 127°05’W; P. Hovingh, personal communi-
cation to Yukon Fish and Wildlife Branch) (Figure 1).
Several adults, juveniles and one egg mass were ob-
served in 2004 and 2005 (Slough 2005b*). The long-
2006 SLOUGH AND MENNELL: AMPHIBIANS OF THE YUKON 89
Key to Map Symbols
Boreal Chorus Frog
Columbia Spotted Frog
Westem Toad
FiGuRE 2. Distribution of amphibians in the Yukon. Refer to text for data sources. Shaded areas are mountain ranges.
90 THE CANADIAN FIELD-NATURALIST
term prospects of this population are unknown as this
small lake has regularly been stocked with Rainbow
Trout (Oncorhynchus mykiss) since 1990, with the lat-
est stocking occurring in 2003 (D. Toews, personal
communication). Rainbow trout are known to prey on
Columbia Spotted Frog tadpoles and juveniles, which
overwinter in lakes (Pilliod and Peterson 2000). Fur-
thermore the heavily forested shoreline habitat is a-
typical for Columbia Spotted Frogs (Ovaska 1999*).
The species has been observed only on human-altered
shoreline which has been cleared of trees and graded
into Irons Creek Lake, creating shallows. In 2004, RLM
observed an adult Columbia Spotted Frog 15 km west
of this site on an ox-bow channel of the Hyland River
(60°O1'N 127°12'W).
The Columbia Spotted Frog localities in the Liard
Basin and Coast Mountain Cordillera are in areas with
early deep snow accumulation which likely prevents
thick ice formation and permits safe aquatic hibernation.
Columbia Spotted Frogs were noticeably abundant
in and around pond outlets, where they were invariably
perched at the waters edge. Breeding and transforma-
tion dates in the Yukon are unknown; however popu-
lations in nearby northwestern British Columbia have
been observed extensively (Mennell 1998*; B. Slough,
unpublished data). Breeding takes place on a single
warm day from 18-22 May (six years of observations)
in traditional shallow basins along the margins of ponds.
Some of the ponds may be partially ice-covered with
melting snow surrounding the ponds. They breed com-
munally with up to 30 individuals congregating in a few
square metres. Males begin calling above water over
a larger area and finally call underwater as the aggre-
gation consolidates and begins amplexis, leaving a mass
of egg clutches unattached in shallow water. Breeding
sites are occasionally compromised by falling water
levels, leaving egg masses landlocked and stranded.
Tadpoles are free-swimming in 14-16 days and meta-
morphs appear on pond edges throughout August (Men-
nell 1998*). Metamorphs have also been observed in
late May, and tadpoles have been observed in mid-
September at higher elevation ponds, suggesting that
some tadpoles overwinter.
Woop FROG (Rana sylvatica)
The Wood Frog ranges from northern Alaska and
the Yukon through most of Canada below treeline east
to the Atlantic and south to the Appalachian Mountains.
Disjunct populations occur in several states south of this
range. Numerous collections and observations reveal
that Wood Frogs are common and abundant in wet-
lands across the Yukon below the arctic and alpine tree-
line (B. Slough, unpublished data) [Figure 1]. The most
northerly location is Ney Khwi Vun (“Frogs in the
Lake’), on the Old Crow Flats (68°01'N 140°05'W;
Gray and Alt 2000*). Museum collections exist in
the University of Kansas Museum of Natural History
(n = 1), the American Museum of Natural History (n
Vol. 120
= 16), the Canadian Museum of Nature (n=56) and
the Auke Bay Laboratory, Alaska Fisheries Science
Center (Hodge 1976; n = 2). One of the museum spec-
imens from mouth of Coal Creek (AMNH 44751) was
originally misidentified as Pseudacris septentrionalis
(now P. maculata) until verification as a Wood Frog
in 1995 by L. S. Ford Curatorial Associate, AMNH
(F. Cook, personal communication to P. Milligan). In
July 2000, BGS obtained Wood Frog samples for phy-
logeographic studies (J. Irwin and J. Lee- Yaw, unpub-
lished data); eight juveniles from Nogold Creek, west
of Mayo (63°26'N 135°06'W), and 10 juveniles from
south of Whitehorse (60°35’N 134°58’W). The CMN
also has tissue from 11 adults and 25 juveniles from
“Paddy’s Pond”, Whitehorse, July and August 1992
by Gerry Whitley.
The Wood Frog specimens and observations (Fig-
ure 1) document an extensive range for the species in
the Yukon. Most observations are from road and river
access corridors, and gaps in the range exist in inac-
cessible areas. We believe that the species occupies
suitable habitats below treeline within these gaps. Fur-
thermore, the distribution is continuous in areas of ex-
tensive wetlands such as the Whitehorse area, where
only a small sample of locations was plotted. Yukon
Wood Frogs were assigned to the Alaskan phenotype
by Martof and Humphries (1959). While we have no
data on most visible characters used to differentiate the
types, there is a north-south gradient in the frequency
occurrence of the mid-dorsal stripe, from approximately
100% in the north and central Yukon to 50% or less in
the south.
Wood frogs breed in the Yukon as soon as shorelines
become ice-free in late April-early May. They breed
over a period of up to two weeks, they are frequently
solitary breeders, and they do not necessarily use tra-
ditional breeding sites, but use any site with shallow
water and sparse emergent grass or sedge vegetation.
Egg masses are attached singly to grass or sedge stalks.
Metamorphs move onto land throughout July. The most
common Wood Frog breeding habitats are shallow,
permanent and fishless ponds.
Neighbouring Species
Three species of salamanders occur near the south-
western Yukon, but have yet to be recorded there. The
Long-toed Salamander (Ambystoma macrodactylum)
ranges as far north as the Taku River (Hodge 1976; P.
Milligan, personal communication; Mennell and
Slough 1998*) and upstream to the junction of the
Nakina and Sloko rivers (K. Heinemeyer, personal
communication), 110 km from the British Columbia-
Yukon border (observation not shown by Matsuda et
al. 2006). The Northwestern Salamander (Ambystoma
gracile) and Roughskin Newt (Taricha granulosa) oc-
cur in Southeast Alaska (MacDonald 2003), but they
occupy Pacific coastal habitats not present in the Yukon.
2006
Future Surveys
Dedicated amphibian surveys and expert observa-
tions have greatly increased our knowledge of the
species and their ranges in the Yukon. We presented
significant new observations of Western Toads, Colum-
bia Spotted Frogs and Wood Frogs for the Yukon, and
the first records of Boreal Chorus Frogs. Continued
efforts should identify additional populations, their
degree of isolation and habitat requirements; all nec-
essary for conservation planning. The ranges of the
Western Toad, Columbia Spotted Frog and Boreal Cho-
rus Frog in the Yukon imply data deficiencies for these
species in northern British Columbia. The Columbia
Spotted Frog is common on Atlin Lake, south of Atlin
(Slough et al. 2002*; not shown by Matsuda et al.
2006).
Acknowledgments
Funding to BGS and RLM for field surveys was
provided by NatureServe Yukon, Yukon Department
of Environment; and by Northern Research Endow-
ment Grants from the Northern Research Institute,
Yukon College. Canadian Parks and Wilderness Soci-
ety- Yukon Chapter kindly supported BGS on several
wilderness trips. We are grateful to the many individ-
uals who reported amphibian sightings. Significant
Wood Frog sightings were reported by D. Mossop, J.
Meikle, L. Karnis, M. Gill, R. Lamb and G. Whitley.
E. Wind and T. Jung reviewed earlier versions of the
manuscript. G. Perrier of Dragonfly Maps provided
digital cartographic design of the distribution map. We
thank P. Milligan for his work as the founding Yukon
DAPCAN coordinator.
Documents Cited [marked * in text citations]
Fournier, M. 1998. Amphibians & reptiles in the Northwest
Territories. A brochure prepared for Ducks Unlimited,
Environment Canada, Indian and Northern Affairs Canada,
and Northwest Territories Resources, Wildlife and Eco-
nomic Development, Yellowknife, Northwest Territories.
Ecology North, Yellowknife.
Gray, D. R., and B. T. Alt. 2000. Resource description and
analysis of Vuntut National Park of Canada. Western Cana-
da Service Centre, Parks Canada, Vancouver, British Colum-
bia. 650 pages.
Mennell, R. L. 1998. The ecology and population status of
the Columbia spotted frog (Rana luteiventris) at their
northern range limit. Prepared for the Northern Research
Institute, Yukon College, Whitehorse. 21 pages.
Mennell, R. L., and B. G. Slough. 1998. Amphibian and
biodiversity inventories of ecoregions in northwestern
British Columbia. Habitat Conservation Trust Fund Pro-
ject TF28056. Ministry of Environment, Lands and Parks,
Victoria, British Columbia. 86 pages.
Ovaska, K. E. 1999. Status report on the Columbia spotted
frog, Rana luteiventris, in Canada. Committee on the Sta-
tus of Endangered Wildlife in Canada. 55 pages.
Slough, B. G. 2004. Western toad inventory in the Chilkoot
Trail National Historic Site, July-August 2004. Parks Cana-
da Species at Risk Inventory Fund Project SARINV04-30.
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Received 10 March 2005
Accepted 30 November 2005
Temporal Variability of Cetaceans near Halifax, Nova Scotia
PETER SIMARD!:2, JENNIFER L. LAWLOR?, and SHANNON GowaAns! 4
' Blind Bay Cetacean Studies, 144 Victoria Road, Dartmouth, Nova Scotia B3A 1V7 Canada
+ University of South Florida, College of Marine Science, 140 7" Avenue South, St. Petersburg, Florida 33701 USA; e-mail
psimard@ marine.usf.edu.
‘Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada
‘Eckerd College, 4200 54" Avenue South, St. Petersburg, Florida 33711 USA
Simard, Peter, Jennifer L. Lawlor, and Shannon Gowans. 2006. Temporal variability of cetaceans near Halifax, Nova Scotia.
Canadian Field-Naturalist 120(1): 93-99.
Annual and seasonal trends in sightings of coastal cetaceans near Halifax, Nova Scotia, were studied using observations from
whale watching and dedicated research vessels from late spring to early fall of 1996 to 2005. Four species of cetaceans rou-
tinely used the area during the summer: White-beaked and Atlantic White-sided dolphins (Lagenorhynchus albirostris and
L. acutus), Harbour Porpoises (Phocoena phocoena), and Minke Whales (Balaenoptera acutorostrata). The dolphin species
were temporally separated, with White-beaked Dolphins being common earlier in the summer than White-sided Dolphins.
White-sided Dolphins were unusually abundant in 1997, and were found in larger groups (mean = 46.5 + 46.19 sd) than White-
beaked Dolphins (mean = 9.1 + 5.19 SD). The area also appears to be an important habitat for dolphin calves and juveniles
of both species. Fin Whales (B. physalus) were commonly observed in relatively large groups in 1997, but were uncommon
or absent in other years. Humpback Whales (Megaptera novaeangliae), Blue Whales (B. musculus) and North Atlantic
Right Whales (Eubalaena glacialis) were uncommon in the area, although Humpback Whales were sighted frequently in
1997. Increased numbers of White-sided Dolphins, Fin and Humpback whales in 1997 may be explained by increased prey
abundance and decreased sea-surface temperatures.
Key Words: White-beaked Dolphin, Lagenorhynchus albirostris, Atlantic White-sided Dolphin, Lagenorhynchus acutus, Har-
bour Porpoise, Phocoena phocoena, Minke Whale, Balaenoptera acutorostrata, Fin Whale, Balaenoptera physalus,
Humpback Whale, Megaptera novaeangliae, seasonality, group size, calves, Nova Scotia.
In temperate ecosystems, it is common for cetaceans
to exhibit seasonal and annual fluctuations in their dis-
tribution and abundance (e.g., Hooker et al. 1999).
These fluctuations are thought to be a reflection of
seasonal changes in the environment, such as sea sur-
face temperature (e.g., Hooker et al. 1999) and prey
distribution (e.g., Kenney et al. 1996). Monitoring
highly mobile, top level predators such as cetaceans
serves not only to identify their habitat use patterns and
track their population sizes, but also to monitor the
health of the overall marine ecosystem.
The area between Halifax and Mahone Bay, Nova
Scotia, is habitat for several species of cetaceans. It is
used extensively by the commercial fishing industry,
commercial shipping (especially near Halifax Harbour)
and recreational boaters, and it is being used increas-
ingly for whale watching. Given the high potential for
anthropogenic influences in the Halifax area, there is
good reason to study and monitor cetaceans there.
Methods
Field Methods
Data were collected from 1996 to 2005, primarily
from 1 June to 30 September. From 1996 to 1999,
12.6 m and 14 m whale watching vessels were used to
make three 3-hour trips daily, weather permitting. In
2000 and 2001, an 8.5 m auxiliary sailing vessel was
used, and in 2002 to 2005 a 4.2 m rigid inflatable boat
was used. The latter two vessels were used only for
research, and cruises in 2000 to 2005 were generally
conducted only in weather conditions suitable for
photo-identification. All surveys were limited to day-
light hours and the area bounded by approximately
44°19' to 44°38'N latitude and 63°15' to 64°10'W lon-
gitude (Figure 1). The path of all vessels was oppor-
tunistic, based on where cetaceans were previously
sighted. In 1996, only date, time, species, group size
and the presence of calves and juveniles were recorded.
In subsequent years, latitude and longitude positions
were recorded (using GPS) for hourly search effort and
species positions and photo-identification data were col-
lected. Only those sightings identified to species level
were included in analysis.
Animals were classified as adults, juveniles or calves.
For odontocetes, first year calves were identified by
having a length of <50% of adult body length, while
juveniles were identified by having 50% — 70% of
adult body length (Reeves et al. 1999; Lien et al. 2001:
Bjorge and Tolley 2002). The identification of calves
was often facilitated by their position near an adult,
with younger calves displaying “chin-slap”™ surfacings,
and in the case of neonatal calves, by fetal folds and
bent-over dorsal fins (Weinrich et al. 2001). For mys-
ticetes, calves were identified by having a length of
94 THE CANADIAN FIELD-NATURALIST
<40%, and juveniles by having a length of 40% — 60%
of adult body length (Aguilar 2002; Clapham 2002;
Perrin and Brownell 2002).
Over the ten-year study period, changes in method-
ology introduced several biases to our data. Data col-
lected from the whale watching boats (1996-1999)
were particularly problematic as the boat returned sev-
eral times per day with new passengers to the same
area where cetaceans had previously been sighted,
potentially causing groups to be double-sampled and
introducing a positive bias to calculations of hourly
sighting rates. Cetacean encounters on the whale watch-
ing vessels were typically shorter in duration, and there-
fore a larger proportion of time was spent searching for
new groups, introducing another potentially positive
bias. However, the whale watching vessels were more
likely to operate in inclement weather than the re-
search vessels, which should negatively bias the
sighting data. Larger groups were preferred on whale
watching cruises, positively biasing group sizes. Ves-
sel cruising speed also varied greatly in the nine year
study period (ranging from 5 to 15 knots). These dif-
ferences in survey speed changed our spatial sampling
greatly, and therefore likely affected the sighting data.
The height of observers above the sea surface also var-
ied greatly, from a minimum 1.25 m to a maximum
5 m. Search effort varied greatly between different
years and months, and as a result time periods with
low effort may have had unreliable sighting rates (see
Table la and Ib).
Calculation of Sighting Indices
Because of these biases, regular group sighting rates
(sightings per hour) would not be directly comparable.
Therefore, the data were restricted in order to calcu-
late a “sighting index” for each species per year and
per month. Potential positive biases from repeated
sampling of the same cetaceans throughout the day
were eliminated by daily scoring each species as pres-
ent or absent. This restriction also reduces biases intro-
duced from differences in vessel speed, and time spent
searching vs. time spent with groups of cetaceans. Cal-
culated sighting indices could range from a maximum
of 1.0 (sightings occurring every day of field effort)
to 0.0 (no sightings). In order to determine if weather
significantly affected sighting rates, the sighting rates
during good weather and bad weather were subjected
to a chi-squared test. Good weather was defined as
visibility = 1 km, wind < Beaufort 3 (< = 19 km/hr),
wave height < 0.75 m, and swells < 1.5 m. Results
from this test indicate that sighting rates were signifi-
cantly lower in poor weather (y? = 0.9999, df = 5,
0.025
75%.
Coniferous Crown closure 2 10%, coniferous species > 75%.
Mixedwood Crown closure 2 10%, neither type > 75%
Habitat type Description
Immature-Pole
Young
Mature-old
0-40 years old stands. Open areas and new stands. Pole stands are
thick stands of trees (7.5 to 12.4 cm diameter at breast height),
usually with little understory.
40-80 year-old forests. Achievement of dominance by some trees
and death of other trees leads to reduced competition that allows
understory plants to become established. The forest canopy has
begun differentiation into distinct layers. Vigorous growth and a
more open and multi-storied stand than in the pole stage.
> 81 years old late successional stands consisting mostly of mature
stands with even canopy of trees, with or without coarse woody
debris down and leaning logs. A few old stands with tall and large
canopy trees, canopy gaps, large snags, large downed woody
debris, and developed understories. A second cycle of shade
tolerant trees may have become established.
to identify animals. When encountering animals, foot-
prints and trails were studied to ascertain criteria used
to differentiate American Marten from Fisher.
Autocorrelation may occur during analysis of track
survey data because of the uncertainty in whether one
or more animals have made the tracks being counted
(Proulx and O’Doherty 2006). It is sometimes difficult
to confirm that a series of tracks along a transect be-
longs to the same animal (de Vos 1951), as home ranges
overlap (Buskirk and Ruggiero 1994), and winter dis-
persal movements are known to occur (Clark and
Campbell 1976). On the basis of track characteristics,
Proulx et al. (2005) deduced that tracks of two different
animals could be as close as 100 m from each other
along the same transect. To minimize spatial autocor-
relation, only tracks >100 m apart within the same for-
est stand were recorded (Bowman and Robitaille 1997;
Proulx et al. 2006).
In western North America, American Martens ap-
pear to be associated with late successional stands.
Consequently, I hypothesized that American Marten
tracks would not be distributed at random among habi-
tat types, but that they would be more frequent in
mature and old-growth stands. My hypothesis was
declared before examining the track survey data. The
proportion of inventory transects within each habitat
type was used to determine the expected frequency of
track intersects/habitat type (i.e., availability) if tracks
were distributed randomly with respect to habitat types
(Proulx et al. 2006). For each survey, I used a one-
tailed Fisher Probability test (Siegel 1956) to compare
the proportion of American Marten tracks in the habi-
tat growth types, i.e., mature/old vs. immature/young.
Data from the three inventories were pooled to increase
sample size. Chi-square statistics with Yates correc-
tion (Zar 1999) were used to compare observed to
expected frequencies of track intersects among habi-
tat type. If the chi-square analysis suggested an over-
all significant difference between the distribution of
observed and expected frequencies, comparisons of
observed to expected frequencies for each habitat
class were conducted using the G test for correlated
proportions (Sokal and Rohlf 1981). Probability values
<0).05 were considered statistically significant.
Results
Temperatures ranged from -21°C to 0°C and snow
depths exceeded 45 cm during each inventory. Inven-
tories were conducted < 24 h after snowfalls or flurries,
thus assuring that tracks were fresh.
In 1999, 41.1 km of transects were inventoried:
27.8 km (67.6%) in immature/young stands, and 13.3
km (32.4%) in mature/old stands. Eight tracks were
recorded: 3 (37.5%) in young stands, and 5 (62.5%) in
mature-old forests. The observed distribution of tracks
was not significantly (P = 0.31) different from a random
distribution of tracks among habitat types.
In 2002, 36.0 km of transects were inventoried: 16.6
km (46%) in immature/young stands, and 19.4 km
(54%) in mature/old stands. Ten tracks were recorded:
1 (10%) in a young stand, and 9 (90%) in mature/old
forests. The observed distribution of tracks was not sig-
nificantly (P = 0.07) different from a random distribu-
tion of tracks among habitat types.
In 2005, 51.1 km of transects were inventoried:
27.1 km (53%) in immature/young stands, and 24 km
(47%) in mature/old forests. Twenty-six tracks were
recorded: 6 (23%) in young stands, and 20 (77%) in
006 PROULX: WINTER-HABITAT USE BY AMERICAN MARTEN 103
20
18
218
5 14
= ee
E10
eo §8
= 6
=
5 4
z= 2
0
1 ~” aw} n Mo} ~ SS no)
o 3 =| = 3 = = 2
Pee a, PS Ove ni pote. (Qive
ae & D c z as = Eas 1 z “eee.
ie) 5 Ga 3 ov 2 o& 2s 23
& jam > ‘S omm.s a oO = owt =) oO =
tS Sap Aes ee
= 2 ee
Habitat types
IGURE 2. Observed and expected number of American Marten tracks per habitat type in Weyerhaeuser’s Grande Prairie Forest
Management Area.
jature-old forests. The observed distribution of tracks
vas significantly (P = 0.002) different from a random
istribution of tracks among habitat types.
A total of 128.2 km of transects were inventoried
rom 1999 to 2005: 39.1 km (30.5%) in immature/pole
tands, 32.5 km (16.9 % in coniferous, 5.8% in mixed
oniferous-deciduous, and 2.7% in deciduous) in young
tands, and 44.2 km (18.8% in coniferous, 13.5% in
nixed coniferous-deciduous, and 11.9% in deciduous)
n mature/old forests. Forty-four tracks were recorded:
one in immature/pole stands, 10 tracks (22.7%) in
oung coniferous forests, and 34 tracks (77.3%) in
nature/old stands (Figure 2). The observed distribution
f American Marten tracks was significantly different
rom a random distribution of tracks among habitat
ypes (x7 = 39, df: 4, P < 0.001) (Figure 2). Tracks were
ignificantly less frequent than expected in immature/
ole stands (G = 17.2, df: 1, P < 0.001), and more
requent than expected in mature/old mixedwood stands
G = 6.42, df: 1, P < 0.02). The distribution of tracks
uggested that American Martens used young forests,
nd mature/old coniferous and deciduous stands, ac-
ording to their availability.
Jiscussion
In 1999 and 2002, because of small sample sizes, no
ignificant difference was found in the distribution of
\merican Marten tracks among habitat types. Howev-
r, during both years, most tracks were in mature-old
orests. Pooling data increased sample size and statisti-
al power, and showed that American Marten tracks
were definitely more frequent in late-successional
stands. This finding is in agreement with previous stud-
ies conducted in western coniferous forests (Spencer et
al. 1983; Wilbert 1992; Koehler et al. 1990; Therrien
2002; Proulx et al. 2006). American Martens prefer
mature and old-growth forests over young and imma-
ture forests because this is where they maximize for-
aging efficiency and finding rest sites that can protect
them from harsh environmental conditions
Powell et al. 2003).
Although young forests are believed to be subopti-
mal due to a reduction of necessary structure and diver-
sity compared to mature and decadent forests (Thomp-
son and Harestad 1994), in this study, nearly a quarter
of American Marten tracks were located in young
coniferous stands. As late/successional stands are
decreasing in importance in Weyerhaeuser’s FMA due
to timber harvest, and oil and gas exploitation, Ameri-
can Martens may have to make greater use of young
forests. However, previous studies showed that Ameri-
can Martens used young forests in winter where suffi-
cient physical structure (Bowman and Robitaille 1997;
Poole et al. 2004) and prey (Potvin et al. 2000) were
present. More research on the characteristics of young
forests used by American Martens is required to better
understand their importance as winter habitat for this
species in boreal ecosystems.
The fact that no tracks were recorded in immature
and pole stands is in agreement with previous studies
that have generally shown that American Martens
make little or no use of early/successional stages
\C.£.,
104
(Steventon and Major 1982; Thompson 1994; Thomp-
son and Harestad 1994; Chapin et al. 1998; Heinemey-
er 2002; Poole et al. 2004). Previous studies have also
shown that American Martens appear intolerant of habi-
tat fragmentation, and they would not tolerate more
than 30-35% cutovers in their home range (Snyder and
Bissonette 1987; Hargis and Bissonette 1997; Chapin
et al. 1998; Potvin et al. 2000; Poole et al. 2004). In
Weyerhaeuser’s FMA, mature and old forests may be
highly fragmented due to extensive clearcuts, and an
important network of seismic lines and roads. Un-
doubtedly, these habitat modifications would have an
impact on the distribution of American Martens, and
cause shifts in home range boundaries (Poole et al.
2004). Forest development plans taking into consid-
eration the winter habitat requirements of American
Marten should therefore be developed locally accord-
ing to the distribution of late/successional forests, cut
blocks, and access networks (e.g., Proulx 2001*).
Acknowledgments
I thank Luigi Morgantini, Wendy Crosina, Dan Beil-
man, and Darren Lapp from Weyerhaeuser Company
Ltd., and Pauline Feldstein from Alpha Wildlife, for
helping with the project logistics. I also thank Vivian
Banci, Valeria Vergara, and Jamie Farkvam for field
assistance, and two anonymous referees for helpful
comments.
Documents Cited (marked * in text)
Alberta Government. 1999. Alberta natural regions. Alberta
Environment Protection, Edmonton, Alberta.
Proulx, G. 2001. Characteristics and management of Amer-
ican marten habitat at stand and landscape levels. Alpha
Wildlife Research & Management Ltd. report prepared
for British Columbia Ministry of Forests, Prince George
Forest District, Prince George, British Columbia.
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Received 9 June 2005
Accepted 17 October 2005
Notes
Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine
J.C. A. BuRCHSTED! and FRED BURCHSTED2
' Department of Biology, Salem State College, Salem, Massachusetts 01970 USA
* Research Services, Widener Library, Harvard University, Cambridge, Massachusetts 02138 USA
Burchsted, J.C. A., and Fred Burchsted. 2006. Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine. Canadian Field-Naturalist
120(1): 106-108.
The Lady Crab (Ovalipes ocellatus), mainly found south of Cape Cod and in the southern Gulf of St. Lawrence, is reported
from an ocean beach on the north shore of Massachusetts Bay (42°28'60"N, 70°46'20"W) in the Gulf of Maine. All previ-
ously known Gulf of Maine populations north of Cape Cod Bay are estuarine and thought to be relicts of a continuous range
during the Hypsithermal. The population reported here is likely a recent local habitat expansion.
Key Words: Lady Crab, Ovalipes ocellatus, Gulf of Maine, distribution.
The Lady Crab (Ovalipes ocellatus) is a common
member of the sand beach fauna south of Cape Cod.
Like many other members of the Virginian faunal
province (between Cape Cod and Cape Hatteras), it
has a disjunct population in the southern Gulf of St.
Lawrence (Ganong 1890). The Lady Crab is of consid-
erable ecological importance as a consumer of mac-
robenthic invertebrates, and is of biogeographic inter-
est as a member of a large disjunct fauna. However,
the distribution of the Lady Crab in the Gulf of Maine
is unclear from the literature. Here we gather and dis-
cuss available reports of the Lady Crab in the Gulf of
Maine and report an additional site that may represent
a recent habitat expansion. This paper is intended as
a basis for further work on Lady Crab distribution
and for monitoring changes.
Lady Crabs have not generally been reported from
the Gulf of Maine. Field guides (Gosner 1978; Mein-
koth 1981; Miner 1950) and a review of tidal flats
(Whitlatch 1982) give their northern limit as Cape Cod
except for the disjunct Gulf of St. Lawrence popula-
tion. Recent distributional studies of Gulf of Maine
crabs show Lady Crabs on the south shore of Cape Cod
and onto Georges Bank but no further north (Williams
and Wigley 1977; Stehlik et al. 1991). This confirms
the older work of Rathbun (1930). Williams (1984),
and the update by Nizinski (2003), give the range
briefly as Prince Edward Island to Georgia, but in this
they may elide the northern New England gap. Pol-
lock (1998) states the range as Virginian and Acadi-
an, but again the range between the southern Gulf of
St. Lawrence and Cape Cod is unspecified. The Lady
Crab is not listed in Bay of Fundy, Maine, and New
Hampshire checklists (Croker 1972; Brinkhurst et al.
no date; Linkletter et al. 1977; Perkins and Larsen
1975) and distributional studies of the fauna of Maine
sand beaches (Larsen and Doggett 1990) and mud
flats (Larsen and Doggett 1991). Lady Crabs were
not found in intensive local studies of western Cape
Cod Bay (Davis and McGrath 1984) or Ipswich Bay
(Dexter 1944).
Berrick (1986) reports Lady Crabs as common on
Cape Cod Bay sand flats (which commonly reach 20°C
in summer). The few other published records of Lady
Crabs in the Gulf of Maine are from warm, estuarine
habitats. Bousfield and Thomas (1975) identify a group
of three warm water species which tend to co-occur
(Balanus improvisus, Urosalpinx cinera, and Ovalipes
ocellatus) but do not specify locations for the Lady
Crab. Members of this group occurred in 1963 at: Cape
Cod Bay, Duxbury and Cape Ann (Massachusetts);
Cape Neddick and Boothbay Harbor (southern Maine);
and the Bay of Fundy. Bousfield found Lady Crabs at
Sesuit Harbor, East Dennis, Massachusetts, in 1964
(Judith Price, personal communication). Lady Crabs
are reported in Salem Sound, Massachusetts, in 1997
by Chase et al. (2002). Although Dexter (1947) found
no Lady Crabs in an intensive study of a tidal inlet at
Annisquam (Gloucester, Massachusetts) in 1933-1940,
he later (Dexter 1985) found them there during what
he terms a period of warm water from 1952 to 1964.
We have found Lady Crab shells washed up on an
ocean beach adjacent to the Essex and Ipswich river
estuaries (Crane Beach, Ipswich, Massachusetts,
42°41'30"N, 70°46'20"W; 1995 and 1996). Fefer and
Schettig (1980) mention, but do not specify, one or
more southern Maine (south of Saco) locations from
unpublished data by Doggett, Larsen, and Sykes. They
were not found in the 1873 U.S. Fisheries Bureau
investigation of Casco Bay which discovered several
other warm water species (Verrill 1874). There are no
records for the northern Maine cold water region (north
of the Sheepscot River estuary) identified by Bousfield
and Laubitz (1972) and confirmed by Larsen and
106
2006 ss
Doggett (1990). In the northern Gulf of Maine, there
are several reports of Lady Crabs in the Bay of Fundy:
Bousfield and Leim (1960) report Lady Crabs in the
Minas Basin and Channel in 1922 (Bass River and
Scotman Bay, noted also in Rathbun 1930) and again
in 1959 (Bass River, Diligent River, and a cheliped
on beach at Kingsport).
Bousfield and Thomas (1975) state that their Gulf
of Maine records of Lady Crabs are from warm (circa
20°C summer temperatures — similar to their range
south of Cape Cod) brackish water pockets or from
estuaries with two-layer circulations where endemic
warm-water populations could be maintained as des-
cribed by Bousfield (1955) and Sponaugle et al. (2002).
Bousfield and Thomas (1975) suggest these estuarine
populations are relicts from the Hypsithermal warm
era (9500-3000 years Before Present). The many Vir-
ginian invertebrates and marine algae now extending
north to Cape Cod with additional disjunct populations
in the southern Gulf of St. Lawrence had continuous
ranges at that time. The short abstract by Dexter (1985)
provides no details, but his 1952-1964 warm period
may correspond to the 1951-1954 warming reported
by Loder et al. (2001), with the crabs becoming local-
ly extinct during the succeeding cold period (1959-
1967). Dexter’s observations suggest the possibility
that the estuarine distribution is dynamic, populations
becoming extinct and reestablished with changing
conditions.
For the ten years we have been keeping records,
Lady Crabs have been commonly found living on an
ocean beach, called Preston Beach in Marblehead and
Phillips Beach contiguously to the south in Swamp-
scott, on the north shore of Massachusetts Bay
(42°28'60"N, 70°46'20"W). There is no nearby estu-
arine habitat and the beach is exposed to heavy surf
(unlike the Cape Cod Bay flats) [Leo et al. 2003].
Salem Sound is about six coastline miles to the north
around the headlands of Marblehead/Marblehead Neck.
This is the first reported exposed ocean beach location
for Lady Crabs in the Gulf of Maine. Phillips Beach is
in an area of wind-driven summer upwelling that keeps
Summer temperatures around 15-16°C (Geyer et al.
1992). Lady Crabs were commonly found swimming
in shallow water and buried in sand just above low
tide line. The habitat is thus very like that of the Lady
Crab south of Cape Cod. Most of the living animals
were found on the southern portion of the beach. This
area is influenced by groundwater seepage on the high
beach. Old faunal lists for Swampscott and vicinity
yield no records of Lady Crabs: Grabau (1898) for
Nahant, Revere, and Phillips Beach; Pearse (1913) for
Nahant. This suggests our Massachusetts population
may mark a local habitat expansion.
Discovery of other ocean beach populations and
determination of the status of the estuarine populations
would help clarify the distribution and habitat associ-
ations of Lady Crabs in the Gulf of Maine.
NOTES
107
Acknowledgments
We thank E. L. Bousfield for information on his col-
lections and Judith C. Price of the Canadian Museum
of Nature for searching the collections for Lady Crab
records.
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Croker, R. A. 1972. Checklist with habitat notes, of some
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Davis, J. D., and R. A. McGrath. 1984. Some aspects of
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Dexter, R. W. 1947. The marine communities of a tidal inlet
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Fefer, S. I., and P. A. Schettig. 1980. An ecological charac-
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lication FWS/OBS-80/29, Biological Services Program,
US. Fish and Wildlife Service.
Ganong, W. F. 1890. Southern invertebrates on the shores of
Acadia. Transactions of the Royal Society of Canada 8:
167-185.
Geyer, W. R., G. B. Gardner, W. S. Brown, J. Irish, B.
Butman, T. Loder, and R. Signell. 1992. Physical ocean-
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Bays. Massachusetts Bays Program, Boston, Massachu-
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Gosner, K. L. 1978. A field guide to the Atlantic seashore.
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Larsen, P. F., and L. F. Doggett. 1990. Sand beach macro-
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Journal of Coastal Research 6: 913-926.
Larsen, P. F., and L. F. Doggett. 1991. The macroinvertebrate
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taceans of Atlantic coastal and continental shelf waters
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of northeastern North America. Rutgers University Press,
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and Xanthidae. U.S. National Museum Bulletin 152. 609
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2006
NOTES
109
Ring-billed Gull, Larus delawerensis, Food Piracy on Diving Ducks
WILLIAM J. WALLEY
222 Bossons Avenue, Dauphin, Manitoba R7N OR2 Canada
Walley, William J. 2006. Ring-billed Gull, Larus delawerensis, food piracy on diving ducks. Canadian Field-Naturalist 120(1)
109-110.
One of four Ring-billed Gulls observed 26 April 2002 at Dauphin Lake, Manitoba attacked a group of diving ducks and
took abandoned food items.
Key Words: Ring-billed Gull, Larus delawerensis, Common Goldeneye, Bucephala clangula, kleptoparasitism, Manitoba.
Kleptoparasitism or piracy is the forceful taking of
food by one species of bird from another (Terres 1980).
According to Bent (1921) Ring-billed Gulls (Larus
delawerensis) have been observed to hover over feed-
ing ducks, notably Red-breasted Mergansers (Mergus
serrator) darting at them as they re-surfaced apparent-
ly to steal their prey although success is not reported.
Although Ring-billed Gulls do not usually pirate other
gull species (Ryder 1993) they have been observed
pirating food from European Starlings, Sturnus vul-
garis (Burger and Gochfeld 1981). Clapp et al. (1983)
state that a variety of species are pirated by the Ring-
billed Gull. Jarvis and Southern (1976), in their study
of the food habits of this gull in the Great Lakes
region, made no mention of piracy or attempted pira-
cy during the breeding season.
Crescent Cove at the south end of Dauphin Lake
(51°10'N; 99°53'W) in west central Manitoba was
windless and sunny but cold (-11°C) at mid morning
on 26 April 2002. There, 15 m from shore, a stretch of
water 70 m in length and 30 to 50 m in width had
opened up parallel to a sandy beach. Small numbers of
grebes and diving ducks, about 5 and 25 respectively,
swam about on the calm surface, some diving for food.
Ring-billed Gulls were also on the water, stood on the
edge of the ice, or flew around in the area.
Upon my arrival on the nearby ridge above the beach
2 Red-necked (Podiceps grisegena), 3 Horned grebes
(Podiceps auritus), 3 Redheads (Aythya americana),
and 5 Canvasbacks (Aythya valisneria), and unhurriedly
swam away leaving a half dozen Common Goldeneyes
(Bucephala clangula) and 3 Greater Scaup (Aythya
marila). With the ducks diving and returning to the
surface, a Ring-billed Gull, only one of four gulls
believed to actually be an attacker, typically took wing
off the surface from among the waterfowl and flew
low toward a duck up to 15 or 20 m away where it
had just surfaced. Upon reaching the targeted bird,
the gull flew up and hovered just above and behind its
head as described by Bent (1921). The duck plunged
out of sight and the gull, using its bill, snatched the
food item surrendered. One attack was initiated from
as close as 1.5 m.
Often a duck surfaced without prey and with sev-
eral ducks diving it was remarkable how quickly and
efficiently the gull identified a successful duck from
the others and attacked it. Two prey items were iden-
tified: crayfish (Decapoda) and small fish which, when
plucked from the water, were eaten on the spot after the
gull alighted on the surface nearby or flew to a shelf of
ice where the item was tossed down, re-grasped, then
swallowed.
During the forty minute observation, attacks on the
diving ducks were initiated 20 — 25 times. With one
exception — a female scaup — all attacks targeted the
goldeneyes. Other than one instance where four male
goldeneyes directed a threat display involving the
thrusting of their heads low over the water toward a
nearby gull, little aggression was shown by the ducks.
Avoidance of contact and hence injury from the gull
attacks appeared to be paramount for the ducks. Frus-
tration on the part of the ducks was not in evidence,
but determination to capture more prey was.
In April at Dauphin Lake, carrion, notably discard-
ed coarse fish from ice fishing activities, would seem
to be an important source of food for early arriving
carnivorous or omnivorous birds from the south such
as gulls, Bald Eagles (Haliaeetus leucocephalus),
American Crows (Corvus brachyrhyncos) and some
species of hawks, in addition to resident populations
of the Common Raven (Corvus corax). A possible
shortage of fish and/or other sources of food in the
Crescent Cove area of the lake and, perhaps, on the
lake ice generally due to competition, may have been
a factor in the gulls’ attacks on the goldeneyes. Ob-
servations of the lake from shore at Crescent Cove with
the use of a 20 power spotting scope that day revealed
only one such food item, the carcass of a Canada Goose
(Branta canadensis) lying on the ice 250 m out. In
immediate succession at this source of carrion, includ-
ed: a Northern Harrier (Circus cyaneus) with two
crows nearby; an adult Bald Eagle that causing the
harrier to fly off and three immature Bald Eagles that
displaced the adult eagle at the carcass; all to the ex-
clusion of any gulls, suggesting intensive competition
for food. This competition may have resulted in the
unusual method of securing food observed in at least
one Ring-billed Gull.
Acknowledgments
I thank Ken Kingdon of Parks Canada, Riding
Mountain National Park, for literature concerning the
110
Ring-billed Gull — Red-breasted Merganser record.
Appreciation is also extended to A. J. Erskine and other
reviewers whose constructive comments were well
received.
Literature Cited
Bent, A. C. 1921. Life histories of North American gulls
and terns. U.S. National Museum Bulletin 113. Washing-
ton, D.C.
Burger, J., and M. Gochfeld. 1981. Age-related differences
in piracy behavior of four species of gulls, Larus. Behav-
ior 77: 242-267.
Clapp, R. B., D. Morgan-Jacobs, and R. C. Banks. 1983.
Marine birds of the southeastern United States and Gulf
of Mexico. Part 3: Charadriiformes. U.S. Fish and Wildlife
THE CANADIAN FIELD-NATURALIST
Vol. 120
Service, Biological Service, FWS-OBS-83/30. Washing-
ton, D.C.
Jarvis, W. L., and W. E. Southern. 1976. Food habits of
ring-billed gulls breeding in the Great Lakes region. Wil-
son Bulletin 88: 621-631.
Ryder, J. P. 1993. Ring-billed Gull. The Birds of North Ameri-
ca. Number 33. The American Ornithologists’ Union, Wash-
ington, D.C. 22 pages
Terres, J. K. 1980. The Audubon society encyclopedia of
North American birds. Alfred A. Knopf, New York. 1109
pages.
Received 28 February 2005
Accepted 2 January 2006
Probable Black Bear, Ursus americana, Retrieval of an Elk, Cervus
elaphus, Carcass from a Small Lake in Riding Mountain National Park,
Manitoba
WILLIAM J. WALLEY
222 Bossons Avenue, Dauphin, Manitoba R7N OR2 Canada
Walley, William J. 2006. Probable Black Bear, Ursus americana, retrieval of an Elk, Cervus elaphus, carcass from a small
lake in Riding Mountain National Park, Manitoba. Canadian Field-Naturalist 120(1): 110-112.
Strong circumstantial evidence indicated that a sow Black Bear (Ursus americana) retrieved an Elk (Cervus elaphus) carcass
from near the middle of a small lake to feed herself and three cubs.
Key Words: Black Bear, Ursus americana, Elk, Cervus elaphus, carcass, small lake, Riding Mountain National Park.
Accounts of observations of Black Bears (Ursus
americana) retrieving carcasses from bodies of water
are difficult to locate, even those based on circumstan-
tial evidence. Studies of food habits of Black Bears
by Hatler (1972) and Machutchon (1989) cite no such
instances. Herein is provided a description of a series
of observations that lead to the conclusion that an Elk
(Cervus elaphus) carcass was brought to shore from
mid-lake by a female Black Bear.
The observation reported was made at Grayling
Lake (50°47"N, 100°W) in Riding Mountain Nation-
al Park, Manitoba. This is a kettle lake (Lang 1974)
and as such is deep (9 m) for its small surface of
approximately 25 hectares. It is situated well below
surrounding rolling hills in mixed forest about 9 km
north of Clear Lake. With the exception of an 80-m
gap to the NW and the lake outlet through sedge and
grass meadows to the north, tall spruce (Picea) trees
encircle and shelter Grayling Lake.
At 0815 on 2 August 2004 under full sunshine a
large object was seen floating just west of the middle
of the lake by the author and two companions from
north-south orientated Highway 10 that passes just to
the west of the lake (Figure 1).
Identification of the brown-gray object was not
made even with 8 x 40 binoculars. At 12:15 that day
the lake was re-visited. Though air movement was
Table 1. Wind direction and speed in km/hr in Riding Moun-
tain National Park 2 August 2004*
Time At Wasagaming At old Kippen’s mill site
08:00 NW 6 SW 3.6
09:00 NW 4 NW 63
10:00 NW 9 NW 8.1
11:00 NW 7 WNW 7.8
12:00 NW 11 NW 8.6
*Recorded by Environment Canada (personal communica-
tion, K. Kingdon, 2005)
almost negligible, the object had been moved close to
the east shore, where two Coyotes (Canis latrans) fed
on it. Some 50 m to the north, a Black Bear was un-
hurriedly walking north away from the carcass and the
Coyotes. It then disappeared into heavy coniferous
forest. Shortly after, the Coyotes moved off and a bear
appeared at a small partial clearing above the shore
and the carcass. In subsequent days the bear, from this
point, repeatedly chased off Coyotes but tolerated the
intensive food collecting activity of a pair of Common
Ravens (Corvus corax). On 3 August a sow led her
three cubs to the carcass, where they fed.
In the next few days the remains of the carcass were
identified as cow Elk (Cervus elaphus) by Ken Kingdon
2006
AN
WIND OUT OF NW
I
NOTES 11]
observation
points
COW ELK
CARCASS
STRANGE "OBJECT" | 1215
A 3
815 oO 50° '
dock /
HWY 10
GRAYLING LAKE
/_—_______
200 m i
3 =
Expected location of
carcass if blown by
wind
FIGURE |. Elk carcass locations relative to wind direction, Grayling Lake, 2 August 2004.
of Parks Canada (personal communication) and inde-
pendently by the author.
It was hypothesized that the bear had retrieved the
carcass from the lake and had to swim out at least
200 m to get it. Veteran Parks Canada Warden Pat
Rousseau (personal communication), an expert on
Black Bears, concurred. In his experience he had not
actually seen bears drag Elk carcasses ashore but, from
an aircraft, had seen a Black Bear swimming halfway
out toward an Elk carcass 40 m from the shore of a lake.
On another occasion he had seen Black Bear and Gray
Wolf (Canis lupus) tracks associated with an Elk car-
cass on the shore of a remote lake in the park. In the
mid-1990s a party of hikers reported an animal with
a black head behind a carcass in deep water of Bead
Lake Four less than 3 km NW of Grayling Lake. The
only scavenger other than a bear that it might have been
mistaken for would be the much rarer Wolverine (Gulo
gulo) (Pat Rousseau, personal communication).
Environment Canada wind data in the critical time
period that day (08:00—12:00) showed only very light
breezes primarily from the NW which almost certainly
eliminated the wind as being a factor in the carcass
being blown to the east shore (Table 1) (Figure 1). On
112
a similarly calm day, 13 August, blocks of wood were
placed in the middle of the lake to see if water currents
would carry them to the east shore.
No definitive proof was obtained that the bear swam
out and hauled the carcass a minimum of 200 m to
shore. However, the negative results of the water cur-
rents experiment coupled with the data on wind direc-
tion and speed that day provided a strong suggestion
that this is what happened. Still, it is possible that anoth-
er species of scavenger (e.g., Wolverine) brought the
carcass to shore, and then abandoned it. Further, al-
though none was seen before 08:15 or after 12:15 there
is a possibility that, while the carcass was unobserved
by the author between these times that day, one or more
people kayaking or canoeing on the lake took the car-
cass to the east shore. Interviews with such recreation-
ists at the lake on subsequent days revealed no knowl-
edge of the carcass being moved. Hence, both of these
scenarios seem very unlikely.
In retrospect, the bloated condition of the carcass
near the middle of Grayling Lake on 2 August doubt-
less produced a powerful olfactory stimulus for the
bear and her cubs.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Acknowledgments
I thank Parks Canada Warden Patrick Rousseau
for his accounts of Black Bears and Elk and Warden
Service Communications Officer of Riding Mountain
National Park, Ken Kingdon, for providing informa-
tion on wind speed and direction. Appreciation is
also extended to the reviewers of the manuscript who
made constructive comments and gave suggestions.
Documents Cited (marked * in text)
Parks Canada. 2004. Riding Mountain National Park esti-
mated elk and moose populations from aerial survey 1976-
2004.
Literature Cited
Hatler, D. F. 1972. Food habits of black bears in interior
Alaska. Canadian Field-Naturalist 86: 17-31.
Lang, A. H. 1974. A guide to the geology of Riding Mountain
National Park. The Geological Survey of Canada. Miscel-
laneous Report 20, Department of Energy, Mines and
Resources, Ottawa. 68 pages.
Machutchon, A. G. 1989. Spring and summer food habits
of black bears in the Pelly River Valley, Yukon. Northwest
Science 63(3): 116-118.
Received 28 February 2005
Accepted 2 January 2006
Book Reviews
ZOOLOGY
rhe Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi
By Terry Stevenson and John Fanshawe. 2006. Princeton
University Press: 41 William Street, Princeton, New Jer-
sey, 08540 USA. 640 pages U.S. $35.00 Paper.
Birds of East Africa: Kenya, Tanzania, Uganda,
Rwanda, and Burundi is one of a number of bird guide
books on this region. Indeed, this publication is a paper-
back version of the award-winning 2001 hardback edi-
‘ion. It has not been updated.
The first of these books I bought was Birds of East
Africa by Williams and Arlott, published in 1963. At
he time I was very pleased at the style and quality of
his work. Some 665 species were illustrated. In 1999
his was superseded by Birds of Kenya and Northern
Tanzania by Zimmerman, Turner and Pearson. This
900k was a giant leap forward with 1084 species illus-
rated (even though this only covered 40% of the area
of East Africa.) The illustrations increased by 419 spe-
>ies despite the smaller area covered. Also the quality
of the artwork and printing was far better and was a
sloser match to European and North American guides.
Now we have the new Birds of East Africa with Norm
Arlott again providing much of the artwork. This vol-
ume illustrates 1388 species and is another, although
smaller leap forward.
Birds of East Africa covers the resident, migrant, and
vagrant bird species of Kenya, Tanzania, Uganda and
the two smaller countries of Rwanda and Burundi (the
area of these two together is less than 3% of the region,
but adds a surprising number of species to the list). The
text is high quality and faces the colour plate page (a
more convenient format). The species accounts de-
scribe identification, status, range, habits, and voice.
The colour distribution maps are small, but clear.
The question now arises, which is the better book
for the visitor going on typical trips to the major parks
in Kenya and northern Tanzania? Birds of Kenya and
Northern Tanzania is slightly smaller (around 5%)
and covers less territory. While this loses about 20% of
the species, most of these are specialist species con-
fined to Uganda, Rwanda and Burundi (about 225
species that are wholly or primarily confined to those
countries ). The remainder are species predominantly
from southern Tanzania.
To my eye the artwork in The Birds of East Africa
is noticeably better both for the colourful species and
the LBJ’s (“Little Brown Jobs”: warblers and Cisti-
colas), but this not a huge difference. I checked sev-
eral species from my slide collection and found Birds
of East Africa: Kenya, Tanzania, Uganda, Rwanda,
Burundi was generally more accurate. The user will
find it very difficult to separate Long-billed from Tawny
pipit using Birds of Kenya and Northern Tanzania, but
will find only a small improvement with Birds of East
Africa as these birds are difficult to separate anyway.
There is not much to choose between the texts (al-
though it is more extensive in Birds of Kenya and
Northern Tanzania) and the coverage of endemics is
very similar. It is easier to have the text, illustration and
map side by side (Birds of East Africa ). The biggest
difference lies in the range maps. The Birds of East
Africa shows only a single range designation, covering
winter, summer and migratory ranges. In comparison,
Birds of Kenya and Northern Tanzania shows summer
and winter ranges separately. Just as important, Birds
of Kenya and Northern Tanzania more clearly defines
where the birds are located. For example waterside
birds like bitterns and herons are show in localities
where water occurs. Similarly plains species are shown
in the plains. The Birds of East Africa maps blanket the
entire area that water birds can occur giving the impres-
sion they can be anywhere, even in dry savannah.
Birds of Kenya and Northern Tanzania includes a
map of region inside its front cover that shows the key
locations mentioned in the text. Although The Birds
of East Africa has maps that show features (like for-
est cover), they do not depict the location of national
parks, cities, etc. Neither book gives alternative names,
which is a bit irritating when making comparisons with
park checklists, older books, trip reports etc. (e.g.
Schoenicola brevirostris is called the Fan-tailed Grass-
bird or Broad-tailed Warbler and not to be confused
with Cisticola juncidis called the Fan- tailed Warbler
but now known as the Zitting Cisticola.)
If you are planning a trip that includes Kenya, Tan-
zania, Uganda, Rwanda or Burundi, The Birds of East
Africa is the more valuable book to take with you.
When I go to Kenya and Tanzania later this year, the
extra information given by the range maps would prove
beneficial (These more detailed range maps will be
particularly useful for the first time visitor.), but | am
taking Birds of East Africa: Kenya, Tanzania, Uganda,
Rwanda, Burundi because of ease of use and higher
accuracy of the illustrations.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
HS
114
The Birding Sites of Nova Scotia
By B. Maybank. 2005. Nimbus Publishing Ltd., 3731
Mackintosh St., Box 9166, Halifax, Nova Scotia B3K
5MB8 Canada. 654 pages, $35.
There is a path that runs from a popular beach in
southern Nova Scotia. If you know exactly where, you
can leave this path and cross several hundred metres
to the start of a more secret path. This second path
runs through a spruce grove along a cliff. If you know
exactly which spruce to slip behind, you will find a
hidden rope that will allow you to drop several metres
down a cliff face. You will be in a very secluded cove
with a golden beach and blue water, sheltered from any
cold wind. This secret spot must be the only place in
Nova Scotia that Blake Maybank has not visited! His
new book is an incredibly detailed set of directions to
(almost) every tiny nook of this beautiful province.
The main purpose of this book is to give sufficient
directions for the visitor or native Nova Scotian to find
the numerous coves and special places that dot the
province. This is not as easy as it sounds as some of the
roads and tracks are obscure and hard to distinguish.
For example, I know native-born people who did not
know of the track to Cherry Hill Beach, even though
they had driven past it many times. This is because it
is tucked between houses and looks like a driveway.
Very sensibly Maybank has given the GPS coordinates
of this location and others where someone might have
trouble.
At each locality the author describes the habitat and
comments on the birds typically to be found there.
He also includes cultural highlights such as churches,
lighthouses, museums and other tourist attractions.
Although the title suggests this book is only about birds,
Maybank includes similar information on mammals,
butterflies, reptiles and amphibians. Where noticeable
he comments on the wildflowers but this is not a focus.
Each section has icons representing the types of birds,
plants or mammals found and other key features like
washrooms to be found on the site.
Giving directions can, of necessity, be dry reading
even (or especially?) when they are given in great detail.
The Complete Fauna of Iran
By Eskandar Firouz. 2005. I. B. Tauris. London, New York.
Xiv + 322 pages. US $90.
This book covers the vertebrate fauna of Iran, mam-
mals, birds, reptiles, amphibians and freshwater fish-
es. It is a revised, English language version of a 2000
publication by the same author in Farsi (reviewed by
Coad and Keivany 2002). Those fluent in Farsi might
prefer that version at the bargain price of $5.63!
Iran is a country that often appears in the political
news for its great economic and strategic geographi-
cal importance. This is mirrored in its vertebrate
THE CANADIAN FIELD-NATURALIST
Vol. 120
The writer’s style is much more upbeat and entertain-
ing in the other sections. I was amused to read (Meat
Cove) is “beyond the radar ... its delights are shared
by a privileged minority” (Meat Cove, despite its name,
is indeed a beautiful place of seabirds, orchids and gor-
geous scenery; naturalists should certainly go there).
Maybank does not give a detailed account of all the
birds likely to be seen at a given locality. He uses gener-
ic terms such as gulls, shorebirds and warblers. Howey-
er he has set up a website where you can access bird
lists and information on the key localities. I assume
he plans to keep this site current, so it is much more
useful then putting a list in the book (which would
soon be dated).
There are a few items that I think need changing.
Page 243B has lost part of its text, an obvious printing
problem. The author states “There are no deer ticks ...
hence no Lyme disease.” It is not this simple. Dog ticks
(Dermacentor variabilis) also carry the Lyme disease
bacteria, Borrelia burgdorferi (technically they are
“infected”, but maybe not “infective”). Whether or not
the D. variabilis tick transmits Lyme disease is not yet
resolved, suggesting caution. I was dismayed to see he
did not mention Grassy Island (and hence Roseate
Terns) in his account of Pearl Island (perhaps the last
pair has been extirpated). The book has no index, mean-
ing you will need a map to get you oriented. (Find the
county you want and use the index to get you to the cor-
rect chapter.)
Any traveler who has gone to the expense of travel-
ing to Nova Scotia needs this book; it will be invalu-
able. They will get to more places and see things they
would have missed. Nova Scotians will also benefit. I
doubt if anyone has traveled so widely and could pro-
vide such a precise record as Maybank. I will be using
this book for my upcoming trip to Ile Madame in Cape
Breton. As my last trip was in winter, I look forward
to more informed and interesting birding this time.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
fauna, which includes such famous economic species
as the caviar-bearing sturgeons and many diverse
species of interest to naturalists close to its location
at the intersection of the Palaearctic, Ethiopian and
Oriental realms. :
The Introductory sections of the book are particu-
larly valuable for readers not familiar with this part of
the world, and encompass 48 pages. A transliteration
system is explained for the Farsi common names of
organisms (a local language name list is always use-
ful as scientific names may be more of a barrier than a
2006
help outside academia), the distribution of Lranian prov-
inces is mapped, major freshwater drainage basins
outlined, and topography summarized in colour. A sum-
mary of zoological work in Iran follows with accounts
of the transition between hunting and falconry to
conservation and protection. [ran now has an impres-
sive list of 16 National Parks, 13 National Nature Mon-
uments, 33 Wildlife Refuges and 92 Protected Areas.
Summaries of physical geography, climate, vegeta-
tion, zoogeography, threatened species and some fos-
sils follow, illustrated by colour habitat photographs.
Appendix I lists all the parks, their area, year estab-
lished and a grid reference for a map in the Introduc-
tory section. Appendix II describes the Pardisan proj-
ect (from the Old Persian for paradise), a 270-ha park
in Tehran intended to replicate Iranian environments
as an innovative educational project and centre for re-
search in all fields related to the natural world. Con-
ceived by Firouz, Pardisan was meant to be an academy
of sciences, a zoo, herbarium, aquarium and museum.
Hopefully it will come to fruition. The Bibliography
encompasses 7 pages and is a general introduction to
the literature on Iran. This literature is surprisingly
diverse and could not be covered in this book — that on
freshwater fishes, for example, numbers over 2900 items
(see www.briancoad.com). The book ends with indices
to Latin and English names of vertebrates and a gen-
eral index.
The mammals occupy 60 pages, birds 88, reptiles
54, amphibians 10 and freshwater fishes 34. The num-
ber of mammal species referred to is 168, birds 514,
reptiles 199, amphibians 20, and freshwater fishes 173,
for a total of 1074. Because of space limitation, not all
species are described. Introductory sections give an
overview of work on each group and an update from
the 2000 edition in Farsi; e.g., six species of mammals
have been added to the fauna. Iranian scientists became
very active in the 1990s and subsequently, as evidenced
by their publications in such journals as Zoology in
the Middle East. Comments are made on conservation
and the destruction of habitats that threaten so many
species, here as in the rest of the world.
As an example, the first family dealt with, the hedge-
hogs, number four species which are listed by English
name, Farsi name in English script, Latin name and a
numbered listing of the provinces in which they occur.
This requires the reader to find the map of provinces
at the front of the book, and these maps are very small
(15 by 10 mm), and would require a knowledge of Iran-
ian geography to interpret easily. Any illustrated species
is indicated by a diamond next to the English name.
A general review of hedgehog biology precedes this
list. No mention is made of the absence of hedges in
Iran (where hedgehogs are found in burrows, bushes
and rock crevices), and the disconcerting habit shown
by some species of hissing loudly when disturbed is
also omitted, an attribute not noted in the more sedate
English hedgehogs.
BooK REVIEWS
Mammals of note, and evidence of the land bridge
that Iran forms between Europe, Asia and Africa, in
clude lions and tigers (both now, sadly, extinct), Egypt-
ian fruit bats, wolves, red foxes and jackals, cheetahs,
leopards and lynx, hyenas and badgers, deer and gaz-
elles, and jerboas and hares. The plan to re-introduce
lions from India to southern Iran is noted but not
whether this actually took place. Rumours in the 1970s
had them introduced, but quickly wiped out by local
villagers who were not ardent conservationists when
faced with a potential predator on their sheep and
goats, not to mention sleepy goatherds and shepherds.
The illustrations are a mix of photographs and colour
drawings.
The birds are illustrated mainly from colour draw-
ings provided by Birdlife International and are a good
summary of that fauna. Necessarily, this section in par-
ticular must be quite abbreviated in its text, given the
bird diversity, but books devoted to birds are widely
available for the fauna of the Middle East. Status sym-
bols are given for each species rather than provincial
distributions as with mammals; e.g., R = common res-
ident, r = scarce resident, again a system that requires
reference back to the page listing 16 possibilities.
These could certainly have been included with each
species’ name in an abbreviate form.
The reptiles are well-illustrated with colour photo-
graphs. The fauna is varied and interesting but not evi-
dent without effort in field situations. I seldom saw
snakes and lizards while living in Iran although small
geckos could be found on house walls (and had a fero-
cious bite). One of the largest lizards readily encoun-
tered is the Bengal Monitor which reaches 2.75 m.
Chasing one of these rapid lizards across the desert
could be quite tiring if they did not try to hide behind
a bush much too small to conceal their length. Marsh
crocodiles live in the southeast of Iran, much decimat-
ed by drought in the late 1990s. They are here reported
as not attacking people, a fact I wish I had known in the
1970s when I might have sampled their home streams
for fish less nervously. Five species of marine turtles
are known from the Persian Gulf, all badly in need of
protection.
The amphibian fauna is small in species although
toads and frogs are common in and near water. Tree
frogs are found on banks of streams, trees being rare
in much of desert Iran.
Extensive critical comment on the freshwater fishes
would be inappropriate given the input by this review-
er. Only freshwater fishes are dealt with, although this
includes Caspian Sea fishes where salinities are about
one-third of sea water. The diverse fauna of the Per-
sian Gulf and Sea of Oman are not covered—a recent
summary work is Assadi and Dehqani Posterudi (1997).
One loss of information is that space does not always
allow for a full explication of evocative Farsi names.
The enormous sturgeon, Huso huso, reaching 1.5 ton-
nes and 6 m in length, is known as fil mahi which
116 THE CANADIAN FIELD-NATURALIST
translates appropriately as elephant fish while the pike,
Esox lucius is ordak mahi, duck fish from its duck-
like snout.
The author is eminently qualified to write on this
topic having been the Director of the Department of
the Environment and vice-president of IUCN. He is
regarded as the father of Iran’s environmental move-
ment. The Farsi version of this book won the Iranian
Publisher’s Association top award.
This book is highly recommended as a well-writ-
ten overview of a fascinating fauna.
Fishes of the World
By Joseph S. Nelson. 2006. John Wiley & Sons, Hoboken,
New Jersey. Fourth Edition. xvii + 601 pages. U.S. $125.
This book covers all the 515 families of fishes in the
world in 62 orders and 5 extant classes. Fossil families
are more briefly treated. Each class, subclass, order,
suborder, family and subfamily receives a brief descrip-
tion. In this way the reader is treated to a tour of the
world’s fishes through their higher classification and
relationships. This being the fourth edition, 30 years
after the first, proves the need, and success, of such a
work. Scientists and students look to this work for an
overview of fish classification and, with an estimated
28 400 species at the end of 2006 compared to about
26 730 tetrapods (mammals, birds, reptiles and amphib-
ians), the fishes need this structure. At the individual
species level, constantly changing as new ones are
described or old ones revised, the online “Catalog of
Fishes” from the California Academy of Sciences is
the update source (www.calacademy.org/research/ich
thyology/catalog/index.htm).
A website, http://members.shaw.ca/fishesoftheworld/
index.htm contains errata, a list of new families post-
1958, student exercises and links to relevant websites.
Changes in the numbers of taxa between the four edi-
tions of the book are listed in the table below.
Variation in the number of families is mostly due
to lumping and splitting as Nelson points out at the
website mentioned above. Discovery of new families
is a rarity in fishes. However, the estimated number of
known fish species for the end of 2006 cited above is
almost 10 000 more than in 1976, showing that this
field of endeavour has been most active.
The text has continued to be expanded over the ear-
liest editions, with families once without a description
now provided with one. Much necessarily remains the
same from the last edition as relationships, anatomy and
species content remain unchanged in certain families.
Vol. 120
Literature Cited
Assadi, H., and R. Dehqani Posterudi. 1997. Atlas-e Mahian-e
Khalij-e Fars 0 Dary-ye Oman/Atlas of the Persian Gulf & the
Sea of Oman Fishes. Iranian Fisheries Research and Training
Organization, Tehran. 10 + 226 + 23 pages. In Farsi and English.
Coad, Brian W., and Yazdan Keivany. 2002. Review of “A Guide
to the Fauna of Iran.” E. Firouz. 2000. Iran University Press (Uni-
versity Publication Centre), Tehran. ISBN: 964-01-0956-8. vi +
491 pages. 45,000 Rls ($5.63) (hardbound)”. Copeia 2002(4):
1164-1166.
BRIAN W. COAD
Canadian Museum of Nature, Ottawa, Ontario K1P 6P4
Canada
Not all genera are listed within each family, although
a couple listing would be a great convenience, and not
all families have a line drawing (about 74% do). Some
families have undergone significant changes, with new
information on species numbers, ecology and fossil
members added, including the new coelacanth found
in a fish market. Several groups (e.g., Characiformes,
Osphronemidae) have been rearranged based on new
systematic studies, and Nelson makes a concerted ef-
fort to follow a cladistic framework. However, as Nel-
son notes in his preface, it is naive to accept the latest
proposals as being the best. Nelson has used his own
considerable judgment in determining which groups
are sufficiently systematically stable to include. This
has also resulted in some families (e.g. Cichlidae) los-
ing some structure, as new studies have shown past
taxonomic groupings to be non-monophyletic.
The literature on fish classification is extensive,
sometimes arcane in its arguments, and scattered in
journals worldwide. Having an author mull over this lit-
erature and provide an interpretation and digest for con-
sideration is a great service to students and to those pro-
fessionals who need to dip into these waters at intervals.
In this new edition, Nelson has included many more
references, providing the most recent literature. The
Bibliography is comprehensive and covers 54 pages.
Some authors may be disappointed that not all their
papers are cited, but Nelson has generally included
good summary papers that will provide, within their
own references, many titles of more specific articles.
The Index at 63 pages is also most useful in locating
particular taxa but certain genera will not be found (as
noted above) and the reader will have to resort to Cat-
alog of Fishes. Additionally, the previous edition used
the names of orders as the running header throughout
the text. The use of the more inclusive class level as
the running header in the new edition is less helpful
in flipping through to find the desired pages.
Marine and
Year Species Freshwater Diadromous* Families Orders Classes
1976 18,818 6851 11,967 450 46 +
1984 21,450 8411 IBS 445 50 4
1994 24,618 9966 14,652 482 57 5
2005 ZU UT 11,952 16,025 515 62 5
2006
The single disappointment of the new edition is the
loss of the chart of categories and relationships of fish-
es from the inside front cover. It is now buried on (un-
numbered) pages xvi-xvii, between the Acknowledg-
ments and the Introduction, making it much harder to
find. Owners of the book may want to place a perma-
nent bookmark in the chart for quick reference. The
new chart, while showing better resolution of groups
in this edition, has also lost all the names between
Class and Order. While some of these names were for
non-cladistic grades, their presence on the previous
edition’s chart was of great benefit, particularly for stu-
dents, to determine exactly what is a “teleost” or “acti-
nopterygian”, or any of the other names that are com-
monly used by ichthyologists.
At the end of the list of errata on the website given
above, and on page 9 of the new edition, Nelson draws
Insects: Their Natural History and Diversity:
North America
By Stephen A. Marshall. 2006. Firefly Books Ltd., 66 Leek
Crescent Richmond Hill, Ontario L4B 1H1 Canada. 720
pages. Can $95. Hardcover.
It has always been said that you can’t judge a book
by its cover, but after reading /nsects: Their Natural
History and Diversity lve learned that this well-worn
axiom isn’t always true. When this book first crossed
my desk, to say that I was instantly enamoured would
be an understatement. It was so beautiful, the cover
adorned with a stunning jewel-toned dogbane beetle
(Chrysochus auratus). 1 almost didn’t want to crack the
binding. However my curiosity finally got the better
of me and I’m glad it did because once I started read-
ing I couldn’t put the book down.
Visually stunning, with over 4000 color photographs
of insects in their natural habitats, nsects: Their Nat-
ural History and Diversity has the look and feel of a
glossy coffee table book while still being full of accu-
rate, well researched information.
As its title implies Insects: Their Natural History
and Diversity focuses on the diversity and natural his-
tory of common families of northeastern North Amer-
ican insects. The book opens with a brief synopsis of
basic insect anatomy and morphology. This is fol-
lowed by chapters covering the diversity of all insect
orders, including all the major families, along with
two chapters on non-insect arthropods and methods
for observing, collecting and photographing insects.
The book’s last 50 pages are dedicated to illustrated
keys to order and family as well as a key to the most
commonly encountered insect larvae. These keys are
designed to facilitate ease of use and therefore empha-
size morphological characters visible to the naked eye
or easily seen with a hand lens. Also peppered through-
out the book are helpful suggestions on where to look
for and find various insect orders/families. For exam-
ple “Depending on your inclination and the weather,
BooK REVIEWS
117
attention to the need for ichthyologists, and the work
that still remains for future generations. He urges us to
support the replacement of retiring ichthyologists to
continue the work. The Introduction of Fishes of the
World provides a sampling of all the rich areas of ich-
thyology to be studied and highlights the importance
of fishes to all of us. Perhaps if the Introduction were
required reading for everyone, Nelson’s enthusiasm
for fishes would be passed on to all, and the worth of
ichthyological research would be clearly visible to
governments and society. This book continues to be a
seminal work, finding an essential place in libraries and
on the bookshelves of anyone interested in fishes.
BRIAN W. CoaD and ALISON M. MURRAY
Canadian Museum of Nature, Ottawa, Ontario KIP 6P4
Canada
With a photographic guide to insects of eastern
a good place to start looking for assassin bugs would
be in your kitchen light fixture. Unless you are much
more fastidious than most, the odds are that among the
crispy critters accumulated there you will find a large
black assassin bug called the Masked Bed Bug Hunter
(Reduvius personatus).”
Considering that Insects: Their Natural History and
Diversity is priced so as not to be cost prohibitive and
is clearly written in plain language makes it highly
accessible to a broad audience including naturalists,
amateur entomologists as well as seasoned profession-
als. The author has also included a dollop of humour
and wit throughout the text. For example this passage
describing the appearance of springtails: “Some are
covered with scales, like those of a butterfly, many are
brilliantly colored and all are morphologically bizarre,
starting from the long, forked tail used to make Her-
culean leaps, and ending with the deeply pocketed
mouth that makes springtails look like they have lost
their dentures and then sucked on a bunch of lemons.”
This book would make a great textbook for a natu-
ral history or general entomology course. Especially
when you consider that the impetus for this book cen-
ters on materials originally gathered in support of the
author’s third-year course “The Natural History of
Insects” at the University of Guelph. With its depth of
scope and true-to-life color photographs /nsects: Their
Natural History and Diversity would be indispensable
in the field; however, due to its size it would be a bit
unruly to have to lug around.
I thoroughly enjoyed this book and would recom-
mend it whole heartedly to anyone who has an interest
in entomology, natural history or a simple curiosity
about the six-legged world that surrounds us.
GINA PENNY
Department of Biology, St. Francis Xavier University, Anti-
gonish, Nova Scotia B2G 2W5 Canada
118
THE CANADIAN FIELD-NATURALIST
Vol. 120
Secret Weapons — Defenses of Insects, Spiders, Scorpions, and Other Many-Legged Creatures
By Thomas Eisner, Maria Eisner, Melody Siegler. Belknap
Press of Harvard University Press, Cambridge. 2005. 372
pages.
This book is dominantly about chemical defences of
terrestrial arthropods, though other types of defences
(camouflage, catapulting, sticky traps) are also men-
tioned. Chapters are short, and typically feature the
defences of one organism, or at most a small group of
related organisms. One or more colour photos, in them-
selves almost worth the cost of the book, enhance both
the aesthetics of the book and the ability to drive home
each chapter’s message. Chemical formulae are illus-
trated in those chapters focusing on chemical defences.
As the subtitle alludes, an array of terrestrial arthro-
pods is dealt with, but by far, the majority of this book
is about insect defences. However, scorpions, centi-
pedes and other non-insects together account for about
15% of the book’s material. Examples include the
well-known spray of the Bombardier Beetles, the less
well-known vomit attacks of noctuid moth caterpillars
and immobilizing wax of oleander aphids.
This book will appeal to a wide array of readers,
the caveat being that those with no (or little) training
in chemistry (like me) will miss out on some of the
nuances of the chemicals themselves; this did not give
Rodents and Lagomorphs of British Columbia
By David W. Nagorsen. 2005. Royal British Columbia Muse-
um, Victoria, British Columbia.
I read this book both with a general interest in mam-
mals overall, and, as a raptor biologist, a desire to more
completely understand the creatures which are the prey
of my birds. I was not disappointed on either account.
From a simply presentational point of view, the art-
work sticks out as being very aesthetically pleasing.
Each species is illustrated with a pencil drawing of the
complete animal — not in a standard three-quarter pose
or profile, but in varying postures that show snapshots
of each animal’s character. Additionally, the skull of
each species is presented in lateral, dorsal and palatal
views — perhaps this is not useful to the general reader,
but I certainly benefited from them; the illustrators
(Michael Hames, Donald Gunn and Bill Adams) must
be commended.
The book has three keys — one each for whole ani-
mals, skulls and chipmunk genital bones. Anyone who
has used an unillustrated key understands the frustra-
tion of trying to interpret the author’s meaning (what is
meant by “relatively large?’”’). A fully illustrated key
has diagrams for each couplet — the keys in this book
are not fully illustrated, but there are plenty of dia-
grams that make the process of keying out an organ-
ism rather straightforward.
An introductory chapter describes the biogeography
of British Columbia, along with conservation strate-
gies, winter survival and several other topics which link
these species together. Maps and good photographs are
featured here.
rise to any problems at all for my enjoyment of this
book. The other caveat being that those not liking those
long, tongue-tripping scientific names of organisms will
have to gloss over them periodically as they read; even
where common names exist for the organism of the
chapter’s focus, the scientific name is typically used
throughout the text. Grade-school children will be able
to use this book though clearly not as completely as
older folk; entomology instructors could easily incor-
porate this material into lectures. Truly, this book can
meet the needs of many.
There is an abundant amount of fascinating mate-
rial that simply makes this well-written book a joy to
read; I have been equally satisfied after having read
Waldbauer’s books, including What Good are Bugs?
(also from Harvard Press). Unlike the latter book, the
chapter bibliographies of Secret Weapons are replete
with papers of the book’s authors, particularly the
Eisners. Their work in chemical ecology, translated by
them from their papers to this book, is a grand exam-
ple of making science more consumable by the non-
specialist.
RANDY LAUFF
St. Francis Xavier University, Antigonish, Nova Scotia
B2G 2W5 Canada
The species accounts dominate this book. Each runs
about six pages and describes physical aspects of the
species, its natural history, geography, and conserva-
tion status. An occurrence map accompanies each ac-
count; I believe British Columbia residents and non-
residents alike would have benefited from place names
being included on the maps — especially those which
are only mentioned once or twice in the whole book.
Similarly, even the introductory maps do not include
all the major sites mentioned in the book.
There are a small number of mistakes, some edito-
rial (e.g. “supraorbital bone” is used instead of “supra-
orbital process”), some factual (e.g. “[The muskrat has]
rooted cheek teeth.”). In the introduction to the lago-
morphs, I think it would have been appropriate to
differentiate rabbits from hares.
I take particular exception to the oft-used expression
that some introduced animal (in this book, the East-
ern Cottontail) “seems to be filling an ecological niche.”
This is at best an empty comment (what introduced
animal would not have a different ecology than the
natives) and at worst could be interpreted as an excuse
to tolerate and perhaps enhance introductions.
Those few shortcomings aside, this is‘a very good
book which serves its purpose admirably. I will refer
to it frequently.
RANDY LAUFF
St. Francis Xavier University, Antigonish, Nova Scotia
B2G 2W5 Canada
2006
Book REVIEWS
119
Tanagers, Cardinals, and Finches of the United States and Canada
By D. Beadle and J. Rising. 2006. Princeton University Press,
41 William Street, Princeton, New Jersey 08540 USA
196 pages, U.S.$29.95 Paper.
The authors use 200 selected photographs to com-
plement text covering the biology, identification, molts,
song and distribution of the tanagers, cardinals, and
finches found north of Mexico. In all, forty-six species
are covered. Of these, one is an introduced species and
31 are native residents of some part of North America.
The remaining 14 are vagrants with five as Alaska spe-
cials and seven creeping north over the southern border.
I have just spent a couple of hours resolving an iden-
tification issue using about 20 photographs off the Inter-
net (out of the 200 I found there) of a single species.
Most of those photos were of limited use. Normally I
find such photographs less useful than artwork because
they are so influenced by light, angle and the condition
of the specimen. However, my jaundiced eye is very
impressed with the photographs in this book. I do be-
lieve that I would be able to separate female House and
Purple finches using the photographs and text given
in this book. Indeed I used this book to confirm that I
had correctly identified a Hoary Redpoll I saw recently
in the Arctic. The typical resident species are represent-
ed by several photographs taken at different angles thus
giving the reader a better understanding of the bird.
The 31 residents each have a range map giving sum-
mer, winter, and permanent ranges. These are about
5.5 cm square and very readable and useful. There is a
written description of range too, which adds more
detail. I had no difficulty deciding I was at the northern
edge of my Hoary Redpoll’s range. The text is clear and
provides great detail without wasted words.
The vagrants are similarly treated, but do not have
a range map. Under distribution their normal range
(outside of North America) is described, followed by
a summary of their occurrence in North America, with
additional details, such as precise locations and dates,
where warranted.
This is a very useful supplementary guide (to the
classical all-species field guides) for providing addi-
tional information on this group of birds. It is conve-
niently sized to carry in the field and will be especially
useful in sorting out look-alikes such as Hoary and
Common redpolls. It provides carefully organized and
well-presented information to those who wish to learn
more about these birds than is given in the typical field
guide. This is the type of book that makes a most wel-
come present!
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada
Wheatears of Palaearctic: Ecology, Behaviour and Evolution of the Genus Oenanthe
By E. N. Panov. Pensoft, Sofia, Moscow. 439 pages, U.S.
$167.50
Wheatears are the quintessential Afro-Eurasian bird.
Found anywhere there is open ground, the more deso-
late the better, wheatears make ideal subjects for orni-
thological study; indeed, according to the author they
are among the most studied groups of passerines. They
are easy to see and follow, they exhibit interesting be-
haviours, and, as a group, pose many riddles. How can
up to four species with seemingly identical foraging
behaviours co-exist in a relatively inhospitable habi-
tat? Why in some species is there high sexual dimor-
phism and in others virtually none? What light can this
group renowned for intra-species variation shed on
the nature of polymorphism and its evolutionary ori-
gins? What are the advantages of a contrasty black and
white plumage? How did one species of this desert
loving family break through the oh so inhospitable for
wheatears boreal forest to successfully colonize much
of the Arctic including not once but twice in North
America?
These and other questions are copiously answered
in this book by E. N. Panov, a Russian scientist with
a passion for this intriguing group. The book is divid-
ed into two parts, the first dealing with general fea-
tures of wheatear biology, the latter providing detailed
monographs on individual species or superspecies
found in Eurasia. Between them, these sections cover
every conceivable angle on wheatears, from the geo-
graphic distribution and origins of the group in its
entirety through habitats, movements, social behav-
iour, reproductive behaviour to predators and parasites.
Each monograph follows a template covering system-
atics, distribution, behaviour, nesting, and movements.
The book is extensively illustrated with black and
white sketches illustrating everything from behaviours,
nest sites to plumage variations and is amply provided
with sonograms, family trees, graphs and maps illus-
trating key points. There is also a section of 40 colour
plates showcasing the array of arid habitats so beloved
of the genus, portraits of selected species, and at least
one picture of one being devoured by a snake. The book
closes with a 14-page bibliography and an index help-
fully divided into a general subject index as well as an
index of animal names.
There are about twenty species of wheatear, with
the exact number hard to pinpoint because of blurry
lines with other turdidae such as saxicola (“stone-
chats’’) and uncertainty about the taxonomy of several
closely related taxa. Taxonomic and genetic evidence
suggests that the genus had its origins in Africa, where
it may have emerged from an ancestor shared with
redstarts. The large amount of variation and frequent
contact zones within the genus suggest that there is
an ongoing evolutionary process. The English name,
which comes from the old English for white rump,
120
refers to the bold white “T” pattern which is character-
istic of many members of the genus. Its Russian name,
Kamenka, refers to the stony habitat they so love; their
ability to cope with extreme conditions is witnessed
by the fact for some 160 000 square kilometres in the
Sahara the White-crowned Black Wheatear is the only
breeding passerine, and the fact that the Northern
Wheatear has successfully colonized arctic tundra.
Given the lack of culinary choice their dietary habitats
are pretty well catch as catch can; recorded food items
include scorpions and bumblebees.
The Northern Wheatear, the only member of the
genus in North America, is remarkable. Each year birds
from Alaska, in the words of the author, “stubbornly
make for their remote African homeland”, an epic voy-
age that would exhaust an Arctic Tern. Birds nesting
in Eastern Canada head in the opposite direction, and
are the only Canadian passerine that winters in Africa.
Being found in places where lots of ornithologists live;
the Northern Wheatear is one of the better known
members of the group. From these studies, we know
that they maintain a frenetic lifestyle during the breed-
ing season that can, in the far north, involve a 24/7 rou-
tine of zealously patrolling its borders while attending
the needs of two females in widely separated territo-
ries. According to Panov, Wheatears are highly aggres-
sive with a “low threshold for territorial aggression”.
BOTANY
Giant Trees of Western America and the World
By Al Carder. 2006. Harbour Publishing Co., Madeira Park,
British Columbia. 138 pages. $18.05.
Al Carder has been researching giant trees for a lot
of his long career, collecting stories of large trees in the
Pacific forest of North America, his home and main
study area. This book is a sequel to his earlier work
trying to record the dimensions of many of the world’s
largest trees now including more facts and more tree
species. The book is ordered by continents and tree
families to give the reader perspective of the variety
of trees which have achieved huge dimensions, but
many of which have disappeared in the past century.
A short anecdotal essay at the beginning puts Carder’s
agenda forward quickly. He has always been awestruck
by the beauty and perspective of being among the lar-
gest trees and wants to preserve the memory of those
trees for future ages. Sadly the largest specimens of
any tree species are gone from our world due to com-
mercial logging in the last two centuries and when sin-
gle giant trees survived the logging, these succumbed
to windstorms, having been left without the surround-
ing forests which protected and supported their growth
over the centuries. In North America, Australia and
New Zealand, the story of logging is the history of
large trees and our most impressive forests, a history
THE CANADIAN FIELD-NATURALIST
Vol. 120
To communicate with one another they have three dif-
ferent song groups and a remarkably varied repertoire
of physical signals involving different postures and
movements. The author also usefully informs us that
they have an unpleasant taste, evidenced by experi-
ments in which hornets were offered wheatear “meat”.
It is disappointing that the author, demonstrably a
competent illustrator and photographer, did not choose
to include a series of systemic plates illustrating the
various taxa and their variations. Another disappoint-
ment is that the author focused the species accounts on
Eurasian taxa; one suspects that this reflects the author’s
research interests but also the fact that African taxa
are relatively less well known. In the review copy the
hatching on many of the maps was blurry, making
them hard to figure out. Nonetheless, there are few
errors in the book, although the fact that the title con-
tains a grammatical error did not inspire much up front
confidence.
This book is a scientific monograph primarily aimed
at ornithologists. It poses, and proposes answers to a
number of interesting ecological and behavioural ques-
tions and will be of great interest to anyone intrigued
by evolutionary biology.
MARK GAWN
Permanent Mission of Canada, 5 avenue de |’ Ariana, Gene-
va, Switzerland.
which has more to do with destruction of forests than
the story of humans interacting with their environment.
The book is short with only one hundred and thirty
pages devoted to describing the trees. Each tree des-
cription has an accompanying page with a diagram of
the tree at its largest reported size and shape, each
species being unique in shape. Fifty of those pages are
devoted to the trees of North America with the other
pages listing and describing the outstanding trees of
other continents. North America is obviously Carder’s
first priority but his love of trees encompasses the
entire world.
Many trees in England and Europe have been avail-
able for study to written history over the centuries
and Carder has researched these written accounts to
give us historical anecdotes from their long life. A Tule
tree in Mexico with an opening in the trunk large
enough to accommodate twelve horsemen when report-
ed 375 years ago, and the Sweet Chestnut in Sicily
whose spread of branches sheltered the Queen of Ara-
gon and her escort of 100 horsemen over 600 years
ago are two of the anecdotes which describe trees still
standing. Whether the tree has records of growing to
over 400 feet as the Douglas Fir or whether its claim
is to have lived over 5000 years as the English Yew,
the grandeur of the trees is given to us by Carder to
2006
be considered, enjoyed and lamented as phenomena
which have existed but sometimes passed us by.
I enjoyed reading about present and past giant trees
and was entertained different times looking for inter-
esting facts and diagrams. It is not a book to be read at
one sitting, even though short, because the tree species
stand alone and reading several is like reading a list
which can go on too long. A short reading of one or
two tree descriptions at each visit is sufficient with
another few minutes reading the next descriptions at
ENVIRONMENT
BOoK REVIEWS 121
another time. Soon the book becomes a reference for
tree species and musings of how much we have missed
and what we might visit. | have shown it to my students
and they spent time with it as a picture and description
book to browse and marvel. Al Carder has given us a
little gem of knowledge and history for entertainment
based on good science and research.
Jim O’ NEILL
28718 Five Mile Road, Livonia, Michigan, 48154 USA
Fire and Avian Ecology in North America — Studies in Avian Biology Number 30
Edited by Victoria A. Saab and Hugh D.W. Powell. 2005.
Cooper Ornithological Society. 193 pages.
This technical work is a collection of 11 papers, ten
of which address the role and effect of fire in one or
more ecosystems (e.g., Boreal Forest, Oak Woodlands,
Interior Chaparral); the first chapter is a cogent sum-
mary of the ten others.
The chapters more or less follow a logical pattern
in their layout. Each paper shows a map of the distri-
bution of the habitat(s) discussed, and normally one
or two other figures. A table of pertinent literature is
presented in each chapter, which also includes respons-
es to fire of a number of bird species. The papers des-
cribe historic fire regimes, including the use of fire by
aboriginals, as well as the effect of fire suppression on
birds (and by default, the plants making up the habi-
tat for them); sections on conservation strategies, in-
cluding the use of prescribed burns and withholding
of fire suppression end each chapter.
The only irritant to me was lumping all the refer-
ences at the end of the book; normally, each paper
should be concluded with its own collection of refer-
ences. Should someone want to photocopy one chapter,
they would have to photocopy the entire compilation
of references, instead of just the pertinent ones. None-
the-less, it is a solid work.
RANDY LAUFF
St. Francis Xavier University, Antigonish, Nova Scotia B2G
2W5 Canada
Fisheries Assessment and Management in Data-Limited Situations
Edited by G. H. Kruse, V. F. Gallucci, D. E. Hay, R. I. Perry,
R. M. Peterman, T. C. Shirley, P. D. Spencer, B. Wilson,
and, D. Woodby. 2005. Alaska Sea Grant, Fairbanks, Alas-
ka. Publication Number: AK-SG-05-02. 958 pages. Price:
$50.00 US, ISBN:156612-093-4.
Maintaining sustainable fisheries resources requires
the combined efforts of scientists, fisheries managers
and policy makers. Balancing fisheries growth and
conservation is a difficult task, particularly in situations
Where available information is limited. Data-limited
fisheries are often plagued by a lack of long-term data
on the basic biology, ecology and productivity of the
species that are relevant to these resources. A number
of successful fisheries have been developed by com-
bining a scientific background with robust policy man-
agement to successfully develop sustainable fisheries,
even in situations where data is limited. However,
newly developing and small-scale fisheries often oper-
ate without sufficient data to develop production mod-
els, assessment techniques and sustainable manage-
ment strategies.
To address these challenges, a symposium entitled
“Assessment and Management of New and Developed
Fisheries in Data-limited Situations” was held in
Anchorage, Alaska, in 2003. The purpose of this sym-
posium was to share knowledge, research and manage-
ment strategies for newly developing and small-scale
fisheries by bringing together fisheries scientists, man-
agers and policy makers from all over the world. The
result is a proceedings book titled “Fisheries Assess-
ment and Management in Data-limited Situations.”
Specifically, this book is a collection of forty-six peer-
reviewed research papers that provide case studies and
management considerations for fisheries with limited
data. The book is divided into seven sections: (1) case
studies of fishery failures and successes; (2) indica-
tors of stock health and productivity from limited sam-
pling programs; (3) involvement of fishermen and use
of local knowledge; (4) multi-species and ecosystem
indicators and models; (5) precautionary management
approaches; (6) stock assessment models; and (7) stock
assessment surveys and applications. The first two sec-
tions focus on science-based assessments of fishery case
studies while the final five sections identify and dis-
cuss management and assessment strategies. Through-
out many of the case studies in this book, several key
management themes emerge that are highly relevant
to fisheries managers in data-limited situations.
Section one provides insight into the failures and
successes of the world’s fisheries. Several case studies
on a variety of species are described, and management
implications are discussed. A key paper in this sec-
122
tion provides a general overview of the world status
of data-limited fisheries using landings statistics with
a Bayesian approach (Vasconcellos and Cochrane).
Each of the subsequent case studies describes tech-
niques for managing both struggling and successful
fisheries. These case studies focus on the development
of new or developed fisheries that have limited data
(Bechtol and Trowbridge; Bureau and Hand; Therri-
ault et al.) and the recovery and management of strug-
gling resources (Bargmann et al.; Kruse et al.; Stephen-
son and Jackson). Despite their geographically and
species-specific nature, the general knowledge con-
tributed by these case studies may be highly relevant
to similar fisheries in data-limited situations in other
parts of the world.
The second section focuses on developing indicators
of stock health and productivity from limited data sets.
Each paper describes a case study where early warning
signs can be detected in order to predict the state of
fisheries resources. The indicators that are described
include biological indicators (LeBlanc et al.; Tribuzio
et al.), performance indicators and stock dynamics
(Hay et al.; Orensanz; Raid et al.; Ziegler et al.) and
management indicators (Howland and Tallman; Scan-
dol; Weyl et al.). A key paper from this section is a
simulated study of Yellowfin Bream (Acanthopagrus
australis) where empirical stock-status indicators were
assessed using quality control methods (Scandol).
The third section focuses on the integration of local
fishermen and traditional ecological knowledge in fish-
eries management. A key paper in this section by Freire
describes the use of data provided by anglers in the
assessment of Brazil’s growing recreational fisheries
industry. This paper represents the first data of their
kind from South America and illustrates the potential
value of recreational anglers as a key data source for
improving fisheries management in data-limited situa-
tions. The fourth section contains papers that are geared
towards the assessment of the effects of fisheries at the
ecosystem level by using ecosystem indicators (Stob-
berup et al.) or models (Ainsworth and Pitcher; Che-
ung and Pitcher; Morato and Pitcher). This section
contains an interesting study by Cheung and Pitcher
that outlines possible strategies for evaluating fish-
eries management policies that aim to conserve bio-
diversity. The fifth section contains a number of stud-
ies that take a precautionary approach to data-limited
fisheries management. The majority of the studies in
this section focus on fisheries uncertainty (A’mar and
Punt; Campbell and Dowling; Punt and Methot; Welch;
Zheng). One key paper involves an evaluation of a
multi-stage approach to data-limited fisheries manage-
ment. Perry et al. assesses the four-stage approach to
new and developing fisheries (i.e., data collection, de-
sign of key studies, implementation of selected strate-
gies and establishment of the new fishery) designed by
Fisheries and Oceans Canada. One of the most chal-
lenging factors in this staged framework, as with the
management of many of the world’s fisheries, is the
THE CANADIAN FIELD-NATURALIST
Vol. 120
contradiction between the demands of access to fish-
eries and the conservation of these resources. This
paper addresses some key points that are relevant to
balancing these challenges.
The sixth section contains several papers that address
the use of models in stock assessment. Stock assess-
ment bias and sensitivity are addressed by two key
papers in this section (Ernst and Valero; Mesnil). Also,
Catch-Survey Analysis (CSA) is discussed by two pa-
pers to estimate stock abundance (Collie et al.) and to
address concerns over sensitivity and bias (Mesnil).
Similarly, the seventh section addresses stock assess-
ment methods and surveys. In this section, several key
methods are described, including population density
assessments using scuba (Byerly and Bechtol) and the
applications of acoustical estimations (Kaljuste et al.).
This section also addresses sampling designs for esti-
mating bycatch (Menon et al.), methods for account-
ing for climate variability in models (Shotwell) and
assessments to address concerns over recreational over-
harvesting (Therriault and Hay). A case study discussing
the modeling of climate variability in relation to fore-
casting Pacific salmon spawner-recruit dynamics (Shot-
well et al.) is particularly relevant to data-limited fish-
eries that require the input of variable environmental
data into forecast models.
This book contains an excellent assemblage of well-
written, highly relevant research papers on data-limit-
ed fisheries. Several management themes emerge from
this book, including (1) the development of key rapid
assessment tools, (2) the integration of traditional knowl-
edge from fishermen and local ecological knowledge,
(3) the adoption of a precautionary approach to fish-
eries management, (4) the evaluation of sensitivity and
bias in stock assessment models, and (5) the develop-
ment of robust stock assessment methods. These key
strategies recur throughout many of these studies and
are relevant to fisheries worldwide. Further, several
novel strategies for managing data-limited fisheries are
discussed, including the development of key modeling
and assessment tools. This book offers one of the first
attempts to assemble key literature on data-limited fish-
eries into a single volume. This is an invaluable resource
for the management of new or developing fisheries.
The intention of this collection is to provide scien-
tific background to new and developing fisheries. Each
research paper attempts to suggest applications of
their case study to other similar data-limited fisheries
resources. However, the seven sections contained in the
book are organized with little or no preface or inter-
pretation of this research. Throughout the book, the
links between the results of each case study to a broad-
er application are often tenuous. For most proceedings
books in this style, it is common to omit interpretations
by the authors or editors in each section. However, the
nature of this research demands a synopsis in each sec-
tion to highlight the applications and extensions of this
research. Managers require the interpretation of fish-
eries experts who are experienced with data-limited
2006
situations to identify how the methods described in this
book can be applied to their own data-limited situations.
In summary, the management themes that emerge from
this collection are highly relevant to all fisheries man-
agers, policy makers and scientists working with data-
limited fisheries. However, this book would be consid-
erably more useful to these stakeholders if the context
Lapland — A Natural History
By D. Redcliff. 2006. Yale University Press, P.O. Box 209040,
New Haven, Connecticut 06520-9040. 352 pages, U.S.
$60. Cloth.
Lapland: a Natural History by the late Derek Red-
cliff, is a remarkable account of the birds and higher
plants of northern Scandinavia. This book begins
with a general description in land use and the geolog-
ical background terms followed by a skillful history
of the earlier naturalist visitors. The book is mainly
given over to a detailed evaluation of the various habi-
tats found in this area. These include Boreal Forests,
Forest Peatlands, Lakes and Rivers, Coastlands, Tun-
dra and Man-influenced habitats. Each of these chap-
ters is divided into vegetation and flora and birds. The
birds section is subdivided into key groups such as
wildfowl, waders (shorebirds), birds of prey, songbirds
and birds that are of particular importance in that habi-
tat (e.g., wildfowl or grouse).
The author writes in a style reminiscent of a profes-
sor giving a lecture to keen students and as a result the
book is easy and pleasant to read. The accounts are
packed with information drawn from many research
sources, backed up by the author’s own observations.
The text does contain references, but not in sufficient
number to impede the flow of ideas. Every few pages
there are four half-page photos. These depict habitats,
plants, birds and bird nests. Particularly impressive are
the almost two dozen photos of shorebird nests (have
you ever tried to find a shorebird nest?). These photos,
most by the author, are of very fine quality. Interspersed
with the text are some engaging black-and-white
illustrations by Mike Unwin, reminiscent of those in
the journal, British Birds.
I particularly enjoyed the way Redcliff linked the
occurrence of a species with the other components of
the habitat. He does a wonderful job of showing the
inter-dependance of each constituent of a habitat and
the consequences of disruptions to the balance, both
natural and man-induced.
Even though I found this a treasure trove of infor-
mation (and wisdom) I did have several problems. I
hoped, while doing this review, I could compare the
plant life of Northern Canada, Svalbard and Lapland.
The author’s inconsistent use of scientific names be-
came a source of frustration for making this compari-
son. The worst case was his discussions of Dwarf Aza-
lea. As he does not include the scientific name it is hard
to be sure which plant he is referring to. Lapland Rose-
Book REVIEWS 123
of each section was clearly described in a one to two
page preface and synopsis summary that outlines how
this knowledge could be applied in practice.
M. R. DONALDSON
Ottawa-Carleton Institute of Biology, Carleton University,
Ottawa, Ontario KIS 5B6 Canada
bay (Canada) or Lapland Rhododendron (Europe) is
Rhododendron lapponica, but the author quotes this
plant separately. Indeed, he includes a beautiful photo
of this lovely plant. He also includes a photo of “Dwarf
Azalea’, but it is a very un-Rhododendron-like plant
(more like a saxifrage or even Moss Campion). After
much searching and consultation with Bill Cody (author
of several books etc. on northern plants) I was still un-
clear until I found a reference to Loiseleuria procum-
bens as Trailing Azalea (and several other English
names) while identifying plants from Cambridge Bay,
Nunavut. Similarly I was never sure if the Finnmark
Primrose was the same as Primula erects or nutans or
if it was a new species or subspecies.
Also the book only covers the higher plants well.
There is relatively scant mention of the lichens and
other more primitive plants. I am sure that there must
be well over 500 species of lichens in this area, yet the
book refers to only a handful. Other groups are even
more poorly represented. On the animal side, birds are
well covered. Mammals get an introduction and other
groups (reptiles, amphibians, butterflies and dragon-
flies) get an honourable mention. Other wildlife such
as beetles and bugs do not get any attention.
I also had problems with many of Redcliff’s terms.
To start with, he does not really define what he means
by Lapland, Fennoscandia, Fennoscandinavia and
Scandinavia. As he uses terms like “Southern Lapland”,
I felt it important to understand the region’s precise
location. His map showing the location is vague so I
thought I would check further. Lapland is somewhat
well defined (northern Norway, Sweden, Finland and
Russia — or the home of the Sami). Actually I have
found Finnmark, Nordland, Nord-Trondelag and Troms
[counties of Norway], Lapland [a province of Finland],
Jamtlands Lan, Norrbottens Lan and Vasterbottens Lan
[counties of Sweden], and Murmansk [an oblast — or
“province” — of Russia] form Lapland. Knowing at
least where the county of Finnmark is located is most
useful when reading the text. However, the other terms
vary from source to source. They could include some
or all of Norway, Sweden, Denmark, Finland, Iceland
and the Faeroes. Similarly I had trouble with the terms
flark, fell, strang, patterned fen and mire. Eventually
I did get them sorted out but a glossary would have
been so helpful.
I have several problems with the index. First, it does
not reference any scientific names, only English ones.
None of the photos are listed. Some of the entries are
124
listed forward (e.g., Lapland Rhododendron and Alpine
Lady Fern) and some entries are listed backward (e.g.,
Primrose, Finnmark and Birch, Downy) making it dif-
ficult to decide which letter to use.
The author includes range maps from other sources.
The maps for plants use the scientific names (If you
did not know Silene acaulis is a Moss Campion [as
opposed to the synonyms: Silene bryoides, Silene exs-
capa] you would be a little lost. The lovely photo of
Parmelia centrifuga is not identified as a concentric
ring lichen). The map captions for birds use English
names. The maps are not in the index.
The Three-toed Woodpecker was split in 2003 into
the American Three-toed (Picoides dorsalis) and Eura-
sian Three-toed woodpeckers (Picoides tridactylus)
using differences in mitochondrial DNA sequences.
The author has not mentioned this or any other recent
splits. Perhaps this is not as significant a issue in a book
focused on ecology rather than identification.
Finally, the book contains two photos of people; one
is the author, although the caption omits this detail and
the other is Jeanette. The book is dedicated to this lady,
but her presence is never explained (she is Mrs. Red-
cliff).
THE CANADIAN FIELD-NATURALIST
Vol. 120
This is a great book for anyone interested in the far
north or who wants to understand the complexities of
life. It is wonderful and fascinating reading if you are
passionate about natural history, but is of more limited
value for research.
Derek Redcliffe was the first to establish the role
of agricultural pesticides with the decline of birds of
prey. He was chief scientist of the Nature Conservan-
cy Council in the UK where he acted as a scientist
and lobbyist for nature conservation. He fearlessly
attacked and changed agricultural and forestry inter-
ests for the damage to wildlife value caused by their
policies and practices in place at the time. Not an
easy or popular task in Maggie Thatcher’s world.
Despite his legendary contribution to UK conserva-
tion he was denied any honour by the state — in a
country that knights pop stars. Redcliff was known
an extraordinary naturalist, good company and friend.
His last book 1s a fitting legacy.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
Fish, Fur and Feathers: Fish and Wildlife Conservation in Alberta, 1905-2005
Fish and Wildlife Historical Society. Federation of Alberta
Naturalists (FAN), Edmonton. $39.50, 418 pages.
This sumptuous, well-illustrated book is an eclectic
mix of articles, reminiscences and interviews. The main
text is on white paper, and the other contributions on
yellowish paper. Only rarely does this confuse the read-
er but the white paper portion of chapter four ends in
the middle of an incomplete sentence.
As expected with any product of a committee, some
chapters are stronger than others. We often know who
wrote a particular piece, but sometimes we don’t. No-
where, not even at the end of the two-page Editor’s
Note, are we told the name of the chief editor (Bill
Wishart, as I learned from a special enquiry to Edmon-
ton). At the end of each chapter are references and a
useful chronological list of the main happenings.
In Alberta, as elsewhere, the responsibility for fish
and wildlife management grew from a handful of vol-
untary game guardians to a diverse group of biologists
and enforcement officers within professional, academ-
ic, and conservation communities. Yet for the better
part of a century the emphasis was on fishing and hunt-
ing. In 1967 the first female biologist (now the presi-
dent of the Canadian Medical Association) was hired,
and only in 1987 was the first non-game biologist hired.
Now there are 14 recovery teams developing and imple-
menting plans for threatened and endangered species.
Some accounts are superb. The account of the change
in upland game numbers associated with harvesting
practices, from threshing machines, grain separators,
and wheat stooks to combine harvesters, is among the
best I have encountered.
Alberta has had some unique successes. The Pere-
grine Falcon restoration program, under the direction
of Richard Fyfe, was immensely important. One week
in the 1960s, biologists taught a three- or four-evening
short course in wildlife biology and management, spe-
cially designed for Members of the Legislative Assem-
bly; more than half of them attended! A M.Sc. thesis
on the fish and wildlife damage inherent in the planned
Highwood Dam helped to influence the shelving of
that project.
Biographical information is generally excellent, giv-
ing credit to world leaders in biology, especially Wil-
liam Rowan. Even though this book is co-sponsored
by the FAN, amateurs are somewhat slighted, except
for Frank Farley, Cam and Joy Finlay, Elsie McAllis-
ter, Myrna Pearman, Dewey Soper, and Kerry Wood.
Edgar Jones is barely mentioned and Otto H6hn,
Robert Lister and Al Oeming are omitted. Informa-
tion resulting from bird banding is provided only as
regards waterfowl. The Beaverhills Bird Observatory is
not indexed. The outstanding studies of Golden Eagle
migration west of Calgary by Peter Sherrington, the
landmark studies of Ferruginous Hawk biology by
Josef Schmutz, and the northern owl banding by the
Edmonton group are not mentioned.
The compilers are frank about some of the bureau-
cratic bungles (“bureaucracies are often much too short-
sighted;’ page 207) and failures, such as legislation
2006
drafted, but never proclaimed. The “biggest mistakes
of the century” are identified as the transfer of diseased
bison to Wood Buffalo Park, game farming, predator
control attempts, “too many fishermen,’ and loss of
habitat.
There are a few errors of fact and interpretation.
The two-page account of the Prairie Farm Rehabilita-
tion Administration is unusually misleading when it
speaks of 87 community pastures by 1999, omitting the
fact that only one (in three blocks within the Suffield
Military Range) of these PFRA pastures is in Alberta.
The American White Pelican was threatened, never
endangered (page 341). Gary Pelchat was born in Pon-
teix, not the never-existent Phoenix, Saskatchewan.
David Munro is misspelled “Munroe”. The first two
Franklin Arctic expeditions were hoping to find an arc-
tic sea route, not an overland route to Asia. The index
is inadequate, failing to index names of any organism
The Gulf of Alaska: Biology and Oceanography
By P. Mundy 2005. Alaska Sea Grant College Program, Uni-
versity of Alaska-Fairbanks, Edited by Exxon Valdez Oil
Spill Trustee Council.
Alaska is perceived “...as a place of wilderness,
beauty and a special way of life”. This book describes
the Gulf of Alaska (GOA), a region with many world
records. Besides covering a huge area, having the third
largest permanent ice field in the world and being a
central place, crossroads, for almost all of the Pacific
salmon, GOA also is among the most productive oceans
in the world.
The introduction is a chapter on general oceanog-
raphy for the lay public; the classic concept of the
‘Conveyor Belt’ is presented. Due to its ecological im-
portance, I like the human footprint statement such
as “Human uses of the GOA are extensive” and“... it
is rare to walk the intertidal zone anywhere in GOA
without seeing evidence of human activity”. The
“...marine pollution and floating refuse from as far
away as Asia, or originating from deliberate deep-ocean
dumping or accidents at sea, can be swept north and
westward around the shelf edge in the GOA. Trash from
the international fishing industry operating 200 miles
offshore is commonly found on beaches”. Further, one
will read that GOA has arguably the best known rocky
tidal zone on earth, and this small tidal zone is cru-
cial for the ocean. This book admits climate change
but leaves it somewhat undefined whether it is a man-
made issue, or not.
The book contributors emphasize that to fully under-
stand the ecology of GOA first one needs to make an
effort to identify of all relevant species, e.g., plankton.
Shrimp, once among the dominant benthic epifauna
in Lower Cook Inlet, Kodiak and along the Alaskan
Peninsula, declined after maximum harvests in 1977,
and today its fishery is virtually nonexistent. Red King
Crab crashed in 1980, and Tanner Crab fisheries are
Book REVIEWS
125
(e.g., pronghorn, ring-necked pheasant, and liver fluke).
For consistency, the book title might better have been
Fin, Fur and Feathers.
With pressure from an increasing urban human
population, highways, and mechanization of farming,
mining, oil extraction, forestry, dams and irrigation, it
will be difficult to maintain biodiversity in the decades
to come. Some hope may be gained from looking at
both the successes and failures of the past.
A large book, it is too heavy for bedside reading.
Caveats aside, this is a wonderful collection of infor-
mation that should be in all major Canadian libraries,
and read by everyone interested in the history of fish
and wildlife conservation and management.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N OJ8
Canada
down as well. The clam fishery in Cordova never recov-
ered from damages brought by an earthquake. Despite
its fishery of global importance, the two short para-
graphs for the Pacific Cod section are rather uninfor-
mative. The fisheries for Walleye Pollock have been
down since 1982. The Herring loss story for Seldovia
Bay and Lower Cook Inlet is reported on as well.
Salmon from Japan, Russia, British Columbia, all of
Alaska and the Pacific North West spend some part
of their life cycle in the GOA. The canned salmon
industry collapsed in 1959, the year when Alaska be-
came a U.S. state.
Despite all of this overwhelming evidences of limit-
ed ocean supplies, some book contributors still claim in
their chapters the traditional views of “unlimited re-
sources”. Other sections blame the “stocks” and anglers
for their impacts (rather than their managers with a
governmental mandate for safeguarding the public
resource). Fortunately, sea floor dredging effects are
explained, since fishery impacts to the soft-bottom
benthic community are a possible driver of commu-
nity change. It is suggested in the book text that this
affects Steller Sea Lions as well.
Throughout, the book shows that for non-commer-
cial species we lack management information. These
species compose the bulk of the biomass. By-catch
of the juvenile cohort occurs in fisheries like Herring,
Pollock, and Salmon. However, for many of the non-
commercial species the magnitude of this phenomenon
is not really known, nor how much it affects these
stocks.
Even apart from all of these severe marine issues
mentioned in the book, the described terrestrial situa-
tion is equally puzzling: The book states that in the
GOA, 24% of the water bodies listed are on the state’s
list of polluted sites attributed to some aspect of log-
ging.
126
The book makes clear that Alaskan “Subsistence
Harvest” is a much-debated issue, a major political
player in Alaska and beyond, and thus, well supported
by the government. However, only 2200 people in the
Exxon Valdez Oil Spill (EVOS) region actually still
practice this form of life style. This must be among
the most subsidized lifestyles in the world. The gov-
ernment uses undercover agents to report on how much
natural resources are taken.
How much do Alaskans value their nature? This
book provides an answer: Anchorage operates under
a Clean Water Act Section waiver, only completing pri-
mary treatment of the sewage. The classic cocktail of
PCB. DDE, DDT and Dioxin gets named several times,
but there are many more pollutants in the Arctic. The
complex toxic pollution of the Copper River is men-
tioned though. I think that most readers will get excited
that the Alaskan government does not choose to moni-
tor for environmental pollutants.
A seabird chapter deals with nesting species, not
with its pelagic, non-breeding populations. Cliff nesting
murtres and Black Kittiwakes are the most well studied:
underground and nocturnal species are hardly known.
The statements “There is no evidence that seabirds in
the GOA have been directly affected by commercial
fisheries” and “ ...do not compete with commercial
fisheries for biomass” give the wrong impression.
The book publishers need to be chastised for this
misleading statement as it sets the stage for inappro-
priate environmental management policies in favour
of economic activities.
Seabird starvations in GOA are mentioned several
times: e.g., related to climatic ENSO (El Ninio/South-
ern Oscillation) events. There is nice documentation of
increased gull predation e.g., Glaucous-winged Gulls
on Black-legged Kittiwakes, due to increased gull
populations. The given Marbled Murrelet estimate of
200 000 nesting birds is likely low when compared
with the authoritative literature. The provided nesting
seabird population table is misleading since non-breed-
ers occur in the region and thus, populations are higher.
The Minerals Management Services, EVOS, NMFS
(National Marine Fisheries Service), and the Alyeska
Consortium did extensive benthos survey and moni-
toring efforts in GOA in the 1970s. The reader is left
wondering where these raw data are and this book does
not help us to learn where to find them. One simply has
to trust the “experts” blindly. This must set the stage
for conflict.
The contaminant issue I find not well handled in
this book either, considering that EVOS was a GOA
event of global attention and recognition. Since this
book is an EVOS publication one has to ask, what
did we really learn from EVOS? There is no chapter
devoted to toxicology or sections contributed by a
professional Alaskan toxicologist. One will have a hard
time to find much oil spill literature. At least we know,
in 2000, 604 spills were reported in the EVOS GOA
THE CANADIAN FIELD-NATURALIST
Vol. 120
region alone, proving we do have an oil and hazardous
substances problem.
The book notes that there is a conflict between in-
creasing development and loss of environmental qual-
ity in the GOA region (as found anywhere else in the
world). Rats, ground squirrels and voles introduced
in most of the Aleutians between 1700 and the 1900s
make for a great example. Other examples include fox
farms starting 1894 onwards, followed by canneries,
the Klondike Goldrush of 1897, a copper mine operated
from 1905 and the Northwestern railway built in 1911.
Marine mammals are elaborated on in greater detail:
seal decline in 1973-2000 and the sea lion decline of
approximately 83% (maybe >90%). The federal fund-
ing for investigating this sea lion decline is between
40 to 60 Million $$; a record in federal spending on
one species. Was it a success?
The infamous Killer Whale story and their blame for
the seal decline gets featured as well: e.g.. in Prince
William Sound. Killer Whales consume 400 seals each
year (somebody has to eat the individuals of poor
health: sharks were suggested as well). Killer Whales
are believed to have suffered through EVOS.
Ecologists might be happy to see that the textbook
example of how Sea Otters and their over-harvest links
with Sea Urchin and kelp populations is mentioned.
The book reports that lead in Sea Otters can now be
traced directly to industrial sources, namely from North
America and Asia, which makes for a great example
for the need of international management in GOA
(none of this is demanded in the book text though).
Overall. it remains unclear to me what the exact
focus of this book is, and why it was published. Readers
must see it as a major oversight that no professional
contacts and addresses are provided for the 13 contrib-
utors or even editors. I found the content editing some-
times poor and not well structured. This becomes obvi-
ous in the multiple mentioning of the study area and its
repetitive definitions and maps throughout chapters.
Some of the chapter sub-headings are virtually similar
to the chapter heading, other sub-headings and themes
re-occur in other chapters. The quality and amount of
citations varies throughout chapters. I like the abbre-
viation and acronym section because it also includes
websites and their URLs. Some of the figures are stun-
ningly made, but unfortunately in black and white.
It becomes quickly obvious that in concert with the
many other signs and evidences there is something
wrong with the natural state of GOA, and subsequently
how it is managed. This book is a great example how
unpublished and non-peer reviewed grey literature
can rather add to confusion and is not achieving its
goals for sustainable management.
FALK HUETTMANN
Biology and Wildlife Department, Institute of Arctic Biolo-
gy. 419 Irving I, University of Alaska, Fairbanks, Alaska
99775-7000 USA
2006
MISCELLANEOUS
Fossil Legends of the First Americans
By Adrienne Mayor. 2005 Princeton University Press, Prince-
ton, New Jersey, USA. 446 pages. illustrated.
The lure of fossils is not a recent venture. For cen-
turies, by accident or pursuit, “sports of nature” as they
once were called have fascinated our cultures. Yet,
while fossils rose from “sports” to conceptions of the
organic past, this was essentially a western perspective,
shaped by culture and scientific advancements. It is a
little surprise, however, that the North American abo-
riginals or First Nations people were also drawn to
these pieces of the past. The reasons varied, to be sure,
but hidden to us were concepts of evolution and a sense
of deep time. It is a great fortune that author Adri-
enne Mayor has investigated and recorded this diver-
sity of cultures, much of it based on the oral, not
written, tradition, and to some extent existing in vari-
ous artifacts.
Mayor’s first foray into fossil lore and history was
The First Fossils Hunters, Paleontology in Greek and
Roman Times (2000) which examined Greek and
Roman period and place with respect to fossils. Of
course these cultures are rich in textual artifacts and
documentation, and were in essence at the root of
western civilization. Mayor’s recent study, Fossil Leg-
ends of the First Americans, is somewhat more of a
challenge. As part of paleontological lore, aboriginal
communities certainly had contact with fossils but to
what extent? Lacking written tradition, painted in broad
strokes, and compounded by our interpretation of their
awareness, what were their concepts of “deep time”,
extinction, or even a notion of evolution?
Mayor argues that aboriginals had more than a cur-
sory view of fossils, and in many respects, and with a
much finer, multi-colored strokes particular to specific
groups, their views are much richer than anyone expect-
ed. By reviewing cultures from south to north, each
North American culture ebbed and flowed territorially,
originating or modifying concepts of their surround-
ings and to some degree of the fossil bones and stones
encountered. Some Navajos in the southwest United
States, for example, believe that it is horrific to disturb
buried bones. “The Navajos envisioned a series of past
worlds that were destroyed before this world” says
Mayor, each with good and evil monsters. By digging
them up, the fear emerges of bringing these past mon-
Sters to our world. The large fossils found in the Nava-
jo territory bespeak of life of great size in the form of
petrified logs from the Petrified Forest National Park
in Arizona, and of other possibilities in the rich, pale-
ontological nest that they could have come across. This
is more than mere tales of monsters.
Fossils were not always considered dire by those
who came across them. “Rock medicine of the Crows
and their traditional enemies, the Sioux, often included
unusual fossils, especially ammonites and baculites”
Book REVIEWS
127
(page 268). They believed that these fossils carried with
them mystical powers. In Alberta, medicine pouches
of the Blackfeet often carried “buffalo-calling stones”,
which are the segmented portions of a Cretaceous bac-
ulite that look the shape, in cross section, of a buffalo.
Aboriginals even attach some significance to the esthet-
ics of fossil-bearing rock, in one case a point made of
Pennsylvanian-age rock containing the ornate designs
of the single- celled marine fusulinids from Kansas.
Throughout Mayor’s work she references the revered
paleontologist the late George Gaylord Simpson (1902-
1984). Like the few earlier records of ethnology and
paleontology, Simpson is often taken to task for his
forays into the history of fossil collecting in North
America. His definition of “discovery” was much nar-
rower than Mayor’s. He believed that “merely seeing
a fossil bone or picking up an idle curiosity is hardly
discovery” (1943, page 26). Therefore, Simpson gave
little weight to beliefs and customs of First Nations
people with regard to fossils in his reviews. He knew
that there was contact with fossils, as in the case with
the abundant fossils from the “Big Bone Lick” of the
Ohio Valley, popular in the lore of American history
as well as paleontology, but that’s about it. Despite the
fact that Simpson’s views were reflective of 1940s
American culture, Mayor’s stance is that the discoveries
“were not scientific in the modern sense, of course, but
there is something akin to scientific inquiry in these
activities” (page 29-30). Many of the aboriginal oral
traditions were handed down from one generation to
another, thus becoming a permanent record in their
culture. This is one of the requisites of scientific tra-
dition.
The 19" century American expansionism included
vast paleontological initiatives like those seeded by
Joseph Leidy (1823-1891), O. C. Marsh (1831-1899),
and E. D. Cope (1840-1897). Ethnological initiatives
also followed, but for some reason, cultural interpre-
tation and fossils never met. Even when aboriginals
were hired as scouts for the paleontologists, the “pale-
ontologists ... [left] folklore to the ethnologists, so the
ethnologists left paleontological topics out of their
conversations with Native Americans” (page 195). As
such, paleontological/ethnological information gathered
today, some if recorded for the first time by Mayor, is
tainted with over a 150 years of invading culture. A
number of these aboriginal cultures have substituted
their original, monstrous, interpretations of fossils with
post-1840s terminology like “dinosaurs”.
Where Mayor falls short, however, is in the extent
of parallelism between aboriginal and western views
of fossils and their meaning. There are a lot of “may
have[s]’”, “could appear[s]” between the two views of
scientific and cultural interpretation, and thus some of
the interpretations seem somewhat forced. For exam-
ple, an Assinoboine tale has “a great river monster”
succumbing to a “violent death by the impact of a
128
“thunder stone”, a black projectile that came whistling
out of the west” at great speed and sound, “a scenario
that seems akin to the modern theory of an asteroid
impact 65 million years ago” (page 289; emphasis
added). The implied connection is not tenable. Likely
an interpretation of a meteor, the connection between
the extinction of a group of species by an extraterres-
trial object is a contemporary idea.
Fossil Legends of North American should have been
written a century ago when contact with the First
Nations people was still fresh. But the exploration of
the cultural/science intertwining is still a relatively
recent phenomenon. In this case, only a few other re-
cent titles address fossils in a cultural context (see
Cohen 2002; and to a lesser extent Semonin 2000).
Still, if it were not for Mayor’s effort, from document-
THE CANADIAN FIELD-NATURALIST
Vol. 120
ing the oral traditions, the searching of archives, muse-
ums and journals, we would be left with a culturally
insensitive, simpleton view of rich and diverse cul-
tures and their varied, unique surroundings.
Literature Cited
Cohen, C. 2002. The fate of the mammoth: Fossils, myth, and history.
University of Chicago Press, Chicago. 297 pages.
Mayor, A. 2000. The First Fossil Hunters. Princeton University
Press, Princeton, New Jersey. 361 pages.
Semonin, P. 2000. American monster: How the Nations first prehis-
toric creature became a symbol of National Identity. New York
University Press, New York. 482 pages.
Simpson, G. G. 1943. The discovery of fossil vertebrates in North
America. Journal of Paleontology, 17: 26-38.
TIM TOKARYK
Box 163, Eastend, Saskatchewan SON OTO Canada
A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s
Drive of Vancouver
By Rick Hudson, 2006. Volume 2 Revised Edition. Harbour
Publishing Co. Ltd., P.O. Box 219, Madeira Park, British
Columbia VON 2HO0 Canada. 368 pages $26.95 Paper.
Rick Hudson has succeeded in expressing his pas-
sion for mineral collecting through his field guide book.
On a second level he also tries to promote a sense of
adventure by teasing the reader to go out and explore
or prospect for nature’s buried treasures.
The author structured the book so the reader can eas-
ily cross-reference required information in the lengthy
location lists. A “How to use this Book” and a “Quick
Reference Guide” helps to quickly locate items of
interest throughout the book.
The book cover illustrates the hobby of mineral col-
lecting as a family activity. Hudson educates the reader
from first principles on how to become a rockhound.
To accomplish this, he describes how to prepare your-
self by choosing proper clothing, safety gear, collect-
ing tools and maps. He warns the future mineral col-
lector about natural hazards they may encounter such
as rattlesnakes, grizzly bears and ticks. Ethics, access
rights and mineral rights are also discussed to keep the
collector within acceptable collecting practices and to
NEw TITLES
+Available for review * Assigned
ZOOLOGY
The Eastern Palaearctic Species of Agnetina (Plecoptera:
Perlidae). Scopolia 56. Edited by I. Sivec, L. Zhiltzova and
B. Stark. 2005. Pensoft Publishers, Geo Milev Street 13a,
1111 Sofia, Bulgaria. 21 pages. EUR 15.00 Paper.
Discovering Amphibians. By J. Himmelman. 2006. Nim-
bus Publishing Ltd., P.O. Box 9166, Halifax, Nova Scotia
B3K 5M8 Canada. 208 pages. $23.95 Paper.
A Field Guide to the Amphibians and Reptiles of Bali.
By J. Lindley McKay. 2006. Krieger Publishing, P.O. Box
9542, Melbourne, Florida USA. US$39.50 Cloth.
avoid unintentional trespassing.
Mineral collecting is attractive to many people be-
cause as you find an interesting mineral specimen, you
want to learn how nature created it and or its geological
history. The author has included short geological
descriptions partitioned by collecting areas as well as
mining histories of local economic importance within
the collecting area.
Colour plates illustrate the minerals found in some
of the collecting sites and detailed occurrence maps
are included in all the collecting area sections through-
out the book. The colour plates represent minerals that
are possible to find in the collecting sites. Government
mineral index files were included in the locality infor-
mation along with detailed road log descriptions.
The author has made a successful merger of person-
al knowledge and technical data to compile an inform-
ative volume which can provide years of enjoyment for
many future rockhounds and their families and friends.
ROGER POULIN
President, Sudbury Rock and Lapidary Society, 3171 Romeo
Street, Val Caron, Ottawa P3N 1G5 Canada
The Amphibians of Belarus. Advances in Amphibian
Research, Volume 10. By S. Drobenkov, R. Novitsky, L.
Kosova, K. Ryzhevich, and M. Pikulik. 2006. Pensoft Pub-
lishers, Acad. G. Bonchev Street Bl. 6, 1113 Sofia, Bulgar-
ia. 176 pages. EUR 34.00 Paper.
The Amphibians and Reptiles of the Honduran Mosquita.
By J. McCranie, J. Townsend and L. Wilson. Krieger Publish-
ing P.O. Box 9542, Melbourne, Florida 32902-9542 USA.
302 pages. $63.50 US. Cloth.
Bolton’s Catalogue of Ants of the World, 1758 — 2005. By
B. Bolton, G. Alpert, P. Ward and P. Naskrecki. 2006. Har-
2006
vard University Press, 79 Garden Street, Cambridge, Mass-
achusetts 02138 USA. US$49.95.
The Beetles of the Galapagos Islands, Ecuador. By Stew-
art B. Peck. 2006. NHBS Environment Bookstore, 2-3 Wills
Road, Totnes, Devon TQ9 SXN United Kingdom. £26.50
Paper. [Available in Canada from publisher Monograph
orders NRC Resarch Press M-SS, National Research Coun-
cil Canada, Ottawa, Ontario KIA OR6 Canada. $39.95 +
GST 7%).
Bird Coloration — Volumes I and II. Edited by G. Hill and
K. McGraw. 2006. Harvard University Press, 79 Garden
Street, Cambridge, Massachusetts 02138 USA. 589 and 477
pages. US$95.00.
* Gibson’s Guide to Birdwatching and Conservation. By
Merritt Gibson. 2006. Nimbus Publishing Ltd., 3731 Mackin-
tosh Street, Box 9166, Halifax, Nova Scotia B3K 5M8 Cana-
da. 224 pages. $24.95.
* Birds of East Africa — Kenya, Tanzania, Uganda, Rwan-
da and Burindi. By T. Stevenson and J. Fanshawe. 2006.
Princeton University Press, 41 William Street, Princeton, New
Jersey 08540-5237 USA. 604 pages. US$35 Paper.
Atlas — Birds of Moscow City and the Moscow Region.
Pensoft Series Faunistica 54. By M. Kalyakin and V. Voltzit.
2006. (In English and Russian (bilingual). Pensoft Publishers,
Geo Miley Street 13a, 1111 Sofia, Bulgaria. 372 pages. EUR
59.90.
The Birds of Malawi — An Atlas and Handbook. By Fran-
coise Dowsett-Lemaire and Robert Dowsett. 2006. Tauraco
Press, Jupille-Liége, Belgium. 556 pages. £25.00 Paper.
Birds of Mexico and Central America. By Ber van Perlo.
2006. Princeton University Press, 41 William Street, Prince-
ton, New Jersey 08540 USA. 336 pages. $US29.95 Paper.
Birds of Moscow City and the Moscow Region. By M.
Kalyakin and O.V. Voltzit. 2006. Pensoft Publishers, Geo
Milev Street 13a, 1111 Sofia, Bulgaria. In English and Russ-
ian (bilingual). 372 pages. EURO 58.00 Cloth.
+ Birds of Prey Rescue. By Pamela Hickman. 2006. Firefly
Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1 Canada. 66 pages. $9.95 Paper.
New Holland Field Guide to the Birds of South-East
Asia. By Craig Robson. 2006. New Holland Publishers (UK)
Ltd., Garfield House, 86-88 Edgware Road, London W2 2EA
United Kingdom. £19.99 Paper.
Birds, Mammals and Reptiles of the Galapagos Islands:
An identification guide. 2"¢ Edition. By Andy Swash and
Robert Still. 2006. Yale University Press, P.O. Box 209040,
New Haven, Connecticut US$25 Paper.
Birds — A Visual Guide. By Joanna Burger. 2006. Firefly
Books Ltd. 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1 Canada. 304 pages. $29.95.
Birds of the World. By Les Beletsky. 2006. UBC Press, 2029
West Mall, Vancouver, British Columbia V6T 1Z2 Canada.
628 pages. $55 Cloth.
The Black Bear Almanac. By D. Smith. 2006. Globe Pequot
Press, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut
0643 USA. US$29.95. Cloth.
Fauna De Burpestidae De La Penisula Iberica Y Baleares
(Coleoptera). Edited by A Verdugo. [In Spanish.] 2005.
Book REVIEWS
129
Pensoft Publishers, Geo Miley Street 13a, 1111 Sofia, Bul
garia. 350 pages. EUR 89.00.
A Catalogue of Butterflies of the Ex-USSR, with Remarks
on Systematics and Nomenclature. By S. Korb. 2005
Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bul
garia. 158 pages. EUR 34.00 Paper.
Butterflies of Vietnam. Volume 1: Nymphalidae: Satyrinae
By A. Monastyrskii. 2005. Pensoft Publishers, Geo Mile
Street 13a, 1111 Sofia, Bulgaria. 198 pages. EUR 45.00 Paper
Butterflies of the World, Part 22, Nymphalidae X, Charax-
es I. By B. Turlin. 2005. Pensoft Publishers, Geo Miley Street
13a, 1111 Sofia, Bulgaria. 40 pages. EUR 38.00 Cloth.
Butterflies of the World, Part 23, Papilionidae XII, Par-
nassius Apollo III Text. By E. Moehn. {In German] 2005.
Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bul-
garia. 34 pages. EUR 25.00 Cloth.
World Butterflies. By Bernard d’ Abrera. 2006. Hill House,
Publishers, 491 Illington Road, Ossining, New York 10562
USA. 272 pages. £18.50 Paper.
Dodo: The Bird Behind the Legend. By Alan Grihault.
2005. Imprimerie & Papeterie Commerciale Ltee. Port-Louis,
Mauritius. 171 pages. £16.99 Paper.
* Fishes of the World. 4" edition. J. Nelson. 2006. John
Wiley & Sons Canada Ltd. 6045 Freemont Boulevard, Mis-
sissauga, Ontario L5R 4J3 Canada. 601 pages. US$125.00,
$162.99 Can.
Grizzlies and Grizzled Old Men : A Tribute to the Men
Who Fought to Save the Great Bear. By Mike Lapinski.
2006, FalconGuide, 246 Goose Lane, P.O. Box 480, Guilford,
Connecticutt 06437 USA. 186 pages. US$14.95 Paper.
Harvestmen. By R. Pinto-de-rocha, G. Machado and G. Giri-
bet. 2006. Harvard University Press, 79 Garden Street, Cam-
bridge, Massachusetts 02138 USA. 516 pages. US$125.
Lemurs of Madagascar. By R. A. Mittermeier, W. R. Kon-
stant, Frank Hawkins, Edward E. Louis et al. 2006. Conser-
vation International ,1919 M Street, NW Suite 600, Washing-
ton, District of Columbia 20036 USA. 520 pages. USS$25.
Paper 2™ edition revised from the original 1994 edition.
Field Guide to North Atlantic Wildlife. By N. Proctor and
P. Lynch. 2006. Yale University Press, P.O. Box 209040, New
Haven, Connecticut USA. 221 pages. US$19.95.
* Birding Sites of Nova Scotia —- A Comprehensive, Year-
Round Guide for Birders and Other Nature Lovers. By
B. Maybank. 2006. Nimbus Publishing Ltd., P.O. Box 9166,
Halifax, Nova Scotia B3K 5M8 Canada. 555 pages. $35.
The World of the Polar Bear. By Norbert Rosing. 2006.
Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1 Canada. 204 pages. $45.
World Catalogue of the Family Sepsidae (Insecta: Diptera).
Zoologicheskie Issledovania (Zoological Studies), Num-
ber 8. By A. Ozerov. 2005 Pensoft Publishers, Geo Milev
Street 13a, 1111 Sofia, Bulgaria. 76 pages. EUR 29.00 Paper.
* Tanagers, Cardinals, and Finches of the United States
and Canada. By D. Beadle and J. Rising. 2006. Princeton
University Press, 41 William Street, Princeton, New Jersey
08540 USA. 196 pages. US$29.95. Paper.
Fauna de Tenebrionidae de La Peninsula Iberica y Balear-
es. Volume 1 Lagriniinae y Pimeliinae. Edited by A. Vinolas.
130
C. Cartagena [in Spanish]. 2005. Pensoft Publishers, Geo
Milev Street 13a, 1111 Sofia, Bulgaria. 428 pages. EUR
110.00.
Whalewatcher — A global guide to watching whales, dol-
phins and porpoises in the wild. By T. Day. 2006. Firefly
Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1 Canada. 160 pages. $19.95.
The Urban Whale. Edited by S. Kraus and R. Rolland. 2006.
Harvard University Press, 79 Garden Street, Cambridge, Mass-
achusetts 02138 USA. 514 pages. US$55.
New Xixuthrina from Indo-Australian Region (Coleoptera,
Cerambycidae, Prioninae). By G. Marazzi, V. Marazzi
and Z. Komiya. 2006 Pensoft Publishers, Geo Milev Street
13a, 1111 Sofia, Bulgaria. 48 pages. EUR 35.00.
BOTANY
Aquatic and Wetland Plants of Eastern North America —
Volumes I and II. By G. Crow and C. Hellquist. 2006. (paper-
back edition) University of Wisconsin Press, 1930 Monroe
Street, Madison, Wisconsin 53711 USA. U.S.$45/volume
Emily Dickinson’s Herbarium (Facsimile Edition) Intro-
duction. By R. Sewall. 2006. Harvard University Press, 79
Garden Street, Cambridge, Massachusetts 02138 USA. 224
pages. US$135.
* Flowers — how they changed the world. By W. Burgur.
2006. Prometheus Books, 59 John Glenn Street, Amherst,
New York, USA. 337 pages. US$23. Cloth.
Plants on Islands Diversity and Dynamics on a Conti-
nental Archipelago. By Martin L Cody. 2006. University
of California Press, 2120 Berkeley Way, Berkeley, Califor-
nia 94720 USA. 315 pages. US$49.95. Cloth.
North American Mushrooms: A Field Guide to Edible
and Inedible Fungi. By Dr. Orson Miller and Hope Miller.
2006. Globe Pequot Press, 246 Goose Lane, P.O. Box 480,
Guilford, Connecticut 06437 USA. U.S.$25.95
The Tree of Life. By G. Lecointre and H. Le Guyader. 2006.
Harvard University Press, 79 Garden Street, Cambridge,
Massachusetts 02138. 560 pages. U.S. $39.95.
ENVIRONMENT
A Complete Guide to Arctic Wildlife. By R. Sale. 2006. Fire-
fly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1 Canada. 400 pages. $49.95.
Wild Borneo. By Nick Garbutt and Cede Prudente. 2006.
New Holland Publishers (UK) Ltd., Garfield House, 86 — 88
Edgware Road, London,W2 2EA United Kingdom. £29.99
Cloth.
* Guide to Deserts. Edited by A. Warren and T. Allan.
2006. Firefly Books Ltd., 66 Leek Crescent Richmond Hill,
Ontario L4B 1H1 Canada. 240 pages. $19.95.
Ecological Nationalisms — Nature, Livelihoods, and Iden-
tities in South Asia. Edited by Gunnel Cederlof and K.
Sivaramakrishnan. 2006. University of Washington Press,
1326 Fifth Avenue, Suite 555 Seattle, Washington 98101-2604
USA. US$50. Cloth.
The Emirates: A Natural History. Edited by Peter Hellyer
and Simon Aspinall. Trident Press. 175 Piccadilly, Mayfair,
London WIJ 9TB United Kingdom. 428 pages. £65.00 Cloth.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Endangered — Wildlife on the brink of extinction. By G.
McGavin and D. Burnie. 2006. Firefly Books Ltd. 66 Leek
Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 192
pages. $35.
Evolutionary Dynamics. By M. Nowak. 2006. Harvard Uni-
versity Press, 79 Garden Street, Cambridge, Massachusetts
02138 USA. 312 pages. U.S.$35.
The Revenge of Gaia — Why the Earth is fighting back —
and how we can still save humanity. By James Lovelock.
2006. NHBS Environment Bookstore, 2-3 Wills Road,
Totnes, Devon TQ9 5XN United Kingdom. 177 pages. £16.99
Cloth.
Galapagos: A Natural History. By John Kricher. 2006.
NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon
TQ9 5XN United Kingdom. US$19.95, £12.95. Paper.
Galapagos Islands Explorer. By Nigel Sitwell. 2006. NHBS
Environment Bookstore, 2-3 Wills Road, Totnes, Devon
TQ9 5XN United Kingdom. £9.99.
Field Guide to Gold Gemstones and Mineral Sites of BC
— Volume 2 Revised Edition (Sites within a Day’s Drive to
Vancouver). By Rick Hudson. 2006. Harbour Publishing,
4437 Rondeview Road, Madeira Park, British Columbia
VON 2HO Canada. 368 pages. $26.95
* A Paddler’s Guide to Killarney and the French River.
By K. Callan. 2006. The Boston Mills Press, 132 Main Street,
Erin, Ontario NOB 1T0 Canada. 168 pages. $19.95.
* Lapland — A Natural History. By D. Redcliff. 2006. Yale
University Press, P.O. Box 209040, New Haven, Connecticut
06520-9040 USA. 352 pages. US$60 Cloth.
The Man-Eaters of Eden: Life and Death in Kruger Na-
tional Park. By Robert Frump. 2006. The Lyons Press, 246
Goose Lane, P.O. Box 480, Guilford, Connecticut 06437
USA. 272 pages. US$24.95 Cloth.
Western Newfoundland and Gros Morne National Park.
By B. Bursey. 2006. Nimbus Publishing Ltd., 3731 Mackin-
tosh Street, Box 9166, Halifax, Nova Scotia B3K 5M8 Cana-
da. 88 pages. $22.95.
New Zealand — a natural history. By Tui De Roy and
Mark Jones. 2006. Firefly Books Ltd., 66 Leek Crescent,
Richmond Hill, Ontario L4B 1H1 Canada. 160 pages. $40.
A View of the River. By L. Leopold. 2006. Harvard Uni-
versity Press, 79 Garden Street, Cambridge, Massachusetts
02138 USA. 298 pages. US$19.95
+ States of Nature. By Tina Loo. 2006. UBC Press, 2029
West Mall, Vancouver, British Columbia V6T 1Z2 Canada.
320 pages. $85 Cloth.
Southeast Asian Biodiversity in Crisis. By Navjot S Sodhi
and Barry W Brook. 2006. Cambridge University Press, The
Edinburgh Building, Shaftesbury Road, Cambridge, CB2 2RU
United Kingdom. £65.00 Cloth.
The Green Tiger — The Costs of Ecological Decline in the
Philippines. By Barbara Goldoftas. 2006. Oxford Universi-
ty Press, 198 Madison Avenue, New York, New York 10016
USA. 264 pages. US$74 Cloth, US$24.95 Paper.
Vietnam: A Natural History. By Eleanor Jane Sterling,
Martha Maud Hurley and Le Duc Minh. 2006. Yale Universi-
ty Press P.O. Box 209040, New Haven, Connecticut 06520-
9040 USA. 448 pages. US$40 Cloth.
2006 Book REVIEWS 13]
Waterfowl Ecology and Management — 2" Edition. By
G. Baldassarre and E. Bolen, 2006. Krieger Publishing P.O.
Box 9542, Melbourne, Florida 32902-9542 USA. 580 pages.
US$112.50.
+ Walk around the Pond. By G. Walbauer. 2006. Harvard
University Press, 79 Garden Street, Cambridge, Massachu-
setts 02138 USA. 286 pages. US$22.95.
Wilderness Predators of the Rockies — The Bond Between
Predator and Prey. By Mike Lapinski. 2006. Falcon Guide,
246 Goose Lane, P.O. Box 480, Guilford, Connecticut 06437
USA. 208 pages. U.S. $17.95 Paperback
The World’s Wild Places. By C. Prior. 2006. Firefly Books
Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1HI
Canada. 224 pages. $49.95,
News and Comment
Ontario Natural Heritage Information Centre Science and Information Newsletter 11(1)
Winter 2006
Contents of this 20-page issue: FEATURE ARTICLE: Trans-
ferring conservation science to practitioners — 2005 PRO-
GRAM HIGHLIGHTS: Plant communities of Ontario upper
great lake marshes — NHIC to track rare disjunct arctic-
alpine plants — 2005 species at risk field survey highlights
— Rare plant surveys of Long Point region — Natural spaces
program — Mapping species at risk with biotics — Two
major dragonfly meetings in Ontario — 2005 PROJECT HIGH-
LIGHTS: NHIC launches Ontario Odonata atlas website —
Aquatic species database — Great Lakes islands conservation
blueprint — Lichen biodiversity project — NEWS AND
Marine Turtle Newsletter (111)
January 2006. 28 pages: EDITORIAL: Does the Mediter-
ranean Green Turtle exist? (N. Mrosovsky) — Comment on
the guest editorial by Paul J. Ferraro (P.C.H. Pritchard) —
Response to Comment by Peter C. H. Pritchard (P. J. Ferraro
— ARTICLES: Report on long-term transmitter harness
retention by a Leatherback Turtle (S. Dowling, R. Solano, A.
Diaz-Merry, J. Ordonez, J. Taylor, D. R. Evans, D. Godfrey,
D. Bagley, L. Ehrhart, and S. Eckert — Marine Turtles on the
Southern Coast of Bioko Island (Gulf of Guinea, Africa),
2001-2005 (H. Rader, M. A. E. Mba, W. Morra, and G. Hearn
— Incidental capture of marine turtles in Marine fisheries
of southern Spain (J.C. Baez, J. A. Camidax, and L. Rueda
— First evidence of Leatherback movement from Africa to
South America (A. Billes, J. Fretey B. Verhage, B. Huijbregts,
B. Giffoni, L. Prosdocimi, D. A. Albareda, J.-Y Georges, and
M. Tiwari — NOTES: Charybdlis hellerii, a non-indigenous
portunid crab from the gastrointestinal contents of a Kemp’s
Ridley Sea Turtle (Lepidochelys Kempi) in Georgia, USA (M.
Frick and K. Williams) — An update on Eritrea’s marine
Wildlife Afield 2(1,2)
A British Columbia semi-annual publication (ISSN 1712-
2880) published by The Biodiversity Centre for Wildlife
Studies twice a year. Additional information at www.wild
lifebc.org.
Volume 2, Number 1, contains FROM THE EDITOR:
Spinning a web — On the covers — FEATURE ARTICLES:
Migratory occurrence and status of select shorebirds in the
vicinity of Fort St. John, British Columbia (Chris Siddle) —
Survey of Painted Turtles in the Creston Valley, British
Columbia, in 2003 and 2004 (Linda M.Van Damme) —
NOTES: Changing status of Ring-necked Ducks wintering
on Salt Spring Island, British Columbia, 1978-2004 (Robert
B. Weeden — Barred Owl feeds on Ban-tailed Pigeon (David
C. Schultz) — First winter occurrences of the Cape May
Warbler for British Columbia (Phil Ranson) — Burdock
entrapment of a Ruby-crowned Kinglet in the Creston Valley
— First nest and eggs the Canada Warbler for British Col-
umbia (R. Wayne Campbell) — Pinyon Jay (Gymnorhinus
cyanocephalus): A new species for British Columbia (Kim
Asquith and Malcolm Dennington) — Incidences of Gopher
NOTES: Summer experience program at the NHIC — Ontario
eastern habitat joint venture — GIS Project — Eastern region
Natureserve conference — NHIC data supports the assess-
ment of species of global conservation concern in Canada —
NHIC joins new biodiversity section of MNR — Publication
notices — NHIC staff information.
Mailing address for Natural Heritage Information Centre,
Ontario Ministry of Natural Resources, 300 Water Street,
2" Floor, North Tower, P.O. Box 7000, Peterborough, Ontario
K9J 8M5, Canada; www.nhic.mnr.gov.on.ca/MNR/nhic.cfm.
turtle programme and first record of Olive Ridley turtle nesting
in the Red Sea (n. Pilcher, S. Mahmud, S. Howe, Y. Tecle-
mariam, S. Weldeyohannes, T.Menggsu, and M. Giotom —
Epibionts associated with Chelonia mydas from northern
Brazil (S. Pereira, E.H. S. M. Lima. L. Ernesto, H. Mathews,
and A. Ventura — Small Leatherback found in Dominica (R.
Byrne and S. A. Eckert) —MTSG UPDATE — ANNOUNCE-
MENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICA-
TIONS.
The Marine Turtle Newsletter is edited by Brendan J.
Godley and Annette C. Broderick, Marine Turtle Research
Group, Centre for Ecology and Conservation, University of
Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ
United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392
263700. Subscriptions and donations towards the production
of the MTN can be made online at or postal mail to Michael Coyne (online Editor)
Marine Turtle Newsletter, 1 Southampton Place, Durham,
North Carolina 27705 USA (e-mail: mcoyne@seaturtle.org).
Snake predation in bluebird nest boxes — Unsuccessful
nesting attempt of Black-necked Stilt in British Columbia
(Ryan Tomlinson) — Status of the Rock Pigeon (Columba
livia) in Nelson, British Columbia (Janice E. Arndt) —
WILDLIFE DATA CENTRE: Featured species — Thinhorn
Sheep (Fred L. Bunnell) — Report of the Wildlife Data
Centre: | January 2005 to 30 June 2005 (Michael I. Pres-
ton) — BRITISH COLUMBIA ROUND-UP: News of Friends
— From the Archives — Final Flight — Publications of In-
terest (Compiled and edited by Chris Siddle) — Announce-
ments and Meetings.
Volume 2, Number 2, contains: FROM THE EDITOR: A
turn of the times —On the covers — FEATURE ARTICLES:
Field Survey of Red-eared Sliders (Trachemys scripta ele-
gans) in the Lower Fraser Valley, British Columbia, in 2005
(Cory G. Bunnell — Origin and current status of the Pacific
Pond Turtle (Actinemys marmorata) in British Columbia
Francis R. Cook, R. Wayne Campbell and Glenn R.
Ryder — Recent range expansion of Sandhill Cranes (Grus
canadensus tabida) in southeastern British Columbia (Doug -
132
2004
las Leighton — Diet of Great Horned Owl in the Creston
Valley, British Columbia, 1998-2005 (Linda M. Van Damme
— NOTES: Anti-predator behaviour by a Mule Deer to Coyotes
in Vernon, British Columbia (Chris Siddle) — Attempted
capture of a Red-legged Frog by an American Robin (Michael
I. Preston) — Evidence of an unusual prey item in a Barred
Owl pellet (Christian W. Gronau) — Northernmost nest record
of the Flammulated Ow! (Otus flammeolus) in Canada (Karen
L. Wiebe — First nesting record of Say’s Phoebe (Sayornis
saya) in the East Kootenay, British Columbia (Robert S.
Ferguson and Mark Dascher) — Current status of the Black-
throated Blue Warbler in British Columbia (Gary S. Davidson
— Field observations of Gray Flycatchers breeding at Sum-
merland, British Columbia, 1988-2005 (I. Laurie Rockwell)
— American Crow predation in an Eared Grebe nesting
colony at Rawlings Lake, British Columbia (Chris Siddle)
Errata The Canadian Field-Naturalist 119(3)
Catling, Paul M. 2005 [2006]. Additions to the flora of the
continental Northwest Territories from the Great Slave
area. Canadian Field-Naturalist 119(3): 437-440.
(page 438) Actaea rubra f. neglecta : This form had previous-
ly been reported by Cody 1961. Canadian Field-Naturalist
75(2): 62 from Fort Simpson. Michael Oldham (Ontario
Ministry of Natural Resources) reports confirming its pre-
sence there in August 2007.
NEWS AND COMMENT
133
— SPECIES OF REGIONAL SIGNIFICANCE: Canvasback
Fort St. John (Chris Siddle) — Bobolink — Nakusp to Edge-
wood (Gary S. Davidson) WILDLIFE DATA CENTRE
Featured species — Turkey Vulture (R. Wayne Campbell,
Michael I. Preston, Linda M. Van Damme, and Diann Mac-
Rae — BRITISH COLUMBIA ROUND-UP: Field notes and
caring people — News of Friends — From the archives —
Final flight — Publications of interest [Compiled and edited
by Chris Siddle}] — Announcements and meetings.
Annual memberships in the Biodiversity Centre for Wildlife
Studies (which included Wildlife Afield are $30 (individual),
$40 (family). $20 (student), $500 (life). Biodiversity Centre
for Wildlife Studies, PO Box 6218, Station C, Victoria,
British Columbia V9P 5LS Canada. Tel/Fax: 250-477-0465;
e-mail: editor@ wildlifebc.org
(page 449) Sonchus arvennsis subspecies uliginosus : Thiert
reported this in 1963 Sida 1(3): 169 as S. arvensisvar.
glabrescens from the Yellowknife Highway (Hwy 3).
My appreciation to Michael Oldham for bringing these to my
attention.
PAUL M. CATLING
Agriculture and Agri-Food Canada
Ottawa, Ontario
134
THE CANADIAN FIELD-NATURALIST
Vol. 119
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cheque or money order paid when the invoice is received.
FRANCIS R. COOK, Editor
TABLE OF CONTENTS (concluded) Volume 120 Number 1
Notes
Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine A.C. A. BURCHSTED and FRED BURCHSTED
Ring-billed Gull, Larus delawerensis, food piracy on diving ducks WILLIAM J. WALLEY
Probable Black Bear, Ursus americanus, retrieval of an Elk, Cervus elaphus, carcass from a
small lake in Riding Mountain National Park, Manitoba WILLIAM J. WALLEY
Book Reviews
ZooLoGyY: The Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi — The Birding Sites of
Nova Scotia — The Camplete Fauna of Iran — Fishes of the World — Insects: Their Natural History and
Diversity: With a Photographic Guide to Insects of Eastern North America — Secret Weapons: Defenses
of Insects, Spiders, Scorpions, and Other Many-legged Creatures — Rodents and Lagomorphs of British
Columbia — Tanagers, Cardinals, and Finches of the United States and Canada — Wheatears of Palaeartic:
Ecology, Behaviour and Evolution of the Genus Oenanthe
Borany: Giant Trees of Western North America and the World
ENVIRONMENT: Fire and Avian Ecology in North America: Studies in Avian Biology Number 50 — Fisheries
Assessment and Management in Data-limited Situations — Lapland: A Natural History — Fish, Fur and
Feathers: Fish and Wildlife Conservation in Alberta, 1905-2005 — The Gulf of Alaska: Biology and
Oceanography
MISCELLANEOUS: Fossil Legends of the First Americans — A Field Guide to Gold, Gemstones & Mineral
Sites of British Columbia, Sites within a Day’s Drive of Vancouver
New TITLES
News and Comment
| Ontario Natural Heritage Information Centre Science and Information Newsletter 11(1) Winter 2006 —
Marine Turtle Newsletter (111) — Wildlife Afield 2 (1,2) — Errata The Canadian Field-Naturalist 119(3)
Advice to Contributors
!
| Mailing date of the previous issue 119(4): 25 April 2007
2006
106
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THE CANADIAN FIELD-NATURALIST Volume 120 Number 1
Articles
Predicting bird oiling events at oil sands tailings ponds and assessing the importance
of alternate waterbodies for waterfowl: a preliminary assessment ROBERT A. RONCONI
A test of interspecific effects of introduced Grey Squirrels, Sciurus carolinensis,
on Douglas’s Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia
YEEN TEN HWANG and SERGE LARIVIERE
Reponses of syrphids, elaterids, and bees to single-tree selection harvesting in
Algonquin Provincial Park, Ontario ERICA NOL, HUME DOUGLAS, and WILLIAM J. CRINS
New nesting records of the Le Conte’s Sparrow, Ammospiza leconteii, from
northeastern Ontario with some notes on nesting behaviour MICHAEL PATRIKEEV
Conditions for sexual interactions between wild Wolves, Canis lupus,
and Coyotes, Canis latrans RICHARD P. THIEL
Multiple scale den selection by Swift Foxes, Vulpes velox, in southeastern Colorado
ANN KITCHEN, ERIC M. GESE, and SARAH G. KuPIS
First records of the Northern Long-eared Bat, Myotis septentrionalis, in the Yukon Territory
THOMAS S. JUNG, BRIAN G. SLOUGH, DAVID W. NAGORSEN, TANYA A. DEWEY, and TODD POWELL
Recent declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces
ANTHONY J. ERSKINE
First record of the Southern Red-backed Vole, Clethrionomys gapperi, in Newfoundland:
Implications for the endangered Newfoundland Marten, Martes americana atrata
BRIAN J. HERN, JOHN T. NEVILLE, WILLIAM J. CURRAN, and DEAN P. SNow
Long-range homing by an adult female Black Bear, Ursus americanus
L. J. LANDRIAULT, M. N. HALL, J. HAme, and F. F. MALLORY
Early ontogenetic diet in Wolves, Canis lupus, of coastal British Columbia
HEATHER M. BRYAN, CHRIS T. DARIMONT, THOMAS E. REIMCHEN, and PAUL C. PAQUET
Extension of Coyote, Canis latrans, breeding range in the Northwest Territories,
Canada H. DEAN CLUFF
Lake Sturgeon, Acipenser fulvescens, movements in Rainy Lake, Minnesota and Ontario
W. EUGENE ADAMS JR., LARRY W. KALLEMEYN, and DAVID W. WILLIS
Northern range expansion and invasion by the Common Carp, Cyprinus carpio, of the Churchill
River system in Manitoba PASCAL H. J. BADIOU and L. GORDON GOLDSBOROUGH
Diversity and range of amphibians of the Yukon Territory
BRIAN G. SLOUGH and R. LEE MENNELL
Temporal variability of cetaceans near Halifax, Nova Scotia
PETER SIMARD, JENIFER L. LAWLOR, and SHANNON GOWANS
Winter habitat use by American Marten, Martes americana, in western Alberta
boreal forests GILBERT PROULX
2006
39
43
93
100
(continued on inside back cover)
ISSN 0008-3550
84
The CANADIAN
FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
Volume 120, Number 2 April—June 2006
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patron
Her Excellency The Right Honourable Michaille Jean
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members
Edward L. Bousfield Ellaine Dickson George F. Ledingham E. Franklin Pope
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2006 Council
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Vice-President: Ken Alison Fenja Brodo Karen McLachalan Hamilton Justin Peter
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this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.
PAP Registration Number 9477. Canada We acknowledge the financial support of the Government of Canada through the
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Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca
Copy Editor: Elizabeth Morton
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mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor.
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Cover: Elk, Cervus elaphus, part of herd near Hartsmore, Ontario. Photographed 1 February 2005 by R. Rosatte. See paper on late!
born calf — pages 188-191.
Volume 120, Number 2
The Canadian Field-Naturalist
—“'BRARY
NOV 28 2007
AR VA
UNE iD
April-June 2006
A Tribute to George Wayne Douglas 1938 — 2005
ener L. PENNY!
-'Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, P.O. Box 9993 Stn Prov Govt,
| Victoria, British Columbia V8W 9R7 Canada.
Penny, Jenifer L. 2006. A tribute to George Wayne Douglas 1938-2005. Canadian Field-Naturalist 120(2): 135-146
Coming from humble beginnings, George Wayne
Douglas, with his determination and strong spirit, estab-
lished himself as one of British Columbia’s most res-
pected botanists. I first met George in 1995 when |
began working with him at the British Columbia Con-
servation Data Centre (CDC). In time, I came to know
him as an adept field botanist, a knowledgeable ecolo-
gist, an accomplished author, a practical taxonomist,
a cunning businessman, a conservationist at heart, and
a generous mentor. During our numerous field trips,
George often talked about writing his memoirs and
enjoyed recounting the stories and adventures that
would go into the chapters. He had a lot of different
experiences throughout his life that would have result-
ed in an interesting read. He was a born leader: he
had a strong character, held his ground on issues, and
had a critical, but practical approach. George had a
vision for botany in British Columbia and he brought
that vision to fruition.
_ George (known to family and close friends as
Wayne) was born in the Royal Columbian hospital in
New Westminster on 22 June 1938. He spent his early
years exploring the bushes around the base of Burna-
by Mountain near Vancouver, British Columbia, this
may have been the root of his inspiration to study
botany and ecology later in his life. He grew up in a
large family with six brothers and sisters: Bonnie, Mar-
‘ilyn, and David who currently live in British Columbia,
‘and Robert, who lives in South Africa (siblings David
‘Charles and Judy predeceased him). His family was
‘not affluent, so George always aspired to do well for
himself, which he certainly accomplished.
As a youth, he excelled at sports but wasn’t that
jinterested in academics. In high school (Figure 1), he
‘ran record sprints and was an excellent basketball play-
ler, as well as an accomplished golfer. He would con-
‘tinue to enjoy golfing throughout his life, playing at
least two to three times a week. When he first started,
vat about 12 years of age, he used to sneak onto the Van-
couver Golf Course, and golf his way to school, hide
'
FIGURE |. George at Como Lake High School (circa 1956).
his clubs and then golf his way home after school.
Later he was on the high school golfing team and was
an amateur champion. He always knew how to have
fun, especially back in high school, but he also worked
hard. One of his favourite jobs at that time was at
Labatt’s Brewery where his Friday afternoons were
spent “sampling” the products.
136
Despite his lack of interest in scholarly pursuits in
high school, George ended up at the University of
Washington (College of Forest Resources) in Seattle,
Washington, on a track and field, basketball and golf
scholarship. George continued to work on his athlet-
ics; however, much to his surprise, he became very
keen on his studies and excelled in his field. Al-
though he never finished high school, during his edu-
cational career he ended up receiving three post-grad-
uate degrees!
George had a scholarship but he still had to work
part-time jobs to finance going to university. Amongst
his occupations were working in a mill, a cannery and
a slaughterhouse. He was also an ambulance and taxi
driver. Later in his schooling he got work that better
suited his career goals like as a forestry consultant,
where he supervised forest inventory, mapping and re-
forestation projects. He also worked as an ecological
consultant while attending University of Washington
and completed inventories in the North Cascades Na-
tional Park focussing not only vegetation, but also
wildlife, soils and geology.
During his bachelor studies, George received tute-
lage from two famous Pacific Northwest botanists,
Dr. C. Leo Hitchcock and Dr. Art Kruckeberg. In later
years, George dedicated one of his books to Dr. Kruck-
eberg (Mountain Plants of the Pacific Northwest), in
which he wrote that Dr. Kruckeberg had inspired him
during his first plant identification attempts and start-
ed him on the road to his lifelong career in botany.
Somehow, George even ended up getting bestowed
with Dr. Hitchcock’s plant press, a true honour for a
botanist! As a result of considerable effort during his
bachelor degree, he was accepted into graduate school
and studied under Dr. Ronald J. Taylor. George re-
ceived a Bachelor of Science degree in forestry in 1968
and a Master’s of Science degree in 1970 from Uni-
versity of Washington.
George’s master’s thesis involved a vegetation study
in the subalpine zone of the western North Cascades,
Washington, which was published in Arctic and Alpine
Research (Douglas 1970). Following his Master’s de-
gree, he moved back to Canada to embark on his Ph.D.
at the University of Alberta (Department of Botany) in
Edmonton, Alberta, with Dr. Larry Bliss. It was a good
time to return to Canada since the United States Army
was rallying young troops for Vietnam. His doctorate
was a study of alpine and high subalpine plant com-
munities of the North Cascades Range, in Washington
and British Columbia, and he published the research
in Ecological Monographs (Douglas and Bliss 1977).
In 1974, following his doctorate at University of Alber-
ta, he received a Postdoctoral Fellowship in the Depart-
ment of Botany to study lichens of the North Cascade
Ranges of Washington (Douglas 1974).
His graduate work took him to the places he loved
— the mountains of the Cascade Range. In fact, there
is a special place on Mount Baker called Grouse Ridge
where he wanted to have his ashes spread, but alterna-
THE CANADIAN FIELD-NATURALIST
Vol. 120
tively he thought that Mount Prevost overlooking his
home in Duncan would be a fine place too, which is
where his family ultimately scattered his ashes.
After finishing his graduate studies, George took
courses in landscape architecture at Pennsylvania State
University, which later enhanced his immense enjoy-
ment of gardening at home, but at the time, made him
more marketable as a consultant. Then he focussed
entirely on developing his consulting firm, Douglas
Ecological Consultants Ltd., based out of Victoria, Bri-
tish Columbia. His consulting work was to take him
all over British Columbia and to the Yukon Territory
and resulted in valuable vegetation interpretations and
guidance to land managers on the issues of environ-
mental impacts and biodiversity monitoring.
In 1973 and 1974, he started working in the Yukon
Territory, where the alpine was full of new species and
breath-taking landscapes. His time there began when
he was contracted by the Canadian Wildlife Service to
examine the vegetation and soils of Kluane National
Park and their vulnerability to land use within the park.
George ended up spending eight summers working in
the park, a good time for the business, but also some
of the best memories of his life.
Parks Canada also hired him to work on many im-
portant projects in the 1970s. In fact, he was awarded
some of Parks Canada’s largest contracts ever. Dur-
ing this time he employed numerous prestigious sci-
entists and colleagues to provide expertise on his proj-
ects. Among other projects, he was asked to evaluate
the use and ecological impacts of chemical fire retar-
dants in North America and to make recommenda-
tions on their future use, to assess the environmental
impact of recreational use (mainly human and horse
trampling) on different vegetation types in Waterton
Lakes National Park, Alberta, and to assess the envi-
ronmental impact of five potential transportation cor-
ridors in Kluane National Park. He also carried out a
four-year biophysical inventory project in Kluane Na-
tional Park, which was a multidisciplinary study that
included ten professional scientists and reported on
landforms, surficial deposits, soils and vegetation.
Douglas Ecological Consultants also worked on
projects involving vegetation reconnaissance of large
tracts of land in British Columbia, examining spruce
hybridization in the province, evaluating the British
Columbia ecological reserve system, air quality mon-
itoring projects using lichens in northern Alberta, and
a variety of environmental impact studies.
In the early 1980s, George started working on con-
tracts for the British Columbia Ministry of Environ-
ment (MOB), for which he would work until his retire-
ment in 2002. He started out at MOE as a consultant,
designing and coordinating a biological monitoring net-
work using lichens in northeastern British Columbia.
He also investigated the occurrence of alpine plant
communities in British Columbia for the Ecological
Reserves unit of MOE, and suggested where new eco-
logical reserves should be established.
2006
PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS
137
eo: , . ~*
a re : he . be
Pea oe
Pa Se ee
alt Se
. ’
FIGURE 2. George (left) and Lloyd Freese above the Tatshenshini River. 1992. Photo credit unknown.
At that time, he also became a Royal British Colum-
bia Museum research associate and started working
on a complete taxonomical and ecological treatment of
approximately 360 taxa in the Asteraceae (sunflower
family) in British Columbia. Unfortunately, only two
of the planned three volumes were ever published due
to funding difficulties. He was also involved in numer-
ous articles and government documents in the 1980s,
and he edited and contributed to an attractive and
informative coffee table book on Kluane Park, Yukon,
called Kluane, Pinnacle of the Yukon (Theberge and
Douglas Editors 1980). His work in the Yukon also
resulted in the Rare Vascular Plants of the Yukon (Dou-
glas et al. 1981).
George also established a strong relationship with
ecologists in the British Columbia Ministry of Forests
(MOF) in the 1980s as well as the 1990s. He worked
on floristic manuals and bibliographies for them, and
annotated several thousand sheets of British Colum-
bia plants. George and his colleagues at MOF (Del
Meidinger, Andy Mackinnon and Jim Pojar) felt that
it was of utmost importance to have a flora for British
Columbia to provide a foundation for all inventory,
floristic and conservation work going on within the
province. Therefore, he collaborated with them on the
very first flora for British Columbia and became the
chief editor of the The Vascular Plants of British Col-
umbia (Douglas et al. 1991-1994). For the first time
ever, botanists had a flora for British Columbia all in
one work. Previously, botanists had to carry a vari-
able number of floras depending on what part of the
province they were working in.
In the 1990s, George was also involved in the fol-
lowing very popular field guides: Plants of Northern
British Columbia (MacKinnon et al. 1992), Plants of
the Southern Interior of British Columbia (Parish et al.
1996), and the second, expanded edition of the Moun-
tain Plants of the Pacific Northwest (Taylor and Dou-
glas 1995). In total, he contributed approximately 45
publications during that decade that greatly enhanced
the knowledge of vascular plants in British Columbia
and the Pacific Northwest.
In addition to his valuable contributions to the botan-
ical literature, George also added thousands of valuable
collections to major herbaria in Canada. During his
career, he collected 13893 specimens, many of which
represented new taxa for the province, new range exten-
sions for known taxa, rare or at-risk species, and other
interesting finds. From his collections in the Cascade
Mountains he and John Packer named a new Erigeron
(fleabane) for the Pacific Northwest, Erigeron salishii
(Salish Daisy) (Douglas and Packer 1988). He also
made the first collections in British Columbia of species
like Orthocarpus barbatus (Grand Coulee Owl-clover),
Erigeron leibergii (Leiberg’s Fleabane), and Tonella
tenella (Small-flowered Tonella).
138
In 1991, George started working for the British
Columbia government as the Program Botanist at the
CDC. This position gave him the opportunity to fur-
ther contribute to floristics in British Columbia through
the CDC mission of identifying, tracking, and monitor-
ing the rarest or most at-risk plants in the province.
To accomplish this goal, George continued to collect
and amass important collections. In addition, through
inventory work, George travelled throughout the prov-
ince chasing the plants that were on the CDC’s track-
ing list and surveying a wide variety of habitats, includ-
ing range lands, wetlands and mountain tops. The year
that I joined the CDC, we toured the peaks of the south-
em Rocky Mountains for five days via helicopter escort.
Because of his background in alpine ecology, mountain
excursions were definitely a focus for George, but also
critical was our inventory work in the other habitats for
rare plants, such as the sand dunes of the west coast of
Vancouver Island, the hot springs in the East Koote-
nays, and the beaches of Osoyoos Lake. These trips
sound like a series of holidays but that was where our
work took us to confirm the conservation status of rare
and at-risk plants in British Columbia.
In order to accurately survey and track the species,
George not only saw the need for meticulous inspec-
tion of the herbaria collections and for inventory work
in poorly surveyed areas as well as in regions under
high development pressure, but he also saw the need
for an updated flora. Therefore, in the late 1990s he
started focusing his efforts on the production of the
Illustrated Flora of British Columbia (Douglas et al.
1998-2002). At that time, he also co-authored the Rare
Native Vascular Plants of British Columbia (Douglas
et al. 2002) so that land use planners and field workers
had an easy-to-use reference to the CDC-listed plants.
The Illustrated Flora of British Columbia (Douglas
et al. 1998-2002) dominated George’s life from the late
1990s to 2002. Since he was so dedicated to its com-
pletion, he worked on it at home as much as he did in
the office. The production of the flora not only involved
coordinating the contributors’ schedules, keeping them
on timelines and reviewing the products, but it also
involved compiling high quality illustrations (some bor-
rowed, others newly commissioned). The maps were
a whole other project (published in Volume 8 of the
series) — the first complete set of distribution maps for
British Columbia plants ever done. It was an enormous
effort but with enormous benefits.
The inventory work that George was involved with
at the CDC not only allowed for more accurate prov-
incial listing of rare and at-risk taxa, but also allowed
him to prepare numerous status reports on the local
southeast Vancouver Island and Okanagan Valley En-
dangered and Threatened plant taxa. As a result of
these studies, he authored and co-authored 40 Commit-
tee on the Status of Endangered Wildlife in Canada
(COSEWIC) status reports and update status reports,
more than any other contributor of reports to the nation-
al species assessment team!
THE CANADIAN FIELD-NATURALIST
Vol. 120
Along the way, George encouraged many of the in-
dividuals that he mentored to co-author status reports.
He often invited enthusiastic students and other pro-
tégés to join him on field outings where he was very
forthcoming about sharing his botanical knowledge.
He also spent time helping the Co-operative Educa-
tion students that worked at the Royal British Colum-
bia Museum and many others that sought his advice.
He encouraged his staff with regular bouts of appre-
ciation. Every year, at Christmas his botanical support-
ers at the CDC were invited to lunch at his favourite
spot, The Royal Colwood golf course clubhouse, where
patrons have a great view of the lovely grounds with
its ancient Douglas-firs and Garry Oaks, manicured
greens and ponds.
Through his work it was obvious that George was
a conservationist at heart. He may not have portrayed
the image of one in the typical sense, but he always
extolled the virtues of preserving natural areas. In the
early 1990s, he participated in the Sierra Club expe-
dition down the Tatshenshini River, northern British
Columbia, part of a successful campaign to have a pro-
tected area established in there (Figure 2).
He became a member of the Garry Oak Ecosystem
Recovery Team Rare Plant Recovery Implementation
Group, and contributed valuable input to the mem-
bership discussions. He was also on the Adiantum
capillus-veneris (Maidenhair Fern) recovery team and
performed an advisory role to the South Okanagan-
Similkameen Conservation Program. In his consulting
work, he contributed to conservation through research
and inventory, but also through work with golf clubs
advising them on how to create environmentally friend-
ly courses.
He also supported an important Nature Conservan-
cy of Canada campaign to purchase one of the last
remaining sizeable stands of Garry Oak in sem1-pris-
tine condition left on southeastern Vancouver Island, a
property owned by the Elkington family, located east
of Duncan. He did early inventories in the Garry Oak
ecosystem, and advised the Nature Conservancy staff
about the biodiversity values of the property during the
raising of funds to purchase it. In 1999, George had the
opportunity to conduct the most intensive and objective
study of a Garry Oak woodland ever done in British
Columbia on this very property, later coined the “Cow-
ichan Garry Oak Preserve”. The study examined the
composition, phenology, stand structure, and soils of
the Garry Oak woodland. Phenological data was ob-
tained on a weekly basis from March until September
and soils were also examined in detail from three soil
pits. In 2004, Douglas Ecological Consultants Ltd.
repeated the vegetation component of the study to mon-
itor changes after five years (Smith and Douglas 2005).
During his government career, George reached out
to many different groups to communicate his message
to protect rare and at-risk plants. In particular, he estab-
lished a relationship with Osoyoos Indian Band, and
befriended Environmental Officer Ron Hall. He spent
2006
many summers doing inventories on Osoyoos band
lands monitoring the special habitats that they have
conserved over the years in the region, while all sur-
rounding properties have been converted to vineyards,
orchards, golf courses, and residential and commercial
developments. Even after he retired from government
in 2002, he continued to work with the band helping
them with invasive species control in sensitive rare
plant habitats.
When George retired from government after 11 years
of service in early 2002, he started up Douglas Eco-
logical Consultants Ltd. full-time once again. He was
very excited about getting out in a consulting capacity
like the “old days” on some field trips since the inven-
tory work had been scant in recent years with the gov-
ernment. In his capacity as a consultant to government,
he became active in a variety of interesting botanical
and ecological projects and produced many reports.
In the last three years of his life, he worked on numer-
ous status reports, four species at risk recovery strate-
gies, and one multi-species recovery strategy. Sadly,
his life and work were cut short by a long battle with
cancer. He died on 10 February, 2005 at Cowichan
District Hospital, Duncan, British Columbia. He left
behind wife Sylvia, son Geordie, daughter-in-law Mary
McPhail, and grandchildren Mackenzie and Ryder, his
four siblings and numerous nieces and nephews.
He has, however, left a truly inspiring legacy. His
botanical contributions are used often by many plant
lovers, botanists and ecologists in British Columbia and
beyond. He will be remembered as a leader, not a fol-
lower, who made things happen. He was hard-working
and passionate about his work. His lengthy bibliogra-
phy and professional experience, which follows, will
attest to this assertion.
Other tributes to George W. Douglas appeared in
Botanical Electronic News (BEN), the first, an obitu-
ary written by his MOF colleagues, Del Meidinger,
Jim Pojar and Andy MacKinnon, and the second, a
tribute to George’s immense bibliography (http://www.
ou.edu/cas/botany-micro/ben/ben344.html; http://www.
ou.edu/cas/botany-micro/ben/ben351.html). The bib-
liography found in this article is an updated version
of the one that was published in October of 2005 in
“BEN # 351” and is found at http://www.ou.edu/cas/
botany-micro/ben/suppl/35 1/gwdouglas.html.
An obituary also appeared in the local Duncan and
Victoria papers that featured one of George’s favourite
pictures with his beloved cat, Garfield, where the two
of them are reading a Czech satirical, antimilitaristic
novel from the First World War given to him many
years ago by a valued colleague (Figure 3; http://www.
obituariestoday.com/Obituaries/ObitSearchList.cfm).
Acknowledgments
Thanks to Sylvia Douglas for important insights into
an era in George’s life that pre-dates me and for editing
help. Thanks also to Brenda Costanzo, Adolf Ceska,
PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS
139
esvesebere the:
7 ee.
FIGURE 3. George and his beloved cat, Garfield. Circa 1993.
Photo by Sylvia Douglas.
Sharon Hartwell, Diana Demarchi, Frank Lomer, Marie
Malo (née Fontaine), Gail Harcombe, Marta Donovan,
and Del Meidinger who graciously provided review
comments. George would have appreciated that they
contributed to the article.
Positions in Scientific and Professional Societies
Member: Northwest Scientific Association; Ottawa
Field-Naturalists’ Club; Rare Plant Consortium; Asso-
ciation of Professional Biologists
Research Associate: Royal British Columbia Muse-
um (1991-2004).
Dissertations and Post-doctoral Fellowships
Douglas, G. W. 1970. A vegetation study in the subalpine
zone of the western North Cascades, Washington. M.Sc.
thesis, University of Washington, Seattle, Washington. 293
pages.
Douglas, G. W. 1973. Alpine plant communities of the North
Cascades Range, Washington and British Columbia. Ph.D.
thesis, University of Alberta, Edmonton, Alberta. 145 pages.
Postdoctoral Fellowship: 1973-1974, Department of Botany,
University of Alberta, Edmonton, Alberta.
Publications
Douglas, G. W., and R. J. Taylor. 1970. Contributions to the
flora of Washington. Rhodora 72: 496-501.
140
Franklin, J. F., W. H. Moir, G. W. Douglas, and C. Wiberg.
1971. Invasion of subalpine meadows by trees in the Cas-
cades Range, Washington and Oregon. Arctic and Alpine
Research 3: 215-224.
Douglas, G. W., and T. M. Ballard. 1971. Effects of fire on
alpine plant communities in the North Cascades, Wash-
ington. Ecology 52: 1058-1064.
Douglas, G. W. 1972. Subalpine plant communities of the
western North Cascades, Washington. Arctic and Alpine
Research 4: 147-166.
Douglas, G. W., and R. J. Taylor. 1972. The biosystematics,
chemotaxonomy, and ecology of Claytonia lanceolata
Pursh. in western Washington. Canadian Journal of Botany
50: 2177-2187.
Douglas, G. W., D. B. Naas, and R. W. Naas. 1973. New
plant records and ranges for Washington. Northwest Sci-
ence 47: 105-108.
Douglas, G. W., and J. Ramsden. 1973. Representation of
leaf orientation data using equal-area projection techniques.
Canadian Journal of Botany 51: 1081-1088.
Taylor, R. J., G. W. Douglas, and L. M. Sunquist. 1973.
Contributions to the flora of Washington: IT. Northwest Sci-
ence 47: 169-179.
Douglas, G. W. 1974. Lichens of the North Cascades Range,
Washington. The Bryologist 77: 582-592.
Douglas, G. W. 1974. Montane zone vegetation of the Alsek
River region, southwestern Yukon. Canadian Journal of
Botany 52: 2505-2532.
Taylor, R. J., and G. W. Douglas. 1975. Mountain wildflow-
ers of the Pacific Northwest. Binford and Mort, Portland,
Oregon. 176 pages.
Douglas, G. W., and D. H. Vitt. 1976. Moss-lichen flora of
St. Elias-Kluane Ranges, southwestern Yukon. The Bry-
ologist 79: 437-456.
Douglas, G. W., K. E. Denford, and I. Karas. 1977. A con-
tribution to the taxonomy of Antennaria alpina var. media,
A. microphylla, and A. umbrinella in western North Amer-
ica. Canadian Journal of Botany 55: 925-933.
Douglas, G. W., and L. C. Bliss. 1977. Alpine and high sub-
alpine plant communities of the North Cascades Range,
Washington and British Columbia. Ecological Monographs
47: 113-150.
Douglas, G. W., and G. Ruyle-Douglas. 1978. Contributions
to the floras of British Columbia and the Yukon Territory
I. Vascular Plants. Canadian Journal of Botany 56: 2296-
2302.
Douglas, G. W., and G. Ruyle-Douglas. 1978. Nomenclatur-
al changes in the Asteraceae of British Columbia. I. Sene-
cioneae. Canadian Journal of Botany 56: 1710-1711.
Taylor, R. J., D. B. Naas, R. B. Naas, and G. W. Douglas.
1978. Contributions to the flora of Washington. III. North-
west Science 52: 220-225.
Taylor, R. J., and G. W. Douglas. 1978. Plant ecology and
natural history of Chowder Ridge, Mt. Baker: A potential
alpine research natural area in the western North Cascades.
Northwest Science 52: 35-50.
Douglas, G. W. 1980a. The flora of the Haines Road region,
northwestern British Columbia. British Columbia Provin-
cial Museum Occasional Paper.
Douglas, G. W. 1980b. Vegetation. Jn Biophysical inventory
studies of Kluane National Park. Parks Canada, Winnipeg.
The flora of the Haines Road region, Northwestern British
Columbia. British Columbia Provincial Museum Occa-
sional Paper.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Douglas, G. W., and W. L. Peterson. 1980. Contributions
to the floras of British Columbia and the Yukon Territory.
II. Mosses and Lichens. Canadian Journal of Botany 58:
2145-2147.
Murray, D. F., and G. W. Douglas. 1980. The Green Man-
tle. Pages 52-63 in Kluane, Pinnacle of the Yukon. Edited
by J. Theberge and G. W. Douglas. Doubleday Canada
Ltd., Toronto, Ontario. 175 pages.
Theberge, J., and G. W. Douglas. 1980. Kluane, Pinnacle of
the Yukon. Doubleday Canada Ltd. Toronto, Ontario. 175
pages.
Douglas, G. W., W. L. Peterson, and A. C. Skorepa. 1981.
A preliminary checklist of mosses and lichens in the Fort
McMurray area, Alberta. Canadian Journal of Botany 59:
1456-1464.
Douglas, G. W., G. W. Argus, L. Dickenson, and D. Brun-
ton. 1981. Rare vascular plants of the Yukon. National
Museum of Canada Syllogeus 28. 61 pages.
Douglas, G. W., and M. J. Ratcliffe. 1981. Some rare plant
collections, including three new records for Canada, from
Cathedral Provincial Park, southern British Columbia.
Canadian Journal of Botany 59: 1537-1538.
Douglas, G. W. 1982. The sunflower family (Asteraceae) of
British Columbia. Volume I. Senecioneae. British Columbia
Provincial Museum Occasional Paper Series 23. 180 pages.
Douglas, G. W., A. Ceska, and G. G. Ruyle. 1983. A floris-
tic bibliography for British Columbia. Land Management
Report Number 15. British Columbia Ministry of Forests,
Victoria, British Columbia. 143 pages.
Straley, G. B., R. L. Taylor, and G. W. Douglas. 1985. Rare
Vascular Plants of British Columbia. Canada National
Museum Syllogeus 59. 165 pages.
Douglas, G. W. 1986. Nomenclatural changes in the Aster-
aceae of British Columbia. II Astereae, Eupatoreae, Inuleae,
Lactuceae and Heliantheae. Canadian Journal of Botany
64: 2726-2727.
Douglas, G. W., and J. G. Packer. 1988. Erigeron salishii, a
new Asteraceae for British Columbia. Canadian Journal
of Botany 66: 414-416.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1989. The
vascular plants of British Columbia. Part 1 - Gymnosperms
and Dicotyledons (Aceraceae through Cucurbitaceae). Spe-
cial Report Series Number 1. British Columbia Ministry of
Forests, Victoria, British Columbia. 208 pages.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1990. The
Vascular Plants of British Columbia. Part 2 — Dicotyledons
(Diapensiaceae through Portulaceae). Special Report Series
Number 2. British Columbia Ministry of Forests. Victoria,
British Columbia. 155 pages.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1991. The
Vascular Plants of British Columbia. Part 3 — Dicotyledons
(Primulaceae through Zygophyllaceae) and Pteridophytes.
Special Report Series Number 3. British Columbia Min-
istry of Forests, Victoria, British Columbia. 177 pages.
Douglas, G. W. 1992. Composites, Ferns and Allies. Pages
95-120 in Plants of Northern British Columbia. Edited by
A. Mackinnon, J. Pojar and R. Coupé. Lone Pine Publish-
ing, Edmonton, Alberta. 351 pages.
Douglas, G. W. 1993. Tracking rare British Columbian vascu-
lar plants with a computerized data bank. Iiamna 2(2): 1.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1994. The
Vascular Plants of British Columbia. Part 4 — Monocotyle-
dons. British Columbia Ministry of Forests, Victoria, Bri-
tish Columbia. 255 pages.
2006
Douglas, G. W., J. Pojar, D. Meidinger, and K. McKeown.
1994. Rare vascular plant collections from the St. Elias
Mountains, northwestern British Columbia. Canadian
Field-Naturalist 108(4): 391-396.
Straley, G. B., and G. W. Douglas. 1994. Rare and endan-
gered vascular plants — an update. Pages 95-112 in Bio-
diversity in British Columbia Our Changing Environment
Edited by L. E. Harding and E. McCullum. Environment
Canada, Canadian Wildlife Service, Pacific and Yukon
Region. Vancouver, British Columbia. 425 pages.
Douglas G. W. 1995. The sunflower family (Asteraceae) of
British Columbia. Volume II. Astereae, Anthemdieae, Eupa-
toreae and Inuleae. Royal British Columbia Museum, Vic-
toria. 393 pages.
Taylor, R. J., and G. W. Douglas. 1995. Mountain plants of
the Pacific Northwest: a field guide to Washington, west-
ern British Columbia and southeastern Alaska. Mountain
Press Publishing Company, Missoula, Montana. 437 pages.
Douglas, G. W. 1996. Asteraceae: sunflower family. Pages
106-145 in Plants of the Southern Interior of British Colum-
bia. Edited by R. Parish, R. Coupé and D. Lloyd. Lone Pine
Publishing, Edmonton, Alberta. 463 pages.
Douglas, G. W. 1996. Endemic vascular plants of British
Columbia and immediately adjacent regions. Canadian
Field-Naturalist 110(3): 387-391.
Douglas, G. W., and J. M. Illingworth. 1997. Status of the
white-top aster, Aster curtus (Asteraceae), in Canada. Cana-
dian Field-Naturalist 111(4): 622-627.
Douglas, G. W., F. Lomer, and H. L. Roemer. 1998. New
or rediscovered native vascular plant species in British
Columbia. Canadian Field-Naturalist 1 12(2): 276-279.
Douglas, G. W., G. B. Straley, and D. V. Meidinger. 1998.
Rare Native Vascular Plants of British Columbia. British
Columbia Ministry of Environment, Lands and Parks, Vic-
toria, British Columbia. 423 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998. Illustrated Flora of British Columbia. Volume 1:
Gymnosperms and Dicotyledons (Aceraceae through
Asteraceae). British Columbia Ministry of Environment,
Lands and Parks and British Columbia Ministry of Forests,
Victoria, British Columbia. 436 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998. Illustrated Flora of British Columbia. Volume 2:
Dicotyledons (Balsaminaceae through Cucurbitaceae).
British Columbia Ministry of Environment, Lands and
Parks and British Columbia Ministry of Forests, Victoria,
British Columbia. 401 pages.
Douglas, G. W., and J. M. Illingworth. 1998. Status of the
Water-plantain Buttercup, Ranunculus alismifolius var.
alismifolius (Ranunculaceae), in Canada. Canadian Field-
Naturalist 112(2): 280-283.
Douglas, G. W., and M. Ryan. 1998. Status of the Yellow
Montane Violet, Viola praemorsa ssp. praemorsa (Vio-
laceae) in Canada. Canadian Field-Naturalist 112(3): 491-
495.
Jamison, J. A., and G. W. Douglas. 1998. Status of the
Coastal Wood Fern, Dryopteris arguta (Dryopteridaceae),
in Canada. Canadian Field-Naturalist 112(2): 284-288.
Penny, J. L., G. W. Douglas, and G. A. Allen. 1998. Status of
the Bearded Owl-clover, Triphysaria versicolor ssp. ver-
sicolor (Scrophulariaceae), in Canada. Canadian Field-
Naturalist 112(3): 481-485.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illustrat-
ed Flora of British Columbia. Volume 3: Dicotyledons
(Diapensiaceae through Onagraceae). British Columbia
PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS 14]
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia
401 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illus
trated Flora of British Columbia. Volume 4: Dicotyledons
(Orobanchaceae through Rubiaceae). British Columbia
Ministry of Environment, Lands and Parks and British Col-
umbia Ministry of Forests. Victoria, British Columbia
401 pages.
Douglas, G. W., and M. Ryan. 1999. Status of the Golden
Paintbrush, Castilleja levisecta (Scrophulariaceae), in Can-
ada. Canadian Field-Naturalist 113(2): 299-301.
Lomer, F., and G. W. Douglas. 1999. Additions to the flora
of the Queen Charlotte Islands, British Columbia. Cana-
dian Field-Naturalist 113(2): 235-240.
Miller, M. T., and G. W. Douglas. 1999. Status of Lyall’s
Mariposa Lily, Calochortus lyalli (Liliaceae), in Canada.
Canadian Field-Naturalist 1 13(4): 652-658.
Penny, J. L., and G. W. Douglas. 1999. Status of the Tall
Bugbane, Cimicifuga elata (Ranunculaceae), in Canada.
Canadian Field-Naturalist 113(3): 461-465.
Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illustrat-
ed Flora of British Columbia. Volume 5: Dicotyledons
(Salicaceae through Zygophyllaceae and Pteridophytes).
British Columbia Ministry of Environment, Lands and
Parks and British Columbia Ministry of Forests, Victoria,
British Columbia. 389 pages.
Douglas, G. W., and J. A. Jamison. 2001. Status of Scouler’s
Corydalis, Corydalis scouleri (Fumariaceae), in Canada.
Canadian Field-Naturalist 1 15(3): 455-459.
Douglas, G. W. 2001. Abronia umbellata ssp. acutalata:
Rarest plant on the planet or the rarest plant in Canada?
Menziesia 6: 4-5.
Donovan, M. T., and G. W. Douglas. 2001. Status of snake-
root, Sanicula arctopoides (Apiaceae), in Canada. Canadian
Field-Naturalist 115(3): 466-471.
Douglas G. W., D. Meidinger, and J. Pojar. 2001. Illustrated
Flora of British Columbia. Volume 6: Monocotyledons
(Acoraceae to Najadaceae). British Columbia Ministry of
Environment, Lands and Parks, British Columbia Min-
istry of Forests, Victoria, British Columbia. 361 pages.
Douglas G. W., D. Meidinger, and J. Pojar. 2001. Illustrated
Flora of British Columbia. Volume 7: Monocotyledons
(Orchidaceae to Zosteraceae). Ministry of Sustainable
Resource Management and British Columbia Ministry of
Forests. Victoria, British Columbia. 379 pages.
Douglas, G. W., and J. Pojar. 2001. Trillium ovatum Pursh
variety hibbersonii (Taylor et Szczawinski) Douglas ef
Pojar, variety nova. Canadian Field-Naturalist 115(2): 343.
Douglas, G. W., and M.W. Ryan. 2001. Status of Deltoid Bal-
samroot, Balsamorhiza deltoidea (Asteraceae), in Canada.
Canadian Field-Naturalist 115(3): 451-454.
Penny, J. L., and G. W. Douglas. 2001. Status of Purple
Sanicle, Sanicula bipinnatifida (Apiaceae), in Canada.
Canadian Field-Naturalist 1 15(3): 460-465.
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare
Native Vascular Plants of British Columbia. Second edi-
tion. Province of British Columbia, Victoria, British Colum-
bia. 359 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2002. Ilustrat-
ed Flora of British Columbia. Volume 8: General Sum-
mary, Maps and Keys. British Columbia Ministry of Sus-
tainable Resource Management and British Columbia
Ministry of Forests. Victoria, British Columbia. 457 pages.
142
Douglas, G. W. 2005. Conservation evaluation of Lemmon’s
Holly Fern, Polystichum lemmonii (Dryopteridaceae), a
Threatened fern in Canada. Canadian Field-Naturalist
118(2): 164-168.
Douglas, G. W., and J. L. Penny. 2005. Conservation eval-
uation of Howell’s Triteleia, Triteleia howellii (Liliaceae),
an Endangered lily in Canada. Canadian Field-Naturalist
118(2): 174-178.
Douglas, G. W., and J. M. Illingworth. 2005. Conservation
evaluation of the Pacific population of Tall Woolly-heads,
Psilocarphus elatior (Asteraceae), an Endangered herb in
Canada. Canadian Field-Naturalist 118(2): 169-173.
Smith, T., G. W. Douglas, and A. Harris. 2005. Conserva-
tion evaluation of Small-flowered Lipocarpha, Lipocarpha
micrantha (Cyperaceae), in Canada. Canadian Field-Nat-
uralist 118(2): 179-184.
Douglas, G. W., and J. Desrosiers. 2006. Conservation eval-
uation of Pacific Rhododendron (Rhododendron macro-
phyllum), in Canada. Canadian Field-Naturalist 120(2):
169-174.
Douglas, G. W., and J. L. Penny. 2006. Conservation eval-
uation of Slender Collomia, Collomia tenella, in Canada.
Canadian Field-Naturalist 120(2): 175-178.
Douglas, G. W., and Jenifer L. Penny. 2006. Conservation
evaluation of the Small-flowered Tonella, Tonella tenella,
in Canada. Canadian Field-Naturalist 120(2): 179-182.
Douglas, G. W., J. L. Penny, and K. Barton. 2006. Conser-
vation evaluation of the Dwarf Woolly-heads, Psilocarphus
brevissimus var. brevissimus, in Canada. Canadian Field-
Naturalist 120(2): 163-168.
Douglas, G. W., J. L. Penny, and K. Barton. 2006. Conser-
vation evaluation of the Stoloniferous Pussytoes, Antennar-
ia flagellaris, in Canada. Canadian Field-Naturalist 120(2):
183-187.
Douglas, G. W., and M. W. Ryan. 2006. Conservation eval-
uation of Prairie Lupine, Lupinus lepidus var. lepidus, in
Canada. Canadian Field-Naturalist 120(2): 147-152.
Douglas, G. W., and M. W. Ryan. 2006. Conservation eval-
uation of the Seaside Bird’s-foot Trefoil, Lotus formosis-
simus, in Canada. Canadian Field-Naturalist 120(2): 153-
156.
Douglas, G. W., and M. J. Oldham. Conservation evaluation
of Scarlet Ammannia, Ammania robusta, in Canada. Cana-
dian Field-Naturalist (submitted).
Douglas, G. W., and M. J. Oldham. Conservation evaluation
of Toothcup, Rotala ramosior, in Canada. Canadian Field-
Naturalist (submitted).
Unpublished Reports
Douglas, G. W. 1971. An ecological survey of potential nat-
ural areas in the North Cascades National Park Complex.
Unpublished report submitted to the Intercampus Educa-
tional and Scientific Preserves Committee. 137 pages.
Douglas, G. W. 1974. A reconnaissance survey of the vegeta-
tion of Kluane National Park. Report submitted to Cana-
dian Wildlife Service, Edmonton, Alberta. 219 pages.
Douglas, G. W. 1974. Ecological impact of chemical fire
retardants: a review. Report submitted to Northern Forestry
Research Centre, Environment Canada, Edmonton, Alber-
ta. NOR-X-109. 33 pages.
Douglas, G. W., and L. J. Knapik. 1974. Montane zone soil
characteristics in Kluane National Park. Report submitted
to Canadian Wildlife Service, Edmonton, Alberta.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Blood, D. A., G. W. Douglas, and L. J. Knapik. 1975.
Soil, vegetation and wildlife resources of five potential
transportation corridors in Kluane National Park, Yukon.
Report submitted to Donald A. Blood & Associates Ltd.,
Nanaimo, British Columbia. 460 pages.
Douglas, G. W. 1975. An evaluation of the examination and
selection of ecological reserves in Priority Area one, west-
central British Columbia. Report submitted to British Col-
umbia Ministry of Forests, Victoria, British Columbia.
14 pages.
Douglas, G. W. 1975. Spruce (Picea) hybridization in west-
central British Columbia. Report submitted to British
Columbia Ministry of Forests, Victoria, British Columbia.
15 pages.
Douglas, G. W., J. A. Nagy, and G. W. Scotter. 1975. Effects
of human and horse trampling on natural vegetation, Wat-
erton Lakes National Park. Report submitted to Canadian
Wildlife Service, Edmonton, Alberta. 135 pages.
Douglas, G. W. 1977. Vegetation. Jn Environmental impact
statement, Shakwak Highway improvement, British Colum-
bia and Yukon, Canada. Two volumes. Report submitted to
Department of Public Works Canada, and United States
Department of Transportation.
Douglas, G. W. 1978. The status of some rare plants in the
Haines Junction area, Yukon Territory. Report submitted to
Department of Public Works, Canada, Whitehorse, Yukon
Territory. 10 pages.
Douglas, G. W. 1979. An initial environmental impact assess-
ment of the vegetation and flora along the Watson Lake-
Haines Junction section of the Alaska Highway. Report
submitted to Department of Public Works, Canada, Van-
couver, British Columbia. 34 pages.
Douglas, G. W. 1979. Vegetation. In Shakwak Highway proj-
ect reclamation program formulation. I. Technical report of
soil and plant community types and reclamation materials
and methods suitable for use in segments one to eight.
Report submitted to Department of Public Works, Canada,
Whitehorse, Yukon Territory.
Douglas, G. W. 1980. Alpine plant communities of British
Columbia and their occurrence in existing or proposed Eco-
logical Reserves. Report submitted to Ecological Reserves
Unit, Ministry of the Environment, Victoria, British Colum-
bia.
Douglas, G. W. 1980. Vegetation. Jn Biophysical inventory
studies of Kluane National Park. Report submitted to Parks
Canada, Winnipeg, Manitoba.
Blood, D.A., E. A. Harding, M. E. Walmsley, and G. W.
Douglas. 1987. A preliminary assessment of resources in
the Strathcona Recreation Area. Report submitted to British
Columbia Parks and Outdoor Recreation Division, North
Vancouver, British Columbia.
Lea, E. C., and G. W. Douglas. 1991. Endangered and
Threatened Plant species of the southern interior of Bri-
tish Columbia — Protecting the insignificant. Pages 59-66
in Community Action for Endangered Species — A public
symposium on British Columbia Threatened and Endan-
gered Species and Their Habitats. Edited by S. Rautio.
Federation of British Columbia Naturalists and Northwest
Wildlife Preservation Society, Vancouver, British Colum-
bia. 238 pages.
Douglas, G. W. 1995. Status Report on the Rabbit-brush
goldenweed (Ericameria bloomeri [A. Gray] J. F. Mc-
Bride). Unpublished report submitted to the Committee
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 10 pages.
2006
Illingworth, J. M., and G. W. Douglas. 1995a. Status report
on the slender woolly-heads, Psilocarphus tenellus Nutt.
var. tenellus. Unpublished report submitted to Committee
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 19 pages.
Illingworth, J. M., and G. W. Douglas. 1995b. Status report
on the tall woolly-heads, Psilocarphus elatior (A. Gray)
A. Gray. Unpublished report submitted to Committee on
the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 19 pages.
Ryan, M., and G. W. Douglas. 1995a. Status report on the
golden paintbrush, Castilleja levisecta, in Canada. Unpub-
lished report submitted to Committee on the Status of En-
dangered Wildlife in Canada, Ottawa, Ontario. 26 pages.
Ryan, M., and G. W. Douglas. 1995b. Status report on the
yellow montane violet, Viola praemorsa var. praemorsa,
in Canada. Unpublished report submitted to Committee
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 22 pages.
Desrosiers, J., and G. W. Douglas. 1997. Status Report on
Pacific rhododendron, Rhododendron macrophyllum D.
Don ex Don (Ericaceae), in Canada. Committee on the Sta-
tus of Endangered Wildlife in Canada, Ottawa, Ontario.
22 pages.
Douglas, G. W., and J. M. Illingworth. 1996. Status report
on the white-top aster, Aster curtus Cronq. In Canada. Un-
published report submitted to Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 26 pages.
Illingworth, J. M., and G. W. Douglas. 1996. Status report
on the water-plantain buttercup, Ranunculus alismaefolius
Geyer ex Benth. var. alismaefolius. Unpublished report
submitted to Committee on the Status of Endangered Wild-
life in Canada, Ottawa, Ontario. 24 pages.
Ryan, M. W., and G. W. Douglas. 1996a. Status report on
the deltoid balsamroot, Balsamorhiza deltoidea Nutt. in
Canada. Unpublished report submitted to Committee on
the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 30 pages.
Ryan, M., and G. W. Douglas. 1996b. Status report on the
seaside birds-foot lotus, Lotus formosissimus in Canada.
Unpublished report submitted to Committee on the Status
of Endangered Wildlife in Canada, Ottawa, Ontario. 25
pages.
Ryan, M., and G. W. Douglas. 1996c. Status report on the
prairie lupine, Lupinus lepidus var. lepidus, in Canada. Un-
published report submitted to Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 26 pages.
Penny, J. L., and G. W. Douglas. 1997. Status report on
bearded owl-clover, Triphysaria versicolor ssp. versicolor,
in Canada. Unpublished report submitted to the Committee
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 29 pages.
Jamison, J. A., and G. W. Douglas. 1998. Status report on
coastal wood fern, Dryopteris arguta, in Canada. Unpub-
lished report submitted to the Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 23 pages.
White, D. J., and G. W. Douglas. 1998. Update COSEWIC
status report on giant helleborine orchid (Epipactis gigan-
tea). Unpublished report submitted to Committee on the
Status of Endangered Wildlife in Canada, Ottawa, Ontario.
15 pages.
White, D. J., and G. W. Douglas. 1998. Update COSEWIC
status report on Southern Maidenhair Fern (Adiantum capil-
lus-veneris). Unpublished report submitted to Committee
PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS 143
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 14 pages.
Douglas, G. W., and M. J. Oldham. 1999. Status report on
scarlet ammannia, Ammannia robusta, in Canada. Unpub
lished report submitted to the Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 26 pages
Douglas, G. W., and M. J. Oldham. 1999. Status report on
toothcup, Rotala ramosior, in Canada. Unpublished report
submitted to the Committee on the Status of Endangered
Wildlife in Canada, Ottawa, Ontario. 25 pages.
Fontaine, M., and G. W. Douglas. 1999. Rare Grassland
Plants. Species and Ecosystems at Risk in British Colum-
bia series. British Columbia Ministry of Environment,
Lands and Parks, Victoria, British Columbia. 4 pages.
Fontaine, M., and G. W. Douglas. 1999. Rare Forest Plants
of Southern British Columbia. Species and Ecosystems
at Risk in British Columbia series. British Columbia Min-
istry of Environment, Lands and Parks, Victoria, British
Columbia. 4 pages.
Donovan, M., and G. W. Douglas. 2000. Status report on
snake-root sanicle, Sanicula arctopoides, in Canada. Un-
published report submitted to the Committee on the Sta-
tus of Endangered Wildlife in Canada, Ottawa, Ontario.
20 pages.
Douglas, G. W., and J. A. Jamison. 2000. Status report on
Scouler’s corydalis, Corydalis scouleri, in Canada. Unpub-
lished report submitted to the Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 18 pages.
Douglas, G. W., and J. L. Penny. 2000. New vascular plant
species to the British Columbia flora. Menziesia 5(4): 5.
Douglas. G. W., J. L. Penny, and M. Donovan. 2000. Sta-
tus reports on two Sanicula species. Menziesia 5(4): 4-S.
Miller, M., and G. W. Douglas. 2000. Status report on Lyall’s
mariposa lily, Calochortus lyallii, in Canada. Unpublished
report submitted to the Committee on the Status of En-
dangered Wildlife in Canada, Ottawa, Ontario. 32 pages.
Penny, J. L., and G. W. Douglas. 2000. Status report on
purple sanicle, Sanicula bipinnatifida, in Canada. Unpub-
lished report submitted to the Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 21 pages.
Penny, J. L., and G. W. Douglas. 2000. Status report on tall
bugbane, Cimicifuga elata, in Canada. Unpublished report
submitted to the Committee on the Status of Endangered
Wildlife in Canada, Ottawa, Ontario. 22 pages.
Penny, J. L., and G. W. Douglas. 2000. Tracking rare native
vascular plants in British Columbia. Pages 75-77 in Pro-
ceedings of a conference on the biology and manage-
ment of species and habitats at risk. Edited by L. M. Dar-
ling. British Columbia Ministry of Environment, Lands
and Parks, Victoria, British Columbia.
Douglas, G. W., J. L. Penny, and R. E. Maxwell. 2001.
Composition, phenology, stand structure and soils of a
Quercus garryana (Garry Oak) woodland community at
Cowichan Garry Oaks Preserve, Quamichan Lake, Van-
couver Island, British Columbia, Nature Conservancy of
Canada. Victoria, British Columbia. 53 pages.
Smith, T. W., G. W. Douglas, and A. Harris. 2001. Update
COSEWIC Status Report on Small-Flowered Lipocarpha,
Lipocarpha micrantha, in Canada. Unpublished report
submitted to the Committee on the Status of Endangered
Wildlife in Canada, Ottawa, Ontario. 23 pages.
Douglas, G. W. 2003. COSEWIC Status Report on Baikal
Sedge, Carex sabulosa, in Canada. Unpublished report sub-
mitted to Committee on the Status of Endangered Wildlife
in Canada, Ottawa, Ontario. 22 pages.
144
Douglas, G. W. 2003. COSEWIC Status Report on Lem-
mon’s holly fern, Polystichum lemmonii, in Canada. In
COSEWIC Assessment and Status Report on Lemmon’s
Holly Fern, Polystichum lemmonii, in Canada. Committee
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. 15 pages (Available online at: http://www.sara
registry.gc.ca/status/status_e.cfm).
Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status
report on Howell’s Triteleia, Triteleia howellii, in Canada.
In COSEWIC Assessment and Status report on Howell’s
Triteleia, Triteleia howellii, in Canada. Committee on the
Status of Endangered Wildlife in Canada, Ottawa, Ontario.
21 pages (Available online at: http://www.sararegistry.gc.
ca/status/status_e.cfm).
Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status
report on Slender Collomia, Collomia tenella, in Canada.
In COSEWIC Assessment and Status report on Slender
Collomia, Collomia tenella, in Canada. Committee on the
Status of Endangered Wildlife in Canada, Ottawa, Ontario.
15 pages (Available online at: http://www.sararegistry.gc.
ca/status/status_e.cfm).
Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status
report on the Small-flowered Tonella, Tonella tenella, in
Canada. In COSEWIC Assessment and Status report on
the Small-flowered Tonella, Tonella tenella, in Canada.
Committee on the Status of Endangered Wildlife in Cana-
da, Ottawa, Ontario. 16 pages (Available online at: http://
Wwww-.Ssararegistry.gc.ca/status/status_e.cfm).
Douglas, G. W., J. L. Penny, and K. Barton. 2003.
COSEWIC Status Report on Dwarf Woolly-heads, Psilocar-
phus brevissimus var. brevissimus, in Canada. In COSEWIC
Status Assessment and Report on Dwarf Woolly-heads,
Psilocarphus brevissimus var. brevissimus, in Canada. Com-
mittee on the Status of Endangered Wildlife in Canada,
Ottawa, Ontario. 21 pages (Available online at: http://www.
sararegistry.gc.ca/status/status_e.cfm).
Douglas, G. W., J. L. Penny, and K. Barton. 2003.
COSEWIC Status Report on Stoloniferous Pussytoes,
Antennaria flagellaris, in Canada. In COSEWIC Assess-
ment and Status Report on Stoloniferous Pussytoes,
Antennaria flagellaris, in Canada. Committee on the Sta-
tus of Endangered Wildlife in Canada Ottawa, Ontario.
19 pages (Available online at: http://www.sararegistry.gc.
ca/status/status_e.cfm).
Douglas, G. W., J. Pojar, P. Sheppard, and S. Walker.
2003. Gwaii Haanas Rare Vascular Plant Survey — Field
Report. Unpublished report submitted to Parks Canada,
Pacific and Yukon Region, Vancouver, British Columbia.
1 page.
Douglas, G. W., and S. J. Smith. 2003. Stewardship account
for foxtail muhly (Muhlenbergia andina). Unpublished
report submitted to Ministry of Water, Land and Air Pro-
tection, Biodiversity Branch. 17 pages.
Douglas, G. W., and S. J. Smith. 2003. Stewardship account
for pale bulrush (Scirpus pallidus). Unpublished report
submitted to Ministry of Water, Land and Air Protection,
Biodiversity Branch. 18 pages.
Douglas, G. W., and S. J. Smith. 2003. Stewardship account
for Southern Maidenhair-fern (Adiantum capillus-vener-
is). Unpublished report submitted to Ministry of Water,
Land and Air Protection, Biodiversity Branch. 18 pages.
Brownell, V. R., and G. W. Douglas. 2004. Draft National
Recovery Strategy for Toothcup, Rotala ramosior (L.)
Koehne. Unpublished report submitted to Recovery of Na-
tionally Endangered Wildlife (RENEW), Ottawa, Ontario.
39 pages.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Douglas, G. W. 2004. A Survey of the Rare Native Vascular
Plants of the Southern Okanagan Grasslands Protected
Area. Unpublished report submitted to Parks Canada,
Pacific and Yukon Region, Vancouver, British Columbia.
7 pages.
Douglas, G. W. 2004. COSEWIC status report on the pink
sand-verbena, Abronia unbellata in Canada. In COSEWIC
assessment and status report on the pink sand-verbena,
Abronia umbellata, in Canada. Committee on the Status of
Endangered Wildlife in Canada, Ottawa, Ontario. 19 pages.
Douglas, G. W. 2004. Draft National Recovery Strategy for
Lemmon’s Holly Fern (Polystichum lemmonii). Unpub-
lished report submitted to British Columbia Ministry of
Water, Land, and Air Protection, Biodiversity Branch, Vic-
toria, British Columbia. 15 pages.
Douglas, G. W., and S. J. Smith. 2004. Draft COSEWIC
Status Report on California Hedge-parsley, Yabea micro-
carpa (Apiaceae) in Canada. Unpublished report submit-
ted to British Columbia Ministry of Water, Lands and Air
Protection, Biodiversity Branch, Victoria, British Colum-
bia. 21 pages.
Douglas, G. W., and S. J. Smith. 2004. Draft COSEWIC
Status Report on Lindley’s Microseris, Microseris lind-
leyi (Asteraceae), in Canada. Unpublished report submit-
ted to British Columbia Ministry of Water, Land, and Air
Protection, Biodiversity Branch, Victoria, British Colum-
bia. 24 pages.
Douglas, G. W., and S. J. Smith. 2004. Draft COSEWIC
Status Report on Slender Popcorn Flower, Plagiobothrys
tenellus (Boraginaceae) in Canada. Unpublished report
submitted to British Columbia Ministry of Water, Land,
and Air Protection, Biodiversity Branch, Victoria, British
Columbia. 22 pages.
Douglas, G. W., and S. J. Smith. 2004. Draft Update
COSEWIC Status Report on Scouler’s Corydalis, Cory-
dalis scouleri (Fumariaceae), in Canada. Unpublished report
submitted to British Columbia Ministry of Water, Land,
and Air Protection, Biodiversity Branch, Victoria, British
Columbia. 18 pages.
Douglas, G. W., and S. J. Smith. 2004. Draft National Multi-
species Recovery Strategy for plants at risk in Garry Oak
woodlands. Unpublished report submitted to British Col-
umbia Ministry of Water, Land, and Air Protection, Bio-
diversity Branch, Victoria, British Columbia. 10 pages.
Douglas, G. W., and S. J. Smith. 2004. Draft National
Recovery Strategy for Scarlet Ammannia (Ammannia
robusta Heer & Regel). Unpublished report submitted to
British Columbia Ministry of Water, Land, and Air Pro-
tection, Biodiversity Branch, Victoria, British Columbia.
23 pages.
Douglas, G. W., and S. J. Smith. 2004. Rare Native Vascu-
lar Plants of the Southern Okanagan Grasslands Protect-
ed Area Field Report. Unpublished report submitted to
The Nature Trust of British Columbia, North Vancouver,
British Columbia. 10 pages.
Harris, A. G., R. F. Foster, G. W. Douglas, and S. J. Smith.
2004. National Recovery Strategy for Small-Flowered
Lipocarpha (Lipocarpha micrantha) in Canada. Unpub-
lished report submitted to the Ontario Ministry of Natural
Resources, Northwest Region, Thunder Bay, Ontario, and
British Columbia Ministry of Water, Land, and Air Pro-
tection, Biodiversity Branch, Victoria, British Columbia.
25 pages.
*COSEWIC. 2005a. COSEWIC assessment and status report
on the Branched Phacelia Phacelia ramosissima in Can-
ada. Committee on the Status of Endangered Wildlife in
-
2006
Canada, Ottawa, Ontario. vi + 18 pages. (Available online
at: http://www.sararegistry.gc.ca/status/status_e.cfm).
*COSEWIC. 2005b. COSEWIC assessment and status report
on the Cliff Paintbrush Castilleja rupicola in Canada.
Committee on the Status of Endangered Wildlife in Cana-
da, Ottawa, Ontario. vii + 18 pages. (Available online at:
http://www.sararegistry.gc.ca/status/status_e.cfm).
*COSEWIC. 2005c. COSEWIC assessment and status report
on the Grand Coulee Owl-clover Orthocarpus barbatus in
Canada. Committee on the Status of Endangered Wildlife
in Canada, Ottawa, Ontario. vi + 17 pages. (Available on-
line at: http://www.sararegistry.gc.ca/status/status_e.cfm).
“COSEWIC. 2005d. COSEWIC Assessment and Status
report on the Mountain Holly Fern, Polystichum scopuli-
num (Dryopteridaceae), in Canada. Committee on the
Status of Endangered Wildlife in Canada, Ottawa, Ontario.
vii + 18 pages (Available online at: http://www.sarareg
istry.gc.ca/status/status_e.cfm).
“COSEWIC. 2005e. COSEWIC assessment and status report
on the Spalding’s Campion, Silene spaldingii, in Canada.
Committee on the Status of Endangered Wildlife in Cana-
da, Ottawa, Ontario. vi + 18 pages. (Available online at:
http://www.sararegistry.gc.ca/status/status_e.cfm).
Smith, S., and G. W. Douglas. 2005. Cowichan Garry Oak
Preserve Monitoring Report: Assessing changes to plant
community composition 1999-2004. Douglas Ecological
Consultants Ltd. Unpublished report submitted to the
Nature Conservancy of Canada, Victoria, British Colum-
bia. 22 pages + appendices.
Professional Experience Summary
1966 — 1967: Forester and Consultant with Rayonier, Cana-
da Ltd., Jeune Landing, BC and Mills, and
Potter and Mills, Vancouver, British Columbia,
where he supervised forest inventory, forest
type mapping, and reforestation projects on
northern Vancouver Island, British Columbia.
Ecological Consultant to the Intercampus Edu-
cation and Scientific Preserves Committee
(Washington Universities) at Western Wash-
ington State College, Bellingham, Washington,
hired to examine and quantitatively sample
more than twenty-five potential natural areas
throughout the State of Washington, and rec-
ommend which ones should be protected.
Ecological Consultant to the United States
National Park Service in Sedro Woolley, Wash-
ington, where he initially provided a biological
inventory (vegetation, wildlife, soils and geol-
ogy) of the newly formed North Cascades
National Park, Washington. He also conducted
vegetation analysis of the entire park (18000
km?) and provided advice and research with
respect to vegetation problems.
Ecological Consultant to Canadian Wildlife
Service, Edmonton, Alberta, and Parks Canada,
Winnipeg, Manitoba. Project co-ordinator and
chief ecologist examining vegetation and soils
of Kluane National Park, Yukon Territory, and
their relation to fragility and land use within
the park. This study was also co-ordinated with
small mammal and bird studies.
1968 — 1970:
1969 — 1974:
1973 — 1974:
PENNY: TRIBUTE TO GEORGE WAYNE
1973 — 1974:
1974 — 1975:
1974 — 1975:
1974 — 1975:
1975 — 1977:
1975 — 2004:
1975 — 1979:
1977 — 1978:
DOUGLAS 145
Ecological Consultant to Canadian Forestry Ser-
vice, Edmonton, Alberta, reviewing and evalu
ating the ecological impact and use of chem
ical fire retardants in North America. He also
made recommendations pertaining to the fu-
ture use of chemical fire retardants in Canada.
Ecological Consultant to Canadian Wildlife
Service, Edmonton, Alberta, and Parks Cana-
da, Calgary, Alberta. Project co-ordinator and
chief plant ecologist for a study designed to
assess the environmental impact of recreation-
al use (mainly human and horse trampling) on
different vegetation types in Waterton Lakes
National Park, Alberta.
Ecological Secretariat, Department of Lands
and Forests, Victoria, British Columbia. Chief
plant ecologist on a three-faceted study in
west-central British Columbia. The study in-
cluded a vegetation reconnaissance of a 1500
square mile area, examination of the spruce
hybridization problem in the region, and an
evaluation of the ecological reserve system
and selection in British Columbia.
Ecological Consultant with Don Blood &
Associates Ltd., Lantzville, British Columbia,
for Parks Canada, Winnipeg, Manitoba. Chief
plant ecologist with a biological and geolog-
ical team assessing the environmental impact
of five potential transportation corridors in
Kluane National Park, Yukon Territory.
Ecological Consultant to Syncrude Canada
Ltd., Edmonton, Alberta. This project involved
setting up a continuous biological monitoring
program for air quality in the tar sands refin-
ery area near Fort McMurray, Alberta. Using
lichens as an indicator, the affect of effluents
(especially SO,) and their environmental im-
pacts will be continuously evaluated.
Research Associate in Biological Collections,
Herbarium, Royal British Columbia Museum,
Victoria, British Columbia. Author of a flora
on the Asteraceae (Sunflower family) of Bri-
tish Columbia. This work includes a complete
taxonomical and ecological treatment of ap-
proximately 350 taxa.
Ecological Consultant to Parks Canada, Win-
nipeg, Manitoba. Project co-ordinator and
chief plant ecologist on four-year Biophysical
Inventory Project in Kluane National Park,
Yukon Territory. This multidisciplinary study
included landforms, surficial deposits, vegeta-
tion, and soils. A total of ten professional sci-
entists were involved in this study.
Ecological Consultant to Thurber Consultants
Ltd., Victoria, British Columbia, with Depart-
ment of Public Works Canada, Vancouver,
British Columbia. Chief plant ecologist on
environmental impact study team. This multi-
disciplinary study included all aspects from
engineering and social to environmental im-
pacts with respect to the paving and relocation
of a large section of the Alaska Highway.
“In the production note, COSEWIC acknowledges the late George W. Douglas and Ms. Shyanne J. Smith for writing the status report.
“In the production note, COSEWIC acknowledges the late George W. Douglas and Jacques Labreque for writing the status report.
146
1978 — 1979:
1978 — 1980:
1979 — 1980:
1980:
1981 — 1986:
1985:
1985 — 1986:
1986 — 1995:
1987:
THE CANADIAN FIELD-NATURALIST
Ecological Consultant to Department of Pub-
lic Works, Canada — Whitehorse, Yukon Terri-
tory. Chief plant ecologist on environmental
impact study. Assessed the environmental im-
pact of Alaska Highway relocation and paving
between Watson Lake and Haines Junction.
Ecological Consultant with Hardy and Asso-
ciates Ltd. for Department of Public Works,
Canada in Whitehorse, Yukon Territory. Chief
plant ecologist on revegetation project. This
multidisciplinary study examined vegetation
and soils and recommended reclamation pro-
cedures for a 175 mile segment of the Alaska
Highway.
Ecological Consultant to British Columbia
Ministry of Environment, Land and Parks,
Victoria, British Columbia. Designed and co-
ordinated a biological monitoring network
(using lichens) in northeastern British Colum-
bia.
Ecological Consultant to Ecological Reserves
Unit, British Columbia Ministry of Environ-
ment, Victoria, British Columbia. Provided an
assessment of alpine plant communities in
British Columbia and adjacent regions and
their occurrence in existing or proposed eco-
logical reserves.
Taxonomical Consultant to British Columbia
Ministry of Forests, Victoria, British Colum-
bia. Provided editorial assistance for floristic
manuals and prepared several reports, includ-
ing a floristic bibliography. Identified and
annotated several thousand sheets of British
Columbia plant material.
Taxonomical Consultant to National Museums
of Canada, Ottawa, and World Wildlife Fund.
Prepared detailed status reports on four En-
dangered vascular plants of British Columbia.
Interpretive Consultant to British Columbia
Provincial Parks, Liard Hotsprings, and Mt.
Robson Provincial Parks. Provided interpre-
tive services for Provincial Parks summer
interpretive programs.
Taxonomical Consultant to British Columbia
Ministry of Forests, Victoria, British Colum-
bia. Chief editor and major contributor for the
four-volume Vascular Plants of British Colum-
bia.
Ecological and Taxonomical Consultant for
British Columbia Provincial Parks, Strathcona
Provincial Park, with Donald A. Blood &
Accepted 11 November 2006
1988 — 1989:
1991 — 2002:
1992:
2001 — 2004:
2002:
2002:
2003:
2003:
2003:
2003 — 2004:
2004:
1974 — 2004:
Vol. 120
Associates, Nanaimo, British Columbia. Pro-
vided ecological and floristic expertise with
respect to a preliminary assessment of natu-
ral resources.
Ecological and Taxonomical Consultant to
British Columbia Ministry of Parks, Ecological
Reserves Unit, Victoria, British Columbia.
Chief co-ordinator for study team (including
Blood & Associates and Westland Resources
Group) formulating a new ecological reserve
system for British Columbia.
Program botanist at British Columbia Con-
servation Data Centre, British Columbia Min-
istry of Environment, Lands and Parks, Vic-
toria, British Columbia. Responsible for the
tracking and inventorying vascular plants of
conservation concern in British Columbia.
Botanist, Sierra Club expedition down the Tat-
shenshini River, northern British Columbia;
part of the campaign to have a protected area
established in the region.
Recovery team member and scientific advisor,
Garry Oak Ecosystem Recovery Team Plants
at Risk Recovery Implementation Group.
Recovery team member and scientific advisor,
Adiantum capillus-veneris Recovery Team.
Ecological Consultant to Parks Canada, Pacif-
ic Rim National Park. Chief botanist involved
in inventory and invasive species management.
Botanical consultant to British Columbia Min-
istry of Environment. Chief botanist on inven-
tory projects in South Okanagan Grasslands
Protected Area and adjacent parcels.
Botanical Consultant to British Columbia Min-
istry of Environment. Chief Botanist hired to
inventory for Silene spaldingii to fill informa-
tion gaps in its current status in the province.
Botanical Consultant to Parks Canada. One of
chief botanists hired to inventory rare plants
on the Queen Charlotte Islands.
Botanical Consultant to Osoyoos Indian Band.
Botanist and Project Coordinator managing
invasive species in sensitive habitats on band
lands. Also trained band members in manag-
ing the vegetation.
Botanical Consultant to British Columbia
Ministry of Environment. Chief botanist hired
to inventory Azolla mexicana to fill informa-
tion gaps in its current status in the province.
Principal of Douglas Ecological Consultants
Ltd. Botanist, Ecologist and Project Director.
Conservation Evaluation of the Prairie Lupine, Lupinus lepidus var.
lepidus, in Canada*
GEORGE W. DouGLas! and MICHAEL RYAN2
' Deceased
Present address: 801 Frayne Road, RR # 1, Mill Bay, British Columbia VOR 1LO Canada
Douglas, George W., and Michael Ryan. 2006. Conservation evaluation of the Prairie Lupine, Lupinus lepidus var. lepidus,
in Canada. Canadian Field-Naturalist 120(2); 147-152.
In Canada, Prairie Lupine, Lupinus lepidus var. lepidus, is restricted to southeastern Vancouver Island. Of the nine sites where
it has been collected, five are extirpated and the status of two of the populations is uncertain. There are two extant popula-
tions; some of the other sites may contain the species in the seed bank. Some of the sites are protected to a certain extent
from direct habitat destruction by their remote location, although introduced herbaceous species may pose a serious threat
by preventing the establishment of the species at other sites. Fire suppression or the lack of other types of disturbance also
likely plays a role in discouraging emergence of Lupinus lepidus.
Key Words: Prairie Lupine, Lupinus lepidus, endangered, distribution, population size, British Columbia.
The Prairie Lupine, Lupinus lepidus Dougl. ex Lindl.
var. lepidus* is a member of a genus of over 100 spe-
cies. Most of these occur in North America but the
- genus occurs on all continents except Australia (Hitch-
cock et al. 1961). It is one of 20 species of Lupinus
_ occurring in British Columbia (Pojar 1999) and of about
27 species in Canada (Scoggan 1978). The genus is of
some economic importance in the horticultural trade.
The Lupinus lepidus complex has been a source of
difficulty to researchers over the years. Hitchcock et
al. (1961) state that although the L. /epidus complex is
distinct, the variation within this group has resulted in
a large number of proposed subspecies and varieties
_ of which some are considered to be distinct species by
others. According to Phillips (1955), the nomenclature
| is even more confusing, and the number of synonyms
_ is greater for L. /epidus than for any other Lupinus spe-
| cies. The work of Detling (1951), Phillips (1955) and
Dunn and Gillett (1966) has improved the nomenclature
and delineation of Lupinus species but many problems
remain.
The type specimen of L. lepidus var. lepidus was
collected by David Douglas from “... Fort Vancouver
' to the Great Falls of the Columbia”. Subsequently,
Hitchcock et al. (1961) recognized five varieties of L.
| lepidus two of which, var. lepidus and var. lobbii (A.
\ Gray) C. L. Hitche., are reported to occur in British
Columbia. The latter variety is considered to be a sep-
arate species (L. lyallii A. Gray) in more recent floras
of Canada and British Columbia (eg., Dunn and Gillett
1966; Taylor 1974; Scoggan 1978; Douglas 1990; Pojar
1999) and occurs on well-drained soils in subalpine and
alpine areas (Douglas 1990; Douglas and Bliss 1977).
Lupinus lepidus var. lepidus as recognized by Hitch-
cock et al. (1961), Douglas (1990), Douglas et al.
(1998a) and Pojar (1999), included L. minimus Doug].
in synonomy but they failed to account for the loca-
tion of the type specimen (Rocky Mountains, Kettle
Falls in the upper Columbia River drainage) or the
range of L. minimus in the western Cordillera east of
the Coast-Cascade Mountains. Lupinus minimus is con-
sidered by other authors to be a separate species (e.g.,
Cox 1973a, 1973b; Dunn and Gillett 1966; Scoggan
1978; Taylor 1974). Douglas et al. (2002a, 2002b)
also now treat L. minimus as a distinct entity. The lat-
ter has been reported to occur in the Columbia River
drainage of southern British Columbia (Dunn and
Gillett 1966; Taylor 1974), but no specimens have been
located. It is also reported to occur in the Crowsnest
area of Alberta (Dunn and Gillett 1966) and, more
recently, in and adjacent to Waterton Lakes National
Park (Argus and White 1978; Kuijt 1982) and at six
locations near the southern Alberta border (Packer
1983). Both Kuijt (1982) and Packer (1983) consider
L. minimus to be synonymous with L. lepidus.
Lupinus lepidus is a multi-stemmed perennial herb
20-45 cm tall (Figure 1; Pojar 1999). The stems are
slender with long, soft hairs. The plant has oblanceo-
late palmate compound leaves that are 5 to 9 foliate
and copiously silky-hairy. The leaves are mostly basal
with only | to 4 leaves on the stem. The petioles are
2-5 times as long as the blade. The racemose, pea-like
“The original field work for the Lupinus lepidus project was funded by the British Columbia Conservation Data Centre. The results appear in
the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis
for a Committee on the Status of Endangered Wildlife in Canada status report (Ryan and Douglas 1996*) and the subsequent assessment of
| endangered (COSEWIC 2000’). This paper includes more recent information that did not appear in the original status report.
‘Taxonomy and nomenclature follow Douglas et al. (1998b, 1999, 200 1a, b) and Pojar (1999).
—————————
}
| 147
148 THE CANADIAN FIELD-NATURALIST
blue flowers are similar to those of other Fabaceae
species. The petals are 8-13 mm long, and the banner
is often lighter or darker and reflexed below the mid-
point. The wings are glabrous, the keel ciliate, and the
calyx silky-hairy. Both lips are 6-7 mm long, the upper
ones of which are bifid for at least half their length.
Peduncles are 9-10 cm long, pedicels are 2 mm long
and petals are 8-13 mm long. The hairy pods are 1-2 cm
long and contain three to five large seeds. In British
Columbia, it is separated from three similar looking
species by its distinct range in the coastal lowlands
on southeastern Vancouver Island. It is distinguished
from most other species of Lupinus by its small stature,
the white appressed hairs covering the stems and leaves
and the thick caudex at the base of the plant.
Distribution
Lupinus lepidus extends from southwestern British
Columbia to north-western Oregon, west of the Cas-
cade Mountains (Douglas et al. 1990; Pojar 1999). In
Canada it is rare on southeastern Vancouver Island,
British Columbia (Figure 2; Douglas et al. 2002a; Pojar
IO).
Habitat
Lupinus lepidus inhabits xeric sites ranging from
grass-dominated meadows to steep rocky slopes where
the vegetation is comprised of scattered clumps of
Douglas-fir (Pseudotsuga menziesii), Arbutus (Arbu-
tus menziesii), and Lodgepole Pine (Pinus contorta var.
contorta). These sites are restricted to southeastern
Vancouver Island, where rainfall is low compared to
other coastal areas in British Columbia, and during the
summer, they often experience an extended period of
drought. Associated species include Scotch Broom
(Cytisus scoparius), Hedgehog Dogtail (Cynosurus
echinatus), Sweet Vernalgrass (Anthoxanthum odora-
tum), several species of Brome (Bromus), and Common
Camas (Camassia quamash). The habitat at the Some-
nos site was unnatural, consisting of gravel deposited
during railroad or highway construction.
TABLE |. Populations of Lupinus lepidus in Canada.
Collection Last
Site Observation
Langford Plains (Victoria) 1908
Observatory Hill (Victoria) 1915
Koksilah River Valley (Duncan) 1960
Cattle Point (Victoria) 1973
Beacon Hill (Victoria) 1993
Somenos Lake (Duncan) 1994
Mount Wells (Victoria) 2006
(2 plants/1 m? in 2003)
Mount Braden (Victoria) 2005
Mount McDonald (Victoria) 2005
Vol. 120
Ficure 1. Illustration of Lupinus lepidus. (Line drawing from
Pojar 1999)
Collector/ Population data
Observer (plants/area in m7?)
Macoun Extirpated
Newcombe Extirpated
Hardy Unknown
Brayshaw Extirpated
Brayshaw Extirpated
Douglas Extirpated
Roemer No plants
Roemer 2 plants
Roemer 113 plants/75 m?
2006
Biology
Little is known regarding the biology of Lupinus
lepidus. However, many species of Lupinus share com-
mon traits, some of which likely characterize L. lep-
idus. It is likely to be associated with nitrogen-fixing
Rhizobium, which may allow it to invade sites where
soils are low in nitrogen. It is suspected that L. lep-
idus is a short-lived colonizer of recently disturbed sites
in Xeric, open, exposed areas where it persists for sev-
eral years. It eventually declines, possibly from compe-
tition with more aggressive species that either directly
compete for resources or prevent the establishment of
seedlings, or from lack of vigour in mature plants. The
seeds may not have any dormancy requirements but
like other legumes, L. /epidus seeds have a hard seed
coat which may delay germination for several months
or years. Nothing is known regarding pollination, seed
set, and survival rates.
Population Attributes
Of the nine Canadian sites from which Lupinus lep-
idus has been collected (Table 1, Figure 2), five pop-
ulations are extirpated. Another two sites (Koksilah
River and Mount Wells) are assumed extant but their
status is unknown. Little is known regarding popula-
tion trends for this species. Monitoring this taxon is
difficult because populations may only be comprised
of a few plants and therefore, easily overlooked, or
they may remain in a seed bank for an unknown period
before reappearing. However, four years of data col-
lected at Somenos Lake (extirpated) showed a decline
before it was destroyed at the site: 250, 1,0, and | in
1991, 1992, 1993 and 1994, respectively. Other sites
with limited data include Mount Wells, which con-
tained nine plants in 2001, only a few plants in 2002,
two plants in 2003, and no plants in 2004, 2005 and
2006 (H. Roemer, personal communication). The
steady decline at this site is attributed to the re-growth
of shrubs following a fire. In addition, in 2005, two
plants were observed on Mount Braden (up from sin-
gle plant in 1996) and 113 plants were observed on
Mount McDonald (where they had only previous been
known from 1915). In 2006, a single plant was seen on
Mount Braden and 54 plants on Mount McDonald,
but a thorough search of all the locations from 2005
was not done and therefore, the population should not
be considered in decline.
Provincial, National, and Global Ranks
The British Columbia Conservation Data Centre
has ranked Lupinus lepidus as S1 and placed it on the
British Columbia Ministry of Environment Red-list
(Douglas et al. 2002a). This is the most critical cate-
gory for imperiled rare native vascular plants in British
Columbia. A rank of S1 is considered “critically imper-
iled because of extreme rarity (5 or fewer occurrences
or very few remaining individuals) or because of some
factors making it especially vulnerable to extirpation or
DOUGLAS AND RYAN: PRAIRIE LUPINE
149
FIGURE 2. The status and location of Lupinus lepidus in British
Columbia. @ recently confirmed; ® present status unknown:
O extirpated
extinction” (Douglas et al. 2002a). Lupinus lepidus is
on the Canadian Species At Risk Act legal list and the
Committee on the Status of Endangered Wildlife in
Canada has also assigned this species to the Endangered
category (COSEWIC 2000’). Globally, it is ranked
G5 or common and secure in its range (NatureServe
2006°).
Threats and Protection
The most direct threat to Lupinus lepidus is habitat
destruction. Grass-dominated meadows, often associ-
150
ated with Garry Oak-Brome (Quercus garryana-
Bromus) stands, commonly occurred on gentle slopes
on the most climatically favourable coastal areas on
south-eastern Vancouver Island and some of the Gulf
Islands prior to European colonization. Since coloniza-
tion both types of vegetation have been subjected to ex-
tensive agricultural and residential development. Their
destruction has continued to the present, resulting in
the elimination of almost all sites occurring outside
parks or ecological reserves. The loss of these habi-
tats, estimated at 95% (Lea 2002), severely reduces the
availability of sites for the establishment of L. lepidus
var. lepidus and imposes severe limitations on the long-
term survival of this species in Canada.
Introduced species including Cytisus scoparius,
Cynosurus echinatus and Anthoxanthum odoratum now
dominate L. /epidus habitats, further threatening the
survival of this species. The introduction of aggressive
European species has resulted in substantial changes to
the grass-dominated meadows associated with Quer-
cus garryana, and rocky xeric sites in the Victoria area.
Cytisus scoparius, in particular, has been one of the
most devastating species. It has become a dominant
shrub on xeric, exposed sites throughout much of south-
eastern Vancouver Island and the Gulf Islands.
Furthermore, the suppression of natural and unnat-
ural periodic fires appears to have resulted in changes
to the vegetation of many sites where this species would
be expected to occur. In the past, aboriginal peoples
may have set fire to these sites to maintain them as an
important habitat for wildlife (Roemer 1972). This
would have destroyed much of the competing vegeta-
tion and created habitats where L. lepidus would have
had the opportunity to become established. However,
since that time, these sites have experienced little dis-
turbance, resulting in invasion and expansion by trees,
shrubs, and especially introductions, which effectively
eliminate many herbaceous species.
In Canada, some populations of L. /epidus are pro-
tected to a certain extent by their location on public
property. This was not the case, however, for the plant
in Beacon Hill Park which was destroyed by mainte-
nance activites along a roadbank in 1994, or for the
population on the highway right-of-way at Duncan,
which was also destroyed and covered in 1994 by
heavy equipment used in a North Cowichan Munici-
pality sewer line project. Extirpation is also a probable
fate at Cattle Point and Observatory Hill. Even though
Cattle Point is located in a municipal park, it is not
protected from trampling by pedestrians and mountain
bike enthusiasts. During the summer months, tour buses
arrive at this park many times each day resulting in
large numbers of people trampling the vegetation along
the shoreline and adjacent rock outcrops. As a result
of these activities, much of the vegetation has been
degraded to the point where the number of potential-
ly suitable habitats for the establishment of L. lepidus
are far fewer today than in the past. The population
THE CANADIAN FIELD-NATURALIST
Vol. 120
could also have been overcome if it occurred in areas
now dominated by shrubs.
Observatory Hill is owned by the federal government
and is home to the Dominion Astrophysical Observa-
tory. Much of the vegetation has been replaced by
exotic species but access is limited to a single road.
Although further development on the hill is unlikely to
occur, many of the suitable habitats in which L. lep-
idus would likely be found are infested with Cytisus
scoparius .
Populations at Mount McDonald and Mount Braden
are protected by their location within either the Greater
Victoria Watershed District (which supplies drinking
water to Victoria and adjacent communities and pro-
hibits public access) or the CRD Sooke Hills Wilder-
ness, respectively. Since these areas are dominated by
steep rocky terrain and since there are few trails, veg-
etation has not been degraded to the same extent as it
has in more accessible areas near Victoria.
The specific site at which L. lepidus was collected
in the Koksilah River Valley is not known, thus it is
not possible to provide any information on the extent
to which this population is protected. However, if it is
located on a steep, rocky, xeric sites similar to those
found on Mount McDonald, Mount Braden, and Mount
Wall, it is unlikely to be threatened in the near future
by development or habitat destruction.
The site on Mount Wells is on private land but is not
easily accessed at the present time. Nearby mountains,
however, have been subject to housing developments
in recent years thus the site of L. lepidus is not neces-
sarily secure.
Lupinus lepidus is not formally protected in British
Columbia, however, it could be in the future since
this species is a potential candidate for listing under
the provincial Wildlife Amendment Act (2004). As part
of its commitment to the National Accord (National
Accord for the Protection of Species at Risk), the
province is required to take measures to protect this
species. It is on Schedule 1 of the federal Species At
Risk Act and a recovery strategy (which treats Lupi-
nus lepidus in addition to several other species-at-risk)
has been proposed (Parks Canada Agency 2005"). The
recovery strategy is a planning document that identi-
fies what needs to be done to arrest or reverse the de-
cline of a species. Detailed planning is done at the
action plan stage. However, an action plan is not cur-
rently available, nor are there any stewardship activi-
ties that are currently being implemented.
Evaluation
The Committee on the Status of Endangered Wildlife
in Canada and the British Columbia Conservation Data
Centre consider Lupinus lepidus to be endangered in
Canada (Douglas et al. 2002a). It has only been col-
lected nine sites in Canada, only three of which have
recently been confirmed, all confined to southeastern
Vancouver Island. It is believed to be extirpated at five
2006
of these sites and has not been confirmed recently at
two sites. The prognosis for this species is not good
considering the threats posed by aggressive competi-
tive species. The loss of suitable habitats at other sites
to residential development severely limits the potential
of this species to become established at new sites. Fur-
thermore, fire, which may have been an important agent
in the continued existence of this species in British
Columbia, is now actively suppressed. With so little
information known on the ecology of L. lepidus, and
on its relationship to the environment and other species,
successful management of the species is hampered.
Acknowledgments
Special thanks to Hans Roemer, whose persistent
searches during recent years continue to reveal Lupi-
nus lepidus sites. Thanks also to Jenifer Penny for office
assistance.
Documents Cited (marked * in text)
COSEWIC. 2000. Canadian Species at Risk. Committee on
the Status of Endangered Wildlife in Canada. www.
cosewic.gc.ca, Ottawa, Ontario. Available Online at: http://
Www.sararegistry.gc.ca/species/speciesDetails_e.cfm?sid
=187.
NatureServe Explorer. 2006. NatureServe Explorer: An
online encyclopedia of life. Version 1.7. Arlington, Vir-
ginia, USA. Web site: http://www.natureserve.org/explorer/
[Accessed 22 March 2006].
Parks Canada Agency. 2005. Recovery Strategy for Multi-
species at Risk in Maritime Meadows Associated with
Garry Oak Ecosystems in Canada (proposed). Jn Species at
Risk Act Recovery Strategy Series. Edited by Parks Canada
Agency. Ottawa, Ontario. 98 pages.
Ryan, M., and G. W. Douglas. 1996. Status Report on the
Prairie Lupine, Lupinus lepidus var. lepidus, in Canada.
Committee on the Status of Endangered Wildlife in Canada.
25 pages.
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of Alberta. National Museums Canada, Syllogeus 17,
Ottawa, Ontario.
Cox, B. J. 1973a. A chemosystematic comparison of the Lupi-
nus lepidus-L. caespitosus complex. Bulletin of the Torrey
Botanical Club 100: 12-17.
Cox, B. J. 1973b. Protein relationships among the perennial
caespitose lupines. Bulletin of the Torrey Botanical Club
100: 153-158.
Detling, L. E. 1951. The caespitose lupines of western North
America. American Midland Naturalist 45: 474-499.
Douglas, G. W. 1990. Fabaceae. Pages 21-45 in The Vascular
Plants of British Columbia. Part 2 — Dicotyledons (Dia-
pensiaceae through Portulacaceae). Edited by G. W. Dou-
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Ministry of Forests, Victoria, British Columbia. 158 pages.
Douglas, G. W., and L. C. Bliss. 1977. Alpine and subalpine
plant communities of the North Cascades Range, Washing-
ton and British Columbia. Ecological Monographs 47: 1 13-
150.
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare
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DOUGLAS AND RYAN: PRAIRIE LUPINE 151
tion. British Columbia Ministry of Sustainable Resource
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Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Ulustrated
Flora of British Columbia. Volume 3. Dicotyledons (Dia-
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Douglas, G. W., D. Meidinger, and J. Pojar. 200/a. Ilus-
trated flora of British Columbia. Volume 6. Monocotyle-
dons (Acoraceae through Najadaceae). British Columbia
Ministry of Sustainable Resource Management and British
Columbia Ministry of Forests, Victoria, British Columbia.
361 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2001b. Ilustrat-
ed flora of British Columbia. Volume 7. Monocotyledons
(Orchidaceae to Zosteraceae). British Columbia Ministry
of Sustainable Resource Management and British Columbia
Ministry of Forests, Victoria, British Columbia. 379 pages.
Douglas G. W., D. Meidinger, and J. Pojar. 2002b. Illus-
trated Flora of British Columbia. Volume 8 — General
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Sustainable Resource Management, British Columbia Min-
istry of Forests, Victoria, British Columbia. 457 pages.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1990.
The Vascular Plants of British Columbia. Part 2 — Dico-
tyledons (Diapengiaceae through Portulaceae). Special
Report Series Number 2, British Columbia Ministry of
Forests, Victoria, British Columbia. 155 pages.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a.
Rare Native Vascular Plants of British Columbia. British
Columbia Ministry of Environment, Lands and Parks, Vic-
toria, British Columbia. 423 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998b. Illustrated flora of British Columbia. Volume 1.
Gymnosperms and Dicotyledons. (Aceraceae through
Asteraceae). British Columbia Ministry of Environment,
Lands and Parks and British Columbia Ministry of Forests,
Victoria, British Columbia. 436 pages.
Dunn, D. B., and J. M. Gillett. 1966. The lupines of Canada
and Alaska. Monograph Number 2. Canada Department of
Agriculture, Ottawa, Ontario.
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1961. Vascular plants of the Pacific Northwest.
Part 3: Saxifragaceae to Ericaceae. University of Washing-
ton Press, Seattle, Washington. 614 pages.
Kuijt, J. 1982. A flora of Waterton Lakes National Park. The
University of Alberta Press, Edmonton, Alberta. 684 pages.
Lea, Ted. 2002. Historical Garry oak ecosystems of Greater
Victoria and Saanich Peninsula. 1:20,000 Map. Terrestrial
Information Branch, British Columbia Ministry of Sustain-
able Resource Management. Victoria, British Columbia.
Available online at: http://www.goert.ca/maps/go-020202.
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Packer, J. G. 1983. Flora of Alberta. Second edition. Univer-
sity of Toronto Press, Toronto, Ontario. 687 pages.
Phillips, L. L. 1955. A revision of the perennial species of
Lupinus of North America exclusive of southwestern United
States and Mexico. Research Studies of the State College
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Pojar, J. 1999. Fabaceae. Pages 64-180 in Illustrated Flora
of British Columbia. Volume 3 — Dicotyledons (Diapen-
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D. Meidinger, and J. Pojar. British Columbia Ministry or
Sy THE CANADIAN FIELD-NATURALIST Vol. 120
Environment, Lands and Parks, British Columbia Ministry Taylor, T. M. C. 1974. The Pea Family (Leguminosae) of
of Forests, Victoria, British Columbia. 423 pages. British Columbia. Handbook Number 12. British Colum-
Roemer, H. L. 1972. Forest vegetation and environments of bia Provincial Museum, Victoria, British Columbia. 251
the Saanich Peninsula, Vancouver Island. Ph.D. thesis, Uni- pages.
versity of Victoria, Victoria, British Columbia. 405 pages.
Scoggan, H. J. 1978. The flora of Canada. Part 3. National
Museum of Natural Sciences Publications in Botany. Num- _— Received 29 April 2002
ber 7. National Museum of Natural Sciences, Ottawa, Accepted 24 July 2006
Ontario 568 pages.
Conservation Evaluation of the Seaside Birds-foot Trefoil. Lotus
. . . *
formosissimus, in Canada
GEORGE W. DouGLas! and MICHAEL RYAN?
'Deceased.
Present address: 801 Frayne Road, RR #1, Mill Bay, British Columbia VOR 1LO Canada.
Douglas, George W., and Michael Ryan. 2006. Status of the Seaside Birds-foot Trefoil, Lotus formosissimus, in Canada
Canadian Field-Naturalist 120(2): 153-156.
- In Canada, Seaside Birds-foot Trefoil, Lotus formosissimus, is restricted to the Victoria area on southeastern Vancouver Island
and two adjacent islands. Populations at five sites are extant and have been recently inventoried, whereas populations at two or
more sites in Victoria are extirpated. Existing populations represent the northern range limits of L. formosissimus. Although
known populations are protected to a certain extent from direct habitat destruction, introduced herbaceous species may pose
| a serious threat to the continued existence of most populations and may prevent the establishment of L. formosissimus at
other sites. Fire suppression, grazing, encroachment and low survivorship are also limiting factors.
Key Words: Seaside Birds-foot Trefoil, Lotus formosissimus, endangered, distribution, population size, British Columbia.
Seaside Birds-foot Trefoil, Lotus formosissimus
Greene, is a member of a genus of about 200 species
occurring throughout the world, primarily in temper-
ate zones. It is one of nine species occurring in British
- Columbia and Canada, four of which have been intro-
duced from the Old World (Pojar 1999; Scoggan 1978).
_ Lotus formosissimus is a sprawling or ascending,
| multi-stemmed perennial that grows from a short root-
| stock and thick taproot to 20-50 cm tall (Figure 1;
' Pojar 1999). It has compound, pinnate leaves 4-8 cm
| long with 3 to 7, ovate to obovate leaflets. The three to
five yellow and pink pealike flowers are 10-15 mm long
and arranged in umbels. The umbels are borne on axil-
lary peduncles 2-6.5 cm long and are usually subtend-
ed by trifoliate bracts. The wings of the flowers are
pinkish to purplish tinged, the keel is purple tipped,
and the banner is yellow and 11-15 mm long. The
wings and clawed keel exceed the calyx considerably.
Pods are 27-36 mm long by 1.5-2 mm wide and en-
close 7 to 15, dark brown to black seeds. Lotus for-
mosissimus is distinguished from other Lotus species
_ by its perennial habit, membranous stipules, trifoliate
' bracts (sometimes unifoliate), and pink and yellow
_ flowers (Ryan and Douglas 1994*, 1996*). Bog Bird’s-
_ foot Trefoil (L. pinnatus) also has yellow but never
pink corollas. Ficure 1. Illustration of Lotus formosissimus. (Line drawing
| from Pojar 1999).
| Distribution
Lotus formosissimus is restricted to the west coast
| of North America, from southeastern Vancouver Island
(S central California (Isely 1993; Pojar 1999). In Cana- and nearby islands (Figure 2; Dunn and Gillet 1966;
da, it is restricted to southeastern Vancouver Island Pojar 1999; Douglas et al. 2002a, 2002b).
OF
Pe an
1
| ‘The original field work for Lotus formosissimus was funded by the British Columbia Conservation Data Centre. The results appear in the
| British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a
| Committee on the Status of Endangered Wildlife in Canada status report (Douglas and Ryan 1996*) and the subsequent assessment of Endan-
| gered. This paper includes more recent information that did not appear in the original status report.
153
154
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Populations of Lotus formosissimus in Canada (in Victoria and west of Victoria at Department of National
Defense — Rocky Point ).
Collection Site Last Observation
Oak Bay, Victoria 1896
Foul Bay, Victoria 1912
William Head, W of Victoria 2002
Bentinck Island, Rocky Point 2004
Rocky Point, E side 2005
Rocky Point, Church Point 2004
Trial Island, Victoria 2004
Collector Population Status (number/area)
Anderson Extirpated
Henry Extirpated
Fairbarns 19/8 m?
Fairbarns 278/110 m?
Fairbarns 36-53/75.5 m?
Fairbarns 209-327/137 m2
Fairbarns 383/5000 m?
Habitat
Lotus formosissimus grows in various coastal mead-
ow and woodland habitats, ranging from open exposed
grass-dominated meadows to exposed steep rocky sites
inhabited by Garry Oak (Quercus garryana), to shad-
ed Garry Oak-Brome (Quercus garryana-Bromus)
forests. Associated species, which typically occur in
the understory of Quercus garryana stands, include
Lance-leaved Stonecrop (Sedum spathifolium), Sea
Blush (Plectritis congesta), Nodding Onion (Allium
cernuum), Menzies Larkspur (Delphinium menziesii),
Orchard Grass (Dactylis glomerata), Sweet Vernalgrass
(Anthoxanthum odoratum), Hairy Cat’s-ear (Hypo-
chaeris radicata), Common Velvet-grass (Holcus lana-
tus), Early Hairgrass (Aira praecox), Ribwort Plantain
(Plantago lanceolata), and several species of brome
(Bromus). Lotus formosissimus also occurs in vernal
seeps. It is suspected that L. formosissimus is a poor
competitor with a number of perennial grasses because
it is absent in a number of grass-dominated micro-sites
that are otherwise apparently suitable habitats.
Biology
Little is known regarding the biology of Lotus for-
mosissimus. It appears that L. formosissimus shoots
emerge in spring followed by flowering between May
and the end of June when moist soil conditions and
warm temperatures are prevalent in Quercus garryana
stands and meadows. By July, when drought conditions
are prevalent, seed maturation and dispersal begin,
followed soon thereafter by die-back of the stems to the
rootstock, which is located several centimetres below
the soil surface. In his demographic studies of Lotus
formosissimus, M. Fairbarns (personal communication)
discovered, based on ex situ experiments, that seeds may
remain in the seed bank for at least two years. He
postulated that this was a likely explanation for his in
situ observations of higher numbers of seedlings than
numbers of seeds set in the previous year.
Flowers likely require cross-pollination to set viable
seed although pollinators have not yet been identified
(Zandstra and Grant 1968). Seeds are believed to be
easy to germinate (Hitchcock et al. 1961) but like other
members of Fabaceae, they may have a hard seed coat,
delaying the germination of seeds for several months
or years. Mature plants may be long-lived and recruit-
ment from germinating seeds is likely a rare event.
It is believed that L. formosissimus, like other le-
gumes, is associated with nitrogen-fixing Rhizobium
that provides the plants with an independent source
of nitrogen from that in the soil.
Population Attributes
Lotus formosissimus has been collected at seven sites
in Canada on southern Vancouver Island or adjacent
islands (Table 1, Figure 2). Populations at two of these
sites are considered extirpated. Other older collections
not listed in Table 1 may or may not represent differ-
ent locations (e.g., Victoria, and Oak Bay, Victoria).
The population on Trial Island has been known since
the 1950s and surveyors reported 28 plants in 1994 and
380 plants in 2004. Likewise, the population at Rocky
Point has also been observed for over 20 years and was
represented by 165 plants in 1993 and 245-380 plants
in 2005. The increase, however, is likely attributable to
different observers and different sampling methods.
It is very unlikely that the populations are actually in-
creasing significantly. The remaining sites have not been
monitored for any length of time and thus, no infor-
mation is available on population trends.
Provincial, National and Global Ranks
The British Columbia Conservation Data Centre has
ranked this species as S1 and placed it on the Ministry
of Environment Red list. This is the most critical cat-
egory for imperiled rare native vascular plants in the
province. A rank of S1 is considered “critically imper-
iled because of extreme rarity (five or fewer occurrences
or very few remaining individuals) or because of some
factors making it especially vulnerable to extirpation
or extinction”. Lotus formosissimus is on the Canadian
Species At Risk Act legal list (COSEWIC 2000*) and
globally, it is G5 or common and secure in its range
(NatureServe Explorer 2006').
Threats and Protection
In the past, the most direct threat to Lotus formo-
sissimus was that of habitat destruction. Grass-domi-
nated meadows, often associated with Quercus gar-
ryana-Bromus stands, commonly occurred on gentle
slopes on the southeastern side of Vancouver Island
and some of the Gulf Islands prior to European colo-
nization. Since colonization, both types of vegetation
have been subjected to extensive agricultural and res-
2006
idential development and have been essentially elimi-
nated outside parks or ecological reserves. Direct habi-
tat destruction, however, is no longer the most impor-
tant threat. It is the loss of suitable sites outside the
current occupied area of L. formosissimus that severely
limits the potential for this species to spread into new
areas and therefore, threatens its long-term survival
in Canada.
Less direct factors which threaten known popula-
tions include the introduction of aggressive European
species. These species have resulted in substantial
changes to the grass-dominated meadows associated
with Quercus garryana and the rocky xeric sites in the
Victoria area. One of the most devastating species is
Scotch Broom (Cytisus scoparius). This species has
become a dominant shrub on xeric, exposed sites
throughout much of southeastern Vancouver Island
and the Gulf Islands and either competes with native
plants such as L. formosissimus for resources or pre-
vents the establishment of their seedlings.
Furthermore, the suppression of natural and unnat-
ural periodic fires may have resulted in changes to the
vegetation of many sites where this species would be
expected to occur. In the past, aboriginal peoples prob-
ably set fire to many of these sites to maintain them as
an important habitat for food plants and wildlife (Roe-
mer 1972). During the past century these sites have
experienced little disturbance resulting in increased
domination of some sites by trees and shrubs, especially
introductions, that effectively eliminate many herba-
ceous species.
Another limiting factor is grazing. M. Fairbarns
(personal communication) observed that grazing prior
to seed set had greatly limited seed production and
caused reduced vigour among established plants on
Bentinck Island.
Finally, demographic constraints also play a role in
the viability of populations. Fairbarns (personal com-
munication) found low rates of juvenile survivorship
and low levels of seed production. He found that they
were related primarily to summer drought and grazing.
All extant populations of L. formosissimus in British
Columbia are protected from heavy recreational pres-
sure and disturbance caused by the general public. Al-
though these sites contain a large number of introduced
species, the latter were not nearly as abundant at Rocky
Point, Bentinck Island, and Trial Island as they are at
other public sites. It is very likely that restricted public
access and lack of disturbance are the primary reasons
for the continued existence of L. formosissimus at these
sites.
The populations at Rocky Point and Bentinck Island
are owned by the Department of National Defense. The
Department of National Defence is aware of the loca-
tion of rare species at Rocky Point and there are no plans
to develop those areas in which L. formosissimus occurs.
Bentinck Island is used by DND to detonate unused
ammunition, but such activities do not coincide with
DOUGLAS AND RYAN: SEASIDE BiIRDS-FOOT TREFOIL
155
VANCOUVER
ISLAND
FIGURE 2. The status and location of Lotus formosissimus
sites in British Columbia. @ — recently confirmed
sites, O — extirpated site.
locations of L. formosissimus on the island. These pop-
ulations are also protected by the Canadian Species at
Risk Act by their occurrence on land under federal
jurisdiction and therefore, are not likely to be directly
harmed. The Canadian Forest Service is also involved
in a number of projects with the goal of managing ap-
propriately for species at risk on federal lands, includ-
ing Scotch Broom removal from sites.
The Lotus formosissimus population in Trial Island
Ecological Reserve is also provided with a high degree
of protection. Despite its proximity to Victoria, it has
156
very limited public access because it is only accessi-
ble by boat, thus preventing excessive disturbance.
A recovery strategy (which treats Lotus formosis-
simus in addition to several other species-at-risk) has
been proposed (Parks Canada Agency 2005"). The
recovery strategy is a planning document that identi-
fies what needs to be done to arrest or reverse the
decline of a species. Detailed planning is done at the
action plan stage. In the absence of an action plan, no
attempts have been made to introduce L. formosissimus
to suitable habitats or to increase the number of indi-
viduals at current locations. However, Scotch Broom
removal at the Bentinck Island site has revealed addi-
tional Lotus plants (M. Fairbarns, personal communi-
cation). The site now offers improved conditions for
growth and reproduction.
Evaluation of Status
The British Columbia Conservation Data Centre con-
siders Lotus formosissimus to be endangered in Cana-
da (Douglas et al. 2002a). The Committee on the Sta-
tus of Endangered Wildlife in Canada has also assigned
this species to the Endangered category (COSEWIC
2000"). Between 925 and 1060 plants are known from
five sites on southeastern Vancouver Island and adja-
cent islands. The long-term survival of this species in
Canada is dubious as a result of a number of limiting
factors: shading by aggressive introduced species, poor
survivorship, grazing, fire suppression and encroach-
ment. Even though known populations are protected
from excessive disturbance on federal lands and in the
ecological reserve, the loss of suitable habitats at other
sites severely limits the potential of this species to
become established at new sites.
Acknowledgments
We thank Matt Fairbarns for population updates for
all extant locations and Jenifer L. Penny for assistance
in the office.
Documents Cited (marked * in text)
COSEWIC. 2000. COSEWIC assessment and status report
on the Seaside birds-foot lotus Lotus formosissimus in
Canada. Committee on the Status of Endangered Wildlife
in Canada. Ottawa, Ontario. vi + 18 pages.
NatureServe Explorer. 2006. NatureServe Explorer: An
online encyclopedia of life. Version 1.7. Arlington, Virgi-
nia, USA. Web site: http://www.natureserve.org/explorer
[Accessed 22 March 2006].
THE CANADIAN FIELD-NATURALIST
Vol. 120
Parks Canada Agency. 2005. Recovery Strategy for Multi-
species at Risk in Maritime Meadows Associated with
Garry Oak Ecosystems in Canada (proposed) in Species at
Risk Act Recovery Strategy Series. Edited by Parks Canada
Agency. Ottawa, Ontario. 98 pages.
Ryan, M., and G. W. Douglas. 1994. Status report on the
Seaside birds-foot lotus, Lotus formosissimus Greene. Un-
published report submitted to the Committee on the Status
of Endangered Wildlife in Canada, Ottawa. British Colum-
bia Conservation Data Centre, Victoria, British Columbia.
25 pages.
Ryan, M., and G. W. Douglas. 1996. COSEWIC status report
on seaside birds-foot lotus Lotus formosissimus in Canada.
Committee on the Status of Endangered Wildlife in Cana-
da. Ottawa, Ontario. 1-18 pages.
Literature Cited
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a.
Rare native vascular plants of British Columbia. Second
edition. British Columbia Ministry of Sustainable Resource
Management, Victoria, British Columbia. 359 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2002b. Illus-
trated flora of British Columbia. Volume 8. General sum-
mary, maps and keys. Ministry of Sustainable Resource
Management, Ministry of Forests, Victoria, British Colum-
bia. 458 pages.
Dunn, D.B., and J. M. Gillett. 1966. The lupines of Canada
and Alaska. Monograph number 2. Canada Department
of Agriculture, Ottawa, Ontario.
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1961. Vascular plants of the Pacific Northwest.
Part 3: Saxifragaceae to Ericaceae. University of Washing-
ton Press, Seattle, Washington. 614 pages.
Isley, D. 1993. Lotus. Pages 616-622 in The Jepson manual:
higher plants of California. Edited by J. C. Hickman. Uni-
versity of California Press, Los Angeles, California. 1400
pages.
Pojar, J. 1999. Fabaceae. Pages 64-180 in Illustrated Flora
of British Columbia. Volume 3. Dicotyledons (Diapensi-
aceae through Onagraceae). Edited by G. W. Douglas, D.
Meidinger, and J. Pojar. British Columbia Ministry of
Environment, Lands and Parks, British Columbia Ministry
of Forests, Victoria, British Columbia. 423 pages.
Roemer, H. L. 1972. Forest vegetation and environments of
the Saanich Peninsula, Vancouver Island. Ph.D. thesis, Uni-
versity of Victoria, Victoria, British Columbia. 405 pages.
Scoggan, H. J. 1978. The flora of Canada. Parts 1-4. Nation-
al Museum of Natural Sciences Publications in Botany 7:
1-1626.
Zandstra, I. I., and W. F. Grant. 1968. The biosystematics
of the genus Lotus (Leguminosae) in Canada. I. Cytotax-
onomy. Canadian Journal of Botany 46: 557-583.
Received 29 April 2002
Accepted 24 July 2006
Conservation Evaluation of the Bog Bird’s-foot Trefoil, Lotus pinnatus,
. *
in Canada
~ Marta Donovan!
Victoria British Columbia V8W 9R7 Canada.
Donovan, Marta. 2006. Conservation evaluation of the Bog Bird’s-foot Trefoil, Lotus pinnatus, in Canada. Canadian Field-
' Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, P.O. Box 9993 St~ Prov Gove.
) Naturalist 120(2): 157-162.
In Canada, Bog Bird’s-foot Trefoil, Lotus pinnatus, is confined to vernally wet meadows, creek margins, ditches and seepages,
where underground water comes to the surface. Seven localities, within a small area on the east coast of Vancouver Island
near Nanaimo, British Columbia, have been confirmed in recent years. At two others it is considered extirpated. The lack of
protection on private land, alterations to hydrology, and erosion arising from intensive recreational use of primary habitat by
off-road vehicles as well as the appearance of invasive, species are the main concerns for British Columbia populations.
Key Words: Bog Bird’s-foot Trefoil, Lotus pinnatus, endangered species, distribution, conservation, British Columbia.
Bog Bird’s-foot Trefoil, Lotus pinnatus Hook’, is a
low-growing, short-lived perennial of the pea family
_ (Fabaceae) with many erect to spreading stems ranging
from 15 to 60 cm long (Figure 1). The pinnately com-
pound leaves are 4-8 cm long, with 5-9 elliptic, oblong
or narrowly egg-shaped leaflets. The inflorescence is
| a compact, stalked, axillary umbel of 3 to 12 flowers.
The corollas are 10-15 mm long with a yellow ban-
ner and keel and creamy-white wings. The keel petals
are fused along one edge that is elongated into a well-
_ defined beak towards the outside of the inflorescence.
The calyx is tubular and lobed, with two upper lobes
that are joined most of their length. The linear seed
pods are 3-6 cm long and 1.5-2 mm wide and contain
5-20 cylindrical, glossy, dark-coloured seeds (Pojar
1999). In the field, several species superficially resem-
ble L. pinnatus. Vetches (Vicia species) are often pres-
“ent in similar habitats, but the leaves are smaller than
those of L. pinnatus and the terminal leaflet in Vicia spp.
_is a tendril. Seaside Birds-foot Trefoil (Lotus formo-
sissimus) looks similar to L. pinnatus except the wing
petals of L. formosissimus are pinkish-purple, while
‘those of L. pinnatus are cream-colored. In addition,
|though a trifoliate bract usually subtends the flowers
of L. formosissimus, bracts were sometimes absent in
field specimens or only a unifoliate bract was present
(Ryan and Douglas 1994*). Lotus pinnatus and L. for-
|mosissimus could be confused if the plants are imma-
‘ture or not in flower, but the two species do not over-
lap in their distribution in British Columbia. Lotus
formosissimus is known only from the Victoria area
and nearby islands, where it occurs in various xeric
habitats, ranging from open grass-dominated meadows
|
i
FiGure 1. Illustration of Lotus pinnatus (line drawing in
Hitchcock et al. 1961; Pojar 1999).
to exposed steep rocky sites with Garry Oak (Quercus
garryana) (Ryan and Douglas 1994*). Lotus pinnatus is
found only in the Nanaimo area on Vancouver Island,
where it occurs in moist soil on exposed, coastal low-
land areas. Bird’s-foot Trefoil (Lotus corniculatus)
could also be confused for L. pinnatus as it occasion-
ally grows in wet places, but it is usually found in drier,
disturbed sites. Also, L. corniculatus has completely
yellow flowers and its leaflets are smaller and more
blunt than the other trefoils.
'* The information, gathered during field surveys in 2003 formed the basis for a Committee on the Status of Endangered Wildlife in Canada
|Status report (Donovan 2004*) and the subsequent assessment of Endangered (COSEWIC 2004*). This paper includes more recent information
ie did not appear in the original status report.
|’ Taxonomy and nomenclature follow Douglas et al. (1999).
lSy7/
158
Distribution
Lotus pinnatus occurs in western North America
from Vancouver Island, south to northwest Washing-
ton, western Oregon and the Columbia River Gorge,
to central California and sporadically eastward to Idaho
(Figure 2; Isely 1993; Pojar 1999). In Canada, Lotus
pinnatus is known from seven extant populations with-
in a small area on the east coast of Vancouver Island
near Nanaimo (Donovan 2004’).
Habitat
The British Columbia populations of L. pinnatus
occur within the moist maritime subzone of the Coast-
al Douglas-fir (CDF) biogeoclimatic zone (Meidinger
and Pojar 1991) in the Nanaimo Lowlands Ecosection of
the Georgia Depression Ecoprovince (Demarchi 1996’).
In this region, the Olympic Mountains in Washington
State to the south and the Insular Mountains on Vancou-
ver Island to the west produce a rain shadow effect
resulting in a Mediterranean-type climate with warm,
dry summers and mild, wet winters. Most of the rain-
fall occurs during the winter months, and limited pre-
cipitation and high temperatures during the summer
months result in pronounced summer moisture deficits
(Meidinger and Pojar 1991).
Lotus pinnatus grows in open, springy meadows,
along the margins of creeks, or in seepages, where
underground water comes to the surface and the plants
are in close physical contact with cool, flowing water
(H. Roemer, personal communication). In all cases, the
soils are shallow (< 15 cm), over gently sloping sand-
stone or conglomerate bedrock with abundant mois-
ture during the growing and blooming period. Lotus
pinnatus is most commonly associated with Yellow
Monkey-flower (Mimulus guttatus), Sea Blush (Plec-
tritis congesta), White Triteleia (Triteleia hyacinthina),
Small-leaved Montia (Montia parvifolia), Scouler’s
Popcornflower (Plagiobothrys scouleri) and American
Speedwell (Veronica beccabunga ssp. americana).
Stands of Douglas-fir (Pseudotsuga menziesii) and
dense thickets of Nootka Rose (Rosa nutkana), Ocean
Spray (Holodiscus discolor), Nine-bark (Physocarpus
capitatus) and willow (Salix spp.) exist on the margin
of some seepages but Lotus pinnatus does not occur in
the shaded understory of these sites, suggesting that the
species is shade-intolerant. Prolonged moisture, edge
habitat along streams and meadows and shallow soils
derived from sedimentary rock are key habitat compo-
nents. Other factors, such as slope and aspect are vari-
able and do not appear to be critical in defining suit-
able habitat. In British Columbia, the elevation for this
species ranges between 40 m and 150 m.
Biology
Zandstra and Grant (1968) studied the biosystem-
atics of native and introduced Lotus species in Cana-
da and reported that although many of the Old World
species are polyploids, none of the North American
THE CANADIAN FIELD-NATURALIST
Vol. 120
FIGURE 2. Distribution of Lotus pinnatus in North America
(from Donovan 2004*).
species examined exhibited the same trait. North Ameri-
can Lotus species have a base chromosome number
of 6 or 7. Of the species native to Canada, L. pinnatus
and L. formosissimus are considered to be the most
closely related, based on morphological, habitat, and
cytological characteristics. A chemotaxonomic study
using thin-layer chromatography further supported the
general relationships among Canadian Lotus species
with L. formosissimus and L. pinnatus demonstrating
the greatest degree of similarity (Grant and Zandstra
1968). These species differed from other native Cana-
dian species in that both were perennial outcrossers
with large flowers on long pedunculate umbels. The
other three native species were self-fertile annuals with
small flowers. Evolution in angiosperm reproduction
has frequently occurred with a decrease in basic chro-
mosome number, a shift from an outcrossing (cross-
pollinating) to an inbreeding (self-pollinating) repro-
ductive system and a shift from a perennial to an
annual habit (Stebbins 1957). On this basis, Zandstra
and Grant (1968) considered L. pinnatus and L. formo-
sissimus to be more primitive than the annual species.
Lotus pinnatus germinates in the late winter or early
spring when soil moisture and temperature conditions
are most favourable, and seedling survival appears to
be dependent on continuous surface moisture during
this period. Though the specific timing of events will
depend on local habitat conditions, flowering in L. pin-
natus generally occurs between May and the end of
June.
2006
The pea-like, bisexual flowers are borne at the end
of a long peduncle arising from the leaf axil, and the
umbellate inflorescences are produced sequentially
throughout the growing season. As the flowers senesce,
some become progressively more reddish (Isely 1981).
During July, when drought conditions are prevalent,
seed maturation and dispersal begin, followed by die-
back of the stems to the rootstock, which is located sev-
eral centimetres below the soil surface. The seeds lack
any strong adaptations for long-distance dispersal by
wind or animal vectors. Most seeds are gravity-dis-
persed and generally land in the immediate vicinity
of the parent plant. The plant’s habitat along stream
channels may permit the legumes and seeds to be trans-
ferred by water during times of seasonal flooding.
However, germination and seedling survival appear to
depend upon continuous surface moisture, and the
species does not appear to be a strong competitor with
native shrub species or with invasive alien plants. The
chances of a healthy population returning to Canada if
local populations become extirpated are highly unlike-
ly. The nearest population in Washington State, from
which collections were made in 1940, is 240 km away
in Bremerton. Whether this population is extant is un-
known.
As with other perennial species in the genus Lotus,
Zandstra and Grant (1968) suggested that the flowers
of L. pinnatus required cross-pollination to set viable
seed. Given the bright yellow appearance of the flower
and the wide keel that serves as a suitable landing plat-
form, flowers of L. pinnatus appear to be adapted to
bee pollination. Although the importance of the bum-
blebee to the flower (and vice versa) is yet to be specif-
ically determined, bumblebees were observed visiting
flowers of L. pinnatus at Harewood Plains during field
observations made in 2003. Although bees are capable
of dispersing pollen over short distances, cross-polli-
nation between most populations in Canada is unlikely
in view of the distances involved.
Like many other legumes, L. pinnatus appears to be
associated with nitrogen-fixing Rhizobium bacteria
that occupy root nodules and provide the plants with
a source of reduced nitrogen in exchange for a supply
of carbon and other nutrients. Specific strains of Rhi-
zobium bacteria are required for effective nodulation
of Lotus species grown as forage crops, such as L.
corniculatus and L. tenuis. To maximize establishment
in areas that have never produced Lotus, inoculation
of seeds with the appropriate rhizobia is necessary
(Blumenthal and McGraw 1999).
An examination of root nodules collected from L.
pinnatus at Harewood Plains indicated the presence
of bacteria, most likely of genus Rhizobium (S. Berch,
personal communication).
Population Attributes
Lotus pinnatus has been reported from nine sites in
British Columbia (Figure 3; Table 1), all in the Nanai-
DONOVAN: BOG BiIRD’S-FOOT TREFOIL
159
BRITISH
, COLUMBIA
=
Ta
Ke
ree
VICTORIA
FiGurE 3. Distribution of Lotus pinnatus in British Colum-
bia (from Donovan 2004*) @ confirmed: a uncon-
firmed.
mo area on Vancouver Island and on nearby Gabriola
Island. Of the nine reported records, seven populations
were verified in recent years. The remaining two sites
have been extensively disturbed and it is unlikely that
these populations currently exist. Between 1500 and
2000 individual plants were counted in Canada in 2003
over an area of 650 square metres. Relatively little is
known of demographic characteristics and population
trends of L. pinnatus due to a lack of long-term mon-
itoring. Estimates of population trends in L. pinnatus
are currently unreliable as the clumped, sprawling habit
makes it difficult to identify separate individuals and
counting methods have varied with different investiga-
tors. Until counting methods are standardized, popula-
tion numbers should be considered rough estimates.
Provincial, National and Global Ranks
The British Columbia Conservation Data Centre
has ranked L. pinnatus as “S1” and placed it on the
“Red list”. A rank of “S1” indicates that the species is
“critically imperiled because of extreme rarity (often
160
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Locations and Population Sizes for Lotus pinnatus in British Columbia.
Populations Observation Collector/ Number of
Date Observer Individual Plants Apparent Status
Harewood Plains North’ 7 July 2003 M. Donovan 1500 in 500 m? Extant
Harewood Plains South 1 12 July 2003 M. Donovan 25 in 4 m? Extant
Harewood Plains South 2 12 July 2003 M. Donovan 10 in 2 m Extant
Woodley Range 1 June 2003 M. Donovan 120-140 in 90 m? Extant
Gabriola Island, off Peterson
Road, on Perry Road 25 May 2003 Donovan/Penny 65-70 in 10.5 m? Extant
Nanaimo, west of Cinnabar Valley 15 June 2003 M. Donovan 40 in 25 m? Extant
Nanaimo, south of Extension 23 June 2003 M. Donovan 30 to 45 in 17 m2 Extant, but declining
Waddington & Departure Bay
Roads Nanaimo 2 June 1939 J.W. Eastham N/A Extirpated
Departure Bay Road and Island
Highway, Nanaimo 20 June 1965 K. Beamish N/A Extirpated
“The collection by W. R. Carter at Mt. Benson in | June 1918 is included with this population as Harewood Plains may be
considered the foothills of Mt. Benson.
5 or fewer occurrences) or because of factors making
it especially vulnerable to extirpation.” This is the most
critical status that can be applied to a species at the
provincial level. Though it currently lacks formal pro-
tection in British Columbia, Lotus pinnatus 1s a poten-
tial candidate for listing under the provincial Wildlife
Amendment Act (2004).
At the national level, Lotus pinnatus has been as-
sessed as Endangered by the Committee on the Status
of Endangered Wildlife in Canada (COSEWIC 2004")
and appears on Schedule 1 of the federal Species At
Risk Act (Environment Canada 2006’).
Globally, Lotus pinnatus has a rank of G4G5 which
indicates that the plant is apparently secure in most
of its range (NatureServe Explorer 2006’).
Threats and Protection
Habitat loss presents a serious and urgent threat to
Lotus pinnatus in Canada. Over 90% of the extant pop-
ulation of L. pinnatus is located on land that is privately
owned, making this species vulnerable to habitat loss
as a result of urban expansion and residential devel-
opment.
In addition to potential commercial and residential
development, the population of L. pinnatus at Hare-
wood Plains is under direct and immediate threat of
habitat degradation and fragmentation resulting from
intensive, unauthorized use of recreational off-road
vehicles; including ATVs, 4 x 4s and dirt bikes (C. Thir-
kill, personal communication). In some areas at this
site, the thin and fragile soils have been rutted down
to bedrock and plants have been dislodged on to bare
rock where they cannot re-establish (Figure 4). There
is high potential for further impacts as vehicles con-
tinue to access the site, despite efforts by the former
landowner (Weyerhaueser Canada) to block access to
off-road vehicles with boulder placements, ditches and
the installation of several cement barriers along the
access road (C. Thirkill, personal communication). In
an effort to minimize the disturbance of sites desig-
nated as “Environmentally Sensitive Areas” the City
of Nanaimo has established a Development Permit Area
on a portion of the property at Harewood Plains (Rob
Lawrance, personal communication). As the habitat at
Harewood Plains supports several other species at risk
such as Dense-spike Evening Primrose (Epilobium den-
siflorum), Green-sheathed Sedge (Carex feta), Foothill
Sedge (Carex tumulicola), Slim-leaf Onion (Allium
amplectens) and Nuttall’s Quillwort (/soetes nuttallii),
a working group comprised of representatives from
industry, non-governmental organizations and local and
provincial governments has been established in order
to develop proactive communication with the different
land owners, managers and stakeholders at this location.
At the site south of Extension, near Nanaimo, at least
some portions of the population have been reported
extirpated due to recreational off-road vehicle use and
residential development (A. Ceska, personal commu-
nication).
Other than habitat destruction, competitive exclu-
sion from native and non-native vegetation represents
the most significant ongoing threat to L. pinnatus at all
sites. Encroachment of native shrub species in poten-
tial habitats may prevent this species from occupying
new sites. Invasive alien grass species that threaten the
persistence of L. pinnatus include Sweet Vernalgrass
(Anthoxanthum odoratum), Orchard Grass (Dactylis
glomerata), Kentucky Bluegrass (Poa pratensis) and
Barren Brome (Bromus sterilis). Scotch Broom (Cytisus
scoparius) is the most dominant invasive alien shrub.
Logging operations close to (within 50 m of) popula-
tions of L. pinnatus at Harewood Plains increase the
potential for the spread of aggressive native and non-
native plant species. By disturbing soils and carrying
seeds, off-road vehicles are also a major factor in the
spread of invasive non-native plants in sensitive areas.
The population located in the Woodley Range Eco-
logical Reserve is protected, to a certain extent, by its
2006
DONOVAN: BOG Birpd’s-FOOT TREFOIL 16]
a ~
Ss
N ay he
if
FiGure 4. Destruction of habitat caused by all-terrain vehicles at Harewood Plains (Photo George Douglas, 2003).
location within an ecological reserve, but this repre-
sents only 7% of the total Canadian population. In
addition, plants at this location are vulnerable to adja-
cent land uses including all-terrain vehicle use, log-
ging and other land clearing activities which could
impact on the hydrological features that provide seep-
age to the location.
A national multi-species strategy that addresses the
recovery of Lotus pinnatus and five other endangered
plant species of vernal pools and other ephemeral wet
areas has been completed. This strategy outlines recov-
ery goals for each species, suggests strategic approach-
es for meeting the recovery goals and recommends that
a draft recovery action plan (RAP) be completed by
October 2009, which includes proposing Critical Habi-
tat for this species under the Species At Risk Act (Parks
Canada Agency 2005*).
Evaluation
The British Columbia Conservation Data Centre
considers Lotus pinnatus to be endangered in British
Columbia (Donovan 2004*) and the Committee on the
Status of Endangered Wildlife in Canada has assessed
the species as Endangered (COSEWIC 2004"). Only
a few small fragmented populations remain, and these
are restricted to wet meadows of limited occurrence
that are considerably disjunct from the main range of
the species in the north-western United States. The
populations are at risk from continued habitat loss, en-
croachment of invasive species and recreational off-
road vehicle use. Development of the habitat supporting
the only sizeable remaining population could result in
significant losses to the population (COSEWIC 2004 ).
Acknowledgments
Thanks to Jenifer Penny, Charles Thirkill and Don
Piggott for help with field surveys. Hans Roemer and
Shannon Berch provided information and technical
assistance. Matt Fairbarns provided strong support
throughout this study and Ted Lea reviewed the man-
uscript. I am indebted to the late George Douglas for
nurturing my interest in plants.
Documents Cited (marked * in text)
COSEWIC. 2004. COSEWIC assessment and status report
on the bog bird’s-foot trefoil Lotus pinnatus in Canada.
Committee on the Status of Endangered Wildlife in Cana-
da. Ottawa, Ontario. 33 pages. www.cosewic.gc.ca
Demarchi, Dennis A. 1996. An Introduction to the Ecore-
gions of British Columbia. Wildlife Branch, Ministry of
Environment, Lands and Parks, Victoria. (Available at http:
srmwww.gov.be.ca/ecology/ecoregions/title_author.html
[Accessed 22 June 2006])
Donovan, M. 2004. COSEWIC status report on the bog
bird’s-foot trefoil Lotus pinnatus in Canada. Jn COSEWIC
assessment and status report on the bog bird’s-foot trefoil
Lotus pinnatus in Canada. Committee on the Status of
Endangered Wildlife in Canada. Ottawa, Ontario. 33 pages.
162
Environment Canada, Schedule 1: List of species at risk.
Website: http://www.sararegistry.gc.ca/species/schedules_e.
cfm?id=1 [Accessed 22 June 2006].
NatureServe Explorer. 2006. NatureServe Explorer: An on-
line encyclopedia of life. Version 1.7. Arlington, Virginia,
USA. Web site: http://www.natureserve.org/explorer [Ac-
cessed 22 March 2006].
Parks Canada Agency. 2005. Recovery Strategy for Multi-
species at Risk in Maritime Meadows Associated with
Garry Oak Ecosystems in Canada (proposed) in Species
at Risk Act Recovery Strategy Series. Edited by Parks
Canada Agency. Ottawa, Ontario. 98 pages.
Literature Cited
Blumenthal, M. J., and R. L. McGraw. 1999. Lotus adap-
tation, use and management. Pages 97-119 in Trefoil: The
science and technology of Lotus. Edited by P. R. Beuse-
linck. Crop Science Society and the American Society of
Agronomy. Madison, Wisconsin.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illus-
trated Flora of British Columbia. Volume 3. Dicoteyle-
dons (Diapensiaceae through Onagraceae). British Colum-
bia Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia.
423 pages.
Grant, W. F., and I. I. Zandstra. 1968. The biosystematics
of the genus Lotus (Leguminosae) in Canada. II. Numer-
ical cytotaxonomy. Canadian Journal of Botany 46: 585-
589.
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1961. Vascular Plants of the Pacific Northwest.
Part 3: Saxifragaceae to Ericaceae. University of Washing-
ton Press, Seattle, Washington. 614 pages.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Isely, D. 1981. Leguminosae of the United States. III. Sub-
family Papilionoideae: tribes Sophoreae, Podalyrieae, and
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Meidinger, D., and J. Pojar (editors). 1991. Ecosystems of
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Pojar, J. 1999. Fabaceae. Pages 64-180 in Illustrated Flora
of British Columbia. Volume 3. Dicotyledons (Diapensi-
aceae through Onagraceae). Edited by G. W. Douglas, D.
Meidinger, and J. Pojar, British Columbia Ministry of
Environment, Lands and Parks, British Columbia Ministry
of Forests, Victoria, British Columbia.
Ryan, M., and G. W. Douglas. 1994. Status report on the
Seaside birds-foot lotus, Lotus formosissimus Greene. Un-
published report submitted to the Committee on the Status
of Endangered Wildlife in Canada, Ottawa. British Colum-
bia Conservation Data Centre, Victoria, British Columbia.
25 pages.
Stebbins, G. L. 1957. Self-fertilization and population vari-
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onomy. Canadian Journal of Botany 46: 557-583.
Received 24 July 2006
Accepted 24 July 2006
Conservation Evaluation of the Pacific Population of Dwarf Woolly-
heads, Psilocarphus brevissimus var. brevissimus, in Canada’
GEORGE W. DouGLas'!, JENIFER L. PENNY’, and KSENIA BARTON?
' Deceased.
2 Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, PO Box 9993 Stn Prov Govt,
Victoria British Columbia V8W 9R7 Canada
4Current address: 204-4272 Albert Street, Burnaby, British Columbia V5C 2E8 Canada
Douglas, George W., Jenifer L. Penny, and Ksenia Barton. 2006, Conservation evaluation of the Pacific Population of Dwarf
Woolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada. Canadian Field-Naturalist 120(2): 163-168
In Canada, Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, is restricted to the Similkameen River valley,
south of Princeton in southwestern British Columbia and the extreme southeast and southwest corners of Alberta and
Saskatchewan, respectively. This paper deals with the three British Columbia populations which represent the northwestern
limit of the species which ranges from south-central British Columbia, southward in the western United States to Montana,
Idaho, Washington, Oregon, Nevada, Utah, Wyoming , California and Baja California, Mexico. In British Columbia, P. bre-
vissimus is associated with calcareous vernal pools and ephemeral pond edges in large forest openings. This habitat is rare
in the area the few existing populations could easily be extirpated or degraded through slight changes in groundwater levels,
-coalbed methane gas drilling, housing development or recreational vehicles.
Key Words: Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, British Columbia, endangered, distribution,
population size.
|
Dwarf Woolly-heads, Psilocarphus brevissimus vat. (\
_ Nutt.', is a member of a genus of five Q®W
species which occur in the Americas (Cronquist 1950).
It is one of three species occurring in British Columbia es Fen Ne Yy
and Canada (Douglas 1998). Two varieties of P. bre-
=e -
_vissimus, var. brevissimus and var. multiflorus, are rec- a Ce . 7, “4
ognized. The var. multiflorus occurs only in California ae, UB SS
‘(Morefield 1993). Psilocarphus brevissimus var. bre- aS
_vissimus was first recorded in Canada in 1997 (Dou-
_gias et al. 1998a).
__ Psilocarphus brevissimus var. brevissimus is a pros-
trate, matted annual herb with a short taproot (Figure 1;
‘Douglas 1998). The plants are from 8 to 20 cm long.
‘The few leaves are restricted to the stems and are oppo-
site, lance-linear to lance-oblong or lance-triangular,
14-15 mm long and white woolly-hairy. The flower heads
|
jare disciform, lack a pappus, and are usually solitary
‘in the leaf axils or at the tips of the branches and lack
involucres. The receptacular bracts are 2.4-4 mm long,
‘hooded and balloonlike. The achenes are more or less
cylindric, glabrous, nerveless and tipped with a small,
_ 1-2 mm long, offset style.
| In British Columbia herbaria, specimens of P. bre-
\vissimus var. brevissimus may be distinguished from
[" elatior by their prostrate and matted habit and much — Fygure 1. Illustration of Psilocarphus brevissimus var. brevis-
narrower leaves. It is separated from P. tenellus var. simus (Line drawing by Elizabeth J. Steven in Douglas
tenellus by its longer (2.5-4 mm versus 1.3-2.5 mm et al. 1998; 2002a).
}
}
|
The field work for Psilocarphus brevissimus was funded by the British Columbia Conservation Data Centre. The results appear in the British
Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002). This information formed the basis for a Commit-
f
tee on the Status of Endangered Wildlife in Canada status report (Douglas et al. 2003") and the subsequent assessment of Endangered
\(COSEWIC 2003°). This paper includes more recent information that did not appear in the original status report.
‘Taxonomy and nomenclature follows Douglas et al. (1998b; 1998c; 1999; 2001).
163
164
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Locations and population sizes for Psilocarphus brevissimus var. brevissimus south of Princeton, British Columbia.
Collection site Observation Collector/Observer | Number of plants/area
Princeton, Tracey Lake, ca. 0.75 km SE of Lake 1996 Lomer 30/unknown
Princeton, Tracey Lake, ca. 1.3 km ESE of Lake 2002 Douglas and Penny 450/392 m?
2004 Douglas and Smith —7200+500/100 m?
Princeton, Stevenson Lake, ca. 0.75 km SE of 2002 Douglas and Penny 900 000+100 000/570 m2
2004 Douglas and Smith —_11 775/155 m?
long) receptacular bracts and its much narrower leaves.
For many years in Alberta and Saskatchewan, this
species was called P. elatior (Packer 1983; Kershaw
2001; Saskatchewan Conservation Data Centre 2004’).
In 2004, Alberta taxonomists examined material from
both provinces and reidentified them as Psilocarphus
brevissimus var. brevissimus (J. Gould, personal com-
munication). It is quite likely that the “species of con-
cern” ranking given by COSEWIC (2004°) for the
Alberta/Sackatchewan populations of P. elatior will
now be applied to the P. brevissimus populations.
North American and Provincial Ranges
Psilocarphus brevissimus var. brevissimus ranges
from south-central British Columbia, southward in the
western United States to Montana, Idaho, Washington,
Oregon, Nevada, Utah, Wyoming, California and Baja
California, Mexico (Morefield 1993; Cronquist 1994;
Douglas 1998). There are also disjunct populations in
Chile and Argentina (Cronquist 1950, 1955). In Cana-
da, the species is known only from along 2.6 km of
Highway #3 in the Similkameen River valley south
of Princeton in south-central British Columbia (Fig-
ure 2; Douglas 1998; Douglas et al. 2002) and in the
south-eastern and south-western corners of Alberta
and Saskatchewan, respectively.
Habitat
Psilocarphus brevissimus var. brevissimus occurs in
the southern interior of British Columbia in the lower
montane zone in the Interior Douglas-fir biogeocli-
matic zone (Hope et al. 1991). Climatic conditions
are continental, characterized by hot, dry summers, a
fairly long growing season and cool winters. A rain-
shadow effect prevails in this area due to the pres-
ence of the Coast-Cascade Mountains to the west.
Within this zone, the sites occur in a matrix of level
to gently sloping Big Sagebrush (Artemisia tridentata)
shrub/grassland with scattered Ponderosa Pine (Pinus
ponderosa) and Douglas-fir (Pseudotsuga menziesii)
[Nomenclature follows Douglas et al. 1998, 1998c,
1999 and 2001]. The area is at the western edge of the
distribution of open shrub/grassland at that elevation.
Specifically, P. brevissimus var. brevissimus occurs in
calcareous clay soils in vernal pools in large forest
openings dominated by Scouler’s Popcornflower (Pla-
giobothrys scouleri) and Close-flowered Knotweed
(Polygonum polygaloides ssp. confertiflorum (Figure
3). Associates in these sites include One-spike Oat-
grass (Danthonia unispicata), Tiny Mousetail (Myosu-
rus minimus), Carolina Meadow-foxtail (Alopecurus
carolinianus), Common knotweed (Polygonum avic-
ulare), Lowland Cudweed (Gnaphalium palustre), and
Annual Hairgrass (Deschampsia danthonioides). Psilo-
carphus brevissimus var. brevissimus also occurs at
the edge of ephemeral ponds in the same type of sub-
stratum (Figure 4). The vernal pools and ephemeral
pond edges are wet in spring and dry and compacted
in the summer.
Psilocarphus brevissimus var. brevissimus is consid-
ered to be a vernal pool specialist (Schlising and San-
ders 1982; Keeley and Zedler 1998; Bauder 2000).
Keeley and Zedler (1998) define vernal pools as “‘pre-
cipitation-filled seasonal wetlands inundated during
periods when temperature is sufficient for plant growth,
followed by a brief waterlogged-terrestrial stage and
culminating in extreme desiccating soil conditions of
extended duration”. The species is able to outcompete
grassland species due to its tolerance of inundation,
and aquatic/wetland species due to its tolerance of soil
dessication and heat during summer drought (Bauder
2000).
Biology
Little information is available on the biology of
Psilocarphus brevissimus var. brevissimus in British
Columbia. Only basic facts about its life cycle and
reproduction are known.
Keeley and Zedler (1998) recognize four stages in
the annual vernal pool cycle: (1) a wetting phase; (2) an
aquatic or inundation phase; (3) a waterlogged-terres-
trial phase; and (4) the drought phase. In vernal pool
Species, germination is typically initiated during the
wetting or inundation phases. Flowering is usually ini-
tiated during the transition to the waterlogged-terrestrial
phase. In the Princeton area of south-central British
Columbia, this phase occurs during June and fruiting
follows in July during the drought period.
Psilocarphus brevissimus var. brevissimus is an an-
nual herb that likely reproduces primarily by self-pol-
lination. The assumption of a selfing breeding system
is based on the taxon’s lack of pollinator attracting struc-
tures and the proximity of the stigmas to the central
pollen-producing flowers. In addition, the abundance
of woolly hairs and lack of achene pappus appear to
limit pollen dispersal by wind. Another possibility is
that plants set seed without pollination, through asex-
ual reproduction (Cronquist 1950).
2006
Animals may be the primary vectors of seed disper-
sal for Psilocarphus brevissimus var. brevissimus. Birds
may disperse seeds over small and large geographic
areas. The use of vernal pools by various migrating
avian species, particularly waterfowl and shorebirds,
is well documented in other regions (Silveira 1998).
Finally, cottontail rabbits (Sy/vilagus spp.) are poten-
tial agents of dispersal of P. brevissimus var. brevis-
simus seeds. A study in California found that P. bre-
vissimus Var. brevissimus seeds were commonly found
in cottontail rabbit pellets. The excreted seeds germi-
nated easily, suggesting that the lagomorphs may be
important agents of seed dispersal between vernal pools
on a small scale (Zedler and Black 1992). British
Columbia sites for P. brevissimus var. brevissimus
may provide habitat for Nuttall’s Cottontail (Sy/vila-
gus nuttallii), a species that frequents grasslands.
Population Attributes
The three populations of Psilocarphus brevissimus
var. brevissimus were discovered in 1997 in the Simil-
kameen River valley, west of Princeton, British Colum-
bia (Table 1; Douglas et al. 1998a). These populations
are between approximately 0.8 and 2.6 km apart (Fig-
ure 2). During the period from 2002 to 2004, two of
the populations decreased in size by about 25%. Plant
numbers increased at one site from 450 to about 7200
plants and decreased at the other from about 900 000
to about 12000 plants.
Trends for these populations are not well known but
it is obvious that they can be expected to vary marked-
ly due to differences in yearly seed germination and
seedling success. Population sizes of this annual herb
are strongly tied to annual precipitation patterns (Baud-
er 2000) and are typical of vernal pool plants (Griggs
and Jain 1983).
Provincial, National and Global Ranks
Globally, Psilocarphus brevissimus var. brevissimus
has a rank of G4T4 indicating that in most of its range
the plant is secure and frequent to common. Since the
species is restricted to British Columbia it has a nation-
al rank in Canada of N1. Provincially, P. brevissimus
var. brevissimus has been ranked S1 by the Conserva-
tion Data Centre and appears on the British Columbia
Ministry of Sustainable Resource Management red
list (Douglas et al. 2002). This is the most critical rank
that can be applied to species at the provincial level and
indicates that the species is “critically imperiled be-
cause of extreme rarity (typically five or fewer occur-
rences or very few remaining individuals) or because
of some factor(s) making it especially vulnerable to
extirpation or extinction”.
Threats and Protection
The major threats to P. brevissimus var. brevissimus
are through drilling for coalbed methane gas, housing
development and off-road recreational vehicles. In 2002
DOUGLAS, PENNY, AND BARTON: DWARF WOOLLY-HEADS
165
™m—
Ly»
\
\
\
a
BRITISH
COLUMBIA
PRINCETON
FIGURE 2. Distribution of Psilocarphus brevissimus var. bre-
vissimus in British Columbia.
three test holes, targeting coalbed methane gas deposits,
were drilled in the Similkameen valley. If brought into
production the drilling, access roads and pumping sta-
tions could cause major degradation in the area.
Other potential threats result from activities that
are permitted in the Agricultural Land Reserve. The
private property where P. brevissimus var. brevissimus
occurs is on the Agricultural Land Reserve, where pri-
mary land use is agriculture. For instance, some types
of fill/soil removal are allowable without application
to the Agricultural Land Commission (Provincial Agri-
166
THE CANADIAN FIELD-NATURALIST
Vol. 120
FIGURE 3. Calcareous vernal pool habitat of Psilocarphus brevissimus var. brevissimus south of Princeton, British Columbia.
Colorado Rush (Juncus confusus), another rare species in British Columbia, is conspicuous on both sides of the dried-
up vernal pool. The several large plants in the habitat are the introduced grass, American Sloughgrass (Beckmannia
syzigachne).
cultural Land Commission 2003°). On these lands,
development pressures do not appear to be an imme-
diate issue at this time. The Agricultural Land Reserve
status may prevent subdivision development, but does
allow other activities that could also potentially threaten
the populations. In recent years many tracts of Agri-
cultural Land Reserve land in southern British Colum-
bia has been converted to housing developments, shop-
ping malls and golf courses, either by decisions of the
Agricultural Land Commission or very rarely by an
“order in council” by the sitting provincial legislature.
Another threat likely to have a negative impact is
recreational off-road vehicle use. The authors observed
evidence of all-terrain vehicles and dirt bike use in the
area of the sites in 2004. The vernal pools and ephemer-
al pond edges are wet in spring and are susceptible to
permanent changes if disturbed at this.
An additional threat to the population of P. brevis-
simus var. brevissimus is its vulnerability to extirpation
due to the extremely small occupied area and popula-
tion size. Such small populations are at risk of inbreed-
ing depression (Primack 1998). Due to lack of genetic
variation, these small populations are vulnerable to
demographic and environmental variation. Further-
more, suitable habitats for P. brevissimus var. brevis-
simus in the southern interior of British Columbia are
extremely limited and thus opportunities for coloni-
sation are also limited.
Psilocarphus brevissimus vat. brevissimus 1s not for-
mally protected in British Columbia, however, it could
be in the future since this species is a potential candi-
date for listing under the provincial Wildlife Amend-
ment Act (2004). As part of its commitment to the
National Accord (National Accord for the Protection
of Species at Risk), the province is required to take
measures to protect this species. It is on Schedule 1 of
the federal Species At Risk Act and a recovery strategy
is expected to be completed by the summer of 2006.
Evaluation
The British Columbia Conservation Data Centre
considers P. brevissimus var. brevissimus to be threat-
ened/endangered in British Columbia (Douglas et al.
2002) and the Committee on the Status of Endangered
Wildlife in Canada has assessed the species as endan-
gered (COSEWIC 2003*). The three populations are
small and vulnerable to a number of threats. With lim-
ited knowledge of the plant’s biological and ecological
requirements, this species is vulnerable to extirpation
in British Columbia. Without research on growth re-
quirements and further demographic information, the
stability of the present populations will remain un-
known. The limited number of plants also reduces the
potential for genetic variation, which may be necessary
to respond to environmental changes in the future.
2006
Acknowledgments
We thank Frank Lomer for providing information
on his original discovery of Psilocarphus brevissimus
var. brevissimus in 1997 and for his assistance in re-
locating the sites in 2002. Shyanne J. Smith assisted
with field work in 2004.
Documents Cited (marked * in text)
COSEWIC., 2003. COSEWIC assessment and status report
on Dwarf Woolly-heads, Psilocarphus brevissimus var.
brevissimus in Canada, Committee on the Status of Endan-
gered Wildlife in Canada. Ottawa, Ontario. 19 pages. www.
cosewic.gc.ca.
COSEWIC., 2004. Canadian Species at Risk. Committee
on the Status of Endangered Wildlife in Canada. Ottawa,
Ontario. 13 pages. www.cosewic.gc.ca.
Douglas, G. W., J. L. Penny and K. Barton. 2003. COSE-
WIC Status report on Dwarf Woolly-heads, Psilocarphus
brevissimus var. brevissimus in Canada. In COSEWIC as-
sessment and status report on Dwarf Woolly-heads, Psilo-
carphus brevissimus var. brevissimus in Canada. Commit-
tee on the Status of Endangered Wildlife in Canada. Ottawa,
Ontario. 1-19 pages. www.cosewic.gc.ca.
Provincial Agricultural Land Commission. 2003. Agricul-
tural Land Reserve — Frequently Asked Questions. Provin-
cial Agricultural Land Commission. 133-4940 Canada Way,
Burnaby, British Columbia V5G 4K6 Canada. http://www.
alc.goy.be.ca/faq/faq_alr.htm.
Saskatchewan Conservation Data Centre. 2004. Tracked
species list for vascular plants. Saskatchewan Conservation
Data Centre, Fish and Wildlife Branch, Saskatchewan
Environment and Resource Management. Regina, Saskat-
chewan.
Literature Cited
Bauder, E. T. 2000. Inundation effects on small-scale plant
distributions in San Diego, California vernal pools. Aquatic
Ecology 34: 43-61.
Cronquist, A. 1950. A review of the genus Psilocarphus.
Research Studies of the State College of Washington 18:
71-89.
Cronquist, A. 1955. Vascular plants of the Pacific Northwest.
Part 5: Compositae. University of Washington Press, Seat-
tle, Washington. 343 pages.
Cronquist, A. 1994. Intermountain flora-vascular plants of
the Intermountain West, U.S.A. Volume 5: Asterales. New
York Botanical Garden, New York, New York. 496 pages.
Douglas, G. W. 1998. Asteraceae. Pages 96-392 in Illustrat-
ed flora of British Columbia. Volume 1. Gymnosperms and
Dicotyledons (Aceraceae through Asteraceae). Edited by
G. W. Douglas, G. B. Straley, and D. Meidinger. British
Columbia Ministry of Environment, Lands and Parks and
British Columbia Ministry of Forests, Victoria, British
Columbia. 436 pages.
Douglas, G. W., F. Lomer, and Hans L. Roemer. 1998a.
New or rediscovered native vascular plant species in
British Columbia. Canadian Field-Naturalist 112: 276-279.
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare
native vascular plants of British Columbia. Second edi-
tion. Province of British Columbia, Victoria, British Colum-
bia. 359 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illustrated
flora of British Columbia. Volume 4. Dicotyledons (Oroban-
DOUGLAS, PENNY, AND BARTON: DWARF WOOLLY-HEADS
167
oa
or
. ; vy. -
aint of,
* | a
:
a
i
FIGURE 4. The ephemeral pond habitat of Psilocarphus bre-
vissimus Var. brevissimus south of Princeton, British
Columbia. Myosurus minimus, Plagiobothrys scouleri
and Polygonum aviculare are also prominent species
on the calcareous clay soils. The adjacent bands of
vegetation consist mainly of Meadow Barley (Hordewmn
brachyantherum) in the foreground and Common Cat-
tail (7ypha latifolia) in the background.
chaceae through Rubiaceae). British Columbia Ministry
of Environment, Lands and Parks and British Columbia
Ministry of Forests, Victoria, British Columbia. 427 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2001. Illustrat-
ed flora of British Columbia. Volume 7. Monocotyledons
(Orchidaceae to Zosteraceae). British Columbia Ministry
of Sustainable Resource Management and British Colum-
bia Ministry of Forests, Victoria, British Columbia. 379
pages.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1998b.
Illustrated flora of British Columbia. Volume 1. Gymno-
sperms and Dicotyledons (Aceraceae through Asteraceae).
British Columbia Ministry of Environment, Lands and
Parks and British Columbia Ministry of Forests, Victoria,
British Columbia. 436 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998c. Illustrated flora of British Columbia. Volume 2.
Dicotyledons. (Balsaminaceae through Cuscutaceae). Bri-
tish Columbia Ministry of Environment, Lands and Parks
and British Columbia Ministry of Forests, Victoria, British
Columbia. 401 pages.
168
Griggs, F. T., and S. K. Jain. 1983. Conservation of vernal
pool plants in California, I]. Population biology of a rare
and unique grass genus Orcuttia. Biological Conserva-
tion 27: 171-193.
Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson,
W. L. Harper, and B. M. Wikeem. 1991. Interior Douglas-
fir zone. Pages 153-166 in Ecosystems of British Columbia.
Edited by D. Meidinger and J. Pojar. British Columbia
Ministry of Forests Special Report Series Number 6, Vic-
toria, British Columbia. 330 pages.
Keeley, J. E., and P. H. Zedler. 1998. Characterization and
global distribution of vernal pools. Jn Ecology, conserva-
tion, and management of vernal pool ecosystems: Proceed-
ings from a 1996 Conference. Edited by C. W. Witham,
E. T. Bauder, D. Belk, W. R. Ferren Jr., and R. Ornduff.
California Native Plant Society, Sacramento, California.
Kershaw, L. 2001. Rare vascular plants of Alberta. The Uni-
versity of Alberta Press, Edmonton, Alberta. 484 pages.
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son manual: Higher plants of California. Edited by J.C.
Hickman. University of California Press. Berkeley, Cali-
fornia. 1400 pages.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Packer, J. G. 1983. Flora of Alberta. Second edition. Univer-
sity of Toronto Press, Toronto, Ontario. 687 pages.
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Schlising, R. A., and E. L. Sanders. 1982. Quantitative analy-
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American Journal of Botany 69: 734-742.
Silveira, J. G. 1998. Avian uses of vernal pools and implica-
tions for conservation practice. Jn Ecology, conservation,
and management of vernal pool ecosystems. Proceedings
from a 1996 Conference. Edited by C. W. Witham, E. T.
Bauder, D. Belk, W. R. Ferren Jr., and R. Omduff. Califor-
nia Native Plant Society, Sacramento, California.
Zedler, P. H., and C. Black. 1992. Seed dispersal by a gen-
eralized herbivore: rabbits as dispersal vectors in a semi-
arid California vernal pool landscape. American Midland
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Received 8 October 2002
Accepted 20 March 2006
macrophyllum, in Canada*
GEORGE W. DouG-Las! and JULIE DEsROSIERS2
' Deceased.
in Canada at this time.
The Pacific Rhododendron, Rhododendron macro-
phyllum D. Don ex G. Don’, is a member of a genus
of almost 1000 species occurring mainly in north-
eastern Asia (Wallace 1993). It is one of three species
| occurring in British Columbia (Meidinger 1999) and
four occurring in Canada (Scoggan 1979).
Rhododendron macrophyllum was first reported by
naturalist Archibald Menzies in 1792 in his “Journal
of Vancouver’s Voyage”, and subsequently described
by David Don in 1834. Joseph Hooker also described
plants collected during Vancouver’s voyages, and in
1855 he published a description of this rhododendron
under the species name californicum. This name was
in common use by botanists for close to a century.
Rhododendron macrophyllum is an erect to spread-
ing, branched shrub, 1-8 m tall (Meidinger 1999; Fig-
ure 1). The evergreen leaves are alternate, often appear-
|ing whorled, ovate or elliptical, 8-20 cm long, 3-6 cm
|wide, leathery, dark green above, paler beneath and
glabrous, or nearly so. The pale-pink to deep rose-
purplish corollas are tubular bell-shaped, spotted red-
|brown and 2.5-4 cm long and up to 5 cm across. The
fruits consist of woody capsules 1.5-2 cm long with
numerous minute seeds. It is easily separated from the
other two Rhododendron species in British Columbia.
Lapland Rosebay (Rhododendron lapponicum) is a
northern plant with much shorter habit and smaller
leaves, while White-flowered Rhododendron (R. albi-
florum) is a higher elevation species with white flowers.
Conservation Evaluation of Pacific Rhododendron. Rhododendron
* 4254 Garden Drive, Burnaby, British Columbia VSG 4G6 Canada
Douglas, George W., and Julie Desrosiers. 2006. Conservation evaluation of Pacific Rhododendron, Rhododendron
macrophyllum, in Canada. Canadian Field-Naturalist 120(2); 169-174.
In Canada, Pacific Rhododendron (Rhododendron macrophyllum) is restricted to the Skagit River drainage and southeastern
_ Vancouver Island in southwestern British Columbia. These populations represent the northern limits of the species, which
ranges south to northern California. In British Columbia, Rhododendron macrophyllum is usually associated with coniferous
forests. Until recently this species was considered to be rare. Results of this study indicate that the populations in the Skagit
River watershed are large, extensive and mostly contained in protected areas, thus the Species is not considered to be at risk
Key Words: Pacific Rhododendron, Rhododendron macrophyllum, distribution, conservation evaluation, British Columbia
North American and Provincial Ranges
Rhododendron macrophyllum is restricted to coastal
habitats from northern California north through Oregon
and Washington to southwestern British Columbia
(Hitchcock et al. 1959). In Canada, it occurs in south-
western British Columbia in the western Cascade Moun-
tains (Skagit River valley) and on southeastern Van-
couver Island (Shawnigan and Rhododendron lakes)
[Meidinger 1999; Douglas et al. 2002; Figure 2}.
Habitat
Skagit River Valley
The Rhododendron macrophyllum populations in
the Skagit River valley occur on well-drained, coarse-
textured, gravelly, dry soils overlain by fibremors. Ele-
vations range from 500 to 870 metres.
About half of the Rhododendron populations in the
Skagit River valley are within the Interior Douglas-fir
(IDF) Zone while the remaining are within the Coastal
Western Hemlock (CWH) Zone (Hope et al. 1991;
Pojar et al. 1991). Summaries of all plant communities
within these zones are taken from Desrosiers and Cad-
rin (1993). The four plant communities within the
Interior Douglas-fir (IDF) Zone are described below:
(a) Douglas-fir/Falsebox/Feathermoss (Pseudotsuga menziesii
Paxistima myrsinites/Pleurozium schreberi) community.
Sites for this community are located in the dry, mid-
dle part of the Skagit River Valley, on the flats of the
The field work for the Rhododendron macrophyllum project was funded by the British Columbia Conservation Data Centre and The Skagit
avironmental Endowment Commission. The results appear in the British Columbia Conservation Data Centre database and a Skagit Envi-
onmental Endowment Commission report (Desrosiers and Cadrin 1993"). This information formed the basis for a Committee on the Status
of Endangered Wildlife in Canada status report (Desrosiers and Douglas 1997") and the current assessment of nor at risk (COSEWIC 2006’).
Taxonomy and nomenclature follows Douglas et al. (1998a, 1998b, 1999a, 1999b, 2000, 2001) and Schofield (1992).
169
170
THE CANADIAN FIELD-NATURALIST
Vol. 120
FiGurE 1. illustration of Rhododendron macrophyllum. (Line drawing by Lora May Richards)
valley floor. These are the driest IDF sites and were
mid-range in terms of their nutrient regime. The 80-
115 year old stands are dominated by Pseudotsuga
menziesii, Paxistima myrsinites, and Pleurozium schre-
beri. Other important species include Rhododendron
macrophyllum, Lodgepole Pine (Pinus contorta), White
Hawkweed (Hieracium albiflorum), Heart-leaved Tway-
blade (Listera cordata) and Cow-wheat (Melampyrum
lineare var. lineare). Rhododendron macrophyllum had
an average cover of 27% in this community.
(b) Douglas-fir — Western Redcedar/Beaked Hazelnut (Pseu-
dotsuga menziesii-Thuja plicata/Corylus cornuta) com-
munity.
The lowest elevations in the lower part of the valley
contained these sites. They tended to be intermediate,
both environmentally and floristically, for the IDF in
this area. Pseudotsuga menziesii, Thuja plicata, and
Corylus cornuta were dominants in these 80-100 year
old stands. Cover values for Thuja plicata, Western
Hemlock (Tsuga heterophylla), Paxistima myrsinites
1
|
'
n
2006
BRITISH
COLUMBIA
Koksilah
River
VICTORIA
DOUGLAS AND DESROSIERS: PACIFIC RHODODENDRON
171]
-=--
as ly
‘MANNING
\. PROVINCIAL
\ PARK
BB Ecological Reserve
---- Boundary
=—— Road
FIGURE 2. The location of Rhododendron macropyhilum sites in British Columbia.
and Dull Oregon-grape (Mahonia nervosa) were high-
est in these sites. Average cover of Rhododendron
macrophyllum was 3%.
(c) Douglas-fir — Western Redcedar/Vine Maple (Pseudot-
suga menziesii-Thuja plicata/Acer circinatum) community.
This community is found at a higher elevation than
other IDF sites, on the east-facing slopes of the Rhodo-
dendron Ecological Reserve. The sites were mid-range
in term of moisture regime and were nutrient-poor.
Pseudotsuga menziesii, Thuja plicata, Acer circinatum
and Step Moss (Hylocomium splendens) were the dom-
inant species. Douglas Maple (Acer glabrum var. dou-
glasii), Thimbleberry (Rubus parviflorus var. parvi-
florus), Oceanspray (Holodiscus discolor), Hooker's
Fairybells (Prosartes hookeri var. oregana), Fireweed
(Epilobium angustifolium), and Lanky Moss (Rhy-
tidiadelphus loreus) were also important. Rhododen-
dron macrophyllum had an average cover of 3%.
(d) Pacific Rhododendron/Westerm Tea-berry/Cladonia (Rho-
dodendron macrophyllum/Gaultheria ovatifolia/Cladonia)
community.
This Rhododendron community occurs on the east-
ern section of the Rhododendron Ecological Reserve
as well as on the west side of the Skagit River. These
sites were among the driest and most nutrient-poor of
all sites sampled. The stands were 105-115 years old.
172
Tree cover in these plots was extremely low. Pseudo-
tsuga menziesii had an average cover of 8% while West-
em White Pine (Pinus monticola) had an average cover
of 7%. Average cover of Rhododendron macrophyl-
lum was the highest of all plots at 70%. Species of
importance in this community included Soopolallie
(Shepherdia Canadensis), Paxistima myrsinites, Twin-
flower (Linnaea borealis), Mahonia nervosa, Kinnikin-
nick (Arctostaphylos uva-ursi) and species of lichen
(Cladonia spp.).
All of the plots sampled in the northern half of the
study area fell within a single community type in the
Coastal Western Hemlock (CWH) Zone. This com-
munity may be summarized as follows:
Douglas-fir — Western Hemlock/Falsebox (Pseudotsuga men-
ziesti-Tsuga heterophylla/Paxistima myrsinites ) community.
Compared to IDF sites the CWH sites have higher
moisture regimes, lower nutrient levels and higher tree
cover values. In addition to Pseudotsuga menziesii,
Tsuga heterophylla and Paxistima myrsinites other
important species include Amabilis Fir (Abies ama-
bilis), Grouseberry (Vaccinium scoparium) and Hylo-
comium splendens. The stands ranged from 80-150
years old. Average cover of Rhododendron macro-
phyllum was 18%.
Southeastern Vancouver Island
The Rhododendron sites on southeastern Vancouver
Island are within the Coastal Douglas-fir Zone (Neuz-
dorfer et al. 1991). The populations east of Shawnigan
Lake were found in the following plant community:
Western Redcedar/Pacific Rhododendron-Alaskan Blueberry/
Step Moss (Thuja plicata/Rhododendron macrophyllum-
Vaccinium alaskense/Hylocomium splendens) community.
This community was located on a moderate, south-
west slope at an elevation of 490 m approximately
16 km west of Shawnigan Lake. The small (550 m7)
area on which Rhododendron occurred was dominated
by 40 m tall Thuja plicata. The shrub layer was well-
developed with Rhododendron macrophyllum, Vac-
cinium alaskense, Red Huckleberry (V. parvifolium)
and Salal (Gaultheria shallon) dominating. A dense
layer of mosses and lichens was dominated by Hylo-
comium splendens.
At Rhododendron Lake three markedly different
communities occur in bands around Rhododendron
Lake. A very wet band of vegetation (about 3-5 m
wide), dominated mainly by Sweet Gale (Myrica gale),
occurs around the immediate edge of the lake. Adja-
cent to the latter band is another, slightly drier but still
very moist 3-10 m band of vegetation containing Myri-
ca gale and Rhododendron macrophyllum. This satu-
rated habitat is atypical for Rhododendron macrophyl-
lum. A wider band (5-25 m wide) of forested vegetation
occurs upslope from the lakeside communities. Approx-
imately nine ha was covered by Rhododendron plants
in the latter two vegetation bands. The two communi-
ties containing Rhododendron may be summarized as
follows:
THE CANADIAN FIELD-NATURALIST
Vol. 120
(a) Pacific Rhododendron — Sweet Gale/Salal (Rhododendron
macrophyllum-Myrica gale/Gaultheria shallon) community.
This community occurs in a 3-10 m wide band that
extends for about 300 m along the south side of the
lake. Soils are constantly wet with the presence of
Skunk Cabbage (Lysichitum americanum) and Sphag-
num sp. indicating the hydric nature of the site. This
site had been logged many years earlier. It appears all
trees cut were Yellow-cedar (Chamaecyparis nootka-
tensis) and would have had about a 40-60% average
cover. At the present time, Rhododendron macrophyl-
lum has an average cover of 47% with only Gaultheria
shallon being more abundant (average cover of 57%).
Bracken Fern (Preridium aquilinum), Labrador Tea
(Ledum groenlandicum) and Myrica gale were also
important species in the community.
(b) Western Hemlock/Pacific Rhododendron/Salal (Tsuga
heterophylla/Rhododendron macrophyllum/Gaultheria shal-
lon) community.
Upslope from the previous community is a forested
community dominated by Tsuga heterophylla. The
site has been partially logged and was previously dom-
inated by Thuja plicata. Other coniferous species pres-
ent, but in low numbers, include Chamaecyparis nootka-
tensis, Thuja plicata, Pinus monticola and Pseudotsuga
menziesii. Important understory species include Gaul-
theria shallon, Rhododendron macrophyllum and
Hylocomium splendens. Rhododendron has an aver-
age cover of 61%.
Biology and Ecology
Research on the biology and ecology of Rhododen-
dron macrophyllum has been limited. The majority
(60%) of populations in the Skagit Valley showed good
to very good vigour. One in three populations had
young genets less than one metre in height. These
appear, for the most part, to have germinated on rot-
ten logs or thick moss mats. Asexual reproduction,
by layering, appears to be much more common than
sexual reproduction.
Many sites showed evidence of fire on the domi-
nant trees and charcoal layers were often found in the
soil profile. This history of fire and its importance to
the maintenance of Rhododendron macrophyllum pop-
ulations was also noted in an Olympic National For-
est study (Henderson et al. 1989).
Population Attributes
Approximately 200 populations of Rhododendron
macrophyllum have been observed at 15 ecosites in
the Skagit River valley over a distance of 25 km (Fig-
ure 2). Of these, 92 were examined in detail at 13 eco-
sites for this study (Table 1). Populations ranged in
size from 4 m? to about 200 000 m°’. The latter area
(part of the Rhododendron Ecological Reserve) had an
extrapolated count of approximately a million canes.
Two populations on Vancouver Island were also exam-
ined. Counts at one of them (Rhododendron Lake)
revealed 400 plants and approximately 9000 canes.
2006
DOUGLAS AND DESROSIERS: PACIFIC RHODODENDRON
173
TABLE |. Locations and population sizes of Rhododendron macrophyllum in south-western British Columbia
Number of
Populations
Map Location Number(s)/
Ecosite Name
Manning Provincial Park
1. Cayuse Flats |
2. Silverdaisy Trail 3
3. Rhododendron Flats 4
Skagit Valley Provincial Park
4. Upper Skagit Area 3
5. Klesilkwa River 6
6. Valley Floor North
7. Pinus contorta Terrace North
8. Pinus contorta Terrace South
9. Shawatum Area
10. Rhododendron Ecological Reserve, East
11. Rhododendron Ecological Reserve, West
12. Nepopekum area
13. Galene Lakes Trail
Vancouver Island
1. Rhododendron Lake
2. Shawnigan Lake |
n
WAwWe LK WY
Approximate Area
of Populations (m*)
Approximate Total
Number of Canes
300 1 000
250 1 750
150 2 400
12 600
360 600
400 2 100
3 450 no count
920 4 000
140 300
200 000 1 000 000
40 550
60 1 000
100 1 250
90 000 9 000
550 no count
' Detailed counts of canes per plant were made for 50 plants at both Cayuse Flats and Rhododendron Lake. Numbers of canes
per plant ranged from 18 to 24 at Cayuse Flats and 20 to 25 at Rhododendron Lake (Desrosiers and Douglas 1997*).
The intensive inventory work conducted by the
British Columbia Conservation Data Centre resulted in
the documentation of thousands of plants in the Skagit
Valley (Table 1). Prior to this work the species had been
reported as a rare plant in British Columbia (Straley
et al 1985; Argus and Prior 1990).
Locations of Rhododendron macrophyllum in the
nearby Chilliwack River valley were mentioned by
Slaney (1971*); however, no references or supporting
information were provided in that document. It is pos-
sible that these populations could be extirpated due
to development. In addition, Rhododendron macro-
phyllum was reported from Lightning Lake in Man-
ning Park (Carl et al. 1952); this population was not
verified by the authors or others working in that area.
Provincial, National and Global Ranks
Provincially R. macrophyllum is ranked S4 (Douglas
et al. 2002) which indicates that it is “rare or uncommon
(typically 21 to 100 occurrences); may be susceptible to
large-scale disturbances; e.g., may have lost extensive
peripheral populations”. Since the species is restricted
to British Columbia, the National rank is N1. Globally,
Rhododendron macrophyllum is ranked GS. This rank-
ing indicates that, on a global scale, the plant is con-
sidered “frequent to common or very common; demon-
strably secure and essentially ineradicable under present
conditions”.
Threats and Protection
Habitat destruction, due to logging and road build-
ing, likely eliminated some populations of Rhododen-
dron macrophyllum, especially in the Skagit River
area. The establishment of Manning Provincial Park
and the more recent Skagit Valley Provincial Park
has eliminated most threats to R. macrophyllum. On
Vancouver Island, where the two populations are locat-
ed within timber harvesting areas, small informal pro-
tected areas have been established.
In the Skagit River valley it was evident that fire had
previously swept through most of the ecosites (Des-
rosiers and Cadrin 1993"). Whenever possible, natural
fires are quickly extinguished in the Cascades Moun-
tains. If this control is successful over long periods of
time, fuel loads may build up resulting in extremely
hazardous conditions. It is, therefore, possible that pop-
ulation sizes may be temporarily reduced from time
to time after large burns.
Rhododendron macrophyllum receives protection
since all but the Vancouver Island populations occur
in provincial parks. The Vancouver Island populations
have informal protection set up by the timber compa-
nies that own the properties
Until recently there was formal protection under the
Dogwood, Rhododendron and Trillium Protection Act.
This act was repealed in 2002 due to our greater knowl-
edge of the abundance of the species involved and the
fact that sufficiently large numbers of their populations
occur in protected areas.
Evaluation
Rhododendron macrophyllum was considered rare
in Canada for many years (Straley et al. 1985; Argus
and Pryor 1990). The present study, however, indicates
that the species is not rare and is much more abun-
dant than previously thought. Most of the extant sites
174
are secure since they are contained either within E.
C. Manning Provincial Park, the adjacent Skagit Val-
ley Provincial Park or in several ecological reserves.
Acknowledgments
We thank Shane Ford for locating and examining
the Shawnigan Lake population and Jenifer L. Penny
for office assistance.
Documents Cited (marked * in text)
COSEWIC. 2004. Canadian Species at Risk. Committee
on the Status of Endangered Wildlife in Canada, Ottawa,
Ontario. www.cosewic.gc.ca.
Desrosiers, J., and C. M. Cadrin. 1993. The Pacific Rho-
dodendron (Rhododendron macrophyllum D. Don ex G.
Don) communities of the Skagit River watershed, British
Columbia. Unpublished Report, The Skagit Environmen-
tal Endowment Commission. 27 pages.
Desrosiers, J., and G. W. Douglas. 1997. Status Report on
the Pacific Rhododendron, Rhododendron macrophyllum
D. Don ex G. Don (Ericaceae), in Canada. Ottawa, Ontario.
22 pages.
Slaney, F. F. 1971. Rhododendrons in the Skagit Valley. The
Canadian Skagit Environmental Newsletter Volume 7, 4
pages.
Literature Cited
Argus, G. W., and K. M. Pryer. 1990. Rare vascular plants
in Canada — our natural heritage. Canadian. Museum of
Nature, Ottawa, Ontario. 191 pages.
Carl, G. C., C. J. Guiguet, and G. A. Hardy. 1952. A natural
history survey of the Manning Park area, British Columbia.
British Columbia Provincial Museum Occasional Paper 9.
Victoria, British Columbia.
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002.
Rare native vascular plants of British Columbia. Second
edition. British Columbia Ministry of Sustainable Resource
Management, Victoria, British Columbia. 359 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus-
trated flora of British Columbia. Volume 3. Dicotyledons
(Diapensiaceae through Onagraceae). British Columbia
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia.
423 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus-
trated flora of British Columbia. Volume 4. Dicotyledons
(Orobanchaceae through Rubiaceae). British Columbia
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia.
427 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illustrated
flora of British Columbia. Volume 5. Dicotyledons (Sali-
caceae to Zygophyllaceae) and Pteridophytes. British
Columbia Ministry of Environment, Lands and Parks and
British Columbia Ministry of Forests, Victoria, British
Columbia. 389 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2001. Illustrat-
ed flora of British Columbia. Volume 7. Monocotyledons
(Orchidaceae to Zosteraceae). British Columbia Ministry
of Sustainable Resource Management and British Columbia
Ministry of Forests, Victoria, British Columbia. 379 pages.
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Douglas, G. W., G. B. Straley, D. Meidinger and J. Pojar.
1998a. Illustrated flora of British Columbia. Volume 1.
Gymnosperms and Dicotyledons. (Aceraceae through
Asteraceae). British Columbia Ministry of Environment,
Lands and Parks and British Columbia Ministry of Forests,
Victoria, British Columbia. 436 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998b. Illustrated flora of British Columbia. Volume 2.
Dicotyledons. (Balsaminaceae through Cuscutaceae). Bri-
tish Columbia Ministry of Environment, Lands and Parks
and British Columbia Ministry of Forests, Victoria, Bri-
tish Columbia. 401 pages.
Henderson, J. A., D. H. Peter, R. D. Lesher, and D. C.
Shaw. 1989. Forested plant associations of the Olympic
National Forest. United States Department of Agriculture,
Forest Service, Pacific Northwest Region, Ecological Tech-
nical Paper Report 001-88, Portland, Oregon. 502 pages.
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1959. Vascular plants of the Pacific Northwest.
Part 4: Ericaceae through Campanulaceae. University of
Washington Press. Seattle, Washington. 510 pages.
Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson,
W. L. Harper, and B. M. Wikeem. 1991. Interior Dou-
glas-fir zone. Pages 153-166 in Ecosystems of British
Columbia. Edited by D. Meidinger and J. Pojar. British
Columbia Ministry of Forests Special Report Series (6)
Victoria, British Columbia. 330 pages.
Meidinger, D. 1999. Ericaceae. Pages 14-55 in Illustrated
Flora of British Columbia. Volume 3 — Dicotyledons (Dia-
pensiaceae through Onagraceae). Edited by G. W Douglas,
D. Meidinger and J. Pojar. British Columbia Ministry or
Environment, Lands and Parks, British Columbia Ministry
of Forests, Victoria, British Columbia. 423 pages.
Nuszdorfer, F. C., K. Klinka, and D. A. Demarchi. 1991.
Coastal Douglas-fir zone. Pages 95-112 in Ecosystems of
British Columbia. Edited by D. Meidinger and J. Pojar.
British Columbia Ministry of Forests Special Report Series
(6), Victoria British Columbia. 330 pages.
Pojar, J.. K. Klinka, and D. A. Demarchi. 1991. Coastal
Western Hemlock Zone. Pages 95-111 in Ecosystems of
British Columbia. Edited by D. Meidinger and J. Pojar.
British Columbia Ministry of Forests Special Report Series
(6), Victoria, British Columbia. 330 pages.
Schofield, W. B. 1992. Some common mosses of British
Columbia. Royal British Columbia Museum. 394 pages.
Scoggan, H. J. 1979. The flora of Canada. Part 4 — Dicotyle-
doneae (Loasaceae to Compositae). National Museum of
Natural Sciences Publications in Botany (7). Ottawa, On-
tario. Pages 1117-1626.
Straley, G. B., R. L. Taylor, and G. W. Douglas. 1985. The :
rare vascular plants of British Columbia. National Muse- -
um of Natural Sciences. Syllogeus (59), Ottawa, Ontario. .
165 pages.
Wallace, G. D. 1993. Ericaceae. Pages 544-567 in The Jep-
son Manual — Higher plants of California. Edited by J.C.
Hickman. University of California Press, Berkeley, Cali-
fornia. 1400 pages.
Received 14 October 2002
Accepted 20 March 2006
Conservation Evaluation of Slender Collomia, Collomia tenella. in
Canada*
GEORGE W. DouG-as'! and JENIFER L. PENNY?
' Deceased.
2 Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, PO Box 9993 Stn Prov Govr,
Victoria, British Columbia V8W 9R7
Douglas, George W., and Jenifer L. Penny. 2006. Conservation evaluation of Slender Collomia, Collomia tenella, in Canada
Canadian Field-Naturalist 120(2): 175-178.
In Canada, Slender Collomia, Collomia tenella, is restricted to the Princeton area in southwestern British Columbia. The single
population represents the northern limits of the species, which ranges from southwestern British Columbia, south in the
western United States to Wyoming, Utah, Nevada and Oregon. In British Columbia, C. tenella is associated with an eroded section
of a steeply sloping, southeast-facing sandy ridge. Population numbers fluctuate and in some years plants fail to appear. The
major threats to C. tenella are through drilling for coalbed methane gas, sand removal for road construction, housing development
and off-road recreational vehicles.
Key Words: Slender Collomia, Collomia tenella, endangered, distribution, population size, British Columbia.
Slender Collomia, Collomia tenella A. Gray’ (Pole-
- moniaceae), is member of a genus of about 13 species
| found in North and South America (Hitchcock et al.
1959). Four species occur in British Columbia and
Canada (Scoggan 1979; Pojar 1999). Collomia tenella
_ was first recorded in Canada by Douglas et al. (1998a).
_ Collomia tenella is an ascending to spreading, freely
branched, annual, tap-rooted herb up to 15 cm tall
| (Figure 1; Pojar 1999). The leaves are alternate, lin-
| ear, entire, 1-5 cm long and 1.5 mm wide. Flowers
are single or in pairs at the branch tips, in the leaf axils
or at the forks of the branches. The pinkish to white
corollas are five-lobed. The calyces, which bow out
and often form purplish knobs at the sinuses, have 1-
_ 2mm long, triangular teeth. The fruits consist of cap-
| sules with 1-seeded chambers; the seeds become sticky
_ when moistened.
}
| North American and Provincial Ranges
Collomia tenella ranges from southwestern British
Columbia, south in the western United States to Wyo-
/ming, Utah, Nevada and Oregon (Hitchcock et al.
'
h|
1959). In Canada, C. tenella is known only from the
| Princeton area in British Columbia (Figure 2; Pojar
| 1999; Douglas 2002a). Ficure |. Illustration of Collomia tenella (line drawing from
Douglas et al. 1999b).
‘ Habitat
| Collomia tenella occurs in the southwestern interi- (Hope et al. 1991). Climatic conditions in this region
\ai of British Columbia, along the Similkameen River are continental, characterized by hot, dry summers, a
_| valley in the Interior Douglas-fir biogeoclimatic zone fairly long growing season, and cool winters. A rain-
|
1
i]
{
1
)
* The field work for the Collomia tenella project was funded by the British Columbia Conservation Data Centre. The results appear in the
| British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis fora
Committee on the Status of Endangered Wildlife in Canada status report (Douglas and Penny 2003*) and the subsequent assessment of
"ae (COSEWIC 2003). This paper includes more recent information that did not appear in the original status report.
‘" Taxonomy and nomenclature follow Douglas et al. (1998b, 1999a, 1999b, 2000, 2001).
iS)
176
shadow effect prevails in this area due to the pres-
ence of the Coast-Cascade Mountains to the west.
Within this region, Collomia tenella occurs on an
eroded, steeply-sloped, southeast-facing section of a
sandy ridge (Figure 3). The sandy ridge, formed by flu-
vial processes during recent glaciation, consists of fine-
textured sands. The eroded sections of the slopes are
sparsely vegetated with about 20% cover. The vegeta-
tion consists of a variety of herbs and shrubs including
Arrow-leaved Balsamroot (Balsamorhiza sagittata),
Timber Milk-vetch (Astragalus miser), Narrow-leaved
Collomia (Collomia linearis), Thread-leaved Phacelia
(Phacelia linearis), Silky Lupine (Lupinus sericeus),
Saskatoon (Amelanchier alnifolia), Dalmation Toad-
flax (Linaria genistifolia ssp. dalmatica), Cheatgrass
(Bromus tectorum), and Bluebunch Wheatgrass (Pseu-
doroegneria spicata). Scattered Douglas-fir (Pseudot-
suga menziesii) and Ponderosa Pine (Pinus ponderosa)
also occur on the ridge (Figure 3).
Biology
There is no information available on the biology and
ecology of Collomia tenella in the literature. Since it is
an annual plant it appears to be absent in some years
if environmental conditions for seed germination and/
or seedling survival are not favourable. Other annual
members of the genus are self-compatible and self-
pollinating (Wilken 1993). Seeds may be dispersed
by animals since they are sticky when moistened.
Population Attributes
The single population of Collomia tenella, covering
an area of less than 60 m?, was discovered in 1997 in
the Similkameen River valley, west of Princeton, British
Columbia (Figure 2; Douglas et al. 1998a). Counts for
the population in 1997, 2000, 2002, 2003 and 2004
were 10, 1,0, 127 and 0, respectively. In 2003, the area
occupied was ca. 56 square metres, with plants in four
groups of 56, 67, 1 and 3 plants. These fluctuations
indicate that seed germination and/or seedling success
of this annual species is highly variable and thus, trends
are difficult to determine. A fairly thorough search in
adjacent areas in southern British Columbia have not
revealed similar habitats or other populations of this
species.
Provincial, National and Global Ranks
The British Columbia Conservation Data Centre
has ranked Collomia tenella as S1 and placed it on
the British Columbia Ministry of Environment Red list
(Douglas et al. 2002a). This is the most critical cate-
gory for imperilled rare native vascular plants in British
Columbia. A rank of S1 is considered “critically imper-
illed because of extreme rarity (5 or fewer occurrences
or very few remaining individuals) or because of some
factors making it especially vulnerable to extirpation
or extinction” (Douglas et al. 2002a). Since the species
is restricted in Canada to British Columbia, the Nation-
THE CANADIAN FIELD-NATURALIST
Vol. 120
} BRITISH
¢ — COLUMBIA
FIGURE 2. Distribution of Collomia tenella in British Columbia.
al rank is N1. Globally, C. tenella is ranked “G4?” or
apparently secure but with a range of uncertainty
which includes the possibility it is more vulnerable
or conversely that is more widespread, abundant, and
secure.
Threats and Protection
The major threats to Collomia tenella are through
drilling for coalbed methane gas, sand removal for road
construction, housing development and off-road recre-
ational vehicles. In 2002, three test holes targeting coal-
bed methane gas deposits were drilled in the Simil-
2006
DOUGLAS AND PENNY: SLENDER COLLOMIA
177
FIGURE 3. Collomia tenella habitat occurs on the eroded section near the top of this sandy ridge south of Princeton, British
Columbia. Prominent trees on the ridge are Pseudotsuga menziesii and Pinus ponderosa.
kameen area. If brought into production, the drilling,
access roads and pumping stations could cause exten-
sive degradation to the natural habitat in the area.
Other potential threats result from activities that
are permitted in the Agriculture Land Reserve. The
private property where C. tenella occurs is in the
Agricultural Land Reserve, where primary land use is
agriculture. For instance, some types of fill/soil removal
are allowable without application to the Agricultural
Land Commission (Provincial Agricultural Land Com-
mission 2003"). On these lands, development pressures
may not appear to be an issue at this time. The Agri-
cultural Land Reserve status may prevent subdivision
development, but does allow other activities that could
also potentially threaten the populations. In recent
years many tracts of Agricultural Land Reserve land
in southern British Columbia has been converted to
housing developments, shopping malls and golf cours-
es, either by decisions of the Agricultural Land Com-
mission or very rarely by an ‘order in council’ by the
sitting provincial legislature.
Another threat likely to have a negative impact is
recreational off-road vehicle use. The authors observed
evidence of all-terrain vehicles and dirt bike use in the
area of the sites in 2002. The steep and relatively unsta-
ble slopes where C. tenella occurs are highly suscep-
tible to disturbance but at the same time, present a
desirable challenge for off-roaders.
Although weeds are not a major threat to C. tenella
populations, weed control activities do constitute a
minor potential threat to populations. Under the Weed
Control Act, an occupier must control noxious weeds
growing or located on land and premises. Marginally
specific chemical weed control compounds that kill
broad-leaved plant species would likely also kill C.
tenella. Noxious weeds such as Dalmation Toadflax
(Linaria genistifolia var. dalmatica) and Cheatgrass
(Bromus tectorum) occur in the C. tenella habitat and
could increase. Currently the habitat is not highly in-
vaded, but a variety of other introductions could also
eventually take root there as they have in adjacent habi-
tats in the surrounding landscape. Therefore, the eco-
logical integrity of the C. tenella habitat could even-
tually become as comprised as some of these other
surrounding habitats are.
An additional threat to the population of C. renella
is its vulnerability to extirpation due to the extremely
small occupied area and population size. Such small
populations are at risk of inbreeding depression (Pri-
mack 1998). Due to lack of genetic variation, these
small populations are vulnerable to demographic and
environmental variation. Furthermore, suitable habitats
for C. tenella in the southern interior of British Colum-
bia are extremely limited and thus, opportunities for
colonisation are also limited.
178
Collomia tenella is on Schedule 1 of the federal
Species at Risk Act. As part of its commitment to the
National Accord (National Accord for the Protection
of Species at Risk), the province is required to take
measures to protect this species. A recovery strategy
will be required under this commitment. Additionally,
C. tenella is a potential candidate for listing under the
provincial Wildlife Amendment Act.
Evaluation
The British Columbia Conservation Data Centre con-
siders Collomia tenella to be Endangered in British
Columbia (Douglas et al. 2002a) and the Committee
on the Status of Endangered Wildlife in Canada has
also assessed the species as Endangered (COSEWIC
2003*; Douglas 2005"). The single population is small
and vulnerable to a number of threats. The small num-
ber of plants also reduces the potential for genetic vari-
ation which may be necessary to respond to environ-
mental changes in the future. Without research on
growth requirements and more demographic informa-
tion, the stability of the present populations will remain
unknown, but the population appears to be quite vul-
nerable.
Acknowledgments
We thank Frank Lomer for providing information
on his original discovery of Collomia tenella in 1997
and for his help in finding the site in 2002. Thanks
also to Shyanne J. Smith who assisted with fieldwork
in 2004.
Documents Cited (marked * in text)
COSEWIC. 2003. COSEWIC assessment and status report
on Slender Collomia, Collomia tenella in Canada. Com-
mittee on the Status of Endangered Wildlife in Canada,
Ottawa, Ontario. 13 pages. www.cosewic.gc.ca.
Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status
report on Slender Collomia, Collomia tenella in Canada,
in COSEWIC assessment and status report on Slender
Collomia, Collomia tenella in Canada. Committee on the
Status of Endangered Wildlife in Canada, Ottawa, Ontario.
13 pages. www.cosewic.gc.ca.
Provincial Agricultural Land Commission. 2003. Agricul-
tural Land Reserve — Frequently Asked Questions. Provin-
cial Agricultural Land Commission, 133-4940 Canada Way,
Burnaby, British Columbia V5G 4K6 Canada (Available
at: http://www.alc.gov.bc.ca/faq/faq_alr.htm).
Literature Cited
Douglas, G. W., F. Lomer, and Hans L. Roemer. 1998a. New
or rediscovered native vascular plant species in British
Columbia. Canadian Field-Naturalist 112: 276-279.
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare
native vascular plants of British Columbia. Second edition.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Province of British Columbia, Victoria, British Columbia.
358 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus-
trated flora of British Columbia. Volume 3. Dicotyledons
(Diapensiaceae through Onagraceae). British Columbia
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia.
423 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus-
trated flora of British Columbia. Volume 4. Dicotyledons
(Orobanchaceae through Rubiaceae). British Columbia
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia.
427 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illus-
trated flora of British Columbia. Volume 5. Dicotyledons
(Salicaceae to Zygophyllaceae) and Pteridophytes. British
Columbia Ministry of Environment, Lands and Parks and
British Columbia Ministry of Forests, Victoria, British
Columbia. 389 pages.
Douglas, G.W., D. Meidinger, and J. Pojar 2001. Illustrat-
ed flora of British Columbia. Volume 7. Monocotyledons
(Orchidaceae to Zosteraceae). British Columbia Ministry
of Sustainable Resource Management and British Columbia
Ministry of Forests, Victoria, British Columbia. 379 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998b. Illustrated flora of British Columbia. Volume 1.
Gymnosperms and Dicotyledons (Aceraceae through
Asteraceae). British Columbia Ministry of Environment,
Lands and Parks and British Columbia Ministry of Forests,
Victoria, British Columbia. 436 pages.
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1959. Vascular plants of the Pacific Northwest.
Part 4: Ericaceae through Campanulaceae. University of
Washington Press. Seattle, Washington. 510 pages.
Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson,
W. L. Harper, and B. M. Wikeem. 1991. Interior Dou-
glas-fir zone. Pages 153-166 in Ecosystems of British
Columbia. Edited by D. Meidinger and J. Pojar. British
Columbia Ministry of Forests Special Report Series (6),
Victoria, British Columbia. 330 pages.
Pojar, J. 1999. Polemoniaceae. Pages 36-60 in Illustrated
flora of British Columbia.Volume 4. Dicotyledons (Oro-
banchaceae through Rubiaceae). Edited by G. W. Douglas,
D. Meidinger; and J. Pojar. British Columbia Ministry of
Environment, Lands and Parks and British Columbia Min-
istry of Forests, Victoria, British Columbia. 427 pages.
Primack, R. B. 1998. Essentials of conservation biology.
Sinauer Associates Inc., Sunderland, Massachusetts.
Scoggan, H. J. 1979. The flora of Canada. Part 4. Dicotyle-
doneae (Loasaceae to Compositae). National Museum of
Natural Sciences Publications in Botany 7 (4): 1117-1710.
Wilken, D. A. 1993. Collomia. Pages 825-826 in The Jepson
manual: Higher plants of California. Edited by J. C. Hick-
man. University of California Press, Berkeley, California.
1400 pages.
Received 4 November 2002
Accepted 10 April 2006
Conservation Evaluation of the Small-flowered Tonella, Jonella
tenella, in Canada’
GEORGE W. DouGLAs! AND JENIFER L. PENNY?
' Deceased.
2 Conservation Data Centre, Ministry of Environment, Ecosystems Branch, P.O. BOX 9993 St~ Prov Govt, Victoria British
Columbia V8W 9R7 Canada
Douglas, George W., and Jenifer L. Penny. 2006. Status of the Small-flowered Tonella, Jonella tenella (Scrophulariaceae) in
Canada. Canadian Field-Naturalist 120(2): 179-182.
In Canada, the Small-flowered Tonella, Tonella tenella, is restricted to the west side of Saltspring Island in the Gulf Islands of
southwestern British Columbia. This population represents the northern limits of the species which is disjunct from its main
range in southern Washington (Columbia River gorge), through Oregon to central California. In British Columbia, Jonella
tenella is associated with rock outcrops and dry, steep, sparsely forested talus slopes at elevations of 50 to 300 m. The population
on Saltspring Island is on private property and not directly imperilled at this time. There is, however, a potential for housing
development in the future on this waterfront site, thus the authors consider the species endangered.
Key Words: Small-flowered Tonella, Zonella tenella, endangered, distribution, population size, British Columbia.
The Small-flowered Tonella, Tonella tenella (Benth.)
Heller’, is a member of a genus of only two species
occurring in western North America (Hitchcock et al.
1959). It is the only species found in British Columbia
and Canada (Pojar 2000). Tonella tenella was first re-
corded in Canada by Douglas and Ruyle-Douglas
(1978).
Tonella tenella is a slender, ascending to prostrate,
annual herb from a delicate taproot (Figure 1; Pojar
2000). The smooth, often branched stems are 5-25 cm
long with opposite leaves 1-2 cm long. The leaves are
stalked, simple, ovate to round, and few-toothed or
lobed below and unstalked and deeply 2-3 lobed above.
The blue or white flowers are small, slightly zygomor-
phic and long-stalked with deeply five-lobed calyces.
The corollas are also five-lobed. There are four, epi-
petalous stamens and one pistil with 2 ovules. Anthers
are 0.4 mm long. The fruits are obtuse to globe-shaped
capsules containing 2 to 4, 1-1.5 cm long, wingless
seeds.
Distribution
The southwestern British Columbia occurrence of
Tonella tenella is disjunct from its main range in south-
ern Washington (Columbia River gorge), through Ore-
gon to central California (Wetherwax 1993; Pojar
2000). In Canada, T. tenella is known only from the
west side of Saltspring Island in the Gulf Islands of
southwestern British Columbia (Figure 2; Pojar 2000;
Douglas et al. 2002a, 2002b).
.
FiGureE 1. Illustration of Tonella tenella (Line drawing from
Douglas et al. 2000)
The field work for Tonella tenella was funded by the British Columbia Conservation Data Centre. The results appear in the British Columbia
Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a Committee on the
Status of Endangered Wildlife in Canada status report (Douglas and Penny 2003”) and the subsequent assessment of Endangered (COSEWIC
2003"). This paper includes more recent information that did not appear in the original status report.
"Taxonomy and nomenclature follows Douglas et al. (1998a, 1998b, 1999a, 1999b, 2000, 2001) and Schofield (1992).
179
180
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Locations and sizes of Tonella tenella subpopulations on Saltspring Island, British Columbia.
Collection site Last Observation
1- Upper slope, south 1976
2- Upper slope, north 2002
3- Mid-slope 2002
4-Lower slope 2002
5- ca. 10 m uphill from beach 2002
Habitat
Tonella tenella sites in British Columbia are found
in a climatic and floristic anomaly in coastal British
Columbia, the dry Coastal Douglas-fir (Pseudotsuga
menziesii) Zone (Nuszdorfer et al. 1991) of southeast-
ern Vancouver Island. This area is in a rainshadow belt
created by the Olympic Mountains to the south, result-
ing in a relatively warm and dry Mediterranean climate.
Within this region, Jonella tenella occurs on west-
facing slopes on stable talus. This talus occurs in open
Big-leaf Maple (Acer macrophyllum)-Arbutus (Arbutus
menziesii) forests, where associates include Cleavers
(Galium aparine), Little Western Bitter-cress (Car-
damine oligosperma), Miner’s-lettuce (Claytonia per-
foliata), Common Vetch (Vicia sativa), Barren Brome
(Bromus sterilis), and Large-flowered Blue-eyed Mary
(Collinsia grandiflora) or in open Douglas-fir — Arbu-
tus — Garry Oak forests with Oregon Beaked Moss
(Kindbergia oregana), heron’s-bill moss (Dicranum
species), Harford’s Melic (Melica harfordii), Galium
aparine, Upright Hedge-parsley (Torilis japonica) and
Barren Brome (Bromus sterilis).
Biology
Species of Tonella, along with its sister genus, Blue-
eyed Mary (Collinsia) of the tribe Collinsieae, are self-
compatible annuals (Armbruster et al. 2002). Tonella
tenella is considered a small-flowered type and accord-
ing to Armbruster et al. (2002), large- and small-flow-
ered taxa appear to have differences in timing of self-
pollination. Large-flowered taxa maintain herkogamy
(spatial separation of anthers and stigmas) early in
anthesis by differential elongation of staminal filaments,
while small-flowered taxa do not show this elongation
pattern. As a result, large-flowered taxa experience a
delay in self-pollination whereas in small-flowered
taxa, anther-stigma contact and self-pollination occur
early. Furthermore, the stigmas are receptive to pollen-
tube growth early in Tonella species. Small-flowered
populations are almost certainly autogamous (Arm-
bruster et al. 2002). Low elevation pollinators for the
tribe include the insects Bombus, Anthophora, Empho-
ropsis, Synhalonia and Osmia (Armbruster et al. 2002).
Population Attributes
Four small subpopulations of Tonella tenella were
recently confirmed in 2002 at the Saltspring Island site.
These subpopulations, plus an earlier (1976) collection
Collector Number of plants/area (m7)
Douglas unknown
Lomer 56+ /40.18
Lomer 100-150/15
Douglas 65+/-15/6
Lomer 30/1
record, occur in a narrow band extending up the moun-
tainside for about 425 m. The subpopulations consist-
ed of 6 to 356 plants with areas of 1 to about 40 m?
(Table 1). Since the plant is inconspicuous and extreme-
ly difficult to detect, it is quite likely other subpopu-
lations occur on the slope.
Provincial, National and Global Ranks
Globally, Tonella tenella has a rank of G5 indicating
that in most of its range the plant is common. Since
the species is restricted to British Columbia it has a
national rank of N1. Provincially, 7. tenella has been
ranked as S1 by the Conservation Data Centre and
appears on the British Columbia Ministry of Environ-
ment Red List (Douglas et al. 2002a). This is the most
critical rank that can be applied to species at the provin-
cial level and indicates that the species is “critically
imperiled because of extreme rarity (typically five or
fewer occurrences or very few remaining individuals)
or because of some factor(s) making it especially vul-
nerable to extirpation or extinction”.
Threats and Protection
The most immediate threat to Jonella tenella in
British Columbia is habitat destruction through a hous-
ing development on waterfront private property. Sup-
pression of both natural and human-induced fires in
the last century may also have had an effect on the sur-
vival of 7. tenella. The vegetation in this region would
naturally be maintained by fires; however, in their
absence, high fuel loads build and catastrophic fires
could result. This may result in unsuitable conditions
for T. tenella. Introduced species are also a problem.
The vegetation characterizing the 7. tenella site has
been altered by the introduction of European species.
Although a large number of native forbs occur in these
areas, much of the vegetation is dominated to a large
extent by introduced species, particularly grasses, in-
cluding a number of species of Bromus. The Tonella
tenella site occurs on private land.
Tonella tenella is not formally protected in British
Columbia; however, it could be in the future since
this species is a potential candidate for listing under
the provincial Wildlife Amendment Act (2004). As part
of its commitment to the National Accord (National
Accord for the Protection of Species at Risk), the
province is required to take measures to protect this
species. It is on Schedule 1 of the federal Species At
Risk Act and a recovery strategy will be required.
2006
Evaluation of Status
The British Columbia Conservation Data Centre con-
siders T. tonella to be Endangered in British Columbia
(Douglas et al. 2002a). The Committee on the Status
of Endangered Wildlife in Canada has also assigned
this species to the Endangered category (COSEWIC
2003"). Only one extant population of Tonella tenella
is known in British Columbia; thus, the loss of the one
site would result in the loss of the species from British
Columbia and Canada. Potential housing developments
threaten 7. tenella in Canada since the one site occurs
on private oceanside property.
Acknowledgments
We thank Frank Lomer for his assistance with field
work and Marilyn Lambert for providing water trans-
portation.
Documents Cited (marked * in text)
COSEWIC 2003. COSEWIC assessment and status report
on the small-flowered tonella Tonella tenella in Canada.
Committee on the Status of Endangered Wildlife in Canada.
Ottawa. vii + 14 pages. Available at: www.sararegistry.gc.
ca/status/status_e.cfm.
Douglas, G. W., and J. L. Penny. 2003. COSEWIC status
report on the small-flowered tonella Tonella tenella in
Canada in COSEWIC assessment and status report on the
small-flowered tonella Tonella tenella in Canada. Com-
mittee on the Status of Endangered Wildlife in Canada.
Ottawa. 1-14 pages.
Literature Cited
Armbruster, W. S., C. P. H. Mulder, B. G. Baldwin, S.
Kalisz, B. Wessa, and H. Nute. 2002. Comparative analy-
sis of late floral development and mating-system evolution
in tribe Collinsieae (Scrophulariaceae s.1.). American Jour-
nal of Botany 89: 37-49.
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare
native vascular plants of British Columbia. Second edition.
Province of British Columbia, Victoria, British Columbia.
359 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus-
trated flora of British Columbia. Volume 3. Dicotyledons
(Diapensiaceae through Onagraceae). British Columbia
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Columbia.
423 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus-
trated flora of British Columbia. Volume 4. Dicotyledons
(Orobanchaceae through Rubiaceae). British Columbia
Ministry of Environment, Lands and Parks and British
Columbia Ministry of Forests, Victoria, British Colum-
bia. 427 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illustrated
flora of British Columbia. Volume 5. Dicotyledons (Sali-
caceae to Zygophyllaceae) and Pteridophytes. British
Columbia Ministry of Environment, Lands and Parks and
British Columbia Ministry of Forests, Victoria, British
Columbia. 389 pages.
Douglas, G. W., D. Meidinger, and J. Pojar 2001. Illustrat-
ed flora of British Columbia. Volume 7. Monocotyledons
(Orchidaceae to Zosteraceae). British Columbia Ministry
DOUGLAS AND PENNY: SMALL-FLOWERED TONELLA 18]
a) BRITISH
‘e COLUMBIA
eS |
~™*©_f VICTORIA
FIGURE 2. Distribution of Jonella tenella in British Columbia.
of Sustainable Resource Management and British Columbia
Ministry of Forests, Victoria, British Columbia. 379 pages.
Douglas G. W., D. Meidinger, and J. Pojar. 2002b. Illustrated
flora of British Columbia. Volume 8 — General Summary,
Maps and Keys. British Columbia Ministry of Sustainable
Resource Management and British Columbia Ministry of
Forests, Victoria, British Columbia. 457 pages.
Douglas, G. W, and G. Ruyle-Douglas. 1978. Contributions
to the flora of British Columbia and the Yukon Territory I.
Vascular plants. Canadian Journal of Botany 56: 2296-2302.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a.
Illustrated flora of British Columbia. Volume 1. Gymno-
sperms and Dicotyledons. (Aceraceae through Asteraceae).
British Columbia Ministry of Environment, Lands and
Parks and British Columbia Ministry of Forests, Victoria,
British Columbia. 436 pages.
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998b. Illustrated flora of British Columbia. Volume 2.
Dicotyledons. (Balsaminaceae through Cuscutaceae). Bni-
tish Columbia Ministry of Environment, Lands and Parks
and British Columbia Ministry of Forests, Victoria, British
Columbia. 401 pages.
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1959. Vascular plants of the Pacific North-
west — Part 4: Ericaceae through Campanulaceae. Universi-
ty of Washington Press, Seattle, Washington. 510 pages.
182
Nuszdorfer, F. C., K. Klinka, and D. A. Demarchi. 1991.
Coastal Douglas-fir zone. Pages 95-112 in Ecosystems of
British Columbia. Edited by D. Meidinger and J. Pojar.
British Columbia Ministry of Forests Special Report Series
(6), Victoria, British Columbia. 330 pages.
Pojar, J. 2000. Scrophulariaceae. Pages 114-218 in Illus-
trated flora of British Columbia.Volume 5. Dicotyledons
(Salicaceae through Zygophyllaceae) and Pteridophytes.
Edited by G. W. Douglas, D. Meidinger, and J. Pojar. Bri-
tish Columbia Ministry of Environment, Lands and Parks
and British Columbia Ministry of Forests, Victoria, British
Columbia. 427 pages.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Schofield, W. B. 1992. Some common mosses of British
Columbia. Royal British Columbia Museum Handbook.
Royal British Columbia Museum, Victoria, British Colum-
bia. 394 pages.
Wetherwax, M. 1993. Tonella. Page 1063 in The Jepson
manual: Higher plants of California. Edited by J.C. Hick-
man. University of California Press, Berkeley, California.
1400 pages.
Received 26 November 2002
Accepted 20 March 2006
|
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‘
Conservation Evaluation of Stoloniferous Pussytoes, Antennaria
. . *
flagellaris, in Canada
GEORGE W. DouGLas!, JENIFER L. PENNY2, and KSENIA BARTON?
' Deceased.
2 Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, P.O. Box 9993, Stn Prov Govt,
Victoria, British Columbia V8W 9R7 Canada
3Current address: 204-4272 Albert Street, Burnaby, British Columbia V5C 2E8 Canada
Douglas, George W., Jenifer L. Penny, and Ksenia Barton. 2006. Conservation evaluation of Stoloniferous Pussytoes, Anten-
naria flagellaris, in Canada. Canadian Field-Naturalist 120(2): 183-187.
In Canada, Stoloniferous Pussytoes, Antennaria flagellaris, is restricted to the Similkameen River valley south of Princeton, in
south-western British Columbia. The three populations represent the northern limits of the species which ranges from south-
western British Columbia, south in the western United States to Idaho, Wyoming, Nevada and California. In British Colum-
bia, Antennaria flagellaris is associated with eroded, unstable, calcareous clay seepage slopes on open, southerly aspects.
This habitat is infrequent in the Similkameen River area and the few existing plant populations could easily be extirpated
through slight changes in drainage through drilling for coalbed methane gas, road-building, or housing development.
Key Words: Stoloniferous Pussytoes, Antennaria flagellaris, endangered, distribution, population size, British Columbia.
Stoloniferous Pussytoes, Antennaria flagellaris (A.
Gray) A. Gray’ is a member of a genus of approximate-
ly 35 species mainly of North America but also found
in South America (Cronquist 1994). Fifteen species
occur in British Columbia (Douglas 1998) and approx-
imately 18 in Canada (Kartesz 1999). Antennaria fla-
gellaris was first recorded in Canada by Douglas et al.
(1998a).
Antennaria flagellaris is a stoloniferous, short-lived
perennial from a fibrous root (Figure 1; Douglas 1998).
The stolons are up to 10 cm long while the few central
flowering stems are up to 3.5 cm tall. The numerous
basal leaves and the few stem leaves are linear to linear- wy
oblanceolate, silky woolly-hairy and 1-3 cm long. The FS
flower heads are terminal with involucres of two types.
The female involucres are 7-13 mm tall bracts that are
tinged brown or reddish-brown, thinly woolly-hairy
below. The male involucres are 4-7 mm tall translucent
bracts that are brownish at the tips. The female flowers
are 5-7 mm tall while the male flowers are 3-4.5 mm tall. é
The single-seeded achenes are elliptic, warty and 2-3 mm a
long with a 6-8 mm tall pappus that has white, hairlike a (\
bristles.
VAN A hl
INIA
4
Distribution
Antennaria flagellaris ranges from southwestern
British Columbia, south in the western United States to Ficure 1. Illustration of Antennaria flagellaris (Line drawing
Idaho, Wyoming, Nevada and California (Cronquist by Elizabeth J. Steven in Douglas et al. 1998, 2002b).
“The field work for Antennaria flagellaris was funded by the British Columbia Conservation Data Centre. The results appear in the British
| Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a Com-
4
}
|
}
;
'
|
|
]
Mittee on the Status of Endangered Wildlife in Canada status report (Douglas and Penny 2004") and the subsequent assessment of Endan-
gered (COSEWIC 2004’). This paper includes more recent information that did not appear in the original status report.
‘Taxonomy and nomenclature follow Douglas et al. (1998b, 1999, 2001).
183
184
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Locations and Population Sizes for Antennaria flagellaris near Princeton, British Columbia.
Collection site Last Observation
1. Stevenson Creek 1997
2. Whipsaw Creek, 2 km N of 1997
3. Stevenson Lake, 300 m NE of 2002
4. Tracey Lake 2003
1994). In Canada, A. flagellaris is known only from
along a 3.2-km length of Highway #3 in the Similka-
meen River valley south of Princeton in southwestern
British Columbia (Figure 2; Douglas 1998; Douglas
et al. 1998a, 2002a, b).
Habitat
Antennaria flagellaris occurs in the southern inte-
rior of British Columbia along the Similkameen River
valley in the lower montane zone in the Interior Dou-
glas-fir biogeoclimatic zone (Hope et al. 1991). Cli-
matic conditions in this region are continental, char-
acterized by hot, dry summers, a fairly long growing
season, and cool winters. A rainshadow effect prevails
in this area due to the presence of the Coast-Cascade
Mountains to the west. Within this zone, A. flagellaris
occurs in calcareous, gravelly-clay soils or gravelly
sands. The sites are characterized by a unique hydrol-
ogy. The slopes, which are moderate (20-30%) with
open southerly aspects, are eroded, unstable, and char-
acterized by ephemeral seepage. These moist microsites
are saturated by underground water in the winter and
dry up in the summer. The slow down-slope movement
that occurs as a result of the unusual hydrology is unique
in the region and excludes many taxa which are not
able to tolerate the conditions. As a result, the slope
is disturbed and sparsely vegetated with A. flagellaris
being the dominant component (Figure 3).
Vegetation in the vicinity of the A. flagellaris sites is
characterized by Big Sagebrush (Artemisia tridentata)
shrub/grassland with scattered Ponderosa Pine (Pinus
ponderosa) and Douglas-fir (Pseudotsuga menziesii)
[Nomenclature follows Douglas et al. 1998b, 1998c,
1999 and 2001]. The sites occur at the western edge
of the distribution of open shrub/grassland at that ele-
vation. Associates of A. flagellaris include Cushion
Buckwheat (Eriogonum ovalifolium var. nivale), Par-
snip-flowered Buckwheat (E. heracleoides var. angus-
tifolium), Swale Desert-parsley (Lomatium ambiguum),
Thread-leaved Sandwort (Arenaria capillaris ssp. amer-
icana), Slender Wheatgrass (Elymus trachycaulus),
One-spike Oatgrass (Danthonia unispicata), Common
Rabbit-brush (Ericameria nauseosus var. speciosa),
and Cut-leaved Daisy (Erigeron compositus var. glabra-
tus). No non-native species were found in association
with Antennaria flagellaris.
Biology
Little information is available on the biology of A.
flagellaris in British Columbia. Only basic reproductive
facts have been compiled about the species.
Collector Number of plants/area (m7)
Lomer 50 +3 or 4
Lomer Unknown
Douglas and Penny 1 400 000 + 100 000/ 2100
Lomer 4-500/84
Antennaria flagellaris is a short-lived perennial, dioe-
cious (male and female structures on separate plants)
and wind-pollinated. This species produces seeds sex-
ually by outcrossing (Bayer 1996). The fruits consist of
single-seeded achenes. The numerous hair-like bristles
of the mature achene facilitate its dispersal by wind.
Antennaria flagellaris also reproduces vegetatively
by producing stolons that terminate in plantlets. Ini-
tially, the mother plant provides the plantlet with nutri-
ents via the stolon. The genetically identical plantlets
eventually become independent plants as stolons are
severed. This mode of reproduction results in very re-
stricted dispersal, as stolons only grow up to 10 cm long
(Douglas 1998).
Population Attributes
There are three recently surveyed populations of
Antennaria flagellaris in the Similkameen River valley,
south of Princeton, British Columbia (Table 1). These
populations occur along a 3.2-km length of Highway
#3 (Figure 2). The populations range from a couple of
square metres to over 2100 m/ in size and number from
50 to approximately 1 400 000 + 100 000 plants.
Short- and long-term trends for these populations are
unknown but can be expected to vary markedly due
to differences in yearly plantlet survival, seed germi-
nation and seedling success. Growing conditions on
the ephemerally moist, clay slopes may vary from year
to year according to normal climatic fluctuations affect-
ing site hydrology.
Provincial, National and Global Ranks
Globally, Antennaria flagellaris has a rank of G5?
indicating that in most of its range the plant is appar-
ently secure to demonstrably widespread, abundant, and
secure. Since the species is restricted to British Colum-
bia it has a national rank in Canada of N1. Provincial-
ly, Antennaria flagellaris has been ranked as S1 by the
Conservation Data Centre and appears on the British
Columbia Ministry of Environment Red list (Douglas
et al. 2002a). This is the most critical rank that can be
applied to species at the provincial level and indicates
that the species is “critically imperiled because of ex-
treme rarity (typically five or fewer occurrences or very
few remaining individuals) or because of some factor(s)
making it especially vulnerable to extirpation or extinc-
tion”.
Threats and Protection
The major threat to populations of Antennaria fla-
gellaris are through drilling for coalbed methane gas,
2006
road construction, housing developments and off-road
recreational vehicles. In 2002 three test holes, target-
ing coalbed methane gas deposits, were drilled in the
Similkameen valley. If brought into production the
drilling, access roads and pumping stations could cause
major degradation in the area.
The southern interior of British Columbia has recent-
ly seen an increase in housing projects and the Prince-
ton area is no exception. Any developments which even
slightly change groundwater levels could be devastat-
ing to this taxon. Furthermore, suitable habitats for
A. flagellaris are extremely restricted in Canada, only
occurring in the Princeton area, thus opportunities for
colonisation are extremely limited.
Although weeds are not a major threat to A. flagel-
laris populations, weed control activities do constitute
a minor potential threat to populations. Under the Weed
Control Act, an occupier must control noxious weeds
growing or located on land and premises; thus margin-
ally specific chemical weed control compounds, that
kill broad-leaved plant species, would likely be fatal
to A. flagellaris.
Recreational off-road vehicle use could also pres-
ent a threat due to the nature of the steep, clay slopes
where A. flagellaris occurs. When moist, this habitat
would be highly susceptible to disturbance and at the
same time, be at its most desirable to off-road users.
Overuse of the habitat in this way could disturb the
sites enough to radically alter the hydrological regime,
and promote the establishment of non-native species.
Other potential threats result from activities that are
permitted in the Agriculture Land Reserve. The private
property where A. flagellaris occurs is on the Agricul-
tural Land Reserve, where primary land use is agri-
culture. For instance, some types of fill/soil removal
are allowable without application to the Agricultural
Land Commission (Provincial Agricultural Land Com-
mission 2003"). On these lands, development pressures
may not appear to be an issue at this time. The Agri-
cultural Land Reserve status may prevent subdivision
development, but does allow other activities that could
also potentially threaten the populations. In recent years
many tracts of Agricultural Land Reserve land in south-
ern British Columbia have been converted to housing
developments, shopping malls, and golf courses, either
by decisions of the Agricultural Land Commission
or, very rarely, by an “order in council” by the sitting
provincial legislature.
An additional threat to the population of A. flagel-
laris is its vulnerability to extirpation due to the ex-
tremely small occupied area and population size. Such
small populations are at risk of inbreeding depression
(Primack 1998). Due to lack of genetic variation, these
small populations are vulnerable to demographic and
environmental variation. Furthermore, suitable habi-
tats for C. flagellaris in the southern interior of
British Columbia are extremely limited and thus
opportunities for colonisation are also limited.
DOUGLAS, PENNY, AND BARTON: STOLONIFEROUS PUSSYTOES
185
BRITISH
COLUMBIA
FiGuRE 2. Distribution of Antennaria flagellaris in British
Columbia.
Antennaria flagellaris is not formally protected in
British Columbia, however, it could be in the future
since this species is a potential candidate for listing
under the provincial Wildlife Amendment Act (2004).
As part of its commitment to the National Accord
(National Accord for the Protection of Species at Risk),
the province is required to take measures to protect this
species. It is on Schedule | of the federal Species At
Risk Act and a recovery strategy is expected to be com-
pleted by the summer of 2006.
186 THE CANADIAN FIELD-NATURALIST
Vol. 120
FIGURE 3. Open habitat of Antennaria flagellaris in the Princeton area. Antennaria flagellaris is abundant in the open areas
between clumps of Eriogonum heracleoides var. angustifolium and Ericameria nauseosus Vat. speciosa.
Evaluation
The British Columbia Conservation Data Centre con-
siders A. flagellaris to be Endangered in British Colum-
bia (Douglas et al. 2002a). The Committee on the Sta-
tus of Endangered Wildlife in Canada has also assigned
this species to the Endangered category (COSEWIC
2004*). The three populations are small and vulnerable
to a number of threats. With limited knowledge of the
plants, biological and ecological requirements, this
species is vulnerable to extirpation in British Columbia.
Without research on growth requirements and further
demographic information, the stability of the present
populations will remain unknown. The limited number
of plants also reduces the potential for genetic variation
which may be necessary to respond to environmental
changes in the future.
Acknowledgments
We thank Frank Lomer for providing information on
his original discovery of A. flagellaris and for his assis-
tance in relocating the sites in 2002 and 2003.
Documents Cited (marked * in text)
COSEWIC. 2004. COSEWIC assessment and status report
on Stoloniferous Pussytoes, Antennaria flagellaris in Cana-
da. Committee on the Status of Endangered Wildlife in
Canada, Ottawa, Ontario. Available at: http://www.sarareg
istry.gc.ca/virtual_sara/files/cosewic/as%5Fstoloniferous%
5Fpussytoes%5Fe%2Epdf.
Douglas, G. W., and J. L. Penny. 2004. COSEWIC Status
report on Stoloniferous Pussytoes, Antennaria flagellaris,
in Canada, in COSEWIC assessment and status report on
Stoloniferous Pussytoes, Antennaria flagellaris in Cana-
da. Committee on the Status of Endangered Wildlife in
Canada, Ottawa, Ontario. 13 pages. Available at: http://www.
sararegistry.gc.ca/virtual_sara/files/cosewic/sr%5Fstolon
iferous%5Fpussytoes%5Fe%2Epdf.
Provincial Agricultural Land Commission. 2003. Agricultur-
al Land Reserve — Frequently Asked Questions. Provincial
Agricultural Land Commission. 133-4940 Canada Way,
Burnaby, British Columbia V5G 4K6 Canada. Available
at: http://www.alc.gov.bc.ca/faq/faq_alr.htm.
Literature Cited
Bayer, R. J. 1996. Phylogenetic inferences in Antennaria
(Asteraceae: Inuleae: Gnaphaliinae) based on sequences
from the nuclear ribosomal DNA internal transcribed spac-
ers (ITS). American Journal of Botany 83: 516-527.
Cronquist, A. 1994. Intermountain flora — Vascular plants of
the Intermountain West, U.S.A. Volume five: Asterales. New
York Botanical Garden, New York, New York. 496 pages.
Douglas, G. W. 1998. Asteraceae. Pages 96-392 in Illustrated
flora of British Columbia. Volume 1. Gymnosperms and
Dicotyledons (Aceraceae through Asteraceae). Edited by
G. W. Douglas, G. B. Straley, and D. Meidinger. British
Columbia Ministry of Environment, Lands and Parks and
British Columbia Ministry of Forests, Victoria, British
Columbia. 436 pages.
Douglas, G. W., F. Lomer, and H. L. Roemer. 1998a. New
or rediscovered native vascular plant species in British
Columbia. Canadian Field-Naturalist 112: 276-279.
«
2006
Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare
native vascular plants of British Columbia, Second edition.
Province of British Columbia, Victoria, British Columbia.
359 pages.
Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illustrated
flora of British Columbia. Volume 4. Dicotyledons (Oro-
banchaceae through Rubiaceae). British Columbia Ministry
of Environment, Lands and Parks and British Columbia
Ministry of Forests, Victoria, British Columbia. 427 pages.
Douglas, G. W., D. Meidinger, and J. Pojar 2001. Illustrat-
ed flora of British Columbia. Volume 7. Monocotyledons
(Orchidaceae to Zosteraceae). British Columbia Ministry
of Sustainable Resource Management and British Columbia
Ministry of Forests, Victoria, British Columbia. 379 pages.
Douglas G. W., D. Meidinger, and J. Pojar. 2002b. Illus-
trated flora of British Columbia. Volume 8 — General Sum-
mary, Maps and Keys. British Columbia Ministry of Sus-
tainable Resource Management and British Columbia
Ministry of Forests, Victoria, British Columbia. 457 pages.
Douglas, G. W., G. B. Straley, and D. Meidinger. 1998b.
Illustrated flora of British Columbia. Volume 1. Gymno-
sperms and Dicotyledons. (Aceraceae through Asteraceae).
British Columbia Ministry of Environment, Lands and
Parks and British Columbia Ministry of Forests, Victoria,
British Columbia. 436 pages.
DOUGLAS, PENNY, AND BARTON: STOLONIFEROUS PUSSYTOES
187
Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar.
1998c, Illustrated flora of British Columbia. Volume 2
Dicotyledons. (Balsaminaceae through Cuscutaceae).
British Columbia Ministry of Environment, Lands and
Parks and British Columbia Ministry of Forests, Victoria,
British Columbia. 401 pages.
Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson,
W. L. Harper, and B. M. Wikeem. 1991. Interior Douglas-
fir zone. Pages 153-166 in Ecosystems of British Columbia.
Edited by D. Meidinger and J. Pojar. British Columbia Min-
istry of Forests Special Report Series Number 6, Victoria,
British Columbia. 330 pages.
Kartesz, J.T. 1999. A synonymized checklist and atlas with
biological attributes for the vascular flora of the United
States, Canada, and Greenland. First Edition. /n Synthesis
of the North American flora, Version 1.0. Edited by J.T.
Kartesz and C. A. Meacham. North Carolina Botanical Gar-
den, Chapel Hill, North Carolina.
Primack, R. B. 1998. Essentials of conservation biology. Sin-
aur Associates Inc., Sunderland, Massachusetts.
Received 30 October 2002
Accepted 20 March 2006
Late-born Elk, Cervus elaphus, Calf Observed Near Bancroft, Ontario
Rick C. Rosatte!? AND JOE NEUHOLD?
‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, DNA Complex,
2140 East Bank Drive, Peterborough, Ontario K9J 7B8 Canada
2 825 Wentworth St., Peterborough, Ontario K9J 8R7 Canada »
3Corresponding author: e-mail: rick.rosatte@ontario.ca
Rosatte, Rick, and Joe Neuhold. 2006. Late-born Elk, Cervus elaphus, calf observed near Bancroft, Ontario. Canadian Field-
Naturalist 120(2): 188-191.
A recently born male Elk (Cervus elaphus) calf was observed and photographed approximately 30 km east of Bancroft,
Ontario, on 5 October 2004. Based on the gestation period for Elk, the estimated conception date was between late Decem-
ber 2003 and early January 2004. This is unusual as the majority of calves are born between late May and early June with
conception having occurred in September/October. This reproductive strategy allows for an optimum growth period for
calves to occur before the onset of winter which increases the probability of survival. An eleven-year-old cow Elk was iden-
tified as the mother of the late born calf. The age of the cow, and her inability to meet the annual energy requirements neces-
sary for normal reproduction, may have contributed to the late conception and parturition dates.
Key Words: Elk, Cervus elaphus, Elk calf, restoration, late-born, Ontario.
In 1998, the Ontario Ministry of Natural Resources
(OMNR) and partner organizations implemented a
plan to restore Elk (Cervus elaphus) to Ontario, Cana-
da (Bellhouse and Broadfoot 1996). Between 1998
and 2001, 443 elk acquired from Elk Island National
Park (EINP), Alberta, were released in four different
areas of Ontario; Lake of the Woods, Lake Huron/North
Shore (near Blind River), French River/Burwash, and
Bancroft/North Hastings (Rosatte et al. 2002). In total,
120 Elk were released during 2000/2001 in the Ban-
croft/North Hastings area at a location near Gin Lake
(UTMC = 304000E 4990000N). Each summer, Elk
calves have been observed and each winter, Elk calf
surveys were conducted using OMNR helicopters to
determine Elk calf survival and estimate the size of
the herd.
On 5 October 2004, a male Elk calf with spots, esti-
mated to be about three weeks of age, was observed
and video taped near Hartsmere, Ontario (UTMC=
300000E 4995000N), approximately 25 km east of
Bancroft, Ontario (Figure 1). The calf weighed about
25-30 kg based on body size; average birth weights
for Elk calves are 15-22 kg (Hudson et al. 2002). On
1 November 2004, the calf was again observed nurs-
ing from a 11-year-old cow Elk which had been ear-
tagged (All-Flex ear-tag # 347), weighed (238 kg),
and radio-collared (Lotek GPS Collar- 148.115 Mhz)
at EINP, Alberta, and released in the Bancroft/North
Hastings, Ontario, area during April 2001. The calf was
observed nursing from the cow during December 2004
and January 2005. The calf, still with spots, was pho-
tographed again on | February 2005, near Hartsmere,
Ontario (Figure 2). It was with a group of 38 Elk com-
prised of 13 bulls {4 mature (>Syr), 4 yearlings, 5 im-
mature (2-4 yrs), 8 calves, and 17 cows (which includ-
ed the calf’s mother — Figure 3)}. Based on observation
during January/February 2005 the late-born calf with
spots was estimated to be 25%-35% smaller than the
other calves (Figure 2). Those calves were of a size —
comparable to Elk calves handled at EINP, Alberta,
where calf weights averaged 120 kg (264 Ib) (n=16)
during January 2001. These weights are comparable to
8-9 mo old Elk calves in other studies that were born
during early spring and summer (Cook 2002; Peek
2003). Unfortunately, the spotted calf was killed on
25 February 2005. Based on tracks in the fresh snow,
injuries to the calf, and blood on the antlers of a mature
bull Elk, the calf was likely gored to death.
Gestation periods for Elk range between 247-262
days with a mean of about 255 days (Bubenik 1982;
Hudson et al. 2002). As this calf was first observed on
5 October 2004, conception must have occurred dur- —
ing late December 2003 or early January 2004. Wishart
(1981) reported a cow Elk in Alberta that was shot in
September 1979 and had a fully developed fetus. In
this case, the conception date was estimated to be Jan-
uary 1979 and the cow was estimated to be 10 years of
age. In Elk, reproductive success generally declines
after 7 years of age (Raedeke et al. 2002).
One adaptive strategy of Elk is the production of
highly developed, rapid growing, large young, so that
calves quickly reach a size that maximizes survival
during winter (Geist 2002). The high energy require-
ments for rapid growth of Elk calves is met through
the energy acquired from maternal milk. Elk calves |
acquire most of their nutritional requirements during |
their first month of life from milk; however, after this ;
neonatal period, requirements are greater and the inges- -
tion of solid forage is required. That is why parturition
throughout most of Elk range is early spring and sum-
mer (late May/early June) when forage is high in pro- -
tein and nutritious (Hudson et al. 2002). The timing |
188
2006
ROSATTE AND NEUHOLD: LATE-BORN ELK CALF
189
FiGurE 1. Late born elk calf with spots photographed by J. Neuhold near Hartsmere, Ontario, on October 5, 2004.
of calving for Elk is such that inclement winter/spring
weather is avoided but is early enough (e.g. June) so
that calves can attain sufficient weight and size to sur-
vive the winter (Hudson et al. 2002). Juvenile Elk have
evolved to reduce or cease growth during fall and win-
ter, when the nutritive value of forage declines. Cook
et al. (1996) reported that calf growth in mid-Novem-
ber was 50% of that found in September and hypoth-
esized that the nutritional strategy of Elk calves during
the late fall/early winter was primarily to maintain con-
dition rather than maximizing growth. Inevitably, ener-
gy acquisition for rapid growth becomes a problem
for late-born calves, as not only is forage less nutri-
tious during the fall and winter, but milk production
in lactating cow Elk also declines during that period
due to poor quality forage (Cook et al. 1996; Cook
2002). In addition, survival was found to be lower for
late-born Elk calves in Yellowstone National Park
than for earlier-born calves (Singer et al. 1997), and
calf mortality during winter was primarily due to mal-
nutrition. The calf in this study was observed nursing
in mid-January 2005 and supplemental feeding by peo-
ple in the area may have provided sufficient energy
for the cow to continue milk production during the
winter period.
Nutrition and lactation can affect the timing of es-
trous in Elk and subsequently the parturition date (Cook
2002). Generally, cows that are suckling a calf during
the rut come into estrous later than cows that do not
have a calf (Hudson et al. 2002). During the early part
of rut, estrous approximates 21 days but is variable
(19-25 days) (Bubenik 1982) and estrous length in-
creases and becomes irregular during the later stages
of the breeding period (Hudson et al. 2002). Three to
four estrous periods during one breeding season are
possible with a refractory period of 62 days between
sequential estrous cycles (Bubenik 1982) and Elk can
be in heat as late as February. Older aged ungulates ~
often cannot support production or reproduce (Cameron
1994; Cook 2002). Generally, reproductive success for
elk 11 years of age is low (Raedeke et al. 2002). The
age of the cow in this study, and her inability to meet
the annual energy requirements necessary for normal
reproduction, may have contributed to the late concep-
tion and parturition dates.
Acknowledgments
The authors wish to thank the numerous partners of
the Ontario Elk Restoration and Research Program:
the Provincial Elk Technical Team, the Local Imple-
mentation Committees, Cambrian College, the Rocky
Mountain Elk Foundation, Elk Island National Park,
the Ontario Federation of Anglers and Hunters, Trent
University, Lakehead University, Laurentian University,
190 THE CANADIAN FIELD-NATURALIST Vol. 120
ai
1
1
PAl
uf
FIGURE 2. Late born elk calf with spots was about 25% smaller than other calves when photographed by R. Rosatte near
Hartsmere, Ontario, 1 February 2005.
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Sil 20 EL ag pes TMT ES ea ay
ee at ai ee ee a
FIGURE 3. Elk calf with spots with 11-year old mother (on right) (photo by R. Rosatte near Hartsmere on 1 February 2005)
————E—————__ sr t——
a a a eS
a
2006
University of Guelph, Sault College, Safari Club Inter-
national (Ontario Chapter), Northern Ontario Tourist
Outfitters, French River Resorts Association, Parks
Canada, Northern Ontario Heritage Foundation, On-
tario Fur Managers Federation, and the Ontario Minis-
try of Natural Resources. C. Davies, manager, OMNR,
Wildlife Research & Development Section, reviewed
the manuscript and provided useful comments.
Literature Cited
Bellhouse, T., and J. Broadfoot. 1996. Plan for the restora-
tion of elk in Ontario. Ontario Ministry of Natural Re-
sources internal report, Ontario, Canada, 55 pages.
Bubenik, A. 1982. Physiology. Pages 125-179 in Elk of
North America: ecology and management. Edited by J.
W. Thomas and D. Toweill. Stackpole Books, Harrisburg,
Pennsylvania.
Cameron, R. 1994. Reproductive pauses by female caribou.
Journal of Mammalogy 75: 10-13.
Cook, J. 2002. Nutrition and food. Pages 259-349 in North
American elk: ecology and management. Edited by D.
Toweill and J. W. Thomas. Smithsonian Institution Press,
Washington, D.C.
Cook, J., L. Quinlan, L. Irwin, L. Bryant, R. Riggs, and
J. Thomas. 1996. Nutrition-growth relations of elk calves
during late summer and fall. Journal of Wildlife Manage-
ment 60: 528-541.
ROSATTE AND NEUHOLD: LATE-BORN ELK CALF 19]
Geist, V. 2002. Adaptive behavioral strategies. Pages 389-433
in North American elk: ecology and management. Edited
by D. Toweill and J. W. Thomas. Smithsonian Institution
Press, Washington, D.C.
Hudson, R., J. Haigh, and A. Bubenik. 2002. Physical and
physiological adaptations. Pages 199-257 in North Ameri-
can elk: ecology and management. Edited by D. Toweill
and J. W. Thomas. Smithsonian Institution Press, Washing-
ton, D.C,
Peek, J. 2003. Wapiti. Pages 877-888 in Wild Mammals of
North America: Biology, Management and Conservation.
Edited by G. Feldhamer, B. Thompson and J. Chapman.
The Johns Hopkins University Press, Baltimore, Maryland.
Raedeke, K., J. Millspaugh, and P. Clark. 2002. Population
characteristics. Pages 449-491 in North American elk:
ecology and management. Edited by D. Toweill and J. W.
Thomas. Smithsonian Institution Press, Washington, D.C.
Rosatte, R., J. Hamr, B. Ranta, J. Young, and N. Cool.
2002. Elk restoration in Ontario, Canada: Infectious dis-
ease management strategy, 1998-2001. Annals of the New
York Academy of Sciences 969: 358-363.
Singer, F., A. Harting, K. Symonds, and M. Coughenour.
1997. Density dependence, compensation, and environ-
mental effects on elk calf mortality in Yellowstone National
Park. Journal of Wildlife Management 61: 12-25.
Wishart, W. 1981. January conception in an elk in Alberta.
Journal of Wildlife Management 45: 544.
Received 17 February 2005
Accepted 7 October 2005
Annual Variation in Habitat Use by White-footed Mice, Peromyscus
leucopus: The Effects of Forest Patch Size, Edge and Surrounding
Vegetation Type
CHRISTINE S. ANDERSON!, DOUGLAS B. MEIKLE’, ALAN B. Capy”, and ROBERT L. SCHAEFER?
' Present address: Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, Columbus, Ohio
43210 USA. e-mail: anderson.1309@osu.edu; formerly C. Schandorsky; corresponding author.
"Department of Zoology, Miami University, Oxford, Ohio 45056 USA
3Department of Mathematics and Statistics, Miami University, Oxford, Ohio 45056 USA
Anderson, Christine S., Douglas B. Meikle, Alan B. Cady, and Robert L. Schaefer. 2006. Annual variation in habitat use by White-
footed Mice, Peromyscus leucopus: The effects of forest patch size, edge and surrounding vegetation type. Canadian
Field-Naturalist 120(2): 192-198.
White-footed Mice (Peromyscus leucopus) were trapped for two years in the exterior matrix, edge, and interior forest habitat
sections of six forests patches in a fragmented agricultural landscape. We used data on the capture locations of P. leucopus
individuals from the two years, which differed in rainfall (1.e., summer of 2000 with 50% more rain than summer of 1999), to
assess how patch size, edge habitat, and surrounding habitat type influence habitat use and movements in populations of this
forest habitat generalist. We found that the proportion of individuals subsequently captured in the forest edge from the exterior
was 16 times greater in the wet year than in the dry year and approximately twice as many P. leucopus were not subsequently
recaptured from the exterior matrix in the dry year compared to the wet year. For each year, captures between habitats did not
differ in relation to patch size, edge forest habitat, or exterior matrix type. These results illustrate the generalist habitat pref-
erences of P. leucopus, but emphasize annual variation in their behavior and distribution.
Key Words: White-footed Mice, Peromyscus leucopus, forest patches, habitat preference, edge habitat, exterior matrix, Ohio,
Indiana.
Human activities have led to a reduction and frag-
mentation of secondary-growth deciduous forests in
North America, as well as changes in the quality of the
remaining habitat (Yahner 1988; Fahrig 1997). The
effects of fragmentation include those related to loss of
connectivity among remnant patches, and increases in
edge habitat, which both can cause local changes in
abundance, distribution, and species’ persistence (An-
dren 1994, 1997; Murcia 1995). In addition, the frag-
mentation of forested habitat results in the creation of
forest patches that vary in size. Studies have shown that
there are taxonomic differences in the relationship be-
tween animal population densities and patch area.
Insects and birds generally show a positive relationship,
likely due to an avoidance of edge habitat, which is
greater in proportion in smaller patches (Connor et al.
2000). The majority of mammal species show fairly
constant population densities with patch area, this
follows the theory of island biogeography (Bowers
and Matter 1997).
In contrast, investigators have found a consistent
negative relationship between density of the White-foot-
ed Mouse (Peromyscus leucopus Rafinesque), a forest
habitat generalist, and forest patch area (Nupp and
Swihart 1996, 1998; Krohne and Hoch 1999; Mossman
and Waser 2001; Schmid-Holmes and Drickamer 2001;
Anderson et al. 2003; Anderson and Meikle 2006).
Although resource availability may influence popula-
tion dynamics of small mammals (e.g., Nupp and Swi-
hart 1996), there are other factors that may be respon-
sible for the negative density-area relationship. One
possibility is that dispersal is inhibited from smaller
forest patches due to lower habitat quality in the matrix
than under continuous forest cover (Brown and Lit-
vaitis 1995; Nupp and Swihart 1998; Krohne and Hoch
1999). We hypothesize in the current study that if inhib-
ited dispersal results in the negative density-area rela-
tionship, then fewer P. leucopus should tend to move
from the forest edge to the agricultural fields surround-
ing smaller compared to those surrounding larger for-
est patches.
In addition to patch area, another consequence of
forest fragmentation is an increase in the proportion
of edge habitat in the landscape since smaller forest
patches have a greater edge: interior ratio than larger
forest patches. Forest edges usually exhibit differences
in microclimate, species composition, and vegetation
structure compared to forest interiors (Harris 1988;
Saunders et al. 1991). These difference may influence
abundance and distribution of animal species. Studies
have found that some forest animals may prefer for-
est edges while others may avoid those areas (Harris
1988; Saunders et al. 1991; Murcia 1995). However,
there is debate as to whether P. /eucopus prefers for-
est edge to forest interior habitat. Some investigators
have reported greater densities of Peromyscus spp.
near edges (Cummings and Vessey 1994; Boyne and
Hobson 1998; Manson et al. 1999), possibly because
192
=
2006
vegetation in edge habitat has been shown to be struc-
turally more complex compared to interior habitat
(Cummings and Vessey 1994; Anderson et al. 2003).
In contrast, others have found fewer mice and lower
maternal survival and litter success in edge habitats,
presumably because parasitism, predation and inter-
specific competition can be higher along edges (Mat-
ter et al. 1996; Morris and Davidson 2000; Wolf and
Batzli 2001, 2004). Since there is no evidence to date of
higher rates of predation on mice in forest edges, then
we predict that mice may prefer the high quality vege-
tation in forest edges such that there would be higher
recapture rates in forest edges than in forest interiors.
Another consequence of forest fragmentation is that
forest patches are surrounded by a matrix of agriculture
or residential development that may inhibit the move-
ment of organisms (Fitzgibbon 1997; Collinge 1996).
Even though the agricultural matrix surrounding forest
patches may be considered “hostile” habitat for some
species (Collinge 1996), generalists such as P. leuco-
pus (Gottschang 1981) may disperse relatively easily
(Lackey 1978) through row crop fields (Mossman and
Waser 2001). However, it is not well understood whe-
ther matrix types (e.g., corn or pasture) differentially
affect capture rates of White-footed Mice in the exterior
matrix. M’Closkey and Lajoie’s (1975) work suggests
that P. leucopus may prefer corn over pasture because
mice were absent from grasslands. Even though grass-
es may provide cover for mice from aerial predators,
Getz (1961) also noted that individual Peromyscus
avoided grassy sites under forest canopy (also see
M’Closkey and Lajoie 1975) likely because it rustles as
the mouse moves (Barnum et al. 1992). If surrounding
habitat type influences movements of mice through-
out the landscape, we predict that they may stay in or
move into the surrounding matrix habitat at greater
rates when the matrix is comprised of corn rather than
pasture.
The purpose of this study was to use data on the
first two captures of P. leucopus individuals in differ-
ent habitats (i.e., exterior matrix, forest edge, and
forest interior habitat) in small and large forest patch-
es for two summers to test the hypotheses related to
patch size, edge habitat, and surrounding habitat type
as described above. In addition, we compared captures
among habitats between two summers differing in
rainfall amounts (i.e., one dry year and one wet year)
because temporal variation in precipitation regulates
rodent population dynamics (Lewellen and Vessey
1998). Utilizing data on habitat use instead of overall
abundance, we assumed that if mice tended to prefer
one habitat over another, then, on average, they would
move into (or stay in) the preferred habitats in rela-
tively high proportions. Since much of the work on the
effects of forest fragmentation has been focused on
birds, this study contributes to our understanding of
how human-dominated landscapes influence habitat
use and movements of populations of small mammals.
ANDERSON, MEIKLE, CADY, and SCHAEFER: WHITE-FOOTED MICE
193
Methods
Study Area
Peromyscus leucopus were trapped in both 1999
and 2000 in the same three “small” (~S-9 ha) and three
“large” (~1 10-150 ha) forest patches in a row-crop agri-
cultural landscape covering approximately 144 km* in
SW Ohio and SE Indiana, USA (study area, 39°S1'N,
84°74'W; Anderson and Meikle 2006; Table 1). We
analyzed data from four transects of six Sherman™
live traps (7.5 x 7.5 x 25.5 cm) that were placed per-
pendicular to the center of one edge of each patch,
with each transect beginning 20 m exterior to and ex-
tending 30 m into the patch (Figure |). The transects
and the traps in each transect were spaced 10 m apart,
resulting in a 30 m by 50 m grid with a total of 24 traps
at each forest patch. The eight traps in the agricultural
field outside of each forest patch were considered “ex-
terior” and the next eight traps were considered “edge”.
The eight traps inside the forest patch were consid-
ered “interior” traps. Based on our previous findings
(Anderson et al. 2003) and corroborated by Burke and
Nol (1998), the width of the edge habitat was defined
based on typical edge floristic features. The surround-
ing habitat type in the exterior matrix was either pas-
ture or corn fields (Table 1).
Data collection
Traps were set on 26 nights in each forest patch be-
tween June and September in 1999 and 2000 for a total
of 7488 trap-nights. When a mouse was captured ini-
tially it was lightly anesthetized with halothane and a
12 mm AVID® microchip (PIT tag) was injected under
the scruff of its neck. To distinguish between P. leu-
copus (primarily a woodland species) and P. manicu-
latus (primarily a grassland species), which are mor-
phologically similar and overlap in the nonforested
habitat within the study area, saliva samples were ana-
lyzed for the migration of the amylase enzyme with
polyacrylamide gel electrophoresis (Aquadro and Pat-
ton 1980; Bruseo et al. 1999). We followed the proto-
col of Bruseo et al. (1999) with minor modifications.
Each patch was trapped for two consecutive nights
every 5-7 d and we recorded the date, identification
number and location of every individual at every trap-
ping event. Based on trap location we determined which
habitat (i.e., exterior, edge, or interior) each individ-
ual was located in for its first capture and then where
it was subsequently captured for its second capture
event. It is possible that the length of time between any
two capture events for a particular individual could be
24 hours apart or more. A fourth category for the sub-
sequent capture was “not subsequently recaptured”,
which was used if the individual was not recaptured
again after tagging. Only the first two capture events
for each individual were analyzed so that each individ-
ual was included only once in the analysis to avoid
pseudoreplication. In order to verify whether a mouse
moving into a particular habitat reflects fidelity to this
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Description of study sites and their location, size, and surrounding habitat type (i.e., exterior matrix habitat) in 1999
and 2000.
Patch names Location
Small patches
ERC (Ecology Research Station) Oxford, Ohio
Indiana Bath, Indiana
Jericho Oxford, Ohio
Large patches
Bachelor Woods and Game Preserve Oxford, Ohio
Hueston Woods State Park
Reily Township Reily, Ohio
Preble/Butler Counties, Ohio
Surrounding habitat type
Size (ha) 1999 2000
5 pasture pasture
8 com com
9 com com
150 pasture pasture
200 com pasture
110 corm soybeans*
*This patch was not included for tests involving surrounding habitat type since it was the only patch with soybeans.
habitat shift, we also analyzed the data for the first year
using all captures of all mice instead of the first two
capture events only (i.e., 188 mice captured a total of
901 times) and found similar results (not presented),
suggesting a permanence of habitat shifts. Although
over winter survival was very low (i.e., four known
individuals) between the two years of this study (An-
derson 2004), we did not include those four individuals
in the data set for the second year. Rainfall amounts
from the Ohio Agricultural Research & Development
Center (OARDC) weather station on Miami Universi-
ty’s Oxford campus (http://jrscience.wcp.muohio.edu/
van tagepro/latestweather.html) were totaled for each
summer month (June — August) in 1999 and 2000. The
OARDC is located within the center of the study area.
Statistical analyses
Data on the proportion of mice trapped for their sec-
ond capture in the exterior, edge, or interior habitats, or
not subsequently recaptured, based on the three initial
capture habitats (i.e., exterior, edge, or interior) were
compiled (see Table 2). The data set was analyzed with
PROC CATMOD using chi-square tests to determine
whether captures among habitats differed in relation to
patch size, surrounding matrix type, and year (SAS
1999-2000). Subsets of the data were analyzed with
PROC FREQ using either chi-square tests or Fisher’s
exact tests when sample size was small (n < 50 or cells
Agricultural field
Forest Interior
Ficure |. The trapping grid design for each of the forest patch-
es in this study. The letter “O” denotes each outside
(1.e., exterior) trap, the letter “E” represents the eight
edge traps, and the “I” denotes the interior trap loca-
tions.
with values <5; SAS 1999-2000). Data from each year
were analyzed separately (see Results; Table 2). In post
hoc analyses, Fisher’s exact tests and chi-square tests
were used where appropriate (Moore and McCabe
1993) with a Bonferroni adjustment to the level of sig-
TABLE 2. The habitat where each Peromyscus leucopus was trapped for its second capture based on where it was first cap-
tured in 1999 and 2000. Values are pooled across patch sizes and number of individuals and row percentages are given in
parentheses.
Habitat where first captured
Habitat where subsequently (i.e., second) trapped
1999 Exterior Edge Interior Not subsequently recaptured
Exterior 9 (28%) tm GZ) 2 (6%) 20 (63%)
Edge 6 (14%) 19 (43%) 10 (23%) 9 (21%)
Interior 2 (4%) 8 (15%) 17 61%) 27 (50%)
2000 Exterior Edge Interior Not subsequently recaptured
Exterior 1 (8%) 6 (50%) 1 (8%) 4 (33%)
Edge ily 1G) 18 (46%) 10 (26%) 10 (26%)
Interior 0 (0%) 7 (13%) 23 (41%) 26 (46%)
|
|
:
|
}
'
|
|
|
2006
nificance at & = 0.05 (Sokal and Rohlf 1981). An un-
paired t-test was used to test for a difference in mean
monthly rainfall in the summer (June — August) of
1999 compared to 2000 (Sokal and Rohlf 1981).
Results
Patch size did not influence captures of P. leucopus
among habitats in either 1999 or 2000 (1999: x7, = 0.9,
P = 0.8; 2000: y°, = 4.7, P = 0.2). Hence, there was
no difference in captures of mice among different habi-
tats (i.e., exterior matrix, edge, or interior habitats, or
not subsequently recaptured) for those individuals that
had been initially trapped in edge habitat in small com-
pared to large patches in either year (1999: Fisher’s
exact test, n = 44, P = 0.5; 2000: Fisher’s exact test,
n= 39, P=0.7). Data were pooled across patch sizes
within each year for all other analyses (Table 2). The
proportion of P. leucopus recaptured in edge habitat
(i.e., both the first and second captures occurred in edge
habitat) compared to the proportion recaptured in inte-
rior habitat did not differ in either year (1999: oa = 1.4)
P=0.2; 2000: x7, = 0.2, P = 0.6), suggesting that mice
did not show a preference for edge or interior forest
habitat. Surrounding habitat type (i.e., corn or pasture)
did not influence the proportion of mice subsequently
captured across different habitats that had been first
captured in the edge in either year (1999: Fisher’s exact
test, nm = 44, P = 0.5; 2000: Fisher’s exact test,
n = 33, P = 0.2). Likewise, surrounding habitat type
did not affect subsequent captures across habitats for
mice that had been first captured in the exterior matrix
habitat in either year (1999: Fisher’s exact test, n = 32,
P=0.3; 2000: Fisher’s exact test, n = 10, P=0.3).
When captures between habitats in 1999 and 2000
were compared, P. /Jeucopus first captured in the exte-
rior were subsequently trapped in different proportions
among habitats in 1999 compared to 2000 (Fisher’s
exact test, = 44, P = 0.002); specifically, the propor-
tion of individuals subsequently captured in the forest
edge from the exterior was 16 times greater in 2000
(6/12 = 50%) than in 1999 (1/32 = 3%; Fisher’s exact
test,n = 44, P=0.0008; Table 2, Figure 2a). Likewise,
a greater proportion of mice stayed in the exterior
matrix compared to moving into the forest (edge and
interior combined) in 1999 (9/12) than in 2000 (1/7;
Fisher’s exact test, n = 20, P = 0.02). Approximately
twice as many P. leucopus were not subsequently re-
captured after their first capture in the exterior matrix
in 1999 (20/32 = 63%) compared to 2000 (4/12 = 33%),
although the test was not significant (Fisher’s exact
test,n = 44, P=0.1; Figure 2b). Mean monthly rain-
fall amounts were over 50% higher in the summer
(June-August) of 2000 (mean + SE = 11.8 + 0.9 cm)
compared to 1999 (5.3 + 1.0 cm; t,= 4.7, P = 0.009).
Discussion
Patch size
Studies have shown that populations of P. leuco-
pus, unlike birds, insects, and most other mammals,
ANDERSON, MEIKLE, CADY, and SCHAEFER: WHITE-FOOTED MICE
195
100
@ Subsequently
80 trapped in exterior
interior, of not
subsequently
60
recaptured
40 @ Subsequently |
trapped in edge
20
0
b
100
gw Subsequently
trapped in extenor
edge, or interior
@ Not subsequently |
recaptured
Percentage of mice initially captured in exterior matrix
0 aa [hoo
199 2000
Year
FIGURE 2. Percentage of mice (a) subsequently captured in
forest edge habitat in 1999 compared to 2000 and (b)
not subsequently recaptured in 1999 compared to 2000
(see Table 2 for sample sizes) that had been initially
trapped in the exterior matrix habitat.
have greater densities in smaller forest patches (Nupp
and Swihart 1998; Krohne and Hoch 1999; Anderson
et al. 2003). One proposed mechanism to account for
this negative density-area relationship is that habitat
quality is lower in the exterior matrix than in woodland
sites (Nupp and Swihart 1996; Krohne and Hoch 1999).
Predation pressure may be elevated in the agricultural
fields surrounding smaller patches because some pred-
ators may attain higher densities in a landscape with
a variety of land uses (Brown and Litvaitis 1995). We
predicted that fewer mice would move from the forest
edge to the agricultural fields surrounding smaller than
larger forest patches, but our results did not support
the inhibited dispersal hypothesis. Diaz et al. (1999)
found no difference in movements of Apodemus syl-
vaticus to the exterior in relation to patch size, where-
as Krohne and Hoch (1999) found that P. leucopus
populations in Indiana had lower dispersal rates from
smaller patches, a factor that may have resulted in
higher densities in those patches (Adler and Levins
1994). Nupp and Swihart (1996) concluded that both
reduced dispersal and greater abundances of food con-
tributed to the high densities of P. leucopus they ob-
served in small forest woodlots.
196
It is unclear why densities of P. leucopus tend to be
higher in small forest patches (Anderson and Meikle
2006). One possibility is that immigration rates are in
fact greater into smaller forest patches, resulting in
higher densities in those patches (Bowman et al. 2002).
However, we did not find that movements from exterior
to the edge or interior habitats were greater in smaller
than larger patches in this study. If smaller and larger
forest patches do not vary in emigration and immigra-
tion rates, as our data suggest, then a second possibility
is that other factors, such as greater food abundance or
cover from predators, may have resulted in greater abun-
dances of mice in smaller patches or in particular the
edges of small patches (Nupp and Swihart 1996; Diaz
et al. 1999: Anderson et al. 2003; Anderson and Meikle
2006). A third possibility is that lower densities in larg-
er patches may be the result of mice dispersing to low
density regions within the heterogeneous forest patch
to alleviate high densities, with fewer sites likely pres-
ent in smaller forest patches (Krohne and Burgin 1990;
Krohne and Hoch 1999).
Edge habitat
Differences in species composition and vegetation
structure in forest edges compared to forest interiors,
regardless of patch size; may influence the distribution
of small mammals within forest fragments (Matlack
1994; Collinge 1996). Currently, there are conflicting
reports about the effects of edge habitat on the abun-
dance and distribution of White-footed Mice (e.g.,
Cummings and Vessey 1994; Anderson et al. 2003;
Wolf and Batzli 2004). We found that P. leucopus were
recaptured in the same proportion in forest edge com-
pared to forest interior habitat in either year. Our data
on initial and second capture habitat locations support
those studies that reported no difference in the relative
abundance of mice in those two habitats (Heske 1995;
Nupp and Swihart 1996, 1998). This result is somewhat
surprising since forest edge habitats have been shown
to have greater structural complexity in the understory
vegetation compared to edge habitat (Anderson et al.
2003) and White-footed Mice have been shown to
prefer areas with more plant cover (Drickamer 1990).
One explanation could be that mice are not moving
into edge habitat but are already present in high abun-
dances with small territory sizes. Perhaps once territo-
ries are established in early spring, when movements
among high (i.e., edge) and low quality (i.e., interior)
habitats are negligible due to competitive exclusion.
Surrounding habitat type
Types of vegetation in the surrounding matrix habi-
tat may differentially affect the population dynamics
of rodent species in a fragmented landscape. For exam-
ple, Wolf and Batzli (2004) found that predation risk
for P. leucopus was higher at edges adjacent to re-
stored prairies than at edges adjacent to agricultural
fields. Our findings showed that P. leucopus moved in
similar proportions from the forest edge into two dif-
ferent matrix habitat types (i.e., corn and pasture). This
THE CANADIAN FIELD-NATURALIST
Vol. 120
is in contrast to studies that have found that P. leucopus
tend to avoid grassy habitats and can be found in corn
fields at densities as high as those in woodlots (Getz
1961; M’Closkey and Lajoie 1975; Wegner and Mer-
riam 1979, 1990). Mice in our study sites may have
simply responded to the seasonal presence of cover and
not to the type of cover (Cummings and Vessey 1994).
Henein et al. (1998) designed a simulation model illus-
trating flexible and opportunistic responses of P. leu-
copus to landscape structure and connectivity via an
expansion of their use of corn and small grain fields.
Similar results have been found for A. sylvaticus, a
mouse species also described as a habitat generalist
(Garcia et al. 1998; Kozakiewicz et al. 1999).
Annual variation
Rainfall may impact rodent populations directly by
influencing their reproductive status and survival, or
indirectly in relation to changes in food sources and
vegetative cover from predators (Kaufman et al. 1995;
Kesner and Linzey 1997). Monthly rainfall data showed
that totals in the summer of 2000 were almost 50%
greater than in 1999. For mice initially captured in the
exterior matrix habitat, we found that a greater propor-
tion moved to the edge habitat and more were subse-
quently recaptured in the exterior matrix in the wet
year compared to the dry year. If rainfall influenced
patterns of capture among habitats, one explanation is
that in 1999 (i.e., the dry summer) a higher proportion
of mice were not subsequently recaptured from the ex-
terior matrix habitat because they died due to poor con-
ditions, or moved throughout the landscape in search of
other better quality habitats (Kesner and Linzey 1997;
Lidicker and Stenseth 1992). In 2000, the shrub and
herbaceous layers in the forest habitat may have had
more or higher-quality food sources compared to the
exterior matrix as a consequence of rainfall which
caused an influx of mice to the forest habitat (Parmenter
and MacMahon 1983; Bowers and Dooley 1999; Hut-
chinson et al. 1999). Our results illustrate the gener-
alist habitat preferences of P. leucopus but emphasize
annual variation in their distribution within the forest
and surrounding habitats. However, year-to-year dif-
ferences in patterns of habitat use may be influenced by
a number of variables, and a longer-term study would
help to elucidate those factors (Kesner and Linzey
1997):
Acknowledgments
We thank the landowners for permission to use
study sites, in addition to the Ohio Department of
Natural Resources, Hueston Woods State Park, Miami
University’s Natural Areas Committee and Miami Uni-
versity’s Ecology Research Center. We thank J. Bruseo,
M. Spritzer, S. Vessey, and three anonymous reviewers
for their valuable advice. K. Fielitz, J. Gliha, C. Har-
rison, R. Lim, A. Lohrey, D. Markert, M. Tarbox, K.
Wolfe and especially R. Young provided valuable sup-
port in the laboratory and in the field. This work was
2006
supported with funding from Sigma Xi and the Depart-
ment of Zoology at Miami University. Research on live
animals was performed in a humane manner and was
approved by the Institutional Animal Care and Use
Committee (IACUC) at Miami University (protocols
#430 and #467).
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Received 13 May 2005
Accepted 12 December 2006
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Salmon to Arrival of Parasitized Wild Adult Pink Salmon
ALEXANDRA Morton! and RoB WILLIAMS2
' Raincoast Research Society, 13 Cramer Pass, Simoom Sound, British Columbia VOP 1SO Canada email:
wildorca@island.net
2 Pearse Island, Box 193, Alert Bay, British Columbia VON 1A0 Canada
Morton, Alexandra, and Rob Williams. 2006. Response of the sea louse Lepeophtheirus salmonis infestation levels on juve-
nile wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, salmon to arrival of parasitized wild adult Pink
Salmon. Canadian Field-Naturalist 120(2): 199-204.
Recent recurring infestations of Sea Lice, Lepeophtheirus salmonis, on juvenile Pacific salmon (Oncorhynchus spp.) and
subsequent annual declines of these stocks have made it imperative to identify the source of Sea Lice. While several studies
now identify farm salmon populations as sources of Sea Louse larvae, it is unclear to what extent wild salmonid hosts also
contribute Sea Lice. We measured Sea Louse numbers on adult Pink Salmon (Oncorhynchus gorbuscha) migrating inshore.
We also measured Sea Louse numbers on wild juvenile Pink and Chum salmon (Oncorhynchus keta) migrating to sea
before the adults returned, and as the two age cohorts mingled. Adult Pink Salmon carried an average of 9.89 (SE 0.90)
gravid lice per fish, and thus were capable of infecting the adjacent juveniles. Salinity and temperature remained favourable
to Sea Louse reproduction throughout the study. However, all accepted measures of Sea Louse infestation failed to show
significant increase on the juvenile salmon, either in overall abundance of Sea Lice or of the initial infective-stage juvenile
lice, while the adult wild salmon were present in the study area. This study suggests that even during periods of peak inter-
action, wild adult salmon are not the primary source of the recent and unprecedented infestations of Sea Lice on juvenile
Pacific Pink and Chum salmon in the inshore waters of British Columbia.
Key Words: Sea Lice, Lepeophtheirus salmonis Pink Salmon, Oncorhynchus gorbuscha Chum Salmon, O. keta, salmon
farm, British Columbia
The sea louse Lepeophtheirus salmonis is a common
salmon-specific parasite (Nagasawa et al. 1993; Kaba-
ta 1973). While their preferred Pacific host is the Pink
Salmon (Oncorhynchus gorbuscha) (Nagasawa 1987;
Nagasawa et al. 1993), L. salmonis is found on all
species of Pacific salmonids, as well as Atlantic Salmon
(Salmo salar), Sea Trout (Salmo trutta trutta), Rain-
bow Trout (Salmo gairdneri) and Artic Charr (Salveli-
nus alpinus). Once considered benign (Boxshall 1974;
Nagasawa 1987; Berland 1993), L. salmonis is now a
significant problem species on farmed salmon (Pike
and Wadsworth 1999; Johnson et al. 2004). Salmon
farms are a series of floating net pens attached to the
shoreline or anchored in the nearshore. Salmon arrive
from hatcheries and are fed in these pens for 18-24
months, then harvested and sold as food. All farm
salmon in our study area were Atlantic Salmon.
Optimal salinity for L. salmonis survivorship and
reproduction is 30%o (Johnson and Albright 1991b),
with copepodids tolerating salinities as low as 15%c
(Pike and Wadsworth 1999). L. salmonis hatch into a
free-swimming naupliar stage directly from egg strings
attached to the female’s body (Johnson and Albright
1991a). The interval from hatching to infective capa-
bility (copepodid stage) is approximately 4 d at 10°C
and 2d at 15°C (Johnson and Albright 1991b). The
copepodid stage is the first attached stage, thus abun-
dant presence of this stage signals a local source of
gravid Sea Lice and a host salmonid population. When
wild salmon enter freshwater to spawn, the attached
L. salmonis experience decay of life-processes leading
to death (Hahnenkamp and Fyhn 1985; McLean et al.
1990; Johnson and Albright 1991b). As one would
expect from an obligate parasite, there have been no
reports of L. salmonis over-wintering off-host in the
nearshore marine environment.
While early studies on juvenile wild Atlantic sal-
monids Salmo salar and Salmo trutta make no mention
of parasitism by L. salmonis (Heuch and Mo 2001):
Sea Lice are now reported in high numbers on wild
European salmonids adjacent to farms (Tully et al.
1993; Birkeland 1996). Historically, researchers did not
look specifically for L. salmonis on juvenile Pacific
salmon in the inshore environment. However, the sem-
inal works on juvenile Pink and Chum salmon (Healey
1978, 1982; Parker 1965; Parker and Vanstone 1966)
did note the presence of the smaller sea louse, Caligus
clemensi (Parker and Margolis 1964). C. clemensi is
a generalist non salmon-specific parasite that has been
reported on 13 species of fish (Jones and Nemec 2004’).
It seems reasonable that if Caligus was noted, then the
larger, more conspicuous L. salmonis would also have
been reported had it been historically present on juve-
nile Pink salmon.
Today, Sea Lice are reported on juvenile Pink and
Chum salmon in some areas of coastal British Colum-
IP)
200
bia and appear associated with salmon farming (Mor-
ton and Williams 2003; Morton et al. 2004; Morton
et al. 2005; Krkosek et al. 2005; Krkosek et al. 2006).
The impact of Sea Lice is host-size dependant (Grimnes
and Jakobsen 1996; Bjgrn and Finstad 1997) and this
is critical to Pink and Chum salmon as they enter the
marine environment at the smallest size of any salmon-
id. Pink and Chum Salmon weigh less than one gram
in the study area (Broughton Archipelago) throughout
March and April (Morton et al. 2005), and are without
protective scales (A.M. personal observations). Juve-
nile Pink and Chum salmon exhibit prolonged inshore
residency from March to September (Healey 1978).
If some adult wild salmonids occur inshore year round,
their numbers are low. The largest wild host popula-
tion for L. salmonis are the mature Pink Salmon that
enter coastal waters in July and run through Septem-
ber (Heard 1991).
Thus, in a natural setting, a system has evolved that
isolates returning adult salmon from the most juvenile
stage conspecifics. Juvenile Pink and Chum salmon
do not share habitat with a significant L. salmonis host
population until July, by which time they have grown
to weigh several grams. Krkosek et al. (2005) report
that after initial infection by salmon farms, juvenile
salmon schools will re-infect themselves some weeks
later as lice mature and being reproducing.
Salmon farming has altered this arrangement by sit-
ing 26 salmon farms in the nearshore environment of
the Broughton Archipelago (Figure 1), each site hold-
ing potentially 1 million Atlantic Salmon (Naylor et al.
2003), a known host for L. salmonis. In July of 2003
there were approximately 6 million Atlantic Salmon
among the 17 farm sites leased by Stolt Sea Farms with
another nine sites leased by another company (Her-
itage) that did not report numbers (Orr in press). Orr
(in press) reports an average of 2.2 — 9.2 gravid Sea
Lice per salmon on the 6 million farm Atlantic Salmon
for 2003-2004.
While salmon farms are now a recognized source
of Sea Lice, this study further illuminates this subject
by measuring the response in Sea Louse numbers on
wild juvenile salmon to the arrival of a large popula-
tion of this Sea Louse’s preferred host, the adult Pink
Salmon. We hypothesise that if wild adult salmon were
a substantial source of Sea Lice on juvenile salmon in
the study area, then the number of copepodid L. sal-
monis should rise in response to the arrival of in-migrat-
ing adult Pink Salmon. To test this, we first measured
lice levels on juvenile Pink Salmon, then looked for
evidence of copepodid Sea Louse recruitment subse-
quent to wild adult Pink Salmon arrival in the area.
Understanding the mechanisms of L. salmonis infes-
tation of inshore juvenile wild salmon and the relative
importance of both wild and farm lice sources has
become crucial to both wild and farm salmon manage-
ment. This study provides analysis on a previously
THE CANADIAN FIELD-NATURALIST
Vol. 120
unreported aspect of the dynamic between adult and
juvenile Pacific salmon and Sea Lice.
Methods
This study took place in the 400 km’ Broughton
Archipelago in British Columbia. There are seven
major Pink Salmon rivers and 26 Atlantic Salmon farm
sites between the rivers and the open waters of Queen
Charlotte Strait (Figure 1). The Broughton Archipel-
ago’s 200 km of inlets and 90 km of passages and
small bays makes it ideal marine rearing habitat for
juvenile Pink and Chum salmon (Healey 1978). With-
in this region, Fife Sound (Figure 1) through Tribune
Channel is considered a primary route used by return-
ing adult Pink Salmon (G. Neidrauer, Fisheries and
Oceans Canada, Fisheries Patrolman, General Deliv-
ery, Simoom Sound, BC VOP 1S0 personal commu-
nication).
Juvenile Pink and Chum salmon were sampled once
each week at the same sampling location from 1 June
2003 through 19 September 2003 (Figure 1). The sal-
mon farm at this site was fallow, and without any farm
salmon. A 50' x 4' beach seine of %" mesh was used in
June and a 120' x 8' beach seine with 4" mesh and a %"
bunt was used for the remainder of the study. Sample
size ranged from 20 to 30 fish per week. The juvenile
salmon were placed individually in Whirlpak™ bags
immediately after capture, laid on ice and then frozen.
They were later examined under 30x magnification and
the number of Sea Lice, their age-class and sex were
determined and recorded using published taxonomic
keys (Johnson and Albright 1991a). Adult female lice
were classified as either virgin or gravid as per the
Norwegian method of counting sea lice (Karin Box-
aspen, Researcher, Institute of Marine Research, Ber-
gen, Norway). Results are reported only for the salmon-
specific Sea Louse, L. salmonis.
We used three independent methods to estimate the
timing of the wild adult Pink Salmon return to the study
area. First, we used measured Pink Salmon catch-per-
unit directly at our juvenile salmon sampling site. We
used two fishing rods with lures known to attract Pink
Salmon in this area (A.M. personal observations). We
fished for 166 hook-hours between 4 June and 19 Sep-
tember.
The second method compared our catch-per-unit-
effort analysis with the timing of the adult return re-
ported by Fisheries and Oceans Canada (DFO) as
determined by their enumeration program on the known
Pink Salmon-bearing streams nearby (Figure 1). The
Fisheries and Oceans Pink Salmon enumeration flights
began in the second week of August and continued
through the end of October. Finally, a tourism operator
in the Glendale River watershed (Figure 1) had guides
on the river for 3 days in May, 11 days in June, 23 days
in July, 28 days in August, and reported to us when
Pink Salmon were first sighted.
2006
Wakeman River
J
Embly River
50°
oO
Queen
Charlotte Strait
MORTON AND WILLIAMS: SEA LOUSE INFESTATION LEVELS 20)
128
Kingcome
River
Ahnuhati
River
Ahta River Kakwetan
Active farm site
@ Fallow farm site
@ Sample site
FiGurE 1. Study area, the Broughton Archipelago, showing major Pink Salmon rivers, salmon farm sites and
status, and study site.
The adult salmon caught in this study were lifted
from the water without a net and kept free from con-
tact with any surface in order to reduce loss of lice.
They were examined immediately for Sea Louse
counts, stages and species identification using a hand
lens. We estimated the infective capability of the return-
ing adult salmon by counting gravid female sea lice
per fish: a standard measure used to trigger de-lousing
treatment of farm salmon to reduce infective capabil-
ity and lower sea lice numbers on adjacent juvenile
wild salmon (Heuch and Mo 2001).
Louse counts were not expected to follow a nor-
mal distribution. Rather than transforming the data,
we made comparisons using Mann-Whitney U tests
(MWU; the non-parametric equivalent of a two-sample
t-test) in GraphPad version 3.05 by InStat (GraphPad
Software Inc., 5575 Oberlin Drive, #110, San Diego,
California 92121 USA).
We used the following standard measures of louse
infestation rates (Margolis et al. 1982): prevalence = the
proportion of fish infested with lice; intensity = the
mean number of lice on each infected fish; and abun-
dance = the mean number of lice on the entire sample.
Results
We fished for 47 “hook-hours” 4 June through
July, without catching one adult Pink Salmon. We
fished for 119 hook-hours in August until 19 Septem-
ber, and beginning on 4 August, caught 50 adult Pink
Salmon at an average of 0.78 Pink Salmon/hook-hour
(SE = 0.10, n = 17 fishing sessions, range 0-1.4 Pink
Salmon/ hook-hour). Given that there was no variance
in the 12 June/July fishing sessions (all 12 failed ses-
sions to catch a fish), a Wilcoxon signed-rank test was
conducted to assess whether the median catch-per-unit-
effort in August and September differed significantly
from the observed zero catch-per-unit-effort in June
and July. The sum of positive ranks was 136, indicating
that the median value was significantly greater than 0
(P < 0.0001).
Fisheries and Oceans Canada estimated that 188 730
Pink Salmon entered the Broughton Archipelago
rivers from mid-August through September 2003 (G.
McEachen, Fisheries and Oceans Canada, chief of
conservation management for the central coast, Central
Coast Area Office, 315-940 Alder Street, Campbell
River, British Columbia VOW 2P8, personal commu-
202
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Mass, length, and sea lice infection of Pink and Chum salmon collected during the two sampling periods with
average sea surface temperature and salinity for both time periods. “Total” signifies all L. salmonis and “cope” signifies
copepodite L. salmonis only.
Period n Grams Sizecm Salt % Temp °C Prevalence % Abundance Intensity
(SE) (SE) (SE) (SE) (SE) Range (SE)
Total Cope Total Cope Total Cope
June — July 623 6.94 8.47 29.1 12.1 ASO ez 1.20 0.079 2.79 1.09
(0.15) (0.03) (0.03) (0.01) (0.12) (0.012) (0.26) (0.018)
0-42 0-2
August — VAS a2 el Gy eel 223.8 30.8 11.1 400 48 1.00 0.048 2.50 1
19 September (0.57) (0.10) (0.11) (0.05) (0.25) (0.018) (058) (0)
0-33 0-1
nication). This would indicate that Pink Salmon were
present seaward of the rivers and within the study area
for 2-3 weeks prior to the end of the second week of
August, when they were first identified in the rivers,
through September. We cannot be certain how many
of these fish passed the sampling site, but Fife Sound
is considered the primary migration corridor for adult
Pink Salmon (Neidrauer, personal communication).
The tour operator and his guides looked for Pink
Salmon on each of their tours, but did not see Pink
Salmon in the largest Pink Salmon river in the study
area, the Glendale River, until 20 August (H. Pattinson,
owner/operator, Tide Rip Tours, 1660 Robb Avenue,
Comox, British Columbia V9M 2W7 Canada, person-
al communication). Again, assuming that adults took
two to three weeks to migrate through the Broughton
Archipelago to reach the Glendale watershed, one
would expect that the adults reached our sampling
location in early August. In fact, these three methods
show very good agreement, and all point to a best esti-
mate return date for large numbers of wild adult Pink
Salmon in the first days of August. Consequently, we
estimate that juvenile Pink and Chum salmon were
introduced to louse-infected adults around the first day
of August.
We divided the pre- and post-exposure stages into
two sampling periods: 1 June to 31 July, and 1 August
to 19 September. As one would expect from a rapidly
growing cohort, the juvenile salmon increased in both
length and weight throughout the study (Table 1).
The average seawater temperature and salinity varied
slightly between the two sampling periods, but re-
mained at all times within optimal range for L. salmo-
nis (Table 1). The mean number of gravid lice on return-
ing adult Pink Salmon was 9.31 (SE 0.93).
No measure of total Sea Louse occurrence varied
between the sampling periods (Table 1) (abundance —
MWU statistic: 43186, P = 0.4088; intensity - MWU
statistic: 7153.5, P = 0.3412). As well, no measure of
copepodite Sea Lice varied significantly (abundance
—MWU 44072; P = 0.6448).
Given that there was no variance in the observed
intensity of copepodid infection on juvenile salmon in
the later sampling period (that is, all seven fish infected
with copepodites carried a single copepodid louse), a
Wilcoxon signed-rank test was used to assess whether
the median early-season intensity differed from the
observed intensity of 1 louse/fish in the later sampling
season. The mean intensity of infection on the 45 fish
infected with copepodites in the earlier sampling sea-
son was 1.09 (SE 0.043, median 1.00, range 1-2). This
was not significantly greater than the | louse/infected
fish observed in the later sampling season (sum of all
signed ranks = 10, after correcting for ties; P = 0.13).
Discussion
The number of returning adult Pink Salmon carry-
ing gravid Sea Lice was on the order of two hundred
thousand, with approximately nine gravid Sea Lice per
fish and thus we estimate that the juvenile Pink and
Chum salmon in Fife Sound were exposed to the infec-
tive capability of close to two million gravid female
Sea Lice. However no marked increase above the pre-
existing background infection levels could be detected
on juvenile salmon after the adult fish arrived in the
area.
Even if the total number of Sea Lice failed to in-
crease on juvenile salmon after the return of adult Pink
Salmon, an increase in the proportion of lice at the
copepodid stage would have indicated arrival of a
source of Sea Lice that exceeded ambient levels. How-
ever, no increased copepodid settlement was observed.
In fact, prevalence of copepodids was slightly lower
after the adult Pink Salmon returned, which is as one
would expect from an epizootic living out the course
of its infestation. In the early, i.e., pre-exposure sam-
pling period, 7.2% of juveniles sampled carried at least
one copepodid, and after the adult Pink Salmon re-
turned, 4.8% of juveniles sampled carried at least one
copepodid. The juvenile salmon were traveling west
out of an area where gravid Sea Lice occurred on farm
salmon.
The decline we recorded in copepodite Sea Lice is
contrary to what Krkosek et al. (2005) found where
infected juvenile salmon re-infected each other as
their farm-origin Sea Lice matured. The difference in
2006
findings might be a result of geography. Krkosek et
al. (2005) was conducted within the confined water-
way of Tribune Channel, whereas this work occurred
in an area of much greater flush at the archipelago
edge. There are no juvenile Pink and Chum salmon
in Tribune Channel in August. The juvenile Pacific
salmon out-migrations and adult salmon in-migrations
are generally timed such that interaction between the
two cohorts occurs in open waters.
Ocean temperature and salinity were unlikely to have
hindered parasite growth or transfer during August and
September. Mean sea-surface temperature fell 1°C dur-
ing the adult spawning in-migration, which had the
potential to slow reproduction, but salinity increased,
which favoured reproduction. Both values, however,
were well within optimal range necessary for Sea Lice
reproduction (Johnson and Albright 1991b; Boxaspen
and Naess 2000; Tucker et al. 2000). Consequently, we
conclude that the juvenile hosts in our study simply
failed to respond measurably to exposure to a large
influx of infested wild adult salmon.
We fully expected louse loads to increase when the
adult Pink Salmon returned. We know that Sea Lice
must be transmitted among wild salmon. The fact that
Louse numbers did not increase in our study was sur-
prising, and leads us to conclude that, compared to
ambient levels of farm-origin parasitism in the Brough-
ton Archipelago, the contribution of wild-source lice
was negligible. We suspect the higher numbers of
salmon-specific Sea Lice on juvenile Pink and Chum
salmon in June and July originated from the popula-
tion of several million, farmed Atlantic Salmon in the
Broughton Archipelago in high-density open-net pens.
Orr (in press) reports an average of several gravid sea
lice per each of 6 million farm salmon during our study
period, which translates into a very large number of
copepodite sea lice occurring to the east of our study
site area, but not at our study site itself which was adja-
cent to an empty salmon farm.
Why we did not find an increase in Sea Lice in
August and September is perplexing. The mandatory
2-4 day dispersal term in the plankton layer likely
limits Sea Louse recruitment rates to allow both para-
site and host to thrive. Perhaps there were not enough
wild adult Pink Salmon in the Broughton Archipelago
for measurable Sea Lice transmission to the juvenile
cohort in the face of such dispersal. Perhaps Sea Louse
recruitment is an infrequent event between free-ranging
salmon, and our sample size was too small to capture it.
Perhaps the duration of exposure to adult Pink Salmon
was not long enough or the dynamic of juvenile and
adult salmon travelling in opposite directions with juve-
niles along the shoreline and adults mid-channel sup-
pressed copepodid recruitment. Perhaps some small
increase in parasite load did occur, but natural variabil-
ity and small sample size left us with insufficient sta-
tistical power to detect this small effect above the ambi-
ent levels in the Broughton Archipelago. Morton et al.
MORTON AND WILLIAMS: SEA LOUSE INFESTATION LEVELS
203
(2004) reported that in areas without salmon farms
juvenile Pink Salmon were not infested with L. salmo-
nis. Therefore, it is possible that in the absence of farm
salmon the 4.8% prevalence we found on juvenile sal-
mon in August and September could have been entirely
or partially in response to arrival of the wild adult Pink
Salmon.
Likely our inability to detect response in Sea Louse
populations to returning adult Salmon was some com-
bination of these. The lack of observed recruitment of
new Sea Lice suggests that some mechanism has
evolved to suppress transfer of lice from returning
adults to out-migrating smolts in a naturally function-
ing ecosystem. Because Sea Louse pathogenicity is
host-size dependant, protecting the smaller salmon from
L. salmonis infestation would have clear evolutionary
benefits, particularly for the Pink and Chum salmon
that go to sea smaller than any other salmonid.
If exposure to abundant Sea Lice infested wild
adults did not trigger a measurable increase in Sea Lice
loads on juvenile Pink and Chum salmon, then this
leaves the other, less abundant (if at all present), over-
wintering inshore wild salmon species poor candidates
for explaining the enormous and recent spring L. salmo-
nis outbreaks in the Broughton Archipelago.
Clearly, more extensive research is urgently needed.
In the meantime, our study suggests that wild Pink
Salmon do not appear to be the primary source of the
high Sea Lice infestations reported on wild juvenile
Pink and Chum salmon in the Broughton Archipelago.
They may, however, be the primary source infecting
the farm salmon and thus pose a risk to the industry.
We conclude the most obvious source of L. salmonis
on juvenile Pink and Chum salmon in the Broughton
Archipelago is the stationary population of several
million, farmed Atlantic Salmon.
Acknowledgments
This research was supported in part by Tides Canada.
RW is supported by the Jane Marcher Foundation. Our
thanks to Ron Turner for assistance in beach seining.
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ResDocsDocRech/2004/2004_105_e.htm.
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Received 17 April 2005
Accepted 9 February 2007
Physical Characteristics, Hematology, and Serum Chemistry of Free-
ranging Gray Wolves, Canis lupus, in Southcentral Alaska
MATTHEW J. BUTLER!, WARREN B. BALLARD!, and HEATHER A. WHITLAW2
'Department of Natural Resources Management, Texas Tech University, Box 42125, Lubbock, Texas 79409-2125 USA
29305 Winston Avenue, Lubbock, Texas 79424 USA
Butler, Matthew J., Warren B. Ballard, and Heather A. Whitlaw. 2006. Physical characteristics, hematology, and serum chemistry
of free-ranging Gray Wolves, Canis lupus, southcentral Alaska. Canadian Field-Naturalist 120(2): 205-212.
Examination of morphometric characteristics and blood parameters has become a widely used tool for assessing the physio-
logical and nutritional status of wild and captive animals. During 1976 through 1984, 155 Gray Wolves (Canis lupus) were
chemically immobilized in south-central Alaska. Of those, we obtained physical measurements from 132 and blood samples
from 121 individuals. Also, 208 carcasses of harvested and dead radiocollared Wolves were weighed and measured. We obtained
blood samples from three of the fresh carcasses. We measured age, body weight, skull length and width, and upper and lower
canine length. We analyzed blood serum for Ca, P, Fe, chlorides, creatinine, glucose, lactic dehydrogenase, alkaline phosphatase,
glutamic oxalic transaminase, triglyceride, beta globulin, serum urea nitrogen, and uric acid. We obtained packed cell volume
and hemoglobin values from whole blood. We classified samples by season, sex, and age. Seasonal differences were observed
for physical measurements, packed cell volume, alkaline phosphatase, and serum urea nitrogen. Age differences were observed
for physical measurements, hemoglobin, packed cell volume, alkaline phosphatase, P, Ca, creatinine, serum urea nitrogen, and
percent femur bone marrow fat. However, differences among sexes were observed for physical measurements only. These
data provide a baseline for physical condition, hematology, and serum chemistry for free-ranging Gray Wolves.
Key Words: Gray Wolf, Canis lupus, blood, chemistry, hematology, measurements, physical, serum, Alaska.
Examination of morphometric characteristics and
blood parameters has become a widely used tool for
assessing the physiological and nutritional status of
wild and captive animals (e.g., LeResche et al. 1974;
Seal and Mech 1983; DelGiudice et al. 1987; Franz-
mann and Schwartz 1988). Although baseline hema-
tology and serum chemistry values for many species
of captive animals exist, baseline values for free-rang-
ing animals are also necessary to properly interpret data.
It is also important to understand how factors such as
age, sex, and season influence hematology and serum
chemistry values.
Hematology and serum chemistry of captive Gray
Wolves (Canis lupus) has been studied and described
by Fox and Andrews (1973) and Constable (1998) in
Alaska, Seal and Mech (1983) and DelGiudice et al.
(1987) in Minnesota, Pospisil et al. (1987) in the Czech
Republic, and Drag (1995) in Missouri. Seal et al.
(1975) and DelGiudice et al. (1991) in Minnesota,
Messier (1987) in Quebec, and Constable et al. (1998)
in Alaska studied and described hematology and serum
chemistry of free-ranging Gray Wolves. Mech et al.
(1984) reported changes in hematology and serum
chemistry of a female Gray Wolf recuperating from
near starvation in Minnesota. However, little baseline
data exist for age, sex, and season for free-ranging
Gray Wolf populations. The purpose of our study was
_ to describe physical characteristics and report base-
line hematology and serum chemistry values for age,
sex, and season in free-ranging Gray Wolves of south-
central Alaska.
Study Area
The study was conducted in an area of approxi-
mately 45 000 km’, known as Alaska Game Manage-
ment Unit 13, located approximately 125 km north-
east of Anchorage, Alaska. The area was bisected by
three major river systems including the Copper, Nel-
china, and Susitna rivers. Primary prey species were
Moose (Alces alces) and Barren-ground Caribou (Ran-
gifer tarandus granti). Alternate prey species includ-
ed Dall Sheep (Ovis dalli), Beaver (Castor canaden-
sis), and Snowshoe Hare (Lepus americanus). Climate,
topography, and other important aspects of the study
area were described by Ballard et al. (1987, 1991b).
Gray Wolf densities during this study ranged from
10.3/1000 km? in autumn 1975 to 2.6/1000 km? in
spring 1982 (Ballard et al. 1987). Mean annual litter
size in November ranged from 3.7 to 7.3 pups. The
Wolf population was heavily exploited by humans (i.e.,
>40% of population) during the course of this study
(Ballard et al. 1987).
The reference values we present here occurred dur-
ing a period when both Moose and Caribou popula-
tions were increasing. Moose densities were highly
variable in the study area and averaged about 603 to
741/1000 km? (Ballard et al. 1991b) and the popula-
tion increased at a finite rate of increase of 1.03 to
1.06 annually (Ballard et al. 1986). Caribou popula-
tions during this time period increased from approxi-
mately 10 000 in 1976 to about 25 000 in 1984 (Ber-
gerud and Ballard 1988). Winter conditions as reflected
by snow depths were relatively moderate during this
205
206
time period except during 1978-1979 which was classi-
fied as severe (Ballard et al. 1991b).
Methods
We captured Gray Wolves for radiocollaring by dart-
ing them from a helicopter during 1976 through 1984
using methods described by Ballard et al. (1982,
1991a). We immobilized Wolves with phencyclidine
hydrochloride, etorphine hydrochloride, or a combina-
tion of zolzepam hydrochloride and tiletamine hydro-
chloride following dosages and methods described by
Ballard et al. (1982, 1991a). Induction times averaged
7.2 minutes (Ballard et al. 1991a). After immobiliza-
tion each Wolf was radiocollared, weighed, sexed, ear-
tagged, bled, and a number of physical measurements
were recorded on standardized forms. Physical meas-
urements included skull length (SL) and width (SW),
and upper (UCL) and lower canine length (LCL). We
measured canine length from the edge of the gum line
to the tip of the tooth. Skull and tooth measurements
were taken with calipers and recorded to the nearest
mm.
Hunters and trappers were paid $10 (U.S.) per
skinned Wolf carcass. Carcasses were weighed (CW)
and then kept frozen until examined. Skulls were not
boiled and cleaned before measuring to maintain sim-
ilarity with live Wolf measurements. Also, the effects
of freezing carcasses on physical measurements were
likely minimal. We determined percent femur bone
marrow fat (MF) by the dry weight method (Neiland
1970). Because this Wolf population was heavily
exploited (Ballard et al. 1987) we were able to do com-
parisons with many individuals that had been radio- .
collared and harvested within the same year.
We collected blood samples from saphenous veins
using the B-D Vacutainer system (Becton-Dickenson
Co., Rutherford, New Jersey, USA). Generally blood
was taken within 15 minutes of induction time and
serum was separated each evening. Only three blood
samples were collected from carcasses. Those samples
were obtained within 15 minutes of harvest. We used
one EDTA tube to preserve whole blood. Packed cell
volume (PCV) was determined with uncoagulated
whole blood using a microhematocrit centrifuge (Triac,
Clay-Adams Co., Parisppany, New Jersey, USA) while
hemoglobin (Hb) was estimated with an AO Hb-Meter
(American Optical, Buffalo, New York, USA). Clot-
ted blood was centrifuged within 36 h of collection and
serum was then stored frozen at -17°C. We sent frozen
serum to Pathologist Central Laboratory, Seattle, Wash-
ington, USA, where it was analyzed for the following
via an auto-analyzer: calcium (Ca), phosphorous (P),
iron (Fe), chlorides (CHL), creatinine (CRE), glucose
(GLU), lactic dehydrogenase (LDH), alkaline phos-
phatase (SAP), glutamic oxalic transaminase (SGOT),
triglyceride (TRI), serum urea nitrogen (SUN), and uric
acid (UA). Protein fractions were determined by stan-
THE CANADIAN FIELD-NATURALIST
Vol. 120
dard protein electrophoresis for beta globulin (BEG).
Because of unbalanced cells in our analyses, we
pooled years, sorted blood and physical measurements
by sex, age, and season, and conducted multiple analy-
ses of variance. We conducted post-hoc comparisons
with a Scheffe test (Sokal and Rohlf 1995). We deter-
mined correlations among variables by Spearman’s
rank correlation (Conover 1980). We classified ages as
pup (<12 months), yearling (13 to 23 months), and
adult (>24 months), while seasons were divided into
autumn (September through November), winter (De-
cember through February), spring (March through
May), and summer (June through August). Differences
were considered significant when P < 0.05 unless oth-
erwise specified.
Results
We immobilized 155 individual Gray Wolves dur-
ing 1976 through 1984. From that total, we obtained
121 blood samples (n = 121 serum samples, n = 117
whole blood samples) and 132 live body weights
(BW). Due to low sample sizes, we excluded yearling
and summer data from the analyses. In addition, we
weighed and measured a total of 208 carcasses. We
obtained three blood samples from fresh carcasses
(n = 3 serum samples, n = 2 whole blood samples).
Physical Characteristics
Mean BW differed by season (F = 26.121, 2 df,
P <0.001), sex (F = 37.084, 1 df, P < 0.001), and age
(F = 147.890, 1 df, P < 0.001), and there were signifi-
cant interactions between season and age (F = 15.665,
2 df, P < 0.001), and among season, sex, and age
(F = 3.231, 2 df, P=0.043). Lowest BW occurred for
pups during autumn, while highest BW occurred for
adult males during autumn and winter (Table 1). Adult
males had higher BW than male pups during autumn
and winter; adult females had higher BW than female
pups during autumn and spring (Table 1). Body weights
were not different between adult males and adult fe-
males among seasons (Table 1). Body weights between
male and female pups was only significantly differ-
ent during spring (i.e., males were heavier; Table 1).
Body weights of male pups increased with seasons
(i.e., over the course of the biological year); both win-
ter and spring BW were higher than autumn weights,
although winter and spring were not different (Table
1). In addition, BW of female pups increased with
season though female pups were significantly heavier
in the spring than during autumn (Table 1).
Mean CW exhibited similar trends by sex, age, and
season as BW although sample sizes were too small
to evaluate post-hoc differences (Table 1). Season
(F = 4.849, 2 df, P = 0.009), sex (F = 6.665, 1 df,
P=0.011), and age (F = 24.849, 1 df, P < 0.001) each
had a significant effect on CW and there were signifi-
cant interactions between sex and age (F = 3.950,
2 df, P = 0.046), and among season, sex, and age
2006
TABLE 1. Mean physical measurements + | standard error by season, sex, and age for free-ranging Gray Wolves in south
central Alaska during 1976 through 1984. Sample size denoted in parentheses.
BUTLER, BALLARD, and WHITLAW: WOLVES IN SOUTHCENTRAL ALASKA
207
Physical Season
measurements Age Sex Autumn Winter Spring
Live-body weight (kg) Pup Male 26.2+2.3 (8) 37.9 + 14 (10) 413+ 14(16)
Female 23.5+2.0 (7) 33.3'4+.25 (5) 33.34 0.9(20)
Adult Male 46.8+1.5 (8) 48.1 + 1.3 (11) 46.2+ 0.9(17)
Female 38.9+1.4(13) 41.141.2 (3) 424+ 1.0(14)
Carcass weight (kg) Pup Male 19.7+2.7 (3) 30.7 + 0.9 (51) 299+ 25 (8)
Female 25.0424 (3) 27.3 + 0.6 (51) 253+ 10 (7)
Adult Male 39.1 (1) 39.8 + 0.9 (36) 335+ 18 (8)
Female 24.8 +13.8 (2) 32.8 + 0.7 (32) 338+ 1.1 (6)
Skull length (cm) Pup Male 24.5+0.8 (3) 26.2 + 0.2 (58) 27.1+ 0.3(18)
Female 24.5+0.7 (5) 25.1 + 0.2 (52) 25.5+ 0.3(14)
Adult Male 27.7+40.3 (3) 27.8 + 0.2 (39) 27.0+ 04(12)
Female 24.8+1.0 (3) 26.5 + 0.2 (31) 26.4+ 0.4(12)
Skull width (cm) Pup Male 12.34+0.5 (3) 13.5+0.1 (58) 14.44 0.2(18)
Female 124+04 (5) 13.1 + 0.2 (52) 14.0+ 0.7(14)
Adult Male 16.0+0.6 (3) 14.8 + 0.1 (35) 14.9+ 0.3(12)
Female 13.4+05 (6) 14.5 + 0.3 (30) 14.34 0.1(11)
Upper canine length (cm) Pup Male 2.1402 (2) 2.5 + 0.1 (38) 2.6+ 0.1(11)
Female 1940.1 (5) 2.340.1 (45) 2.34 0.1(10)
Adult Male 2.8 + 0.04 (6) 2.9+0.1 (32) 3.0+ 0.1 (7)
Female 23-033) 2.5+0.1 (25) 25+ 0.04(6)
(F = 4.105, 1 df, P=0.018). Lowest CW occurred for
pups during autumn, while highest weights were for
adult males during autumn and winter (Table 1).
There were significant differences in average SL by
season (F = 5.008, 2 df , P = 0.007), sex (F = 23.574,
1 df ,P <0.001), and age (F = 22.331, 1 df ,P < 0.001),
and there were significant interactions between season
and age (F = 3.769, 2 df , P = 0.024) and among sea-
son, sex, and age (F = 3.874, 2 df , P = 0.022). Low-
est mean SL was for pups during autumn and highest
mean SL was for adult males during autumn and win-
ter (Table 1). Males had higher mean SL than females
during each season (Table 1). Mean pup SL increased
from autumn through spring (Table 1).
Mean SW differed by season (F = 3.831, 2 df ,
P=0.023), sex (F = 7.692, 1 df , P= 0.006), and age
(F = 32.160, 1 df , P < 0.001), and there was a signif-
icant interaction between season and age (F = 4.582,
2 df ,P =0.011). Mean male SW was greater than fe-
male SW for all seasons and ages (Table 1). There were
no differences in average adult SW among seasons
(Table 1). Also, mean pup SW increased by season dur-
ing the first year of life (Table 1).
Mean UCL differed by season (F = 5.257, 2 df ,
P = 0.006), sex (F = 19.804, 1 df , P < 0.001), and
age (F = 26.872, 1 df , P < 0.001), and there were no
significant interactions. Mean UCL was lower during
autumn than during either winter or spring (Table 1).
Mean male UCL was greater than that of females (Table
1). Mean UCL for adults was greater than that for pups
(Table 1). There was no significant difference in mean
LCL by season (F = 0.002, 1 df , P = 0.964), sex
(F = 1.707, 1 df , P=0.218), or age (F = 0.828, 1 df,
P = (0.382), and there were no significant interactions.
Mean LCL was 2.3 + 0.07 cm (n =18). Body weight
was also correlated with the other physical character-
istics, MF, and PCV (Table 2).
Hematology
Mean Hb concentration differed by age (F = 7.671,
1 df ,P =0.007) and there was a significant interaction
between age and season (F = 3.876, 2 df , P =0.024):
differences were between pups and adults during win-
ter and autumn (Table 3). Mean pup Hb concentrations
increased throughout the year, whereas adult Hb con-
centrations remained relatively constant during the year
(Table 3).
Average PCV differed by season (F = 6.015, 2 df,
P =0.003) and age (F = 29.083, 1 df, P < 0.001), and
there was a significant interaction between season and
age (F = 10.375, 1 df, P < 0.001). Adults during
autumn had higher mean PCY than all other groups by
season and age (Table 3). Mean PCV did not differ
between adults and pups in winter and spring (Table 3).
Packed cell volume was also correlated with Hb and
BW (Table 2).
Serum Chemistry
Mean SAP concentration differed by season (F = 6.557,
2 df, P= 0.002) and age (F = 19.621, 1 df, P< 0.001),
and there was a significant interaction between age and
season (F = 5.189, 2 df, P = 0.007). Mean SAP con-
centration appeared greater in pups than adults dur-
ing autumn, winter, and spring (Table 3). However,
only during autumn were SAP concentrations signifi-
cantly different between pups and adults (Table 3).
Among pups, autumn and winter SAP concentrations
were higher than spring, but only the autumn—spring
comparison was significant. Among adults, seasonal
208
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 2. Correlations among physical characteristics, hematology, and serum chemistry values for free-ranging Gray Wolves
in south-central Alaska during 1976 through 1984.
Characteristics
Body weight Carcass weight
Skull length
Skull width
Upper canine length
Marrow fat
Packed cell volume
Hemoglobin
Carcass weight
Skull length
Skull width
Upper canine length
Marrow fat
Serum alkaline phosphatase Age
Phosphorous
Calcium
Beta globulin Lactic dehydrogenase
Packed cell volume Hemoglobin
iP n P
0.954 23 <0.001
0.665 67 <0.001
0.639 62 <0.001
0.583 47 <0.001
0.725 14 0.003
0.271 97 0.007
0.362 82 0.001
0.365 108 <0.001
0.281 108 0.003
0.494 108 <0.001
0.316 79 0.005
—0.516 91 <0.001
0.600 140 <0.001
0.511 142 <0.001
0.500 91 <0.001
0.527 102 <0.001
TABLE 3. Mean blood characteristics + | standard error by season and age for free-ranging Gray Wolves in south-central
Alaska during 1976 through 1984. Sample size denoted in parentheses.
Blood
characteristics Age Autumn
Alkaline phosphatase (IU/L) Pup 177.8 + 33.1
Adult 493+ 6.1
Urea nitrogen (mg/dL) Pup 262+ 2.5
Adult 454+ 42
Hemoglobin (g/dL) Pup 179+ 04
Adult 199+ 0.0
Packed cell volume (%) Pup 478+ 2.2
Adult 63.6+ 2.0
SAP concentrations were not significantly different
(Table 3). Serum alkaline phosphatase was also cor-
related to age, P, and Ca (Table 2).
Mean SUN concentration differed by season
(F = 6.746, 2 df, P = 0.002) and age (F = 10.382, 1 df,
P = 0.002), and there were no significant interactions.
Mean SUN concentration was higher during spring
than in winter; autumn SUN concentration was not
different from spring or winter (Table 3). Mean adult
SUN concentration was higher than that of pups,
regardless of season or sex (Table 3).
Mean Pi — 6595s lidine 0.012) ia (i 6.036:
1 df, P=0.016), and CRE (F = 4.189, 1 df, P = 0.043)
concentrations differed by age with no significant inter-
actions. Pup P concentrations were higher than adults,
regardless of season or sex, as were Ca concentrations
(Table 4). Mean adult CRE concentration was higher
than that of pups, regardless of season or sex (Table
4). There were no significant differences in mean val-
ues of BEG, CHL, GLU, Fe, LDH, SGOT, TRI, or UA
by season, sex, or age, and there were no significant
interactions (Table 5). Measures of BEG were also
correlated with LDH concentrations (Table 2).
Season
Winter Spring
(10) 141.9+ 32.0 (15) 65.0+ 6.9 (29)
(21) 90.0 + 38.8 (12) 49.8+ 6.7 (37)
(10) 254+ 60 (15) 46.1+ 4.1 (29)
(21) 376+ 74 (12) 52.9+ 34 (37)
(8) 18.0+ 0.8 (10) 19.14 0.2 (26)
7 (21) 189+ 0.6 (11) 190+ 0.3 (29)
(7) 442+ 1.2 (12) 490+ 1.1 (32)
(23) 55.14 1.8 (11) 497+ 13 (34)
Bone Marrow Fat
Mean MF differed by age (F = 5.062, 1 df, P=0.027).
Mean pup MF was less than adult MF (Table 4). There
were no differences in mean MF by season (F =0.351,
1 df, P = 0.555) or sex (F = 0.000, | df, P = 0.989)
and there were no significant interactions. Percent bone
marrow fat was also correlated to BW, CW, SL, SW,
and UCL (Table 2).
Discussion
Average BW of free-ranging Gray Wolves in south-
central Alaska was greater than captive Gray Wolves
in Minnesota (VanBallenberghe and Mech 1975; Seal
and Mech 1983; Mech 2006) probably reflecting sub-
species differences (Mech 1970). The relationship be-
tween CW and BW may allow managers to gain reli-
able biological data from harvested individuals. Also,
the relationship between BW and MF may allow biol-
ogists to infer condition based upon BW.
Pup and adult Hb concentrations were in the upper
range of values reported for free-ranging (Seal et al.
1975; Messier 1987; DelGiudice et al. 1991) and cap-
tive (Seal and Mech 1983; DelGiudice et al. 1987;
2006
Pospisil et al. 1987) Gray Wolves. In addition, we ob-
served a correlation between Hb and BW and it has
been suggested Hb concentration is a good indicator
of physical condition (Franzmann and LeResche 1978;
Messier 1987; Franzmann and Schwartz 1988). How-
ever, Harlow and Seal (1981) observed little change
in Hb concentrations in food-deprived captive Bad-
gers (Taxidea taxus). Perhaps, lower Hb concentrations
in pups are indicative of a dietary difference due to
social status. Seal and Mech (1983) observed low Hb
concentrations during July and August in captive Gray
Wolves. They also noted reduced BW during this peri-
od. Also, lower Hb concentrations in winter than in
spring were reported by Thomas and Kittrell (1966)
for German shepherds. Changes in Hb concentrations
by season have also been observed in female Black
Bears (Ursus americanus) (Franzmann and Schwartz
1988) and female White-tailed Deer (Odocoileus vir-
ginianus) (DelGiudice et al. 1992). However, our Hb
data were unaffected by season and we observed com-
parable Hb concentrations in summer (18.6 + 1.4 g/dL,
n= 5). Franzmann and LeResche (1978) did not ob-
serve a seasonal effect on Hb concentrations in Moose.
Perhaps, the reduced BW and Hb concentrations ob-
served by Seal and Mech (1983) during summer can be
attributed to a physiological response to greater sum-
mer heat in Minnesota. However, the elevation of Alas-
ka Game Management Unit 13 lies above 1,220 m and
the research conducted by Thomas and Kittrell (1966)
suggested increased elevation resulted in increased
Hb concentrations in German shepherds. Thus, per-
haps, high Hb concentrations in south-central Alaskan
Gray Wolves may have been due to higher elevation.
Thomas and Kittrell (1966) also observed increased
elevation resulted in increased PCV levels in German
shepherds. Perhaps, this explains why mean PCV in
south-central Alaskan Wolves was greater than values
reported by DelGiudice et al. (1991) in free-ranging
Minnesota Wolves. Similar PCV values have been ob-
served in free-ranging (Smith and Rongstad 1980) and
captive (Gates and Goering 1976; Rich and Gates 1979)
Coyotes (Canis latrans).
Franzmann and LeResche (1978) and Franzmann
and Schwartz (1988) suggested PCV was a good indi-
cator of physical condition. We also observed a sea-
sonal effect on PCV. Autumn PCV (59.9 + 2.0% ,n = 30)
was greater than spring (49.4 + 0.8%, n = 66) and
winter (49.4 + 1.6%, n = 23). This suggests Wolves
in Alaska Game Management Unit 13 were in their
best physical condition in autumn. However, differ-
ent seasonal patterns were observed by Thomas and
Kittrell (1966) in German shepherds: lower PCV in
winter than spring. Seasonal changes in PCV have also
been observed in female Black Bears (Franzmann and
Schwartz 1988), female White-tailed Deer (Bahnak et
al. 1979; DelGiudice et al. 1992), and Moose (Franz-
mann and LeResche 1978). Female White-tailed Deer
in Minnesota exhibited two peaks (February-March
BUTLER, BALLARD, and WHITLAW: WOLVES IN SOUTHCENTRAL ALASKA
209
TABLE 4. Mean characteristics + | standard error by age for
free-ranging Gray Wolves in south-central Alaska during
1976 through 1984. Sample size denoted in parentheses
Characteristics Age Mean + standard error (n)
Phosphorous (mg/dL) Pup 58+ 03 (54)
Adult 44+ 04 (69)
Calcium (mg/dL) Pup 96+ 03 (54)
Adult 89+ 03 (70)
Creatinine (mg/dL) Pup 09+ 0.04 (54)
Adult 11+ 0.1 (70)
Marrow fat (%) Pup 79.6+16.9 (64)
Adult 87.0+ 9.2 (44)
TABLE 5. Mean serum chemistry values + | standard error
for free-ranging Gray Wolves (season, sex, and age com-
bined) in south-central Alaska during 1976 through 1984.
Sample size denoted in parentheses.
Serum chemistry values Mean + standard
error (71)
474+ 45 (71)
161.1+ 4.0 (124)
118.7+ 7.4 (122)
306.7 + 22.7 (123)
Iron (umol/L)
Chlorides (mmol/L)
Glucose (mg/dL)
Lactic dehydrogenase (IU/L)
Glutamic oxalic
transaminase (IU/L) 226.5 + 23.8 (120)
Triglyceride (mg/dL) 518+ 5.8 (124)
Beta gobulin (g/dL) 10+ 0.06 (93)
Uric acid (mg/dL) 13+ 0.09(114)
and October-November) in BW and PCV (DelGiudice
et al. 1992). However, PCV levels in Michigan female
White-tailed Deer reached their lowest levels in July
through September when those animals reached slight-
ly lower weights (Buhnak et al. 1979). Female Black
Bear exhibited a decline in PCV during summer and,
though not significant, an increase during autumn
(Franzmann and Schwartz 1988). Franzmann and Le-
Resche (1978) observed greater PCV during June
through October in Moose.
Our SUN data was greater than or in the upper range
of values reported for free-ranging Wolves (Seal et al.
1975; Messier 1987), captive Wolves (Seal and Mech
1983; Drag 1995), captive Coyotes (Rich and Gates
1979; Dunbar and Giordano 2002), and free-ranging
Coyotes (Smith and Rongstad 1980). However, other
studies have reported concentrations similar to this
study. DelGiudice et al. (1987) reported average SUN
concentrations of well-fed captive Gray Wolves be-
tween 37.5 and 44.4 mg/dL. However, during fasting
those concentrations were reduced to a range of 12.0 to
19.8 mg/dL. Another study conducted on free-ranging
Wolves (n = 11) in the Yukon-Charley Rivers National
Preserve, Alaska (Constable et al. 1998), reported SUN
concentrations (46.2 mg/dL) similar to our observa-
tions.
Higher dietary intake of protein is indicated by high-
er SUN concentrations as described for Dogs (Lane
210
and Robinson 1970; Bressani and Braham 1977),
White-tailed Deer (Seal et al. 1972; Bahnak et al.
1979), and Pronghorn (Antilocapra americana) (Seal
and Hoskinson 1978). We observed lower SUN con-
centrations in pups, suggesting a dietary difference,
perhaps, due to social status. We also observed season-
al differences in SUN concentrations. Urea nitrogen
was greatest during spring, intermediate in autumn,
and lowest in winter, suggesting greatest intake of pro-
tein during spring. This may be indicative of increased
Moose vulnerability due to reduced Moose group size
and increased number of calves (Ballard et al. 1991b).
Though sample size was low (n = 5), SUN concentra-
tion during summer was relatively high (41.8 + 6.1 mg/
dL) which was comparable to well-fed captive Wolves
(DelGiudice et al. 1987). Seasonal variation has also
been observed in Black Bears (Franzmann and Schwartz
1988), female White-tailed Deer (Bahnak et al. 1979;
DelGiudice et al. 1992), Santa Cruz Island Spotted
Skunks (Spilogale gracilis amphiala) (Crooks et al.
2003), Santa Cruz Island Foxes (Urocyon littoralis)
(Crooks et al. 2000), and Son Joaquin Kit Foxes (Vul-
pes macrotis mutica) (McCue and O’Farrell 1992). For
each of these species, peak SUN concentrations ap-
peared to coincide with the most productive seasons.
Phosphorus concentrations were similar to those
reported for free-ranging (Seal et al. 1975; Messier
1987; Constable et al. 1998) and captive (Drag 1995;
Constable et al. 1998) Wolves. Our analyses suggest-
ed pup P concentrations were higher than adults.
Messier (1987) observed lower P concentrations in
adults as well. Calcium concentrations were similar
to reported values (DelGiudice et al. 1987 in addition
to those cited above). However, no other Wolf study
reported pup Ca values but we observed pup Ca con-
centrations were higher than adults. We also observed
higher SAP concentrations in pups than adults. This
has been observed in Gray Wolves (Messier 1987),
Coyotes (Smith and Rongstad 1980), and Dogs (Pick-
rell et al. 1974). High SAP concentration is indicative
of bone formation and osteoblast differentiation (Searcy
1969). Serum alkaline phosphatase was also correlated
to age, P, and Ca. Interestingly, of adult SAP values,
11.4% (all male and sampled in different seasons)
were above the lower 95% confidence interval for pup
SAP values. High SAP concentration has been asso-
ciated with mammary tumors in dogs (Karayannopou-
lou et al. 2003), mast cell disease in humans (Pardanani
et al. 2002), pregnancy in animals (Stockham and Scott
2002; Bain 2003), healing of broken bones (Searcy
1969), and other diseases in Dogs (Cornelius 1980).
Thus, SAP concentration may be used to identify in-
jured or diseased individuals.
Conclusions
New patterns in physical condition, hematology, and
serum Chemistry were identified for Gray Wolves. Pups
appeared to be under greater nutritional stress than
adults as suggested by reduced MF, PCV, and SUN and
THE CANADIAN FIELD-NATURALIST
Vol. 120
Hb concentrations in pups. We also confirmed estab-
lished differences in SAP concentrations between adults
and pups. However, abnormally high SAP concentra-
tions were observed in a portion of the adult male
Wolves, perhaps, identifying injured or diseased indi-
viduals. High PCV in autumn suggested Gray Wolves
of this population were in their best physical condition
during autumn. High SUN concentrations in spring
suggested these Wolves were under little nutritional
stress during spring and appeared to be on a good
nutritional plane during summer, as suggested by rel-
atively high summer SUN concentrations. Also, high
Hb concentrations and PCV in this population were
probably a result of higher elevation.
These data, classified by season, sex, and age, pro-
vide a baseline physical condition, hematology, and
serum chemistry for free-ranging Gray Wolves with a
relatively high and increasing prey base in south-cen-
tral Alaska. However, more information on yearlings
and summer is needed to properly understand physical
condition, hematology, and serum chemistry of this
Gray Wolf population. Although we found several cor-
relations among BW, Hb, PCV, MF, and several other
variables, a number of measurements should be used
to assess physical condition.
Acknowledgments
The study was funded by several Alaska Federal
Aid to Wildlife Restoration Projects and the Alaska
Power Authority. Publication costs were paid by the
Department of Natural Resources Management, Texas
Tech University. We gratefully acknowledge L. Au-
miller, T. Balland, A. W. Franzmann, C. L. Gardner, P.
Hessing, L. Metz, R. Toby, J. Westlund, J. Whitman,
and many other employees of the Alaska Department
of Fish and Game for their assistance with the collec-
tion and processing of specimens. Also, two anonymous
reviewers provided valuable comments and suggestions
on earlier drafts of the manuscript. This is Texas Tech
University, College of Agricultural Science and Natur-
al Resources technical publication T-9-1021.
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Received 20 September 2005
Accepted 28 November 2006
Are Lesser Snow Geese, Chen caerulescens caerulescens, Exceeding the
Carrying Capacity of the Fraser River Delta’s Brackish Marshes?
MIKE W. DEMARCHI
LGL Limited, environmental research associates, 9768 Second Street, Sidney, British Columbia V8L 3Y8 Canada; e-mail
demarchi@1g1.com
Demarchi, Mike W. 2006. Are Lesser Snow Geese, Chen caerulescens caerulescens exceeding the carrying capacity of the
Fraser River delta’s brackish marshes? Canadian Field-Naturalist 120(2): 213-224.
Brackish marshes of the Fraser River delta provide important habitats for such high-profile animals as White Sturgeon
(Acipenser transmontanus), Pacific Eulachon (Thaleichthys pacificus), Pacific salmon (Oncorhynchus spp.), Western Sand-
piper (Calidris mauri), and Lesser Snow Goose (Chen caerulescens caerulescens), the latter comprising the “Fraser-Skagit™
segment of the Wrangel Island (Russia) population. This study assessed whether the current numbers of Snow Geese are
exceeding the carrying capacity of brackish marshes in the Fraser River delta. Simulation modelling predicts that those
marshes are presently capable of supporting ~17 500 Snow Geese—a value that is greatly exceeded by the numbers of
geese that have over-wintered there in recent years (~80 000 in 2004-2005). The Pacific Flyway Council's target 3-y aver-
age population and segment sizes of 120 000 and 50 000 - 70 000, respectively, were set without considering the carrying
capacity of natural wintering habitats, the potential impacts of too many geese on upland agriculture, or implications for
hazards to civilian aircraft at Vancouver International Airport. The modelled results of the present study suggest that the
Fraser River delta can sustain the current numbers of Snow Geese that stage or winter there only if those birds also forage in
agricultural and refuge fields—a relatively recent phenomenon that likely bolstered the Snow Goose population. Over-use
by Snow Geese can degrade the productivity and habitat quality of marshes. There is documented evidence that some key
plant species (e.g., Scirpus americanus) of the brackish marshes of the Fraser River delta are well below their biomass
potential (~15%), primarily because of grubbing by Snow Geese. Other species that depend on this brackish environment as
well as human interests in the Fraser River delta may be adversely affected by an overabundance of Snow Geese. The future
effectiveness of hunting as a primary means of population regulation is questioned.
Key Words: Lesser Snow Goose, Chen caerulescens caerulescens, American Three-square Bulrush, Scirpus americanus, brack-
ish marsh, carrying capacity, Fraser River delta, bird strike, British Columbia.
The recent dramatic increase in numbers of the mid-
continent (North America) Lesser Snow Goose (Chen
caerulescens caerulescens) has attracted considerable
attention— mainly because of the habitat degradation
it has caused (e.g., Kerbes et al. 1990; Williams et al.
1993; Ankney 1996). Consumption of agricultural
foods (which may be more abundant and contain high-
er fractions of metabolizable energy than native brack-
ish-marsh plants) and a network of refugia are thought
to have contributed to substantial increases in some
North American populations of Lesser Snow Geese by
improving overwinter survival (Ankney 1996; Jefferies
1997). Although the bird community that winters on
the Fraser River delta contains many species of graz-
ing waterfowl, the large amount of rhizome biomass
consumed annually by Lesser Snow Goose (Boyd
1995) means it is potentially a keystone species there
as it is elsewhere (e.g., Kerbes et al. 1990).
Many Snow Geese that breed on Wrangel Island
(Russia) use the Fraser River delta as staging or win-
tering habitat between October and April (Subcom-
mittee on White Geese 1992*; Boyd 1995; Pacific Fly-
way Council 2006*). They forage in brackish marshes
of the Fraser River delta, primarily on rhizomes of
American Three-square Bulrush (Scirpus americanus)
and shoots of Lyngbei Sedge (Carex lyngbyei). J. P.
Hatfield (personal communication, cited in Campbell
et al. 1990) noted that Snow Geese began foraging in
agricultural fields on the Fraser River delta in 1977.
The number of Snow Geese using those fields rose
from 2000 in the early 1980s to between 20 000 and
30 000 in the 1990s (Boyd 1995; Canadian Wildlife
Service et al. 1999*). The Canadian Wildlife Service
(CWS) and local farmers actively enhance the Alaksen
National Wildlife Area (ANWA) on Westham and
Reifel islands in the Fraser River delta for wintering
Snow Geese and other waterfowl. Fields within the
ANWA are planted with crops to provide the birds with
supplemental feed (Pacific Flyway Council 2006*).
Because Snow Geese and other grazing waterfowl!
can affect private agricultural fields outside the ANWA
by consuming grass, crops, or crop residue such as
corn and potatoes, and by altering surface-water drain-
age, the “Greenfields Program” administered by the
Delta Farmland and Wildlife Trust (DFWT) was estab-
lished in 1990. The goal of that program is to spread
waterfowl impacts over a larger area, thereby lessen-
ing their severity in any one given area (Smith 1996*).
The program encourages farmers to plant cover crops
for soil conservation and provide forage for over-win-
o>)
214
tering waterfowl. The costs of these cropping practices
are shared among farmers, conservation societies, and
government bodies.
Snow Geese that overwinter on the Fraser River delta
are managed according to the Pacific Flyway Council’s
2006 management plan (Pacific Flyway Council 2006*)
which is a revised version of the 1992 plan (Subcom-
mittee on White Geese 1992*). Although the overall
Wrangel Island population target of 120 000 birds (3-
year average) remains the same between the 1992 and
2006 plans, the target size of the Fraser-Skagit seg-
ment (also referred to as “subpopulation” or “flock’”’)
target was increased to a range of 50 000-70 000 birds
in the 2006 plan. According to the 1992 plan, the seg-
ment target was a minimum 3-year average of 35 000
birds. To achieve that, the ideal segment size had been
identified to be between 30 000 and 60 000 birds (S.
Boyd, personal communication). The historical size of
the Wrangel Island population was considered when
the goal was set, but some key factors were not: the
carrying capacity of the Fraser and Skagit deltas, the
possible impacts of Snow Geese on habitat quality and
ecosystem function of the Fraser River delta, nor the
possible impacts of Snow Geese on agriculture or air-
safety. Under the 2006 plan, regulation of the Wrangel
Island population is to occur by way of sport and sub-
sistence hunting.
Burton (1977) concluded 30 years ago that the brack-
ish marshes of the Fraser River delta would be damaged
if the Fraser-Skagit segment exceeded 20 000 Snow
Geese. The purpose of this study was to evaluate, by
way of simulation modelling, recent numbers of Snow
Geese in light of the carrying capacity of the brackish
marshes in the Fraser River delta.
Study Area
The Fraser River sustains the largest delta on Can-
ada’s Pacific coast. The delta covers ~680 km? of aquat-
ic and terrestrial areas, stretching 30 km from New
Westminster westward to Sand Heads Lighthouse and
from Point Grey southward to the international bound-
ary (Butler and Campbell 1987). Brackish marshes
occupy ~3% of the delta.
The Fraser River delta is a critical link in a series
of migratory bird habitats along the Pacific Flyway
between the arctic breeding grounds of North America
and northeast Asia and wintering areas in southwestern
North America, Central America, and South America.
The delta supports some of the highest densities of
wintering waterfowl, shorebirds, and raptors in Cana-
da (Butler and Campbell 1987).
Brackish marshes of the Fraser River delta have
evolved as a result of the interactions between the
maritime conditions of the Pacific Ocean and the fresh
water and sediment load of the Fraser River (Hutch-
inson 1982). Local tides are predominantly semidiurnal
with a maximum range of ~5 m. The Fraser River is
the largest river in British Columbia, with a mean an-
THE CANADIAN FIELD-NATURALIST
Vol. 120
nual flow of 3500 m?/s. Eighty percent of the annual
flow occurs during March to July (Hoos and Packman
1974, cited in Hutchinson 1982). Hutchinson (1982)
divided the present brackish marshes of Lulu Island
into three elevational zones: (1) low marsh—dominat-
ed by Scirpus americanus and S. maritimus; (2) middle
marsh—dominated by Carex lyngbyei, Triglochin mar-
itimum, and S. maritimus; and (3) high marsh—a com-
munity of Agrostis exerata, Potentilla pacifica, Dis-
tichlis spicata, and Typha latifolia.
Prior to dyking, the low elevations of much of the
land near the western limit of the delta were subject to
periodic flooding during spring freshet. Such flood-
ing would have affected the delta’s vegetation commu-
nities. Habitats of this area have been, and are heavily,
modified by human activities. Although much of the
Fraser River delta is used for agriculture, residential,
commercial, and industrial developments are rapidly
increasing at the expense of agricultural land-uses.
Commercial greenhouse operations are increasingly
replacing traditional field-based agriculture.
Much of the upland and foreshore areas of Reifel
Island and Westham Island make up a wildlife sanc-
tuary that is managed by the CWS. The sanctuary com-
prises 586 ha, of which 300 ha includes the federally
owned ANWA that was designated a Ramsar site in
May 1982 (Wetlands International, no date*). Approx-
imately 410 ha of the sanctuary are in cultivated
farmland. The adjacent George C. Reifel Refuge, ini-
tiated in the 1960s, consists of 400 ha of Crown land
designated as a Migratory Bird Sanctuary under the
Migratory Birds Convention Act of 1917.
Nearly 6000 ha in the municipality of Delta is used
to produce vegetable crops and livestock. Of the crop-
land, nearly two-thirds is planted in vegetables such as
potatoes, peas, beans, corn, pumpkins, cabbage, rutaba-
gas. Hay fields, pasture, and corn silage represent more
than one-quarter of the cropland on the delta. The re-
mainder of the cropland on the delta is allocated to
cereals and berries.
Methods
A map of the brackish marshes was prepared by
digitally scanning colour air photos (1:24 000 scale)
at high resolution and ortho-rectifying each photo by
tying control points to those on a TRIM base-map
(1:20 000 scale). Air photos from 18 September 1991
were deemed the best in terms of resolution, scale,
date, season, and tide height. I. Hutchinson (personal
communication) indicated that the brackish marsh
profile has likely undergone very minor net changes
since that time. Habitat polygons were digitized man-
ually on-screen while viewing habitat features at var-
ious magnifications. The digital map upon which habi-
tat calculations were made had a minimum resolution
of 1:10 000. Small patches of Scirpus spp. near south-
western Sea Island were amalgamated because it was
believed that the distribution of those plants had in-
2006
creased since the photos were taken (S. Boyd, per-
sonal communication). Interpretations of habitat type
and zonation in the Fraser River delta were aided by
information from Yamanaka (1975), Moody (1978),
and Hutchinson (1982). Although changes in the marsh
communities have undoubtedly occurred since maps
in those reports were produced, the visual patterns of
intertidal plant zonation were expected to have changed
insignificantly. All mapping was done using the GIS
programs MapInfo and ArcView.
Estimates of Snow Goose numbers since 1987-1988
were obtained from Boyd (1995) and Pacific Flyway
Council (2006*). The numbers of Snow Geese using
agricultural fields were obtained from Boyd (1995) for
the period 1987-1992, and from Canadian Wildlife Ser-
vice et al. (1999*) for the period 1995-1998. Data on
the Greenfields Program were obtained from Smith
(1996*).
Because of variation and error inherently associated
with parameters used in the analyses, a stochastic sim-
ulation model was developed. The model, run in MS
Excel 98, incorporated published and unpublished data
from several sources. Data in Imperial units (e.g., kcal)
were converted to SI units (e.g., kJ). Stochasticity was
incorporated in the model by randomly selecting val-
ues that were within the 95% confidence limits, or
hypothesized range of some parameters.
The model in Appendix | was used to estimate the
carrying capacity of the brackish marshes in the first
half of the overwintering period (“autumn”, versus
the second half of the overwintering period: “spring”).
Because the Fraser River delta experiences substan-
tially more Snow Goose-days during autumn than in
spring (Boyd 1995) and has less autumn habitat than
spring habitat (as mapped in this study), I assumed that
any habitat-imposed limitations on the geese would
occur in autumn. Each simulation first calculated the
metabolizable caloric value of the available S. ameri-
canus. From that value, the number of supportable
Snow Goose-days was determined. In turn, Snow
Goose-days (y) were used to estimate the size of the
Fraser-Skagit segment (x) by using the inverse of the
linear relation: y = 47.764(x) (P = 0.028; r = 0.89;
df = 5; y-intercept = 0) between “autumn” Snow
Goose-days on the Fraser River delta and the size of
the Fraser-Skagit segment as calculated from data
presented in Boyd (1995). The y-intercept of this equa-
tion was set at 0 to account for segment sizes lower
than those presented in Boyd (1995). Five thousand
simulations were run.
A lack of data on two issues may have implications
for the accuracy of the model’s results, but because
the parameters represented by these issues would not
affect the model in a unidirectional manner, their impli-
cations are not viewed as necessarily troublesome.
First, Burton et al. (1979) suggested that Snow Geese
feed almost exclusively on rhizomes of S. americanus
during autumn. In fact, however, Burton (1977, page
DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA
17) stated that Snow Geese strongly prefer both S.
americanus and §, paludosus (the latter also known as.
S. maritimus). Unfortunately, as observed using giz-
zard contents, Burton (1977) could not differentiate
between the cuticle reticulations of the rhizomes of
the two species, thereby casting doubt on the accuracy
of his conclusion about a dual food preference. Abun-
dance of S. maritimus is about one-third that of S.
americanus on the Fraser River delta (Yamanaka 1975).
Second, the model assumes that all biomass grubbed
by the Snow Geese is consumed by them. MclIlhenny
(1932; cited in Burton 1977) guessed that over-win-
tering Snow Geese in the southern United States re-
jected up to 10 times as much grubbed vegetation as
they consumed. Burton (1977) felt that although some
food rejection occurred on the Fraser River delta it
did not likely approach those proportions.
Results and Discussion
Distribution, Abundance, and Carrying Capacity of
Brackish Marshes in the Fraser River Delta
Autumn marsh habitat that is suitable for Snow
Geese occurs in a narrow band along the western edge
of the Fraser River delta’s westernmost islands and
near Brunswick Point (Figure 1). Most spring marsh
habitat is similarly distributed. According to habitat
mapping, the Fraser River delta contains 917.1 ha of
“low marsh” habitat and 946.3 ha of “mid-high marsh”
habitat in areas used by Snow Geese. Low marsh habi-
tat is used by Snow Geese for “grubbing” bulrush
rhizomes in the autumn period and mid-high marsh
habitat is used in the spring for foraging on emergent
sedges (Boyd 1995).
The results of autumn carrying capacity simulations
of the Fraser River delta’s brackish marshes are sum-
marized in Figure 2. During autumn, brackish marshes
can presently support a number of Snow Goose-days
that is commensurate with a mean Fraser-Skagit seg-
ment size of 17 600 geese (range 7400-35 000). Num-
bers larger than this will exceed the calculated carry-
ing capacity of brackish marshes during most years.
According to the model there was only a 1.4% chance
that >30 000 geese could be sustained by the brackish
marshes (Figure 2). Those observations, and the conclu-
sion that §. americanus biomass in the Fraser River
delta is only ~15% of what it might support in the ab-
sence of Snow Goose grubbing (Boyd 1995), suggest
that brackish marshes are either already over-grubbed
or are at considerable risk of being over-grubbed.
Distribution and Abundance of Snow Geese on the
Fraser River Delta
Numbers of Snow Geese in the Fraser-Skagit seg-
ment have been estimated from aerial photos of geese
on the Fraser and Skagit river deltas (Subcommittee
on White Geese 1992*; Boyd 1995; Pacific Flyway
Council 2006*). Most use of the Fraser River delta oc-
curs during early October to mid-January (“autumn”)
and from late February to mid-April (“spring”). Al-
216 THE CANADIAN FIELD-NATURALIST Vol. 120
: Mainland Ser i 130° } 125° 120° 115°
: LOE NANO =
‘ ss (Vancouver) | \af. a
Point, Grey WS8\ British
(6 km) : pe.
55°
, Columbia | ajperta
|
|
I
Sea Island
a YVR Rich d)
ZIG (Richmon
t e(Richmoend
Rete
aS Low Marsh
Lulu Island 4 Mid-High Marsh
(Richmond)
yued uoebinis
2
; 4 a —————
Sand : q Kilometres
Heads ae ~ 9%
Lighthouse
Woodward Island
re ae ;
Complex
Westham Island
(Delta)
Mainland
International Y Brunswick Point (Delta)
Boundary (~7 km) —
Figure 1. Distribution of intertidal marsh habitats used by Snow Geese during the autumn and spring periods on the Fraser
River delta, British Columbia. Runways of the Vancouver international Airport (Y VR) are indicated.
2006 DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA 217
Frequency
S
Fraser-Skagit Sub-Population Size (birds)
FiGuRE 2. Simulation model results of the size of the Fraser-Skagit segment that could be supported by the brackish marshes of
the Fraser River delta for Snow Geese during the autumn. 95% confidence interval on mean: 17 606 +131 (n = 5000).
120000: _——_<_—__—$_——————____—__———_____“_-
100 000 ° 0
® 30000 ,
o se
® P :
oO : 2
el
© 60000 - b—0
—_
ro)
iS)
5
> 40 000 -
20 000 -
fo} o N a \o co fom) N wv \o oo o N vT
~ io) co co foe) oo fo aD oa a a —) So =
aN fon) fon) lon an an fo fo) a an nN —) o oC
ee Maa) ap eh oet AE aad i = = = a A Q
Year /_o— Wrangel Island Population
oe Fraser-Skagit Segment |
FiGurE 3. Estimates of the Wrangel Island Lesser Snow Goose Population size (total, spring) and size of the Fraser-Skagit
segment (overwintering period beginning in year shown). Source: Pacific Flyway Council (2006*).
218
6 000 000 -
5 000 000
4 000 000
3 000 000
Goose Days
2 000 000 | ~
1 000 000
measured
1987-88
1988-89
1969-90
1990-91
1991-92
1992-93
1993-94
THE CANADIAN FIELD-NATURALIST
1994-95 -
1995-96
1996-9?
Vol. 120
O Autumn & Spring
predicted
1997-98 -
1998-99
1999-00 -
2000-01
2001-02
2002-03 -
2003-04
2004-05
Year
FiGuRE 4. Stacked area chart showing measured and predicted number of Snow Goose-days on the Fraser River delta since
1987-1998. Data from 1987-1988 through 1991-1992 were measured by Boyd (1995). The remaining data were pre-
dicted using linear regression equations developed from data for the 1987-1988 through 1991-1992 period (Boyd 1995),
and using population estimates (x values) from Pacific Flyway Council (2006*). Regression equations are as follows:
(1) Autumn: y = 36.4(x) + 448 977 where y = autumn goose-days, and x = annual estimate of Fraser-Skagit segment
(P =0.016, r = 0.94, df = 4). (2) Spring: y = 22.7(x) - 221 987 where y = spring goose-days, and x = annual estimate
of Fraser-Skagit segment (P = 0.116, 7 = 0.78, df = 4).
though geese occur on both deltas simultaneously
during autumn and spring, most or all geese use the
Skagit River delta for the four to five weeks between
these periods. Since the winter of 1978-1979 (when the
first photo counts on the Fraser and Skagit deltas were
done) the Fraser-Skagit segment grew from ~27 000
birds (Pacific Flyway Council 2006*). Owing to high
juvenile production and survival rates of Snow Geese,
the Fraser-Skagit segment increased 3- to 4-fold in
the late 1970s (Boyd 1995). Numbers of Snow Geese
fluctuated in the 1980s then increased thereafter (Fig-
ure 3). Figure 3 shows an upward trend in the size of
the Fraser-Skagit segment from 1978 through spring
2005 (P < 0.001; r = 0.76; df = 26). Recent surveys
indicate that during the winter of 2004-2005 the seg-
ment was the highest ever recorded at ~80 000 birds
(Pacific Flyway Council 2006*). From 1987-2005
the Fraser River delta sustained an estimated 2-5 mil-
lion goose-days annually (Figure 4).
Farm fields apparently play an increasingly impor-
tant role in the over-wintering ecology of the Fraser-
Skagit segment of Snow Geese. The percentage of
Snow Goose days spent on upland fields between 1987-
1988 and 1991-1992 varied among years (Figure 5),
but autumn-use, ranging between 7% and 30%, was
typically much greater than spring use, which ranged
from 0% to 20% (Boyd 1995). Although surveys of use
of agricultural fields on Westham Island were conduct-
ed via ground counts from autumn 1995 through spring
1998 (cf. photo counts up to that point), the use of farm
fields has increased in recent years. From 1995-1996
through 1997-1998, the use of farm fields on Westham
Island (representing ~90% of all use of upland fields
(S. Boyd, personal communication) ranged between
17% and 46% of the total number of goose days spent
on the Fraser River delta in the autumn, and increased
from 13% to 70% of such days in the spring (Figure
5). If foods in brackish marshes were being fully uti-
lized, such a pattern would be expected if more goose-
days were spent on the Fraser River delta. Indeed,
Figure 4 shows a predicted increase in goose days over
time and the linear relation between annual goose days
on agricultural fields and size of the Fraser-Skagit
segment is significant (P = 0.002, r = 0.83, df = 7); the
lowest proportion of field-use occurred when the least
number of goose days were spent on the Fraser River
delta (1989-1990; Figures 4 and 5).
Snow Geese and the Brackish Marsh
The brackish marshes of the Fraser River delta are
probably being over-utilized by Snow Geese. Model
results indicate that the Fraser-Skagit segment exists
at current levels only because the geese can also for-
age in upland fields. Maintaining numbers of Snow
Goose above the carrying capacity of brackish marshes
by way of providing supplemental feed on refuges and
agricultural fields may degrade or continue to suppress
the productivity of the brackish marshes. Boyd (1995)
suspected that grubbing pressure by Snow Geese on
2006
photo counts
Percent of Season Use
BS
S
=!
1987- 1988- 1989-
88 89 90 9]
DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA
bhi al
1990- 1991-
219
ground counts
@ Autumn
0 Spring
1995- 1996- 1997-
G2 796 SOR WOR
Year
FiGurE 5. The estimated percentage of total Snow Goose-days on the Fraser River delta spent on agricultural fields during the
“autumn” and “spring” seasons. Source: 1987-1992, Boyd (1995); 1995-1998, Canadian Wildlife Service et al. (1999*).
+Spring use in 1995-1997 is likely underestimated because surveys ended several weeks before spring migration
(departure).
parts of the Fraser Delta was contributing to a reversal
in marsh succession. Boyd (1995) also speculated that
the reason that production in the brackish marshes of
the Skagit River delta was even lower (i.e., 10%) than
the 15% figure reported for the Fraser River delta was
because the Skagit River delta supported about twice
as many goose-days as did the Fraser River delta dur-
ing 1987-1992. Further, there is some concern that in
recent years a greater proportion of the Wrangel Island
population has been overwintering on the Fraser and
Skagit deltas (Pacific Flyway Council 2006*).
If Snow Geese were excluded from all agricultural
fields, including the sanctuary on Reifel and Westham
islands, the segment would likely decline to a habitat-
limited maximum size of ~15 000-20 000 birds. Al-
though that scenario may be unrealistic, it is possible
that changing land-uses and agricultural practices
throughout the Wrangel Island population’s winter
range, in addition to changes in farmers’ tolerance of
Snow Geese in their fields, could direct even more for-
aging pressure onto brackish marshes. Regardless of
farm-field availability, brackish marshes will always
be vital for Snow Geese because during extended
periods of freezing, the marshes are used exclusively
(Boyd 1995). Use of marshes during cold weather
likely reflects difficulties geese experience in obtaining
food from the frozen or snow-covered soil of agricul-
tural fields.
Snow Geese and Agricultural Fields
Forage from agricultural fields has contributed to
the improved survivorship and subsequent population
growth of Snow Geese in other parts of North America
(Ankney 1996; Jeffries 1997). Possible reasons for the
onset of, and increased use of, upland fields by Snow
Geese on the Fraser River delta are not known with
certainty, but it is plausible that rapid increases in the
Snow Goose population during the late 1970s were
largely responsible (Figure 3).
Burton (1977) predicted that if the Fraser-Skagit
Snow Goose segment expanded beyond ~20 000 birds,
the brackish marshes of the Fraser River delta would
be adversely affected. Although estimates of S. amer-
icanus biomass in the mid 1970s (Burton 1977) and
in the early 1990s (Boyd 1995) are not directly com-
parable due to different sampling strategies, a general
review of the datasets suggests that rhizome densities
during the early 1990s likely approximated those dur-
ing the mid 1970s. This suggests that the brackish
marshes were already being used to capacity by Snow
Geese prior to the onset of farm-field use. Unfortunate-
ly, there are no data documenting trends in S. ameri-
canus biomass immediately following the rise in Snow
220
Goose numbers and onset of field use in the late 1970s
and early 1980s so it is not known whether S. ameri-
canus biomass declined under more foraging pressure
then merely rebounded as incremental foraging pres-
sure on the delta shifted to upland fields.
The increased use of agricultural fields by the geese
supports the hypothesis that the brackish marshes alone
could not sustain the increased foraging pressure that a
rising Fraser-Skagit segment was putting on the Fraser
River delta. If the geese could not meet their energetic
demands in brackish marshes, they would have been
forced to explore alternate food sources such as those
in agricultural fields. Several lines of evidence support
this hypothesis.
First, from the perspective of Snow Goose forag-
ing habitat, agricultural production suitable for Snow
Geese in the local area has probably declined since the
1970s, both in areal extent and crop type (Boyd 1995).
So, contrary to what might be expected, little or no use
occurred when agricultural habitats were better for
Snow Geese. According to W. Temple (DWFT, person-
al communication), there is no evidence that agricul-
tural changes on the Fraser River delta influenced the
initial use of upland habitats by Snow Geese in the late
1970s and early 1980s. W. Temple (personal communi-
cation) indicated that crops that Snow Geese presently
use were grown for many years before the birds started
using them. In 1993, farming of corn, peas, and beans
by commercial processors on the Fraser River delta
was largely halted (W. Temple, personal communica-
tion); thereby further diminishing the area of agricul-
tural fields suitable for foraging Snow Geese. This like-
ly put increased foraging pressure on other fields, and
perhaps, on the brackish marshes.
Second, S. americanus stands in the Fraser River
delta are at about 15% of their biomass potential in the
absence of grubbing by Snow Geese (Boyd 1995).
This suggests that the brackish marshes are not pro-
ducing vast amounts of unutilized rhizomatous foods.
Assuming a “steady state” between rhizome produc-
tion and consumption, as reported by Boyd (1995),
brackish marshes are being fully utilized (even over-
utilized) by consumers such as Snow Geese.
Third, Snow Geese on the Fraser River delta exhibit
a high degree of site fidelity (Burton 1977; Boyd 1995).
Thus, the full or partial abandonment of an area could
indicate decreased habitat suitability. A lack of food in
the traditionally used brackish marshes is a plausible
reason why Snow Geese explored upland feeding areas.
Such exploratory behaviour was predicted by Burton
(1977) before the segment increased and before agri-
cultural fields were used by the geese.
Finally, speculation that changes to hunting seasons
and hunting areas in the Fraser River delta forced Snow
Geese to use upland fields beginning in the late 1970s
is not supported. The existence of a waterfowl refuge
on Reifel Island has conceivably provided Snow Geese
with upland foraging opportunities free of hunting
pressure since the 1960s. Further evidence that hunting
THE CANADIAN FIELD-NATURALIST
Vol. 120
did not precipitate field use by Snow Geese is provided
by Trumpeter Swans (Cygnus buccinator); a protected
(i.e., unhunted) species since the 1930s. Trumpeter
Swans and Snow Geese have similar diets during win-
ter (Boyd 1995; Carter 1997). Trumpeter Swans have
also experienced substantial increases in over-winter-
ing populations in recent years. Carter (1997) noted
that as numbers of Trumpeter Swans wintering near
southwestern British Columbia have increased since
the early 1970s, there has been a recent shift in habitat
use from brackish marshes to agricultural areas. G.
Fowler (CVWMBP, personal communication) indicated
that the use of agricultural fields by Trumpeter Swans
in the Comox Valley on Vancouver Island, British
Columbia, began in the mid-1970s. The onset of swan
use of upland habitats was unlikely a response to any
refuge effects provided by fields, as hunting for other
waterfowl continues to this day on the same farm fields
used by swans. Further, if swans were seeking refuge,
the Courtenay River estuary (their traditional brackish-
marsh wintering ground that has been largely aban-
doned as a foraging area in favour of upland sites)
should still be used for foraging because it has been a
no-shooting area since 1994. The swans roost in the
estuary, but minimal feeding occurs there (G. Fowler,
personal communication).
Conclusions
Currently, the increasing numbers of Snow Geese
wintering on the Fraser River delta (~80 000 in 2004-
2005; Pacific Flyway Council 2006*) exceed the mod-
elled autumn carrying capacity of the area’s brackish
marshes (~17 500) by a large margin. As a result, the
geese are highly dependent on forage provided in near-
by agricultural and refuge fields. Despite compelling
evidence that Snow Geese were grubbing S. ameri-
canus biomass to ~15% of its biomass potential since
at least the early 1990s (Boyd 1995) and a doubling of
the Fraser-Skagit segment from 1995-2005, the 2006
management plan (Pacific Flyway Council 2006*) only
states that increasing numbers of Snow Geese may be
reducing the biomass and extent of S. americanus.
The Pacific Flyway Council (2006* page 13) recom-
mends researching the “trends in biomass and growth
dynamics of American Bulrush, impacts of grubbing
by Snow Geese on intertidal marshes, and the carrying
capacity of the Fraser-Skagit.” The current plan relies
on hunting to regulate the Wrangel Island population
once it exceeds a 3-y average of 160 000 birds (Pacific
Flyway Council 2006* page 32). As society’s values
and interests have shifted, sales of migratory bird hunt-
ing permits and participation in waterfowl hunting in
British Columbia have declined steadily from 1974-
2003 (Environment Canada 2005*). Hunting oppor-
tunity on the Fraser River delta has also diminished
over time. Consequently, it is questionable that man-
agers will be able to rely solely on hunting as a tool to
effectively manage Snow Geese on the Fraser delta
in the future.
2006
Given that the brackish marshes are currently being
used to (present) carrying capacity, any further increas-
es in the Fraser-Skagit segment will likely require:
(1) a substantial increase in the production of S. amer-
icanus rhizomes, (2) a concomitant increase in the use
of agricultural fields by the geese, (3) natural selec-
tion for lower adult body weights, and thus lower gross
energetic demands, or (4) a combination of these.
Recognizing the reduced production of the brackish
marsh, Boyd (1995) recommended that consideration
be given to increasing the productivity of S. ameri-
canus by applying fertilizer. However, unless fertiliza-
tion results in a yield of rhizomes exceeding that which
Snow Geese could potentially consume, there is no rea-
son to believe that Snow Geese would not consume
fertilizer-induced growth back down to the “steady-
state”. The current management plan (Pacific Flyway
Council 2006*) does not identify a need to research
ways of increasing the carrying capacity of the marsh
as was recommended in the 1992 plan (Subcommittee
on White Geese 1992*).
By serving as food for herbivores and detritivores
and as structural habitat (cover) for fish and inverte-
brates, healthy brackish marshes in the Fraser River
delta are integral to an ecosystem that supports such
key animals as White Sturgeon, Pacific Eulachon,
Pacific salmon, shorebirds, and Snow Geese among
many others. Large numbers of Snow Goose have neg-
ative implications not only for the brackish marsh eco-
system, but also for human safety. Bird hazard evalua-
tions have concluded that Snow Geese pose an extreme
hazard to civilian aircraft at the Vancouver Interna-
tional Airport on Sea Island (Demarchi and Searing
1995*). Consequently, an aggressive Snow Goose con-
trol program is necessary each winter near Sea Island
to mitigate that hazard.
A prudent approach to managing the Wrangel Island
population of Lesser Snow Geese would, at a mini-
mum, be sensitive to Snow Goose conservation, poten-
tial adverse effects of Snow Geese on the brackish
marsh ecosystem, and the interactions between Snow
Geese and agriculture, air-traffic safety, and other land
uses in the Fraser River delta.
Acknowledgments
Many people made important contributions to this
study and manuscript. Their assistance is greatly appre-
ciated, but such assistance does not necessarily connote
full agreement with the manuscript’s conclusions. S.
Boyd, Canadian Wildlife Service (CWS) provided
important data and reviewed drafts of this manuscript.
K. Moore and M. Porter, CWS, also provided impor-
tant data (Canadian Wildlife Service et al. 1999). S.
Smith, Delta Farmland and Wildlife Trust (DFWT),
provided data. W. Temple, DFWT, provided informa-
tion about agriculture. C. Levings, Department of Fish-
eries and Oceans and I. Hutchinson, Simon Fraser Uni-
versity, provided information and advice on brackish
DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA
marshes in the Fraser River delta. R. Alisauskas, CWS,
provided information on the current status of bioen
ergetic studies of Lesser Snow Geese and gave con-
structive feedback on the simulation model. R. Trost,
U.S. Fish and Wildlife Service, provided information
about the Pacific Flyway Management Plan. G. Fowler,
Comox Valley Waterfowl Management Project, provid-
ed information about Trumpeter Swans in the Comox
Valley. D. Kraege, U. S. Fish and Wildlife Service,
reviewed drafts of this manuscript. S. Johnson and G.
Searing, LGL Limited, contributed to the design of
this assessment and reviewed a draft of this manu-
script. R. Tamasi, LGL Limited, assisted with GIS and
mapping. G. Glova, LGL Limited, reviewed a draft
of this manuscript as did A. Erskine and an anony-
mous reviewer.
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Environment Canada. 2005. Data accessed via the INTER-
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data_e.cfm#Graph2 Last updated 14 June 2005.
Pacific Flyway Council. 2006. Pacific Flyway management
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Revised Draft. White Goose Subcommittee, Pacific Flyway
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Smith, S. 1996. The Greenfields project 1995/96 historical
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1993. Environmental degradation, food limitation and re-
productive output: juvenile survival in Lesser Snow Geese.
Journal of Animal Ecology 62: 766-777.
Yamanaka, K. 1975. Primary productivity of the Fraser River
delta foreshore: yield estimates of emergent vegetation.
M.Sc. thesis, University of British Columbia, Vancouver,
British Columbia. 133 pages.
Received 10 November 2005
Accepted 20 September 2006
OF THE FRASER RIVER DELTA
JEESE
: SNOW
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224
ADDENDA
Last Flight?
My third, and supposed final, flight of 10 Novem-
ber 2005 was in a De Havilland Dash 8 that was to take
me home from a journey that began a few hours earlier
in a 6-seat Piper Navaho that departed from a snow-
covered gravel strip in northern B.C., then continued in
a Bombardier CRJ from Prince George to Vancouver
(YVR). The westbound departure from YVR’s runway
26L would take us over the marsh along the foreshore
of Sea Island. It was like so many others I’ve been on
since 1994 when I began studying bird-hazards to air-
craft at YVR and helped develop their 24/7 wildlife
control program to mitigate those hazards—many of
which I had witnessed first-hand. There I was, wonder-
ing what critters might be flying around in the flight
path this time. For night-time flights such as this, my
usual thoughts turn to some poor, unsuspecting barn
owl out patrolling the airfield in search of a plump
vole. But this time a far more serious hazard was pon-
dered: snow geese that breed in the Russian arctic and
winter here by the tens of thousands. By chance, I had
just submitted a manuscript for publication earlier in
the week, addressing adverse effects of a burgeoning
snow goose population on the Fraser Delta. And two
days earlier on final approach I had spied a flock of
geese loafing in the marsh near the flight path. I had
heard recently that the wintering flock this year was
bigger than ever.
With plenty of runway lights still below us, we levi-
tated steeply into the darkness. By the time we were
over the marsh, we’d reached an altitude that, based on
my numerous hours of field-observations from a ground
station near the runway’s end, led me to believe that we
had cleanly escaped the main bird-hazard zone yet
again. Still, I peered out the right side at the faintly
illuminated propeller hauling us into the blackness.
At some 300+ km/h in the dark there was no chance
for more than a glimpse of three or four whitish blurs,
accompanied by subtle impact vibrations so transient
that I can’t remember if they were heard, felt, or both.
Snow geese? Gulls? “Shit!” seemed like the right thing
to say, so I did as I sat there with a helpless feeling
wondering what, if any, damage had been done, and on
tenterhooks bracing for more white blurs that, luckily,
didn’t appear.
THE CANADIAN FIELD-NATURALIST
Vol. 120
As the craft began its usual roll toward Victoria I
said to myself, “Okay, maybe I’m making this out to
be a bigger deal than it is,’ thinking that my height-
ened awareness of bird hazards probably causes me
to fret more than I should. Except this time, the turn
continued a bit longer so that instead of heading over
the Strait of Georgia, we were heading southeast over
the lights of Delta. I knew something was amiss. Then
the captain announced that we’d hit some birds and that
we were going back to YVR for an “inspection”. Well,
at least the plane sounded okay and I couldn’t see any
evidence of severe damage like fire or smoke. Touch-
down, taxi, stop, shutdown, back home—sort of.
Sitting on the apron and watching an aircraft
mechanic inspect the wing, cowling, and what looked
to be a bloodstained propeller (I surmised that blood
does not show up so clearly on a black background)
some questions arose. Can his flashlight look-see really
pronounce the plane safe? Should I just grab my gear,
walk off this plane and board another? Is it too late to
catch a ferry? I sat there knowing that bird-strikes are
not that uncommon and that by choice or not, the sim-
ple fact that I choose to fly means that I must have some
faith in our air industry’s standards and its technicians
and mechanics. Besides, what could I know of the con-
dition of any other aircraft in the fleet? When the cap-
tain announced that the plane was damaged and that
we would have to deplane and board another, well,
let’s just say that I was not put out.
As deftly handled by the gate crew on what was
already a very busy evening for regional air-travel out
of Gate C38, within an half an hour 37 of the original
50 of us were lining up to board a shorter Dash 8. The
others might seek the calming benefit of a pint or two
before catching their ride. On the way out of the cov-
ered walkway I engaged the ground crewman’s humour
about this time we should avoid the “shredded tweet”;
taking it as a reference to the fact that this mustn’t be
a particularly unique event if it’s got its own pun. Yet I
must say that his humour was tempered somewhat by
my second-hand knowledge of the deadly havoc a few
birds can wreak when they strike the “wrong” parts of
a fully loaded, outbound plane.
I’m happy to report the most exciting thing about the
fourth of my supposed three plane rides that day was
the “free” candy. Home safe (but a little shaken) in
time for dinner. My next trip to YVR will be on Thurs-
day. Faith be with me.
MIKE W. DEMARCHI
12 November 2005
——————————— LL Lt s—S
Predicting Raccoon, Procyon lotor, Occurrence Through the Use of
Microhabitat Variables
RoGeR A. BALDWIN! 4, ALLAN E. HousSTON?, MICHAEL L. KENNEDY!, and Pin SHUO Liv?
' Department of Biology, The University of Memphis, Memphis, Tennessee 38152 USA
* Ames Plantation, The University of Tennessee, P.O. Box 389, Grand Junction, Tennessee 38039 USA
§ Department of Environmental Science and Geography, William Patterson University, Wayne, New Jersey 07470 USA
* Current address: Department of Fishery and Wildlife Sciences, P.O. Box 30003 MSC 4901, New Mexico State University,
Las Cruces, New Mexico 88003 USA
Baldwin, Roger A., Allan E. Houston, Michael L. Kennedy, and Pin Shuo Liu. 2006. Predicting Raccoon, Procyon lotor, occur-
rence through the use of microhabitat variables. Canadian Field-Naturalist 120(2): 225-231.
Recent increases in Raccoon (Procyon lotor) abundance have been implicated for decreased nesting success of songbirds and
transmission of rabies. Understanding the relationship between occurrence and microhabitat factors should be helpful in
managing this species, though our current understanding of this relationship is inadequate. Therefore, we conducted a study in
western Tennessee during 2000-2002 to determine this association. Occurrence (capture) data were assessed from results of
live trapping at 176 and 112 trap sites during winter and summer, respectively, at three sites. A maximum of 26 habitat variables
were measured at each trap location; all grids were combined for statistical analyses to account for varying relationships
between occurrence and microhabitat factors across different landscapes. Univariate and stepwise logistic-regression analyses
were used to assess associations among microhabitat variables and occurrence. Resulting models were validated through the
jackknife procedure. Predictive equations were constructed from logistic-regression models to compute capture probabilities.
Univariate analyses yielded numerous significant variables with those representing forest characteristics and proximity to water
generally the most significant. Strong concordance was observed between winter and summer seasons for most variables though
several differed (number of large hardwood snags, ground dens, and plant food species, distance to potential water and roads).
Such temporal variability was expected due to seasonal differences in habitat components and biological needs of Raccoons.
Variables included in derived models were similar to those scoring highest in univariate analyses; classification rates for models
(winter = 72%; summer = 78%) were among the highest recorded for generalist species. By accounting for landscape attributes
and replicating across sites, more accurate and useful models were developed. Such models should provide the information
required to effectively manage this species.
Key Words: habitat partitioning, logistic regression, mesopredator, microhabitat, Procyon lotor, Raccoon, western Tennessee.
Raccoons (Procyon lotor) and their associated habi- | Kennedy 1992; Pedlar et al. 1997) indicate various asso-
tats have been the subject of several biological investi-
gations (see Lotze and Anderson 1979; Kaufmann 1982;
Broadfoot et al. 2001; Gehrt 2003). At a landscape level,
this medium-sized predator (mesopredator) is thought
to be most abundant in aquatic associated habitats
(Johnson 1970; Minser and Pelton 1982). However,
Raccoons can thrive in an array of habitats although
| seeming to depend on particular features to reach high
_and stable populations (Broadfoot et al. 2001; Zeveloff
/ 2002). Several investigators (i.e., Morris 1987; Pedlar
vet al. 1997; Chamberlain et al. 2002) have noted that
knowledge of spatial scale and landscape composition
i is integral to understanding habitat associations of spe-
‘cies, though at present, much of the understanding of
‘the relationship of Raccoons to their habitat is based on
assumptions drawn from investigations (e.g., Sanderson
1987; Minser and Pelton 1982; Oehler and Litvaitis
1996) conducted at the landscape level (macrohabitat).
The relationship between the occurrence (defined as
presence at a location) of Raccoons and microhabitat
‘factors has not been studied at multiple sites collec-
tively and findings of previous investigations (Leberg
and Kennedy 1988; Kennedy et al. 1991; Kissell and
—————
i)
ciations depending on season and location. Currently
the relationship between the occurrence of Raccoons
and habitat variables appears to be unclear and in need
of additional study (Kissell and Kennedy 1992; Cham-
berlain et al. 2003), and models allowing for the pre-
diction of occurrence based on microhabitat factors are
lacking from the published literature.
Previous studies (Noss et al. 1996; Oehler and Lit-
vaitis 1996; Rogers and Caro 1998) have indicated that
due to a combination of factors (e.g., removal of top
predators, altered land use, reduced hunting), abun-
dance of mesopredators has increased in recent years.
Estes (1996) noted that increased mesopredator popu-
lations could influence numerous aspects of ecosystems.
For example, high densities can have negative effects
on populations of ground-nesting birds (Crabtree and
Wolf 1988; Vickery et al. 1992; Schmidt 2003) and, by
enhancing the spread of diseases, can impact negative-
ly the health of populations (Carey and McLean 1983;
Hill et al. 1993; Schuburt et al. 1998; Rosatte 2000).
Predators play an important role in structuring biologi-
cal communities (Meffe et al. 1997), and an increase in
abundance of these species requires strong management
25
226
and conservation planning to limit the negative
impact of these taxa on ecosystems. However, sound
management and conservation plans are difficult to
derive without a clear understanding of specific habitat
factors critical to target species. At this time, managers
of natural resources are faced with increasing popula-
tions of Raccoons but limited habitat information on
which to make management decisions (see Rosatte
2000 for the consideration of habitat factors in con-
trolling rabies). In particular, models allowing for the
prediction of Raccoon occurrence are lacking. For
example, certain factors are important components in
determining Raccoon occurrence (e.g., den sites and
aquatic habitats, Pedlar et al. 1997; Broadfoot et al.
2001; Henner et al. 2004). However, the exact rela-
tionship among these components is not known, there-
by requiring individuals to make management deci-
sions without the necessary information (i.e., containing
rabies outbreaks, Rosatte et al. 2001). Therefore, the
purpose of this study was to assess at a microhabitat
scale the relationship of occurrence of Raccoons (based
on capture frequency) with selected habitat factors
from multiple sites representing a mosaic of habitat
types. Specifically, the following predictions were
assessed: (1) there is an association between occur-
rence (based on capture) and selected (individual) habi-
tat variables; and (2) selected habitat variables can be
used to construct models predictive of species occur-
rence.
Study Area
This study was conducted in temperate deciduous
forest in western Tennessee characterized by a frag-
mented landscape consisting of various levels of for-
est, early successional and agricultural fields, residen-
tial buildings, and road systems at three sites. Site 1 was
located at the Edward J. Meeman Biological Station
(Meeman), which was located approximately 20 km
north of Memphis, Tennessee (35°33'N, 90°09'W). This
location was comprised primarily of hardwood forests
with old-field and Kudzu (Pueraria lobata) habitats
interspersed throughout. Old-fields were dominated
by goldenrod (Solidago spp.), fescue (Festuca spp.)
and Switch Grass (Panicum virgatum). Topography
of the site was characterized by numerous drainages
that resulted in a gently rolling terrain throughout the
area. Upland and bottomland forests included various
oaks (Quercus spp.), hickories (Carya spp.), and maples
(Acer spp.) as well as Tulip Poplar (Liriodendron
tulipifera) and Sweet Gum (Liguidambar styraciflua;
Maris 1998). Several smali ponds and intermittent
streams occurred throughout the site.
Sites 2-3 were located at the Ames Plantation
(Ames; 35°06'N, 89°12'W), which was a 7462 ha farm
located northwest of Grand Junction in Fayette and
Hardeman counties, and was operated by The Hobart
Ames Foundation in cooperation with the University
THE CANADIAN FIELD-NATURALIST
Vol. 119
of Tennessee. Ames was located approximately 79 km
southeast of Meeman. Site 2 was located adjacent to
the North Fork of the Wolf River. It was comprised pri-
marily of agricultural fields and bottomland hardwood
forest with some upland forest present. Site 3 was
composed of upland and bottomland forest, cropland,
and old-field habitats. In general, agricultural crops
included Soybeans (Glycine max), Corn (Zea mays),
and cotton (Gossypium spp.). Typical upland tree spe-
cies were Loblolly Pine (Pinus taeda), oaks, and hick-
ories; typical bottomland species included oaks, maples,
Cottonwood (Populus deltoides), and Sweet Gum
(Gabor 1993). Old-field habitats included native warm
season grasses and were similar to those at Meeman.
Topography of upland forest sites was characterized
by gently rolling slopes, whereas bottomland forest,
old-field, and agricultural areas constituted a flatter
topography. Ponds and intermittent streams were num-
erous, and drainages were interspersed throughout all
of these sites.
Methods
Trapping grids were established at each site. Site 1
followed a 5 x 10 trap configuration with traps locat-
ed approximately 150 m apart. Sites 2—3 followed an
8 X 8 trap configuration with traps located approxi-
mately 230 m apart though two trap locations were
excluded from analysis for site 2 as habitat factors were
not measured. Difference in trap configuration for site
1 was due to the limited size of the area. Collectively,
the three sites represented most habitat types occurring
in western Tennessee.
The association of microhabitat variables and occur-
rence of Raccoons was assessed during two periods
that coincided with times when leaves were present
or absent (winter and summer, respectively) on most
woody and herbaceous vegetation (Kolowski and Woolf
2002). During winter, sites were operated on selected
nights from 3 November—7 April, 2000-2002; sum-
mer trapping included only sites 1-2 and were oper-
ated on selected nights from 7 May—12 October, 2001—
2002. An approximate total of 2000 trap nights (one
trap night = one trap set for one night) were observed
for each site during both seasons.
Raccoon-size Tomahawk (Number 108; Tomahawk
Live Trap Co., Tomahawk, Wisconsin, USA) and
Havahart (Woodstream Corporation, Lititz, Pennsyl- -
vania, USA) live traps were used. Traps were baited
with canned cat food in winter and a combination of
cat food, dog food, and doughnuts in summer. Initial-
ly captured individuals were anesthetized with a mix-
ture of ketamine hydrochloride (Ketaset; Bristol Lab-
oratories, Syracuse, New York, USA) and acepromazine
maleate (PromAce; Ayerst Laboratories, New York,
New York, USA) at a 10:1 ratio with 0.1 cc of ketamine
hydrochloride used per estimated kg of live weight
(Bigler and Hoff 1974). Raccoons were tagged in both |
2005
ears with Number 3 Monel (National Band and Tag
Company; Newport, Kentucky, USA) ear tags to deter-
mine recaptures.
For the winter season, 19 habitat variables were
measured to determine the influence of habitat at the
micro-scale on raccoon occurrence. “Slope” represent-
ed the average percentage slope for a 32 m radius
around the trap site as measured by a clinometer. “Total
basal area” [25 cm diameter at breast height (dbh)],
“basal area of small trees” (5-35 cm dbh) and “basal
area of large trees” (>35 cm dbh) represented the
amount of area (m*) covered by trees per hectare and
were determined through use of a prism sweep (10
_ basal area factor prism) conducted at the trap site and
at two additional sites 11.4 m in two random cardinal
' directions. Heights were recorded for each tree meas-
_ ured during the basal area estimation using a haga alti-
+ meter (Forestry Suppliers, Inc., Jackson, Mississippi,
_ USA) with the mean recorded as “average height”.
_ “Number of fallen logs” (£10 cm in diameter) “number
of total snags” (210 cm dbh), “number of small hard-
_ wood snags” (10-35 cm dbh), “number of large hard-
_ wood snags” (>35 cm dbh), “number of pine snags”
(210 cm dbh), “number of ground dens”, “number of
, tree dens”, and “number of total dens” were counted
within a 32 m radius of trap site. An opening of 25 cm
' in diameter was required to be considered a potential
_ den. The “number of plant food species” represented
_ the number of plant species present within a 32 m
_ radius of the trap site that could be utilized as food
+ sources. The “number of stems” 1—5 cm in diameter
- was counted for a 3.2 m radius around the trap site.
_ The procedure was repeated in two random cardinal
_ directions 11.4 m from the trap site and the average
used. “Distance to potential water” represented the
Nearest distance to a water source that held water
_ 230 days a year. “Distance to permanent water” was
1 _ the minimum distance to a water source that held water
/211 months a year. “Distance to road” was measured
to the nearest road or man-made vehicular trail, while
| ieee to open area” represented the nearest dis-
| ba to a non-forested patch. All distance measure-
ments were in meters and were measured using digital
i _ orthophoto quarter-quadrangles georeferenced in Arc-
| View software.
)
Seven additional variables were assessed during the
Bsepiner season yielding a total of 26. “Depth of leaf
j litter” was averaged from 10 random measurements
jtaken within a 32 m radius around the trap site. “Verti-
| Wal cover’ was assessed through the use of a2 mx 0.2m
| ‘cover board checkered with 0.1 x 0.1 m black and
. ‘white squares. The percentage of board uncovered was
i pggorded at the trap site in all four cardinal directions.
‘The percentage of squares uncovered on a spherical
‘densiometer (Forestry Suppliers, Inc., Jackson, Missis-
; ! sippi, USA) was recorded to determine * ‘canopy cover’.
‘Readings were taken in all 4 cardinal directions at the
trap site. “Grass cover”, “woody cover”, “forb cover”
,
|
BALDWIN, HOUSTON, KENNEDY, and LIU: PREDICTING RACCOON OCCURRENCE
nN
nN
~
and “bare soil cover” were visually estimated for a
3.2 m radius around the trap site. All procedures for
measuring summer variables (except “depth of leaf lit-
ter”) were repeated in two random cardinal directions
11.4 m from trap site and the average used.
Because the purpose of this investigation was to
assess at a microhabitat scale the relationship of occur-
rence of Raccoons from multiple sites, statistical analy-
ses were conducted only on data from all sites com-
bined for both winter and summer seasons. A natural
log transformation was applied to all continuous vari-
ables; percentage variables were arcsine transformed
to approximate a normal distribution (Zar 1999). Uni-
variate logistic regression was used to assess associa-
tions between single habitat variables and Raccoon
occurrence with a non-adjusted @ = 0.05 to indicate
significance following suggestions by Moran (2003).
In stepwise logistic-regression analyses, an @ = 0.15
was used as a minimum threshold for inclusion into
the regression function to reduce the data set (Hosmer
and Lemeshow 2000). Multicollinearity effects be-
tween two significant variables were addressed by
assessing correlations among habitat variables. If two
significant variables were correlated at r > 0.70, only
the more significant variable of the pair was included
in further analyses to reduce redundant variables (Ag-
resti 1996).
Variables remaining after univariate logistic regres-
sion were included in a backward stepwise logistic-
regression function and were removed from the model
at P > 0.15. Subsequent models often contained a
large number of variables. Therefore, for practicality
and management purposes, these models were reduced
further by forcing exclusion of variables with lowest
t ratios resulting in a minimum variable model. The ¢
ratio represents the ratio of each regression coefficient
to its standard error. Relative importance of variables
included in the final models was ascertained through
t ratios, with maximal f ratios reflecting the best vari-
able to predict occurrence (Hacker and Coblentz 1993;
Hosmer and Lemeshow 2000; Kolowski and Woolf
2002).
Percentage correct classification of trap sites was
determined using logistic regression models. Accuracy
of these models was determined using the jackknife
procedure as a pseudo-validation technique (Morri-
son 1976; Kolowski and Woolf 2002). This procedure
tested percentage correct classification by removing one
trap site at a time and then classified that site based on
the model built from all other sites combined, result-
ing in a less-biased percentage classification (Hacker
and Coblentz 1993; Kolowski and Woolf 2002). All
statistical procedures were conducted using SYSTAT
10.0 (SPSS 2000).
Results
Trapping resulted in 209 total captures of 112 indi-
vidual Raccoons obtained from 176 trap sites in winter
228 THE CANADIAN FIELD-NATURALIST Vol. 119
TABLE 1. Resulting ¢ ratios and P values from univariate logistic regression of captures of Raccoons (Procyon lotor) com-
pared to habitat variables at three sites during winter and two sites during summer 2000-2002 in western Tennessee. All
variables are considered significant at P < 0.05. Basal area measurements are per ha. Height and distance measurements are
in m. See text for explanation of variables.
Winter Summer
(n = 12454) (n = 8406)
Variable t ratio P value t ratio P value
Slope 2.517 0.012 1.878 0.060*
Total basal area 4.986 <0.001 3.593 <0.001?
Basal area of small trees 4.252 <0.001 3.400 0.001
Basal area of large trees 5.516 <0.001 3.447 0.001°
Average height 4.052 <0.001 2.433 0.015
Number of fallen logs 4.548 <0.001? 2.653 0.008?
Number of total snags 2.677 0.007 1.753 0.080?
Number of small hardwood snags 1.506 0.132 1.647 0.100
Number of large hardwood snags 2.786 0.005? -0.205 0.838
Number of pine snags 2.579 0.010 1.494 0.135
Number of ground dens 1.947 0.052 0.412 0.680
Number of tree dens 3.399 0.0014 2.556 0.011?
Number of total dens 3.410 0.001? 2.265 0.024
Number of plant food species 4411 <0.001? -0.046 0.963
Number of stems 2.657 0.008? 2.140 0.032?
Distance to potential water -6.045 <0.001° -0.170 0.865
Distance to permanent water -2.045 0.041° -3.158 0.002
Distance to road 0.908 0.364 2.645 0.008*
Distance to open area 3.669 <0.001° 2.023 0.043
Depth of leaf litter 0.699 0.485
Vertical cover -0.843 0.399
Canopy cover -3.018 0.003?
Grass cover 0.182 0.855
Woody cover 1.353 0.176
Forb cover -0.742 0.458
Bare soil cover 3.052 0.002
“Variable included in stepwise logistic-regression function.
(site 1 = 100 total captures of 49 individuals; site 2 = 40
total captures of 26 individuals; site 3 = 69 total cap-
tures of 37 individuals) and 173 total captures of 117
individuals from 112 trap sites in summer (site 1 = 93
total captures of 52 individuals; site 2 = 80 total cap-
tures of 65 individuals).
Winter.— Sixteen habitat variables were significant
statistically when compared to Raccoon occurrence
using univariate logistic regression; greatest positive
associations occurred for basal area of large trees and
total basal area, whereas the strongest negative asso-
ciation was for distance to potential water (Table 1).
The only habitat variable not closely related to Rac-
coon occurrence was distance to road. Through step-
wise procedures and model construction, six variables
(four positive, basal area of large trees, number of
plant food species, number of tree dens, basal area of
small trees; two negative, distance to potential water,
number of total dens) were selected (Table 2). Per-
centage of trap sites correctly classified was 72%.
Summer.—Thirteen habitat variables were signifi-
cantly associated to Raccoon occurrence based on
univariate logistic regression; strongest positive asso-
ciations were for total basal area, basal area of large
trees, basal area of small trees, and bare soil, whereas
distance to permanent water and canopy cover had the
strongest negative associations (Table 1). In general,
variables associated to basal area, height of stand, num-
ber of logs, den sites, canopy cover, number of stems, .
bare soil, and distances to permanent water, roads, and |
open areas were associated to occurrence; variables
not associated to occurrence included snags, leaf lit-
ter, food species, vertical density, vegetative ground |
cover, and potential water sources. Three significant |
variables (two positive, total basal area, distance to )
road; one negative, distance to permanent water) were °
selected through subsequent stepwise procedures and |
model construction resulting in the correct classifica-
tion of 78% of trap sites (Table 2).
Discussion
Univariate analyses yielded numerous significant
variables indicating those selected for the current in-
vestigation influenced Raccoon occurrence. These vari-
ables were similar to those correlated in previous inves- |
tigations (e.g., association with forest characteristics,
|
|
|
2005
BALDWIN, HOUSTON, KENNEDY, and LIU: PREDICTING RACCOON OCCURRENCE
TABLE 2. Regression coefficients, ¢ ratios, and P values (P) for Raccoons (Procyon lotor) derived from logistic-regression
functions, as well as percentage correct classifications using the jackknife procedure. Data included in analyses were as fol
lows: winter = three sites totaling 12454 trap nights; summer = two sites totaling 8406 trap nights. See text for explanation
of variables and sampling locations.
Logistic regression
Season Variable Coeff
Winter Constant -4.605
Distance to potential water -0.251
Basal area of large trees 0.194
Number of plant food species 0.491
Number of total dens -0.389
Number of tree dens 0.381
Basal area of small trees 0.149
Summer Constant -4.605
Distance to permanent water -0.338
Total basal area 0.355
Distance to road 0.230
* Percentage correct classification of sites with no captures.
» Percentage correct classification of sites with captures.
© Percentage correct classification of all sites combined.
Leberg and Kennedy 1988; Kennedy et al. 1991; Ped-
lar et al. 1997; Dijak and Thompson 2000; associa-
tion with water sources, Leberg and Kennedy 1988;
_Dijak and Thompson 2000) and suggest that a strong
component of forested areas with large trees and close
proximity to water sources are important factors in
_ predicting Raccoon occurrence. Additional variables
significant in the present investigation were not sig-
nificant in other studies (e.g., average height, slope,
_and distance to roads — Leberg and Kennedy 1988;
Kissell and Kennedy 1992). This lack of concordance
may be attributed to differences in landscape compo-
sition and a lack of replication (Maurer 1986; Temple
and Wilcox 1986; Oehler and Litvaitis 1996). Bald-
win (2003) found similar results when addressing in-
dividual sites for Virginia Opossums (Didelphis vir-
'giniana) in western Tennessee. However, by assessing
multiple sites, noticeable trends were observed. The
lack of concordance for individual sites was attributed
to differing landscape compositions. By assessing mul-
tiple sites of similar landscape composition, the dif-
ference i in significant habitat factors dissipated. Simi-
lar results were observed for Striped Skunks (Mephitis
jmephitis — Baldwin et al. 2004) and suggest that land-
‘scape composition is an important factor when con-
seetng models predictive of occurrence (Temple and
) Wilcox 1986; Dijak and Thompson 2000).
Differential use of habitat by Raccoons across sea-
sons has been documented (e.g., Lotze and Anderson
1979; Kaufmann 1982; Chamberlain et al. 2002), al-
though quantitative assessments of these shifts among
seasons have received little study. Nevertheless, con-
Le nce of significant variables across seasons was
generally high although a few exceptions were noted.
For example, large hardwood snags and ground dens
Were important factors during winter but were non-
i
i
\\
Jackknife classification
t ratio P No capture* Capture” “Combined
-10.656 <0.001 70 We 72
-4.671 <0.001
2.732 0.006
2.583 0.010
-2.249 0.024
2.081 0.037
2.046 0.041
-7.034 <0.001 57 88 78
-4.177 <0.001
3.322 0.001
2.656 0.008
significant during summer. Both variables provide ther-
mal shelter for Raccoons during winter (Stains 1961;
Sanderson 1987) but may be less important during
warmer seasons. Likewise, number of plant food
species and distance to roads were significant factors
during winter but not in summer. The non-signifi-
cance of these variables may be related to the abun-
dance of food and the lack of hunter use of roads dur-
ing summer. This non-use of roads as travel corridors
may have resulted in increased use of stream banks
during winter (Hilty and Merenlender 2004), thus
yielding different results between seasons for dis-
tance to potential water. Ultimately, temporal vari-
ability should be expected due to seasonality in vari-
ables measured as well as in the biological needs of
Raccoons and should be considered when modeling
Raccoon occurrence.
Most previous investigations have attempted to deter-
mine those factors most important in influencing Rac-
coon occurrence at the micro-scale but did not include
modeling techniques in their analyses (Leberg and
Kennedy 1988; Kennedy et al. 1991; Kissell and Ken-
nedy 1992; but see Pedlar et al. 1997 for different mod-
eling strategy). Such modeling strategies allow for the
prediction of heavy-use areas by Raccoons and are
particularly useful to managers of natural resources.
Therefore, such procedures were incorporated into the
current investigation and yielded models that gener-
ally included the strongest variables from univariate
analyses (i.e., forest and water characteristics). Result-
ing models maintained high classification rates while
allowing managers to focus only on those variables
most important for predicting Raccoon occurrence.
Such techniques have yielded slightly higher classifi-
cation rates for habitat specialists [Fisher (Martes pen-
nanti) = 79% — Carroll et al. 1999; Iberian Lynx (Lynx
230
pardinus) = 83% — Palma et al. 1999] likely due to
their need for more specific habitat components. None-
theless, classification rates observed in the current study
are generally higher than those reported for other hab-
itat generalists [Bobcat (Lynx rufus) = 59-70% —
Kolowski and Woolf 2002; Striped Skunk = 56-75%
— Baldwin et al. 2004]. Therefore, models that uti-
lize replicated sites and account for differences in land-
scape composition could serve as a blueprint for future
investigations involving habitat generalists ultimately
resulting in models more useful for wildlife managers.
Unfortunately, construction of such models can be
expensive. Care must be taken to develop an appro-
priate sampling strategy to maximize results.
Acknowledgments
Partial funding for this project was provided by the
Tennessee Wildlife Resources Agency, the Ames Plan-
tation, and The University of Memphis. Thanks are
extended to S. B. Franklin for critical review of the
manuscript, and to M. Biernacki, S. B. Franklin, and
T. A. Wasklevicz for assistance with statistical analy-
sis and GIS application. Special thanks go to B. D.
Carver, J. R. Hisey, J. B. Jennings, S. J. Mahady, and
numerous students from The University of Memphis,
as well as several individuals from the University of
Tennessee and the Ames Plantation for their assis-
tance in data collection.
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Notes
Scoter, Melanitta spp., Migrations Interrupted by Confederation Bridge:
An Update
KATHERINE BUNKER-POPMA
40 Weldon Street, Sackville, New Brunswick E4L 4N4 Canada
Bunker-Popma, Katherine. 2006. Scoter, Melanitta spp., migrations Interrupted by Confederation Bridge: An update. Canadian
Field-Naturalist 120(2): 232-233.
Continued monitoring over 10 years of scoter migrations through Northumberland Strait confirmed that these birds continue
to perceive the Confederation Bridge (completed in 1997) as an obstacle. Such problems — and alternatives not causing
them — need serious consideration when “strait crossings” are contemplated elsewhere.
Key Words: Scoters, Melanitta spp., migrations, Northumberland Strait, bridge, New Brunswick.
While the Confederation Bridge (46°15'N, 63°40'W)
linking Prince Edward Island to New Brunswick was
being completed in 1997, Hicklin and Bunker-Popma
(2001) examined the possible influence of the bridge
upon migration by marine birds. They concluded that
migrating scoters (Melanitta spp., seaducks) were un-
willing to fly under the (40 metre high) bridge and often
reluctant to fly over it.
Earlier information, recently supported by several
studies (e.g. Wilson et al. 2003*), suggested that, in
recent years at least, most migrating scoters passing
through the Maritime Provinces in spring cross the
Chignecto Isthmus 40-50 km. further west and do not
approach the Confederation Bridge. Much less is
known about their fall migrations.
TABLE 1. Ten years of scoter movement at Confederation Bridge, New Brunswick/Prince Edward Island. Spring (S) and fall
(F) totals shown separately for each year.
Year Observation Scoters seen to Scoters seen to pass
time (hr) approach bridge over bridge* (%)
1997 S 54 731 243 (33)
F 102 876 338 (39)
1998 S 15 70 29 (41)
F 49 923 388 (42)
1999 S 4 16 0
F 78 1495 881 (59)
2000 S 35 389 67 (17)
F 74 1675 859 (51)
2001 S 16 212 115 (54)
F 72 906 431 (48)
2002 S 18 461 150 (33)
F 70 1368 725 (53)
2003 S 27 604 129 (21)
F 50 1824 1274 (70)
2004 S 28 SS) 275 (53)
F 47 1253 696 (56)
2005 S 11 46 6 (13)
F 22 435 156 (36)
2006 S 14 124 68 (55)
F 40 1038 547 (53)
Totals S 222 3168 1082 (34)
F 604 11793 6295 (53)
Grand Totals 826 14961 7377 (49)
* No scoters were seen to pass under the bridge
May,
2006
I monitored seaduck migrations at Confederation
Bridge each year since 1997, using procedures of our
original study (Hicklin and Bunker-Popma 2001) to
see if these birds habituated to the perceived obstacle.
Results of these observations, summarized in Table 1,
suggest that the majority of scoters still, after 10 years,
have not learned a simple, general, way of passing the
bridge.
Nearly half of those seen (34% in spring, 53% in
fall) circled upwards and passed extremely high over
the bridge without landing. Others landed on the
water, and remained there, often for hours. Still oth-
ers followed the bridge towards the opposite shore,
and sometimes were lost to sight while still flying. As
my observations do not suggest that birds in the two
latter categories stay near the bridge indefinitely, pre-
sumably they continue their migrations later, but tim-
ing of their departures and the routes followed remain
unknown.
Given the small fraction of regional scoter popula-
tions that approach it, this bridge cannot be considered
a major obstacle to their migrations, at a flyway level.
Nevertheless, the continuing problem posed to the birds
that encounter this bridge needs to be borne in mind
whenever and wherever bridges are proposed in loca-
NOTES
tions traversed by major portions of migratory species
populations.
Acknowledgments
A. J. Erskine edited my original draft and encour-
aged publication of this update.
Document Cited (marked * in text)
Saint John Naturalists Club, Inc. 2003. Mystery migra-
tion: Secret of the Black Scoter (bilingual). Video (23:45
English, 24:15 French); filmed and edited by Rowan Ridge
Productions Ltd. (This video refers to several unpublished
studies on scoter migrations in the Maritime Provinces).
Literature Cited
Hicklin, P. W., and K. Bunker-Popma. 2001. The spring and
fall migrations of scoters, Melanitta spp., at Confederation
Bridge in the Northumberland Strait between New Bruns-
wick and Prince Edward Island. Canadian Field-Naturalist
115: 436-445.
MacKinnon, C., and A. Kennedy. 2006. An observation of
the spring 2006 migration of Black Scoter, Melanitta nigra,
in Northumberland Strait interrupted by the Confederation
Bridge. Canadian Field-Naturalist 120(2): 233-234.
Received 14 December 2004
Accepted 26 February 2007
An Observation of the Spring 2006 Migration of Black Scoter
Melanitta nigra, in Northumberland Strait, Interrupted by the
Confederation Bridge, New Brunswick — Prince Edward Island
COLIN MACKINNON and ANDREW KENNEDY
Canadian Wildlife Service, Atlantic Region, P.O. Box 6227, Sackville, New Brunswick E4L 1G6 Canada
MacKinnon, Colin, and Andrew Kennedy 2006. An observation of the spring 2006 migration of Black Scoter Melanitta
nigra, in Northumberland Strait, interrupted by the Confederation Bridge, New Brunswick — Prince Edward Island.
Canadian Field-Naturalist 120(2): 233-234.
An observation from the bridge of a flock of Black Scoters suggests that, nine years after construction, the 12.9 km Confed-
eration Bridge may still be a partial barrier to bird migration. Only 3 of 18 scoters (16.7%) crossed over the structure during
the observation period.
Key Words: Black Scoter, Melanitta nigra, seaducks, migration, Confederation Bridge, Northumberland Strait, New Brunswick,
Prince Edward Island, Cape Jourimain National Wildlife Area.
Construction of the Confederation Bridge began in
1995 and was completed in 1997. This 12.9 km con-
crete structure spans the Northumberland Strait at its
narrowest point between the Cape Jourimain Nation-
al Wildlife Area, New Brunswick, and Borden, Prince
Edward Island. As part of collecting pre-construction
baseline data in 1990, MacKinnon et al. (1991) mon-
itored seabird and seaduck movement through the
Northumberland Strait. Twenty-two hours of surveys,
during spring and fall, produced 5214 observations on
25 species. Of these, 2041 (39.1%) were seaducks,
1400 (26.9%) were three species of scoters and 641
(12.3%) were Common Eider, Somateria mollisima
(MacKinnon et al. 1991).
During the early 1990s, concerns were expressed
over possible effects the Confederation Bridge may
have on seabird migration. The closest comparable
existing structure was the Canso Causeway, connect-
ing Cape Breton Island with mainland Nova Scotia.
Erskine and Smith (1986) reported that, although the
causeway was a comparatively low and short structure,
Common Eider were believed to have treated the struc-
ture as a barrier to flight. A flock of eiders flew parallel
to the causeway, gained altitude, and eventually crossed
over. The question was, would seaducks do the same
with the Confederation Bridge?
In 1997, Hicklin and Bunker-Popma (2001) con-
ducted surveys at Confederation Bridge during spring
234
migration, similar to those reported by MacKinnon et
al. (1991). The Confederation Bridge was then opera-
tional, and their observations suggested that it formed
a partial barrier to scoter migration. Of 3986 and 1441
scoters observed by them during respective spring and
fall migration, only 12.8% of scoters crossed over the
bridge during the spring and slightly more, 21.9%, in
the fall. Bunker-Popma (2006) subsequently reported
additional years of observation.
The Confederation Bridge utilizes a multi-span con-
crete box girder structure and is composed of three main
components: the approaches (east and west), a 40 m-
high deck and 60 m-high ship-navigation span. The east
approach is 0.6 km over 7 piers and the west approach
is 1.3 km over 14 piers. The main bridge is 11.0 km (44
piers) in length. In the middle of the bridge is a 2.50 km
navigation section. The typical span, between piers, is
250 m.
The Confederation Bridge has been in operation for
nine vears. During the morning of 13 April 2006, we
observed the behaviour of a flock of Black Scoters,
Melanitta nigra, during their northward migration
(46°13'N, 63°44'W). This observation proved more
interesting as the authors were travelling across the
bridge, by vehicle, at the time and were able to track
the flock’s activities over most of the crossing. At the
time of observation, visibility was excellent, no clouds,
+6°C and wind light SSW.
Shortly after we entered the Confederation Bridge
from Borden, Prince Edward Island, at 08:10, a flock
of 18 Black Scoters was observed flying parallel with
and approximately 100 m south of the Bridge at a
height of approximately 40 m above the water. The
flock was observed to rise and approach the bridge
deck on three occasions. On two occasions, the flock
approached the deck and then veered back to its origi-
nal flight path, parallel with the Bridge. On the third
occasion, the flock gained altitude and, similar to the
previous occasions, aborted its attempted flight over
the bridge after coming within approximately 50 m of
the structure. However, this time the flock completed a
clockwise circle, towards the bridge, in order to regain
their original flight path. At the instant in which the
flock began this circle, three birds broke from the main
group and promptly crossed over the Bridge, behind
our vehicle, at a height of approximately 10 m above
the bridge deck. This crossing occurred approximate-
ly 4.5 km from the Prince Edward Island side of the
THE CANADIAN FIELD-NATURALIST
Vol. 120
bridge. The velocity of the flock was constant with
our driving speeds of about 80 km/h, until it could no
longer be viewed in the vicinity of the navigation
channel. While rising over the navigational channel,
the scoters remained out of sight below us. We main-
tained a constant speed of 80 km/hr and the flock was
immediately re-sighted once we descended from the
elevated navigation span. Upon re-locating the flock,
we noted that the group had decreased in size, while
out of sight, from 15 to 8 birds. We did not observe any
scoters coming into view on the north side of the Bridge
and assume the missing seven birds did not cross the
structure. The remaining eight birds eventually landed
in the water approximately 100 m south of the bridge
and 500 m from the New Brunswick coastline (46°10'N,
63°49'W). Observations ceased at 08:25. It is notewor-
thy that the Confederation Bridge is built in a general
“S” curve, thus the flight of the scoter flock was fol-
lowing the contours of the bridge and not a straight line.
This observation demonstrates that nine years after
the Confederation Bridge was built, some seaducks,
such as Black Scoters, still may perceive the structure
as a partial barrier to their northward migration through
the Northumberland Strait.
Acknowledgments
We thank P. Hicklin, K. Davidson and A. J. Erskine
for helpful reviews of the manuscript.
Literature Cited
Bunke-Pompa, K. 2006. Scoter, Melanitta spp., migrations
interrupted by Confederation Bridge: an update. Canadi-
an Field-Naturalist 120(2): 232-233.
Erskine, A. J., and A. D. Smith. 1986. Status and movements
of Common Eiders in the Maritime Provinces. Pages 20-29
in Eider Ducks in Canada. Edited by A. Reed. Canadian
Wildlife Service, Report Series Number 47. 177 pages.
Hicklin, P., and K. Bunker-Popma. 2001. The spring and
fall migrations of scoters Melanitta spp., at Confederation
Bridge in the Northumberland Strait between New Bruns-
wick and Prince Edward Island. Canadian Field-Naturalist
115: 436-445.
MacKinnon, C. M., R. W. Daury, and R. J. Hicks. 1991.
Seabird and seaduck movement through the Northumber-
land Strait, 1990. Technical Report Number 130. Canadian
Wildlife Service — Atlantic Region. 86 pages.
Received 25 April 2006
Accepted 6 February 2007
2006
NOTES
First Record of a River Otter, Lontra canadensis, Captured on the
Northeastern Coast of Alaska
SHAWN P. HASKELL!
'LGL Alaska Research Associates, Inc., 1101 E 76th Avenue, Suite B, Anchorage, Alaska 99518 USA; e-mail: shawn haskell @
ttu.edu
Haskell, Shawn P. 2006. First record of a River Otter, Lutra canadensis, captured on the northeastern coast of Alaska. Canadian
Field-Naturalist 120(2): 235-236.
A River Otter (Lontra canadensis) was captured on the Beaufort Sea coast of northeastern Alaska during a coastal fish study
in late July 2001. The otter was dead upon retrieval and of adult size. River Otter distribution is not known to extend north
of Alaska’s Brooks Range. This occurrence may have represented a one-time spring dispersal event. However, fluviomor-
phological features of some river systems may provide suitable habitat for an overwintering otter in this region of Alaska’s
North Slope.
Key Words: River Otter, Beaufort Sea, Lutra canadensis, fyke net, distribution, Alaska.
In late July 2001, a River Otter (Lontra canaden-
sis) of adult size was retrieved from a shoreline fyke
net in Lions Bay, a barrier island lagoon system on
the Beaufort Sea coast of northeastern Alaska. The
live-capture fishing net was located at 70°10.024'N,
146°12.557'W, approximately 82 km east of Prudhoe
Bay and 11 km west of the Canning River; the nearest
non-industrial areas of human habitation are Barter
Island, 100 km to the east, and the Colville River delta,
170 km to the west. A westerly wind event had caused
coastal water levels to rise, completely submerging the
trap-end of the net in about 1.5 m of water, drowning
the otter. Physical samples or measures were not taken
of the specimen. The animal was discarded to the water
at the capture site. About two weeks after the capture,
the author retrieved the carcass for brief examination.
Marine arthropods of the family Mysidae had entered
all openings in the otter’s body, expediting decompo-
sition.
River Otter distribution is commonly noted as ex-
tending far north up the Mackenzie River drainage of
northwestern Canada but then falling south of the
Alaskan Brooks Range to the west (Rue 1981; Whitak-
er 1996; Forsyth 1999; Wilson and Ruff 1999). Bee and
Hall (1956) reported a River Otter sighting in the cen-
tral Brooks Range at Kanayut Lake and tracks observed
near the Beaufort Sea coast at the Kuparuk River.
Manville and Young (1965) also displayed a point loca-
tion near the mouth of the Kuparuk River, but given
Survey methods, that point may stem from the same
observation of tracks documented by Bee and Hall
(1956). A commercial winter fishery operated in the
Colville River Delta since the early 1950s has never
captured a River Otter (James Helmericks, homestead-
er and naturalist, personal communication). There are
only two state sealing records for River Otters in Alas-
ka’s North Slope region; one was caught near Wain-
wright in northwestern Alaska in 1990 and the other
near Anaktuvuk Pass in the central Brooks Range in
1985 (Geoff Carroll, Alaska Department of Fish and
Game Area Biologist, personal communication). Biol-
ogists have reported seeing small numbers (i.e., one or
two) and tracks of otters north of Anaktuvuk Pass but
south of the Arctic Coastal Plain on the Anaktuvuk and
Chandler rivers, both tributaries to the north-flowing
Colville River, where freshwater upwellings provide
open water throughout winter and habitat for fishes
such as Arctic char (Salvelinus alpinus; Geoff Carroll,
Alaska Department of Fish and Game Area Biologist,
personal communication).
It is unknown whether the 2001 otter capture in
Lions Bay signified an anomalous extralimital occur-
rence or a case of regular periodic use by otters of this
area seldom occupied by humans. Leffingwell (1919)
noted that larger rivers in the region supported popu-
lations of fish year-round. Relatively small catches of
juvenile amphidromous fishes during the summers of
1999 and 2001 in Lions Bay indicated that these fish,
and likely some adults of the same species, did over-
winter in the area (Griffiths et al. 2002"). Craig and
McCart (1974°) identified groundwater springs in the
nearby Kavik River and Canning River drainage, as
well as a field of aufeis in the Canning River Delta, that
supported overwintering fish. Some of these areas
could potentially provide suitable habitat for an over-
wintering otter on Alaska’s North Slope.
Acknowledgments
This capture occurred during the daily operation of
a remote coastal fish study operated by LGL Alaska
Research Associates, Inc. The study was funded by
the Exxon Owners Group of Point Thomson, admin-
istered and reviewed by the Environmental Studies
Group of BP Exploration (Alaska) Inc.
Documents Cited (marked ° in text)
Craig, P. C., and P. McCart. 1974. Fall spawning and over-
wintering areas of fish populations along routes of the pro-
posed pipeline between Prudhoe Bay and the Mackenzie
Delta. Chapter 3 in Fisheries research associated with pro-
posed gas pipeline routes in Alaska, Yukon, and Northwest
236
Territories. Edited by P. J. McCart. Arctic Gas Biological
Report Series 15(3). 36 pages.
Griffiths, W. B., L. R. Martin, S. P. Haskell, W. J. Wilson,
and R. G. Fechhelm. 2002. Nearshore Beaufort Sea fish
studies in the Point Thomson area, 2001. Report for BP
Exploration (Alaska) Inc. by LGL Alaska Research Asso-
ciates, Inc., Anchorage, Alaska. 55 pages + appendices.
Literature Cited
Bee, J. W., and E. R. Hall. 1956. Mammals of northern Alas-
ka on the Arctic Slope. University of Kansas, Lawrence,
Kansas.
Forsyth, A. 1999. Mammals of North America: temperate and
arctic regions. Firefly Books Ltd., Willowdale, Ontario.
Leffingwell, E. K. 1919. The Canning River region, northern
Alaska. Government Printing Office, Washington D.C.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Manville, R. H., and S. P. Young. 1965. Distribution of
Alaskan mammals. Bureau of Sport Fisheries and Wildlife.
Circular 211. U.S. Government Printing Office, Washing-
ton, D.C.
Rue, L. L., IM. 1981. Furbearing animals of North America.
Crown Publishers, Inc., New York.
Whitaker, J. O., Jr. 1996. National Audobon Society field
guide to North American mammals. Alfred A. Knopf, Inc.,
New York.
Wilson, D. E., and S. Ruff. 1999. The Smithsonian book of
North American mammals. UBC Press, Vancouver/Toronto.
Received 4 August 2002
Accepted 24 December 2004
Osprey, Pandion haliaetus, Depredates Common Eider, Somateria
mollissima, Duckling
BRENDA M. BLINN!, VICKY VIOLETTE!~, and ANTONY W. DIAMOND!
' Atlantic Cooperative Wildlife Ecology Research Network, University of New Brunswick, P.O. Box 45111, Fredericton,
New Brunswick E3B 6E1 Canada
> Present address: Department of Fish and Game, Bay-Delta Region, P.O. Box 47, Yountville, California 94599 USA; e-
mail: bblinn@dfg.ca.gov
3 Mailing address: 80, Notre-Dame Nord, Louiseville, Québec JSV 1X5 Canada
Blinn, Brenda M., Vicky Violette, and Antony W. Diamond. 2006. Osprey, Pandion haliaetus, depredates Common Eider,
Somateria mollissima, duckling. Canadian Field-Naturalist 120(2): 236-237.
To our knowledge, we present the first published account of a successful attack on a Common Eider (Somateria mollissima)
duckling by an Osprey. Ospreys were sighted periodically during 940 hours of Common Eider observations in June-August,
2001 on Grand Manan Island, New Brunswick. One duckling (<1 week old) was observed depredated by an Osprey. Vulner-
ability of ducklings to predation may promote such opportunistic behaviour. The rarity of observations of such incidents
suggests depredation on ducklings by Osprey is uncommon.
Key Words: Common Eider, Somateria mollissima, Osprey, Pandion haliaetus, predation, Grand Manan Island, New Brunswick.
The diet of Ospreys (Pandion haliaetus) consists al-
most exclusively of live fish (Poole et al. 2002). Coastal
populations may switch prey species depending on the
pattern and timing of fish migration (Greene et al. 1983).
Ospreys are known to feed occasionally on rodents,
birds, reptiles, amphibians, and invertebrates (see Wiley
and Lohrer 1973 for literature review). Birds reported
include grebes, gulls, ducks, including a 2-week old
Wood Duck (Aix sponsa; Wiley and Lohrer 1973),
and passerines (Chubbs and Trimper 1998).
From 7 June to 3 August 2001, a study of Common
Eiders (Somateria mollissima) was conducted on Grand
Manan Island located 35 km from the New Brunswick
south shore. It is the largest (25 km long, 11 km wide)
of 17 other islands in the archipelago. A total of 940
hours of observations were made at eight 1 km‘? sites,
typically between 06:00 and 21:00 h. Duckling num-
bers and ages were recorded at 20-minute intervals
using scan sampling (Altman 1974). Predatory attacks
were recorded as they occurred. Eider broods occupied
the intertidal zone which was covered with brown
fucoid algae (Ascophyllum nodosum and Fucus vesi-
culosus) with some rock, sand or mud. A mixed forest
of Balsam Fir (Abies balsama), spruce (Picea sp.) and
alder (Alnus sp.), as well as shrubs (Rubus sp.) and var-
ious grasses (Graminae) were usually present within
10-50 m of the high tideline. A vertical tidal range of
up to 8.4 m can be expected in this area. Approxi-
mately 3370 Common Eider pairs (Ronconi and Wong
2002) and 15 other Anatidae species breed in the archi-
pelago (Dalzell 1991).
On 4 July 2001, an Osprey of unknown breeding sta-
tus captured and carried off a 0-7 day-old eider duck-
ling approximately 100 m offshore. The researcher (V.
V.) was situated near the waterfowl sanctuary of Cas-
talia Marsh (44°42'N, 66°48'W). Just prior to the attack,
an eider créche of 25 adult females and 62 ducklings
was observed in the area dabbling for invertebrates.
Duckling ages and counts (number) in this group were
0-7 days old (18), 8-13 days old (38), and 14-21 days
old (6). At 20:00, the créche stopped foraging and
became alert when an Osprey flew nearby before leav-
2006
ing the site. It returned at 20:03, swooped down, took
an eider duckling without any defensive behaviour by
the hens, and flew out of sight. The presence of the
duckling in the raptor’s talons was verified using a
Bushnell Spacemaster 45-60 x 200 spotting scope.
Ospreys were not observed at Castalia for the remainder
of the evening. Ospreys were recorded at Castalia and
at five other sites during visits both before and after the
attack. Only lone Ospreys were seen at any given time,
and no nests were present within any of the study sites.
Bald Eagles (Haliaeetus leucocephalus) also took three
ducklings at Castalia and two ducklings at another site
approximately 10 km away.
One of the reasons suggested to explain Ospreys tak-
ing non-fish prey is reduced visibility caused by turbid
water (Chubbs and Trimper 1998). Machmer and Yden-
berg (1990) found that capture rate by Ospreys de-
creased greatly when winds were above 25.2 km/h. On
the evening of the attack, moderate winds (~13 km/h),
medium-sized waves and clear skies probably did not
contribute to poor fishing conditions, although incom-
ing tide and floating algae could have made fish less
detectable.
Ospreys may take easily-captured (usually weak or
captive) birds when available (Dement’ev and Glad-
kov 1951). Eider ducklings <3 weeks old have poorly
developed diving skills (Hamilton 2001). This makes
them vulnerable to predation, especially in the first
10 days of life, with reported mortalities exceeding
90% from gull attacks (Mawhinney and Diamond
1999). Because Ospreys cannot dive deeper than one
meter into water (Prevost 1982), they forage in shallow-
water environments (Poole et al. 2002). Shallow-water
areas are often near brood-rearing habitat of eiders
(McAloney 1973).
In spite of many hours of observation of eider créch-
es in waters surrounding Grand Manan Island, we ob-
served only one instance of duckling depredation by
an Osprey. Ducklings less than one week old, which
are vulnerable to predation, were observed during five
out of eight observation periods (63%) and during 32 out
of 309 scans (10%) at Castalia. Therefore, it seems un-
likely that this Osprey was a “specialist”; i.e., fed more
frequently on ducklings than the average rate for the
species. Predation on eider ducklings by Ospreys is
evidently a rare event everywhere.
Acknowledgments
Financial, logistical and collaborative support that
made this observation possible was provided by the
Atlantic Cooperative Wildlife Ecology Research Net-
NOTES
237
work (ACWERN) at the University of New Brunswick,
the New Brunswick Wildlife Trust Fund, Acadian Sea-
plants Ltd. and the New Brunswick Department of
Natural Resources and Energy. Our sincere gratitude
goes to Greg McGuire for his assistance in the field.
André Breton provided valuable comments on earlier
versions of the manuscript. This is ACWERN Publi-
cation Number UNB-59.
Literature Cited
Altmann, J. 1974. Observational study of behavior: sampling
methods. Behaviour 49: 227-267.
Chubbs, T. E., and P. G. Trimper. 1998. The diet of nesting
Ospreys, Pandion haliaetus, in Labrador. Canadian Field-
Naturalist 112: 502-505.
Dalzell, B. 1991. Grand Manan Birds..A checklist with
occurrence graphs and a site guide. Third Edition. Brian
Dalzell, Grand Manon, New Brunswick.
Dement’evy, G. P., and N. A. Gladkovy. Editors. 1951. Birds of
the Soviet Union. Volume |. Sovietskaya Nauka, Moscow.
Greene, E. P., A. E. Greene, and B. Freedman. 1983. For-
aging behaviour and prey selection by Ospreys in coastal
habitats in Nova Scotia, Canada. Pages 257-267 in Biology
and management of Bald Eagles and Ospreys. Edited by
D.M. Bird. Harpell Press, Ste. Anne de Bellevue, Quebec.
Hamilton, D. J. 2001. Feeding behaviour of Common Eider
ducklings in relation to availability of rockweed habitat
and ducklings age. Waterbirds 24(2): 233-241.
Machmer, M. M., and R. C. Ydenberg. 1990. Weather and
Osprey foraging energetics. Canadian Journal of Zoology
68: 40-43.
Mawhinney, K., and A. W. Diamond. 1999. Using radio-
transmitters to improve estimates of gull predation on Com-
mon Eider ducklings. Condor 101: 824-831.
McAloney, R. K. 1973. Brood ecology of the Common Eider
(Somateria mollissima dresseri) in the Liscombe area of
Nova Scotia. M.Sc. thesis, Acadia University, Wolfville,
Nova Scotia.
Poole, A. F., R. O. Bierregaard, and M.S. Martell. 2002.
Osprey (Pandion haliaetus). In The Birds of North Amer-
ica, Number 683. Edited by A. Poole and F. Gill. The
Birds of North America, Inc., Philadelphia, Pennsylvania.
Prevost, Y. A. 1982. The wintering ecology of Ospreys in
Senegambia. Unpublished Ph.D. dissertation, University
of Edinburgh, U.K.
Ronconi, R. A., and S. N. P. Wong. 2002. Seabird colonies of
the Grand Manan Archipelago: 2001 census results and
guidelines for surveys and future monitoring. Grand Manan
Whale and Seabird Research Station. Bulletin Number 4.
Wiley, J. W., and F. E. Lohrer. 1973. Additional records of
non-fish prey taken by Ospreys. Wilson Bulletin 85: 468-
470.
Received 14 October 2005
Accepted 23 April 2007
238
THE CANADIAN FIELD-NATURALIST
Vol. 120
Extension de |’aire de distribution de 1 Umbre de vase, Umbra limi,
dans le nord-est du Québec
JEAN-FRANCOIS DESROCHES!
'Cégep de Sherbrooke, Département des Techniques de bioécologie, 475 rue du Cégep, Sherbrooke, Québec JIE 4K1 Canada;
e-mail: Jean-Francois.Desroches @cegepsherbrooke.qc.ca
Desroches, Jean-Frangois. 2006. Extension de |’aire de distribution de ’Umbre de vase, Umbra limi, dans le nord-est du Québec.
Canadian Field-Naturalist 120(2): 238-239.
Le 26 juillet 2005, plusieurs Umbres de vase (Umbra limi) ont été capturés au secteur Les Buissons de Pointe-aux-Outardes,
sur la Céte-Nord au Québec. Ces mentions élargissent l’aire de distribution de ce poisson de 300 km vers le nord-est.
On 26 July 2005, we caught many Central Mudminnows (Umbra limi) at the Les Buissons sector of Pointe-aux-Outardes,
on the Céte-Nord region of Québec. These records extend the known distribution of this fish 300 km to the northeast.
Mots-Clés: Umbre de vase, Mudminnow, Umbra limi, distribution, aire de distribution, Québec
L’Umbre de vase (Umbra limi) se retrouve dans le
centre de I’ Amérique du Nord, du centre-sud des Etats-
Unis jusqu’au sud du Canada, vers le nord (Page et
Burr 1991). Au Canada, il est réparti d’ouest en est
depuis |’ Alberta jusqu’au Québec (Scott et Crossman
1974). Au Québec, l’espéce est confinée au sud-ouest
de la province, se rendant a l’est jusque dans la région
de la ville de Québec selon les données actuellement
disponibles (Scott et Crossman 1974; Bergeron et
Brousseau 1981; Bernatchez et Giroux 2000).
En juillet 2005, dans le but de capturer des amphi-
biens aquatiques, deux nasses a ménés ont été placées
dans un étang a castors situé dans le secteur Les Buis-
sons de Pointe-aux-Outardes (49°7'47"N; 68°21'36"0).
Les nasses ont été relevées le 26 juillet 2005. Dans la
premiere nasse, il n’y avait aucune capture, mais quel-
ques Umbres de vase ont été capturés en fouillant dans
l'eau avec une épuisette. Neuf spécimens, soit quatre
jeunes (longueur totale 27,0-30,7 mm) et cing adultes
(longueur totale 54,5-62,7 mm) ont été gardés et sont
conservés dans la collection de poissons du Musée
canadien de la nature (CMNFI 2005-0032). Dans la
seconde nasse se trouvaient une jeune Anguille d’ Amé-
rique (Anguilla rostrata) vivante et les restes de deux
Umbres de vase, probablement dévorés partiellement
ou régurgités par l’anguille. Des fouilles effectuées
avec une épuisette ailleurs dans |’étang, la méme
journée, ont permis la capture de quelques autres
Umbres de vase et de tétards de grenouilles.
Cet étang a castors est relié a un lac plus gros, le lac
Lagacé, qui a une superficie d’environ 6 ha (850 m x
85 m). Le contour de l’étang est irrégulier, l’eau est
teintée de brun, et on y retrouve quelques arbres morts.
Il présente certains méandres vaseux ot |’on retrouve
un peu de végétation aquatique submergée. La forét
bordant l’étang est mixte et composée notamment
d’épinettes (Picea sp.), de Sapin baumier (Abies bal-
samea) et de Bouleau blanc (Betula papyrifera). Des
aulnes (Alnus sp.) bordent I’étang.
L’Umbre de vase est une espéce tolérante reconnue
pour vivre dans des eaux chaudes et riches en végéta-
tion, généralement peu oxygénées (Scott et Crossman
1974). Dans le sud-ouest du Québec, on le trouve
dans les baies riches en végétation des lacs et riviéres
lentes, dans les étangs 4 castors et les étangs de tour-
biéres, et dans les fossés (observation personnelle). Les
plans d’eau retrouvés sur la Cote-Nord sont en général
des lacs clairs typiques a l1’Omble de fontaine (Salve-
linus fontinalis), habitat peu propice a l’?Umbre de
vase. On y retrouve également des étangs a castors et
des tourbiéres, lesquels sont plus favorables a l’espéce.
L’Umbre de vase pourrait donc se retrouver ailleurs
sur la Céte-Nord.
La découverte de |’;Umbre de vase sur la C6te-
Nord au Québec, a 300 km au nord-est de sa distribu-
tion actuellement connue, revét un intérét particulier.
Il apparait probable que ce poisson soit plus répandu
qu’on le croyait au Québec et qu’il ait été sous-inven-
torié. Il s’agit d’une petite espéce ne présentant aucun
intérét pour la péche sportive ou commerciale et vivant
le plus souvent dans des milieux peu propices aux
poissons convoités pour la péche. Ces facteurs con-
tribuent sans doute a la méconnaissance de sa distribu-
tion dans le nord du Québec, et possiblement sur la rive
sud du Saint-Laurent vers l’est. Il est également pos-
sible que cette population d’Umbres de vase soit le
fruit d’une introduction volontaire ou accidentelle
par des pécheurs les ayant prélevés plus au sud, dans
le but de s’en servir comme appats de péche. Des
inventaires mieux ciblés permettraient de préciser la
distribution de cette espéce au Québec.
Remerciements
L’auteur remercie Julie et Félix Soucy pour leur
aide sur le terrain.
Littérature citée
Bergeron, J. F., et J. Brousseau. 1981. Guide des poissons
d’eau douce du Québec. Ministére du Loisir, de la Chas-
se et de la Péche, Québec. 217 pages.
Bernatchez, L., et M. Giroux. 2000. Les poissons d’eau
douce du Québec et leur répartition dans |’est du Canada.
Broquet inc., Boucherville, Québec. 350 pages.
2006 NOTES 239
Page, L. M., et B. M. Burr. 1991. A field guide to freshwa- — Scott, W. B., et E. J. Crossman. 1974. Poissons d'eau douce
ter fishes of North America, North of Mexico. Houghton du Canada. Ministére de l'Environnement, Service des
Mifflin Company, Boston et New York. 432 pages. péches et des sciences de la mer, Ottawa. 1026 pages
Regu 24 aoit 2005
Acceptée 19 février 2007
The Mink Frog, Rana septentrionalis, in Southeastern Labrador
JEAN-FRANCOIS DESROCHES!, ISABELLE PICARD, and JOHN E. MAUNDER?
' Cégep de Sherbrooke, Département des Techniques de bioécologie, 475 du Cégep, Sherbrooke, Québec JIE 4K 1 Canada;
e-mail: jean-francois.desroches@ cegepsherbrooke.qc.ca
* P.O. Box 250, Pouch Cove, Newfoundland and Labrador AOA 3L0 Canada
Desroches, Jean-Francois, Isabelle Picard, and John E. Maunder. 2006. The Mink Frog, Rana septentrionalis, in Southeastern
Labrador. Canadian Field-Naturalist 120(2): 239-240.
Mink Frogs (Rana septentrionalis) were collected at three new localities in southeastern Labrador in 2003 and 2005, extend-
ing the known range of the species about 200 km to the east, to the limits of the coast.
La Grenouille du Nord (Rana septentrionalis) a été trouvée dans trois nouvelles localités dans le sud-est du Labrador en 2003
et 2005. Ces mentions élargissent |’aire de répartition connue de |’espéce d’environ 200 km vers lest, aux limites de la c6te
de l’Océan Atlantique.
Key Words: Mink Frog, Rana septentrionalis, range, Labrador.
The Mink Frog (Rana septentrionalis) has the most
northerly southern limit of any North American anuran
(Hedeen 1986). This southern limit ranges from south-
ern Manitoba and northern Minnesota, south and east
through Wisconsin, northern Michigan and southern
Ontario, then northeast through northern New York,
northern New England, New Brunswick, and Nova
Scotia (Conant and Collins 1998; Cook 1984; Hedeen
1986). The species has recently been introduced on the
island of Newfoundland (Warkentin et al. 2003). Its
northern limit remains poorly known, largely because
the species is particularly aquatic (Hedeen 1986) and its
call and appearance are unfamiliar to many observers.
In Labrador, the Mink Frog has been documented from
several scattered localities, though not from the extreme
north or southeast (Maunder 1983, 1997; Cook 1984;
Desroches and Rodrigue 2004).
In 2003, during a primarily botanical and malaco-
logical field trip for the Provincial Museum of New-
foundland and Labrador (JEM), and in 2005, during
a vacation trip (J-FD and IP), we surveyed ponds and
brooks along all of the main roads and smaller town
access roads of southeastern Labrador, as far to the
northeast as the town of Cartwright. FicurE |. The Mink Frog in Labrador. Previous records: dots
On 3 August 2003 a single adult interface round (Maunder 1983) and squares (Maunder 1997). Question
=) o°
mark: Questionable record (see Maunder 1983: 42, 44).
Gray zone: proposed distribution based on previous
records (Bleakney 1958; Cook 1984; Desroches and
freshly dead from unknown causes, was collected
(NFM HE-250) from the shoreline of a medium-sized
pond, located beside the access highway running be- Rodrigue 2004). Stars: New records in southeastern
tween the Trans-Labrador Highway and the town of Labrador (1: southwest of Paradise River, 2: Charlot-
Cartwright, about 39.5 km southwest of the road tetown, 3: St. Lewis, 4: east of Churchill Falls). Bro-
junction to the town of Paradise River (53°09'52"N; ken line: Labrador/Québec border.
57°31'S6"W). This locality was, at the time, about
100 km east of the known range of the species (num-
ber | in Figure 1). The water of the pond was clear.
240,
The dominant emergent plants were Yellow Pondlily
(Nuphar variegata), Narrowleaf Bur-reed (Spargani-
um angustifolium), and Water Lobelia (Lobelia dort-
manna). The shoreline of the pond was mainly gravel-
ly, over a boulder substrate, and was well-vegetated.
The surrounding habitat was Kalmia scrub heath, with
patches and strips of low Black Spruce (Picea mari-
ana) woods in more sheltered areas.
In 2005, Mink Frogs were observed at two addition-
al localities in southeastern Labrador, about 200 km
east of the former known range of the species (num-
bers 2 and 3 in Figure 1). On 11 July, a subadult Mink
Frog was collected (CMN-35805) from a small pond
located beside a gravel road in the town of Charlotte-
town (52°46'09"N; 56°06'59"W). The pond was about
60 x 15 m in size, and had clear water, a muddy bot-
tom, and grassy and rocky shores. Trees at the site were
mostly Black Spruce. On 18 July, two Mink Frog tad-
poles were collected (CMN-35810) by dipnetting at
Blue Hill Pond, St. Lewis (52°24'55"N; 55°55'45"W).
The pond had clear water, a sandy and rocky bottom,
and some emergent plants near the shore. It was bor-
dered by Sweet Gale (Myrica gale) and alder (Alnus
sp.), and the adjacent forest was dominated by Black
Spruce. Two Threespine Sticklebacks (Gasterosteus
aculeatus) were caught in the same habitat, but re-
leased.
In 2004, an additional observation of the Mink Frog
was recorded at St. Lewis by Christian Fortin (Fortin
2005) but no voucher specimen was collected (C. Fortin,
personal communication).
In 2005, Mink Frogs were also observed in west-
ern Labrador, near the site of an old record, along the
Trans-Labrador Highway, east of the town of Churchill
Falls (53°29'29"N; 63°40'06"W) (number 4 in Figure
1). Two adults and one subadult were collected (CMN-
35798). Although this observation is not a range exten-
sion it is included here because it is a new collection
record.
During our searches for amphibians in southeastern
Labrador, the only other species observed was Amer-
ican Toad (Bufo americanus). Tadpoles of this species
were seen and/or collected at five locations along the
roads.
THE CANADIAN FIELD-NATURALIST
Vol. 120
The new Mink Frog collections recorded in this
paper show that the species is more widespread in
southeastern Labrador than has been previously doc-
umented. Further Labrador surveys should emphasize
areas north of the presently known distribution of the
species.
Acknowledgments
The authors thank Frederick W. Schueler for com-
ments on the manuscript before submission, and
Francis R. Cook for useful comments on the submit-
ted manuscript.
Literature Cited
Bleakney, J.S. 1958. A zoogeographical study of the amphib-
ians and reptiles of eastern Canada. National Museum of
Canada Bulletin 155. 119 pages.
Conant, R., and J. T. Collins. 1998. A field guide to the rep-
tiles and amphibians, Eastern and Central North America.
Revised third edition. The Peterson Field Guide Series,
Houghton Mifflin Company. Boston and New York. 616
pages.
Cook, F. R. 1984. Introduction to amphibians and reptiles of
Canada. National Museum of Natural Sciences, and Nation-
al Museums of Canada, Ottawa. 211 pages.
Desroches, J.-F., and D. Rodrigue. 2004. Amphibiens et
reptiles du Québec et des Maritimes. Editions Michel
Quintin, Waterloo. Québec. 288 pages.
Fortin, C. 2005. Sortie herpétologique sur la Cote-Nord et au
Labrador: de Natashquan a St. Lewis. Le Naturaliste can-
adien 129: 50-56.
Hedeen, S. E. 1986. The southern geographic limit of the
Mink Frog, Rana septentrionalis. Copeia 1986: 239-244.
Maunder, J. E. 1983. Amphibians of the province of New-
foundland. Canadian Field-Naturalist 97: 33-46.
Maunder, J. E. 1997. Amphibians of Newfoundland and
Labrador: status changes since 1983. Herpetological Con-
servation 1: 93-99.
Warkentin, I. G., C. E. Campbell, K. G. Powell, and T. D.
Leonard. 2003. First record of Mink Frog, Rana septen-
trionalis, from insular Newfoundland. Canadian Field-
Naturalist 117: 477-478.
Received 15 August 2005
Accepted 15 March 2007
2006
NOTES 24
First Record of the Great Barracuda, Sphyraena barracuda
from Canada
DARLENE BALKWILL!, BRIAN W. Coap!, ISMAEL GALVEZ2, and JOHN GILHEN
‘Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada
21219 Cromwell Road, Halifax, Nova Scotia B3H 4L1 Canada
4Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada
Balkwill, Darlene, Brian W. Coad, Ismael Galvez, and John Gilhen. 2006. First record of the Great Barracuda
Spn Vraena
i
barracuda, from Canada. Canadian Field-Naturalist 120(2); 241-242.
We report the collection, and provide a description, of the first Great Barracuda, Sphyraena barracuda, in Canadian water
based on a partial skeleton from Country Island, Nova Scotia.
Key Words: Great Barracuda, Sphyraena barracuda, new
record, fish fauna, Nova Scotia, Canada.
The cranium, lower jaw and attached trunk verte-
brae of a Great Barracuda, Sphyraeana barracuda
(Family Sphyraenidae), were found dried on a beach
on Country Island, Guysborough County, Nova Scotia
at 45°06'00"N, 61°32'37"W in May 1999 by Ismael
Galvez. This is the first record for this species from
Canada (Scott and Scott 1988; Coad et al. 1995). The
specimen is catalogued in the Nova Scotia Museum of
Natural History, Halifax, under NSM 85385.
Description
Three species of barracuda are reported from the
western Atlantic Ocean north of Florida, Sphyraena
barracuda (Walbaum, 1792), S. borealis DeKay, 1842,
and §. guachancho Cuvier in Cuvier and Valenciennes,
1829. Only S. borealis has been recorded from Cana-
da, a single specimen from Halifax Harbour, Nova
Scotia, caught on 5 October 1928 (Scott and Scott
1988). Carpenter (2002) records S. barracuda and §
guachancho from Massachusetts (rare) southwards.
Our specimen is neither S. borealis nor S. guachan-
cho on the basis of length, these two species general-
ly reach maximums of 46-50 cm, (commonly) 35 cm
and 50-60 cm, respectively (De Sylva 1984; Robins
and Ray 1986; Carpenter 2002). Murdy et al. (1997)
give | m for the latter species. Our specimen is close
in size to a disarticulated specimen of S. barracuda
in the Royal Ontario Museum (ROM R6281; pur-
FicureE |. Photograph of skeletal remains of a Great Barracuda from Country Island, Guysborough County, Nova Scotia
(NSM 85385)(bottom) with skull of ROM R6281 above.
242
THE CANADIAN FIELD-NATURALIST
Vol. 120
1 2 3
CENTIMETRES
FIGURE 2. Photograph of skeletal remains of a Great Barracuda from Country Island, Guysborough County, Nova Scotia
(NSM 85385) flanked by the upper and lower jaws of ROM R6281.
chased from Kensington Fish Market, Toronto, and
thought to be from the Caribbean area) that has a re-
corded total length of 86.0 cm. The length of the dor-
sal skull surface (from the anterior tip excluding the
upper jaw to the origin of the supraoccipital crest) is
135.1 mm in NSM 85385 and 116.8 mm in ROM
R6281. These measurements make the Nova Scotian
specimen ca. 99.5 cm total length. NSM 85385 has a
head length of ca. 181.7 mm (the opercular bone is
somewhat curved) and FishBase (www.fishbase.org)
states that head length is 17.7% total length, giving a
total length for the Nova Scotian specimen of 1.03 m.
These two forms of measurement agree, making the
Nova Scotian specimen about 1 m long in life. This
unequivocally eliminates S. borealis and probably S.
guachancho.
Other, osteological, characters also strongly sug-
gest the Nova Scotian specimen is a Great Barracuda.
The specimen has the area between eyes flat (convex in
S. guachancho), teeth are contiguous, vertically flat-
tened and erect (conical and widely spaced in S. bore-
alis, angled backwards in S. guachancho), and the tip
of the maxilla reaches past the anterior orbit margin
(not reaching the orbit in S. borealis) (De Sylva 1984;
Murdy et al. 1997; Carpenter 2002; comparative mate-
rial of S. guachancho ROM R1837 and R2706).
We consider it unlikely that this specimen came
from a local fish market and was imported. One of us
(JG) has not seen this species on sale locally. The shore
of Country Island is remote from commercial sites. It
probably represents a stray that died in Canadian
waters and was washed ashore.
Acknowledgments
We are indebted to Kevin Seymour, Department of
Palaeobiology, Royal Ontario Museum, Toronto, for
the loan of comparative material. Photographs were
taken by Alison Murray, Canadian Museum of Nature,
Ottawa, who also provided osteological advice.
Literature Cited
Carpenter, K. E. Editor. 2002. The Living Marine Resources
of the Western Central Atlantic. Volume 3. Bony fishes
part 2 (Opistognathidae to Molidae), sea turtles and marine
mammals. Food and Agriculture Organization, Rome. vi +
1375-2127.
Coad, B. W., with H. Waszczuk, and I. Labignan. 1995.
Encyclopedia of Canadian Fishes. Canadian Museum of
Nature, Ottawa, and Canadian Sportfishing Productions,
Waterdown, Ontario. viii + 928 pages, 128 colour plates.
De Sylva, D. P. 1984. Sphyraenoidei: development and rela-
tionships. Pages 534-540 in Ontogeny and Systematics of
Fishes. Special Publication Number 1, American Society
of Ichthyologists and Herpetologists, Lawrence, Kansas.
ix + 760 pages.
Murdy, E. O., R. S. Birdsong, and J. A. Musick. 1997.
Fishes of Chesapeake Bay. Smithsonian Institution Press,
Washington. xi + 324 pages.
Robins, C. R., and G. C. Ray. 1986. A Field Guide to the
Atlantic Coast Fishes of North America. Houghton Mif-
flin Co., Boston. xi + 354 pages.
Scott, W. B., and M. G. Scott. 1988. Atlantic Fishes of
Canada. Canadian Bulletin of Fisheries and Aquatic Sci-
ences 219: xxx + 731 pages.
Received 18 July 2005
Accepted 16 January 2007
A
oncmoms
See
I
Book Reviews
ZOOLOGY
Animal Skulls — A Guide to North American Species
By Mark Elbroch. 2006. Stackpole Books, 5067 Ritter Rd.,
Mechanicsburg, PA 17055. 727 pages, $34.65 Paper.
As an instructor of comparative vertebrate anatomy,
and a self-proclaimed enthusiast of skulls, | very much
looked forward to reading and reviewing this book. I
learned a fair amount from this book, mostly regard-
ing animals that don’t live in my area, so overall, this
book was for me, a valuable read.
There is an enormous amount of material synthe-
sized in this book. There are the expected figures
illustrating the skull bones from a number of species;
neighbouring figures illustrate the measurements most
commonly used. However, the figure for reptiles has
several errors, and those for the birds and mammals
also have a few...not a good start for fundamental
material near the beginning of the book.
Skulls of each species are illustrated in three views
within the species accounts. Elsewhere, montages of
dorsal views (mammalian and herp crania) and lateral
views (mammalian mandibles, birds) are quite useful,
and where practical, are illustrated at life-size. Even
though the skulls in these montages show less detail
than in the species accounts, I’ve always found this
set-up useful (as in the Golden Field Guide to Birds,
where warblers and sparrows are illustrated in a simi-
lar manner).
There are a number of issues that I found annoying.
_ Why, for instance, is the Mantled Howler Monkey des-
cribed (and illustrated along with two other species
of non-human primate)? They are native to Central
and South America; Are they feral in North America?
We are never told. We are never told of the distributions
of any of the animals, a shortcoming if someone is try-
ing to identify a skull they’ve found based on this book.
Secondly, the author is somewhat free with his use of
the word bone. “Occipital condyles”, for instance, are
processes of bones, not bones unto themselves, though
that is how they’re treated. In a book that thankfully
retains appropriate terminology, this oversight will be
undetected by the novice, but irritating to the experi-
enced. I now need to shout...““Molars are not flat!” I
have to remind my students of this constantly. In Eng-
lish, we have the two words, “flat” and “broad”, and
they are not interchangeable. Molars adapted for grind-
ing vegetation are broad...they are anything but flat
as erroneously described numerous times in this book.
There are a few other annoying errors in this vein.
Given the number of prominent, pre-publication re-
viewers mentioned in the acknowledgements, one must
wonder about their diligence to this project.
The title is awkward, Animal Skulls (as opposed to
part two in the series, Plant Skulls?)...and mislead-
ing...a guide to North American Species: fish are not
covered at all, amphibians and reptiles are given a
token amount of space (12 species, fewer than 3% of
those in North America), birds (71 species, 10% of
breeders) are allotted only a fraction of the coverage
of mammals (143 species, one-third of known species),
this despite the fact that the number of species of birds
alone outnumbers those of mammals in North Ameri-
ca. I do understand that there is very little variation in
skulls amongst warblers or sparrows, for example, and
illustrating each species could quickly become re-
dundant. However, the mammalian bias is still over-
whelming.
The mammalian dominance in this book is also
reflected in the amount of text devoted to each species
account — one line for each herp, one paragraph for
each bird, four or more paragraphs for mammals.
About 100 pages of measurement data are provided for
mammals (which is wonderful), though none is pro-
vided for the others. One is forced to ask, “Why bother
with the birds and herps at all?” The same amount of
space could have been used to make a more complete
book focussing on mammal skulls alone. Then, subse-
quent volumes addressing other taxa could complete
the series.
I particularly liked the early chapters on interpret-
ing form, marks on skulls (from gnaw marks to builet
holes) and preparing skulls. These chapters were in-
formative and well-written. For me, a large part of the
worthiness of this book was in these early chapters.
That is not to diminish the worthiness of the mam-
malian species accounts, they too are thorough and
instructive.
RANDY LAUFF
Department of Biology, St. Francis Xavier University, Antigo-
nish, Nova Scotia Canada B2G 2W5
244
A Field Guide to North Atlantic Wildlife
By N. Proctor and P. Lynch. 2006. Yale University Press
P.O. Box 209040, New Haven, Connecticut 06520-9040
USA 221 pages. U.S. $19.95 Paper.
This book arrived on the morning I left for Svalbard
in the Norwegian arctic. While the area covered by this
book, offshore from North Carolina to Newfoundland,
is some distance from Norway, the two areas do share
a lot of wildlife. So I took the book along to make
direct field comparisons.
I like the author’s concept for this guide. He has
selected the most likely species you have some possi-
bility of seeing on a pelagic trip or ferry crossing in the
northwestern Atlantic. This includes the large “plank-
ton’ and algae, fish, turtles, whales, seals and seabirds.
Only the more common species are included, enabling
the authors to keep the book small [18 x 11 x 1.5 cm]
and portable. This meant I was able to identify Beroe’s
Comb Jelly, Fin Whale and Ivory Gull carrying only
one guide. The guide does not include coastal species
such as crabs, molluscs and inshore fish.
The descriptions are well written. For example, the
text for Fin Whale is a version of the classic descrip-
tions found in many guides. I compared this to a whale
that circled the ship and the information given was
accurate down to the details of the back chevrons.
I had far more problems with the illustrations. First
there is no attempt to show scale. A 10-inch flying fish
is shown as the same size as an 18-inch species. This
is a critical characteristic for look-alike species such as
Glaucus and Iceland gulls. You need to check the text
to see that there is a 5-inch difference in size. Similarly
the Northern Seal plate shows the tiny 4 foot Ringed
Seal as the same size as the 9 foot Hooded Seal. I found
this disconcerting.
Some of the species shown are really cold water
arctic animals. The North Atlantic is the southern fringe
of their range. This is not mentioned for animals like
Beluga, Dovekie or Harp Seal. I think giving their typ-
A Complete Guide to Arctic Wildlife
By Richard Sale. 2006. Firefly Books, 66 Leek Crescent,
Richmond Hill, Ontario L4B 1H1 Canada. 400 pages.
$49.95 Paper.
This is a comprehensive field guide to the birds
and mammals of the Arctic. Author Sale describes
the ecology and people of the region along with some
Arctic history. He cannot avoid discussing both cli-
mate change and the threat of pollution. Short chap-
ters on geology, geography and biogeography set the
background. These are followed by the main text: a
field-guide style section on the Arctic’s mammals and
birds. The information provided covers identification,
size, voice, distribution, diet, breeding and taxonomy.
The guide is illustrated with photographs supported
THE CANADIAN FIELD-NATURALIST
Vol. 120
ical range would add perspective and could be said in
few words.
The non-bird illustrations were very good. The artist
has captured the short face of the Ringed Seal and
the longer-nosed head of the Harp Seal very well. The
fish portraits from the warm-water Dolphin [the fish]
to the cold-water Cod are accurate.
The bird illustrations are another story. For example,
the illustration of the Arctic Tern looks almost identi-
cal to that of the Common Tern. The Arctic has a short-
er head and a longer tail giving it a different fore-and-
aft look from the Common. The wing pattern is also
quite different, having less black and giving the Arctic
a much paler appearance. I had similar issues with other
confusing pairs [Leach’s and Wilson’s storm-petrels,
Glaucous and Iceland gulls, Audubon’s and Manx
shearwaters, etc.] Many other species lacked the sub-
tle plumage differences that help confirm identification.
For example, the two-toned grey on the wing of a Kit-
tiwake and the armpit marks on the murres. Many birds
are shown in winter plumage only. This is a reason-
able approach as many birds disappear inland or north
to breed. However, the winter Atlantic Puffin is very
odd and the loon illustrations do not capture these birds
at all well. It appeared that the artist did not have field
experience with these species.
This is, however, a useful little book. If you are a
novice I would recommend you take this book plus a
modern bird guide on your pelagic trips. For more ex-
perienced bird watchers this book will suffice to remind
you of the key points for what you may see and will
be useful for all the non-bird species.
I thank Dr. Tom Smith, a fellow traveller, for sharing
his immense knowledge of arctic wildlife, and thereby
contributing to my review.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
by some field-guide style plates. Each species has a
range map showing their circumpolar ranges.
I am always delighted to find a book that shows birds
and mammals from a bio-regional perspective. Last
summer I saw eight Snow Geese on the west coast of
Svalbard — an unusual sighting. When you look at the
distribution of the birds it is not hard to see how their
expanding population could spill over the now unfrozen
ocean a short distance eastward. This kind of perspec-
tive is difficult to reach when you use guides based
on a political boundary (birds of Europe, Russia etc.)
The author’s first problem was the difficultly of de-
fining the boundary of the Arctic. He uses, in my opin-
ion, the most sane choice. He has taken the 10°C July
2006
isotherm, modified by extreme winter temperatures.
This still leaves out some areas that I think of as “Arc-
tic’ such as James Bay and its Polar Bears and the
northwest coast of Alaska with Belugas and Yellow-
billed Loons. This, perhaps, proves that nature will
never entirely fit into our human need for definitions.
Following his definition, the author then chose the
species to be included as arctic birds and mammals.
Here | am a little perplexed. He has, for example in-
cluded Carrion Crow but not Black-billed Magpie
(which I have seen at 63°N). He has only an introduc-
tory mention of the Gray Jay, which ranges to the shores
of the Arctic ocean. Tree Swallow is given an individual
account while the more northerly Cliff Swallow (I have
seen it at 73°N) is merely mentioned as a rarity. | am
also confused by the species he has decided to split and
those he leaves as sub-species. For, example he has
Iceland and Thayer’s gulls as separate species but not
Mew and Common gulls. There seems to be better evi-
dence to split the latter. Similarly you can question hav-
ing separate entries for Black and Common scoters.
These are minor points and not as significant to my
two major concerns. The species descriptions are de-
tailed and clear and are some of the best I have read.
While they are first rate, the author gives exceedingly
spartan information on confusing species. This is in
contrast to the descriptions, which often allude to an
important field mark. For example, he states the three
central tail feathers of the jaegers are “distinctive.” In
the field separating a Parasitic Jaeger with a 9 cm tail
extension from a Long-tailed with a 12 cm extension
is not that easy. I well remember a controversial bird
The Black Flies (Simuliidae) of North America
By P. Adler, D. Currie, and D. Monty Wood. 2004. Royal
Ontario Museum, 100 Queen’s Park, Toronto, Ontario
MSS 2C6 Canada, and Cornell University Press. 941
pages. $99.95 US.
This is a huge book. It is huge in size, filling 941
pages. It is huge in scope, treating the biology, man-
agement and identification of the entire black fly fauna
of Canada, the United States, and Greenland. And it
is huge intellectually, the culmination of many decades
of work by an enthusiastic international community
of researchers, among whom the authors of this book
are prominent members.
In the preface, Peter Adler states that the book is
“geared to a general readership.’ I would not have
guessed that this was true, at first glance, but after some
perusal it seemed to me that the book would be useful
from about the undergraduate level on up. The writing
is clear and precise, but the vocabulary will be daunting
for most people without biological training. I suspect
that the book will find a place in almost all scholarly
libraries, and that the specialists who will use it most
probably ordered a copy before it came into print. But
there is a wider audience for the book as well, and in
BooK REVIEWS
245
that needed the late great Earl Godfrey's talents to re-
solve as a Long-tailed Jaeger. Similarly the informa-
tion on the Common and Hoary redpolls is not enough
for field separation. There is no mention, for example,
of the stubby bill of the Hoary — very evident in the
nice accompanying photo.
My second point is that this is not a complete guide
to wildlife. Unlike such books as Les Beletsky and
Dennis Paulson’s Ecotraveller's Wildlife Guide to Alas-
ka it does not contain any information on plants, insects,
marine invertebrates and fish or amphibians. Admit-
tedly Beletsky only includes the commoner, more vis-
ible species although his title does say it is a complete
guide, but he does include a broad spectrum of the wild-
life, other than birds and mammals, visible to the naked
eye.
While this in not meant as a coffee table book the
photographs are really eye-catching. The bird plates
are quite good, but I was less taken with the mammal
plates. My chief reservation about all these illustrations
is they frequently show only adult males.
I will take this book with me on my upcoming trip
to the Pribiloffs. For the most part I will be seeing
wildlife that I am familiar with and I will not carry a
conventional field guide to the birds or mammals. This
book will provide me with more encyclopaedia-style
information about the creatures I am seeing. | will also
take Beletsky and Paulsen’s book and a flower guide.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
this regard I predict that it will be a required reference
for many decades to come. The authors mention its rel-
evance to medical and veterinary entomologists, aquat-
ic biologists, environmental consultants, systematists,
naturalists, pest management specialists and students,
but I suspect the real list will be longer than that.
The text is arranged in four parts, covering back-
ground information, biology of black flies, economic
concerns, and systematics and taxonomy. The first part
provides an overview of the subject, a very readable
history of the study of black flies (with lovely historical
photographs) and a chapter on techniques for collecting
and curating black fly specimens. As an entomologist
(with no particular interest in black flies, 1] admit) I
nonetheless found this section very interesting.
The second part of the book deals with the biology
of black flies, and consists of a fine treatment of the
structure of these insects (and their immature stages),
as well as a well-written overview of their cytology.
Black fly cytology was pioneered by the late Klaus
Rothfels, a man deeply missed by the authors, and who
they refer to as “friend, mentor, and phenomenon.” A
chapter summarizing the life history and behaviour
246
of black flies is also offered, along with an exhaustive
summary of blood feeding host records, and records
of black fly diseases and parasites.
For most readers, it is likely that the third part of
the book, which treats the management of black fly
pests, will be the most interesting. I can see this part
of the book providing fodder for any number of term
papers, research paper introductions, and consulting
and government reports. I was surprised, however, to
find that the chapter on management (Chapter 8) was
largely historical in nature, without a summary of con-
trol recommendations. On careful reading, however, I
could see the reason for this approach. Oil-based pes-
ticides, DDT, and other chemical controls for black
flies have not stood the test of time. The authors sum-
marize the use of the bacterial toxin Bi, but give it only
a qualified nod, with the caution that it too might lose
its effectiveness over time. The chapter ends in an inter-
esting discussion of repellants, and repellant clothing.
The fourth and final part of the book is clearly the
section closest to the authors’ own interests. It treats
the 254 species of “North American” black flies, thor-
oughly. The focus is not simply on identification, it is
also deeply phylogenetic, and includes a very careful
evolutionary justification for each and every level of
the classification. This is followed by a superb species
by species treatment of the entire fauna. Significantly,
there is additional information on the economic sig-
nificance (if any) of each and every species, adding
greatly to the summary in part three of the book.
As a non-specialist, I have to admit that the most
impressive aspects of the book to me were the illus-
THE CANADIAN FIELD-NATURALIST
Vol. 120
trations. They begin on page 436 (well before the
halfway mark) and they are so masterfully executed
that I couldn’t help but come away thinking that there
was something deeply beautiful and elegant about
black flies, their larvae and pupae, and the fine details
of their anatomy. More full-body illustrations of the
adults would have been nice, but the overall sameness
of the other life stages illustrated convinced me that
side-by-side comparisons of the adults might not be
all that useful. Maps follow the illustrations, and treat
the United States and Canada on a county level, but
they do not show Greenland. And, as one might expect
in a book of this nature, the reference section and the
indices are exhaustive.
This is a magnum opus, and a tremendous labour
of love. It ranks, in my opinion, right up there with
Holldobler and Wilson’s The Ants. Having said that,
it seems to me a shame that the book appears to have
been presented as a somewhat impenetrable scholar-
ly tome, without a showy slip cover, and with little or
no fanfare for the “general readership” to which it
was addressed. Don’t be scared off by the size, or the
complexity of this book. It is a masterwork, and if any
aspect of your interests overlaps with the subject of
black fly biology, do consider adding this fine vol-
ume to your library.
JOHN ACORN
University of Alberta, Edmonton, Alberta TST 5L7 Canada
Literature Cited:
Hollidobler, Bert, and E. O. Wilson. 1990. The ants. Belknap Press
(Harvard University Press), Cambridge, Massachusetts. 732 pages.
Insects: Their Natural History and Diversity: With a Photographic Guide to Insects of Eastern
North America
By Stephen A. Marshall. 2006. Firefly Books Ltd., 66 Leek
Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 720
pages. CDN $95. Hardcover
It has always been said that you can’t judge a book
by its cover, but after reading Insects: Their Natural
History and Diversity I’ve learned that this well-worn
idiom isn’t always true. When this book first crossed
my desk, to say that I was instantly enamoured would
be an under-statement. It was so beautiful, the cover
adorned with a stunning jewel-toned dogbane beetle
(Chrysochus auratus). 1 almost didn’t want to crack the
binding. However, my curiosity finally got the better
of me and I’m glad it did because once I started read-
ing I couldn’t put the book down.
Visually stunning, with over 4000 colour photo-
graphs of insects in their natural habitats, Insects: Their
Natural History and Diversity has the look and feel
of a glossy coffee table book while still being full of
accurate, well researched information.
As its title implies, Insects: Their Natural History
and Diversity focuses on the diversity and natural his-
tory of common families of northeastern North Ameri-
can insects. The book opens with a brief synopsis of
basic insect anatomy and morphology. This is followed
by chapters covering the diversity of all insect orders,
including all the major families, along with two chap-
ters on non-insect arthropods and methods for observ-
ing, collecting and photographing insects. The book’s
last 50 pages are dedicated to illustrated keys to order
and family as well as a key to the most commonly en-
countered insect larvae. These keys are designed to
facilitate ease of use and therefore emphasize morpho-
logical characters visible to the naked eye or easily
seen with a hand lens. Also peppered throughout the
book are helpful suggestions on where to look for
and. find various insect orders/families. For example,
“Depending on your inclination and the weather, a
good place to start looking for assassin bugs would be
in your kitchen light fixture. Unless you are much more
2006
fastidious than most, the odds are that among the crispy
critters accumulated there you will find a large black
assassin bug called the Masked Bed Bug Hunter (Redu-
vius personatus).”
Considering that Insects: Their Natural History and
Diversity is priced so as not to be cost prohibitive and
is clearly written in plain language, it will be highly
accessible to a broad audience, including naturalists,
amateur entomologists as well as seasoned profession-
als. The author has also included a dollop of humour
and wit throughout the text. For example, this passage
describing the appearance of springtails: “Some are
covered with scales, like those of a butterfly, many are
brilliantly colored and all are morphologically bizarre,
starting from the long, forked tail used to make Her-
culean leaps, and ending with the deeply pocketed
mouth that makes springtails look like they have lost
their dentures and then sucked on a bunch of lemons.”
Book REVIEWS
247
This book would make a great textbook for a natu-
ral history or general entomology course. Especially
when you consider that the impetus for this book cen-
tres on materials originally gathered in support of the
author’s third-year course “The Natural History of
Insects” at the University of Guelph. With its depth of
scope and true to life colour photographs /nsects: Their
Natural History and Diversity would be indispensable
in the field; however due to its size, it would be a bit
unruly to have to lug around.
I thoroughly enjoyed this book and would recom-
mend it whole-heartedly to anyone who has an interest
in entomology, natural history or a simple curiosity
about the six-legged world that surrounds us.
GINA PENNY
Department of Biology, St. Francis Xavier University,
Antigonish, Nova Scotia B2G 2W5 Canada
The Northern Goshawk: A Technical Assessment of its Status, Ecology and Management
By Micheal L. Morrison. 2006. Cooper Ornithological Soci-
ety, Pennsylvannia.
The Northern Goshawk: A Technical Assessment
of Its Status, Ecology and Management, is just as the
title suggests, a highly technical assessment of the
goshawk. I do not recommend it to those looking for
a light read. Micheal L. Morris, editor, has compiled
22 recent journal articles that study the Northern Gos-
hawk, focussing on ecology and population. Morris
outlines two objectives for the assessment; the first is
to summarize existing knowledge of the goshawk into
one book, and the other is try to provide a framework
for the development of future regional conservation
and management strategies for the species.
Due to the legal issues surrounding the Northern
Goshawk and its proposed listing as a Species at Risk,
particularly in the United States, this book is a useful
step for establishing what we do and what we do not
know about this Accipiter. Unfortunately, it appears that
the latter predominates.
Morris structures this assessment in an appropriate
way, Starting with an article that extensively covers the
ecology of the goshawk, which helps provide the read-
er with a general knowledge of the bird. After this first
article, the volume is divided into three distinct sec-
tions: Regional, Ecology, and Management, each of
which contains a number of independent studies that
Morris has gathered from several sources. Each of
these studies take a highly regionally specific look at
certain aspects of the Northern Goshawk. The goshawk
has an extensive range over North America, Europe
and Asia. As a result of this huge regional coverage
there is extensive continental/regional variation amongst
populations of these birds. This makes it difficult as a
reader to distil any major trends between the articles.
I found it hard to link many of the articles together
because of the different styles by which the studies
were carried out, as well as some differences of opin-
ion. For example, Squires and Kennedy suggest that
“(impacts from forest management] can either enhance
or degrade goshawk habitat depending on type and
extent of habitat alterations” (page 61). This is not so
much a weakness of the assessment but rather the real-
ity of goshawk management, and emphasizes the need
for regional management strategies instead of an all
encompassing global strategy, which is one of Morris’s
initial goals for the assessment.
Overall, this publication is a building block. It estab-
lishes where we have been in terms of goshawk man-
agement and conservation and makes clear where we
need to head in the future. Morris has done a good job
of selecting appropriate articles that clearly depict
the challenges faced by researchers and the Northern
Goshawk. I would recommend this book only to those
who want to critically study goshawk ecology.
JARED B. CLARKE
323 Habkirk Drive, Regina, Saskatchewan S4S 6A9 Canada
248
Turtles of the World
By Franck Bonin, Bernard Devaux, and Alain Dupré (Trans-
lated by Peter C.H. Pritchard). 2006. Johns Hopkins Uni-
versity Press. 416 pages. Illus., $50 (US).
First published in 1996 in French, this guide to the
turtles of the world has been translated into English by
eminent turtle biologist Peter Pritchard. In the trans-
lator’s note, Pritchard acknowledges that he has also
made some corrections and updates to the text. As such,
the book includes recently discovered species such as
Pelusios cupulatta, only described in 2003.
The book is divided into two sections: General
Biology and Identification. General Biology provides
a brief (nine pages) overview of the evolutionary his-
tory of turtles, their biology and conservation issues.
Identification provides information on the approxi-
mately 330 recognized species of turtles. The species
are grouped by families within the two suborders Pleu-
rodira and Cryptodira. In general, each species account
follows the same standardized format: distribution,
description, natural history, protection. The species
accounts vary from about half a page to three pages in
length.
Each species also has a small regional map showing
which part of the world the species occurs in and a
larger, close-up map showing the distribution as shad-
ed areas. These distribution maps also show major river
systems, as well as some political boundaries. In addi-
tion, almost every species has one or more colour pho-
tographs.
Any attempt to list the names of all the species in
the world of a certain group is a daunting task of sort-
ing out sometimes conflicting and constantly chang-
ing taxonomy. The authors have accepted the recent
splitting of the genus Clemmys and duly note this
change in the appropriate species accounts. In con-
trast, the entire genus Sternotherus has been subsumed
within Kinosternon without any explanation. While
there has been debate over the validity of Sternotherus
as a distinct genus, lumping the two genera goes against
the current standard North American taxonomy. The
authors are also somewhat inconsistent in assigning
species status. They do not provide species accounts
for Chelodina mccordi or C. pritchardi (although they
are mentioned under the species account for C. no-
vaeguineae), yet both these species are widely accept-
ed. On the other hand, they elevate Hardella indi, a
subspecies of H. thurjii, to full species status without
any justification. This may explain why Hardella indi
has the shortest species account in the book — three
sentences.
THE CANADIAN FIELD-NATURALIST
Vol. 120
There are some strange oversights in this book. The
IUCN Red List of Threatened Species is the standard
international ranking system for species at risk of be-
coming extinct. It designates status for these species
from Lower Risk up to Critically Endangered. Turtles
of the World occasionally mentions if a species has
been “red listed” but not consistently, even within the
same genus. For example, all four species within the
genus Acanthochelys are on the Red List, yet the spe-
cies accounts mention this for only one of the species.
The book says one of the other species is on the
“IUCN priority list” and the “IUCN is worried” about
another. None of the species accounts provides the
IUCN status, so the reader will not learn that A. pal-
lidipectoris is considered the most endangered of the
four species.
A thorough reference section is also lacking. The
references for this book do not even fill two pages and
almost all of the references are to other books on tur-
tles. Accordingly, most of the information in the book
is presented without any reference to a specific primary
source. However, occasionally a statement is referenced
and in almost every case that reference is absent from
the reference section.
Of the Canadian species, the map for Graptemys
geographica (Common Map Turtle) excludes the Ot-
tawa area and all of the Quebec part of the range, and
maps for both Glyptemys insculpta (Wood Turtle) and
Emydoidea blandingii (Blanding’s Turtle) omit Nova
Scotia. Terrapene carolina (Eastern Box Turtle), a
species never confirmed to have been historically pres-
ent in Canada at all, is shown as ranging across all of
southern Ontario as far north as Toronto. The range is
also over-extended in Michigan, Ohio, Pennsylvania
and New York.
Despite these criticisms this is a significant book in
turtle biology. The photographs are often excellent. The
natural history section, while sometimes too short, pro-
vides insight into an incredible variety of life history
strategies. For many species, the contents of the protec-
tion section provide important information on threats,
as well as a wide diversity of conservation measures.
This volume is not the definitive statement on any of
these 300+ species but it is a good introduction to the
diversity of turtles.
DAVID SEBURN
Seburn Ecological Services, 2710 Clarenda Street, Ottawa,
Ontario K2B 7S5 Canada
2006
Book REVIEWS
249
The Wolves of Algonquin Park: A 12-year Ecological Study
By John B. Theberge and Mary T. Theberge. Department of
Geography, University of Waterloo, Waterloo, Ontario
N2L 3G1 Canada. 163 pages. Available from bkevans@
fes.uwaterloo.ca. $23.50 Canadian, $20 U.S.
John and Mary Theberge, a husband and wife team,
have put together information available on wolves and
their prey in one of North America’s great ecosystems.
Research results from the twelve year period (1987-
1999) was used as a core for the publication. Student
projects were as follows: | Ph.D. thesis, 7 MSc. theses,
11 BSc. theses). Overall, scientific publication of 55
papers is impressive. In the end, it was the Theberge’s
that pulled the material together into this single mono-
graph. For that they need to be congratulated.
The results of the study are also based on previous
work begun in the 1960s. The Theberges reached back
in time and reanalyzed and re-evaluated some of the
data collected earlier. A major portion of the review
centred on the taxonomic study of Wolves in the park.
They concluded that these canids were larger than the
Coyote-Wolf hybrids to the southwest, smaller than
wolves to the northwest. It reinforced earlier conclu-
sions of the existence of “Algonquin type”, “tweed
type” and “boreal type” Wolves. New is the interpre-
tation that these Wolves are taxonomically connected
to the Red Wolf taxon, once more common on the con-
tinent to the southeast. In the “canid soup of genet-
ics” these are many theories and assertions. So far there
have been no clear indications of the “ultimate” posi-
tion. Maybe none will ever emerge that will satisfy
everyone who is interested in the subject. The The-
berges have presented a convincing case.
Other portions of the monograph are equally well
presented. There is much in the way of natural history
— basic insight into the ecology of the predator within
the prey dynamics dictated by geographical and botan-
ical (habitat) setting. In an age of computers, abstract
BOTANY
Flowers — How They Changed the World
By William C. Burger. 2006. Prometheus Books, 59 John
Glen Drive, Amherst, New York. 14228-2197 USA. 337
pages. US $23.00. Cloth.
Why are flowers important? They are not just for the
viewing pleasure of humans, although they are spec-
tacular — but why? This book will answer these ques-
tions. “Without the gift of flowers ... man might still
be a nocturnal insectivore gnawing a roach in the dark.”
It is often true that the more we know about some-
thing the more we appreciate it and the more we enjoy
it. A book that leads to enjoyment is worthwhile, but
this book also provides a much needed perspective
on the flowering plants at a time when the world faces
a biodiversity crisis. It contributes to an improved
analysis and spurious mathematical modelling, this
work stands out as a major contribution to science. It
combines solid field work with a realistic quantitative
assessment of the results, and | would recommend this
as a case study for first-year graduate students who are
contemplating a career in wildlife research. Beyond
that, the study has had a real impact on conservation.
In 2001, a total year-round Wolf killing ban in areas
bordering Algonquin Park was announced by provin-
cial authorities. Political responses were based on re-
search results. It is a rare occurrence that this happens
and it is a response to the well executed research. By
any standard, that is a remarkable outcome to many
years of dedicated work by scientists whose meticulous
research and commitment to science were also cou-
pled to a dedication to conservation and to bioethics.
This is a scholarly monograph that likely will not be
surpassed for many years to come.
At the outset the Theberges posed a series of ques-
tions that they wanted to find answers to. Likely some
were listed after the fact, when intensive research
results came in from the field work and data analysis.
That too has strength. I cannot see much that has been
left out. Appropriate photos and illustrations are found
throughout the text. Some questions arose in my mind
as to alternate explanations for some of the conclusions
reached. Such results are inevitable and provide a use-
ful basis for future scholars. This monograph is an
exceptional document and has set a very high standard.
I highly recommend it to anyone just starting field
research on mammals, or for those veterans, who have
spent much of their time in the pursuit of knowledge
in the field of wildlife science.
Lu CARBYN
137 Wolf Willow Crescent, Edmonton, Alberta TST 1T1
Canada
understanding of the natural world and a concern for
the future. It is a very valuable contribution as well as
a very enjoyable book that will appeal to anyone as a
result of the friendly writing style.
William Burger, curator emeritus in the Department
of Botany at Chicago’s Field Museum, takes us on a
journey to the discovery of how flowering plants have
transformed the world. He begins with “What exactly
is a flower?” which is a delightful and comprehensive
introduction to flower morphology or what makes a
flower a flower. Floral parts and their role are described
in different kinds of flowers. It comes as a surprise to
many people that the four bright white petals of a dog-
wood flower are not petals and the flower is actually an
250
aggregation of many flowers, that a dandelion flower
is made up of dozens of flowers, that grasses do have
very special flowers with floral parts analogous to
those of tulips.
The book is full of fascinating anecdotes. Why are
Wolves friends of plants? By 1926 Wolves had been
extirpated in Yellowstone and Yogi, Booboo, and the
other bears were of course more interested in picnic
baskets than controlling the Elk so there were soon too
many Elk and the vegetation began to change. Seven-
ty-five thousand Elk were removed between 1926 and
1968 but it was not enough to get things back to the
way they were — although it did maintain some plant
diversity and kept the park green! Finally, in 1995 a
human brain triumphed and thirty-one Wolves were
reintroduced to Yellowstone and that quickly got things
back to the way they were. There was less browsing
pressure as “fear factor” came into play with Elk avoid-
ing places where they were more likely to be ambushed
by Wolves. Browsing was not only reduced, it was spa-
tially variable, creating more niches for plants. The
entire Chapter 2 is a very entertaining voyage though
the annals of ecology indicating how plants are assist-
ed and influenced by other organisms.
Flowering plants have a vast array of enemies, the
herbivores, which are everything that eats plants, but
they have developed chemical and mechanical de-
fences. This is the subject of Chapter 3 “flowers and
their enemies.” It is full of the interesting stories that
are known well to biologists but not to others — such
as the central American Acacias that have a police force
of stinging ants that emerge from the hollow thorns to
attack intruders but also destroy the surrounding veg-
etation that competes with the Acacia. This is a mutu-
ally beneficial relationship. As well as a home in hollow
thorns the ants are supplied protein by specialized leaf
glands and sugar from other glands on the Acacia stems.
Chapter 5 “how are flowering plants distinguished”
outlines the evolution of the distinctive features includ-
ing tissues adapted to prevent water loss and to allow
transport of water and nutrients and particularly the
seed; which allowed land plants to achieve reproduc-
tion without external water, unlike ferns and frogs.
Did you know that the simple, 120 million-year-old
flower of Archaefructus has cast some suspicion on
the evolution of flowers by reduction from Magnolia-
like ancestors (contrary to what we told and regurgi-
tated for the final exam a few years ago). The classifica-
tion system, double fertilization and the evolution of
fruit and many other complex phenomena are intro-
duced in a clear and very interesting way. There is a
wealth of information in this chapter and although it
may be the most tedious chapter in the book, if you
want to know what a plant is all about, you will not
find a better general presentation of this information.
Chapter 6, “what makes flowering plants so special?”
points out that flowering plants were not present in
THE CANADIAN FIELD-NATURALIST
Vol. 120
the forests of 300 million years ago (that made coal)
but appeared 120 million years ago, were apparently
little affected by the great extinction that wiped out
the dinosaurs and other lineages, and became 87% of
all plants today. The estimated 100-fold increase in
the number of plant species since the Carboniferous
period is mostly a result of the proliferation of flow-
ering plants. The chapter explains why they are so suc-
cessful and special in sustaining 99% of life on the
planet. It may seem strange to find an explanation of
photosynthesis in the same chapter as an explanation
for the similarity of the floras of eastern North Amer-
ica and Japan, but the reader is definitely left with an
idea of how special flowering plants are.
With the excellent introduction in these chapters the
reader has developed a comprehensive knowledge of
botany and ecology which provides a basis for the last
two chapters, which indicate how flowering plants
changed the world. Chapter 7, “primates, people and
the flowering plants” explains how flowering plants
created a world in which humans could live. The nec-
tar, pollen, foliage and fruits together set the stage for
a proliferation of insects and this led to the evolution of
many new groups of insect eaters including the earliest
primates, — but how did flatter faces, dextrous digits,
upright stance and larger brains develop? It is all here.
The chapter concludes with an explanation of how
flowering plants and people came together to develop
agriculture and how major civilizations rely upon flow-
ering plants.
Of course flowering plants did not just create a world
for humans, they also changed the world in many other
ways and this is dealt with in Chapter 8, “how flow-
ers changed the world.” As they vastly increased in
numbers they increased overall biodiversity. Biochem-
ical specialization is explained with fascinating exam-
ples. Effect on the weather is considered including
gradual cooling through carbon sequestration and the
effect of grassland expansion. The chapter concludes
with an outline of the effect of human actions on flow-
ering plants and their impact on the world. An epilogue
outlines the responsibility of humans with regard to
protecting and managing plant diversity and preserv-
ing the world.
Following the epilogue is a series of notes 17 pages
in length providing additional information sources for
each chapter. This is an extremely useful section, vast-
ly improving the educational value of the work, and
it serves as an indication of the very extensive knowl-
edge of the author. The book also has a-colour insert
of 12 attractive plates. It concludes with a very useful
glossary that defines terms such as carpel, caryopsis
and cleistogamous. The index is very helpful.
Burger’s book is definitely a journey. It pulls togeth-
er a broad array of concepts into a developing theme
of how flowering plants got us to where we are today.
It offers a biological perspective including the basics
2006
of botany, ecology and evolution combined into a
picture of plants and people and ends with an evalua-
tion of where we are and where we are going. If every-
one read and understood this book — if everyone had
the knowledge and the respect for life and the inquir-
ing and sensitive mind of its author — we would be in
a better position to save the world.
This book is an example of outstanding communi-
cation. It provides biologists with what they need to
know to teach, but it is not just for the biologist, nor
just for Americans who spend nineteen billion dollars
ENVIRONMENT
Guide to Deserts
By Andrew Warren and Tony Allan (General Editors). 2006.
Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1 Canada. 240 pages. $19.95 Paper.
After reading this book I realised I have never been
to “see” a desert. I searched for Scimitar-horned Oryx
(northern Sahara). I marvelled at the marine life off
the Peruvian-Chilean coast (Sechura-Atacama desert).
I have visited the impressive Abu Simbel temples
(Libyan desert). But I have not really looked at the
desert for its own sake. This is a mistake I will not
repeat. While I have not been completely unobservant
there are many subtleties I have missed. In future I will
look for zengens, desert pavement, inselbergs and yar-
dangs.
Deserts is a neat, small book in which the author
defines deserts and describes the climatic conditions
that shape their landscapes. He gives examples of the
different types of desert and illustrates the impact wind
and — yes — water have had on the land and its inhabi-
tants. There are six chapters on aspects of deserts and
an atlas of the major desert areas. The chapters cover
wildlife such as plants, invertebrates and mammals and
how they are adapted to thrive in an arid ecosystem.
The people of the desert, their food, clothes, homes,
customs and religions are described. This is an impor-
tant chapter with the current focus of news from the
deserts of Badiyat Ash Sham (Iraq) and Dasht-E-Mar-
gow (Afghanistan). The authors explain the riches of
the desert. Not just oil, but farming and mining. They
also look at conservation issues and the damage done
by human activity. After reading this material I am
more convinced we need to re-assess the use of water
for irrigation.
Book REVIEWS
251
annually on leisure-time activities related to flowers
and plants — it is for anyone. Behind the provocative
title is a beautifully written book describing what flow-
ering plants are, how they have influenced the evolu-
tion of life, contributed to the origin of humans and
enabled humans to become the masters of our planet
with the responsibility to manage its resources prop-
erly. It is so far from being just another book!
PAUL CATLING
170 Sanford Avenue, Ottawa, Ontario K2C OE9 Canada
The atlas is really a field guide of the world’s 14
major arid zones [Canada’s minuscule desert in British
Columbia is not mentioned]. In addition to a map there
are descriptions of each area’s key characteristics. This
includes some history, the people and key issues as
well as a physical description. The authors have writ-
ten a special chapter on visiting deserts to encourage
people to see these special places.
There are plenty of photos and illustrations. These
have been chosen not so much as pretty pictures, but
to illustrate the points in the text. They range from
the stark beauty of Monument Valley, through the rock-
cut ruins at Petra to simple depictions of dunes. They
show the faces and costumes of Tuareg, Berber and
Bedouin (but not Navajo or Apache). The author has
not missed mine sites, luxury hotels or aerial views in
making his selection. One photo supposedly contain-
ing Giraffe, Ostriches and Eland shows six Oryx, some
distant gazelles (presumably Springbock) but no Eland.
This book is very interesting to read just because it
puts deserts in perspective. For someone like me who
goes to places looking for something special, this book
gives added value to a trip. For all of us it gives a won-
derful background on where desert’s ecosystem fits
in the world’s biosphere. The authors have convinced
me that deserts are important and I will pay more atten-
tion in the future. Buy this book for yourself.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
Day
THE CANADIAN FIELD-NATURALIST
Vol. 120
Endangered: Wildlife on the Brink of Extinction
By George C. McGavin. 2006. Firefly Books Ltd. 192
pages. U.S. $35.00 Cloth
An ecologist could not help but be drawn to a book
with the title Endangered. Wildlife on the brink of
Extinction. seeking an update on the current state of
the world’s species at risk. Unfortunately McGavin’s
book does not provide a succinct analysis of current
status, the impacts we should be most concerned about,
geographic localities of concern and the likely future
trends. Rather this coffee table book provides a piece-
meal of “information bites” for fifty animal groups
ranging from dogs and cats to salmon and songbirds.
The lengthy (50 of 192 pages) two chapter intro-
duction on the “Nature of Extinction” and “The Sixth
Extinction” could be a grade nine biology course cov-
ering the basics of evolution and human development
from our hunter-gather roots, through our agrarian and
industrial revolutions to an assessment of supply and
demand harvesting of nature. Instead of a discussion
of the ecological implications of the loss of species to
ecosystem integrity, extinction is considered more in
terms of the extent of human mourning and the likely
curiosity of future generations about species that once
existed.
The “Lost and the Last” chapter constitutes the cen-
tral text starting with Monotremes and Marsupials. Typ-
ical treatment of species groups include general state-
ments such as: “Platypus eggs are eaten by introduced
red foxes and rats as well as by native predators such
as lizards. Many echidna eggs suffer the same fate.”
This text provides no meaningful information on the
status of even these most unique of creatures for which
clear trends must surely be known. No consistent for-
mat is used to cover each animal group, for example
within the sub-section Insectivores Insectivora the sub-
titles used could have been chosen at random: Sense
of Smell; Desmans in Danger; Desert Moles; Rare and
Restricted; and Solendon Capture. Often the informa-
tion presented begs the question; What has this to do
with wildlife on the brink of extinction? If general eco-
logical information is to be presented it would have
been better in a text box or sidebar as a special case
study.
MISCELLANEOUS
Much more information could have been conveyed
by following a consistent format for each animal group
with sub-headings that allowed the reader to move
from one group to another building an understanding
of the most threatened species, the number of species
threatened within each group, the ecological implica-
tions of species loss, the key causes of species decline,
likely future trends and actions required to reverse neg-
ative trends. Summary graphs, tables or pictorial rep-
resentations are virtually absent, yet they too could have
been used to convey large amounts of information in
a simple repeatable format for each species group.
The most impressive aspect of Endangered is sure-
ly the figures but, while there are captions, the text
makes no direct reference to the figures and text and
figures are often unrelated. In the Prosimians group
the text states: “The strangest prosimian is surely the
aye-aye. Its wide eyes, pointed ears and thin bony fin-
gers lend it a goblin-like appearance”. No picture of
the aye-aye is included, but pictures of the sifaka and
loris are included with no information on their status.
There is considerable variability in the treatment of
each species group with some written in a more author-
itative style, Bats Chiroptera is one example where
there is a logical organization of sub-sections and more
information on extinction. Variability is also seen in
how comprehensively each species group is covered
and this appears unrelated to the extent of information
likely available. For example, the monkeys species
group includes less than one page of text while the
amphibians group has four pages of text.
In summary, Extinction provides an enormous
amount of anecdotal information that is interesting to
read, but which is likely forgotten soon after having
read it. The book leaves the reader with a sense of
disaster for all creatures as a result of human activity
(which may or may not be far off the truth) and attempts
to deal with all the ills of humankind from switching
off light bulbs to buying local produce. Unfortunately
readers that might be inclined to read Endangered will
likely come away knowing little more than they start-
ed with.
BRENT TEGLER
North-South Environmental Inc., 35 Crawford Crescent, Unit
U5, P.O. Box 518, Campbellville, Ontario LOP 1B0 Canada
The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His
Theory of Evolution
By David Quammen. 2006. Atlas Books/ W. W. Norton &
Company, New York. 304 pages. $30.00 Cloth.
As naturalists, we never cease to be amazed by the
myriad wonderful life forms in nature, be it the flow-
ering plants that suddenly appear following snowmelt,
the wide variety of birds that return north every spring,
or the colourful mushrooms found each fall in our
forests. However, the source of this very diversity may
easily be perplexing: why are there so many species
and how did they originate? The theory of evolution has
gone a long way towards explaining these and related
questions by means of a wealth of empirical and the-
oretical studies.
2006
Evolution by natural selection as [formulated] pro-
posed by Charles Darwin is one of the most impor-
tant and far-reaching ideas ever formulated, as it pro-
vides the basic mechanism whereby evolution has
created the enormous diversity of life. David Quam-
men’s objectives with his book were to offer a con-
cise yet insightful overview of the background, birth
and development of Darwin’s idea of natural selec-
tion, and to consider the reasons for Darwin’s reluc-
tance in making his theory public. Indeed, there were
many reasons why Darwin might have wanted to keep
his evolutionary views to himself. These included the
fear of rejection from a science corps whose natural
history was based largely on teleological ideals with
foundations in religious agendas, as well as his person-
al relationship with his beloved wife Emma, a pious
Christian. In the end, it was Darwin’s receipt of a man-
uscript from Alfred Russel Wallace, who at the time was
collecting beautiful bird and insect specimens some-
where in the Malay Archipelago, which prompted the
eventual publication of On The Origin of Species in
1859.
Quammen outlines this momentous piece of histo-
ry concisely and does not attempt to compete with
other more exhaustive biographies that provide a
more detailed and scholarly account of Darwin’s
idea. The result is a very readable book that, despite
the fact that Quammen is not an academic scientist,
is based on thorough research of original background
material. Hence, this book follows in the tradition of
Quammen’s previous successes, most notably The
Song of the Dodo (1996), his outstanding and award
Book REVIEWS 253
winning volume on island biogeography. Several
interesting aspects of Darwin's life are also high-
lighted, such as his mysterious illness, characterized
by dizziness, stomach pain, and bouts of illness,
often flaring up under stressful conditions. Darwin
spent weeks on end at health clinics, surrendering his
body to coldwater treatments, hoping to rid himself
of the persistent ailment. The book also describes a
number of the small natural history experiments that
Darwin performed to understand the mechanisms of
evolution, such as submerging common vegetable
seeds in brine solution to investigate the possibility
of over water dispersal, or dangling duck feet in
water to see if gastropods would climb aboard on a
flight destined for other ponds. With these sorts of
amusing passages highlighting the development of
the most important idea in biology, The Reluctant Mr.
Darwin should be of great interest to naturalists
regardless of whether they are deeply aware of evolu-
tionary theory or simply familiar with Darwin’s
work. The good quality of the material in this vol-
ume, along with Quammen’s command of the Eng-
lish language, makes this book well worth the read.
Literature Cited
Quammen, D. 1996. The Song of the Dodo — Island Biogeography
in an Age of Extinctions. Hutchinson/Pimlico/Random House of
Canada Limited, Mississauga, Ontario.
JOHAN LINDELL
35 Charles Street West, Apartment 1408, Toronto, Ontario
M4Y 1R6 Canada
A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s
Drive of Vancouver
By Rick Hudson. 2006. Volume 2. Revised Edition. Harbour
Publishing Co. Ltd., P.O. Box 219, Madeira Park, British
Columbia VON 2HO Canada. 368 pages. 8 illustrations
$16.95 CAD
Rick Hudson has succeeded in expressing his pas-
sion for mineral collecting through his field guide
book. On a second level he also tries to promote a
sense of adventure by teasing the reader to go out
and explore or prospect for nature’s buried treasures.
The author structured the book so the reader can
easily cross-reference required information in the
lengthy location lists. A “How to Use This Book” and
a “Quick Reference Guide” help to quickly locate items
of interest throughout the book.
The book cover illustrates the hobby of mineral
collecting as a family activity. Hudson educates the
reader from first principles on how to become a Rock-
hound. To accomplish this, he describes how to pre-
pare yourself by choosing proper clothing, safety gear,
collecting tools and maps. He warns the future min-
eral collector about natural hazards they may encounter
such as; rattlesnakes, grizzly bears and ticks. Ethics,
access rights and mineral rights are also discussed to
keep the collector within acceptable collecting prac-
tices and to avoid unintentional trespassing.
Mineral collecting is attractive to many people be-
cause as you find an interesting mineral specimen, you
want to learn how nature created it and or its geologi-
cal history. The author included short geological des-
criptions partitioned by collecting areas as well as min-
ing histories of local economic importance within the
collecting area.
Colour plates illustrate the minerals found in some
of the collecting sites and detailed occurrence maps
are included in all the collecting area sections through-
out the book. The colour plates represent minerals that
are possible to find in the collecting sites. Government
mineral index files were included in the locality infor-
mation along with detailed road log descriptions.
The author has made a successful merger of person-
al knowledge and technical data to compile an inform-
ative volume which can provide years of enjoyment for
many future rockhounds and their families and friends.
ROGER POULIN
President, Sudbury Rock and Lapidary Society, 3171 Romeo
Street, Val Caron, Ontario P3N 1G5 Canada
254
History of the Russian Zapovedniks 1985-1995
By F. Shtilmark. (Translated from Russian by G. H. Harper).
2003. Russian Nature Press (www.rusnatpress.org.uk). 307
pages, £30.95
This book makes a major contribution to the envi-
ronmental history of Russia, its influence zone and
also to the global nature heritage. The Zapovedniks -
Russia’s protected nature reserve system — have with-
stood the test of time, although it has been distorted
by social and political factors. Due to the great Eng-
lish translation by G. H. Harper, the Russian Nature
Press offers us new views on conservation, as well as
solutions to the current problems with protected areas
world-wide. The original text is written by one of the
leading Russian environmental writers and fascinat-
ingly energetic activists providing for superb inside
knowledge unknown to the western world. His visions
are guided by principles such as “we have a duty to
make our zapovedniks and national parks the pride of
Russia...”.
In this great book the reader will learn about Rus-
sia’s nature, and foremost, about how to protect it dur-
ing such fascinating and complex times like Tsarism,
the rules of Lenin and Stalin, and Gorbachovs’ glas-
nost and perestroika (the book manuscript was fin-
ished 1996). Throughout the history of the Zapovedniks
(in short called Zap’s) the role of the Academy of Sci-
ence proves very significant. The composition and atti-
tude of its board members towards protection vs.
exploitation play a crucial role for Zap-Management
(and species survival).
Most readers will likely be surprised to learn that
the Russian Zapovednik system actually acted as a
model for setting up the global network of biosphere
reserves. But to this very day there exists a funda-
mental confusion in the western world regarding the
terminology and meaning of Russian protected nature
reserves. As described over the 307 pages, western-
style National Parks represent only one component in
the complex Russian system of Zapodniks, Zapoved-
nosts, Zapovednayes and Zakazniks. A critical view
of the author refers to National Parks simply as pseu-
do-zapovedniks because “An absolute Zapovednik is
an area of land or water that together with all products
of nature on and in it (plants, animals, soils, rocks, min-
erals) is preserved in perpetuity and in a state of invi-
olability”.
Many Zapovednys were originally used for the Tsars
or as a prince’s hunting estate, e.g., the famous Kuban
Hunting Reserve. So originally, these systems go back
to land grants from the aristocrats. But the author takes
great care in explaining that Qualified Zapovedniks
have a weaker definition in their protection status,
allowing for human use. A Zapovednost implies the
complete withdrawal of an area of land from any eco-
nomic use. The biological theory of Zapovednosts rests
on the idea of an ecological equilibrium, and the capac-
THE CANADIAN FIELD-NATURALIST
Vol. 120
ity of organisms to regulate themselves in the absence
of direct human influence. Zapovedaniye means a with-
drawal of a certain area of nature from the sphere of
ordinary economic activity for the attainment of par-
ticular non-traditional economic, social and ecological
purposes. It represents samples of “virgin nature select-
ed for comparison with exploited land”. Zapovedny
Management relates to a method of managing protect-
ed territories and water bodies. Further, one can find
Zapovednik Museums, as well as Game Zapovedniks
and even Spa-Zapovedniks. Only the Zakazniks are
areas set aside for hunting. Finally, the Etalons are pro-
tected areas, model examples, representative for a spe-
cific ecosystem type and completely free from human
influence.
Several black/white maps, portraits, tables and dia-
grams are presented, and including with some paint-
ed Russian scenery, turn this publication into a pre-
cious book to own.
The book makes a nice case that fur-bearing ani-
mals were considered as Russia’s “Soft Gold”; they
were one of the chief resources for foreign income.
This wealth created the influential Moscow Fur Insti-
tute. Besides the earlier over-exploitation of Russian
beaver resources, the 20" century showed also a sharp
decline of sable, and a subsequent nation-wide harvest
moratorium was not really effective. Therefore, sable
reserves were requested, and a Sable Zakaznik was set
up by local people in Kamchatka as early as 1882.
These Game Zapovedniks were called Superior Zap-
ovedniks in which hunting was not allowed, and breed-
ing programs were initiated.
As elsewhere in the world, the tensions around pro-
tected areas were intense. Borders of Zapovedniks were
often not followed or defined even. Poaching was com-
mon in taiga Zapovedniks or where “old believers”,
Chinese bandits, and fur-traders lived. In Crimea and
elsewhere, the designated Zap-guards and zoo-staff
were tortured or even executed.
Besides the many day to day struggles in Zapoved-
niks, Lenin’s revolution then had a massive restruc-
turing in place for them, resulting in the “Socialisa-
tion of the Land”. It converted the entire country in a
gigantic “national park”; ownership rights were removed
with one stroke. Lenin was a hunter himself and had
a great interest in nature affairs. That proved to be for-
tunate for Russia’s nature, as well as for some aspects
of the global environment. From that point on, the
Russian Soviet Federal Socialist Republic (RSFSR,
founded 25" October 1917) played a crucial role in
the protected areas, and started its well known enthu-
siasm for regulations.
Some Russian Zapovednik administrators used the
guiding principle “Nature is not a cathedral, but a work-
shop”. And typical Zapovednik tasks dealt with the
introduction (“Acclimatisation”) and control of animal
populations. Culling was part of most Zapovedniks,
2006
and the culled game meat was contributed to the social
food-supply or else was sold to Zapovednik staff. The
elimination of crows, and cormorants as fish-eating
birds took place Russia-wide. Bears, marten, gulls,
skuas, crows, hawks and eagles got killed en masse.
Extensive campaigns against wolves by poison and air
were carried out for almost a century shaping the Rus-
sian culture against predators to this very day.
The text reads like a crime story. As felt in many
other places in the world, the Stalinist revolution fur-
ther led to more and major changes. Namely, a shift
from nature protection to exploitation in the interest
of the socialist construction. Protecting nature for the
sake of nature was not en vogue anymore. Further,
“Bourgeois” professors who comprised a large pro-
portion of the conservation’s cadre in the Soviet state
in the 1930s were eliminated, and many members of
the Academy of Science got arrested. From then on,
Zapovedniks were explored for minerals, coal and oil
deposits, timber resources and fur bearing resources,
culminating in slogans like “We must take the Urals
apart’. The fetish of inviolability as a bourgeois theory
got sacrified. Protection was now defined as exploita-
tion. Under comrade Stalin, Zapovedniks turned into
multi-purpose economic enterprises. It led to the view
that planned exploitation of Natural Resources would
be the main pre-condition of Nature Protection.
From the 1960s onwards the Zapovednik system
experienced a modernization. A. G. Bannikov and G. P.
Dementey started to participate in the IUCN. Russian-
Japanese international migratory bird conventions got
signed. In 1970-71 the UNESCO biosphere reserves
“Man and Biosphere” were established. In 1972 a U.S.-
Russian agreement on protecting the natural environ-
ment got accomplished, and the USSR Red Data book
came into place.
As early as 1975 multidisciplinary monitoring was
demanded in Russia, a theme that is very relevant for
today even, and not achieved by most countries yet.
When cheap fuel got provided for that purpose to the
Zapovedniks by the state, they got quickly swamped
with motorboats, snow tractors and motorcycles. Heli-
copters can be added to the list as well since flying was
very common by then. The author explains that cater-
pillars tracked and wheeled cross-country, and vehi-
A Field Guide to the Identification of Pebbles
By E. Van der Flier-Keller. 2007. Harbour Publishing, Box
219 Madeira Park, British Columbia VON 2HO Canada.
9 inch by 8 inch pamphlet, $7.95
Professor Van der Flier-Keller has produced a neat
pebble guide for children. She has condensed funda-
mental pebble geology into a single page. Then she
portrays 34 types of rock as they appear when they
are transformed into pebbles. She adds in a few man-
made objects that also appear on beaches, such as glass.
The pamphlet is a single long sheet of plasticized paper
Book REVIEWS 25
a)
cles ploughed up the tundra on Vrangel and Taymyr.
In the 1990s, the U.S. American Soros and Mac-
Arthur Foundations with their foreign funds gained
considerable influence over Russia’s nature; WWF,
Evraziya, the Japanese Society for the Protection of
Birds, the Global Ecological Facility, and the Interna-
tional Bank for Reconstruction & Development were
other international players accounting for as much as
one quarter of Western money on the expenditure for
protecting Russia’s biodiversity. This foreign influence
made many Russian politicians extremely uneasy.
However, despite all of these gigantic efforts, and
similar to the case in North America, nothing remains
of the many original habitats such as the steppe. “We
now need to introduce the specially-protected areas
system almost forcibly to counterbalance irrational
natural-resource-use, and to prevent deterioration of
landscapes and valuable sites”.
It becomes clear from the text that Russia and its
Zapovedniks etc. are not set up for the information
society at all, and thus Russian high quality online (raw)
data are hard to come by still. Much crucial biodiver-
sity information is published though in hardcopy and
journals, or exists in filing cabinets, private ownerships
and cabinets, and as expert knowledge. Otherwise,
check the Russian Conservation News found online
WWwW.russianconservation.org.
One cannot find a better and centralized overview
of Russian conservation history issues than this book;
its detailed Appendices are further assets consisting
of current and former Zapovednik lists, a Kyrillic ref-
erence list and in-depth annotations for each chapter.
I find that this book is a little too friendly on the wide-
spread view in Russia that traditional Zapovednik man-
agement caters primarily to the Russian race but leaves
natives and other ethnical groups out. Marine protect-
ed areas were apparently not really part of Russia's
nature protection system either.
However, “Come back in hundred years, then let’s
talk” (And in the meantime, I highly suggest buying
and reading this important book).
FALK HUETTMANN
Institute of Arctic Biology, Biology & Wildlife Department,
University of Alaska-Fairbanks, Alaska 99775 USA
with clear, “life” size photographs of pebbles. It is easy
to use and is ideal for children as it does not get over-
complicated and therefore overwhelming. For example,
it only shows two forms of granite — pink and white.
This is quite sufficient for an 8-year old to absorb.
While the author has reduced the text to a simple ex-
planation she does use the correct technical terms
throughout. Younger children may need help with some
of the terms (plutonic) or concepts (glacial deposits),
but then this is what parents are for. The pamphlet folds
256
to 4.75 inches by 9 inches and can readily slip into a
pocket or glove compartment.
Although aimed at children this is a useful tool for
the non-geologist adult who does not have the time
(or inclination) to master the complex technology of
geological science. I have a habit of picking up small
NEw TITLES
Prepared by Roy John
+ Available for review * Assigned
ZOOLOGY
The Amphibians of Belarus. By S. Drobenkov, R. Novitsky,
L. Kosova, K. Ryzhevich, M. Pikulik. 2006. Pensoft Pub-
lishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 176 pages.
EURO 34.00 Paper.
The Amphibians and Reptiles in Bulgaria. By V. Beshkov
and K. Nanev. 2006. Pensoft Publishers, Geo Milev Street
13a, 1111 Sofia, Bulgaria. 120 pages. EURO 38.90 Cloth.
* Animal Skulls, A Guide to North American Species. By
Mark Elbroch. 2006. Stackpole Books, 5067 Ritter Road,
Mechanicsburg, Pennsylvania 17055 USA. 448 pages. $44.95
PB 740 pages.
Atlas of Bird Migration. By J. Elphick (Editor) 2006. Fire-
fly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1 Canada. 176 pages. $35 Cloth.
Birder’s Companion. By S. Moss. 2006. Firefly Books, 66
Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada.
192 pages, $16.95 Paper.
* Bird Conservation Implementation and Integration in
the Americas: Proceedings of the Third International Part-
ners in Flight Conference [on CD]. By C. J. Ralph and T.
Rich. 2007. USDA, Redwood Sciences Laboratory, 1700
Bayview Drive, Arcata, California 955231 USA.
Birds of the Dominican Republic and Haiti. By S. Latta,
C. Rimmer, A. Keith, J. Wiley, H. Raffaele, K. McFarland and
E. Fernandez. 2006. Princeton University Press, 41 William
Street, Princeton, New Jersey 08540-5237 USA. 360 pages.
US. $35.
Birds: A Visual Guide. By J. Burger. 2006. Firefly Books,
66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada.
304 pages. $29.95 Cloth.
Birds of the World. Edited by A. Gossler. 2006. Firefly
Books, 66 Leek crescent, Richmond Hill, Ontario L4B 1H1
Canada. 384 pages. $39.95 Cloth.
A Birdwatcher’s Companion to North American Birdlife.
By C. Leahy. 2006. Princeton University Press, 41 William
Street, Princeton, New Jersey 08540-5237 USA. 1072 pages.
US. $19.95
Southern African Birdfinder. Callan Cohen, Claire Spot-
tiswoode and Jonathan Rossouw. 2006. New Holland Pub-
lishers (UK) Ltd., Garfield House, 86-88 Edgware Road,
London W2 2EA, U.K. 456 pages, £19.99 Paper
100 Butterflies and Moths — Portraits from the tropical
forests of Costa Rica. By J. Miller, D. Janzen and W.
THE CANADIAN FIELD-NATURALIST
Vol. 120
rocks and pebbles from the places I visit. I have not
been consistent in identifying them, but I am sure
this pamphlet will help me improve.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
Hallwachs. Harvard University Press, 79 Garden St., Cam-
bridge, Massachusetts 02138, USA. 260 pages. U.S. $39.95.
*A Field Guide to North Atlantic Wildlife. By N. Proctor
and P. Lynch. 2006. Yale University Press, P.O. Box 209040,
New Haven, Connecticut 06520-9040 USA. 221 pages. U.S.
$19.95.
+ Conservation of the Black-tailed Prairie Dog. Edited by
J. Hoogland. 2006. Island Press, 1718 Connecticut Avenue,
NW, Suite 300, Washington D.C. 20009. 368 pages. U.S.
$35 Paper.
+ The Northern Goshawk: A technical Assessment of its
Status, Ecology and management. Edited by M. Morrison.
2006. Cooper Ornithological Society, c/o Western Founda-
tion of Vertebrate Zoology, 439 Calle San Pablo, Camarillo
California 93012-8506, USA. 369 pages. U.S. $23 Paper.
* Migrating Raptors of the World — Their Ecology and
Conservation. By Keith L. Bildstein. 2006. Cornell Universi-
ty Press, 512 East State Street, Ithaca, New York 14850
USA. 344 pages. $35.00 Cloth.
Rattlesnake Adventures: Hunting with Oldtimers. By J.
Kemnitzer, Jr. [Editor]. 2006. Krieger Publishing Company,
P.O. Box 9542, Melbourne, Florida 32902-9542. 236 pages.
US. $32.50.
Atlas of the Millipedes (Diplopoda) of Britain and Ire-
land. By P. Lee. 2006. Pensoft Publishers, Geo Milev Street
13a, 1111 Sofia, Bulgaria. 216 pages. EURO 32.00 Cloth.
An Atlas of the Reptiles of North Eurasia. Taxonomic
Diversity, Distribution, Conservation Status. By N. Anan-
jeva, N. Orlov, N. Khalikov, R. Darevsky, I. Ryabov, I. Bara-
banov. 2006. Pensoft Publishers, Sofia-Moscow. 250 pages.
EURO 75.00 Cloth.
* Songbird Journeys: Four Seasons in the Lives of Migra-
tory Birds. By Mary Chu. 2006. Walker & Company, 104
Fifth Avenue, New York, New York 10011 USA. 312 pages.
U.S. $23.00 Cloth.
Snake Venoms and Evenomations. By J-P. Chippaux. 2006
English Edition. Krieger Publishing, P.O. Box 9542, Mel-
bourne, Florida 32902 USA. 300 pages. U.S. $58.50.
+ Tracks. By D. Jackson. 2006. University Press of Missis-
sippi, 3825 Ridgewood Road, Jackson, Mississippi 39211
USA. $36.25 279pages. Cloth.
The Alligator Snapping Turtle. By P. Pritchard. 2006 re-
print. Krieger Publishing, P.O. Box 9542, Melbourne, Flori-
da 32902 USA. 152 pages. U.S. $42.
2006
* Turtles of the World. By F Bonin, B Devaux and A Dupre.
2006. The Johns Hopkins University Press, 2715 North Charles
Street, Baltimore, Maryland 21218-4363, USA. 416 pages.
U.S. $50 Cloth.
Whales, Dolphins and Seals — A Field Guide to the Marine
Mammals of the World. By Hadoram Shirihai and Brett
Jarrett. 2006. NHBS Environment Bookstore, 2-3 Wills Road,
Totnes, Devon TQ9 5XN, U.K. 384 pages, £12.99 Cloth.
BOTANY
Botanical Riches — Stories of Botanical Exploration. By
R. Aitken. 2007. Ashgate Publishing, Box 2225, Williston
Vermont 05495. 256 pages. U.S. $ 50 Cloth.
Wildflowers of the Rocky Mountains. By G. Scotter. 2006.
Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1 Canada. 256 pages. $29.95 Paper.
* Wild Orchids of the Canadian Maritimes and Northern
Great Lakes Region. By Paul Martin Brown. 2006. Uni-
versity Press of Florida, 15 Northwest 15" Street, Gaines-
ville, Florida 32611-2079 USA. 366 pages. U.S. $ 29.95
* Wild Orchids of the Pacific Northwest and Canadian
Rockies. By Paul Martin Brown. 2006. University Press of
Florida, 15 Northwest 15" Street, Gainesville, Florida 326] 1-
2079 USA. 304 pages. U.S. $ 29.95
ENVIRONMENT
Algonquin Souvenir. By M. Runtz. 2006. Firefly Books,
66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada.
96 pages. $19.95 Cloth.
An Artist’s and Photographer’s Guide to Wild Ontario.
By R. Stimson and C. Thompson. 2006. Firefly Books, 66
Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada.
224 pages. $29.95 Paper.
The Enchantment of Birds. By R. Cannings. 2007. Grey-
stone Books, Suite 201, Quebec Street, Vancouver, British
Columbia VST 4S7 Canada. 168 pages. $29.95 Cloth.
The International Politics of Bird Conservation. By R.
Boardman. 2006. Edward Elgar Publishing, 9 Dewey Court,
Northampton, Massachusetts 01060 USA. 288 pages. U.S.
$110 Cloth.
* The Atlas of Climate Change. By Kirstin Dow and
Thomas E. Downing. 2006. NHBS Environment Bookstore,
2-3 Wills Road, Totnes, Devon TQ9 SXN, U.K. £12.99, Paper.
Principles of Population Genetics — 4" edition. By D.
Hart and A. Clark. Sinauer Associates, Box 407, Sunderland,
Massachusetts 01375 USA. 565 pages. U.S. $93.95.
An Inconvenient Truth. By Al Gore. 2006. NHBS Environ-
ment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN,
UK. £14.99.
The Rough Guide to Climate Change. By Robert Henson.
2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes,
Devon TQ9 5XN. U.K. £9.99.
Book REVIEWS
* The Reluctant Mr. Darwin: An Intimate Portrait of
Charles Darwin and the Making of His Theory of Evolu-
tion. By David Quammen. 2006. Atlas Books, 10 East 53rd
Street, 36th Floor, New York, New York 10022, USA. 304
pages. U.S. $22.95.
+ Marshes. By W. Burt. 2007. Yale University Press. P.O. Box
209040, New Haven, Connecticut 06520-9040 USA. 192
pages. U.S. $35 Paper.
Nature’s Engraver — A life of Thomas Bewick. 2007. Far-
rar Straus Giroux, North Point Press, 19 Union Square West,
New York, New York 10003. 480 pages. $27.50 Cloth.
Wild Nova Scotia. By L.Wagg. 2006. Nimbus Publishing,
Box 9166, Halifax, Nova Scotia B3K 5M8 Canada. 96 pages
$29.95.
People, Fish and Whales — The Vancouver Aquarium
Story. By Dr. Murray Newman. 2006. Harbour Publishing,
P.O. Box 219, Madeira Park, British Columbia VON 2HO
Canada. 120 pages. $19.95 Paper.
* Pilgrimage to Vallombrosa — From Vermont to Italy in
the Footsteps of George Perkins Marsh. By John Elder.
2006. University of Virginia Press, 210 Sprigg Lane, Char-
lottesville, Virginia 22903 USA. U.S. $33.99 Cloth.
Sarcophaginae et Paramacronychiinae du Proche Orient
(Insecta, Diptera, Sarcophagidae). By A. Lehrer. 2006. Pen-
soft Publishers, Sofia-Moscow. [In French.}, 264 pages. EURO
64.00 Cloth.
Tales from the Thébaide- Reflections of a Turtleman. By
P. Pritchard. 2006. Krieger Publishing Company, P.O. Box
9542, Melbourne, Florida 32902-9542 USA. 340 pages. U.S.
$44.50.
Wild Borneo. By N. Garbutt and J. Prudente. 2006. The MIT
Press, Five Cambridge Center, 4" Floor, Cambridge, Mass-
achusetts 02142-1493 USA. 176 pages. $34.95 US.
The End of the Wild. By S. Meyer. 2006. The MIT Press,
Five Cambridge Center, 4“ Floor, Cambridge, Massachusetts
02142-1493 USA. 96 pages. U.S. $14.95.
+ Wildfire in the Wilderness. By C. Czajkowski. 2006. Har-
bour Publishing, P.O. Box 219, Madeira Park, British Colum-
bia VON 2HO Canada. 224 pages. $19.95.
Endangered Wildlife on the Brink of Extinction. By G.
McGavin. 2006. Firefly Books Ltd., 66 Lark Crescent, Rich-
mond Hill, Ontario L4B 1H1 Canada. 142 pages. $35. Cloth.
YOUNG NATURALISTS
Tale of a Great White Fish — A sturgeon story. 2007. Grey-
stone Books, Suite 201 Quebec Street, Vancouver, British
Columbia VST 4S7 Canada. 48 pages. $10.95 Cloth.
SPECIAL NOTE
Sale of the Herpetological Library of Adrian Crane (800 items
including antiques). PDF catalogue from Breck @herplit.com
News and Comment
Marine Turtle Newsletter (112)
April 2006. 28 pages: ARTICLES: Nesting of Green Turtles
in Saint Leu, Reunion Island (S. Ciccione and J. Bourjea)
— Sea Turtles and Fishery Interactions in Brazil: Identifying
and mitigating potential conflicts (M. A. Marcovaldi, G.
Sales, J. C. A. Thome, A. C. C. Dias da Silva, B. M. G.
Gallo, E. H. S. M. Lima, E. P. Lima, C. Bellini) — First
Report of Leatherback Turtle Entanglement in Trap Lines in
the Uraguayan Continental Shelf (M. Laporta, P. Miller, S.
Horta, and G. Riestra) — Loggerhead Turtle Nesting
Activity in Kuriat Islands (Tunisia): Assessment of Nine
Years Monitoring (I, Jribi, M. N. Bradai and A. Bouain) —
Hawksbill Turtles on the Pacific Coast of Costa Rica (A.
Gaos, R. Arauz and I. Yanez) — NOTES: Oceanic Movement
of Benthic Foraging Juvenile Hawksbill Turtle from The
Cocos (Keeling) Islands (S. D. Whiting and A. U. Koch) —
Use of Marine Turtles in Zootherapy in Northeast Brazil (R.
Romeu da N. Alves) — MEETING REPORTS — ANNOUNCE-
MENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICA-
TIONS.
The Marine Turtle Newsletter is edited by Brendan J.
Godley and Annette C. Broderick, Marine Turtle Research
Group, Centre for Ecology and Conservation, University
of Exeter in Cornwall, Tremough Campus, Penryn TR10
9EZ United Kingdom; e-mail MTN @seaturtle.org; Fax +44
1392 263700. Subscriptions and donations towards the pro-
duction of the MTN can be made online at or postal mail to Michael Coyne (online
Editor) Marine Turtle Newsletter, 1 Southampton Place,
Durham, North Carolina 27705 USA (e-mail: mcoyne@
seaturtle.org).
Herpetological Survey 2006 Atlas of Amphibians and Reptiles of Quebec
Instructions for participants in the 19" herpetological sur-
vey for the Quebec Atlas of amphibians and reptiles, coordi-
nated by the St. Lawrence Natural History Society in col-
laboration with the Ministére des Ressouces naturelles de la
Faune du Québec. This provincial inventory is done every
year with the important cooperation of volunteers. Observation
cards to fill out and details of information needed can be ob-
tained from David Rodrique, Director, Research and Con-
servation, Program Coordinator, or Mathieu Oulette, Research
and conservation, St. Lawrence Valley Natural History Society,
Ecomuseum, 21 125 chemin Saint-Marie, Saint-Anne-de-
Bellevue, Quebec H9X 3Y7; phone 514-457-9449, extension
105; fax 514-457-0769; e-mail: info@herpetofaune.org. Cur-
rent information on The Atlas of Amphibians and Reptiles
of Quebec is now available online at the St. Lawrence
Natural History Society website: www.herpetofaune.org.
Bill Cody Receives Yukon Biodiversity Awareness Award
From 1946 to 1987 Bill (W.J.) Cody was employed
with Agriculture and Agri-Food Canada as plant tax-
onomist and curator of Canada’s largest collection of
plants. From 1988 to 2006 he has continued to come in
to work every day as an Honourary Research Associate
and he has made a series of remarkable contributions
to our knowledge of the plants of northwestern North
America. His 643 page textbook on the Flora of the
Yukon was published in 1996. It included current
information on status, distribution, ecology, classifica-
tion and identification. In 1997 this outstanding work
was recognized with the prestigious Lawson Medal
awarded by the Canadian Botanical Association. An
updated second edition of the book was published by
the National Research Council in 2000. Since his
retirement Bill has published 14 articles on the flora
of the Yukon in scientific journals. He has provided
an enormous amount of information on plants to the
agricultural sector, natural resources staff, wildlife biol-
ogists, native people and landscape planners. Always
willing to help, he has identified a thousand plants
from the Yukon every year. It is therefore not a sur-
prise that Bill has been recognized for his outstanding
service with the Yukon Biodiversity Awareness Award.
The award honours those who have made major con-
tributions to educating people about biodiversity and
its importance.
258
)
2004
Bill started as an assistant with the Canada Depart-
ment of Agriculture in Ottawa. Twenty-one years later
in 1967 he was made a Research Scientist. This clas-
sification was generally reserved for people who had
a Ph.D., but Bill’s outstanding accomplishments at
that time were judged by his colleagues and the sci-
ence arm of the federal civil service to warrant treat-
ment at the Ph.D. level. The awards that Bill has
received since then are numerous, but one in partic-
uar draws attention his broad contribution. In No-
vember 2002 he received a Queen’s Golden Jubilee
Commerorative Medal. These medals were awarded
to a limited number of people who have made a sig-
nificant contribution to Canada, in this case “especial-
ly for his work on The Canadian Field-Naturalist,
Canada’s foremost scientific journal for field biology.”
Bill has served as the business manager, article and
book review contributor, and manuscript reviewer for
this journal for 60 years, and his influence on its devel-
opment, support, improvement and content is beyond
question.
Bill also served as curator of the largest dried plant
collection in Canada from 1959 to 1988 and was large-
ly responsible for its development. This collection of
now over | million specimens has become a major tool
of Agriculture and Agri-food Canada for plant identi-
fication. This collection provides a wealth of informa-
Errata The Canadian Field-Naturalist 120(1)
NEWS AND COMMENT
259
tion that is needed to implement Agriculture and Agri-
food Canada’s Biodiversity Strategy and to respond
to the International Convention on Biodiversity. It is
particularly valued as a tool for improved plant clas-
sification studies used by researchers in other institu-
tions both in Canada and worldwide. It is also impor-
tant in protecting Canada’s borders and enforcing
federal regulations. Many thousands of specimens
that Bill collected in the northern wilderness under
extreme and dangerous conditions are part of this
collection and serve as vouchers for his numerous
publications and books.
Bill was born in Hamilton on 2 December 1922. His
father was a doctor and his mother a nurse at Hamilton
General. He grew up in Hamilton and received his
B.A. from McMaster University in 1946.
The Yukon Biodiversity Awareness Award plaque
that Bill received included a photo of one of Yukon’s
rarest plants, McBride’s Phacelia (Phacelia mollis)
which is a Beringian endemic (confined to the ungla-
ciated area of Alaska and Yukon).
Text slightly modified from that provided courtesy
of Paul Catling, Canada Agriculture and Agri-food,
Ottawa. Photograph of the award plaque courtesy
Stephen Daryshire, Canada Agricuture and Agri-food,
Ottawa.
Book Review. Lapland a Natural History, pages 123-124: replace “Redcliff’ and Redcliffe” with Ratcliffe, throughout.
Articles.
Diversity and range of amphibians and reptiles of the Yukon Territory. Brian G. Slough and R. Lee Menell in
Literature Cited page 91 “Matsurla” should read Matsuda.
Recent Declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces. Anthony J. Erskine.
Page 48 insert following Dunn et al.
Erskine, A. J. 1980. A House Sparrow die-off. Nova Scotia Bird Society Newsletter 22: 183-184.
For reference following Erskine 1992b insert Erskine before initials.
Editor’s Report for Volume 119 (2005)
Mailing dates for issues in volume 119: (1) 6 June
2006, (2) 13 June 2006, (3) 30 November 2006, (4)
17 April 2007. A summary of membership and sub-
scriber totals 2004 is given in Table 1. The number of
articles and notes in volume 119 is summarized in
Table 2 by topic; totals for Book Reviews and New
Titles are given in Table 3, and the distribution of con-
tent by page totals per issue in Table 4.
Council continued to authorize 40% of membership
dues for publication of the journal and all of subscrip-
tions (both individual and institutional). Council also
has allocated 80% of the annual interest from the Man-
ning Fund and other capital funds to The Canadian
Field-Naturalist. Use of the Manning fund portion was
broadened on an issue-by-issue basis to offset the publi-
cation costs where author and institutional contributions
were insufficient to cover page charges.
The journal was printed at Gilmore Printers, Ottawa,
and thanks are due business representative Tom Smith,
customer representative Chuck Graham for overseeing
production, and Wendy Cotie for typesetting and cor-
rections. Leslie Cody prepared the Index for volume
119; Elizabeth Morton proofed the galleys. Long-time
(60 years) Business Manager for the journal, Bill Cody,
gradually relinquished much of his active role during
the year and some of his duties were shouldered by
treasurer Frank Pope and assistant treasurer Jim Ward.
Roy John processed books sent by publishers, assigned
reviewers, edited book reviews and prepared the
New Titles listing. Sandra Garland the Ottawa Field-
Naturalists’ Club webmaster, posted contents and ab-
stracts from each issue as it appeared, and sent pdfs
of articles and notes ordered by authors.
Manuscripts (excluding book reviews, notices, and reports)
submitted to The Canadian Field-Naturalist totalled 75 in
2005, equalling the total for 2004. The following reviewed
papers submitted in 2005 (with number of manuscripts re-
viewed in parentheses if more than one):
TABLE 2. Number of articles and notes published in The
Canadian Field-Naturalist Volume 119 (2005) by major field
of study.
Subject Articles Notes Total
Mammals 19 17 36
Birds 14 5 19
Amphibians + reptiles 2 1 3
Fish 6 1 7
Invertebrates 7 2 9
Plants 13 1 14
Other Ie 0) 1
Totals 62 Dl 89
Associate Editors:
R. Anderson, Canadian Museum of Nature, Ottawa, On-
tario (2); C. D. Bird, Erskine, Alberta (9); R. R. Campbell,
St. Albert, Ontario (3); P. M. Catling, Agriculture and Agri-
food Canada, Ottawa (7); B. W. Coad, Canadian Museum of
Nature, Ottawa (8); A. J. Erskine, Sackville, New Brunswick ||
(12); D. F. McAlpine, New Brunswick Museum, Saint John,
New Brunswick (9); D. W. Nagorsen, Mammalia Biological
Consulting, Victoria, British Columbia (9); W. O. Pruitt, Jr.,
University of Manitoba, Winnipeg (16).
Additional Reviewers: W. B. Ballard, Texas Tech University,
Lubbock, Texas (3); L. Bernatchez, Laval University, Quebec,
Quebec; J. R. Bider, Baie o’ Urfe, Quebec; G. Blouin-Demers,
University of Ottawa, Ontario; D. Boyd, Helena, Montana;
M. Brigham, University of Regina, Saskatchewan; L. Carbyn.
Canadian Wildlife Service, Edmonton, Alberta (3); J. Cayou-
ette, Agriculture and Agri-Food Canada, Ottawa; J. Cebek.
Trent University, Peterborough, Ontario; T. Chubbs, Happy
Valley — Goose Bay, Labrador, Newfoundland & Labrado1
(3); H. D. Cluff, Resources, Wildlife & Economic Develop-
ment, Yellowknife, Northwest Territories (2); M. Crete, Mini-
stre de l’Environment, Quebec, Quebec; A. W. Diamond
University of New Brunswick, Fredericton, New Brunswick
(2); L. C. Dickamer, Northern Arizona University, Flagstaff
Arizona; P. Dobson, School of Veterinary Medicine, Phila-)
delphia, Pennsylvania; J. Eger, Royal Ontario Museum, Tor:
TABLE 1. The 2005 circulation of The Canadian Field-Naturalist (2004 in parenthesis). Compiled by Robin Murphy, bmrgroup
Ottawa.
Canada USA Other Totals
Memberships
Family & individual 624 (717) 26 (26) 6 (6) 656 (749)
Subscriptions
Individuals 130 (165) 63 (63) 4 (5) O33)
Institutions 156 (164) 229 (246) 34 = (30) 419 (440)
TOTALS 910 (1046) 318 (335) 44 (41) 1272 (1422)
Note: 22 countries are included under “Other” (outside Canada and United States): Austria, Belgium, Brazil, Denmark (2), Unite
Kingdom (9: including 1 to Scotland), Finland (2), France (3: including 1 to St. Pierre & Miquelon), Germany (2), Icelanc
Ireland, Japan, Mexico, Netherlands (3), New Zealand, Norway (4), Poland, Russia, South Africa, Spain (3), Sweden (2 |
Switzerland (2), Trinidad and Tobago.
260
2005
TABLE 3. Number of reviews and new titles published in Book
Review section of The Canadian Field-Naturalist Volume 119
by topic.
——— — — _ _ _— —_—_ — — EEEEEns=gQTeaeaEs«q@nanana=anaaa———
Reviews New Titles
Zoology 33 104
Botany 5 19
Environment 17 10
Miscellaneous 6 20
Young Naturalists 0 2
Totals 61 lis)
onto, Ontario; W. Foissner, Institut fur Zoologie, Salzburg,
Austria; B. Freedman, Dalhousie University, Halifax, Nova
Scotia; W. A. Fuller, Athabasca, Alberta; D. M. Green, Red-
path Museum, McGill University, Montreal, Quebec (4); P.
T. Gregory, University of Victoria, British Columbia (3); E.
Haber, National Botanical Services, Ottawa, Ontario; F. H.
Harrington, Mt. St. Vincent University, Halifax, Nova Scotia;
-Y.L. Harms, University of Saskatchewan, Saskatoon; T. Hax-
ton, Ministry of Natural Resources, Kemptville, Ontario (2);
| S. Herrero, University of Calgary, Alberta (2); G. Holroyd, Can-
cadian Wildlife Service, Edmonton, Alberta; E. Holm, Royal
Ontario Museum, Toronto; S. Hooker, University of St. An-
‘drews, Fife, United Kingdom; C. S. Houston, Saskatoon, Sas-
katchewan; R. James, Sutherland, Ontario (2); M. Johnson,
Game & Fish Department, Bismarck, North Dakota; T. S. Jung,
‘Environment, Fish and Wildlife Branch, Whitehorse, Yukon
(2); R. W. Knapton, Edmonton, Alberta; K. Larsen, University
‘College of the Cariboo, Kamloops, British Columbia; J. Lien,
‘Memorial University of Newfoundland, St. John’s; R. Mac-
‘Culloch, Royal Ontario Museum, Toronto, Ontario; F. F. Mal-
‘lory, Laurentian University, Sudbury, Ontario (2); J. Maunder,
‘Pond Cove, Newfoundland (3); L. D. Mech, U.S. Geological
Survey, The Raptor Center, University of Minnesota, St.
‘Paul (3); R. Meisterfeld, Institute for Biology II (Zoology),
‘Department of General Biology, Aachen; D. Naughton, Cana-
‘tian Museum of Nature, Ottawa; J. S. Nelson, University of
Alberta, Edmonton; J-P. Ouellet, Universite du Quebec a Ri-
‘nouski; P. Paquet, Meacham, Saskatchewan (3); D. Paulson,
Seattle, Washington; G. Parker, Canadian Wildlife Service,
sackville, New Brunswick; S. Petrie, Bird Studies Canada,
! Port Rowan, Ontario; G. Proulx, Alpha Research & Manage-
ment Ltd., Sherwood Park, Alberta (2); Raine, Golder Asso-
viates Ltd., Calgary, Alberta; R. Reeves, Okapi Wildlife Asso-
Epiror’s REPORT
26)
TABLE 4, Number of pages per section published in The Cana
dian Field-Naturalist Volume 119 (2005) by issue
(1) (2) (4) Total
Articles 128 132 118 95 473
Notes 12 K 21 16 57
Book Reviews’ 15 20 1] 19 65
CFN/OFNC Reports” 0 2 x 4 14
News and Comment 2 2 | | 6
Index 0 0 0 32 32
Advice to Contributors | 0 | |
Totals 158 164 160 168 650
* Total pages for book review section include both reviews and
new titles listings.
™ Includes CFN Editor's report in (2), OFNC Annual Business
Meeting (3) and OFNC Awards (4).
ciates, Hudson, Quebec; J. D. Rising, University of Toronto,
Ontario; L. Rogers, Ely, Minnesota; R. Rosatte, Ministry of
Natural Resources, Trent University, Peterborough, Ontario;
A. Russell, University of Calgary, Alberta; S. Sampson, Utah
Museum of Natural History, Sault Lake City; Fred Scott,
Acadia University, Wolfville, Nova Scotia; F. W. Schueler,
Bishops Mills Natural History Center, Oxford Mills, Ontario
(6); F. E. Schwab, College of the North Atlantic, Labrador City,
Newfoundland and Labrador; D. Smith, Southern Connecticut
State College, New Haven; K. W. Stewart, University of
Manitoba, Winnipeg, Manitoba; J. Theberge, Oliver, British
Columbia (3); I. Thompson, Canadian Forest Service, Sault
Ste. Marie, Ontario (2); P. M. Youngman, Ottawa, Ontario.
I am also indebted again to the President of the Ot-
tawa Field-Naturalists’ Club Gary McNulty and the Club
Council for continuing support of the journal; Chair-
man Ron Bedford and the Publications Committee of
the OFNC for editorial encouragement and support,
the finance committee for input and encouragement to
speed things up, the Canadian Museum of Nature for
access to its library and the facilities at the Natural
Heritage Building, 1740 Pink Road, Aylmer, Quebec,
and to Joyce for continuing support.
FRANCIS R. COOK
Editor
Advice for Contributors to The Canadian Field-Naturalist
Content
The Canadian Field-Naturalist is a medium for the publi-
cation of scientific papers by amateur and professional natu-
ralists or field-biologists reporting observations and results
of investigations in any field of natural history provided that
they are original, significant, and relevant to Canada. All read-
ers and other potential contributors are invited to submit for
consideration their manuscripts meeting these criteria. The
journal also publishes natural history news and comment items
if judged by the Editor to be of interest to readers and sub-
scribers, and book reviews. Please correspond with the Book
Review Editor concerning suitability of manuscripts for this
section. For further information consult: A Publication Policy
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian
Field-Naturalist 97(2): 231-234. Potential contributors who
are neither members of The Ottawa Field-Naturalists’ Club
nor subscribers to The Canadian Field-Naturalist are encour-
aged to support the journal by becoming either members or
subscribers.
Manuscripts
Please submit by post to the Editor, in either English or
French, three complete manuscripts written in the journal
style. Manuscripts may also be submitted (one copy) by e-
mail. The research reported should be original. It is recom-
mended that authors ask qualified persons to appraise the
paper before it is submitted. All authors should have read and
approved it. Institutional or contract approval for the publica-
tion of the data must have been obtained by the authors. Also
authors are expected to have complied with all pertinent leg-
islation regarding the study, disturbance, or collection of ani-
mals, plants or minerals. The place where voucher specimens
have been deposited, and their catalogue numbers, should be
given. Latitude and longitude should be included for all indi-
vidual localities where collections or observations have been
made.
Manuscripts should be printed on standard-size paper, dou-
blespaced throughout, generous margins to allow for copy
marking, and each page numbered. For Articles and Notes
provide a bibliographic (citation) strip, an abstract, and a list
of key words. Generally, words should not be abbreviated but
use SI symbols for units of measure. The names of authors of
scientific names may be omitted except in taxonomic manu-
scripts or other papers involving nomenclatural problems.
“Standard” common names (with initial letters capitalized)
should be used at least once for all species of higher animals
and plants; all should also be identified by scientific name.
The names of journals in the Literature Cited should be
written out in full. Unpublished reports and web documents
should not be cited here but placed in a separate Documents
Cited section. List the captions for figures numbered in arabic
numerals and typed together on a separate page. Present the
tables each titled, numbered consecutively in arabic numerals,
and placed on a separate page. Mark in the margin of the text
the places for the figures and tables.
Check recent issues (particularly Literature Cited) for
journal format. Either “British” or “American” spellings are
acceptable in English but should be consistent within one
manuscript. The Oxford English Dictionary, Webster’s
’ New International Dictionary and le Grand Larousse
Encyclopédique are the authorities for spelling.
Illustrations
Photographs should have a glossy finish and show sharp
contrasts. Electronic versions should be high resolution. |
Photographic reproduction of line drawings, no larger than
a standard page, are preferable to large originals. Prepare
line drawings with India ink on good quality paper and let-
ter (don’t type) descriptive matter. Write author’s name, title
of paper, and figure number on the lower left corner or on ©
the back of each illustration.
Reviewing Policy
Manuscripts submitted to The Canadian Field-Naturalist
are normally sent for evaluation to an Associate Editor (who |
reviews it or asks another qualified person to do so), and at |
least one other reviewer, who is a specialist in the field, cho-
sen by the Editor. Authors are encouraged to suggest names |
of suitable referees. Reviewers are asked to give a general |
appraisal of the manuscript followed by specific comments
and constructive recommendations. Almost all manuscripts
accepted for publication have undergone revision—some-
times extensive revision and reappraisal. The Editor makes |
the final decision on whether a manuscript is acceptable for —
publication, and in so doing aims to maintain the scientific
quality, content, overall high standards and consistency of
style, of the joumal.
Special Charges — Please take note
Authors must share in the cost of publication by pay-
ing $90 for each page, plus $20 for each illustration (any size
up to a full page), and up to $90 per page for tables (depend-
ing on size). Authors may also be charged for their changes in
proofs. Reproduction of color photos is extremely expensive;
price quotations may be obtained from the Editor. If grant or
institutional funds are not available, club members and sub-
scribers may apply for a waiver of charges for the first five —
pages.
Limited joumal funds are available to help offset publi- |
cation charges to authors without grants or institutional sup- |
port. Requests for financial assistance should be made to the |
Editor when the manuscript is submitted.
An order form for the purchase of repents will accompa-
ny the galley proofs sent to the authors. Invoices for publi- |
Reprints |
cation costs will be sent when the submission is published. |
FRANCcIs R. Cook, Editor |)
RR 3 North Augusta, Ontario KOG IRO Canada ||
262
TABLE OF CONTENTS (concluded) Volume 120 Number 2
Osprey, Pandion haliaetus, depredates Common Eider, Somateria mollissima, duckling
BRENDA M. BLINN, VICKY VIOLETTE, and ANTONY W. DIAMOND
Extension de l|’aire de distribution de vase, Umbra limi, dans le nord-est du Québec
JEAN-FRANCOIS DESROCHES
The Mink Frog, Rana septentrionalis, in southeastern Labrador
JEAN-FRANCOIS DESROCHERS, ISABELLE PICARD, and JOHN E. MAUNDER
First record of the Great Barracuda, Sphyraena barracuda, from Canada
DARLENE BALKWILL, BRIAN W. COAD, ISMAEL GALVEZ, and JOHN GILHEN
Book Reviews
Zoo.LoGy: Animal Skulls: A Guide to North American Species — A Field Guide to North Atlantic
Wildlife — A Complete Guide to Arctic Wildlife — The Black Flies (Simultidae) of North America —
Insects: Their Natural History and Diversity: With a Photographic Guide to Insects of Eastern North
America — The Northern Goshawk: A Technical Assessment of its Status, Ecology and Management
— Turtles of the World — The Wolves of Algonquin Park: A 12-Year Ecological Study
Borany: Flowers: How They Changed the World
ENVIRONMENT: Guide to Deserts — Endangered: Wildlife on the Brink of Extinction
MISCELLANEOUS: The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of
His Theory of Evolution — A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia,
Sites Within a Day’s Drive of Vancouver — History of the Russian Zapovedniks 1985-1995 — A Field
| Guide to the Identification of Pebbles
NEW TITLES
News and Comment
Marine Turtle Newsletter (112) — Herpetological Survey 2006 Atlas of Amphibians and Reptiles of
Quebec — Bill Cody Receives Yukon Biodiversity Awareness Award — Errata: The Canadian Field-
Naturalist 120(1)
Editor’s Report for Volume 119 (2005)
po to Contributors
i
Mailing date of the previous issue 120(1): 16 July 2007
2006
258
260
262
THE CANADIAN FIELD-NATURALIST Volume 120 Number 2
Articles
George Wayne Douglas 1938-2005 JENIFER L. PENNY
Conservation evaluation the Prairie Lupine, Lupinus lepidus var. lepidus,
in Canada GEORGE W. DOUGLAS and MICHAEL RYAN
Conservation evaluation of Seaside Bird’s-foot Trefoil, Lotus formosissimus, in Canada
GEORGE W. DOUGLAS and MICHAEL RYAN
Conservation evaluation of Bog Bird’s-foot Trefoil, Lotus pinnatus, in Canada
MARTA DONOVAN
Conservation evaluation of Dwarf Wolly-heads, Psilocarphus brevissimus var. brevissimus,
in Canada GEORGE W. DOUGLAS, JENIFER L. PENNY, and KSENIA BARTON
Conservation evaluation of Pacific Rhododendron, Rhododendron macrophyllum,
in Canada GEORGE W. DOUGLAS and JULIE DESROSIERS
Conservation evaluation of Slender Collomia, Collomia tenella, in Canada
GEORGE W. DOUGLAS and JENIFER L. PENNY
Conservation evaluation of Small-flowered Tonella, Zonella tenella, in Canada
GEORGE W. DOUGLAS and JENIFER L. PENNY
Conservation evaluation of Stoloniferous Pussytoes, Antennaria flagellaris, in Canada
GEORGE W. DOUGLAS, JENIFER L. PENNY, and KSENIA BARTON
Late-born Elk, Cervus elaphus, calf with spots observed near Bancroft, Ontario
Rick ROSATTE and JOE NEUHOLD
Annual variation in habitat use by White-footed Mice, Peromyscus leucopus, in forest fragments:
the effects of forest patch size, edge, and surrounding vegetation type
CHRISTINE S. ANDERSON, DOUGLAS B. MEIKLE, ALAN B. CApDy, and ROBERT L. SHAEFER
Response of the Sea Louse Lepeophtheirus salmonis infestation levels on juvenile
wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, salmon, to arrival of parasitized
wild adult salmon ALEXANDRA Morton and Ros WILLIAMS
Physical characteristics, hematology, and serum chemistry of free-ranging Gray Wolves,
Canis lupus, in southcentral Alaska
MATTHEW J. BUTLER, WARREN B. BALLARD, and HEATHER A. WHITLAW
Are Lesser Snow Geese, Chen caerulescens caerulescens, exceeding the carrying capacity
of the Fraser River Delta’s brackish marshes? MIKE W. DEMARCHI
Predicting Raccoon, Procyon loter, occurrence through the use of microhabitat variables
ROGER A. BALDWIN, ALLAN E. HousTON, MICHAEL L. KENNEDY, and PIN SHUO LIU
Notes
Scoter, Melanitta spp., interrupted migrations by Confederation Bridge: An update
KATHERINE BUNKER-POPMA
An observation of the spring 2006 migration of Black Scoter, Melanitta nigra, in Northumberland
Straight, interrupted by the Confederation Bridge, New Brunswick—Prince Edward Island
CoLIN MACKINNON and ANDREW KENNEDY
First record of a River Otter, Lutra canadensis, captured on the northern coast of Alaska
SHAWN P. HASKELL
2006
133
147
153
158
163
169
le
179
183
188
192
192
235,
(continued on inside back cover)
ISSN 0008-3550
(7)
1S aH
The CANADIAN
FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
Volume 120, Number 3 July-September 2006
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Her Excellency The Right Honourable Michaélle Jean
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members
Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope
Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr.
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson
2006 Council
President: Mike Murphy Ronald E. Bedford = Diane Kitching Dan Millar
Vice-President: Ken Allison Fenja Brodo Karen McLachalan Hamilton Justin Peter
Recording Secretary: Susan Laurie-Bourque Julia Cipriani David Hobden Stanley Rosenbaum
Treasurer: Frank Pope William J. Cody Diane Lepage Henry Steger
Past President: Gary McNulty Francis R. Cook Ann Mackenzie Chris Traynor
Susan Howell Gillian Marston Eleanor Zurbrigg
To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069,
Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc@achilles.net.
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.
We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward
our mailing costs. PAP Registration Number 9477. Canada
Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca
Copy Editor: Elizabeth Morton
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LIBRA™®
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The Canadian Field-Naturalist{,\veRs'”
Volume 120, Number 3 July-September 2006
An Ethogram Developed on Captive Eastern Coyotes Canis latrans
JONATHAN G. Way!, DEAN-LORENZ M. SZUMYLO?, and Eric G. STRAUSS?
'Boston College, Environmental Studies Program and Lynch School of Education, Higgins Hall, Chestnut Hill, Massachusetts
02467 USA; e-mail: jw9802@yahoo.com
Urban Ecology Institute and Environmental Studies Program, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA
3Boston College, Biology Department and Urban Ecology Institute, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA;
e-mail: strausse@be.edu
4Present address: 64 Cranberry Road, Marstons Mills, Massachusetts 02648 USA; Corresponding author: e-mail: szumylo@
be.edu
Way, Jonathan G., Dean-Lorenz M. Szumylo, and Eric G. Strauss. 2006. An ethogram developed on captive eastern Coyotes
Canis latrans. Canadian Field-Naturalist 120(3): 263-288.
We studied capture Eastern Coyotes (Canis latrans) from 27-585 days of age and compiled an ethogram on them. A total of
72247 15-sec samples were taken, amounting to 301 h of field time varying between 59.4—61.3 h per Coyote. A total of 540
behavioral patterns was observed amongst the 16 behaviour categories ranging from 9 (miscellaneous) to 72 (explore
investigate) action patterns per parent category. The 16 parent categories that we believed best described and appropriately
sorted the behavioural actions were resting, sitting, sitting], sitting2, standing, traveling, explore/investigating, hunting, feed-
ing, infantile, greeting, self play, play initiating, playing, agonistic, and miscellaneous. Exploring accounted for >31% of all of
the behaviours observed with resting and sitting (combined), standing, traveling, and play as categories decreasing in order
of most to least frequent. Despite some omissions in our ethogram and drift associated with its ongoing development, we
believe that the large amount of data collected made it rigorous enough to be a useful guide for the species. We argue that
although future research will no doubt add to and/or modify components of it, its ease of use in the field (in captivity or in
the wild) and it being the first complete ethogram described for the species, make it a useful tool for future researchers.
Key Words: canid, Canis latrans, eastern Coyote, ethogram, behavioural study.
The concept of an ethogram, also called action
system (Makkink 1936), behavioural inventory (Bekoff
1972a, 1978), or behaviour pattern (Scott and Fuller
1965), is a durable tool for analyzing vertebrate be-
haviour and dates back to Jennings (1906), Makkink
(1936), Tinbergen (1959), and Lorenz (1971). Although
there have been direct observation behavioural studies
of Coyotes (Canis latrans) in captivity (Snow 1967;
Silver and Silver 1969: Bekoff 1972a, 1978: Parks
1979; Ryon 1986) and in the wild (Camenzind 1978;
Bekoff and Wells 1981; Gese et al. 1996), there has yet
to be a standardized and complete ethogram published
for the species. Except for Goodmann and Klingham-
mer’s (1990) non-peer reviewed but detailed manual
which presents a Wolf (Canis lupus) ethogram and
Scott and Fuller’s (1965) ethogram on Domestic Dogs
(Canis familiaris) (which they compared with literature
on Wolves, Coyotes and Red Foxes [Vulpes vulpes]),
we could not find any other study that described a full
behavioural repertoire of any other of the approximate-
ly 35 canid species (Ewer 1973; Wozencraft 1989).
Most studies examining behavioural data have focused
on specific time periods (e.g., young pups/behaviour-
al development — Bekoff 1972a, 1978, 1989), specific
actions (e.g., social interactions — Fox 1970; Bekoff
1974; Zimen 1982), lumping behaviours into general
categories (Silver and Silver 1969; Parks 1979; Gese
et al. 1996), or describing many behavioural patterns
but providing no summary or chart showing a complete
ethogram (Mech 1970; Lehner 1978). Schleidt et al.
(1984) termed these “partial ethograms” and stated the
importance of publishing complete ethograms on as
many species as possible. In addition, Macdonald et
al. (2000) underscored the significance of developing
ethograms on different species by stating that without
one, meaning is not fixed. As such, ethograms are vital
reference material for assessing a study’s conclusions
(Macdonald et al. 2000).
The objective of this paper is to describe, in detail,
the full range of behavioural acts that we have ob-
served in a captive group of five Eastern Coyotes. This
ethogram is intended to be: (1) reader-friendly in for-
mat and applied and used in future direct observation
behavioural studies of wild and captive Coyotes: and
(2) compared with ethograms of closely related species
(e.g., Wolves, Jackals [Canis sp.] and foxes) and taxa
(e.g., ursids, procyonids, and mustelids: Ewer 1973).
263
Methods
On 12 April 2002, five (2 M; 3 F) of an original lit-
ter of nine (3 M; 6 F) estimated 25 day old (Parks
1979), wild-born sibling Coyote pups were removed
from under a shed in a residential area in Falmouth
(Cape Cod), Massachusetts, and were reared at JGW’s
house (note: the other four pups [1M; 3F] were placed
back underneath the shed for the wild parents to raise).
The five Coyotes taken into captivity were contained
within a 53 m? area both inside and outside of JGW’s
house until 13 May 2002, when the Coyotes were trans-
ferred to a 16 m? quarantine facility for one month at
the Franklin Park Zoo, Boston, Massachusetts. On 13
June 2002 the five Coyotes were moved to their perma-
nent 403 m? public viewable exhibit at the Stone Zoo
located in Stoneham, Massachusetts, and remained
there for the duration of this study until late October
2003 — pup age 585 days). We chose the endpoint of
our study as the last day of data collection prior to the
Coyotes being permanently separated due to intrapack
aggression (Late and Trans [See Table 1] were re-
moved). The area of their exhibit was described by
Frank (1987) as a suitable size for a long-term study
of canid behaviour. Although the Coyotes were hand-
raised, we made no attempt to interfere with or disci-
pline their activities and therefore gave the Coyotes
free access, at all times, to their exhibit/living facilities.
The Coyotes were provided with puppy milk (Esbilac,
PetAg, Inc., Hampshire, Illinois) via bottle or bowl up
until 15 May and were given access to water and dry
dog chow (commercially available dog foods until
March 2003 then Mazuri exotic canine chow/diet [PMI
Nutrition International, LLC., Brentwood, Missouri]
thereafter) at all times. They were group fed (i.e., all
five at once) 0.75 — 1.6 kg (varying with their age) of
Nebraska Brand chopped frozen canine meat (Central
Nebraska Packing Co., North Platte, Nebraska) mixed
with dog chow on a daily basis and were given frozen
or thawed laboratory rats, guinea pigs and mice (do-
nated from a rodent breeding facility), and/or bones
2-4 times per week. Within their exhibit, the Coyotes
commonly hunted (at least | prey item 2-3 times daily)
and captured (about 1-2 prey items per week) Starlings
(Sturnus vulgaris), House Sparrows (Passer domesti-
cus), Chipmunks (Jamias striatus), Brown/Norway
Rats (Rattus norvegicus), Meadow Voles (Microtus pen-
nsylvanicus), and Gray Squirrels (Sciurus carolinen-
sis), but did not appear to eat much or any of these prey
items. Care and use of animal subjects was approved
by Zoo New England’s Institutional Animal Use and
Care Committee (letter dated 23 January 2002 to JGW),
by Boston College’s Institutional Animal Care and Use
Committee Protocol Number 01-03, and by the Massa-
chusetts Division of Fisheries and Wildlife Permit #052.
02LP.
We developed an ethogram that noted basic behav-
ioural patterns (Scott and Fuller 1965; Bekoff 1972a,
1972b, 1978; Parks 1979). To avoid influencing Coy-
THE CANADIAN FIELD-NATURALIST
Vol. 120
ote behaviour with different observers only JGW inter-
acted with and conducted activity budgets on them. Us-
ing this technique, we acknowledge that we did not
have interobserver reliability as described by Bekoff
(1974) but the large amount of data collected attempt-
ed to offset intraobserver reliability. We used focal indi-
vidual animal sampling (i.e., one Coyote per 30 min
bout of data collection), took instantaneous point or
scan samples (Martin and Bateson 1986; Macdonald
et al. 2000) every 15 sec for 30 min on the target Coy-
ote, recorded the date, time, and weather before each
observation bout took place, and noted important con-
textual information in between each 15 sec sampling
period. By using frequent (i.e., every 15 sec) instanta-
neous samples, we tried to obtain an accurate approxi-
mation from continuous recording (Martin and Bateson
1986). Thus, 120 samples per 30 min behavioural bout
on a Coyote were ideally recovered. However, there
were two reasons why we occasionally did not obtain
120 samples per bout: (1) a Coyote was momentarily
out of sight during a particular 15 sec sampling period
(usually this happened at most, once per 30 min bout);
and (2) we had to stop a sampling bout earlier (e.g.,
rain, darkness, or some kind of disturbance that forced
JGW to abandon an activity budget). We randomly
chose which Coyote to conduct observations on before
entering the field but attempted to evenly sample all
Coyotes (1.e., each Coyote was scored every fifth time)
during the study. We typically recorded behavioural
data about 5 days/week and took between | and 4 (us-
ually 1-2) 30 min behavioural bouts/day during day-
light hours. Behaviours were also recorded on digital
still and videocameras and 35-mm film cameras week-
ly for the duration of the juvenile period of the pups
(Parks 1979) then ad lib after pups reached adulthood
(one yr of age — Bekoff and Jamieson 1975). We con-
ducted most of the behavioural bouts during early to
mid-morning (ca. 07:00-11:00 h) or between late-after-
noon to early-evening hours (ca. 16:00-19:30 h); these
were the times with the fewest number of people around
the zoo (the zoo opened at 10:00 and closed at 16:30-
1800 h depending on the time of year). Because the
Coyotes were habituated to and did not react negative-
ly to JGWs presence, he followed them around the
exhibit similar to the description of Goodall’s (1986)
“follows” of wild Chimpanzees (Pan troglodytes) in
Gombe, Tanzania. JGW made an effort not to influ-
ence the movements and/or behaviour of Coyotes by
minimizing his movement in the exhibit. This was es-
pecially important because the Coyotes would follow
him around the exhibit when he interacted with them
but would generally ignore him when he was stand-
ing erect and writing on a clipboard (J. Way, unpub-
lished data). “Following” was a necessary technique to
use on the Coyotes because there was not a single ob-
servation spot outside the exhibit where the Coyotes
could reliably be seen all of the time (i.e., many times
trees and shrubs concealed the Coyote under observa-
2006
tion). Besides the senior author’s presence, other human
contact was kept to a minimum before and after each
sampling period to avoid humans affecting Coyote
behaviour. Context was a critical variable with respect
to the Coyotes’ response to human behaviour. For ex-
ample, they generally ignored people (unless very loud)
on the public path but would intently watch and/or bark
at staff members that were behind (i.e., the opposite side
of the public path that was off-limits to non-employees)
or approaching their exhibit. Accordingly, zookeepers
did not enter the exhibit to feed the Coyotes when JGW
interacted with them. Despite these precautions there
were no doubt many instances where Coyotes changed
their activity in response to a person’s (public or staff)
presence — sometimes even when a person simply
walked by their exhibit area. Similar instances of Coy-
otes shifting their behaviour because of the presence
of people have been documented in areas where wild
Coyotes inhabit urbanized areas (Way 2001). We at-
tempted to mitigate these factors by increasing the total
size of the pool of sample bouts.
We entered all of the data into an excel spreadsheet.
First, we entered the raw data into the spreadsheet.
Then we alphabetized that data in order to group sim-
ilar behaviours for each budget. Next we summed the
frequency of each distinct behaviour observed. Finally
we entered those summed values into a separate file
(for each Coyote) that had our developing ethogram.
Due to the large amount of data and behaviour se-
quences in our ethogram (Table 1), properly entering
and compiling data were critical parts of this ethogram’s
creation. Behavioural actions or patterns (denoted as
subcategories in Table 1), as defined by Scott and Ful-
ler (1965), were grouped into “parent” categories, or
behavioural complexes as described by Schleidt et al.
(1984), or behavioural systems, as noted by Scott and
Fuller (1965), based on motivational context and activ-
ity observed (Parks 1979).
In our ethogram, we organized the categories and
the behavioural actions within each of the complexes
generally on a scale from less to more active or intense
an activity, then from solitary to group-oriented activ-
ities. Notable exceptions were feeding, which was
grouped next to hunting for obvious reasons, and self
play, which was grouped close to social play but was
of a solitary endeavor. The 16 parent categories that we
believed best described and appropriately sorted the
behavioural actions were resting, sitting, sitting], sit-
ting2, standing, traveling, explore/investigating, hunt-
ing, feeding, infantile, greeting, self play, play initiat-
ing, playing, agonistic, and miscellaneous. There were
instances (especially early in our study when we were
working on differentiating different behavioural ac-
tions) where a Coyote was engaged in an activity but
we could not describe, other than in a general category,
what the Coyote was doing. When this occurred (e.g.,
when a Coyote was behind trees and was only par-
tially observed) we simply classified the behaviour in
its parent category, so that we could at least provide a
Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES
265
coarse description of a Coyote’s behaviour al a given
15 sec interval (Table 1).
We calculated the percentage that each behaviour
pattern was observed and added all within a parent cat-
egory to examine the relative frequency that each behavy-
ioural complex/system occurred (Table 1). In addition,
we used the ages (in days) of the Coyotes to denote
the first time that each behaviour was observed, with
day 27 (14 April 2002) being the beginning point —i.c.,
the date we first did an ethogram on a Coyote. We want
to stress here that although we report both the percent-
age of each behaviour observed and the age that each
behaviour was first recorded for comparison purposes
(Table 1), the purpose of this manuscript is to describe
the ethogram that we developed. We will more thor-
oughly analyze these data in future manuscripts.
Results and Discussion
The ethogram that we developed on the Eastern
Coyote is presented as Table 1. A total of 72247 15-sec
samples were taken amounting to 301 h of field time,
varying between 59.4-61.3 h per Coyote. A total of
540 behavioural patterns were observed amongst the
16 behaviour categories ranging from 9 (Miscellaneous)
to 72 (Explore and Investigate) acts per parent category
(Table 1; Figures 1-25).
Resting (RE) occurred when a Coyote was lying
down with its head on the ground. Sitting (SIT) was
classified as sitting down either on rear or all four legs;
this category was subsequently divided into sit] (SIT1)
and 2 (SIT2). SIT1 was noted when a Coyote was sitting
down on all four legs; in other words, it was almost ly-
ing down but it had its head up. SIT2 was when a Coy-
ote sat on its rear legs but its front legs were up in
standing-like position. It was difficult to always reli-
ably determine what the motivation was for a Coyote
in the aforementioned positions. For example, a Coy-
ote frequently sat in SIT1 position to sniff the ground.
Because the animal was in a resting-like position we
believed that it was important to separate these activi-
ties from explore/investigate activities where the ani-
mal was active (1.e., at least in a standing position).
Similarly, it was sometimes difficult to understand the
motivational context when an animal was just stand-
ing (ST) and performing routine activities. Thus, we
developed a category for just standing activities.
Any of these categories could always be lumped
together or put into different or new categories if the
need arose — but the opposite cannot occur if the be-
haviours are not split into discrete actions at the outset
(Bekoff 1972b). For example, SIT1 Alt and SIT1 Alt Pt
were very similar except for the panting involved. Al-
though we could have easily combined these categories
we thought that keeping them separated would allow
us to potentially analyze this difference in the future
(e.g., the level of panting on hot versus cold days).
Resting and sitting behaviours were observed quite
frequently and accounted for >23% of all of the etho-
gram despite the Coyotes generally being much more
Vol. 120
THE CANADIAN FIELD-NATURALIST
266
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SFO T80 OF0 110 600 09 79 +9 79 G&L 960 09 BUNS o[IYM Joolgo ue UO SUIMAYD [qo Moy)
r00 910 LOO SOO OO I8 OL +9 O8 62 800 +9 SUIS BITYM Joalgo ue sung [qo ag
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000 000 100 000 000 0 0 r9 0 O 000 8369 J[as SuPyor] pure yoye
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000 100 000 000 000 0 s8 0 0 0 000 =s8 Ayfeoytoads SuryjAue uo Sursnd0} Jou suns a[TYM punose SuryooT] Ie yoo]
000 100 000 000 000 0 242 0 0 0 000 LL J[a8 Suryory] pue suyyiug MOI] 2 Us
100 SOO 000 100 00 6F +9 O I91 SL 700 3066 BUNS OLY J[as SULIT J[98 YOUT
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100 100 100 +00 600 SsI III OSI El $8 £00 8 oye a[rYyM (are ay} Ayjensn) suyyrug ye 2 us
Vol. 120
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THE CANADIAN FIELD-NATURALIST
CIN WE) GSO VN) TE) LG AG CO Sor x5 L990 ©96 SUNIS I[IYM SULYIOWIOS UO posndo04 WV
100 100 100 000 000 811 Lrl 6€€ 0 OO 000 8tt SUIS ofA (uado soso) SuryJAue UO pasndoy jou Jnq Suu (id) queg
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€10 €00 I10 €00 €00 OIF OIr 801 P9r O6r LO0 ~=801 surued a[tym oaoge sv ouieg jd Ie yoo]
a GEO 120 C0 CW Ge Sh wh Gs Gs 6S0 6b SSUIY) JUDIOJJIP 298 0} pray SurAow ‘joalqo Aue uO pasndoy JON Ie YOoT
OW OOO Cod COO ao @ © © @ 6s 000 6¢ BUNS OLY YINOW YIM SuIyJouWOs Sury[Ng {qo [nd
000 000 000 000 100 0 0 0 0 80r 000 80r SUIS O[IYM SUTyJOWOS Je SuIMed pure suYsIUS [qo med 2 ug
000 100 700 100 100 0 £66 96 SOI FII 100 96 SUNS B[IYM punoss ye SIp 10 Meg SIp/Med
ZO;0 400 €00 100 €00 LZ 08 96 9€S 66 €00 6S SUIS OILY (JOM JI '3"9) Peay SuLyeys ayeys peoy
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100 000 000 000 100 ss 0 0 0 000 6P uortsod [Is UL J[as SULYST] pue Suryoyesog —_ FAS Yor] yoe.1og
00 900 600 S00 SOO €rl €6 OIL OO ZI 900 £6 uor}isod [}Is Ul Jas Suryoye.19g J[9s yoqesog
00:0 000 100 000 000 0 O ist O O01) “ZEI UOTISOd [IIS Ul B[IYM SUIJOWIOS SUIMEYO puk Y}aNgS [qo mayo yoIeng
OO 100 COO COO COO M Cd O @ 7 000 §=€ZI suruMed YIM aAoge sv ouleg —- UMBA 29 YO}ANS
OOO 1700 100 100) 100 @ Ser Sil Gus eel GO irk uontsod [Is Ur afIyM BUIYo}ONS yoang
000 000 000 400 000 0 OO 0 OO! 0 100 001 (SBIMJ/SsvIs °3"9) SUNS O[IYM SUIYJOWIOS SuNIg ust) SUYyIUS aq 2 Ug
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p00 600 STO 100 910 6S £€6 6S 801 96 600 6S SUIS ay Joalgo uv sunrg [go og
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100 000 €00 000 100 9€1 0 Gi O Ws 100 Sol Woye (TS) SUNS a[TYM SuIyJOWIOS SUIMaYD [qo mayo VW
OFT GEC IS Gi pil Ger LL Gr Be 6S WE = Sy joafgo ue SulmayD [qo May)
000 000 100 100 100 0 0 6€€ 16h LOS 100 6€€ SuruMed jou yng yNou sutusdO yjnow usdO
KOO CO) COO COO lO @ ~—© © @ ei 000 §8LEI joafgo ue Suryor] pur suyyius YOM 3 US
OOO 100 COO OOO OO) © Wr OM 0 0 000 9r Sunued oplym Jas Suryory id Jas Yor]
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100 100 000 100 7200 9SI €91 0 9€I SEI 100 = SEI SUIYJOWOS UO pasnooj a]IYM (Are A][eNsn) SuryJowlos SuYyIUS US 2 WV
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4
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100 000 000 000 OOO Lb O 0 0 0 000 L6z Sunued pur woe Surpurys opyM ysry prey, dn jrey id apy
100 soo OOO 100 100 sds Its 0 Lt Lt cOO 806Le dunued apy aaoge se aug id jie) dem yy
90:0 670 #00 OTO FIO OE FS 87S OLI ILI p10 80S Sa]0A0D J9yIO punose
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té60 TO $60 S60 TSO 6 COL OO! 6 PFII SE0- 6 dunued apiym Loy jued jy
SSS O0S2 80S S285 WSS 6E& ve Ov LE &E OL vé SUIYJOWIOS UO SuIsndo4 Qe) wary
roo 100 100 100 000 +6 OI 8Ol Ost 0 100 = t6 sutyiAue UO Sulsndoy Ajjvad ou ing sunurd (ad) jurg
100 100 100 100 O00 S82 IeS 196 6Fl O 100 82 SuIpurys siyM Surysnoy ysnoy
000 000 000 100 000 O 0 OAs (0) 000 LEE JUDWA}I9XO SutAduut fre} BursseM pur SULUMEA [ie] BRM UME
100 000 000 100 O00 sss 0 0 LSE 0 000 LIE SuIpuR}S pure LOTR O[IYM SUIUMEA ye UME,
900 F000 900 SIO 900 Ft FLI LLI LOT TLZ L400. =I SUIPURs JIIYM SUIUMPA UMP,
co0 000 000 000 000 10s 0 0 0 0 loo ~—-10€ Sulpurys ap1yM soko Suryury¢ yun
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100 000 000 000 000 I10€ O 0 0 0 00°0 10€ uonisod Suipur)s vB Ul a[IYM SUIZOG aZ0C
SO SFO trO £0 EFO Cf PE PvE HE €E TPO = FE SAMOSAILS JOYIO OFUT IY JOU PfNod Jey} sIOLAvYaq SurUUOJ1od pur surpurig (LS) Surpurig
be Sti Set G60. P30 pel ‘ZS IO} adRJUsdIAd [RIO], GANAS
000 100 000 000 000 0 61S 0 0 0 000 61s uontsod Zils ur afiym SuIssip pur suysiug 3Ip US
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000 000 SOO 000 000 0 OO 6% 0 0 100 6b uontsod Zs Ur SUT|MOH [MOH
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r00 €10 910 100 S00 66 OZI OL 88€ 9FI 800 66 SUIS BIYM SuryJoWOs UO SuIpady AT[ROI JOU Ing SuIMeYyD [qo Mey)
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000 000 000 000 wo OO 0 0 oO ZOI 00;0 #8 ZOl uado safa yng SuryjAue UO pasndoj jOU pure SUIIIIS uado |
000 100 000 000 000 O wc, S055 0 0 000 FIZ uonisod Zs ul LoTe a[IyM (Are Ayensn) Suryjouros Suyyrus us 1V
£00 400 900 £00 sO0 46 901 LOF EIS OSI soo 8 86L6 sunued ary aaoge se sues id atv
THE CANADIAN FIELD-NATURALIST Vol. 120
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SOO PEDO OOO NOOO SOO OM SLI 010 9TI 2.0 Oe Oo Ssurpurys opty dn Mosyy/SurspNAUOD WULOA,
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000 000 100 100 070 0 0 Or SOS (LIKE 700 LIE SUTYJOWOS UO PasNoo} a[IYM dAoge sv owWeG 4B SBYIO 1OAQ
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100 100 000 000 000 8t€ 666 0 OO 0 000 8€€ Surpueys aptyM YNoW wory Joolqo uv SuIsva[ay yoalqo doiq
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OOO C0) COW COO OOO gal © © 9 0) 000 ri sunestsaaut AT[eor Jou :A[poyiwougyey SuryJOWOS Je SuIME yoalqo Med
700 €00 900 LOO €00 6rl S6% 6%b 87S LZ 00 = 6rl yoou pure peay oy} isnf ayeYs 10 ySTAMT, peoy ayeysg
AO ClO UO WhO SIO Gs Sz Gn ms 3s Glo LE J[98 JJO Isnp 393 10 Arp 0} — Apog suryeys Apog axeys
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CGO OGOR USO G0i0e G00) [8= 16. “El ESi- 9eI 810.—s*CT8 uontsod Surpurys ut (20.3 ‘aouay) joofgo ue jsulese Apog surqqny Apog qny
TOO) OY) WO) COW Onn 0 Me 0 0) 000 LOE SUIPUL)S O[IYM Olu OWS Je SuTeULIN-Jenbs pure suryojang 9JPULIN YOON
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GOO CeO SO TOO Mo cy Wh yer OSL Gs 710 66 Surpueys opty Apog Suryojang Yyo}aNg
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000 000 100 100 100 0 0 (GON (KG AE 100 I Loye STI 9Aoge sv sues ye may
GSC EXC CEC GEG Wee inp 8S ip hip ip YC — TAP SUIpUR}S d[IYM Jago Ue SUIMEYD [qo MayD
b10 0€0 €00 £00 900 €ZI 981 86 9SI S9 LEO, “S9 BUIpUe}S J]IYM Jogo Ue SUTYOTT yoalqo Yor]
€00 100 100 100 100 S87 66% ZL 66€ LZS 100 ZL MOrAvyaq o[HUPJUT JOU Os SUTIOOIS Jou ng AOAOD Jotjour suryory Ayog KOO JOT
ISO (OO £70 GO S/O SS OL 8) @) ‘Wil 090 ss J[es SuPyor] SurUOOIDH F128 YT
000 000 000 100 000 0 0 0 O8€ 0 000 O8€ — parejas AyjeIs0s ‘snorxue sreadde a0A0D ‘]1e} SuIsSeM aI OAoge sv oureg — [Iz] SBM OUTYM ITV
suviy, odnqy ayey uorwd curd suviy odny ayy uorwg soured CIV SIV
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(panuyuor) sajoko_ Uia\see UO WeIsOUY ‘| ATAVL
271
4THOGRAM ON CAPTIVE EASTERN COYOTES
Way, SZUMYLO, and STRAUSS: AN E
2006
ooo OOO 100 000 O00 O 0 SIs 0 0 00';O ~=8ts SuLypeM [IM Sunroayac] aRIOJOP YA
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moo £00 £00 TO 100 TWH 6Ol te 90S Its c0'0 pt SUPE JIU UPUINY 10 aJOAOD kB Buryoroiddy 1M Vv
£00 100 100 FOO 100 897 O8l C7S Lel O6t wOO 83 LI SuruMmevd pur SULyTeA uMvd 1A
000 OOO 000 100 000 0 O O Sb O 000 = Sb suryfe opty surduary dunt] 4A
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000 OOO TOO 900 000 O 0 98e TIE O ZOO 3 Z9E S9}0A0F5 aaAtsstuigns Aq auop {Moy Apog IM SUTRA, MO] Apog AW
100 100 TOO 000 100 LOZ S6Z. TEI 0 F6z 100 = ZEl Jam Udy auOp Ajjensn :Xpog SuLjeys puv duryjeA — 98 O4PYS AM
ooo 000 OOO 100 000 0 O O- E£LE 0 000 €LE MOKOD & Ie pajdastp Jou Ing paytoxe Asa :Apog Surddem pur SuryyeA Apog SBM 4M
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000 100 000 000 000 0 Sel Ge: OF: 0 000 §€€l SUIYIONS Pur SULYyLPAA YIO.S YA
000 080 100 000 000 O 0 91€ 0 0 000 9IE (yoursg *aouay) algo Isuresde Apog surqqns puv SuryyeAA —MA/OYBAOS YMA
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100 £00 L400 TOO +00 trl Chl Cel EMI CCl £00 CTI SUIYJOWIOS UO SUIMOYO puR SULYTEAA [go Mayo AA
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000 000 100 000 000 O 0 19s 0 0 000 198 paonpo.d punos ou jnq UOHOUL ay!]-[MoY B SuUPyeU pur suryTeA [MOY OARS AM
000 000 100 100 000 0 OO - 6% 87S 0 000 6t>P suljMmoy dnoss pue sux YM 2 13 [MOH
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THE CANADIAN FIELD-NATURALIST
276
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THE CANADIAN FIELD-NATURALIST
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282
active in JGW’s presence than other times of the day
(Stone Zoo Keepers, personal communication).
Traveling T was defined as a Coyote moving with
purpose from one place to another (i.e., from point A
to point B). A Coyote, when traveling, would move in
a straight line and would waste little energy investigat-
ing other objects until it reached its destination. This
differed from explore/investigate E/I activities, where
animals were characterized by actively sniffing objects
or other Coyotes, digging, and moving around in a
non-direct way in order to explore their surroundings.
E/I behaviour is typically not described in the litera-
ture for canids (but see Scott and Fuller 1965) yet was
an important activity that our Coyotes engaged in, by
far the most frequently of any parent category (Table 1;
32% of activity). The difference between explore/inves-
tigate and traveling activities would be interesting to
compare between wild and captive canids. The captive
Coyotes that we studied were very active but did more
exploring then traveling because of the relatively small
area that they lived in. Conversely, Coyotes in the wild
routinely have to traverse their home ranges (Camen-
zind 1978; Gese et al. 1996; Way 2000) and potential-
ly might do more traveling than exploring.
Hunting (HU) behaviour was classified when a Coy-
ote was actively trying to catch/kill another object. Many
small mammals and birds entered their enclosure, giv-
ing the Coyotes the opportunity to hunt (although we
occasionally gave them live chicken chicks, we never
scored the Coyotes when they were killing them). Ad-
ditionally, when a Coyote was attempting to ambush a
conspecific, we also classified this as hunting because
its intentions were very similar to hunting prey: L.e.,
body very alert and focused, in a crouched position,
and rushing at its intended target. Furthermore, differ-
ent results occurred after an ambush; e.g., no social inter-
actions occurred (i.e., both individuals walked away
from one another), a play initiation was given once the
ambusher got within close range of the Coyote, play
immediately started, or fighting or agonistic behaviour
ensued. Parks (1979) classified “ambushing” as a sep-
arate category in her thesis but perhaps the larger size
of the exhibit in which we studied our Coyotes allowed
us to detect the similarity to hunting.
Feeding was simply associated with ingesting some-
thing. Although we documented our Coyotes feeding
(FE) we attempted to avoid conducting behaviours
around any type of feeding time (e.g., we did not do a
behavioural bout when dog chow was just placed out
or when meat, rats or mice were just given to them)
to avoid biasing the data.
Infantile (IN) behaviours were associated with neo-
natal contact as described by Parks (1979), but we
also included licking/grooming one another. Because
the Coyotes were in contact with other Coyotes, we
believed that this was an important division for a par-
ent category even though other behavioural complex-
es, such as resting, sit] and sit 2, are included in this
category. These behaviours were intermittently seen
THE CANADIAN FIELD-NATURALIST
Vol. 120
me : 2 bt S
FiGurE 1. JGW (first author) conducting a focal animal sam-
pling bout on the Coyote to the right of him.
nt
FIGURES 2 and 3. Coyotes sleeping in rest position.
FiGure 4. Coyote in Sit! Alert position.
2006 Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES
FIGURE 9. Coyote in Stand Alert position.
FIGURE 10. Coyote Standing and Howlin
.¢ \ipee : % rs
p in sit2 Alert position in JGW’s house. FiGuRE 11. Coyotes Standing and Group Howling.
<—" Se
Sis < ut .
Rud
FiGuRE 8. Coyote on left in Sit] Alert Pant position; Coyote © FIGURE 12. Coyote on the right and in the back are Traveling
on right in Sit2 Alert position. and Trot Following the Coyote in the front.
284 THE CANADIAN FIELD-NATURALIST Vol. 120
Signi i pe is bs E 3 : Wrle . she
US core "9 adi naar ac == FIGURE 16. Coyote in Hunting mode, chasing and outing a
4 bird.
FiGuRE 15. Coyote in foreground in Hunt Ambush Crouch
(Sitl) position. Coyote in background in Ambush-
rush Conspecific position. This behavior can quickly
turn into an Approach (Travel trot towards) when
the Coyote is not in a crouched hunting mode and is
moving towards another Coyote.
FIGURE 18. Coyote Greeting Person (JGW).
2006 Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES 285
€
FIGURE 19. Coyote in a true Greeting with JGW.
FIGURE 24. Coyote trying to Play Initiate with JGW by Lying
, on back and Pawing.
FiGurE 25. Agonistic display. Coyote on top is in an Agonistic
Dominance Display while the Coyote pinned to the
ground is in Agonistic SuP (passive submission).
throughout the 18 months of observation when Coy-
otes would occasionally sleep on one another (espe-
cially on rainy days) or one Coyote would lick anoth-
er but was not greeting it. A licking Coyote appeared
to be grooming the recipient, which made it hard to
place into any other category (i.e., it was doing more
FIGURE 22. Coyotes Play Group Wrestling. than simply greeting another Coyote).
286
Greetings (GR) were classified when non-agonistic
and/or non-play interactions occurred between =2 Coy-
otes. Zimen (1982) classified many of these behaviours
as neutral social contacts but we separated Zimen’s
descriptions of sniffing other Coyotes and left them
in the explore/investigate category because the motiva-
tion was not always clear (e.g., was sniffing another
Coyote a social contact or just a Coyote curious to
smell another one?). Simply put, when both Coyotes
licked and/or had muzzle contact with each other and/
or wagged their tails in excitement to see one another,
then we classified this as a greeting.
We used Bekoff’s (1972a,b) definition of “play” to
define our category, where play was the behaviour that
was performed during social interactions in which there
was a decrease in social distance between the interac-
tants, and no evidence of social investigation or of ago-
nistic or passive-submissive behaviours on the part of
the members playing, although these actions may occur
as derived acts during play (e.g., passive submission
during a play bout). Bekoff (1974) also classified play
as: (1) incorporating various contexts into unpredictable
temporal sequences; (2) preceded by a metacommu-
nicative signal (which we have a separate category for:
i.e., play intitiation); (3) certain actions are repeated
and performed in an exaggerated manner; and (4) the
activity appears pleasurable to the participants: a play
face (wide open jaws and eyes) is apparent.
We also included the numerous role reversals in this
category where more dominant Coyotes allowed lower
ranked Coyotes to pin them to the ground; this was
never documented during agonistic displays. Bekoff
(1974) includes a description of this occurring amongst
western Coyotes but does not delineate it as part of clas-
sifying play. Additionally, we separated play initiation
from play because the attempter was not always suc-
cessful in initiating play during an attempt; 1.e., they
appeared to be two separate categories. We observed
many play initiation-like (e.g., approach/withdraw and
general movements) behaviours occurring during play,
but because they happened during play we kept them
in this category; i.e., play was already initiated. Self
play (PS) was similar to play but when a Coyote was
by itself. Altogether, play-like sequences accounted for
ca. 10% of all observations (Table 1).
Agonistic (AG) behaviours were classified as aggres-
sive acts associated with conflict where there were clear
acts of dominance and submission or where growling
or fighting was observed (Fox 1969; Scott and Fuller
1965; Zimen 1982). About 2% of all activities were
aggressive, especially towards the end of the study
when Coyotes were separated. Finally, a miscellaneous
(MISC) category was one where we could not classify
the behaviour pattern into any of the other parent sys-
tems; these were very infrequent (Table 1).
Despite the large number of behavioural patterns
described herein, there are some important omissions
in our ethogram. First, because the Coyotes were all
siblings and were young adults (18 months) when the
THE CANADIAN FIELD-NATURALIST
Vol. 120
study ended, we did not notice any sexual/reproductive
behaviours (although it was witnessed during their sec-
ond winter: J. Way, unpublished data). Future etho-
grams should add this behaviour as a separate (par-
ent) category much as Scott and Fuller (1965), Zimen
(1982), and Goodmann and Klinghammer (1990) did
with their respective ethograms. Second, because the
Coyotes were taken from the wild at 25 days of age,
we missed many basic neonatal behaviours, such as
crawling and nursing (Scott and Fuller 1965; Scott
1967; Parks 1979). Third, because the Coyotes were all
siblings and of the same age, we could not document
any adult-pup interactions such as epimeletic behay-
iours, i.e., the giving of care (Scott and Fuller 1965).
However, many times we did document et-epimeletic
behaviours where the Coyotes were soliciting or call-
ing for care where they would whine, wag their tails
for a greeting (with JGW and/or another Coyote), and
lick one another. Fourth, the full range of hunting be-
haviours (e.g., killing/biting prey) was not documented
mainly because of the rarity of observing a kill — espe-
cially during the 15 sec intervals when we recorded
data. Finally, there were some different behavioural
actions that we could have added but simply did not
because of the sheer volume of our ethogram. For ex-
ample, under infantile behaviour we have sit] and sit2
although we do not note if their eyes are open or closed.
Because both examples were rarely observed, this was
a minor detail in the overall scheme of Table 1, but it
does illustrate the arbitrarily defined nature of any
ethogram (Schleidt et al. 1984), no matter how simple
(i.e., the parent categories) or complex (i.e., the behav-
ioural actions) it is. Also, we thought that sit] and sit2
were logical separations due to the noticeably different
body positions; however, we theoretically could have
had many more sitting positions based on a combina-
tion of head, body, and leg positions.
Although we attempted to be consistent with our
behavioural classifications, there was no doubt that
drift occurred in our categorizations (MacDonald et al.
2001). For example, many of the behaviours occurred
well before first noted on Table 1. Although usually
very similar to other categories (for example E/I wk
sn and E/I st sn versus E/I step sn; Table 1) a new cat-
egory randomly created during the study certainly
introduces a bias to the finally tally of percentages in
Table 1. However, the goal of this study was to pro-
duce an end product (i.e., the ethogram) even at the
expense of being consistent throughout — 1.e., there
was no available model of a typical ethogram on wild
canids. Also, a number of simple, but newly recorded,
behaviours occurred after the study concluded. For
example, E/I step sniff became much more frequent
no doubt because of the new category described not
because a Coyote just started doing that action pattern.
Also, stand step alert and stand alert move head were
described soon after this ethogram was finalized (J.
Way, unpublished data). Although these were com-
monly recorded after the Coyotes were separated they
2006
were not described herein. That being said, we believe
that the ethogram is largely complete and that these
slight nuances are offset by our rationale of produc-
ing the ethogram when all five Coyotes were housed
together (i.e., before they were permanently separat-
ed) and the fact that the main categories would stay
virtually identical even with these changes.
The categories that we created are intended for ease
of use in the field where a researcher can simply note
the major behavioural system observed and the behay-
ioural pattern associated with it. One advantage of this
ethogram is that it is in discrete units (i.e., the behav-
ioural patterns). However, data could always be lumped
together (i.e., into general behavioural systems or =2
behavioural patterns could potentially be combined
into one action) if the data/list are unmanageably large
(e.g., when studying wild Coyotes). The disadvantage
of merging behaviours into more broad groupings is
that parent categories or lumped behavioural patterns
can never be split back into more distinct behavioural
patterns if not done at the outset; as Bekoff (1972b)
indicated, this would cause data to be irreversibly lost.
The previous studies that have described canid be-
haviours were quite variable (Scott and Fuller 1965;
Silver and Silver 1969; Bekoff 1972a; Parks 1979:
Zimen 1982) so we basically combined data/categories
from all of these studies and made modifications or
additions as needed in order to try to create an accurate
yet practical guide for researchers studying canids in
the field. Future research should try to generate similar
ethograms for other species and should continue to
modify and/or expand our list if new behavioural ac-
tions are found with Coyotes. Analyzing ethograms of
closely related species may be one technique to effec-
tively discriminate between them (e.g., Western Coy-
otes, Eastern Coyotes and Wolves — Silver and Silver
1969; Bekoff 1978).
Acknowledgments
This study would not have been possible without the
support from the staff at Zoo New England, including
the keepers, management, veterinarians and their tech-
nicians, and security. In-kind benefits from the Way
family were invaluable. Peter Auger and the Biology
Department and Urban Ecology Institute at Boston
College provided support. Katie-Jo Glover and Kelly
Holland assisted with data entry. The Town of Fal-
mouth’s Animal Control and Department of Natural
Resources were instrumental in helping JGW obtain
the wild-born pups.
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First Observations of an Eastern Screech-Owl, Megascops asio,
Population in an Apple-Producing Region of Southern Quebec
NGAIo L. RICHARDS!, PIERRE MINEAU2, DAVID M. Birp*, PIERRE WERY*, JACQUES LARIVEE®, and
JASON DUFFE®
‘Anglia Ruskin University, East Road, Cambridge CB1 IPT United Kingdom; e-mail: n.richards@anglia.ac.uk
Canadian Wildlife Service / National Wildlife Research Centre, Environment Canada, 2 Raven Road, Carleton University,
Ottawa, Ontario KIA OH3 Canada
‘Avian Science and Conservation Centre, Department of Natural Resource Science, McGill University, Macdonald Campus, 21,
111 Lakeshore Road, Ste-Anne-de-Bellevue, Québec H9X 3V9 Canada
4Parc national du Mont-Saint-Bruno, 330 rang des 25 Est, Saint-Bruno-de-Montarville, Québec J3V 4P6 Canada
Etude des populations d’ oiseaux du Québec, 194 Ouellet Street, Rimouski, Québec GSL 4R5 Canada
°Cégep de Rimouski, 60 Evéché ouest, Rimouski, Québec GSL 4H6 Canada
Richards, Ngaio L., Pierre Mineau, David M. Bird, Pierre Wery, Jacques Larivée, and Jason Duffe. 2006. First observations
of an Eastern Screech-Owl, Megascops asio, population in an apple-producing region of southern Quebec. Canadian
Field-Naturalist 120(3): 289-297.
Baseline information was collected on a local Eastern Screech-Owl population found in the apple-producing region of
Saint-Hilaire and Rougemont, Quebec, as part of a larger study on pesticide exposure conducted between 2000 and 2003.
Screech-Owls visited or occupied 41 of 89 nest boxes installed in 12 orchards and 2 control locations. The mean height of
occupied nest boxes was 3.83 m (2.00 — 5.80 m). Squirrels, Sciurus and Tamiasciurius sp., and chipmunks, Tamias striatus,
Northern Flickers, Colaptes auratus, and wasps were the owls’ primary competitors for the boxes. Intact Screech-Ow] pel-
lets retrieved from nest boxes (1 = 82) had a mean length and width of 3.57 and 1.44 cm, respectively, and weighed a mean
of 1.77 g. Screech-Owls in the study area consumed a variety of small mammal, avian, insect and aquatic prey. Of these,
Meadow Voles, Microtus pennsylvanicus, were the predominant prey item identified in pellets, and Mourning Doves, Zenai-
da macroura, were the primary avian prey found in nest boxes. In Canada, the owl’s trend status remains largely unknown.
The species is currently listed as “Not at Risk”, based on an assessment conducted for COSEWIC in 1986. Given that a lim-
ited amount of information exists on the natural history and ecology of the species in Quebec, we also generated a map of
the owl’s distribution in the province, using data from ornithological databases and rehabilitation facilities. Potential risks to
the species within the province, particularly pesticide exposure and habitat loss, are briefly addressed and follow-up studies
are discussed.
Key Words: Eastern Screech-Owl, Megascops asio, natural history, ecology, apple orchard, Quebec, range, distribution, census, pel-
let analysis, prey inventory, nest box, wing chord, population management, pesticide exposure, habitat loss, develop-
ment, conservation.
The Eastern Screech-Owl, Megascops asio, has been
extensively studied in the United States (i.e., VanCamp
and Henny 1975; Gehlbach 1994a). However, it has
received considerably less attention north of the bor-
der, where it occupies a much smaller area. Within
Quebec, the Screech-Owl’s range is vaguely delineated
and general knowledge about the species is limited. As
part of a study assessing the exposure of the Screech-
Owl to pesticides currently and previously used in
apple orchards of southern Quebec (see Richards et
al. 2004, 2005), we collected baseline information on
the species. Here, we present observations on a popu-
lation of Eastern Screech-Owls (Figures 1, 2, 3 that
tesides in one of the primary apple-producing regions
of Quebec.
Methods
Mapping the range of the Eastern Screech-Owl in Quebec
We obtained case files from the Clinique des oiseaux
de proie (COP), based in Saint-Hyacinthe, and the Cen-
ter for the Rehabilitation of Injured Birds of the Mon-
térégie (CRIB), based in Otterburn Park, which admin-
ister care to the majority of injured or orphaned owls
in the province. The locations from which adult and
juvenile Screech-Owls were admitted were tabulated
and mapped. These data harbour certain biases and
so were used only to illustrate the overall, rather than
density, distribution of the species within Quebec. For
example, the number of owls admitted from an area
is positively correlated with its proximity to the near-
est rehabilitation facility (Richards et al., unpublished
data). Visibility and predisposition to capture can arti-
ficially inflate admission numbers; in this case facili-
ties tend to receive an influx of orphaned Screech-Ow]
babies and fledglings in the late spring and summer,
largely from suburban settings, where they are highly
conspicuous (Richards et al., unpublished data). Thus,
each admission location was given one data point,
and multiple admissions from any given area were
not considered an indication of a large Screech-Ow]
population or that it was especially suitable for the
species.
289
290
We also obtained Screech-Owl locations from the
Etudes des populations d’oiseaux du Québec (EPOQ)
database, which is managed and continually updated by
members of ornithological clubs and societies belong-
ing to the Association québécoise des groupes d’ orni-
thologues (AGQO). Finally, we contacted personnel
from the Ontario Breeding Bird Atlas project (OBBA)
and from the New Brunswick division of Bird Studies
Canada to ensure that our map represented the species’
range at the eastern and western borders of Quebec
as precisely as possible.
Site selection and censusing
All fieldwork was carried out in orchards of Rouge-
mont and Saint-Hilaire (45:28"73°W to 4532"73°N),
two adjacent apple-producing regions of southern Que-
bec. The study region was first surveyed in the sum-
mer of 2000 for areas containing habitat requirements
for Screech-Owls; i.e., mature and mixed forest with
old cavity-ridden trees and conifers for shelter, as well
as running water. Potential orchards and control sites
were then censused between November and March
2000 and 2002. By November, most first-year individ-
uals should have dispersed from their natal habitat
and established a territory. Thus, owls that respond are
likely to be established, rather than transient, individ-
uals. This provides a “true sense” of occupancy. We
have found, based on our experience censusing Screech-
Owls in southern Quebec, that this timeframe also
corresponds to the period when individuals are rea-
sonably responsive to broadcast calls.
We walked or snowshoed between the edge of the
orchard and the adjacent forest, broadcasting a com-
bination of “bounce” (territorial) and “whinny* (defen-
sive) calls into the forest from a small, hand-held tape
player. Between 3 and 7 minutes of silence followed,
and then the calls were repeated. If a response was not
obtained, we then proceeded around the perimeter and
repeated the process roughly 10 minutes later. When a
response was obtained, the presence of an individual
was visually confirmed with a headlamp or flashlight
and its colour phase was noted. Two individuals ob-
served in the immediate vicinity of one another (i.e.,
perched in the same tree) were considered a pair. We
also noted responses from Barred Owls, Strix varia,
or Great Horned Owls, Bubo virginianus. The pres-
ence of these potential predators in an area may cause
Screech-Owls to suppress vocalizations (Fuller and
Mosher 1981), restrict the size of their territory, or deter
them from establishing themselves altogether (Gehl-
bach 1994a).
The locations were then re-censused every winter
to verify whether or not they were still occupied. The
12 orchard and 2 control sites where Screech-Owls
were observed in the winter were censused again in
June and July of 2001 and 2002. Given that Screech-
Owls are not always particularly cooperative even
under optimal conditions, locations were repeatedly
censused (at least twice) if a response was not obtained
THE CANADIAN FIELD-NATURALIST
Vol. 120
during the first censusing effort. We did not census
when wind speed exceeded 15 km/h, or during rain or
snow, to ensure that censusing was carried out under
the best possible conditions.
Nest box installation and inspection
Between 2000 and 2002, 89 nest boxes were in-
stalled in the forests where Screech-Owls responded
during our initial censusing efforts. All orchards were
assigned a letter code to maintain orchard-owner con-
fidentiality. Boxes were constructed of pine, Pinus sp.,
or Eastern Hemlock, Tsuga canadensis, with dimen-
sions for American Kestrel, Falco sparverius, as per
Henderson (1992). Material does not appear to influ-
ence nest box selection by Screech-Owls (Gehlbach
1994b). One caveat is that boxes made of hemlock must
be stored under conditions of relatively uniform heat or
they will crack and warp. Pine boxes are heavier and
more challenging to transport but much more resistant
to temperature fluctuations, both inside during storage
and in the field.
Between 1.0 and 2.0 inches of dust-free wood chips
were placed in the bottom of each box. The distances
from the base of the box to the ground and from the
box to the nearest orchard edge were recorded. Nest
box covers were secured on each side with spring-
loaded latches to facilitate inspections while minimiz-
ing entry by mammalian predators, particularly Rac-
coons, Procyon lotor. Though most locations were
fairly remote, boxes installed in areas regularly ac-
cessed by the public were painted with a brown, non-
toxic paint, and placed a greater distance from the
ground to reduce visibility and the possibility of distur-
bance or vandalism (as described in Gehlbach 1994b).
Boxes were initially inspected for pellets and prey
remains 3 to 4 times per year: in the spring, in mid to
late summer, in the late autumn and in the early winter,
depending on ease of accessibility to each site espe-
cially during periods of heavy snowfall. However, we
soon stopped inspecting in the spring and summer be-
cause of the disturbance posed to other species occupy-
ing the boxes; unfortunately, we unwittingly caused
nuthatch, Sitta sp., and European Starling, Sturnus vul-
garis, parents to abandon their nests. From an ethical
and logistical perspective, the late autumn and early
winter were deemed the best times to inspect nest box-
es. Thus, between 2003 and 2004, boxes were inspect-
ed once or twice per year in the late autumn and/or
early winter. When prey items were found, boxes were
emptied and replenished with wood chips to prevent
duplication of results on subsequent visits. Recovered
wood chips were carefully sifted for pellets or prey
items.
Pellets, prey remains and prey inventory
Pellets were air-dried prior to dissection for prey
identification. Length, width and weight of intact pel-
lets were recorded. Small mammal species were iden-
tified by lower mandible (Cahn and Kemp 1930; Rac-
2006
zynski and Ruprecht 1974), using a reference collection
of known skulls and mandibles. Vole species were fur-
ther distinguished as Clethrionomys or Microtus sp.
using Maisonneuve et al. (1997). Avian prey were iden-
tified to species from feather remains with the help of
two local wildlife rehabilitators, and referring to El-
broch and Marks (2001). An inventory was then drawn
up for each of the locations where prey items were
found,
Screech-Owl captures
Screech-Owls were initially captured between June
and August of 2001 and 2002 as part of an unsuccess-
ful attempt to radio-track them and evaluate their use
of orchards in relation to pesticide applications. We set
up a mist net and a bal-chatri trap baited with a white
mouse within orchards but near adjacent forest (as per
permits no. 2001-05-07-103-16-SF and 2002-06-10-
120-16-SF). We then broadcast defensive and territori-
al calls into the forest to attract owls. Capture attempts
were initiated just prior to or soon after dusk.
All captured adults and young were processed on-
site. First, they were placed headfirst into a clean
Pringles® chips canister and weighed with a 300 g
spring-loaded Pesola scale. After the owl was removed
from the canister, the keel region was gently palpated
and a keel index was assigned on a scale of 0 to 5, with
1 being “emaciated”, 3 being of “medium” weight, and
5 being “obese”. The wing chord was also measured.
Owls were aged as adults based on the presence of
blood feathers and on other evidence of moulting. It
was not possible to sex individuals. All owls were band-
ed and released.
Results
Range map
One hundred and sixty-nine locations were plotted:
20 from the COP (records spanning 1998 to 2001),
17 from the CRIB (records spanning 1988 to 2003),
7 from the OBBA (2001 to 2004 data) and 125 from
the EPOQ database (1940 to 2004). According to the
personnel at the New Brunswick division of Bird Stud-
ies Canada, the species is incidental at best in the Mar-
itime provinces, and they were unable to provide us
with definitive sightings.
Censusing
Between the winters of 2000 and 2002, we censused
27 orchards and 3 potential control sites. We received
a response from Screech-Owls in 12 orchards and 2
control sites, one a wooded area near a trailer camp-
ground and the other in the research sector of the Mont-
Saint-Hilaire Biosphere Reserve. Four pairs and 7 indi-
viduals were observed during censusing and all were
grey-phased.
Great Horned Owls or Barred Owls were heard or
seen in the vicinity of 6 of the 14 locations occupied
by Screech-Owls and 6 of the 16 remaining locations.
It is likely that some or all of the locations where we
did not obtain a response are frequented by one or both
RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC 29)
ppg //~ Brunswick ~ /
FIGURE 1. Distribution of the Eastern Screech-Owl (Meegas-
cops asio) in Quebec.
species; the habitat observed is suitable and they are
all either quite close to the Mont-Saint-Hilaire Bios-
phere Reserve or to Mont Rougemont, both of which
have Great Horned Owl and Barred Ow! populations
(C. Cormier, personal communication 2003).
Nest box occupation
A nest box was considered occupied if pellets, prey
remains or an owl were observed in it during inspec-
tions (Table 1). To determine whether height of nest
box or distance from box to nearest habitat/forest edge
influenced its selection or use, we first considered the
height and distance of those boxes occupied ar least
once by a Screech-Owl. Then, we further refined our
analysis to nest boxes for which we had evidence of
multiple uses, defined as three visits or more (Table
2): either evidence of occupancy on at least three sep-
arate nest box inspections or discovery of at least three
pellets or prey remains, given that Screech-Owls gen-
erally regurgitate one pellet per resting period.
We measured the distance from the tree on which
nest boxes were installed to the nearest “true’ edge”,
an important distinction as a network of paths criss-
crossed most of the forests in which boxes were in-
stalled. True edge was defined as an expanse of open
area immediately adjacent to, rather than contained
within, the forest itself, such as a field or an orchard.
Box distance ranged from 0 to 68.0 m from edge.
TABLE 1. General nest box occupancy of Eastern Screech-
Owls, by location, in Saint-Hilaire and Rougemont, Quebec
areas (2000 — 2004).
Location Number of Percent
boxes installed occupation
Campground 9 89
Biosphere Reserve 6 100
A/AI 7 14
B 10 30
K 8 25
Cc 5) 20
D/F 10 100
G 7 14
I 11 0
M 7 29
N/N1 9 78
Total 89
A variety of other species occupied the nest boxes
(see Table 3). To determine the overall frequency of
nest box use, we evaluated the total number of times
boxes were occupied by Screech-Owls at each site over
the course of the study relative to other species. Table
4 shows occupation by the most commonly observed
species in relation to that by Screech-Owls.
Pellets and prey remains
Nest boxes were installed at the 12 orchards and 2
control locations where Screech-Owls responded to
censusing. However, in some of these locations, there
was little or no evidence (1.e., pellets or prey remains)
that Screech-Owls ever occupied these boxes. Only
one pellet was found at Orchards A/A1, C and M, so
these were excluded from the prey inventory analysis
(Table 5). Intact pellets were retrieved from 8 of the
orchards, including C and M, which were included in
the analysis of pellet dimensions.
TABLE 3. Nest box occupancy by other species.
Mammal occupant
Eastern chipmunk
(Tamius striatus)
Southern Flying squirrel
(Glaucomys volans)
Grey squirrel
(Sciurus carolinensis)
THE CANADIAN FIELD-NATURALIST
Avian occupants
Great Crested Flycatcher
(Myiarchus crinitus)
Downy Woodpecker
(Picoides pubescens)
European Starling
(Sturnus vulgaris)
Vol. 120
TABLE 2. Mean height from ground of nest boxes (n = 41)
occupied by Eastern Screech-Owls in Saint-Hilaire and
Rougemont areas, Quebec (2000 — 2004).
Height from
ground (m) Range
Unoccupied boxes (n = 48) 4.22 2.40 — 6.82
Occupied boxes (n = 41) 3.83 2.00 — 5.80
“High-use”’ nest boxes (n = 33) 3.67 2.00 — 4.72
Screech-Owl captures
Despite repeated censusing, a response was never
obtained in Orchards G or M. These locations were
therefore excluded from capture attempts. We did ob-
tain a response in Orchard C, but the ow! never deigned
to approach our traps.
Nine individuals were captured in the summers of
2001 and 2002. On occasion, Screech-Owls began to
respond to broadcast calls while some daylight re-
mained. However, owls were always captured under
cover of darkness, between 21:15 to 03:10. Though
we were not always able to capture all the individuals
observed, we did record their presence (see Table 6).
Of the nine captured individuals and six observed, but
not captured, only one was red-phased; the rest were
grey.
Discussion
Eastern Screech-Owl range in Quebec
The Eastern Screech-Owl’s northernmost North
American range coincides with the southern tip of
Quebec (Gauthier and Aubry 1996). To the west of
Quebec, the species is commonly reported along the
southern edge of Ontario in the Carolinian zone along
Lake Erie and Ontario. More local and uncommon
populations extend up another 100 to 200 kilometres
north. In eastern Ontario, there appears to have been
Other
Hymenoptera:
ants, bees*, wasps
Eupplexoptera:
Earwigs
Unknown snake**
Jumping Mouse Hairy Woodpecker Arachnid:
Zapodidae (Picoides villosus) Spiders
Red squirrel Northern Flicker Coleoptera:
(Tamiasciurus hudsonicus)
(Colaptes auratus)
Unknown beetles
White-breasted Nuthatch
(Sitta carolinensis)
*could also have been cached prey item
**snakeskin found, could also have been part of nesting material
2006
an overall expansion of the range in the last 20 years
(M. Gahbauer, personal communication, 2004). To the
east of Quebec, the range may extend into the extreme
southwestern part of New Brunswick (Gauthier and
Aubry 1996); however, there are no confirmed breed-
ing records for the species there. If Screech-Owls do
in fact breed in New Brunswick, it is likely to be in very
low numbers (B. Whittam, personal communication,
2004). Screech-Owls have also been observed in Nova
Scotia and in Prince Edward Island, but their presence
there is thought to be incidental (ibid). Elsewhere in
Canada, the range is defined as extending from south-
western Saskatchewan to southern Manitoba (Panak
1988*). All of the above suggests that an important
proportion of the species’ range in Canada occurs in
Quebec and in Ontario.
Censusing
For our purpose, which was to confirm the presence
of resident Screech-Owls at a given location, we found
the optimum census period to be between November
and February. There are several logistical advantages
to censusing towards the beginning of this period.
During the early winter months, pairs jointly defend
their territory and are very responsive to broadcast calls.
As a result, there is a greater likelihood of obtaining
a response from both the male and the female, which
eliminates the need for speculation as to whether or
not a location is occupied by a pair. Later on, heavy
snowfall can lengthen the time required to census each
location considerably, and render some virtually inac-
cessible.
Censusing can have adverse effects. In late Febru-
ary and early March, females select a suitable nesting
location, then remain in or near their cavity. Broadcast
calls may elicit a response from territorial males, but
RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC
Ig?
FIGURE 2: A grey-phase Screech-Owl discovered during a
mid-winter census at the Réserve écologique Marcel-
Raymond, Henryville, Quebec, near the Richelieu
River. Photographed by Léopold Gaudreau, Assistant
Deputy Minister, Sustainable Development Branch,
Ministére du Développement durable, de |’Environ-
nement et des Parcs. 9 December 2006.
TABLE 4. Frequency of nest box occupation in orchards and control sites of Saint-Hilaire and Rougemont areas, Quebec
(2000 — 2004)
Location Number of CH/
boxes inspected** ESOW SQ NOFL WASP OTHER NO
Biosphere Reserve 38 26 2 l 0 0 9
Campground 116 48 26 3 5 5 29
A/A1* 60 8 4 5 | 7 35
B/K* 173 16 39 9 i) 9 93
(c 47 4 10 6 l | 25
D/F* 103 20 33 2 3 4 4]
G 51 5 3 2 l 5 25
I 67 2 29 0 l 9 26
M 39 4 22 0 2 0 1]
N/N1* 59 13 24 l 3 7 1]
Total 753 146 202 29 24 47 305
*Orchards are side by side or adjacent
**Indicates number of boxes inspected at Iccation over duration of the study
ESOW: number of boxes occupied by Screech-Owl
CH/SQ: number of boxes occupied by chipmunk or squirrel
NOFL: number of boxes occupied by a Northern Flicker
OTHER: number of boxes occupied by other inhabitants (see Appendix 1)
NO: number of unoccupied boxes
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 5. Eastern Screech-Ow! prey inventory based on pellets and prey remains found in nest boxes at 6 orchards and 2
control sites in Saint-Hilaire and Rougemont areas, Quebec (2001 — 2004)
Scientific name
Blarina brevicauda
Microtus pennsylvanicus
Peromyscus sp.
Sorex sp.°
Zapodidae®
Aegolius acadicus
Bombycilla cedrorum
Cardinalis cardinalis
Carduellis tristis
Colaptes auratus
Columba livia
Cyanocitta cristata
Junco hyemalis
Molothrus ater
Poecile atricapillus
Picoides villosus
Quiscalus quiscula
Sayornis phoebe
Sturnus vulgaris
Toxostoma rufum
Turdus migratorius
Zenaida macroura
Sittae
Turdidae
Tyrannidae
Unidentified bird*
Miscellaneous artifacts and prey
Apis mellifera
Rana clamitans
Unidentified membrane!
Corn, seeds
Eggshell fragments
* data spans 2001-2003 only
> White-footed or Deer Mouse
° Masked or Smoky Shrew
Common name
Short-tailed Shrew
Meadow Vole
Northern Saw-whet Owl
Cedar Waxwing
Northern Cardinal
American Goldfinch
Northern Flicker
Rock Dove
Blue Jay
Dark-eyed Junco
Brown-headed Cowbird
Black-capped Chickadee
Hairy Woodpecker
Common Grackle
Eastern Phoebe
European Starling
Brown Thrasher
American Robin
Mourning Dove
unidentified nuthatch
unidentified thrush
unidentified flycatcher
Honey Bee
Green Frog
4 Meadow or Woodland Jumping Mouse
© down remains unidentifiable
‘ may be the remains of a frog
D\F*
SN = tS
B\K N\NI
fs 3
5 31
1
3
1
1
1
1
1 5)
D
1
Biosphere
Reserve Campground
3 5
53 19
5
1
13 1
1
2
1
1
3
1 1
1
1
1
1
1
1 8
1
1
12
2
3
1 2
1
1 2
TABLE 6. Eastern Screech-Owls (n = 9) captured and banded in one control and six orchard locations of Saint-Hilaire and
Rougemont areas Quebec (2001-2002), by month of capture. See methods for keel index categories.
Location
Orchard I
Campground Control
Orchard K>
Orchard A/A1°4
Orchard D/F**
Capture
date
29 August 2002
22 July 2001 1 adult*
13 July 2002 1 adult
3 juvenile
31 July 2002 1 adult
9 August 2002 1 adult
| juvenile
1 juvenile
Captured
individual
Keel index Wing chord
Weight (g) (0-5) (mm)
2.5 to 3.0 170
78.0 3 177
78.0, 94.0, 96.0 52h 182,184,194
160 3 176
159 + 176
155 3 185
171 4 178
* red-phased individual
> | adult and 1 juvenile observed but not captured at B in June 2002
“ orchards are adjacent to one another and considered to comprise one territory
* 1 other juvenile observed but not captured
* | adult and 2 juveniles observed in June 2001 but not captured
2006
RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC 295
FiGureE 3. An Eastern Screech-Owl peers sleepily from a cavity at Pare Summit on the Island of Montreal. As development
within the province increases, so too does the importance of the parks and protected areas within the species range.
Photo by Jean-Sébastien Guénette, 17 April 2005.
given the necessity of devoting their energy to provi-
sioning their mate and themselves, and to defending
their territory against real rather than perceived intrud-
ers, censusing at this time can impose an unnecessary
and potentially harmful stress. Predators of the Screech-
Owl can also be attracted to the broadcast calls, and
to respondents, especially if illuminated by a flash-
light or headlamp. We are aware of one instance, for
example, where a Long-eared Owl, Asio otus, was
drawn to a Screech-Owl during censusing and was illu-
minated in the process of closing in on it. Since this
study, a “minimal-disturbance” censusing protocol has
been developed for the Screech-Owl and for other owl
Species (see Balej 2006).
Nest Box inspections
Nest boxes (” = 89) were installed in the vicinity of
where Screech-Owls responded during censusing. Ordi-
narily, an owl’s presence in an area would go largely
undetected because they are very inconspicuous and
occupy natural cavities. As the study progressed, our
primary concern became to place the nest boxes at a
height that would reduce the possibility of drawing
untoward human attention to them. Overall nest box
occupation at each location varied from 0% to 100%.
Forty-one boxes were occupied at least once by an
owl, and these boxes were a mean of 3.8 m (2.0 to
5.8 m) from the ground (Table 2). Gehlbach (1994b)
found owls nesting, on average, 3.7 m from the ground
in natural cavities (m = 23).
A number of species may compete with Screech-
Owls for nest boxes. The four most frequent occu-
pants were squirrels, Sciurus and Tamiasciurius sp.,
and chipmunks, Jamias striatus, Northern Flickers,
Colaptes auratus, and Hymenoptera, especially wasps.
During pre-snowfall inspections in October and No-
vember, we frequently removed nesting material left
by other species, and during inspections the following
spring found Screech-Owl pellets and prey remains.
Occasionally, Screech-Owls deposed chipmunks and
squirrels, judging from pellets or prey remnants deposit-
ed over nesting material. Northern Flickers have been
reported to destroy Screech-Owl eggs (Penak 1986*).
However, the Screech-Owls in our region infrequently
(22 of the 146 recorded times) occupied nest boxes in
the summer and early autumn months, when Flickers
occupied them. Late autumn inspections enabled us to
remove nesting material left by other species (or by
Screech-Owls during the winter). We also could replen-
ish the boxes with wood chips during these visits with-
out having to disturb any of the species that had pre-
viously occupied the boxes and make any necessary
repairs.
296
Pellets
The intact pellets retrieved from nest boxes (n = 82)
ranged widely in length from 1.0 to 8.5 cm and from
0.8 to 3.2 cm in width, with a mean weight of 1.77 g.
Elbroch and Marks (2001) recovered Screech-Ow1 pel-
lets (n = 18) from nearby Vermont and New Hampshire
which ranged from 1.6 to 3.8 cm length and 0.9 to
1.4 cm width. Fifty-six of our pellets exceeded the up-
per length and width reported by Elbroch and Marks.
Most of the intact pellets we measured were retrieved
from nest boxes, where they were protected from the
elements and from breaking. This, and our larger sam-
ple size, may explain the discrepancy in our upper
reported length.
Prey Consumption
Meadow voles were the predominant intact prey
item found in our nest boxes. Screech-Owls are also
highly insectivorous (Ritchison and Cavanagh 1992),
and although we repeatedly found evidence of insect
consumption, we were unable to fully assess or quan-
tify it. Only insects with hard parts, such as beetles,
Coleoptera, can be distinguished in pellets. For the
most part, we found only a dust of legs and carapaces
intermingled with the wood chips and with other arti-
facts. Mourning Doves were the avian prey most fre-
quently found in nest boxes, but few avian prey remains
were found. Screech-Owls may simply prefer to pluck
feathers on a perch or in a deeper tree cavity rather than
in a nest box which feathers could potentially fill quite
quickly. Hence, our prey inventory underestimates
avian and insect consumption. As well, several cached
frogs and shrews, and a decapitated vole, were also
found in nest boxes at the Campground control site.
On a number of occasions, remnants of crayfish and
fish scales were recovered from this site, as well as
pieces of a greenish membrane, which may have been
a remnant from the frogs. The greatest diversity in prey
items seems to have been consumed at the Biosphere
Reserve.
Conclusion
The Screech-Owl is found in suburban, rural and
agricultural habitats. It is considered to be one of the
most ubiquitous owls in North America (Gehlbach
1995). There is, however, very little information avail-
able about the species’ natural history and ecology, or
about its density and overall distribution in Canada,
where its status was last evaluated for the Committee
on the Status of Endangered Wildlife in Canada over
20 years ago (by Penak 1986*). Paradoxically, the
species is currently designated as “Not at Risk”, even
though there are no current data on its population
trend status (http://www.cws-scf.ec.gc.ca/mgbe/trends/
index .cfm?lang=e& go=info.bird&speciesid=3730).
An important part of the species’ range in Canada
occurs in southern Quebec, where it coincides with an
agricultural belt and with the province’s major apple-
producing region. Screech-Owls are known to favour
THE CANADIAN FIELD-NATURALIST
Vol. 120
orchard habitats (Gauthier and Aubry 1996) where
pesticides are intensively applied. Our study evaluating
the exposure of Screech-Owls to pesticides currently
and previously used in one of the largest apple-pro-
ducing regions of southern Quebec provided an oppor-
tunity to collect information on the species’ regional
diet, record physiological parameters, evaluate occupa-
tion response to nest boxes and gain a sense of the den-
sity and distribution of the local population. As far as
we know this is the first time that these types of data
have been collected on the Screech-Owls of the area.
More baseline information of this type must be gath-
ered for future monitoring and population management
efforts throughout the owl’s Canadian range. Changes
in development and land use patterns must also be
examined. While we found that exposure to DDE and
organophosphorus pesticides warranted further inves-
tigation (see Richards et al. 2004, 2005), we also ob-
served a marked increase in clearing and development
of forested lands and orchards over the course of the
study. We believe that loss of habitat and overall de-
crease in habitat quality currently pose a far greater
threat to the local population than exposure to pesti-
cides.
Our colleagues at the Quebec environment ministry
(Ministere du Développement durable, de |’Environ-
nement et des Parcs) have been conducting autumn
and winter censuses in ecological reserves within the
known range of the species since 2005. This is part of
an ongoing study initiated to evaluate the relative risks
that pesticide exposure and habitat loss/deterioration
pose to Screech-Owls in southern Quebec (if interest-
ed in collaborating on this study, please contact N.
Richards).
Even if the North American Screech-Owl popula-
tion may not be at risk as a whole, we are concerned
that habitat loss and modification may be adversely
affecting regional populations. At the very least, a re-
examination of the Screech-Owl’s distribution and pop-
ulation status throughout its Canadian range, espe-
cially in Quebec and Ontario, is long overdue. If this
adaptable and opportunistic species is being adverse-
ly affected, how does this bode for the more vulnera-
ble species which occupy the same habitats?
Acknowledgments
We are grateful to the orchard owners of the study
area, without whose collaboration this study could not
have taken place. We thank Terry Ryan and Jean Gou-
geon for building industrial quantities of nest boxes.
Doctors Guy Fitzgerald and Claude Lacasse of the
Clinique des oiseaux de proie and Annemarie Roth
of the Centre for the Rehabilitation of Injured Birds
of the Montérégie provided us with Screech-Owl ad-
mission records. Stéphane Deshaies and Annemarie
Roth helped identify avian prey remains. Ambroise
Lycke, Christine Bourque, Patrick MacFarlane, Anke
Roth, Gavin McMorrow, Jessus Karst, Hugo Gee, Mar-
2006
cel Gahbauer, Tina Newbury, Amy Stevenson, Mar-
isha Futer, Jean-Sébastien Guénette, Joanna Coleman
and Anneli Jokela all provided invaluable assistance
and/or training in the field. We thank Nicole Kopysh
at the Ontario Bird Banding Atlas project and Becky
Whittam at the New Brunswick division of Bird Stud-
ies Canada for information related to the Screech-
Owl’s distribution in these provinces. Caroline Cormi-
er of Nature-Action, Québec Inc. and Gina Rossini
provided information on the whereabouts of birds of
prey in Rougemont and Saint-Hilaire. We are pleased
to acknowledge the collaboration and hard work of
Rodolph Balej and Marcelle Ruest at the Ministére de
Development durable de l’Environnement et des parcs.
Funding for this study was provided to N. L. Richards
by the Fonds Québécois de la recherche sur la nature
et les technologies and by the Canadian Wildlife Ser-
vice, Environment Canada. Additional funding was
provided through a Natural Sciences and Engineering
Research Council of Canada (NSERC) grant to P.
Mineau. Initial stages of the project were funded by
Environment Canada’s Science Horizons for Youth
programme, in collaboration with World Wildlife Fund
Canada. In particular, we gratefully acknowledge the
continued funding and support provided by Bird Pro-
tection Quebec (BPQ) (formerly the Province of Que-
bec Society for the Protection of Birds — PQSPB).
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tion on the osteological composition of ow! pellets. Acta
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cides and anticoagulant rodenticides in apple orchards of
southern Québec, Canada. Pages 389-408 in Raptors World-
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ceedings of the Sixth World Conference on Birds of Prey
and Owls. May 18-23, Budapest, Hungary.
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assessment approach to DDE exposure based on the case
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Ritchison, G., and P. Cavanagh. 1992. Prey use by Eastern
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Received 26 July 2005
Accepted 4 June 2007
Effects of Timber Harvesting and Plantation Development on
Cavity-nesting Birds in New Brunswick
STEPHEN J. WOODLEY!, GREG JOHNSON”, BILL FREEDMAN?, and Davip A. KirK*
‘Ecological Integrity Branch, Parks Canada, 25 Eddy Street, Hull, Québec K1A 0M5 Canada
"Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada; Present address: Jacques-Whitford
Environment Ltd., Fredericton, New Brunswick E3B 5C2 Canada
>Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada
*Aquila Applied Ecologists, C.P. 87, Carlsbad Springs, Ontario KOA 1KO Canada
Woodley, Stephen J., Greg Johnson, Bill Freedman, and David A. Kirk. 2006 Effects of timber harvesting and plantation
development on cavity-nesting birds in New Brunswick. Canadian Field-Naturalist 120(3): 298-306.
We studied the abundance of cavity-nesting birds in forestry-related habitats in a region of Acadian forest in New Brunswick.
We examined five reference stands of natural forest, a chronosequence of conifer plantations up to 19 years old (the oldest
in the study area), two selectively harvested stands, and a 30-year-old naturally regenerated clear-cut. The species richness
and abundance of cavity-nesting birds were higher in reference forest (average 10.0 species per stand; 5.3 territories per 10 ha)
than in plantations (2.3/stand; 1.0/10 ha), selectively harvested stands (7.0/stand; 3.8/10 ha), or the naturally regenerated
clear-cut (5.0/stand; 2.5/10 ha). A cluster analysis segregated the “community” of cavity-nesting birds of natural forest from
those of other treatments. Of the various harvested stands and plantations, five with a relatively large number of residual
snags clustered similarly in the cluster analysis, while those with no or very few snags also clustered together. We used
arrays of nest boxes (12 per stand) to examine whether the availability of cavities was limiting the use of habitats otherwise
suitable for foraging by cavity-dependent species. Nest-box use for nesting and roosting was much higher in the seven plan-
tations examined (average 4.0/10 ha for nesting and 2.9/10 ha for roosting) than in three reference stands (each 0.3/10 ha),
suggesting that the plantations were deficient in this critical-habitat element. Our results suggest that certain mitigations,
such as leaving residual snags and living cavity-trees, would help maintain populations of some cavity-dependent birds in
clear-cuts and plantations. However, some cavity-dependent species might not be accommodated by these mitigations and
are potentially at risk in intensively managed areas, unless landscape-scale management plans ensure the survival of suffi-
cient areas of older mixed-wood forest.
Key Words: Acadian forest, cavity-nesting birds, conifer plantations, critical habitat, forestry, management recommendations,
nest boxes, New Brunswick.
Freedman et al. 1996; Hobson and Schieck 1999;
Schieck and Hobson 2000). Intensive forestry prac-
tices generally decrease the richness and abundance
of cavity-dependent birds by causing the following
habitat changes to occur (Niemi et al. 1998; Hobson
and Bayne 2000; Flemming and Freedman 1998; Imbeau
et al. 2001; Kirk and Machtans 2004; McRae et al.
2001):
e the conversion of natural forest having complex
biological and physical structure (i.e., multi-
species stand composition and intricate spectra of
age and size) into silvicultural plantations domi-
nated by a monospecific cohort of similar-sized
and -aged trees
e the reduction in quantity of cavity trees, dead
snags, and coarse-woody debris by intensive man-
The effects of forestry on biodiversity range over a
wide continuum, depending largely on the intensity of
harvesting and subsequent management (Hunter 1990;
Freedman et al. 1994; Angelstam and Milkusinski
1994; Freedman 1995; Hagan et al. 1997; Niemi et al.
1998; Imbeau et al. 2001; McRae et al. 2001; Kimmins
2003). For instance, clear-cutting followed by intensive
management to develop an even-aged monocultural
plantation has relatively extreme effects, while the
effects associated with selective-harvesting and natural
regeneration are much smaller. In boreal Europe, for
example, natural pine (Pinus sylvestris) forest has been
converted extensively into commercially preferred,
short-rotation conifer plantations (Esseen et al. 1997),
resulting in large declines of birds dependent on older
natural forest (Virkkala 1991). Similar changes are
occurring in temperate- and boreal-forest regions of
Canada, where older mixed-species forest is being con-
verted extensively into conifer plantations (Niemi et al.
1998; Imbeau et al. 2001; McRae et al. 2001).
Cavity-dependent birds in temperate and boreal
forest are generally most abundant and species-rich
in older, uneven-aged, mixed-species stands (Hunter
1990; Schreiber and Decalesta 1992; Newton 1994;
agement practices (in Canada and elsewhere snag
felling may even be required under health and
safety regulations; e.g., Naylor et al. 1999)
e the truncation of stand age-at-maturity by short-
rotation management (typically 40-60 years),
which precludes the regeneration of large cavi-
ty-trees and coarse-woody debris
298
2006
Moreover, at the level of landscape, forestry typi-
cally decreases the average age and complexity of
habitat “patches,” while isolating remnants of older
natural forest (Hunter 1990; Freedman et al. 1994;
Hagan et al. 1997; McRae et al. 2001). Landscape-
scale forest management must accommodate species
with extensive and specialized habitat needs, includ-
ing some cavity-nesting birds dependent on older
forest (Renken and Wiggers 1989, 1993; Bull and
Holthausen 1993; Freedman et al. 1996; Bonar 2000).
Because of the specialized need of cavity-users for
critical habitat, they are often considered indicators
of the ecological sustainability of forest management
(Angelstam and Mikusinski 1997). It is particularly
important to maintain the habitat used by “keystone”
primary excavators, such as older heart-rotten trees
and snags, because a diverse group of secondary users
depends on the cavities they create (Freedman et al.
1996; Martin and Eadie 1999). The Pileated Woodpeck-
er (Dryocopus pileatus) is one example of a keystone
excavator whose abandoned cavities are used by other
species for nesting or roosting (Bull and Jackson 1995;
Bonar 2000).
The effects of forestry on cavity-users are relatively
wel known for forest types of the eastern and north-
western United States (Raphael and White 1984;
DeGraaf and Shigo 1985; Zarnowitz and Manuwal
1985; Renken and Wiggers 1989; Connor et al. 1994).
However, much less information is available on which
to base management decisions in the north-temperate
and boreal regions of eastern Canada. This deficiency
is important because habitat needs of cavity-nesters
vary geographically and are likely to be different in
boreal and Acadian forest than in other regions and
biomes (Parker et al. 1999). In recognition of this dearth
of information, studies were initiated in the mid-1990s
of cavity-nesting birds in eastern Canada, with the in-
tent of informing management guidelines to conserve
their populations (Doyon et al. 1999; Naylor et al.
1999; Bonar 2000; Flemming et al. 2000). This advice
is greatly needed, because of the increasing areas of
natural forest that are being intensively managed for
timber production in eastern Canada.
Within this context, our study examines effects of
forestry on cavity-dependent birds in a region of Aca-
dian forest in southern New Brunswick. We assessed
the degree to which cavity-nesting birds use conifer
plantations and less-intensively managed stands, com-
pared with natural forest. This was done by conducting
breeding surveys of these habitats and by comparing
occupancy of nest boxes as an indicator of limitation
by natural cavities.
Study Area
The study area is in southeastern New Brunswick,
Canada. It is located in the Atlantic Maritime Ecozone
(Ecological Stratification Working Group 1995), with-
in the Fundy Plateau Ecodistrict of the Acadian For-
WOODLEY, JOHNSON, FREEDMAN, KIRK: EFFECTS ON CAVITY-NESTING BIRDS
299
est Region (Loucks 1962; Rowe 1972). The climate is
humid temperate and the natural forest is dominated
by mixed-species stands of Red Spruce (Picea rubens),
White Spruce (P. glauca), Balsam Fir (Abies bal-
samea), Sugar Maple (Acer saccharum), Red Maple
(A. rubrum), Yellow Birch (Betula alleghaniensis),
White Birch (B. papyrifera), and Mountain Birch (B.
cordifolia). Extensive natural disturbances, including
irruptions of native Spruce Budworm (Choristoneura
fumiferana), have affected the natural forest, which
tends to have a mixed-species canopy with gap-phase
regenerating patches. Superimposed on the natural dis-
turbance regime is a complex of anthropogenic influ-
ences. This began with the selective harvesting of large
trees for lumber and ship-building, followed by exten-
sive deforestation for agricultural development, then
abandonment of most poorer-quality farmland begin-
ning in the 1920s, and more recently the widespread
establishment of conifer plantations.
Our study area is within the Greater Fundy Ecosys-
tem (GFE), a region consisting of Fundy National Park
and its surrounding area. The GFE was defined to
study ecological consequences of the insularization of
Fundy National Park, a 204 km? protected area embed-
ded within a landscape whose matrix is being trans-
formed by the conversion of natural forest into conifer
plantations (Woodley et al. 1993, 1998; Freedman et
al. 1994). The GFE itself is within the Fundy Model
Forest (FMF), one of ten “model forests” established
to demonstrate “sustainable forestry” in Canada. One
declared indicator of sustainability in the FMF is that,
at the landscape level, native biodiversity must not be
compromised by forest management (Parker et al.
1999).
Methods
Stand selection
We selected 18 stands for study (see Table 1), includ-
ing representative natural (or “reference”) forest and
habitats resulting from various intensities of manage-
ment, including:
¢ five reference stands of unmanaged mature for-
est, including three mixed woods (dominated by
Red Spruce (Picea rubens), Balsam Fir (Abies
balsamea), and White Birch (B. papyrifera), and
Mountain Birch (B. cordifolia) and two soft-
wood dominated by Red Spruce:
¢ ten stands that had been clear-cut and then in-
tensively managed to develop conifer plantations
ranging from 5 to 19 years in age (the oldest avail-
able in the study area); eight were planted with
Black Spruce (Picea mariana), one with Jack Pine
(Pinus banksiana), and one with Norway Spruce
(Picea abies):
* two selectively harvested stands (1-year-old and
12-years-old);
* one 30-year-old naturally regenerated clear-cut.
300
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. General habitat characteristics of stands surveyed for cavity-nesting birds.
Survey Area Stand
Site Year (ha) description
Reference mixedwood forest
Ra 1994 10 mature/mixedwood forest; Red Spruce, Balsam Fir, White Birch
Rb 1995 20 mature/mixedwood forest; Red Spruce, Balsam Fir, White Birch
Re 1994 10 mature/mixedwood forest; Red Spruce, Balsam Fir, White Birch
Rd 1994 10 mature/Red Spruce-dominated forest
Re 1995 20 mature/Red Spruce-dominated forest
Plantations
P5a 1994 10 5-year-old Black Spruce plantation
P5b 1994 10 5-year-old Norway Spruce plantation
1Py/ 1995 20 7-year-old Black Spruce plantation with residual snags
P8 1994 10 8-year-old Black Spruce plantation
P9a 1994 10 9-year-old Black Spruce plantation
P9b 1994 10 9-year-old Jack Pine plantation
IIS) 1995 8 15-year-old Black Spruce plantation with residual snags
P17 1994 10 17-year-old Black Spruce plantation
P18 1995 10 18-year-old Black Spruce plantation
P19 1994 10 19-year-old Black Spruce plantation
Selectively harvested stands
$12 1995 20 12-year-old selection harvested stand
S1 1995 20 1-year-old selection harvested stand
Naturally regenerated clear-cut
NR30 1995 20 30-year-old naturally regenerated clear-cut with residual snags
Note that selected study stands were not always
paired because of the lack of availability of suitable
stands that were given the same management. The
study plots affected by forestry were essentially opera-
tional stands embedded in a dissimilar habitat matrix
generally of mixed-wood forests, and they were sur-
veyed in their entirety. The reference plots were of a
size and shape comparable to forestry-affected plots.
The study stands were no more than 20 km from each
other. Fieldwork was done during 1994 and 1995.
Habitat description
Trees (>5 cm diameter at breast height (DBH) and
>1.5 m tall) and snags (standing dead trees) were sam-
pled in twelve quadrats (20 m x 20 m) arranged as a
grid across the study plot. However, in plantations
15 years and older, smaller quadrats (10 m x 10 m)
were used because of the uniform structure of tree-
sized vegetation. Shrubs (<5 cm DBH) were measured
for diameter at 25 cm in two 5 m x 5 m sub-quadrats
in opposite corners of each tree-quadrat. Length and
diameter of coarse-woody debris (CWD; >5 cm diam-
eter) were also measured in tree-quadrats. The field data
were used to calculate the average DBH and basal area
by species, including snags, and the volume of CWD.
The cover of species of ground vegetation was deter-
mined in 30 randomly located 1 m? quadrats per site.
The cover of overhead canopy was estimated as the
percentage area obscured by foliage when sighting
upward through a 4.2-cm diameter cylinder. Here we
present summary information; detailed data on woody
habitat are in Flemming and Freedman (1998) and on
ground vegetation are in Veinotte et al. (2004).
Abundance of birds
Cavity-nesting birds were surveyed by modified
spot-mapping to identify territories (Bibby et al. 1999).
Habitat use was designated as “nesting territory” if we
found evidence of a nest, and as “present” if a nest site
could not be located. The surveys were conducted in
plots with areas of 8 to 20 ha (Table 1). Ten stands were
surveyed in 1994 and eight in 1995, and each was
examined seven times from early June through to early
July. The locations of calling or singing male birds
were mapped in the study plots and also in adjacent
habitat, and territorial boundaries were estimated using
standard spot-mapping procedures (Bibby et al. 1999).
Where possible, we recorded the sex, age (adult/juve-
nile), and relevant activity (nesting, feeding young,
foraging, drumming, singing, calling) of individuals.
This information contributed to the assignment of
breeding territories.
Nest-box survey
To indicate whether cavities were a limiting factor
for dependent species, we installed arrays of 12 nest
boxes in each of seven plantations and three reference
stands. The nest boxes were made of sections of White
Cedar (Thuja occidentalis) with natural heart rot and
were divided into three sizes: small (3.5-cm diameter
entrance hole; ca. 6.5-cm internal width by 16 cm
height), medium (6-cm hole, 11 x 34 cm), and large
(9-cm hole; 15 x 48 cm). Because the site location
and orientation of the entrance hole can affect occu-
pancy (Rendell and Robertson 1994), these factors
were standardized. Each nest box was mounted on an
aspen pole supported by three steel cables fixed to
2006 WOODLEY, JOHNSON, FREEDMAN, KIRK: EFFECTS
rebar pegs, and was placed at a predetermined height
(small cavities at 2.5 m, medium at 3 m, and large at
5 m) with the entrance hole facing south. A total of
12 nest boxes was placed in each stand; two of each
of size were positioned at 30 m and again at 100 m
from a stand edge. The nest boxes were surveyed seven
times during each of the 1994 and 1995 breeding sea-
sons, and once late in the summer. We recorded the
numbers of eggs, hatchlings, and fledglings of any
species occupying the boxes, as well as roosting birds
and non-avian species.
Data analysis
Because the study plots were unique habitats in
terms of location, site and habitat attributes, and dis-
turbance history, they were not treated as true statisti-
cal “replicates” (Hurlbert 1984; Heffner et al. 1996).
Moreover, many of our data did not meet assumptions
of normality or homegeneity of variance. Consequent-
ly, we restricted our analyses to simple comparisons
among Classes of habitat types. Relationships among
stands were examined using multivariate analyses, with
data input being matrices of species abundance by site
(Kovach 1995). A cluster analysis was used to identify
groupings of stands (or “communities”), using an un-
weighted pair-group procedure with arithmetic aver-
ages (UPGMA,; Sokal and Rohlf 1995). Chi-square
contingency tests were used to test for differences in
occupancy rate of nest boxes between (a) unmanaged
and managed stands and (b) boxes located at 30 m or
100 m from a forest edge.
Results and Discussion
Habitat
The stands of natural (reference) forest had a large
basal area (an indicator of biomass) of trees compared
with the other habitats studied (Table 2). Although a
high tree density occurred in older plantations and the
naturally regenerated site, these were smaller trees than
in natural forest. The species composition of the nat-
ural forest was mixed, whereas the plantations are more
strongly dominated by the planted conifer species. The
natural forest also had relatively abundant snags. Of
the various managed stands, only the 7-year-old and
15-year-old plantations (P7 and P15, respectively) had
many snags, because the pre-harvest snags had not all
been removed or felled during the clear-cut, and a sub-
sequent herbicide treatment killed surviving hardwood
trees. The 30-year-old naturally regenerated clear-cut
also had abundant snags, as did the 12-year-old selec-
tively harvested site. Coarse-woody debris was abun-
dant in the stands of reference forest, where it origi-
nated as natural deadfall. Coarse-woody debris was
also abundant in the plantations and selectively har-
vested stands, where it mostly originated as logging
slash and, in older plantations, trees cut and left dur-
ing a non-commercial thin. Similar but more detailed
observations have been made by other studies of nat-
ural forest and plantations in our study area (Flem-
ming and Freedman 1998).
TABLE 2. Summary of key habitat data (presented as mean area).
ON CAVITY-NESTING BIRDS
a
c
°
p=)
SS
=
c
Ss
oO
Reference Stands
Sl
Rd
Re
Rb
Ra
Tree Basal Area (m2/ha)
Conifers
18.3 19.0 30.3 30.2
18.3
10.9
10.7
6.4
11.4
10.9
10.9
Hardwoods
Snags
5.0
Sal
3.6
6.1
Tree Density (10°*/ha)
Conifers
0.9
0.5
0.6 1.0 0.9
0.6
0.5
Hardwoods
Snags
ala
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arse = —i—
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0.5
0.5
0.7
0.2
().2
Shrub Basal Area (m2/ha)
301
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Vol. 120
302
THE CANADIAN FIELD-NATURALIST
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WOODLEY, JOHNSON, FREEDMAN, KIRK: EFFECTS ON CAVITY-NESTING BIRDS 303
FIGURE 1. Cluster analysis of stands based on their species and abundances of cavity-nesting birds. See Table | for explana-
tion of site codes.
The abundance and cover of shrubs and ground veg-
etation were highly variable among stands (Table 3).
In general, these vegetation elements were more abun-
dant in the plantations and other harvested stands,
which were in younger stages of secondary succes-
sion than the more mature natural forest. Similar, but
more detailed observations of forest and plantations
in our study area have been made by Veinotte et al.
(2004).
Cavity-nesting Birds
A total of 16 species of cavity-nesting birds was
observed (Table 3). The most abundant species were
the Black-capped Chickadee (24% of the territories;
see Appendix | for avian binomials), Boreal Chick-
adee (23%), Red-breasted Nuthatch (16%), and Win-
ter Wren (14%). The species richness was higher in the
natural forest (average of 10.0 species per stand; total
of 14 species present in the habitat type) than in plan-
tations (2.3/stand; 8 species present), selectively har-
vested stands (7.0/stand; 9 species present), or the nat-
urally regenerated clear-cut (5.0/stand). Abundance was
also higher in the reference forest (average 5.3 terri-
tories/10 ha) than in plantations (1.0/10 ha), selective-
ly harvested stands (3.8/10 ha) or the naturally regen-
erated clear-cut (2.5/10 ha). Among the ten plantations
studied, the highest density of cavity-nesting birds was
in P15 (4.0 territories/10 ha), which also had a rela-
tively large number of snags (2.1 m?/ha compared with
an average of 0.4 m’/ha among the other nine planta-
tions).
A cluster analysis segregated the “community” of
cavity-nesting birds of the natural forest from those of
the other treatments (Figure 1). In the cluster analy-
sis, the five reference stands were arranged together on
the right-hand side. The selectively harvested stands
(S1 and $12) and naturally regenerated stand (NR30)
had relatively abundant cavity-nesting birds and clus-
TABLE 4. Cavity-using species encountered in this study.
American Kestrel
Barred Owl
Northern Saw-whet Owl
Northern Flicker
Yellow-bellied Sapsucker
Downy Woodpecker
Hairy Woodpecker
Black-backed Woodpecker
Pileated Woodpecker
Tree Swallow
Black-capped Chickadee
Boreal Chickadee
Brown Creeper
White-breasted Nuthatch
Red-breasted Nuthatch
Wood Duck
Winter Wren
Falco sparverius
Strix varia
Aegolius acadicus
Colaptes auratus
Sphyrapicus varius
Picoides pubescens
Picoides villosus
Picoides arcticus
Dryocopus pileatus
Tachycineta bicolor
Poecile atricapillas
Poecile hudsonicas
Certhia americana
Sitta carolinensis
Sitta canadensis
AIx sponsa
Troglodytes troglodytes
304
tered close to the reference stands. Of the plantations,
those with relatively abundant and species-rich cavity-
nesting birds clustered together (1.e., P7, P15, P19, and
P17), while the most depauperate ones formed another
cluster (i.e., P5b, P8, P9a, P9b, P5a).
The observation that cavity-nesters are more abun-
dant in mature mixedwood forest than in harvested
stands or plantations is consistent with research else-
where, although the specifics vary depending on the
forest region and species of birds present (e.g., Raphael
and White 1984; DeGraaf and Shigo 1985; Zarnowitz
and Manuwal 1985; Hansen et al. 1995; Kirk and Nay-
lor 1995; Niemi et al. 1998; Hobson and Bayne 2000).
Of the 16 species of cavity-nesting birds recorded
during our study, eight were not recorded in any of the
plantations: Saw-whet Owl, Barred Owl, Yellow-bellied
Sapsucker, Downy Woodpecker, Black-backed Wood-
pecker, Pileated Woodpecker, White-breasted Nuthatch
and Winter Wren. One species, the Tree-Swallow, was
only recorded as breeding in plantations in which nest
boxes were present.
Nest-box Occupancy
Nest boxes were used by five species for nesting
and by four for roosting (Table 4). Tree Swallow ac-
counted for 41% of the nests, American Kestrel 31%,
and Boreal Chickadee 17%. The most frequently roost-
ing species were Tree Swallow (48%) and Northern
Flicker (33%). The use of nest-boxes for nesting was
much greater in plantations (average 4.0/10 ha) than
in the natural forest (0.3/10 ha), and similarly for roost-
ing (average 2.9/10 ha in plantations and 0.3/10 ha in
reference stands). Overall, 33% of the nest boxes in
plantations were used for nesting or roosting (28 of
84 boxes with nests), compared with only 3% in nat-
ural forest (1 of 36 boxes with nests). These observa-
tions support the notion that suitable cavities are a lim-
iting factor in plantations in the study area, compared
with the natural forest. The plantations we studied
THE CANADIAN FIELD-NATURALIST
Vol. 120
appear to provide suitable habitat for foraging and
other purposes, but can only be used for nesting and
roosting by cavity-dependent birds if their need for
cavities is met. There were no recorded cases of a failed
nest in the nest boxes.
Of the 28 nest boxes used for nesting in plantations,
7.5% were located 30 m from an intact forest edge
and 15.8% were 100 m from such an edge (marginally
significant difference; y* = 3.78, 1 df; >0.05 P <0.1).
The apparent reluctance to use nest-boxes located clos-
er to a habitat edge may be associated with a greater
risk of predation (Paton 1994; Derochers and Hannon
199M):
Implications for Management
It appears that conifer plantations in our study region
can recover bird populations rather quickly after estab-
lishment, but the species composition is different from
that of natural forest, and cavity-dependent species are
present in low abundance (Freedman et al. 1994; Freed-
man and Johnson 1999; Johnson and Freedman 2002).
Moreover, observations and models of stand develop-
ment suggest that intensively managed plantations are
depauperate in snags, cavity-trees, and coarse-woody
debris and are likely to remain so over subsequent rota-
tions because the harvesting removes all large-dimen-
sion tree biomass (Flemming and Freedman 1998).
These changes do not augur well for cavity-depend-
ent species in stands and landscapes extensively con-
verted into forestry plantations.
Clearly, the extensive conversion of mixedwood
Acadian forest into conifer plantations will result in a
decline in the abundance and species richness of cavity-
dependent birds at the scales of both stand and land-
scape. Certain site mitigations might help some species,
particularly the retention of some snags and large liv-
ing trees within plantations, or much less preferably,
the provision of nest boxes to provide local nesting
and roosting habitat (Welsh and Capen 1992; Newton
TABLE 5. Use of nest boxes in plantations and reference forest during two study years. Each site had an array of 12 nest boxes,
each made of a section of a hollow log of White Cedar. All plots were 10 ha in area. Note some stands from Table 3 had no
nest boxes installed, so the number of stands differs between Table 3 and 4.
Reference
Species
Nesting
Wood Duck 1
American Kestrel 2;
Northern Flicker
Tree Swallow D)
Boreal Chickadee 1
Total nests I 0 5
Roosting
American Kestrel
Pileated Woodpecker
Northern Flicker
Tree Swallow
Total roosting
Plantation
2 2 2 1
1 1
3 3 1
2 2
6 6 5 1 3 2
2
1 3 2 1
2 5 3
4
2006
1994; Flemming et al. 2000). It is likely, however, that
these site mitigations will not be sufficient to maintain
some cavity-dependent species, particularly those with
large home ranges that encompass a mosaic of habitat
patches, such as Pileated Woodpecker (Bonar 2000;
Flemming et al. 2000). These species will require large
protected areas of unmanaged forest to function as
population centres.
Without sympathetic management of this sort, it is
likely that the population viability of some cavity-
dependent birds in regions extensively managed for
forestry will become compromised. Some jurisdictions
have established guidelines to help these species — in
New Brunswick, for example, timber companies oper-
ating on Crown Land are to required to retain at least
10% of conifer-dominated forest in mature or older suc-
cessional stages, in patches of 500 or more ha (Sulli-
van 1996). However, these requirements may not be
adequate to maintain critical habitat for all species of
cavity-nesting birds, particularly keystone primary exca-
vators (Flemming et al. 2000). It is essential that these
conservation guidelines be monitored to ensure they
achieve their intended ecological purpose, including
the maintenance of viable populations of all cavity-
dependent species. In the meantime, a precautionary
approach to ecologically sustainable forestry would
require that additional large areas of natural forest be
set aside from intensive economic use, as parks or other
kinds of protected areas.
Acknowledgments
We are grateful to staff at Fundy National Park for
logistical support. We thank Irving Forest Products
Ltd. for allowing access to their plantations and for
providing poles and cedar sections for constructing
nest boxes. Research funding was provided by the
Fundy Model Forest, Natural Resources Canada, Parks
Canada, and the Natural Sciences and Engineering
Research Council of Canada.
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Received 12 April 2005
Accepted 5 May 2007
Recent Invasion, Current Status and Invasion Pathway of European
Common Reed, Phragmites australis subspecies australis, in the
Southern Ottawa District
PAUL M. CATLING! and SUSAN CARBYN?
'Biodiversity, National Program on Environmental Health, Agriculture and Agri-food Canada, Wm. Saunders Building,
Central Experimental Farm, Ottawa, Ontario K1 A 0C6 Canada; e-mail: catlingp@agr.ge.ca
Biodiversity, National Program on Environmental Health, Agriculture and Agri-food Canada, 32 Main Street, Kentville,
Nova Scotia B4N IJS Canada; e-mail: carbyns@agr.gce.ca
Catling, Paul M., and Susan Carbyn. 2006. Recent invasion, current status and invasion pathway of European Common Reed,
Phragmites australis subspecies australis, in the southern Ottawa district. Canadian Field-Naturalist 120(3): 207-312.
A survey of populations of native North American Common Reed (Phragmites australis subsp. americanus) and the introduced
invasive alien, European Common Reed (Phragmites australis subsp. australis) was conducted in four eastern Ontario town-
ships in 2003. The historical status of the two taxa in the region was evaluated through reference to herbarium specimens
collected over the past century. The introduced subspecies appears to have entered the district in the 1970s and subsequently
assumed dominance but was not recognized as an alien until 2003. It is now widespread in the four townships where ninety-five
populations were recorded during the 2003 survey. Of these, 25 were referable to the native subspecies which was localized
and 70 were referable to the introduced subspecies which was widespread. The native subspecies occurs in natural wetlands
and also spreads to roadsides, but the introduced subspecies is much more strongly associated with roads, where the rhizomes
extend onto gravel shoulders and are broken and transported by construction equipment, graders, ploughs, mowers, and in
the treads of many kinds of vehicles. Sensitive wetlands should have buffer zones exceeding 1000 m to prevent colonization
of subsp. australis. Monitoring of the two subspecies will be essential to the protection of native biodiversity, since early
detection of the alien subspecies in a sensitive wetland will offer the best opportunity for control.
Key Words: North American Common Reed, Phragmites australis subsp. americanus, European Common Reed, Phragmites
australis subsp. australis, invasion pathway, alien, invasive species, Ottawa, Canada.
One of the most productive plants in temperate
regions, Common Reed, Phragmites australis (Cav.)
Trin. ex Steud. has economic and agricultural values
and is under-exploited (Small and Catling 2001).
Domestication and introduction of non-native races
however, pose some potential risks to both agricul-
ture and environment. It is a major pest of irrigation
and flood control channels in some parts of the world
and, being able to grow in water 2 m deep or on dry
ground, it is not readily controlled by drawdown.
It was only recently realized that both native and
introduced races existed in North America (Catling et
al. 2003). The native plants have been designated sub-
species americanus Saltonstall, P.M. Peterson & Soreng
(Saltonstall et al. 2004) and the introduced plants iden-
tified as subspecies australis (Catling 2006, 2007, Cat-
ling et al. 2007). It is believed that the introduced sub-
species australis originated from southern Europe. It
has been recently recognized as a major threat to North
American biodiversity (Chambers et al. 1999; Meyer-
son et al. 2000; Catling et al. 2003; Robichaud and Cat-
ling 2003; Catling 2005). In a recent survey of the St.
Lawrence River wetlands it was found to have a greater
impact on native plant diversity than most other inva-
sive aliens including Purple Loosestrife, Lythrum sali-
caria, (Lavoie et al. 2003). The subspecies australis is
currently spreading in Canada from areas of concen-
tration in southern Ontario and southern Quebec and
has only recently been reported from western Canada
(Martin 2003; Schueler et al. 2003) and the maritime
provinces (Catling et al. 2004).
In eastern Ontario the alien subspecies australis was
first recognized in 2003, but it apparently invaded the
area a few decades earlier. Local botanists noticed the
rapid spread of dense stands of Phragmites beginning
in the 1970s and the rapid change from the status of a
locally rare plant (Dore 1959) created suspicion about
the possibility of an introduced race. Preliminary obser-
vations suggested that it spread into the region along
roads very recently and became widespread. Here we
analyse its apparent recent spread into the region along
roads by providing information on its history of inva-
sion, current status and invasion pathway in the south-
ern part of the Ottawa district based on identification
of herbarium specimens and a current survey of popu-
lations in four adjacent townships. This work will also
provide benchmark data for analysis of future changes
in status of native and introduced subspecies, thus con-
tributing information needed to predict rate of spread
and impact.
Methods
A recent study of plants of Phragmites australis sen-
su lato in eastern Ontario suggested that plants with
lower glumes exceeding 4.2 mm long were referable to
the native subspecies americanus (Robichaud and Cat-
307
308
subsp. australis subsp. americanus
Ficure |. Glumes of Phragmites australis. The smaller on the
right of each attached pair is the lower glume, whereas
the longer glume on the left of each pair is the upper
glume. The subspecies australis on the left has short-
er lower glumes as represented by three examples
from an inflorescence of a specimen from Dundas
County (Catling and Robichaud, 10 Oct. 2002, DAO).
On the right are three pairs from an inflorescence of
subspecies americanus from Leeds and Grenville
County (Catling and Kostiuk, 30 Aug. 2003, DAO)
showing its characteristically longer lower glumes.
ling 2003). More recent work based on correlation with
basal stem color and date of collection (Saltonstall et
al. 2004; Catling and Mitrow in press) has supported the
separation using 4.2 mm for length of longer glumes,
this separation being correct (1.e. correlated with other
diagnostic features) 97.3% of the time (Figure 1). Most
recently (Catling 2006) the following key has been rec-
ommended for distinguishing the two subspecies:
Basal internodes red or reddish-purple; longer lower glumes
Sirf Molen MON ooo ecpe ease uc subspecies americanus
Basal internodes pale yellow; longer lower glumes 2.6-4.2
(Giss})) oo ONS gh odudovogec vee subspecies australis
For the study reported here undertaken in 2003, we
used a length of 4.2 mm or more to identify the native
subspecies americanus and 4.1 mm or less to identify
the introduced subspecies australis in a survey of
four adjacent geographic townships (Figure 2): Rus-
sell (45.2833°N, 75.2833°W), Osgoode (45.2333°N,
75.5000°W), North Gower (45.1500°N, 75.6833°W)
and Marlborough (45.0666°N, 75.8166°W). Hybrid-
like or intermediate clones were not observed during
this study but several were located subsequently in
townships to the north.
The survey was conducted in May 2003 at which
time the persistent flowering stalks of the previous year
were readily visible on the landscape and the persist-
ing inflorescences were not so damaged as to prevent
measurement of first glume lengths. The survey in-
cluded driving all roads in each township, checking
known locations and searching larger open wetlands
in Rideau Township on foot. All small wetlands visi-
ble from roads were examined with binoculars. Loca-
tion of populations was recorded with GPS and size of
THE CANADIAN FIELD-NATURALIST
Vol. 120
patches and number of flowering stalks and distance
from road were also recorded. Populations were defined
as less than 500 m apart or 500 m in length. A voucher
inflorescence from one of the tallest flowering stems
was collected from each population and from these
vouchers the length of the longest first glume of 25
examined was recorded. Voucher specimens for some
the collections were deposited in the AAFC [Agricul-
ture and Agri-Food Canada] vascular plant collection
(acronym DAO [Department of Agriculture, Ottawa”’]).
All flowering culms within individual stands were sim-
ilar and were assumed to represent single clones.
Collections in Canadian herbaria from Ottawa area
and Carleton and Prescott counties were identified to
race (using criteria described above) in order to deter-
mine status of the two subspecies in the region in
the past. The historical information was compared to
maps of the occurrence of the native and introduced
Eastern Ontario
Townships surveyed
1. Marlborough
2. North Gower
3. Osgoode
4. Russell
FiGurRE 2. Location of four adjacent townships in eastern
Ontario where native and introduced subspecies of
Phragmites australis were surveyed and mapped in
May 2003. Inset map above shows eastern Ontario
and Great Lakes with Eastern Ontario region shown
in black.
2006
20 km
CATLING AND CARBYN: EUROPEAN COMMON REED IN OTTAWA DISITRICT
309
Phragmites australis
subsp. australis
Phragmites australis
subsp. americanus
FiGurE 3. Distribution of native and introduced subspecies of Common Reed Phragmites australis in four eastern Ontario town-
ships based on a survey in May 2003. Aove, introduced Eurasian subspecies australis. Below, native North American
subspecies americanus.
subspecies in the four townships based on the results
of the 2003 survey.
The relationship to roads was analysed by compar-
ing the distribution of both subspecies by plotting on
township maps showing roads using histograms show-
ing frequency of various distances from roads. In order
to further elucidate the history of spread, the size of
patches was compared on major and minor roads, all
at least 20 years old, in Russell township.
Results
History of Invasion
In Canadian herbaria there are 45 specimens of
Phragmites australis collected in the southern portion
of the Ottawa district (Ottawa-Carleton Regional Muni-
cipality and Prescott-Russell counties). Of these, 21
collected prior to 1970 are referable to the native sub-
species. The earliest collection of the introduced sub-
species was in 1976 at Manotick (Hanes, DAO). Of
the 24 specimens collected after 1970, only five are
referable to the native subspecies, the remainder hav-
ing the relatively short first glumes of the introduced
subspecies. Thus the introduced subspecies appears to
have entered the district in the 1970s and subsequently
assumed dominance but was not recognized as an alien
until 2003 (Catling et al. 2003).
Present Status
Ninety-five populations were recorded in the four
townships during the 2003 survey. Of these, 25 were
referable to the native subspecies and 70 were refer-
able to the introduced subspecies. Both subspecies
310
THE CANADIAN FIELD-NATURALIST
Vol. 120
Ficure 4. Patch of pale stems of P. australis subsp. australis on gravel portion of a roadside and extending in a large clone in
the adjacent ditch and swamp. Highway 417, Russell Township, May 2003.
occurred in all four townships, but the native sub-
species was much more localized, with most popula-
tions in a single wetland (Figure 2).
Invasion Pathway
As seen in the distribution maps (e.g. Figure 3) there
is a strong association of the introduced subspecies with
roads (Figure 4). This is confirmed in the histogram
(Figure 5) where the vast majority of the populations
of the introduced subspecies are within 100 m of a road.
In fact 52% were in the roadside gravel (Figure 4) and
84% were within 3 m of the boundary between vege-
tation and gravel shoulder. Populations of the native
subspecies occurred at greater distances from a road
(Figure 5) with 36% on the shoulder and 60% within
3 m. Although occurrence of the native subspecies on
roadsides was much less, it is clearly spreading from
natural habitats to roadsides. Since both subspecies
may come into close contact on roadsides, there is an
opportunity for the subspecies to hybridize. The intro-
duced subspecies was most frequent in the eastern
townships that have the most roads (Figure 3). The
native subspecies was most frequent in the far western
township which has the least extensive road network
and the most extensive natural wetlands.
Major concentrations of the introduced subspecies
occurred along Highway 417 (upper right in Figure 3),
the busiest and oldest major highway in the region, and
the largest patches in Russell township occurred along
this road, suggesting that this is one of the places where
it first entered the region (Figures 3 and 6). The asso-
ciation of the introduced subspecies with roads sug-
gested spread by dispersal of rhizomes by road vehi-
cles rather than wind-blown seed which would have
resulted in more occurrences at greater distances from
roads. The tough rhizomes extend into the gravel shoul-
ders and even onto the asphalt (Figure 4). They are
readily broken and transported by ploughs, graders and
in the tire treads of many kinds of vehicles (personal
observation). This mechanism of dispersal is believed
to be largely responsible for the rapid spread and pres-
ent strong association with roads.
There is a possibility that spread along roads is
facilitated by relatively high sodium salt (NaCl) con-
centrations, as noted for many other vascular plants
spreading along roadsides (e.g. Catling and McKay
1980, 1981; Reznicek and Catling 1987). Salt concen-
trations may reduce competition by eliminating other
species, thus promoting salt-tolerant species. Although
the alien subspecies may be tolerant of relatively high
salt concentrations, reliance on salty habitats is prob-
ably much less than for other rapidly spreading road-
side plants such as Carex praegracilis (e.g., Reznicek
and Catling 1987). Some patches of the alien sub-
species of Common Reed do occur at great distances
from roads and some roadside patches extend up to
100 m from roads beyond the influence of sodium salt.
2006
80
60
40 | Phragmites australis
subsp. americanus
20
>
6)
i 8
WW
> 0
CG — 1 1 1 as ee a |
wa
~ 80
Ww
60 F
40 Phragmites australis
subsp. australis
20
~The sn a
0) 500 1000 1500 2000
DISTANCE FROM ROAD (m)
FiGurE 5. Histograms showing frequency of distances from
major and minor roads of native and introduced sub-
species of australis in four eastern Ontario townships.
Further evidence for a lack of strong dependence on
sodium salt is the observation that the alien subspecies
of Common Reed occurred along many minor and
gravel roads that do not receive applications of de-
icing salt.
The introduced subspecies has colonized two natu-
ral wetlands in the study area without spreading from
a roadside. One of these stands is 500 m from a road
along a pond shore used extensively as a staging area
by waterfowl. This colonization may have been the
result of transport of rhizomes by the waterfowl, which
occurs with many aquatic plants, but is unusual. Once
in a wetland system the rhizomes are likely to be trans-
ported further by aquatic mammals and birds for build-
ing habitations, platforms and nests. Successful ger-
mination of wind-dispersed seed is also a possibility,
but strong association with roads suggests that it hap-
pens infrequently. Both native and introduced popula-
tions produce apparently viable seed with quantities
varying between poplulations (personal observation,
CATLING AND CARBYN: EUROPEAN COMMON REED IN OTTAWA DISITRICT 31)
10 }
8 |
|
6 | Phragmites australis
} ssp. australis
i
4 minor roads
9 |
>
a5 UF
Z |
PE | Of yess. ee |
is}
= r
uw 8
5
6 Phragmites australis
ssp. australis
4 hwy 417
| |
OF I i
fy a at a fo Se
4000
Tt
T
0 1000 2000 3000
POPULATION SIZE (square m)
FIGURE 6. Histograms showing frequency of clone sizes of
the introduced Phragmites australis subspecies aus-
tralis in relation to major and minor roads in Russell
Township.
Catling). As alien Common Reed increases on the
landscape the frequency of unusual dispersal mecha-
nisms is likely to increase, and spread through wet-
lands may increase substantially.
Future Spread
Considering that it is being spread by road vehi-
cles along corridors of essentially continuous habitat,
including new and disturbed habitats, the rate of spread
can be anticipated to be rapid. Carex praegracilis W.
Boott is believed to have spread the same way through-
out much of eastern North America from the midwest
in only 30 years (e.g., Reznicek et al. 1976; Reznicek
and Catling 1987). Since the introduced subspecies of
Phragmites australis has apparently spread from road-
sides to distances of up to 500 m from roads (Figure
5), it appears that it is not confined to the near road
environment. Although it seems less likely that the
introduced subspecies will invade sensitive wetlands
through introduction by waterfowl, the chances will
increase as it becomes more abundant. There are also
a2
opportunities for spread into isolated wetlands through
transport along tracks of off-road vehicles including
motorcycles, ATVs and snowmobiles.
The introduced subspecies has extensively spread
into both saline and non-saline wetlands and along both
dry and wet roadsides in parts of the United States.
Within another 10-20 years it is expected to line many
major roadways in eastern Canada and it will have
entered many wetlands from adjacent or nearby roads.
As it spreads it is most likely to become established
first along major roads, providing an early warning
of future impact on the nearby landscape.
The introduction of invasive aliens into wetlands by
roads provides another reason to have extensive buffer
zones around wetlands with high protection priority.
Considering that the vegetative spread of P. australis
can be very rapid, and mammals and birds transport
rhizome fragments over substantial distances for nest-
ing, buffers exceeding 1000 m are desirable. Monitor-
ing of the two subspecies will be essential to the pro-
tection of native biodiversity, since early detection of
the introduced subspecies in a sensitive wetland will
offer the best opportunity for control.
Acknowledgments
J. Cayouette, W. J. Cody and F. W. Schueler pro-
vided useful comments.
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tion Bulletin 34(2): 21-26.
Received 26 April 2005
Accepted 17 February 2007
Pack Size of Wolves, Canis lupus, on Caribou, Rangifer tarandus,
Winter Ranges in Westcentral Alberta
GERALD W. Kuzyk!“, JEFF KNETEMAN2, AND FIONA K. A. SCHMIEGELOW!
' Department of Renewable Resources, University of Alberta, Edmonton, Alberta, T6G 2H1 Canada
* Alberta Sustainable Resource Development, Fish and Wildlife Division, Hinton Alberta, T7V 2E6 Canada
3 Present address: Ministry of Environment, Wildlife Science Section, P.O. Box 9338, Provincial Government, Victoria,
British Columbia V8W 9M1 Canada; e-mail: Gerald. Kuzyk@gov.be.ca
Kuzyk, Gerald W., Jeff Kneteman, and Fiona K.A. Schmiegelow. 2006. Pack size of Wolves, Canis lupus, on Caribou, Rangifer
tarandus, winter ranges in westcentral Alberta. Canadian Field-Naturalist 120(3): 131-318.
We studied pack size of Wolves (Canis lupus) on Woodland Caribou (Rangifer tarandus caribou) winter ranges in westcentral
Alberta. These Caribou winter ranges are experiencing increasing pressure from resource extraction industries (forestry, energy
sector) and concerns have been raised regarding increased Wolf predation pressure on Caribou in conjunction with landscape
change. Thirty-one Wolves, from eight Wolf packs, were fitted with radiocollars on two Caribou winter ranges in the Rocky
Mountain foothills, near Grande Cache, Alberta (2000-2001). There was a mean of 8.2 Wolves/pack and between 30 and 39
Wolves on each of the RedRock/Prairie Creek and Little Smoky Caribou ranges. The average pack size of Wolves in this region
does not appear to have increased over that recorded historically, but the range (5-18) in the number of Wolves per pack varied
considerably over our study area. Wolves preyed predominately on Moose (Alces alces), averaging one Moose kill every three
to five days. There was some indication that pack size was related to prey size, with the smallest pack preying on Deer (Odo-
coileus spp.). It was clear that Caribou could not be the primary prey for Wolves, due to their low numbers, and relative to
the pack size and Wolf kills we observed.
Key Words: Wolf, Canis lupus, Caribou, Rangifer tarandus, Moose, Alces alces, White-tailed Deer, Odocoileus virginianus,
predation, pack size, colour phase, kills, Alberta.
Pack size of Wolves (Canis lupus) can provide an
important indication of ungulate abundance (Fuller
1989; Schmidt and Mech 1997; Hayes et al. 2003)
and human impacts on the landscape (Mech 1995).
The number of Wolves in each pack can be related to
prey availability (Fuller 1989) and may be regulated by
dispersal, pup survival and disease prevalence (Mech
1970; Mech et al. 1998). The amount of food available
to each Wolf has been inversely related to pack size
(Thurber and Peterson 1993; Schmidt and Mech 1997),
with smaller pack sizes found in areas of low ungulate
density (Messier 1985). In contrast, pack size may re-
main stable when the amount of food per Wolf decreas-
es, possibly due to a genetic advantage for the breeding
pair when sharing limited food with their offspring
(Schmidt and Mech 1997). Maximum Wolf pack size
may be regulated by behaviors within the pack, such as
social dominance, which can facilitate dispersal. Pack
size has also been related to prey size. Average pack
size of Wolves hunting White-tailed Deer (Odocoileus
virginianus) is generally less than seven (Mech 1970;
Fuller 1989), whereas packs preying on Moose (Alces
alces) are generally larger, having ten or more Wolves
per pack (Peterson et al. 1984; Mech et al. 1998; Hayes
et al. 2003). Vucetich et al. (2004) demonstrated that
larger packs are better able to compete with ravens and
do accrue a foraging advantage. Reports of large Wolf
Packs with over 20 members have been documented in
regions where there is little Wolf harvest (Hayes and
Harestad 2000) or in protected areas (Carbyn et al.
1993; Peterson 1995; Mech et al. 1998; Smith et al.
2004).
Predators may negatively affect endangered prey
species when landscapes become impacted by human
disturbance (Schneider 2001). Landscapes in west-
central Alberta are facing increased pressures from
forest harvesting and the energy sector (oil and gas).
Forest harvesting may alter the movements and distri-
bution of Wolves (Kuzyk et al. 2004) and ungulates
(Smith et al. 2000), and in combination with linear cor-
ridors (roads, seismic lines) from oil and gas activities,
can enhance human access to once remote areas and
possibly increase human-caused Wolf mortality from
shooting and trapping. Enhanced predation by Wolves,
facilitated by landscape change such as increased travel
efficiency on linear corridors (James and Stuart-Smith
2000) and associated increases in alternate prey pop-
ulations (James et al. 2004), is thought to be the pri-
mary factor for Caribou declines in Alberta (Edmonds
1988; McLoughlin et al. 2003). In Alberta, Woodland
Caribou (Rangifer tarandus caribou) are classed as a
threatened species under the provincial Wildlife Act
and on the Species at Rist Act Public Registry
[SARA, http://www.sararegistry.gc.ca/species/Species
Details_e.cfm?sid=636] and special management con-
siderations are necessary to maintain numbers and
habitat (Edmonds 1998; Dzus 2001). More informa-
tion is required on Wolf populations, such as pack sizes,
to be better able to predict long-term changes in the
dynamics of Wolves and Caribou associated with land-
Sule)
314
scape change (Weclaw and Hudson 2004). The objec-
tive of this paper is to present current information on
pack size and associated predation by Wolves that can
be used in decision-making processes for long-term
Caribou conservation.
Study Area
The study area is approximately 5000 square kilo-
metres, located in the foothills of west-central Alberta,
near the town of Grande Cache (54°N 119°W). The
area is classed into subalpine and boreal natural sub-
regions (Beckingham and Archibald 1996), and con-
tains several main rivers but lakes are scarce. Elevations
range from 1300-1800 metres, and the climate is sub-
arctic, with short wet summers and long cold winters.
Temperatures average 16°C in July and -13.5°C in
December (Beckingham and Archibald 1996). The
forests are primarily Lodgepole Pine (Pinus contorta)
and some White Spruce (Picea glauca). The wetlands
support mostly Black Spruce (Picea mariana) and
some Tamarack (Larix laricina). Some south facing
slopes have Aspen (Populus tremuloides) and willow
(Salix sp.).
The study area contains three major Caribou herds:
the Red Rock/Prairie Creek and Al La Peche are moun-
tain herds and represent a migratory ecotype, while
the Little Smoky herd is boreal or sedentary ecotype
(Edmonds 1988). Population size for the mountain
herds is estimated at 600-750 Caribou (Edmonds 1998),
and the Little Smoky herd is estimated at fewer than
100 animals (Smith 2004). The study area also sup-
ports a high diversity of other large mammals. Moose
are found at densities ranging from 0.12 to 0.25 Moose
per square kilometer (Alberta Fish and Wildlife unpub-
lished data). Other ungulates occurring in the area are
Elk (Cervus elaphus), Mule Deer (Odocoileus hemi-
nous), White-tailed Deer, Bighorn Sheep (Ovis cana-
densis), Mountain Goats (Oreamnos americanus) and
Wild Horses (Equus cabalus). Wolves (Canis lupus),
Coyotes (Canis latrans), Grizzly Bears (Ursus arctos),
Black Bears (Ursus americanus) and Cougars (Felis
concolor) also exist throughout the study area.
Wolf Captures and Monitoring
Wolf captures were conducted on two Caribou win-
ter ranges in the winters of 2000 and 2001. Four Wolf
packs were located on the winter range of a migratory
mountain Caribou population (Red Rock/Prairie Creek
herd) and four packs located on the winter range of a
sedentary boreal Caribou population (Little Smoky
herd) (Figure 1). Ungulate baits were placed in strate-
gic locations seven to ten days prior to the initiation of
the Wolf capture operation, to enhance detection of
Wolves. Wolves were located by following trails in the
snow from a fixed-wing aircraft (Mech 1966; Mech
et al. 1998). All Wolf handling was approved by the
Faculty of Agriculture, Forestry and Home Economics
THE CANADIAN FIELD-NATURALIST
Vol. 120
Animal Care Policy (Number 96-99D), subject to the
protocols of the Canadian Council of Animal Welfare.
Wolf captures were accomplished by either helicopter
darting (Ballard et al. 1991) or netgunning, then phys-
ically restraining the Wolf with restraining forks, and
hand-injecting 1-2 mls of telazol at 200mg/ml (Kuzyk
2002). Adults were distinguished from pups by tooth
eruption patterns (Van Ballenberghe and Mech 1975)
whereas yearlings were classed on subjective physical
criteria (size), as there is no definitive method to cat-
egorize yearling Wolves (Mech et al. 1998). Adult
Wolves were fitted with store-aboard GPS (Global
Positioning System) collars (Lotek Engineering Sys-
tems, Newmarket, Ontario or Televilt GPS-Simplex,
Lindesberg, Sweden) or VHF (Very High Frequency)
radiocollars (Lotek Engineering). Pups were fitted
with VHF collars only. All Lotek GPS collars were
equipped with remote dropoff units, intended to release
when signaled from the air. All radiocollared Wolves
were relocated by aerial radiotracking (Mech 1974)
within one to four days of capture to determine if they
had rejoined their packs. Color phases of Wolves were
recorded as gray, black, or white following Dekker
(1986), and adding a category of blue.
Wolf pack size was recorded during aerial radio-
tracking in February and March of each year, when
pack size would be at a minimum (Mech 1970), thus
yielding a conservative estimate. Most packs were
intensively monitored in March of 2000, increasing the
chances of a good count. The best estimates of Wolf
pack size were made when the Wolves were traveling
in single file on a linear corridor or river. Lone Wolves
were assumed to account for 10% of the total popula-
tion (Fuller 1989). We did not extrapolate our pack
size data to determine Wolf density, due to insufficient
temporal data required to adequately determine annual
territory sizes. In addition, our broader project objec-
tives entailed studying Wolves in late winter, corre-
sponding with winter range occupancy by migratory
Caribou. It is the winter range of these caribou that is
currently undergoing industrial development, and thus
of primary conservation concern.
Wolf kills were determined during a two-week peri-
od in March 2000 by aerially locating radiocollared
Wolf packs and finding their ungulate kills (Mech
1974). Flights were conducted twice daily in hopes
of detecting Wolf-killed deer (Fuller 1989). When a
Wolf pack was located, Wolves were counted and the
area searched for ungulate carcasses. A kill was as-
sumed to be caused by Wolves if there were blood-
trails indicating a successful chase and a disarticulated
carcass (Hayes et al. 2000). At each kill site, the number
and behavior of Wolves were recorded (Mech 1966).
All Wolf-killed ungulates that could not be identified to
species, gender and age (adult-calf) from the air were
later ground inspected.
2006
7 eee
ES
Alberta
* Grande Prairie
LYNX CREEK PACK
Grande Cache
Wolf locations (GPS)
Wolf Locations (VHF)
Wolf pack territory
Preliminary Wolf Pack Territroy |
Caribou Management Zones
Major Waterways
Town of Grande Cache
KUZYK, KNETEMAN, AND SCHMIEGELOW: PACK SIZE OF WOLVES
ilometers |
Ficure 1. Distribution of eight Wolf packs on two Caribou winter ranges in west-central Alberta during 2000 and 2001.
TABLE 1. Sex ratio, age class and color phase of Wolves
(n=31) captured in west-central Alberta during 22 January —
13 March, 2000 and 15—17 February, 2001.
Sex Ratio Age Class Color Phases
(n=31) (n=31) (n=31)
14 Males 15 adults 19 black (61%)
17 Females 8 yearlings 8 gray (26%)
8 pups 3 white (11%)
1 blue (2%)
Results
Thirty-one Wolves were captured on two Caribou
ranges: 18 captured in 2000 and 13 in 2001 (Table 1).
Nine Wolves were aerially darted and 24 were netted
(two recaptures). There were no capture-related Wolf
mortalities during this study. A total of 16 GPS (14
Lotek and 2 Televilt) and 17 VHF radiocollars were de-
ployed. There were seven known mortalities of radio-
collared Wolves (23%) over fifteen months: two were
shot and five died of natural or unknown causes. Colors
phases of the radiocollared Wolves were 61% black,
26% gray, 11% white and 2% blue.
Eight Wolf packs were located on the two Caribou
ranges (Tables | and 2; Figure 1). There were between
54 and 77 (mean = 66) Wolves on the Little Smoky
and Red Rock/Prairie Creek Caribou ranges, equating
TABLE 2. Pack size of Wolves on the Red/Rock Prairie Creek
(RPC) and Little Smoky (LSM) Caribou winter ranges in
west-central Alberta in late winters of 2000 and 2001.
Wolf Pack Estimated pack size Caribou herd
Lynx Creek 12-18 RPC
Cutbank 7-8 RPC
Prairie Creek 5-6 RPC
Sheep Creek 6-7 RPC
Simonette 7-11 LSM
Little Smoky 7 LSM
Deep Valley 9 LSM
Berland 8-11 LSM
Total (Range) 54-77
Total (Mean) 65.5
3)
io)
Mean
TABLE 3. Ungulate kills by Wolf packs in west-central Alber-
ta in March 2000.
Wolf Days Moose Deer Ungulate
Pack Monitored kills kills __ kills/day
Lynx Creek 9 (Mar 2-10) 2 1/45
Cutbank 14 (Mar 2-15) 3 1/4.7
Simonette 14 (Mar 2-15) 4 2 1/18
Note: Prairie Creek pack not entered as only one Elk kill was
located.
316
to 8.2 Wolves/pack. Each Caribou range had between
30-39 Wolves with Wolf pack sizes ranging from 5-18
(Table 2). Twelve ungulate kills were recorded from
four Wolf packs during 14 days of monitoring in March
2000 (Table 3). Ungulate kills consisted of seven cow
and two calf Moose, two deer (unknown species) and
one cow Elk. Wolves preyed predominately on Moose,
averaging one Moose-kill every three to five days.
Discussion
We recorded a mean pack size of 8.2 Wolves/pack
for eight Wolf packs on the RedRock/Prairie Creek
and Little Smoky Caribou ranges. This is similar to
the 8.7 Wolves/pack averaged over five other Alberta
Wolf studies conducted between 1975 and 1985 (Gun-
son 1992) and marginally higher than the 6.8 Wolves
per pack recorded in January 1995 in west-central
Alberta during the Yellowstone-Idaho Wolf relocation
(Kneteman 1995*). Pack size of Wolves in our study
area varied considerably, ranging from 5 to 18 Wolves/
pack. The larger Wolf packs (Lynx Creek n = 18; Si-
monette n = 11) preyed primarily on Moose, whereas
the smallest pack (Prairie Creek n = 5) preyed pre-
dominately on deer, consistent with results from other
nearby studies (Carbyn 1974; Weaver 1994). A con-
founding factor in estimating numbers of Wolf-killed
ungulates in west-central Alberta is the difficulty of
detecting Wolf-killed deer (Kuzyk 2002; Kuzyk et al.
2005), due to the small size and cryptic color of deer,
and the short time required for Wolves to consume
deer carcasses (Fuller 1989). All Wolf packs in this
study were observed either hunting deer or at deer
kills. During intensive monitoring in March 2000, the
Prairie Creek pack was seen hunting deer, and was
thought to have made deer kills, but those were never
detected from the aircraft. This resulted in somewhat
ambiguous information, as data indicate this pack
killed only one Elk in nine days of monitoring. But
further analysis of GPS data collected during the nine-
day monitoring period suggested the Prairie Creek pack
had made a minimum of three deer kills (Franke et al.
2006). The importance of deer to Wolves in this study
area should not be underestimated. Kuzyk et al. (2005)
found Wolves in this study area traveled 4.2 times less
when near ungulate carcasses then when away from
them, and suggested that Wolf packs preying primarily
on deer, as opposed to Moose, may pose a greater pre-
dation risk to Caribou due to associated increase travel
and encounter rates. Further research to quantify the
importance of deer in this Wolf-ungulate system should
be initiated such as measuring Wolf encounter rates
with deer (Kunkel et al. 2004).
No Caribou kills were detected during this study,
probably due to the low numbers of Caribou in the
region (Smith 2004) and the short time it takes Wolves
to consume a carcass (Hayes et al. 2000). Nevertheless,
Caribou could not be the primary prey for Wolves in
westcentral Alberta, as the numbers of Caribou could
THE CANADIAN FIELD-NATURALIST
Vol. 120
not support the numbers of Wolves we recorded. For
example, there are fewer than 100 Caribou in the Lit-
tle Smoky herd (Smith 2004), and four Wolf packs
overlap their range, with each pack killing an esti-
mated 40-85 ungulates each winter.
Humans may impact Wolf pack size when access to
remote areas increases, and human-caused mortality
to Wolves increases from shooting, trapping and road
collisions (Mech 1995). Landscapes in west-central
Alberta are being altered by resource extraction indus-
tries which have increased human access to previously
remote areas. We recorded 23% mortality for radio-
collared Wolves over fifteen months, similar to annual
mortality rates (20-27%) found in other studies (Plet-
scher et al. 1997; Mech et al. 1998). This level of mortal-
ity is thought to be low and would not influence over-
all Wolf population size due to the high reproductive
and dispersal rates of Wolves (Mech et al. 1998; Hayes
et al. 2003). Forest harvesting can increase human ac-
cess and change the amount and spatial distribution
of habitats for Wolves and their prey. Wolf packs in our
study area had a seven-fold difference in the amount
of forest harvested within their territories (Kuzyk et al.
2004). The Cutbank pack (n = 7-8) had 36% of the
forest in their territory harvested whereas the Lynx
Creek (n = 12-18) pack had 7% harvested. The lowest
level of forest removal corresponded with the largest
pack size of 18 Wolves. In 2001, this pack contained
12 black and 6 gray Wolves and was observed traveling
on a road in an open forest cutblock (Kuzyk 2001).
This pack size exceeds the largest recorded from
nearby studies: 12 Wolves in the Simonette River area
(Bjorge and Gunson 1989) and 10 Wolves in northern
Jasper National Park (Weaver 1994).
The color phase of a Wolf may influence its detec-
tion by humans (Mech et al. 1998) and subsequent mor-
tality. Radiocollared Wolves in this study were pre-
dominantly black (61%), similar to those recorded in
nearby Jasper National Park (53% black) (Dekker
1986). It is noteworthy that black Wolves may change
to a lighter color possibly due to aging or physiologi-
cal stress (Gipson et al. 2002). Black Wolves may be
more easily detected by hunters, especially during win-
ter and on fragmented landscapes. The two Wolves shot
during our study were black, and of the seven total
mortalities (2 shot, 5 natural or unknown), five Wolves
were black (71%).
Lone and small groups of Wolves are also an im-
portant component of this population. During the two
years of our monitoring, several Wolves dispersed from
their natal territories, as individuals or in small groups
(<3 Wolves), consistent with other research (Gese and
Mech 1991). These lone Wolves, or small groups, could
represent 10-30% of the Wolf population (Fuller 1989;
Mech et al. 1998) and may be an important factor when
assessing predation risk to Caribou. Such Wolves would
be travelling great distances to establish new territories
(Mech 1970), and may use human trails as travel routes
2006
(Kuzyk and Kuzyk 2002), thereby increasing their
chances of encountering Caribou. In addition, pairs of
Wolves may have proportionately higher kill rates than
larger packs (Hayes et al. 2000). Further, if Wolf packs
generally avoid Caribou habitats due to a lack of Moose
(James et al. 2004), dispersing Wolves may select these
habitats to avoid being killed by resident pack Wolves
defending their territories, a primary cause of natural
Wolf mortality (Mech 1994). Conducting research on
single or small groups of Wolves would be logistically
difficult but resulting information could lead to impor-
tant insights into Caribou predation risk from Wolves.
Documenting current pack size and prey relations
of Wolves in our study area is an important step in
understanding the potential implications of landscape
change and resultant alteration of predator/prey sys-
tems for Caribou in the area. However, as pack size
could theoretically remain constant while overall num-
bers increase, determining Wolf density is necessary.
In combination with additional information on kill rates
and prey availability, this would permit evaluation of
the numerical and functional responses of Wolves in
this system to landscape alteration, and provide a foun-
dation for examining alternative management strategies
aimed at long-term Caribou conservation.
Acknowledgments
Funding for this research was provided by the West-
Central Alberta Caribou Standing Committee; the Al-
berta Sport, Recreation, Parks and Wildlife Foundation;
a University of Alberta Challenge Grant in Biodiversity
(supported by the Alberta Conservation Association);
and the Foothill Model Forest. We acknowledge the
safe piloting conducted by C. Wilson from Bighorn
Helicopters and D. Dennison from Coyote Air during
Wolf captures and monitoring. S. Shirkoff and K. Lisgo
provided logistical and data management support. The
spatial data on the figure is published with permis-
sion of Alberta Sustainable Resource Development,
Natural Resource Canada and the Spatial Data Ware-
house. Thanks to K. Smith for helpful comments on
earlier drafts of this paper.
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Received 3 May 2005
Accepted 5 March 2007
Bur Buttercup, Ranunculus testiculatus, New to Eastern Canada
MICHAEL J. OLDHAM!, CLIVE E. Goopwin2, and SEAN BLANEY?
' Ontario Natural Heritage Information Centre, Ministry of Natural Resources, 300 Water Street, P.O. Box 7000, Peterbor-
ough, Ontario K9L 1C8 Canada
* | Queen Street, Suite 405, Cobourg, Ontario K9A 1M8 Canada
4 Atlantic Canada Conservation Data Centre, P.O. Box 6416, Sackville, New Brunswick E4L 1C6 Canada
Oldham, Michael J., Clive E. Goodwin, and Sean Blaney. 2006. Bur Buttercup, Ranunculus testiculatus, new to eastern
Canada. Canadian Field-Naturalist 120(3): 319-322.
Bur Buttercup (Ranunculus testiculatus) is newly reported for eastern Canada based on two collections from campgrounds
in southern Ontario. This vernal, annual, Eurasian weed is widespread in western North America and is expanding its range
in the east; it should be expected elsewhere in eastern Canada. Bur Buttercup is known to be toxic to livestock.
Key Words: Bur Buttercup, Ranunculus testiculatus, Ranunculaceae, first records, weed, range expansion, campground,
Ontario, eastern Canada.
On 17 April 2004, Blaney found an unusual butter-
cup growing on a campsite in Long Point Provincial
Park, on the north shore of Lake Erie, Norfolk County,
Ontario (42.58°North, 80.41° West). A few days later
on 22 April 2004, Goodwin found the same species
growing in a municipal campground in Cobourg, on
the north shore of Lake Ontario, Northumberland Coun-
ty, Ontario (43.96°North, 78.16° West). This distinctive
plant, called Bur Buttercup (Ranunculus testiculatus
Crantz), has not previously been reported from Ontario
(Morton and Venn 1990; Newmaster et al. 1998) or
eastern Canada (Scoggan 1978; Gleason and Cron-
quist 1991; Whittemore 1997; Kartesz 1999).
Ranunculus testiculatus is a small scapose, some-
what tomentose, annual in the buttercup family (Ranun-
culaceae) which flowers early in the season. The leaves
are entirely basal and are 1.5 to 4 cm long, ternate to
biternate, with narrow linear divisions. The several leaf-
less peduncles are 2 to 8 cm tall, and each bears a sin-
gle, inconspicuous pale yellow flower (Figure 1). The
fruiting heads are bur-like when dry and mature, giving
the plant its common name, Bur Buttercup. Superfi-
cially the plant is very different in appearance from
other buttercups and is often placed in a separate genus
as Ceratocephalus testiculatus (Crantz) Roth by some
authors (e.g., Cody 1988; Cusick 1989) or Cerato-
cephala testiculata (e.g., Tutin et al. 1993; Mitchell
and Tucker 1997; Kartesz 1999). However, most recent
North American floras now include it within Ranuncu-
lus (e.g., Gleason and Cronquist 1991; Whittemore
1997). A related species, R. falcatus L., has been report-
1
\
ed from North America, but all reports are apparently
\ based on misidentified material of R. testiculatus (Whit-
_ temore 1997). Common names used for R. testicula-
tus in North America include Bur Buttercup or Bur-
' buttercup (e.g., Barkworth 1982; Pohl 1984; Cusick
_ 1989: Gleason and Cronquist 1991; Swink and Wilhelm
_ 1994), Hornseed Buttercup (e.g., Cody 1988; Douglas
et al. 1999), Sage Buttercup (e.g., Mitchell and Tucker
1997), and Curve-seed-butterwort (e.g., Kartesz 1999).
The first North American report of Ranunculus tes-
ticulatus was from Utah in 1932, followed by discover-
ies in other western states soon after: Oregon in 1938,
Washington in 1940, and Colorado in 1948 (Barkworth
1982). It continues to spread in western North America
(Hitchcock and Cronquist 1973; Taylor 1983; Weber
1990; Whittemore 1997). In Canada, Bur Buttercup has
been known from southern British Columbia for some
time (Boivin 1966; Scoggan 1978), where it is infrequent
in south-central and south-eastern parts of the province
in dry disturbed clearings and sagebrush slopes in the
steppe zone (Douglas et al. 1999). Cody (1988) report-
ed the species new to Saskatchewan based on a collec-
tion from Assiniboia in 1987. Cody (1988) also mapped
the Canadian distribution of R. testiculatus.
Cusick (1989) was the first to report Bur Buttercup
in eastern North America, based on a 1977 collection
from South Bass Island in western Lake Erie, and pre-
dicted it would be found elsewhere in the Great Lakes
region. In New York State it was first collected in
1990 from the parking lot of a campground near Har-
riman in Orange County by D. J. DeLaubenfels (speci-
men in the New York State Museum, NYS A18183,
personal communication C. Sheviak and T. Weldy).
The first Chicago area collection was made in 1991 and
with concerted searching it was found in 12 additional
Chicago region counties in 1992 and 1993 (Swink and
Wilhelm 1994). Other recent new state records are from
Nebraska in 1970 (Macgrath and Weedon 1974),
Iowa in 1984 (Pohl 1984), and Missouri in 1987 (Ladd
and Schuette 1990). Bur Buttercup is considered a
significant toxic weed of grazing lands in the western
United States and has been responsible for the death
of sheep (Olsen et al. 1982, 1983).
Several authors have noted the affinity of this species
for campgrounds, particularly in eastern North Amer-
319
320
THE CANADIAN FIELD-NATURALIST
Vol. 120
Ficure |. Bur Buttercup (Ranunculus testiculatus) plants. Long Point Provincial Park, Ontario, 6 May 2004. Photograph by
Michael Oldham.
ica, for example the first reports from Iowa, Missouri,
New York, Ohio, and the Chicago Region are all from
campgrounds (Pohl 1984; Cusick 1989; Ladd and
Schuette 1990; Swink and Wilhelm 1994; C. Sheviak
personal communication). It seems likely that this
species is dispersed by camping vehicles and associ-
ated equipment. The bur-like fruiting heads of the
buttercup may readily attach to blankets and tents, and
are thus carried to campsites that appear to provide
ideal disturbed conditions. Other vernal, ephemeral
Eurasian weedy species have found a niche in camp-
grounds and are presumed to have a similar dispersal
mechanism, such as Aira caryophyllea (Silvery Hair-
grass), Aira praecox (Early Hairgrass), Apera interrup-
ta (Interrupted Windgrass), Cerastium pumilum (Cur-
tis’ Mouse-ear Chickweed), Cerastium semidecandrum
(Small Mouse-ear Chickweed), Erophila verna (Spring
Whitlow-grass), Myosotis stricta (Blue Scorpion-grass),
Poa bulbosa (Bulbous Bluegrass), and Veronica verna
(Spring Speedwell) (Crins et al. 1987; Oldham et al.
1995; Goltz 1996; Oldham and Zinck 1997). Both
Ontario records are from campgrounds, a municipal
campground in Cobourg, Northumberland County, and
a provincial park campground at Long Point, Norfolk
County. At both Ontario locations Bur Buttercup is
well established but quite local. At Long Point Provin-
cial Park at least 500 plants occur, but they were found
on only a single campsite, possibly suggesting a rela-
tively recent introduction. In addition, the Long Point
peninsula has been relatively well botanized in the
past (Reznicek and Catling 1989). At Cobourg, Bur
Buttercup grew in open, disturbed ground on at least
four campsites with Erophila verna and Taraxacum
officinale (Common Dandelion), while at Long Point
it was found in sandy soil beneath planted Scots Pine
(Pinus sylvestris) with Cerastium semidecandrum, Stel-
laria pallida (Lesser Chickweed), Poa annua (Annual
Bluegrass), Myosotis stricta, and Taraxacum offici-
nale (Figure 2).
Collection data for the Ontario records are as follows:
Ontario, Norfolk County, Long Point Provincial Park,
42.58°North, 80.41°West, 6 May 2004, M. J. Oldham
29983 (DAO, MICH, NHIC, TRTE, UWO).
Northumberland County, Cobourg Municipal Campground,
43.96°North, -78.16°West, 9 May 2004, M. J. Oldham
29996 (DAO, MICH, NHIC, TRTE).
Acknowledgments
Troy Weldy and Charles Sheviak provided infor-
mation on Bur Buttercup in New York state. The
comments of two anonymous reviewers were helpful
in improving the manuscript.
2006
OLDHAM, GOODWIN, and BLANEY: BUR BUTTERCUP 32)
FIGURE 2. Bur Buttercup (Ranunculus testiculatus) habitat at Long Point Provincial Park, Ontario, on 6 May 2004. Bur But-
tercup plants appear as a light coloured area in front of the closest Scots Pine tree in the foreground. Photograph by
Michael Oldham.
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ty Press, New York.
Received 16 May 2005
Accepted 8 February 2007
I
Assessment of Effects of an Oil Pipeline on Caribou, Rangifer tarandus
granti, Use of Riparian Habitats in Arctic Alaska, 2001—2003
LYNN E. Nogv!, MATTHEW K. BUTCHER2, MATTHEW A. CRONIN! “, and BILL STREEVER*
'ENTRIX, Inc., 1600 A Street, Suite 304, Anchorage, Alaska 99501 USA; e-mail: Inoel|@entrix.com
*ENTRIX, Inc., 2701 First Avenue, Suite 500, Seattle, Washington 98121 USA
4Present address: University of Alaska Fairbanks, Palmer Research Center, 533 East Fireweed Avenue, Palmer, Alaska 99645
USA
4BP Exploration (Alaska) Inc., 900 East Benson Boulevard, Anchorage, Alaska 99519 USA
Noel, Lynn E., Matthew K. Butcher, Matthew A. Cronin, and Bill Streever. 2006. Assessment of effects of an oil pipeline on
Caribou, Rangifer tarandus granti, use of riparian habitats in arctic Alaska, 2001-2003. Canadian Field-Naturalist
120(3): 325-330.
Elevated oil field pipelines may alter Caribou (Rangifer tarandus granti) movements and delay or prevent access to insect
relief habitat. In an attempt to determine if the 40-km elevated Badami pipeline in northern Alaska changed Caribou use of
riparian habitats at the three river crossings where the pipeline is buried, we quantified Caribou habitat use at all three crossings
using time-lapse video cameras and aerial distribution surveys over three summers. We compared habitat use, behavior and
duration of observations among pipeline and non-pipeline sites. We used a block experimental design with cameras at four
sites at the three river crossings to evaluate differences in numbers of Caribou per day at pipeline and non-pipeline sites. At
each crossing, four cameras were positioned, with one pair of cameras next to the pipeline (pipeline sites) and one pair of
cameras |.8-3.2 km upstream from the pipeline (non-pipeline sites); where cameras monitored the river bank and channel
(river habitat) and the tundra within about 200 m of the river (tundra habitat). Peak numbers of Caribou per day occurred
during early July 2003 and mid-July 2001 and 2002. Large numbers of Caribou recorded north of the pipeline during aerial
surveys did not usually correspond with increased number per day recorded by cameras suggesting Caribou probably also
crossed the pipeline outside of the riparian areas. We assessed local changes in riparian habitat use by comparing the numbers
of Caribou per day in river and tundra habitats at pipeline and non-pipeline sites and found no difference. We assessed regional
changes in riparian habitat use by comparing numbers of Caribou per day at pipeline sites and at non-pipeline sites and found
no difference. Caribou groups spent an average of | minute longer at tundra pipeline sites and groups spent 30 seconds longer
feeding and trotting at pipeline sites, but these differences were not significant.
Key Words: Caribou, Rangifer tarandus granti, aerial survey, insects, Badami pipeline riparian habitat, summer, time-lapse
video, Alaska.
In northern Alaska, Caribou (Rangifer tarandus
granti) regularly use insect relief habitats during the
summer (Cameron and Whitten 1979; Murphy and
Curatolo 1987; Walsh et al. 1992; Pollard et al. 1996a, b:
Young and McCabe 1998). This frequently involves
north-south movements from inland feeding habitats
to coastal insect relief habitats (Cameron and Whitten
1979; Pollard et al. 1996b). Oil transport pipelines tra-
verse open tundra and riparian corridors, often in an east-
west direction, so there is the potential for obstruction
of movements to and from insect relief habitats (Nation-
al Research Council 2003). Previous work has shown
that pipelines associated with roads can have an impact
Caribou movements (Murphy and Lawhead 2000);
however, impacts from pipelines not associated with
roads and elevated = 1.5 m above the tundra, a height
designed as a mitigation to allow Caribou passage
(Cronin et al. 1994"), have not been documented.
One such pipeline on Alaska’s north slope runs 40 km
from the Badami oil production site to the oil fields at
Prudhoe Bay. Most of the Badami pipeline is elevated
about 1.5 m above the tundra, but it is buried under-
neath the three rivers it crosses (Figure | and 2). Cari-
bou are common in this area during the summer, and
aerial surveys document animals on the north and south
sides of the pipeline, indicating that they cross the pipe-
line (Jensen and Noel 2002°: Jensen et al. 2003°; Noel
and Cunningham 2003"). However, the elevated sections
of this pipeline could alter Caribou movement and delay
access to coastal habitats (National Research Council
2003). If this occurs, Caribou should be more likely to
cross the pipeline in the riparian zones (including the
river and adjacent tundra) where the pipeline is buried.
We recorded Caribou by using time-lapse video and
aerial distribution surveys over three years to assess
whether Caribou use of riparian habitats was influ-
enced by the pipeline.
Methods
Study Area
A 40-km elevated pipeline extends from the Badami
facility (70°9'2.71"N, 147°1'25.93"W) across undevel-
oped terrain to the Endicott pipeline near the Prudhoe
Bay oil field (70°15'18.25"N, 148°1'32.75"W, Figure
323
Vol. 120
10
—— ; ? = jap Area
---- Pipelines Led
— Roads i
Bl Gravel pads and mines
(5) Aerial Survey Area
THE CANADIAN FIELD-NATURALIST
FIGURE 1. Study area, North Slope, Alaska. Arrows indicate study sites.
1). The Badami pipeline is elevated = 1.5 m above the
tundra surface for > 99% of its length. This pipeline
is buried beneath the East Channel of the Sagavanirk-
tok, Shaviovik, and Kadleroshilik rivers. At the Saga-
vanirktok and Shaviovik rivers, the pipeline is buried for
about 1000 m; at the Kadleroshilik River the pipeline
is buried for about 400 m. The Sagavanirktok River
crossing is split into two channels by a large vegetat-
ed river bar; requiring two sites to be monitored. The
tundra between these river channels is a gently rolling
thaw-lake plain landscape with elevation rises of 6 to
8 m (Walker and Acevedo 1987).
Time-Lapse Videography
We used time-lapse video camera assemblies to
monitor Caribou movements between 19 June and 26
August in 2001, 2002, and 2003. Each camera assem-
bly consisted of aGYYR™ TLC1800-DC time-lapse
videocassette recorder and a Panasonic™ WV-CL 322
color CCTV digital camera equipped with a Compu-
tar™ APC auto-iris 8.5 mm semi wide-angle lens (Pol-
lard and Noel 1994"; Noel et al. 1998). Each assembly
was powered by four 12-volt, 80-amp sealed lead acid
batteries, charged by four Solarex™ SX-56 photovolta-
ic panels. The video recorder, camera, and batteries
were housed in insulated aluminum casings to protect
them from weather and animals. The time-lapse video
recorders were set to record at 34 second intervals.
Markers were placed 100 m from each camera to assist
kilometers
Badami Pipeline y a
MIKKELSEN
BAY
BADAMI
in determining visibility and to standardize the area
sampled. Cameras had a 72° view angle resulting in a
7260-m*? field of view out to 100 m (i.e., the sampling
area). Videotapes were changed at 10—13 day intervals.
The cameras were arranged as follows at four river
crossings (Figure 1): (1) the east side of the buried East
Channel Sagavanirktok crossing (E Sag), (2) the west
side of the buried East Channel Sagavanirktok cross-
ing (W Sag), (3) the east side of the Shaviovik River
crossing (Shav), and (4) the east side of the Kadlero-
shilik River crossing (Kad). At each crossing, four cam-
eras were positioned, with one pair of cameras next to
the pipeline (pipeline sites) and one pair of cameras
1.8-3.2 km upstream from the pipeline (non-pipeline
sites) (Figure 2). One of each pair of cameras moni-
tored the river bank and channel (river habitat), while
the other monitored the tundra within about 200 m of
the river (tundra habitat).
Non-pipeline sites with river channel width and con-
figuration and tundra habitats similar to those of the
corresponding pipeline sites were selected to reduce
variability. This arrangement allowed us to collect Cari-
bou data in four settings at each river crossing (Fig-
ure 2): (1) along the pipeline corridor where the pipe-
line was buried under the river (buried pipeline site/river
habitat), (2) along the pipeline corridor where the pipe-
line was elevated above the tundra (elevated pipeline
site/tundra habitat), (3) 1.8-3.2 km upstream from the
2006
pipeline at the river’s edge (non-pipeline site/river habi-
tat), and (4) upstream from the pipeline on the tundra
(non-pipeline site/tundra habitat).
Videotapes were viewed and the number of Caribou
groups seen each day by each of the cameras was re-
corded. Caribou groups were defined by lapses of 15-
30 seconds of tape with no Caribou observations. For
each group observed within 100 m of the camera, data
for the number of individuals, sex/age category, pre-
dominant behavior, and direction of movement were
recorded. Caribou group behaviors included: feeding,
resting (laying), standing, walking, trotting, running,
and swimming.
Aerial Surveys
Eleven systematic aerial strip-transect surveys were
completed from a Cessna 206 fixed-wing aircraft
(Caughley 1977) to document the number of Caribou
within a 1043 km? area surrounding the Badami pipe-
line between 25 June and | August 2001-2003 (Fig-
ure 1) (Jensen and Noel 2002*; Jensen et al. 2003*;
Noel and Cunningham 2003*). Transect centerlines
were spaced at 1.6-km intervals, oriented north-south,
and centered on township and section lines from | :
63360 scale U.S. Geological Survey (USGS) topogra-
phic maps. Transects were flown at 90 m altitude and
130-180 km/h. Two observers, each searching an 800-
m wide area on their side of the transect centerline,
provided 100% coverage of the survey area. Aircraft
wing struts were marked to enable visual control of
transect strip width and estimation of distance between
Caribou groups and the survey aircraft (Pennycuick
and Western 1972). Global positioning system (GPS)
receivers were used for navigating the aircraft along
transects and for estimating the location of the aircraft
when animals were observed. The locations of ani-
mals were recorded using a GPS receiver linked to a
notebook computer. For each sighting, species, group
size, group composition, and perpendicular distance
from the aircraft were recorded. Coordinates of animal
sightings were later calculated using the visual esti-
mates of distance from the aircraft to offset the GPS
aircraft positions. Geographic Information System (GIS)
software was used to complete geographic summaries.
Weather Data
An automated weather station was established near
the East Channel Sagavanirktok River. Temperature
(T100 probe with radiation shield) and wind (Gill 3-
cup anemometer, Gill low threshold vane) sensors trans-
mitted readings at 5-min intervals to a data recorder
which averaged and stored values as mean hourly wind
speed and mean hourly air temperature data (Dryden
R2® data logger, Anchorage, Alaska).
Data Analysis
The cameras were arranged to create a complete
block experimental design (Steel and Torrie 1980).
Each river location (E Sag, W Sag, Kad, or Shav) was
a complete block containing all four settings: buried
NOEL ET AL: EFFECTS OF OIL PIPELINE ON CARIBOL 325
FIGURE 2. Top: An elevated pipeline on Alaska’s North Slope.
Bottom: Schematic showing the positions of four
cameras relative to a river and the Badami Pipeline. As
shown, the pipeline is buried at river crossings. (Draw-
ing by Daniel King not to scale.)
pipeline site/river habitat, elevated pipeline site/tundra
habitat, non-pipeline site/river habitat, and non-pipe-
line site/tundra habitat, resulting in two cameras at
the pipeline site and two cameras at the non-pipeline
site (Figure 2). The potential for the pipeline to change
Caribou habitat use was assessed by comparing mean
numbers of Caribou per day (averaged for each year)
among the four settings, and by evaluating the direc-
tion, duration and behavior of Caribou observed by the
time-lapse cameras. We summarize the aerial survey
data to compare the numbers of Caribou recorded us-
ing time-lapse cameras to the numbers of Caribou with-
in the survey area, north of the pipeline and within ripar-
ian habitats. Riparian habitats were defined as a 200 m
area surrounding the Sagavanirktok, Kadleroshilik,
and Shaviovik rivers (Figure 1). Observation duration,
direction of movement, behavior, temperature, and
wind speed were compared among habitats and with
and without the pipeline using a general linear model
and analysis of variance (ANOVA).
The block design allowed for the evaluation of
local (within the riparian zone) and regional changes
in Caribou use of riparian habitats. At a local scale, if
Caribou changed habitat use due to the elevated pipe-
line within the riparian zone (i.e., within 200 m of the
river bank), we would expect more Caribou to use river
habitat at the pipeline sites compared to river habitat
at the non-pipeline sites, as Caribou would avoid the
elevated portion of the pipeline by crossing at the river
where the pipeline was buried. At a regional scale, if
Caribou moved toward the riparian corridors along
the pipeline corridor, we would expect to see more
Caribou at the pipeline sites (both river and tundra habi-
tats) than at the comparable non-pipeline sites (both
river and tundra habitats), as blockage of direct north-
south movements by the elevated pipeline would lead
to east-west Caribou movements along its length.
We used complete block ANOVA to assess local and
regional changes in Caribou use of riparian habitats.
Because of the lack of independence between counts
of Caribou on consecutive days, we averaged the num-
ber of Caribou per day across each year, and used this
mean as the response variable, giving an overall sam-
ple size of n=47. The ANOVA model was:
Caribou per day = Constant + Location + Year + Pipeline
+ Habitat + (Pipeline x Habitat),
where:
Constant = overall mean Caribou per day
Location = a block for each river crossing (W Sag, E Sag,
Kad, or Shav),
Year = each year of the study (2001, 2002, and 2003)
Pipeline = pipeline site or non-pipeline site, and
Habitat = river or tundra.
All statistical analyses were conducted with SYS-
TAT® Version 10.2 (SYSTAT® Software Inc., Rich-
mond, California).
Results and Discussion
Peak numbers of Caribou per day were recorded dur-
ing early July in 2003 and mid July in 2001 and 2002
(Figure 3). Peaks in mean daily temperature coincided
with increased numbers of Caribou per day recorded
by cameras during late June and early July, but trends
were not consistent. During the 11 aerial surveys be-
tween 25 June and | August 2001-2003, 36% of the
Caribou were distributed north of the pipeline within
13% of survey area and 17% of the Caribou were with-
in riparian habitats representing 9% of survey area
(Table 1). Generally, when few Caribou were record-
ed within the survey area, no Caribou were recorded
by the time-lapse cameras (Table 1, Figure 3).
The sex-age distribution of Caribou occuring within
the survey area was dominated by adult cows (61%-—
70%) based on the aerial survey data (Table 1). These
sex-age distributions are estimates and likely under-
represent calves based on the results of fall composi-
tion surveys for this herd of 28% bulls, 42% cows and
THE CANADIAN FIELD-NATURALIST
Vol. 120
30% calves (Lenart 2003*). Sex-age distributions based
on time-lapsed video were more similar to fall com-
position results (Lenart 2003*), but nearly half of the
individuals were unclassified (Table 1).
Mean annual Caribou per day by camera ranged
from 0.3—20.4 across habitats, pipeline configurations,
river locations, and years (Table 2). Least squares means
from the ANOVA show that Caribou were more abun-
dant in tundra habitats (mean = 8.9 Caribou per day)
than in river habitats (mean = 3.3 Caribou per day)
(Table 3). This difference between habitats was the
only significant factor (P = 0.02, Table 4).
If local habitat use changed because of the presence
of the elevated pipeline, more Caribou would have
been seen in river habitat than in tundra habitat at the
pipeline sites. This local effect was reflected in the
ANOVA Habitat x Pipeline interaction term, which
was not significant (P = 0.64, Table 4). The difference
between mean Caribou per day in tundra and river
habitats (7.6 and 3.2 Caribou per day, respectively) was
smaller for pipeline sites than for non-pipeline sites
(10.4 and 3.5 Caribou per day, respectively).
Because we detected no effect on numbers of Cari-
bou per day, we evaluated the duration that Caribou
were recorded at pipeline and non-pipeline sites on the
tundra (Table 5). Blockage of northward or southward
movements across the pipeline corridor could result in
delays at pipeline sites, which would be reflected in
duration and behavior at these sites. Groups moving
north appeared to spend an average of one minute longer
at pipeline sites in tundra habitats, while groups mov-
ing south spent about two minutes longer at non-pipe-
line sites (Table 5). A general linear model was used
to test whether the presence of the pipeline affected
observation duration for Caribou moving northward
potentially seeking refuge from insects. Neither the
presence of the pipeline nor the explanatory variables
habitat (river versus tundra), temperature, and wind
speed (atmospheric conditions may have an effect on
the severity of insect harrassment) had statistically
significant (P = 0.05) effects on the observation dura-
tion for Caribou moving northward. Duration of group
behaviors indicated that Caribou spent about 30 sec-
onds longer feeding and trotting on the tundra at pipe-
line sites than non-pipeline sites (Table 6). Separate
ANOVAs were applied to the duration of feeding, trot-
ting, and walking activities in different habitats (river
and tundra) and in the presence or absence of the pipe-
line. For trotting and walking, neither habitat nor pipe-
line or their interaction were significant explanatory
variables (P > 0.2). For feeding Caribou, the duration
within tundra (mean = 00:01:09) was significantly
longer than within river (mean = 00:06:35, P = 0.002).
If the pipeline caused regional changes in Caribou
riparian habitat use, we would expect to see more Cari-
bou at the pipeline sites than at the non-pipeline sites.
However, the difference between mean Caribou per
day at the pipeline and non-pipeline sites (5.4 and 6.8
Caribou per day, respectively, Table 3) was not signifi-
2006 NOEL ET AL: EFFECTS OF OIL PIPELINE ON CARIBOI 327
- Caribou per day
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FiGureE 3. Comparison of numbers of Caribou recorded per day for all time-lapse cameras, numbers of Caribou recorded in
riparian habitats north of the Badami pipeline corridor during aerial surveys (Figure 1), and mean daily temperature
(°C), 19 June (day 170) 26 August (day 238) 2001-2003, North Slope, Alaska.
328
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Summary of Caribou sex-age classes, numbers, and distribution within the aerial survey area and observed in riparian
areas at pipeline and non-pipeline sites using time-lapse video 19 June—26 August 2001—2003, North Slope, Alaska. For
bulls, cows, and calves % is per cent of classified. Unclassified % is per cent of total.
Total Total Percent
Bulls Cows Calves Unclassified Caribou Groups _ of Total
Aerial Surveys
Survey Area (1043 km?) 2846 12% 16001 65% 5775 23% 5798 19% 30420 7715
North of Pipeline (136 km?) 1479 14% 7492 70% 1807 17% 287 3% 11065 134 36%
Riparian Zone (98 km?) 811 21% 2392 61% 722 18% 1222 24% 5147 111 17%
Time-Lapse Video
All Sites (1293 days) 1352 36% 1352 36% 1005 27% 4165 53% 7874 635
Pipeline Sites (660 days) 664 37% 659 37% 457 26% 1816 51% 3596 352 46%
Non-Pipeline Sites (633 days) 688 36% 693 36% 548 28% 2349 55% 4278 283 54%
TABLE 2. Total Caribou, days of camera operation, and mean Caribou per day by camera with standard deviation (SD) for
all three years pooled, 19 June—26 August 2001—2003, North Slope, Alaska.
Camera Site
Grand Total
Habitat
Tundra
River
Tundra
River
River
Tundra
River
Tundra
River
Tundra
River
Tundra
River
Tundra
River
Tundra
Pipeline
Elevated
Buried
Non-pipeline
Non-pipeline
Buried
Elevated
Non-pipeline
Non-pipeline
Buried
Elevated
Non-pipeline
Non-pipeline
Buried
Elevated
Non-pipeline
Non-pipeline
Total Days of camera Caribou per
Caribou operation day (SD)
562 89.88 5.8 (35.32)
DD 87.62 0.2 (1.47)
488 86.88 Sli@5e70)
8 88.67 0.1 (0.82)
243 78.60 2.8 (14.56)
1122 77.88 12.9 (63.03)
320 81.23 3.6 (19.21)
596 77.78 6.7 (29.63)
151 79.67 1.6 (7.19)
234 88.42 2.5 (13.54)
591 84.98 6.5 (36.29)
1222 59.81 19.1 (77.81)
629 80.20 7.1 (27.53)
633 78.00 13 (C2)
DD 76.32 2.6 (14.29)
828 76.93 9.5 (48.67)
7874 1292.87
TABLE 3. Caribou per day by pipeline and habitat with sites
pooled (buried pipeline site/river habitat, elevated pipeline
site/tundra habitat, non-pipeline site/river habitat, non-pipeline
site/tundra habitat), 19 June—26 August 2001-2003, North
Slope, Alaska.
Habitat
Pipeline River Tundra Total
Pipeline 3.2 7.6 SD)
Non-pipeline 3.5 10.4 6.8
Total Shes) 8.9 6.1
cant (P = 0.57, Table 4). Peaks in the number of Cari-
bou north of the pipeline in 2002 and 2003 did not
coincide with large numbers of Caribou recorded at
cameras, suggesting that Caribou crossed the pipeline
outside of riparian areas (Figure 3).
The results of this study support the conclusion that
pipelines elevated =1.5 m above the tundra did not
cause changes in Caribou riparian habitat use at buried
river crossings or delay north-south movements to and
from coastal insect relief habitats (Cronin et al. 1994*,
Murphy and Lawhead 2000).
TABLE 4. Results of ANOVA for Caribou per day along three rivers crossed by the Badami pipeline during 19 June—26
August 2001-2003, North Slope, Alaska.
Source Sum-of-Squares df Mean-Square F-ratio IP
Pipeline (Present, Absent) 22.08 1.00 22.08 0.33 0.57
Habitat (River, Tundra) 415.86 1.00 415.86 6.29 0.02
Location (i.e., Block) (E Sag, W Sag, Kad, Shav) 170.90 3.00 56.97 0.86 0.47
Year (2001, 2002, 2003) 29.50 2.00 14.75 0.22 0.80
Interaction (Pipeline x Habitat) 14.40 1.00 14.40 0.22 0.64
Error 2512.10 38.00 66.11
329
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330
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the range of the Central Arctic Herd. Part A: Aerial surveys
in the Milne Point Unit, Prudhoe Bay Oilfield, Badami, and
Bullen Point to Staines River study areas, summer 2001.
Chapter 2A in Arctic Coastal Plain caribou distribution,
summer 2001. Edited by M. A. Cronin. Unpublished report
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pages plus appendices.
Lenart, E. A. 2003. Unit 26A and B Caribou management
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survey and inventory activities 1 July 2000-30 June 2002.
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parasitic insect abundance in the Prudhoe Bay oil field,
summer 1993. Report to BP Exploration (Alaska) Inc. by
LGL Alaska Research Associates, Inc., Anchorage, Alaska.
70 pages.
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Received 17 August 2005
Accepted 19 April 2007
First Records of the Southern Red-backed Vole, Myodes gapperi, in
the Yukon
THOMAS S. JUNG!, AMY M. RUNCK?2, DAvip W. NAGORSEN?, BRIAN G. SLOUGH*, and Topp PoweLL>”
'Yukon Department of Environment, Fish and Wildlife Branch, P.O. Box 2703, Whitehorse, Yukon Y!1A 2C6 Canada;
e-mail: thomas.jung@ gov.yk.ca
Department of Biological Sciences, Idaho State University, 650 Memorial Drive, Pocatello, Idaho 83209-8007 USA
4Mammialia Biological Consulting, 4268 Metchosin Road, Victoria, British Columbia V9C 324 Canada
435 Cronkhite Road, Whitehorse, Yukon Y1A 5S9 Canada
‘Yukon Department of Environment, Fish and Wildlife Branch, P.O. Box 194, Watson Lake, Yukon YOA 1CO Canada
°Current address: Alberta Department of Sustainable Resource Development, Fish and Wildlife Division, P.O. Box 9915,
Fort McMurray, Alberta T9H 2K4 Canada
Jung, Thomas S., Amy M. Runck, David W. Nagorsen, Brian G. Slough, and Todd Powell. 2006. First records of the Southern
Red-backed Vole, Myodes gapperi, in the Yukon. Canadian Field-Naturalist 120(3): 331-334.
Twenty Southern Red-backed Voles, Myodes gapperi, were collected in July 2004 in the LaBiche River valley of southeast-
ern Yukon. Specimens were identified using morphological characteristics and analysis of cytochrome b gene sequences.
These are the first records of this species in the Yukon. No Northern Red-backed Voles, M. rutilus, were collected and it is
not known whether the two species are sympatric or parapatric in the Yukon. Further survey work is needed in southeastern
Yukon to better delineate the extent of the northwestern range of this species and the extent, if any, of introgression with M.
rutilus.
Key Words: Southern Red-backed Vole, Myodes gapperi, distribution, Yukon.
The Southern Red-backed Vole (Myodes [formerly
Clethrionomys see Musser et al. 2005] gapperi) is
broadly distributed across the boreal, montane, Pacific
coastal, and other conifer-dominated forests of North
America (Banfield 1974; Merritt 1981; Batzli 1999).
Youngman (1975) did not report M. gapperi from the
Yukon, in part, because he was of the opinion that M.
gapperi and the Northern Red-backed Vole (M. rutilus)
were conspecific. Cook et al. (2004), however, provided
molecular data that reinforced the separate status of M.
gapperi and M. rutilus. North American species of
Myodes, including M. gapperi and M. rutilus, are cryp-
tic and field identification is highly problematic where
congeners come into contact (Merritt 1981; McPhee
1984; Runck 2001, Nagorsen 2002, 2005; Cook et al.
2004). Therefore, we considered that M. gapperi may
have been overlooked in the Yukon due to the difficulty
in distinguishing specimens from M. rutilus in the field,
and a general paucity of mammalian diversity surveys
in most of the Yukon. Based on the distribution of M.
gapperi in adjacent northeastern British Columbia and
southwestern Northwest Territories (Banfield 1974;
Nagorsen 2005) we suspected the greatest likelihood
of finding the species in the Yukon was in the Liard
River Watershed. In summer 2004, we undertook a
brief field survey of mammalian diversity in the bore-
al forest of southeastern Yukon, partially with an aim
to procuring specimens of M. gapperi. Here, we pro-
vide the first records of the Southern Red-backed Vole
in the Yukon.
Methods
On 28 July — 2 August 2004, we used pitfall and
snap traps to collect shrews and rodents in the LaBiche
River Valley (60.126°N, 124.064°W) of southeastern
Yukon. Seven habitats were sampled: wet meadows and
shrub thickets adjacent to Beaver (Castor canadensis)
ponds; xeric grassy meadows; riparian old-growth
White Spruce (Picea glauca) forest; lowland Black
Spruce (Picea mariana) forest; subalpine old-growth
Spruce-fir (Abies lasiocarpa) forest; second-growth
mixedwood forest; and regenerating clearcut forest.
All habitats were sampled with snap traps. We estab-
lished 14 variable length traplines (ca. 100 m— 250 m)
in each of the seven habitat types. Trap stations were
set 10 m apart on the traplines. One Museum Special
snap trap and one Victor snap trap (Woodstream, Lititz,
Pennsylvannia) were set at each trapping station. Traps
were baited with oats and peanut butter, run for 2-3 days
and checked each morning. In addition to the snap
traps, we established three traplines of metal conical
pitfall traps in the wet meadow habitats, primarly to
capture soricids. Pitfall traps were 25 cm in height and
15 cm in diameter at the top. Pitfall traps were also
arranged in traplines with one trapline in each of the
wet meadows sampled. Traps were spaced about 10 m
apart, with 15 or 20 traps per trapline, and installed
flush with the ground surface. Pitfall traps were not
baited and they were left open for 3-5 days. Each mom-
ing we checked the traps and processed the captures.
331
332
Captures were tentatively identified to species, mea-
sured, aged, weighed, sexed, and examined for repro-
ductive condition. Whole carcasses were frozen on
site. Species were tentatively identified in the field us-
ing morphological, pelage, and dental characteristics,
following Nagorsen (2002) and van Zyll de Jong
(1983). Because of the difficulty in distinguishing M.
rutilus from M. gapperi in the field, we undertook sub-
sequent laboratory investigations at Idaho State Uni-
versity (Pocatello, Idaho) to confirm our field identifi-
cations. All skulls were examined for fusion of the
post-palatal bridge (Merritt 1981; Nagorsen 2002) and
the first 600 base pairs (bp) of mitochondrial cyto-
chrome b gene were examined to confirm species iden-
tification. Restriction enzyme screening and sequencing
of cytochrome b followed established protocols for
Myodes (Runck and Cook 2005). All individuals un-
derwent restriction enzyme digestion using the restric-
tion enzyme ALU I to determine the presence of the
restriction site AGCT which is found in the 600 bp frag-
ment of the cytochrome b gene in M. rutilus but not
M. gapperi (Runck 2001). Five random individuals
were sequenced for the first 600 bp of the cytochrome
b gene and were used in a phylogenetic reconstruction
using published sequences of Myodes (Cook et al.
2004; Runck and Cook 2005), representing M. gapperi
(n= 10), M. rutilus (n = 2), M. californicus (n= 1), M.
glareolus (n= 1), and M. rufocanus (n = 1). Sequences
for specimens not collected in LaBiche (Figure 1) were
taken from GenBank. Phylogeographic reconstruction
was conducted using neighbor-joining framework in
PAUP b10 (Swofford 2002) using the Tamura and Nei
(1993) model of sequence evolution as determined
through Modeltest (Posada and Crandall 2000).
Results and Discussion
We captured 20 Myodes during sampling. Field
identification, based on morphological (tail and pelage)
characteristics, suggested that all captures were of M.
gapperi. Further, 15 of the 20 specimens had complete
post-palatal bridges, which is a primary diagnostic
characteristic of M. gapperi (Merritt 1981; Nagorsen
2002). Two specimens had incomplete post-palatal
bridges and the bridges of a further three were broken
by the traps and not usable as a diagnostic character.
Genetic analyses using restriction enzyme screening
and sequencing of cytochrome b gene confirmed that
all 20 specimens of Myodes collected in the LaBiche
River Valley were M. gapperi (Figure 1).
The Southern Red-backed Vole is now added to the
list of mammals in the Yukon. This is the first addition-
al rodent found in the Yukon since Youngman (1975).
These 20 specimens were deposited at the Museum of
Southwestern Biology at the University of New Mex-
ico (Albuquerque, New Mexico).
Capture rates were 0.28 per 100 trap nights (TN)
and 1.32 per 100 TN in pitfall and snap traps, respec-
THE CANADIAN FIELD-NATURALIST
Vol. 120
tively. Captures of M. gapperi represented the range of
sex and age classes (5 adult females, 7 adult males,
5 juvenile females, and 3 juvenile males), confirming
that a breeding population was present. M. gapperi were
taken in all of the habitats sampled. Species caught in
association with M. gapperi in the LaBiche River Val-
ley included: Meadow Voles (Microtus pennsylvani-
cus, n = 14), Deer Mice (Peromyscus maniculatus,
n = 13), Meadow Jumping Mice (Zapus hudsonius,
n= 10), Eastern Heather Vole (Phenacomys ungava,
n= 1), Masked Shrew (Sorex cinereus,n = 1), Pygmy
Shrew (Sorex hoyi, n = 1), and Dusky Shrews (Sorex
monticolus, n = 2).
We do not know the extent of the range of the South-
ern Red-backed Vole in the Yukon. They have been
collected, however, near Fort Liard, Northwest Terri-
tories (S. Carriére, unpublished data; Runck and Cook
2005). Specimens from near Watson Lake, Yukon (ca.
270 km W of the present study; n = 43), were also
analyzed in the lab using skull and genetic characters
and they were determined to be M. rutilus (T. S. Jung
and A. M. Runck, unpublished data), suggesting that
M. gapperi may not be widely distributed in southern
Yukon. Perhaps M. gapperi is restricted in the Yukon
to the extension of the Boreal Plain ecoregion in the
extreme southeast. Further surveys from nearby river
valleys, and subsequent DNA analysis, are needed to
better document the ranges and contact zone of M. gap-
peri and M. rutilus in southeastern Yukon and adjacent
British Columbia and Northwest Territories. Our dis-
covery of M. gapperi in extreme southeastern Yukon,
along with that of the Yukon’s only records of Northern
Long-eared Bats (Myotis septentrionalis; Jung et al.
2005) and other taxa (e.g., some species of neotropi-
cal migrant passerines: Eckert et al. 1997), suggests
that this ecoregion may be a zoogeographically unique
part of the Yukon.
Acknowledgments
The staff of Devon Canada Corporation’s LaBiche
River Gas Plant kindly went out of their way to accom-
modate us and provide logistical support. Funding was
provided by the Yukon Department of Environment,
NatureServe Yukon, and a Northern Research Endow-
ment Grant from the Northern Research Institute, Yukon
College, to B. Slough. Thoughtful comments from S.
Cannings, and five anonymous reviewers improved
the manuscript.
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2006 JUNG ET AL: SOUTHERN RED-BACKED VOLE IN THE YUKON 333
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Fort Liard, Northwest Territories AY952147
Swan Lake, British Columbia AY952158
85 LaBiche DQ152249
LaBiche DQ152250
LaBiche DQ152251
Fort Liard, Northwest Territories AY952146
High Latitude
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93
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M. glareolus Finland AY119272
400 M. rutilus Alaska AY309426
M. rutilus Alaska AY309427
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Received 12 September 2005
Accepted 4 September 2007
ee
A Conservation Evaluation of Smooth Goosefoot, Chenopodium
subglabrum (Chenopodiaceae), in Canada
DIANA BIZECK!I ROBSON
The Manitoba Museum, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2 Canada
Robson, Diana Bizecki. 2006. A conservation evaluation of the Smooth Goosefoot, Chenopodium subglabrum (Chenopodi-
aceae), in Canada. Canadian Field-Naturalist 120(3): 335-341.
Smooth Goosefoot (Chenopodium subglabrum) is restricted to North America and reaches its northern distribution limit in
Alberta, Saskatchewan and Manitoba. The habitat of Chenopodium subglabrum contains some element of active sand. It is
commonly found on the stabilizing edges of active dunes as well as dune blowouts, and occasionally on bare or recently dis-
turbed sand plains. Chenopodium subglabrum is a sexually reproducing annual species with seeds that may remain in the
seed bank for several years, waiting for the conditions it favours for germination. Germination of this species is erratic, pos-
sibly in response to moisture, making the overall population trend difficult to determine. The estimated population in Cana-
da is likely between 5000 and 10000 individuals. There has been considerable loss of habitat as dunes become vegetated.
The processes of dune stabilization and fire control threaten survival of this species.
Key Words: Chenopodium subglabrum, Smooth Goosefoot, distribution, population size, rare, Alberta, Saskatchewan, Manitoba.
Chenopodium subglabrum (S. Wats.) A. Nels., com-
monly called Smooth Goosefoot, is a rare, shallow-
rooted annual plant (Figure 1) found in the prairie
provinces of Canada. Prior to 1990 only small num-
bers of this species had been observed at 20 sites in 4
Canada (Smith and Bradley 1990*). Search efforts Jf 2 OP \ (/ '
( Za
a
from the mid-1990s on resulted in the discovery of
new populations in five Sand Hill regions in Saskat-
chewan and Manitoba. In 2004, the highest number
of individuals ever recorded in Canada was observed,
suggesting that this species has a strong temporal com-
ponent to its rarity. The objective of this paper is to doc-
ument these recent findings and determine their con-
servation implications.
a
Habitat
Chenopodium subglabrum populations occur in the
Aspen Parkland, Moist Mixed Grassland, and Mixed
Grassland Ecoregions of the Prairie Ecozone (Acton et
al. 1998). Within this ecological area, C. subglabrum
is found in Mixed Grasslands with sandy soils. The
populations in Saskatchewan and Alberta are isolated
from each other as the land in between them is largely
cultivated. The Manitoba populations are particularly
isolated, being approximately 500 km from the near-
est C. subglabrum population in Saskatchewan and
about 350 km from the nearest population in North
Dakota (Schmoller 2002*). This fragmentation is nat-
ural since C. subglabrum does not generally grow on
the silty and clayey soils that occur in between the var- 4
ious sand hills. The populations that occur along the \
South Saskatchewan River are less isolated than the
ones growing in the sand hills to the south since habi- 2
tat along the river is largely undisturbed. ,
Chenopodium subglabrum is an early successional — Ficure 1. Illustration of Chenopodium subglabrum. Line
habitat specialist growing typically in the stabilizing drawing by D. B. Robson.
335
336
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 1. Nationally and provincially rare plants associated with Chenopodium subglabrum.
Associated Species
Provincially rare *
Bur Ragweed
(Ambrosia acanthicarpa)
Annual Skeleton-weed
(Shinnersoseris rostrata)
Hairy Prairie-clover (Dalea villosa var. villosa)
Sand-dune Wheatgrass (Elymus lanceolatus
Sand Nut-sedge
(Cyperus schweinitziii)
Ambrosia acanthicarpa
Wingless Bugseed
(Corispermum villosum)
Shinnersoseris rostrata
Ambrosia acanthicarpa
Corispermum villosum
Shinnersoseris rostrata
Indian Rice-grass (Achnatherum
hymenoides)
Cyperus schweinitziti
Purple Ball Cactus
(Escobaria vivipara)
Louisiana Broomrape
(Orobanche ludoviciana)
Shinnersoseris rostrata
Winged Pigweed
Province Sand Hill Nationally rare !
Alberta Dominion Nodding Umbrella-plant
(Eriogonum cernuum)
Sand-verbena (Tripterocalyx micranthus)
Saskatchewan Dundurn
ssp. psammophilus)
Elbow Western Spiderwort (Tradescantia
occidentalis)
Great Eriogonum cernuum
Pelican Lake Dalea villosa var. villosa
Seward Elymus lanceolatus ssp. psammophilus
Manitoba Routledge Sand Bluestem (Andropogon hallii)
Tradescantia occidentalis
Brandon Dalea villosa var. villosa
(Carberry)
1According to Argus and Pryer 1990
2 According to provincial Conservation Data Centres
sand at the edges of active dunes and blowouts, and
along eroding, sandy river banks and coulees (Smith
and Bradley 1990*; Robson 1997a; Lamont and Gerry
1998*). It is generally found on south- or west-facing
actively eroding slopes, at the edge of stabilizing sand
and sometimes in dune slacks. Populations tend to
occur in discrete units clustered around active dunes
and are typically highest in areas of finer and more
compacted sand. Given that C. subglabrum was found
historically along the South Saskatchewan River, it is
possible that much of this species’ habitat disappeared
when Gardiner Dam was built on the South Saskatche-
wan River in the late 1960s, flooding the sand bars and
terraces. Dynamic factors such as grazing, erosion and
fire may aid in destabilizing sand, resulting in more
habitat for C. subglabrum (Robson 1999).
Commonly associated species include the following:
Indian Rice-grass (Achnatherum hymenoides), Sand
Grass (Calamovilfa longifolia), Narrow-leaved Goose-
foot (Chenopodium pratericola), Canada Wild Rye
(Elymus canadensis), Northern Wheat-grass (Elymus
lanceolatus ssp. lanceolatus), Prairie Sunflower (Heli-
anthus petiolaris ssp. petiolaris), Hairy Golden-aster
(Heterotheca villosa var. villosa), Skeleton-weed (Ly-
godesmia juncea) and Lance-leaved Psoralea (Psora-
(Cycloloma atriplicifolium)
Achnatherum hymenoides
lidium lanceolatum) (Smith and Bradley 1990*; Rob-
son 1997a). Shrub cover often includes: Common
Bearberry (Arctostaphylos uva-ursi), Hoary Sagebrush
(Artemisia cana ssp. cana), Creeping Juniper (Junipe-
rus horizontalis) and Sandbar Willow (Salix exigua
ssp. interior) (Smith and Bradley 1990*; Robson
1997a). Chenopodium subglabrum is also associated
with six nationally rare and eight provincially rare
plants in various locations (Table 1).
Chenopodium subglabrum is an annual plant with
bisexual flowers (Clements and Mosyakin 2004).
Flowering occurs from June to August and seed pro-
duction in August and September (Wallis and Wershler
1988). Although no studies on C. subglabrum pollina-
tion have been performed, other species in this genus
are wind- or self-pollinated (Johnson and Ward 1993;
Royer and Dickinson, 1999). The distances that C. sub-
glabrum pollen travels is unknown. Since the wind-
pollinated C. pratericola often occurs in the same habi-
tat with C. subglabrum, hybridization may be possible,
although it has not been observed (Bassett and Cromp-
ton 1982).
The fruits and seeds of C. subglabrum lack structures
that would aid in wind (1.e., hairs) or animal (i.e., burs
or fleshy fruits) dispersal although the seeds may ad-
2006
here to wet fur. The seeds likely fall close to the par-
ent plant and may be buried by shifting sand, or travel
short distances in the winter over snow crust. This
means that the main exchange of genetic material be-
tween populations would occur via pollination. Lam-
ont and Gerry (1998*) asserted that dry weather could
limit its germination. The dramatic increase in the num-
ber of individuals observed at most sites in Saskatche-
wan in 2004 coupled with the observation that late sum-
mer was unusually wet and cool suggests that maximum
seed germination occurs under moist conditions. The
seeds can likely remain dormant for several years (Rob-
son 1999). This dormancy can last at least eight years;
at the Beaver Creek site plants were observed in 1996
and 2004 but not in the years in between. There may
be dormant seeds present in stabilized areas that would
germinate under appropriate climatic conditions if the
area were denuded of its vegetative cover.
No attempts to propagate the species for seed pro-
duction have been attempted. If seeds were harvested
from the wild and grown in a greenhouse the seeds
from those plants could be introduced back into the
wild. However, given the species’ sensitivity to climate
conditions, the seeds may not germinate in the wild the
following year making assessment of the introduction
success difficult to ascertain.
Chenopodium subglabrum is both a halophyte
(i.e., salt-loving plant) and a xerophyte (1.e., dry-lov-
ing plant) making it a stress-tolerant species (Grime
1979). Plants in the Chenopodiaceae are generally
non-mycorrhizal (Mukerji et al. 2000). Since C. sub-
glabrum grows in areas of active sand where mycor-
rhiza are usually not abundant, it is probably normally
non-mycorrhizal.
At several sites in Alberta and Saskatchewan there
were specimens of C. subglabrum that had been grazed.
Grazing may have been by cattle or wildlife. Some
grazed plants produced side shoots to compensate for
stem loss. A report on Ord’s Kangaroo Rat (a nation-
ally rare mammal) ecology noted that there were seeds
of C. subglabrum found in their pouches and food
caches so seed predation occurs (Lamont and Gerry
1998*). The growth of several plants observed ap-
peared to be inhibited by an unidentified purplish fun-
gus infecting the leaves.
The Grassland Natural Region is one of the most
threatened natural regions in Canada. Over two-thirds
of the Mixed Grassland has been lost to cultivation or
other development (Wallis 1987). Although some sandy
plains have been cultivated, the hilliest areas have not
been. The main limiting factors affecting Chenopodi-
um subglabrum are its natural narrow preference for
unstabilized sites within dune fields and loss of natu-
ral habitat through succession. Large areas of once
active sand have become stabilized over the last fifty
years. The active sand surface of some dunes in the
Milk River Sand Dunes of Alberta has been reduced by
50% to 75% over 40 years (Wallis 1987). In the Middle
ROBSON: SMOOTH GOOSEFOOT IN CANADA
337
Sand Hills of Alberta, 90% of the sand that was active
in 1950 is now stabilized (Wallis 1987). Hugenholtz
and Wolfe (2005) estimate that stabilization rates in
the Great Sand Hills of Saskatchewan have ranged
from 7.2 to 10.5 ha/yr in the northern portion and
|.3-1.4 ha/yr in the southern portion since 1946. Sta-
bilization rates from 1944 to 1996 in the Elbow Sand
Hills have ranged from 0.4 to 1.9 ha/yr (Hugenholtz
and Wolfe 2005). In the Seward Sand Hills stabiliza-
tion rates from 1970 to 1991 were estimated to be 1.2
to 3.8 ha/yr (Hugenholtz and Wolfe 2005). The Tun-
stall Sand Hills stabilization rates from 1969 to 199]
were estimated to be 0.6 to 3.6 ha/yr (Hugenholtz and
Wolfe 2005). In Manitoba only one of the six sand hills
still has active dunes (Wolfe 2001). The Brandon (Car-
berry) Sand Hills stabilization rate from 1928 to 1990
has ranged from 1.8 to 17.7 ha/yr (Hugenholtz and
Wolfe 2005; Wolfe et al. 2000).
While the exact mechanisms are unclear, a combi-
nation of drought conditions and land use appears to
be influential in dune stabilization (Hugenholtz and
Wolfe 2005; Wolfe et al. 2001, 2002). If current trends
continue, rare native plants that now have dangerously
low populations could be eliminated entirely. How-
ever, Wolfe and Thorpe (2005) speculate that climate
change may result in a potential increase in the suscep-
tibility of sand hills to erosion, possibly reversing this
stabilization trend.
Exotic invasive weeds are becoming more common
in dune areas and have the potential to usurp C. subgla-
brum habitat (Robson 1997a). Exotic plants observed in
the same habitats with C. subglabrum include: Crested
Wheat-grass (Agropyron cristatum), Lamb’s-quarters
(Chenopodium album), Leafy Spurge (Euphorbia esula
var. esula), sweet-clover (Melilotus spp.) and Russian-
thistle (Salsola tragus) (Wallis and Wershler 1988;
Schmoller 2002*; Robson et al. 2005). Oil and gas ex-
ploration and extraction, and to a lesser extent recre-
ation are also potential threats (Wallis 1987; Robson
1997a; Lamont and Gerry 1998*).
Distribution
Chenopodium subglabrum is found in the prairie
provinces of Canada, south to Utah and Colorado, west
to Washington and Nevada, and east to the Dakotas,
Nebraska and Iowa (Figure 2) (Clements and Mosya-
kin 2004). The populations in Manitoba, Washington,
Nevada, Utah, Colorado and Iowa appear to be dis-
junct from the species’ main range (Clements and
Mosyakin 2004).
In Alberta there are populations of C. subglabrum
in five different regions of sand hills: Grassy Lake,
Middle, Rolling Hills Lake, Dominion and Pakowki
Lake Sand Hills. In Saskatchewan, C. subglabrum has
been collected in 11 different sand hill regions: Big-
stick-Crane Lakes, Birsay, Burstall, Cramersburg, Dun-
dur, Elbow, Great, Seward, Pelican Lake, Piapot and
Tunstall Sand Hills. In southwestern Manitoba Che-
eS)
eS)
oo
nopodium subglabrum populations are found in the
Routledge (White and Johnson 1980; Robson et al.
2005) and Brandon (Carberry) Sand Hills.
Due to the annual nature of this species, the popula-
tion numbers fluctuate widely. Data collected in 2004-
2005 supports this hypothesis. At several Saskatche-
wan locations fewer than 50 individuals were found in
the late 1990s but several hundred were observed in
2004. In the Routledge Sand Hills of Manitoba, 19
plants were observed in 2004 but 68 were seen in 2005.
Population fluctuations, possibly in response to cli-
mate, make it difficult to accurately estimate the num-
ber of plants and the vulnerability of populations. In
2004 there were an estimated 8400 plants growing in
Canada; in other years only a few hundred to several
thousand plants were observed. However, even though
plants may not be growing at a site in a given year,
seeds are likely present in the seed bank.
The largest populations occur in the Great and
Seward Sand Hills of Saskatchewan, and in the Grassy
Lake Sand Hills of Alberta. Based on the observed
fluctuations in population size and recognition that at
any given year there are some C. subglabrum seeds
dormant in the seed bank, there are likely between
5 000 and 10 000 plants in Canada. The following is
an overview of populations in the three provinces of
occurrence.
Alberta
The only Alberta site visited recently (in 2004) was
Lost River. Most population research on C. subglabrum
in Alberta was conducted in the late 1980s (Table 2).
Since such a long time has passed since these sites
were visited, the dunes may have stabilized further,
reducing available habitat.
Saskatchewan
Extensive C. subglabrum survey work was done in
the mid to late 1990s in the sand hills of Saskatchewan
(Robson 1997a; Lamont and Gerry 1998*). Since the
THE CANADIAN FIELD-NATURALIST
Vol. 120
FiGurE 2. Distribution of Chenopodium subglabrum in North
America.
first status report was written in 1990 (Smith and
Bradley 1990*) new populations have been discov-
ered in the Bigstick-Crane Lakes, Seward, Great and
Tunstall Sand Hills (Robson 1997b; Nelson Dynes &
Associates 1998*; Lamont and Gerry 1998*) (Table
3). This is balanced somewhat by the apparent loss of
five populations where no plants have been observed
for at least four decades. The Dundurn, Elbow, Great
and Seward Sand Hills locations were visited as
recently as 2004.
TABLE 2. Locations and population data for Chenopodium subglabrum in Alberta.
Location Population Last observation! Population size
Dominion Sand Hills Lost River 2004 5-50
Grassy Lake Sand Hills Barnwell 1988 8
Purple Springs 1988 30
Turin 1986 100-200
Medicine Lodge Coulee Medicine Lodge Coulee 1995 ?
Middle Sand Hills Cavendish 1987 ?
Hilda 1988 3
Pakowki Lake Sand Hills Pakowki Lake North 1980 4
Rolling Hills Lake Sand Hills Lonesome Lake 1988 1
Estimated size of the population in Alberta 7 200-300
' Data obtained from Smith and Bradley (1990*)
* Calculated by adding the minimum number of plants observed or the maximum number of plants estimated at each site
and rounding up or down.
2006 ROBSON: SMOOTH GOOSEFOOT IN CANADA 339
TABLE 3. Locations and population data for Chenopodium subglabrum in Saskatchewan.
Location Population Last observation’ Population size
Bigstick-Crane Lakes Sands Hills Bigstick Sand Hills 1998 21-80
Crane-Lake Sand Hills 1997 311
Tompkins 1997 21
Birsay Sand Hills Dunblane 1961 4
Broderick Broderick 1989 )
Burstall Sand Hills Empress 1985 )
Burstall 1997 202
Cramersburg Sand Hills Cramersburg 1997 179
Dundurn Sand Hills Beaver Creek 2004 1-4
Beaver Creek East 1952 ?
CFB Dundurn 1997 6
Elbow Sand Hills Bridgeford 1968 ?
Elbow 2004 288-500
Head of Qu’appelle 1879 ?
Great Sand Hills Sandhill Stockman’s Association 2004 2018-37967
Diamond Ranch 1997 435
Heck Stockman’s Association 2004 18-3307
Signal Stockman’s Association 2004 36-3307
Watson’s 2004 156 + 3307
McMahon McMahon 1949 ?
Pelican Lake Sand Hills Caron 2002 <10
Piapot Sand Hills Piapot 1983 ?
Patience Lake Patience Lake 1986 ?
Seward Sand Hills SSH 1 1996 55-5407
SSH 2 2004 39-1000
SSH 3 2004 13-330
SSH 4 2001 90-5407
SSH 5 2001 10-540?
SSH 6 2001 2
SSH7 2001 l
Tunstall Sand Hills Bitter Lake 1997 1]
Estimated population in Saskatchewan + 5000-9700
' Data obtained from Harms (1990); Smith and Bradley (1990*); J. & W. Resource Management Consulting (1997*); Robson
(1997a); Lamont and Gerry (1998*); Nelson Dynes & Associates (1998*).
? Estimated number of plants in 2004; determined by multiplying the number of dune fields present in the area by the aver-
age number of plants found at other dunes in the same area.
3 Calculated by adding the minimum number of plants observed or the maximum number of plants estimated at each site
and rounding up or down.
TABLE 4. Locations and population data for Chenopodium subglabrum in Manitoba.
Location Population Last observation Population size
Routledge Sand Hills Routledge 2005 68-75
Oak Lake 1959 ?
Brandon (Carberry) Sand Hills Spruce Woods 2005 9-25
Estimated size of the population in Manitoba ! 77-100
' Calculated by adding the minimum number of plants observed or the maximum number of plants estimated at each site
and rounding up or down.
Manitoba
Although the species was found near Oak Lake in
1959 it was not observed again until 2004 when it was
rediscovered (The Manitoba Museum voucher speci-
men catalogue # 37136) near Routledge (Robson et
al. 2005) (Table 4). The active dunes in the Brandon
(Carberry) Sand Hills were also searched for C. sub-
glabrum in 2004 but no plants were found. However,
in 2005 a small population of nine plants (The Mani-
toba Museum voucher specimen catalogue # 37859)
was located. Two plants were found in the same area
in 2006. This is the first record ever for this location
and represents a range extension for this species of
approximately 100 km east.
340
Evaluation
Significance
Chenopodium subglabrum plays a role in dune sta-
bilization, being one of the few plants that can colonize
active sand. Several related species are important eco-
nomically, including Quinoa (C. quinoa), grown for its
seeds, and Good King Henry (C. bonus-henericus),
grown for its leaves (Everett 1981). The aboriginal
peoples of the western United States consumed seeds
of Chenopodium pratericola and Fremont’s Goosefoot
(C. fremontii) (Smith and Bradley 1990*).
Rarity Status
Chenopodium subglabrum was listed by the Com-
mittee on the Status of Endangered Wildlife in Canada
in 2006 as being a threatened species in Canada. Legal
protection of this species via the Species at Risk Act,
2003 will likely occur in the next few years. This spe-
cies is considered rare in all three prairie provinces
(Packer and Bradley 1984; Maher et al. 1979; White
and Johnson 1980) but is not yet protected by any
provincial legislation. This species is not protected by
any legislation in the United States.
The NatureServe ranks are Global G3G4 (globally
vulnerable to apparently secure), Canada N2 (nation-
ally imperiled), Alberta S1 (provincially critically im-
periled), Saskatchewan S2 (provincially imperiled),
and Manitoba S1 (provincially critically imperiled).
The United States ranks are: Montana S1 (critically
imperiled in the state), North Dakota S1 (critically
imperiled in the state), Nebraska S3 (vulnerable in
the state) and Wyoming S3 (vulnerable in the state)
(NatureServe 2005*). In the remainder of the states
where it occurs it is has either not been ranked (e.g.,
Colorado, Idaho, Kansas, Nevada, Oregon, South Da-
kota, Utah and Washington) or is considered unrank-
able due to a lack of information (e.g. Michigan)
(NatureServe 2005*).
Future Outlook
Chenopodium subglabrum is restricted to sandy
habitats in the prairie provinces that are geographical-
ly isolated from one another. Due to C. subglabrum’s
annual nature, it experiences large fluctuations in pop-
ulation size from year to year. Dune stabilization has
been occurring at a high rate in all of the sand hill
habitats where this species occurs. Increased pressure
on these ecosystems from the oil and gas industry,
and encroachment of exotic weeds further threaten
this species.
Acknowledgments
The Canadian Wildlife Service (Environment Cana-
da), Manitoba Conservation, and the Manitoba Museum
Foundation Inc., provided funding for this research.
Some of the data in this article were obtained from the
original status report on this species prepared by Bonnie
Smith and Cheryl Bradley. Thanks are extended to the
many scientists, particularly Candace Elchuck and Joyce
THE CANADIAN FIELD-NATURALIST
Vol. 120
Gould, who provided information for this report, the
landowners for allowing access to their land, and Janis
Klapecki, Cary Hamel and Cathy Foster for participat-
ing in the field surveys.
Documents Cited (marked * in text)
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ogy, University of Saskatchewan, Saskatoon, Saskatche-
wan.
J. & W. Resource Management Consulting. 1997. Rare
plant survey of Dundurn Military Training Reserve, Sas-
katoon, Saskatchewan.
Lamont, S., and A. Gerry. 1998. Species status report
Chenopodium subglabrum (S. Wats.) A. Nels. Saskatche-
wan Conservation Data Centre, Regina.
NatureServe. 2005. NatureServe Explorer: An Online Ency-
clopaedia of Life. Web site: http://www.natureserve.org/
explorer. [accessed June 2005].
Nelson Dynes & Associates Inc. 1998. Beverley-Cantuar
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Schmoller, D. 2002. Chenopodium subglabrum surveys
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Dakota.
Smith, B., and C. Bradley. 1990. Status report on the Smooth
Goosefoot Chenopodium subglabrum (S. Wats.) A. Nels.
Committee on the Status of Endangered Wildlife in Cana-
da. Ottawa. 27 pages.
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Canada. Canadian Museum of Nature, Ottawa. 274 pages.
Bassett, I. J., and C. W. Crompton. 1982. The genus Cheno-
podium in Canada. Canadian Journal of Botany 60: 586-
610.
Clements, S. E., and S. L. Mosyakin. 2004. Flora of North
America Volume 4: Magnoliophyta: Caryophyllidae, Part
1, Chenopodiaceae, Chenopodium. Oxford University
Press, Toronto.
Everett, T. H. 1981. Illustrated encyclopedia of horticulture.
Volume 3. Garland Publishing Inc., New York.
Grime, J. P. 1979. Plant strategies and vegetation processes.
John Wiley & Sons, Toronto.
Hugenholtz, C. H., and S. A. Wolfe. 2005. Recent stabiliza-
tion of active sand dunes on the Canadian prairies and rela-
tion to recent climate variations. Geomorphology 68: 131-
147.
Johnson, D. L., and S. M. Ward. 1993. Quinoa. Pages 219-
221 in New Crops. Edited by J. Janick and J. E. Simon.
John Wiley & Sons, New York.
Maher, R. V., G. W. Argus, V. L. Harms, and J. H. Hudson.
1979. The rare vascular plants of Saskatchewan. National
Museum of Natural Sciences Syllogeus 20. National Muse-
ums of Canada, Ottawa. 81 pages.
Mukerji, K. G., B. P. Chamola, and J. Singh. 2000. Myc-
orrhizal biology. Kluwer Academic, New York.
Packer, J. G., and C. E. Bradley. 1984. A checklist of the
rare vascular plants in Alberta. Provincial Museum of Al-
berta Natural History Occasional Paper 5. Alberta Culture,
Edmonton. 112 pages.
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Robson, D. B. 1997a. Ecology of rare vascular plants in
southwestern Saskatchewan. M.Sc. dissertation, University
of Saskatchewan, Saskatoon, Saskatchewan. 128 pages.
Robson, D. B. 1997b. Smooth Goosefoot discovered in the
Webb sandhills. Blue Jay 55: 106-108.
Robson, D. B. 1999. Reasons for prairie plant rarity. Pages
92-95 in Proceedings of the Fifth Prairie Conservation
and Endangered Species Conference, Saskatoon, Saskat-
chewan. Edited by J. Thorpe, T. A. Steeves, and M. Gollop.
Provincial Museum of Alberta, Edmonton.
Robson, D. B., J. Greenall, C. Hamel, and C. Foster. 2005.
Smooth Goosefoot rediscovered in Manitoba. Blue Jay 63:
81-84.
Royer, F., and R. Dickinson. 1999. Weeds of Canada. Lone
Pine Publishing, Edmonton.
Wallis, C. A. 1987. Critical, threatened and endangered
habitats in Alberta. Pages 49-63 in Proceedings of the
Workshop on Endangered Species in the Prairie Provinces,
Edmonton, Alberta. Edited by G. L. Holroyd et al. Provin-
cial Museum of Alberta Natural History Occasional Paper
9.
Wallis, C. A., and C. Wershler. 1988. Rare wildlife and plant
conservation habitats of southern Alberta. Albert Forestry,
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White, D., and K. Johnson. 1980. The rare vascular plants of
Manitoba. National Museum of Natural Sciences Syllo-
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Wolfe, S. A. 2001. Eolian deposits in the Prairie Provinces
of Canada. Natural Resources Canada, Ottawa.
Wolfe, S.A., D. J. Huntley, P. P. David, J. Ollerhead, D. J.
Sauchyn, and G. M. MacDonald. 2001. Late 18 century
drought-induced sand dune activity, Great Sand Hills, Sas-
katchewan. Canadian Journal of Earth Science 38: 105-
117.
Wolfe, S. A., D. R. Muhs, P. P. David, and J. P. McGeehin.
2000. Chronology and geochemistry of late Holocene
eolian deposits in the Brandon Sand Hills, Manitoba,
Canada. Quaternary International 67: 61-74.
Wolfe, S.A., J. Ollerhead, and O. B. Lian. 2002. Holocene
eolian activity in south-central Saskatchewan and the south-
ern Canadian prairies. Geographie physique et Quaternaire
56 (2-3): 215-227.
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123-142.
Received 14 September 2005
Accepted 5 February 2007
Summer Movements and Impact of Individual Striped Skunks,
Mephitis mephitis, on Duck Nests in Saskatchewan
SERGE LARIVIERE!*, LYLE R. WALTON’, and FRANCOIS MESSIER?
'Delta Waterfowl Foundation, R.R. 1, Box 1, Site 1, Portage La Prairie, Manitoba RIN 3A1 Canada. Present address: Cree
Hunters and Trappers Income Security Board, 2700 boulevard Laurier, Champlain # 1110, Québec (Québec) G1V 4K5
Canada
*Wildlife Research and Development Section, Ontario Ministry of Natural Resources, Ontario Government Complex, Hwy
101 E., P.O. Bag 3020, South Porcupine, Ontario PON 1HO Canada
> Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada
Lariviére, Serge, Lyle R. Walton, and Frangois Messier. 2006. Summer movements and impact of individual Striped Skunks,
Mephitis mephitis on duck nests in Saskatchewan. Canadian Field-Naturalist 120(3): 342-346.
Striped Skunks (Mephitis mephitis) are important predators of duck eggs in the Canadian prairies. We estimated the impact
of individual Striped Skunks on duck nests by intensive observations of foraging movements and depredation of duck nests
in southern Saskatchewan, 1993-1994. Nightly, skunk movements were variable (range 0-20 km per night), and did not dif-
fer among seasons for females, or between males and females during the parturition/rearing season. Overall, nightly move-
ment of Striped Skunks averaged 7.4 km for females (SD = 5.9 km, n = 20 females) and 6.7 km for males (SD = 3.2 km,n=5
males). During 1 873 h of radio-tracking, we observed depredation of 10 duck nests by 8 skunks (7 F, 1 M). Using our
observed estimate of one depredation for every 187 h, and averaged nightly activity of 8-10 h per night, we estimated that indi-
vidual skunks find one duck nest every 19-23 nights. Thus, during the 60-day nesting season for ducks (mid-May to mid-July),
individual skunks probably find 2-3 duck nests. These observations reinforce the growing evidence that, at normal duck nest
densities (<2.5 nests/ha), depredation of eggs by Striped Skunks is opportunistic, and the impact of Striped Skunks on duck
nests is a direct function of Striped Skunk abundance.
Key Words: Striped Skunk, Mephitis mephitis, duck, movement, nest predation, prairie, predator ecology, Thickwood Hills,
Saskatchewan.
Predation on eggs is an important limiting factor for
many bird populations (Ricklefs 1969; Martin 1995).
In the grasslands of North America, population decline
of many species of ground nesting birds is attributed to
increased predation on eggs during the breeding sea-
son (BOhning-Gaese et al. 1993). Of these birds, ducks
(Anatidae) nesting in the prairies of Canada and the
United States experience high predation rates on their
nests. The primary mammalian predators are Striped
Skunks (Mephitis mephitis), Red Foxes (Vulpes vulpes),
Raccoons (Procyon lotor), American Badgers (Taxidea
taxus) and Coyotes (Canis latrans). The impact of these
predators on duck nest success is well recognized (Klett
et al. 1988; Johnson et al. 1989; Pasitschniak-Arts and
Messier 1995). However, the contribution of individ-
ual animals is poorly understood, yet is critical for
management and mitigation of nest predation.
In the parklands of Saskatchewan, the Striped Skunk
is an important predator of duck eggs (Pasitschniak-
Arts and Messier 1995), yet it depredates duck nests
opportunistically (Lariviére and Messier 1997a, 1998a).
This suggests that the impact of skunks on duck nest
success is the result of “chance” encounters with duck
nests as skunks forage for other foods such as small
mammals and insects (Greenwood et al. 1999). There-
fore, the impact of Striped Skunks on duck nests should
be a function of (a) habitat selection patterns (do skunks
use habitats where ducks nest?), (b) movement rates
(the longer the movements by skunks, the greater the
chances of discovering a duck nest), and (c) density
of skunks (the more skunks, the greater the impact on
duck nests). A recent experiment suggested that indi-
vidual predators were unlikely to key on duck nests at
densities <2.5 nests/ha (Lariviére and Messier 1998a).
Furthermore, partial depredation of duck nests by
Striped Skunks commonly occurs (Lariviére and Mes-
sier 1997a; Ackerman 2002), suggesting that duck eggs
may not be a preferred food of skunks or that their
importance may be marginal to the overall nutrition of
Striped Skunks. An estimate of the impact of individual
Striped Skunks on duck nests would help further our
understanding of the role that duck eggs play in the for-
aging ecology of this carnivore. Herein, we estimate the
impact of individual Striped Skunks on duck nests by
combining intensive radio-tracking and observations
of free-ranging Striped Skunks in the Prairie Pothole
Region of Saskatchewan, Canada.
Methods
We conducted this study ca. 80 km northwest of
Saskatoon in the Thickwood Hills of southcentral Sas-
katchewan, Canada (52°45'N, 107°08'W). In 1993, the
area (ca. 220 km?) was dominated by farmland and
interspersed with many wetlands and stands of trem-
342
2006
bling aspen (Populus tremuloides). Cropland repre-
sented 66% of the area, and was primarily used for the
production of grain (e.g., wheat, barley, rye, and oats),
and oil crops (e.g., canola and flax). Other less common
crops included peas, lentils, buckwheat, and canary seed.
Woodland and areas managed for nesting waterfowl]
(mostly dense nesting cover) occupied 11% and 9%
of the landscape, respectively. Managed nesting areas
consisted mostly of dense nesting cover, hay land, and
idle pastures. Fields of dense nesting cover were sown
with various tame and wild grasses to create prime
habitat for nesting waterfowl. Small or linear cover
types such as wetlands, rights-of-way, farmsteads, and
miscellaneous habitats represented 8%, 4%, 1%, and
1% of the land available, respectively. Topography was
gently rolling, and an extensive network of roads divid-
ed the land.
In 1993, Striped Skunks were live-trapped using
opaque-sided livetraps (Dura-Poly livetraps, Ram Sup-
plies, Birtle, Manitoba). Captured skunks were anes-
thetized with halothane and Telazol® (Lariviére and
Messier 1996a, 1996b), and equipped with motion-
sensitive radio-collars (Telonics Inc., Mesa, Arizona,
USA). Skunks were handled and released at site of
capture.
Radio-collared animals were located and tracked by
a single observer on foot from 18:00 to 06:00 h, dur-
ing the period of greatest activity (Lariviére and Mes-
sier 1997b). During tracking, a single observer fol-
lowed the foraging skunk while remaining downwind,
and usually within 15-50 m of the animal (Lariviére
and Messier 1998b). This distance allowed precise
location of focal animals every 15 min, and allowed
observation of foraging behavior. For short movements
(<100 m), distance from previous location was esti-
mated by observer. Locations immediately following
an observer-induced defensive posture (n = 198: Lar-
iviére and Messier 1996c) were discarded from analy-
ses because of observer disturbance (Lariviére and
Messier 1998b). At dark, we used night-vision equip-
ment (AN-PVS 5C, Bill’s Electronics, Mildmay,
Ontario) to continue observations (Lariviére and Mes-
sier 1998b). Encounters with duck nests typically were
easy to detect because of the flushing hen (Lariviére
and Messier 1997a) or repeated visits to the nest (Lar-
iviére and Walton 1998). Depredated nests were in-
vestigated immediately following departure of the
skunk, and usually reexamined the following day.
For each location, we recorded position of the focal
skunk using a Global Positioning System (GPS) unit.
We estimated accuracy of our GPS as + 250 m. To
overcome biases associated with the inaccuracy of our
GPS on short movements, we only considered the total
distance traveled by each skunk during the 12-h track-
ing period (sum of all 15-min movements). We also
only considered complete 12-h blocks for which onset
and cessation of activity were known. Missing values
(typically caused by delays of the GPS unit to obtain
LARIVIERE, WALTON, MESSIER: IMPACT OF SKUNKS ON DUCK NESTS
343
fixes in forested habitats) for <2 intervals per night
were approximated by the nightly mean of 15-min trav-
el distances. Sessions with >2 missing values were dis-
carded from analyses. Biological seasons for Striped
Skunks were identified as pre-parturition (1 April-14
May), parturition/rearing (15 May- 30 June), pre-dis-
persal (1-31 July), and dispersal (1-31 August).
Each summer (May-August), >50% of radio-col-
lared animals died or dispersed outside the study area
(Lariviére and Messier 1998c). Thus, the resulting data
set was not balanced, and precluded use of repeated-
measures analyses, and comparison among seasons.
Therefore, we used a one-way analysis of variance to
test for effects of season on movements. To avoid bias-
es caused by animals tracked >1 night in a given sea-
son, we used the mean distance traveled for each skunk
in each season (weighted mean, only one value per
animal per season). Because some skunks were tracked
during more than one season and they may not have
been independent, we used an alpha value of 0.01 to
remain conservative and reduce the chance of a Type
1 error.
Results
During 1993 and 1994, we obtained complete move-
ment information for 63 12-h tracking sessions (11 in
1993, and 52 in 1994) on 25 Striped Skunks (5 M, 20
F). Because no skunks were radio-tracked during both
years, we pooled data from 1993 and 1994 for analyses.
Nightly movements of female Striped Skunks during
pre-parturition (mean = 4.2 km, SD = 4.9 km, n = 4),
parturition/rearing (mean = 7.2 km, SD = 6.1 km,
n = 19), pre-dispersal (mean = 5.6 km, SD = 3.0 km,
n= 8), and dispersal (mean = 3.0 km, SD = 3.1, = 2)
were not statistically different Ga 0:70; P:=056):
Distance traveled during 12-h tracking sessions were
variable and ranged from 0.0 to 20.3 km per night
(mean = 6.2 km, SD = 5.2 km, n = 33 female-season).
When we considered only one value for each female
(weighted mean; average of all seasons), nightly move-
ments averaged 7.4 km (SD = 5.9 km, n = 20 females).
During the parturition/rearing season, movements
of males (mean = 6.7 km, SD = 3.2 km, n = 5) did not
differ (t-test, df = 22, P = 0.84) from movement of
females (mean = 7.2, SD = 6.1 km, n = 19). Move-
ments of all skunks during the parturition/rearing peri-
od averaged 7.1 km (SD = 5.6 km, n = 24 skunks).
During spring and summer 1993 and 1994, we radio-
tracked Striped Skunks for 1873 h (858 h in 1993, and
1015 h in 1994). During radio-tracking, we observed
depredation of 10 duck nests (4 in 1993, 6 in 1994),
thus averaging one depredation event for every 187 h
of radio-tracking (1 for 215 h of radio-tracking in 1993,
and | for 169 h in 1994). Observed depredation events
were caused by 8 Striped Skunks (7 females, 1 male).
Observed depredation events occurred between 9 June
and 9 July in 1993, and between 15 May and 20 July
in 1994.
344
From our observations, we extrapolated the number
of nests depredated by a skunk during the duck nest-
ing season. First, using our rate of discovery of | nest
every 187 h of tracking, and allowing an average 8-
10 h of foraging movements per night (Lariviere and
Messier 1997b), we estimated that individual skunks
found 1 duck nest for every 19-23 nights of foraging.
Considering the duck nesting season lasts approxi-
mately 60 d (mid-May until mid-July), then individual
skunks may find 2-3 duck nests per season. However,
because skunks foraging in dense vegetation may have
depredated duck nests without us noticing, this esti-
mate represents a minimum number, but our close-
range observations (typically <50 m) and use of night-
vision equipment suggests that we did not miss many
depredation events.
Discussion
Our results clearly show that the foraging behavior
of Striped Skunks consists of highly variable foraging
movements and opportunistic, but infrequent, depreda-
tion of duck nests. The variable movements we observed
within and among seasons, or between sexes, preclud-
ed detection of statistical differences. However, the
trend suggested that movements were shorter during
the pre-parturition season, higher during parturition
season, and declined temporally thereafter. Interesting-
ly, the decline in skunk movements over time coin-
cides with the steady increase in availability of both
small mammals and insects over time (Lariviére and
Messier 2000). Although we could not statistically link
movements of skunks and food abundance because of
only four sampling periods, we nonetheless observed
apparently reduced skunk movements over time as prey
abundance increased. An experiment done with sim-
ulated nests in our study area in 1995 indicated that
increases in availability of food (chicken eggs) caused
a reduction of activity and home ranges in female
Striped Skunks (Lariviére and Messier 2001a).
Movement rates observed in our study are the largest
reported for this species. However, most previous stud-
ies of skunk movements relied on distance between
consecutive den sites (Shirer and Fitch 1970), maxi-
mum distance from the den during any given night
(Verts 1967), or distance between recapture sites
(Bjorge et al. 1981). Our study is one of three that
reported actual distance moved from telemetry loca-
tions. In Illinois, daily movements of Striped Skunks
averaged 1.7 km for males (n = 4) and 1.4 km for
females (n = 9) (Storm 1972). In North Dakota, night-
ly movements ranged from 0.0 to 9.3 km for 26 females
(mean = 2.7 km), and from 0.4 to 10.5 km for males
(mean = 3.3 km) (Greenwood et al. 1985). We believe
our values are higher because we collected a more
accurate description of movement paths by collecting
locations at closer intervals (i.e., every 15 min) where-
as animals were located every 2 h in North Dakota
(Greenwood et al. 1985), and at intervals varying
THE CANADIAN FIELD-NATURALIST
Vol. 120
between 10 and 120 minutes in Illinois (Storm 1972).
Two additional explanations may explain the long
movements of Striped Skunks in our study area. The
first may be the composition of the landscape. Our
study area was primarily cropland (66% of study area),
a habitat that provides low abundance and diversity
of foods for skunks (Larivieére and Messier 2000).
Not surprisingly, Striped Skunks in our study area
avoided cropland when foraging (Lariviere and Mes-
sier 2000). Thus, large amounts of cropland may con-
tribute to increased movements as skunks were forced
to travel across large fields to access prime habitats
embedded therein. Nightly movements of Striped
Skunks may be lower in large patches of continuous
foraging habitats, a phenomenon that would help ex-
plain the typically higher nest success of ground-nest-
ing birds in large patches of non-fragmented habitat
(Pasitschniak-Arts and Messier 1996; Ball et al. 1995).
Another explanation may be related to the amount
of food in the landscape. Possibly, food abundance
may have been lower in our study area compared to
previous studies, thus contributing to longer move-
ments. Food supplementation experiments on Striped
Skunks and other species typically yield lower activi-
ty, smaller ranges, and shorter foraging bouts (Boutin
1990; Lariviére and Messier 1998a). However, because
we have no way of comparing food abundance among
studies, we can only speculate on whether differences
existed between our study area and those used in pre-
vious studies.
Managers attempting to mitigate predation on duck
nests need to know whether high predation rates occur
because of specialized foraging by a few individuals
depredating numerous nests, or simply by a high abun-
dance of individuals each depredating only a few nests.
Previous observations of skunk foraging behavior sug-
gested that predation is incidental to foraging for other
foods (Vickery et al. 1992; Lariviére and Messier
1997a), and that greater predation rates are explained
by a greater abundance of nest predators (Lariviere
and Messier 1998a).
Our study suggests that individual Striped Skunks
in the Canadian prairies find between 0 and 3 duck
nests per year. In fact, we observed only two skunks
depredate more than one duck nest in any given year
(both cases of females depredating two nests). Because
our study area was located where nesting cover was
common (managed nesting areas represented 9% of the
study area), we believe our estimates probably are
high because most areas of the Canadian prairies do
not have 10% of the land as cover available for ducks,
and hence probably hold fewer duck nests. Nonethe-
less, this suggests that availability of duck nests in the
landscape is not high enough to warrant special for-
aging efforts; this reinforces the notion that nest depre-
dation by Striped Skunks is strictly incidental (Lariv-
iére and Messier 1997a, 1998a, 2001b; Vickery et al.
1992). Finally, the rate of encounter suggests that the
2006
impact of skunks on duck nests is directly related to
skunk abundance, as was previously suggested (Lar-
iviere and Messier 1998a).
The impact of individual Striped Skunks on duck
nests provides insight into the low success of manage-
ment techniques that aim at educating individual pred-
ators. For example, conditioned-taste aversion had been
unsuccessful under field conditions when mammals
such as skunks and Raccoons were the main predators
(Clark et al. 1996). We believe our data suggest that
encounter rates with duck nests by individual preda-
tors are too low for such techniques to be effective,
notwithstanding problems in causing aversion (Clark
et al. 1996). At normal nest densities (<1 nest/ha),
predators cannot subsist on eating duck eggs alone,
and duck eggs only represent a marginal food item
(Greenwood et al. 1999). Not surprisingly, foraging
activity of Striped Skunks is directed at habitats where
base prey such as insects and small mammals are most
abundant, and not in areas managed for nesting ducks
(Lariviere and Messier 2000). Thus, management tech-
niques that aim to increase nest success should be
directed at reducing predator abundance locally (Gar-
retson and Rohwer 2001), or reducing overall habitat
suitability for predators with implementation of large-
scale habitat programs through agricultural policies
(e.g., Conservation Reserve Program in the United
States, Reynolds et al. 2001).
Acknowledgments
This study was supported by the Prairie Habitat
Joint Venture through a research grant from Ducks Un-
limited (Institute for Wetland and Waterfowl Research),
the Canadian Wildlife Service, as well as support from
the Delta Waterfowl Foundation, the National Science
and Engineering Research Council through and oper-
ating grant to SL, as well as postgraduate and post-
doctoral fellowships from the Fonds pour la Forma-
tion de Chercheurs et |’ Aide a la Recherche (FCAR),
Québec. Field and laboratory assistance was provid-
ed by B. Dinter, E. Howard, G. Poon, B. Fry, K. Eade,
H. Royer, P. Toner, S. Daviduik, and D. Anderson. J.
Pitt, Y. T. Hwang, H. Nelson, and D. Murray reviewed
earlier drafts of this manuscript. We followed a uni-
versity-approved animal welfare protocol (#920091)
while conducting this research.
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activity patterns of striped skunks (Mephitis mephitis) in
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346
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ton, and B. D. J. Batt. 2001. Impact of the Conservation
THE CANADIAN FIELD-NATURALIST
Vol. 120
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Received 18 August 2005
Accepted 23 April 2007
Life History Phenology and Sediment Size Association of the
Dragonfly Cordulegaster dorsalis (Odonata: Cordulegastridae)
in an Ephemeral Habitat in Southwestern British Columbia
LAURIE B. MARCZAK'!, JOHN S. RICHARDSON!, and MARIE-CLAIRE CLASSEN2
' Department of Forest Sciences, University of British Columbia, 3041-2424 Main Mall, Vancouver, British Columbia V6T 174
Canada; Corresponding author: e-mail: laurie@interchange.ubc.ca
? Department of Biological Sciences, University of Alberta, CW405, Biological Sciences Centre, University of Alberta,
Edmonton, Alberta T6G 2E9 Canada
Marezak, Laurie B., John S. Richardson, and Marie-Claire Classen. 2006. Life history phenology and sediment size associ-
ation of the dragonfly Cordulegaster dorsalis (Odonata: Cordulegastridae) in an ephemeral habitat in southwestern
British Columbia. Canadian Field Naturalist 120(3): 347-350.
The life cycle of the dragonfly Cordulegaster dorsalis was studied over one year by systematic sampling of larvae in three
intermittent headwater streams in southwestern British Columbia. We determined that larvae normally take three years to
reach maturity, emerging throughout July and August. There is limited evidence suggesting a split cohort development, with
early emergence after two years. Additionally, we tested whether larval instars were distributed randomly or if they occupied
different sediment microhabitats. Smaller animals tend to be associated with smaller grained organic sediments, although
there was high variation between the streams.
Key Words: Odonata, dragonfly, Cordulegaster dorsalis, ephemeral stream, life history, phenology, British Columbia.
Dragonfly larvae are often conspicuous and impor-
tant predators in stream ecosystems. In British Colum-
bia, the family Cordulegastridae is represented by a
single species, Cordulegaster dorsalis Hagen. Cord-
ulegastrids occur in lotic waters, usually in small
streams, where larvae live as shallow burrowers in sedi-
ment (Cannings 2002). In southwestern coastal British
Columbia, larvae of C. dorsalis are often found in
small headwater streams that are subject to extreme
episodic drying. In the interior of the province, the
species is less common and is mainly associated with
small, spring-fed streams. Adults of C. dorsalis nor-
mally fly from mid-May to early September, peaking in
July (Cannings 2002). Published accounts of the life
history, habitat associations or ecological requirements
of C. dorsalis in its larval stage are sparse.
Kennedy (1917) estimated C. dorsalis larvae in cen-
tral California may require as many as four years to
develop. The larval development of cordulegastrids
in Europe ranges from 2 to 5 years and may include
split cohorts (Ferreras-Romero and Corbet 1999). Split
cohort timing has been documented in other species in
the genus (C. boltonii [Donovan] in Ferreras-Romero
and Corbet 1999). Corbet (1999: 230) maintains it is
_ possible to identify a winter critical size, below which
larvae in the first year of development will enter dia-
_ pause during the subsequent summer period, requiring
_ another year’s growth to reach metamorphosis. Such
:
differential growth rates within a single cohort may be
produced by staggered egg-laying times, food limita-
_ tion or variability in the environment. Given the tran-
sitory nature of the streams used by C. dorsalis, it is
possible that it exhibits plasticity in growth rates and
emergence times in response to continual shrinking
and rewetting of the streams.
Dragonfly larvae increase in size by three to four
orders of magnitude as they develop. Microhabitat
selection may have much to do with the physical con-
straints of size. Different species of burrowing odo-
nates prefer certain sediment-size categories (Corbet
1999). C. boltonii has sediment-size preferences that
vary with specific larval size classes (Prodon 1976
unpublished dissertation referenced in Corbet 1999).
In contrast, C. maculata Sélys in Virginia is a habitat
generalist, occuring in equal densities in both silts and
sands (Burcher and Smock 2002). In general, small
larvae of C. boltonii prefer much finer sediments than
do larger larvae — larvae tend to select a particle size
that offers the least obstruction to burrowing (Corbet
1999).
We report life-history observations (morphological
measurements) of larval C. dorsalis over a one year
period in three small headwater streams subject to late
summer drying. We observed that large and small lar-
vae did not occur together within pools. At the same
time, we observed surprisingly high densities of odo-
nate larvae in small, shrinking pools during late sum-
mer. We tested whether small and large larval instars
occupy different sediment microhabitats and report life-
history observations (morphological measurements) of
larval C. dorsalis over a one-year period in three small
headwater streams subject to late summer drying.
347
348
Head width (mm):
THE CANADIAN FIELD-NATURALIST
Vol. 120
Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul
BR 100%
FiGurE |. Life history patterns of Cordulegaster dorsalis shown for 2002-2003 — in G Creek (n = 175). Continuous lines
ascending from left to right indicate inferred boundaries between successive hatching cohorts. The emergence period
during late summer is shown (dotted line). Size ranges of the last three instars are indicated. The broken line indicates
an alternate interpretation and represents the inferred boundary between the two components that could result from
cohort-splitting in the two-year-old hatching cohort, thus distinguishing larvae that grow fast (above line) and slowly
(below line). Numbers below columns indicate sample sizes for each sampling date. Widths of black bars for each
interval of headwidth on a given date represent the relative fraction of the total sample that fell in that interval; total
possible width for any given sampling date is represented by the black legend box labelled 100%.
Methods
Study Site
This study took place in the 5157 ha Malcolm
Knapp Research Forest, located in Maple Ridge,
British Columbia. The forest is located in the Pacific
coastal rain forest, with a maritime climate of cool dry
summers and warm wet winters (Naiman et al. 2000).
We sampled Cordulegaster dorsalis larvae for life
history characteristics and sediment associations in
three streams within the forest (I, F and G streams;
49°18'40"N, 122°32'40"W), which were part of an on-
going riparian management experiment (see Kiffney et
al. (2003) for site descriptions). Both F and G streams
retained a 10 m forest buffer while I stream flowed
through a 5-year-old clearcut. The dominant vegetation
surrounding these creeks included Red Alder (Alnus
rubra) and Vine Maple (Acer circinatum) with a canopy
composed largely of Western Hemlock (Tsuga hetero-
phylla), Western Redcedar (Thuja plicata) and Dou-
glas-fir (Pseudotsuga menziesii). Vegetation adjacent
to I stream consisted of shrubs, particularly Salmon-
berry (Rubus spectabilis) and Huckleberry (Vaccinium
ovatum).
Life-history characteristics
We surveyed approximately 200 m each of I, F,
and G streams in August, October and December of
2002 and April, July and August of 2003, collecting
larvae using dip nets and hand sorting. We measured
head width (Hdw max. distance between the lateral
margins of the compound eyes), total body length (Tbl;
max. distance between the mouthparts and the end of
the cerci measured along the dorsal surface) and wing-
pad length to the nearest 0.1 mm with vernier-scale
callipers. Using regression analysis we determined the
scaling coefficient (reported as the slope of the line) for
head width and body length. Size ranges of late instar
classes were determined from the relationship between
wingpad length and head width. Year class separation
was analysed using wingpad to head width relation-
ships together with visual assessment of size frequen-
cy distributions (after Ferreras-Romero and Corbet
1999). The level of significance was set at P=0.05.
Sediments
The habitat associations of different size classes of
larvae were characterized from August 2002 through
April 2003. Sediment samples were collected at each
point of larval capture and sieved into six size frac-
tions (63 tm, 250 Um, 500 um, | mm, 2 mm, 4 mm).
Each sediment fraction was air-dried and weighed,
then ashed at 550°C for 2 h and reweighed to obtain
weights of both inorganic and organic fractions. Cumu-
lative percent curves were used to determine the medi-
an particle size of each sample. We used linear regres-
sion to investigate the data for relationships between
larval head width, median sediment size and organic
fraction.
2006
head width (mm)
0 10 20 30 40 50
% organic matter
MARKZAK, RICHARDSON, AND CLASSEN: DRAGONFLY IN EPHEMERAL HABITAT
349
0 |
nN
we
=
median particle size (mm)
FIGURE 2. Relationship between larval head width of Cordulegaster dorsalis and sediment characteristics — (A) percentage
of organic matter; and (B) median particle size. Regressions were significant for F Creek (closed circles) but not for
G Creek (open squares) or I Creek (open triangles).
Results and Discussion
During August 2003 when sample sizes were lar-
gest, head width of F-0 (7.5 mm — 9.2 mm) and F-1
(6 mm — 7.8 mm) larvae overlapped, but wingpad
lengths were discrete (3 mm — 5.5 mm in F-2, 6.9 mm
— 10 mm in F-1 and 11.9 mm — 15.5 mm in F-0). Total
body length and head width were correlated (R? = 0.86,
P<0.001, TBL = 4.49 HW — 0.59).
In G Creek, larvae of Cordulegaster dorsalis — de-
rived from eggs that likely hatched in early summer —
grew through late summer and early autumn (Figure
1). At the beginning of their second year most larvae
had a head width between 2 mm and 5 mm. At the end
of their second year of growth, larvae achieved head-
widths between 5 mm and 8 mm. Emergence occurred
at head widths greater than ca. 8 mm during July and
August of the third year.
The pattern of development that we observed in G
Creek (where sample size was greatest, n = 180) indi-
cated that, in general, C. dorsalis larvae require three
years to achieve metamorphosis. It is possible to inter-
pret these data as evidence of a split-cohort in the sec-
ond year with a “fast” group of larvae emerging fol-
lowing accelerated late summer to early fall growth
at the end of their first year (Figure 1, dashed line).
This interpretation would suggest a winter critical size
of ca. 6 mm (Hdw; Figure 1). Sample sizes were too
small in I Creek (n = 30) and F Creek (n = 21) to bol-
Ster a split-cohort interpretation; the Hdw of collected
larvae generally support a three-year development.
Many Odonata retain a synchronized emergence
period through the use of photoperiod cues (e.g. Aoki
1999; Burcher and Smock 2002). Some species of
odonates appear to have both “slow” and “fast” com-
ponents of the same cohort (Martin et al. 1991; John-
son et al. 1995; Ferreras-Romero and Corbet 1999).
We present two possible development schedules for
C. dorsalis: a split cohort with a “fast” component
emerging after two years and a “slow” component re-
quiring three years to complete development. Cordule-
gastridae are noted for their protracted larval develop-
ment, a pattern that might be anticipated of shallow
burrowers occupying small, cool streams (Corbet
1999). Two years would represent a remarkably short
development time within this family, particularly given
the known four-year development time for cordule-
gastrids in northern California (Kennedy 1917). A
more complete record of larval growth rates is required
before evidence of a slow and fast development in C.
dorsalis can be dismissed; in the meantime, the data
most strongly support a simple three-year development
schedule.
During our surveys for larval development, we ob-
served that small pools seldom contained both large
and small larvae. Larvae were most abundant at medi-
an particle sizes between 1.5 mm and 2.5 mm. The
head width of larvae was negatively associated with
percent total organic content in F Creek (Figure 2a:
P <0.01; R? = 0.451) but no relationship was evident
in either G or I creeks. Similarly, the head width of
larvae was positively associated with median particle
size in F Creek (Figure 2b; P < 0.05; R? = 0.446) while
no relationship was evident in either G or I creeks. It
should be noted that median particle and percent
350
organic matter are not independent measures; there is
a negative correlation (r = -0.699). Post-hoc power
analysis suggested that these results must be interpreted
cautiously (power = 0.5134 organic; 0.5072 median
particle).
All odonate larvae are opportunistic predators that
will eat almost any type of prey they can detect and
subdue (Burcher and Smock 2002). Since larger lar-
vae will eat smaller larvae when they encounter them,
it seems plausible that some form of habitat partition-
ing occurs to minimize intraspecific competition and
cannibalism. How larvae are distributed at the micro-
habitat scale will largely determine the frequency of
their interactions with conspecifics, affecting the prob-
ability of agonistic interactions. The presence or ab-
sence of other predators, such as fish, may also affect
the microdistribution of larvae. Suhling (1999) demon-
strated that small larvae of the gomphid, Onychogom-
phus uncatus (Charpentier), were significantly more
abundant in sand where fish predation was also lowest
while medium-sized and large larvae were most abun-
dant in gravel and stones, respectively.
In our study, smaller larvae appeared to be associ-
ated with finer sediments with a higher organic frac-
tion. There was a trend supporting larval size sorting
relative to sediment microhabitats; however, these re-
sults must be interpreted conservatively since this
pattern was evident in only one of the three streams
surveyed and overall sample sizes were low. In previ-
ous habitat surveys of this genus, larvae of C. biden-
tata Sélys were most abundant at a mean grain size
of 2.04 mm while C. heros Theischinger were most
abundant at 2.79 mm (Lang et al. 2001). This is con-
sistent with our finding that larvae of C. dorsalis were
most abundant within medium sand sediments with a
median grain size between 1.5 mm and 2.5 mm.
Although C. dorsalis larvae prefer smaller sediments
as a group, our evidence is inconclusive regarding
microhabitat partitioning amongst larval size classes.
There are several alternative explanations for the size
sorting we observed between small pools. Larvae may
become trapped in these shrinking habitats in a nearly
random fashion with respect to size, and the apparent
sorting occurs post hoc through intraspecific predation.
Smaller larvae would only survive in pools that have a
low proportion of late instar larva when these habitats
become isolated. Alternatively, the observed size sort-
ing of large and small larvae may relate to a factor that
is strongly correlated with sediment grain sizes such
as Current speed or oxygen levels.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Acknowledgments
Robert Cannings provided helpful commentary and
review. The authors would like to thank Trent Hoover
and Conan Phelan for assistance with fieldwork, and
members of the Stream and Riparian Research (StaRR)
lab at the University of British Columbia for comment.
The authors acknowledge funding support from NSERC
and the Forest Investment Initiative Account.
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regulation in Asiagomphus pryeri (Selys) (Odonata: Gom-
phidae). Hydrobiologia 394: 179-192.
Burcher, C. L., and L. A. Smock. 2002. Habitat distribution,
dietary composition and life history characteristics of
odonate nymphs in a blackwater coastal plain stream.
American Midland Naturalist 148: 75-89.
Cannings, R. A. 2002. Introducing the dragonflies of British
Columbia and the Yukon. Royal British Columbia Muse-
um, Victoria.
Corbet, P. S. 1999. Dragonflies: behaviour and ecology of
Odonata. Cornell University Press, New York.
Ferreras-Romero, M., and P. S. Corbet. 1999. The life
cycle of Cordulegaster boltonii (Donovan, 1807) (Odona-
ta: Cordulegastridae) in the Sierra Morena Mountains
(southern Spain). Hydrobiologia 405: 39-48.
Johnson D. M., T. H. Martin, M. Mahato, L. B. Crowder,
and P. H. Crowley. 1995. Predation, density dependence,
and life histories of dragonflies: A field experiment in a
freshwater community. Journal of the North American
Benthological Society 14(4): 547-562.
Kennedy, C. H. 1917. Notes on the life history and ecology
of the dragonflies (Odonata) of central California and
Nevada. Proceedings of the U.S. National Museum 52:
483-635.
Kiffney, P. M., J. S. Richardson, and J. P. Bull. 2003.
Responses of periphyton and insects to experimental
manipulation of riparian buffer width along forest streams.
Journal of Applied Ecology 40: 1060-1076.
Lang, C., H. Miiller, and J. A. Waringer. 2001. Larval habi-
tats and longitudinal distribution patterns of Cordulegaster
heros Theischinger and C. bidentata Sélys in an Austrian
forest stream (Anisoptera: Cordulegastridae). Odonatolog-
ica 30: 395-409.
Martin, T. H., D. M. Johnson, and R. D. Moore. 1991. Fish-
mediated alternative life-history strategies in the dragonfly
Epitheca cynosura. Journal of the North American Ben-
thological Society 10: 271-279.
Naiman, R. J., R. E. Bilby, and P. A. Bisson. 2000. Riparian
ecology and management in the Pacific coastal rain forest.
BioScience 50: 996-1011.
Suhling, F. 1999. Effects of fish on the microdistribution of
different larval size groups of Onychogomphus uncatus
(Odonata: Gomphidae). Archiv fiir Hydrobiologie 144:
229-244.
Received 14 September 2005
Accepted 15 March 2007
An Analysis of the Vascular Flora of Annapolis Heathlands,
Nova Scotia
S. CARBYN!, P. M. CATLING2, S. P. VANDER KLOET®, and S. BASQUILL4
‘Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, 32 Main Street, Kentville, Nova Scotia B4N 1J5
? Agriculture and AgriFood Canada, Environmental Health, Biodiversity, Saunders Bldg., Central Experimental Farm,
Ottawa, Ontario KIA 0C6 Canada; e-mail: catlingp@agr.ge.ca
*Department of Biology, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada
‘Atlantic Canada Conservation Data Centre, PO Box 6416, Sackville, New Brunswick E4L 1G6 Canada; e-mail: sbasquill@
mta.ca
Carbyn, S., P. M. Catling, S. P. Vander Kloet, and S. Basquill. 2006. An analysis of the vascular flora of Annapolis Heath-
lands, Nova Scotia. Canadian Field-Naturalist 120(3): 351-362.
A description and analysis of the vascular plant composition of heathlands in the Annapolis valley were undertaken to pro-
vide a basis for biodiversity preservation within a system of protected sites. Species presence and abundance were recorded
at 23 remnant sites identified using topographic maps, air photos, and Nova Scotia Department of Natural Resources
records. A total of 126 species was recorded, of which 94 were native and 31 introduced. The Annapolis heathland remnants
are strongly dominated by Corema conradii with Comptonia peregrina, Vaccinium angustifolium and Pteridium aquilinum var.
latiusculum. A number of species, including Solidago bicolor, Carex tonsa var. rugosperma, Dichanthelium depauperatum,
Lechea intermedia, Melampyrum lineare, and Rubus hispidus, were characteristic of heathland remnants, although they usu-
ally contributed little to the total cover. The most frequent alien species were Hieracium pilosella and Festuca filiformis, but
Pinus sylvestris, present at 7 of 18 sites, appeared to have the greatest impact in displacing native species. Species listed as
at risk and sensitive in Nova Scotia, including Helianthemun canadense, Hudsonia ericoides and Viola sagittata var. ovata,
occur in open disturbed sand in the Corema heathlands. Distinctive patterns of variation occur in several species and varia-
tion in crop relatives is noted with particular reference to the genera Rubus (blackberries), Amelanchier (Juneberries, Saska-
toon) and Vaccinium (Blueberries). The available evidence suggests that the heathlands and sandy barrens in the Annapolis
valley differ from those further west in Canada and from anthropogenic and coastal heathlands of Nova Scotia in their
species composition including particularly the presence of Corema conradii, Hudsonia ericoides and Amelanchier lucida.
The need to protect representative examples is supported.
Key Words: Heathlands, Annapolis valley, barrens, vascular plants, flora, Corema, Nova Scotia, Canada.
In 1921, legendary Harvard botanist Merritt Lyndon
accords. The only descriptions of Annapolis heathlands
Fernald visited the Annapolis valley. He found exten-
currently available (Fernald 1921; Roland 1946; Catling
sive open heathlands. He noted: “near Berwick and
from there to Wilmot were vast uncultivated plains
carpeted, wherever dry enough, with a close growth
of the New Jersey Pine barren Corema conradii, and,
... remnants of them near Middleton” (Fernald 1921).
As recently as the 1960s open heathlands with scat-
tered Red Pines (Pinus resinosa) occurred for many
miles along the Evangeline Trail (Figure 1). It has been
estimated that in pre-settlement times the actual area
of heathland encompassed approximately 200 km”.
Today less than 3% of the original heathland vegeta-
tion remains in the Annapolis Valley (Catling et al.
2004), and even that is threatened by loss of natural
ecological processes, invasive species and conversion
of the landscape (Catling et al. 2004; Catling and
Carbyn 2004). Protection of this ecosystem is impor-
tant for the protection of (1) insect pollinators of adja-
cent crops; (2) protection of wild relatives of crops for
crop improvement; (3) benchmark research examples;
(4) teaching examples; (5) nature-related recreational
opportunities; and (6) protection of biodiversity gen-
erally in connection with national and international
et al. 2004) are brief, non-quantitative, and insufficient
as a basis for protection of biodiversity. Here we pro-
vide a description of remnants of natural heathland in
the Annapolis Valley along with an indication of dom-
inant species, rare and significant species, variation
between sites and relationship to other eastern Cana-
dian sand barrens. This is designed to provide a basis
for further study and for the establishment of a system
of protected sites.
Methods
Sites and data
The study area consists of 23 sites in the Annapolis
Valley (Table 1, Figure 2). Sites are defined as areas
surveyed separated by at least 0.5 km. Information
from the Nova Scotia Department of Natural Resources
and topographical maps were used to determine the
most probable locations of heathland vegetation occur-
ring on sandy soil.
At sites 1-19 abundance of vascular plants was
recorded. Areas surveyed varied from approximately
0.561 hectares in extent. The majority of these 19 sites
351
B52
THE CANADIAN FIELD-NATURALIST
Vol. 120
FiGurE |. Open heathland dominated by Corema conradii (Broom Crowberry) with Pinus resinosa (Red Pine). Photo by P.
M. Catling in 2004 at site 8.
were visited on at least two occasions in 2003, with
one visit in early summer and another in autumn. Ap-
proximately two hours were spent at each site on each
visit. Species lists were made at each site (Table 2).
An abundance value ranging from 0-5 was assigned
to each species (1 = rare, 2 = uncommon, 3 = com-
mon, 4 = frequent and locally dominant, 5 = dominant)
based on consensus of two or three observers. Although
some larger sites were visited more often, almost all
species recorded were recorded in the first hour of two
visits. Time spent at sites was therefore considered to
be adequate and the lists are thought to be essentially
TABLE 1. Site number and location of heathland study sites in the Annapolis Valley, Nova Scotia.
Site Number, Name
Evangeline Heathland E of 201
Evangeline Heathland W of 201
Trail Heathland
Evangeline Main Heathland
Brooklyn Street Wildlife Sanctuary
Caribou Bog Heathland remnant
Trailer Park Heathland
Exit 17 Heathland, NE corner
Exit 17 Heathland, NW corner
4 km W of Middleton
W side of Middleton
Evangeline Cemetary
13 E side of Greenwood Military Base
Aldershot Military Base behind shooting range
Greenwood section
Exit 17 far east
Greenwood Military Base main site
Greenwood Military Base satellite
201 East near bog
Plot — W of Whitman Road
Plot — E of route 201
Plot — N of route 201
23 Plot — S of Ward Road
BSR K OMNAAUNHKWN
Ne Oo
a
OMADNFE
~~)
ie)
(=)
i)
N
Latitude ° Longitude °
45.0059 —64.8882
45.0048 —64.8930
45.0005 —64.9004
45.0094 —64.8705
45.0872 —64.5245
45.0377 —64.783
45.0219 —64.8255
44.9992 —64.9411
44.9962 —64.9509
44.9204 —65.1048
44.9607 —64.9999
44.9675 —64.9866
44.9896 —64.8955
45.0966 —64.5333
44.9833 —64.8873
44.9974 —64.9470
44.9801 —64.9381
44.9853 —64.9308
44.9791 —64.9366
45.0109 —64.8404
44.9992 —64.8854
44.9472 —65.0203
44.9681 —64.9367
ree
a ——————eeeesss
2006
Heathland Sites Surveyed in
CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS
353
Annapolis and Kings Counties Nova Scotia
20 Kilometres
65°0'W 64°50'W
Legend
A Sites Inventoried (19)
@ Plots Evaluated (4)
Major Roads
——— Minor Roads
—™- County Boundary
DW sandy soil
64°40'W 64°30'W 64°20'W
FiGuRE 2. Heathland sites surveyed in the Annapolis Valley in 2004 (solid triangles) and location of plots evaluated for
cover in 2005 (dots). The sandy soils of the valley (shaded) suggest the maximum extent of heathland.
complete. Voucher specimens collected at various sites
were deposited in the vascular plant herbarium at
Agriculture Canada in Ottawa (DAO). Status of plant
species was determined by using Maher et al. (1978),
the recent listing of species at risk (Nova Scotia De-
partment of Natural Resources 2001*) and informa-
tion and lists on file at the Atlantic Conservation Data
Centre in Sackville.
At sites 20—23, plant association sampling methods
were employed following provincial plot standards
(Quigley et al. 2005*) and Canadian vegetation clas-
sification conventions (CNVC Technical Committee
2004*). In May of 2005, at each of the four sites, a four
hundred square meter plot was subjectively placed in a
homogeneous tract of heathland vegetation, where can-
opy tree cover exceeded 10 percent. The sampled asso-
ciation was the open woodland stage of the Annapolis
Valley sand plain. Species and cover values, represent-
ing percentage of plot area within a vegetation layer,
were visually assessed for five life forms (Table 3).
Lichen names follow Esslinger (1997*), bryophyte
names follow Anderson et al. (1990), and vascular plant
names are from Kartesz and Meachum (1999).
Although many voucher specimens were collected
(and deposited at DAO), a sufficient number of speci-
mens of the genus Aronia were not collected to enable
a determination of which species (or hybrid) was pres-
ent at a particular site. Thus only the genus name ap-
pears in Table 2. Although Rosa carolina and Rosa vir-
giniana were noted, both may have been present at any
site where either was recorded, and some intermediates
were noted. Rubus hispidus may be over-represented
in the survey and Rubus arenicola may be under-rep-
resented due to difficulties in distinguishing these, and
hybrids between them may also have been present.
Results and Discussion.
Native species and limitations of the vegetation
description
Although 104 native species are recorded from the
23 sites surveyed, and although these sites covered a
rather extensive area, they were mostly drier examples
of dry, open sandy habitats (Figure 1). The relatively
short-lived wetter examples and those associated with
natural disturbances, such as fire, are now much less
common than the drier examples. Thus as a reconstruc-
Vol. 120
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2006 CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS 357
Tne a eee tion of the Annapolis sand barren flora, the mesic and
LE
3
5 — - successional stages of dry barrens are to a large ex-
a AN Ee eae tent omitted. This may explain the lack of some species
8g = = = in the survey, such as Agalinis neoscotica (Greene)
a Sale cence Si-estel | {ual Fern. (Middleton False Foxglove), which was des-
Mt; cribed from Annapolis heathlands near Middleton,
i Ren ete!) fy ule=te| and also Bartonia virginica (L.) BSP., Carex atlantica,
5 Polygala sanguinea L. and Sisyrinchium fuscatum
be Sia) Pad be tw Bickn. All of these species occur on heathlands near
| Middleton (personal observation), but were not seen
s are tet. Tec aril during the present survey. Their abundance in the area
q in the past (Fernald 1921, page 138) suggests that the
e SMT ott [last evo | more mesic and naturally disturbed areas were much
3. x more prevalent only a short time ago. Fernald (1921)
& al mureretaworce, hel (lah referred to these as the “the damper Polytrichim-car-
At lc Ae ere peted areas.” He noted that “Bartonia virginica was
| ‘i everywhere,” but it was not recorded in any of the bar-
é SAE IP cont cle legen rens in our survey and only our site 10 approached this
S damper sand barrens habitat.
i Praline gh edt Te Me Lipeile? 4) In addition to the lack of mesic sites it is of interest
a Mra ety cca. eee that in two days in 1920 Fernald (1921) recorded two
S species from the drier heathlands near Middleton that
: NOM iaetor lh cetier re | were not seen by us. These were Potentilla tridentata
2 || 2 and Pyrola rotundifolia. Although our data indicate the
general and dominant composition of the Annapolis
Sees ACO ily lett aly wilieel heathlands, the differences with Fernald’s brief survey
suggest that it may never be possible to have a com-
ah ee plete knowledge of their former floriste composition.
ase bet eit (Cea ah T SN lal val Dominant vascular plants
Relatively few of the 126 species (94 were native
le de et and 31 introduced — Table 2) present in the 19 com-
ME od Pasian gett es pletely surveyed sites were dominant in the vegetation.
Species present at all sites included Corema conradii,
Qu} Sp th tssh st Danthonia spicata, Pinus strobus and Solidago bicolor.
Species present at 18 of the 19 sites were Amelanchier
laevis, Carex tonsa var. rugosperma, Comptonia peregi-
na, Dichanthelium depauperatum, Pteridium aquilinum
var. latiusculum, Vaccinium angustifolium and Populus
tremuloides. Species with the highest overall cover
values included Corema conradii with 91 followed by
Comptonia peregrina with 59, Preridium acquilinum
var. latiusculum with 57, and Vaccinium angustifolium
with 56. A number of species, including Solidago bi-
TABLE 2. (continued) Species abundance and status at 19 heathland sites in the Annapolis Valley. 1 = rare, 2
=
ie)
tal
2 ¥ : color, Carex tonsa var. rugosperma, Dichanthelium
= ue S depauperatum, Lechea intermedia, Melampyrum lin-
iS) S & = eare and Rubus hispidus, were characteristic of heath-
M 2 eS > land remnants, although they were rarely dominant.
2 = aS 5 The most frequent alien species were Hieracium pilo-
g See e & = S sella and Festuca filiformis, both at 17 and 16 (respec-
‘@ oe 5 3 S Ws tively) of 19 sites. The most serious invasive of the
= igs 2 SE oS Se SND heathlands in terms of displacing native species was
1 cae SS Sa PaaS Pinus sylvestris, present at 8 sites (Catling and Car-
a Beeess2 ss byn 2004).
3|}/2| S888 = a8 "s S “s 3 The more wooded plot sites (20-23) gave a very sim-
3 > S 3 S SESES SS 25 ilar picture of the Corema-dominated heathland, but
eels 5 s g§ : gs Soe : with scattered trees, mostly Pinus resinosa (Table 3,
15 || SRS RSS SSO) iss Figure 3). Three species more often associated with
Tee = g S S § Sees S85 woodland than with barrens (Clintonia borealis, Lyco-
*HalRRSSSSSSS5 BAe =
358 THE CANADIAN FIELD-NATURALIST Vol. 120
TABLE 3. Cover values for vegetation layers in single 400 square meter plots at each of 4 sites in the Annapolis valley, based
on data collected by Sean Basquill in May 2005. * = introduced, + = not listed in Table 2.
Species Site 20 Site 21 Site 22 Site 23
Trees
Amelanchier laevis Wieg. 2 = 2 =
Betula populifolia Marsh 3 2 5 5
Picea mariana (P. Mill.) B.S.P. = ES 5 £
Pinus resinosa Ait. 20 25 20 10
Pinus strobus L. — = 5 8
Pinus sylvestris L. * = = = 10
Prunus pensylvanica L. f. - 1 = =
Quercus rubra L. = 4 0.1 Es
Shrubs and small trees
Amelanchier laevis Wieg. = ~ :
Betula populifolia Marsh 3 2 Sal 3
Comptonia peregrina (L.) Coult. 1 1
Juniperus communis L. var. depressa Pursh ~ - _ 0.1
Picea mariana (P. Mill.) B.S.P. - 0.1 -
Pinus resinosa Ait. 4 + 1
Pinus strobus L. 0.5 1
Pinus sylvestris L.* - - -
Quercus rubra L. — 1 - _
Vaccinium angustifolium Ait. 0.1 0.5 3 6
Herbs and dwarf shrubs
Arctostaphylos uva-ursi (L.) Spreng. 30 5) 10 8
Betula populifolia Marsh =
Carex tonsa (Fern.) Bickn. (sensu lato) 0.01 0.1 1 1
Clintonia borealis (Ait.) Raf. + - = 0.01 -
Corema conradii (Torr.) Torr. ex Loud. 58 40 60 60
Deschampsia flexuosa (L.) Trin. — - 1 1
Dichanthelium depauperatum (Muhl.) Gould - - 5 0.1 -
Epigaea repens L. — - 0.01 0.1
Festuca filiformis Pourret * 0.1 -
Festuca trachyphylla (Hackel) Krajina * - 0.001 - -
Gaultheria procumbens L. 0.5 0.01 -
Hieracium piloselloides Vill. * - - 0.01 -
Hudsonia ericoides L. 0.01 0.1 - 0.3
Kalmia angustifolia L. 0.1 0.01 - =
Lechea intermedia Leggett 0.001 - = =
Lycopodium obscurum L. + = =
Lycopodium tristachyum Pursh 0.1 0.01 -
Maianthemum canadense Desf. - - 0.01 -
Oryzopsis asperifolia Michx. + — 0.1 0.2 =
Poa pratensis L. - 0.1 0.1 0.1
Potentilla simplex Michx. - = - 0.2
Pteridium aquilinum (L.) Kuhn
var. latiusculum (Desv.) Underw. 10 1 0.02 -
Rubus hispidus L. - 0.001 - 0.01
Solidago bicolor L. - 0.2 = =
Solidago nemoralis Ait. = 0.001 0.2 0.1
Taraxacum officinale G. H. Weber ex Wiggers -
Trientalis borealis Raf. 0.01 = 0.01
Vaccinium angustifolium Ait. =
Mosses
Ceratodon purpureus (Hedw.) Brid.+ - 0.001 — 2
Dicranum scoparium Hedw.+ ~ 1 — =
Pleurozium schreberi (Brid.) Mitt.+ 0.5 0.1 0.5 =
Polytrichum commune Hedw.+ = — 0.1 1
Polytrichum juniperinum Hedw.+ - 0.01 0.3 3
Lichens
Cladina rangiferina (L.) Nyl.+ 18 40 i 10
Cladina stellaris (Opiz) Brodo + 2 10 3
a
a ————————————————————E
2006
Se ae
nr %
CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS
359
FIGURE 3. Open wooded heathland dominated by Corema conradii and Cladina rangiferina with scattered Pinus resinosa.
Although there is much open area, the site exceeds the threshold 10% tree cover required for classification as a treed
association under Canadian vegetation classification conventions (CNVC Technical Committee 2004*). Photo by S.
Basquill on 25 May 2005 at site 20.
podium obscurum and Oryzopsis asperifolia) were
present a these more wooded sites. Although vascular
plants were the focus of the descriptive work, the plot
data included lichens and bryophytes, suggesting the
former (Cladina rangiferina and C. stellaris) to be a
significant component. This suggestion is supported by
general observations at the 19 other sites where quan-
titative data on bryophytes and lichens was not ob-
tained.
Although Roland (1946) described the Annapolis
sand barrens and heathlands 60 years ago, his descrip-
tion, which recorded 35 species, corresponds very
closely to the composition seen in the relicts that re-
main today. The successional processes are possibly
also the same as described generally by Roland, but
succession may have been much more rapid in some
areas than in others where barrens and heathlands may
have existed as a subclimax lasting for centuries, even
without fire. Roland’s article was written at a time when
sand barrens, heathlands and savanna were regarded
as wasteland rather than as special places for native
biodiversity, but his strong forestry theme is accompa-
nied by many astute observations regarding the native
flora. At the time of early settlement the barrens and
heathlands may have expanded due to cutting and burn-
ing of pine forests and abandonment of sandy land
cleared for agriculture as suggested by Roland. How-
ever there is little doubt that they also existed in pre-
settlement times.
Rare and significant species
The only species considered to be at risk in Nova
Scotia that occurs in the Annapolis heathlands is
Helianthemum canadense (Long-Branch Frostweed,
Figure 4). It was found at 6 of the sites, always in dis-
turbed habitats with some bare sand. Both Viola sagit-
tata var. ovata and Hudsonia ericoides are listed as
sensitive (Nova Scotia Department of Natural Re-
sources 2001*), and these also occurred in disturbed
sandy areas (Figure 5). The provincially rare (Maher
et al. 1978) Sisyrinchium fuscatum Bickn. (Coastal-
Plain Blue-Eyed-Grass) was reported from “sandy areas
near Middleton” (sub Sisyrinchium arenicola Bickn.,
Roland and Smith 1969), but was not seen during the
present survey.
In addition to the rare and threatened species there
is a suggestion of some distinctive patterns of varia-
tion due to taxonomic recognition of several variants.
Among these is a sand barren ecotype of Amelanchier
lucida (personal observation). Taxa of blackberries
(Rubus particeps, R. arenicola), although not recog-
nized as discrete in some recent literature, have been
reported from the Annapolis heathlands suggesting at
360
least the presence of distinctive genetic variants with-
in broadly defined species. Further study may indi-
cate both of these to be worthy of taxonomic recog-
nition. A restricted glabrous variant of Viola sagittata
var. ovata (f. glabrata) occurs on the dry open sand with
the more typical form. Three taxa have been described
from Annapolis heathlands: Amelanchier lucida Fer-
nald, Dichanthelium (sub Panicum) depauperatum vat.
psilophyllum f. cryptostachys Fernald, and Rubus par-
ticeps Bailey.
The heathland ecosystem is particularly valuable as
a reservoir of genetic variation in crops and crop rel-
atives. Fernald (1921) commented on the remarkable
variation in wild blueberries (Vaccinium angustifolium
and V. myrtilloides) in Annapolis heathland remnants
near Middleton. In addition to blueberries (Table 2)
there are potential genotypes of Aronia (Aronia sp.),
of blackberries (Rubus hispidus and others), juneber-
ries (Amelanchier lucida and A. laevis), huckleberries
(Gaylussacia baccata and G. dumosa), cherries (Pru-
nus pensylvanica, P. virginiana, and P. serotina) and
a strawberry (Fragaria virginiana).
Variation between sites
Although all sites shared dominant species (Table
2) they varied in diversity from 26 to 57 native species.
Much of the variation between sites appeared to be
attributable to disturbance and soil moisture. Those
sites with lower lying and periodic moist areas and
open sand had the highest vascular plant diversity.
Site 10 for example was the most unlike the other
sites in native species composition, possibly a result
of parts of it having a relatively high water table and
disturbed areas of periodically moist sand where dry
ground species such as Carex tonsa var. rugosperma
and Danthonia spicata occurred with wetland species
such as Drosera intermedia.
Characteristic native species and “at risk” or “sen-
sitive” species were present at some sites but absent
from others (Table 2). Sites also varied in the extent to
which alien species were present and dominant (Table
2). For example parts of sites 1 and 4 had extensive
and spreading stands of Pinus sylvestris, but displace-
ment of native vegetation by this introduced tree was
either less extensive or not observed at other sites.
Festuca filiformis was a co-dominant at site 5 but not
elsewhere.
Species presence, diversity and extent of impacts
all require consideration in selecting sites for protec-
tion. The variation between sites in composition and
abundance, including that of rare and/or characteristic
species, suggests that protection of a number of sites
will be necessary to protect representative ecosystem
components.
Relationship to other heathlands and barrens
In Canada, Corema-dominated barrens are charac-
teristic of the maritime region. Corema conradii has
a restricted distribution extending from the Gulf of
THE CANADIAN FIELD-NATURALIST
Vol. 120
~ af
FIGURE 4. Helianthemum canadense (Long-Branch Frost-
weed), a native plant at risk in Nova Scotia, with its
main Nova Scotian occurrence on the Annapolis valley
heathlands. Photo by P. M. Catling in 2004 at site 8.
St. Lawrence region south to New Jersey. It does not
occur in sand barrens further to the west in Canada, for
example in the Ottawa valley, where Vaccinium angus-
tifolium is the dominant heath shrub accompanied by
other shrubs such as Prunus susquehanae and Comp-
tonia peregina (Carbyn and Catling 1995). Other spe-
cies present in the Annapolis heathlands but absent in
Ottawa valley sites were Amelanchier lucida, Des-
champsia flexuosa, Rubus hispidus and Solidago bi-
color. Among the prevalent species in Ottawa valley
sand barrens but absent in the Annapolis sites were
Carex siccata, Carex lucorum, Dichanthelium sabu-
lorum var. thinium, Polygonella articulata and Prunus
susquehanae (Carbyn and Catling 1995).
Within Nova Scotia, “barrens” with heath vegeta-
tion occupied a large part of the western portion of the
province (Strang 1972). Some of these barrens are a
short-lived successional stage following fire, whereas
in other cases they are long persisting. Those of short
duration have in some cases been produced by cut-
ting and then maintained by fires. Two such barrens
were described by Hall and Aalders (1968). Such bar-
rens, produced by human activities, have been consid-
ered a degraded landscape of little economic value. The
existence and importance of apparently natural bar-
rens have only recently become apparent (Catling et al.
2004). The sites described by Hall and Aalders (1968)
differ from those in the Annapolis Valley in lacking
Corema conradii, the dominant of the Annapolis bar-
rens. They were also without several other species,
including Hudsonia ericoides. On this basis the dif-
ferences between man-made and natural barrens seem
2006
CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS 36)
FIGURE 5. Open sandy ground in a disturbed area of heathland with Comptonia peregrina (Sweet Fern) and Viola sagittata
var. ovata (Arrow-leaved Violet). Photo by P. M. Catling in 2004 at site 8.
pronounced, but with only two anthropogenic sites, a
more detailed comparison is unnecessary.
The apparently longer persisting natural barrens in
Nova Scotia are readily divided into two major vegeta-
tion types based on either granitic or sandy substrates
(personal observation). Sandy heathlands like those
of the Annapolis Valley were also well developed in the
interior of Yarmouth County and near Debert, Spring-
hill, Parrsboro, and on coastal dunes in Guysborough
and Kings counties and on Sable Island. The coastal
heathlands on sand differ from the interior sites in both
species composition and species abundance, and the
differences suggest adaptation to different conditions.
Differences also exist at the infraspecific level. For
example Lechea intermedia vat. intermedia occurs in
interior heathlands while L. intermedia var. juniperina
occurs on the coastal sandy heathlands. The heathlands
on Sable Island appear distinct (Catling et al. 1985).
The interior sandy heathlands also appear to vary re-
gionally, presumably due to differences in climate. The
climate in southern Yarmouth and Shelburne differs
from that of the Annapolis Valley (personal observa-
tion). The data suggest that the Annapolis heathlands
are distinctive in their floristic composition.
Acknowledgments
Mark F. Elderkin, Species at Risk biologist with the
Nova Scotia Department of Natural Resources, assisted
with current information on status ranks. Lawrence
Benjamin and Randy Milton, also of the Nova Scotia
Department of Natural Resources, provided informa-
tion on site locations for rare plant species in the An-
napolis Valley. Steve Javorek and M. Grant assisted
with field studies (site 1). Geoff Mercer assisted with
the inventory of sites on military bases at Aldershot
(site 14) and Greenwood (sites 17 and 18).
Documents Cited (marked * in text)
CNVC Technical Committee. 2004. Plant association con-
cept for the Canadian National Vegetation Classification.
NatureServe Canada. 5 pages. http://www.cnve-cnvc.ca.
Esslinger, T. L. 1997. A cumulative checklist for the lichen-
forming, lichenicolous and allied fungi of the continental
United States and Canada. North Dakota State University:
http://www.ndsu.nodak.edu/instruct/esslinge/cheklst
cheklst7.htm
Nova Scotia Department of Natural Resources. 2001.
General status ranks of wild species. Vascular plants. http:
www.gov.ns.ca/natr/wildlife/genstatus/specieslist.asp
Quigley, E., P. Neily, K. Keys, and B. Stewart. 2005. Nova
Scotia Department of Natural Resources Forest Ecosys-
tem Classification; Methods for Plot Establishment and
Vegetation Inventory. Unpublished Internal Document.
Literature Cited
Anderson, L. E., H. A. Crum, and W. R. Buck. 1990. List
of the mosses of North America north of Mexico. Bryol-
ogist 93: 448-499.
362
Carbyn, S. E., and P. M. Catling. 1995. Vascular flora of
sand barrens in the middle Ottawa valley. Canadian Field-
Naturalist 109: 242-250.
Catling, P. M., and S. Carbyn. 2004. Invasive Scots Pine
(Pinus sylvestris) replacing Corema heathland in the Anna-
polis valley. Canadian Field-Naturalist 119(2): 237-244.
Catling, P. M.,S. Carbyn, S. P. Vander Kloet, K. MacKen-
zie, S. Javorek, and M. Grant. 2004. Saving Annapolis
Heathlands. Canadian Botanical Association Bulletin 37(1):
12-14.
Catling, P. M., B. Freedman, and Z. Lucas. 1985. Vegeta-
tion and phytogeography of Sable Island, Nova Scotia.
Proceedings of the Nova Scotia Institute of Science 24:
181-248.
Fernald, M. L. 1921. The Gray Herbarium expedition to
Nova Scotia. Rhodora 23: 130-171.
Hall, I. V., and L. E. Aalders. 1968. The botanical compo-
sition of two barrens in Nova Scotia. Le Naturaliste cana-
dien 95: 393-396.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Kartesz, J. T., and C. A. Meachum. 1999. Synthesis of the
North American Flora, version 1.0. North Carolina Botan-
ical Garden, Chapel Hill, North Carolina.
Maher, R. V., D. J. White, G. W. Argus, and P. Keddy.
1978. The rare vascular plants of Nova Scotia. Syllogeus
(National Museums of Canada, Ottawa) 18.37 pages.
Roland. A. 1946. The vegetation of the Annapolis valley I.
— well drained sand areas. Acadian Naturalist 2(7): 1-20.
Roland, A. E., and E. C. Smith. 1969. The flora of Nova
Scotia. Nova Scotia Museum. Reprinted from Proceedings
of the Nova Scotian Institute of Science 26: 1-238, 277-
743.
Strang, R. M. 1972. Ecology and land use of the barrens of
western Nova Scotia. Canadian Journal of Forest Research
2: 276-290.
Received 25 September 2005
Accepted 17 February 2007
Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei,
and Scavenger Activity on Tethered Carrion in Subtidal Benthic
Communities off Western Vancouver Island
SARAH Davies!, ALI GriFFiTHs!, and T. E. REIMCHEN?
'Bamfield Marine Sciences Centre, Bamfield, British Columbia VOR 1BO Canada
*Department of Biology, P.O. Box 3020, University of Victoria, Victoria, British Columbia V8W 3N5 Canada; e-mail: reimchen@
uvic.ca (corresponding author)
Davies, Sarah, Ali Griffiths, and T. E. Reimchen. 2006. Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei,
and scavenger activity on tethered carrion in subtidal benthic communities off Western Vancouver Island. Canadian
Field-Naturalist 120(3): 363-366.
The influence of pelagic carrion food falls on marine benthic scavenging communities was investigated at two depths (10 m,
50 m) in Barkley Sound, west Vancouver Island, British Columbia from 12 May to 4 June, 2003. A remotely operated vehicle
(ROV) equipped with video cameras was used to monitor anchored carrion (15 kg pig leg) during daylight and darkness.
The videos were subsequently analyzed for species diversity, abundance and the intensity of scavenging. At 10 m, Redrock
Crab (Cancer productus) and Kelp Greenling (Hexagrammos decagrammus) dominated, while at 50 m, Spot Shrimp (Pandalus
platyceros), Spotted Ratfish (Hydrolagus colliei) and Pacific Hagfish (Eptatretus stoutii) were the dominant species, most of
which were nocturnal. Hagfish were the major consumers of the carrion and after 23 days, no soft tissues remained at 50 m
while 40% remained at 10 m. Within 24 hours of the carrion deployment, two of eleven ratfish succumbed, probably due to
the direct clogging effects of hagfish mucus on the respiratory apparatus of the ratfish. These field observations are consistent
with laboratory results suggesting high efficacy of hagfish mucus in competitive interactions.
Key Words: Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, Spot Shrimp, Pandalus platyceros, Redrock
Crab, Cancer productus, Kelp Greenling, Hexagrammos decagrammus, marine scavengers, carrion, remotely operated
vehicle (ROV), nutrient cycling, Vancouver Island, British Columbia.
Vertebrate carcasses provide an episodic but highly
_ important food source for benthic organisms, particu-
- larly at depths beyond the zones of primary produc-
tivity (Lalli and Parsons 2001). Such carcasses have
_ recently become of interest due to increasing discards
_ in fisheries as this may shift benthic community struc-
ture (Collins et al. 1999). Species such as ratfish and
_ benthic sharks as well as crustaceans may benefit from
such food falls (Collins et al. 1999; Tamburri and Barry
_ 1999). There are few descriptions of the scavenging
community or the interactions among species around
_ these carcasses.
We anchored carrion at two depths (10 m, 50 m) in
_ Barkley Sound, west Vancouver Island, British Colum-
bia and monitored the carrion with a remotely-operated
vehicle (ROV). We provide here an examination of
' scavenger diversity, abundance and interactions among
» species during daylight and darkness.
\
‘Methods
Two soft-substrate sites (10 m, 50 m: 48°50'N,
-125°10'W) in Mackenzie Anchorage, Barkley Sound,
British Columbia were chosen and single 15 kg pig
legs (frozen with intact skin) were anchored at each
‘site. Initial deployment occurred at 1030 h on 12 May
2003. An ROV (Deep Ocean Engineering Phantom
HD2+) equipped with 5mW 635 nm lasers (10 cm
Standard for scaling) and 250W Tungsten-Halogen
lights were used to monitor the carrion. The ROV was
stationed beside the carrion for 30 minutes during night
trials and 15 minutes (due to less activity) during day
trials. We switched the ROV lights off for 1-2 minutes
on several occasions during the night monitoring phase
but observed no changes in activity. Before surfacing
with the ROV, areas within 5 m of carrion were scanned
for individuals. Day trials of the ROV were carried out
near 1400 h and replicated on three occasions (13, 14,
16 May) while night trials occurred near 2230 h and
were replicated on four occasions (12, 13, 14, 23 May).
The experiment was terminated on 4 June and the two
carcasses returned to the surface and relative tissues
were estimated. Twelve hours of videotape were ana-
lyzed. Percent of remaining carrion and sizes of scav-
engers were estimated from the laser standards. Species
were identified and all individuals within several metres
of the carrion were counted. General behavior of the
species was noted. Paired sampled t-tests were used to
test for significant differences for species and abun-
dance between sites and within sites.
Results
Species Abundance
Carrion anchored at the two depths attracted a dif-
ferent assemblage of species (Figure 1). At 10 m,
Redrock Crab (Cancer productus) (7-20 cm), and Kelp
Greenling (Hexagrammos decagrammus) (11-23 cm)
363
364
Cumulative Abundance: Depth 10m
Cumulative Abundance
Species
THE CANADIAN FIELD-NATURALIST
Vol. 120
Cumulative Abundance: Depth 50m
Cumulative Abundance
Species
FIGURE 1. Cumulative species abundances of scavengers observed at carrion anchored at 10 m and 50 m in Barkley Sound,
Vancouver Island.
were the dominant species, while at 50 m, Long-rayed
Star (Stylasterias forriei) (20-60 cm), Spot Shrimp
(Pandalus platyceros) (12-20 cm), Spotted Ratfish
(Hydrolagus colliei) (18-25 cm), Pacific Hagfish (35-
65 cm) and Rocksole (Lepidosetta bilineata) were
dominant. Only two of 16 species (Spotted Ratfish,
Kelp Greenling) were common to both sites (Figure 2).
Although all species were in direct contact or imme-
diately adjacent to the carrion, only six species active-
ly foraged off the carrion (Pacific Hagfish, Redrock
Crab, Dungeness Crab (Cancer magister), Sunflower
Star (Pycnopodia helianthoides), Long-rayed Star
(Orthasterias koehleri), and Spot Shrimp). Hagfish
undertook extensive burrowing and excavations and
were the major consumers of tissues. Ratfish swam
commonly in close vicinity to the carrion but we
observed no foraging activity. Daily consumption
rates at 10 m were initially high and then decreased
while at the 50 m site, the rate increased throughout
the duration of sampling. On the last ROV survey
(Day 23), 40% of the soft tissues remained at the 10
m site while only bone remained at the 50 m site.
Diel variations
The scavenger community differed during daylight
and darkness (Figure 3). At both sites there were in-
creased numbers of species observed during darkness
(10 m Day, 5 , Night, 11; 50 m Day, 5, Night, 9). At
10 m depth, nocturnal scavengers were Redrock Crab,
Spotted Ratfish, Roughback Sculpin (Chitonotus
pugetensis), and Dungeness Crab. At 50 m, Long rayed
Star was present in both day and night trials and was
seen continually in all surveys. Northern Ronquil (Ron-
quilus jordani), Rocksole and Kelp Greenling were
seen only during day trials. Pacific Hagfish, Spotted
Ratfish, and Spot Shrimp were dominant at night.
Behavioural interactions within and among scavenger
species
Diverse interactions occurred among species. Red-
rock Crabs defended the carrion by direct attacks
against conspecifics as well as against Kelp Green-
ling. Pacific Hagfish excavated burrows within the
carrion by rasping, full-body spinning and occasional
knotting. Spotted Ratfish swam in close vicinity to
the carrion and in several instances directly contacted
the hagfish but we saw no evidence of agonistic
behaviour. However, during the second ROV flight,
we observed two freshly deceased ratfish lying with-
in several meters of the carrion. There was no evi-
dence of external damage to either ratfish but there
was extensive subsurface haemorrhaging around the
mouth on one ratfish and strands of mucus on the
second and we infer rapid respiratory failure from the
hagfish mucus that was prevalent in the water column
around the carrion. During the ROV survey, hagfish
did not feed on dead ratfish. Strands of mucus that
separated from hagfish when feeding on the carrion
were very adhesive as the strands disabled thruster
motors of the ROV.
Discussion
High productivity and the recycling of nutrients sus-
tain large populations of benthic organisms in the
waters of the continental shelf (Nybakken 2001). We
examined an experimental food fall at two (10 m, 50 m)
natural habitats in coastal British Columbia and found
that species assemblages differed between depths and
with diel variation. The intensity of scavenging was
greater at increased depth and increased during night,
which corroborates well with other observations on
scavenging communities (Collins et al. 1999). Shifts in
diel patterns to nocturnal activity are common in marine
2006 DAVIES, GRIFFITHS, and REIMCHEN: SCAVENGER ACTIVITY 365
!Om/ 50m Abundance Difference
FiGuRE 2. Relational abundance of scavenger species at carrion anchored at 10 m and 50 m depth (number of individuals at
10 m — number of individuals at 50 m depth), Barkley Sound. Positive values show greater occurrence at 10 m
depth. Negative values show greater occurrence at 50 m depth.
10m
Day/Night Abundance Difference
Species
FiGuRE 3. Relational abundance of scavenger species due to diel variation (number of individuals during daylight — number
of individuals during night). Positive values show greater occurrence during daylight. Negative values show greater
occurrence during night.
benthic habitats (Lalli and Parsons 2001) and may
reflect predator avoidance or reduction in competition.
The agonistic encounters observed among crabs and
fish indicate that carrion represents significant local
energy enrichment to the benthic community (Stock-
ton and DeLaca 1982).
50m
—s
Day/ Night Abundance Difference
Species
Spotted Hagfish were the major scavengers at 50 m
and were largely nocturnal, consistent with the find-
ings of Fernholm (1974). These fish play an important
role in the turnover and cycling of nutrients (Lesser
et al. 1996: Collins et al. 1999). We observed a much
higher rate of tissue loss at the 50 m site where hagfish
366
were prevalent. Since hagfish are dependent on food
falls, specialized adaptations to minimize competition
are present. Knotting is considered to be an important
feeding behavior in which the hagfish attaches to prey
with tooth plates and passes a knot from the caudal
region anterior towards the head allowing tissues to be
ripped off (Collins et al. 1999). We also observed this
behavior, but only on a single occasion. Rather, hagfish
primarily used full body spinning to pull pieces of tis-
sue from the carrion. Such full body rotation is also
present in sharks feeding on large prey and may allow
an increase in their effective gape (Helfman et al.
1997).
Hagfish produce copious amounts of slime when
feeding, agitated or threatened (Strahan 1963; Tam-
burri and Barry 1999). There are probably diverse
functions to this slime including burrowing, defense
and competitive exclusion (Collins et al. 1999). Hag-
fish burrowed into the carcass up to % of their total
body length. While the mucus may facilitate entry into
the carrion, the burrowing would also cover the gills
and this would limit gas exchange of the hagfish. This
suggests the potential for posterior cutaneous respira-
tion. Lesser et al. (1996) noted that hagfish skin was
well-vascularized and hagfish are therefore able to sur-
vive with plugged nostrils and no respiratory current
across the gills. Slime produced by hagfish has high
strength (Fudge et al. 2003) and can also obstruct
mouthparts and gills of other species attempting to feed
at food falls (Tamburri and Barry 1999; Collins et al.
1999). Our ROV observations are consistent with these
suggestions. Two of the ratfish swimming in close prox-
imity to the foraging hagfish quickly succumbed, pre-
sumably from loss of respiratory ability. The close prox-
imity suggests high efficacy and rapid effects of the
hagfish mucus in natural conditions even under high
dilution effects. If these observations are representa-
tive, it suggests that hagfish may not only have a sub-
stantial competitive advantage over other fish species
in these episodic carrion fallouts but also operate as
predators through the lethal influences of mucus on
other scavengers.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Acknowledgments
We thank Nimrod Levy for discussion, James Mor-
timer, John Richards and Janice Pierce for ROV and
vessel support, J. Yakimishyn and J. P. Danko for lab
and technical support, and the Bamfield Marine Sci-
ences Center for funding this project. TER acknowl-
edges support (A2354) from the Natural Sciences and
Research Council of Canada.
Literature Cited
Collins, M. A., C. Yau, C. P. Nolan, P. M. Bagley, and I. G.
Priede. 1999. Behavioral observations on the scavenging
fauna of the Patagonian Slope. Journal of the Marine Bio-
logical Association of the United Kingdom 79: 963-970.
Fernholm, B. 1974. Diurnal variation in the behavior of the
hagfish Eptatretus burgeri. Marine Biology 27: 351-356.
Fudge, D.S., K. H. Gardner, V. T. Forsyth, C. Riekel, and
J. M. Gosline. 2003. The mechanical properties of hy-
drated intermediate filaments: Insights from hagfish slime
threads. Biophysical Journal 85: 2015-2027.
Helfman, G.S., B. B. Collette, and D. E. Facey. 1997. The
diversity of fishes. Blackwell Science, London.
Lalli, C. M., and T. R. Parsons. 2001. Biological Oceanog-
raphy: An Introduction. 2? Edition. Butterworth/Heine-
mann, Great Britain.
Lesser, M. P., F. H. Mar’tini, and J. B. Heiser. 1996. Ecolo-
gy of the hagfish, Myxine glutinosa in the Gulf of Maine.
I. Metabolic rates and energetics. Journal of Experimental
Marine Biology and Ecology 37: 215-225.
Nybakken, J. W. 2001. Marine biology: An ecological ap-
proach. 5" edition. Benjamin Cummings, Montreal.
Stockton, W. L., and T. E. DeLaca. 1982. Food falls in the
deep sea: occurrence, quality and significance. Deep-Sea
Research 29: 157-169.
Strahan, R. 1963. The behaviour of Myxine and other Myxi-
noids. Pages 22-33 in Biology of Myxine. Edited by A.
Broda and R. Fange. Universitetsforlaget, Oslo, Norway.
Tamburri, M. N., and J. P. Barry. 1999. Adaptations for
scavenging by three diverse bathyal species, Eptatretus
stoutii, Neptunea amianta and Orchomene obtusus. Deep
Sea Research I 46: 2079-2093.
Received 26 October 2005
Accepted 16 February 2007
Notes
A New Record Size Wolf, Canis lupus, Group for Ontario
Liv S. Vors! and Puitie L. WILSON?
Trent University, Peterborough, Ontario Canada
'Current address: Department of Biological Sciences, University of Alberta, Edmonton, Alberta. T6G 2E9, Canada; e-mail:
vors@ ualberta.ca
Current address: c/o Sir Sanford Fleming College, Fish and Wildlife Department, Frost Campus, Albert Street South, P.O. Box
8000, Lindsay, Ontario Canada
Vors, Liv S. and Philip L. Wilson. 2006. A New Record Size Wolf (Canis lupus) Group for Ontario. Canadian Field-Naturalist
120(3): 367-369.
This report documents a group of 19 Wolves (Canis lupus) in northwestern Ontario. This is the largest group observed since
record keeping in the Moose Aerial Inventory commenced in 1995. This large group may be a response to a high Moose
(Alces alces) population in the Red Lake area.
Key Words: Gray Wolf, Canis lupus, social organization, pack size, Moose, Alces alces, Ontario.
The close association between Gray Wolves (Canis
lupus) and their ungulate prey greatly influences the
ecology of this cursorial predator, which is capable of
killing prey much larger than itself (Kolenosky and
Standfield 1975). Pack size is believed to be influ-
enced by numerous factors. Pack sizes do not neces-
sarily differ among Wolf populations whose major prey
species are different. For example, average sizes of
Wolf packs mainly depredating on Moose (Alces alces)
are not necessarily larger than those feeding on White-
tailed Deer (Odocoileus virginianus), but Wolf packs
tend to be largest where Wolves prey on the largest
ungulates (Fuller et al. 2003). This note documents one
large group of Wolves observed in Ontario, Canada.
Observation
A group of 19 wolves was observed on 15 February
2005 between 1300 and 1400 hours on a frozen lake
west of the settlement of Red Lake, Ontario, at approx-
imately 51°OSN and 94°05W. Red Lake (51°04N,
93°47W) is a settlement located in the boreal forest of
northwestern Ontario. This sighting took place during
a transect flight to map areas of Woodland Caribou
(Rangifer tarandus caribou) habitat activity. The air-
craft, a De Havilland Turbo Beaver, was flying at a
speed of 90 knots and an altitude of 400 feet when the
group was observed. Ontario Ministry of Natural
Resources (OMNR) pilot Carl Hansson, Trent Univer-
sity graduate student Liv Vors, Sir Sanford Fleming
College Fish and Wildlife Technologist graduate Philip
Wilson, and Red Lake OMNR employee Claire Que-
wezence observed and counted the Wolves.
The aircraft made several circles over the animals,
which were loosely congregated near the western shore
of the lake. The Wolves ran around the ice, evidently
disturbed by the aircraft, but made no attempts to flee
into the neighbouring forest. No kill was visible. The
animals were counted and confirmed by all observers.
After about two minutes of observation, the aircraft
resumed its transect flight.
Discussion
We believe this group to be the largest on record for
Ontario, and possibly all of Canada east of the Rocky
TABLE |. Records of maximum Canadian Wolf pack sizes west of the Rocky mountains, 1969-2005.
Location
_ Riding Mountain National Park, Manitoba
Jasper National Park, Alberta
Southeastern Quebec
| Northwestern Alberta
Algonquin Provincial Park, Ontario
Southwestern Quebec
Parry Sound Forest District, Ontario
_ Pukaskwa National Park, Ontario
Size Source
16 Carbyn (1980)
14 Carbyn (1974)
10 Potvin (1988)
10 Bjorge and Gunson (1985)
9 K. Mills, personal communication, 2005
8 Messier (1985)
8 Kolenosky (1972)
6 Forshner (2000)
367
368
Mountains. Wolf pack records are kept in the Moose
Aerial Inventory Database 1995—2006*. Before this
sighting the largest pack on record was 13, and was
sighted in WMU (Wildlife Management Unit) 15A in
1998. Reported average pack sizes in Ontario range
from 3.8 (Bergerud et al. 1983) to 5.9 (Pimlott et al.
1969) to 6.0 (Forbes and Theberge 1996), with maxi-
mums of 7,9 and 13, respectively. Our observation of
19 wolves is significantly higher than most packs re-
ported in Canada (Table 1).
Gray Wolves found in the North American land-
scape represented in their northern Ontario range pre-
dominantly prey upon ungulates and Beaver (Castor
canadensis) (Carbyn 1987; Gauthier and Theberge
1987). Large packs likely provide increased hunting
efficiency when prey is scarce, or may alternatively be
a response to abundant prey (Schmidt and Mech 1997).
We suggest the large group we observed is a response
to the latter. Since 1975, Moose populations in the Red
Lake area have increased from a density of 0.059-
0.209 animals/km? to 0.359-0.509 animals/km? (Mc-
Kenney et al. 1998). Current Moose densities in the
Red Lake area are lower than McKenney’s estimates.
Moose in the Red Lake WMU were estimated to be
0.265 animals/km? in 2004 (unpublished data, Ontario
Ministry of Natural Resources, Bracebridge, Ontario).
White-tailed Deer are also expanding their range north-
ward, providing additional prey (Red Lake OMNR,
personal communication). Wolf density is positively
correlated with ungulate numbers (Fuller 1989; Mes-
sier 1994). Although Moose density has declined since
1998, the ungulate biomass may still be high enough
to support a large Wolf population.
The relationship between pack size and prey size,
however, is imperfectly understood. While food ac-
quired per Wolf may be positively correlated with pack
size (Nudds 1978), this relationship is not consistent.
A review of prey use by Wolves in several geographic
areas suggested a negative relationship between pack
size and food acquisition (Schmidt and Mech 1997).
In addition, prey size and pack size may not be closely
linked. Mech (2000) observed a pack of 22-23 wolves
in Minnesota, and this group preyed primarily on
White-tailed Deer. Consequently, prey size and food
acquisition are unlikely to be the only factors explain-
ing the large group we observed. An additional caveat
is the group we observed may have consisted of sev-
eral packs that converged at the time of observation.
An additional point of interest was the colour of the
wolves. All were dark brown or black, with the excep-
tion of one silvery-white individual. Most wolves in
eastern Canada are gray, with black and white indi-
viduals constituting approximately four and less than
one per cent of the population, respectively (Gipson
et al. 2002).
THE CANADIAN FIELD-NATURALIST
Vol. 120
Acknowledgments
The aerial surveys were funded by the Ontario Min-
istry of Natural Resources, The Wildlands League, The
Wildlife Conservation Society, and The National Geo-
graphic Society. Many thanks to Brent Patterson for
his comments on the manuscript.
Documents Cited (marked * in text)
Moose Aerial Inventory Database. 1995-2006. Ontario Min-
istry of Natural Resources, Bracebridge, Ontario.
Literature Cited
Bergerud, A. T., W. Wyett, and J. B. Snider. 1983. Role of
wolf predation in limiting a moose population. Journal of
Wildlife Management 47: 977-988.
Bjorge, R. R., and J. R. Gunson. 1985. Evaluation of wolf
control to reduce cattle predation in Alberta. Journal of
Range Management 38: 483-487.
Carbyn, L. N. 1974. Wolf predation and behavioural inter-
actions with elk and other ungulates in an area of high prey
density. Canadian Wildlife Service, Edmonton, Alberta.
233 pages.
Carbyn, L. N. 1980. Ecology and management of wolves in
Riding Mountain National Park, Manitoba. Canadian Wild-
life Service, Edmonton, Alberta. 184 pages.
Carbyn, L. N. 1987. Gray wolf and red wolf. Pages 358-
376 in Wild Furbearer Management and Conservation in
North America. Ontario Ministry of Natural Resources.
Published by The Ontario Trappers Association under the
authority of the Licensing Agreement with the OMNR.
Printed in Ontario, Canada. 1150 pages.
Forbes, G. J., and J. B. Theberge. 1996. Response by wolves
to prey variation in central Ontario. Canadian Journal of
Zoology 74: 1511-1520.
Forshner, A. 2000. Population dynamics and limitations of
wolves (Canis lupus) in the greater Pukaskwa Ecosystem,
Ontario. M.Sc. Thesis, University of Alberta, Edmonton.
Fuller, T. K. 1989. Population dynamics of wolves in north-
central Minnesota. Wildlife Monographs 110, The Wildlife
Society, Bethesda, Maryland. 37 pages.
Fuller, T. K., L. D. Mech, and J. F. Cochrane. 2003. Wolf
population dynamics. Pages 161-191 in Wolves: Behavior,
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Gauthier, D. A., and J. B. Theberge. 1987. Wolf predation.
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servation in North America. Published by The Ontario
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Agreement with the OMNR. Printed in Ontario, Canada.
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Cluff, D. W. Smith, and M. D. Jiminez. 2002. Color pat-
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east-central Ontario. Journal of Wildlife Management 36:
357-369.
Kolenosky, G. B., and R. O. Standfield. 1975. Morpholog-
ical and ecological variation among gray wolves (Canis
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Their systematics, behavioral ecology, and evolution. Edited
by M. W. Fox. Nostrand Reinhold, New York.
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McKenney, D. W., R. S. Rempel, L. A. Vernier, Y. Wang,
and A. R. Bisset. 1998. Development and application of a
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Nudds, T. D. 1978. Convergence of group size strategies by
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957-960.
NOTES
369
Pimlott, D. H., J. A. Shannon, and G. B. Kolenosky. 1969
The ecology of the timber wolf in Algonquin Provincial
Park, Ontario. Research Report (Wildlife) 87. Ontario
Department of Lands and Forests, Toronto. 92 pages.
Potvin, F. 1988. Wolf movements and population dynamics
in Papineau-Labelle reserve, Quebec. Canadian Journal of
Zoology 66: 1266-1273.
Schmidt, P. A., and L. D. Mech. 1997. Wolf pack size and
food acquisition. American Naturalist 150: 513-517.
Received 27 September 2005
Accepted 10 May 2007
A Tribute to Nicholas Stephen Novakowski 1925-2004
JOSEPH E. BRYANT
447 Thessaly Circle, Ottawa, Ontario K1H 5W7 Canada
Bryant, Joseph E. 2006. A tribute to Nicholas Stephen Novakowski 1925-2004. Canadian Field-Naturalist: 120(3):
370-378.
Nicholas Stephen Novakowski was an enthusiastic
and very knowledgeable wildlife and fisheries scien-
tist. Although his list of published works is modest, he
was well known and respected across Canada and in
several international fora. He has been credited as “the
person who saved the Wood Bison as a discrete sub-
species” (W. E. Stevens, personal communication)
and was highly engaged in the early years of the sur-
veys and recovery of the Whooping Crane (Novakows-
ki, 1965b, 1966). Following those early days at Fort
Smith, North West Territories, Nick was transferred
first to the Edmonton office of the Canadian Wildlife
Service and then to CWS headquarters in Ottawa
where he was appointed Staff Specialist, Mammalogy,
and took on national and international responsibilities.
He retired in 1983 but continued his strong support for
endangered wildlife and their habitats until his death
on 3 July 2004 in Ottawa. He is survived by his wife,
Amy, and two sons, Kent, a professor of hydrogeolo-
gy and environmental engineering at Queen’s Univer-
sity and Erin Nicholas, a professor of geography and
environmental studies at Memorial University.
Nick was born 28 November 1925 in Mundare, Al-
berta (about 70 km east of Edmonton), the second child
and eldest son in a family of 10 children. His father
owned a Chrysler dealership and garage and from an
early age Nick learned about all things mechanical —
a fact that would stand him in good stead in later years
as a field biologist. He also developed an abiding loy-
alty to Chrysler products and, briefly, owned only one
automobile that didn’t bear that company’s mark. Find-
ing the alternate car highly unreliable, he soon returned
to his original choice and did not waver again. In his
younger years he was a superb athlete. He and his
brothers Archie and Van were well known in central
Alberta hockey circles as a high-scoring powerhouse.
Nick, who was ambidextrous, could play either wing
and was a sharp goal tender (Ken Novakowski, person-
al communication). Also in those early years he became
an ardent angler and as often as possible visited mem-
bers of his extended family who lived in towns north of
Edmonton. One of his favourite spots was Siebert Lake.
Just a couple of weeks before his death he very happi-
ly returned there with his sister, Sharon, and her hus-
band to renew those earlier experiences — with consid-
erable success. Time had not dulled his expertise.
Nick entered the University of Alberta in 1947 and
in the following two summers worked on forest fire
suppression crews based in Fort Smith, North West
Territories. There he became acquainted with Bill Ful-
ler, the resident mammalogist for the Canadian Wildlife
Service. He graduated with a B.Sc. in Chemistry in
1950 and that summer returned north but this time he
was hired by Northern Transportation Company Limit-
ed (NTCL) for summer work on the docks at the com-
pany’s shipping base near Fort Smith. At that time
NTCL gave hiring priority to young men who could
enhance its baseball team and Nick “was a very good
ball player” (W. A. Fuller, personal communication).
Nick returned to the University of Alberta that autumn
to begin a Master’s program and went back to Fort
Smith in the spring of 1951. That autumn Dr. Fuller
was assigned to conduct research of Wolves in Wood
Buffalo National Park (WBNP) and adjacent areas of
the Mackenzie District and employed Nick as an assis-
tant (see Fuller and Novakowski 1955). Nick also
helped Fuller with his studies of Beaver and with the
“autopsies of lone, crippled, bison we encountered from
time to time” (W. A. Fuller, personal communication).
It was the start of Nick’s wildlife and fisheries career,
and perhaps of equal importance, the start of the
romance that led in June 1952 to his marriage to the
love of his life, Amy Swartz, a Fort Smith school
teacher from Saskatchewan.
Also in 1952, with Fuller’s recommendation, Dr.
D. S. Rawson accepted Nick for graduate studies in
fisheries at the University of Saskatchewan. Over the
next few years he worked very happily at his studies
and as a fisheries biologist for the Government of Sas-
katchewan. Although only one published paper appears
to have resulted from that work (Atton and Novakows-
ki 1956), he authored or co-authored many in-house
reports for the Saskatchewan government dealing with
fisheries research and resource management. Amy No-
vakowski tells me that those were the happiest years
of Nick’s life — helped, no doubt, by the fact that Amy
was able to accompany him on many of his extended
field studies in northern Saskatchewan. The photograph
accompanying this article was taken at that time.
After completing his M.Sc., (Novakowski 1955c*)
Nick was hired by the Canadian Wildlife Service in
1956 and sent to Fort Smith. His mentor, Dr. Fuller, was
transferred that year to Whitehorse, Yukon, and Nick
dived immediately into Dr. Fuller’s continuing studies
of Beaver populations in the southwestern Northwest
Territories, Wolves in Keewatin District (Flook and |
370
2006 A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004
FIGURE 1. Nick Novakowski, fisheries researcher, Lac des Iles. Saskatchewan, 1955.
3/72
Novakowski 1956*; see also Novakowski 2003) and
Bison and Whooping Cranes in Wood Buffalo Nation-
al Park.
In 1957, in the course of an aerial survey of Bison in
WBNP, he discovered the presence of an isolated herd
of Bison in the Nyarling River area in the northwest
corner of the Park (Banfield and Novakowski 1960).
Subsequent aerial and ground studies indicated that the
herd might be Wood Bison genetically uncontaminated
by the Prairie Bison that had been transferred to WBNP
in the 1920s. In 1959, Nick was able to obtain speci-
mens which were confirmed by the National Museum
to be Wood Bison (Banfield and Novakowski 1960). In
the following three years he arranged for over 70 live
specimens to be trapped and tested for tuberculosis and
brucellosis. Of those, 18 that were disease-free were
moved in February 1963 to a large holding corral north-
west of Fort Smith. Three calves arrived that spring
and in early summer all of the animals were again test-
ed for disease. All were clear. Nick had previously
scouted out appropriate habitat north of Fort Provi-
dence, North West Territories (Novakowski, 1959d*),
and 18 disease-free Wood Bison, including the new
calves, were transported there in August 1963. That
area, now known as the Mackenzie Bison Sanctuary,
proved to be an excellent site and the small nucleus
soon swelled. Subsequently other specimens were taken
from the Nyarling River site, transferred to Elk Island
National Park, and after a rigorous clean-up protocol,
produced another disease-free herd of Wood Bison
(Gates et al. 2001). Some of Nick’s less well-known
Bison research included “the first use of dental annuli
to estimate the age of bison” (Christianson et al. 2005).
The same authors also credit him with being the first
to measure and to demonstrate the usefulness of other
Bison dentition measurements in aging large samples.
In 1958, Nick was also in charge of stocking Pine
Lake and a few smaller lakes in WBNP with Rainbow
Trout. He established and operated a small hatchery for
that purpose, putting his Saskatchewan fisheries experi-
ence to good purpose (Novakowski 1958c*, 1959c*).
In 1962, when anthrax struck the Bison population
just outside WBNP, it was the first recognized instance
in Canada of anthrax in wildlife and all concerned were
faced with a very steep learning curve. Nick worked
closely with a team of scientists from the Canadian
Wildlife Service, the Health of Animals Branch of the
federal Department of Agriculture, and medical per-
sonnel to bring it under control (Novakowski et al.
1963; Choquette et al. 1972).
Dr. Fuller had engaged the interest of a long-time
trapper in the southern portion of WBNP, Solomon
Lacaille, to help with Muskrat studies (Fuller 1951).
Nick followed through and set up a multi-year study on
Mr. Lacaille’s trapline to study many aspects of beaver
biology and ecology, leading to his Ph.D. from the Uni-
versity of Saskatchewan in 1965 (Novakowski 1965a*,
1967a, 1969a).
THE CANADIAN FIELD-NATURALIST
Vol. 120
The nesting grounds of the Whooping Crane, long
an international mystery, were discovered in 1954 in
the northern portion of Wood Buffalo National Park.
In the subsequent two years the site was monitored by
Dr. W. A. Fuller. Nick took over those studies in 1956
and monitored the small breeding population for sever-
al years (Novakowski 1966, 1967d). He was also res-
ponsible for capturing an injured juvenile whooper in
1964 and for ensuring its safe conduct to the breeding
facility of the United States Fish and Wildlife Service
near Washington, D.C. (Novakowski 1965b, 1965e*).
That bird, a male named “Canus” for its dual nationali-
ty, sired “over 180” (E. Kuyt, personal communication)
Whooping Cranes over its long career. It died in Jan-
uary 2003. The mounted body was repatriated to Cana-
da and is now installed at the Northern Life Museum in
Fort Smith, N.W.T. (E. Kuyt, personal communication).
Nick’s work on Wood Bison and Whooping Cranes
introduced him to the burgeoning field of rare and
endangered species. It was a field that would occupy
much of his attention for the remainder of his life.
The Canadian Wildlife Service moved Nick to a Staff
role in Ottawa in 1966. J. Anthony Keith, who was
Nick’s immediate superior in the Canadian Wildlife
Service for much of his time in Ottawa, reported that
“Nick Novakowski was the founding voice in CWS
headquarters for wide-scale collaborative action on
endangered species, beginning in 1967 when he was
Staff Specialist, Mammalogy. While his own earlier
work on Whooping Cranes became the trademark fed-
eral endangered species project, there was a continuing
debate amongst headquarter staff about how far CWS
should get involved in endangered species work. Nick
had to argue against those who saw it as a diversion of
limited resources from urgent wildlife conservation
issues. He worked with Ted Mosquin to produce the
first set of scientific articles in Canada on endangered
vertebrates (The Canadian Field-Naturalist 84(1)
1970), writing the article on mammals.” [See Nova-
kowski 1970b].
In 1973, Nick represented the Canadian Wildlife
Service at the international negotiations in Washing-
ton of what became the Convention on International
Trade in Endangered Species of Wild Fauna and Flora
(CITES). While John Heppes joined CWS to estab-
lish the Canadian management authority for CITES,
Nick chaired the scientific authority until he retired
in 1983 (Burnett 1999, page 162).
Nick Novakowski and John Heppes worked well
together. Their first task was to convince provincial gov-
ernments that CITES would not interfere with provin-
cial authority over wildlife resources. John dealt with
the administrators and enforcement personnel while
Nick met with the wildlife managers and scientists.
“Nick showed an amazing understanding and ability
in persuading all concerned to fully cooperate ...” (J.
Heppes, personal communication). Canada ratified the
Convention in early 1975. Every two years there was
2006
a “Conference of the Parties” at which member nations
negotiated changes. “It was at these Conferences that
Nick really proved his worth. In his role as Scientific
Advisor he was always able to present the Canadian
viewpoint in a clear and concise manner. While usually
a calm and collected person, Nick could at times be
very forceful in getting the Canadian opinion across
... he was respected by many international authorities
who often sought his advice on biological status of en-
dangered species and his assistance in presenting their
positions.” (J. Heppes, personal communication).
Eugene Lapointe, the first Secretary General of
CITES, remembers Nick’s contributions for their sci-
entific rigour and said, “My recollection of Nick is that
he played a key role at the CITES meeting held in Gab-
arone, Botswana in 1983. The international commu-
nity had an opportunity to realize what real and cred-
ible science was all about, and that thanks to Nick. I
always perceived him as one of the most candid scien-
tists. He was highly respected by his peers from the
CITES community” (E. Lapointe, personal communi-
cation).
I recall one meeting in Ottawa where Canadian Non-
Government Organizations (NGOs) were presenting
their concerns about an upcoming Conference of the
Parties. Two delegates were particularly vociferous in
their assault on a proposed Canadian position. Nick as
usual was thoroughly prepared and met both scientific
and emotional arguments with calm, well-reasoned,
non-confrontational responses which won the day.
Nick was also front and centre in the discussions lead-
ing up to the establishment of the Committee on the
Status of Endangered Wildlife in Canada (COSEWIC).
Ted Mosquin recalled that as he was preparing draft
recommendations for an upcoming symposium on
Canada’s Threatened Species and Habitats in 1975 Nick
suggested that he look into the UK model that had “a
unique committee that addressed the matter of the
status of endangered species in the UK. [Nick] came
over to the CNF office to fill me in on how the UK
system worked. The result was a draft recommenda-
tion ... which led to almost immediate establishment
of COSEWIC by agreement of CWS and provincial
agencies via the Federal Provincial Wildlife Confer-
ence” (T. Mosquin, personal communication).
“Throughout Nick’s work as a senior scientist in
headquarters, he maintained a network of colleagues in
provincial and territorial governments. Nick’s credibil-
ity with these colleagues was vital in their governments’
agreement in 1977 to the initiative that became the
Committee on the Status of Endangered Wildlife in
Canada (COSEWIC), with Nick elected as its first per-
manent secretary. During the first formative years of
COSEWIC Nick did outstanding work helping to forge
a new way of doing business, helping to bring federal,
provincial, territorial and non-government agency peo-
ple to agree on publicly-released status reports. Making
public the facts about endangered species was what
Nick had been calling for since 1967, believing that this
A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004
373
would lead to remedial actions supported by the public
COSEWIC continued to grow in the spirit of Nick's
early aspirations, and recently was established as a for-
mal entity under the Species at Risk Act.” (J. A. Keith,
personal communication).
In addition to Nick’s work as COSEWIC secretary,
he maintained his avid interest in species biology. He
wrote the original Wood Bison status report and con-
tributed to others. He also chaired the Marine Mammal
and Terrestrial Mammal subcommittee in 1980 and,
when it split, he chaired the Terrestrial Mammal sub-
committee from 1981 to 1984.
Dr. Novakowski was involved in a much broader
scope of activities than this review can encompass but
three items may help to round out the story. For a num-
ber of years in the 1960s and 1970s he was involved
with the search for more humane traps for capturing
furbearers. His field experience with trappers proved
very useful. He thoroughly understood the trappers’
needs (Novakowski 1958d*) and also the political pres-
sure being created to do away with the leg-hold trap. In
1967 he gave a brief but cogent review of the problem
to the Federal-Provincial Wildlife Conference (Nova-
kowski 1967e) and continued to be involved with the
resulting program to develop humane traps until his
retirement. Another example of his varied work in-
volved blocking the introduction by an Ontario fur
rancher of several dozen Raccoon Dogs (Nyctereutes
procyanoides) from Finland. When the news arrived on
Nick’s desk, he nearly blew a fuse! He was one of the
very few people in Canada who had the knowledge
to foresee a disastrous outcome if those animals were
allowed to stay. He knew from the literature and from
his many European colleagues that if the animals ever
got loose the species could have devastating ecological
effects. And he knew that no matter what precautions
were taken, sooner or later some of them would get
loose. He sought the assistance of Dr. Eric Broughton,
a veterinarian with the CWS, to have Agriculture Cana-
da deny the import permit. No dice — there was no basis
for denial on environmental grounds. Nevertheless,
based on knowledge of the ecology of the species, he
convinced Canadian authorities that the animals posed
an unacceptable threat to Canadian wildlife and ob-
tained authorization to have all of the imports killed,
the pelts processed by the Hudson’s Bay Company and
compensation made to the owner. But he didn’t leave
it there. Nick pursued the problem domestically and
with the United States authorities resulting in a formal
Canada—USA agreement to bar the future importation
or Raccoon Dogs into either Canada or the USA (E.
Broughton, personal communication). A third dimen-
sion of Nick’s professional work involved teaching a
variety of courses, including biology, terrestrial ecol-
ogy, and wildlife management at University of Ottawa,
the University of Guelph, and Sir Sandford Fleming
College in Lindsay, Ontario.
After retiring from the Canadian Wildlife Service in
1983, Dr. Novakowski became active as a scientific
374
consultant and engaged in a wide range of projects.
In some cases he worked independently. In most he
served as a member of a consulting team. In one of his
early projects, he worked with Ted Mosquin to con-
solidate and review many earlier reports on Caribou
and fur-bearers in Pukaskwa National Park (Novakows-
ki and Mosquin 1986a*, b*, c*, d*). Dr. Mosquin
recently commented: “Nick provided the publications/
reports, guidelines, policies, appendices, survey data,
illustrations, etc. as well as some text. I basically organ-
ized and wrote the reports based on the material and dis-
cussions. He was the expert. We worked well together
and I learned a lot! ... We sometimes sat for more than
a few hours trying to figure out what needed to be said
and how to best say it.”
Several of Dr. Novakowski’s post-retirement proj-
ects dealt with major Environmental Impact Assess-
ments of such projects as the Military Low-Level Fly-
ing program in Labrador, uranium mining in northern
Saskatchewan, the Rafferty-Alameda dams proposal
in southern Saskatchewan, military base proposals in
the arctic, and bison management in Wood Buffalo
National Park. He became very knowledgeable about
environmental impact assessments in Canada and con-
tributed behind the scenes on a variety of policy issues.
He also contributed to a review paper on the evolution
of environmental impact assessments of watershed mod-
ification projects (Dirschl et al. 1992).
In 1982, a year before Nick retired from the Cana-
dian Wildlife Service, Dr. Husain Sadar arrived at the
Federal Environmental Assessment Review Office
(FEARO). In a letter following Nick’s death, Dr. Sadar
wrote that one of his “major responsibilities was to
organize teams of highly recognized, accomplished and
credible independent experts and specialists to assist
and advise [the independent review panels] for analyz-
ing and addressing often complex scientific, technical
and socio-economic issues ... Nick served as an inde-
pendent expert for several panels [dealing with] some
of the most difficult, controversial and complex mega
project proposals ... the panels benefited immensely
from Nick’s unique experiences and expertise in wild-
life management and habitat protection. Whether the
issue was protection of caribou herds, moose, large car-
nivores, furbearers, waterfowl, seabirds, birds of prey,
marine birds or deer, Nick’s critical analysis, evalua-
tion and resolution of all such issues was simply out-
standing ... But the most important quality of Nick’s
wonderful character was his ability to interact and
work with other professionals in a highly profession-
al, civilized and friendly fashion. He truly was a real
gentleman.”
Dr. Herman Dirschl, with whom Dr. Novakowski
collaborated on many contracts, recalled their joint
efforts on some of those major projects and comment-
ed, inter alia that, “Nick could be very passionate in his
views, particularly when political decisions seemed to
often favour short term economic gains at the expense
THE CANADIAN FIELD-NATURALIST
Vol. 120
of the long term environmental health. However, he
managed to always stay calm and collected in the face
of such adversity. I admired him for this ability and will
greatly miss him.” Dr. Dirschl also recalled their work
“on an assessment of priority habitat sites throughout
Canada to be considered for protection by the Canadi-
an Wildlife Service in concordance with the Federal
Government’s ‘Green Plan’ goals. Nick took on the res-
ponsibility for reviewing the site records for a total of
262 sites in all provinces and territories; designing a
classification system and defining latitude/longitude
centroids for each site.”
Even Canada Post sought out Dr. Novakowski’s ad-
vice in planning for a commemorative postage stamp
featuring Beaver — the 150" anniversary of the origi-
nal 3¢ Beaver stamp (Novakowski 1997*).
Nick the scientist was also Nick the practical envi-
ronmentalist. Early in their stay in Ottawa he and his
wife, Amy, bought 150 acres of forested land in near-
by Lanark County. It was covered primarily with sec-
ond growth hardwoods but also had a small plantation
of Red Pine which Nick carefully nurtured. He became
very interested in the “Model Forest” concept and was
a very keen member of the “Eastern Ontario Certified
Forest Owners.”
I leave the closing words to his good friend, car pool
partner and long-time professional colleague, Gerry
Lee: “On a house visit one day, Nick invited me into
his garage to see his latest pride and joy. There sat a
beautiful cedar strip canoe, like a collector’s piece of
rare furniture, hand built by canoe builder Nick for one
of his offspring. Apart from the odd comment previous-
ly about “building a boat someday, I had no inkling of
his interest or talent in this area. So much for thinking
you know someone! To sum up, Nick to me was the
consummate good guy; respected professional, devot-
ed family man, equally devoted environmentalist, polite,
humble and honest. A good friend I'll never forget.”
Acknowledgments
Nick’s wife, Amy, was very supportive, provided
copies of important papers, and authorized the use of
the photograph used in this tribute. His brother, Ken,
provided details of Nick’s early family life. Several
librarians were especially helpful in assembling the
lists of Dr. Novakowski’s publications and unpublished
reports: Terri Fraser, Canadian Wildlife Service, Edmonton;
Jean-Francois Bélanger, Environment Canada Library,
Gatineau, Quebec; Carol Schurr, former librarian, and
Rob Wallace, Fisheries Ecologist, Saskatchewan Re-
source Stewardship Branch, Saskatoon; Staff, Library
and Archives Canada, Ottawa, and Keith Van Cleave,
Librarian, Northern Prairie Wildlife Research Centre,
Jamestown, North Dakota. Many former colleagues of
Dr. Novakowski provided assistance: I thank in par-
ticular Bill Fuller, Tony Keith and Gerry Lee for their
contributions and for their very helpful comments on
an earlier draft of this paper. Additional thanks are due
2006
Eric Broughton, Herman Dirschl, Don Flook, Andy
Hamilton, John Heppes, Ernie Kuyt, Eugene Lapointe,
Alan Loughrey, Ted Mosquin, Doug Pollock, Dick
Russell, Husain Sadar, Ward Stevens, John Tener and
Don Thomas. I also thank my wife, Mary Bryant, for
her constant support and assistance.
Literature Cited (exclusive of items listed in Bibliography)
Burnett, J. A. 1999. A passion for wildlife: A history of the
Canadian Wildlife Service, 1947-1997. The Canadian
Field-Naturalist 113(1) 1-214.
Christianson, D. A., P. J. P. Gogan, K. M. Podruzny, and
E. M. Olexa. 2005. Incisor wear and age in Yellowstone
bison. Wildlife Society Bulletin 33(2): 669-676.
Fuller, W. A. 1951. Natural history and economic importance
of the muskrat in the Athabasca-Peace delta, Wood Buffalo
Park. Canadian Wildlife Service, Wildlife Management Bul-
letin Series 1, Number 2, 82 pages, charts, photographs.
Gates, C. C., R. O. Stephenson, H. W. Reynolds, C. G.
vanZyll de Jong, H. Schwantje, M. Hoefs, J. Nishi, N.
Cool, J. Chisholm A. James, and B. Koonz. 2001.
National Recovery Plan for the Wood Bison (Bison bison
athabascae). National Recovery Plan Number 21. Recov-
ery of Nationally Endangered Wildlife (RENEW). Ottawa,
Ontario. 50 pages.
Munro, D. A. 1973. Introductory remarks. Pages 28-29 in
Convention on International Trade in Endangered Species
of Wild Fauna and Flora. Transactions, 37" Federal-Provin-
cial Wildlife Conference, Ottawa.
Bibliography
This bibliography is divided into published and un-
published sections. While many of Nick’s reports were
never published, their listing helps to demonstrate the
breadth of his professional activities. Most reports pre-
pared for the Federal Environmental Assessment and
Review Office (FEARO) are not listed, in line with the
terms of the contracts under which they were produced.
Listings are chronological irrespective of whether Nick
was the sole author, lead author or junior author.
Publications
Fuller, W. A., and N. S. Novakowski. 1955. Wolf control
operations, Wood Buffalo National Park, 1951-52, Canadi-
an Wildlife Service. Wildlife Management Bulletin, Series
1, Number 11. [2] + 20 pages, photographs.
Novakowski, N.S. 1956. Additional records of bats in Sas-
katchewan. Canadian Field-Naturalist 70: 142.
Atton, F. M., and N. S. Novakowski. 1956. The value of
freshwater fish as a mink food. Fur Trade Journal of Cana-
da 33 (11): 12-13, 38, 40-42.
Banfield, A. W. F. and N. S. Novakowski. 1960. The survival
of wood bison (Bison bison athabascae Rhodes) in the
Northwest Territories. Natural History Paper Number 8,
National Museum of Canada. 6 pages, illustrations, map,
tables.
Novakowski, N. S., J. H. G. Cousineau, G. B. Kolenosky,
G.S. Wilton, and L. P. E. Choquette. 1963. Parasites and
diseases of bison in Canada. II. Anthrax epizootic in the
Northwest Territories. Pages 233-239 in the Transactions
of the 28" North American Wildlife and Natural Resources
Conference. Wildlife Management Institute, Washington,
D.C. [CWSC 1357].
A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004
375
Novakowski, N.S. 1965a. Cemental deposition as an age cn
terion in bison, and the relationship of incisor wear, eye-lens
weight, and dressed bison carcass weight to age. Canadian
Journal of Zoology 43: 173-178. [CWSC 1426}
Novakowski, N.S. 1965b. The day we rescued a whooping
crane. Audubon Magazine 67: 230-233.
Novakowski, N.S. 1966. Whooping crane population dynam-
ics on the nesting grounds, Wood Buffalo National Park,
Northwest Territories, Canada. Canadian Wildlife Service,
Report Series Number 1. 19 pages, illustrations, tables,
maps.
Novakowski, N. S. 1967a. The winter bioenergetics of a
beaver population in northern latitudes. Canadian Journal
of Zoology 45 (6): 1107-1118.
Novakowski, N.S. 1967b. Conservation of rare and endan-
gered species of mammals in Canada. Pages 73-75 in the
Transactions of the 31" Federal-Provincial Wildlife Con-
ference, Ottawa.
Novakowski, N.S. 1967c. Humane trapping. Transactions of
the 31* Federal-Provincial Wildlife Conference, Ottawa.
Pages 67-68.
Novakowski, N.S. 1967d. Whooping crane. Modern Game
Breeding 3 (6): 19-25, 35-41, 43. [A reprint of most of
Novakowski 1966]
Novakowski, N.S. 1967e. Beaver — Wood Buffalo National
Park. Page 48 in Canadian Wildlife Service ‘66. Queen's
Printer, Ottawa. Catalogue Number RR66-3766.
Novakowski, N.S. 1967f. Bison — Wood Buffalo National
Park. Page 48 in Canadian Wildlife Service ‘66. Queen’s
Printer, Ottawa. Catalogue Number RR66-3766.
Novakowski, N.S. 1968a. The National Committee on Wild-
life Land Meeting, May 28-30, 1968. Pages 22-23 in the
Transactions of the 32 Federal-Provincial Wildlife Con-
ference, Ottawa.
Novakowski, N. S. 1968b. The Export-Import Convention.
Page 27 in the Transactions of the 32 Federal-Provincial
Wildlife Conference, Ottawa.
Novakowski, N. S. 1968c. Humane Trapping. Pages 27-29
in the Transactions of the 32 Federal-Provincial Wildlife
Conference, Ottawa.
Novakowski, N.S. 1969a. The influence of vocalization on
the behavior of beaver, Castor canadensis Kuhl. American
Midland Naturalist 81: 198-204.
Novakowski, N.S. 1969b. Report on the Administrative Com-
mittee for Polar Bear Research and Management in Canada.
Page 22 in the Transactions of the 33 Federal-Provincial
Wildlife Conference, Ottawa.
Novakowski, N.S. 1970a Endangered wildlife —- mammals.
Pages 6-7 in Endangered Wildlife in Canada. Canadian
Wildlife Federation, Ottawa, Ontario.
Novakowski, N. S. 1970b. Endangered Canadian mam-
mals. Canadian Field-Naturalist 84(1): 17-23.
Jeffrey, W. W., C.S. Brown, M. Jurdant, N.S. Novakow-
ski, and R. H. Spilsbury. 1970. Foresters and the challenge
of integrated resource management. The Forestry Chron-
icle, June: 196-199.
Choquette, L. P. E., E. Broughton, A. A. Currier, J. G.
Cousineau, and N. S. Novakowski. 1972. Parasites and
diseases of bison in Canada. II. Anthrax out-breaks in the
last decade in northern Canada and control measures. Cana-
dian Field-Naturalist 86: 127-132.
Novakowski, N. S. 1973a. Natural resources in the Northwest
Territories and the Yukon. Pages 91-97 in Natural Resource
Development in Canada. Edited by Phillippe Crabbe and
Irene Spry. University of Ottawa Press.
376
Novakowski, N.S. 1973b. Report of the Administrative Com-
mittee for Caribou Preservation. Page 75 in the Transactions
of the 37" Federal-Provincial Wildlife Conference, Ottawa.
Novakowski, N.S., and V. E. F. Solman. 1975. Potential of
wildlife as a protein source of food. Journal of Animal
Science 40(5): 1016-1019.
Novakowski, N.S. 1976. Endangered species. Pages 28-31 in
the Transactions of the 39" Federal-Provincial Wildlife
Conference, Ottawa.
Novakowski, N.S. 1977. [an edited version of Dr. Novakow-
ski’s statements as a witness on behalf of the Canadian
Wildlife Federation before the National Energy Board hear-
ings into a Mackenzie Valley Pipeline.] Pages 24A-24E
in Wildlife Report. The Canadian Scene. May-June 1977.
Canadian Wildlife Federation.
Choquette L. P. E., E. Broughton, J. G. Cousineau, and
N.S. Novakowski. 1978. Parasites and diseases of bison
in Canada. IV. Serologic survey for brucellosis in bison
in northern Canada. Journal of Wildlife Diseases 14(3):
329-332.
Novakowski, N. S., and Geoff Robins. Editors. 1982. Pro-
ceedings, Symposium on Natural Resources Use and Native
Rights in Canada. Canadian Society of Environmental Biol-
ogists, Ottawa. 105 pages. [EC-E] [EC-O]
Novakowski, N. S. 1984. Control of mammals at airports.
Pages 183-186 in Proceedings, Wildlife Hazards to Aircraft.
Conference and Training Workshop, Edited by M. J. Har-
rison, S. A. Gauthreaux and L. A. Abron-Robinson. U.S.
Department of Transportation; Federal Aviation Admin-
istration; Office of Airport Standards, Charleston, North
Carolina.
Hill, E. P., and N. S. Novakowski. 1984. Beaver manage-
ment and economics in North America. Acta Zoologica
Fennica 172: 259-262.
Novakowski, N.S. 1985. Beaver management in Canada; a
review. Proceedings of the 4" International Theriological
Congress, Edmonton, Alberta. [Abstract] Session S08.
Dirschl, H. J., N.S. Novakowski, and M. H. Sadar. 1993.
Evolution of environmental impact assessment as applied
to watershed modification projects in Canada. Environ-
mental Management 17(4): 545-556.
Novakowski, N.S. 2003. The Wilson River Escapade. Pages
8-10 in Then: Friends sharing memories. Edited and pub-
lished by Mary and Joe Bryant, Ottawa. vii + 256 pages,
illustrations. ISBN 0-9733372-0-6.
Unpublished papers (marked * where cited) [EC-E] =
Canadian Wildlife Service Library, Edmonton. [EC-O] =
Environment Canada Library, Ottawa (Gatineau, Quebec).
[ES-S] = Environment Saskatchewan, Saskatoon.
Atton, F. M., and N. S. Novakowski. 1953a. Utilisation of
fish on mink ranches in the Buffalo Narrows Region. Fish-
eries Technical Report 53-1. 17 pages. [ES-S]
Atton, F. M., and N. S. Novakowski. 1953b. The value of
freshwater fish as mink food. Fisheries Technical Report
53-5. 10 pages. [ES-S] (See also published version, Atton
and Novakowski, 1956.)
Atton, F. M., and N.S. Novakowski. 1954a. Analysis of sam-
pling the commercial fishery of Lake Athabasca, 1951-1953.
Fisheries Technical Report 54-4. 11 pages. [ES-S]
Atton, F. M., and N. S. Novakowski. 1954b. Analysis of
sampling the commercial fishery of Lake Athabasca, 1954.
Fisheries Technical Report 55-3. 7 pages. [ES-S] [EC-E]
THE CANADIAN FIELD-NATURALIST
Vol. 120
Novakowski, N. S. 1955a. Report on biological and fish-
eries survey of Reindeer Lake, 1954. Fisheries Technical
Report 55-1. 62 pages. [ES-S]
Novakowski, N.S. 1955b. The ecology of Reindeer Lake with
special reference to fish. Fisheries Manuscript Report M-4.
62 pages. [ES-S]
Novakowski, N.S. 1955c. The ecology of Reindeer Lake with
special reference to fish: a thesis submitted to the Faculty
of Graduate Studies in partial fulfillment of the require-
ments for the degree of Master of Science in the Depart-
ment of Biology, University of Saskatchewan. 99 pages,
2 fold-out maps, figures.
Atton, F. M., and N. S. Novakowski. 1956a. Biological sur-
vey and fisheries management of Waterhen Lake and adjoin-
ing lakes. Fisheries Technical Report 56-4. 50 pages. [ES-S]
Atton, F. M., and N.S. Novakowski. 1956b. Biological sur-
vey and fisheries management of Lac des Iles. Fisheries
Technical Report 56-3. 34 pages. [ES-S]
Atton, F. M., and N. S. Novakowski. 1956c. Biological
survey and fisheries management of Jackfish and Murray
Lakes. Fisheries Technical Report 56-2. 66 pages. [ES-S]
Atton, F. M., and N. S. Novakowski. 1956d. The value of
freshwater fish as mink food. Fisheries Technical Report.
2 pages. [ES-S] (See also published version, Atton and
Novakowski 1956.)
Novakowski, N. S. 1956a. Analysis of autopsy records of
wolves taken in the Eskimo Point area 1956. Canadian
Wildlife Service, CWSC 51: 12 pages, 2 tables. [EC-E]
[EC-0]
Novakowski, N.S. 1956b. Resurvey of beaver habitat in the
Fort Simpson, Fort Liard and Wrigley areas, 1956. Cana-
dian Wildlife Service, CWSC 753: 4 pages, maps, illustra-
tions. [EC-E] [EC-0]
Flook, D. R., and N. S. Novakowski. 1956. An experiment
in poisoning wolves on the arctic tundra. Part A. CWSC
49: 9 pages, 2 maps. [EC-O]
Novakowski, N.S. 1957a. Aerial resurvey of bison in Wood
Buffalo National Park and surrounding areas, 1957. Cana-
dian Wildlife Service, CWSC 216: 12 pages, maps. [EC-E]
[EC-O]
Novakowski, N. S. 1957b. Report on tagging, testing and
slaughtering of bison in the Lake Claire area Wood Buf-
falo National Park, October, November 1957. Canadian
Wildlife Service, CWSC 728: 14 pages. [EC-E] [EC-O]
Novakowski, N.S. 1958a. Report on the testing, tagging and
slaughter of bison in Wood Buffalo National Park, October-
November, 1958. Canadian Wildlife Service, CWSC 764:
18 pages, illustrations, charts. [EC-E] [EC-O]
Novakowski, N.S. 1958b. Fur resources survey of Wood Buf-
falo National Park. Canadian Wildlife Service, CWSC 749:
45 pages, tables. [EC-E] [EC-0]
Novakowski, N.S. 1958c. Report on the hatching and plant-
ing of rainbow trout Fort Smith, N.W.T., June 10-26, 1958.
Canadian Wildlife Service, CWSC 1050: 5 pages. [EC-E]
[EC-O]
Novakowski, N.S. 1958d. Report on the testing of killer traps
as a substitute for leghold traps. Canadian Wildlife Service.
CWSC 772: 8 pages, photographs.
Novakowski, N.S. 1959a. Aerial resurvey of the higher den-
sity bison areas in Wood Buffalo Park and the N.W.T. Cana-
dian Wildlife Service, CWSC 883: 5 pages, maps. [EC-E]
[EC-O]
2006
Novakowski, N.S. 1959b. Analysis and appraisal of a three
year aerial survey of beaver habitat in the southern and cen-
tral areas of the Mackenzie District, N.W.T., 1956, 1957,
1958. Canadian Wildlife Service, CWSC 884: 17 pages,
tables, maps. [EC-E] [EC-O]
Novakowski, N.S. 1959c. Fish hatchery operations at Pine
Lake — June 1959. Canadian Wildlife Service, CWSC 1051:
7 pages. [EC-E] [EC-O]
Novakowski, N. S. 1959d. Report on the investigation of
probable Wood Bison in the Nyarling River area and an
investigation of historical Wood Bison range in the Fort
Providence area. Canadian Wildlife Service, Fort Smith,
CWSC 3493: 12 pages, tables, photographs. [EC-E] [EC-O}]
Novakowski, N. S. 1961a. Cemental deposition as an age cri-
terion in bison, and the relationship of incisor wear, eye-
lens weight and dressed bison carcass weight to age. Cana-
dian Wildlife Service, Fort Smith, N.W.T. CWSC 1426:
10 pages [EC-E] [EC-O] See also published version, Nova-
kowski 1965a.
Novakowski, N.S. 1961b. Estimates of the bison population
in Wood Buffalo National Park and the Northwest Territo-
ries based on transect and total counts. Canadian Wildlife
Service, CWSC 882: 6 pages, maps. [EC-E] [EC-O]
Novakowski, N.S. 1961c. Total counts of bison in the higher
park density areas of Wood Buffalo Park. Canadian Wildlife
Service CWSC 889: 11 pages, illustrations, map. [EC-E]
[EC-O]
Novakowski, N.S., and L. P. E. Choquette. 1961. Report on
the testing and slaughter of bison at Sweetgrass, Wood Buf-
falo National Park. Canadian Wildlife Service, CWSC 1102:
10 pages. [EC-E] [EC-O]
Novakowski, N.S. 1962. Proposals for a bison management
policy. Canadian Wildlife Service: 3 pages. [EC-E]
Novakowski, N.S. 1963a. Report on anthrax in bison in the
Hook Lake and Grand Detour areas, N.W.T., 1963. Cana-
dian Wildlife Service: 4 pages. [EC-E]
Novakowski, N. S. 1963b. Report on the transfer of Wood
Bison, 1963. Canadian Wildlife Service: 5 pages, maps.
[EC-E]
Novakowski, N.S. 1963c. Wood bison transfer: completion
report. Canadian Wildlife Service: 4 pages. [EC-E]
Novakowski, N.S., and L. P. E. Choquette. 1963. Bison test-
ing and slaughter at Hay Camp, Wood Buffalo National
Park 1963. Canadian Wildlife Service, CWSC 1103: 9
pages. [EC-E] [EC-O]
Novakowski, N.S. 1964a. The effect of disease syndromes
on the ecology of the bison in Wood Buffalo National Park.
Canadian Wildlife Service: 9 pages. [EC-E]
Novakowski, N. S. 1964b. Report and recommendations con-
cerning the occurrence and control of anthrax in bison in
Wood Buffalo National Park and vicinity. Canadian Wild-
life Service: 5 pages, maps. [EC-E]
Novakowski, N.S. 1964c. Report on the assessment of pro-
posed bison range in the Saskatchewan River delta. Cana-
dian Wildlife Service: 11 pages, illustrations, table, maps.
[EC-E]
Novakowski, N. S. 1964d. Slaughter report - Hay Camp,
November 1964. Canadian Wildlife Service, CWSC 1104:
9 pages. [EC-E] [EC-O]
Novakowski, N.S. 1964e. The use of eye-lens weight, tooth
replacement, cementum deposition and dressed weight as
aging criteria in the bison. Canadian Wildlife Service: 11
pages, illustrations. [EC-E]
Novakowski, N.S. 1965a. Population dynamics of a beaver
population in northern latitudes. Ph.D. thesis, University of
A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004
377
Saskatchewan, CWSC 1019: x + 154 pages, figures. [EC-E]
[EC-0}
Novakowski, N.S. 1965b. The effects of disease upon bison,
Mackenzie District. Canadian Wildlife Service, CWSC
1696: 2 pages. [EC-E] [EC-O]
Novakowski, N.S. 1965c. Possible bison range, Cumberland
marshes. Canadian Wildlife Service: 2 pages, maps. [EC-E}
Novakowski, N.S. 1965d. Slaughter report, Grand Detour,
1964-65. Canadian Wildlife Service, CWSC 1105: 2 pages
[EC-E] [EC-O}
Novakowski, N.S. 1965e. Wilderness rescue of an injured
whooping crane: parents try and [sic] fend off an invad-
ing helicopter in the wild muskegs of northern Canada.
Canadian Wildlife Service: 5 pages. [EC-E}
Novakowski, N. S., and W. E. Stevens. 1965. Survival of
wood bison (Bison bison athabascae Rhodes) in Canada.
Canadian Wildlife Service, Edmonton, 5 pages. [EC-E}
[Paper delivered to the 45" annual meeting of the American
Society of Mammalogists, Winnipeg, 20-26 June, 1965. An
abstract is filed under CWSC 1020 in EC-O.]
Novakowski, N.S. 1967a. Anticipated ecological effects of
possible changes in the water levels of the Peace River-
Athabasca River delta as a result of the damming of the
Peace River. Canadian Wildlife Service, CWSC 1086: 17
pages, maps. [EC-E] [EC-O]
Novakowski, N. S. 1967b. Bighorn sheep study meeting,
Edmonton, February 28, 1967. Canadian Wildlife Service:
10 pages. [EC-E]
Novakowski, N.S. 1967c. Concepts of multiple-use and inte-
grated resource management. Canadian Wildlife Service:
10 pages. [EC-E]
Novakowski, N.S. 1967d. Investigation of the Killdeer-Val
Marie area in southern Saskatchewan as possible bison
range. Canadian Wildlife Service, CWSC 1085: 9 pages,
[+2] , maps. [EC-E] [EC-O]
Novakowski, N. S. 1967e. Rare and endangered mammals
in Canada. Canadian Wildlife Service, CWSC 2991: 141
pages. [EC-E] [EC-O]
Novakowski, N. S. 1967f. Wood Buffalo National Park.
Canadian Wildlife Service: 8 pages, map. [EC-E]
Novakowski, N.S., and L. G. Blight. 1967. Investigation of
Athabasca delta to assess feasibility, cost of constructing a
rock plug and subsequent effects resulting from restricting
the flow of water from the Lake Claire area. Canadian
Wildlife Service. CWSC 1357: 5 pages, photographs, chart,
map. [EC-E] [EC-O]
Novakowski, N.S., and G. More. 1967. Mammalian pest and
predator problems in Canada. Canadian Wildlife Service,
Ottawa. CWSC 1703: 67 pages. [EC-E] [EC-O]
Novakowski, N. S., and G. More. no date. Contributions
toward a national pest control policy for Canada — mam-
mals. Canadian Wildlife Service, CWSC 3444: 56 pages.
[EC-E] [EC-O]
Novakowski, N.S. 1970. Fire priority report, Wood Buffalo
National Park. Canadian Wildlife Service, CWSC 1305:
6 pages, maps. [EC-E] [EC-O]
Novakowski, N.S. 1971a. An overview study of the tourist
potential of the Northwest Territories — wildlife. Canadian
Wildlife Service, CWSC 1370: 8 pages. [EC-E] [EC-O]
Novakowski, N. S. 1971b. Report to the National Energy
Board on staffing requirements for an Environmental Divi-
sion. CWSC 1369: 5 pages, 10 appendices, chart. [EC-O]
Novakowski, N. S. 1975. Evidence concerning impacts of
pipelines on rare and endangered species and aquatic and
terrestrial animals. Paper presented to the Mackenzie Pipe-
378
line enquiry at the request of the Canadian Arctic Resources
Committee. Canadian Wildlife Service, CWSC 2143: 33
pages, appendices. [EC-E] [EC-O]
Novakowski, N. S., G. More, and P. Reilly. 1975. Exotic
mammals and birds in Canada — a historical review. Cana-
dian Wildlife Service, CWSC 1704: 47 pages. [EC-E]
[EC-O]
Novakowski, N. S., and T. Mosquin. 1986a. Caribou Man-
agement Plan; Pukaskwa National Park, Volume I. 53
pages. Natural Resource Conservation, Ontario Region,
Environment Canada, Ottawa. Consultant Report to Parks
Canada by Mosquin Bio-Information Limited, Box 279
Lanark, Ontario.
Novakowski, N. S., and T. Mosquin. 1986b. Caribou Man-
agement Plan; Pukaskwa National Park, Volume II (Imple-
mentation Strategy). 32 pages. Natural Resource Conser-
vation, Ontario Region, Environment Canada, Ottawa.
Consultant Report to Parks Canada by Mosquin Bio-
Information Limited, Box 279 Lanark, Ontario.
Novakowski, N. S., and T. Mosquin. 1986c. Furbearer
Management Plan; Pukaskwa National Park, Volume I.
69 pages. Natural Resource Conservation, Ontario Region,
Environment Canada, Ottawa. Consultant Report to Parks
Canada by Mosquin Bio-Information Limited, Box 279
Lanark, Ontario.
Novakowski, N. S., and T. Mosquin. 1986d. Furbearer Man-
agement Plan; Pukaskwa National Park, Volume II (Imple-
mentation Strategy). 69 pages. Natural Resource Conser-
vation, Ontario Region, Environment Canada, Ottawa.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Consultant Report to Parks Canada by Mosquin Bio-
Information Limited, Box 279 Lanark, Ontario.
Novakowski, N.S. 1989. Historical origin of National Park
Plains Bison. Environment Canada, Canadian Park Service,
Natural Resources Branch, Ottawa. 31 pages.
Novakowski, N.S. 1990. Northern Diseased Bison Environ-
mental Assessment Panel compendium of supplementary
submissions, March 1990. Federal Environmental Assess-
ment Review Office, Ottawa. 308 pages, maps. [EC-E]
Novakowski, N.S., and H. J. Dirschl. 1992. Assessment of
priority sites for the establishment of a Network of Con-
servation Areas by the Canadian Wildlife Service. Environ-
mental Social Advisory Services (ESAS) Inc., Ottawa,
Ontario.
Dirschl, H., N.S. Novakowski, and L. C. N. Burgess. 1992.
An overview of the biophysical environmental impact of
existing uranium mining operations in northern Saskat-
chewan. vi + 90 pages, appendices. Report prepared for
the Saskatchewan Uranium Mine Development Review
Panel. Environmental Social Advisory Services (ESAS)
Inc., Ottawa, Ontario.
Novakowski, N.S. 1997. The beaver in Canada. A textual and
visual exposition of the life history of the beaver, Castor
canadensis. A report commissioned by the Stamp Products
Division of the Canada Post Corporation. ii + 35 pages,
illustrations.
Received 13 January 2006
Accepted 3 May 2006
Book Reviews
ZOOLOGY
The Birder’s Companion
By Stephen Moss. 2007. Firefly Books Ltd., 66 Leek Cres-
cent, Richmond Hill, Ontario L4B 1H1 Canada. 208
pages. U.S. $16.95
This book is an entry in the “everything you ever
wanted to know about birds” category. Each of its ten
chapters deals with some aspect of birds — where do
birds live? How do birds eat? Where do birds go?
And, yes, each chapter does consist of a series of ques-
tions and answers — over 450 in all — together with
side-bars giving what can best be described as “gee-
whiz” facts relevant to the chapter’s topic, all in rough-
ly 20 pages or less each.
Maybe a disclaimer is appropriate at this point — this
kind of book turns me off. With that out of the way,
Moss has done a creditable job in amassing a formi-
dable series of facts and presenting them in a simple,
colloquial style. In the introduction the author quotes
a friend who suggests the book would appeal to “intel-
ligent, inquiring eleven-year-old boys”, and while the
author clearly hopes to appeal to a much wider audi-
ence, he makes no apology for his approach.
In spite of its disarming level of simplicity, the book
is very well thought-out, up-to-date, and quite accurate.
I imagine I could find an error or two in all this mass
of data if I tried hard enough, but the fact remains that
I didn’t. He even — glory be — gives the correct origin
for the word “jizz” , with no inanities about “general
impression of size and shape” usually parroted as the
origin nowadays. My main quarrel with the text is the
inevitably high level of simplification, which constant-
ly left me feeling “yes, but...” Some of the author’s
generalizations seemed very broad, and I wondered
how he arrived at them; but again the ones I could
check seemed within reasonable bounds.
The author is British, and this can sometimes be a
problem when writing for a North American audience.
Birds of the Dominican Republic and Haiti
By Steven Latta, Christopher Rimmer, Allan Keith, James
Wiley, Herbert Raffaele, Kent McFarland, Eladio Fernan-
dez. Princeton Field Guides, Princeton University Press.
2006. 258 pages. U.S. $35.
Hispaniola, the second largest island in the Carib-
bean, has a wealth of habitats and of birds, making it
an emerging destination for birdwatchers. It also has
immense development challenges, most acutely felt
in Haiti, the poorest country in the western hemisphere.
However, in this case the book is very carefully tailored
to North America. One minor area where it could be
misleading was in references to “buzzards”, the Euro-
pean name for Buteo hawks, but also a term colloqui-
ally used in parts of North America for vultures. Not
only are we not told what buzzards are, but there are
no buzzards in the index. One reference, to them being
eaten by the European Eagle Owl [also not in the
index]; was a particularly poor choice, given that Great
Horned Owls will attack Red-tailed Hawks similarly.
The weaknesses in the index are by no means con-
fined to these two examples. It seems to place most
of its emphasis on the material in the side-bars, and
the text as a whole is far less well covered. This is
particularly unfortunate because for me, at least, this
is not a book to read from cover to cover, but is best
dipped into from time to time, as some topic of interest
arises. For such uses, a thorough index is very impor-
tant.
There is a one-page bibliography, although again
some transatlantic emphasis creeps in, as many of the
references are British — it’s hard to see the relevance of
the UK breeding bird atlas — but more importantly they
may be hard to find on this continent. The line illus-
trations are not one of the strengths of the book.
My carping aside, this book would appeal to novice
birders or anyone with a mild interest in birds. Its
appeal to more experienced birders is perhaps more
limited, but it’s an easy read, best taken in small doses,
and you'll probably find some things you didn’t know.
And I seriously think it could be a very good gift for
intelligent, inquiring eleven-year-olds.
CLIVE E. GOODWIN
1 Queen Street, Suite 405, Cobourg, Ontario K9A 1M8
Canada
These development challenges translate into conser-
vation challenges, putting Hispaniola’s ecological rich-
ness at peril. This book, the latest in several Caribbean
guides issued by Princeton, is thus doubly important,
both as an aid to identifying the island’s birds, but also
as a valuable conservation tool which maps the cur-
rent avian diversity of the island and communicates the
value and beauty of the island’s birds to local popula-
tions.
379
380
The book will be instantly familiar to anyone who
has seen any of Princeton’s earlier Caribbean guides,
such as A guide to the birds of the West Indies by Raf-
faele et al. Indeed, some of the plates have been taken
from that earlier guide. The book follows a standard
formal, with front pieces speaking to the topography
and habitats of the island (from the highest mountains
in the Caribbean, at 3098 m, to the lowest lake, at 44 m
below sea level), endemic species (31) and subspecies
(50), avian conservation, and the island’s ornithological
history. This is followed by detailed species accounts
with information on similar species, vocalizations, hab-
itat preferences, status, nesting, and global range, as
well as a “comments” section containing interesting
anecdotes such as the place of the bird in local culture.
Each species account is accompanied by an up-to-date,
detailed map of the bird’s distribution on the island.
This information is complemented by 56 colour plates,
most of which group up to 10 similar species however,
pride of place is given to the island’s endemic species
and one endemic family (Palmchat), which merit larg-
er portraits. The book closes with descriptions of 14
birding sites and a checklist.
The species accounts are accurate and provide suf-
ficient information to identify most species. Particular
attention is paid to the resident species most likely to be
of interest to visitors, thus ensuring that one is indeed
able to safely differentiate Long-billed and Narrow-
billed Todys, the vireos, and other potentially confus-
ing groups. Canadian readers will find the discussion
of Bicknell’s Thrush of interest as Hispaniola is the
chief wintering ground of this enigmatic breeding spe-
cies. In general, the treatment of migrants is better than
in many other regional guides; while you might want
to lug around a good North American guide for some
of the shorebirds and basic plumaged warblers, the
level of detail in the plates is impressive and will usu-
ally suffice (e.g, if you look closely you can see the
differently marked tertials in the illustrations of Long-
billed and Short-billed dowitchers). In general the
plates have the rather washed out and sombre tones of
the earlier Princeton guides, and in some of the por-
traits the feathers have a rather messy look, not unlike
old museum specimens, but this is a stylistic quibble;
on the whole the plates are very good. While the deci-
THE CANADIAN FIELD-NATURALIST
Vol. 120
sion to illustrate all 306 species which have been re-
corded on the island has considerable merit, having
three illustrations of a bird which has only occurred
once (Swainson’s Hawk) would seem an inappropriate
use of always scarce space.
The authors largely follow standard taxonomy, but
have jumped the gun on American Ornithologists’ Union
orthodoxy by accepting recently proposed revisions,
such as treating the Hispaniolan Nightjar as distinct
from the Cuban Nightjar, both traditionally combined
as one species known as Greater Antillean Nightjar
(confusingly, the latter name is used in the similar spe-
cies account under Least Pauraque, one of relatively
few editorial errors in the book). Similarly, the authors
have abolished the names of two endemics, Green-
tailed Ground-Warbler and White-winged Warbler
which have now morphed into Green-tailed Ground-
Tanager and Hispaniolan Highland-Tanager, respec-
tively. Some other incipient splits and lumps are sug-
gested in the text, reflecting the fact that yet more work
needs to be done on the island. Likewise, the species
accounts indicate that several species have been added
to the island’s avifauna in the last decade, suggesting
a recent increase in coverage and the possibility that
the island’s bird list will grow: the local occurrence of
several pelagic species in particular seems to be poor-
ly known.
Who should buy this book? Someone with a gen-
eral interest in Caribbean birds may be better served
by A guide to the Birds of the West Indies by Raffaele
et al., which covers Hispaniola along with the rest of
the islands. But anyone planning to travel to this
beautiful island, or with a keen interest in Caribbean
ornithology, would be well advised to acquire a copy;
the range maps and site guides alone are worth the
price if you are planning a self-guided visit (but bet-
ter still, go with one of the organizations cited in the
book). Finally, and most importantly, there should be
a copy in every school and public library on the island:
one hopes that Spanish and French versions are in
the works.
MARK GAWN
Mission of Canada, Chemin de |’Ariana 5, Geneva 1202,
Switzerland
Atlas of Bird Migration — Tracing the Great Journeys of the World’s Birds
Edited by J. Elphick. 2007. Firefly Books Ltd., 66 Leek Cres-
cent, Richmond Hill, Ontario L4B 1H1 Canada. 176 pages.
$35.
Firefly Books has come out with another decent
book. They’ve claimed the target audience to be, “bird
enthusiasts, naturalists and the inquiring reader” and
those who “wish to comprehend the perilous voyages
that constitute one of the great wonders of the natural
world.” The book is written to be consumable by peo-
ple in grade school and perhaps almost fully under-
stood by those as young as 12 or 14 years of age; it will
be an enjoyable, and not too taxing to read for adults
who have a passing interest in the topic. Serious birders
will want to go elsewhere for more in-depth descrip-
tions of the topics.
The book starts out with 15 two-to-four page chap-
ters on the biology and environmental conditions of
migration, including timing, flight styles, genetics and
weather. For the most part, these are well-illustrated
2006
synopses of the topics. The subsequent chapters are
organized into sections covering major geographic
areas (e.g, Eurasia, southern hemisphere, oceans), with
two to eighteen chapters per section. Each chapter cov-
ers an order or family of birds, using one to four species
as examples. There is a map as well as a calendar show-
ing migration and breeding times for most of the illus-
trated birds.
Although the book is worldwide in scope, it is dom-
inated by a British point of view; after the typical migra-
tion pattern of a species is noted, the accidental occur-
rences in Britain (but nowhere else) are described (if
such has occurred). The global perspective is worth-
while, though only a small fraction of the planet’s
migratory birds can be described.
There are a few mistakes scattered throughout the
book; these include classifying the Great Horned Owl
Handbook of the Birds of the World Volume II
By Josep del Hoyo, Andrew Elliott and David A. Christie
(Editors). 2006. Lynx Edicions, Montseny, 8, 08193 Bel-
laterra, Barcelona, Spain. 800 pages. 199 Euros Cloth.
Handbook of the Birds of the World. Volume IT cov-
ers eight families containing 733 species each with its
own account and distribution map. The first of these is
the Old World Flycatchers which, with the prejudices
of youth, conjure up visions of Little Brown Jobs
(LBJs). LBJs are the birders bane; small obscure birds
that are difficult to see and identify. My first encounter
with an Old World Flycatcher was the Spotted Fly-
catcher. This is an LBJ where “spotted’ is an over-
statement. They are only faint marks on the breast. This
is a Sweeping and somewhat unfair assessment as more
than 40 percent of Old World Flycatchers are colour-
ful. Indeed, the 22 members of Cyorrus are as colour-
ful as North American Bluebirds.
In the next family, the Batises are small and largely
black-and-white, whereas Wattle-eyes are somewhat
more colourful with some orange and yellow. Many of
the Monarch-flycatchers are a smart blue, but others are
black and brown or black-and-white. However, the
members of the small family of Paradise Flycatchers
are among the most spectacular of the world’s birds.
One of my life’s highlights was seeing the display of
the black-and-white morph of the Madagascar Paradise
Flycatcher. Similar and equally impressive species live
in Africa and India. Kinglets and Firecrests is a family
that is split between the old and new worlds. They are
woodland sprites that can be surprisingly “cute” des-
pite an initial dull impression. Gnatcatchers is a fami-
ly of small, grey, long-tailed birds and is the only one
in this volume that is confined to the new world. To
my mind their counterparts are the Prinias of Africa
and Asia.
Cisticolas and allies are a bit of a nightmare. Most
of them are true LBJs being streaky or dull brown.
Knowing their songs and distribution helps. Even the
BOOK REVIEWS
38]
as an irruptive species, labelling Lake Winnipeg and
its neighbours as the Great Lakes and describing the
furcula as a pit (it is the wishbone). Many folks will
gloss over these, but the more demanding reader will
catch them; mistakes are never desirable, but most of
these are relatively minor.
In addition to covering distance migrants, the book
pays some attention to altitudinal migrants. When ap-
propriate, the authors have included conservation mes-
sages on topics such as introduced species, habitat loss
and the implications of commercial fisheries on birds.
Overall, this wouldn’t be a book I'd give one of my
serious birder friends as a gift, but it would do nicely
for the beginner who is just migrating into the hobby.
RANDY LAUFF
St. Francis Xavier University, Antigonish, Nova Scotia
— Old World Flycatchers to Old World Warblers
easier to recognise birds like the Common Jery of
Madagascar and the long-billed Tailor Bird of Asia can
be a problem. They are so active that critical charac-
teristics can be hard to see. Whenever you think you
have seen all the relevant characteristics, after the bird
has flown, you often discover there was something
critical you missed!
The 270 species of Old World Warblers in the Sylvi-
idae (with 14 species confined to the antique island of
Madagascar) are mostly LBJs. They are some of the
most difficult birds to identify in the field. This night-
mare is composed of several genera with hyphenated
names like bush-warbler, reed-warbler, leaf-warbler and
swamp-warbler. While there are discernable differences
between, say, leaf- and swamp-warblers within each
group there are many look-alike species which vary
only by minuscule differences in eye-stripe or bill
length. I spent every lunch hour for two weeks beside
a Willow Warbler’s nest so that I could identify this
species with confidence. This meant I could better see
the subtle difference between Willow Warblers and Chif-
fchafs [greener versus browner, etc.]. Unfortunately these
birds are variable. The reader need only compare the
photo of a pale, greyish Willow Warbler on page 551
with the brownish-yellow one on page 514 to under-
stand this difficulty. It gets even worse with birds like
the newly split Booted Warbler [Hippolais caligata] and
Syke’s Warbler [H. rama] Syke’s Warbler winters all
over India whereas H. caligata is found only on the
eastern coast between the Ganges and Pondicherry. So
the two species overlap in the east and are almost iden-
tical. As they do not sing in winter it will take a DNA
test to separate them!
So how did Handbook of the Birds of the World.
Volume II fare in dealing with birds that are not as mag-
nificent as albatross or wildly coloured like parrots?
It actually does remarkably well. Once again we have a
well-organized and researched volume that adds to the
growing status of this project. As each new volume is
produced it adds to the benchmark status of the exist-
ing volumes as a primary source of information on the
world’s birds. The format has remained stable for the
last several volumes, which makes it easy to use and
compare. The photos are top rate despite the small and
obscure nature of these species. I know how difficult
Conservation of the Black-tailed Prairie Dog:
Edited by John Hoogland. 2006. Island Press, Washington,
DC. 350 pages paperback U.S. $35.
It is amazing how the conservation of an animal as
interesting and entertaining as the Black-tailed Prairie
Dog can be so controversial. But controversial it has
been in the last few decades as environmental inter-
ests have woken up to the fact that the campaign of
Prairie Dog poisoning and habitat loss has resulted in
a range contraction to less than 2% of the historical
geographic distribution. To this day many ranchers
actively dislike Prairie Dogs for their competition with
livestock for rangeland resources. In contrast, some
conservation practitioners think that enough other spe-
cies rely on the habitat created on Prairie Dog colonies
that Prairie Dogs should be considered a keystone
species. In my own practice as a biologist at Grass-
lands National Park, Saskatchewan, I have frequently
been taken aback by how polarized the opinions on
Prairie Dogs can be between individuals and groups
that often share many similar core values.
The book, Conservation of the Black-tailed Prairie
Dog: Saving North America’s Western Grasslands is
the brain child of Dr. John Hoogland who has re-
searched the behaviour of Prairie Dogs at Wind Cave
National Park in South Dakota since the mid-1970s.
Dr. Hoogland felt that there was a need to summarize
the extensive scientific literature on the Black-tailed
Prairie Dog in a non-technical format designed to high-
light the information’s relevance to conservation. To
accomplish this Dr. Hoogland engaged 30 other spe-
cialists and challenged them to draw out the lessons
for conservation from their areas of specialty. These
lessons are explored through 18 chapters on topics such
THE CANADIAN FIELD-NATURALIST
Vol. 120
it is to get any photographs of such little and active
sprites, let alone artistic ones of birds singing or dis-
playing. Handbook of the Birds of the World has now
reached biblical reference status.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
Saving North America’s Western Grasslands
as natural history, social behaviour, competition with
livestock, keystone species, human attitudes, and how
to establish new colonies of Prairie Dogs. The chapter
authors are all respected specialists and do an excel-
lent job of relating their topic area to the central theme
of the book: the conservation of Black-tail Prairie
Dogs.
The only significant criticism I have is that Dr. Hoog-
land repeatedly states in Chapter 2 that Black-tailed
Prairie Dogs do not hibernate. This is odd because pub-
lished studies have shown that Black-tailed Prairie Dogs
use facultative torpor (Lehmer et al. 2001). In fact, in
southwestern Saskatchewan, at the northern edge of
their distribution, Black-tailed Prairie Dogs spend as
much as 95 days a year in winter torpor bouts lasting
7.6 to 13.6 days with minimum core body temperatures
ranging from 7.1 to 11.6°C (Gummer 2005). Clearly,
this behaviour deserved to be recognized and discussed
as to how it might relate to efforts to conserve this spe-
cies. Despite this small failing, I fully recommend this
book to anyone interested in the conservation of this
fascinating social rodent or who wants to become better
informed on conservation issues on the Great Plains.
Literature Cited
Gummer, D. L. 2005. Geographic variation in torpor patterns: The
northernmost populations of prairie dogs and kangaroo rats. Ph.D.
dissertation, University of Alberta. 210 pages.
Lehmer, E. M., B. Van Horne, B. Kulbartz, and G. L. Florant.
2001. Facultative torpor in free-ranging black-tailed prairie dogs
(Cynomys ludovicianus). Journal of Mammology, 82(2): 551-557.
Pat FARGEY
Grasslands National Park, PO Box 150, Val Marie, Saskat-
chewan SON 2T0 Canada
Migrating Raptors of the World: Their Ecology and Conservation
By Keith L. Bildstein. Cornell University Press, Sage House,
512 East State Street, Ithaca, New York 14850 USA. 332
pages. U.S.$35.00 Cloth.
Raptors and bird migration both hold great fascina-
tion throughout the world, so the combination of the
two topics makes for an irresistible book. Bildstein is
a foremost world authority. Based at Hawk Mountain,
the world’s first site dedicated to the conservation of
hawks and the watching of their migration, he is the
Sarkis Acopian Director of Conservation Science.
Bildstein tells us that “a bad day at Hawk Mountain
Sanctuary is better than a good day anywhere else.”
His wide knowledge derives in part from his travels
to each of the world’s main hawk-watching sites. Fig-
ure | displays the five major hawk pathways through-
out the world and Figure 7 shows the 12 locations where
100 000 or more raptors pass by. In addition to stud-
ies at Hawk Mountain, banding of thousands of raptors
has occurred at Cedar Grove, Wisconsin; Cape May,
New Jersey; Hawk Ridge on Lake Superior, Minneso-
ta; the Goshute Mountains, Nevada; the Manzano
Mountains, New Mexico, and Golden Gate Observa-
tory, California. Notable Old World banding sites are
Chokpak Pass, Kazakhistan, and Elat, Israel. Switching
2006
to birds counted overhead, 851 600 Honey Buzzards
flew over Elat in the spring of 1985; there the spring
flights are much larger than in fall. Elsewhere large
numbers are counted in the fall: an average of 730 O00
at Corpus Christi, Texas; 5 200 000 near Cardel in the
state of Veracruz, Mexico; | 950 000 at Kekéldi Indi-
genous Reserve, Costa Rica, and 380 000 over the
Strait of Gibraltar at the western end of the Mediter-
ranean. In Veracruz, sometimes dubbed the “river of
raptors,’ 2 677 355 Turkey Vultures were counted in
2003, 2 389 323 Broad-winged Hawks in 2002 and
1 197 850 Swainson’s Hawks in 2003, including
782 653 in one day, 17 October that year.
Recent advances in technology have provided a big
boost to raptor migration studies. Application of satel-
lite radios to 117 Ospreys and 51 Golden Eagles added
immeasurably to our understanding of the speed and
variability of migration. Nine satellite-tracked Honey
Buzzards averaged 270 km/day across the Sahara
Desert, with brief stopovers suggesting that most fast-
ed during at least this leg of their trip south. Since rap-
tors often reduce energy requirements by soaring in
thermals, Bildstein has included a map and three dia-
grams of the major deflection/updraft corridors. Rap-
tors are able to orient and navigate simultaneously (set-
ting a direction and then achieving it), using internal
magnetic cues, visible landmarks, the sun by day, the
stars for the few species that migrate by night. Despite
a great deal of research, their navigational prowess still
exceeds human comprehension. In at least nine species,
including harriers, accipiters and two falcons, the juve-
niles precede the adults in southward migration, yet
come spring the adults often head north first. In the
Osprey and a few other species, juveniles spend an extra
year in the tropical wintering habitat before returning
north at two years of age.
A positive feature is that a number of hawk watch-
ing stations, including Hawk Mountain, were found-
Land Snails of British Columbia
By Robert G. Forsyth. 2004. Royal British Columbia Museum,
Victoria, British Columbia, Canada. 188 pages. $35. Paper.
The study of snails has certain advantages for a nat-
uralist. For one thing, unlike birds or butterflies, they
move slowly enough to be identified! However, anyone
who has ever tried to identify snails will know that
there is a dearth of readily accessible information out
there to help with the task. This is a lack that Robert
Forsyth has set out to remedy with his guidebook, pro-
viding help with the identification of 92 species of
land snails and slugs found in British Columbia. This
book is similar in layout and format to other recent
guidebooks from the Royal British Columbia Museum,
a handy 5.5” x 8.5” soft cover book, easy to slip into
a backpack or daysack.
Each taxon is provided with a “species account” that
consists of a description of the animal, its distribution,
Book REVIEWS
383
ed as conservation measures. Incensed by the car-
nage along the Kittatinny Ridge at Hawk Mountain,
Rosalie Edge purchased the property and hired Mau-
rice Broun to patrol it and count the raptors. Rachel
Carson, in Silent Spring in 1962, used the 25-year
Hawk Mountain counts, especially of declining Bald
Eagles, to support her arguments concerning the im-
pact of organochlorine pesticides, especially DDT, on
the reproductive success of birds of prey.
Bildstein’s final chapter is perhaps the strongest. It
explains the widespread effects of bounties, not com-
pletely eliminated in Pennsylvania until 1969. From
1917 to 1952, Alaska territory paid bounties on 128 000
Bald Eagles. Today, the main threats to raptors are habi-
tat loss and environmental contaminants, an example
being a painkiller, diclofenac, fed to cattle in India, that
has led to population declines of over 90 per cent for
three species of vulture that eat dead cows.
My only criticisms are small ones. Because owls are
also raptors, the absence of any mention of their no-
madic movements, cyclical irruptions and migrations
is somewhat contrary to the title of this book that is
restricted to diurnal raptors. Readers of the main text
might believe that DDT has long been banned every-
where, but in fact it is still being used in tropical coun-
tries to combat malaria and yellow fever. Only in the
Glossary does Bildstein inform his readers that DDT
is “still used by public health organizations to control
malarial mosquitoes.”
The writing is clear. Each chapter (I have only
touched on a few highlights) ends with a synthesis and
summary of the main points. This book is a treasure.
It belongs in every high school and college library
and in the library of every raptor enthusiast.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N OJ8
Canada
and its natural history (basically its habitat preferences).
Each account also includes information on the etymol-
ogy of the animal’s name, some remarks, usually deal-
ing with taxonomic issues or species with which it may
be confused, and references. The species are arranged
in taxonomic order. These accounts form the bulk of
the book (128 pages). Most taxa are illustrated by line
drawings or black-and-white photographs. For the pla-
nispiral snails, the images generally include the upper
(apical) surface, lower (umbilical) surface, and apertur-
al view. For the conispiral snails, the images generally
consist of an apertural view and a distal view. There
are also 33 colour images, of which 23 are of slugs.
These colour images are generally much crisper and
more useful than the black and white photos. Forsyth
includes some brief discussion of eleven other taxa that
he considers of doubtful occurrence in British Colum-
384
bia or that he judges to have equivocal taxonomic sta-
tus. These are often species that were perhaps only
recorded from one locality or for which the record was
made many decades ago and has not been confirmed
or supported by subsequent observations.
These species accounts highlight some interesting
points about the land molluscs of British Columbia,
as they are known at present. Of the species discussed,
about one third (24) are slugs. Of these, more than half
of them (13) are introduced. Many of these slug species
are described as being common in gardens, which per-
haps suggests that eagle-eyed gardeners may be respon-
sible for drawing attention to them. Forsyth notes that
many of these exotic species are significant agricul-
tural pests, in contrast to members of the native fauna,
which are not. He points out (page 19) that most may
have been accidental introductions, brought in on
plants or associated soil. Most of the snail species, in
contrast, are native (53) with less than a quarter (15)
identified as introduced. This striking pattern raises
the question as to whether it is a consequence of the
method of introduction (that is, on horticultural or
agricultural plants) or simply a lower level of investi-
gation of the mollusc fauna outside urban and inten-
sively farmed areas. Despite its focus on British
Columbia, many native species described by Forsyth
have wider distributions. Some are found in other areas
in the Pacific Northwest, from Oregon to Alaska. Oth-
ers are more widespread and some occur east of the
continental divide in other areas of western North
America. Despite its focus on British Columbia, there-
fore, this book has wider applicability in western Can-
ada.
The remainder of the book offers useful supplemen-
tal information. An extensive introduction (19 pages)
describes the biology of the animals, including, for the
snails, some discussion of shell characteristics, which
are important for species identification. There is also
a checklist of the species described in the volume,
and two dichotomous keys, one for snails and one for
slugs, to help narrow down the selection to genus. Ten
other keys are interspersed among the species accounts
and focus on genera, such as Vallonia, Vertigo, and
Arion, in which there are more than a couple of species.
All the snails described are dextral coiling species, as
are most land snails. Following the species accounts,
there is a lengthy (14 pages) reference list, a glossary
of terms, and a species index.
I obtained this book primarily to help with the iden-
tification of mollusc shell remains recovered from fine-
fraction analysis of sediment samples from archaeolog-
ical sites and other postglacial depositional contexts.
THE CANADIAN FIELD-NATURALIST
Vol. 120
In these situations, the soft parts of the animals are not
preserved and only the shells remain. Hence, identifi-
cations rely primarily on the morphology and struc-
ture of the shells. So I was particularly interested in the
shell descriptions. In this regard, the book has been very
helpful although I have noticed some limitations. My
main complaint is that the line drawings are not repro-
duced well. This is a significant problem. On many
drawings, the lines are faint and details of shell sur-
face features and ornamentation are extremely diffi-
cult to discern. Moreover, Forsyth only provides one
shell measure for each taxon. For the planispiral mol-
luscs, he provides the width of shells, but not the spire
height. For the conispiral molluscs, he provides the
height of the shells, but not the width. The complemen-
tary measures can be approximated from the drawings,
but it would have been useful to have them included
in the text. The width/spire height ratio is an easily
acquired descriptive character that helps discriminate
different taxa.
Two additional sets of information would have made
this book even more helpful. First, I thought each
species could have been illustrated by a “dot map”
showing the localities in British Columbia where it
has been collected or observed. I imagine that the dots
would have been very sparse for many species! Nev-
ertheless, such maps could highlight areas that may
well repay survey and collecting effort. Second, it
would have been useful to have a list of the accession
numbers for the specimens that were studied to com-
pile this volume. This is important so that in future
scholars could re-examine the specimens to confirm
and refine the taxonomy or acquire additional meas-
urements and images. Because the Royal British Col-
umbia Museum is the publisher, I assume that it may
be the institution housing these collections. However,
this information (that is, the institutional repository
and accession numbers) could have easily been in-
cluded, for example, in the species checklist, making
such detective work unnecessary.
Compiling this volume obviously involved a formi-
dable amount of painstaking work and study. I heartily
commend Forsyth’s commitment to a neglected but
rewarding faunal group. This book is a fine introduc-
tion to these animals and deserves to be on all west-
ern Canadian naturalists’ bookshelves. Anyone who
pays attention to the information in this volume will
never look at a snail on a rockery or a slug on a cab-
bage leaf in quite the same way again!
ALWYNNE B. BEAUDOIN
Royal Alberta Museum, Edmonton, Alberta TSN 0M6 Canada
2006
Book REVIEWS
385
Songbird Journeys: Four Seasons in the Lives of Migratory Birds
By Miyoko Chu. 2006, Walker & Company, 104 Fifth Avenue,
New York, New York 10011 USA. 312 pages. U.S. $23.
Cloth,
This is a welcome, attractive, and highly recom-
mended book. Each chapter is preceded by a well-
chosen quotation, often an appropriate full-page poem.
Miyoko Chu has combined the best of investigative
journalism with a passionate account of bird migra-
tion that holds the reader’s attention. Four chapters
deal with spring migration and two with fall migration.
She also must deal with where these migrating birds
summer and winter, so five chapters deal with these
intervening seasons. Not only does she tell of the lat-
est research into bird migration, but she interviews at
length some of the leading authorities. Thus, we read
fascinating stories about exceptional people who are
studying interesting birds. I will mention a few high-
lights.
Her opening chapter deals with a long-unsolved
question: do large numbers of birds make the seem-
ingly impossible long-distance flight over the Gulf of
Mexico, as George Lowery fervently believed, or was
this a preposterous fiction of people’s imaginations,
as George Williams posited? Lowery began by stud-
ies through a telescope, and admitted that trans-Gulf
migration was “fraught with enigmas and strained one’s
credulity.” Williams accused Lowery of jumping from
minute observations to sweeping conclusions, of extrap-
olating from the 12 birds he had seen through the tel-
escope to claim a migration of 21 million birds. Sid
Gauthreaux, when 17 years old in Louisiana, inveigled
his way into one of the first weather radar stations and
persuaded the meteorologists to let him look at the
fuzzy dots that he thought must be movements of birds.
He then persuaded Lowery to accept him as a graduate
student, embarking on a master’s thesis and then a
doctoral dissertation that proved to doubtful ornithol-
ogists everywhere that radar was a superb modality
for studying many aspects of migration. In so doing,
he proved beyond doubt that Lowery was correct.
Frank Moore’s group at Johnsons Bayou, Louisiana,
studied weights and condition of migrants as they first
reached landfall, after crossing the Gulf. Almost half
the birds arrived with their fat reserves totally depleted.
Richard Graber, at Urbana, Illinois, was an incred-
ibly early pioneer, four full decades ahead of the pack,
in placing tiny radio transmitters on small birds. His
friend, Bill Cochran, made him a number of tiny, 3 g
transmitters, which he glued to the backs of thrushes
that weighed about 30 g each. He tuned his receiver
to the appropriate frequency and then followed the
signals by truck, and once by aeroplane for 400 miles
in eight hours. Cochran then took over the study and
with special equipment he logged 150 000 miles by
truck to follow individual birds for a total of 30 000
miles. He followed one Peregrine Falcon from Wiscon-
sin to Tampico, Mexico, a Sharp-shinned Hawk from
Cedar Grove, Wisconsin, to Huntsville, Alabama, and
a Common Nighthawk from Illinois to South Carolina.
Another hero 1s a bird, a Yellow Warbler, nicknamed
“Wally,” that carried band 1750-17109, and returned
for nine years in a row, to sing near the Long Point
Bird Observatory in Ontario. At Hubbard Brook in
New Hampshire, where Richard Holmes and his col-
leagues have studied birds since 1969, bird numbers
have declined by about sixty percent. There, Black-
throated Blue Warblers produce more young in years
when caterpillars are common and in La Nifia years.
When Olin Sewall Pettingill was a boy in Maine, his
mother showed him a Bobolink nest. But they faced
hazards. About 720 000 Bobolinks were shipped as
game from one place in South Carolina in 1912 alone.
In Argentina, Pettingill learned in 1978, Bobolinks
were now being shot as pests for eating rice crops. It
is not surprising that the number of breeding bobolinks
has dropped by 75 percent in 25 years.
A new branch of science, studying the calls of noc-
turnal migrants to recognize individual species, was
pioneered particularly by Bill Evans in 1985, who pro-
duced a cassette tape of the night calls of the thrush-
es. His compilations were joined by those of Michael
O’Brien in 1991. Four years later they had “nailed
down” the final two species that had eluded them, the
night calls of Canada and Wilson’s Warblers. By 2001,
they produced the landmark guide that identified the
flight calls of 211 species of migrating landbirds in
eastern North America. Collaboration with researchers
and software programmers at the Cornell Laboratory of
Ornithology allowed analysis of different bird species
from their individual characteristics. There are now
thirty rooftop stations monitoring nocturnal migra-
tion with these new methods. Although Chu fails to
list the Evans/O’Brien CD in her references, it became
available in 2002 and can be ordered through http:
oldbird.org.
Only in 1995 was Bicknell’s Thrush recognized by
the American Ornithologists’ Union as a separate
species from the similar Gray-cheeked Thrush. Chris
Rimmer studied this thrush, one of the rarest migratory
birds in North America, in its breeding habitat among
the stunted firs near the top of Mount Mansfield in
Vermont. But where did they winter? Rimmer’s crew
found them in the Sierra de Bahoruco in the Dominican
Republic and, miracle of miracles, in 1995 captured in
a mistnet one of the banded birds they had tagged in
Vermont the previous summer — and then recaptured
the same bird again in Vermont during the next two
summers!
The reader is also given detailed directions to North
American birding migration hotspots, in both spring
and fall, and mid-winter hotspots in Yucatan, Belize.
Costa Rica, and Panama. She provides instructions for
amateurs to help contribute to knowledge, by partici-
386
pating in Nest Record Card programs and Project Feed-
erWatch.
Chu concludes by calling attention to the many haz-
ards facing bird populations, such as loss of habitat, in-
creasing parasitism by cowbirds, deaths from striking
windows, communication towers, pesticides, carbon
dioxide emissions, and acid rain. Twenty-nine species
BOTANY
Wildflowers of the Rocky Mountains
By G. Scotter and H. Flygare. Whitecap Books, 351 Lynn
Avenue, North Vancouver, British Columbia V7J 2C4
Canada. 255 pages. $ 29.95 Paper.
Plants of Alberta
By F. Rover and R. Dickinson. 2007. Lone Pine Publishing.
10145 — 81 Avenue, Edmonton, Alberta T6E 1W9 Cana-
da. 527 pages. $ 29.95 Paper.
It is most interesting that these two books have been
published within such a short space of time. They were
of immediate interest to me as I visit family in Alberta
and usually manage a trip or two to the Rockies or the
prairies. While they cover very different areas they
include many of the same species. Furthermore, the
authors have taken significantly different approaches.
Wildflowers of the Rocky Mountains [Wildflowers]
contains 350 species of selected plants in six sections
by flower colour [white, yellow and cream, green, pink,
red, orange, and brown, purple and blue.] Each page
depicts typically two species, all with a description, a
photograph and a range map. The authors say they
have chosen the most common species (typically the
pretty species) likely to be encountered during visits to
the Rockies, plus a few rarities. The area covered is the
Rocky Mountain zone from northern British Columbia
to New Mexico. This is a complete revision and expan-
sion of a 1986 guide, that covered 228 species. The authors
state that over the years they have learned to use non-
technical language. They have been successful in apply-
ing this talent and has produced a very readable book.
I have a few problems with Wildflowers. I have a dif-
ferent view on the choice of colours. For example, I
consider Mountain Sorrel to be reddish-brown not pink
as the authors contend. Similarly, I think of Flame-
coloured Lousewort (Pedicularis flammea) as red, but
the authors have placed this plant in the cream section
(because the flowers have a cream base). To further add
to my confusion this plant is called Oeder’s Lousewort
(P. oederi) but P. flammea is given as the synonym.
(However a web search of P. oederi showed images
of cream-only flowers, while an image search for P.
flammae showed cream-based flowers with significant
amounts of red on the tips. A search for synonyms of
P. oederi did not include P. flammae.) The alternative
names are not included in the index. Many of the Eng-
lish, and some of the scientific names are not the ones
I am used to. As an example, this book’s Four-part
THE CANADIAN FIELD-NATURALIST
Vol. 120
have declined by more than fifty percent in the last
40 years. Despite these losses, the arrival of songbirds
every spring remains a cause for celebration.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N O0J8
Canada
Dwarf Gentian or Felwort is called Alpine Gentian in
other texts.
The range maps are coloured in for the entire state
or province regardless of how widespread the plant is.
So Bear Grass is shown as occurring in Alberta, where-
as it can only be found in the vicinity of Waterton Lakes
National Park. This greatly reduces the usefulness of
these maps.
I thought the approach used by Wildflowers — arrang-
ing the species by flower colour — was useful only for
beginners. After a couple of years most people know
the major families, like vetch, violet, saxifrage and so
forth and I thought this is the way people progressed.
Recently I was told that women prefer books organ-
ized by flower colour and I found my wife agreed.
The photographs are very good and will be useful in
identifying blooms in the field. The descriptions in
Wildflowers are written in prose, often in a poetic style
that makes for pleasant reading. There is an excellent
introduction on the eco-zones and a wonderfully clear
glossary.
The second book is Plants of Alberta (Plants) which
covers over 1500 native plants that can be found in
Alberta. It is arranged by family and includes aquatic
plants, grasses, ferns and trees (so not just the pretty
flowers). Certainly I prefer the Plants method of arrang-
ing by families so I can reference all the orchids, vio-
lets, louseworts, etc. together. It too has a description, a
photograph and a range map also shown two to a page.
There is a short introduction and a decent glossary.
Plants’ descriptions are cryptic and scientific. Com-
pare “While other saxifrages may dress in flowers of
white or yellow, Purple Mountain Saxifrage is garbed
with flowers of the richest rose-purple to royal purple”
(in Wildflowers) with the terse “Flowers: Purple (occa-
sionally pinkish or white.)” (in Plants)
The photos in Plants average about twice the size of
those in Wildflowers — a considerable benefit. This is
achieved by having no margins, giving 20% extra space.
Also the headers giving the plant names and family are
60% smaller [but less artistic!]. As the text is shorter
the text size is bigger — great advantage for older eyes.
The larger script comes at the loss of poetic text to terse
notes. So despite the book’s slightly smaller pager size
(10%), by the creative use of space the publishers have
created a more appealing looking book.
2006
The range maps cover Alberta and show the area
within the province where you can see the plant. This is
useful because it shows over 35% of Alberta’s species
are confined to the southern border with British Colum-
bia and the United States. As the maps are small (1 x
2 cm) they require some interpretation. | photographed
a Striped Coralroot in Bow Valley Provincial Park, so
I verified that the range map showed this plant at this
location. It does, but it required careful examination
and analysis due to the size. The other distributions
are equally valid.
There is also a key arranged by leaf structure and
flower colour. This uses thumbnail (4") photos of the
MISCELLANEAOUS
BooK REVIEWS
387
families. So Violas appear in the white, yellow and pur-
ple sections. Is this enough to satisfy the ladies?
Despite my clear preference for Plants as a book, |
would say that both of these books are good guides.
The photographs are excellent and the text, while very
different in style, gives solid information. Your choice
should be dictated by your intentions or location. Al-
ternatively, and this I suggest is the best idea, you could
buy both as both, are good value for money.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
Pilgrimage to Vallombrosa: From Vermont to Italy in the Footsteps of George Perkins Marsh
By John Elder. 2006. University of Virginia Press, Charlot-
tesville, Virginia, USA. xv + 282 pages. U.S. $29.95 Cloth.
This book is a story of pilgrimage, of memory, of
hope. It’s a beautiful book, a book of life, of family, of
nature and culture. John Elder, a professor of English
at Vermont’s Middlebury College, invites the reader on
a pilgrimage. It is a story of connections; from the sec-
ond and third growth sugar bush hills of Vermont to the
centuries-old, culturally prestigious forests of Vallom-
brosa in the Tuscany hills; from the sugar bush to the
olive grove; from Robert Frost to Dante.
Vermont-born George Perkins Marsh was Abra-
ham Lincoln’s diplomat to the new Kingdom of Italy
from 1861 until his death in 1882. Considered to be
America’s first environmentalist, Marsh was the author
of the celebrated Man and Nature (1864). Inspired by
his study of the rise and fall of Mediterranean cultures
and his extensive experience of the region, Marsh’s
manifesto focused on environmental decline as central
to the collapse of these once powerful cultures. Marsh
was the first to describe and document the drastic,
long-term impact of deforestation on civilization. His
Man and Nature stirred the American imagination as
America itself experienced the wave of deforestation
flowing from east to west across the American land-
scape.
A sabbatical year permitted Elder and his wife to set
out on a pilgrimage across France and Italy. Elder’s pur-
pose was to pursue the footsteps of Marsh who died in
Vallombrosa in 1882. A pilgrimage usually has a set
route, but novelty and surprise are to be expected. A
pilgrimage on foot always becomes a pilgrimage of the
heart and of the mind. Elder skillfully invites the read-
er on a pilgrimage of ideas, one that examines our place
in nature.
Elder divides his pilgrimage into three landscapes,
each connected to Marsh’s vision of conservation. First
is the journey to and within Tuscany, the crucible of
Marsh’s thinking in the city of Florence, the surround-
ing hill towns and the celebrated forests of Vallombrosa.
Second, is the literary landscape where Elder situates
Marsh within the lineage of William Wordsworth, Mat-
suo Basho and Robert Frost. The final section returns to
Vermont, to Marsh’s native Woodstock, the sugar bush
of the Elder family, and to the Vermont-wide conser-
vation initiatives.
Those who set out on a pilgrimage usually have a
destination in mind. But that is often the only sure thing
about a pilgrimage. It’s what happens in between that
counts and that often surprises. Nothing is hard and fast
for Elder, nothing black-or-white. He writes skillfully
of what he terms the “middle landscape,” that ecotone
where wilderness and civilization meet. It is upon that
middle ground that Elder chooses to focus, weaving
the theme of “stewardship” throughout his work.
The term “stewardship” has fallen on hard times in
certain intellectual circles. Its anthropocentric conno-
tations are too aristocratic, too old-fashioned for some.
Not so for Elder, who calls for a mutuality of human
history and natural history. He calls for preservation
of not only biodiversity but also human values and
community, a place where technology and religious
faith both have their own special role to play. For Elder,
the environmental crisis is ultimately a crisis of culture,
a crisis of the human soul, a crisis of the human heart.
Stewardship, brought into proper dialogue with con-
temporary ecological, feminist, religious and social
justice thought, can engender the cultural and moral
resources necessary to address the environmental needs
of our time.
Elder’s attempt to re-frame the discussion is wel-
come. Too often has environmental conversation frac-
tured into dualistic dichotomies, into opposing camps
of nature and culture. We need both voices to forge a
social ecology, where ecosystems are not considered
in the absence of culture and vice-versa. It’s that mid-
dle ground that provides the vital connection between
388
remote regions of wilderness and the places were we
live, work and play.
We must thank Elder for eschewing sectarian lan-
guage. His goal takes much more effort. It seeks a com-
mon ground that does not reduce to the lowest common
denominator. The built environment, the cultivated envi-
ronment, the wilderness environment; they all have a
voice that needs to be heard. Elder speaks in the tradi-
tion of Aldo Leopold, René Dubos and Wendell Berry,
voices that seek a dialogue between culture and the
wild. For all three, the steward is the facilitator of such
a needed task. The steward lives in that “boundary zone
where the wilderness ethic may engage with recent dev-
elopments of environmental history, and where the ideal
of preservation transcending our narrow utilitarianism
may engage with the tradition of stewardship.” (page
218)
This is marvellous meditation. It’s no wonder, since
Elder’s trade and tackle are words and stories. He
weaves a story, a meditation, a contemplation that opens
up our mind and heart to new possibilities for our rela-
tionship to forests, to those wild and wonderful places.
It is a story of people tending the olive groves under
the warm Tuscany sun, and of his own family’s stew-
arding of the sugar maple groves of their Vermont
Where to Watch Birds — World Cities
By Paul Milne. 2007. Yale University Press P.O. Box
209040, New Haven, Connecticut 06520-9040 USA. 496
pages. U.S. 28.00 Paper.
This type of book is a reviewer’s dream. Before you
open it you know the author, Paul Milne, must make
choices and these will not be the same as your choic-
es. This means you know you will have the opportu-
nity to criticize. So how well do his choices match
those I would have made? First he has included only
one Canadian city, Toronto. It is Canada’s largest city
but it is not the capital nor the best for the naturalist.
One out of the 60 cities covered in the book seems a
little thin for second largest country in the world [even
if we rank 36" for population]. Putting my obvious bias
aside, I noted that other key cities are missing. Oslo,
with its great royal park, and Kiev, a city of wonderful
chestnut trees, are two evident examples. Indeed 60%
of the cities are in Europe and Asia; a disproportionate
number for these two continents over North and South
America and Australia.
Accepting the author’s choices, how useful is this
book? First this book covers more than the city itself.
When I visit Victoria, British Columbia, I get up at
dawn and walk from my downtown hotel through Bea-
con Hill park to the coast. I return before breakfast and
then go to my meeting. These two hour jaunts have net-
ted me some very interesting birds [e.g. Ancient Mur-
relet]. These activities were all within the downtown
core of Victoria and did not involve any travel. The
author includes in his version of “city” many areas that
THE CANADIAN FIELD-NATURALIST
Vol. 120
home. Landscape and culture provide the nexus of dis-
cussion. Forgetting one or the other dehumanizes us
and sets the stage for either pillage or misanthropism.
In this era of apocalyptic predictions, environmen-
tal and social haemorrhaging, a widening gap between
culture and nature, between rich and poor, urban and
rural, Elder sets out a path of reconciliation, a path that
seeks the common good. It’s a path laced with com-
munity, dialogue, a sense of the sacred, good work, jus-
tice, stewardship, care and respect. These virtues don’t
come easily.
A final word to Elder: “I have come to believe that
without the stories that integrate the face of nature with
the drama of our human lives, society will not have the
power to restrain our appetite and respect the larger
balance of nature.” (page 67) If you accept this conclu-
sion, read this book. It will draw you into a world of
memory and elegance. If you don’t agree with Elder,
read this book anyway. The full, inclusive fare offered
by Elder will call you into a respectful dialogue.
JOHN MCCARTHY
Ecology Project, Ignatius Jesuit Centre of Guelph, P.O. Box
1238, Guelph, Ontario N1H 6N6 Canada
require significant travel — you will need a car. The
equivalent of me driving to Goldstream Park, 17 kilo-
metres from downtown Victoria, and more. Indeed
some sites are over 100 km away (distances are given
in the measurement used in the country; e.g., they are
in miles in the U.S. and kilometres in Canada.)
Accepting the author’s broad boundaries how useful
is this book? I would say very useful indeed. I have
birded in almost half of the cities mentioned [gener-
ally without a car] and have visited many of the sites he
describes. He has an introductory section that explains
the city in birder’s terms and has a good section on trans-
portation. Where there is a good public transportation to
suitable birding locations (e.g., London, Moscow) the
author provides enough information to get the traveller
started. The site descriptions are clear and accurate and
I wish I had owned this book years ago. There are loca-
tor maps where they are appropriate. There is a list of
birds typically seen at each location. This worked well
for all places and I verified with my notes that I too had
seen a similar list of species.
I can quibble on some minor points. For example,
under Johannesburg he includes Suikerbosrand [upland
species — about 70 km from Johannesburg] but omits
the nearby Pilansburg — perhaps the best park to visit
[lowland species — about 100+ km from Johannes-
burg]. His information on travel omits to say it would
be almost suicidal for a tourist to drive in Mumbai.
Taxis are cheap and you can get a car and driver, sym-
pathetic to your needs, from the Bombay Natural His-
2006
tory Society for reasonable cost. If you take a photo
booth shot of yourself plus a glue stick to Austria, you
can get a multi-day pass to Vienna’s excellent public
transit; a most economical way to travel.
All in all this is a great and useful book. My next
trips are to Halifax and then Anchorage, but neither of
these is included by the author. However, I look for-
ward to using it on my next trip to one of the cities in
Marshes: The Disappearing Edens
By W. Burt. 2007. Yale University Press, P.O. Box 209040
New Haven, Connecticut 06520-9040 USA. 192 pages.
U.S. 35.00 Cloth.
William Burt has a very deserved reputation. His
photographs are amazing. As a naturalist who also takes
photos I have some idea of the skill and the patience
needed to get a good shot. Burt goes beyond good. Not
only are the photos amazing in their quality, but he spe-
cialises in birds that are typically difficult to see! Take
Black Rail as an example. I spent several hours at night
wading in a swamp with an ardent group of seekers
before I had my first glimpse of a Black Rail. It ran
over my foot. It took another hour to actually get my
binoculars [and flashlight] on one of the cute but elu-
sive beasts. The idea of a photograph never entered my
head. I do have some slides of other rail species I took
mostly by good luck. They are nice, but they are not
artistic and they all have little flaws. Burt’s photos are
technically crisp and clear. They are also artistic; the
kind you would frame and hang on the wall.
In addition to birds, the author has also included a
number of pictures of marsh vegetation and some of
the more picturesque flowers. Actually this book is
over 50% photographs.
Once you have finished drooling over the illustra-
tions you could read the text. The author describes his
visits to wetlands throughout North America. Start-
ing with his home base in Connecticut river marshes he
travels to Maryland (Elliot Island), Manitoba (Sewall
Lake), Saskatchewan (Crane Lake), Oregon (Malheur)
and California (Klamath). He also takes us on a trip
Book REVIEWS
389
this book, which will likely be Moscow. | would encour-
age Paul Milne to travel more in North and South Amer-
ica. Then I can hope he plans a second volume to cover
some of the cities he has missed.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
through wetlands in Texas, Louisiana, Florida and Vir-
ginia. At each site he describes the value that each place
brings and some of the issues it now faces.
Burt make an emotional, almost poetic appeal for
marshes. He describes his favourite haunts along the
east coast of the United States with such verve that I
realised, somewhat for the first, that I too had experi-
enced the same feelings. I tend to look at life more clin-
ically, but Burt is more passionate and has shown that
I have similar emotions to him below the surface.
I believe you should learn something new every day.
I began to learn as soon as | started to read. For the
first time I properly understand the issue with Phrag-
mites. The ones I see are more likely the aggressive —
and therefore dangerous — European plant and not the
look-alike native version. Alien invaders are a key threat
and Phragmites and loosestrife lead that charge.
His description of the work of the photographers
Walter Finley and Herman Bohlman in the 1800s is
enlightening on the persistence and dedication of these
pioneers plus their resounding contribution to conser-
vation.
This a lovely book and would make a wonderful
present for both naturalists and non-naturalists. The
beautiful writing style and powerful messages might
even convert some folks to be conservationists.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario,
K1J 6K5 Canada
390
NEw TITLES
Prepared by Roy John
+ Available for review * Assigned
ZOOLOGY
Albatrosses, Petrels and Shearwaters of the world. By D.
Onley and P. Scofield. 2007. Yale University Press, P.O.
Box 209040, New Haven, Connecticutt 06520-9040 [North
America] or Christopher Helm Publishers Ltd./A&C Black
Publishers Ltd., 38 Soho Square, London W1D 3HB [Europe
and the Commonwealth]. 256 pages. U.S. $29.95.
Catalogue of Aleocharine Rove Beetles of Canada and
Alaska (Coleoptera, Staphylinidae, Aleocharine). By N.
Gouix and J. Klimaszewski. Pensoft Publishers, Geo Milev
Street 13a, 1111 Sofia, Bulgaria. 166 pages. EURO 60 hard-
back.
Atlas of Bird Migration. Edited by J. Elphick. 2007. Firefly
Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1 Canada. 176 pages. $35.
An Enchantment of Birds. By R. Cannings. 2007. Greystone
Books, #201 — 2323 Quebec Street, Vancouver, British Colum-
bia VST 487 Canada. 211 pages. $29.95.
Aves amenazadas de Espana. By J. Varela. [In Spanish].
2007. Lynx Edicions, Passeig de Gracia, 12, 4rt. 2a, 08007
Barcelona, Spain. 325 pages. U.S. $38.40.
The Lapwing. By Michael Shrubb. 2007. T & AD Poyser
(A&C Black Publishers Ltd, 38 Soho Square, London W1D
3HB). 320 pages. $100 (approximately).
Lista Comentada de las Aves de Guatamala — Annotated
Checklist of Birds of Guatamala [bilingual English Span-
ish]. By K. Eiserman and C. Avendano. Lynx Edicions, Passeig
de Gracia, 12, 4rt. 2a, 08007 Barcelona, Spain. U.S. $15.60
* The Birder’s Companion. By Stephen Moss. 2007. Firefly
Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1
Canada. 239 pages.
All the Birds of Brazil — An Identification Guide. Second
Edition. By D. De Souza. 2006. Editora Dall/Subbuteo Nat-
ural History Books, The Rea, Upton Magna, Shrewsbury SY4
4UR UK. 325 pages. U.S.$60.
500 Butterflies. By K. Preston-Mafham. 2007. Firefly Books
Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1
Canada. 528 pages. $29.95 Cloth.
Ecology and Behaviour of Chickadees and Titmice. Edit-
ed by K. Otter. 2007. Oxford University Press, 198 Madison
Avenue, New York, New York 10016 USA. 352 pages. U.S.
$102. Cloth.
*Deep Alberta — Fossil Facts and Dinosaur Digs. By John
Acorn. 2007. University of Alberta Press, Ring House 2,
Edmonton, Alberta T6G 2E1 Canada. 186 pages. $26.95 Paper.
* Ladybugs of Alberta. By J. Acorn. 2007. University of
Alberta Press Ring House 2, Edmonton, Alberta T6G 2E1
Canada. $ 29.95
Lost Land of the Dodo. By A. Cheke and J. Hume. 2007.
Christopher Helm Publishers Ltd./A&C Black Publishers
Ltd., 38 Soho Square, London W1D 3HB. 504 pages. U.S.
$84.
THE CANADIAN FIELD-NATURALIST
Vol. 120
300 Frogs. By C. Mattison. 2007. Firefly Books Ltd., 66 Leek
Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 528
pages. $29.95 Cloth.
Pocket Guide to the Insects of Britain and Western Europe.
By M Chinery. 2007. Christopher Helm Publishers Ltd./
A&C Black Publishers Ltd, 38 Soho Square, London W1D
3HB UK. 320 pages. U.S. $26.20.
The Red Kite. By I. Carter. 2007. Arlequin Press, The Rea,
Upton Magna, Shrewsbury SY4 4UR U.K. 200 pages. U.S.
$44. Cloth.
The Lepidoptera of Israel. Volume II: Noctuidae. By V.
D. Kravchenko, M. Fibiger, A. Hausmann and G. C. Muel-
ler. 2007. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia,
Bulgaria. In English, 320 pages. EURO 110. Hardback.
Where to Watch Mammals in Britain and Ireland. By R.
Moores. 2007. Christopher Helm Publishers Ltd /A&C Black
Publishers Ltd, 38 Soho Square, London W1ID 3HB U.K.
304 pages. U.S. $29.60. Paper.
Manipulative Monkeys — the Capuchins of Lomas Barbu-
dal. By S. Perry and J. Manson. 2007. Harvard University
Press, 79 Garden Street, Cambridge, Massachusetts 02138
USA. 326 pages. U.S. $45.
A Field Guide to New Zealand Birds. By B. Parkinson.
2007. New Holland Publishers (UK) Ltd., Garfield House, 86-
88 Edgware Road, London W2 2EA U.K. 136 pages. U'S.
$22.75.
Know Your New Zealand Birds. By L. Moon. 2007. New
Holland Publishers (UK) Ltd., Garfield House, 86-88 Edg-
ware Road, London W2 2EA U.K. 126 pages. U.S. $22.75.
Collins Field Guides — Birds of the Palearctic: Passerines.
By N. Arlott. 2007. Harper Collins Publishers Ltd., 2 Bloor
Street East, 20th Floor, Toronto, Ontario M4W 1A8 Canada.
384 pages. U.S $46.
Photographic Guide: Birds of Peru. By C. Byers. 2007. New
Holland Publishers (UK) Ltd., Garfield House, 86-88 Edgware
Road, London W2 2EA U.K. 144 pages. U.S. $15.95.
The Sand Wasps — Natural History and Behavior. By H.
Evans and K. O’ Neill. 2007. Harvard University Press, 79 Gar-
den Street, Cambridge, Massachusetts 02138 USA. 340 pages.
+ At Sea Distribution and Abundance of Seabirds off
Southern California: A 20 Year Comparison. Studies in
Avian Biology Number 33. By J. Mason et al. Cooper Ornitho-
logical Society, c/o Western Foundation of Vertebrate Zoology,
439 Calle San Pablo, Camarillo, California 933012-8506 USA.
101 pages. U.S. $15.
Sharks of the Pacific Northwest. By A. De Maddalena, A.
Preti and T. Polansky. 2007. Harbour Publishing, P.O. Box
219, Madeira Park, British Columbia VON 2HO Canada.
160 pages. $21.95 Paper.
The Smaller Majority. (Insects) By P. Nasrecki. 2007. Har-
vard University Press, 79 Garden Street, Cambridge, Massa-
chusetts 02138 USA. 288 pages. U.S. $24.95.
2006
The Birds of Thai-Malay Peninsula — Volume 2. By D.
Wells. 2007. Christopher Helm Publishers Ltd./A&C Black
Publishers Ltd, 38 Soho Square, London WID 3HB U.K.
704 pages. U.S. $120.
Oceanic Wilderness. By Roger Steene. 2007. Firefly Books
Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1
Canada. 340 pages. $59.95 Cloth.
Shells. By P. Starosta and J. Senders. 2007. Firefly Books
Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1
Canada. 384 pages. $85. Cloth.
Whales and Dolphins of the North American Pacific. By
G. Cresswell, D. Walker and T. Pusser. 2007. Harbour Pub-
lishing, P.O. Box 219, Madeira Park, British Columbia VON
2HO Canada. 216 pages. $21.95 Paper.
BOTANY
* Plants of Alberta. By F. Rover and R. Dickinson. 2007.
Lone Pine Publishing. 10145 — 81 Avenue, Edmonton, Al-
berta T6E 1W9 Canada. 527 pages. $ 29.95. Paper.
Comparative Plant Ecology. J. Grime, J. Hodgson and R.
Hunt. 2007. Springer Science+Business Media Deutschland
GmbH, Heidelberger Platz 3, 14197 Berlin, Germany. 752
pages. $168. Cloth.
Pocket Guide to the Trees of Britain and Northern Europe.
By A Mitchel. 2007. Christopher Helm Publishers Ltd. /A&C
Black Publishers Ltd, 38 Soho Square, London W1D 3HB
U.K. 288 pages. U.S. $26.20
* Wild Flowers of the Rocky Mountains. By G. Scotter
and H. Flygar. 2007. Whitecap Books Ltd, 351 Lynn Avenue,
North Vancouver. British Columbia V7J 2C4 Canada. 255
pages. Illustrations $29.95.
OTHER
Digital Wildlife Photography. By D. Tipling. 2007. Firefly
Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1 Canada. 160 pages. $24.95 paper.
Drawing and Painting Plants. By Christina Brodie. 2007.
Timber Press, 133 SW 2"™ Avenue, Ste. 450, Portland, Oregon,
USA. 144 pages. C$27.95. Paper.
Book REVIEWS
39]
Where to Watch Birds in Donana. By F. Chiclana and J
Garzon. 2006. Lynx Edicions, Passeig de Gracia, 12, 4rt. 2a,
08007 Barcelona, Spain. 160 pages. U.S. $28.
* Gilbert White. By Richard Mabey. 2007. University of Vir-
ginia Press, P.O. Box 400318, Charlottesville, Virginia 22904-
4318 USA. 230 pages.
Hispaniola. Edited by E. Fernandez. 2007. Harvard Univer-
sity Press, 79 Garden Street, Cambridge, Massachusetts 02138
USA. 400 pages. U.S. $60.
* Pioneering Conservation in Alaska. By Ken Ross. 2007.
University Press of Colorado, 5589 Arapahoe Avenue, Suite
206C, Boulder, Colorado 80303 USA. 540 pages. $34.95.
Cloth.
This Tender Place — The Story of a Wetland Year. By Lau-
rie Lawlor. 2007. The University of Wisconsin Press, 1930
Monroe Street, Third Floor, Madison, Wisconsin 53711-2059
USA. 166 pages. U.S. $19.95. Paper.
* Marshes: The Disappearing Eden. By W Burt. 2007. Yale
University Press P.O. Box 209040New Haven, Connecticut
06520-9040 USA. 192 pages. U.S. $35.
Oregon’s Dry Side — Exploring East of the Cascade Crest.
By Alan D. St. John. 2007. Timber Press, 133 SW 2° Avenue,
Ste. 450, Portland, Oregon 97204 USA . 324 pages. U.S.
$29.95.
The Ornitholologist’s Dictionary. By J. Erritzoe, K. Kampp,
K. Winker and C. Frith. 2007. Lynx Edicions, Montseny, 8,
08193 Bellaterra, Barcelona, Spain. U.S. $25. Paper.
* A Paddlers’ Guide to Quetico and Beyond. By K. Callan.
2007. Firefly Books, 132 Main Street, Erin, Ontario NOB 1TO
Canada. 192 pages. $ 24.95.
Intelligent Courage: Natural Resource Careers that Make
a Difference. By M. Fraideburg. 2007. Krieger Publishing
Company, P.O. Box 9542, Melbourne, Florida 32902-9542
USA. 220 pages. U.S. $32.50.
YOUNG NATURALISTS
* Gracie, the Public Gardens Duck. By Judith Meyrick.
2007. Nimbus Publishing, 3731 Mackintosh Street, Halifax,
Nova Scotia B3K SAS Canada. 32 pages. $16.95.
News and Comment
Marine Turtle Newsletter (113)
July 2006. 24 pages: EDITORIAL: New editors for
MTN — ARTICLES: Hawksbill and Olive Ridley nest-
ing on Masirah Island, Sultanate of Oman: an update
(A. F. Rees and S. L. Baker) — Preliminary data from
an increasing Olive Ridley population in Sergipe,
Brazil (J. C. Castilhos and M. Tiwari) — NOTES: First
report of Green and Kemp’s Ridley Turtle nesting on
Wassaw, Georgia, USA (K. L. Williams, M. G. Frick,
and J. B. Pfaller) — LETTERS — MEETING REPORTS
— MST UPDATE — ANNOUNCEMENTS — NEWS &
LEGAL BRIEFS — RECENT PUBLICATIONS.
The Marine Turtle Newsletter is edited by Brendan
J. Godley and Annette C. Broderick, Marine Turtle Re-
search Group, Centre for Ecology and Conservation,
University of Exeter in Cornwall, Tremough Campus,
Penryn TR10 9EZ United Kingdom; e-mail MTN @sea
turtle.org; Fax +44 1392 263700. Subscriptions and
donations towards the production of the MTN can be
made online at or postal
mail to Michael Coyne (online Editor) Marine Turtle
Newsletter, 1 Southampton Place, Durham, North Caro-
lina 27705 USA (e-mail: mcoyne @seaturtle.org).
The Boreal Dip Net/L’Epuisette Boréale January 2006
Newsletter of the Canadian Amphibian and Reptile
Conservation Network (CARCNET) 10(1). CON-
TENTS: Editor’s Note — Faces and Places/In the Bag
— Snake habitat manipulation — A request for infor-
mation — CARCNET field trip — Natural Heritage
Building of the Canadian Museum of Nature & Parc
de la Gatineau, QC — Presentations made at the 2005
annual meeting of CARCNET/RECCAR — Reintro-
duction of Leopard Frogs at Bummer’s Flats — North-
ern Leopard release marks fifth year — Lost Leopard?
— A significant range extension for the Northern
Alligator Lizard in British Columbia — Faces and
Places/Out of the bag.
Membership information can be obtained from
Bruce D. Pauli, Canadian Wildlife Service, National
Wildlife Research Centre, Carleton University, Raven
Road, Ottawa, Ontario K1A 0H3 Canada. Additional
information is at website http://www.carcnet.ca.
392
Chairperson:
Attendance:
Mike Murphy, President
1. Minutes of the Previous Meeting
Under Report of the Nominating Committee/Com-
mittee Chairs: The Chair of the Education and Publicity
Committee should be corrected to read Gillian Marston
not John Cameron.
It was moved by Gillian Marston seconded by Fenja
Brodo that the minutes be accepted.
(Motion Carried)
2. Business Arising from the Minutes
There was no business arising from the Minutes.
'3.Communications Relating to the Annual
| Business Meeting
There were no communications relating to the Annual
' Business Meeting.
4. Treasurer’s Report
Frank Pope reviewed the financial report for the year
ending 30 September 2005, noting that the Club’s net
) assets had increased by approximately $86,350. This
was due in large part to a bequest from the estate of
| Mildred Groh of $62,250. There was an amendment to
| the Report.
Moved by Frank Pope and seconded by Diane Lepage
that the Financial Report be accepted as amended.
(Motion Carried)
5. Committee Reports
_ Mike Murphy introduced each of the Committee
\reports and a representative of the appropriate Commit-
tee and asked for questions and comments. He thanked
_ the committee chairs and committee members for their
/work over the previous year.
Moved by David Hobden seconded by Gillian
Marston, that the reports be accepted with suggested
‘amendments.
| (Motion Carried)
at
'6. Nomination of the Auditor
Moved by Frank Pope, seconded by Bill Cody, that
Janet Gehr continue as Auditor for another year.
| (Motion Carried)
{
Minutes of the 127" Annual Business Meeting of
The Ottawa Field-Naturalists’ Club 10 January 2006
Place and time: Canadian Museum of Nature, Ottawa, Ontario, 7:30 p.m.
Forty-two persons attended the meeting.
Attendees spent the first half-hour reviewing the minutes of the previous meeting, the Treasurer’s report and the
Report of Council. The meeting was called to order at 7:40 p.m. with some opening remarks from the President.
7. Report of the Nominating Committee
President Mike Murphy
Vice President Ken Allison
Secretary Susan Laurie-Bourque
Treasurer Frank Pope
Past President
Business Manager
Gary McNulty
Bill Cody
Editor, CFN Francis Cook
Editor, T&L Karen McLachlan Hamilton
Committee Chairs
Birds Chris Traynor
Computers Dan Millar
Conservation Stan Rosenbaum
E&P Gillian Marston
E&L Fenja Brodo
Finance Ann MacKenzie
FWG David Hobden
Macoun rep. Diane Kitching
Membership Henry Steger
Publications Ron Bedford
ON Rep Eleanor Zurbrigg
Members at large
Susan Howell
Julia Cipriani
Justin Peter
Chairs not on Council
Awards Ernie Brodo
Macoun Rob Lee
Nominations Fenja Brodo
Retiring from the council: Kathy Conlan,
Louise Schwartz,
Dave Smythe
Ken Allison, Julia Cipriani,
Susan Howell, Ann
MacKenzie, Dan Millar
and Justin Peter.
Moved by Fenja Brodo, seconded by Frank Pope,
that the slate of nominations for the 2006 Council be
accepted.
New on the council:
(Motion Carried)
393
394
8. New Business
There was no new business.
9. Presentation of Awards for the 2005 OFNC
Photo Contest
There was a presentation of the awards to the win-
ners of the 2005 OFNC Photo Contest by Suzanne
Deschene and Gillian Marston. The grand prize winner
received a digital camera and photo printer.
Amphibians: Ruth Allison
Birds: Lois Naggs
Insects and spiders: Christine Hanrahan
Mammals and vegetation: Jim Robertson
Scenery: Stephen Darbyshire
Honorable mention —
Birds and vegetation:
Grand prize winner:
Gwen Williams
Dave Sankster
THE CANADIAN FIELD-NATURALIST
Vol. 120
The Education and Publicity committee was already
planning the next photo contest, tentatively titled “The
Tale of the Trail”. Details to be published in Trail &
Landscape.
The presentation of the awards was followed by a
slide show of the contest entries. This was viewed
with admiration and enjoyment by the members in
attendance.
10. Adjournment
Moved by Henry Steger/Ron Bedford that the meet-
ing be adjourned at 9:35 pm.
(Motion Carried)
SUSAN LAURIE-BOURQUE
Recording Secretary
The Ottawa Field-Naturalists’ Club Committee Reports for 2005
Awards Committee
At the club’s Annual Soirée, on 30 April 2005 at St. Basil’s
Church in Ottawa, awards were once again given to members,
and two non-members, who distinguished themselves by ac-
complishments in the field of natural history and conservation,
or by extraordinary activity within the club. There isn’t al-
ways a winner for every potential award, and this year, once
again, the Anne Hanes Natural History Award was not given.
On the other hand, a new award was approved by Council and
awarded for the first time this year: the Mary Stuart Educa-
tion Award. This new award has been established for members,
non-members or organizations in recognition of outstanding
achievements in the field of natural history education in the
Ottawa Region. The new Education Award was appropriately
named in honour of Mary Stuart, who was so passionately in-
terested in establishing a new generation of young naturalists.
The winners of the 2004 Awards were:
HONORARY MEMBER:
Charles D. Bird — a long-time member of the OFNC and
one of Canada’s most outstanding naturalists in the fields of
cryptogamic botany and entomology. He has been an associate
editor of The Canadian Field-Naturalist for over 25 years.
MEMBER OF THE YEAR:
Marilyn Ward — for her efforts in handling the club phone,
helping to make sure lecture and excursion events are suc-
cessful, and keeping the Fletcher Wildlife Garden spic and
span.
GEORGE MCGEE SERVICE AWARD:
Martha Camfield — for her many years of service with the
Macoun Field Club both at meetings and in the field, and for
her botanical work on the Leitrim wetlands.
Mary STUART EDUCATION AWARD:
Andrea Howard — for her extraordinary, innovative work
in teaching natural history at the Eastern Ontario Biodiversity
Museum in Kemptville and her “Museum in a Suitcase” school
programs.
CONSERVATION AWARD — MEMBER:
Ghislaine Rozon — for her extraordinary efforts in trying
to save the Larose Forest from destruction by developers.
CONSERVATION AWARD — NON-MEMBER:
Barbara Barr — for attending municipal and regional meet-
ings and hearings regularly, mostly representing the Green-
space Alliance, gathering information, presenting briefs, and
being a source of sage advice to policy makers on conservation
issues.
The full text of the citations for each of these awards was
read at the annual soirée and will be published in vol. 119,
no. 4 of The Canadian Field-Naturalist.
IRWIN BRODO
Chair, Awards Committee
Birds Committee
The Birds Committee participated with the Club des Orni-
thologues de |’Outaouais in both the Christmas Bird Count
and the Fall Bird Count. The compilation dinners for both
events were very well attended. The committee again orga-
nized the Peregrine Falcon Watch at the nest site downtown.
Donations to the watch in 2005 totalled $275. While both
chicks fledged, one disappeared and its fate remains unknown.
Sadly, the adult female, Horizon, was injured and subsequently
euthanized at the request of the Ministry of Natural Resources.
We continued to provide seasonal bird summaries for Trail &
Landscape. Several members have also enhanced our com-
mittee’s presence on the OFNC’s web site. The 5" and final
year of the Ontario Breeding Bird Atlas has been completed
and a report summary has been prepared. The Bird Record
Sub-committee met several times during the year to review
rare bird records. The committee also provided several leaders
for OFNC bird trips. The committee has also added a new
bird feeder, provided by the National Capital Commission,
near the new picnic shelter at the Mer Bleu. The 2005 seed-a-
thon raised over $300 in pledges. We continue to operate the
rare bird alert and Ottawa’s bird status line which provides
updates and information on both rare birds and regular report
summaries.
CHRIS TRAYNOR
Chair, Birds Committee
Computer Management Committee
During 2005, the Club purchased two new computers,
one for use by the Treasurer’s Assistant; old equipment has
been retired.
2006
The Computer Management Committee reviewed how
club volunteers manage key information holdings of the club,
to ensure that club data are secure and accessible. Key files
include the membership database, financial records, Trail &
Landscape electronic archives, and Canadian Field-Naturalist
files. The committee concluded that current data management
practices are adequate.
There were no requests to the Computer Management
Committee from other committees during the year for com-
puter support, as they had expertise available in-house.
The club upgraded the capabilities of its web site consid-
erably when it moved to a new service provider. These im-
provements are allowing the Web Master to continue to improve
the content of our popular web site.
ELEANOR ZURBRIGG
Chair, Computer Management Committee
Conservation Committee
Alfred Bog
Frank Pope continues to chair the Alfred Bog Committee,
while planning to hand over to a new person who has pro-
visionally accepted this position.
Planning Reform
In follow-up to our 2004 brief and letters to Ontario Min-
isters, additional letters were sent on 19 May 2005, including
a joint letter to the Premier and four Ministers, and individual
covering letters to each. Frank Pope and Eleanor Zurbrigg
presented our position on planning reform at the Ontario
Nature/Federation of Ontario Naturalists regional meeting
at Alfred College on 15 October. A meeting to discuss the
issues with the ON/FON regional Board Member was set for
6 December.
South Gloucester (5309 Bank Street)
We sent notification by registered letter to both the Minis-
try of Natural Resources and R. W. Tomlinson, stating our
continuing opposition to the granting of an aggregate extrac-
tion licence to which we had previously objected. We pointed
out that R. W. Tomlinson’s proposed mitigation measures for
damage to significant plant species were dismissed in short
work by Brunton Consulting Services in a second Peer
Review specially ordered by the City. The issue was presently
on hold pending hearings at the Ontario Municipal Board of
appeals by the Greenspace Alliance and others.
Pesticide Use on Lawns and Gardens
Email letters were sent to the Mayor of Ottawa and Coun-
cillors, supporting a campaign that is trying to get the pre-
sent (ineffective) policy of voluntary reduction in pesticide
use replaced by a ban on the cosmetic use of pesticides. City
Council rejected a ban, but the Mayor and some Councillors
in favour of a ban stated that they would make this an elec-
tion issue in 2006.
STAN ROSENBAUM
Chair, Conservation Committee
Education and Publicity Committee
Kiosks, DISPLAYS AND OTHER EVENTS
The committee enhanced the material available for displays
and kiosks by adding several light weight portable displays.
Committee members participated with displays at the Ottawa-
Carleton Elementary Teachers’ Federation Publishers’ Dis-
play, National Wildlife Week’s Wildlife Festival, Environment
Week Fair and Eco Fair at Friends of the Farm and the OFNC
Soirée. Once again the OFNC participated in the Ottawa Sci-
MINUTES OF THE 127™ ANNUAL BUSINESS MEETING
395
ence Fair sponsoring a prize for a project judged by Education
and Publicity Committee member Kathy Conlan. The com-
mittee also arranged speakers for outside groups.
OFNC MEemBeERrSHIP Drivt
The portable kiosks were used for approximately 10 out-
ings for a successful spring club membership drive
DIGITAL LIBRARY
The committee has initiated the OFNC’s ‘digital photo
library and catalogue’ as a successor to the slide collection that
the committee currently manages.
OFNC Nature PHOTOGRAPHY CONTEST
The committee ran a members’ photography contest to en-
courage discovery of nature and enhance the club’s digital
library. Over 200 entries were received and catalogued in
the digital photo library.
BROCHURES
The committee continued to distribute the club brochure
to area libraries and nature shops.
GILLIAN MARSTON
Chair, Education and Publicity Committee
Executive Committee
No report
Excursions and Lectures Committee
In 2005 this committee arranged 29 events, ten monthly
meetings (including the Annual Business Meeting) plus the
Soirée. The majority of our trips (twelve) were of general inter-
est, two were devoted to plants, two to geology, four to mam-
mals and invertebrates and nine were birding trips. The latter
included our club’s biannual bus trip to Pt. Pelee which was
very well attended, and two winter birding trips in conjunc-
tion with the Kingston Field-Naturalists Club. There have
been fewer birding trips the last five years because potential
leaders were heavily involved with the Breeding Bird Atlas.
Something new in 2005 was a winter wildlife tracking ex-
cursion, on snowshoes, in the Gatineau. Also a first, club mem-
bers were invited to join members of the Dragonfly Society
of America, meeting in Armprior, on their scheduled field trips.
Unfortunately a woodworking workshop had to be cancelled
for lack of interest.
Our policy remains to arrange meeting places that are ac-
cessible by bus. The signed waiver forms allow us to track
the popularity of events and help leaders to recognize parti-
cipants.
Our monthly meetings cover a great diversity of subjects
and most of our speakers are OFNC members.
FENJA BRODO
Chair, Excursions and Lectures Committee
Finance Committee
The Finance Committee met four times in 2005. Highlights
include:
1. Increase in membership fees for the calendar year 2007
Fees had not been increased for at least four years. and
some concerns were noted about gradually declining mem-
bership and operating deficits experienced over the previous
few years. The committee recommended to Council that fees
not be increased for calendar year 2006, but that the member-
ship fees increase by at least $5 in 2007. These recommen-
dations were approved by OFNC Council.
2. Meeting with the OFNC Auditor
The Treasurer, Frank Pope, and the Chair of the Finance
Committee met with the OFNC Auditor, Janet Gehr. She
advised that the financial health of the club was good com-
pared to similar non-profit organizations and that we had
ample reserves. She also advised that our computer hardware
and software were adequate, as well as the club’s record keep-
ing and documentation. It was recommended that future finan-
cial statements include a note on potential liabilities related to
the backlog of publication of the Canadian Field-Naturalist.
The amount of deferred revenue which would cover a part of
this financial commitment should also be noted. This recom-
mendation was acted upon by the Treasurer.
3. Budget for 2005-06 Fiscal Year
A proposed budget for the OFNC and the Canadian
Field-Naturalist for the period 1 October 2005 to 30 Septem-
ber 2006 was submitted to Council. A deficit of over $18,000
was forecast, and if realized would be a draw on the unrestrict-
ed reserve. Council approved this proposed budget.
4. Other Items
During the year the committee discussed the impact of
changes in the federal government’s Publication Assistance
Program on mailing costs for The Canadian Field-Natural-
ist and Trail & Landscape, the increase to the audit fee from
$1,000 to $2,000, a credit card for club business (not ap-
proved), the bequest from the estate of M. Groh, and reviewed
the draft financial statements for 2004-05 before finalization.
LOUISE SCHWARTZ
Chair, Finance Committee
Fletcher Wildlife Garden Committee
The Garden completed another successful year with over
2000 volunteer hours contributed. We held or participated in
a number of events. In February, it was the Great Backyard
Bird Count and then an information table at the Pesticides
Conference. April brought the Wildlife Festival with a dis-
play at Billings Bridge Shopping Centre, joint sponsorship of
a seminar at the Canadian Museum of Nature and an event at
our Centre for Annual Migratory Bird Day. Our annual plant
sale in June raised over $2000. We also participated in a Cen-
tral Experimental Farm Open House held in September.
Without a grant we were only able to have a part-time
employee who worked on the interpretation program. Our
preferred employee was unable to start immediately, so the
program began later and ran through September.
Remodelling of the Backyard Garden continued. Some beds
have been reshaped, a new entrance sign added and the patio
re-laid. In 2006 most of the beds will contain only regional
native flowers.
CANADIAN
Type Local Other
Family 312 (301) 23 (19)
Individual 307 (310) 101 (107)
Honorary 14 (14) 11 (10)
Life 21 (21) 20 (20)
Sustaining 10 (11) 2 (4)
Total 664 (657) 157 (160)
THE CANADIAN FIELD-NATURALIST
Vol. 120
Our work on invasive plants continues. There is some suc-
cess with buckthorn and garlic mustard, but swallow-wort
continues to be a problem. All methods of removing it end up
destroying everything else as well. We are also working on
burdock, Canada thistle and amur maple which have become
too common. Flowering rush and cat-tails are being removed
from the pond. The pond itself is showing decreased biodi-
versity with very few insect larvae observed in 2005. The pri-
mary water source is agricultural run-off so this trend may
be difficult to reverse.
In the spring we published a new general brochure in En-
glish, using a grant from TD Friends of the Environment
Foundation and intend to seek another grant to publish the
French version. Information sheets on burdock and garlic
mustard were also produced. For the Central Experimental
Farm Open House a bilingual miniguide was prepared to give
an overview of what the Fletcher Wildlife Garden has to offer.
This is now available to all visitors.
We have renewed the main entrance sign, enhanced the
boundary sign system and the direction signs along the Bill
Holland Trail. Other improvements include a small shed and
gravel on heavily used parts of the trail system. During the
winter there was considerable rabbit damage. We must expect
this problem to continue until the local fox population recovers.
DaviD HOBDEN
Chair, Fletcher Wildlife Garden Committee
Macoun Club Committee
The committee met once during the year, with most of the
planning being coordinated by telephone and e-mail. Commit-
tee members supervised or gave presentations at 18 indoor
meetings (down from 44 last year) and led 14 field trips and
one camping trip (roughly the same).
The much reduced number of meetings stems from the
failure of a Senior (high-school-age) group to form this year,
apparently for the first time in the Club’s 57-year history. In
part, this follows from weak participation over a period of
years in the Intermediate (middle-school) group, which would
normally feed into the Seniors. But it also appears to be a
social phenomenon of a proliferation of competing activities,
and ever more tightly scheduled lives.
Participation in the Junior group remains strong, although
the same social pressures have become evident in the fami-
lies of younger children, too. With no Senior members, the
Macoun Field Club’s annual publication (The Little Bear)
had to be edited and produced by a committee member for
the first time since 1988. The content, however, continues to
be dominated by the children’s work.
ROBERT E. LEE
Chair, Macoun Club Committee
FOREIGN
USA Other Total
2 (1) 1 (1) BS 8ma(622)
18 (19) 5 (4) 431 (440)
0 (0) 0 (0) 25 (24)
7 (6) 1 (1) 49 (48)
0 (0) 0) (0) 12 S))
27 (26) 7 (6) (855) (849)
2006
Membership Committee
The distribution of memberships for 2005 is shown in the
table on previous page, with the comparable numbers for 2004
in parentheses. These statistics do not include the 23 affiliate
organizations which receive complimentary copies of Trail
& Landscape.
During 2005, the club lost a long time member with the
death of Eileen Evans, who joined the club in 1974 and served
on Council as Corresponding Secretary from 1990 to 1996.
Eileen was an important organizer and contributor to the social
part of the monthly meetings at the Canadian Museum of
Nature and the annual Soirée for many years.
DAVE SMYTHE
Chair, Membership Committee
Publications Committee
The Publications Committee met three times in 2005.
Only two issues of The Canadian Field-Naturalist were
published in 2005: Volume 118, numbers two and three. These
MINUTES OF THE 127™ ANNUAL BUSINESS MEETING
397
two issues contained a total of 336 pages; 32 articles; 15
notes; 41 book reviews; 71 new titles; | commemorative trib-
ute; 14 pages of News and Comments; and 2 pages of miscel-
lany. The journal fell further behind schedule largely because
of some problems with the (relatively new) printer. These
problems seem to have been solved, and the next four issues
were near to being ready for production. The possibility of
eventual electronic publication was considered at some length,
and is continuing. For the present, the Abstracts of CFN
papers will be posted on the OFNC website (to accompany
the Tables of Contents which have been appearing there for
some time). None of the Manning funds were drawn upon
in 2005.
Volume 39 of Trail & Landscape was published in four
issues containing a total of 200 pages having the usual excel-
lent mix of articles.
RONALD E. BEDFORD
Chair, Publications Committee
398 THE CANADIAN FIELD-NATURALIST
Auditor’s Report
To The Members of THE OTTAWA FIELD NATURALISTS’ CLUB
I have audited the balance sheet of THE OTTAWA
FIELD NATURALISTS’ CLUB as at 30 September 2005, the
statement of changes in net assets, and the statements
of operations. These financial statements are the res-
ponsibility of the organization’s management. My res-
ponsibility is to express an opinion on these statements
based on my audit.
Except as explained in the following paragraph, I
conducted my audit in accordance with Canadian gen-
erally accepted auditing standards. Those standards re-
quire that I plan and perform an audit to obtain reason-
able assurance whether the financial statements are
free of material misstatement. An audit includes exam-
ining evidence supporting the amounts and disclosures
in the financial statements. An audit also includes as-
sessing the accounting principles used and significant
estimates made by management, as well as evaluating
the overall financial statement presentation.
In common with many non-profit organizations,
the Ottawa Field-Naturalists’ Club derives some of its
revenue from donations and fund raising activities.
These revenues are not readily susceptible to complete
audit verification, and accordingly, my verification was
limited to accounting for the amounts reflected in the
records of the organization.
In my opinion, except for the effect of the adjust-
ments, if any, which I might have determined to be
necessary had I been able to satisfy myself concern-
ing the completeness of the revenues referred to in the
preceding paragraph, these financial statements present
fairly, in all material respects, the financial position of
the OFNC as at September 30, 2005, and the results
of its operations and changes in net assets for the year
then ended in accordance with Canadian generally
accepted accounting principles.
JANET M. GEHR
Chartered Accountant
North Gower, Ontario
3 January 2006
The Ottawa Field-Naturalists’ Club
Balance Sheet
September 30, 2005
ASSETS
CURRENT
Cash (Note 1)
Investment certificates (Note 1)
Marketable securities (Note 2)
Accounts receivable
Prepaid expenses
LAND — ALFRED BOG
2005
$ 27,819
29 401
0
21,837
1,000
80,057
3,348
MARKETABLE SECURITIES (Note 2) 338,834
LIABILITIES AND FUND BALANCES
CURRENT
Accounts payable and
accrued liabilities
Deferred revenue
LIFE MEMBERSHIPS
NET ASSETS
Unrestricted
Club reserve
Manning principal
Manning interest - OFNC
- CFN
Seedathon
Anne Hanes memorial
de Kiriline-Lawrence
Macoun Baillie Birdathon
Alfred Bog
$ 422,239
$ 3,500
12,049
15,549
14,079
147,823
100,000
100,000
1,014
20,490
810
870
16,619
1,180
3,805
392,611
$ 422,239
Vol. 120
2004
$ 22,306
29,175
69,310
16,703
1,000
138,494
3,348
191,740
$ 333,582
$ 2,000
11,714
13,714
13,607
62,667
100,000
100,000
2.457
15,805
1,281
870
18,429
1,052
3,700
306,261
$ 333,582
2006 MINUTES OF THE 127™ ANNUAL BUSINESS MEETING
The Ottawa Field-Naturalists’ Club
Statement of Operations
for the Year Ended September 30, 2005
2005
REVENUE
Memberships $ 13,635
Trail and Landscape 220
Interest 2,129
GST rebate 720
Other nae OLE
17,326
OPERATING EXPENSES
Affiliation fees 652
Computer 1,846
Membership 1,363
Office Assistant 1,000
Telephone 1,733
Insurance WS)
Audit 2,500
GST ISS)
Other 1,072
12,646
CLuB ACTIVITY EXPENSES
Awards 390
Birds 618
Education and Publicity 1,149
Excursions and Lectures (1,805)
Macoun Field Club 466
Origins and History of OFNC 3,800
Soiree 228
Trail and Landscape 8,887
Fletcher Wildlife Garden (Note 4) 3,667
Other 0
17,400
Excess EXPENSES OVER
REVENUE $ (12,720)
2004
$ 13,969
280
1,835
768
373
17,225
670
1,294
1,168
1 000
1,640
U25
1,000
1,214
952
9,663
150
540
775
(508)
606
0
270
9578
(1,734)
84
9,761
$ (2,199)
The Ottawa Field-Naturalists’ Club
399
The Canadian Field-Naturalist — Statement of Operations
for the Year Ended September 30, 2005
2005
REVENUE
Memberships $ 9,090
Subscriptions 25,135
Reprints 6578
Publication charges 23,270
Interest and exchange 9 824
GST rebate 2,503
Other 1 23t
77,631
EXPENSES
Publishing 20,777
Reprints 4,658
Circulation 8532
Editing 2394
Office Assistant 5,000
Honoraria 9 000
GST rebate 2,693
Other 191
53,245
EXcCEss EXPENSES OVER
REVENUE $ 24386
2004
$ 9313
23 936
5 574
23,685
8 290
3,889
1,477
76,164
76,823
$ (659)
400
THE CANADIAN FIELD-NATURALIST
The Ottawa Field-Naturalists’ Club Notes to the Financial Statements
September 30, 2005
1. CASH
Chequing
Savings
Nesbitt Burns
Fletcher Wildlife Garden
Investment Certificates: 2005
2. MARKETABLE SECURITIES 2005
Investment Certificates:
Province of Newfoundland Coupon
Province of Ontario Coupon
Province of Manitoba Coupon
Res CIBC Int BB6
CMHC Global Debs
Province of Ontario Bond
Province of Newfoundland Bond
Government of Canada Coupon
Province of New Brunswick Bond
Province of New Brunswick Bond
3. CAPITAL ASSETS
Equipment at a cost of $16,748 is fully amortized.
4. FLETCHER WILDLIFE GARDEN
REVENUE
Human Resources and Skills Dev. Canada
TD Friends of the Environment Fund
Taverner Cup
Sales
GST
Donations
EXPENSES
Program
Backyard
Habitats
Interpretation centre
Administration
Publications
GST
Library
Maturity
Value
$ 29,963
Maturity
Value
$ 44,782
5237/16
45,740
70,827
52,000
30,000
20,000
30,167
60.000
20,000
Maturity
Date
05/01/06
Maturity
Date
10/17/11
12/02/12
09/05/13
10/31/14
12/01/06
09/12/07
10/07/08
12/01/09
12/03/15
06/15/10
2005
$ 16,144
6,963
459
4,253
$ 27,819
Yield
1.90%
Yield
4.525%
4.591%
4.694%
4.144%
5.250%
6.125%
6.263%
5.605%
3.965%
6.231%
1,479
388
112
“aan
$(3,667)
Vol. 120
2004
$ 11,899
4,49]
0
5,916
$ 22,306
Book
Value
$ 29.401
Book
Value
$ 34,509
11,106
31,909
48 934
53,539
31,187
20,538
23,534
62,510
21,068
$338,834
8460
$(1,734)
2006 MINUTES OF THE 127™ ANNUAL BUSINESS MEETING 40)
The Ottawa Field-Naturalists’ Club Statement of Changes in Net Assets
for the Year Ended September 30, 2005 (Note 5)
Net Beginning Excess Excess Other Other Ending
Assets Balance Revenue CFN —_ Expenses OFNC Revenue Expenses _—_ Balance
Unrestricted $ 62,667 $ 24,386 $ (12,720) $ 73490a $ $ 147,823
Club reserve 100,000 ~ ~ : 100,000
Manning Principal 100,000 - ~ 100,000
Manning - CFN 15,805 - - 4,685 20,490
Manning - OFNC 2,457 - - 1,172 (2,615 b) 1014
Seedathon 1,28] - - 807 (1,278 c) 810
Anne Hanes Memorial 870 - - ~ ~ 870
de Kiriline-Lawrence 18,429 - - 190 (2,000 d) 16,619
Macoun Baillie Birdathon 1,052 - - 128 - 1,180
Alfred Bog 3,700 - - 105 ~ 3,805
$ 306,261 $ 24,386 $ (12,720) $ 80,577 $ (5,893) $ 392,611
5. STATEMENT OF CHANGES IN NET ASSETS
a) Unrestricted other revenue includes a bequest of $62,250
from the Estate of Mildred Groh.
b) Manning OFNC expenses: Innis Point Bird Observatory
$2,000, Peregrine Falcon Watch $649.
c) Seedathon Expenses: birdseed for club feeders, $809.
d) Louise de Kiriline-Lawrence Expenses: Contribution
to the defence of the Larose Forest at a hearing of the
Ontario Municipal Board.
6. PUBLICATION LIABILITY
An annual membership fee or subscription entitles the
member/subscriber to four issues of The Canadian Field-
Naturalist based on a calander year. As the year end of the
club is September 30, the Club incurs a liability for publishing
the fourth issue of each publication.
At this time, however, the publication of The Canadian
Field-Naturalist is ranning late. At September 30, 2005, the
club owes members/subscribers of 2005 issue number 4 of
2004 and three issues of 2005. Although most of the work
preparing these publications is done by volunteers, the club
must pay for printing and mailing. Based upon recent costs,
it is estimated that the club has a liability of $68,000 for the
outstanding issues. This amount will be reduced by page charg-
es to the authors in the amount of 40% of the printing costs.
The Ottawa Field-Naturalists’ Club Summary of Significant Accounting Policies
September 30, 2005
1. Nature of Business
The organization is non-profit and incorporated under the
laws of Ontario (1884). The organization promotes the appre-
ciation, preservation, and conservation of Canada’s natural
heritage. It encourages investigation and publishes the results
of the research in all fields of natural history and diffuses infor-
mation on these fields as widely as possible. It also supports
and cooperates with other organizations engaging in pre-
serving, maintaining or restoring environments of high quality
for living things.
2. Financial Instruments
The organization’s financial instruments consist of cash,
accounts receivable, marketable securities, and accounts pay-
able. Unless otherwise noted, it is the management’s opinion
that the organization is not exposed to significant interest, cur-
tency, or credit risks arising from these financial instruments.
The fair value of these instruments approximate their carry-
ing values, unless otherwise noted.
3. Capital Assets
Capital assets in excess of $4,000 cost are recorded as assets
at cost and amortized on a straight-line basis. These assets have
been fully amortized.
4. Statement of Changes in Financial Position
A statement of changes in financial position has not been
provided as it would not provide additional meaningful infor-
mation.
5. Foreign Currency
Transactions during the year in U.S. dollars have been con-
verted in the accounts to Canadian dollars at the exchange
rate effective at the date of the transaction. All monetary assets
in U.S. dollars at year end have been converted to Canadian
dollars at the rate effective on Sept. 30, 2005.
Gains or losses resulting therefrom are included in revenue
or expenses.
Advice for Contributors to The Canadian Field-Naturalist
Content
The Canadian Field-Naturalist is a medium for the publi-
cation of scientific papers by amateur and professional natu-
ralists or field biologists reporting observations and results
of investigations in any field of natural history provided that
they are original, significant, and relevant to Canada. All read-
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journal also publishes natural history news and comment items
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Review Editor concerning suitability of manuscripts for this
section. For further information consult: A Publication Policy
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian
Field-Naturalist 97(2): 231-234. Potential contributors who
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nor subscribers to The Canadian Field-Naturalist are encour-
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subscribers.
Manuscripts
Please submit by post to the Editor, in either English or
French, three complete manuscripts written in the journal
style. Manuscripts may also be submitted (one copy) by e-
mail. The research reported should be original. It is recom-
mended that authors ask qualified persons to appraise the
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islation regarding the study, disturbance, or collection of ani-
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“Standard” common names (with initial letters capitalized)
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Check recent issues (particularly Literature Cited) for
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ter (don’t type) descriptive matter. Write author’s name, title
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FRANCIS R. Cook, Editor
RR 3 North Augusta, Ontario KOG IRO Canada
402
TABLE OF CONTENTS (concluded) Volume 120 Number 3 2006
Book Reviews
Zoo.oGy: The Birder’s Companion — Birds of the Dominican Republic and Haiti — Atlas of Bird
Migration: Tracing the Great Journeys of the World’s Birds — Handbook of the Birds of the World
Volume II — Old World Flycatchers to Old World Warblers — Conservation of the Black-tailed Prairie
Dog: Saving North America’s Western Grasslands — Migrating Raptors of the Word: Their Ecology
and Conservation — Land Snails of British Columbia — Songbird Journeys: Four Seasons in the Lives
of Migrating Birds 379
Borany: Wildflowers of the Rocky Mountains — Plants of Alberta 386
MISCELLANEOUS: Pilgrimage to Vallombrosa: From Vermont to Italy in the Footsteps of George Perkins
Marsh — Where to Watch Birds: World Cities — Marshes: The Disappearing Edens 387
New TITLES
News and Comment
Marine Turtle Newsletter 113 — The Boreal Dip Net/L’Epuisette Borale January 2006 392
inutes of the 127" Annual Business Meeting of The Ottawa Field-Naturalists’ Club 393
Advice to Contributors 402
Mailing date of the previous issue 120(2): 1 November 2007
THE CANADIAN FIELD-NATURALIST Volume 120 Number 3
Articles
An ethogram developed on captive eastern Coyotes, Canis latrans
JONATHAN G. WAY, DEAN-LORENZ M. SZUMMMYLO, and ERIC G. STRAUSS
First observations of an Eastern Screech-Owl, Megascops asio, population in an apple-producing
region of southern Quebec NGAIO L. RICHARDS, PIERRE MINEAU, DAVID BIRD,
PIERRE WERY, JACQUES LARIVEE, and JASON DUFFE
Effects of timber harvesting and plantation development on cavity-nestng birds in New Brunswick
STEPHEN J. WOODLEY, GREG JOHNSON, BILL FREEDMAN, and DAVID A. KIRK
Recent invasion, current status, and invasion pathway of European Common Reed,
Phragnites australis subspecies australis, in the southern Ottawa District
PAUL M. CATLING and SUSAN CARBYN
Pack Size of Wolves, Canis lupus, on Caribou, Rangifer tarandus, winter ranges in
west-central Alberta GERALD W. KUZYK, JEFF KNETEMAN, and FIONA K. A. SCHMIEGELOW
Bur Buttercup, Ranunculus testiculatus , new to eastern Canada
MICHAEL J. OLDHAM, CLIVE E. GOODWIN, and SEAN BLANEY
Assessment of effects of an oil pipeline on Caribou, Rangifer tarandus granti, use of
riparian habitats in arctic Alaska, 2001-2003 LYNN E. NOEL, MATTHEW W. BUTCHER,
MATTHEW A. CRONIN, and BILL STREEVER
First records of the Southern Red-backed Vole, Myodes gapperi, in the Yukon Territory
THOMAS S. JUNG, ANY RUNCK DAvID W. NAGORSEN, BRIAN G. SLOUGH, and TODD POWELL
A conservation evaluation of Smooth Goosefoot, Chenopodium subglabrum (Chenopodiaceae),
in Canada DIANA BIZECKI ROBSON
Summer movements and impact of individual Striped Skunks, Mephitis mephitis, on duck nests in
Saskatchewan SERGE LARIVIERE, LYLE R. WALTON, and FRANCOIS MESSIER
Life history phenology and sediment size association of the dragonfly Cordulegaster dorsalis
(Odonata: Cordulegasteridae) in an emphemeral habitat in southwestern British Columbia
LAURIE B. MARCZAK, JOHN RICHARDSON, and MARIE-CLAIRE CLASSEN
An analysis of the vascular flora of Annapolis heathlands
S. CARBYN, P. M. CATLING, S. P. VANDER KLOET, and S. BASQUILL
Pacific Hagfish. Eppptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and scavenger activity
on tethered carrion in subtidal benthic communities off western Vancouver Island
SARAH DAVIES, ALI GRIFFITHS, and T. E. REIMCHEN
Notes
A new record size Wolf, Canis lupus, pack for Ontario Liv S. Vors and PHILIP L. WILSON
Tributes
A tribute to Nicholas Stephen Novakowski 1925-2004 JOSEPH E. BRYANT
ISSN 0008-3550
2006
263
(continued on inside back cover)
CAIN
IS AY
The CANADIAN
FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
1
Volume 120, Number 4 October-December 2006
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Her Excellency The Right Honourable Michaélle Jean
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members
Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope
Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr.
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson
2006 Council
President: Mike Murphy Ronald E. Bedford —_ Diane Kitching Dan Millar
Vice-President: Ken Allison Fenja Brodo Karen McLachalan Hamilton Justin Peter
Recording Secretary: Susan Laurie-Bourque Julia Cipriani David Hobden Stanley Rosenbaum
Treasurer: Frank Pope William J. Cody Diane Lepage Henry Steger
Past President: Gary McNulty Francis R. Cook Ann Mackenzie Chris Traynor
Susan Howell Gillian Marston Eleanor Zurbrigg
To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069,
Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net.
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.
We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward
our mailing costs. PAP Registration Number 9477. Canada
Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca
Copy Editor: Elizabeth Morton
Honorary Business Manager: William J. Cody
Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwgsc.gc.ca
Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen
Charles D. Bird Brian W. Coad Donald F. McAlpine
Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr.
Chairman, Publications Committee: Ronald E. Bedford
All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail
or e-mail.
Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor.
Subscriptions and Membership
Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per year
(volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) and $500 (life)
includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must add an addi-
tional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club events.
It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $33 per year.
Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The
Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement
number 40012317. Return Postage Guaranteed. Date of this issue: October- December 2006 (January 2008).
————- >
|
Cover: Incubating female Northern Hawk Owl Surnia ulula, remained seated on the nest for long periods of time, changing her
position infrequently (nest number 1). Photo by Michael Patrikeev. 30 May 2001. See article pages 433-437.
The Canadian Field-Naturalist | ™ 04 2008
Volume 120, Number 4 October—Decémber 2906
A Reevaluation of Sexual Dimorphism in the Postcranium of
the Chasmosaurine Ceratopsid Chasmosaurus belli (Dinosauria:
Ornithischia)
JORDAN C. MALLON! and ROBERT B. HoLmMeEs2+
'Department of Earth Sciences, Carleton University, Ottawa, Ontario K1S 5B6 Canada; e-mail: jmallon@ucalgary.ca
Research Division, Palaeobiology, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario KIP 6P4 Canada
Current address: Faculty of Veterinary Medicine, University of Calgary, 3330 Hospital Drive, Calgary, Alberta T2N 4N1
Canada; e-mail: jmallon@ucalgary.ca
‘Current address: Department of Biological Sciences, University of Alberta, CW 405 Biological Sciences Building, Edmonton,
Alberta T6G 2E9 Canada; e-mail: holmes! @ualberta.ca
Mallon, Jordan C., and Robert B. Holmes. 2006. A reevaluation of sexual dimorphism in the postcranium of the chasmosaurine
ceratopsid Chasmosaurus belli (Dinosauria: Ornithischia). Canadian Field-Naturalist 120(4): 403-412.
The sexual dimorphism attributed to Chasmosaurus belli by Sternberg (1927) is revisited and reevaluated. A reexamination
of the two specimens originally considered by Sternberg reveals that they are less complete than first suggested, with only a
moderate amount of overlapping material between them. Only a few of the postcranial elements (humeri, sternal plates, and
presacral vertebrae) show evidence of dimorphism, the significance of which is either doubtful or equivocal. Instead of repre-
senting sexual dimorphs, it is likely that the two specimens belong to separate species, C. belli and C. russelli, as evidenced
by their distinct frill morphologies and by their stratigraphic segregation within the Dinosaur Park Formation of Alberta.
These findings emphasize the need to remain sceptical about claims advocating sexual dimorphism in the fossil record in the
absence of statistical significance or stratigraphic control.
Key Words: Chasmosaurus, Ceratopsia, Dinosaur Park Formation, sexual dimorphism, variation, Alberta.
Chasmosaurus are modest-sized (~1500 kg) ceratop-
sid dinosaurs known from the upper Campanian-aged
beds (~75 Ma [million years ago]) of western North
America. They are characterized by the possession of a
premaxillary flange, recurved supraorbital horncores,
and a broad parietosquamosal frill with a straplike pos-
terior border (Forster et al. 1993). The type species,
Chasmosaurus belli (Figure 1), was discovered in the
middle strata of the Dinosaur Park Formation in Alber-
ta and was originally named Monoclonius belli (Lambe
1902). Two additional species of Chasmosaurus, C.
russelli and C. irvinensis, are currently recognized from
elsewhere in the formation (Godfrey and Holmes 1995;
Holmes et al. 2001; Ryan and Evans 2005). A further
species, C. mariscalensis, has been described from the
Aguja Formation in Texas (Lehman 1989), though it
was recently given the new genus name Agujaceratops
(Lucas et al. 2006) and may prove to be more closely
related to Pentaceratops than to Chasmosaurus (Holmes
et al. 2001; Diem and Archibald 2005).
Charles M. Sternberg (1927) reported on two spec-
imens of Chasmosaurus, both identified as C. belli, on
display at the National Museum of Canada (now the Figure 1. Life restoration of Chasmosaurus belli, based on
Canadian Museum of Nature) in Ottawa (Figure 2). CMN 2245. Illustration by JCM.
403
404
the Canadian Museum of Nature).
As restored, the skeletons measured “approximately
the same length but the bones of one... are much lighter
in construction than those of the other” (Sternberg
1927: 67). According to Sternberg, the total length of
the ‘gracile’ skeleton measured 4.93 m, while that of the
“robust” skeleton measured 4.95 m. Unfortunately, their
supports have since been disassembled and these
original measurements cannot be verified. Sternberg
attributed the difference in robustness between the two
specimens to sexual dimorphism, stating, “it is thought
that they represent male and female as it is common,
among reptiles, for the female to be larger than the
male” (Sternberg 1927: 67). This assumption has gone
largely unchallenged in the literature (e.g., Lull 1933;
Carpenter and Currie 1990; Dodson 1996), except for
an implication by Godfrey and Holmes (1995) that
the reported differences between the specimens do not
reflect sexual dimorphism because they pertain to two
separate species of Chasmosaurus.
The recognition of sexual dimorphism in a fossil
species must begin with the establishment that the vari-
ation observed within that species does represent dimor-
phism. With only two reasonably complete skeletons
of Chasmosaurus belli at his disposal, Sternberg could
not have established this with certainty, and indeed,
he neither quantified nor qualified the differences he
observed between the specimens. It is also important
to note that many of the bones were highly reconstruct-
ed with plaster and subsequently painted over so that
the reconstructed portions cannot be easily identified.
THE CANADIAN FIELD-NATURALIST
Vol. 120
FIGURE 2. CMN 2280 (left) and CMN 2245 (right) on display at the National Museum of Canada (circa 1926). (Courtesy of
As chronicled by C. H. Sternberg (C. M. Sternberg’s
father) regarding the restoration of one of the skulls:
“we had colored our plaster to resemble the fossil bone
—no small task, by the way, as we had to learn to mix
colors as well as do the work of a sculptor — with
wax” (Sternberg 1917: 83). As such, the possibility
remains that the difference in robustness between the
two specimens might in part be attributable to the
extensive restoration of certain elements. With these
concerns in mind, we set out to identify and describe
the variation noted between the skeletons, and con-
ducted a reevaluation of Sternberg’s original attribu-
tion of sexual dimorphism to C. belli.
Materials
The two Chasmosaurus belli skeletons (Figure 3)
described by Sternberg (1927) reside in the collec-
tions of the Canadian Museum of Nature in Aylmer,
Quebec. The “gracile” specimen was designated CMN
2245; the “robust,” CMN 2280. CMN 2245 was col-
lected from quarry Q037, and CMN 2280 from QO10,
of the Dinosaur Park Formation (upper Campanian,
~75 Ma) in Dinosaur Provincial Park, Alberta (origi-
nally referred to by Sternberg [1927: 67] as the “Belly
River series’).
CMN 2245 (Figure 3A) is represented by the pos-
terior half of a skull and mandibles (missing only the
predentary), complete presacral series, synsacrum com-
plete to the sixth sacral vertebra, twenty-four caudal
vertebrae, most cervical and thoracic ribs, pectoral gir-
2006 MALLON AND HOLMES: SEXUAL DIMORPHISM IN CHASMOSAURINE CERATOPSID 405
[ ’ . Utter re... =
: SETAE Vii" cee
FIGURE 3. Material (in grey) attributed to (A) CMN 2245 and (B) CMN 2280. Scale bar = 1 m. Modified from original
skeletal drawing by Gregory S. Paul. Used with permission.
406
dle (missing only the right coracoid), both humeri, left
ulna, pelvic girdle, both femora, right tibia and fibula,
and several carpal and tarsal elements.
CMN 2280 (Figure 3B) consists of a complete skull
and mandibles, complete presacral series, synsacrum
complete to the fourth sacral vertebra, most cervical
and thoracic ribs, pectoral girdle (missing only the left
coracoid), both humeri, anterior halves of both ilia, both
pubes, and the proximal third of the left femur. A return
visit to the quarry has yielded more hindlimb material,
but it was in poor condition and not collected (D. H.
Tanke, personal communication).
Overlapping material between the two specimens
therefore consists of the posterior region of the skull,
presacral and anterior sacral vertebrae, most ribs, most
of the pectoral girdle, humeri, anterior ilia, pubes, and
left proximal femur.
Methods
In order to facilitate distinction between the original
bone and plaster, the brown paint was removed from
many of the bones using acetone and a stiff-bristled
brush. In some instances, the removal of the paint was
not warranted when it was obvious that the element of
interest had been created entirely of plaster (in which
case a simple tap with the knuckle would confirm sus-
picion). In this way, the original elements common to
CMN 2245 and CMN 2280 could be identified.
Three hundred and fourteen measurements were then
taken from the postcranium of each specimen to the
nearest millimetre, primarily according to the standards
of Chinnery (2001). For those elements not considered
by Chinnery, such as those belonging to the axial skele-
ton, one of us (JCM) devised our own measurement
parameters. Measurements below 300 mm were gen-
erally taken with Mitutoyo SD type dial callipers and
measurements above 300 mm were taken either with
simple outside callipers and measuring tape or with large
(1.04 m) Helios brand Vernier callipers. Circumferences
were measured using a tailor’s measuring tape. Angles
were measured from photographs using a protractor.
All available postcranial elements were measured, but
only those bones deemed reliably complete and shared
between the two skeletons were considered in this study.
Comparative Osteology
Description of Overlapping Material
Although CMN 2245 and CMN 2280 exhibit mod-
est variability in the dimensions of their overlapping
elements (Figure 4), much of this appears to be due to
the extensive restoration and occasional postmortem
deformation of the skeletons. Despite this, the best pre-
served elements usually differ in size by only a few
percent, which is insufficient to produce visually dis-
cernable dimorphism, and is probably within the realm
of individual variation that might be expected in tet-
rapods (e.g., Rising and Somers 1989; Zaaf and Van
Damme 2001; Kelly et al. 2006). The coracoids of
THE CANADIAN FIELD-NATURALIST
Vol. 120
CMN 2245 and CMN 2280 are noticeably different in
shape, although the left and only coracoid preserved
in the former specimen has been badly weathered and
highly reconstructed as a result, and is therefore of
limited use. Only a few bones differ considerably in
several dimensions and are described below.
Humerus
The humerus (Figure 4A) of CMN 2280 is consid-
erably more robust than that of CMN 2245 in most
dimensions (Table 1), particularly those of the humer-
al head, deltopectoral crest, and midshaft circumfer-
ence. Besides proportional differences in the humerus,
the insertional scar for the /atissimus dorsi muscle on
the deltopectoral crest is much less pronounced in
CMN 2245.
Sternal Plate
Although the sternal plate (Figure 4B) is quite sim-
ilar in size and shape between the two specimens, the
process at the posterior end of the element extends
42-55% further laterally in CMN 2280 than in CMN
2245.
Presacral Vertebrae
The presacral vertebrae (Figure 4C) of these speci-
mens are uniform in most dimensions, with most appar-
ent differences being attributable to the addition of
plaster. However, in the region of the withers (anteri-
or thoracics), the vertebrae of CMN 2280 average
21% taller (Figure SA) and their neural spines 19°
more erect (Figure 5B) than in CMN 2245. Similarly,
the transverse processes of the cervical and anterior
thoracic vertebrae of CMN 2280 average 17° more
erect than in CMN 2245 (Figure 5C), although this is
difficult to state with confidence given the artificial
coalescence of the vertebrae and the incompleteness
of many of their spinous processes in CMN 2245. It
seems, too, that the transverse widths of the vertebral
centra anterior to the sixth thoracic average 10% wider
in CMN 2280 (Figure 5D).
Discussion
Preservation of CMN 2245 and CMN 2280
While CMN 2245 is the most complete skeleton of
Chasmosaurus known to date, it is less complete and
less well preserved than generally supposed (e.g., Dod-
son 1996: 107). Some of the elements touted by Stern-
berg (1927) as being complete, such as the femora and
caudal vertebrae, are actually only partially represent-
ed by nondescript scraps of bone buried in plaster,
causing confusion among later authors (e.g., Lull 1933).
Many of the presacral vertebrae have also been artifi-
cially “fused” together with plaster, obscuring much
of the detail. CMN 2280 is by far the better preserved
of the two specimens, although it is less complete.
Consequently, there is only a moderate amount of mate-
rial shared between the two specimens, and additional
overlapping material would be desirable to make a con-
vincing case for dimorphism in C. belli.
2006 MALLON AND HOLMES: SEXUAL DIMORPHISM IN CHASMOSAURINE CERATOPSID 407
FiGurE 4. Dimorphic overlapping material of (A) CMN 2245 and (B) CMN 2280. i, right and left humeri; ii right and left
sternal plates; iii posterior thoracic vertebrae in anterior (left) and left lateral (right) views. Note that, because of the
poor preservation of the overlapping vertebrae, the eighth thoracic vertebra is figured for CMN 2245 and the tenth
thoracic vertebra is figured for CMN 2280. Scale bar = 10 cm.
A Reevaluation of Sexual Dimorphism in Chasmosaurus __ et al. 2005). Chinnery (2001, 2004) attempted to discern
belli morphological variation, including sexual dimorphism,
Recognizing sexual dimorphism in a fossil species in the appendicular skeleton of the Ceratopsia, but
is rarely a straightforward task, especially given small _ could find none. The dimorphism originally ascribed by
Sample sizes where subtle but statistically significant Sternberg (1927) to CMN 2245 and CMN 2280 was
dimorphic characters are difficult to resolve (Padian _ presumably based on variation in the postcranial skele-
408
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Selected measurements (after Chinnery 2001) for left humeri of CMN 2245 and CMN 2280, showing the large
differences between their dimensions. Dimensions in square brackets are estimated.
Measurement parameter
Width of deltopectoral crest
Deltopectoral crest length, from external tuberosity to distal
end of deltoid muscle scar
Craniolateral view; proximal width
Head height
Head width
Circumference about midshaft*
* = parameter not used in Chinnery (2001).
ton, although distinct differences in frill morphology
are also present (see below). These differences cannot
be attributed to either allometry or ontogeny since both
specimens are of comparable size. Nor are they like-
ly due to geographic variation since both specimens
were found less than nine kilometres from one anoth-
er. Post-mortem distortion cannot account for the dif-
ferences either, as most of the elements are relatively
uncrushed.
A review of the overlapping material reveals that the
most striking differences between CMN 2245 and
CMN 2280 lie with the humerus. The disparity in the
robustness of the humerus and in the size of the inser-
tional scar for the Jatissimus dorsi muscle seems to
follow the prediction made by Chapman et al. (1997)
that sexual dimorphism in ceratopsians would be ex-
pressed in the limb bones. It is also reminiscent of the
condition seen in the humerus of many pachypleu-
rosaurid sauropterygians (e.g., Sander 1989; Cheng
et al. 2004) and of the sauropod Camarasaurus (Ike-
jiri 2005). However, if these humeral characters alone
separate male from female, then other specimens of
Chasmosaurus belli should exhibit the same disparity
in humeral robustness as well. A bivariate plot derived
from a Principle Components Analysis by Chinnery
(2001: 134) revealed that the humerus of CMN 2245
plots as an outlier from the other eleven Chasmosaurus
humeri included in the study. As the deviation lies along
the y-axis, the differences are likely due to shape rather
than size. Thus, while the shape differences between
the humeri of CMN 2245 and CMN 2280 are real, the
highly asymmetrical sex ratio implied is very unlikely.
The inability of Lehman (1990) to identify dimorphism
in the humeri from a population of C. mariscalensis,
and M. J. Ryan’s (personal communication) failure to
find the same in several populations of Centrosaurus
apertus, further supports this position. Sternberg (1927)
specifically states that “‘[t]here have been no bones of
other individuals used in the mounts”, so the differ-
ences cannot be ascribed to the incorporation of mate-
rial from a smaller individual into the mount. It seems
that the humeri of CMN 2245 are simply unusually
small and gracile, although the reason for this is unclear.
The reasons for the disparity in the dimensions of the
Dimensions (mm) Disparity
CMN 2245 CMN 2280 (%)
95 72 24
225 Dil, IES
44 76 42
[56] 78 28
55 86 36
216 275 21.5
sternal plates and of the presacral vertebrae are likewise
uncertain in the absence of more material. The appar-
ently longer and more erect neural spines and transverse
processes of CMN 2280 coincide with observations
made by Tereshchenko (2001) of protoceratopsids, in
which the spinous processes of the vertebrae were
reputedly shown to be longer and more erect in males
than in females. If this apparent dimorphism is sexual,
however, it would imply the “robust” specimen (CMN
2280) was male rather than female (contra Sternberg
1927). This type of “normal” sexual dimorphism, in
which the male is more massive than the female, is typ-
ical of most reptiles (Fitch 1981; Shine 1989; Brochu
2002).
Thus, undoubted differences (albeit fewer than orig-
inally implied by Sternberg) between the two speci-
mens do exist. The marked difference in the size and
shape of the humerus may simply be anomalous, unless
we accept the unlikely hypothesis that of the twelve
Chasmosaurus humeri measured by Chinnery (2001),
eleven are of one sex and only one is of the other.
Other differences, such as the specific shape of the
sternal plates and the relative size and orientation of
the spinous processes of the presacral vertebrae, might
represent sexual dimorphism or intraspecific variation.
It is also possible that all of the postcranial discrep-
ancies noted here are functionally related, reflecting a
developmental compensation of the extrinsic muscles
of the forelimb in response to the atrophied humeri
of CMN 2245. However, in the absence of a statisti-
cally significant sample, it is impossible to resolve this
question. Whatever the significance of the variation,
there seems little choice but to reject the hypothesis
of sexual dimorphism.
Two-Species Hypothesis
It is possible that the few differences documented
here within the postcranial skeletons of CMN 2245 and
CMN 2280 may simply reflect that these two speci-
mens represent separate species. In a review of the
systematics of the genus Chasmosaurus (Godfrey and
Holmes 1995), two species were diagnosed based on
differences in parietosquamosal frill structure. C. belli
was defined as possessing a “parietal frill with nearly
straight transverse posterior bars, each bearing one large
2006
vertebra (mm)
ee A =
SBSBSSBSERREFKRKE
Vertebra
T8
Tg
T10
mT
T12
—t— CMN 2245 —@- CMN 2280
Vertebra
—*— CMN 2245 —®- CMN 2280
MALLON AND HOLMES: SEXUAL DIMORPHISM IN CHASMOSAURINE CERATOPSID
409
70
Ls 8) A A
hee é /
/ e “~
be / —. ya Poe
» tn | \™* Oe =
3 : . /
low © ;
2 en ae
i]
S8BSBSFFFFRF REF RFP SES
Vertebra
—e- CMN 2245 —@- CN 2280
120
- 110 = aN
gp ete
3E \ tA s
2 § ee = 2" VA
se J \_~<
2s 70
ce a ——-
50
SS8SSBSFFFFFRFERFSSES
FiGuRE 5. Graphical depictions of vertebral disparity between CMN 2245 and CMN 2280. A, Differences between maxi-
mum heights of vertebrae. The anterior thoracic vertebrae of CMN 2280 are consistently taller than those of CMN
2245. B, Differences between angles of neural spines. In the anterior thoracic region, the neural spines of CMN
2245 are angled further posteriorly than in CMN 2280. C, Differences between angles formed by transverse
processes. The transverse processes of the cervical and anterior thoracic vertebrae of CMN 2280 are more erect than
those of CMN 2245. D, Differences between transverse widths of centra. The cervical and anterior thoracic centra of
CMN 2280 are consistently wider transversely than those of CMN 2245. Abbreviations: C, cervical vertebra; T, tho-
racic vertebra.
triangular epoccipital on its posterolateral corner; other
parietal epoccipitals variable in number and degree of
coossification with the parietal, but always much small-
er. The lateral bar of the parietal completely encloses
the parietal fenestra.” C. russelli was characterized as
having a frill in which the posterior margin is “broadly
arched on either side of [the] median emargination.
Each side bears three low triangular, roughly equal-
sized epoccipitals. The lateral ramus of the parietal is
reduced and does not completely encircle the fenes-
tra in all but one specimen, permitting the squamosal to
form a part of its lateral border.’ Under these amended
diagnoses, CMN 2245 was retained within the hypo-
digm of C. belli, but CMN 2280 was reassigned to C.
russelli. This distinction was challenged by Lehman
(1998), who argued that the discrete characters used to
distinguish C. belli from C. russelli actually represent
end-members of a gradational spectrum of frill mor-
phologies. In support of his argument, Lehman (1998:
figure 9) figured seven skulls attributed to C. belli and
C. russelli, illustrating the range of frill morpholo-
gies he observed. However, the lateral parietal bars of
AMNH 5402 used by Lehman are not illustrated as
being complete as they are in the original specimen,
which creates the false impression that the frills exhibit
graded variation. Correcting for this splits the series
into two discrete groupings of frill morphologies (Fig-
ure 6), the contents of which agree with the original
hypodigms of Godfrey and Holmes (1995).
Further support for the distinction between C. belli
and C. russelli stems from their stratigraphic segrega-
tion within the Dinosaur Park Formation (Figure 7).
This observation was made previously by Godfrey and
Holmes (1995) and Holmes et al. (2001), and more
recent quarry data support this claim, with C. russelli
confined to the lower part of the formation and C. belli
located much higher in the section (Ryan and Evans
2005). These independent lines of evidence strongly
suggest that the variation between CMN 2245 and
CMN 2280 is interspecific in nature, negating Stern-
berg’s case for sexual dimorphism. Stratigraphic sep-
aration has also been noted recently among the suppos-
edly sexually dimorphic lambeosaurine hadrosaurids
of the Dinosaur Park Formation (Evans et al. 2006),
410
viii
THE CANADIAN FIELD-NATURALIST
Vol. 120
q
FiGureE 6. Parietosquamosal frills of (A) Chasmosaurus belli and (B) C. russelli in dorsal view. i, CMN 491 (holotype of C.
belli); ii, YPM 2016; iii, CMN 2245 (paratype of C. belli); iv, AMNH 5402; v, ROM 843; vi, CMN 8803 (paratype
of C. russelli); vii, CMN 2280; viii, RTMP 83.25.1; ix, AMNH 5656. Frills not to scale.
suggesting a need for more critical consideration of
sexual dimorphism in the fossil record.
Conclusion
After a careful review of CMN 2245 and CMN 2280,
it seems that the postcranium of neither specimen is
quite as complete or as well preserved as originally
described by Sternberg (1927). Consequently, the case
for dimorphism (sexual or otherwise) is weaker than
previously thought. Most of the variation exhibited by
the shared elements is attributable either to poor recon-
struction, postmortem distortion, or individual variation.
Exceptional cases involve the humeri, sternal plates,
and presacral vertebrae.
While the disparity in the shape of the humerus is
indeed genuine, the same dimorphism is not seen in
other specimens of Chasmosaurus, so this feature is
likely anomalous rather than dimorphic. The sternal
plates of CMN 2245 and CMN 2280 differ in the lengths
of their posterolateral processes, and the presacral ver-
2006
tebrae differ in the lengths and orientations of their spin-
ous processes. However, analysis of additional Chas-
mosaurus material — preferably derived from some yet
undiscovered monodominant bonebed (Eberth and
Getty 2005) — would be necessary to determine the
significance of this variation with regards to sexual
dimorphism. The results presented here are therefore
only preliminary, pending further investigation of indi-
vidual variation in the ceratopsian postcranial skeleton.
Finally, previous detailed reviews of the frill mor-
phology of either specimen seem to suggest that CMN
2245 and CMN 2280 may, in fact, belong to separate
species (C. belli and C. russelli, respectively), as first
put forth by Godfrey and Holmes (1995). This view is
further supported by the stratigraphic segregation of
the two specimens within the Dinosaur Park Formation
of Alberta. Whether the postcranial differences iden-
tified here represent interspecific distinctions remains
to be demonstrated. Therefore, although there do seem
to be a few legitimate differences between the two
specimens, the weight of the evidence so far accumu-
lated does not support sexual dimorphism as Stern-
berg (1927) first suggested it, and there remains little
choice but to reject this hypothesis. In the future, sex-
ual dimorphism should be attributed to fossil species
only when statistical significance and stratigraphic
control have been demonstrated. Claims made in the
absence of such evidence should be regarded with
scepticism.
Acknowledgments
We would like to thank Gilles Danis, Margaret
Feuerstack and Kieran Shepherd for offering access to
specimens in their care. Philip Currie and Hans Larsson
provided valuable resources and discussion. Andrew
Farke, Michael Ryan, and two anonymous reviewers
provided constructive criticism on earlier versions of
this manuscript. This research stems from a B.Sc. Hon-
ours thesis written by JCM under the supervision of
RH.
Institutional Abbreviations
AMNH, American Museum of Natural History, New
York, New York; CMN, Canadian Museum of Nature,
Ottawa, Ontario; RTMP, Royal Tyrrell Museum of
Palaeontology, Drumheller, Alberta; YPM, Yale Peabody
Museum, New Haven, Connecticut.
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Received 1 November 2005
Accepted 5 June 2007
Use of Saltmarsh by Dragonflies (Odonata) in the Baie des Chaleurs
Region of Quebec and New Brunswick in Late Summer and Autumn
PAUL M. CATLING!, RAYMOND HUTCHINSON?, and PAUL M. BRUNELLE?
'170 Sanford Avenue, Ottawa, Ontario K2C OE9 Canada; e-mail: catlinp@agr.ge.ca
? 12 Ch. de la Savane, Appartment 12, Gatineau, Québec J8T 1P7 Canada; e-mail: Raymond.hutchinson@sympatico.ca
44 Hilltop Terrace, Dartmouth, Nova Scotia B2Y 3T1 Canada; e-mail: pmb@ns.sympatico.ca
Catling, Paul M., Raymond Hutchinson, and Paul M. Brunelle. 2006. Use of saltmarsh by dragonflies (Odonata) in the Baie
des Chaleurs region of Quebec and New Brunswick in late summer and autumn. Canadian Field-Naturalist |20(4):
413-420.
During late summer and autumn, in the Baie des Chaleurs region of Quebec, 18 species of adult dragonflies were recorded
during one or more visits of at least 2 hours each to 14 saltmarshes. Three species, Aeshna canadensis, Sympetrum danae
and S. internum, were present in more than half of the sites. The most abundant species was S. internum with over 100 seen
at some locations. Adults of several species, including Aeshna canadensis, A. umbrosa, Enallagma civile, E. hageni, Ischnura
verticalis, Lestes disjunctus, Libellula quadrimaculata, Sympetrum danae, S. internum and S. obtrusum, occurred in relatively
high frequencies in both Baie des Chaleurs saltmarshes and in those elsewhere in Acadia. Within Baie des Chaleurs observations
of emergence and/or presence of larvae, as well as regional abundance, were recorded for Aeshna canadensis, A. umbrosa,
Ischnura verticalis, Sympetrum costiferum, S. internum and S. vicinum. Oviposition in saltmarsh pools was recorded for Aeshna
canadensis, Enallagma civile, E, hageni, Ischnura verticalis, Lestes congener and Sympetrum danae. The saltmarsh dragon-
fly fauna of Baie des Chaleurs is significantly different from that of the rest of Acadia based on frequencies predicted from
the latter region. To a large extent this difference is a result of significantly increased use of saltmarsh habitat by adults of
six species including Lestes congener, Sympetrum danae, Aeshna canadensis, Sympetrum costiferum, Lestes disjunctus, and
Sympetrum internum (in order of decreasing significance) in Baie des Chaleurs in comparison with saltmarshes elsewhere in
Acadia. Local amelioration of salty conditions in certain saltmarshes, superimposed on regional amelioration as a result of
protection from storms and saltwater dilution in the Baie des Chaleurs estuary, may contribute to an environment where
adaptation can occur or where already tolerant species can exist. Dragonflies use saltmarsh habitat on the northeast coast of
North America more extensively than is currently documented.
Key Words: Odonata, dragonflies, saltmarsh, Baie des Chaleurs, Quebec, New Brunswick.
Dragonflies are generally not highlighted as a group
of insects that inhabits saltmarshes (e.g. Cheng 1976).
Foster and Treherne (1976) outline the difficulty of
characterizing marine insects because many species
wander in from adjacent habitats. They define marine
insects as having a “habitual requirement for some part
of the saltmarsh environment at some stage of their life
cycle.” They show Odonata species comprising 3% of
saltmarsh insect fauna which is dominated by Diptera,
Coleoptera and Hemiptera which collectively make
up 75% of the species. Corbet (1999, page194) notes
the only truly marine dragonfly is Erythrodiplax ber-
enice, but that there is also a “heterogenous assemblage
of Zygoptera and Anisoptera that occupy brackish
waters, usually of relatively low salinity compared with
seawater.”
Dragonflies have been reported to prey on saltmarsh
tabanid flies and mosquitos, both groups of aggressive
biting insects that interfere with human activities. As
a consequence, questions periodically arise as to the
extent of use of saltmarsh by dragonflies. Very little
information on this subject is available for the maritime
tegion of eastern Canada. Hutchinson and Bélanger
(1999, 2004) have reported on adults of Sympetrum
danae and Enallagma civile (respectively) using salt-
marsh in the Baie des Chaleurs region. These obser-
vations suggest that use of saltmarsh by dragonflies
may be more extensive than suspected. Preliminary
observations in the Baie des Chaleurs region of Quebec
and New Brunswick supported this view and sug-
gested the use of saltmarsh by widespread species of
dragonflies.
The objective of the work reported here was to sum-
marize information on the late summer and fall use of
saltmarsh by dragonflies in this region and to provide
a basis for further analysis of evolutionary and eco-
logical aspects of occurrence.
Methods
The study area — Baie des Chaleurs
Baie des Chaleurs is bounded on the north by the
Gaspé Peninsula and on the south by northern New
Brunswick (Figure |). Flowing into it at its far western
end is the Restigouche River, famous for fishing of
Atlantic Salmon, Salmo salar. The tides are approxi-
mately 2 m high in the western basin and seawater is
encountered in the estuary where it expands at Camp-
bellton, New Brunswick. More information on Baie des
413
414
THE CANADIAN FIELD-NATURALIST
Vol. 120
0 100 Kilometers | a
eT - : FL id a-0eeer
-| & Baie des Chaleurs locations
@ Locations outside Baie des Chaleurs
163 61
FicureE |. Region of Acadia showing saltmarshes visited in Baie des Chaleurs (triangles) and elsewhere (dots).
Chaleurs is available from Gauthier (2005), but there
is apparently little information other than this recent
reference (National Wetlands Working Group 1988).
Data gathering
Summarized below for the Baie des Chaleurs region
are our late summer and fall observations of dragon-
flies in saltmarshes as well as some from the Atlantic
Dragonfly Inventory Project (ADIP 2007°) for the same
region. These observations cover the period from 22
July to 1 October over the past decade. The period of
observation at any of these sites was at least 1-2 hours
on any particular date. A list of observations at each
site is followed by an annotated list of species and
additional summary comments. In discussing the ob-
servations, the term “frequent” indicates a species seen
often (e.g. at many sites) whereas “abundant” refers
to a relatively large number of individuals at one or
more Sites.
All of the sites noted were dominated by plant
species considered to be obligate halophytes, including
two species of Spartina (S. alterniflora and S. patens),
and Ruppia maritima was present in pools at most
sites. The presence of these and other obligate halo-
phytes indicates clearly that the sites were saltmarsh-
es subject to periodic inflow of saltwater. Voucher
specimens for the records of dragonflies for Baie des
Chaleurs and elsewhere in Acadia are at the Canadi-
an National Collection at the Agriculture and Agri-food
(AAFC) Central Experimental Farm in Ottawa (Cana-
dian National Collection CNC), and in the insect col-
lections of the Nova Scotia Museum (NSMNH), the
New Brunswick Museum (NEM) and the personal
collection of P. M. Brunelle.
Comparisons
To explore differences between dragonfly occur-
rence in Baie des Chaleurs and elsewhere on the
North Atlantic coast, the number of sites recorded for
species of dragonflies in Baie des Chaleurs was com-
pared with similar data (ADIP 2007*) for the rest of
Acadia (Canadian Maritime Provinces and the state
of Maine) outside of Baie des Chaleurs. Only records
for the period fromm 22 July until the end of October
are included in the comparison to represent late sum-
mer and autumn use of saltmarshes, and all records are
for the past decade. Number of sites where a species
was present was expressed as a percentage of total salt-
marsh sites visited. The extent to which the Baie des
Chaleurs dragonfly fauna is different was explored us-
2006
CATLING, HUTCHINSON, and BRUNELLE:
USE OF SALTMARSH BY DRAGONFLIES 415
none 2 Bcickish pool in CS at St. Omer (site 4) wbere two re of Lay Totee were seen ovipositing in
the foreground bed of floating algae and Ruppia maritima. The pool is surrounded by Scirpus spp. , Juncus gerardii
and Spartina alterniflora. Eight male Sympetrum danae patrolled the edge of this
saltwater during high spring and fall tides.
ing a Chi-square test and percentage frequency else-
where in Acadia to predict percentage frequency in Baie
des Chaleurs. The test also allows the comparison of
Chi-square values for individual species, but consider-
ing the sample size, the values are most useful to iden-
tify species contributing substantially to differences be-
tween Baie des Chaleurs and elsewhere, rather than
to indicate a specific level of significance.
Results
Sites and species present
For each site, observation dates are given followed
by species in alphabetical order and number of indi-
viduals seen in brackets. The total for each site and
the size and nature of the area surveyed are also given.
In cases where we are not the observers, the observers
are listed at the end. A total of 18 species of Odonata
was encountered in 14 saltmarshes in the Baie des
Chaleurs in late summer and autumn.
(1) Bathurst (Queen Elizabeth and Youghall streets), New
Brunswick (47.6648°N, 65.6328°W), 14 August 2005:
Aeshna canadensis (25), Enallagma hageni (40), Libellula
quadrimaculata (2), Somatochlora elongata (4), Sympetrum
25 m? pool, which is inundated by
costiferum (5), S. internum (25), S. vicinum (4). Total: 7
species. Size: The site included a brackish % acre pool.
(2) 4 km E of Pointe-a-la-Croix, Quebec (48.0520°N,
66.6528°W), 15 August 2005: Aeshna canadensis (30), Lestes
congener (4), Lestes disjunctus (2), Sympetrum costiferum
(1), S. danae (9), S. internum (200), S. obtrusum (2). Total:
7 species. Size: 50 acres traversed, including creeks, pools
and a river.
(3) Pointe-a-la-Garde, Quebec (48.0773°N, 66.5425°W), 15
August 2005: Aeshna canadensis (50), Enallagma hageni (43).
Lestes congener (10), L. disjunctus (5), Sympetrum costiferum
(1), S. danae (15), S. internum (150). Total: 7 species. Size:
10 acres traversed with many pools and creeks.
(4) Saint-Omer, Quebec (48.1087°N, 66.2514°W), 15 August
2005: Aeshna canadensis (18), Lestes congener (5), L. dis-
junctus (1), Enallagma hageni (1), Sympetrum danae (15).
Total: 5 species. Size: 3 acres with pools beside creek (Fig-
ure 2).
(5) Charlo, New Brunswick (48.0182°N, 66.3666°W), 16
August 2005: Aeshna canadensis (1), Lestes congener (3).
Sympetrum danae (1), Sympetrum internum (2). Total: 4 spe-
cies. Size: 4 acre with one small pool.
(6) Nash Creek, New Brunswick (47.9229°N, 66.0840°W),
16 August 2005: Aeshna canadensis (6), Sympetrum danae
416 THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Locations of saltmarshes visited in Baie des Chaleurs and elsewhere in Acadia over the past ten years within the
period 22 July to 1 October. The order is increasing latitude from left to right and top to bottom.
Baie des Chaleurs
47.6648, 65.6328 48.0520, 66.6528
48.0182, 66.3666 47.9229, 66.0840
48.1000, 66.1348 48.0500, 65.4833
47.64, 65.62 47.65, 65.61
Acadia outside Baie des Chaleurs
43.5694, 65.4139
44.15, 68.25
43.5702, 65.3747
44.2518, 68.3398
44.3889, 68.7667 44.4417, 68.8833
44.479, 68.013 44.5556, 67.8722
44.7194, 67.45 44.72, 67.41
45.1699, 66.3135 45.19, 66.32
46.1635, 63.8258 46.2083, 60.6333
46.6532, 64.0748 46.732, 63.9946
48.0773, 66.5425
47.9166, 66.0241
48.1333, 66.3166
44.0667, 64.6417
44.2535, 68.3406
44.45, 67.9
44.605, 67.48
44.8111, 66.9806
45.7321, 64.6765
46.4, 63.86
47.1667, 65.0493
48.1087, 66.2514
47.0657, 65.0750
47.7846, 65.0841
44.1483, 64.5106
44.35, 68.35
44.4587, 64.3215
44.6194, 67.7472
45.107, 61.7266
46.06, 62.95
46.5403, 63.56
47.1721, 65.0372
(1), S. internum (4), S. obtrusum (1). Total: 4 species. Size:
5 acres with pools and creeks.
(7) Jacquet River Marsh, New Brunswick (47.9166°N,
66.0241°W), 16 August 2005: Aeshna canadensis (30 adults
1 larva probably this species), Aeshna umbrosa (1 larva prob-
ably this species), Lestes congener (50), L. disjunctus (1),
Ischnura verticalis (200 adults, 4 larvae), Sympetrum danae
(1), S. internum (5). Total: 7 species. Size: traversed 10 acres
with pools and creeks.
(8) North of Eel River, New Brunswick (48.0657°N,
66.4000°W), 16 August 2005: Aeshna canadensis (22), A.
interrupta (3), Enallagma civile (26), Enallagma hageni
(1), E. ebrium (1), Ischnura verticalis (100 ), Lestes congener
(10 ), L. disjunctus (4 ), L. forcipatus (2), Sympetrum internum
(70), S. obtrusum (40). Total: 11 species. Size: at least SO acres
traversed with numerous creeks and pools, all brackish, at least
periodically.
(9) Carleton, Quebec (48.1000°N, 66.1348°W), 29 Septem-
ber—1 October 2004: Aeshna umbrosa (5+), Enallagma civile
(3), Sympetrum danae (7), S. internum (1), S. vicinum (1);
S. costiferum (1); 10 September 1999, Enallagma civile (2),
Sympetrum danae (4); 4 September 1995, Sympetrum danae
(3), Sympetrum sp. (1 reddish specimen). Total: 6 species.
Size: pools within an area of 3 acres.
(10) Bonaventure, Quebec (48.0500°N, 65.4833°W), 3 Sep-
tember 1995: Aeshna sp. (2), Pantala flavescens (1), Sym-
petrum costiferum (1), S. danae (10). Total: 4 species. Size:
3 acres.
(11) Nouvelle (saltmarsh of the Nouvelle River Basin), Que-
bec (48.1333°N, 66.3166°W), 4 September 1995: Lestes con-
gener (1), Sympetrum danae (1), Sympetrum sp. (1, reddish).
Total 3 species.
(12) Caraquet Bay, New Brunswick (47.7846°N, 65.0841°W)
saltmarsh, 12 August 2002: Sympetrum danae (3), Sympetrum
internum (1), Sympetrum obtrusum (1). Observer: Dwayne
L. Sabine. Total: 3 species.
(13) Daly Point Marsh, Bathurst Harbour, New Brunswick
(47.64°N, 65.62°W), 12 August 1993: Sympetrum danae
(1). Observer: Reginald P. Webster. 21 July 1999, Libellula
quadrimaculata (2), Aeshna canadensis (2). Total 3 species.
(14) Carron Point Marsh, Bathurst Harbour, New Brunswick
(47.65°N, 65.61°W), 22 July 1999: Aeshna canadensis (1),
Libellula quadrimaculata, Sympetrum danae (2), Sympetrum
obtrusum (1). Total: 4 species.
Annotated List of Species (alphabetical order)
Aeshna canadensis: at 10 sites and frequent at some, appar-
ently uses saltmarsh regularly as an adult hunting on salty
pools of the outer marshes which are influenced by tides reg-
ularly, as well as inner brackish pools; seen to oviposit in
saltmarsh pools. A larva 20 mm long that was probably (but
not definitely, due to difficulty of identifying young larvae)
referable to this species was retrieved from a periodically
brackish pool dominated by Ruppia maritima (site 7).
Aeshna interrupta lineata: only one at one site but of interest
because this subspecies exists at low frequencies in the Atlantic
provinces and is largely confined to the Gulf of St. Lawrence
area. The specimen was referred to subspecies /ineata on the
basis of uninterrupted lateral stripes. This species is reported
to tolerate saline waters (Dunkle 2000).
Aeshna umbrosa: only from two sites but five adults seen
patrolling saltmarsh ponds; at the other site two young larvae
probably referable to this species (21 and 23 mm long) were
found in a brackish pool dominated by Ruppia maritima
(site 7) — due to their small size, identification is tentative;
mature larvae of this species were found in water-filled pock-
ets within the storm splash zone of a sea cliff near Port-au-
Saumon, Quebec (Hutchinson 1999); and D. Sabine (personal
communication) found larvae in a small depression pool in
a seaside boulder within the tidal splash zone on the Fundy
shore of New Brunswick.
Enallagma civile: two sites; twenty-six captured (most re-
leased), many paired, were flying over a periodically brackish
creek in saltmarsh with mostly Ruppia maritima, also Zan-
nichellia palustris, Potamogeton pectinatus, lined by Spartina
alterniflora (site 8). Enallagma civile was first reported from
this region by Hutchinson and Bélanger (2004), and their re-
port was from a saltmarsh habitat at Carleton (site 9), where
they observed oviposition behaviour (but did not actually see
eggs). Osborn (1906) reported a tolerance of brackish water.
Enallagma ebrium: only one at one site, so likely vagrant.
Enallagma hageni: At four sites and abundant at two; 40 seen
and several ovipositing on Ruppia maritima around a brackish,
4 acre pool with dense beds of Ruppia maritima surrounded
by Spartina alterniflora, S. patens, S. pectinata, and Carex
2006
CATLING, HUTCHINSON, and BRUNELLE: USE OF SALTMARSH BY DRAGONFLIES
417
TABLE 2. Comparison of species of dragonflies recorded from Acadia outside of Baie des Chaleurs and Baie des Chaleurs
based on number (left) and percentage (right) of total saltmarsh sites visited. Acadia includes the Maritime Provinces (Nova
Scotia, New Brunswick and Prince Edward Island) of Canada and the state of Maine. Records from Acadia are from the
ADIP database (2007). Records from Baie des Chaleurs are those reported here and are also in ADIP 2007. Only records for
the period from 22 July until the end of October are included in the comparison to represent late summer and autumn use of
saltmarshes. Chi-square values for each species based on comparison of frequency elsewhere in Acadia to predict frequency
in Baie des Chaleurs are included to highlight species most different in their occurrence in these two regions in bold. Names
are taken from Catling et al. (2005).
Species
Lestes congener Hagen, Spotted Spreadwing
Lestes disjunctus Sélys, Northern Spreadwing
Lestes forcipatus Rambur, Sweetflag Spreadwing
Lestes inaequalis Walsh, Elegant Spreadwing
Lestes rectangularis Say, Slender Spreadwing
Lestes unguiculatus Hagen, Lyre-tipped Spreadwing
Enallagma aspersum (Hagen), Azure Bluet
Enallagma civile (Hagen), Familiar Bluet
Enallagma durum (Hagen), Big Bluet
Enallagma ebrium (Hagen), Marsh Bluet
Enallagma hageni (Walsh), Hagen’s Bluet
Ischnura verticalis (Say), Eastern Forktail
Aeshna canadensis Walker, Canada Darner
Aeshna constricta Say, Lance-tipped Darner
Aeshna eremita Scudder, Lake Darner
Aeshna interrupta Walker, Variable Darner
Aeshna umbrosa umbrosa Walker 1908, Shadow Darner
Anax junius (Drury), Common Green Darner
Dorocordulia lepida (Hagen in Sélys), Petite Emerald
Somatochlora elongata (Scudder), Ski-tipped Emerald
Somatochlora kennedyi Walker, Kennedy’s Emerald
Celithemis elisa (Hagen), Calico Pennant
Erythrodiplax berenice (Drury), Seaside Dragonlet
Leucorrhinia frigida Hagen, Frosted Whiteface
Libellula luctuosa Burmeister, Widow Skimmer
Libellula pulchella Drury, Twelve-spotted Skimmer
Libellula quadrimaculata Linnaeus, Four-spotted Skimmer
Pantala flavescens (Fabricius), Wandering Glider
Plathemis lydia (Drury), Common Whitetail
Sympetrum costiferum (Hagen), Saffron-winged Meadowhawk
Sympetrum danae (Sulzer), Black Meadowhawk
Sympetrum internum Montgomery, Cherry-faced Meadowhawk 1
Sympetrum obtrusum (Hagen), White-faced Meadowhawk
Sympetrum semicinctum (Say), Band-winged Meadowhawk
Sympetrum vicinum (Hagen), Autumn Meadowhawk
Acadia Chaleurs Chi-square
n= 32 n=14
| 3.) Te, 500 352.8872
SG Sau 21.4439
| 3.1 | a] 2.6130
l 3.1 - _ 1.5500
4 12.5 - = 10.0160
2 6.2 - - 4.1333
| 3.1 - 1.5500
ht Z29( 0, Dr NAS 4.1125
] 3.1 - - 1.5500
2 6.2 | Tal 0.0920
LOW S12 4 28.6 0.2125
6 18.7 2 43 0.9084
hin eels 10 =—71.4 98.2049
3 9.3 - - 6.9759
2 6.3 _ = 4.2223
Sy ISG" ] Tel 3.8409
42-5 2 143 0.1963
1 3.1 - 1.5500
2 6.2 - - 4.1333
3 9.3 Tk 0.3815
1 3.1 = 1.5500
1 3.1 = - 1.5500
8 25.0 - - 22.2420
1 Bel -- - 1.5500
1 Bel - 1.5500
2 6.2 _ - 4.1333
9 28.1 2) yAlee! 1.4495
3 9.3 l Ted 0.3815
2 6.2 - - 4.1333
3 9.3 By SY 55.8876
5) ISHS [2a S57 261.2840
22 SieD 9 64.3 17.4604
OF 28el Ss) SPH 1.8149
Sp els!6 - 13.0139
2) 6.2 — - 4.1333
paleacea, flooded by high tides in fall according to local res-
idents (site 1); 43 around pools and creeks with Ruppia mar-
itima, Zannichellia palustris, Potamogeton pectinatus and
Enteromorpha sp. (site 3).
Ischnura verticalis: abundant at each of two locations; more
than 200 seen around pools with Potamogeton pectinatus and
Ruppia maritima with Spartina pectinata fringe, some ovi-
positing, and at the the same location four larvae 17-21 mm
long referable to this species were collected from the pools
(site 7); more than 100 were seen around spring-fed pools sur-
rounded by Spartina pectinata, Spartina alterniflora, Spartina
patens and species of Scirpus (site 8). In both cases the pools
were fed by freshwater springs but were definitely subject
to saltwater during high tides.
Lestes congener: at seven sites and at least 82 individuals
overall; given the number of sites and the number of individ-
uals this species may be a resident of saltmarsh; several pairs
seen ovipositing on Juncus balticus and Juncus gerardti around
brackish pools with Potamogeton pectinatus and Ruppia mari-
tima (site 7).
Lestes disjunctus: at five sites; but not as abundant as L.
congener.
Lestes forcipatus: only one at one location.
Libellula quadrimaculata: at three sites: adults patrolled a
brackish pool at site 1; said to tolerate saline waters (Dunkle
2000).
Pantala flavescens: only one at one location and probably a
migrant.
Somatochlora elongata: only found at one location, but four
seen over pool with Ruppia maritima surrounded by Spartina
alterniflora, S. patens, S. pectinata, and Carex paleacea, flood-
418
ed by high tides in fall according to local residents (site 1).
This species has also been reported from four other salt-
marshes in Acadia (Table 2).
Sympetrum danae: at 12 sites, and abundant on the north side
of the bay, patrolling saltmarsh pools very much like Ery-
throdiplax berenice does in saltmarshes further south on the
Atlantic coast. Two females were seen ovipositing in a pool
25 m* with a thick bed of Ruppia maritima surrounded by
Scirpus spp., Juncus gerardii and Spartina alterniflora (site
4, Figure 2). The pool was said by local people to be flooded
by high tides in spring and fall. Ovipositing behaviour was
also observed in saltmarsh ponds at Carleton but eggs were
not actually seen. The species appears to be less frequent in
the saltmarshes on the south side of the bay. Hutchinson and
Bélanger (2004) have previously drawn attention to the uti-
lization of saltmarsh by adults of this species and suggested
the possibility that larvae may mature in brackish pools. High
salinity of spring and fall high tides may be avoided by the
resistant egg stage that overwinters. Larval development may
be relatively rapid (Corbet 1999) and allow use of saltmarsh
pools during periods of minimal tidal impact. Although the
larval habitat of this species is widely reported in both Europe
and North America as oligotrophic pools and small lakes on
bogs and heathlands, it also inhabits very alkaline fens in east-
ern and northern Canada. Consequently saltmarsh pools would
not be so remarkable a habitat. Interestingly in Ireland, males
are said to be non-territorial (Nelson and Thompson 2000’),
contrary to some of our observations on saltmarsh pools. In
some places within its holarctic range, S. danae is reported to
migrate (e.g. Corbet 1999, page 420). Ability to utilize brackish
habitats may be beneficial to migrants over water in the mar-
itime regions where landfall may have only brackish habitat.
Sympetrum internum: at nine sites and abundant at some.
One emerging from a pool surrounded by Spartina pectina-
ta in upper marsh (site 2) and tenerals in several sites; uses
saltmarsh extensively as an adult and the upper marsh at
least to some extent as larval habitat.
Sympetrum costiferum: at five sites; two emerging from brack-
ish 4 acre pool with dense beds of Ruppia maritima surround-
ed by Spartina alterniflora, S. patens, S. pectinata and Carex
paleacea, flooded by high tides in fall according to local resi-
dents (site 1); one emerging from sandy bottom creek with
Ruppia maritima and Enteromorpha sp. (site 3). Apparently
uses Saltmarsh habitats for development but in low numbers,
possibly due to limited amount of sandy larval habitat in most
marshes. According to Walker and Corbet (1975), S. costiferum
appears to be more tolerant of saline waters than other spe-
cies of Sympetrum, but they are not clearly referring to sodium
salts. Dunkle (2000) also refers to tolerance of saline waters.
Sympetrum obtrusum: at five sites but no strong evidence of
use of saltmarsh as larval habitat by this widespread and gen-
erally abundant species.
Sympetrum vicinum: at two locations but at one of these two
were emerging from brackish 4 acre pool with dense beds of
Ruppia maritima surrounded by Spartina alterniflora, S.
patens and S. pectinata, and Carex paleacea, flooded by high
tides in fall according to local residents (site 1).
Discussion
Comparison with saltmarshes elsewhere in Acadia
A total of 35 species has been recorded from saltmarsh-
es in Acadia (Table 2). Several occurred in relatively
THE CANADIAN FIELD-NATURALIST
Vol. 120
high frequencies in both Baie des Chaleurs saltmarshes
and saltmarshes elsewhere in Acadia suggesting some
degree of salinity tolerance. Included in this category
are Aeshna canadensis, A. umbrosa, Enallagma civile,
E. hageni, Ischnura verticalis, Lestes disjunctus, Libel-
lula quadrimaculata, Sympetrum danae, S. internum
and S. obtrusum.
With a Chi-square value of 912.707, the saltmarsh
dragonfly fauna of Baie des Chaleurs is significantly
(well below the 0.005 level) different from that of the
rest of Acadia based on frequencies predicted from
the latter region. Based on Chi-square values exceed-
ing 10.0 (significance below the 0.005 level), Baie des
Chaleurs saltmarshes differed from sites elsewhere in
having significantly more sites for six generally wide-
spread species that occur throughout Acadia (Table 2,
Donnelly 2004a, b, c) including Lestes congener, Sym-
petrum danae, Aeshna canadensis, Sympetrum costi-
ferum, Lestes disjunctus, and Sympetrum internum (in
order of decreasing significance). Three species were
significantly less frequent in Baie des Chaleurs includ-
ing Erythrodiplax berenice, Sympetrum semicinctum,
and Lestes rectangularis all of which do not extend as
far north as Baie des Chaleurs (Donnelly 2004b, c).
Erythrodiplax berenice for example does not occur
north of 45°N, whereas Baie des Chaleurs is situated
at 48°N.
General Observations
With 35 species of adult dragonflies recorded in salt-
marshes in Acadia and 18 in Baie des Chaleurs, and in
both cases, approximately half of them occurring in
more than 15% of sites, substantial use by dragonflies
is suggested. This is further supported by records of
emergence, presence of larvae, ovipositing behaviour
and relatively large numbers of individuals.
In a number of saltmarsh habitats further south on
the coast of New England, the only abundant dragon-
fly species that we have observed using the saltmarsh
habitat in late summer is the saltmarsh specialist, Ery-
throdiplax berenice. It is often present in large numbers
(to 30 per acre), the males patrolling pools, the females
along the marsh edges but pairs frequent over the marsh
in early to mid-morning. Other Odonatists have also
noted that in many saltmarshes the only dragonfly
present is E. berenice (D. Paulson, personal commu-
nication). Other species fly over the marshes includ-
ing the migratory Anax junius, Tramea spp., and Pan-
tala spp. All of these may hunt over the marsh but seem
much more often to hunt and congregate in open shrub-
by areas around the marsh edge. We have seen species
of Enallagma, Lestes and Sympetrum only occasion-
ally in these more southern saltmarshes. There may
be a trend toward more extensive use of saltmarsh by
dragonflies in a northerly direction. Possibly saltmarsh-
es further south are less utilized by widespread drag-
onfly species because salt may be more concentrated
in pools of upper marsh due to higher evaporation.
2006
Baie des Chaleurs
The fact that Baie des Chaleurs saltmarshes dif-
fered from sites elsewhere in Acadia in having very
significantly more sites for six generally widespread
species, including Lestes congener, Sympetrum danae,
Aeshna canadensis, Sympetrum costiferum, Lestes dis-
junctus, and Sympetrum internum (in order of decreas-
ing significance), suggests either that there has been
some adaptation to saltmarsh conditions by these wide-
ranging species in this region, or that saltmarshes in
this region are more readily colonized as a result of
unique characteristics, or both. Baie des Chaleurs is
home to an endemic and endangered saltmarsh butter-
fly, the Maritime Ringlet (Coenonypha tullia nipisiq-
uit, McDunnough 1939). Recent studies have sug-
gested that this butterfly has evolved physiological
adaptation to saline wetland conditions including tol-
erance of tidewater submergence for up to 24 hours
(Webster 1998; Sei 2004). The endemic but somewhat
more widespread Maritime Copper Butterfly (Lycae-
na dospassosi McDunnough) is also confined to the
saltmarshes of northern New Brunswick and Gaspé
(Thomas 1980).
Unique charaterisitics of Baie des Chaleurs salt-
marshes may be widespread or local, or both. Parts of
saltmarsh are prohibitively salty for non-adapted spe-
cies only during high tides in the spring and fall, and
even then saltwater may be somewhat diluted due to
the estuary effect. There are distinctive east and west
basins and in the west basin, where most of our obser-
vations were made and where the Maritime Ringlet
occurs, the tides are higher and salt concentrations
lower than in the east basin (Gauthier 2005). It has
been suggested that saltmarshes can be divided into
two kinds, estuarine such as Baie des Chaleurs, and
coastal (National Wetlands Working Group 1988). Al-
though the difference between the two types is not
perfectly clear, conditions of inundation and salt con-
centrations in coastal saltmarsh would be more severe.
Saltmarshes have periodic inundation in common,
but the timing and duration may vary as well as the salt
concentrations. Furthermore, occasional lack of ice
development in the bay may result in severe impact
of high tide associated with winter storms on some
saltmarshes, while others may be protected by barrier
beaches (Webster 1994", 1998). Local amelioration of
salty conditions in certain saltmarshes, superimposed
on regional amelioration, may provide an environ-
ment where tolerant species can exist and adaptation
can occur.
Tolerance of salt
Although the places where dragonflies were record-
ed in Baie des Chaleurs saltmarshes may not have
been consistently high in salt concentration, they were
definitely brackish, were part of a tidal salt marsh sub-
ject to periodic inundation with salt water and subject
to salt to the extent of excluding non-halophytic flora.
Species of dragonflies differ in their ability to cope with
CATLING, HUTCHINSON, and BRUNELLE:
USE OF SALTMARSH BY DRAGONFLIES 419
salt (Cannings and Cannings 1987). Of the 18 species
found in Baie des Chaleurs, Aeshna interrupta, Libel-
lula quadrimaculata, Sympetrum danae, and S. costi-
ferum are reported by Dunkle (2000) to tolerate saline
water. However, Walker (1953, 1958), Walker and Cor-
bet (1975), Pilon and Lagacé (1998) do not report any
of the species listed here as occurring in saltmarsh.
Observations in Baie des Chaleurs suggest that drag-
onflies occupy saltmarshes to a greater extent than is
currently documented. Ecological and evolutionary as-
pects, including extent to which local adaptation has
occurred, require more study, particularly directed to-
ward larval occurrences and comparisons with other
habitats in the surrounding landscape.
Acknowledgments
Brenda Kostiuk and Gilbert Bélanger assisted with
field work. Reginald P. Webster and Dwayne L. Sabine
contributed information to the Atlantic Dragonfly In-
ventory Project, which was used here for comparison
with Baie des Chaleurs. Robert Anderson of the Can-
adian Museum of Nature provided helpful comments.
Documents Cited
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Webster, R. P. 1994. The life history and ecology of the Mar-
itime Ringlet butterfly, Coenonympha inornata nipisiquit
McDunnough. Report for the Endangered Species Recoy-
ery Fund, World Wildlife Fund Canada and the New Bruns-
wick Department of Natural Resources and Energy, Fish
and Wildlife Branch. 18 pages
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Received 30 November 2005
Accepted 29 March 2007
First Records of the Yellow Bullhead, Ameiurus natalis,
a Loricariid Catfish, Panaque suttonorum, and a Silver Pacu,
Piaractus cf. P. brachypomus, in British Columbia
G. F. HANKE!?, M. C. E. MCNALL!, and J. RoBERTS*
'Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8W 9W2 Canada
2e-mail ghanke @royalbcmuseum.be.ca; corresponding author
‘Fish and Wildlife — Science and Allocation, Ministry of Environment, Lower Mainland Region, 10470-152 Street, Surrey,
British Columbia V3R OY3 Canada
Hanke, G. F., M. C. E. McNall, and J. Roberts. 2006. First records of the Yellow Bullhead, Ameiurus natalis, a \oricariid
catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia. Canadian Field-
Naturalist 120(4): 421-427.
In Canada, there are no native catfish west of the continental divide and until recently, the list of extant exotic catfishes in
British Columbia only included introduced Black Bullhead (Ameiurus melas) and Brown Bullhead (Ameiurus nebulosus).
We report that a single Yellow Bullhead (Ameiurus natalis) was collected from Silvyermere Lake in the Lower Fraser River
drainage. This represents the first record of the Yellow Bullhead in western Canada, and its introduction likely was accidental
with a shipment of Largemouth Bass (Micropterus salmoides) rather than dispersal from Washington. Warm, eutrophic, weedy
habitat in the Fraser Delta provides ample habitat for Yellow Bullheads and other exotic fishes. A Blue-eyed Panaque
(Panaque suttonorum), a loricariid catfish found in 1995 in Shawnigan Lake, Vancouver Island, probably represents a single,
illegally released aquarium fish, as does a large Silver Pacu (Piaractus cf. P. brachypomus), which was found in Green Lake
on Vancouver Island in 2004.
Key Words: Yellow Bullhead, Ameiurus natalis, Blue-eyed Panaque, Panaque suttonorum, Silver Pacu, Piaractus sp., exotic
introductions, British Columbia.
Most populations of native species were established
in British Columbia during and after deglaciation,
but this slow process of natural colonization and dis-
persal pales in comparison to the wave of accidental
and intentional species introductions in the province
since the arrival of Europeans. Escaped pets, inten-
tionally released pets, the illegal and intentional release
of exotic game fishes by local fishermen, intentional
telease of game fishes by government agencies, con-
taminants in poorly sorted shipments of game fish, and
northward dispersal of fishes introduced in Washing-
ton, now are the principal sources of exotic freshwa-
ter fishes in British Columbia. The pathways used to
enter the province and the dispersal potential once
present, varies with each species.
Carl and Guiguet (1957), Carl et al. (1967), Scott
and Crossman (1973), Hart (1973), Crossman (1991),
Coad (1995), Moyle (2002), and Wydoski and Whitney
(2003) detail the exotic fish species (or hybrids) known
to have been introduced into British Columbia prior
to this report (Table 1). Approximately 7.4% of the
exotic fishes in Table 1 are ornamental (including Mos-
quitofish [Gambusia affinis] and Fathead Minnows
[Pimephales promelas] introduced for mosquito con-
trol; Fathead Minnows also may be introduced illegal-
ly as live bait, or dispersed into the province via the
Peace River drainage [Smith and Lamb 1976]), 24.1%
are non-game fishes (e.g., stickleback, Redside Shin-
ers [Richardsonius balteatus]), and 68.5% are game/
food fishes and hybrids between game fish species. In
many cases, fishes that are native to the province (or
their hybrids) are legally stocked in new locations to
enhance local fisheries and satisfy anglers (e.g., Wall-
eye [Sander vitreus|, Rainbow Trout [Oncorhynchus
mykiss], West-slope Cutthroat [O. clarki lewisi], and
Pacific salmon [O. gorbuscha, O. nerka, O. keta, O.
kisutch, O. tshawytscha}), and in others, game fishes
are illegally introduced to new waterbodies by private
individuals (e.g., Northern Pike [Esox /ucius], Small-
mouth Bass [Micropterus dolomieu]) (McPhail and
Carveth 1994). While most fishes released in this pro-
vince are game fishes, it is likely that the number of
exotic aquarium fishes released in British Columbia
will grow with the burgeoning pet trade (see Padilla
and Williams 2004).
Species Accounts
Silver Pacu (Piaractus cf. PR. brachypomus) — Family
Characidae (tetras)
On 4 June 2004, a large pacu was caught by a
Federal Fisheries Officer angling in Green Lake
(~49°13'50"N, 124°03'39"W) near Nanaimo (Figure 1).
This fish likely is a Silver Pacu (Figure 2), although
initially it was thought to be a piranha (Serrasalmus
sp./Pygocentrus sp.) and subsequent media reports
called the fish a Red-bellied Pacu (a commonly used
name in the pet trade). The species identity as used in
this report cannot be verified unless the body is located
421
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Fishes introduced British Columbia according to Wydoski and Whitney (2003), Moyle (2002), Coad (1995), Crossman
(1991), Hart (1973), Scott and Crossman (1973), Carl et al. (1967), and Carl and Guiguet 1957, and Royal British Columbia
Museum (RBCM) collection records; EX = exotic introductions, NA = native populations also in BC, MM = man-made
hybrids or hybrids in nature due to stocking of exotics.
Family
Petromyzontidae
Acipenseridae
Amiidae
Hiodontidae
Clupeidae
Cyprinidae
Catostomidae
Ictaluridae
Salmonidae
Esocidae
Gasterosteidae
Gadidae
Percopsidae
Poeciliidae
Moronidae
Centrarchidae
Percidae
Cichlidae
Species
species not mentioned
Acipenser transmontanus
Amia calva
Hiodon alosoides
Alosa sapidissima
Couesius plumbeus
Platygobio gracilis
Notropis hudsonius
Pimephales promelas
Richardsonius balteatus
Tinca tinca
Cyprinus carpio
Carassius auratus
Catostomus catostomus
Catostomus commersoni
Ameiurus melas
Ameiurus nebulosus
Ictalurus punctatus
Oncorhynchus mykiss
Oncorhynchus mykiss aguabonita
Oncorhynchus clarki lewisi
Oncorhynchus clarki lewisi x O. mykiss
Oncorhynchus keta
Oncorhynchus nerka
Oncorhynchus tshawytscha
Salvelinus alpinus
Salvelinus fontinalis
Salvelinus malma
Salvelinus fontinalis x S. malma
Salvelinus fontinalis x S. confluentus
Salvelinus fontinalis x S. namaycush
Salvelinus namaycush x S. malma
Salvelinus namaycush x S. confluentus
Salmo salar
Salmo trutta
Thymallus arcticus
Coregonus clupeaformis
Esox lucius
Culaea inconstans
Gasterosteus aculeatus
Lota lota
Percopsis omiscomaycus
Gambusia affinis
Poecilia reticulata (?)
Xiphophorus helleri
Morone saxatilis
Micropterus dolomieu
Micropterus salmoides
Lepomis macrochirus
Lepomis gibbosus
Pomoxis nigromaculatus
Perca flavescens
Sander vitreus
Tilapia sp.
Common Name
White Sturgeon
Bowfin
Goldeye
American Shad
Lake Chub
Flathead Chub
Spottail Shiner
Fathead Minnow
Redside Shiner
Tench
Common Carp
Goldfish
Longnose Sucker
White Sucker
Black Bullhead
Brown Bullhead
Channel Catfish
Rainbow Trout
California Golden Trout
West-slope Cutthroat
Chum Salmon
Sockeye Salmon
Chinook Salmon
Arctic Charr
Brook Trout
Dolly Varden
Atlantic Salmon
Brown Trout
Arctic Grayling
Lake Whitefish
Northern Pike
Brook Stickleback
Threespine Stickleback
Burbot
Troutperch
Western Mosquitofish
Guppy
Swordtail
Striped Bass
Smallmouth Bass
Largemouth bass
Bluegill
Pumpkinseed
Black Crappie
Yellow Perch
Walleye
Tilapia
Status
2006
and examined in detail. Furthermore, only estimates of
size can be taken from images and notes provided by
staff of British Columbia, Ministry of Water, Lands,
and Air Protection (now Ministry of Environment). The
fish measured about 40 cm total length, and 35 cm fork
length, and had been feeding on aquatic vegetation
based on its gut contents (milfoil was the dominant item
in the gut) [M. deLaronde, personal communication,
2005].
Blue-eyed Panaque
(Panaque suttonorum) — Family Loricariidae (suck-
ermouth armoured catfishes)
The Royal British Columbia Museum fish collection
contains a single Blue-eyed Panaque (RBCM 996-
190-001) (Figure 3), which was collected in 1995 from
a ditch where Royce Road crosses Shawnigan Creek
(approximately at 48°40'05", 123°37'32") (Figure 1).
The creek leads into the north end of Shawnigan Lake,
Vancouver Island. The specimen is 21.8 cm total, and
19.2 cm fork length. This specimen originally was
misidentified as Hypostomus plecostomus when it was
received in 1996.
Yellow Bullhead
(Ameiurus natalis) — Family Ictaluridae (North
America Freshwater Catfishes)
The first Yellow Bullhead found in British Colum-
bia (Figure 4) was collected on 21 July 2005, by elec-
trofishing the southeast corner of Silvermere Lake
(49°10'01"N, 122°24'27"W) (Figures | and 5). The
Yellow Bullhead was in stained but clear water about
10 cm deep, and emerged from a clump of submerged,
algae-covered terrestrial grass which was approximate-
ly 30 cm offshore; the water in the bay was 29.5°C at
the surface. The specimen (RBCM 005-079-001) was
anaesthetized/euthanized with 2-phenoxy ethanol, prior
to fixing in 10% formaldehyde. It is possible that other
Yellow Bullheads have been caught in this lake, but
were not recognized as different from Black or Brown
Bullheads by local anglers. The specimen is 12.9 cm
total length and 10.7 cm standard length. The follow-
ing features from identification keys in Scott and Cross-
man (1973, page 589), Becker (1983, pages 143-145),
Moyle (2002, pages 85-86), and Wydoski and Whit-
ney (2003, pages 153-154), confirm the identification
of the Yellow Bullhead from Silvermere Lake: body
lacks spots or blotches; lower jaw does not project be-
yond upper jaw; distance between isthmus and lower
jaw notch is less than the distance between the lower
jaw notch and the tip of the lower jaw (see illustra-
tions in Scott and Crossman 1973, page 589); mental
barbels yellow-white without melanophores; depressed
anal fin overlaps leading edge of caudal fin; and cau-
dal fin with round trailing margin.
Yellow Bullheads are native to Atlantic and Gulf
Slope drainages from New York to Mexico, the St.
Lawrence River, the Great Lakes, and the Mississippi
River Basin from North Dakota to the Gulf of Mexi-
HANKE, MCNALL, AND ROBERTS: BULLHEAD, CATFISH, PACT
423
IN BRITISH COLUMBIA
e
4
Shawnigan Lake ae
“|
1) Panaque suttonorum
2) Piaractus cf P. brachypomus
3) Ameiurus natalis
FiGuRE |. A map showing the general location of the three new
records mentioned in this report, relative to southern
Vancouver Island and the Fraser River delta; scale
bar = 10 km.
FIGURE 2. The single Silver Pacu (Piaractus cf. P. brachypo-
mus) taken from Green Lake, Vancouver Island, 4 June
2004; surrounding parts of image (stainless-steel table-
top) cropped to simplify the illustration. Scale bar =
5 cm.
Scom
FiGuRE 3. A photograph of the preserved Panaque suttono-
rum specimen (RBCM 996-190-001) collected from
the Shawnigan Lake area, Vancouver Island; sur-
rounding parts of image (dissection tray) cropped to
simplify the illustration. Scale bar = 5 cm.
co (Scott and Crossman 1973; Lee et al. 1980), and
they have been introduced to at least 14 states outside
of their native range (Fuller et al. 1999). They also are
known from headwaters of the Hudson Bay drainage
in Minnesota but have not been found downstream in
Manitoba to date (Koel and Peterka 1994; Stewart et
al. 2001; Stewart and Watkinson 2004).
424
Discussion
Two of the three species documented in this report
are not threatening to the ecology of British Columbia,
especially since they are tropical and appear to result
from the release of single fish rather than groups, which
could form breeding populations. However, all three
species are unnecessary additions to the provincial ich-
thyofauna, and there always is a risk that exotic intro-
ductions may carry unknown pathogens which may
infect native fishes (e.g., Dove and Ernst 1998; Yama-
moto and Tagawa 2000; Font 2003).
In British Columbia, a large portion of the north-
er waterways are fast-flowing, cool to cold glacier-fed
streams and oligotrophic lakes running through mon-
tane areas, and as such, are inhospitable to warm water
adapted exotic organisms. However, many exotic spe-
cies can survive in the southwestern region for a sum-
mer or longer, and fortunately, cold meltwater from
higher elevation may limit their long-term survival
and dispersal. In some regions (e.g., the Okanagan,
the Lower Fraser River Valley, and southern Vancou-
ver Island), there are many ponds, ditches, reservoirs,
and slowly flowing low-gradient streams running
through agricultural land that may support a range of
exotic species, and in places, may be warm enough in
summer to allow tropical species to breed. The mild
climate of southwestern British Columbia already is
hospitable to a wide range of organisms presently in
the pet trade (e.g., fishes such as Lepisosteus osseus,
L. platostomus, L. oculatus, Ictalurus punctatus, Clar-
ias sp., Tanichthys albonubes, Rhodeus sp., Cyprinella
lutrensis, Carassius auratus, Cyprinus carpio [koi],
Myxocyprinus asiaticus, Misgurnus anguillicaudatus,
Gastromyzon sp., and some hillstream loaches [Homa-
lopteridae]) or fishes imported live to Canada for human
consumption (e.g., Tilapia sp., Sarotherodon sp., Ore-
ochromis sp., Channa sp., Hypophthalmichthys sp., and
Monopterus sp.). Given predictions of warming aquatic
environments in North America (Mandrak 1989; Meis-
ner 1990; Rahel et al. 1996; Eaton and Scheller 1996;
Schindler 2001; Stefan et al. 2001; Jackson and Man-
drak 2002; Vander Zanden et al. 2004) it is likely many
more exotics could become established in the water-
ways of this province; those that already are present in
Canada likely will expand their range northward where
habitat and water flow permits.
Characid Fishes
The large size of most pacu and related “silver dol-
lar” species limits their suitability for home aquaria,
and likely is the reason why they are occasionally re-
leased in North America. Fuller et al. (1999) discuss
a variety of pacu and silver dollars (Colossoma sp.,
Metynnis sp., Myleus sp., Piaractus sp.), which have
been found outside of captivity in many of the United
States including Washington. Coad (1995, page 11)
noted that a specimen of Colossoma bidens was found
in Ontario (although the location was not given), and
Renaud and Phelps (2001) mention another pacu
THE CANADIAN FIELD-NATURALIST
Vol. 120
which was found in the Rideau Canal, Ontario. It is
not known if there are any wild/naturalized reproduc-
ing populations of pacu and silver dollars in North
America (Fuller et al. 1999).
Apparently the Silver Pacu found in Green Lake
on Vancouver Island was the second pacu to be col-
lected in British Columbia; the first was taken from
the Lower Mainland a few years earlier (R. Dolighan,
personal communication, 2005), but no detailed reports
or literature references to this fish exist. It has been
suggested that the Silver Pacu from Green Lake was
from a large display aquarium in a local restaurant. The
other pacu in the display aquarium also may have been
released when the restaurant closed down, but attempts
to locate other specimens in Green Lake were unsuc-
cessful. The intentional release of large, unwanted
aquarium pets (including large piranha [probably Py-
gocentrus nattereri| which may have been released
into Langford Lake on Vancouver Island [T. Duce, per-
sonal communication, 2005]) is a continuing threat to
British Columbia’s waterways, and a threat which is
difficult if not impossible to control. Piranhas such as
P. nattereri have been found in the wild as far north
as Minnesota, Idaho, and the Columbia River in Wash-
ington (Fuller et al. 1999). Fortunately pacu and pira-
nha usually are kept in small numbers or singly, and
so any released specimens are unlikely to find mates
and reproduce, even if they survive in British Colum-
bia’s waters.
Loricariid Catfishes
Some loricariid fishes (“‘plecostomus”) can tolerate
cool water aquaria and commonly are kept with gold-
fish in home aquaria, as are weatherloaches (Misgur-
nus anguillicaudatus), and surprisingly, it is the expen-
sive Panaque species, not the inexpensive Hypostomus,
which have been found in the wild in Canada. In addi-
tion to the single Blue-eyed Panaque from British
Columbia, a single specimen of the Royal Panaque or
Royal “Pleco” (Panaque nigrolineatus), is known from
southern Ontario (Coad 1995, page 11). Both the Royal
Panaque and the Blue-eyed Panaque are native to South
America and are unlikely to survive long in Canada’s
present climate, unless they are released downstream of
hotsprings or near warm industrial effluent (the same
can be said for most tropical fishes, including pacu).
Illegally released tropical aquarium fishes persist in
Alberta downstream of hotsprings despite cold win-
ters in that province (Nelson and Paetz 1992), and the
possibility of tropical fish introductions is a persistent
threat to hotsprings in British Columbia.
Ictalurid Catfishes
Washington state waterways contain several ictalurid
species, including: Blue Catfish (/ctalurus furcatus),
Channel Catfish (/. punctatus), Tadpole Madtom (Notu-
rus gyrinus), Flathead Catfish (Pylodictis olivaris),
White Catfish (Ameiurus catus), Yellow Bullhead (A.
natalis), Black Bullhead (A. melas), and Brown Bull-
2006
head (A. nebulosus) (Wydoski and Whitney 2003).
None of these fishes are native to the state and exhibit
varying degrees of success in Pacific slope drainages.
Black, Brown, and Yellow Bullhead catfishes now are
found all over Washington (Wydoski and Whitney
2003), but since casual observers find the three species
difficult to differentiate, it is likely that the true range
of each is underestimated.
Carl et al. (1967), Scott and Crossman (1973),
McPhail and Lindsey (1986), McPhail and Carveth
(1994), and Coad (1995) detail the distribution of cat-
fishes in British Columbia and until this year, re-
searchers thought that only the Black Bullhead and
Brown Bullhead existed in this province. As in Wash-
ington, the range of Black and Brown bullheads like-
ly is underestimated because of the difficulty of iden-
tification, and limited survey sampling in appropriate
habitat.
Yellow Bullheads are hardy, and given their pres-
ence in the Columbia River system and the Olympic
Peninsula in Washington (Wydoski and Whitney 2003),
it was only a matter to time before they appeared in
British Columbia. It is possible that Yellow Bullheads
were introduced to British Columbia as a contaminant
in an unsorted or poorly sorted shipment of Large-
mouth Bass. The electrofishing sample from Silver-
mere Lake taken on 21 July 2005 contained the single
Yellow Bullhead, | Brown Bullhead, 5 Pumpkinseeds
(Lepomis gibbosus), 28 Prickly Sculpins (Cottus asper),
30 Black Crappie (Pomoxis nigromaculatus), and 57
Largemouth Bass (Micropterus salmoides). Note that
only one species (Cottus asper) in the six caught in
Silvermere Lake is native to the Fraser River drainage,
and that single native species made up only 23% of
the catch.
The date of introduction of Brown Bullheads into
British Columbia is unknown, but they were stocked in
Washington from 1874 onwards, and those on Van-
couver Island date back to 1906 (Carl and Guiguet
1957; Carl et al. 1967; Wydoski and Whitney 2003).
According to anecdotal evidence, the initial stocking in
Beaver Lake on Vancouver Island was accomplished
from the window of a passing train, and these few
bullheads were from an aquarium displayed in a local
restaurant (Carl and Guiguet 1957; Carl et al. 1967).
Langford Lake and other lakes were stocked soon after,
from the Beaver Lake population. It is possible that
Black Bullheads were a contaminant in the early stocks
of Brown Bullhead since the two species can be diffi-
cult to differentiate, especially when young (Becker
1983; Lever 1996). However, if present, Black Bull-
heads failed to reproduce and were extirpated while
the Brown Bullhead survived on Vancouver Island.
Bullhead catfish have not spread far in British
Columbia because the warm, eutrophic conditions they
require are found only in lakes and ponds of southern
Vancouver Island, the Fraser River Delta, and the
southern portions of the Columbia River drainage in
HANKE, MCNALL, AND ROBERTS: BULLHEAD, CATFISH, PACU IN BRITISH COLUMBIA
425
FIGURE 4. Photographs of (A) the live Yellow Bullhead
(Ameiurus natalis) just after capture, and (B) the
same specimen once preserved (RBCM 005-079-
001); ruler in (B) is in centimetres.
FiGuRE 5. The southeast bay on Silvermere Lake just north
of Highway 7 (A) and detail of shoreline habitat (B)
where the Yellow Bullhead was collected.
this province. Where they have been introduced, bull-
heads are reputed to be effective nest predators, and
have been implicated in the elimination of Three-
spine Stickleback (Gasterosteus aculeatus) popula-
tions in smaller lakes along the coast (McPhail and
Lindsey 1986). Male stickleback cannot repel intrud-
ing adult bullheads, and in one lake near Mission, a
large population of sticklebacks went extinct only two
years after bullheads were introduced (McPhail and
Lindsey 1986; Wydoski and Whitney 2003). The addi-
tion of the Yellow Bullhead (Ameiurus natalis) to
426
British Columbia’s aquatic biota is not to be celebrat-
ed, and probably represents one more case of a con-
taminant species accidentally released with a transplant
of game fish.
Acknowledgments
Initial information on the pacu from Green lake
was provided by Craig Wightman, Fish Biologist, and
Randy Dolighan, Ecosystem Biologist, both with the
Ministry of Water, Land and Air Protection, Environ-
mental Stewardship Division, 2080-A Labieux Road,
Nanaimo V9T 6J9. Mike deLaronde, Conservation
Officer, Industrial Investigations, 2080-A Labieux
Road, Nanaimo V9T 6J9, provided the photograph of
the pacu from Green Lake. Thanks to Kelly Sendall
for information on the Panaque in RBCM collections
and changing the data record to reflect the new and cor-
rect species identity. Financial support for field work
in 2005 was provided by the Royal British Columbia
Museum. Ken Stewart (University of Manitoba, Depart-
ment of Zoology), and James Cosgrove (Royal BC
Museum, Department of Natural History), and two
anonymous individuals reviewed and greatly improved
an earlier, rushed draft of this manuscript; their com-
ments and suggestions are greatly appreciated.
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427
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Received 14 November 2005
Accepted 10 April 2007
Regional Variation in Amelanchier in the Whitewood area of
Southeastern Saskatchewan and the First Saskatchewan Records
of Amelanchier sanguinea
PauL M. CATLING and G. MITRow
Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Saunders Building, Central Experimental Farm, Ottawa,
Ontario K1A 0C6 Canada; e-mail: catlingp @agr.gc.ca
Catling, Paul M., and G. Mitrow. 2006. Regional variation in Amelanchier in the Whitewood area of southeastern Saskatchewan
and the first Saskatchewan records of Amelanchier sanguinea. Canadian Field-Naturalist 120(4): 428-432.
Based on measurements and evaluation of 111 specimens from 51 localities in the Whitewood area of southeastern Saskat-
chewan, most Amelanchier alnifolia from the prairie parkland is represented by short, stoloniferous shrubs, less than 2 m tall
with 5 — 100+ stems. In the Moose Mountain area, A. alnifolia is represented by shrubs that are taller, sometimes to 7 m, not
stoloniferous and usually having fewer than five stems. Although many of these plants have the inflorescence characteristics
of A. alnifolia, some are referable to the eastern A. sanguinea, not previously reported from southeastern Saskatchewan.
Such plants appear to be confined to the wooded environment of Moose Mountain. Amelanchier alnifolia is not as clearly
differentiated by habit or leaf tip characteristics as the literature suggests. Inflorescence characteristics, habit and actual
genetic variation may be associated with biogeographic patterns, especially historically wooded islands within the prairie
ecozone. These patterns are an important consideration in classification studies and in protecting and utilizing Amelanchier
germplasm.
Key Words: Juneberry, Serviceberry, Shadbush, Saskatoon, Amelanchier alnifolia, A. sanguinea, status, biogeophaphic vari-
ation, hybrid, agamospermy, Moose Mountain, Saskatchewan.
At least three species of Amelanchier are known to
occur in the prairie region of Canada. Amelanchier alni-
folia (Nutt.) Nutt. is widespread, Amelanchier florida
Lindl. is confined to ravines in the Cypress Hills (re-
ported by some authors and not by others) and A. san-
guinea (Pursh) DC. occurs in southeastern Manitoba
(e.g. Boivin 1967; Looman 1979). Reports of A. san-
guinea from the northern Great Plains (Stephenson
1973; McGregor et al. 1977) are based on A. humilis
Wieg. (McGregor et al. 1986), whereas Boivin’s (1967)
report of A. sanguinea in southeastern Manitoba may
be based on material of both A. sanguinea and A. hu-
milis since he combined the latter with the former and
both occur in the general region and in northern Min-
nesota (Nielsen 1939; Jones 1946).
In general, flowering material of A. sanguinea 1s dis-
tinguished from A. alnifolia by having flowering pedi-
cels more than 15 mm long instead of less than 10 mm
long (Boivin 1967; Cinq-Mars 1971; Gleason and
Cronquist 1991) and petals 11-22 mm long instead of
6-10 mm long (e.g., Jones 1946). In addition, A. san-
guinea is characterized as a tree with oblong, pointed
leaves and A. alnifolia is generally characterized as a
stoloniferous shrub with truncate or rounded leaves
(e.g., Nielsen 1939; Jones 1946; Fernald 1950). Ame-
lanchier humilis is distinguished from A. alnifolia by
its acute or obtuse instead of truncate or broadly round-
ed leaves (Cinqg-Mars 1971; McGregor et al. 1986).
McGregor et al. (1986) have indicated that it is diffi-
cult to separate A. alnifolia and A. humilis in the north-
ern Great Plains and they have suggested that some
colonies “appear as hybrid swarms and the two could
easily be considered as variants of one species.” The
concept of including A. humilis with A. alnifolia is
not new since it was the basis for McKay’s (1973,
1975, 1976) transfer of Nielsen’s A. humilis var. com-
pacta to A. alnifolia (A. alnifolia var. compacta (Niel-
sen) McKay).
During a survey of Amelanchier in the Whitewood
and Moose Mountain areas of southeastern Saskatche-
wan in early June 2005, it appeared that plants in the
prairie-parkland region (ecodistricts 752, 756, 760,
Agriculture and Agri-Food Canada 1995) differed from
those on the elevated Moose Mountain plateau (eco-
district 761) which was forested in presettlement times
(Catling and Kostiuk 2006). This rolling plateau, al-
most 400 km? in extent with numerous lakes and well-
drained wooded slopes with aspen (Populus) and birch
(Betula) forest, is 100 — 150 m above the surrounding
plains.
The objective of the work reported here was to doc-
ument the variation and to provide a context for the
first report of Amelanchier sanguinea in Saskatchewan.
This information is relevant to systematic studies of
Amelanchier and is also of interest with regard to the :
protection of native Amelanchier germplasm. Although
Saskatoon (Amelanchier alnifolia) is one of Canada’s
fastest growing native crops, its variation is not well
understood and its domestication has yet to take advan-
tage of the full range of variability available (Catling
and Small 2003).
428
2006
103° 00°
CATLING AND MITROW: AMELANCHIER IN SASKATCHEWAN
429
50° 00°
{ w 49° 30°
102° 00°
FiGurE |. Saskatchewan (left) showing location of Whitewood study area (right). Collection localities of Amelanchier used
in the study are indicated by dots. The Moose Mountain ecodistrict is shaded. The collection site of A. sanguinea is
indicated by “S” and dashed line connected to the dot.
Methods .
In June 2005, during late flowering, 111 specimens
of Amelanchier were collected and measured from
51 sites in the Whitewood — Moose Mountain area of
southeastern Saskatchewan (Figure 1). No more than
five specimens were collected at each site and sites are
defined as at least 0.1 km apart. Specimens were select-
ed to represent the extremes of variation present at a
site. For each specimen, lengths of lowest pedicel in an
inflorescence of average size, average petal length in
lowest flower in the inflorescence, average sepal length
in lowest flower in the inflorescence, the plant habit
(stoloniferous or not, height, number of stems), leaf
pubescence (scale 1-4, 1 = glabrous, 4 = densely pubes-
cent) and shape of the tip of a larger leaf (scale 1-5 with
| = truncate, 5 = pointed). Stoloniferous shrubs were
defined as having more than 6 separated stems and
being less than 2 m in height. Non-stoloniferous shrubs
were defined as having 5 or fewer stems together and
exceeding 2 m in height.
The data were analysed using Principal Components
Analysis (PCA), regression and histograms. PCA re-
duced dimensionality of six characters (Table 1), to
two allowing for an evaluation of grouping and indi-
cation of pattern of variation. Histograms evaluated
pedicel length with respect to habitat for different eco-
districts. F-ratios based on the comparison of the mean
Square variance of the regression model to the mean
Square of the residual indicated relationship and sig-
nificance of individual characters noted above. Stat-
graphics software (www. statgraphics.com) was used
to perform all the statistical analyses.
Results and Discussion
On the PCA plot (Figure 2) that reduces the dimen-
sionality of six characters, the first two components
account for 32.53% and 25.75% of the sample varia-
tion and the third and fourth account for 16.90% and
11.68%, respectively. On the first component, pedicel
length, sepal length and height are heavily weighted
whereas on the second component, leaf tip and leaf
shape are heavily weighted. Leaf pubescence was not
heavily weighted on either of the first two components.
With a cumulative 58.28% of the sample variation
accounted for in the diagram, the plot is marginally
useful as an indication of potential groups, and groups
are unclear. However, the plants from historically wood-
ed Moose Mountain area occupy a specific part of the
dispersion suggesting a different pattern of morpho-
logical variation from that represented elsewhere in
the region.
The plants from Moose Mountain tend to have longer
pedicels, and they are mostly non-stoloniferous (Fig-
ure 3). Variation in leaf tip shape, overall leaf shape,
leaf pubescence and the inflorescence characters is
extensive. Most plants on Moose Mountain that are
presumably A. sanguinea on the basis of height and
non-stoloniferous habit have to be treated as A. alni-
430
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°
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° 20 e
9 ©
= i o 2 roe)
R fo) Go %
°
= oo oem
o oO oe
nN eAlihs coe oe °,
= 9
7 = o° od 3 OU
= fo} oe) , Cy 0
a fo) o 02a Ce ee er) e
aeie I oP i
O nics ogo %e
°
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3 2
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Component 1 (32.53%)
FIGURE 2. Plot of 111 specimens used in the study on the first
two principal components derived from correllation
among six morphological characters. The specimens
from Moose Mountain are shown with solid dots and
these occupy a specific portion of the dispersion sug-
gesting distinctive variation within this region.
folia on the basis of pedicels less than 15 mm long
and petals less than 10 mm long. The ten plants poten-
tially referable to A. sanguinea on the basis of pedicels
over 15 mm (Figure 3) had leaves with variable tips.
For example five shrubs had more or less truncate
leaves, four had most leaves broadly rounded and two
had many pointed leaves. Leaf shapes also varied.
Eight of these are moderately pubescent but two are
more or less glabrous. All of these ten had petals over
9 mm but under 12 mm long. The two specimens ac-
cepted as A. sanguinea (e.g., Figure 4 left) had pedicels
17 mm, petals 11-12 mm, pointed, moderately pubes-
cent, oblong leaves. These two plants were distinctive
in blooming later than other plants nearby that pos-
sessed fewer A. sanguinea characteristics. The speci-
men label data for these are: SASKATCHEWAN: Moose
Mountain: White Bear Lake, 49.7550°N, 102.2565°W,
8 June 2005, P. M. Catling 05-27, B. Kostiuk (DAO);
Good Bird Point, White Bear Lake, 49.7639°N,
102.2328°W, 6 June 2005, P. M. Catling 05-16, B.
Kostiuk (DAO). Amelanchier sanguinea has not been
reported previously from Saskatchewan (Harms 2003).
Amelanchier alnifolia from the prairie parkland is
represented mostly by short, stoloniferous shrubs, less
than 2 m tall with 5—100+ stems, but taller, non-stolonif-
erous shrubs also occur in poplar groves. The smaller
stoloniferous plants often have distinctively short and
dense racemes with pedicels little if at all longer than
the petals (Figure 4, right). The leaves may be truncate
or more or less pointed.
The regressions indicated that pedicel length had the
most significant relationships with other characteris-
tics and was significantly correlated with ecodistrict
(Table 1). Sepal length was also significantly correlat-
THE CANADIAN FIELD-NATURALIST
Vol. 120
=
o
=a NO © Fa DN © ©
stoloniferous
non-stoloniferous
=a O&O
Prairie Region
Number of Specimens
stoloniferous
non-stoloniferous
Oy os C5 hy ey fy oes
Moose Mountain i
Mindi ilk anise
6 SU One omen 4G
We AD ee!
Pedicel length (mm)
FIGuRE 3. Histograms showing pedicel lengths and growth
habits for specimens of Amelanchier from the prairie
region and the Moose Mountain region of southeastern
Saskatchewan.
ed with other characters, but was not significantly asso-
ciated with leaf tip which is regarded as a significant
character for distinguishing A. alnifolia by some authors
(see above). In overall addition of significance values,
pedicel length was three time more significant than
sepal length. Futhermore, sepals are more often dam-
aged or shrunken (as in fruit) whereas pedicels are
more often intact. This represents the first quantitative
support for the use of pedicel length in defining taxa.
Leaf shape was only highly correlated with leaf tip and
was not significantly related to ecodistrict suggesting
limited use in defining taxa. These suggestions are
based on the operational concept of taxa being defined
by a number of correlated morphological characters and
by some ecological discreteness. Thus the more recent-
ly defined (see above) characters are supported and
these are included in the following key which will dif-
ferentiate the four species known from the prairie
provinces.
la. Pedicels short, less than 10 mm long; petals
4-10 mm long; mostly stoloniferous shrubs
2a. Leaves glabrous or sparsely tomentose at
flowering, truncate or broadly rounded ... .A. alnifolia
2b. Leaves persistently pubescent at flowering,
OIE TORUS go cabcoccccscuapnocoune A. humilis
1b. Pedicels longer, over 15 mm long; petals
11-22 mm long; non-stoloniferous shrubs .......... 3
3a. Leaves glabrous at flowering ............. A. florida
3b. Leaves more or less tomentose
atiflowering Wier feared antec A. sanguinea
2006
CATLING AND MITROW: AMELANCHIER IN SASKATCHEWAN 43]
‘Reserva Brasil, Foz do Iguacu 85851-970 Brazil; e-mail: annesophie @reservabrasil.org.br
2Ontario Puma Foundation, Beeton, Ontario LOG 1A0 Canada
3Kouchibouguac National Park of Canada, Kouchibouguac, New Brunswick E4X 2P1 Canada
4 Associate Vice-President, Research, Laurentian University, Sudbury, Ontario P3E 2C6 Canada
°Fundy National Park of Canada, Alma, New Brunswick E4H 1B4 Canada
Bertrand, A.-S., S. Kenn, D. Gallant, E. Tremblay, L. Vasseur, and R. Wissink. 2006. MtDNA analyses on hair samples confirm
Cougar, Puma concolor, presence in southern New Brunswick, eastern Canada. Canadian Field-Naturalist 120(4):
438-442.
For the last 40 years, the presence of Cougars (Puma concolor) in eastern Canada has been highly controversial. The purpose
of this study was to collect physical evidence of Cougars using a passive detection method. Baited hair-traps combined with ©
camera-traps were installed in New Brunswick and Nova Scotia, Canada. DNA analyses on two hair samples confirmed that
the species was present in southern New Brunswick in 2003. A footprint photographed after an observation of a Cougar by
reliable observers was examined by experts and was consistent with a Cougar footprint. Additional data are required to deter-
mine the status of Cougars in the northeastern part of its historical range.
Key Words: Cougar, Puma concolor, hair-traps, camera-traps, physical evidence, mtDNA analyses, tracks, eastern Canada.
Historically, the Cougar (Puma concolor) was dis-
tributed all across the American continent, from south-
eastern Alaska to southern Argentina and Chile (Park-
er 1998). It was the most widely distributed terrestrial
mammal in the western hemisphere (Godin 1977). The
subspecies referred to as Eastern Cougar (Puma con-
color couguar) was known to occur in Nova Scotia
[NS], New Brunswick [NB], Quebec [QC], Michi-
gan, Tennessee and South Carolina (Goldman 1946).
While 32 subspecies were initially listed by Goldman
(1946), a recent DNA study by Culver et al. (2000)
showed that only six genetically distinguishable sub-
species are now believed to occur in the western hemi-
sphere, with only one for North America; 1.e., P. c.
couguar. Populations in northeastern America drasti-
cally declined at the beginning of the last century
(Parker 1998). Weaver et al. (1996) argued that con-
flicts for resources and land uses may be the main
causes of most Cougar extirpation in North America.
Previous studies across North America have indicated
that road density, urbanization, agriculture and timber
harvesting may also limit Cougar expansion (van Dyke
et al. 1986; Maehr et al. 2002; Taylor et al. 2002; Dick-
son et al. 2005).
The status of the Cougar in eastern North America
has always been a highly controversial topic (Cumber-
land and Dempsey 1994). Due to the secretive habits of
this species and the highly fragmented regions where
few scattered individuals possibly remain, solid evi-
dence (i.e., a dead animal, DNA collected from scat or
hair, a clear photograph or indisputable confirmation
of authenticated tracks) is difficult to collect, making
decision and species-at-risk designation difficult. Prov-
incially, Cougars are listed as undetermined in NS, and |
endangered in NB. At the federal level, the status of the
eastern cougar population was changed from endan-
gered to data deficient in 1998 (Scott 1998"). No reli-
able estimate of the number of Eastern Cougars has
ever been made because authentic, scientifically-based
evidence is lacking (Cumberland and Dempsey 1994).
However, numerous credible sightings and physical
evidences have since then reopened the Eastern Cougar
debate. An extensive review carried out by the Com-
mittee on the Status of Endangered Wildlife in Canada
[COSEWIC] in 1978 leaves little doubt as to the his-
torical presence of Cougars in eastern Canada (van
Zyll de Jong and van Ingen 1978"). Today, the major
concern of provincial wildlife agencies is whether the
reported animals are natives or feral escaped or re-
leased captives (Scott 1998’).
While extensive, logistically demanding fieldwork)
is usually required to search for such a wide-ranging '
species, this paper focuses on new data obtained for’
NB through non-invasive detection methods (i.e., bait-
ed hair-trap and camera-trap).
Material and Methods
Study Areas
This project was conducted in NB and NS with 12.
traps set up in three national parks, 1.e., Fundy Nation-)
al Park of Canada [FNP], Kouchibouguac National Park’
of Canada [KNP] in NB, and Cape Breton Highlands
National Park of Canada [CBHNP] in NS. Conserva-
tion units being the largest wilderness areas available,
they are more likely to shelter wide-ranging top pred-
ators than urban or agricultural lands. Two additional,
438
|
2006
er eae
Quebec
DE
me ‘ j #
; New Brunswick
| “t
F “Frederfcfon, X
Saint, ohn
’ +
p Nova
A Scotia
BERTRAND ET AL.: MTDNA ANALYSES ON HAIR CONFIRM COUGAR
439
a
bs Prince Edward Island S re
\ (3
Se
Halifax Ap N
FiGuRE 1. Baited hair-trap (n = 14) locations (triangles) in New Brunswick and Nova Scotia, eastern Canada.
traps were installed in remote areas in the Miramichi
region (northeastern NB) because sightings from reli-
able observers (biologists and trappers) were fre-
quently reported to provincial wildlife agents. All
traps were located according to the most recent credi-
ble sighting report or physical evidence (Figure 1).
Four traps were set up in FNP in forested habitats on
the rolling upland plateau. FNP (205.9 km?) is char-
acteristic of the Maritime Acadian Highlands region
and is part of the Appalachian Mountain range (Wood-
ley et al. 1998). Three other traps were installed in KNP
(239.2 km?) in northeastern NB. Two traps were in-
stalled in coniferous stands, one in a relatively remote
area near Black River in KNP’s central region, and the
other, less than | km from human habitations along the
southeastern boundary of the park. The last KNP trap
was installed in a hilly mixed wood stand 3 along the
Major Kollock creek. Finally, five traps were installed
in CBHNP (948.0 km?), the largest protected wilder-
ness area in NS, protecting 20% of northern Cape Bre-
ton. Traps were installed based on recent convincing
Cougar sightings in deciduous, mixed wood and dead
conifer forests. All trap locations had a closed canopy,
an important amount of coarse woody debris or a thick
leaf litter, and were close to water sources.
Non-invasive detection techniques
Field work began during the fall of 2003. The pas-
Sive detection methods used entail collecting hairs,
tracks, and scats in a non-invasive manner. Baited hair-
traps (Figure 2), consisting of 2-m-high posts sur-
rounded by 3 x 3 m? squared enclosures, were installed
in areas where credible sightings had recently been
reported (Figure 1). Two barbed wire strands were
stretched around the wooden posts delimiting the enclo-
sure, at 45 cm and 75 cm from the ground respectively.
This was intended to catch hair as the animal enters the
enclosure, attracted by the species-specific lure. Drip-
ping lure (i-e., Cougar urine obtained from captives,
sex unknown) was hung inside the perforated central
post and fresh lure was added every month.
In 2004, in an attempt to get photographic records
of animals entering the enclosure, we equipped each
station with a camera-trap unit which consists of either
a 35-mm or a digital camera triggered by an infrared
motion sensor. Each camera was fastened to a tree
near the scent post, at a height of 1.8 m (Figure 2). Any
motion up to 7 m away within an angle of 120 degrees
activated the camera. Traps were checked year-round
every 4-5 weeks.
DNA analyses
Samples were submitted to a sequence-based analy-
sis of 16S mitochondrial DNA (mtDNA: Johnson and
O’Brien 1997). DNA was extracted from hair samples
using QIAGEN DNeasy Tissue kits. In order to deter-
mine whether DNA samples were from felid or other
mammal species (i.e., felids, canids, cervids, 4 mustelids,
procyonids, and ursids), 300 base pairs of 16S mtDNA
were amplified using Polymerase Chain Reaction (PCR).
440,
FIGURE 2: Experimental design for Cougar detection, a passive method combining a baited hair-trap (central post and
THE CANADIAN FIELD-NATURALIST
cae "le §=enclosure
Vol. 120
ed hair-tr
my So
: q Barbed-wired
PR LAS = as
barbed-wired enclosure) protocol and a camera-trap (upper left corner, strapped on a tree).
A volume of 15uL contained 100-500 ng template
DNA, |x reaction buffer, 1.5 mmol (millimoles) MgCl,
0.3 mmol each primer, 250 mmol dNTPs and 1 U Tag
polymerase. After initial incubation at 94°C for 2 min-
utes, 40 cycles of PCR were performed at 94°C for
1 minute, 53°C for 1.5 minutes, 72°C for 1.5 minutes,
and 72°C for 10 minutes. Results were then visualized
on a 2.0% agarose gel and compared to 20 species from
the six families cited above. A second amplification
of 600 bp of a specific region of the 16S mtDNA was
then conducted according to the protocol detailed by
Mills et al. (2000). Restriction enzymes Haelll, Hpall,
and Rsal then digested PCR products at 37°C during
12 hours, and results were visualized on 2.5% agarose
gel. Since restriction enzymes produce species-specific
patterns, it was then possible to distinguish Cougar from
other felid species.
Results
Baited hair traps
A total of 207 samples was collected from hair
traps (Table 1). MtDNA analyses conducted on hair
samples collected in 2003 revealed that two samples
collected in FNP were Cougar hair. The first hair sam-
ple was found near the main road entering the park
TABLE |. Number of hair samples collected per location and per
year for 2003-2006.
Location Sampling periods
2003 2004 2005 2006
KNP 0 6 15 7
FNP 29 34 49 23
CBHNP 1 3 17 16
MIRAM —* - 6 1
Total 30 43 87 47
*Baited hair-traps and camera-traps were installed in
the Miramichi (MIRAM) area during the fall of 2004.
(i.e., Kinnie Brook), in a young Red Spruce (Picea
rubens) and Balsam Fir (Abies balsamea) stand (6 to
12 m high, canopy closure of 70%). Ground cover
mostly consisted of Sphagnum spp. and Bunchberry
(Cornus canadensis). The other hair sample was found
in the northern area of the park, along an old logging
trail (i.e., Big Dam trail) relatively overgrown with Red
Spruce, Balsam Fir and birch (Betula spp.). In this
case, trees were taller (12-20 m) but canopy closure was |
only 40-50%. A 6-m-high understory composed of
Balsam Fir, Red Spruce and White Birch (Betula papy-
rifera) allowed no ground cover other than birch leaves.
2006
BERTRAND ET AL.: MTDNA ANALYSES ON HAIR CONFIRM COUGAR 44]
FIGURE 3. Footprint found west of Blackville (GPS coordinates (NAD 1983): 20N0724671; UTM 5188486), New Brunswick,
Canada (22 September 2004). Picture provided by P. Boucher. Box (to left of footprint) diameter: 8.5 cm.
Further DNA sequencing revealed that the two hair
samples came from two distinct animals, i.e., a North
American specimen and a South American specimen.
Camera-traps were only installed in 2004, so there is
no photographic record of these two individuals. No
Cougar hair or picture were collected after the cameras
had been installed.
Additional Observation
Sets of tracks can also be used to identify species
(van Dyke and Broke 1987; Cumberland and Dempsey
1994). A footprint (Figure 3) was photographed in 2004
by three reliable observers in Blackville (southwest
of Miramichi, New Brunswick) 15 minutes after they
had seen the Cougar at 12:25 PM and watched it for
several seconds as it was crossing a dirt road 45 m from
them. This track was examined by experts and declared
not to be of a Coyote (F. Scott and P. G. Crawshaw Jr.,
personal communication).
Discussion
Cougar detection program for the Maritimes
Hair-traps and camera-traps are alternative and con-
venient non-invasive methods that are cost- and time-
efficient techniques for animal surveys (Carbone et al.
2001). After four years of permanent cougar monitor-
ing in the Maritimes, the amount of field data keeps
increasing. As suggested by Cardoza and Langlois
(2002), it would be valuable to establish a systematic
monitoring program supervised by an expert team for
the Eastern Cougar population, especially for the prov-
inces where the status of this species is indeterminate.
This would allow collection of a series of undisputable
physical evidence (Cardoza and Langlois 2002; Maehr
et al. 2003). As mentioned by others, the probability
of proving the presence of a species which has no es-
tablished viable populations in a given area is almost
nil (Broke and van Dyke 1985).
DNA hair analyses confirmed that two Cougars oc-
curred in southern NB in 2003. Additional data (e.g.,
individual genetic haplotype) are nonetheless crucial to
better understand the status of the species in this part
of its range. One of the individuals detected in Fundy
turned out to be a South American animal, recalling the
Chilean specimen shot in Abitibi, Québec, in May 1992
(Jolicoeur et al. 2006). This may indicate that a few
scattered escapees from zoos or captives released by
private owners remain (Stocek 1995). However, the
other positive hair sample caught in the Big Dam Trail
in FNP was from an animal of North American origin.
This opens up a range of possibilities. In eastern North
America, estimates of the number of escaped captives
kept increasing during the last 30 years. In Pennsyl-
vania for instance, McGinnis (1996) reported only 31
cougars held in captivity in 1979, among which four
or five eventually escaped. Then, less than 20 years
later, according to J. Seidensticker (cited in Scott
1998°*), there was an increase of 640% in the number
of licensed private Cougar owners. This dramatic in-
crease in the number of captive individuals probably
suggests many more escapees of North and South
American origins. Further DNA identifications will en-
able researchers to shed more light on this question.
The fact that cougars are efficient colonizers (Nero and
Wrigley 1977) with large home ranges (Seidenticker
et al. 1973) ensures that the debate about the poten-
tial presence of animals in eastern Canada continues.
Reported cougars may be transients, escapees or mem-
bers of a remnant population, but they could act as dis-
persers for the (re)establishment of viable populations
in eastern Canada (Scott 1998"; Maehr et al. 2002). The
442
determination of these cats’ origin should help define
the protection status they deserve.
Acknowledgments
The authors acknowledge the information provid-
ed by F. Scott, M. Mazzolli, P. G. Crawshaw Jr., and
J. Dalponte, technical assistance by P.-E. Hébert (KNP),
G. Sinclair (FNP), S. Lambert, J. Hudson and L. A.
Reeves (CBHNP). Comments by J. Tischendorf, H.
McGinnis, J. Bridgland, G. T. Campbell and two anony-
mous reviewers significantly increased the quality of
this paper. This project was supported by Parks Cana-
da — Species at Risk Program Grant to L. V.
Documents Cited (marked * in text)
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Committee on the Stauts of Endangered Wildlife in
Canada (COSEWIC), Ottawa.
van Zyll de Jong, G. G., and E. van Ingen. 1978. Status
report on eastern cougar Felis concolor couguar in Canada.
COSEWIC, Ottawa.
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Received 19 December 2005
Accepted 27 December 2006
The Leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its
North America Distribution and Additional Notes on Species Description
PETER HOVINGH
721 Second Avenue, Salt Lake City, Utah 84103 USA; e-mail: phovingh@xmission.com
Hovingh, Peter. 2006. The leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its North American distribution and
additional notes on species description. Canadian Field-Naturalist 120(4): 443-451.
The geographic range of Haemopis lateromaculata Mathers 1963 (Hirudinea: Haemopidae) is extended across North Amer-
ica. Its distribution in the coastal region of Alaska and British Columbia suggests a coastal Pleistocene refugia separate from
the populations in the lower United States and suggests that H. /ateromaculata and the Eurasian H. sanguisuga Linnaeus
1758 are sister taxa. Support of the identification and geography is based on the anatomical positions of the reproductive
organs in H. lateromaculata and H. marmorata Say 1824. The variations within these species are described, noting that no
specific variation was confined to a geographical region.
Key Words: leech, Haemopis laterormaculata, description, taxonomy, distribution, North America.
In an effort to understand the limits of leech (Hi-
rudinea) fauna distribution in western North America,
a geographical study defined by hydrological basins
was undertaken and is still in progress. The report
herein describes a taxonomic problem with Haemopis
marmorata (Say 1824) in western North America, a
species known to be difficult to distinguish by external
characteristics from H. grandis (Verrill 1874) (Sawyer
1972) and H. lateromaculata. Four populations of
Haemopis were found without teeth and these popu-
lations were initially identified as H. grandis (Hovingh
1993). Examination by dissection revealed that the
penis sheaths of these populations were H. marmorata
and that these specimens had jaws. As a result of this
identification problem, the penis sheaths of all mature
Haemopis were examined.
In examining the reproductive organs of Haemopis,
it became apparent that H. /ateromaculata specimens
in museum collections had often been misidentified as
H. marmorata. A study of Utah (Beck 1954) and Alas-
kan (Moore and Meyer 1951) Haemopis vouchers de-
posited in the United States National Museum (USNM)
contained several lots identified as H. lateromaculata.
Three lots of specimens that were collected from British
Columbia and Newfoundland, two of which were iden-
tified by J. Madill as H. lateromaculata plus one un-
identified due to poor conditions are deposited in The
Canadian Museum of Nature (CMN) in Ottawa, On-
tario, Canada. The two lots at CMN are the first obser-
vations of H. lateromaculata outside of the type local-
ities of Minnesota and Iowa (Mathers 1963). A specimen
from Michigan was utilized in a leech phylogenetic
studies (Borda and Siddall 2004).
The reproductive organs of Haemopis marmorata
and H. lateromaculata specimens collected from west-
ern North America examined are presented in this
paper. Based on accurate diagnosis of the species, the
external characteristics (pigmentation patterns) were
determined to distinguish these two species from one
another. The results herein describe the geographical
distribution of H. lateromaculata, and support some
taxonomic and paleogeographic hypotheses concern-
ing the family Haemopidae.
Methods
Specimens of Haemopis were collected during the
last 20 years from localities in western North America.
The surveys consisted of visual encounters and hand
picking from the underside of stones, logs, and aquatic
vegetation. The specimens were relaxed with 10% eth-
anol, blotted by paper towel to removed the mucous
excreted from the relaxation process, fixed with 10%
formalin in phosphate buffered saline between two
sheets of paper towels overnight, and placed in 70%
ethanol for preservation.
Identification of all large Haemopis specimens was
determined by examining the reproductive organs
through a mid-dorsal cut from the annulus with the
female gonopore posteriorly approximately 20 annuli.
If the penis sheath was not located with this examina-
tion, the dorsal cut was extended anteriorly for some
10 annuli. Teeth and jaws were examined by a mid-
ventral cut of the lower lip. Posterior sucker width, pig-
mentation, and gonopore positions were noted. Small
specimens were identified by pigmentation. Identifi-
cations followed original descriptions and taxonomic
keys (Mathers 1963; Klemm 1985; Sawyer and Shelley
1976).
The facilities and specimens of the United States
National Museum (USNM), Washington D.C. and the
Canadian Museum of Nature (CMN), Ottawa, Ontario
were utilized. The western specimens of Haemopidae
in these collections were confirmed by dissection if the
length was greater than 40 mm, and pigmentation pat-
terns and the presence of teeth were examined. The
eastern specimens of Haemopis marmorata were select-
443
+44
ed by pigmentation with dissection on random samples.
Specimens were also examined from water quality
studies in Wyoming (Wyoming Department of Water
Quality), now residing in the Albertson College of
Idaho Orma J. Smith Museum of Natural History
(ALBRCIDA), Boise, Idaho.
Taxonomic status
There is presently full acceptance of the haemopisine
species (Davies 1991; Klemm 1985; Sawyer 1986).
The higher classification of leech groupings, and leech-
es within the Annelida, is presently contentious, how-
ever. Historically, all the haemopisine species were
placed in the genus Haemopis including the original
description by Mathers (1954, 1963) of H. kingi Math-
ers 1954 and H. lateromaculata. Richardson (1969)
revised the family Hirudinidae, established the family
Haemopidae, and revised the genus classification by
introducing the genera Percymoorensis including P.
terrestris (Forbes 1890), P. marmorata, P. lateromac-
ulata, and P. kingi), Mollibdella grandis, and Bdellaro-
gatis plumbeus Moore 1912). He retained Haemopis
for the Eurasian H. sanguisuga Linnaeus 1758. Rich-
ardson (1971) suggested that Percymoorensis and
Haemopis (the Eurasian species) were associated in
the subfamily Haemopinae, and that Mollibdella and
Bdellarogatis were in the subfamily Mollibdellinae;
he also identified a Mexican species Percymoorensis
caballeroi Richardson 1971 in that paper.
Davies (1991); Davies and Govedich (2001); and
Sods (1969) accepted the generic nomenclature of
Richardson (1969) but retained the earlier familial clas-
sification of Hirudinidae. Sods (1969) expressed reser-
vation about Richardson’s classification of Hirudinidae
(including Haemopidae) based on his use of type-speci-
mens, and that “long standing genera are strongly het-
erogeneous in content and cannot any longer be sepa-
rated one from the other entirely by single factors, or
contained within single divisions in the key”. Klemm
(1995), Manoleli et al. (1998), and Sawyer (1986) re-
jected the generic nomenclatural changes of Richard-
son but accepted the family Haemopidae classification.
A new species, Haemopis septagon Sawyer and
Shelley 1976, was described from specimens collected
from North Carolina in eastern United States. In their
description Sawyer and Shelley noted that if the classi-
fication of Richardson (1969) was accepted, H. septa-
gon warranted placement in a new genus. Davies (1991)
placed this species with the genus Percymoorensis. The
uncertainty of Richardson’s classification with respect
to H. kingi and H. lateromaculata was revealed when
Richardson (1969, 1971) stated that these two species
most likely belong to a new genus. He did not exam-
ine either species but relied on Mathers (1954, 1963)
for his paper (Richardson 1969). Borda and Siddall
(2004) examined six species of Haemopis for phylo-
genetic studies involving anatomical characteristics,
12 S mitochondrial rDNA, 18S and 28S nuclear rDNA,
THE CANADIAN FIELD-NATURALIST
Vol. 120
and cytochrome c oxidase subunit I gene fragments. Of
the 24 anatomical criteria that were utilized, 23 were
common to all six species, thus not significant in deter-
mining genera or species relations. Borda and Siddall
(2004) confirmed the family Haemopidae. I maintain
use of the genus Haemopis and the family Haemopi-
dae until additional and more conclusive phylogenet-
ic studies have been reported for all the hemopisine
species, using population samples representing the
entire range of these widespread species.
Results
Distribution of Haemopis lateromaculata in
North America
The Nearctic distribution of Haemopis lateromac-
ulata is shown in Figure | and listed in Table 1. This
distribution is based on museum specimens (USNM,
ALBRCIDA, and CMN), from my (PH) collections
from the geographical studies in western North America
and from water quality studies in Wyoming (ALBR-
CIDA). Anatomical dissections distinguished H. lat-
eromaculata from H. marmorata in western North
America when the plain ventrum and plain, cream, or
yellow colored lateral margins were prominent, and
thus could be used to distinguish these two congeners
from one another. Based on this pigmentation distinc-
tion, juvenile specimens (< 40 mm) and museum spec-
imens in eastern North America H. marmorata, were
examined and those with no ventral pigmentation were
considered H. lateromaculata. Some specimens with-
out pigmentation were dissected and the species deter-
mined. The following are the locations of Haemopis
lateromaculata (underlined accession numbers refer
to museum specimens catalogued as H. marmorata or
as Haemopis). Most of these misidentified specimens
had been catalogued and accessioned into the museum
collections before the work of Mathers (1963) had been
completed and published.
Alaska: USNM 19230 Yes Bay, 20732 Afognak Is-
land, 27255 Yes Bay, 21256 Sanak Island, 43455
Revillagigedo Island, CMNA 1982-0746 Esther Island,
and PH collections from Susitna River drainage, Kenai
Peninsula, and Kodiak Island.
Western Canada: Yukon Territory: PH collection
from Pelly River drainage (Yukon River drainage);
British Columbia: USNM 1/923] North Fork Moose
River (at least two Moose Rivers occur in British Col-
umbia), and USNM 38352 “D” [Dee] Lake; CMNA
1985-0289 and CMNA 1990-0056 (identified by J.
Madill, but was not found in the collection) Fraser
River drainage; and from PH collections from the Fras-
er River drainage (CMNA 2006-0036) and Haida Gwaii
(Queen Charlotte Islands) (CMNA 2006-0037); Alber-
ta: CMNA 1978-0426 Mackenzie River drainage; and
from PH collection Peace River drainage (Mackenzie
River drainage) (CMNA 2006-0034).
Lower western United States: Idaho: from PH col-
lection in the Snake River drainage (Columbia River
2006
0 1000
Kilometers
HOVINGH: HAEMOPIS LATEROMACULATA IN NORTH AMERICA
2008
Ficure |. The distribution of Haemopis lateromaculata in North America. Locations are shown by (a) squares,
museum specimens (NMNH, CMN); (b) circles, western North America survey sites (preliminary results)
and identifications from Wyoming water quality studies; and (C) diamonds, Mathers (1963) locations.
drainage); Oregon: from PH collection in the Klamath
River drainage and the Great Basin; Wyoming: from
PH collection in the Snake River drainage (Columbia
River drainage) and Water Quality Studies in Missouri
River drainage (ALBRCIDA 61050-61052); California:
from PH collection in the Pit River drainage (Sacra-
mento River drainage); Nevada: from PH collection in
the Great Basin; Utah: USNM 38308, 39056, USNM
38624, 38627 (the latter two identified to genus), and
from PH collection in the Great Basin and Colorado
River drainage; Colorado: from PH collection in the
South Platte River drainage (Missouri River drainage);
New Mexico: USNM 50/92 Rio Grande drainage (did
not dissect) .
Eastern North America: Canada: Saskatchewan:
USNM 38353; Manitoba: USNM 38347, CMN 1985-
0292, 1985-0301; Ontario: USNM 38360, 38348,
CMN 1985-0253, 1985-259, 1985-279, 1987-0670,
1988-0036, 1988-0163; Newfoundland: CMN /982-
0654, 1985-0378 (both identified by J. Madill): lower
United States: Michigan: USNM 3833/; Missouri:
USNM 2068/; District of Columbia: USNM 30223;
Virginia: USNM 42650.
446
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE |. Distribution of Haemopis lateromaculata in North America, from the field collections in western North America and
from the Orma J. Smith Museum of Natural History (ALBRICIDA), Canadian Museum of Nature (CMNA), and the United
States Museum of Natural History (USNM). (1) In poor condition or not found in the collection. (2) At least two Moose Rivers
in British Columbia. (3) Haemopis unidentified. PH, Peter Hovingh collection work in progress.
Province or State
Alberta
Alberta
British Columbia
British Columbia
British Columbia
British Columbia
British Columbia
British Columbia
Manitoba
Manitoba
Manitoba
Newfoundland
Newfoundland
Ontario
Ontario
Ontario
Ontario
Ontario
Ontario
Ontario
Ontario
Ontario
Saskatchewan
Yukon Territory
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
Alaska
California
Colorado
District of Columbia
Idaho
Michigan
Missouri
Nevada
New Mexico
Catalogue
MNA 1978-0426
MNA 2006-0034
MNA 1985-0289
MNA 1990-0056
MNA 2006-0036
MNA 2006-0037
SNM 19231
SNM 38352
MNA 1985-0301
MNA 1985-0292
SNM 38347
MNA 1982-0654
MNA 1985-0378
MNA 1985-0259
MNA 1985-0262
MNA 1985-0253
M
M
Aeoeeoee@e
G©@
NA 1985-0279
NA 1988-0036
MNA 1988-0163
MNA 1987-0670
USNM 38348
USNM 38360
USNM 38353
CMNA 2006-0035
CMNA1982-0746
CMNA 1985-0297
CMNA 1978-0360
USNM 19230
USNM 21225
USNM 20732
USNM 21256
USNM 43455
PH 1671-1677
PH 1660
PH 1661-1664
PH 1667, 1669
PH 1665, 1666
PH 1668, 1669
PH 2029
PH (0000)
PH (0000)
USNM 30223
PH (0000)
USNM 38331
USNM 20681
USNM (PH 848)
USNM 50192
USNM (PH 852)
PH (0000)
USNM 38308
USNM 38624
USNM 38627
USNM 39056
USNM (PH 497)
USNM (PH 498)
USNM (PH 499)
USNM (PH 489)
USNM (PH 491)
USNM (PH 487)
USNM (PH 488)
USNM (PH 486)
AOeeee eee
Date
1907
Locator
Beaver Lake
Slave River drainage
150 Mile House
Cariboo District
Chilcotin River drainage(1)
Haida Gwaii
North Fork Moose River (2)
[Dee] “D” Lake
Vita
LaSalle River
Placenta Bay
St Marys Bay
Abitibi River
Missinaibi River
Ottawa River
Albany River
Thunder Bay
Rainy River
Kenora
Lake Abitibi
Lake Nipissing
Waskasov
Pelly River drainage
Esther Island
Hinchinbrook Island (3)
Chuckagof Island (3)
Lake Mc Donald, Yes Bay
Lake Mc Donald, Yes Bay
Afognak Island
Sanak Island
Revillagigedo Island
Kodiak Island
Cook Inlet
Cook Inlet
Kenai Peninsula
Kenai Peninsula
Kenai Peninsula
Cold Bay, Alaska Peninsula
Pit River
Great Plains
Easter Branch
Snake River drainage
Mecosta County
Sikeston
Ruby Marsh
Pecos River
Catlow Valley
Klamath Lake
Provo
Mapleton
Provo
Provo River
Provo River
Provo River
Sevier River
Sevier River
Weber River
Weber River
Bear River
2006
TABLE |. continued
HOVINGH: HAEMOPIS LATEROMACULATA IN NORTH AMERICA
447
Province or State Catalogue
Virginia USNM 42650
Wyoming PH (0000)
Wyoming ALBRICIDA 61050
Wyoming ALBRICIDA 61051
Wyoming ALBRICIDA 61052
Identification of leech specimens: external
characteristics
TEETH. All of the Haemopis lateromaculata con-
tained teeth (denticles). This characteristic would sep-
arate these specimens from H. plumbea and H. grandis.
Four populations (Utah, Nevada, Wyoming, Montana)
of H. marmorata were without teeth. One population
(Utah) was collected in 1941 (USNM 50/95) and again
in 1986 suggesting that the loss of denticles may be due
to environmental or genetic factors resulting from col-
onization after the dessication of Lake Bonneville after
13 000 years ago.
POSTERIOR SUCKER. Although the posterior sucker
is large (about % the width of the body) in Haemopis
lateromaculata and small (less than % the width of the
body) in H. marmorata (Klemm 1985), the state of
body relaxation was highly variable. This variability
made the relative size of the posterior sucker to body
width rather arbitrary.
PIGMENTATION. The dorsal surface of Haemopis lat-
eromaculata was patterned with darker blotches or
mottled, or large to small speckles, of frequent to rare
numbers. The lateral margins were cream or yellow
(shared with H. terrestris) and interrupted with irreg-
ular dark intrusions dorsally (a very diagnostic charac-
ter), visible from both the dorsal and ventral surface
[specimens from 14 sites: the Great Basin (8), Col-
orado River drainage (2), Great Plains (1), Columbia
River drainage (2), and the Yukon River drainage (1)];
or with the lateral margins the same shade as the ven-
trum, without cream or yellow coloring [specimens
from 21 sites: the Great Basin (2), Pacific Coast drain-
ages (2), Fraser River drainage (1), Mackenzie River
drainage (1), coastal islands and their adjacent shores
of British Columbia and Alaska (17)]. The external
characteristic that identified all of the specimens was
the plain ventral surface. Mathers (1963: page 170)
noted “with a few indistinct black and yellow blotches”
on the ventral surface. I observed one specimen with an
occasional speckle on the ventral surface.
The pigmentation of H. marmorata in western North
America ranged from heavily blotched or mottled to
heavily speckled or lightly speckled on both the dor-
sum and ventrum (61 sites), with the ventrum having
subdued pigmentation of the same pattern. If either H.
lateromaculata or H. marmorata were of the light- and/
or dark-colored immaculate phase (Klemm 1985: page
155), pigmentation could not be used to identify these
two species. The light-colored immaculate phase was
not found in the living state in western North America,
but only in preserved specimens in museums. The lack
Date
Locator
1957 Dismal Swamp
199] Snake River drainage
1994 Johnson County
1997 Crook County
1997 Niobrara County
of a dorsal stripe excluded H. terrestris, H. kingi, and
H. septagon from further consideration in this study.
MAXIMUM LENGTH (MM). Haemopis lateromaculata:
Great Basin (90), Colorado River drainage (67), Colum-
bia River drainagae (31), Mackenzie River drainage
(83), Fraser River drainage (30), Queen Charlotte Is-
lands (81), Yukon River drainage (82), Alaska (112),
and Great Plains (50). Haemopis marmorata: Great
Basin (112), Colorado River drainage (95), Columbia
River drainage (121), Pacific Coast drainages (98),
Fraser River drainage (98), and Great Plains (94).
Length is highly variable, dependent upon both the age
of living specimens and their state of relaxation dur-
ing fixation.
GONOPORE POSITIONS. Haemopis lateromaculata:
Male gonopore XI b6 anterior 4; female gonopore XII
b6 anterior 4, as determined on two specimens with
developed clitellum from Nevada and Yukon Territory.
All gonopore positions were separated by 5 annuli and
occurred on the anterior 4 of the annulus. Haemopis
marmorata: Male gonopore XI b6 anterior 4; female
gonopore XII b6 anterior 4, as determined on three
specimens with developed clitellum from Nevada and
Colorado. Most gonopore positions were separated by
5 annuli and occurred on the anterior 4 of the annulus.
Three exceptions: male XI 66 mid-annulus and female
XII 56 mid-annulus, and male XI 66 mid-annulus and
female XII b5 posterior 4 both from Arizona; and male
and female on mid-annulus from British Columbia.
Sawyer (1972) noted additional variations of the gono-
pores from mid-annulus to the furrow. The male and
female gonopore separations excluded H. septagon
from further consideration during this study.
Identification of leech specimens by reproductive organs
Figure 2 illustrates the mean measurements of the
reproductive organ position and Table 2 lists the mean
+ standard deviation. The posterior position of the left
and right ejaculatory bulbs, the posterior position of the
penis sheath loop and the vagina organs, and the ante-
rior position of the prostate organ (anterior end of male
atrium) were selected for their defined shape, largely
attributed to the muscular nature of the organs (Rich-
ardson 1969). The organ positions were determined
by vertical positioning of the organ with the annulus,
and hence counting the number of annuli to the annu-
lus with the female gonopore. There is a possible error
of annuli count of +1 annulus due to the positioning in
individual specimens. The nomenclature of the organs
follows that of Klemm (1985). Left and right positions
are determined from the dorsal side. Comparison to
other species is based on literature (Klemm 1985; Mann
1954; Richardson 1971).
PENIS SHEATH LOOP. Haemopis lateromaculata: The
penis sheath loop was located at annulus XIII b5 (XI
b6 to XIV a2). The penis sheath was generally U-
shaped and distinguished H. lateromaculata from H.
marmorata, H. plumbea, H. terrestris, H. septagon,
and H. caballeroi. Haemopis marmorata: The penis
sheath loop was located at annulus XVII b/ (XVbI to
XVIII 5S), a range comparable to Sawyer (1972). The
penis sheath was generally J-shaped and on the left side.
The ranges did not overlap between H. lateromacula-
ta and H. marmorata. The average length of the penis
sheath of H. lateromaculata was 21 annuli and of H.
marmorata 38 annuli between the male gonopore and
the prostate gland. Abnormal variations for the penis
sheath of H. lateromaculata include position on the
right side (4 specimens from Yukon Territory, Alaska,
and Colorado) and highly contorted (1 specimen from
Alaska). Abnormal variations for the penis sheath of
H. marmorata include: positioned on the right side (11
specimens from Great Basin, Colorado River Basin,
and Great Plains), and highly contorted (6 specimens
from Great Basin, Great Plains) with one specimen in
which the terminal loop turns medially and is twisted
to form a topless “8”.
PROSTATE GLAND. Haemopis lateromaculata: The
anterior end of the male atrium is covered by the pros-
tate gland and was located at annulus XI a2 (X a2 to
XII 56), positions that are anterior to the female gono-
pore and for the most part anterior to the male gono-
pore. These locations are associated with the U-shaped
penis sheath. Haemopis marmorata: The anterior end
of the prostate gland was located at annulus XIV b5
(XIII b/ to XVI 55), defining the J-shaped penis sheath.
EJACULATORY BULBS. Haemopis lateromaculata: The
posterior end of the left ejaculatory bulb (seminal vesi-
cle) was located at annulus XIII b2 (XII b2 to XIV
THE CANADIAN FIELD-NATURALIST
Vol. 120
a2) whereas the posterior end of the right ejaculatory
bulb was at annulus XIII b/ (XII b/ to XIV a2). Al-
though the locations show left-right symmetry (Figure
2), large variations were found in which the left bulb
was 10 annuli posterior to 7 annuli anterior of the right
bulb. The distribution of the positions of the ejaculato-
ry bulbs was: right anterior to the left, 13 specimens;
right even with the left, 13 specimens; and right pos-
terior to the left, 8 specimens). Haemopis marmorata:
The posterior end of the left ejaculatory bulb was locat-
ed at XIV b2 (XIII b/ to XV b5) whereas the posterior
end of the right ejaculatory bulb was at annulus XIV b/
(XII a2 to XV b6). The left bulb ranged from 9 annuli
posterior to 5 annuli anterior of the right bulb. The
distribution of the ejaculatory bulbs was: right anteri-
or to the left, 19 specimens; right even with the left,
16 specimens; and right posterior to the left, 17 spec-
imens), a pattern noted by Sawyer (1972). Two speci-
mens had only one ejaculatory bulb.
VAGINA. Haemopis lateromaculata: The posterior end
of the vagina (vaginal bulb) was found at XIV b6 (XIII
a2 to XVII a2). The vagina was always posterior to
the posterior reach of the penis sheath loop and was
generally in the median region. Haemopis marmorata:
the posterior end of the vagina was found at XVI b/
(XIV b2 to XVII a2), comparable to the findings of
Sawyer (1972). The vagina was always anterior to the
posterior reach of the penis sheath loop and posterior
to the anterior end of the prostate gland, and generally
located in the medial region. Three specimens had the
vagina location anterior to the anterior end of the pros-
tate gland. The position of the vagina of H. caballeroi
had a relative position to the penis sheath loop and the
prostate gland (atrium) as in H. marmorata, but the two
specimens of H. caballeroi were highly contracted and
hence not typical (Richardson 1971). The extensive
penis sheath and its relation to the vagina suggests
that these two species deserve a separate genus — in
TABLE 2. Annuli location of selected reproductive organs in Haemopis marmorata and H. lateromaculata. Annulus X b/
was number | and annuli posterior to this annulus were sequentially numbered. Organ nomenclature was taken from
Klemm (1985). Figure 2 illustrates the organs and the points of measurement.
H. lateromaculata
N=31
Penis sheath loop
Mean + standard deviation 18.8 + 2.6
Range 1493
Anterior prostate gland
Mean + standard deviation DIED ES
Range 2-14
Posterior left ejaculatory bulb
Mean + standard deviation 16.7 + 3.0
Range 11-23
Posterior right ejaculatory bulb
Mean + standard deviation Sys) se Stl
Range 10 — 23
Posterior vagina
Mean + standard deviation 24.6 + 4.7
Range 17-38
H. marmorata
N=55
35.8 + 4.6
25 —44
23.9 + 3.7
16 —35
2006
36
EAN ine ran er
HOVINGH: HAEMOPIS LATEROMACULATA IN NORTH AMERICA
449
FiGurE 2. Measured reproductive organs in Haemopis lateromaculata (A) and H. marmorata (B), viewed from the dorsum.
The segments are shown in Roman numerals starting with segment X and extend to segment XVIII. Each segment
has five annuli as denotated names shown on the right side of segment X. For the purposes of this paper, the first
annulus is numbered 1| and is sequenced posteriorly as noted on the left side with only the first annulus of each seg-
ment numbered (see Table 2). Male gonopore on IX b6 and female gonopore on XII 56. Denotations: Elf, posterior
reach of the left ejaculatory bulb; Ert, posterior reach of the right ejaculatory bulb; P, posterior reach of the penis
sheath loop; V, posterior reach of the vagina; and A, the prostate gland at the anterior end of the male atrium. The
vertical lines represent one standard deviation (thick lines) and total range (thin lines) (see Table 2).
this case Percymoorensis, if one follows the distinct
genera classification in Erpobdellidae of Nephelopsis,
Mooreobdella, and Erpobdella, each having morpho-
logical distinct male atriums and sperm ducts.
Discussion
Haemopis lateromaculata is now considered to be
widely distributed throughout North America; prior
to this study, its known distribution had been limited to
two states (Figure 1). The presence of this leech on the
Alaskan Peninsula and adjacent islands, the Cook Inlet
region, the Alexander Archipelago of southeast Alaska,
and Haida Gwaii (Queen Charlotte Islands) of British
Columbia suggest that these leeches occupied off-shore
coastal refugia during the Pleistocene and were sepa-
rated from those populations south of Canada’s cordil-
450
leran and continental glaciers. These northwest refugia
were limited by extensive glaciers from Haida Gwaii
to Umnak Islands (Coulter et al. 1965; Clague 1989;
Kaufman and Manley 2004) with coastal cordilleran
glaciers occurring to 12 000 years ago (Peltier 1994;
Dyke 2004; Fulton et al. 2004). While the occurrence
of small terrestrial refugia have been postulated from
Kodiak to the Queen Charlotte Islands (Heusser 1989;
Ramsey et al. 2004), present day freshwater habitats
(ponds, lakes, slow moving and meandering streams)
from which leeches have been collected add a new
dimension to the terrestrial refugia.
The Pleistocene-Holocene environments were chang-
ing due to eustasy (the sea level rising at least 120 m)
and glacio-isostatic surface adjustments involving the
advance and recession of the glaciers. Hetherington et
al. (2003, 2004) mapped this environment for the Haida
Gwaii and described a land bridge between the islands
and the British Columbia mainland that existed between
11 700 and 11 200 years ago as a result of the presence
of a forebulge from the retreating cordilleran glacier.
This process, if applied throughout the glaciated region
of coastal Alaska, may have allowed island by island
transfer of leeches from coastal Pleistocene refugia,
thus accounting for the modern day presence of H.
lateromaculata populations on the coastal northwest-
ern North America mainland (Reimchen and Byun
2005). Upstream movement of H. marmorata has been
noted (Herrmann 1970; Richardson 1942; Sawyer
1970), and active movement (to Prince Edward Island)
must have occurred by direct means, and not as pas-
sive movement by birds or other agents (Richardson
1943). It is assumed herein the cordilleran glaciers and
marine habitats prevented any leeches from continen-
tal populations colonizing the coastal habitats.
Lukin (1976) suggested that Haemopis originated in
the Nearctic and penetrated the Palaearctic from the
northeast [Siberia] when northern Asia had a warmer
climate. He further suggested that H. sanguisuga
emerged at that time and is now found across central
Eurasia to the Amur Basin and near Vladivostok in east-
ern Asia, but not in Kamchatka Peninsula. The center
of biodiversity of Haemopis is in eastern North Amer-
ica, supporting Lukin’s postulate. Only two species (H.
lateromaculata and H. marmorata) moved out of this
region to occupy most of North America, and only H.
lateromaculata has been identified in Alaska and the
coastal islands, suggesting range expansion in an earli-
er epoch before the range expansion of H. marmorata.
From the Late Cretaceous (80 million years ago) to the
middle Pliocene (3 million years ago), North America
and eastern Asia were connected between Alaska and
eastern Siberia (Repenning and Brouwers 1992; Smith
et al. 1994). For much of this time, the climate in this
region was comparable to that of the southeastern Unit-
ed States today (Bassinger 1991).
I propose that H. lateromaculata and H. sanguisuga
are sister species. Two anatomical features of H. lat-
THE CANADIAN FIELD-NATURALIST
Vol. 120
eromaculata and H. sanguisuga support this relation-
ship: (1) the vaginal bulb lies posterior to the penis
sheath loop (shared with H. grandis and H. kingi) and
(2) the prostate gland (anterior end of the male atrium)
occurs mostly anterior to the male gonopore (shared
with H. kingi and H. caeca) (Mann 1954; Klemm
1985; Manoleli et al. 1998). Note that H. kingi occurs
in both groups, a confirmation of Richardson’s (1969,
1971) views that H. kingi and H. lateromaculata are
closely related.
The distribution of a species is as important as the
original description of a species. As noted by Newton
(2003: page 90) with respect to birds, “how species are
formed is central to understanding their distributions”
and “genealogy and distribution are not separate issues:
they are simply different sides of the same biological
coin”. This paper contributes to the definition of H. lat-
eromaculata by the addition of geographical limits of
its distribution and its evolutionary implications.
Acknowledgments
I thank Jacqueline Madill and Jean-Marc Gagnon at
the Canadian Museum of Nature, Bill Moser at United
States National Museum (Smithsonian Institution), and
John Keebaugh at the Orma J. Smith Museum of Nat-
ural History for assistance and for providing facilities at
these museums which greatly contributed to this study;
Robert Wisseman (Aquatic Biology Associates) and
Kurt King (Wyoming Department of Environmental
Quality) for furnishing leeches for identification from
western United States. Early versions of this document
were reviewed by Bill Moser and Donald Klemm, and
Jacqueline Madill and Mark J. Wetzel reviewed the
manuscript for the journal. I thank these reviewers for
their suggesting and improvements.
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Received 17 January 2006
Accepted 21 April 2007
Predation on Artificial Nests of Northern Bobwhites, Colinus virginianus,
by Mammalian Mesopredators: Does the Problem-Individual Paradigm Fit?
J. B. JENNiNGs!*, M. L. KENNEDY!, A. E. Houston”, and B. D. CARVER?
‘Ecological Research Center and the Department of Biology, The University of Memphis, Memphis, Tennessee 38152 USA
?Ames Plantation, The University of Tennessee, P. O. Box 389, Grand Junction, Tennessee 38039 USA
3Department of Biology, Freed-Hardeman University, Henderson, Tennessee 38340 USA
4Department of Natural Sciences, Unit 208, Southwest Tennessee Community College, 737 Union Avenue, Memphis, Tennessee
38103 USA
Jennings, J. B., M. L. Kennedy, A. E. Houston, and B. D. Carver. 2006. Predation on artificial nests of Northern Bobwhites, Colinus
virginianus, by mammalian mesopredators: Does the problem-individual paradigm fit? Canadian Field-Naturalist 120(4):
452-456.
Using mark/recapture procedures, predation on artificial nests of Northern Bobwhites (Colinus virginianus) by mammalian
mesopredators (Raccoons, Procyon lotor; Virginia Opossums, Didelphis virginiana; and Striped Skunks, Mephitis mephitis) was
assessed in relation to the “problem-individual” paradigm. The paradigm, which is untested among mammalian mesopredators,
predicts that most predation on a prey species is by a small number of individuals repeatedly involved. By examining number
of captures and recaptures on artificial nests during non-nesting and nesting periods in 2000-2003, predation within and among
species were gauged. Results varied by species, sampling period, and year; however, predation was by a small percentage of
individuals and only within the population of Striped Skunks were individuals (2 of 49) captured on an artificial nest more
than once. Raccoons, Virginia Opossums, and Striped Skunks were responsible for 10, 2, and 12% of the loss of all nests avail-
able to predators, respectively. Based on low occurrences of individuals repeatedly involved in predation on nests, the problem-
individual paradigm was unsubstantiated. Although Raccoons, Virginia Opossums, and Striped Skunks (as individual predators)
had only moderate impact on the population of Northern Bobwhites, they had a much greater impact collectively (preying
upon 24% of all nests available). To maximize Northern Bobwhite success, the most productive management is probably
best directed toward a vertebrate guild that includes mammalian, avian, and reptilian predators, and management strategies
that use multiple techniques.
Key Words: Raccoons, Procyon lotor, Striped Skunks, Mephitis mephitis, Virginia Opossums, Didelphis virginiana, Northern
Bobwhites, Colinus virginianus, artificial nests, problem individuals, predation, Tennessee.
Due to several factors (e.g., release of predators and
reduced hunting; Young and Ruff 1982; Amarasekare
1993; Mosillo et al. 1999), populations of Raccoons
(Procyon lotor), Virginia Opossums (Didelphis virgini-
ana), and Striped Skunks (Mephitis mephitis) have in-
creased in abundance throughout much of their range.
These increases have caused widespread removal to be
attempted as one means of controlling populations of
these species (United States Department of Agriculture
2004"). Linnell et al. (1999), in discussing carnivore-
livestock conflicts, pointed out that individuals or
demographic groups within a predator population can
show different behavioral traits where only a small pro-
portion of individuals is accountable for taking sig-
nificant numbers of a prey species. They refer to the
phenomena as the “problem-individual” paradigm. Al-
though this paradigm appears to fit some larger carni-
vores (Rabinowitz 1986; Ross et al. 1997; Sacks et al.
1999; Stahl et al. 2001), the hypothesis rarely has been
tested (Linnell et al. 1999). Presently, the existence of
individuals or a demographic group within mid-sized
mammalian predators that prey upon a disproportionate
number of nests of ground-nesting birds is unknown.
Different rates of predation, by individuals or species,
could allow for targeted removal of individuals rather
than widespread removal as a means of controlling nest
predation which may have broad conservation and
management application as fewer individual predators
would need to be removed.
Mammalian mesopredators are among the known
predators of Northern Bobwhites (Colinus virginianus;
Leimgruber et al. 1994; Fenske-Crawford and Nieme
1997), and populations of this game bird have declined
throughout most of its range (Brennan 1991). However,
causes of mortality for Northern Bobwhites are not well
understood (Burger et al. 1995), and rates of predation
within and among species of mammalian predators on
Northern Bobwhites are uncertain. Additionally, mid-
sized mammalian predators have been successfully
captured and recaptured in live traps (Johnson 1970;
Baldwin et al. 2004) and have been shown to prey upon
artificial nests of Northern Bobwhites (Leimgruber et
al. 1994; Fenske-Crawford and Niemi 1997); therefore,
they make a good model for assessing the “problem-
individual” paradigm using capture/recapture proce-
dures. ;
The purpose of the present study was to examine the
“problem-individual” paradigm in light of predation by
populations of mammalian mesopredators on artificial
nests of Northern Bobwhites. Specifically, the predic-
tion was tested that predation on artificial nests of
Northern Bobwhites by Raccoons, Virginia Opossums,
452
2006
and Striped Skunks is not widespread within and
among species. This work should assist in developing
sound conservation and management programs for
Northern Bobwhites and control of mammalian meso-
predators.
Methods
This investigation was conducted on the Ames Plan-
tation (Ames; 35°4'N, 89°13'W), an agricultural exper-
iment station of The University of Tennessee and home
to the National Field Trial Championship for bird dogs,
located in Hardeman and Fayette counties, Tennessee.
Ames was approximately 7500 ha and consisted of a
mosaic of habitat types that included agricultural fields,
pastures, upland hardwoods, bottomland hardwoods,
and pine plantations. On the site, about 2000 ha were
managed extensively for Northern Bobwhites. The site
provided sufficient bird populations (autumn densities
of ca. 2-3 birds/ha; Seckinger 2004) and associated
predator suite (Leberg and Kennedy 1988; Baldwin
2003; Baldwin et al. 2004) to allow study of a ground
nesting bird. For a more detailed description of Ames
see Baldwin et al. (2004).
Two study sites (separated by 1.6 km) were sampled.
Each was 1.6 km? and was similar in habitat (frag-
mented, upland hardwood forest interspersed with early
successional fields). Because results obtained at each
site were similar, results are combined and are present-
ed here. Trapping during the non-nesting season (Octo-
ber-December 2000-2002) was conducted using Rac-
coon-size live traps (81.3 x 25.4 x 30.5 cm; Tomahawk
Live Trap Co., Tomahawk, Wisconsin, or Havahart Live
Traps, Woodstream Corporation, Lititz, Pennsylvania),
which were baited with cat food and placed in an 8 x 8
configuration with traps spaced at about 230-m inter-
vals. Sampling was conducted for 32 nights/season (for
a total of 2000 trap nights/season). Once captured, sex
and age were determined, and individuals were tagged
(National Band and Tag Company, Newport, Kentucky)
in both ears. Additionally, to confirm that individuals
remained on the study site during the nesting season,
17, 3, and 3 Raccoons; 10, 9, and 5 Virginia Opossums;
and 7, 8, and | Striped Skunks were fitted with radio-
collars in 2000, 2001, and 2002, respectively. Animals
were verified as being on the study site using a three-
element hand-held antenna and a portable receiver.
Telemetry equipment was from Advanced Telemetry
Systems, Inc., Isanti, Minnesota. All individuals were
released at the site of capture. Work during this period
provided a known number of marked individuals that
could be compared to those individuals captured dur-
ing the nesting trapping season.
Results obtained during the nesting trapping season
(June-September 2001-2003) were derived using arti-
ficial nests (10 cm wide by 6 cm deep) and baited with
two eggs of Northern Bobwhites (Hernandez et al.
2001). The nest itself, constructed from wicker, was
commercially purchased. Before placement, nests were
JENNINGS ET AL: PREDATION ON ARTIFICIAL NESTS
453
left outdoors in an undisturbed state for 5-7 days to
minimize human odors as suggested by Donalty and
Henke (2001). Predators can cue on human odors at
artificial nests (Whelan et al. 1994), but this can be min-
imized by rain or scents (Donalty and Henke 2001). To
address this concern, eggs were rinsed with distilled
water and handled with rubber gloves when placed on
nests (Small and Hunter 1988). By comparing individ-
uals of the known population (non-nesting season cap-
tures) that also were captured on artificial nests during
the nesting season, the percentage of the population
actively involved in predation on artificial nests was
determined.
Artificial nests were placed inside live traps. At each
site, 80 traps were established; 50 were in actual use
at any one time. Traps were placed either selectively
or randomly which resulted in an irregular distribu-
tion over each of the 1.6 km? study areas. Because of
this placement of traps, artificial nests occurred in a
variety of habitat types which can be typical of North-
ern Bobwhites (Stribling 1996). Traps that were with-
out nests were wired open. All traps were camouflaged
with available vegetation. Once a nest was preyed upon,
another trap (randomly or selectively placed) in anoth-
er location had an artificial nest placed inside. Traps
with a preyed-upon nest were closed and no longer
used. This provided 50 nests on each of the two sites to
be in continuous use. Traps in operation were open for
90 nights/season (for a total of 9000 trap nights/season),
and were checked daily. To minimize animals associat-
ing traps with nests, 25 pseudo-nests were established
on each site. These were made of chicken wire in the
shape and size of the actual traps used to contain artifi-
cial nests. Pseudo-nests were not baited and were ran-
domly placed in the study areas.
Results
For comparisons between nesting and non-nesting
seasons, radiotelemetry procedures confirmed that 9
Raccoons, | Virginia Opossum, and 2 Striped Skunks,
which were radiocollared during the non-nesting sea-
son (2000), were present during the nesting-trapping
season (2001). Three Raccoons collared in the non-
nesting sessions (2000) along with 1 Striped Skunk
were present on the study area for the nesting-trapping
session (2002). There also was one Raccoon and one
Striped Skunk collared during the non-nesting-trap-
ping session (2002) present on the study area for the
nesting-trapping session (2003).
Capture data derived from the two sampling periods
varied somewhat by species, year, and sampling period
(Table 1). Sampling during the non-nesting period re-
vealed the presence of all target species, with the great-
est number of individuals varying by species each year.
Sampling during the nesting period also showed pres-
ence of all targeted species during each year but in
smaller numbers and with less variation in data across
years than observed in the non-nesting period.
454 THE CANADIAN FIELD-NATURALIST Vol. 120
TABLE |. Capture data derived in a study of predation by mammalian mesopredators (Raccoons, Virginia Opossums, Striped
Skunks) on artificial nests of Northern Bobwhites during two trapping sessions (non-nesting, nesting) on the Ames Plantation
in western Tennessee 2001-2003. C = number of individuals captured; R = number of individuals recaptured at least one
time during the same trapping session of the same year; %R = percentage of marked individuals recaptured (R/C x 100); M
= number of individuals captured > 2 times; %COL = the percentage of radio-collared animals captured (number of collared
animals recaptured/total number collared x 100%); T = number of individuals captured during the non-nesting period that were
recaptured during the nesting period; %K = percentage of individuals captured during the non-nesting period that were recap-
tured during the nesting period (T/C of the Non-nesting period x 100%).
Nesting
Non-nesting Captured during
(October-December) (June—September) both seasons
Year Species C R %R M %COL GIR Gar IM Gao al %K
2001 Raccoon 41 24 3) © 18 13) OQ" @ 0 11 4 10
VirginiaOpossum 33 = 13 39 8 10 3 0 O 0 0 0 0
Striped Skunk 12 5 42 4 43 22 Ae’ 5 if 0 1 8
2002 Raccoon 19 8 42 1 0 14 0 O OSS) 0 0
Virginia Opossum 19 9 47 6 0 3 ORO 0 0 0 0
Striped Skunk DAT 2 SO 7 0 11 1 9 0 0 1 4
2003 Raccoon 38) le 45 4 0 lO OO 0 0 3 8
Virginia Opossum 89 24 27 9 20 4 0 O 0 0 1 1
Striped Skunk 46 $18 39 8 0 1610. .O 0 0 0 0
' This reflects predation by the same individual.
The total number of nests available to predators was
433. Raccoons were responsible for 43 nests preyed
upon (10% of all nests available to predators). Virginia
Opossums were responsible for predation on 10 nests
(2%), and Striped Skunks destroyed 52 nests (12% of
all nests available). Collectively, these predators preyed
upon 24% of all available nests. No significant pattern
existed in the age or sex of the predators involved in
predation on artificial nests.
Discussion
Results of this study documented that at most only
10% of a known predator population (Table 1) was
actively involved in predation on artificial nests of
Northern Bobwhites. Additionally, individual predator
species were involved in predation on artificial nests to
varying levels (Table 1). Based on these results, the
prediction that predation on artificial nests of North-
ern Bobwhites by Raccoons, Virginia Opossums, and
Striped Skunks is not widespread within and among
species, is supported, and the “problem-individual” par-
adigm is unsubstantiated. All predators assessed had
members captured multiple times during the non-nesting
season. These results were similar to those reported in
other studies of mesopredators conducted during autumn
and winter (non-nesting seasons) in the region (Ladine
1995; Baldwin 2003). Such results demonstrate that
individuals can be captured and recaptured in suitable
numbers to explore the “problem-individual” paradigm.
However, only two Striped Skunks were captured on
artificial nests more than once. Given that these indi-
viduals represented only 2 of 49 Striped Skunks cap-
tured and were only captured 2 and 3 times, respective-
ly, in the present study and that Vickery et al. (1992)
reported predation by Striped Skunks on nests of birds
was not targeted, these results do not corroborate the
“problem-individual” paradigm. Additionally, in com-
parison with larger mammalian predators, this para-
digm has been associated, generally, with much higher
levels of predation and by greater numbers of individ-
uals repeatedly involved in taking prey (Claar et al.
1986; Stander 1990; Ross et al. 1997; Sacks et al. 1999;
Stahl et al. 2001). Scalet et al. (1996) pointed out that
predator-prey interactions influence both prey and pred-
ator populations. However, based on results of the pres-
ent study, these interactions appear to be complex and
differ among mammalian taxa.
Lack of support for the “problem-individual” para-
digm among target predators could be explained by
distribution of nests of Northern Bobwhites on the land-
scape and the predators involved. Stribling (1996)
noted that nests of Northern Bobwhites are located in a
number of habitat types. Linnell et al. (1999) suggest-
ed that a scattered distribution of prey, such as the dis-
tribution of artificial nests in this study, would increase
encounter rates by predators without any search be-
havior required, and hypothesized that, under these con-
ditions, problem individuals are less likely to appear
because most individuals have opportunity to take the
target prey without developing specialized behaviors.
Raccoons, Virginia Opossums, and Striped Skunks
are omnivorous and opportunistic predators with strong
preferences for seasonally available foods (McManus
1974; Gardner 1982; Godin 1982; Kaufmann 1982;
Schwartz and Schwartz 2001). Wiens (1976) discussed
population responses to patchy environments and noted
the importance of prey density to responses of popula-
tions to patchiness. It appears likely that the habitat
usually occupied by Northern Bobwhites (forest open-
ings, open woods, fallow fields, and edges of cultivat-
ed fields; Stribling 1996) provides favorable and abun-
dant foods (e.g., insects and other invertebrates, wild
fruits, and small mammals; Merritt 1987; Schwartz and
Schwartz 2001) for mammalian predators. Given that
2006
number of individuals known alive for each species of
predator was greater on the study site during the non-
nesting period than the nesting period, as well as over-
all recapture rate and number of individuals captured
more than 2 times (Table 1), it seems that the presence
of eggs of Northern Bobwhites and Northern Bobwhites
themselves had no unusual attraction for mesopreda-
tors. Neither Raccoons nor Virginia Opossums had a
significant affinity for fields, which are managed inten-
sively for Northern Bobwhites on Ames, and, because
of the lack of individuals recaptured on artificial nests,
the problem-individual hypothesis is not supported for
these species. However, based on the number of cap-
tures in the present study, there was a distinct associ-
ation between fields and Striped Skunks, which reflects
a preference for this habitat (Nowak 1991). These habi-
tats also are good for Northern Bobwhites throughout
the year and not just during their breeding season (Stri-
bling 1996). These mesopredators apparently used the
study site and adjacent areas during the nesting peri-
od of Northern Bobwhites for foods other than those
associated with Northern Bobwhites.
Being opportunistic, mesopredators may prey upon
nests of Northern Bobwhites when they encounter them.
Because of increased protein intake during formation
and laying of eggs (Rosene 1969), eggs and Northern
Bobwhites themselves represent foods of high nutri-
tional value. However, because Northern Bobwhites
on Ames provide only a limited source of food, opti-
mal foraging is likely focused on more abundant and
easily captured food items. Eggs and birds probably
serve as a valuable supplemental food to more abundant
food items. For example, Vickery et al. (1992) reported
a positive correlation between predation by Striped
Skunks on bird eggs and nestlings when that species
was foraging for invertebrates. Such foraging patterns
could partly explain reports that birds are preyed upon
by Raccoons, Virginia Opossums, and Striped Skunks,
although their occurrence in diets of these predators is
small (McManus 1974; Kaufmann 1982; Godin 1982:
Schwartz and Schwartz 2001).
In the present investigation, there was only one year
where the most abundant species captured during the
non-nesting season was responsible for the greater num-
bers of artificial nests preyed upon during the nesting-
trapping season (Table 1). It appears that Raccoons,
Virginia Opossums, and Striped Skunks, as individual
species, had only a moderate impact on artificial nests
(Table 1). However, as a subset of the community or
mesopredator guild—a collection of species that use
common resources in similar ways (Root 1967)—they
had a much greater impact. In fact, Nicolaus (1987)
stated that Raccoons, along with other mammals and
birds, belonged to a guild of nest predators. Of artificial
nests preyed upon in this study, Raccoons, Virginia
Opossums, and Striped Skunks accounted for 10, 2,
and 12% of losses, respectively. Collectively, there was
a 24% loss of artificial nests attributable to this meso-
predator guild.
JENNINGS ET AL.: PREDATION ON ARTIFICIAL NESTS
455
Overall, the pattern of predation on artificial nests
of Northern Bobwhites is apparently driven by a larg-
er vertebrate-predator guild similar to the granivore
guild (birds, rodents, and insects) reported in deserts
of the southwestern United States (Brown and David-
son 1977). This vertebrate-predator guild, including
both mammalian and reptilian predators, also has been
documented by Staller et al. (2005). Because of the
collective predation rates on artificial nests, most pro-
ductive management (using multiple techniques; Jimi-
nez and Conover 2001) for ground-nesting birds prob-
ably is best directed toward a vertebrate guild that
includes not only mammalian predators but avian and
reptilian predators as well. Because of the dynamic
nature of community interactions (Meffe et al. 1997),
it is not likely that single actions will resolve complex
predator-prey relationships, which reflect the dynamic
nature of the ecological world. At present, interaction
among mammalian mesopredators is uncertain (Ladine
1997; Kissell and Kennedy 1992) and the impact of
large vertebrate-predator guilds on prey species remains
unclear. Future investigations are needed to more clear-
ly understand these ecological issues.
Acknowledgments
Thanks are extended to T. L. Best, S. B. Franklin, J.
W. Grubaugh, and B. A. Simco who provided a critical
review of an earlier draft of the manuscript. R. A. Bald-
win, B. D. Carver, J. R. Hisey, and R. C. Whittington
contributed to numerous aspects of the study. Gratitude
also is extended to the Ames Plantation for allowing
access to lands under their control. This investigation
was a cooperative project that included contributions
from the Hobart Ames Foundation, Tennessee Wildlife
Resources Agency, and The University of Memphis.
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Wiens. J. A. 1976. Population responses to patchy environ-
ments. Annual Review of Ecology and Systematics 7: 81-
120.
Young, B. F.,, and R. L. Ruff. 1982. Population dynamics and
movements of black bears in east central Alberta. The
Journal of Wildlife Management 46: 845-860.
Received 3 January 2006
Accepted 17 February 2007
Reactions of Narwhals, Monodon monoceros, to Killer Whale.
Orcinus orca, Attacks in the Eastern Canadian Arctic
KRISTIN L. LAtpRE!*, MADS PETER HEIDE-JORGENSEN2Z, and JACK R. ORR?
'Polar Science Center, Applied Physics Laboratory, University of Washington, 1013 NE 40" Street, Seattle, Washington 98105
USA
*Greenland Institute of Natural Resources, Box 570, DK-3900 Nuuk, Greenland
Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Canada
Laidre, Kristin, L., Mads Peter Heide-Jgrgensen, and Jack Orr. 2006. Reactions of Narwhals, Monodon monoceros, to Killer
Whale, Orcinus orca, attacks in the eastern Canadian Arctic. Canadian Field Naturalist 120(4): 457-465.
A Killer Whale attack on Narwhals was observed at Kakiak Point in Admiralty Inlet, Nunavut, Canada, in August 2005.
Behavioral responses of both Narwhals and Killer Whales were documented by direct observation. Data collected from
Narwhals instrumented with satellite-linked transmitters 5 days prior to the arrival of Killer Whales were used to examine
changes in Narwhal movement patterns (e.g., dispersal and clumping) five days before the attack, during the attack, and five
days after Killer Whales left the area. A minimum of four Narwhals were killed by 12-15 Killer Whales in a period of 6 hours.
Narwhals showed a suite of behavioral changes in the immediate presence of Killer Whales including slow, quiet movements,
travel close to the beach (<2 m from shore), use of very shallow water, and formation of tight groups at the surface. These
behavioral changes are consistent with Inuit accounts of Killer Whale attacks on Narwhals. During the attack, Narwhals dispersed
broadly, the groups were less clumped (standard deviation of inter-whale mean latitudes and longitudes), Narwhal space-use
doubled from pre-attack home ranges of 347 km? to 767 km? (kernel 50% probability), and Narwhals shifted their distribution
further south of the attack site. After the disappearance of Killer Whales, north-south dispersal of Narwhals contracted and was
similar to pre-attack levels, total space use decreased slightly (599 km), yet west-east dispersal remained high. Narwhals were
distributed significantly (P < 0.001) more broadly offshore in areas not used before the occurrence of Killer Whales. In general,
short-term reactions of Narwhals to Killer Whale presence were obvious; yet normal behavior (as observed from shore) resumed
shortly after Killer Whales left the area. Long-term (five day) Narwhal behavioral responses included increased dispersal of
Narwhal groups over large offshore areas. This is among the few reports of eyewitness Killer Whale attacks on Narwhals in
the high Arctic and is the first time changes in Narwhal behavior have been documented in response to a predation event
through the use of satellite telemetry.
Key Words: Killer Whale, Orcinus orca, Narwhal, Monodon monoceros, predation, satellite telemetry, Admiralty Inlet, Nunavut,
Canada.
Killer Whales (Orcinus orca) occur widely in Arc-
tic waters, yet their abundance, movements, site fidelity,
and specific distribution are poorly known (Forney
and Wade 2006). Generally, Killer Whales are sighted
during ice-free months and often near areas with high
densities of other marine mammals, such as summer-
ing grounds for Belugas (Delphinapterus leucas) and
Narwhals (Monodon monoceros). In the eastern Cana-
dian High Arctic, this includes eastern Lancaster Sound
and associated inlets or fjords around Baffin Island
(e.g., Eclipse Sound, Admiralty Inlet, and Prince Regent
Inlet) (Markham 1874; Reeves and Mitchell 1988).
Few eyewitness accounts are available of Killer
Whale attacks on Narwhals (Steltner at al. 1984). This
is because Killer Whales occur over large, sparsely
inhabited Arctic areas and the timing and location of
attacks are unpredictable. Thus, the majority of des-
criptions, both of Killer Whale attacks themselves and
of the response of Narwhals, come from Inuit obser-
vations (Degerbgl and Freuchen 1935; Freuchen and
Salomonsen 1961) or from scattered opportunistic re-
ports (Campbell et al. 1988). Generally these accounts
made by locals are limited in that observations extend
only as far as what can be observed from shore and
generally cover short time scales (hours). Also, Inuit
observations of Killer Whale occurrence or Killer
Whale attacks may not always be reported. There is
little information about the natural mortality of Nar-
whals and generally, Killer Whales are assumed not
to significantly depress Narwhal populations (Davis
et al. 1980). Furthermore, the extent to which Killer
Whales affect the behavior of Narwhals (e.g., move-
ment patterns) is unknown.
In August 2005, a Killer Whale attack on Narwhals
was observed from a camp operating a month-long
Narwhal satellite tagging operation in Admiralty Inlet,
Canada. Several kills of Narwhals by Killer Whales
were inferred from vigorous surface and diving activity
of Killer Whales within large oiled areas on the sur-
face and congregations of seabirds. In combination
with eyewitness observations of behavior of Narwhals
and Killer Whales, data collected from satellite tags
deployed on Narwhals up to five days before Killer
Whales arrived in the area were used to examine clus-
tering and dispersal of Narwhals prior to and follow-
ing the predation events. These are the first quantita-
tive movement data obtained from Narwhals subjected
to Killer Whale predation.
457
458 THE CANADIAN FIELD-NATURALIST
Visual Observations of Attack
Killer Whales were reported by Inuit hunters in a
camp about 3 km north of Kakiak Point (72°31'N
86°38'W), in Admiralty Inlet, Nunavut, Canada, on
19 August 2005. On 20 August, we witnessed a Killer
Whale predation event on Narwhals just offshore of
Kakiak Point. Approximately 12-15 Killer Whales
arrived in a single group at Kakiak Point at 12 noon
local time. At least several hundred Narwhals had been
seen daily in the area for at least two weeks preceding
the arrival of the Killer Whales and also on the morn-
ing of 20 August. On several occasions thousands of
Narwhals had been seen moving past Kakiak Point
over periods of 2-3 hours.
The Killer Whale group consisted of one adult male,
7-10 adult females, and several juveniles of undeter-
mined sex. Narwhals were attacked less than 1 km
from shore. At least two Narwhals were killed during
the first observed attack between 12:00-13:00 local
time. After this event, the Killer Whales left the vicin-
ity of Kakiak Point for approximately 3 hours and
returned at 16:00, when at least two more Narwhals
were killed ~0.5 km from the coast. Killer Whales were
not seen or reported again in the area until 25 August.
Active subsistence hunting for Narwhals was practiced
at several hunting camps along the shore north and
south of the research camp.
The four independent kill events occurred over a 6
hour period in daylight hours. Narwhal kills were in-
ferred from large oil/blubber slicks at the surface, con-
gregations and diving of Fulmars (Fulmarus glacialis)
in surface waters, and focal movements of Killer Whales
diving in the center of oiled areas for 15-30 minutes
at a time (Figure 1). No body parts or pieces of Nar-
whals were directly seen.
Narwhal Movements
Narwhal satellite tagging operations were ongoing
prior to the Killer Whales’ arrival at Kakiak Point. Nar-
whals were captured using nets set perpendicular to
the shoreline 50-100 m from shore (details described
in Dietz et al. 2001) and instrumented with SPOT4 and
SPOTS satellite tags made by Wildlife Computers
(Redmond, Washington). Tags recorded daily geogra-
phic positions based on Doppler shift of tag transmis-
sions, received by polar orbiting satellites through
Vol. 120
Service Argos (Harris et al. 1990). Transmitters were
attached to Narwhals on the dorsal ridge with three
6-10 mm polyethylene pins. The Narwhals were usu-
ally released after less than 30 minutes (Table 1).
Only data collected from Narwhals instrumented
prior to the Killer Whales’ arrival were used in the
analysis to control for differences in sample size (num-
ber of individuals monitored before and after the pre-
dation event). Since the Narwhals were instrumented
and released on different occasions, they were assumed
to represent different groups or pods. Average daily
geographic positions were calculated for each Narwhal
based on good quality Argos locations (location quali-
ty >=0). Average daily positions for each whale were
averaged to create inter-whale means. The standard
deviation of daily average inter-whale positions was
calculated for each of three time periods: five days
before Killer Whales arrived (14-18 August), during
the immediate Killer Whale observations or attacks
(19-20 August), and five days after the Killer Whale
attack (21-25 August) when no Killer Whales were
seen in the area. Narwhals were distributed along a
north-south gradient along the west coast of Admiralty
Inlet; therefore the variation in latitude was interpreted
as coastwise dispersion and the variation in longitude
was interpreted as inshore-offshore dispersion. Narwhal
space-use patterns were quantified in each of the three
time periods with a kernel home range polygon (prob-
abilistic measure of space use) [Arc View 3.1 Environ-
mental Systems Research Institute] based on average
daily positions from each whale.
Results
Pre-attack Narwhal behavior (> 5 days)
Narwhals observed passing Kakiak Point before the
Killer Whale attack moved in groups of 3-8 whales at
a distance of 20-200 m from the beach. Most whales
were heading south. Between 14 and 17 August, seven
Narwhals were instrumented with satellite tags before
the Killer Whales arrived in the vicinity of Kakiak
Point (Table 1). These Narwhals moved along the west-
ern side of Admiralty Inlet south of Kakiak Point and
less than 1-2 km from shore. No tagged Narwhals were
located in the center or eastern side of the inlet and the
linear home range along the west coast was <100 km.
TABLE |. Seven Narwhals instrumented with satellite transmitters prior to Killer Whale occurrence and attack at Kakiak
Point, Admiralty Inlet, Canada, 2005. Calf accompanying female on 17 September 2005 was not tagged.
ID Tagging date Time of release Sex Body length (cm)
20685 14 September 2005 15:06 F 360
20686 14 September 2005 15:56 M 483
20689 17 September 2005 06:25 F 360
37235 17 September 2005 10:45 F + calf 358
37236 17 September 2005 21:20 F 380
37280 17 September 2005 QeDi, F 380
37282 17 September 2005 21:45 F 364
2006
—e —
LAIDRE, HEIDE-JORGENSEN, and Ork: REACTIONS OF NARWHALS
459
ae
SF
Ficure 1. Oil film on the surface created by Killer Whales feeding on Narwhals in Admiralty Inlet, Canada, August 2005.
Large aggregations of Fulmars appeared in the area shortly after each kill. Photo by M. P. Heide-Jorgensen.
The standard deviation of average daily positions
indicated that dispersal of Narwhal groups varied ap-
proximately 19.9 km north-south, and 13 km west-east
before Killer Whales arrived (Table 2). The core of
the pre-attack kernel home range (50% probability
area) was concentrated just offshore of Kakiak Point
and distributed along the west coast of the inlet encom-
passing 347 km? (Table 2).
Immediate Reactions of Narwhals to Killer Whale
Attack
On 19 August, Killer Whales were reported by Inuit
hunters north of the camp, yet no detailed observations
of predation events or behavior were collected. On that
day, we observed Narwhals passing very close to the
shore (within the surf zone approximately 2-3 m from
land) and remaining still at the surface in large groups.
TABLE 2. Dispersal of Narwhals before, during, and after a Killer Whale attack in August 2005 as measured by the distance
between average daily satellite positions obtained from seven Narwhals instrumented at Kakiak Point prior to the arrival of
the Killer Whales. The standard deviation (SD) in latitude and longitude is reported and also converted to north-south dis-
persal and west-east dispersal with distance units of km.
Latitude SD Longitude SD N-S dispersal (km) W-E dispersal (km) Kernel core (50%) km?
Before attack (5 days) 0.18 0.12 19.9 13.0 347
During attack (2 days) 0.33 0.29 36.4 32.8 767
After attack (5 days) 0.18 0.25 19.7 27.6 599
460
ies Sees wk cot = ca . kon
FIGURE 2.
the shore lying still. Photo M. P. Heide-Jgrgensen.
On 20 August, Killer Whales were visually observed
arriving at Kakiak Point traveling a northbound route
along the west side of the inlet. Narwhals were already
present in the coastal area (<500 m from the shore)
around Kakiak Point and in a small bay just behind
the point. When the Killer Whales were within 2-4 km,
Narwhals suddenly moved closer to the shore in shal-
low water (<2 m). Some Narwhals formed tight groups
THE CANADIAN FIELD-NATURALIST
Vol. 120
St a Se
When Killer Whales arrived in the vicinity of Kakiak Point, Narwhals formed tight groups and remained close to
and others moved slowly or lay very still at the surface
(Figure 2). One Narwhal stranded on a flat gravel
beach (<0.5 m of water) and made violent tail thrash-
es for >30 seconds (Figure 3), either as a warning sig-
nal or in attempts to remove itself from the beach.
Satellite telemetry data indicated instrumented
Narwhals clearly responded to the presence of Killer
Whales (Figure 4). During the attack, both the north-
FiGurE 3. During the Killer Whale attack, Narwhals beached themselves in sandy areas and made tail slaps. Coastline can
be seen in lower left. Photo by K. L. Laidre.
2006 LAIDRE,
110°0'0"W
100°0'0"W
70°0'0"N
65°0'0"N
90°0'O"W 80°0'0"W
Kakiak Point
HEIDE-JORGENSEN, and Orr: REACTIONS OF NARWHALS
70°0'0"
J
46]
90°0'O"W 80°0'0"W
\
?
75°0'0"N J
it
AC Admiralty nll
}
@ 8/14 - 8/18
[} 8/19 - 8/20
A\ 8/21 - 8/25
FiGuRE 4. Average daily positions received from seven Narwhals instrumented with satellite tags before the Killer Whale attack.
Average positions are shown in 3 time periods: the five days prior to Killer Whale arrival (14-18 August), the two
days Killer Whales were observed (19-20 August), and the five days after the Killer Whale departure (21-25
south and west-east dispersal (as measured by inter-
whale mean latitude and longitude standard deviations)
doubled (Table 2). When Killer Whales were in the
area, the core kernel home range shifted south by ap-
proximately 80 km and doubled in size (767 km/’) (Fig-
ure 5, Table 2). Generally, the groups of Narwhals were
more widely dispersed in all directions during the
attack than before the Killer Whales arrived.
Post-Attack Narwhal Behavior (>5 days)
Narwhals resumed their normal swimming behay-
ior and distance from the coast within an hour after the
Killer Whales left the locality. No whales were ob-
462
served closer than 20 m from the beach and tight groups
at the surface broke-up and dispersed.
The instrumented Narwhals moved offshore and
utilized a wider area after the attack (ANOVA on longi-
tude, F, ,=17.6, P < 0.001). Standard deviations of the
mean inter-whale latitude (19.7 km) were reduced by
half after the departure of the Killer Whales and were
nearly identical to pre-attack values (19.9 km), demon-
strating a contraction of the north-south dispersal
(Table 2). The mean inter-whale longitude standard
deviation was slightly lower than during the Killer
Whale attack (27.6 km), yet did not return to pre-attack
value, indicating a general inshore-offshore dispersal
from the attack site.
Post-attack kernel home range core area (599 km?)
was less than that during the attack, yet the core area
was located east of the pre-attack core in the center of
Admiralty Inlet extending across to the eastern shore-
line (Figure 5, Table 2). In general the Narwhal groups
maintained a wider offshore dispersal and ranged more
widely than before the Killer Whale attack.
Discussion
Narwhals exhibited clear reactions to the immedi-
ate presence of Killer Whales. Their suite of behav-
iors included slow, quiet movements, travel very close
to the beach in the surf zone, movement into very shal-
low water less than 2 m, and formation of tight groups
at the surface. All of these observations are consistent
with Inuit accounts. Within an hour after the Killer
Whale attack and when Killer Whales had left the area,
direct observations suggested Narwhals resumed nor-
mal behavior (e.g., distances from the coast increased
and tight groups dispersed).
Satellite telemetry data revealed long-term (multi-
day) observations of Narwhal movements before and
after the attack, in addition to observations during the
attack. Average daily positions indicated that Narwhal
groups were more dispersed during and after the Killer
Whale attack. Dispersal primarily increased in the off-
shore (west-east) direction as measured by longitude
standard deviations and home ranges. Cores of area
use calculations demonstrated that the range of Nar-
whal movements increased two-fold during the Killer
Whale attack, shifted south in a direction away from
the attack point, and then expanded to offshore areas
afterwards (where Narwhals had not been observed
previously in 2005).
Later in the season, Killer Whales were reported in
“fairly large numbers” in Admiralty Inlet and vicinity
(especially Adams Sound) until October 2005 (Niore
Annie Igqalukjuak, Arctic Bay, personal communica-
tion). The exact location and movements of these Killer
Whales between August and September are not known,
nor is it known whether other attacks on Narwhals oc-
curred. Satellite tagging data showed that Narwhals
remained in Admiralty Inlet until the third week of
October, after which the southbound fall migration was
underway. Thus, it appears that even on longer time
THE CANADIAN FIELD-NATURALIST
Vol. 120
scales (~2 months) Narwhals do not alter their site fideli-
ty to the summering grounds (see Heide-Jgrgensen et
al. 2003) or depart early in response to the presence
of Killer Whales.
Killer Whales were also reported in Admiralty Inlet
in late August 2004 but no data or specific observa-
tions were available. It is unknown if the occurrence
observed in 2004 involved the same group of Killer
Whales as observed in 2005. On 25 August 2004, one
of the authors (J.R.O.) observed a young Bowhead
Whale (Balaena mysticetus) killed by Killer Whales
at Kakiak Point, and bones were found on the beach
less than 1 km away from the attack site the following
year in 2005. Furthermore, on the same date as the at-
tack reported here, several Killer Whales were observed
attacking Narwhals in Repulse Bay, Hudson Strait,
Canada (Joani Kringayark, personal communication).
Therefore, at least two separate groups of Killer Whales
feed on Narwhals on their summering grounds in the
eastern Canadian Arctic in August.
When Killer Whales attacked Narwhals, there was
limited action at the surface with no struggle or splash-
ing observed. Killer Whales consumed the Narwhals
below the surface of the water, and made multiple dives
in the center of oiled areas (presumed kills) for 15-20
minutes. An eyewitness account of a Killer Whale
attack on Narwhals in Inglefield Bredning, West Green-
land in August 2004 noted Killer Whales jumping on
top of Narwhals and holding them under the surface
until they drowned (Gretchen Freund, personal com-
munication).
If the predation level observed at Kakiak Point (4
Narwhals over 6 hours) is representative of the daily
predation level of a similar sized pod of Killer Whales
in the High Arctic, then consumption of Narwhals dur-
ing a two-month stay in Admiralty Inlet would amount
to approximately 200-300 Narwhals. A photographic
survey estimated 5,556 (CV=0.22) Narwhals in Admi-
ralty Inlet in 1984 (Richard et al. 1994). If the photo-
graphic survey results are corrected for whales that
were diving using a correction factor of 0.38 (0.06)
developed by Heide-Jgrgensen (2004), then the abun-
dance would have been in the order of 14,621(0.23).
A mortality of 200-300 Narwhals from Killer Whale
predation would represent <3% of this abundance esti-
mate. It is possible that predation constitutes a larger
proportion of the annual natural mortality of Narwhals
than previously expected; however, it is not known if
this natural mortality is fully compensatory. In this case,
a continued high level of Killer Whale predation in
combination with a population reduction by subsistence
harvest could reinforce a decline. Killer Whale preda-
tion has been responsible for the decline of sea otters
in Alaska (Estes et al. 1998; Doroff et al. 2003) and has
been hypothesized as the driver for a decline of other
top predators in the North Pacific (Springer et al. 2003).
The reason Narwhals occupy deep and narrow fjords
during summer is unknown and it has been hypothe-
sized that their summer distribution may be related to
463
IN, and ORR: REACTIONS OF NARWHALS
:IDE-JORGENSI
LAIDRE, HE
2006
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464
potential predation by Killer Whales (Kingsley et al.
1994). Narwhals feed very little during the summer
season (Laidre and Heide-Jgrgensen 2005a), and sum-
mering grounds do not appear to be related to calving
needs given calves are born in the spring. The summer
refuge from Killer Whales hypothesis lacks conclusive
evidence as Narwhal distribution in other seasons is
clearly driven by biological needs, such as sea ice for-
mation, open water, or access to prey resources (Laidre
and Heide-Jgrgensen 2005b) and Narwhals did not seek
refuge or depart early in response to the observed at-
tack. Little is known about what proportion of different
types of Killer Whales occurs in Arctic waters (i.e.,
mammal-eating vs. fish-eating). Stomach contents of
30 Killer Whales harvested in Disko Bay, West Green-
land in February 2003 contained only Lumpsucker fish
(Cyclopterus lumpus), despite the fact these Killer
Whales were taken in an area with a large abundance
of Bowhead Whales, Ringed Seals (Phoca hispida),
Narwhals, Belugas, and other potential marine mam-
mal prey items (Greenland Institute of Natural Re-
sources, unpublished data).
Few observations have been collected of Killer
Whales during winter in Arctic ice conditions. It is gen-
erally assumed that Killer Whales avoid the Arctic pack
ice (Heide-Jorgensen 1988) despite the fact many Kil-
ler Whales occur and thrive in the dense pack-ice of the
Antarctic. It is possible that changes in sea ice (lighter
sea ice cover and earlier break-up) will alter (or have
already altered) the occurrence of Killer Whales in
Arctic waters. These changes may facilitate increased
or longer visits by Killer Whales to ice-free Arctic
areas.
The predicted reduction of annual sea ice, together
with a longer open water season, will likely lead to an
increase in Killer Whale predation on Narwhals. At
the same time, reduced sea ice will also decrease the
probability that Narwhals succumb in ice entrapments
(also known as Sassats), another important source of
natural mortality (Laidre and Heide-Jgrgensen 2005b).
It is unclear how the interplay between these two ele-
ments of natural mortality will evolve under chang-
ing sea ice regimes.
Acknowledgments
Research was funded by the Greenland Institute of
Natural Resources, Fisheries and Oceans Canada, and
the Nunavut Wildlife Management Board. The Polar
Continental Shelf Project in Resolute supported logis-
tics for field work in August 2005. The work was also
supported by the Arctic Bay Hunters and Trappers
Organization. Narwhal tagging was conducted under
permits from Fisheries and Oceans Canada and Animal
Care Permit FWISLACC#013. Robert Pitman, Randall
Reeves, and an anonymous reviewer improved earlier
drafts.
THE CANADIAN FIELD-NATURALIST
Vol. 120
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Received 9 February 2006
Accepted 18 June 2007
Urine-marking and Ground-scratching by Free-ranging Arctic Wolves,
Canis lupus arctos, in Summer
L. Davip MeEcu!?
'U. S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 — 37" Street, SE, Jamestown, North Dakota 58401-
7317 USA
Mailing address: The Raptor Center, 1920 Fitch Avenue, University of Minnesota, St. Paul, Minnesota 55108 USA
Mech, L. David 2006. Urine-marking and ground-scratching by free-ranging Arctic Wolves, Canis lupus arctos, in summer.
Canadian Field Naturalist 120(4): 466-470.
Urine-marking and ground-scratching were observed in an Arctic Wolf (Canis lupus) pack on Ellesmere Island, Nunavut,
Canada, during 16 summers between 1986 and 2005. All previously known urination postures and ground-scratching by
breeding males and females were seen, and incidence of marking and scratching was greatest when non-pack wolves were
present. Observations of urine-marking of food remains supported the conclusion from a captive Wolf study that such mark-
ing signals lack of edible food.
Key Words: Arctic Wolf, Canis lupus arctos, ground-scratching, odor, scent-marking, territoriality, urine-marking, Ellesmere
Island, Nunavut, Canada.
Scent-marking by Wolves (Canis lupus) includes
both urine-marking and ground-scratching, usually by
dominant breeding males and females (Peters and
Mech 1975; Harrington and Asa 2003). Most infor-
mation about Wolf urine-marking has been collected
during winter by tracking Wolves in the snow (Peters
and Mech 1975; Rothman and Mech 1979; Paquet and
Fuller 1990; Paquet 1991; Zub et al. 2003) or during
other seasons by studying captive Wolves (Harring-
ton 1981; Raymer et al. 1984, 1986; Asa et al. 1986,
1990; Mertl-Millhollen et al. 1986; Ryon and Brown
1990; Barja and Miguel 2004). Only scattered men-
tions can be found of free-ranging Wolves urine-mark-
ing during summer (Clark 1971; Haber 1977; Mech
1991, 1995), and only one study of Wolf ground-
scratching in summer has been done (Zub et al. 2003).
Detailed information about urine-marking in wild
Wolves during summer is lacking because, with a few
exceptions (Murie 1944; Clarke 1971; Haber 1977;
Mech 1988), it has been difficult to observe Wolves
in the wild during summer because of their usual fear
and avoidance of humans.
I observed free-ranging Wolves in the Canadian
High Arctic over a period of 20 summers, where the
Wolves are highly tolerant of humans (Parmelee 1964;
Miller 1978; Mech 1988). There I obtained detailed
data on their urine-marking and ground-scratching
behavior.
Study Area
The study area includes about 2600 km? of the
Fosheim Peninsula east, north and west of Eureka on
Ellesmere Island (80°N latitude, 86°W longitude),
Nunavut, Canada. During summer, daylight is constant.
The area includes shoreline, hills, lowlands, creek bot-
toms and the area around Blacktop Ridge. Contrary to
much of the surrounding region, this area is generally
snow- and ice-free in summer, and contains rock, grav-
el, bare soil and scattered tundra and northern wetland
vegetation. Wolves, Muskoxen (Ovibos moschatus) and
Arctic Hares (Lepus arcticus) have long been common
in the area (Tener 1954), and Wolves have denned there
over decades or possibly centuries (Parmelee 1964;
Grace 1976; Mech 1988; Mech and Packard 1990).
Aside from intermittent scavenging around a weather
station and military base, the main foods of the Wolves
I studied were Muskoxen and Arctic Hares (Tener
1954), although seals (Phoca spp.) and Lemmings (Dic-
rostonyx groenlandicus) are also taken occasionally.
Methods
The Wolves I studied live far enough from exploita-
tion and persecution by humans that they are relatively
unafraid of people (Mech 1988, 1995). During 1986,
I habituated a Wolf pack to my presence and reinforced
the habituation each summer I observed them. The pack
frequented the same area each summer and usually used
the same den or nearby dens. The habituation allowed
me and an associate to remain with the Wolves daily, to
recognize them individually, and to watch them regular-
ly from 10 to 200-m away, and often as close as | m
(Mech 1988, 1995). This pack had disappeared by
2001, but by 2003 another pack, similarly habituated,
was present (Mech 2005).
Although I made no attempt to systematically or
completely record every urination or scratching made
by each Wolf, I did note as many such events as I could,
usually those made by breeding males and females,
while I was observing general behavior. I identified
breeding females by observing them nursing pups or
by the presence of obvious teats. Breeding males were
identified by their dominance over other males and their |
466
2006
MECH: URINE-MARKING AND GROUND SCRATCHING BY WOLVES
467
TABLE |. Yearly distribution of observed urine marks made by breeding male (RLUs and STUs) and breeding female (FLUs
and SQUs) Arctic Wolves, Ellesmere Island, Nunavut, Canada, 1986-1996, including only years in which observations cov
ered at least four weekly periods.!
Number of Male
Weekly Marks
Summers Periods Marks Per Period
1986 4 7 1.8
1987 5 7 1.4
1988 7 9 eS
1989 8 5) 0.6
1990 8 11 1.4
199] 8 1] 1.4
1992 6 29 4.8
1993 6 12 2.0
19942 5 3 0.6
1996 6 27 4.5
Female Total
Marks Marks
Marks Per Period Marks Per Period
0 0 7 1.8
0 0 7 1.4
0 0 9 1.3
5 0.6 10 hes
18 2.3 29 3.6
22, 2.8 33 4.1
14 25 43 V2
13 ZR, 25 4.2
5 1.0 8 1.6
14 PS, 4] 6.8
' Because of potential biases in times and methods of data collection, fine comparisons are not necessarily valid.
2 1995 covered only two periods.
overall initiative and control of pack activities (Schen-
kel 1947; Mech 1970, 1999; Peterson et al. 2002).
I collected data over periods of 1-8 weeks from
June through early August 1986-2005, except 1997,
1999, 2000, and 2001. Most of the longest data col-
lection took place 1986-1996 (Table 1). Because study
periods, pack behavior, and observation emphasis var-
ied over the summers and years (Table 2), biases in the
collection of urine-marking and ground-scratching
data over both the total duration of the study and over
the seasonal extent of the study each summer proba-
bly affected the data collected. Thus only the data on
ratios of marking postures should be unbiased enough
to warrant statistical testing, and I make no attempt
to draw detailed conclusions from the untested data.
However, these data are still useful to make broader
conclusions and to establish several facts that hereto-
fore were unknown or poorly documented in free-
ranging Wolves during summer.
Results
I identified three breeding female and three breed-
ing male Wolves, plus one pair without pups, that trav-
eled, hunted and marked together (Mech 1995, 2005,
and unpublished). I recorded males making 121 raised-
leg urinations (RLU) 22 June — 8 August and 30 stand-
ing urinations (STU) 15 June — 10 August; and females,
47 flexed-leg urinations (FLU) 15 June — 3 August, and
70 squat urinations (SQU) 15 June — 29 July (Table 2).
The proportion of male RLUs was significantly high-
er than the proportion of female FLUs (x? = 45.01;
P < 0.0001; df. = 1). The number of both male and
female marks I observed per weekly observation period
appeared to decrease over the summer (Table 2). The
number of recorded urinations per observation period
varied by year from 1.3 to 7.2. (Table 1).
In addition to FLUs by breeding females, I also
observed a pre-breeding female (bred the next year)
FLU twice on 6 August. An associate had also seen
this animal do “slight FLUs” three times on 13 July
(see Discussion.)
I observed double marking (Rothman and Mech
1979) 24 times between 26 June and 3 August. Double
marking is marking of a single location by both mem-
bers of a mated pair within a few minutes of each other
and can be initiated by either member. Of eight pos-
sible combinations of double-marking postures (RLUs
or STUs initiated by males, FLUs or SQUs initiated
by females) I observed five of the combinations, with
those involving RLUs and FLUs significantly pre-
dominating (Table 3).
I observed ground-scratching by both male and fe-
male breeding Wolves or by either individual as early
as 15 June and as late as 31 July. The pre-breeding
female (above) scratched as late as 6 August. Besides
the breeders’ and pre-breeder’s ground-scratching, a
non-breeding female was observed scratching during
two different summers, the same Wolf each year. Of
16 summers when I observed breeding or mated Wolves,
I recorded no scratching by them during seven of those
years and three or less scratching events during six
other summers.
Only during 1992, 1993, and 1996 did I see Wolves
do much scratching, and both male and female breed-
ers did so. During the above years, the breeding pair I
studied, the same individuals each year, scratched reg-
ularly. Those two individuals had also been the breeders
since 1989, but I saw little scratching during 1989-
1991, 1994 or 1995. During 1992, the breeding pair
scratched (and urine-marked — Table 2) much more
than I had seen them in the past. They scratched (and
urine-marked) especially (1) near a Muskox they had
killed and at which a non-pack Wolf had also been
feeding a few minutes before, (2) near a garbage dump
where at least one non-pack Wolf had been feeding
regularly and (3) along the easternmost location (ter-
468
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 2. Weekly distribution of observed urine marks made by breeding male (RLUs and STUs) and breeding female
(FLUs and SQUs) Arctic Wolves, Ellesmere Island, Nunavut, Canada, 1986-2005.!
Male
Number Marks
Summers Periods Marks Per Period
15-22 June 5 3 0.6
23-30 June 9 39 43
1-7 July 11 31 2.8
8-14 July 13 36 2.8
15-21 July 10 16 1.6
22-28 July 9 20 DD
29 July—4 August 9 4 0.4
5-10 August 6 2 0.3
Female Total
Marks
Marks Per Period Marks Per Period
9 1.8 12 2.4
18 2.0 Si 6.3
31 2.9 62 5.6
36 2.8 72 5.5)
16 1.6 32 39)
4 0.4 24 De
3 0.3 7 0.7
0 0) 2 0.3
' Because of potential biases in times and methods of data collection, fine comparisons are not necessarily valid.
ritory boundary?) where we saw them that summer.
During 1996, the breeding female scratched and urine-
marked while trying to force a Muskox away from the
den. In another study area, in the Canadian Northwest
Territories, I also observed wolves scratching several
times at a Red Fox (Vulpes fulva) den (Mech unpub-
lished) during August.
I observed ground-scratching in various elimina-
tion contexts: (1) with no other marking behavior, (2)
followed by FLU or RLU, (3) preceded by FLU, RLU
or SQU, (4) preceded and followed by FLU, or (5) pre-
ceded by defecation, similar to the findings of Peters
and Mech (1975) and Mertl-Millhollen et al. (1986).
I also observed urine-marking in relation to food
remains as Harrington (1981) also did. Breeding males
and females used all four kinds of urine postures in
marking uneaten food remains or prey odors including
the following (1) Arctic Hare intestines; (2) chewed
bones; (3) Arctic Hare stomach contents; (4) locations
where regurgitated or other food had recently been
eaten; (5) old kills; (6) the head of a recent Muskox
kill; (7) a putrid Muskox carcass which they sometimes
fed on but often passed up; (8) where another Wolf had
tried but failed to catch a lemming; and (9) at caches
of which they or another Wolf (including a pup) had
just consumed the contents.
Discussion
Captive breeding male and female Wolves urine-
mark at all times of the year, with the peak frequency
during the winter breeding season, and with all studies
except that by Ryon and Brown (1990) observing that
males tend to mark more year around than females
(Peters and Mech 1975; Mert!-Millhollen et al. 1986;
Asa et al. 1990; Barja and Miguel 2004).
Information on urine-marking frequencies during
summer in captive wolves varies considerably. In one
case, where the breeding pairs had pups, both males
and females increased their marking rate during sum-
mer (Mertl-Millhollen et al. 1986). In another, where
“reproductive pairs” were studied, but no pups were
born, the males increased their rate of RLU during
TABLE 3. Order and type of urination postures in double urine
marks! by arctic Wolves observed during summers 1986-2005
on Ellesmere Island, Nunavut, Canada. (Males do raised-leg
[RLU] and standing [STU] urinations, and females, flexed-leg
[FLU] and squat [SQU] urinations.)
Male Initiated
RLU-FLU 6
RLU-SQU 1
STU-FLU 2
STU-SQU 0
Female Initiated
FLU-RLU 9
FLU-STU 0
SQU-RLU 6
SQU-STU 0)
' Urination sequences involving RLUs or FLUs occurred sig-
nificantly more often than expected by chance (x7 = 6.86;
P=0.03; d.f. = 1).
summer, but the females did not increase their FLU
rate (Barja and Miguel 2004). In two others, where no
pups were involved, the RLU and FLU rates of domi-
nant males and females reached their minimum in
summer (Asa et al. 1990; Ryon and Brown 1990).
Testosterone tends to decrease to a minimum during
summer (Asa et al. 1990), as does testis size (Mech
2006). This relationship suggests that urine-marking,
which depends on testosterone (Asa et al. 1990), would
also decrease then. However, the variations found in
the studies cited above suggest that the social milieu
can affect the relationship between testosterone and
urine-marking, so that the relationship might not al-
ways be so tight, as Asa et al. (1990) also found. Be-
cause of possible biases mentioned earlier, my data can
add little to this subject, although conceivably the ex-
treme variation seen in my data reflect actual behavioral
variation similar to that reported in captive Wolves. In
that respect, my data on ground-scratching are instruc-
tive.
This study documents for the first time that wild
Wolves urine-mark throughout summer, using all
known urine postures, including double-marking, but
2006
that the proportion of RLUs and FLUs to STUs and
SQUs is lower then than in captive Wolves during
winter (Asa et al. 1990) when RLUs and FLUs tend
to predominate (Peters and Mech 1975).
The fact that a pre-breeding female was seen FLU-
ing (“RLU” in Mech 1995) is of special interest. Based
on my observation of this pack the previous three
summers, I inferred that she had been born two years
earlier. This pre-breeder began dominating the breed-
ing female, her apparent mother, about the time the
pre-breeder began FLUing. The pre-breeder bred the
next year, apparently with her father, and remained
bonded to him for the next six years, producing pups
during four of them while the mother remained with
the pack for two more years without being seen FLU-
ing, and then disappeared (Mech 1995 and unpub-
lished). This behavior of a young offspring starting to
FLU or RLU and challenging its same-sex parent for
dominance is similar to that seen in a captive colony
(Asa et al. 1990).
Previous information about free-ranging Wolves
ground-scratching in summer comes only from a single
study which concluded that between April and Septem-
ber, ground-scratching was low and stable (Zub et al.
2003). I could make no similar seasonal comparison,
but it was clear from my multi-summer observations
that the degree of ground-scratching varied consider-
ably from summer to summer, with no scratching seen
some summers and regular scratching during others. In
1992, the year of most scratching, the presence of non-
pack Wolves seemed to be the pertinent stimulus. It was
the only year I observed the Wolves I had been study-
ing chase or attack outsiders on three occasions (Mech
1993). It was also the year when I recorded the high-
est amount of urine-marking (Table 1), when the
Wolves usually were extra alert and looking around
intently when they marked and scratched, and when
they marked their easternmost locations, suggesting a
territory boundary. My impression during these times
was that the Wolves were in a high state of arousal
and aggressiveness. Both breeding males and females
scratched about equally, contrary to Mertl-Millhollen
et al. (1986) who found primarily females scratching.
Regarding marking of food remains, my observa-
tions confirm those of Harrington (1981) on captive
Wolves and support his conclusion that marking of
recently emptied food caches (and by extension from
my observations, other inedible food remains) signals
that a site contains no more edible food despite lin-
gering odors.
In summary, this study documents that much infor-
mation learned about Wolf urine-marking behavior in
Captivity during summer applies to free-ranging
Wolves; it extends information about ground-scratch-
ing; and it places information from captive studies
about urine-marking of food remains in its natural
context.
MECH: URINE-MARKING AND GROUND SCRATCHING BY WOLVES
469
Acknowledgments
This research was funded by the U. S. Geological
Survey. Polar Continental Shelf Project (PCSP), Atmos-
pheric Environment Services of Canada, and USDA
North Central Research Station provided logistical sup-
port. I also thank numerous field assistants, including
L. G. Adams, F. H. Harrington, W. Medwid, J. Sanders,
G. Breining, and H. D. Cluff. The following reviewed
the manuscript and offered helpful suggestions for its
improvement: L. G. Adams, S. M. Barber, and H. D.
Cluff, and M. E. Nelson. This is PCSP/EPCP paper
00905.
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Notes
Double Marking in Arctic Wolves, Canis lupus arctos: Influence of
Order on Posture
FRED H. HARRINGTON
Department of Psychology, Mount Saint Vincent University, Halifax, Nova Scotia B3M 2J6 Canada
Harrington, F. H. 2006. Double marking in Arctic Wolves, Canis lupus arctos: Influence of order on posture. Canadian-
Field Naturalist 120(4): 471-473.
Double marking by Arctic Wolves (Canis lupus arctos) was recorded by Mech (2006) from a pack on Ellesmere Island,
Nunavut, Canada, during 16 summers between 1986 and 2005. Using his data on the frequency of occurrence for each of the
four postures used by Wolves for urine marking (males — raised leg and stand urinations; females — flexed leg and squat uri-
nations), the probabilities of occurrence for each of eight possible double mark sequences were determined and compared
with observed frequencies. Females were somewhat but not significantly more likely to initiate double mark sequences.
There was no evidence for any bias in the posture used to initiate a double mark sequence, but assertive postures by both
males and females nearly always completed the sequence, occurring much more often than expected by chance.
Key Words: Arctic Wolves, Canis lupus arctos, scent-marking, double marking, pair bonding, mate guarding, urine-marking,
Ellesmere Island, Nunavut, Canada.
Double marking, in which a mated pair urine marks
a single site in quick succession, is commonly observed
in Wolves (Canis lupus) (Harrington and Asa 2003),
although evidence from wild Wolves is often inferred
from the pattern of urine marks on snow (Peters and
Mech 1975; Rothman and Mech 1979). Direct observa-
tions, such as those of Mech (2006), permit the identi-
fication of the postures used in marking, from which
underlying motivation might be inferred. Males may
use either raised-leg urinations (RLUs) or standing
urinations (STUs) and females either flexed-leg urina-
tions (FLUs) or squat urinations (SQUs), with the for-
- mer posture for either sex considered to be more reflec-
_ tive of aggressive or assertive motivation (reviewed by
_ Harrington and Asa 2003). Thus there are eight pos-
_ sible forms that a double marking sequence can take.
_ Mech (2006) has presented data on urine-marking in
_ Wolves, recorded by direct observation from Arctic
_ Wolves, Canis lupus actus, during the summer. Among
_ those data are 24 cases of double marking (Table 1),
_ and the reported finding that the more assertive pos-
_ tures (RLUs and FLUs) occur more frequently in dou-
ble marking. The purpose of this note is to extend the
analysis of those data in order to offer more insight
) into the motivation behind double marking.
Methods
A dominant adult Wolf may urine-mark using either
posture appropriate for its sex. This analysis seeks to
determine whether there is a non-random pattern in the
_ use of either posture, by either sex, in double marking
_ compared to urine-marking in general. Therefore I first
_ determined the overall frequency of each urination pos-
TABLE |. Urination postures used during double marking by
Arctic Wolves during summers 1986-2005 on Ellesmere
Island, Nunavut, Canada. Male postures: raised-leg [RLU]
and standing [STU] urinations; female postures: flexed-leg
[FLU] and squat [SQU] urinations. (Data from Mech
2006).
Male Initiated Double Marks
RLU—-FLU 6
RLU-SQU l
STU-FLU 2
STU-SQU 0
Female Initiated Double Marks
FLU-RLU 9
FLU-STU 0)
SQU-RLU 6
SQU-STU 0
ture that occurred exclusive of double marking (Table
2) to yield two base rates, one for the probability of
occurrence for each of the postures overall and anoth-
er for the probability of the occurrence of each form
within a sex. From these probabilities, one can calcu-
late expected frequencies for each form of double mark,
under the assumption that the postures used in double
marking simply represent random combinations of the
postures utilized otherwise. This is done by multiply-
ing the value from row | (the random probability the
posture will occur overall and thus initiate a sequence)
with the value from row 2 (the random probability the
other sex will use one of its two alternative postures
to complete the double mark). Thus, for example, the
47]
472
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 2. Calculation of the probability of occurrence for each of the urination postures used by Arctic Wolves on Ellesmere
Island, Nunavut, Canada during summers 1986-2005.
Urination posture
RLU STU
Total marks 121 30
Minus double marks -22 -2
Non-double mark urinations 99 28
(1) Probability of posture (overall) 0.45 0.13
(2) Probability of posture (within sex) 0.78 0.22
FLU SQU Total marks
47 70 268
-17 -7 -48
30 63 220
0.14 0.28 1.00
0.32 0.68 1.00/sex
TABLE 3. Expected and observed frequencies of occurrence for each of the eight different forms of Wolf double marking,
for Arctic Wolves on Ellesmere Island, Nunavut, Canada during summers 1986-2005. Expected frequencies are based on
the combined probability of occurrence for each posture in the sequence (from data in Table 2), under the assumption of
independence of occurrence for each posture type and based on a sample size of 24.
Double urination mark sequence
Male initiated sequences RLU-FLU RLU-SQU STU-FLU STU-SQU
Expected frequency 3.5) eS 1.0 Dell
Observed frequency 6 1 2 0
Probability of occurrence 0.144 0.306 0.042 0.088
Female initiated sequences FLU-RLU FLU-STU SQU-RLU SQU-STU
Expected frequency 2.6 0.7 2 NES)
Observed frequency 9 0 6 0
Probability of occurrence 0.109 0.031 0.218 0.062
probability for an RLU/SQU double mark to occur at
random is 0.45°0.68=0.306, whereas a double mark
in the reverse order (SQU/ RLU) is expected less
often (0.28°0.78=0.218), primarily because males are
more likely to use dominant postures overall than are
females (Table 3).
Results
Three questions can be addressed using the proba-
bilities generated in Table 3. First, is one sex more
likely to initiate a double marking sequence? Al-
though females initiated 63% of the double marks
observed, compared to the expected frequency of 42%,
this increase was not greater than expected by chance
(x? = 2.10, df = 1, ns). Second, is the first mark more
likely to be from an assertive posture than otherwise?
Again, although twice as many double marks were
initiated using the more assertive RLU and FLU pos-
tures, this was expected overall, given the high rate of
assertive posturing (59%) among dominant Wolves
(x? = 0.322, df = 1, ns). Finally, is the second mark
which completes the double mark, more likely to be
from an assertive posture? Here, the data indicate a high-
ly significant deviation from expectations (x7 = 13.90,
df = 1, P < 0.001). All but one of the second marks
involved either RLU or FLU postures. The one excep-
tion was an RLU by the male followed by a SQU by
the female. Males increased their use of RLUs from a
base rate of 78% in other situations to 100% when com-
pleting a double mark, while females increased their
use of FLU postures even more dramatically, from
32% overall to 89% in double marks.
Discussion
Several captive studies have found that dominant
female Wolves are usually the initiators of double
marks (93% of double marks, n=23, Rothman and
Mech 1979; 64%, n=36, Mertl-Millhollen et al. 1986).
In a sample of over 700 double marks by free-ranging
Coyotes (Canis latrans), Gese and Ruff (1997) found
that double marking was initiated by females 75% of
the time. Unfortunately, none of the previous studies
tested observed initiation rates against expected rates,
based on the overall frequency of marking by indi-
viduals of each sex, as I have done here. However, the
females observed by Mech (2006) did initiate double
marking more often than expected, at a rate (63%)
comparable to previous studies, so the lack of statistical
significance for this result may be an artifact of small
sample size. Thus it seems likely that double marks are
more likely to begin as urinations by dominant females.
Mertl-Millhollen et al. (1986) found. that 94% of
double marks in their captive pack were initiated by
RLUs (they combined RLUs and FLUs in their
analyses). Unfortunately, although they indicate that
RLUs comprised the majority of the marks they re-
corded, they do not present their data in a manner to
allow testing of expected versus observed rates for
the postures. In the present study, although the initial
posture used in a double mark was most often an
2006
assertive one (RLU or FLU), this is to be expected,
given the relatively high rate of these postures in dom-
inant individuals. This analysis suggests that the ini-
tial mark of a double mark occurs as an independent
event, influenced by a combination of internal factors
(arousal, hormones, urine load, etc.) and features of
the environment (smells of old marks, food odors, con-
spicuous visual objects, etc.), but not with the delib-
erate intent to initiate a double mark sequence. For
example, when Wolves are traveling, the lead Wolf is
typically the initiator of double marks (Mech 1999);
as it encounters sites which elicit its marking, the op-
portunity for double marking is then created for the
trailing Wolf.
The second mark, however, is highly influenced by
the presence of the first, and either its freshness, indi-
vidual identity, chemical composition, or its mere pres-
ence at a specific location induces the second individ-
ual to use the more assertive RLU or FLU posture.
Mertl-Millhollen et al. (1986) likewise found that all
but one second mark was either an RLU or FLU; the
one exception was a SQU (they combined SQUs and
STUs) that followed another SQU. Thus the second
mark of a double mark likely represents an increased
level of arousal on the part of the marker. Whether that
arousal represents generalized excitement, increased
sexual motivation or increased aggressive (protective)
motivation is not possible to determine here. A vari-
ety of hypotheses have been advanced to account for
double marking: pair bonding, reproductive synchro-
nization, mate guarding, and territorial defense being
among the most common (Rothman and Mech 1979;
Mertl-Millhollen et al. 1986; Gese and Ruff 1997).
The tight relationship between the frequency of dou-
ble marking and successful reproduction in Wolves
(Rothman and Mech 1979: Asa et al. 1986; Mertl-Mill-
hollen et al. 1986; Peterson et al. 2002) suggests that
mating related functions, including mate guarding by
the male, are most likely.
NOTES
473
Acknowledgments
This note grew out of a manuscript review. | would
like to thank L. D. Mech and F. R. Cook for the invi-
tation to prepare this paper and for their willingness
to expedite its consideration. I also thank my stu-
dents for their patience in receiving their final grades
while I worked on the manuscript.
Literature Cited
Asa, C. S., U. S. Seal, E. D. Plotka, and L. D. Mech. 1986.
Effect of anosmia on reproduction in male and female
Wolves (Canis lupus). Behavioral and Neural Biology 46:
272-284.
Gese, E. M., and R. L. Ruff. 1997. Scent-marking by Coy-
otes, Canis latrans: the influence of social and ecological
factors. Animal Behaviour 54: | 155-1166.
Harrington, F. H., and C. S. Asa. 2003. Wolf communica-
tion. Pages 66-103 in Wolves: Behavior, ecology, and con-
servation. Edited by L. D. Mech and L. Boitani. University
of Chicago Press, Chicago, U.S.A. 428 pages.
Mech, L. D. 1999. Alpha status, dominance, and division of
labor in Wolf packs. Canadian Journal of Zoology 77: 1196-
1203.
Mech, L. D. 2006. Urine-marking and ground-scratching by
free-ranging Arctic Wolves, Canis lupus arctos, in sum-
mer. Canadian Field-Naturalist 120: 475-479.
Mertl-Millhollen, A. S., P. A. Goodmann, and E. Kling-
hammer. 1986. Wolf scent marking with raised-leg urina-
tion. Zoo Biology 5: 7-20.
Peters, R., and L. D. Mech. 1975. Scent-marking in Wolves:
A field study. American Scientist 63: 628-637.
Peterson, R. O., A. Jacobs, T. D. Drummer, and L. D.
Mech. 2002. Leadership behavior in relation to domi-
nance and reproductive status in gray Wolves. Canadian
Journal of Zoology 80: 1405-1412.
Rothman, R. J., and L. D. Mech. 1979. Scent-marking in
lone Wolves and newly formed pairs. Animal Behavior 27:
750-760.
Received 24 December 2006
Accepted 24 December 2006
474 THE CANADIAN FIELD-NATURALIST Vol. 120
Use of “Micro”-Corridors by Eastern Coyotes, Canis latrans, in a
Heavily Urbanized Area: Implications for Ecosystem Management
JONATHAN G. Way! and Davip L. EATOUGH?
‘Biology Department, Boston College, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA. e-mail: jw9802 @ yahoo.com.
Present address: 64 Cranberry Ridge Road, Marstons Mills, Massachusetts 02648 USA.
*Science Department, Revere High School, 101 School Street, Revere, Massachusetts 02151 USA
Way, Jonathan G., and David L. Eatough. 2006. Use of “micro”-corridors by eastern Coyotes, Canis latrans, in a heavily
urbanized area: implications for ecosystem management. Canadian Field-Naturalist 120(4): 474-476.
We document the use of very narrow, linear corridors (termed “micro-corridors’’) that facilitated movements by both a transient
and a resident group of eastern Coyotes (Canis latrans) in a heavily urbanized area in north Boston, Massachusetts. Two
corridors are discussed: one, a railroad line through downtown Boston; and two, a hole in a cemetery fence giving access to
two separated cemeteries in a region of intense human development. Coyotes can be good subjects to illustrate the use of
fragmented landscapes because they are common and thus are abundant enough to study yet are wary and avoid novel
things and generally avoid people.
Key Words: Eastern Coyote, Canis latrans, corridor, fragmentation, Massachusetts, micro-corridor, urbanization.
The use of corridors is becoming increasingly rec-
ognized as an important conservation tool in frag-
mented landscapes (Beier 1995; Gilbert et al. 1998;
Meffe and Carroll 1994). Corridors are strips of habi-
tat connecting otherwise isolated habitat patches and
have been promoted as important features of reserve
design that allow movement among high(er) quality
habitats (Meffe and Carroll 1994). Connecting land-
scapes rather than maintaining a large unfragmented
core ecosystem is becoming the only alternative in
many urbanized areas, especially where unregulated
sprawl is occurring (e.g., Beier 1995). Yet, there is a
paucity of data on animal use of corridors and how
effective they are in connecting landscapes (Gilbert
et al. 1998; Meffe and Carroll 1994). Because of the
natural history of predators (e.g., large territories, long-
distance movements), a knowledge of carnivore biol-
ogy can predict minimum areas where ecosystems can
function relatively naturally (Beier 1993; Gittleman
et al. 2001; Mech and Boitani 2003; Meffe and Carroll
1994; Way et al. 2002a). However, the effectiveness
(positive or negative) of corridors could greatly affect
the size needed to maintain predators in certain sized
refuges.
Coyotes are common in North America (Parker
1995), yet are elusive and difficult to capture (Way et
al. 2002b); they are known to avoid novel objects and
structures that are dangerous to them, such as box traps
(Way et al. 2002b) or foothold traps (Conner et al.
1998; Sacks et al. 1999). Because Coyotes are ubiqui-
tous (Parker 1995), neophobic and wary (Sequin et al.
2003), and have relatively large home ranges and move-
ment rates (Way et al. 2004), they are a good species to
use in the determination of corridor use, especially in
urban areas. In other words, although they are common
and frequently use human-dominated areas (Grinder
and Krausman 2001; Way et al. 2004), they are natural-
ly shy and avoid people by being nocturnal and travel-
ing quickly in and out of human-dominated areas. For
example, Way et al. (2004) found that Coyotes travel
extensive distances on linear pathways such as pow-
erlines, railroad tracks and golf courses in urbanized
areas of Cape Cod, Massachusetts. Findings of Coyote
use of corridors in highly urbanized and fragmented
areas can potentially be used by managers to help pro-
tect more rare species and/or better design reserves
for more common species. In this note, we describe
the use of very narrow, linear (also called line corri-
dors by Meffe and Carroll 1994) “micro”-corridors.
Study Area and Methods
Coyotes were captured for an ecological study on
the north edge of Boston (42.43°N, 71.06°W), in east-
ern Massachusetts, in the bordering cities of Everett
(4345.0 people/km7), Malden (4290.5 people/km7), and
Revere (3089.0 people/km’) (U.S. Census Bureau,
2000 estimates). Coyotes were captured by box trap
(Way et al. 2002a) or by ground-based netlauncher
(one coyote; Coda Enterprises, Mesa, Arizona) and
outfitted with radio-collars (Mod-400, Telonics, Mesa,
Arizona, USA and M1900, Advanced Telemetry Sys-
tems, Isanti, Minnesota, USA). Tracking protocols were
described by Way et al. (2002a) and Way et al. (2004).
Portable receivers (Custom Electronics, Urbana, Ilh-
nois, USA) and hand-held 3-element Yagi antennas
were used to radio-track Coyotes both on foot and from
a vehicle. Because of the highly developed landscape
with many roads we mostly restricted our activities to
automobiles as Coyotes did not react to them as much
as to people (e.g., by running away; J. Way, unpub-
lished data); occasionally we approached radio-col-
lared Coyotes as close as possible on foot without
disturbing them. We used binoculars and video-cam-
eras when observing Coyotes, and city street lights,
nightscopes, and occasionally headlights when fol-
lowing Coyotes at night with a vehicle (Way et al.
2002a; Way et al. 2004).
2006
Observations
On 13 April 2004 we box-trap captured a dispers-
ing 13.6 kg female Coyote (“Fog” — ID #BN0402) in
a wooded section of Revere. Based on her behavior,
she seemed to have been dispersing from the north to
the south when she reached our study area. Her move-
ments the week after capture were mostly to the south,
including traveling >1 km through densely populated
neighborhoods and streets. She localized in four dif-
ferent areas for 2-3 days each before moving to a new
location. On 26 April 2004 we located her in a fenced-
in abandoned field of ca. 4 ha behind a shopping mall
at the north edge of the city limits of Boston. She
reached that location by either swimming a 200 m
river or traveling along railroad tracks over that same
river. Based on data taken earlier that night we sus-
pect that she went on the railroad tracks. For 3 days
she remained in the fenced-in area and ate many Nor-
way Rats (Rattus norvegicus) and Cottontail Rabbits
(Sylvilagus floridanus) (J. Way, unpublished data).
On the night of 29 April she traveled 1.8 km south-
west on the railroad line (which was bordered by
industrial buildings on both sides), going under Inter-
state 93 and the Zakim Bridge, and entered an aban-
doned railroad yard in Boston where she spent the
following day in a sparsely vegetated 200 x 50 m area.
At 02:00 h on 30 April she followed the railroad tracks
west through the Cambridge part of Boston and found
the first wooded area available 4.9 km from her pre-
vious day’s location. She was inactive through 23:37 h
on 30 April 2004 and could not be located until 19
November 2004 when she was found alive 100.5 km
south of her capture location (and 88.9 km south of her
last location in Cambridge) in the town of Dartmouth,
Massachusetts, near the Rhode Island border. To make
that voyage Fog had to have crossed (over or under)
six major interstate highways, including route 93 des-
cribed above.
From 17 May 2004 to 3 April 2005 we monitored a
pack of Coyotes (“The Cemetery Pack’’) in the border-
ing towns of Everett, Malden, and Revere. Four Coy-
otes were captured and radio-collared in this pack:
one, “Maeve” (#BN0404), a 14.5 kg lactating female,
captured 17 May 2004, was the breeding female; two,
“Jet” (#BN0403), a 15.9 kg breeding male, captured
by netlauncher on 29 June 2004, was Maeve’s mate;
three, “Jem” (#BN0406), a 10.0 kg 4.5-month-old pup,
was captured on 26 August 2004; and four, “Cour”
(#BN0405), a 12.3 kg 5-month-old pup, was captured
on 15 September 2004. The pack consisted of two to
three adults (i.e., one additional uncollared coyote was
occasionally sighted in the pack’s territory — its status
Was never determined but it was probably a helper
Coyote [Way et al. 2002a] to Jet and Maeve) and four
pups (two of which were not collared). The group
went from six to seven members to four individuals
by mid-winter 2004-2005 when it was presumed that
some of the pack members (two of the pups [includ-
NOTES
475
ing Jem, last successfully located on 11 December
2004] and probably the uncollared adult) dispersed.
The group resided almost exclusively in a green area
(including some thicker woods) surrounding four
large connected cemeteries. The entire area was about
2.5 km? (J. Way, unpublished data) and aside from
inactive railroad tracks (i.e., no trains used them) the
pack’s territory was surrounded by high-density hous-
ing units and/or commercial spaces (malls) on all sides.
Two roads transected their range, including a straight
east-west road (Fuller Street) in the central part of
their territory that connected the two cemeteries that
they most frequently used.
Most of the cemeteries were surrounded by fencing
consisting of vertical metal bars spaced 10 cm apart.
The narrowness of this space prevented the Coyotes
from crossing through the fence at random locations.
However, a small corridor connected two cemeteries
where a 34 cm opening occurred at a height of 55 cm
in the south cemetery because of a missing metal post
and two bent poles, one on each side. After crossing
Fuller Street diagonally for about 15 m the Coyotes
could then go through an open 305 cm x 198 cm gate
(this door was never shut) followed by six steps that
lead down into the north cemetery. Besides that cross-
ing, there were two roads (one open to cars, the other
gated) approximately 50 m east of the main corridor
that the Coyotes could use (by diagonally crossing
Fuller Street) to access either cemetery. There was
one other opening (< 20 cm) in the south cemetery
fence about 400 m west of the main corridor that led
into a residential neighborhood (after crossing Fuller
Street) bordering the west part of the north cemetery.
Cour was the only coyote to use the road crossing cor-
ridor regularly and Maeve was the only one observed
to use the neighborhood crossing corridor.
We radio-tracked the Coyotes six to seven days per
week on average and, when tracking them at night
(street lights illuminating the area), usually made sight-
ings (range: one to ten sightings per night) from our
parked, turned off vehicle of one to five Coyotes togeth-
er crossing the main corridor (exceptions were during
a heavy snowstorm [ca. 60 cm] when they did not use
the south cemetery at all). They often also crossed
the road in daylight during the early morning (06:00-
08:00). During their first 6 months (i.e., April to Octo-
ber) sightings included some of the pups going under
the fence (17 cm from fence to dirt) about | m west
of the opening in the fence leading to the south ceme-
tery, often during the same crossing (i.e., they were
together) as other Coyotes went through the opening
in the fence. This section of Fuller Street was straight.
thus the Coyotes could see in both locations and we
often watched them, especially the adults, standing on
a hill in the south cemetery watching traffic and cross-
ing when there were not any cars going by. Except for
two instances when the Coyotes were almost hit by cars
(the cars had to brake) all crossings were successful.
476
Monitoring of this group ended after the non-dispers-
ing members (Maeve, Jet, Cour and one uncollared)
were illegally poisoned.
Discussion
These data indicate that very small areas, if posi-
tioned in the right place, can be very important for
Coyotes. These micro-corridors gave access to city
habitats with few to no trees (along the railroad tracks)
and also connected fragmented areas (cemeteries with
fences) in already very urban landscapes. While not
ideal habitat for Coyotes, these corridors were cer-
tainly better than nothing, and echoing the statement
of Beier (1995: 235) when discussing Cougar (Puma
concolor) dispersal, “any connection between two
isolated patches is better than no connection.” Man-
agers can use these data for =3 purposes: one, to pro-
vide better habitat connectivity in already fragment-
ed landscapes (e.g., opening sections of fence where
wildlife is likely to cross); two, proactively establish-
ing these types of corridors (ideally larger than des-
cribed in this paper) where development is planned;
and three, more regional planning where important
habitat exists and maintaining linkages between sep-
arated core habitats.
The importance of sub-marginal habitats can not be
overstated to facilitate animal movement. For exam-
ple, Beier (1995) noted that Cougars can use corri-
dors 100 m wide if the distance is < 800 m and 400 m
wide if 1-7 km, yet adult Cougars have enormous home
ranges in the hundreds of km? (Beier 1993, 1995).
Likewise, species previously thought to inhabit only
wilderness such as Wolves (Canis lupus) can often
live at least at the edge of human-dominated areas if
not persecuted (Mech and Boitani 2003) and no doubt
corridors facilitate travel in and out of these landscapes.
While preserving larger areas (e.g., Beier 1995) is
preferred, this is not always possible in the real world
and it is increasingly being discovered that animals
are adaptable and can often respond to human-induced
changes (e.g., Mech and Boitani 2003). Therefore, it
is imperative to at least link what is remaining of our
wild lands and this can start at the very specific micro-
corridor landscape scale. For example, bridges that
exist because of waterways (common in many cities)
could also promote wildlife movement if cover and
some space are provided.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Acknowledgments
We thank the Way and Cifuni families, N. Lima, S.
Cifuni, E. Strauss, Environmental Studies Program,
and Urban Ecology Institute at Boston College, B.
Binder and Saugus Animal Hospital, and Revere High
School.
Literature Cited
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tat corridors for cougars. Conservation Biology 7: 94-108.
Beier, P. 1995. Dispersal of juvenile cougars in fragmented
habitat. Journal of Wildlife Management 59: 228-237.
Conner, M. M., M. M. Jaeger, T. J. Weller, and D. R.
McCullough. 1998. Effect of coyote removal on sheep
depredation in northern California. Journal of Wildlife
Management 62: 690-699.
Gilbert, F., A. Gonzalez, and I. Evans-Freke. 1998. Corri-
dors maintain species richness in the fragmented land-
scapes of a microecosystem. Proceedings of the Royal
Society of London 265: 577-582.
Gittleman, J. L., S. M. Funk, D. Macdonald, and R. K.
Wayne. Editors. 2001. Carnivore conservation. Cambridge
University Press, New York. 675 pages.
Grinder, M. I., and P. R. Krausman. 2001. Home range, habi-
tat use, and nocturnal activity of coyotes in an urban envi-
ronment. Journal of Wildlife Management 65: 887-898.
Mech, L. D., and L. Boitani. Editors. 2003. Wolves: behav-
ior, ecology, and conservation. University of Chicago Press,
Chicago, Illinois. 448 pages.
Meffe, G. K., and C. R. Carroll. 1994. Principles of con-
servation biology. Sinauer Associates, Sunderland, Mas-
sachusetts. 601 pages.
Parker, G. R. 1995. Eastern coyote: the story of its success.
Nimbus Publishing, Halifax, Nova Scotia. 254 pages.
Sacks, B. N., K. M. Blejwas, and M. M. Jaeger. 1999. Rel-
ative vulnerability of coyotes to removal methods on a
northern California ranch. Journal of Wildlife Management
63: 939-949.
Sequin, E. S., M. M. Jaeger, P. F. Brussard, and R. H.
Barrett. 2003. Wariness of coyotes to camera traps rela-
tive to social status and territorial boundaries. Canadian
Journal of Zoology 81: 2015-2025.
Way, J. G., I. M. Ortega, and P. J. Auger. 2002a. Eastern
coyote home range, territoriality and sociality on urbanized
Cape Cod, Massachusetts. Northeast Wildlife 57: 1-18.
Way, J. G., I. M. Ortega, P. J. Auger, and E. G. Strauss.
2002b. Box-trapping eastern coyotes in southeastern Mas-
sachusetts. Wildlife Society Bulletin 30: 695-702.
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Received 4 October 2005
Accepted 14 March 2007
2006 NOTES 477
Bald Eagle, Haliaeetus leucocephalus, Preying on Maritime Garter
Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova
Scotia
STORRS L. OLSON
Division of Birds, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 USA
Olson, Storrs L. 2006. Bald Eagle, Haliaeetus leucocephalus, preying on Maritime Garter Snake, Thamnophis sirtalis pallidulus,
on Cape Breton Island, Nova Scotia. Canadian Field-Naturalist 120(4): 477.
At 1053 hours on 23 July 2006 a Bald Eagle, Haliaeetus leucocephalus, was observed in flight transporting in its talons a
live Maritime Garter Snake, Thamnophis sirtalis pallidulus, at South Harbour (46°52'01.7"N, 60°26'45.8" W), Cape Breton
Island, Victoria County, Nova Scotia.
Key Words: Bald Eagle, Haliaeetus leucocephalus, predator, Maritime Garter Snake, Thamnophis sirtalis pallidulus, South
Harbour, Cape Breton Island, Nova Scotia.
The Bald Eagle, Haliaeetus leucocephalus, is a large
North American raptor that usually frequents aquatic
environments where it feeds predominantly on cap-
tured or scavenged fish. Eagles may be seen daily in
South Harbour, at the northern end of Cape Breton,
Nova Scotia. This area appears to be occupied by a
resident pair based on an observation of two adults
seen standing very close together on a mud flat at the
mouth of the harbour on 16 July 2006. South Harbour
is a Shallow, brackish, nearly landlocked estuary with
an artificially maintained channel through the sandbar
across its mouth providing exchange of water with
the Atlantic Ocean.
On 23 July 2006 at 10:53 h, the screams of eagles
drew my attention to two adult birds wheeling and skir-
mishing in the air over the southern shore of the har-
bour (46°52'01.7"N, 60°26'45.8"W). When two birds
eventually flew directly overhead I saw that the lead
bird was carrying a snake in its talons. The grayish-
brown coloration and very attenuated, pointed tail
writhing in characteristic ophidian fashion precluded
any possibility of the prey being an American Eel
(Anguilla rostrata). Two other witnesses (Johanna R.
Humphrey, Katherine Allen) also confirmed that the
prey, which was very much alive, was a snake. The
individual carried by the eagle appeared to be some-
what in excess of 600 mm in total length. The only
snake of this size in Nova Scotia would be a Maritime
Garter Snake, Thamnophis sirtalis pallidulus, females
of which range from 412 to 916.9 mm, whereas males
do not exceed 602 mm (Gilhen 1984). Because most
of the area surrounding South Harbour, especially the
southern shore, where the observation was made, is
very heavily vegetated, the eagle most likely captured
the snake while the latter was swimming across the
surface of the water. Maritime Garter Snakes are “excel-
lent swimmers” and have been seen on “freshwater lakes
100 metres or more from shore” (Gilhen 1984: 132).
The Bald Eagle feeds almost exclusively on verte-
brates, paticularly on fish, followed by birds, and then
mammals (Stalmaster 1987; Buehler 2000). Although
Stalmaster (1987: 93) emphasized the eagle’s “vast
diversity of prey’, he considered that snakes “are an un-
usual dietary item” (pages 95-96) and cited only the
report of McEwan and Hirth (1980) of the remains of a
single Black Swamp Snake, Seminatrix pygaea, in a
nest in central Florida. Bald Eagles are notorious scay-
engers, and large numbers of snakes, including Semi-
natrix, are victims of vehicular traffic in central Florida
(Smith and Dodd 2003), so there is a strong possibility
that that food item was obtained as road kill. I have
found no other references to Bald Eagles feeding on
snakes, although they probably do so opportunistical-
ly wherever snakes and eagles occur together.
Literature Cited
Buehler, D. A. 2000. Bald Eagle Haliaeetus leucocephalus. In
The Birds of North America (506: 1-40). Edited by A. Poole
and F. Gill, The Birds of North America, Inc., Philadelphia,
Pennsylvania.
Gilhen, J. 1984. Amphibians and reptiles of Nova Scotia.
Nova Scotia Museum, Halifax.
McEwan, L. C., and D. H. Hirth. 1980. Food habits of the
Bald Eagle in north-central Florida. Condor 82: 229-231.
Smith, L. L., and C. K. Dodd. 2003. Wildlife mortality on
U.S. Highway 441 across Paynes Prairie, Alachua County,
Florida. Florida Scientist 66: 128-140.
Stalmaster, M. V. 1987. The Bald Eagle. Universe Books,
New York.
Received 8 August 2006
Accepted 16 February 2007
478 THE CANADIAN FIELD-NATURALIST Vol. 120
Rat Poison Kills a Pack of Eastern Coyotes, Canis latrans, in an
Urban Area
JONATHAN G. Way!, STEPHEN M. CiFuN?, Davip L. EatouGH?, and Eric G. STRAUSS *
‘Environmental Studies Program and Lynch School of Education, Boston College, Higgins Hall, Chestnut Hill, Massachusetts
02467 USA; e-mail: jw9802@yahoo.com Present address: Eastern Coyote Research, 64 Cranberry Ridge Road,
Marston Mills, Massachusetts 02648 USA
232 Maple Street, Revere, Massachusetts 02151 USA
3Science Department, Revere High School, 101 School Street, Revere, Massachusetts 02151 USA
4Urban Ecology Institute, Environmental Studies Program, and Biology Department, Boston College, Higgins Hall, Chestnut
Hill, Massachusetts 02467 USA
Way, Jonathan G., Stephen M. Cifuni, David L. Eatough, and Eric G. Strauss. 2006. Rat poison kills a pack of eastern Coyotes,
Canis latrans, in an urban area. Canadian Field-Naturalist 120(4): 478-480.
We document the death of a pack of Eastern Coyotes (Canis latrans) from high levels of brodifacoum, a second generation
poison that is the active ingredient in some forms of rat poison (e.g., d-Con®). The Coyotes died within a week of each
other during late March/early April 2005. This incident indicates the vulnerability of wild animals to commercial over-the-
counter rodenticides.
Key Words: Canis latrans var., Eastern Coyote, anticoagulant, brodifacoum, poison, rat poison, Massachusetts.
Coyotes (Canis latrans) live successfully in a vari-
ety of habitats ranging from rural to urbanized areas
(Gese et al. 1996; Harrison et al. 1991; Patterson and
Messier 2001; Riley et al. 2003; Way et al. 2001, 2004).
However, in almost all of these settings (except nation-
al parks; Gese et al. 1996) people constitute the major
source of mortality for Coyotes usually via trapping,
shooting and automobile strikes (Grinder and Kraus-
man 2001; Parker 1995). Poison was historically used
to kill predators but was banned in 1973 in the United
States (Mech 2000; Mech and Boitani 2003). Mech
(1970) noted that “poison is no doubt the most effec-
tive and efficient method of controlling or exterminat-
ing Wolves (Canis lupus).’ However, the use of poisons
is controversial because of their relative non-selectivity
and reputation for inhumaneness (Cluff and Murray
1995). Most poisons, such as strychnine, cyanide, and
sodium fluoroacetate (compound 1080), are not readily
obtainable today. Because these poisons are illegal,
many canid populations have greatly increased in the
past 30 years (Mech and Boitani 2003; Parker 1995).
Anticoagulants are present in urban areas (for rat con-
trol), and Riley et al. (2003) found them to be a signifi-
cant cause of death for Coyotes in southern Califor-
nia. This paper details the poisoning deaths of a family
group of Coyotes in urban north Boston, Massachu-
setts, most likely deliberately poisoned by someone .
Study Area and Methods
Coyotes were captured for an ecological study on
the north edge of Boston (42.43°N, 71.06°W), in east-
ern Massachusetts, in the bordering cities of Everett
(4345.0 people/km7), Malden (4290.5 people/km?), and
Revere (3089.0 people/km*) (U.S. Census Bureau,
2000 estimates). Box traps were used to capture Coy-
otes (Way et al. 2002a) except for one (“Jet”) cap-
tured via a ground-based netlauncher (Coda Enterpris-
es, Inc., Mesa, Arizona). Four Coyotes were radio-col-
lared in this pack: “Maeve” (#BN0404), a 14.5 kg
lactating female, captured 17 May 2004, was the breed-
ing female; “Jet” (#BN0403), a 15.9 kg breeding male,
captured 29 June 2004, was Maeve’s mate; “Jem”
(#BN0406), a 10.0 kg 4.5 month-old pup, was captured
on 26 August 2004; and “Cour” (BN0405), a 12.3 kg
5 month-old pup, was captured on 15 September. The
behavior of Maeve and Jet (i.e., frequently being locat-
ed with each other and with the pups, including obser-
vations of them feeding the pups) indicated that they
were the parents of these pups (see Way et al. 2001).
The Coyote pack consisted of two to three adults (1.e.,
one additional uncollared Coyote occasionally sight-
ed in the pack’s territory — its status was never deter-
mined but it was probably a helper Coyote [see Way
et al. 2002b] to Jet and Maeve) and four pups (two of
which were not collared). The group was named the
Cemetery Pack as it resided almost exclusively at one
green area (including some thicker adjacent woods)
surrounding four large connected cemeteries. The entire
area was about 2.5 km? and on the north side, aside
from unused railroad tracks the pack’s territory was
surrounded by high-density housing units and/or com-
mercial spaces (malls) on all sides.
Results and Discussion
The group went from six or seven members in the
fall of 2004 to four individuals by mid-winter 2004-
2005 when it was presumed that some of the pack
members (two of the pups [including Jem] and proba-
bly the uncollared adult) dispersed. Jem was last suc-
cessfully located on 11 December 2004. Snow track-
ing and sighting data indicate that only one uncollared
Coyote (a light yellowish-brown animal) remained
along with the breeding pair and Cour. Similarly, Way
et al. (2002b) found three to four individuals to be a
2006
typical winter pack size of Coyotes in eastern Massa-
chusetts.
The winter of 2004-2005 was harsh with much snow
yet the Coyotes remained in their small territory. Res-
idents often reported to us that they fed the Coyotes
and people noticed some of the animals were collared
(J. Way, unpublished data), indicating that the group
obtained food from people living in and/or around the
cemeteries. Aside from their abnormally small home
range (see Way et al. 2002b), they behaved much like
Coyotes studied in other locations, including the avoid-
ance of people by being nocturnal (Way et al. 2004),
crossing streets quite often, denning in wooded or rel-
atively undisturbed (including under a gravestone)
regions of their territory (Way et al. 2001), and acting
territorial (Way et al. 2002b), including observations
of Jet scent marking on the railroad tracks at the north
edge of his pack’s home range. There was no sign of
the Coyotes’ ill-health until just prior to them dying.
On 27 March 2005 Maeve was found dead in the
middle of a cemetery in the central part of the pack’s
territory. She was an emaciated 12.7 kg despite ap-
pearing normal when sighted on 24 March. A gross
necropsy revealed significant internal bleeding and no
fetuses, indicating that she was not pregnant. A labo-
ratory (Idexx Veterinary Services, www.vetconnect.
com) diagnosis indicated erosive acute gastritis, severe
necrotizing hemorrhagic endometritis with retained
placental decidual tissue, and subacute suppurative
endocarditis and myocarditis.
On 31 March 2005 Jet was found dead, 100 m from
where Maeve died, and was also emaciated (14.3 kg).
Radio-telemetry data indicates date of death was 30
March. He was observed moving normally 2-3 days
before his death. Because of massive internal bleeding
and similar gross necropsy results as Maeve, we only
tested for poisoning (specifically for common chemi-
cals found in household rat poisons) on Jet. Brodifa-
coum was detected in the liver at 0.733 parts per mil-
lion (ppm), and the laboratory (Idexx) indicated that
the results supported a diagnosis of brodifacoum poi-
soning.
On 3 April 2005 Cour was found dead in a shallow
(< 1 m deep) canal that he frequently (i-e., daily) crossed
prior to his death. He appeared healthy and weighed
17.3 kg, heavier than both of his parents. His relatively
robust physique support observations from residents
indicated that Cour (with a red ear tag) was the radio-
collared Coyote most commonly seen eating food left
by people. He was previously observed up-close and,
besides limping on his right hind leg, appeared healthy
on | April 2005. Not having obtained the results from
Jet or Maeve at the time, we had a full necropsy per-
formed at Tufts University (Grafton, Massachusetts).
Internal bleeding (subcutaneous hemorrhage) was noted
and based on autopsy findings and toxicological analy-
sis of the liver (brodifacoum = 0.542 ppm), Cour died
from an anticoagulant rodenticide. Because of the way
NOTES
479
Jet and Cour died, and the similar necropsy findings
from Maeve (i.e., massive internal bleeding), we con-
clude that Maeve also died from brodifacoum poi-
soning.
Because all three of the Coyotes’ behavior seemed
normal prior to their death and that they all died close
together (< 1 week), it appears that someone purpose-
fully poisoned them at high concentrations rather than
the coyotes having eaten enough poisoned prey to have
died (i.e., from bioaccumulation; Riley et al. 2003).
Most likely Maeve and Jet were poisoned around the
same time, and then Cour was given a later dose(s)
judging by a sighting of him traveling alone after
Maeve and Jet were documented as dead. However,
Cour’s healthier condition might have allowed him to
survive longer than his parents. We extensively searched
for the source of the poisons (especially near where
the coyotes died and where they spent the majority of
their time when they were alive), including informal-
ly talking to numerous people, but we never managed
to locate any substantial leads.
Throughout summer 2005 only a couple of sightings
were made by residents and cemetery staff indicating
that either the uncollared Coyote survived or, more
likely, a new Coyote (i.e., formerly not part of this
pack) dispersed into this location. Regardless of those
individual sightings, this pack was decimated in a short
period of time via rodenticides.
The public should be better informed of the dangers
that common household poisons present for wildlife,
especially in urbanized areas, and the potential health
threat to humans and pets. For example, about two
months later, pet dogs (Canis familiaris) died or were
injured from rat poison in neighboring towns (S. Cifu-
ni, personal communication). The slow, painful death of
an animal internally bleeding is not pleasant and should
not be acceptable in our society. These animals were
diagnosed because they were part of an ecological
study; due to the expense of testing and difficulty of
finding non-radio-collared animals that die in the
woods, it would be difficult to estimate how many non-
target (i.e., not rats or mice) animals die from antico-
agulants. We recommend that these poisons be strictly
controlled either through making the over the counter
sale of them illegal to all but licensed, professional
exterminators and/or through required public education
campaigns explaining the dangers of these poisons.
Acknowledgments
This study would not be possible without the sup-
port of the Way and Cifuni families, N. Lima, the
Saugus Animal Hospital and Dr. Bob Binder, Fiore’s
Meat Market and The Meat Market (in Winthrop), D-
L. Szumylo, the Environmental Studies Program and
Urban Ecology Institute at Boston College. Numer-
ous private individuals donated to have the necrop-
sies performed and numerous residents in the area of
the poisonings helped out in myriad ways. Care and
480
capture of free-ranging Coyotes was approved by
Boston College’s Institutional Animal Care and Use
Committee Protocol Number 01-02 (renewed in 2005)
and by the Massachusetts Division of Fisheries and
Wildlife permit #003.04LP.
Literature Cited
Cluff, H. D., and D. L. Murray. 1995. Review of wolf con-
trol methods in North America. Pages 491-504 in Ecology
and conservation of wolves in a changing world. Edited by
L. N. Carbyn, S. H. Fritts, and D. R. Seip. Canadian Cir-
cumpolar Institute, Occasional Publication Number 35,
Edmonton, Alberta, Canada.
Gese, E. M., R. L. Ruff, and R. L. Crabtree. 1996. Foraging
ecology of coyotes (Canis latrans): the influence of extrin-
sic factors and a dominance hierarchy. Canadian Journal
of Zoology 74: 769-783.
Grinder, M., and P. R. Krausman. 2001. Morbidity-mor-
tality factors and survival of an urban coyote population
in Arizona. Journal of Wildlife Diseases 37: 312-317.
Harrison, D. J., J. A. Harrison, and M. O’Donoghue. 1991.
Predispersal movements of coyote pups in eastern Maine.
Journal of Mammalogy 72: 756-763.
Mech, L. D. 1970. The wolf: the ecology and behavior of an
endangered species. 1995, Reprint. University of Minneso-
ta Press, Minneapolis, Minnesota, 384 pages.
Mech, L. D. Editor. 2000. The wolves of Minnesota: howl in
the heartland. Voyageur Press, Stillwater, Minnesota. 128
pages.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Mech, L. D., and L. Boitani. Editors. 2003. Wolves: behav-
ior, ecology, and conservation. University of Chicago Press,
Chicago, Illinois. 448 pages.
Parker, G. R. 1995. Eastern coyote: the story of its success.
Nimbus Publishing Halifax, Nova Scotia, Canada, 254
pages.
Patterson, B. R., and F. Messier. 2001. Social organization
and space use of coyotes in eastern Canada relative to prey
distribution and abundance. Journal of Mammalogy 82:
463-477.
Riley, S. P. D., R. M. Sauvajot, T. K. Fuller, E. C. York,
D. A. Kamradt, C. Bromley, and R. K. Wayne. 2003.
Effects of urbanization and habitat fragmentation on bob-
cats and coyotes in southern California. Conservation Biol-
ogy 17: 566-576.
Way, J. G., P. J. Auger, I. M. Ortega, and E. G. Strauss.
2001. Eastern coyote denning behavior in an anthropogenic
environment. Northeast Wildlife 56: 18-30.
Way, J. G., I. M. Ortega, P. J. Auger, and E. G. Strauss.
2002a. Box-trapping eastern coyotes in southeastern
Massachusetts. Wildlife Society Bulletin 30: 695-702.
Way, J. G., I. M. Ortega, and P. J. Auger. 2002b. Eastern
coyote home range, territoriality and sociality on urbanized
Cape Cod, Massachusetts. Northeast Wildlife 57: 1-18.
Way, J. G., I. M. Ortega, and E. G. Strauss. 2004. Movement
and activity patterns of eastern coyotes in a coastal, subur-
ban environment. Northeastern Naturalist 11: 237-254.
Received 4 October 2005
Accepted 7 April 2007
A New Record of Deepwater Sculpin, Myoxocephalus thompsonii,
in Northeastern Alberta
M. STEINHILBER! and D. A. NEELY
'Royal Alberta Museum, 12845 102nd Avenue, Edmonton, Alberta TSN 0M6
"Department of Biology, St. Louis University, St. Louis, Missouri 63103 USA
Corresponding author: e-mail: dave.neely @ gmail.com. Present address: Department of Ichthyology, California Academy of
Sciences, 875 Howard Street, San Francisco, California 94103 USA
Steinhilber, M., and D. A. Neely. 2006. A new record of Deepwater Sculpin, Myoxocephalus thompsonii, in northeastern
Alberta. Canadian Field-Naturalist 120(4): 480-482.
We present the first documented records of Deepwater Sculpin, Myoxocephalus thompsonii, from northern Alberta, and the sec-
ond record for the province. Three specimens of Deepwater Sculpin were taken in gill nets set at 17 to 20 m depth in Colin Lake,
Alberta, on 15 September 2001. Colin Lake, located in the Canadian Shield region of northeastern Alberta about 125 km north-
east of Fort Chipewyan, drains into Lake Athabasca via the Colin River. The only other known Alberta population of Deepwater
Sculpin inhabits Upper Waterton Lake in the southwestern corner of the province. This record is approximately 300 km SSE
of the nearest verified record in the Northwest Territories and 400 km NW of the nearest verified record in Saskatchewan.
Key Words: Deepwater Sculpin, Myoxocephalus thompsonii, distribution, Colin Lake, Alberta
Three individuals of Deepwater Sculpin, Myoxoce-
phalus thompsonii, were collected in two separate gill
net sets in Colin Lake, Alberta (59°34'N, 110°08'W)
on 15 September 2001. One set was in 17 m of water,
and fished for a period of 15 hours. The other was set
at 18-20 m depth for 14 hours. Each net set consisted
of one 60 m multi-mesh net with six 10 m panels rang-
ing from 10 to 25 mm bar mesh and one 60 m net with
six 10 m panels of 10 to 50 mm bar mesh. All nets were
1.8 m in depth. The two nets were fished in series on
the bottom of the lake. Both collecting sites were locat-
ed on the gently sloping periphery of the two deep
basins in the lake (Figure 1). The maximum depth of
the lake is approximately 25 m. The composition of
the substrate at the sampling sites was not determined.
Species taken syntopically with the sculpins included
Lake Whitefish (Coregonus clupeaformis), Cisco (Co-
regonus artedi), Burbot (Lota lota), Northern Pike
2006
Colin Lake
1 2%, #3
kilometers
Colin River NN
(drainage to Lake Athabasca) *
FiGuRE |. Bathymetric map of Colin Lake, with Deepwater
Sculpin capture localities indicated. Depths given in
metres.
(Esox lucius), Lake Trout (Salvelinus namaycush), and
Longnose Sucker (Catostomus catostomus). Lake
Whitefish, Cisco, and Longnose Suckers were the most
common species.
All three specimens of Deepwater Sculpin are gravid
females, ranging from 62-74 mm SL (Standard Length),
with four prominent, large, and straight preopercular
spines, a large gap between the dorsal fins, and the gill
covers free at the isthmus. Meristic counts of these
specimens are consistent with data presented by McAI-
lister (1961) for adjacent populations in Saskatchewan
and the Northwest Territories. Dorsal fin spines were
(number of specimens in parentheses) 7(2) or 8(1);
dorsal fin rays 14(1), 16(1), or 17(1); anal fin rays 14(1),
15(1), or 17(1); pelvic rays 3(6); pectoral rays 16(2) or
17(4). Two of the three had an incomplete lateral line
(ending under the last dorsal fin ray) with 32 pores;
in the remaining specimen the lateral line continued
posteriad to the caudal peduncle and had 36 pores.
These specimens represent the first record of Deep-
water Sculpin from northern Alberta, and only the sec-
ond record for the province (Roberts 1988). The only
other known population in Alberta occupies Upper
Waterton Lake (49°02'N, 113°54'W) on the Alberta/
Montana border (McAllister and Ward 1972) (Figure
2). Nelson and Paetz (1992) suggested the likely pres-
ence of Deepwater Sculpin in northeastern Alberta,
given the proximity of populations in adjacent Saskat-
chewan and the Northwest Territories. The closest pop-
ulations supported by voucher specimens are Great
Slave Lake, Northwest Territories (~300 km NNW)
and Wollaston (400 km SE) and Reindeer Lakes (520
km SE), Saskatchewan (COSEWIC 2006*). Records
from Lake Athabasca in northwestern Saskatchewan
(~100 km ESE, COSEWIC 2006*) are not supported
by voucher specimens. The specimens from Colin Lake
fill this gap, and further suggest that additional popula-
tions of this enigmatic cottid remain undiscovered in the
northern portions of its range. New populations con-
NOTES
451
Re ho
100"W
FIGURE 2. Distribution of Deepwater Sculpin in western
Canada (data from Scott and Crossman 1973, and
Murray et al. 2003). Colin Lake records indicated by
open circle.
tinue to be discovered as deepwater habitats are system-
atically sampled (i.e., Murray et al. 2003; COSEWIC
2006*).
Although typically encountered at depths greater
than 44 m (Scott and Crossman 1973), this species is
occasionally found at much shallower depths, espe-
cially as larvae and juveniles (COSWEIC 2006 ); Raw-
son (1951) collected specimens in | m of water in Great
Slave Lake, while McPhail and Lindsey (1970) docu-
mented specimens from 4-6 m in Great Bear Lake. Alex
Peden (in McPhail and Lindsey 1970) suggested that
this fish is probably concentrated in deep water during
the summer and moves inshore in the fall. The discoy-
ery of Deepwater Sculpin in Colin Lake is thus some-
what unusual in that the lake is not particularly deep,
with a maximum depth of only 25 m. Further studies
to determine seasonal distribution patterns within the
lake and any unique biological characteristics of this
population would be valuable. It is worth noting that
Colin Lake is connected via the Colin River to the
Saskatchewan side of Lake Athabasca (approximate-
ly 20 km to the SE) where depths can reach 124 m. It
is likely that trapping efforts in deepwater habitats of
Lake Athabasca will provide additional records. How-
ever, there are currently no data to support the use of
the Colin River as a post-glacial movement corridor
between Colin Lake and Lake Athabasca.
The only populations of Deepwater Sculpin current-
ly given protective status in Canada are the Great Lakes
— Western St. Lawrence populations, which were list-
ed as Threatened by the Committee on the Status of
Endangered Wildlife in Canada [COSEWIC] (Parker
1988) but have recently been downgraded to Special
Concern (COSEWIC 2006 ), and are on schedule 2 of
482
the Canadian Species At Risk Act (SARA). Western
populations (which include the new population dis-
cussed herein) were designated Not at Risk in 2006
(COSEWIC 2006’).
The gill nets used to capture these Deepwater Scul-
pins were deployed as part of a Shortjaw Cisco (Core-
gonus zenithicus) survey in the Canadian Shield region
of Alberta (Steinhilber 2004"). Our recent experience
suggests that deployment of cyalume light stick-bait-
ed wire minnow traps in deepwater habitats is more
effective at sampling Deepwater Sculpin than are small-
mesh gill nets. The effectiveness of this technique has
not been previously discussed in the literature. Long-
term surveys of Lake Saganaga on the Ontario/Min-
nesota border have primarily used small-mesh gill nets
(Etnier and Skelton 2003). Between 1986 and 2003,
only one specimen of M. thompsonii was taken. Use
of light stick-baited cylindrical wire minnow traps
during summer 2004 per our request resulted in cap-
ture of 10 specimens (D. A. Etnier, personal commu-
nication). Similarly, during 2003 we provided Fish-
eries and Oceans Canada personnel on Lake Nipigon
with light sticks and traps that were set simultaneously
with their gill nets, with similar results (A. Van Ogtrop,
personal communication). Other researchers working
on Deepwater Sculpins have had similar experiences
(T. Sheldon, personal communication; COSEWIC
2006"). We thus strongly recommend the use of this
technique over small-mesh gill nets for assessing pres-
ence/absence of Deepwater Sculpin.
Acknowledgments
We thank B. Meagher for field assistance. Collect-
ing was conducted under permit 401 FR issued to M.
Steinhilber. D. Neely was supported by a grant from
the British Columbia Ministry of Fisheries.
Materials Examined
All specimens are deposited in the holdings of the
Royal Alberta Museum, Edmonton. PMA L01.33.319,
Myoxocephalus thompsonii, n=1 (74.0 mm SL),
Colin Lake, 59°33'12"N, 110°9'30" W. 15 September
2001. Collectors: M. Steinhilber, B. Meagher. PMA
LO1. 33.311, M. thompsonii, n=2 (62.7-73.0 mm SL),
Colin Lake, 59°33'28"N, 110°5'8" W. 15 September
2001. Collectors: M. Steinhilber, B. Meagher.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Documents Cited (marked * in text)
COSEWIC. 2006. COSEWIC assessment and update status
report on the deepwater sculpin Myoxocephalus thomp-
sonii (Western and Great Lakes-Western St. Lawrence pop-
ulations) in Canada. Committee on the Status of Endangered
Wildlife in Canada. Ottawa. 39 pages. (www.sararegistry.
gc.ca/status/status_e.cfm).
Steinhilber, M. 2004. Shortjaw cisco species at risk assess-
ment 2001. Alberta Sustainable Resource Development,
Fish and Wildlife Division, Alberta Species at Risk Report
Number 82. Edmonton, Alberta. 21 pages.
Literature Cited
Etnier, D. A., and C. Skelton. 2003. Analysis of three Cisco
forms (Coregonus, Salmonidae) from Lake Saganaga and
adjacent lakes near the Minnesota/Ontario border. Copeia
2003: 739-749.
McAllister, D. E. 1961. The origin and status of the deepwa-
ter sculpin, Myoxocephalus thompsonii, a Nearctic glacial
relict. National Museum of Canada Bulletin 172: 44-65.
McAllister, D. E., and J. C. Ward. 1972. The deepwater
sculpin, Myoxocephalus quadricornis thompsoni, new to
Alberta, Canada. Journal of the Fisheries Research Board
of Canada 29: 344-345.
McPhail, J. D., and C. C. Lindsey. 1970. Freshwater fish of
northwestern Canada and Alaska. Bulletin 173, Fisheries
Research Board of Canada, Ottawa, Ontario. 381 pages.
Murray, L., M. H. Papst, and J. D. Reist. 2003. First
record of the Deepwater Sculpin, Myoxocephalus thomp-
sonii, from George Lake in Whiteshell Provincial Park,
Manitoba. Canadian Field-Naturalist 117: 642-645.
Nelson, J. S., and M. J. Paetz. 1992. The fishes of Alberta.
2™4 edition. The University. of Alberta Press, Edmonton.
437 pages.
Parker, B. J. 1988. Status of the deepwater sculpin, Myoxo-
cephalus thompsoni, in Canada. Canadian Field-Naturalist
102: 126-131.
Rawson, D. S. 1951. Studies of the fish of Great Slave
Lake. Journal of the Fisheries Research Board of Canada
8: 207-240.
Roberts, W. E. 1988. The sculpins of Alberta. Alberta Nat-
uralist 18: 121-127; 153. Addendum 19:35.
Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes
of Canada. Bulletin 184, Fisheries Research Board of Cana-
da, Ottawa, Ontario. 966 pages.
Received 29 December 2005
Accepted 27 March 2007
Book Reviews
ZOOLOGY
The Clements Checklist of Birds of the World 6th Edition
By James F. Clements. 2007. Cornell University Press, Sage
House, 512 East State Street, Ithaca, New York 14850 USA.
864 pages. U.S.$59.95. Cloth.
If you travel outside of North America [or any other
home territory] you will have to face the question. Is
this bird species one I have seen before? There are so
many look alike birds that are different species in some
areas and the same species in others. You can wonder
about crows, oystercatchers, stilts and the plethora of
wagtails. Or is this bird with the same or a similar
name to a look-alike home species, the same or a differ-
ent bird. For example are Willow Grouse, Red Grouse
and Willow Ptarmigan the same species? It can be real-
ly confusing.
Enter Dr. James Franklin Clements [1927-2005], an
orphan who served in the U.S. merchant marine and
navy before going to college. In the Korean war he was
in the air force, before finally settling to get his Ph.D.
in 1975. His thesis became the first edition of a check
list of the world’s birds. Because of ever-evolving tax-
onomy we are now at the 6th edition.
Clements has produced a list, simply a list, of all the
known bird species and subspecies. Producing such a
simple list is a very complex task indeed. It must have
felt like trying to resolve the unresolvable on many
occasions. So here we have this simple list and can
ask how useful is it? Theoretically, I should be able
to check out a bird that I am unsure about. Looking it
up in Clements should allow me to clarify the bird’s
status and decide whether it can be added to my life
list or not.
How well does this work? The first problem is you
must use the same English name as Clements. Nowhere
is there a list of alternative names. If you look for Melba
Finch in the index you will be disappointed. You will
need to use either Green-winged Pytilia or look for the
genus Pytilia. The North American bias is shown by
Circus cyaneus being called Northern Harrier as op-
posed to Hen Harrier [the older name].
Now we come to the sticky issue of splits and lumps.
The North American Magpie is split from its European
counterpart, as is Hooded and Carrion Crow. Clements
lists the Somali Ostrich as a subspecies, Struthio cam-
elus molybdophanes, whereas most recent literature
gives this distinctive blue-necked bird separate species
status [Struthio molybdophanes}. Similarly the bright-
ly coloured barn swallow found only in the Nile delta
is not split from its more common cousin. Again this
is listed by some as a separate species, Egyptian Swal-
low, Hirundo savignii. The vegae Herring Gull is sep-
arated as East Siberian Gull. This is open to question
and recent work suggests more research is needed.
Certainly I have found it difficult to see a difference
in the field. Is it a little darker than a Herring or was I
being deluded by poor light?
Each entry is accompanied by a description of the
species range. These ranges are useful when compar-
ing species. For example, my list for Mongolia includes
Pink-footed Goose, Anser fabalis. The range for Pink-
footed Goose in Clements does not include Mongolia,
whereas the Bean Goose, correctly Anser fabalis, is
present. Clearly this is an error in the Mongolian list,
probably due to translation problems, that was quick-
ly resolved.
However, the ranges given for other species are less
useful. Clements can be forgiven for minor lapses such
as the range of the Northern Wheatear or Crested Auk-
let — given as Siberia. The birds I saw on the eastern-
most coast of Russia this year were technically in the
Russian Far East — not Siberia. Siberia does not extend
to the east coast. This is minor compared to Houboura
and Macqueens Bustards. Clements splits these two
birds and gives Macqueens a range of the Nile to ....
Pakistan. The Handbook of the Birds of the World,
Volume 3, cited as the major reference, does not split
this species but lists it as a subspecies, Chlamydotis
undulata macqueenii, and correctly gives the range as
NW Kazakhstan to .... Mongolia and Northern China.
Similarly Clements does not mention the range exten-
sion of Cardinal and House Finch into eastern Canada.
I have checked to see if some of the recent discover-
ies are included in this check list. Scarlet-banded Bar-
bet and Long-whiskered Owlet are, but the two most
recent discoveries, Large-billed Warbler [the 1867 refer-
ence is given but not the recent March 2007 capture of
two birds] and Gorgeted Puffleg, [May 2007] are not.
Having one coherent checklist is extremely valu-
able for the travelling birder. It still does not solve all
the issues with a changing taxonomy, but at least you
can use it as a solid foundation for your records. You
will still have to research current taxonomy and recent
discoveries and decide which birds you will accept as
a full species.
Incidentally, I make it a point never to directly mark
books. I use post-it notes or equivalent. I will make an
exception for this one book. Not only is it set up to be
483
A84
ticked, but it is a good idea to add information on
species as new data become available. That is until
version seven is printed.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
Birds of Europe, Russia, China and Japan: Passerines, Tyrant Flycatchers to Buntings
By Norman Arlott. 2007. Princeton University Press, 41 Wil-
liam Street, Princeton, New Jersey 08540-5237 USA. 240
pages. U.S.$29.95. Paper.
My wife and I recently decided to visit northern
China and Mongolia [the Silk Road]. This left me
with a dilemma; which bird book or books would I
need to take to get adequate coverage. I did not want
to pay $100 or more for a paperback copy of A Field
Guide to the Birds of China (John MacKinnon and
Karen Phillips, Oxford University Press, New York).
My Birds of Russia (Flint et al.) gives fair coverage
but the illustrations are not as precise as I would like
them, particularly for warblers, etc. Birds of China (De
Schauensee) only illustrates some of the species and
gives descriptions of range only and you need a thor-
ough knowledge of China’s geography to follow these
descriptions. Birds of India (Grimmett et al.) is a much
better guide with a lot of species covered [as winter
visitors], but the coverage is far to the south so the
range maps are useless.
Then Princeton announced Birds of Europe, Russia,
China and Japan and I thought it would be just right for
my purposes. While the book covers only the passer-
ines (from Tyrant Flycatchers to Buntings), it is exactly
these groups for which I needed a better, more up-to-
date book. The non-passerines, like ducks and gulls,
tend to be more distinctive and are suitably covered in
previous books.
Technically Princeton calls this book an illustrated
checklist. Each species has an illustration or two with
very brief facing-page text. At the back of the book are
range maps. While the text presents little about each
species but key features, it is perfect for the traveller.
It measures only 5" x 7" x 4" [12.5 x 17.5 x 2 cm]
and weighs 14 oz [100 g or around half a normal field
guide].
The range maps are 2" x %" [5 x 2 cm] and cover
from Britain to Kamchatka, from Svalbard to North-
ern India. As there are no country boundaries [these
would make the little map too cluttered] I found it
helpful to create a mask with a piece of clear plastic
sheet. I traced the outline of the map in blue, then, using
Google Earth as a guide [Google Earth has a similar
orientation to these maps] I shaded in transparent yel-
low the region I plan to visit. I can now place the mask
over each map and quickly see if I will be in any bird’s
range. This was somewhat satisfactory. I compared my
list created with the mask to actual trip lists and I was
in error by around 15 percent.
Although the book’s title suggests it covers Europe,
Russia, China and Japan, the maps go much further.
For example it includes the Nile Valley Sunbird of the
Middle East and the Purple Sunbird of India, as well
as the geographically restricted Sinai Rosefinch and
Sub-Sahara’s Blandford’s Lark. I am not sure of the
author’s rationale for including birds like the Rosy-
patched Shrike, a purely African species [but I am hap-
py he did so].
The critical illustrations are those of difficult to iden-
tify birds such as warblers and some finches. Arlott
has been illustrating bird books for many years [my
first Arlott book is over 40 years old]. His skill has
grown dramatically and his most recent work is top
quality. My copy has all the illustrated species look-
ing a little dark. [Technically the hue is correct but
the illustrations are over saturated and this is part of
the printer’s quality control. North American readers
will see this when looking at the Cedar Waxwing and
Varied Thrush illustrations. Both are way more intense
than the real birds I have seen this year.] In detail
though the artwork is excellent. When I looked at Arc-
tic Warbler and try to separate it from it congener leaf
warblers or the Beautiful Rosefinch from its relatives
then the relevant field marks are apparent. North Amer-
icans will have no difficulty identifying wood warblers
using this guide, even with the intense colours. [I was
surprised to find over 40 “North American” species,
such as Wood Warblers and Icterids, have been seen
in this region as vagrants. ]
This book solved my problem and will be ideal for
other travelers to this large region. I look forward to a
similar book on the non-passerines. I hope Princeton
does well with their checklist series [I believe they have
a dozen titles] as they are so much more convenient
when luggage weight is critical.
Literature Cited
Flint V. E., R. Boehme, Y. Kostin, and A. Kuznetsov. A
Field Guide to Birds of Russia and Adjacent Territories.
Princeton University Press, New Jersey.
De Schauensee, Rodolphe Meyer. The Birds of China.
Smithsonian Institution Press, Washington.
Grimmett, Richard, Carole Inskipp, and Tom Inskipp. Birds
of India, Pakistan, Nepal, Bangladesh, Bhutan, Sri Lanka and
the Maldives. Princeton University Press, New Jersey.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
2006
Deep Alberta. Fossil Facts and Dinosaur Digs
By John Acorn. 2007. The Royal Tyrrell Museum and The
University of Alberta Press, Edmonton, Alberta Canada.
XII + 186 pages. $26.95.
Early on the recognition of our country’s rich re-
sources was often a reactive response, rather than proac-
tive endeavours. It was in response to the U.S. activities,
primarily those of Barnum Brown (1873-1963) of the
American Museum of Natural History, New York, that
the Canadian government in early the 20" century
began their vertebrate paleontological activities in the
Canadian west. First under the Geological Survey of
Canada, followed by the National Museum of Canada
(now, Canadian Museum of Nature, Ottawa). These
activities were central to what has been called “The
Great Canadian Dinosaur Rush” (see David Spalding’s
Into the Dinosaurs’ Graveyard, Doubleday Canada,
Toronto, 1999).
Despite these efforts, and the bountiful treasures col-
lected and studied, it wasn’t until the late 1960s that
the Province of Alberta, host of the majority of these
activities, was able to begin to take control of its own
paleontological resources. Provincialism — restriction
of outside exploitation — emerged. This occurred with
the materialization of the University of Alberta, Edmon-
ton, establishing its own paleontological program of
education and research, supported by the Vertebrate
Paleontology In Alberta conference in 1963 (Univer-
sity of Alberta, 1965) in addition to the establishment of
the Provincial Museum of Alberta (Edmonton), which
opened in 1967. What followed was a slow eruption
of discovery, research, tourism and marketing savvy.
John Acorn’s Deep Alberta skims the surface of all that
is sexy in paleontology in Alberta. And as most will see,
there is a lot to witness.
Acorn’s guide, based on his CKUA radio series by
the same name, is not quite similar to the other popular
guides to Alberta’s paleontological wealth, The Land
Before Us, The Making of Ancient Alberta by the Royal
Tyrrell Museum of Palaeontology (Lone Pine Press,
Edmonton, 1994) is more of a complement to its own
gallery (somewhat similar to the Royal Saskatchewan
Museum’s Earth Sciences gallery guide, Geological
History of Saskatchewan by John E. Storer (Govern-
ment of Saskatchewan, Regina, 1989)). Nor is it obvi-
ously as in-depth as Dinosaur Provincial Park (edited
by P. J. Currie, Indiana University Press, Blooming-
ton, 2005), a symposium volume summarizing all the
Book REVIEWS
485
major taxonomic groups found within the park. Deep
Alberta is designed similar to a field-guide, with one-
page descriptions and colorful illustrations of species,
or non-taxonomic themes.
Alphabetical in its listings of themes, creatures,
and places, each item listed is given usually a page of
text, accompanied by a full page illustration, often a
photograph of an exhibit at the Royal Tyrrell Muse-
um of Palaeontology. These range from mosasaurs,
those snake/lizard derivatives from the marine environ-
ment of the Cretaceous Period, the Chasmosaurus,
the horned-dinosaur familiar to the fields of Dinosaur
Provincial Park, to the singularly rare Atrociraptor, the
“savage robber” skull found near Drumheller — only
one partial skull of this is known.
Of place, from the badlands near Drumheller to the
Milk River regions are few. The thematic questions in-
clude “How do you Know Where to Dig?”, an often
made public query. Yet, surprisingly, a hot topic like
extinction is not addressed. Of people, the likes of
Barnum Brown and the Sternberg family (three gen-
erations of which have worked in Alberta) are briefly
brought to the fore. Appended is a list of most of the
Ph.D.’s who are currently at play in the paleontological
field of Alberta. Those who have contributed research
but lack graduate documentation, or who have provided
discoveries without institutional support, are not rec-
ognized in this summation.
Since the discovery by Joseph B. Tyrrell (1858-
1957) over a century ago of an Albertosaurus jaw, later
culminating with erection of a provincial museum spe-
cifically dedicated to paleontology (The Royal Tyrrell
Museum of Palaeontology) and the designation of
Dinosaur Provincial Park as a World Heritage Site, the
Province of Alberta has ample reason to show off its
fossil resources. Acorn’s Deep Alberta is a primer of a
primer, a very first introduction to this wealth. Acorn
has much to choose from in his summation of pale-
ontology of Alberta and will likely be another useful
marketing tool to maintain the visibility of this science
in the minds of the public. The treatment of these re-
sources by other provincial jurisdictions should take
note. Alberta is not the sole place for paleontology.
There is a lot more to see.
TIM T. TOKARYK
Box 163, Eastend, Saskatchewan SON OTO Canada
Gibson’s Guide to Bird Watching & Conservation
By Merritt Gibson. 2007. Nimbus Publishing Limited, Hali-
fax, Nova Scotia, Canada. 214 pages. $16.46.
This small bird watching guide to the maritime prov-
inces is a tribute to the birds of Nova Scotia, Prince
Edward Island and New Brunswick. It is also a trib-
ute to the ornithologists and bird-watchers of the area,
the unsung heroes who for the past several decades have
been the backbone of bird conservation and study to
preserve habitat, protect endangered species and pro-
486
mote birding. The reader is introduced to a variety of
birds and a variety of birders following the colourful
pages of anecdotes, life histories and illustrated birds.
The book does not try to describe all of the birds
which regularly visit the maritime provinces of Cana-
da. It is not a field guide to all birds. It is a book for
bird-watchers and conservationists and introduces the
reader to several birds, each of the sea, the coast, the
shore, freshwater, the forest, agricultural lands, and
finally towns and gardens, in that order. The book is
divided into the seven sections named above with an
introduction to each section and ending comment intro-
ducing birders amateur and professional studying with-
in each of the sections and involved in the steward-
ship of habitat. Three to seven birds are described in
some detail in each section. The description includes
anecdotal stories of the birds, their habitats, origins of
their common name, behaviour, nesting and egg-laying
behaviours and chick hatching and rearing behaviour.
Whether one is ready to brave the wind and waves to
spot birds, or maybe just sit in an armchair by the win-
dow, the text leads you to appreciate the diversity of
birds present in the Maritime Provinces and the pos-
sibilities of discovering or re-acquaintance with species
which have always been present and probably have
been visible many times.
Conservation is one important focus of the book and
the end of each chapter gives some detail on the work
of amateurs and professionals who research, monitor,
THE CANADIAN FIELD-NATURALIST
Vol. 120
and care for the different populations of each bird spe-
cies mentioned. Throughout Nova Scotia, New Bruns-
wick, Prince Edward Island, and Newfoundland, many
different bird populations are in decline, especially but
not exclusively, species mentioned in the book. Differ-
ent programs are in place to monitor and conserve lands
inhabited by these birds, and programs are being devel-
oped to make the public aware of the trends in bird pop-
ulations. This book is one of those awareness projects.
I read the book quickly and then went back over the
chapters more slowly to savour the stories of birds I
know and the birds mentioned which I would like to
see one day. It is an exciting book for a part-time birder
and one which taught me not just about the birds but
even more about the state of the shoreline, the wet-
lands, the forests and the agricultural fields. To the vis-
itor, the land changes little, but to the resident, there
are problems and potential solutions. The solution is
often awareness and information. I applaud the re-
searchers for their work and Merritt Gibson for bring-
ing it together. The book also serves as a delightful
picture book illustrated by Twilia Robar-DeCoste with
drawings of birds, birds in their natural settings, land-
scapes and seascapes to show perspective and the beau-
ty involved in the study and enjoyment of birding on
Canada’s East Coast.
JIM O’ NEILL
28718 Five Mile Road, Livonia, Michigan 48154 USA
Ladybugs of Alberta: Finding the Spots and Connecting the Dots
By John Acorn. 2007. University of Alberta Press, Edmon-
ton, Alberta, Canada. 169 pages. $29.95.
This is an important book about insects for natural-
ists. It has field credibility. Acorn reviewed and under-
stands the writings of the experts on lady beetle identi-
fication and ecology in North America. Acorn studied
the specimens that collectors in Alberta deposited in
collections. But most importantly, he was in the field in
Alberta chasing and watching and photographing lady
beetles. The result is a guide, written by an expert,
which will be valuable before someone starts chasing
lady beetles and will continue to provide insights as a
person’s expertise grows. And it is accessible; virtu-
ally anyone will be able to use it.
What can you do with this book? First and foremost
it is possible to identify the lady beetles (a.k.a. lady-
bugs, ladybird beetles, Coccinellidae) that live in Al-
berta. The combination of excellent colour drawings,
photographs, plus key features highlighted in text make
it possible. In addition the range maps, both range in
Alberta and much of North America, notes on food and
habitat preferences give some guidance, to reduce the
probability of an embarrassing misidentification. The
difficult identifications are not glossed over. The neces-
sity of dissecting genitalia to distinguish some Hippo-
damia (e.g., glacialis and quinquesignata) and the sig-
nificance of chromosomal differences between Chilo-
corus stigma and C. hexacyclus are covered clearly,
with humour and insight.
Most authors would have taken the easy route and
restricted themselves to the big showy lady beetles with
which most naturalists are familiar. However Acorn
tackled the “lesser ladybugs”, a daunting task to make
what is known about Scymnus and Hyperaspis and
Brumoides widely available, including both features
that make species level identifications possible and
some natural history. Acorn deserves credit for perse-
vering and including the lesser species, especially con-
sidering the extra work, almost double the number of
species, and that they are not as charismatic to gener-
al naturalists and that little is known about them. He
engenders interest in these little beetles.
In addition to identification and species accounts,
there are substantial sections on ecology, behaviour,
and history of coccinellid study in Alberta. The hot
topic of the influence of introduced species on native
species is attacked head on. It starts with a perceptive
critique of the functionally useless Nature Canada at-
tempt to coordinate a citizen science project on lady
beetles in the 1990s. His says that a prejudgment, intro-
duced species are bad, led to many of the problems.
In my view, the other key problem was that there was
little or no quality control that compromised the value
2006
of the data. What was needed was Acorn’s book, or
listening to field entomologists, during the planning
process. The discussion continues with an assessment
of the Evans’ hypothesis that native species have con-
stricted their range to ancestral habitats, that is have
become less common in extensively human altered
habitats (farmland and suburbia) after the arrival of
introduced species such as the seven-spotted lady bee-
tle. Refreshingly, he is able to present his own data to
support his views. His views about ecological change,
conservation and invasion biology are clear, logical and
worthy of discussion.
The style deserves special mention. It is highly cred-
ible, yet at the same time there is a breath of whimsy
and fireside chattiness. A fine example is the descrip-
tion of taste-testing. You will not look at a brightly
coloured lady beetle the same after you read this.
What bothered me during my initial readings? The
first was that I wanted more detail, more data. But, this
is a book for generalists and not the appropriate place
for the level of detail I want. Second were those cou-
plets. Each species account includes a quirky, rhyming
couplet. I did not understand many at first, but they did
grow on me. Perhaps they are like beer, an acquired
taste. Third were the common names. The Entomolog-
ical Society of Canada is working to provide consis-
tent, standard common names for insects, just like we
Book REVIEWS
487
have for birds. However, many were ignored, for exam-
ple Mulsantina picta here is the Painted Ladybug, not
pine lady beetle. However, some of the newly coined
names hit the nail on the head and deserve to be main-
tained. My favourite was the Once-squashed Lady-
bug, a cryptic species in the same genus as the Twice-
stabbed Ladybug. Yes, it needs to be squashed on a
microscope slide to examine its chromosomes in order
to be identified. Brilliant. These are minor complaints.
The identification sections are worth the whole price
of the book. The discussion of introduced species is
worth the whole price of the book. The description of
how to taste a ladybug to assess its palatability is worth
the whole price of the book.
Our understanding of foods, habitat use and phenol-
ogy (seasonal activity) is not as well developed in North
America as it is in Great Britain and Europe. People
armed with this guide, will have the tools to be at the
forefront in remedying this situation. Compilations
often function as a catalyst for a quantum leap in inter-
est and new findings by curious naturalists. I predict
this book will provide another great example in west-
ern Canada.
DaviID MCCORQUODALE
Department of Biology, Cape Breton University, Sydney,
Nova Scotia B1P 6L2 Canada
A New World of Animals: Early Modern Europeans on the Creatures of Iberian America
By Miguel de Asua and Roger French. 2005. Ashgate Pub-
lishing Ltd., Aldershot, England. 235 pages. $89.95
How would you describe a beaver to someone who
has never seen a beaver? Is it simply a 20 kg rat with
thick fur and with something that has the texture and
consistency of an elephant’s ear sticking out of its back-
side? That person to whom you are giving the descrip-
tion might be forgiven for disbelieving you. What about
a whole forest of new animals including muskrats,
otters, woodchucks, skunks and any of the larger for-
est-dwelling species? In North America we have an
abundance of animals which were unknown in the Old
World before European exploration. Combine these
animals with their South American counterparts and
add birds, reptiles, amphibians, and insects known only
to the New World and you have the basis for the book
given to us from Miguel de Asua.
For the history buff and the technophile who wants
to track down the classic literature of the conquest of
the Americas, this book is a new door to obscure liter-
ature of natural history. Miguel de Asua is an Argentin-
ian professor at the Universidad Nacional de San Mar-
tin and Roger French was English, from the department
of History and Philosophy of Science at Cambridge
University, UK. Their work focuses on the animals
found by early explorers of the Americas, particularly
South America though reference is made to animals
occurring in North America as well. The reader is
introduced to many of the early writers describing the
newly discovered lands, and emphasis is placed on the
animals, birds, reptiles, amphibians and insects which
the authors noted at the time.
From the earliest explorers, Columbus and his fol-
lowers, the discovery of the New World has been a dis-
covery of the beauty and ferocity of many new and
exciting creatures. Columbus returned with birds, skins
and many stories trying to describe the richness of
nature which he found and impress his audiences with
tales of strange and exciting beasts. Written accounts
which survived from his time sought to identify the
animals, birds and reptiles using the experience of
known animals and the descriptions are interesting
associations with European and Asian animal forms
put together to construct forms which would be with-
in the experience of the audience. So bison looked like
camels or cows, armadillos looked like striped foxes
and manatees looked like large puppies.
For two hundred years the animals of the New World
were exotic, unique and objects of wonder in Europe.
Even as the years rolled on with more and more litera-
ture being accumulated from explorers, naturalists and
then colonists with animals imported into Europe as
exotic pets and zoo specimens, the earliest descrip-
tions survived and were repeated. De Asua takes us on
a literature search, naming writers, quoting their writ-
488
ings and showing the parallels as information, disin-
formation and wild speculation was accumulated, re-
peated and refuted.
This is a new study of scientific literature because of
its origins and perspectives. First of all it is centred on
South America with only some brief mentions of North
American mammals. Secondly it is restricted to the
writers first in Spanish then in central Europe, all of
whom wrote in their own language for audiences in
their local areas. Only toward the end of the literature
covered in the book are we introduced to English
sources though the names are obscure here as well. A
good conclusion is added where the ideas of the sailors,
soldiers, functionaries, friars and adventurers each had
their day and their utility in telling a part of the natural
Penguins of the World
By Wayne Lynch. 2007. Firefly Books, 66 Leek Crescent,
Richmond Hill, Ontario L4B 1H1 Canada. 175 pages.
U.S.$24.95. Paper.
If you take one of the most photogenic birds in the
world, living in the most pristine habitat and send forth
a remarkable photographer, can you fail? No!
The hard part about penguins is getting to where
they live. Then anyone can point- and-shoot and get a
fine photo. The appearance of penguins is so charming
that failure is not an option. A quick thumb through the
photos in this book will show how Lynch can take such
a good photo opportunity and make the result mag-
nificent.
But let us look at the text first. In a smooth flowing
narrative the author takes us through the lives of the
world’s 17 penguin species. He covers the origins and
ancestry of this long-lived family. Most intriguing is
Lynch’s description of a five foot seven inch prehistoric
bird. Now that would have been a sight! The author ex-
plains why penguins are shaped the way they are and
the uniqueness of their adaptation to sea life. He des-
cribes their habitat. You may be surprised to realise that
most species live in the temperate [not polar] region.
With humour, he explains their sex lives and the advan-
tages of their marriage and divorce systems. He follows
their lives from egg to adulthood. In all of this he make
these birds sound human. Or is it that we humans are like
penguins — after all they have a much longer lineage?
From sex to family life, from feeding to predators
the author weaves a delightful tale of the delightful
creatures. In addition you will get to know some of the
penguins, sometimes less-than-delightful, neighbours.
Only a photographer as good as Lynch could make a
THE CANADIAN FIELD-NATURALIST
Vol. 120
history story. The opinions and wisdom of the native
peoples of the Americas are not given much consider-
ation because of course they did not write to Europe
and the conquerors formed their own opinions based
on the utility of the natural wildlife to their own needs,
not the inherent worth of the ages of native wisdom.
This is neither an easy book to read nor a simple his-
torical storybook. It is a history of intellectual thought
and discovery. There are many gems of thought in-
cluded but the reader needs to be attentive to the the-
sis of the writer not simply to the many repetitions of
the details.
Jim O’ NEILL
28718 Five Mile Road, Livonia, Michigan 48154 USA
pair of Sheathbills look cuddly. He also adds a chap-
ter on the northern counterpart of penguins, the alcids.
An appendix lists all the penguin species, with a
range map and photo for each one. This is not a field
guide as there is no species description. There is, how-
ever, a small photo of an adult. The range maps and
distribution notes are clear and useful. Lynch notes
the population size and the species status. These range
from a scant 1000 or more pairs of Galapagos Pen-
guins to 10 million Macaronis.
The photographs are wonderful. Many of them are
so balanced in format as to look posed. This speaks of
the infinite patience and possibly some good luck that
Lynch had on his trips. One notable point is the sun-
shine. In my trips to polar regions I rarely see the sun,
yet many of his photos bask in glorious light — more
patience I think. There is a photo of a braying penguin
showing the backward-facing spikes on his tongue in
remarkable detail. There are numerous portraits of
chicks, with and without parents, that are irresistibly
charming.
I have seen 10 out of the 17 living species. To see six
of the remaining species I must visit New Zealand and
the islands that lie to its south, a difficult and very ex-
pensive area to reach. But the lure for a penguinophile
is undeniably very strong. However, if you have never
seen wild penguins and are not likely to do so, then this
book is the next best thing. Buy it and enjoy it, for only
the smell is missing.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6K5 Canada
2006
BOOK REVIEWS
489
Raptors of Western North America and Raptors of Eastern North America
By Brian K. Wheeler. Princeton University Press, Prince-
ton, 41 William Street, Princeton, New Jersey 08540
USA. Hardcover. 544 pages and 430 pages, respectively.
U.S.$29.95.
The hardcover editions of these two volumes were
reviewed in Canadian Field-Naturalist 118(1): 142-143
by Houston and John. These books have now been is-
sued as paperbacks, reducing their cost [U.S. $49.50
Cloth] and weight. There are no updates or additions.
Roy JOHN
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario
K1J 6KS Canada
The Sand Wasps: Natural History and Behavior
By Howard E. Evans and Kevin M. O’ Neill. 2007. Harvard
University Press. Cambridge, Massachusetts, USA. 340
pages. U.S.$49.95 Cloth.
As a baby-boomer “bugster’, two writers had a tre-
mendous effect on my early enthusiasm for insects:
Edwin Way Teale and Howard Ensign Evans. Sadly,
both have passed on, but their legacies live on, and it
was with pleasure that I agreed to review a book on
the sand wasps (the subfamily Bembicinae of what is
now the family Crabronidae, formerly included in the
Sphecidae) coauthored by Evans, and created from his
notes, ably revised and updated by his former student
and colleague Kevin O’ Neill.
As one might expect, this is indeed a book for spe-
cialists. However, it is also clearly written, and a model
for studies of how comparative natural history and be-
haviour can be informed by phylogeny and brought
together in a synthetic fashion. If you have read Wasp
Farm, Evans’ 1963 classic of popular science writing,
you understand what this is all about. If not, the first
two paragraphs of the introduction to Sand Wasps
provide a concise, if not downright brilliant, justifica-
tion for this sort of work. Later, on page 5 in the section
on “Sand Wasp Natural History/Sand Wasp Science”
the authors provide a very fine defense of natural his-
tory in general. The overall purpose of the book is to
update Evans’ previous works on the sand wasps, and
it’s clear that to appreciate all of the details of the pres-
ent volume, one really should have read the works that
preceded it. But having skipped that step, I still found
the book a pleasant and interesting read.
The book begins by putting the sand wasps in con-
text, both with respect to their biology and their clas-
sification. After that, chapter-length summaries are
provided of what is known about the wasps, on a tribe-
by-tribe basis. I found myself getting a bit bogged
down in details during Chapters 2-7, but a student of
sand wasp biology would find them riveting, I’m sure.
My only criticism of this part of the book, which lays
out the evidence to be synthesized in the remaining
chapters, is that no summary is provided of the clas-
sification used, to which one could refer back while
working through the tribal accounts. I also found my-
self wishing for a gallery of illustrations, showing the
various genera of sand wasps side by side. For general
readers from eastern North America, the most useful
portion of these chapters will probably be the account
of cicada killer wasp biology (genus Sphecius), in
Chapter 3.
The synthetic portion of the book begins in Chapter
8, “Comparative Ethology of Sand Wasps.” Patterns
begin to emerge, but so does an overall impression of
great diversity, tremendous complexity, and many un-
answered questions. The theme of phylogenetic recon-
struction enters strongly at about page 260, at which
point I became quite nostalgic, since this was always
my favourite aspect of Evans-school wasp studies in
the past. I found myself wishing that I were reading an
updated account of the evolution of all solitary wasp
behaviour, not just that of one subfamily, but the focus
here is clearly on the sand wasps alone. My interest
was strongly piqued again, however, by the section on
sand wasp conservation. The preservation and promo-
tion of open, erosional habitats is becoming increasing-
ly important to those of us who study organisms that
require such places (yes, I’m thinking here of tiger
beetles), and it is encouraging that sand wasp biology
adds further impetus to this cause.
As a work of entomology, this book is superb. I
doubt many people outside the hymenopterists’ guild
will read it cover to cover, but I do think that various
portions of the text have important implications for
evolutionary biology, entomology, natural history, and
conservation biology. And as a final offering from one
of the great naturalists of the last century, it is a wel-
come book indeed.
JOHN ACORN
Department of Renewable Resources, University of Alberta,
Edmonton, Alberta TST 5L7 Canada
490)
Tracks
By D.C. Jackson. 2006. University Press of Mississippi, 3825
Ridgewood Road, Jackson, Mississippi USA. 279 pages.
U.S.$25. Paper.
Tracks is a collection of 23 short essays, each rang-
ing from 2 to 26 pages. Each essay is built around a
personal story of hunting, fishing, trapping or ram-
blings, and Jackson uses these narratives to provoke
thought and contemplation regarding our interactions
with and place within nature. While I think his goal is
very worthwhile and of great value to anyone interested
in understanding our relationship with the environment,
I found that he was only sometimes successful. Often
the stories are simply entertaining anecdotes, and some
essays miss the mark entirely. This is not to detract from
the compilation however, as a range of success is to be
expected in such a compilation of wide-ranging indi-
vidual stories. When successful, the author is very in-
sightful and provides much food for thought.
Given this objective however, the musings and
philosophies are somewhat confined and repetitive in
that all are focused around stories of exploitative nature
(hunting, fishing, trapping) whereas some of a non-
exploitative nature (camping, hiking, canoeing, drifting,
etc.) would be welcome. Such a broader approach would
illustrate the connection between a contemplative man
and nature in a way that does not require the taking of
life. As is, | am left with the vague unease that to “con-
nect” with nature it is necessary to kill something. Dr.
Jackson is a professor at Mississippi State University
and, given that, I would have appreciated some stories
and deep reflections arising from his time in the field on
his research. It would show the natural environment giv-
ing rise to musings in some other aspect of his life be-
sides the exploitative areas of fishing/hunting/trapping.
THE CANADIAN FIELD-NATURALIST
Vol. 120
Tracks is an interesting book, though I would not
recommend it as a must-read. Sometimes the “philos-
ophy” or musings felt forced, but at other times they
were very appropriate, flowing from the personal anec-
dote. The first chapter “Wilderness Before Dawn”,
really captured my attention and made me hope that I
was in for great insight and original perspective. This
hope, while not entirely fulfilled, was not entirely de-
feated either. These essays are clearly intended for a
broad audience of readers and in many ways refreshed
exquisitely enjoyable past memories for me. I grew up on
stories from the magazines Outdoor Life and Field and
Stream, and the stories here are a cross between that
type of article and a primer on environmental ethics.
As a bridge between outdoor adventure stories and a
contemplative approach to nature it is very successful.
In addition, while I may not have gleaned profound
new insight from the writings, I did learn, and think I
have a better appreciation, and respect for, the Deep
South of the United States over that when I began the
book.
All-in-all Tracks is a pleasant read but I would not
recommend it as delivering particular insight or a
message that we have not heard previously (with the
exception of “Wilderness Before Dawn’). For those
interested in the rural culture of the Deep South (at
least several decades ago), it would be a very useful
book. It is the type of book; I think, which is best read
in front of a warm fireplace on a cold winter night.
SEAN MITCHELL
St. Francis Xavier University, Antigonish, Nova Scotia B2G
2W5 Canada
Why Don’t Woodpeckers Get Headaches? And Other Bird Questions You Know You Want To Ask
By Mike O’ Connor. 2007. Beacon Press, Fitzhenry & White-
side Limited, 25 Beacon Street Boston, Massachusetts
02108 USA. 209 pages. U.S.$14.95.
The book Why Don’t Woodpeckers Get Headaches?
is a collection of letters submitted to The Cape Codder
in the “Ask the Bird folks” column, and answered by
Mike O’Connor who is a bird watcher, the owner of a
birding store, and a born comedian. The letters ask
questions of interest to any backyard birder, ranging
from how to attract Bluebirds, to buying the bird bath
that’s right for you (and your birds) and how to get rid
of a squirrel infestation.
The book is divided into eight chapters, each related
to a particular area of backyard birding. The first sec-
tion deals with bringing birds into your yard. O’Connor
gives several important tips to attract various birds to
your yard, such as having a proper nest box, leaving
food that is suited to the birds you hope to attract, hav-
ing suitable habitat, and most importantly, if you want
these birds to stay in your yard, offering birdhouses
with low rent, an example of the author’s humourous
approach.
Other chapters of the book deal with food issues such
as: Peanut Butter is it safe? What birds can you attract
with oranges? Should you replace your rotten seeds?
Another chapter discusses providing some basic com-
forts for your avian friends: selecting a quality bird
house is obviously a key issue, and is talked about
extensively. It is safe to conclude from the various
entries that, in housing birds, simplicity is key, and
while the four story summer villa with the small pick-
et fence and lawn included may look great at the store,
your birds would probably prefer the cheaper wooden
box. Also, providing a heated birdbath is quite impor-
tant; contrary to popular belief, birds don’t enjoy a chilly
swim in the middle of December any more than we do.
Even though this book mostly deals with attracting
birds, and encouraging their visits, there is an entire
section devoted to bird-related problems. People write
in to ask: Why are some woodpeckers more attracted
2006
to metal chimney caps than the peanut feeder? Why
do birds insist on chirping in a new day at 4:30 in the
morning, right outside their bedroom window? And
of course, the timeless question; How to stop their new
bird feeder from becoming an all-you-can-eat diner for
the cunning gray squirrel? O’Connor provides excel-
lent and thorough advice for his readers, even if he
hasn’t found a sure-fire solution to the squirrel problem.
Why Don’t Woodpeckers Get Headaches is a fasci-
nating collection of people’s backyard bird triumphs
and tragedies. O’Connor is a talented writer, whose
quick wit and edgy humour had this reader crying with
laughter. Not only does he answer readers’ queries with
accurate and detailed opinions, he also provides a brief
BOTANY
Alaska Trees and Shrubs. Second Edition
By L. Viereck, and E. L. Little (with contributions by D. F.
Murray and E. L. Little, Jr.). 2007. Snowy Owl Books,
University of Alaska Press, Fairbanks, Alaska. 265 pages.
US$24.95. Paper.
Measured in Cranberries, Alaska must be among the
wealthiest states in the world. Although this Alaskan
hardcopy publication was printed in China, it was pro-
duced with major U.S. governmental support. A long-
awaited second edition, its underlying data exist else-
where in digital geo-referenced format online. For
some of the most abundant Alaskan species (willows,
over 37 Salix species occur in Alaska) the authors
present text that simplifies relevant details and identi-
fications.
These complex details are all part of Alaska’s wild
reality and a management that is mainly oriented on
commercial goals. The lack of a GAP project in Alas-
ka (otherwise found throughout all states within the
U.S.; see http://gapanalysis.nbii.gov/portal/server.pt)
leaves it a large and harmful heritage; e.g., lack of rel-
evant biodiversity management information and digital
administrative culture. At least the digital data from
this book are available in an older GIS format (ArcGIS
9.0 + Metadata) at the Bonanza Creek LTER website
(http://www.lter.uaf.edu) raising hopes of great things
to come in the future. This matters a lot, because as the
book outlines: “An increasing number of people look
to Alaska for wilderness that is no longer present in
the more developed areas of the world”. It therefore
represents a global natural resource leadership issue
and role model on how we deal and publish on Alas-
ka’s biodiversity and wilderness management compo-
nents that are so crucial to business and human welfare
of the global village.
Together with contributions by D. F. Murray and
G. W. Argus, the authors summarize in this book (first
edition published in 1972) their life-time achievement
presenting up-to-date information on over 14 tree and
115 shrub species. The book offers four identification
Book REVIEWS
49]
background look into the bird's appearance and behav-
iour. The illustrations by Catherine E. Clark are well
done and the birds are easily identifiable.
This book is aimed at readers of all ages, and all
birding-skill levels, from novice to expert. It is well
written and gives a new and often amusing perspective
to the world of back yard birding. Even people who are
not interested in birds will have a great time reading
this book and I would recommend it to anyone. So why
don’t woodpeckers get headaches? Well, if you want
to know, buy the book!
JESSICA GAWN
c/o Mission of Canada, 5 Ave de |’Ariana, Geneva 1202,
Switzerland
keys (Key to Alaska Trees and Tall Shrubs Based Main-
ly on Leaves, Winter Key to Deciduous Trees and Tall
Shrubs of Alaska, Key to Genera of Alaska Shrubs,
Winter Key to Alaska Shrubs). For each of the 132 spe-
cies, distribution maps, a descriptive text (usually coy-
ering leaves, twigs, bark, wood, flowers, capsules, hy-
brids, habitat, distribution, uses and notes), paintings
and photos (44 color plates overall) are presented. |
enjoyed the numerous drawings (mostly taken from
Forest Service publications). The plant distribution
maps present traditional-style, expert-drawn “polygon
monsters” with mathematically exact shapes and mar-
gins but are unlikely to represent biological reality or
relevant landscape gradients. Some of the general spe-
cies text information one might find in other botanical
key references. However, readers interested in Alaska-
nia will appreciate the many species details provided.
For instance “Alaska Athabascans eat the fruits raw
or cooked in moose fat” (for Silverberry), or “Alaska
Indians used the wood for totem poles, dugout canoes,
and houses, and made mats, baskets, and ropes from
the stringy bark” (for Western Red Cedar).
The reader of this book will appreciate the incredi-
ble diversity (and abundance) of willows, birches, and
berries (e.g., Blueberry, Bearberry, Cranberry, Huckle-
berry, Snowberry) in Alaska. But environmental prob-
lems, widely cited in geo-botanical publications, such
as climate change, endemic species loss, road impacts,
overcutting, nitrogen input and general human foot-
print found throughout Alaska are sparsely treated).
Oil development, or pipeline impacts, key features in
Alaska’s landscape and recent history, and discussed
almost worldwide with high-profile books devoted to
this issue (see for instance National Research Coun-
cil of the National Academies 2003), are hardly even
mentioned.
The authors maintain a dubious and ecologically
non-sensitive view that “...the distribution of trees and
shrubs have changed little” since the 1970s. However,
492
at least the changes for Greene Mountain Ash and
Creeping Juniper are described. The concepts of (Land-
scape) Ecology, Biogeography, Fragmentation, Bio-
diversity, and related loss of plant DNA are virtually
omitted in this book. With great regret one has to see
once more that the impact of the Intensive Manage-
ment Law (signed 1994 by Governor Hickel; maximiz-
ing moose numbers through non-science based pred-
ator control, and thus severely affecting browsing on
shrubs and vegetation Alaska-wide) remains virtually
unmentioned. But the impact of ANILCA (Alaska Na-
tional Interests Land Conservation Act) in 1980, and
the number one problem of the Alaskan National Parks,
invasive species, is mentioned (over 100 invasive spe-
cies are referred to, but only 9 are described in this
book; for a better description the reader is advised to
visit websites such as http://www.ucsusa.org/invasive_
species/state-invasion-portfolios.html and http://www.
uaf.edu/ces/ipm/plants/index.html). Many of these spe-
cies were introduced with governmental help, even
by agencies that now have to spend huge amounts of
their budget to deal with these earlier sins. The book
section on Transplanted Native Species I found very
informative (e.g., for Sitka Spruce: “During World War
II, the U.S. military had an active program of plant-
ing trees near bases in the Aleutian Islands.”). Other
environmental topics mentioned are facts like “Much
Alaska cedar is exported to Japan in log form...”, or
for Sitka Spruce: “Low-grade lumber is made into
packing boxes for the Alaska salmon industry”. Keep
in mind that the Tongass and Chugach Forest is the
largest temperate rainforest in the world, with an ex-
tensive network of forest roads reaching more than
10 times around the world! Fortunately, the authors
leave no doubt that more forest exploitation will occur
in Alaska. Such an outlook makes one shiver for Alas-
ka’s future well-being; plants included.
But Cranberry juice apart, “Alaska is a land of con-
trasts in climate, physical geography, and vegetation”
providing a great country for botanists with a thirst to
explore. Besides many endemics, it harbours the Quak-
ing (Trembling) Aspen (most widely distributed tree
in North America) as well as the largest cottonwood
known until 1965. And so, the book is full of fascinat-
ing Alaskan landscape details such as nunataks, ice
fields, permafrost, and some fire history. The promi-
nent botanist Robert Marshall and his historic seedling
experiments are also mentioned. But relevant Russ-
ian, Chukotkan, Kamchatkan and Bering Sea botany
are not further dealt with, nor the links to the Yukon,
British Columbia or the circumpolar Arctic.
It appears a botanical religion that a vegetation clas-
sification always has to be carried out in such botani-
cal books and landscapes. Therefore, this publication
THE CANADIAN FIELD-NATURALIST
Vol. 120
offers 8 main vegetation types and their leading species
for Alaska (published by the U.S. Forest Service 1971;
but relevant quantitative analysis details are not pro-
vided, and one has to trust the experts blindly once
more).
The underlying botanical species taxonomy of this
book is still in flux, as can be seen in the large num-
ber of hybrids, or in the alder, birch and willow species
complexes. Such dynamic and complex taxa are not
easy to deal with by using classic governmental hard-
copy black-and-white publication philosophies (the
authors followed for instance the Flora of North Amer-
ica, Furlow 1997 and Argus 2004). It is here where
readers need more leadership from the experts. Per-
haps presenting links with Genbank (http://www.ncbi.
nim.nih.gov/Genbank/GenbankOverview.html) and ITIS
(http://www.itis.gov/) would start this concept?
The general audience will appreciate the two Alaskan
botanical maps and the four page glossary of botanical
terms. A seven page index of Common and Scientific
Names, and the six pages of literature are of further
interest.
The authors state the contributions from several
agencies and funding institutions, including the U.S.
Forest Service and the Museum of the North Herbari-
um Database. Unfortunately, botany is often equated
with timber, and, as elsewhere, a governmental forest
service focus and its cultural bias can be found in this
book. For instance, from an Arctic tundra perspective,
I am disappointed not to find any relevant references
from S. Walker in this book; the reader is advised to
see online for CAVM http://www.geobotany.uaf.edu/
cavm. But despite the forestry involvement, the authors
still make a nice case that “...the forests of Alaska
provide more than timber to the people of Alaska, and
the rest of the United States”. Well spoken.
This book well describes what Alaska currently has
to offer: wide terrain, many new botanical findings that
are waiting to be put in order (a la: “we sort ’em post-
mortem’’), and much relevant ecology and sustainable
management waiting to be done. Curious naturalists
might also want to compare the state of the art in Alas-
ka with the Flora of adjacent British Columbia (http://
www.eflora.bc.ca). One assumes that the future will
likely see digital online species pages, such as already
started in Wikipedia and coming forward in GAP and
GBIF, with a Biology Google (“Boogle”, or Poogle for
Plants) just being around the corner. This books makes
for a great step towards this achievement for Alaska,
its cranberries and beyond.
FALK HUETTMANN
Biology and Wildlife Department, Institute of Arctic Biology,
University of Alaska, Fairbanks Alaska 99775-7000 USA
2006
ENVIRONMENT
BOOK REVIEWS
493
The Atlas of Climate Change: Mapping the World’s Greatest Challenge
By Kirstin Dow and Thomas E. Downing. 2006. University
of California Press, Berkeley, California, USA. 112 pages.
US$23.95 Paper.
An atlas suggests maps, and maps there are in abun-
dance in this volume, though it also contains many
graphs, charts, diagrams, and informative explanatory
text. Although short, this book is packed with useful
information and well repays careful study. The atlas
is divided into seven sections, each focusing on a dif-
ferent aspect of the climate change story and each pre-
ceded by a one-page introduction. For the first five sec-
tions, topics are illustrated primarily by mapped data,
usually at a global scale, though occasionally focusing
on a specific region, such as the Antarctic. The maps
are usually supplemented by a few paragraphs provid-
ing some additional background or explanation for the
displayed data. The atlas is sprinkled with quotes,
though these are mostly undated, which rather dimin-
ishes their urgency or relevance to the issues.
Part 1, Signs of Change, highlights some warning
signs, such as the European heat wave of 2003, and
then covers three aspects of climate change that set
the context for the rest of the book. These three aspects
comprise glacier retreat, weather-related disasters, and
changes in the polar regions, areas that have been iden-
tified as having particular sensitivity to climate change.
These topics are likely to be familiar to Canadian read-
ers through various recent media stories. Part 2, Forc-
ing Change, focuses more directly on the climate sys-
tem, covering four topics, mostly through charts and
diagrams. Two aspects of the climate system are high-
lighted, explanations for the greenhouse effect and how
climate operates in terms of heat transfer across the
globe. This section also provides a long-term perspec-
tive on climate change, showing the record of carbon
dioxide fluctuations for the last 400 000 years derived
from ice-core records. Finally, there are some scenar-
ios, based on various model projections, showing pro-
jected carbon dioxide increases and related tempera-
ture increases.
The following three sections form the bulk of the
atlas. With six topics, Part 3, Driving Climate Change,
examines greenhouse gas production in more detail,
highlighting several critical aspects of modern indus-
trial activity and linkages to fossil fuels, especially
transportation and agriculture. Covering seven topics,
Part 4, Expected Consequences, paints a grim picture,
showing, amongst other consequences, the impacts of
rising sea levels, water shortages, and threats to human
health. Africa stands out as the region likely to be most
severely affected in the last category. One of the most
interesting maps in this section deals with cultural loss-
es, including historical and archaeological sites. These
kinds of impacts are not often highlighted in discus-
sions of climate change. On the other hand, biodiver-
sity losses and biogeographic changes are given very
little attention. Since alterations in plant and animal
distributions are some of the more obvious and visible
signs of climate change, I found this rather surprising.
For Canadian readers, these sections may induce both
complacency and embarrassment. Complacency be-
cause the maps show that on an absolute basis, Canada
does not contribute as much to the problems as other
more populous industrialized countries. Embarrassment
because on a per capita basis we consume so much
compared to so many countries of the Third World.
Such reflections form a natural transition to the last
sections of the book.
Having provided a gloomy outlook in the previous
section, Part 5, Responding to Change, shows what we
can do about it, or how we might be able to modify the
outcomes at a macro-scale. Eight topics are presented,
most of which are focused on institutional or adminis-
trative responses, such as carbon trading or develop-
ment of alternate renewable energy sources. Finally,
Part 6, Committing to Solutions, shifts the focus from
the global to the local, with some charts showing how
individual actions might affect the trajectory of change.
For those who want the “nitty-gritty” details, Part 7,
Climate Change Data, tabulates various measures of
economic activity and consumption on a country-by-
country basis. These data underpin the maps and graph-
ics in the rest of the book.
Generally the book concentrates on the more obvi-
ous impacts of climate change and the ones that are
often the focus of media attention, such as potential
limits on water and food supplies. As such, most read-
ers are likely to have at least some passing familiarity
with the issues. This book therefore serves as a use-
ful source or companion book for those seeking more
background on current issues. More subtle effects of
climate change, such as water quality degradation, or
cascading effects, such as wildfire frequency changes
or aquifer depletion, are not shown, perhaps because
these are less amenable to measurement or estimation
and mapping. Moreover, the emphasis throughout is
on impacts to the terrestrial realm; other than sea-level
changes, very little attention is paid to marine sys-
tems. Again, this is perhaps surprising given the link-
ages that have been found between, for example, the
ENSO (EI Nino-Southern Oscillation) with rainfall
and droughts.
Cartographically, the book is attractive, clear, and
well-designed. It is apparent that a great deal of thought
and care has been paid to the layout, presentation, and
overall design. In my view, the design is one of the
book’s greatest strengths. Many people are “visual
learners” and this book will definitely appeal to anyone
that prefers graphics to text. Most of the global presen-
494
tations use the same map projection, with Europe and
Africa in the centre and the Americas and Australasia
at the extreme left and right. For North American
readers, this projection can seem strange, since we
are more used to seeing the Americas displayed cen-
trally with other continents on the periphery. Much
of the base information is quite complicated but the
maps show good use of colour and symbols to con-
vey information and summarize complex ideas. One
of the more interesting maps shows greenhouse gas
emissions, with countries’ size distorted according the
amounts emitted. The effect is quite startling, with the
dominance of the industrialized regions (USA, Europe,
CIS, and East Asia) standing out. Perhaps more telling
is the minuscule contribution from Africa, which shows
up as a small polygon, despite its large land area and
population.
Even with the references and sources, I did find my-
self wondering exactly how some of the figures under-
lying these maps are derived, especially because so
many of the data are presented at a regional level, where
regions may include several countries, and are “binned”
into a few categories to simplify cartographic presenta-
tion. Uncertainties or error estimates are not provided
for the data. Inevitably, there must be some fairly broad
assumptions and generalizations underlying the figures.
To take one example, what kinds of measurements
could be available to allow an estimate of methane
emissions on a per capita basis? Are the same or com-
parable measurements, that is, the same data quality,
really available for all areas of the world? How are data
integrated even within one country, given that there
THE CANADIAN FIELD-NATURALIST
Vol. 120
are different ecoregions and uneven population distri-
bution? How much of the disparity shown is real and
how much attributable to varying levels of data avail-
able for different areas of the world? In fact, how far
are the numbers behind the maps realistic estimates
based on reliable instrumentation or simply informed
“guesstimates”? Notwithstanding the beguiling pres-
entation, I did find myself asking many questions about
the methodology behind the mapping.
Perhaps the greatest drawback with this book is that,
because it deals with current and rapidly changing is-
sues, the information presented will quickly become
dated. The authors note that no new material was added
after spring 2006, but they also indicate that they in-
tend to update the volume, especially after the next
Intergovernmental Panel on Climate Change (IPCC)
report, expected in 2007 (page 13). It will be interest-
ing to see how future editions compare to this one and
whether detectable changes in some of the indicators
show up on that time scale. One could argue that such
information is better presented on a website, where it
could be updated as needed. However, having the
information gathered together in book form makes it
easier to consult and compare the different maps. The
book’s greatest advantage is its accessibility, with its
clear presentation and distillation of an enormous
amount of complex information into a readily under-
standable format.
ALWYNNE B. BEAUDOIN
Royal Alberta Museum, Edmonton, Alberta T5N OM6
Canada
An Inconvenient Truth: The Planetary Emergency of Global Warming and What We Can Do
About It
By Al Gore. 2006. Rodale, New York, USA. 328 pages.
US$28.95 Paper.
I freely admit that I picked up this book already pre-
disposed to dislike it. After all, it’s supposedly a book
about climate science by a politician, so how good
could that possibly be? I am relieved to report that this
volume pleasantly surprised me; the book is consider-
ably more readable and worthwhile than I expected it to
be. Surprisingly, it also projects, an air of intelligence
and careful consideration. Yes, the book is a call to
action and a strong statement of a particular point of
view, but the hype and hyperbole are at a lower volume
than I had feared. More importantly, the book is largely
based on solid science and the presentation of that sci-
ence is done in a remarkably straightforward fashion.
In his introduction, Gore says that the book was
developed from a slide show and, judging by the lay-
out and design, that presentation mode has carried over
and set the book’s style. It is arranged as a continuous
narrative, with clear graphs and graphics, some on fold-
out pages, colourful eye-catching photographs, short
sentences and paragraphs, and large type. Graphics are
sourced, though complete citations are not provided.
Text and explanatory captions are generally written in
plain conversational language. The book surveys the
evidence for global warming and outlines the main
forcing factors, highlighting the importance of green-
house gases and the role of human activities in the in-
creasing rate of, especially, carbon dioxide content in
the atmosphere.
Gore drives home the message that human activities,
in particular industrial activities, are responsible for
much of the recent increase in greenhouse gas concen-
trations in the atmosphere and therefore the increasing
rate of global warming. He sets these trends in context
by showing, for example, recent rates compared to the
carbon dioxide trend inferred from Antarctic records
extending back to 600 000 years before present. Other
long-term records, such as tree-rings and Greenland
ice-cores, are also used to provide perspective. Several
times, the point is made that it is not simply the absolute
amount of increase that is important but the increasing
rate of increase. Taking it as given that global warming
will continue, even if actions are taken now to reduce
rates of carbon dioxide emissions, Gore then exam-
2006
ines what the implications are likely to be during the
next few decades.
The book presents many different types of evidence
for global warming and covers a lot of territory. It
draws on data and images from across the world. Gore
amasses a formidable array of information to support
his argument. Some of the topics he features include:
droughts and shrinking lakes (including an intriguing
sequence of images showing the reduction in Lake
Chad), reduction in Arctic ice, shrinking of large ice
sheets (notably on Greenland and Antarctica), effects
on temperatures and seasonality, impact of rising sea-
levels (including the inundation of islands), damage
to coral reefs, effects of climate on human health and
diseases rates, frequency and occurrence of extreme
weather events (including, not surprisingly, hurricanes),
and climate impacts on biodiversity and distribution of
plants and animals. The evidence for global warming
includes some striking “then and now” photographs of
glacier forefields showing recent ice retreat. Two issues
that also stand out are the effects of exceptionally high
summer temperatures, as seen in recent years in Europe
especially, and increased storminess, especially on low-
lying islands and coastal areas. The evidence and impli-
cations are shown from a global and not just a U.S.
perspective. This worldwide coverage underlines the
urgency of the issues presented, since some of the im-
pacts are not just hypothetical but are being felt by
people across the world now.
This leads Gore into a consideration of the roots of
the problem, in particular the main sources of green-
house gas emissions. Some striking graphics visually
emphasize the large contribution to greenhouse gas
emissions made by the U.S. Gore points out some of
the global inequities in consumption that have led to
this situation and militate against any remedial action.
He identifies energy production as the main culprit and
highlights the need to move to alternate energy sources,
such as wind power. Here, the text gets more overtly
political with Gore taking aim at the Bush administra-
tion for obfuscating the issue and refusing to act, for
example, by not ratifying the Kyoto Protocol.
Interspersed within the text are eight short essays,
most of which recount anecdotes and incidents from
Gore’s life. These stand out from the rest of the narra-
tive by being printed in smaller font on tan-coloured
pages. The essays highlight some of the reasons for
his passionate concern about global warming. What
they show is that this is not a recent preoccupation but
something that has engaged his attention for many
years, growing out of life-long interest in environmen-
tal issues. Clearly, he wants to let readers know that
he is not simply “jumping on a bandwagon” and taking
BOOK REVIEWS
495
up a trendy issue out of opportunism, but is following
an interest that has underpinned his political career.
Here, I did feel that I was reading a political manifesto
at times. There’s a lot of name-dropping and there are
rather too many “warm and fuzzy” pictures of Gore
and his family for my taste. Nevertheless, these essays
bring the issue from the abstract to the personal, which
does sharpen the sense that these concerns matter to
individuals. By being set off from the rest of the text,
it is clear that the message in these essays differs from
the rest of the book.
The final section of the book, offset by being print-
ed on light-green paper, is the call to action, outlining
some things that individuals can do to make a differ-
ence. Most of these actions will already be familiar to
most readers: buy a hybrid car, bag your groceries in
re-usable shopping bags, replace incandescent light-
bulbs with compact fluorescent bulbs, and so on. Gen-
erally, these actions are an appeal to a sense of guilt
in affluent people living in western countries, where,
Gore maintains, most of the problem originates any-
way. Some of the suggestions run counter to many
driving forces in the economy. When the TV screen is
filled with ads encouraging us to be tourists and travel
to sunny beachside resorts, how realistic is it to tell peo-
ple to reduce air-travel? Interspersed among these ex-
hortations are ten common misconceptions about glob-
al warming and Gore’s recommended responses. These
misconceptions include the ones you might expect,
such as the assertion that “it’s all just natural variation”,
though there was one I hadn’t heard before, attributing
global warming to after-effects of the Tunguska event.
In such a brief statement of the global warming
issue, Gore obviously does not get into any topic in
detail. There’s clearly a lot more that could be said
about all lines of evidence he presents but, unfortu-
nately, there is no reading list for readers to follow up
on any topics raised in the book. Nevertheless, Gore
does assemble a persuasive argument that global warm-
ing is a problem and we need to do something about it
immediately. However, this is very much a book that
is “preaching to the converted”. I can’t see many peo-
ple who are not already convinced there is a problem
reading it. On the other hand, Gore is a well-known
figure so perhaps some people may take a look at this
book because of name-recognition. As a politician,
Gore has to overcome a good deal of cynicism in order
to reach people. This is a pity because this book is an
articulate and thought-provoking presentation of a
global phenomenon that affects us all.
ALWYNNE B. BEAUDOIN
Royal Alberta Museum, Edmonton, Alberta TSN OM6 Canada
496
MISCELLANEOUS
THE CANADIAN FIELD-NATURALIST
Vol. 120
Gilbert White: A Biography of the Author of The Natural History of Selborne
By Richard Mabey. University of Virginia Press, Char-
lottesville. 2006. 239 pages. U.S.$16.50 Paper.
In his introduction, Mabey admits that, at his first
reading of Gilbert White’s The Natural History and
Antiquities of Selborne, he could barely cope “with its
rambling disorder.” Only on visiting Selborne in per-
son did Mabey change his view decisively. I marvel,
as Mabey did, that White “could produce something
so wholly original and appealing out of such unpromis-
ing ingredients.”
In this Whitbread Prize-winning biography, the pain-
staking research of Mabey — whom the Times has called
‘Britain’s foremost nature writer’ — results in a remark-
ably detailed account of the life of Gilbert White. Born
in the tiny village of Selborne, White obtained his uni-
versity education in theology at Oriel College, Oxford,
beginning in 1739. Here he met John Mulso, in the
year behind him. Although “constitutionally lazy and
hypochondriacal,’ Mulso’s life-long correspondence
to and from White became a major source for Mabey.
Perhaps it was equally providential that White was not
particularly successful in his first three appointments to
curacies at a distance from his home. When he returned
to Selborne he began, somewhat inauspiciously, to com-
mit his nature observations to paper in 1756.
White was patient and inquisitive. His early descrip-
tion in 1761 of the life of crickets was both “vivid and
sensuous.” In 1767 he began writing to the naturalist
and author, Thomas Pennant; two years later he began
A Paddler’s Guide to Quetico and Beyond
By Kevin Callan. 2007. The Boston Mills Press, Erin, Ontario
Canada. 192 pages. U.S.$24.95 Paper.
It’s the dead of winter on a Friday evening when
friends lay out the maps on the floor in front of a burn-
ing fire. Over wine, possible routes are discussed and
commitments are made for a canoe trip six months
hence into one of the most intact, accessible and largest
protected wilderness areas in North America: Quetico
Provincial Park. This is a book that will lead you beyond
your usual canoe routes; in fact, it may inspire you to
plan a trip celebrating Quetico’s 100" anniversary in
2009!
Kevin Callan is no stranger to writing books on
wilderness canoeing with seven “‘Paddler’s Guides” for
Ontario canoe routes to his name. Kevin reminds us
that there is a great story to be told after every canoe trip
we make, as a result of spending quality time with a
group of family or friends on a journey through the
natural world of wind, water, wildlife and plants, com-
plete with portages, bannock and fireside stories. Each
of the sixteen canoe routes described in this book is a
well-written informative story, weaving important prac-
his original observations that helped prove that birds
migrate. In 1774, White’s lively letters to Sir Daines
Barrington included one about house martins, which
Barrington read to a meeting of the Royal Society. The
result was publication of these “models of lucidity and
insight” in Britain’s earliest scientific journal, Philo-
sophical Transactions.
Finally, late in 1788, at age 68, White had the sat-
isfaction of seeing his book in print. He died in June
1793, a month short of his 73" birthday, modestly un-
aware of the prominence his book would achieve in
the centuries to come. Second and third editions did not
appear until 1802 and 1813, respectively, but there have
been more than two hundred subsequent editions, one
of the most published books in the English language.
I appreciated Mabey’s delightful biography all the
more because I, too, have visited Selborne. By learn-
ing more about Gilbert White, we can better appreci-
ate White’s impact on all subsequent nature writing. I
congratulate the University of Virginia Press for mak-
ing this inexpensive reprint, first published in hard
cover in 1986, readily available to North Americans.
I heartily recommend it to anyone with the slightest
interest in English literature or the history of natural
history.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N 0J8
Canada
tical information such as the take-out and put-in spots
for portages, special sites to see, and danger spots —
into historical accounts of those who were here long
before our forays using Kevlar canoes and Gortex.
The stories also convey the hardships and joys shared
by the group traveling with Kevin; the “real stuff”
that makes canoe trips memorable.
The book does not overwhelm the reader with un-
necessary detail on the gear to bring, what kind of canoe
paddle is better than another, etc., rightly assuming
someone heading to Quetico is not a novice. At the
same time, simple, straightforward descriptions of
canoe routes make them immensely doable for the aver-
age paddler. Exceptional pictures and tidy maps are
also included, with references for the “proper” maps a
canoeist must purchase in order to undertake these trips
Sixteen canoe routes are described in Quetico and
Beyond; eleven in Quetico Park and five within neigh-
boring areas of northern Ontario. Most routes require
about one week to complete, although some are as
short as 2-4 days and others as long as 28 days. Sum-
maries provided for each of the canoe routes includes
2006
number of days required to complete the route, the
number of portages, the longest portage, route difficul-
ty, access points, and the maps needed. Scanning the
summaries provides a quick method to narrow down
which routes look the most interesting within the time
frame and abilities of the group.
Quetico is a book that provides very readable pleas-
ant conversation that, in a personal way, brings canoe-
BooK REVIEWS
497
ing to life through its past and more recent adven-
tures. The many reminders of the subtle rewards of
canoeing that are conveyed provide a real enticement
to begin planning your next trip.
BRENT TEGLER
North-South Environmental Inc., 35 Crawford Crescent,
Unit BUS, Campbellville, Ontario LOP 1B0O Canada
More than Kin and Less than Kind: The Evolution of Family Conflict
By Douglas W. Mock. 2004. Belknap Press of Harvard Uni-
versity Press, Cambridge, Massachusetts, USA. 267 pages.
U.S.$27.95.
If you wish to understand how complicated and
how variable reproductive behaviours can be, this is
the book! Doug Mock offers a wide vista of nature in
all its infinite variability and complexity. To explain
bird behaviour, Mock invokes human examples, as
diverse as his three older brothers, chess, Wall Street,
sports teams, and vote counting.
Mock offers creative chapter headings and begins
each chapter with a brilliant quotation. He simplifies
difficult-to-understand concepts in an instructive and
often entertaining manner. As he says, this book is a
“mixture of theory and data.” He is especially inter-
ested in brood size and in siblicide. I will offer a few
highlights, some of which mention Canadian research.
Mock attempts to explain both the “hows” and
“whys” of concepts that include Darwinian fitness,
evolutionary game theory, inclusive fitness theory, evo-
lutionarily stable strategy, parent-offspring conflict,
parental manipulation, optimal clutch size, replacement
offspring, and the insurance egg hypothesis.
Reproduction is costly. When Collared Flycatcher
broods were enlarged artificially, there were fewer
breeding adults the following season, and when two
eggs were removed, females laid larger clutches the
following year. Seychelles Warblers produced 77%
male chicks in poor habitat and 88% female chicks in
good habitat.
Lifetime monogamy is uncommon among birds but
there are notable exceptions. A sample of 919 pairs of
Bewick’s Swans showed not a single divorce, but if a
mate died the survivor would re-pair. From a sample
of over 6 000 banded birds, 99.6% of Barnacle Goose
pairs were socially monogamous.
Mock discusses Hamilton’s rule, promulgated in
1964, whereby “two full siblings should be the evo-
lutionary equivalent of one Self’ — or eight cousins —
based on the amount of shared genetic material. Altru-
istic behaviour, such as nest helpers, relates to the mix
of relatedness.
What happens in nature often seems counter-intu-
itive if not outright bizarre. Biologists have difficulty
understanding, much less explaining, such events. What
conceivable evolutionary advantage could result from
a surplus egg or surplus young? Why do some species
of eagle and pelican regularly lay two eggs, yet raise
only one young? Why does a parent bird passively
watch one of its nestlings kill another, even when sur-
plus food is within reach? Why does a hawk nestling
sometimes eat its sibling after killing it, but on other
occasions does not?
At Delta Marsh, Manitoba, Spencer Sealy video-
taped nests with a single cowbird and a single warbler;
the cowbird out-hustled, out-begged and out-stretched
its warbler nestmate. In British Columbia, Bruce Lyon
studied American Coots, which lose some chicks to
starvation in nearly half the broods. Adult coots make
a point of getting more food to the youngest, but when
Lyon clipped the bright ornamental plumage of the tiny
chicks, the clipped birds survived less often.
The late Roger Evans of the University of Manitoba
did experiments to study the role of the extra or “insur-
ance” egg in the American White Pelican. He and his
student Kevin Cash carefully marked the first-hatching
or A egg, and the second-hatching or B egg. Twenty
percent of the B eggs hatched. The parent pelican would
brood its young, but as soon as the parent stood up, the
older young attacked its smaller sibling.
Wahlberg’s Eagle, which ranges across central Africa,
lays a single egg; not more than three percent of pairs
lay a second egg and then the larger chick regularly
kills the smaller one.
The Verraux’s (Black) Eagle lays two eggs, but an
observed first chick began assaulting the second chick
soon after it hatched, giving 1 569 blows with its beak
to kill its sibling, even though food was plentiful in the
nest. The Crested Penguin lays two eggs, but the first
egg is up to 40 percent smaller; four days later the sec-
ond, larger egg is laid and is given the optimal brood
patch position beneath the mother, and is usually the
sole survivor.
Mock does not restrict himself to birds. He writes
about plants, insects and fish, using the firefly as an
example of signalling. He tells how, in a pig litter, ante-
rior teats produce more milk than posterior teats, hence
the colloquialism, “sucking hind teat.”
As Marlene Zuk says on the dust cover, this book
about family conflict “is a model of how behavioral
ecology can and should be done ... Just don’t take it
to family reunions.”
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N OJ8
Canada
498
To See Every Bird on Earth
By Dan Koeppel. 2005. Hudson Street Press, New York,
New York, USA. 278 pages. $24.95 U.S.$35.
Dan Koeppel tells the story of his father, Dr. Richard
Koeppel, a chronically unhappy medical doctor with
two overwhelming obsessions: to see as many species
of birds as possible, world-wide, and to read, cover to
cover, every novel that was short-listed for or won the
Booker Literary Prize.
Each chapter begins engagingly with a postage
stamp that features a bird from the collection of Chris
Gibbins who has amassed stamps depicting 2950
species (a record list!), and a paragraph from Richard
Koeppel’s notebooks telling of a numbered addition
to his life-list.
One learns about the methodology of listers, par-
ticularly the rules laid down by the American Birding
Association (pages 195-197). Dan explains, in layper-
son’s terms, “lumping” and “splitting” of species.
Despite the ever-escalating CPB (cost per bird), Dan
fully realizes that the listing process is “intensely
arcane, fascinating, and absurd.”
This is a quick and easy read, but in no way uplift-
ing. Dan’s brutally frank, very personal psychoanaly-
sis of his father and mother is somewhat excessive and
distracting. Dr. Richard Koeppel, it seems, entered med-
icine to please his parents, and sadly failed to find a
fulfilling niche in medicine. His marriage failed early.
He became a morose, rather pathetic man who found
solace in excessive use of marijuana, alcohol and nico-
tine. Most of his career was in emergency medicine,
where roughly half his time could be spent birding.
When his bad habits caught up with him in the form
of larynx cancer, Richard Koeppel’s personal list was
at 7080 species.
I most enjoyed the behind-the-scenes accounts of
Jim Clements, who compiled the first one-volume list
of birds of the world, and has amassed a personal life
list of 7200; Victor Emmanuel, who began the first
field trips designed to add maximum numbers of birds
to each customer’s life-list; and Bret Whitney, who
lacks formal postgraduate education in ornithology but
has succeeded the late Ted Parker as a finder and des-
criber of new species in South America.
One also meets Richard’s competitors. Within our
region, North and Central America, Dan Canterbury
Wildfire in the Wilderness
By Chris Czajkowski. 2006. Harbour Publishing, Madeira
Park, British Columbia, Canada. 221 pages. $19.95 Paper.
A woman who lives a solitary life in the wilderness
of the British Columbian mountains, dozens of kilo-
metres from her nearest neighbours, has not isolated
herself from her community. Her life is a heart-warm-
ing Set of tales explaining the connection she has to the
community of mountain dwellers, outfitters, and bush
pilots in her immediate neighbourhood as well as
THE CANADIAN FIELD-NATURALIST
Vol. 120
has seen a record 1731 species. Peter Kaestner, who
traveled widely in the diplomatic service, was the first
person to have seen a representative of all 159 avian
families. Others in competition with Richard Koeppel
for the world list have been Harvey Gilston of Britain,
who, without ability to identify the bird himself, would
check off a bird when a guide called out its name (he
reached nearly 8000); Michael Lambarth and Sandra
Fisher of England (Michael quit when his beloved part-
ner died); Joel Abramson, a medical doctor who hired
top birders to lead endurance-test expeditions; Stuart
Keith, founder of the American Birding Association,
who was recognized by the Guinness Book of World
Records when his list reached 4300; Roger Tory Peter-
son, who reached the “half-way total” of 4800 in the
1980s; Bill Rapp, nearing 7000; Jim Plyler, a retired
oil executive at 7200; Peter Winter and John Danzen-
backer, military men, at 7800 and about 7750, respec-
tively; Peter Kaestner, with 7958; Tom Gullick, leader
of those still alive, with 8114; Phoebe Snetsinger, still
tops, who reached nearly 8500 before she died in a
car accident in Madagascar in 1999.
What are the downsides to this book? The title is a
bit misleading, since Dan Koeppel knows full well that
no human will ever see every bird (of about 9600
species) on earth. An index to birder’s names would
have been helpful. Worse, Richard Koeppel, the protag-
onist of this story, is a sad-and pathetic person, lacking
enthusiasm for his profession, with complicated mar-
ital and family situations. He is a perfectionist but
only on rare occasions does he show much feeling,
even for birds. But Richard has made amends with his
son, Dan, and has recently turned his attention to but-
terflies.
All readers of this review, especially the writer, are
sinners. We claim to be environmentalists, yet many or
perhaps most of us drive gas-guzzling vehicles to pur-
sue our sometimes obsessive hobby. At least we fall
short of the obscenely high travel expenses of certain
of the Big Listers.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N O0J8
Canada
friends, publishers, fans of her books and international
tourists in the wider community. The reader is intro-
duced to her dogs, her history of publications, her inge-
nuity in building and maintaining a wilderness site and
even the state of her bank account while following her
stories of maintaining solitude in the mountains and
contacts with the outside world.
A hiker, camper and outdoorsman would certainly
relate to Chris Czajkowski’s stories and her descriptions
2006
which make the mountains a home. The well-being of
the environment becomes close to the heart of any
dweller of the wilderness and the threat of nature tak-
ing a turn for the worst can send dwellers of the back-
woods to worry over their personal charge, the nature
and landscape around them. Sustained inclement
weather, freak storms, too much snow in a season, frost
at the wrong time, drought in the wrong season or pro-
longed drought in any season, are all causes for worry
but a natural disaster like a wildfire is feared above all.
Though the title tells us only of one dramatic event in
the history of her lake, cabins and near environment,
the reader is drawn into her whole lifestyle of pro-
tecting, nurturing and living within the environment
which she has come to call home.
As the story unfolds, Czajkowski leads us into her
world, travelling long distances, keeping up with mod-
ern technology, camping and hiking with her tourist
helpers. She is home in her particular isolated part of
the world a long way from her nearest neighbours but
connected with them by regular radio conversations,
listening to radio broadcasts and occasionally eaves-
dropping on her neighbours’ conversations, and finally
email as that technology became available. At the same
time, she is host to the world by operating a small guest
house for people willing to travel the distance to spend
a few days with her, or international wanderers who
pay by passing a few weeks working and helping her
with projects.
Finally the reader is led to the hot dry summer of
2004 and the devastating fire, the Lonesome Lake Fire
which was well reported to the Canadian public as one
of many fires in a hot dry summer, but to people of
YOUNG NATURALISTS
Tale of a Great White Fish
By Maggie de Vries. Greystone Books, Douglas & McIntyre
Publishing Group, Vancouver, British Columbia, Canada.
34 pages. $12.95.
The book Tale of a Great White Fish follows the
story of a giant white sturgeon. Through its long life,
the sturgeon overcomes many obstacles in its battle to
survive and reproduce. The story deals with the inter-
action between sturgeons and people, along the way
providing the reader with a wealth of information and
insight into the natural history of the sturgeon.
The story starts in 1828, when the sturgeon, named
“Big Fish,” was a small and defenceless egg hatched
in the Fraser River. She matures into a larva and then
into a fry. As a fry she must eat thousands of other
small water creatures to grow. Fry are also especially
vulnerable to predators, and most of the young fry
never reach adulthood. The fry that survive mature into
adults before returning to their spawning ground to lay
their own eggs.
There are many dangers that the sturgeon must over-
come. For most of their long existence sturgeon have
BooK REVIEWS
499
that remote region, a disaster which shaped their envi-
ronment for years to come. The reader is invited to
travel with Czajkowski through the fear and uncertain-
ty of watching gathering smoke and distressing news
from various watchers in neighbouring valleys. We are
inspired to evacuate with her to safer areas, but with-
out any assurance that home or lifestyle could be pre-
served. Czajkowski leads us along her tale with inter-
esting anecdotes and humorous stories about her dogs,
friends, and people who come to visit her. She is an
engaging story-teller and keeps the reader’s attention
for hours or just a few minutes at a time as the book
is filled with short stories, portions of her diary and
longer accounts of specific incidents.
The story builds in an entertaining way through her
history of building the cabins, receiving visitors, includ-
ing facts about making one’s life in the wilderness at
the same time maintaining connections firmly planted
in the outside world. The fire is the high point of the
book and gets the reader to the excitement of living her
experience of the fear, doubt, loneliness and response
of care for her own animals and the natural dwellers of
her wilderness world. The reader enters into the story
with her and can live the excitement she feels. In our
sheltered world, our decisions are not always the same
as the author’s but she puts her concern and care for
her environment in front of the reader and shows us a
different world to enjoy for a little while then look
forward, like her globe-trotting visitors, to the next
visit we might have with her.
Jim O° NEILL
28718 Five Mile Road, Livonia, Michigan 48154 USA
been around since the time of the dinosaurs, the threats
have been natural: predators, natural calamities and
disease. Today, human fishermen and the caviar indus-
try are one of the greatest perils. Big Fish was hooked
by a fisherman, but fortunately she was freed by some
young boys and was able to escape. During the Hells
Gate Slide in 1913 many migrating sturgeon were
killed. Big Fish survived, but she was forced to find a
new place to spawn. Later, Big Fish had to find a new
home because the lake she was living in was drained.
Later still, an unknown disease swept through the Fras-
er River and many sturgeon died and were washed up
on the shores of the lake. When the story closes, almost
a hundred years later, Big Fish is still surviving, but
continues to face many challenges.
Scientists study sturgeons to help them survive. They
keep track of their weight and size, and try to stop over-
fishing. There are now laws which prevent the fishing
of these magnificent animals.
The book is written as a story and at the end there is
a useful list of sturgeon facts, a labelled diagram of a
500
White Sturgeon, a glossary, and a letter from Rick
Hansen, a famous Canadian para-athlete. Rick Hansen
is the chairman of the Fraser River Sturgeon Conser-
vation Society. The drawings by Renné Benoit are done
in water colours and they provide a magnificent glimpse
into the beautiful and dangerous world of the great
White Sturgeon.
The book is very informative. In it, the reader will
learn a great deal about the White Sturgeon’s envi-
ronment and behavior and appearance. The Sturgeon
Facts and a glossary at the book of the book provide
NEw TITLES
Prepared by Roy John
+ Available for review * Assigned
ZOOLOGY
Bird — The Definitive Visual Guide. Edited by Peter Frances
et al. 2007. Dorling Kindersley, DK Enquiries, Tourmaline
Editions Inc., 662 King Street West, Suite 304, Toronto,
Ontario, MSV 1M7. 512 pages. GBP 25.99 Cloth.
The Black Bear Almanac. By David Smith. 2007. Globe
Pequot Press, 246 Goose Lane, P.O. Box 480, Guilford, Con-
necticut 06437 USA. 288 pages. U.S.$29.95.
* The Clements Checklist of Birds of the World (6" edi-
tion). By James F. Clements. 2007. Cornell University Press,
Sage House, 512 East State Street, Ithaca, New York 14850
USA. 864 pages. U.S.$59.95 Cloth.
The Biology of African Savannahs. By Bryan Shorrocks.
2007. Oxford University Press, New York, New York. 268
pages. GBP 27.50 Paper.
Arctic Flight - Adventures Amongst Northern Birds. 2007.
James McCallum. Langford Press, 10 New Road, Langtoft
Peterborough, UK. 180 pages. GBP 37.99 Cloth.
Endangered. By George C. McGavin. 2007. Cassell Ilustrat-
ed, 2-4 Heron Quays, London E14 4JP. 192 pages. U.S.$35.
* Birds of Europe, Russia, China, and Japan: Passerines:
Tyrant Flycatchers to Buntings. By Norman Arlott. 2007.
Princeton University Press, 41 William Street, Princeton, New
Jersey 08540 USA. 240 pages. U.S.$29.95 Paper.
Feathers: Identification for Bird Conservation. By Marian
Cieslak and Bolestaw Dul. 2007. Natura Publishing House,
ul. Olbrachta 118c/20, 01-373 Warszawa, Poland. 320 pages.
GBP 25.62.
Forensic Entomology: An Introduction. By Dorothy Gen-
nard. 2007. John Wiley & Sons Canada Ltd., 6045 Freemont
Boulevard, Mississauga, Ontario LSR 4J3 Canada. 244 pages.
U.S.$55 Cloth.
Homalopsid Snakes, Evolution in the Mud. By J. Murphy.
2007. Krieger Publishing, P.O. Box 9542, Melbourne, Florida
32902-9542 USA. 260 pages. U.S.$68.50 Cloth.
The Inner Bird — Anatomy and Evolution. By G. Kaiser.
2007. The University of Washington Press, P.O. Box 50096,
Seattle, Washington 98145-5096 USA. 464 pages. U.S.$85
Cloth.
Insects of Britain and Western Europe. By Michael Chinery.
2007. A & C Black Publishers Ltd. (United Kingdom), 37 Soho
Square, London, W1D 3QZ. 320 pages. GBP 14.99 Paper.
THE CANADIAN FIELD-NATURALIST
Vol. 120
a very useful summary, and is well presented.
The book is aimed at a child’s reading level. Older
readers would find this book less interesting; howey-
er, it provides a basic insight into the world of a great
white fish and is nicely illustrated. This book is rec-
ommended for any child with an interest in the out-
doors and in nature.
JESSICA GAWN
c/o Mission of Canada, 5 Avenue de |’ Arianna, Geneva
1202, Switzerland
Insect Conservation Biology. Edited by A. J. A. Stewart, T.
R. New and O. T. Lewis. 2007. Oxford University Press, 198
Madison Avenue, New York, New York 10016 USA. 480
pages. U.S.$190 Cloth.
Australia’s Mammal Extinctions. By Chris Johnson. 2007.
Cambridge University Press, 40 West 20" Street, New York,
New York 10011-4221 USA. 278 pages. GBP 34.99 Paper.
Owls of the United States and Canada. By W. Lynch. The
University of Washington Press, P.O. Box 50096, Seattle,
Washington 98145-5096 USA. 256 pages. U.S.$44.95 Cloth.
Spotted Owls. By J. Hobbs. 2007. Greystone Books, Suite
201, Quebec Street, Vancouver, British Columbia V5T 4S7
Canada. 144 pages. $39.95 Cloth.
Birds of Pakistan. By Richard Grimmett and Tim Inskipp.
2007. Helm Field Guides. 256 pages. GBP 24.99 Paper.
Birds of Peru. By Daniel F. Lane, John P. O’ Neill, Theodore
A. Parker II, Thomas S. Schulenberg, and Douglas F. Stotz.
2007. Helm Field Guides. 656 pages. GBP 29.99 Paper.
Raptors: A Field Guide to Survey and Monitoring. By J.
Hardey, H. Crick, C. Wernham, H. Riley, B. Etheridge and D.
Thompson. Stationery Office (TSO), 71 Lothian Road, Edin-
burgh EH3 9A2 UK. Scotland. 300 pages. email: esupport@
tso.co.uk
*Reptiles and Amphibians of Canada. By Chris Fisher
and Ronald Brooks. 2007. Lone Pine Publishing, 10145 — 81
Avenue, Edmonton, Alberta T6E 1W9 Canada. 208 pages.
$18.95.
* Rodent Societies. By J. Wolff and P. Sherman (Eds). 2007.
University of Chicago Press, 1427 East 60" Street, Chicago,
Illinois 60637 USA. 610 pages. U.S.$49 Paper, $125 Cloth.
The New Encyclopaedia of Snakes. By Christopher Matti-
son. 2007. Cassell Illustrated, 2-4 Heron Quays, London
E14 4JP. 272 pages. GBP 30 Cloth.
Native Trees of the Southeast — An Identification Guide.
By L. Katherine Kirkman, Claud L. Brown, and Donald J.,
Leopold. 2007. Timber Press, 133 SW 2nd Avenue, Suite
450, Portland, Oregon 97204 USA. 372 pages. $45.95.
Waterbirds Around the World. By Gerard Boere, Colin
Galbraith and David Stroud [Editors]. 2007. TSO (The Sta-
tionery Office), 71 Lothian Road, Edinburgh EH3 9A2 UK.
Scotland. 940 pages. GBP 50.
2006
The Last Wild Wolves. By I. McAllister. 2007. Greystone
Books, Suite 201, Quebec Street, Vancouver, British Colum-
bia VST 487 Canada. 192 pages. $45 Cloth.
+ Whales and Seals — Biology and Ecology. By P. H.
Fontaine. 2007. Schiffer Publishing Ltd., Lower Valley Road,
Atglen, Pennsylvania. 448 pages. U.S.$34.95.
* Why Don’t Woodpeckers Get Headaches? By M.
O'Connor. 2007. Fitzhenry and Whiteside Limited, 195 All-
state Parkway, Markham, Ontario L3R 4T8 Canada. 209
pages. $14.95.
ENVIRONMENT
The Sacred Balance. By D. Suzuki. 2007. Greystone Books,
Suite 201 Quebec Street, Vancouver, British Columbia VST
487 Canada. 320 pages. $22.95 Cloth.
Scaling Biodiversity. By David Storch, Pablo Marquet and
James Brown. 2007. Cambridge University Press, The Edin-
burgh Building, Shaftesbury Road, Cambridge, CB2 8RU.
GBP 38.
Climate Change. Edited by J. DiMento and P. Doughman.
2007. The MIT Press, 55 Hayward Street, Cambridge, Mas-
sachusetts 02142 USA. 232 pages. U.S.$19.95 Paper.
What We Know About Climate Change. By K. Emmanuel.
2007. The MIT Press, 55 Hayward Street, Cambridge, Mas-
sachusetts 02142 USA. 96 pages. U.S.$14.95 Cloth.
BooK REVIEWS
501
Saving Planet Earth. By Tony Juniper. 2007. BBC Books,
Random House, 1745 Broadway, 3rd Floor, New York, New
York 10019 USA. 256 pages. $39.95 Cloth.
+ Foraging — Behavior And Ecology. By B. Stephens, J.
Brown and R. Ydenberg. 2007. University Chicago Press,
1427 E. 60" Street, Chicago, Illinois USA. 608 pages. U.S.
Cloth.
The Great Lakes — The Natural History of a Changing
Region. By W. Grady. 2007. Greystone Books, Suite 201,
Quebec Street, Vancouver, British Columbia VST 487 Cana-
da. 320 pages. $49.95 Cloth.
MISCELLANEOUS
* Beneath My Feet: the Memoirs of George Mercer Daw-
son. By Phil Jenkins. 2007. McClelland & Stewart Ltd., 75
Sherbourne Street, 5th Floor, Toronto, Ontario MSA 2P9
Canada. 368 pages. $34.99 Cloth.
Modelling for Field Biologists and Other Interesting Peo-
ple. 2007. By Hanna Kokko. Cambridge University Press,
32 Avenue of the Americas, New York, New York. 230 pages.
GBP 27.99.
The World Without Us. By Alan Weisman. 2007. St. Martin’s
Press, 175 Sth Avenue New York, New York 10010 USA.
324 pages. U.S.$24.95 Cloth.
News and Comment
Marine Turtle Newsletter (114)
October 2006. 32 pages: EDITORIALS: Editorial
(H. H. Godfrey and L. M. Campbell) — Revision of
the Kemp’s Ridley Recovery Plan (C. W. Caillouet, Jr.)
— Sea turtle subpopulations and the IUCN Red List:
A complementary role for conservation genetics (E.
Naro-Maciel and A. Formia) — Commentary: What is
a subpopulation? (N. Mrosovsky) — ARTICLES:
Transatlantic migration of juvenile Loggerhead Turtles
from the Strait of Gibraltar (D. Cejudo, N. Varo-Cruz,
A. Liria, J. J. Castillo, J. J. Bellido, and L. F. Lopez-
Jurado) — On the occurrence of Columbus Crabs
(Planes minutus) from Loggerhead Turtles in Florida,
USA (M. G. Frick, K. L. Williams, M. Bressette, D.
A. Singewald, and Richard M. Herren) — Hawksbill
Turtle tracking as part of initial sea turtle research and
conservation at Groote (S. D. Whiting, S. Hartley, S.
Lalara, D. White, T. Bara, C. Maminyamunja, and L.
Wurramarrba) — Lower nesting success of Flatback
Turtles caused by disorientation (A. U. Koch and M. L.
Guinea) — IUCN-MTSG Quarterly report — MEET-
ING REPORTS — LETTER — BOOK REVIEWS — |
ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — —
RECENT PUBLICATIONS.
The Marine Turtle Newsletter is now edited by Lisa |
M. Campbell, Nicholas School of Environment and
Earth Sciences, Duke University, 135 Duke Marine Lab
Road, Beaufort, North Carolina 28516 USA; and Mat- |
thew H. Godfrey, North Carolina Wildlife Resources
Commission, 1507 Ann Street, Beaufort, North Caro-
lina 28516 USA. The Managing Editor is Michael |
Coyne, A321 LSRC, Box 90328, Nicholas School of
the Environment and Earth Sciences, Duke University, |
Durham, North Carolina 27708-0328 USA. Fax +44
1392 263700. Subscriptions and donations towards the
production of the MTN can be made online at www.
seaturtle.org/mtn or postal mail to Michael Coyne (Man-
aging Editor) Marine Turtle Newsletter, | Southampton
Place, Durham, North Carolina 27705 USA (e-mail:
mcoyne @seaturtle.org).
Canadian Association of Herpetologists/Association Canadienne des Herpetologistes Bulletin
Fall 2006
14(1) 28 pages. CONTENTS: Instructions for authors
— EDITORIAL: On the return of CAH/ACH — A mes-
sage from the editor — MEETINGS: CARCNet 2006
Meeting Announcement (Pat Gregory) — JMIH 2006
(Leslie Lowcock) — CAH/ACH Membership and
Executive meetings (Jacqueline Litzgus) — FEATURE
ARTICLE: On the origins and dispersal of Neotropical
rattlesnakes in South America (Adrian Quijada-Mas-
carenas and Wolfgang Wuster) — FIELD NOTES:
Interesting nesting behaviour of Wood Turtles (Glyp-
temys insculpta) (William Greaves) — NEW FACUL-
TY IN HERPETOLOGY AT CANADIAN UNIVERSITIES:
Gabriel Blouin-Demers, University of Ottawa; David
Lesbarreres, Laurentian University; Jacqueline Litzgus,
Laurentian University — THESIS ABSTRACTS IN
CANADIAN UNIVERSITIES: Karine Beriault, M.Sc.
Wildlife Afield 3 (1 and Supplement to 1)
Volume 3, Number 1, contains FROM THE EDITOR:
Coming together — On the covers — FEATURE ARTI-
CLE: Supplement to Wildlife Afield: Roads and wildlife
— NOTES: American Avocets breeding at Bechers
Prairie near Riskke Creek, British Columbia in 2005
and 2006 (Phil Ranson) — Manx Shearwater in British
Columbia: comments on a pioneering seabird (Michael
Force, Ken Morgan, and Jukka Jantunen) — A note-
worthy record of the Pigmy Short-horned Lizard (Glenn
R. Ryder, R. Wayne Campbell, and Lawrence Powell)
2005; Joseph Crowley M.Sc. 2006; Amanda Duffus
M.Sc. 2006; Raymond Saumure Ph.D. 2004; Sara Wick |
M.Sc. 2004 — RECENT PUBLICATIONS IN CANADIAN
HERPETOLOGY: Citations 2005-06 — ANNOUNCE-
MENTS: Former CAH/ACH President Michael Rankin
remembered (Francis Cook) — MEMBERSHIP FORM.
Editor Dr. Jacqueline D. Litzgus, Laurentian Uni-
versity, Sudbury, Ontario P3E 2C6; e-mail: jlitzgus@
laurentian.ca; phone 705-675-1151 extension 2314.
Membership in CAH/ACH is $20.00 regular member,
$10.00 student member payable to the Canadian Asso-
ciation of Herpetologists. Mail to Dr. Patrick Gregory,
President and Treasurer of CAH/ACH, Department!
of Biology, University of Victoria, British Columbia :
V8W 2Y2.
— Cactus traps California Myotis in British Columbia ‘
(Christian W. Gronau) — Whip-poor-will Caprimulgus\\
vociferus: a new species for British Columbia, 1968-':
2006 (Adrian Dorst) — Status of the Yellow-breasted:
Chat in the Creston Valley, British Columbia, 1968-
2006 (R. Wayne Campbell and Edward McMackin)
— Unusual mortality of a male Rufous Hummingbird
in Burton, British Columbia (Arthur Schoeddert) —
Common Raven steals and caches eggs of the Double-
crested Cormorant in the Creston Valley, British Col-
502
2006
umbia (Linda M. Van Damme) — Impact of Cooper’s
Hawk predation on a nesting colony of Brewer’s Black-
birds on Vancouver Island, British Columbia (R. Wayne
Campbell and Barbara Begg) — Golden Eagle attacks
and kills yearling Mountain Goat (Barbara Zettergreen)
— Skeletal remains of the Northern Flicker with a
gross bill deformity discovered in a popular tree cavity
(Sherry L. Lidstone) — WILDLIFE DATA CENTRE: —
Featured species — Common Nighthawk (R. Wayne
Campbell, Martin K. McNicholl, R. Mark Brigham,
and Janet Ng) — Report of the Wildlife Data Centre:
1 January 2006 to 30 June 2006 (Michael I. Preston)
— BRITISH COLUMBIA ROUND-UP: Tips for the field
— Field notes for caring people — Errata — News of
friends — Life membership — a personal commitment
to wildlife — From the Archives — Final Flight —
Announcements, Publications of Interest — Volume
3(1), (and Supplement to 3(1), contains: FROM THE
EDITOR: At a crossroads — On the covers — ROADS &
WILDLIFE: Collisions with wildlife: An overview of
major wildlife vehicle collision data collection sys-
tems in British Columbia and recommendations for
the future (S. Gayle Hesse) — A comparison of roads
and rivers as barriers to animal movements: implica-
tions for roads in protected areas (Joanna Preston) —
High incidence of vehicle-induced ow] mortality in the
lower mainland and central Fraser Valley, British Col-
umbia (Michael I. Preston and Gerry A. Powers) —
Errata The Canadian Field-Naturalist 120(2)
Table of contents outside back cover:
NEWS AND COMMENT
503
A study on the incidence of amphibian road mortality
between Ucluelet and Tofino, British Columbia (Barbara
Beasley) — Mitigation efforts to reduce mammal! mor-
tality on roadways in Kootenay National Park, British
Columbia (Michael I. Preston, Larry Halverson and
Gayle Hesse) — Using wildlife vehicle collision data,
expert opinion and GPS technology to more accurately
predict and mitigate vehicular collisions with wildlife
in northern British Columbia (Roy V. Rea, Eric K.
Rapaport, Dexter P. Hodder, Michael V. Hurley and
Nicole A. Klassen) — Road watch BC: A program for
collecting, centralizing, and synthesizing information
from vehicle-induced wildlife mortalities in British
Columbia (R. Wayne Campbell and Michael I. Pres-
ton) — Estimating the probability of potential vehi-
cle collision from birds cross roads in interior British
Columbia (Michael I. Preston and Joanna Preston)
— USEFUL RESOURCES: Books — Symposia, Pro-
ceedings, Handbooks, and Reports — Websites — Up-
coming meetings — Book review: Road ecology —
Road watch BC — Wildlife collision prevention: Hints
for the highway.
Details on membership can be obtained from the
Biodiversity Centre for Wildlife Studies, PO Box
6218, Station C, Victoria, British Columbia V9P 5L5
Canada. Tel/Fax: 250-477-0465; e-mail: editor@ wild
lifebc.org; web site: www.wildlifebc.org.
Conservation evaluation of Dwarf Wolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada
GEORGE W. DOUGLAS, JENIFER L. PENNY, and KSENIA BARTON
“Wolly-heads” should read Woolly-heads.
Article page 235:
First record of a River Otter, Lontra canadensis, captured on the northern coast of Alaska
SHAWN P. HASKELL
In abstract and citation “Lutra“ should be Lontra.
Errata The Canadian Field-Naturalist 120(3)
Table of contents outside back cover:
Recent invasion, current status, and invasion pathway of European Common Reed, Phragnites
australis subspecies australis, in the southern Ottawa District
PAUL M. CATLING and SUSAN CARBYN
“Phragnites* should read Phragmites.
Pacific Hagfish. Eppptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and scavenger activity on
tethered carrion in subtidal benthic communities off western Vancouver Island
SARAH DAVIES, ALI GRIFFITHS, and T. E. REIMCHEN
“Eppptatretus* should read Eptatretus
The Ottawa Field-Naturalists’ Club Awards for 2005
IRWIN M. BRobo, CHRISTINE HANRAHAN, BEVERLY MCBRIDE, and ELEANOR ZURBRIGG
At the Club’s Annual Soirée, held on 29 April 2006,
at St. Basil’s Church in Ottawa, awards were once again
given to members, and two non-members, who distin-
guished themselves by accomplishments in the field of
natural history and conservation, or by extraordinary
activity within the Club. For the first time in several
years, we had a recipient of the Anne Hanes Natural
History Award, and the Mary Stuart Education Award
was given for the second time. The following citations
for those who received an award were read to the mem-
bers and guests assembled for the event.
Gillian Marston and Suzanne Deschenes — Members of the Year
The OFNC’s Member of the Year award recognizes
the member(s) judged to have contributed the most to
the club during the previous year. Education and Pub-
licity Committee members Gillian Marston and Suz-
anne Deschenes have made stellar accomplishments
over the past year in developing educational and pub-
licity materials for OFNC, and leading a successful
membership drive through publicity efforts. They have
also brought their considerable computer skills to mod-
ernize the Club’s photo library and displays.
Gillian and Suzanne are the driving force behind the
OFNC’s new photography contest which began in 2005
and is set for a second round in 2006. Meant to encour-
age people’s appreciation of nature through photogra-
phy, and to boost the club’s library of digital photo-
graphs on natural themes, the first contest reaped 275
images, all of which have the potential to be used in
displays or slideshows. The contest resulted in a profes-
sional, stunning slide show for the 2006 Annual Busi-
ness Meeting. All these new images are now catalogued
in a digital library, also developed and managed by
Suzanne and Gillian. This library is the successor to the
club’s long-standing and well used slide collection.
Suzanne and Gillian together bring a special synergy
of creativity and management skills. They further ap-
plied their talents in 2005, taking the club’s new image
collection to the public by designing and constructing
several portable exhibits to highlight Ottawa nature, the
club’s activities, and to attract new members. Members
and non-members alike will recall seeing volunteers
ee, ee
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|
staffing these exhibits at the entrance to popular nature _
walk areas such as the Britannia Conservation Area.
Y
They also appeared on site for various club excursions \_
and conferences where they successfully attracted a)
good number of new members and caught the eye of
'}
i
educators. Gillian and Suzanne and the rest of the com- ©
mittee continue to develop more portable exhibits and
to refine the design. For example, they put together a —
set of slide shows on different themes that can be shown) —
at exhibits using a table-top projector or laptop com-
puter.
many special contributions during 2005. Designating
you as members of the year is our way of saying “thank
you”!
Christine Hanrahan — George McGee Service Award
Most successful organizations have a handful of
members that devote themselves, body and soul, to the
workings of the group, serving on committees, on
Council, and in any way that they are needed to make
the Club succeed. Christine Hanrahan is one of these
people. The Ottawa Field-Naturalists’ Club has bene-
fited from her efforts and devotion for over 25 years,
and we recognize these efforts with the George McGee
Service Award. This award honours members who have
volunteered their time to the betterment of the Club
over several years. The award commemorates George
McGee, who for more than three decades actively
devoted much of his spare time to teaching people
about birds and natural history through his talks and
numerous outings.
Christine has been a member of the OFNC since the
late 1970s. She has been an active member of many
Club committees, including the Fletcher Wildlife Gar-
den Committee, Conservation Committee, Bird Com-)
mittee, and Awards Committee. On top of this, she has:
recently coordinated the local group working on they
Gillian and Suzanne, the club is grateful for your |
|
|
{
Ontario Breeding Bird Atlas, which is now drawing to’ |
a close. Co-ordinating over a hundred volunteers for
5 years (plus an additional start-up trial year, plus the) |
final data compilation/coordination in 2006) is no small
feat. Not only was there all the coordination between’
Atlas headquarters and the region’s 24 volunteers.:
which included distributing information packages.’
forms, CDs and other material, but also there were the)
annual meetings to coordinate, the annual reports te!
write, the articles for Trail & Landscape, and hours anc
hours spent on-line writing e-mail messages and check-
ing the data of the central database. And this is the
second time Christine has done this! She was the leac
504
i
q
>:
r
2005
coordinator in region 24 for the first Atlas project 20
years ago. After that she coordinated the Loggerhead
Shrike survey in this area.
But our members probably know Christine best
through her work on the Fletcher Wildlife Garden Com-
mittee over many years. She has diligently kept long-
term and short-term lists of many of the FWG occu-
pants and visitors (e.g., birds and butterflies). This in-
cludes preparing the current sightings board and the
checklists. She has published many articles in Trail &
Landscape about the activities and happenings at FWG.
She has prepared posters, many pamphlets, checklists,
nesting reports, etc., if not solely, then as the main driv-
ing force. She has organized many important activities
at the Centre to draw in and educate visitors. Every
year she has prepared the FWG display and organized
the hosting (including herself) at the National Capital
Region Wildlife Festival. She has also prepared displays
at other events such as the Health Canada-sponsored
Environment Day.
Christine has represented the OFNC and the FWG
on the National Capital Wildlife Festival Committee
for many years and puts a huge amount of time into
organizing workshops and conservation awards. The
workshops she has organized have been on topics of
great interest to the OFNC, such as Invasive Alien Spe-
cies, conservation (Protecting Wild Places) and bird
conservation.
Another of Christine’s favourite activities is the
OFNC Conservation Committee. She has participated
in many committee activities and battles including, most
recently, the Larose Forest issue. Here she worked on
developing checklists of the flora and fauna with vari-
ous specialists. She also worked for hours and hours
with Ghislaine Rozon reviewing documents, preparing
strategy, briefs, writing letters, attending all kinds of
meetings in an effort to save the forest from develop-
CLUB AWARDS
505
ment. Her work on conserving Petrie Island also had a
major impact. Working with The Friends of Petrie Is-
land, she compiled lists, wrote articles for Trail &
Landscape, organized field trips, as well as attending
endless meetings, reviewing documents, and preparing
OFNC responses. There have been many other conser-
vation issues around Ottawa that have received Chris-
tine’s attention and hard work, including: Moffat Farm,
Montfort Woods, Nortel Woods, Leitrum wetlands,
5309 Bank Street (Tomlinson quarry) and Ferguson
Forest. The list seems endless.
And who among us has not read, enjoyed and learned
from Christine’s many articles in Trail & Landscape.
She has published on birds, plants, habitats and a wide
range of conservation issues. Her short article on snags
and old logs is a well known, authoritative and influ-
ential document. It has been reproduced and widely
distributed within the Ontario Ministry of Natural Re-
sources and has been used and cited by people even
in other parts of the world. Her article on hedgerows
and their importance to wildlife is another gem and
has become an important resource for ecologists. Since
the OFNC has been on the Internet, Christine has been
an active contributor to our website with informative
and interesting material on a variety of natural history
subjects.
Because Christine has also served as a knowledge-
able member of the Awards Committee for a number
of years, recognizing all her achievements and work
with an appropriate prize “slipped through the cracks”.
Not until her nomination came from outside the com-
mittee by a number of members all wondering how she
could have been overlooked so long could we make this
presentation. The Club is therefore pleased to present
Christine Hanrahan with this long overdue and well-
deserved Service Award.
Friends of the Carp River — Conservation Award (Non-Member)
The OFNC Conservation Award, non-member, is
given to a group or individual who has made an out-
standing contribution in the cause of natural history
conservation in the Ottawa Valley, with particular em-
phasis on activities within the Ottawa District.
The Friends of the Carp River have worked for many
years to protect and restore the ecosystem of the Carp
River, and this dedication makes them very worthy
recipients of this award.
The Friends of the Carp River grew out of the West
Carleton Township’s Environmental Advisory Commit-
tee which began an extensive study of the Carp River
in 1993. The organization was formed in 1997 as a citi-
zen’s group whose goal was to help restore and improve
the health of the Carp River. The river has its headwa-
ters just south of the Corel Centre and flows 42 km to
_ the Ottawa River at Fitzroy Harbour, and is “the only
river that flows entirely within the boundaries of the
_ City of Ottawa”. Over the years the river has suffered
from shoreline erosion, siltation, flooding, low oxygen
levels and subsequent loss of important wildlife habitat.
Working with various groups, including landowners,
businesses, government agencies and recreational users,
the Friends are developing some substantial, achievable
goals to continue rehabilitating the “ribbon of life”,
their wonderfully descriptive term for the Carp River.
Studies show that the Carp River has suffered overall
degradation, including silting problems due in part to
agricultural tilling practices and channelization. Shore-
line erosion further contributes to siltation, which puts
at risk the 40 species of fish inhabiting the river. Tribu-
taries of the Carp River have severely degraded water
quality due largely to agricultural fertilizers and ani-
mal wastes. In spring 2006, the threat of development
in the floodplain could bring more problems such as
future flooding as well as damaging restoration projects.
The Friends of the Carp River, an entirely volunteer-
run organization, is dedicated to reversing the decline
506
of the river and the Friends have already made tremen-
dous improvements. The Friends have planted over
20 000 trees and shrubs since 1997 with the help and
cooperation of local landowners living along the river
and many volunteers, and this will have a substantial
positive impact on controlling erosion and siltation.
With assistance from provincial foresters, site plans for
tree planting have been created.
The Friends have worked with the City of Ottawa
on its Watershed study, and gathered support from the
Ottawa Stewardship Council and the Rural Clean Water
Program. Their work was recognized with a Trillium
Foundation grant of $50 000 to help them achieve their
goal of rejuvenating the Carp River. One of their objec-
tives for this grant was to commission a study of the
river, which they did, and the Carp River Remediation
Project report was prepared by consultants in April
2003.
The Friends of the Carp River have produced a pam-
phlet which details ways and means for landowners to
achieve best management practices. The Friends are
available to help steer farmers and others to specific
programs, including funding, to enable them to imple-
ment improvements that benefit both landowners and
the river.
The Friends have been ingenious, creative and tire-
less in coming up with ways to garner support for the
river. Believing that an “image change” for the river
was a necessary part of creating a positive public per-
ception, they held a year-long photography contest that
encouraged people to submit photos of the Carp River,
and also persuaded them to look at the river as a thing
Iola Price — Conservation Award (Member)
The OFNC Conservation Award is given in recogni-
tion of a club member who has made an outstanding
contribution towards protecting our natural environ-
ment. Iola Price is a very worthy recipient of this award
in recognition of her effective membership in the City
of Ottawa’s Forest and Greenspace Advisory Committee
(OFGAC) to promote conservation of trees and forests.
As the current Chair of the Ottawa Forests and
Greenspace Advisory Committee, and co-chair before
that, Iola provides an articulate and credible voice on
issues related to conservation of the trees, forests and
greenspace of our Nation’s capital.
The Ottawa Forests and Greenspace Advisory Com-
mittee was established in the fall of 2001 to advise
Ottawa City Council and provide a forum for citizens
on issues related to trees and forests. Iola’s accom-
plishments include reviewing OFGAC position papers
before transmission to appropriate authorities, prepa-
ration of numerous written briefs and making power-
ful submissions to city committees. For example, Iola
recently exhorted City Council at a budget planning
meeting to “think of trees as green infrastructure, as
essential to a well-ordered city as sewers, lighting and
roads”.
THE CANADIAN FIELD-NATURALIST
Vol. 120
of beauty . Winning entries were used to create a Mil-
lennium calendar, Flowing into the next Millennium:
A year in the life of the Carp River, which was sold at
cost and sold out in three months. The photos were also
used to create sets of notecards, again sold at cost.
A benefit concert with Terry Tufts and Kathryn
Briggs was held on Earth Day, 2001 to help raise funds
for continued work on the river. A CD of a song, The
Mighty Carp by Dan Mayo, inspired by the river and
dedicated to the Friends was released in 2002.
The Friends explain that “shorelines are breeding
grounds, nurseries, food sources, shelter, and hiding
places for many species. Ninety per cent of all living
things in a lake or river are found along the shoreline.
Shorelines are effective natural filters and buffer zones,
acting as the ‘kidneys’ for the watershed. Shorelines
help to improve the quality and quantity of ground
water. Polluted run-off is trapped and absorbed by the
vegetation, thus protecting water quality. The complex
interplay of plants, animals, land, and water combine
to make the shorelines the single most important part
of the river ecosystem.”
The Friends have been remarkably successful in
drawing the community into active involvement with
the health of the river as evidenced not only by the many
volunteers who turn out for tree-planting activities but
by the tremendous support they have received from local
businesses and granting agencies. For these reasons and
more, the OFNC is very pleased to give the Friends of
the Carp River this year’s Conservation Award for non-
members. ;
On many occasions, Iola has made a powerful and
persuasive intervention during the City’s planning pro-
cess in support of the importance of greenspace and
protection of sites of urban natural ecological signifi-
cance. Her recent interventions regarding the Nepean
Creek Corridor as well as her efforts for improving
the criteria utilized for determining appropriate urban
density development levels underscores this dedica-
tion towards the maintenance and protection of trees
and greenspace in the National Capital Region (NCR). .
Iola has provided recommendations to strengthen 1
ecological provisions in urban design studies such as $
Riverside South Community Design and Leitrim Com- -
munity Design. She spearheaded opposition to devel- -
opment projects such as the highly controversial Carp )
Ridge housing project and the Phase 2 Leitrim Wetlands
development, which, if approved, would extend hous-
ing into the wetland itself.
Iola is recognized as a very effective and valuable
“networker”, regularly liaising with other tree and
greenspace-oriented community groups operating in the ©
NCR. This includes our Club through the Conserva-
tion Committee and Fletcher Wildlife Garden. Members ©
of these committees wholeheartedly appreciate that Iola
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2005
is very quick and efficient in replying to queries and in
exploring issues and ideas, which helps so much when
it allows one to continue one’s work without delay.
CLUB AWARDS
507
Through this award, OFNC members recognize the
outstanding efforts towards protecting our natural envi-
ronment by one of our members — lola Price.
Macoun Field Club — Anne Hanes Natural History Award
The Anne Hanes Natural History Award, named in
honour and memory of one of the Club’s most dedi-
cated amateur naturalists, is made in recognition of a
member who, through independent study or investiga-
tion, has made a worthwhile contribution to our knowI-
edge, understanding and appreciation of the natural his-
tory of the Ottawa Valley. The award is designed to es-
pecially recognize work that is done by amateur natu-
ralists.
The OFNC feels that there can be no more worthy
recipient than the Macoun Field Club in recognition of
the remarkable studies the members of the MFC have
made of the Macoun Study Area in Bells Corners over
a 35 year period, documenting rare plants and animals,
tracking changes to plant and animal populations as well
as the environment, producing and up-dating vegetation
maps of the area, and much more. The MFC, spon-
sored jointly by the OFNC and Canadian Museum of
Nature, has been a haven for youngsters interested in
nature since 1948, and accepts members from grades
4 through 12.
The idea of having a special area of natural land des-
ignated as a “study area” for members of the Macoun
Field Club came from the senior (high school) mem-
bers, not from the adult leadership. The first study area
in the Bells Corners area was actually where the Wild
Bird Centre is now, but because the OMNR was cutting
out the dead and dying elms at the time, they suggest-
ed an area across the road in a less disturbed site west
of Moodie Drive. Within two years, the members had
drawn a vegetation map of most of the 800 acres mak-
ing up the site. This valuable base map was based on
aerial photographs supplemented by field trips to deter-
mine what was there. There have been several versions
made to keep pace with changes: 1972, 1986, 1992, and
2000. A colour version was created for the Club’s web
site (www.magma.ca/~rel/mfc/studyarea/msa.html).
In this way, the MFC members traced the changes that
occurred in a significant part of the Ottawa Greenbelt
brought on by urbanization, beaver activity, tree dis-
eases and people. The MFC members, on their bi-week-
ly field trips over many years, mostly led by Rob Lee,
created lists of trees (54 species), wildflowers, birds,
mammals (29 species), reptiles and amphibians, and
lichens (215 species). In the process, they discovered
many rarities and have documented the disappearance
of certain species. Their studies continue on every-
thing from “the sand grains in the soil” to “Leather-
wood Treehoppers.”
Because each member is encouraged to “adopt a
tree” and study its growth and health, its inhabitants in
the crown, on the trunk and at its base, the MFC has
amassed an amazing amount of information about sea-
sonal fluctuations in a suburban forest system.
Much of this information has been organized on the
MFC’s amazing website designed and maintained by
Rob Lee, but the information itself was gathered by the
members. It is improbable that any urban greenspace
in Canada has such a well-documented history of its
flora and fauna.
Observations on the Macoun Study Area have been
regularly published by the members in their annual
magazine, The Little Bear, and, because of the Internet,
is now readily available to the general public as well.
Most important, the experience that the members have
gained in doing careful field research has encouraged
many past members to embark on a career in conser-
vation biology, botany, zoology, park management or
teaching.
The OFNC feels that the Macoun Field Club mem-
bers have done a remarkable job, under the leadership
of Rob Lee, and well deserves the Anne Hanes Natural
History Award.
Michael Léveillé — Mary Stuart Education Award
The Mary Stuart Education Award was established
to recognize members, non-members or organizations
for their outstanding achievements in the field of nat-
ural history education in the Ottawa Region. Michael
Léveillé, a science teacher at the Educarium, an Ottawa
private school, is without doubt a natural history edu-
cator par excellence, and a most worthy recipient of
this Award.
Michael has taught at the Educarium since 1996.
His passion for nature is infectious and inspirational
and is observed most keenly by his students. Michael
brings an impressive background to his role as science
_ teacher. He founded the Ottawa Paleontology Society
in 1991 to advance an understanding of fossils. He has
long been associated with the Canadian Museum of
Nature where he created a number of popular programs,
most notably the “Collector's Corner’, a museum dis-
covery centre. At the museum he worked with people
of all ages, instilling an awareness of, and appreciation
for, the natural world. However, there is no doubt that
Michael excels at working with young people, encour-
aging them to be attuned to the rhythms of the natural
world around them.
Michael is also an accomplished artist with a degree
in Visual Arts from the University of Ottawa. His work
has been exhibited at local universities, museums, as
well as at the Educarium. Michael was the artist for the
children’s storybook series “Fergi the Frog”.
508
Under Michael’s knowledgeable direction, in 2004
his students designed and implemented the “Macoun
Marsh Project” at Beechwood Cemetery, to demon-
strate how a thriving and diverse ecosystem can exist
inside a large urban centre. This became one of their
most ambitious and high profile projects, and is ongo-
ing. Because the marsh was unnamed when they began
working on it, they held a “Marsh Naming Contest” and
encouraged neighbours and the community at large to
join in. Overwhelming support led to naming it after
John Macoun (1831-1920) who is buried at Beech-
wood. The contest, and the work done by the students,
led to increased public awareness of the site and even-
tually resulted in the Directors of Beechwood Cemetery
making a firm commitment to protect and maintain it.
The Macoun Marsh project encourages students to
study the marsh over different seasons. They are taught
to record scientific observations in journals, identify
and study the different species found, and learn the fun-
damentals of ecology. They also design, construct, install
and maintain bird feeders at the site. Under Michael’s
THE CANADIAN FIELD-NATURALIST
Vol. 120
careful instruction, the Educarium students have doc-
umented nearly 900 species at the marsh.
The project has grown beyond their wildest expec-
tations. Funding to continue with studies and enhance-
ments has been obtained from a variety of sources in-
cluding the Wetland Habitat Fund and the City of
Ottawa. In conjunction with the Beechwood Cemetery
Foundation, plans are underway for habitat enhance-
ment, a boardwalk, and an outdoor classroom in spring
2006. The students continue to keep nature journals
and will present this work at various venues around the
city over the coming year. Michael invited the neigh-
bouring Jean Vanier Catholic School to join his students
on the project, and in May 2006 both schools sent rep-
resentatives to Géteborg, Sweden, for the Volvo Adven-
ture International contest for natural history projects.
Michael’s inspired teaching guarantees that his stu-
dents will continue to be engaged by the world of
nature, and so it is for this most important work, that
OFNC is pleased to present Michael Léveillé with the
Mary Stuart Education Award for 2005.
\
Index to Volume 120
Compiled by Leslie Cody
Abies amabilis, 172
balsamea, 100,236,238,299,433,440
lasiocarpa, 100,331
Accipiter striatus, 46
Acer spp., 226
circinatum, 11,171,348
glabrum var. douglasii, 171
macrophyllum, 11,180
rubrum, 299,354
saccharum, 299
Achillea millefolium, 354
Achnatherum hymenoides, 336
Acipenser fulvescens, 71
transmontanus, 422
Acipenser fulvescens, Movements in Rainy Lake, Minnesota
and Ontario, Lake Sturgeon, 71
Acorn, J., Reviews by, 245,489
Adams, W.E. Jr., L.W. Kallemeyn, and D.W. Willis. Lake
Sturgeon, Acipenser fulvescens, Movements in Rainy
Lake, Minnesota and Ontario, 71
Adiantum capillus-veneris, 138
Aegolius acadicus, 294,303
Aeshna sp., 416
canadensis, 415
constricta, 417
eremita, 417
interrupta, 416
interrupta lineata, 416
interrupta lineata ssp. lineata, 416
umbrosa, 416
umbrosa umbrosa, 417
Agalinis neoscotica, 357
Agriotes collaris, 19
stabilis, 19
Agropyron cristatum, 337
Agrostis exerata, 214
tenuis, 354
Agujaceratops, 403
Aira caryophyllea, 320
praecox, 154,320
Aix sponsa, 236,303
Alaska, First Record of a River Otter, Lontra canadensis, Cap-
tured on the Northeastern Coast of, 235
Alaska, Physical Characteristics, Hematology, and Serum
Chemistry of Free-ranging Gray Wolves, Canis lupus,
in Southcentral, 205
Alaska, 2001-2003, Assessment of Effects of an Oil Pipeline
on Caribou, Rangifer tarandus granti, Use of Riparian
Habitats in Arctic, 323
Alberta, A New Record of Deepwater Sculpin, Myoxocephalus
thompsonii, in Northeastern, 480
Alberta Boreal Forests, Winter Habitat Use by American
Marten, Martes americana, in Western, 100
Alberta, Pack Size of Wolves, Canis lupus, on Caribou, Ran-
gifer tarandus, Winter Ranges in Westcentral, 313
Alces alces, 205,313,367
alces americana, 5|
Alder, 236,240
Red, 11,348
Speckled, 22
Allium amplectens, 160
cernuum, 154
Alnus sp., 236,238,240
incana, 22
rubra, 348
rubus, \\
Alopecurus carolinianus, 164
Alopex lagopus, 36
Alosa sapidissima, 422
Ambystoma gracile, 90
macrodactylum, 90
Ameiurus catus, 424
melas, 422
natalis, 421
nebulosus, 422
Ameiurus natalis, a Loricariid Catfish, Panaque suttonorum,
and a Silver Pacu, Piaractus cf. P. brachypomus, in
British Columbia, First Records of the Yellow Bull-
head, 421
Amelanchier, 428
alnifolia, 176,428
alnifolia var. compacta, 428
florida, 428
humilis, 428
humilis var. compacta, 428
laevis, 354
laevis x lucida, 354
lucida, 354
sanguinea, 428
Amelanchier in the Whitewood area of Southeastern Saskat-
chewan and the First Saskatchewan Records of Ame-
lanchier sanguinea, Regional Variation in, 428
Amelanchier sanguinea, Regional Variation in Amelanchier
in the Whitewood area of Southeastern Saskatchewan
and the First Saskatchewan Records of, 428
Amia calva, 422
Ammodytes americanus, 96
Ammospiza leconteii, 22
Ammospiza leconteii, from Northeastern Ontario, with Some
Notes on Nesting Behaviour, New Nesting Records
of the Le Conte’s Sparrow, 22
Anaphalis margaritacea, 354
Anas clypeata, 5
platyrhynchos, 6
Anax junius, 417
Anderson, C.S., D.B. Meikle, A.B. Cady, and R.L. Schaefer.
Annual Variation in Habitat Use by White-footed Mice,
Peromyscus leucopus: The Effects of Forest Patch
Size, Edge and Surrounding Vegetation Type. 192
Andrena, 19
509
510
Anguilla rostrata, 238,477
Anguille d’ Amérique, 238
Antennaria flagellaris, 183
howellii ssp. neodioica, 354
Antennaria flagellaris, in Canada, Conservation Evaluation
of Stoloniferous Pussytoes, 183
Anthophora, 180
Anthoxanthum odoratum, 148,154,160
Antilocapra americana, 210
Apera interrupta, 320
Apis mellifera, 294
Apocynum androsaemifolium vat. incanum, 354
Apodemus sylvaticus, 195
Apple-Producing Region of Southern Quebec, First Obser-
vations of an Eastern Screech-Owl, Megascops asio,
Population in an, 289
Aralia hispida, 354
Arbutus menziesii, 148,180
Arbutus, 148,180
Arctostaphylos uva-ursi, 172,336,354
Arenaria capillaris ssp. americana, 184
Aronia sp., 353
Aronia, 360
Artemisia cana ssp. cana, 336
tridentata, 164,184
Ascophyllum nodosum, 236
Asio otus, 295
Aspen, 314,428
Trembling, 22,100,342
Astragalus miser, 176
Athous acanthus, 19
Auditor’s Report, 398
Aulnes, 238
Aythya sp., 6
affinis, 5
americana, 109
collaris, 6
marila, 109
valisneria, 109
Baccha elongata, 17
Badger, 209
American, 31,342
European, 52
Badiou, P.H.J. and L.G. Goldsborough. Northern Range Expan-
sion and Invasion by the Common Carp, Cyprinus car-
pio, of the Churchill River System in Manitoba, 83
Balaena mysticetus, 462
Balaenoptera acutorostrata, 95
borealis, 97
musculus, 95
physalus, 95
Balanus improvisus, 106
Baldwin, R.A., A.E. Houston, M.L. Kennedy, and P.S. Liu.
Predicting Raccoon, Procyon lotor, Occurrence
Through the Use of Microhabitat Variables, 225
Balkwill, D., B.W. Coad, I. Galvez, and J. Gilhen. First Record
of the Great Barracuda, Sphyraena barracuda from
Canada, 241
Ballard, W.B., 205
Balsamorhiza sagittata, 176
Balsamroot, Arrow-leaved, 176
Barley, 343
Barracuda, Great, 241
Barracuda, Sphyraena barracuda from Canada, First Record
of the Great, 241
THE CANADIAN FIELD-NATURALIST
Vol. 120
Barton, K., 163,183
Bartonia virginica, 357
Basquill, S., 351
Bass, Largemouth, 422
Smallmouth, 421
Striped, 422
Bat, Big Brown, 39
Little Brown, 39,51
Long-legged, 39
Northern Long-eared, 39,51,332
Silver-haired, 39
Western Long-eared, 39
Bat, Myotis septentrionalis, in the Yukon Territory, First
Records of the Northern Long-eared, 39
Bdellarogatis, 444
plumbeus, 444
Bear, American Black, 51
Black, 57,68,110,209,314
Brown, 57
Grizzly, 314
Bear, Ursus americana, Retrieval of an Elk, Cervus elaphus,
Carcass from a Small Lake in Riding Mountain Na-
tional Park, Manitoba, Probable Black, 110
Bear, Ursus americanus L., Long-range Homing by an Adult
Female Black, 57
Bearberry, Common, 336
Beaudoin, A.B., Reviews by, 383,493,494
Beaver, 51,62,205,331,368
Bee, 15
Honey, 294
Beetle, 296
Click, 15
Ground, 16
Beluga, 457
Bertrand, A.-S., S. Kenn, D. Gallant, E. Tremblay, L. Vasseur,
and R. Wissink. MtDNA Analyses on Hair Samples
Confirm Cougar, Puma concolor, Presence in South-
ern New Brunswick, Eastern Canada, 438
Betula spp., 428,440
alleghaniensis, 299
cordifolia, 299
papyrifera, 22,50,238,299,354,433,440
populifolia, 354
Birch, 428,440
Mountain, 299
Paper, 433
White, 22,50,299,440
Yellow, 299
Bird, D.M., 289
Bison bison athabascae, 68
Bison, Wood, 68
Bitter-cress, Little Western, 180
Bizecki Robson, D. A Conservation Evaluation of Smooth
Goosefoot, Chenopodium subglabrum (Chenopodi-
aceae), in Canada, 335
Blackberry, 359
Blaney, S., 319
Blarina brevicauda, 294,436
Blastocerms dichotomus, 64
Blinn, B.M., V. Violette, and A.W. Diamond. Osprey, Pandion
haliaetus, Depredates Common Eider, Somateria mol-
lissima, Duckling, 236
Blue-eyed Mary, Large-flowered, 180
Blue-Eyed-Grass, Coastal-Plain, 359
Blueberry, 360
Alaskan, 172
2007
Bluegill, 422
Bluegrass, Annual, 320
Bulbous, 320
Kentucky, 160
Bluejoint, Canada, 22
Bluet, Azure, 417
Big, 417
Familiar, 417
Hagen’s, 417
Marsh, 417
Blush, Sea, 154,158
Bobcat, 230
Bobwhite, Northern, 452
Bobwhites, Colinus virginianus, by Mammalian Mesopreda-
tors: Does the Problem-Individual Paradigm Fit?,
Predation on Artificial Nests of Northern, 452
Bombus, 180
perplexus, 19
vagans, 19
Bombycilla cedrorum, 294
Boreal Dip Net/L’Epuisette Boréale January 2006, The, 392
Bouleau blanc, 238
Bowfin, 422
Brachyopa, 20
notata, 17
Brachypalpus oarus, 17
Branta canadensis, 109
British Columbia, A Test of Interspecific Effects of Introduced
Eastern Grey Squirrels, Sciurus carolinensis, on Dou-
glas’s Squirrels, Tamiasciurus douglasii, in Vancou-
ver, 10
British Columbia, Early Ontogenetic Diet in Gray Wolves,
Canis lupus, of Coastal, 61
British Columbia, First Records of the Yellow Bullhead,
Ameiurus natalis, a Loricariid Catfish, Panaque sut-
tonorum, and a Silver Pacu, Piaractus cf. P. brachy-
pomus, in, 421
British Columbia, Life History Phenology and Sediment Size
Association of the Dragonfly Cordulegaster dorsalis
(Odonata: Cordulegastridae) in an Ephemeral Habitat
in Southwestern, 347
Brome, 148,154
Barren, 160,180
Bromus, 148,154
sterilis, 160,180
tectorum, 176
Broom, Scotch, 148,155,160
Brunelle, P.M., 413
Bryan, H.M., C.T. Darimont, T.E. Reimchen, and P.C. Paquet.
Early Ontogenetic Diet in Gray Wolves, Canis lupus,
of Coastal British Columbia, 61
Bryant, J.E. A Tribute to Nicholas Stephen Novakowski
1925-2004, 370
Bubo virginianus, 290
Bucephala albeola, 6
clangula, 5,109
Buckwheat, 343
Cushion, 184
Parsnip-flowered, 184
Budworm, Spruce, 50,299
Bufflehead, 6
Bufo americanus, 240
boreas, 87
Bullhead, Black, 422
Brown, 422
INDEX TO VOLUME 120 S11
Yellow, 421
Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque sut-
tonorum, and a Silver Pacu, Piaractus cf. P. brachy-
pomus, in British Columbia, First Records of the
Yellow, 42]
Bulrush, American Three-square, 213
Bunchberry, 440
Bunker-Popma, K. Scoter, Melanitta spp., Migrations Inter-
rupted by Confederation Bridge: An Update, 232
Bur-reed, Narrowleaf, 240
Burbot, 422,480
Burchsted, F., 106
Burchsted, J.C.A. and F. Burchsted. Lady Crabs, Ovalipes
ocellatus, in the Gulf of Maine, 106
Butcher, M.K., 323
Butler, M.J., W.B. Ballard, and H.A. Whitlaw. Physical Char-
acteristics, Hematology, and Serum Chemistry of
Free-ranging Gray Wolves, Canis lupus, in Southcen-
tral Alaska, 205
Buttercup, Bur, 319
Buttercup, Ranunculus testiculatus, New to Eastern Canada,
Bur, 319
Butterfly, Maritime Copper, 419
Cabbage, Skunk, 172
Cady, A.B., 192
Calamagrostis sp., 354
canadensis, 22
Calamovilfa longifolia, 336
Calidris spp., 5
Caligus, 199
clemensi, 199
Camarasaurus, 408
Camas, Common, 148
Camassia quamash, 148
Canada, A Conservation Evaluation of Smooth Goosefoot,
Chenopodium subglabrum (Chenopodiaceae), in, 335
Canada, Bur Buttercup, Ranunculus testiculatus, New to East-
ern, 319
Canada, Conservation Evaluation of Pacific Rhododendron,
Rhododendron macrophyllum, in, 169
Canada, Conservation Evaluation of Slender Collomia, Col-
lomia tenella, in, 175
Canada, Conservation Evaluation of Stoloniferous Pussytoes,
Antennaria flagellaris, in, 183
Canada, Conservation Evaluation of the Bog Bird’s-foot Tre-
foil, Lotus pinnatus, in, 157
Canada, Conservation Evaluation of the Pacific Population of
Dwarf Woolly-heads, Psilocarphus brevissimus vat.
brevissimus, in, 163
Canada, Conservation Evaluation of the Prairie Lupine,
Lupinus lepidus var. lepidus, in, 147
Canada, Conservation Evaluation of the Seaside Birds-foot
Trefoil, Lotus formosissimus, in, 153
Canada, Conservation Evaluation of the Small-flowered Tonel-
la, Tonella tenella, in, 179
Canada, Extension of Coyote, Canis latrans, Breeding Range
in the Northwest Territories, 67
Canada, First Record of the Great Barracuda, Sphyraena
barracuda trom, 241
Canada, MtDNA Analyses on Hair Samples Confirm Cougar,
Puma concolor, Presence in Southern New Brunswick,
438
Canada’s Maritime Provinces, Recent Declines of House Spar-
rows, Passer domesticus, in, 43
2
Canadian Arctic, Reactions of Narwhals, Monodon monoc-
eros, to Killer Whale, Orcinus orca, Attacks in the
Eastern, 457
Canadian Association of Herpetologists/Association Cana-
dienne des Herpetologistes Bulletin, 502
Canadian Field-Naturalist, Advice for Contributors to The,
134,262,402,534
Canadian Field-Naturalist 119(3), Errata The, 133
Canadian Field-Naturalist 120(1), Errata The, 259
Canadian Field-Naturalist 120(2), Errata The, 503
Canadian Field-Naturalist 120(3), Errata The, 503
Cancer magister, 364
productus, 363
Canis sp., 263
familiaris, 263479
latrans, 27,31,40,50,67,110,209,263,314,342,472,474,
478
lupus, 27,57,61,67,111,205,263,313,367,466,47 1,476,
478
lupus arctos, 466,471
lupus beothucus, 50
lycaon, 27
rufus, 27
Canis latrans, An Ethogram Developed on Captive Eastern
Coyotes, 263
Canis latrans, Breeding Range in the Northwest Territories,
Canada, Extension of Coyote, 67
Canis latrans, Conditions for Sexual Interactions Between
Wild Grey Wolves, Canis lupus, and Coyotes, 27
Canis latrans, in a Heavily Urbanized Area: Implications for
Ecosystem Management, Use of “Micro’-Corridors
by Eastern Coyotes, 474
Canis latrans, in an Urban Area, Rat Poison Kills a Pack of
Eastern Coyotes, 478
Canis lupus, and Coyotes, Canis latrans, Conditions for
Sexual Interactions Between Wild Grey Wolves, 27
Canis lupus arctos: Influence of Order on Posture, Double
Marking in Arctic Wolves, 471
Canis lupus arctos, in Summer, Urine-marking and
Ground-scratching by Free-ranging Arctic Wolves,
466
Canis lupus, Group for Ontario, A New Record Size Wolf,
367
Canis lupus, in Southcentral Alaska, Physical Characteristics,
Hematology, and Serum Chemistry of Free-ranging
Gray Wolves, 205
Canis lupus, of Coastal British Columbia, Early Ontogenetic
Diet in Gray Wolves, 61
Canis lupus, on Caribou, Rangifer tarandus, Winter Ranges
in Westcentral Alberta, Pack Size of Wolves, 313
Canola, 343
Canvasback, 109
Carassius auratus, 422
Carbyn, L., Review by, 249
Carbyn, S., 307
Carbyn, S., P.M. Catling, S.P. Vander Kloet, and S. Basquill.
An Analysis of the Vascular Flora of Annapolis Heath-
lands, Nova Scotia, 351
Cardamine oligosperma, 180
Cardinal, Northern, 294
Cardinalis cardinalis, 294
Carduellis tristis, 294
Carex atlantica, 357
cumulata, 354
deflexa var. deflexa, 354
emonsii, 354
THE CANADIAN FIELD-NATURALIST
Vol. 120
feta, 160
lucorum, 360
lyngbyei, 213
nigra, 354
oligosperma, 22
paleacea, 416
praegracilis, 310
scoparia, 354
siccata, 360
stipata, 22
tonsa, 358
tonsa vat. rugosperma, 354
tonsa var. tonsa, 354
tumulicola, 160
Caribou, 313,323
Barren-ground, 205
Woodland, 50,313,367
Caribou, Rangifer tarandus granti, Use of Riparian Habitats
in Arctic Alaska, 2001—2003, Assessment of Effects
of an Oil Pipeline on, 323
Caribou, Rangifer tarandus, Winter Ranges in Westcentral
Alberta, Pack Size of Wolves, Canis lupus, on, 313
Carp, Common, 83,422
Carp, Cyprinus carpio, of the Churchill River System in Mani-
toba, Northern Range Expansion and Invasion by the
Common, 83
Carver, D.B., 452
Carya spp., 226
Castor canadensis, 62,205,331,368
canadensis caecator, 51
Cat, Domestic, 46
Cat’s-ear, Hairy, 154
Caterpillar, Forest Tent, 433
Catfish, Black, 425
Blue, 424
Brown, 425
Channel, 422
Flathead, 424
Loricariid, 421
White, 424
Yellow, 425
Catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf.
P. brachypomus, in British Columbia, First Records
of the Yellow Bullhead, Ameiurus natalis, a Loricariid,
421
Catling, P., Review by, 249
Catling, P.M., 351
Catling, P.M. and G. Mitrow. Regional Variation in Amelanchi-
er in the Whitewood area of Southeastern Saskatche-
wan and the First Saskatchewan Records of Amelanchi-
er sanguinea, 428
Catling, P.M. and S. Carbyn. Recent Invasion, Current Status
and Invasion Pathway of European Common Reed,
Phragmites australis subspecies australis, in the
Southern Ottawa District, 307
Catling, P.M., R. Hutchinson, and P.M. Brunelle. Use of Salt-
marsh by Dragonflies (Odonata) in the Baie des Cha-
leurs Region of Quebec and New Brunswick in Late
Summer and Autumn, 413
Catostomus catostomus, 422,481
commersoni, 422
Cedar, White, 300
Celithemis elisa, 417
Centaurea nigra, 354
Centrosaurus apertus, 408
Cerastium pumilum, 320
2007
semidecandrum, 320
Ceratocephalus testiculatus, 319
Ceratodon purpureus, 358
Certhia americana, 303
Cervus elaphus, 110,188,314
Cervus elaphus, Calf Observed Near Bancroft, Ontario, Late-
born Elk, 188
Cervus elaphus, Carcass from a Small Lake in Riding Moun-
tain National Park, Manitoba, Probable Black Bear,
Ursus americana, Retrieval of an Elk, 110
Chalcosyrphus, 20
nemorum, \7
Chamaecyparis nootkatensis, 172
Chamaedaphne calyculata, 23
Channa sp., 424
Char [Charr], Arctic, 199,235,422
Chasmosaurus, 403
belli, 403
humeri, 408
irvinensis, 403
mariscalensis, 403
russelli, 403
Chasmosaurus belli (Dinosauria: Ornithischia), A Reevalu-
ation of Sexual Dimorphism in the Postcranium of
the Chasmosaurine Ceratopsid, 403
Cheatgrass, 176
Cheilosia rita, 17
tristis, 17
Chen caerulescens caerulescens, 213
Chen caerulescens caerulescens, Exceeding the Carrying
Capacity of the Fraser River Delta’s Brackish Marsh-
es?, Are Lesser Snow Geese, 213
Chenopodium album, 337
bonus-henericus, 340
fremontii, 340
pratericola, 336
quinoa, 340
subglabrum, 335
Chenopodium subglabrum (Chenopodiaceae), in Canada, A
Conservation Evaluation of Smooth Goosefoot, 335
Cherry, 360
Pin, 50
Chickadee, Black-capped, 294,302
Boreal, 302
Chickweed, Curtis’ Mouse-ear, 320
Lesser, 320
Small Mouse-ear, 320
Chimpanzee, 264
Chipmunk, 264,295
Eastern, 51
Chitonotus pugetensis, 364
Choristoneura fumiferana, 50,299
Chub, Flathead, 422
Lake, 422
Cifuni, S.M., 478
Circus cyaneus, 109
Cisco, 480
Shortjaw, 482
Cladina rangiferina, 358
stellaris, 358
Cladonia spp, 171
Cladonia, 171
Clarias sp., 424
Clarke, J.B., Review by, 247
Classen, M.-C., 347
Claytonia perfoliata, 180
INDEX TO VOLUME 120
513
Cleavers, 180
Clethrionomys sp., 291
gapperi, 50,331
Clethrionomys gapperi, in Newfoundland: Implications for
the Endangered Newfoundland Marten, Martes amer-
icana atrata, First Record of the Southern Red-Backed
Vole, 5O
Clintonia borealis, 357
Cluff, H.D. Extension of Coyote, Canis latrans, Breeding
Range in the Northwest Territories, Canada, 67
Clupea harengus, 96
Coad, B.W., 241
Coad, B.W. and A.M. Murray, Review by, 116
Coad, B.W., Review by, 114
Cody Receives Yukon Biodiversity Awareness Award, Bill, 258
Coenonypha tullia nipisiquit, 419
Colaptes auratus, 294,303
Coleoptera, 296
Colinus virginianus, 452
Colinus virginianus, by Mammalian Mesopredators: Does
the Problem-Individual Paradigm Fit?, Predation on
Artificial Nests of Northern Bobwhites, 452
Collinsia, 180
grandiflora, 180
Collomia linearis, 176
tenella, 175
Collomia, Narrow-leaved, 176
Slender, 175
Collomia, Collomia tenella, in Canada, Conservation Evalu-
ation of Slender, 175
Collomia tenella, in Canada, Conservation Evaluation of Slen-
der Collomia, 175
Colorado, Multiple Scale Den Site Selection by Swift Foxes,
Vulpes velox, in Southeastern, 31
Colossoma sp., 424
bidens, 424
Columba spp., 57
livia, 294
Comptonia peregrina, 354
Coot, 2
Cordulegaster bidentata, 350
boltonii, 347
dorsalis, 347
heros, 350
maculata, 347
Cordulegaster dorsalis (Odonata: Cordulegastridae) in an
Ephemeral Habitat in Southwestern British Columbia,
Life History Phenology and Sediment Size Association
of the Dragonfly, 347
Coregonus artedi, 480
clupeaformis, 422,480
zenithicus, 482
Corema, 357
conradii, 351
Corn, 226,294
Cornus canadensis, 354,440
Corvus brachyrhyncos, 109
corax, 109,110
Corylus cornuta, 170
Cotton, 226
Cottontail, Nuttall’s, 165
Cottonwood, 226
Cottus asper, 425
Couesius plumbeus, 422
Cougar, 40,314.438.476
Eastern, 438
514
Cougar, Puma concolor, Presence in Southern New Bruns-
wick, Eastern Canada, MtDNA Analyses on Hair
Samples Confirm, 438
Cow-wheat, 170
Cowbird, Brown-headed, 294
Coyote, 27,31,40,67,110,209,314,342,472
Eastern, 50,263,474,478
Coyote, Canis latrans, Breeding Range in the Northwest
Territories, Canada, Extension of, 67
Coyotes Canis latrans, An Ethogram Developed on Captive
Eastern, 263
Coyotes, Canis latrans, Conditions for Sexual Interactions
Between Wild Grey Wolves, Canis lupus, and, 27
Coyotes, Canis latrans, in a Heavily Urbanized Area: Impli-
cations for Ecosystem Management, Use of “Micro”-
Corridors by Eastern, 474
Coyotes, Canis latrans, in an Urban Area, Rat Poison Kills
a Pack of Eastern, 478
Crab, Dungeness, 364
Lady, 106
Redrock, 363
Crabs, Ovalipes ocellatus, in the Gulf of Maine, Lady, 106
Crane, Sandhill, 68
Crappie, Black, 422
Crataegus cf. chrysocarpa, 354
Creeper, Brown, 302
Crepis tectorum, 354
Crins, W.J., 15
Crocuta croenta, 52
Cronin, M.A., 323
Crow, American, 109
Ctenicera cruciatus, 20
hieroglyphica, 19
insidiosa, 19
resplendens, 19
spinosa, 19
triundulata, 18
vulnerata, 18
Cudweed, Lowland, 164
Culaea inconstans, 422
Curran, W.J., 50
Cutthroat, West-slope, 421
Cyanocitta cristata, 294
Cyclopterus lumpus, 464
Cygnus buccinator, 220
Cynomys ludovicianus, 31
Cynosurus echinatus, 148
Cyprinella lutrensis, 424
Cyprinus carpio, 83,422
Cyprinus carpio, of the Churchill River System in Manitoba,
Northern Range Expansion and Invasion by the Com-
mon Carp, 83
Cypripedium acaule, 354
Cytisus scoparius, 148,155,160
Dactylis glomerata, 154,160
Daisy, Cut-leaved, 184
Salish, 137
Dalopius spp., 19
cognatus, 19
fuscipes, 19
Dandelion, Common, 320
Danthonia spicata, 354
unispicata, 164,184
Darimont, C.T., 61
Darner, Canada, 417
THE CANADIAN FIELD-NATURALIST
Vol. 120
Common Green, 417
Lake, 417
Lance-tipped, 417
Shadow, 417
Variable, 417
Dasysyrphus pauxillus, 17
Davies, S., A. Griffiths, and T.E. Reimchen. Pacific Hagfish,
Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei,
and Scavenger Activity on Tethered Carrion in Subtidal
Benthic Communities off Western Vancouver Island,
363
Deer, 68
Black-tailed, 63
Marsh, 64
Mule, 40,314
Pampas, 64
Sitka Black-tailed, 61
White-tailed, 29,40,64,68,209,3 13,367
Delphinapterus leucas, 457
Delphinium menziesii, 154
Delphinus delphis, 97
Demarchi, M.W. Are Lesser Snow Geese, Chen caerulescens
caerulescens, Exceeding the Carrying Capacity of
the Fraser River Delta’s Brackish Marshes?, 213
Deschampsia danthonioides, 164
flexuosa, 354
Desert-parsley, Swale, 184
Desroches, J.-F. Extension de |’ aire de distribution de 1’ Umbre
de vase, Umbra limi, dans le nord-est du Québec, 238
Desroches, J.-F., I. Picard, and J.E. Maunder. The Mink Frog,
Rana septentrionalis, in Southeastern Labrador, 239
Desrosiers, J., 169
Dewey, T.A., 39
Diamond, A.W., 236
Dianthus armeria, 354
Dichanthelium (sub Panicum) depauperatum vat. psilophyl-
lum f. cryptostachys, 360
acuminatum vat. fasciculatum, 354
depauperatum, 354
linearifolium, 354
sabulorum var. thinium, 360
Dicranum sp., 180
scoparium, 358
Dicrostonyx groenlandicus, 466
Didelphis virginiana, 229,452
Diervella lonicera, 354
Distichlis spicata, 214
Doellingeria umbellata var. umbellata, 354
Dog, 479
African Wild, 52
Domestic, 263
Dogtail, Hedgehog, 148
Dollar, Silver, 424
Dolphin, Atlantic White-sided, 95
Common Bottlenose, 97
Dusky, 97
Short-beaked Common, 97
White-beaked, 95
White-sided, 95
Donaldson, M.R., Review by, 121
Donovan, M. Conservation Evaluation of the Bog Bird’s-foot
Trefoil, Lotus pinnatus, in Canada, 157
Dorocordulia lepida, 417
Douglas, G.W. and J. Desrosiers. Conservation Evaluation of
Pacific Rhododendron, Rhododendron macrophyllum,
in Canada, 169
2007
Douglas, G.W. and J.L. Penny. Conservation Evaluation of
Slender Collomia, Collomia tenella, in Canada, 175
Douglas, G.W. and J.L. Penny. Conservation Evaluation of the
Small-flowered Tonella, Tonella tenella, in Canada,
179
Douglas, G.W. and M. Ryan. Conservation Evaluation of the
Prairie Lupine, Lupinus lepidus var. lepidus, in Cana-
da, 147
Douglas, G.W. and M. Ryan. Conservation Evaluation of
the Seaside Birds-foot Trefoil, Lotus formosissimus,
in Canada, 153
Douglas, G.W., J.L. Penny, and K. Barton. Conservation Eval-
uation of Stoloniferous Pussytoes, Antennaria fla-
gellaris, in Canada, 183
Douglas, G.W., J.L. Penny, and K. Barton. Conservation Eval-
uation of the Pacific Population of Dwarf Woolly-
heads, Psilocarphus brevissimus var. brevissimus, in
Canada, 163
Douglas, H., 15
Douglas 1938 — 2005, A Tribute to George Wayne, 135
Douglas-fir, 148,158,164,169,176,184,348
Coastal, 180
Dove, Mourning, 294
Rock, 294
Dragonflies (Odonata) in the Baie des Chaleurs Region of
Quebec and New Brunswick in Late Summer and
Autumn, Use of Saltmarsh by, 413
Dragonfly, 347
Dragonfly Cordulegaster dorsalis (Odonata: Cordulegastri-
dae) in an Ephemeral Habitat in Southwestern British
Columbia, Life History Phenology and Sediment Size
Association of the, 347
Dragonlet, Seaside, 417
Drosera intermedia, 354
Dryocopus pileatus, 299
Duck, 2
Common Eider, 236
Ring-necked, 6
Wood, 236,303
Duck Nests in Saskatchewan, Summer Movements and Impact
of Individual Striped Skunks, Mephitis mephitis, on,
342
Ducks, Ring-billed Gull, Larus delawerensis, Food Piracy
on Diving, 109
Duffe, J., 289
Eagle, Bald, 109,237,477
Eagle, Haliaeetus leucocephalus, Preying on Maritime Garter
Snake, Thamnophis sirtalis pallidulus, on Cape Bre-
ton Island, Nova Scotia, Bald, 477
Eatough, D.L., 474,478
Eel, American, 477
Editor’s Report for Volume 119, 260
Eider, Common, 233
Eider, Somateria mollissima, Duckling, Osprey, Pandion hali-
aetus, Depredates Common, 236
Elk, 110,188,314
Elk, Cervus elaphus, Calf Observed Near Bancroft, Ontario,
Late-born, 188
Elk, Cervus elaphus, Carcass from a Small Lake in Riding
Mountain National Park, Manitoba, Probable Black
Bear, Ursus americana, Retrieval of an, 110
Elymus canadensis, 336
lanceolatus ssp. lanceolatus, 336
repens, 355
trachycaulus, 184
INDEX TO VOLUME 120
515
Emerald, Kennedy's, 417
Petite, 417
Ski-tipped, 417
Emphoropsis, 180
Enallagma aspersum, 417
civile, 413
durum, 417
ebrium, 416
hageni, 415
Enteromorpha sp., 417
Epigaea repens, 355
Epilobium angustifolium, \7\
densiflorum, 160
Epinettes, 238
Epistrophe nitidicollis, \7
Eptatretus stoutii, 363
Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and
Scavenger Activity on Tethered Carrion in Subtidal
Benthic Communities off Western Vancouver Island,
Pacific Hagfish, 363
Eptesicus fuscus, 39
Equus cabalus, 314
Ericameria nauseosus var. speciosa, 184
Erigeron, 137
compositus var. glabratus, 184
leibergii, 137
salishii, 137
strigosus, 355
Eriogonum heracleoides var. angustifolium, 184
ovalifolium var. nivale, 184
Eristalis dimidiatus, 17
Erophila verna, 320
Erpobdella, 449
Erskine, A.J. Recent Declines of House Sparrows, Passer
domesticus, in Canada’s Maritime Provinces, 43
Erythrodiplax berenice, 413
Esox lucius, 421,481
Eubalaena glacialis, 95
Eupeodes americanus, 17
perplexus, 17
pomus, 17
Euphausiid, 96
Euphorbia esula var. esula, 337
Fairybells, Hooker’s, 171
Falco columbarius, 46
sparverius, 290,303
Falsebox, 169
Fargey, P., Review by, 382
Feathermoss, 169
Felis catus, 46
concolor, 314
Fern, Bracken, 172
Maidenhair, 138
Fescue, 226
Festuca spp., 226
filiformis, 355
trachyphylla, 355
Fir, Amabilis, 172
Balsam, 50,100,236,299,433,440
Douglas, 11
Subalpine, 100
Fireweed, 171
Fisher, 62,101,229
Flax, 343
Fleabane, 137
516
Leiberg’s, 137
Flicker, Northern, 294,302
Forktail, Eastern, 417
Fox, Arctic, 36
Red, 36,51,68,263,342,468
San Joaquin Kit, 52,210
Santa Cruz Island, 210
Swift, 31,52
Foxes, Vulpes velox, in Southeastern Colorado, Multiple Scale
Den Site Selection by Swift, 31
Foxglove, Middleton False, 357
Fragaria virginiana, 355
Freedman, B., 298
Frog, Boreal Chorus, 87
Columbia Spotted, 87
Green, 294
Mink, 239
Western Spotted, 87
Wood, 87
Frog, Rana septentrionalis, in Southeastern Labrador, The
Mink, 239
Frostweed, Long-Branch, 359
Fucus vesiculosus, 236
Fulmar, 458
Fulmarus glacialis, 458
Gale, Sweet, 172,240
Galium aparine, 180
Gallant, D., 438
Galvez, I., 241
Gambusia affinis, 421
Gasterosteus aculeatus, 240,422
Gastromyzon sp., 424
Gaultheria ovatifolia, 171
procumbens, 355
shallon, 11,172
Gawn, J., Reviews by, 490,499
Gawn, M., Reviews by, 119,379
Gaylussacia baccata, 355
dumosa, 360
Geese, Chen caerulescens caerulescens, Exceeding the Car-
rying Capacity of the Fraser River Delta’s Brackish
Marshes?, Are Lesser Snow, 213
Gese, E.M., 31
Gilhen, J., 241
Glider, Wandering, 417
Globicephala melas, 97
Glycine max, 226
Gnaphalium palustre, 164
Goat, Mountain, 314
Golden-aster, Hairy, 336
Goldeneye, Common, 5,109
Goldenrod, 226
Goldeye, 422
Goldfinch, American, 294
Goldfish, 422
Goldsborough, L.G., 83
Gomphid, 350
Good King Henry, 340
Goodwin, C.E., 319
Goodwin, C.E., Review by, 379
Goose spp., 5
Canada, 109
Lesser Snow, 213
White, 214
Goosefoot, Fremont’s, 340
THE CANADIAN FIELD-NATURALIST
Vol. 120
Narrow-leaved, 336
Smooth, 335
Goosefoot, Chenopodium subglabrum (Chenopodiaceae), in
Canada, A Conservation Evaluation of Smooth, 335
Gossypium spp., 226
Gowans, S., 93
Grackle, Common, 294
Grass, Orchard, 154,160
Sand, 336
Switch, 226
Grayling, Arctic, 422
Grebe, 2
Horned, 109
Red-necked, 109
Greenling, Kelp, 363
Griffiths, A., 363
Grouseberry, 172
Grus canadensis, 68
Gull, 2
Bonaparte’s, 5
Ring-billed, 109
Gull, Larus delawerensis, Food Piracy on Diving Ducks,
Ring-billed, 109
Gulo gulo, 111
Gum, Sweet, 226
Guppy, 422
Haemopis, 443
caeca, 450
grandis, 443
kingi, 444
lateromaculata, 443
marmorata, 443
plumbea, 447
sanguisuga, 444
septagon, 444
terrestris, 447
Haemopis lateromaculata (Hirudinea: Haemopidae): Its
North America Distribution and Additional Notes on
Species Description, The Leech, 443
Hagfish, Pacific, 363
Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus col-
liei, and Scavenger Activity on Tethered Carrion in
Subtidal Benthic Communities off Western Vancou-
ver Island, Pacific, 363
Hairgrass, Annual, 164
Early, 154,320
Silvery, 320
Haliaeetus leucocephalus, 109,237,477
Haliaeetus leucocephalus, Preying on Maritime Garter Snake,
Thamnophis sirtalis pallidulus, on Cape Breton Is-
land, Nova Scotia, Bald Eagle, 477
Hall, M.N., 57
Hamr, J., 57
Hanke, G.F., M.C.E. McNall, and J. Roberts. First Records of
the Yellow Bullhead, Ameiurus natalis, a Loricariid
Catfish, Panaque suttonorum, and -a Silver Pacu,
Piaractus cf. P. brachypomus, in British Columbia,
421
Hare, Arctic, 51,466
Snowshoe, 51,68,205
Harrier, Northern, 109
Harrington, FH. Double Marking in Arctic Wolves, Canis
lupus arctos: Influence of Order on Posture, 471
Haskell, S.P. First Record of a River Otter, Lontra canadensis,
Captured on the Northeastern Coast of Alaska, 235
2007
Hawk, Sharp-shinned, 46
Hawkweed, White, 170
Hazelnut, Beaked, 170
Hearn, B.J., J.T. Neville, W.J. Curran, and D.P. Snow. First
Record of the Southern Red-Backed Vole, Clethri-
onomys gapperi, in Newfoundland: Implications for
the Endangered Newfoundland Marten, Martes amer-
icana atrata, 50
Hedge-parsley, Upright, 180
Heide-Jgrgensen, M.P., 457
Helianthemum canadense, 355
Helianthus petiolaris ssp. petiolaris, 336
Helophilus fasciatus, \7
Hemlock, Coastal Western, 172
Eastern, 290
Western, 11,170,348
Herring, 96
Heterotheca villosa var. villosa, 336
Hexagrammos decagrammus, 363
Hickory, 226
Hieracium albiflorum, 170
pilosella, 355
piloselloides, 355
Hiodon alosoides, 422
Holcus lanatus, 154
Holmes, R.B., 403
Holodiscus discolor, 158,171
Horse, Wild, 314
Houston, A.E., 225,452
Houston, C.S., Reviews by, 124,382,385,496,497,498
Hoverfly, 15
Hovingh, P. The Leech Haemopis lateromaculata (Hirudinea:
Haemopidae): Its North America Distribution and
Additional Notes on Species Description, 443
Huckleberry, 348,360
Red, 11,172
Hudsonia ericoides, 355
Huettmann, F., Reviews by, 125,254,491
Hutchinson, R., 413
Hwang, Y.T. and S. Lariviére. A Test of Interspecific Effects
of Introduced Eastern Grey Squirrels, Sciurus caro-
linensis, on Douglas’s Squirrels, Tamiasciurus dou-
glasii, in Vancouver, British Columbia, 10
Hydrolagus colliei, 363
Hydrolagus colliei, and Scavenger Activity on Tethered Car-
rion in Subtidal Benthic Communities off Western
Vancouver Island, Pacific Hagfish, Eptatretus stoutii,
Spotted Ratfish, 363
Hyena, Spotted, 52
Hylaeus elliptica, 19
Hylocomium splendens, 171
Hymenoptera, 295
Hypericum perforatum, 355
Hypochaeris radicata, 154
Hypophthalmichthys sp., 424
Hypostomus, 424
Ictalurus furcatus, 424
punctatus, 422
Ischnura verticalis, 416
Tsoetes nuttallii, 160
Jackal, 263
Jay, Blue, 294
INDEX TO VOLUME 120
517
Jennings, J.B., M.L. Kennedy, A.E. Houston, and B.D. Carv-
er. Predation on Artificial Nests of Northern Bob-
whites, Colinus virginianus, by Mammalian Meso-
predators: Does the Problem-Individual Paradigm
Fit?, 452
John, R., Reviews by, 113,114,119,123,244,251,255,38 1 386,
388,389,483,484,488 489
Johnson, G., 298
Junco hyemalis, 294
Junco, Dark-eyed, 294
Juncus balticus, 417
effusus, 355
gerardii, 417
Juneberry, 360
Jung, T.S., A.M. Runck, D.W. Nagorsen, B.G. Slough, and
T. Powell. First Records of the Southern Red-backed
Vole, Myodes gapperi, in the Yukon, 331
Jung, T.S., B.G. Slough, D.W. Nagorsen, T.A. Dewey, and
T. Powell. First Records of the Northern Long-eared
Bat, Myotis septentrionalis, in the Yukon Territory,
39
Juniper, 32
Creeping, 336
Juniperus communis var. depressa, 355
horizontalis, 336
monosperma, 32
Kallemeyn, L.W., 71
Kalmia, 240
angustifolia, 355
Kenn, S., 438
Kennedy, A., 233
Kennedy, M.L., 225,452
Kestrel, 302
American, 290,303
Kindbergia oregana, 180
Kinnikinnick, 172
Kirk, D.A., 298
Kitchen, A.M., E.M. Gese, and S.G. Lupis. Multiple Scale
Den Site Selection by Swift Foxes, Vulpes velox, in
Southeastern Colorado, 31
Kneteman, J., 313
Knotweed, Close-flowered, 164
Common, 164
Kogia breviceps, 97
Koi, 424
Kudzu, 226
Kuzyk, G.W., J. Kneteman, and F.K.A. Schmiegelow. Pack
Size of Wolves, Canis lupus, on Caribou, Rangifer
tarandus, Winter Ranges in Westcentral Alberta, 313
l’Umbre de vase, 238
l’Umbre de vase, Umbra limi, dans le nord-est du Québec,
Extension de l’aire de distribution de, 238
Labrador, The Mink Frog, Rana septentrionalis, in South-
eastern, 239
Lactuca canadensis var. longifolia, 355
Lagenorhynchus acutus, 95
albirostris, 95
obscurus, 97
Laidre, K.L., M.P. Heide-Jorgensen, and J.R. Orr. Reactions
of Narwhals, Monodon monoceros, to Killer Whale,
Orcinus orca, Attacks in the Eastern Canadian Arctic,
457
518
Lamb’s-quarters, 337
Lambdina fiscellaria fiscellaria, 50
Landriault, L.J., M.N. Hall, J. Hamr, and F.F. Mallory. Long-
range Homing by an Adult Female Black Bear, Ursus
americanus, 57
Larch, American, 433
Larivée, J., 289
Larivieére, S., 10
Lariviére, S., L.R. Walton, and F. Messier. Summer Move-
ments and Impact of Individual Striped Skunks,
Mephitis mephitis, on Duck Nests in Saskatchewan,
342
Larix laricina, 314,355,433
Larkspur, Menzies, 154
Larus delawerensis, 109
Philadelphia, 5
Larus delawerensis, Food Piracy on Diving Ducks, Ring-
billed Gull, 109
Lasioglossum admirandus, 19
divergens, 19
laevissimus, 19
nymphaearum, 19
quebecensis, 19
rohweri, 19
versans, 19
zephyrus, 19
Lasionycteris noctivagans, 39
Latissimus dorsi, 406
Lauff, R., Reviews by, 118,121,243,380
Lawlor, J.L., 93
Leatherleaf, 23
Lechea intermedia, 355
intermedia var. intermedia, 361
intermedia vat. juniperina, 361
Ledum groenlandicum, 172
Leech, 443
Leech Haemopis lateromaculata (Hirudinea: Haemopidae):
Its North America Distribution and Additional Notes
on Species Description, The, 443
Lejota aerea, 17
cyanea, 17
Lemming, 466
Lentil, 343
Leontodon autumnalis, 355
Lepeophtheirus salmonis, 199
Lepeophtheirus salmonis Infestation Levels on Juvenile Wild
Pink, Oncorhynchus gorbuscha, and Chum, O. keta,
Salmon to Arrival of Parasitized Wild Adult Pink
Salmon, Response of the Sea Louse, 199
Lepidosetta bilineata, 364
Lepisosteus oculatus, 424
osseus, 423
platostomus, 424
Lepomis gibbosus, 422
macrochirus, 422
Lepus americanus, 68,205
americanus struthopus, 51
arcticus, 466
arcticus bangsii, 51
Lestes congener, 415
disjunctus, 415
forcipatus, 416
inaequalis, 417
rectangularis, 417
unguiculatus, 417
Leucanthemum vulgare, 355
THE CANADIAN FIELD-NATURALIST
Vol. 120
Leucorrhinia frigida, 417
Libellula luctuosa, 417
pulchella, 417
quadrimaculata, 415
Lichen, 172
Limonius aeger, 19
confusus, 19
Linaria canadensis, 355
genistifolia ssp. dalmatica, 176
Lindell, J., Review by, 252
Linnaea borealis, 172
Lion, 52
Liquidambar styraciflua, 226
Liriodendron tulipifera, 226
Listera cordata, 170
Liu, P.S., 225
Lobelia dortmanna, 240
Lobelia, Water, 240
Lomatium ambiguum, 184
Lontra canadensis, 235
canadensis degener, 51
Lontra canadensis, Captured on the Northeastern Coast of
Alaska, First Record of a River Otter, 235
Loon, 2
Looper, Hemlock, 50
Loosestrife, Purple, 307
Lota lota, 422,480
Lotus corniculatus, 157
formosissimus, 153,157
pinnatus, 153,157
Lotus formosissimus, in Canada, Conservation Evaluation of
the Seaside Birds-foot Trefoil, 153
Lotus pinnatus, in Canada, Conservation Evaluation of the
Bog Bird’s-foot Trefoil, 157
Louse, Sea, 199 ,
Louse Lepeophtheirus salmonis Infestation Levels on Juvenile
Wild Pink, Oncorhynchus gorbuscha, and Chum, O.
keta, Salmon to Arrival of Parasitized Wild Adult Pink
Salmon, Response of the Sea, 199
Lumpsucker, 464
Lupine, Prairie, 147
Silky, 176
Lupine, Lupinus lepidus var. lepidus, in Canada, Conservation
Evaluation of the Prairie, 147
Lupinus lepidus, 147
lepidus var. lepidus, 147
lepidus var. lobbii, 147
lyallii, 147
minimus, 147
sericeus, 176
Lupinus lepidus var. lepidus, in Canada, Conservation Eval-
uation of the Prairie Lupine, 147
Lupis, S.G., 31
Lutra canadensis, 62
Luzula multiflora, 355
Lycaena dospassosi, 419
Lycaon pictus, 52
Lycopodium dendroideum, 355
obscurum, 357
tristachyum, 355
Lygodesmia juncea, 336
Lynx lynx susolanus, 51
pardinus, 229
rufus, 230
Lynx, Canada, 51
Iberian, 229
2007
Lysichitum americanum, 172
Lythrum salicaria, 307
MacKinnon, C. and A. Kennedy. An Observation of the Spring
2006 Migration of Black Scoter Melanitta nigra, in
Northumberland Strait, Interrupted by the Confeder-
ation Bridge, New Brunswick — Prince Edward Island,
233
Madtom, Tadpole, 424
Mahonia nervosa, 171
Maianthemum canadense, 355
Maine, Lady Crabs, Ovalipes ocellatus, in the Gulf of. 106
Malacosoma disstria, 433
Mallard, 6
Mallon, J.C. and R.B. Holmes. A Reevaluation of Sexual
Dimorphism in the Postcranium of the Chasmosaurine
Ceratopsid Chasmosaurus belli (Dinosauria: Ornithis-
chia), 403
Mallory, F.F., 57
Malus pumila, 355
Manitoba, Northern Range Expansion and Invasion by the
Common Carp, Cyprinus carpio, of the Churchill
River System in, 83
Manitoba, Probable Black Bear, Ursus americana, Retrieval
of an Elk, Cervus elaphus, Carcass from a Small Lake
in Riding Mountain National Park, 110
Maple, 226
Big Leaf, 11,180
Douglas, 171
Red, 299
Sugar, 299
Vine, 11,171,348
Marczak, L.B., J.S. Richardson, and M.-C. Classen. Life
History Phenology and Sediment Size Association of
the Dragonfly Cordulegaster dorsalis (Odonata: Cor-
dulegastridae) in an Ephemeral Habitat in South-
western British Columbia, 347
Marine Turtle Newsletter, 132,258,392,502
Marten, 62
American, 100
Newfoundland, 50
Pine, 52
Marten, Martes americana atrata, First Record of the South-
ern Red-Backed Vole, Clethrionomys gapperi, in
Newfoundland: Implications for the Endangered New-
foundland, 50
Marten, Martes americana, in Western Alberta Boreal Forests,
Winter Habitat Use by American, 100
Martes americana, 62,100
americana atrata, 50
martes, 52
pennanti, 62,101,229
Martes americana atrata, First Record of the Southern
Red-Backed Vole, Clethrionomys gapperi, in New-
foundland: Implications for the Endangered New-
foundland Marten, 50
Martes americana, in Western Alberta Boreal Forests, Win-
ter Habitat Use by American Marten, 100
Mary, Blue-eyed, 180
Maunder, J.E., 239
McCarthy, J., Review by, 387
McCorquodale, D., Review by, 486
MeNall, M.C.E., 421
Meadow-foxtail, Carolina, 164
Meadowhawk, Autumn, 417
Band-winged, 417
INDEX TO VOLUME 120
519
Black, 417
Cherry-faced, 417
Saffron-winged, 417
White-faced, 417
Mech, L.D. Urine-marking and Ground-scratching by Free-
ranging Arctic Wolves, Canis lupus arctos, in Sum-
mer, 466
Meganyctiphanes norvegica, 96
Megaptera novaeangliae, 95
Megascops asio, 289
Megascops asio, Population in an Apple-Producing Region
of Southern Quebec, First Observations of an East-
ern Screech-Owl, 289
Meikle, D.B., 192
Melampyrum lineare, 357
lineare var. lineare, 170
Melanitta spp., 232
nigra, 233
Melanitta nigra, in Northumberland Strait, Interrupted by the
Confederation Bridge, New Brunswick — Prince Ed-
ward Island, An Observation of the Spring 2006
Migration of Black Scoter, 233
Melanitta spp., Migrations Interrupted by Confederation
Bridge: An Update, Scoter, 232
Melanostoma, 19
mellinum, 17
Melanotus castanipes, 19
Melanpyrum lineare, 355
Meles meles, 52
Melic, Harford’s, 180
Melica harfordii, 180
Melilotus spp., 337
Meliscaeva cinctellam 17
Mennell, R.L., 87
Mephitis mephitis, 229,342,452
Mephitis mephitis, on Duck Nests in Saskatchewan, Summer
Movements and Impact of Individual Striped Skunks,
342
Merganser, Red-breasted, 109
Mergus serrator, 109
Merlin, 46
Messier, F., 342
Metynnis sp., 424
Mice, Peromyscus leucopus: The Effects of Forest Patch Size,
Edge and Surrounding Vegetation Type, Annual
Variation in Habitat Use by White-footed, 192
Micropterus dolomieu, 421
salmoides, 422
Microtus sp., 52,291
pennsylvanicus, 264,294,332,436
pennsylvanicus terraenovae, 50
Milk-vetch, Timber, 176
Mimulus guttatus, 158
Mineau, P., 289
Miner’s-lettuce, 180
Mink, 51,62
Minnesota and Ontario, Lake Sturgeon, Acipenser fulvescens,
Movements in Rainy Lake, 71
Minnow, Fathead, 421
Misgurnus anguillicaudatus, 424
Mitchell, S., Review by, 490
Mitrow, G., 428
Mollibdella, 444
grandis, 444
Mollugo verticillata, 355
Molothrus ater, 294
520
Monkey-flower, Yellow, 158
Monoclonius belli, 403
Monodon monoceros, 457
Monodon monoceros, to Killer Whale, Orcinus orca, Attacks
in the Eastern Canadian Arctic, Reactions of Nar-
whals, 457
Monopterus sp., 424
Monotropa hypopithys, 355
Montia parvifolia, 158
Montia, Small-leaved, 158
Mooreobdella, 449
Moose, 51,205,313,367
Morone saxatilis, 422
Morton, A. and R. Williams. Response of the Sea Louse
Lepeophtheirus salmonis Infestation Levels on Juve-
nile Wild Pink, Oncorhynchus gorbuscha, and Chum,
O. keta, Salmon to Arrival of Parasitized Wild Adult
Pink Salmon, 199
Mosquitofish, 421
Western, 422
Moss, Heron’s-bill, 180
Lanky, 171
Oregon Beaked, 180
Step, 171
Mouse, Deer, 50,57,332
House, 51
Meadow Jumping, 332
White-footed, 192
Mousetail, Tiny, 164
Mus musculus domesticus, 51
Muskox, 466
Muskrat, 51
Mustela ermina, 62
erminea richardsonii, 51
vison, 51,62
Myleus sp., 424
Myodes californicus, 332
gapperi, 331
glareolus, 332
rufocanus, 332
rutilus, 331
Myodes gapperi, in the Yukon, First Records of the Southern
Red-backed Vole, 331
Myosotis stricta, 320
Myosurus minimus, 164
Myotis evotis, 39
keenii, 40
lucifugus, 39
lucifugus lucifugus, 51
septentrionalis, 39,51,332
volans, 39
Myotis septentrionalis, in the Yukon Territory, First Records
of the Northern Long-eared Bat, 39
Myoxocephalus thompsonii, 480
Myoxocephalus thompsonii, in Northeastern Alberta, A New
Record of Deepwater Sculpin, 480
Myrica gale, 172,240
pensylvanica, 355
Myxocyprinus asiaticus, 424
Nagorsen, D.W., 39,331
Narwhal, 457
Narwhals, Monodon monoceros, to Killer Whale, Orcinus
orca, Attacks in the Eastern Canadian Arctic, Reac-
tions of, 457
Neely, D.A., 480
THE CANADIAN FIELD-NATURALIST
Vol. 120
Neoascia distincta, 17
Neohypdonus tumescens, 19
Nephelopsis, 449
Neuhold, J., 188
Neville, J.T., 50
New Brunswick, Eastern Canada, MtDNA Analyses on Hair
Samples Confirm Cougar, Puma concolor, Presence
in Southern, 438
New Brunswick, Effects of Timber Harvesting and Plantation
Development on Cavity-nesting Birds in, 298
New Brunswick in Late Summer and Autumn, Use of Salt-
marsh by Dragonflies (Odonata) in the Baie des
Chaleurs Region of Quebec and, 413
New Brunswick — Prince Edward Island, An Observation of
the Spring 2006 Migration of Black Scoter Melanitta
nigra, in Northumberland Strait, Interrupted by the
Confederation Bridge, 233
Newfoundland: Implications for the Endangered Newfound-
land Marten, Martes americana atrata, First Record
of the Southern Red-Backed Vole, Clethrionomys
gapperi, in, 50
Newfoundland Marten, Martes americana atrata, First Record
of the Southern Red-Backed Vole, Clethrionomys
gapperi, in Newfoundland: Implications for the En-
dangered, 50
Newt, Roughskin, 90
Nine-bark, 158
Noel, L.E., M.K. Butcher, M.A. Cronin, and B. Streever.
Assessment of Effects of an Oil Pipeline on Caribou,
Rangifer tarandus granti, Use of Riparian Habitats
in Arctic Alaska, 2001-2003, 323
Nol, E., H. Douglas, and W.J. Crins. Responses of Syrphids,
Elaterids and Bees to Single-tree Selection Harvesting
in Algonquin Provincial Park, Ontario, 15
North America Distribution and Additional Notes on Species
Description, The Leech Haemopis lateromaculata
(Hirudinea: Haemopidae): Its, 443
Northwest Territories, Canada, Extension of Coyote, Canis
latrans, Breeding Range in the, 67
Notropis hudsonius, 422
Noturus gyrinus, 424
Nova Scotia, An Analysis of the Vascular Flora of Annapolis
Heathlands, 351
Nova Scotia, Bald Eagle, Haliaeetus leucocephalus, Prey-
ing on Maritime Garter Snake, Thamnophis sirtalis
pallidulus, on Cape Breton Island, 477
Nova Scotia, Temporal Variability of Cetaceans near Halifax,
93
Novakowski 1925-2004, A Tribute to Nicholas Stephen, 370
Nuphar variegata, 240
Nuthatch, 290
Red-breasted, 302
White-breasted, 302
O’Neill, J., Reviews by, 120,485,487,498
Oak, 226
Garry, 150,154,157,180
Oat, 343
Oatgrass, One-spike, 164,184
Oceanspray, 171
Odocoileus sp., 68
heminous, 40,314
hemionus sitkensis, 61
virginianus, 29,40,64,68,209,3 13,367
Oldham, M.J., C.E. Goodwin, and S. Blaney. Bur Buttercup,
Ranunculus testiculatus, New to Eastern Canada, 319
2007
Olson, S.L. Bald Eagle, Haliaeetus leucocephalus, Preying
on Maritime Garter Snake, Thamnophis sirtalis pal-
lidulus, on Cape Breton Island, Nova Scotia, 477
Omble de fontaine, 238
Oncorhynchus clarki lewisi, 421
gorbuscha, 199,421
keta, 199,421
kisutch, 421
mykiss, 90,421
mykiss aguabonita, 422
nerka, 421
tshawytscha, 421
Oncorhynchus gorbuscha, and Chum, O. keta, Salmon to
Arrival of Parasitized Wild Adult Pink Salmon, Res-
ponse of the Sea Louse Lepeophtheirus salmonis In-
festation Levels on Juvenile Wild Pink, 199
Oncorhynchus keta, Salmon to Arrival of Parasitized Wild
Adult Pink Salmon, Response of the Sea Louse Lep-
eophtheirus salmonis Infestation Levels on Juvenile
Wild Pink, Oncorhynchus gorbuscha, and Chum, 199
Ondatra zibethicus obscurus, 51
Onion, Nodding, 154
Slim-leaf, 160
Ontario, A New Record Size Wolf, Canis lupus, Group for,
367
Ontario, in 2001, Observations on the Nesting of the Northern
Hawk Owl, Surnia ulula, near Timmins and Iroquois
Falls, Northeastern, 433
Ontario, Lake Sturgeon, Acipenser fulvescens, Movements
in Rainy Lake, Minnesota and, 71
Ontario, Late-born Elk, Cervus elaphus, Calf Observed Near
Bancroft, 188
Ontario Natural Heritage Information Centre Science and
Information Newsletter 11(1) Winter 2006, 132
Ontario, Responses of Syrphids, Elaterids and Bees to Sin-
gle-tree Selection Harvesting in Algonquin Provin-
cial Park, 15
Ontario, with Some Notes on Nesting Behaviour, New Nest-
ing Records of the Le Conte’s Sparrow, Ammospiza
leconteii, from Northeastern, 22
Onychogomphus uncatus, 350
Opossum, Virginia, 229,452
Orcinus orca, 457
Orcinus orca, Attacks in the Eastern Canadian Arctic, Reac-
tions of Narwhals, Monodon monoceros, to Killer
Whale, 457
Oreamnos americanus, 314
Oregon-grape, Dull, 171
Oreochromis sp., 424
Orr, J.R., 457
Orthasterias koehleri, 364
Orthocarpus barbatus, 137
Oryzopsis asperifolia, 358
Osmia, 180
atriventris, 19
Osprey, 236
Osprey, Pandion haliaetus, Depredates Common Eider, Soma-
teria mollissima, Duckling, 236
Ottawa Field-Naturalists’ Club Awards for 2005, The, 504
Ottawa Field-Naturalists’ Club 10 January 2006, Minutes of
the 127th Annual Business Meeting of The, 393
Otter, 51
River, 62,235
Otter, Lontra canadensis, Captured on the Northeastern Coast
of Alaska, First Record of a River, 235
INDEX TO VOLUME 120
52]
Ovalipes ocellatus, \06
Ovalipes ocellatus, in the Gulf of Maine, Lady Crabs, 106
Ovibos moschatus, 466
Ovis canadensis, 314
dalli, 205
Owl, Barred, 290,302,
Great Horned, 290
Long-eared, 295
Northern Hawk, 433
Northern Saw-whet, 294,303
Saw-whet, 302
Owl, Surnia ulula, near Timmins and Iroquois Falls, North-
eastern Ontario, in 2001, Observations on the Nesting
of the Northern Hawk, 433
Owl-clover, Grand Coulee, 137
Ozotocerus bezarticus, 64
Pacu, Red-bellied, 421
Silver, 421
Pacu, Piaractus cf. P. brachypomus, in British Columbia, First
Records of the Yellow Bullhead, Ameiurus natalis, a
Loricariid Catfish, Panaque suttonorum, and a Silver,
42]
Pan troglodytes, 264
Panaque sp., 424
nigrolineatus, 424
suttonorum, 421
Panaque, Blue-eyed, 423
Royal, 424
Panaque suttonorum, and a Silver Pacu, Piaractus cf. P.
brachypomus, in British Columbia, First Records of
the Yellow Bullhead, Ameiurus natalis, a Loricariid
Catfish, 421
Pandalus platyceros, 364
Pandion haliaetus, 236
Pandion haliaetus, Depredates Common Eider, Somateria
mollissima, Duckling, Osprey, 236
Panicum virgatum, 226
Pantala spp., 418
flavescens, 416
Panthera leo, 52
Paquet, P.C., 61
Parasyrphus sp., 17
semiinterruptus, 17
Passer domesticus, 43,264
Passer domesticus, in Canada’s Maritime Provinces, Recent
Declines of House Sparrows, 43
Patrikeev, M. New Nesting Records of the Le Conte’s Spar-
row, Ammospiza leconteii, from Northeastern Ontario,
with Some Notes on Nesting Behaviour, 22
Patrikeev, M. Observations on the Nesting of the Northern
Hawk Owl, Surnia ulula, near Timmins and Iroquois
Falls, Northeastern Ontario, in 2001, 433
Paxistima myrsinites, 169
Pea, 343
Pennant, Calico, 417
Penny, G., Reviews by, 117,246
Penny, J.L., 163,175,179,183
Penny, J.L. A Tribute to George Wayne Douglas 1938 — 2005,
135
Pentaceratops, 403
Perca flavescens, 422
Perch, Yellow, 422
Percopsis omiscomaycus, 422
Percymoorensis, 444
327) THE CANADIAN FIELD-NATURALIST
caballeroi, 444
kingi, 444
lateromaculata, 444
marmorata, 444
terrestris, 444
Peromyscus sp., 294
leucopus, 192
maniculatus, 50,57,332
Peromyscus leucopus: The Effects of Forest Patch Size, Edge
and Surrounding Vegetation Type, Annual Variation
in Habitat Use by White-footed Mice, 192
Phacelia linearis, 176
Phacelia, Thread-leaved, 176
Phenacomys ungava, 332
Phoca spp., 466
hispida, 464
vitulina, 62
Phocoena phocoena, 95
Phoebe, Eastern, 294
Phragmites australis, 307
australis subspecies americanus, 307
australis subspecies australis, 307
Phragmites australis subspecies australis, in the Southern
Ottawa District, Recent Invasion, Current Status and
Invasion Pathway of European Common Reed, 307
Physalis alkekengi, 355
Physocarpus capitatus, 158
Piaractus sp., 424
brachypomus, 421
Piaractus brachypomus, in British Columbia, First Records
of the Yellow Bullhead, Ameiurus natalis, a Loricari-
id Catfish, Panaque suttonorum, and a Silver Pacu,
Piaractus cf., 421
Piaractus cf. P. brachypomus, in British Columbia, First
Records of the Yellow Bullhead, Ameiurus natalis, a
Loricariid Catfish, Panaque suttonorum, and a Silver
Pacu, 421
Picard, I., 239
Picea sp., 20,110,236,238,431
abies, 299
engelmannii, 100
glauca, 100,299,314,331,355
mariana, 22,100,240,299,3 14,33 1,355,433
rubens, 299,355,440
sitchensis, \1
Picoides arcticus, 303
pubescens, 303
villosus, 294,303
Pigeon, 57
Pike, Northern, 421,480
Pimephales promelas, 421
Pine, 290
Jack, 299,433
Loblolly, 226
Lodgepole, 100,148,170,314
New Jersey, 351
Pinyon, 32
Ponderosa, 164,176,184
Red, 351
Scots, 320
Western White, 172
White, 20
Pinus sp., 290
banksiana, 299,355,433
contorta, 100,170,314
contorta var. contorta, 148
Vol. 120
edulis, 32
monticola, 172
ponderosa, 164,176,184
resinosa, 351
strobus, 20,355
sylvestris, 298,320,355
taeda, 226
Pipiza femoralis, 17
Piranha, 421
Plagiobothrys scouleri, 158,164
Plantago lanceolata, 154
Plantain, Ribwort, 154
Plathemis lydia, 417
Platycheirus, 19
confusus, 17
obscurus, 17
Platygobio gracilis, 422
Pleco, Royal, 424
Plectritis congesta, 154,158
Pleurozium schreberi, 169,358
Poa annua, 320
bulbosa, 320
compressa, 356
pratensis, 160,356
Podiceps auritus, 109
grisegena, 109
Poecile atricapillus, 294,303
hudsonicas, 303
Poecilia reticulata, 422
Polygala sanguinea, 357
Polygonella articulata, 360
Polygonum sp., 356
aviculare, 164
polygaloides ssp. confertiflorum, 164
Polytrichum commune, 358
Juniperinum, 358
Pomoxis nigromaculatus, 422
Pondlily, Yellow, 240
Popcornflower, Scouler’s, 158,164
Poplar, 433
Balsam, 22
Tulip, 226
Populus, 428,433
balsamifera, 22
deltoides, 226
grandidentata, 356
tremuloides, 22,100,314,343,356
Porpoise, Harbour, 95
Potamogeton pectinatus, 416
Potentilla argentea, 356
canadensis, 356
pacifica, 214
recta, 356
simplex, 356
Poulin, R., Reviews by, 128,253
Powell, T., 339,331
Prairie Dog, 36
Black-tailed, 31
Primrose, Dense-spike Evening, 160
Prince Edward Island, An Observation of the Spring 2006
Migration of Black Scoter Melanitta nigra, in North-
umberland Strait, Interrupted by the Confederation
Bridge, New Brunswick —, 233
Procyon lotor, 57,225,290,342,452
Procyon lotor, Occurrence Through the Use of Microhabitat
Variables, Predicting Raccoon, 225
2007
Pronghorn, 210
Prosartes hookeri var. oregana, \7\
Proulx, G. Winter Habitat Use by American Marten, Martes
americana, in Western Alberta Boreal Forests, 100
Prunus pensylvanica, 50,356
serotina, 356
susquehanae, 360
virginiana, 356
Pseudacris maculata, 88
septentrionalis, 90
triseriata maculata, 88
triseriata septentrionalis, 88
Pseudoroegneria spicata, 176
Pseudotsuga menziesii, 11,148,158,164,169,176,180,184,348
Psilocarphus brevissimus var. brevissimus, 163
brevissimus var. multiflorus, 163
elatior, 163
tenellus var. tenellus, 163
Psilocarphus brevissimus vat. brevissimus, in Canada, Con-
servation Evaluation of the Pacific Population of
Dwarf Woolly-heads, 163
Psoralea, Lance-leaved, 336
Psoralidium lanceolatum, 336
Pteridium acquilinum var. latiusculum, 357
aquilinum, 172
aquilinum var. latiusculum, 356
Pueraria lobata, 226
Puma concolor, 40,438,476
concolor couguar, 438
Puma concolor, Presence in Southern New Brunswick, East-
ern Canada, MtDNA Analyses on Hair Samples Con-
firm Cougar, 438
Pumpkinseed, 422
Pussytoes, Stoloniferous, 183
Pussytoes, Antennaria flagellaris, in Canada, Conservation
Evaluation of Stoloniferous, 183
Pycnopodia helianthoides, 364
Pygocentrus sp., 421
nattereri, 424
Pylodictis olivaris, 424
Quebec and New Brunswick in Late Summer and Autumn,
Use of Saltmarsh by Dragonflies (Odonata) in the
Baie des Chaleurs Region of, 413
Québec, Extension de l’aire de distribution de 17? Umbre de
vase, Umbra limi, dans le nord-est du, 238
Quebec, First Observations of an Eastern Screech-Owl,
Megascops asio, Population in an Apple-Producing
Region of Southern, 289
Quebec, Herpetological Survey 2006 Atlas of Amphibians
and Reptiles of, 258
Quercus spp., 226
garryana, 150,154,157
rubra, 356
Quillwort, Nuttall’s, 160
Quinoa, 340
Quiscalus quiscula, 294
Rabbit, Cottontail, 475
Rabbit-brush, Common, 184
Raccoon, 57,225,290,342,452
Raccoon, Procyon lotor, Occurrence Through the Use of
Microhabitat Variables, Predicting, 225
Rana clamitans, 294
luteiventris, 88
INDEX TO VOLUME 120
WwW
Nm
w
septentrionalis, 239
sylvatica, 90
Rana septentrionalis, in Southeastern Labrador, The Mink
Frog, 239
Rangifer tarandus, 313
tarandus caribou, 313,367
tarandus granti, 205,323
tarandus terraenovae, 5\
Rangifer tarandus granti, Use of Riparian Habitats in Arctic
Alaska, 2001-2003, Assessment of Effects of an Oil
Pipeline on Caribou, 323
Rangifer tarandus, Winter Ranges in Westcentral Alberta,
Pack Size of Wolves, Canis lupus, on Caribou, 313
Ranunculus falcatus, 319
testiculatus, 319
Ranunculus testiculatus, New to Eastern Canada, Bur Butter-
cup, 319
Raphanus raphinistrum, 356
Rat, 478
Brown, 264
Norway, 51,264,475
Rat Poison Kills a Pack of Eastern Coyotes, Canis latrans,
in an Urban Area, 478
Ratfish, Spotted, 363
Ratfish, Hydrolagus colliei, and Scavenger Activity on Teth-
ered Carrion in Subtidal Benthic Communities off
Western Vancouver Island, Pacific Hagfish, Eptatretus
stoutii, Spotted, 363
Rattus norvegicus, 264,475
norvegicus norvegicus, 51
Raven, Common, 109,110
Redcedar, Western, 11,170,348
Redhead, 109
Reed, Common, 307
European Common, 307
Reed, Phragmites australis subspecies australis, in the South-
ern Ottawa District, Recent Invasion, Current Status
and Invasion Pathway of European Common, 307
Reimchen, T.E., 61,363
Rhingia nasica, 17
Rhizobium, 154,159
Rhodeus sp., 424
Rhododendron albiflorum, 169
californicum, 169
canadense, 356
lapponicum, 169
macrophyllum, 169
Rhododendron, Pacific, 169
White-flowered, 169
Rhododendron macrophyllum, in Canada, Conservation Eval-
uation of Pacific Rhododendron, 169
Rhododendron, Rhododendron macrophyllum, in Canada,
Conservation Evaluation of Pacific, 169
Rhytidiadelphus loreus, 171
Rice-grass, Indian, 336
Richards, N.L., P. Mineau, D.M. Bird, P. Wery, J. Larivée, and
J. Duffe. First Observations of an Eastern Screech-
Owl, Megascops asio, Population in an Apple-Pro-
ducing Region of Southern Quebec, 289
Richardson, J.S., 347
Richardsonius balteatus, 421
Ringlet, Maritime, +19
Roberts, J., 421
Robin, American, 294
Rocksole, 364
524
Ronconi, R.A. Predicting Bird Oiling Events at Oil Sands
Tailings Ponds and Assessing the Importance of Alter-
nate Waterbodies for Waterfowl: a Preliminary Assess-
ment, |
Ronquil, Northern, 364
Ronquilus jordani, 364
Rosa carolina, 353
nutkana, 158
virginiana, 353
Rosatte, R.C. and J. Neuhold. Late-born Elk, Cervus elaphus,
Calf Observed Near Bancroft, Ontario, 188
Rose, Nootka, 158
Rosebay, Lapland, 169
Rubus sp., 236
allegheniensis, 356
arenicola, 353
hispidus, 353
idaeus vat. strigosus, 18
particeps, 359
parviflorus var. parviflorus, 171
spectabilis, 11,348
vermontanus, 356
Rudbeckia hirta, 356
Rumex acetosella, 356
Runck, A.M., 331
Ruppia maritima, 414
Russian-thistle, 337
Ryan, M., 147,153
Rye, 343
Canada Wild, 336
Sagebrush, Big, 164,184
Hoary, 336
Salal, 11,172
Salamander, Long-toed, 90
Northwestern, 90
Salix sp., 158,314
exigua ssp. interior, 336
humilis, 356
Salmo gairdneri, 199
salar, 199,413,422
trutta, 199,422
trutta trutta, 199
Salmon, Atlantic, 199,413,422
Chinook, 422
Chum, 199,422
Pacific, 421
Sockeye, 422
Wild Pink, 199
Salmon, Response of the Sea Louse Lepeophtheirus salmonis
Infestation Levels on Juvenile Wild Pink, Oncorhyn-
chus gorbuscha, and Chum, O. keta, Salmon to Arrival
of Parasitized Wild Adult Pink, 199
Salmonberry, 11,348
Salmonella, 46
Salsola tragus, 337
Salvelinus alpinus, 199,235,422
confluentus, 422
fontinalis, 238,422
malma, 422
namaycush, 422,481
Sander vitreus, 421
Sandlance, 96
Sandwort, Thread-leaved, 184
Sapin baumier, 238
THE CANADIAN FIELD-NATURALIST
Vol. 120
Sapsucker, Yellow-bellied, 302
Sarotherodon sp., 424
Saskatchewan and the First Saskatchewan Records of Ame-
lanchier sanguinea, Regional Variation in Amelanchi-
er in the Whitewood area of Southeastern, 428
Saskatchewan Records of Amelanchier sanguinea, Regional
Variation in Amelanchier in the Whitewood area of
Southeastern Saskatchewan and the First, 428
Saskatchewan, Summer Movements and Impact of Individual
Striped Skunks, Mephitis mephitis, on Duck Nests in,
342
Saskatoon, 176
Sayornis phoebe, 294
Scaup, 6
Greater, 109
Lesser, 5
Schaefer, R.L., 192
Schmiegelow, F.K.A., 313
Scirpus sp., 214,295,417
americanus, 213
maritimus, 214
paludosus, 215
Sciurus carolinensis, 10,264
vulgaris, 10
Sciurus carolinensis, on Douglas’s Squirrels, Tamiasciurus
douglasii, in Vancouver, British Columbia, A Test of
Interspecific Effects of Introduced Eastern Grey Squir-
rels, 10
Scorpion-grass, Blue, 320
Scoter, 232
Black, 233
Scoter Melanitta nigra, in Northumberland Strait, Interrupt-
ed by the Confederation Bridge, New Brunswick —
Prince Edward Island, An Observation of the Spring
2006 Migration of Black, 233
Scoter, Melanitta spp., Migrations Interrupted by Confedera-
tion Bridge: An Update, 232
Screech-Owl, Eastern, 289
Screech-Owl, Megascops asio, Population in an Apple-Pro-
ducing Region of Southern Quebec, First Observa-
tions of an Eastern, 289
Sculpin, Deepwater, 480
Prickly, 425
Roughback, 364
Sculpin, Myoxocephalus thompsonii, in Northeastern Alberta,
A New Record of Deepwater, 480
Seal, 466
Harbour, 62
Ringed, 464
Seburn, D., Review by, 248
Sedge, Awl-fruited, 22
Few-seeded, 22
Foothill, 160
Green-sheathed, 160
Lyngbei, 213
Sedum acre, 356
spathifolium, 154
Selatosomus pulcher, 19
Seminatrix pygaea, 477
Sericomyia chrysotoxoides, 17
Serrasalmus sp., 421
Shad, American, 422
Sheep, Bighorn, 314
Dall, 205
Shepherdia canadensis, 172
2007
Shiner, Redside, 421
Spottail, 422
Shoveler, Northern, 5
Shrew, Dusky, 332
Masked, 50,332
Northern Short-tailed, 436
Pygmy, 332
Short-tailed, 294
Shrimp, Spot, 364
Silene antirhina, 356
Simard, P., J.L. Lawlor, and S. Gowans. Temporal Variability
of Cetaceans near Halifax, Nova Scotia, 93
Sisyrinchium arenicola, 359
fuscatum, 357
Sitta sp., 290
canadensis, 303
carolinensis, 303
Skeleton-weed, 336
Skimmer, Four-spotted, 417
Twelve-spotted, 417
Widow, 417
Skunk, Santa Cruz Island Spotted, 210
Striped, 229,342,452
Skunks, Mephitis mephitis, on Duck Nests in Saskatchewan,
Summer Movements and Impact of Individual Striped,
342
Slough, B.G., 39,331
Slough, B.G. and R.L. Mennell. Diversity and Range of
Amphibians of the Yukon Territory, 87
Snake, Black Swamp, 477
Maritime Garter, 477
Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island,
Nova Scotia, Bald Eagle, Haliaeetus leucocephalus,
Preying on Maritime Garter, 477
Snow, D.P., 50
Solidago spp., 226
bicolor, 356
canadensis, 356
graminifolia, 356
Juncea, 356
nemoralis, 356
puberula, 356
Somateria mollissima, 233,236
Somateria mollissima, Duckling, Osprey, Pandion haliaetus,
Depredates Common Eider, 236
Somatochlora elongata, 415
kennedyi, 417
Soopolallie, 172
Sorex sp., 294
cinereus, 50,332
cinereus acadicus, 51
hoyi, 332
monticolus, 332
Soybean, 226
Sparganium angustifolium, 240
Sparrow, House, 43,264
Le Conte’s, 22
Sparrow, Ammospiza leconteii, from Northeastern Ontario,
with Some Notes on Nesting Behaviour, New Nest-
ing Records of the Le Conte’s, 22
Sparrows, Passer domesticus, in Canada’s Maritime Provinces,
Recent Declines of House, 43
Spartina, 414
alterniflora, 414
patens, 414
INDEX TO VOLUME 120
pectinata, 356,416
Speedwell, American, 158
Spring, 320
Spermophilus spp., 31
Sphaerophoria novaeangliae, \7
Sphagnum sp., 172,440
Sphegina brachygaster, \7
campanulata, \7
flavomaculata, \7
keeniana, \7
Sphyraena barracuda, 24}
borealis, 241
guachancho, 241
Sphyraena barracuda from Canada, First Record of the Great
Barracuda, 24]
Sphyrapicus varius, 303
Spilogale gracilis amphiala, 210
Spiraea alba, 356
Spiranthes lacera, 356
Spray, Ocean, 158
Spreadwing, Elegant, 417
Lyre-tipped, 417
Northern, 417
Slender, 417
Spotted, 417
Sweetflag, 417
Spruce, 20,110,236,431
Black, 22,100,240,299,314,331,433
Englemann, 100
Norway, 299
Red, 299,440
Sitka, 11
White, 100,299,314,331
Spruce-fir, 331
Spurge, Leafy, 337
Squirrel, 295
Douglas’s, 10
Eastern Grey, 10
European Red, 10
Gray, 264
Ground, 31
Red, 10,51,57
Squirrels, Sciurus carolinensis, on Douglas’s Squirrels, Tami-
asciurus douglasii, in Vancouver, British Columbia,
A Test of Interspecific Effects of Introduced Eastern
Grey, 10
Squirrels, Tamiasciurus douglasii, in Vancouver, British
Columbia, A Test of Interspecific Effects of Intro-
duced Eastern Grey Squirrels, Sciurus carolinensis,
on Douglas’s, 10
Star, Long-rayed, 364
Sunflower, 364
Starling, 264
European, 109,290
Steinhilber, M. and D.A. Neely. A New Record of Deepwater
Sculpin, Myoxocephalus thompsonii, in Northeastern
Alberta, 480
Stellaria graminea, 356
pallida, 320
Stickleback, 421
Brook, 422
Threespine, 240,422
Stonecrop, Lance-leaved, 154
Strauss, E.G., 263,478
Strawberry, 360
526
Streever, B., 323
Strix varia, 290,303
Sturgeon, Lake, 71
White, 422
Sturgeon, Acipenser fulvescens, Movements in Rainy Lake,
Minnesota and Ontario, Lake, 71
Sturnus vulgaris, 109,264,290
Stylasterias forriei, 364
Sucker, Longnose, 422,481
White, 422
Sunflower, Prairie, 336
Surnia ulula, 433
Surnia ulula, near Timmins and Iroquois Falls, Northeastern
Ontario, in 2001, Observations on the Nesting of the
Northern Hawk Owl, 433
Swallow, Tree, 302
Swan, Trumpeter, 220
Sweet-clover, 337
Swordtail, 422
Sylvilagus spp., 165
floridanus, 475
nuttallii, 165
Sympetrum sp., 416
costiferum, 415
danae, 413
internum, 415
obtrusum, 415
semicinctum, 417
vicinum, 415
Symphyotrichum laterifloum, 356
Synhalonia, 180
Syrphus rectus, 17
Szumylo, D.-L.M., 263
Tachycineta bicolor, 303
Tamarack, 314
Tamias striatus, 264,295
striatus lysteri, 51
Tamiasciurius sp., 295
douglasii, 10
hudsonicus, 10,57
hudsonicus ungavensis, 51
Tamiasciurus douglasii, in Vancouver, British Columbia, A
Test of Interspecific Effects of Introduced Eastern
Grey Squirrels, Sciurus carolinensis, on Douglas’s
Squirrels, 10
Tanichthys albonubes, 424
Taraxacum officinale, 320,358
Taricha granulosa, 90
Taxidea taxus, 31,209,342
Tea-berry, Western, 171
Tea, Labrador, 172
Tegler, B., 496
Tegler, B., Review by, 252
Temnostoma balyras, 17
Tench, 422
Thamnophis sirtalis pallidulus, 477
Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova
Scotia, Bald Eagle, Haliaeetus leucocephalus, Prey-
ing on Maritime Garter Snake, 477
Thiel, R.P. Conditions for Sexual Interactions Between Wild
Grey Wolves, Canis lupus, and Coyotes, Canis latrans,
27
Thimbleberry, 171
Thrasher, Brown, 294
THE CANADIAN FIELD-NATURALIST
Vol. 120
Thuja occidentalis, 300
plicata, 11,170,348
Thymallus arcticus, 422
Tilapia sp., 422
Tilapia, 422
Tinca tinca, 422
Toad, American, 240
Western, 87
Toadflax, Dalmation, 176
Tokaryk, T., Reviews by, 127,485
Tonella tenella, 137,179
Tonella, Small-flowered, 137,179
Tonella tenella, in Canada, Conservation Evaluation of the
Small-flowered Tonella, 179
Tonella, Tonella tenella, in Canada, Conservation Evaluation
of the Small-flowered, 179
Torilis japonica, 180
Toxomerus geminatus, 17
Toxostoma rufum, 294
Tramea spp., 418
Trefoil, Bird’s-foot, 157
Bog Bird’s-foot, 153,157
Seaside Bird’s-foot, 153,157
Trefoil, Lotus formosissimus, in Canada, Conservation Eval-
uation of the Seaside Birds-foot, 153
Trefoil, Lotus pinnatus, in Canada, Conservation Evaluation
of the Bog Bird’s-foot, 157
Tremblay, E., 438
Trientalis borealis, 356
Trifolium arvense, 357
procumbens, 357
Triglochin maritimum, 214
Tringa flavipes, 5
Triteleia hyacinthina, 158
Triteleia, White, 158
Troglodytes troglodytes, 303
Trout, Brook, 422
Brown, 422
California Golden, 422
Lake, 481
Rainbow, 90,199,421
Sea, 199
Troutperch, 422
Tsuga canadensis, 290
heterophylla, 11,170,348
Turdus migratorius, 294
Tursiops truncatus, 97
Twayblade, Heart-leaved, 170
Twinflower, 172
Typha latifolia, 214
Umbra limi, 238
Umbra limi, dans le nord-est du Québec, Extension de |’ aire
de distribution de 1’ Umbre de vase, 238
Urocyon littoralis, 210
Urosalpinx cinera, 106
Ursus americanus, 57,68,110,209,314
americanus hamiltoni, 51
arctos, 57,314
Ursus americanus, Retrieval of an Elk, Cervus elaphus, Car-
cass from a Small Lake in Riding Mountain National
Park, Manitoba, Probable Black Bear, 110
Ursus americanus L., Long-range Homing by an Adult Female
Black Bear, 57
2007
Vaccinium alaskense, 172
angustifolium, 357
angustifolium X corymbosum, 357
myrtilloides, 357
ovatum, 348
parvifolium, 11,172
scoparium, 172
Vander Kloet, S.P., 351
Varden, Dolly, 422
Vasseur, L., 438
Velvet-grass, Common, 154
Vernalgrass, Sweet, 148,154,160
Veronica beccabunga ssp. americana, \58
officinalis, 357
verna, 320
Vetch, Common, 180
Viburnum nudum var. cassinoides, 357
Vicia sp., 157
cracca, 357
sativa, 180
tetrasperma, 357
Viola sagittata var. ovata f. glabrata, 360
Sagittata var. ovata, 357
Violette, V., 236
Vole, Eastern Heather, 332
Meadow, 50,264,294,332,436
Northern Red-backed, 331
Southern Red-backed, 50,331
Vole, Clethrionomys gapperi, in Newfoundland: Implications
for the Endangered Newfoundland Marten, Martes
americana atrata, First Record of the Southern Red-
Backed, 50
Vole, Myodes gapperi, in the Yukon, First Records of the
Southern Red-backed, 331
Volucella bombylans, 17
Vors, L.S. and P.L. Wilson. A New Record Size Wolf, Canis
lupus, Group for Ontario, 367
Vulpes fulva, 468
macrotis, 52
macrotis mutica, 210
velox, 31
vulpes, 36,68,263,342
vulpes deletrix, 51
Vulpes velox, in Southeastern Colorado, Multiple Scale Den
Site Selection by Swift Foxes, 31
Walley, W.J. Probable Black Bear, Ursus americanus, Retrieval
of an Elk, Cervus elaphus, Carcass from a Small Lake
in Riding Mountain National Park, Manitoba, 110
Walley, W.J., Ring-billed Gull, Larus delawerensis, Food
Piracy on Diving Ducks, 109
Walleye, 421
Walton, L.R., 342
Wasp, 295
Waxwing, Cedar, 294
Way, J.G. and D.L. Eatough. Use of “Micro’-Corridors by
Eastern Coyotes, Canis latrans, in a Heavily Urban-
ized Area: Implications for Ecosystem Management,
474
Way, J.G., D.-L.M. Szumylo, and E.G. Strauss. An Ethogram
Developed on Captive Eastern Coyotes Canis latrans,
263
Way, J.G., S.M. Cifuni, D.L. Eatough, and E.G. Strauss. Rat
Poison Kills a Pack of Eastern Coyotes, Canis latrans,
in an Urban Area, 478
INDEX TO VOLUME 120
527
Weasel, Short-tailed, 51,62
Weatherloach, 424
Wery, P., 289
Whale, Blue, 95
Bowhead, 462
Fin, 95
Humpback, 95
Killer, 457
Long-finned Pilot, 97
Minke, 95
North Atlantic Right, 95
Pilot, 97
Pygmy Sperm, 97
Sei, 97
Whale, Orcinus orca, Attacks in the Eastern Canadian Arctic,
Reactions of Narwhals, Monodon monoceros, to Kil-
ler, 457
Wheat, 343
Wheat-grass, Bluebunch, 176
Crested, 337
Northern, 336
Slender, 184
Whiteface, Frosted, 417
Whitefish, Lake, 422,480
Whitetail, Common, 417
Whitlaw, H.A., 205
Whitlow-grass, Spring, 320
Wildlife Afield, 132,502
Williams, R., 199
Willis, D.W., 71
Willow, 158,314
Sandbar, 336
Wiison, P.L., 367
Windgrass, Interrupted, 320
Wissink, R., 438
Wolf, 27,57,67,263,313,367,466,47 1,476,478
Arctic, 466,471
Eastern, 27
Eastern Timber, 51
Gray [Grey], 27,61,111,205,367
Newfoundland, 50
Red, 27
Timber, 27
Wolf, Canis lupus, Group for Ontario, A New Record Size,
367
Wolverine, 111
Wolves, Canis lupus arctos: Influence of Order on Posture,
Double Marking in Arctic, 471
Wolves, Canis lupus, and Coyotes, Canis latrans, Conditions
for Sexual Interactions Between Wild Grey, 27
Wolves, Canis lupus arctos, in Summer, Urine-marking and
Ground-scratching by Free-ranging, 466
Wolves, Canis lupus, in Southcentral Alaska, Physical Char-
acteristics, Hematology, and Serum Chemistry of Free-
ranging Gray, 205
Wolves, Canis lupus, of Coastal British Columbia, Early Onto-
genetic Diet in Gray, 61
Wolves, Canis lupus, on Caribou, Rangifer tarandus, Winter
Ranges in Westcentral Alberta, Pack Size of, 313
Woodley, S.J., G. Johnson, B. Freedman, and D.A. Kirk.
Effects of Timber Harvesting and Plantation Devel-
opment on Cavity-nesting Birds in New Brunswick,
298
Woodpecker, Black-backed, 302
Downy, 302
528
Hairy, 294,302
Pileated, 299,302
Woolly-heads, Dwarf, 163
Woolly-heads, Psilocarphus brevissimus var. brevissimus, in
Canada, Conservation Evaluation of the Pacific Popu-
lation of Dwarf, 163
Wren, Winter, 302
Xiphophorus helleri, 422
Xylota, 20
confusa, 17
quadrimaculata, 17
Index to Book Reviews
Botany
Burger, W.C. Flowers How They Changed the World, 249
Carder, A. Giant Trees of Western America and the World, 120
Rover, F. and R. Dickinson. Plants of Alberta, 386
Scotter, G. and H. Flygare. Wildflowers of the Rocky Moun-
tains, 386
Viereck, L. and E.L. Little. Alaska Trees and Shrubs. Second
Edition, 491
Environment
Dow, K. and T.E. Downing. The Atlas of Climate Change:
Mapping the World’s Greatest Challenge, 493
Fish and Wildlife Historical Society. Fish, Fur and Feathers:
Fish and Wildlife Conservation in Alberta, 1905-2005,
124
Gore, A. An Inconvenient Truth: The Planetary Emergency of
Global Warming and What We Can Do About It, 494
Kruse, G.H., V.F. Gallucci, D.E. Hay, R.I. Perry, R.M.
Peterman, T.C. Shirley, P.D. Spencer, B. Wilson, and,
D. Woodby. Fisheries Assessment and Management
in Data-Limited Situations, 121
McGavin, G.C. Endangered: Wildlife on the Brink of Extinc-
tion, 252
Mundy, P. The Gulf of Alaska: Biology and Oceanography,
125
Redcliff, D. Lapland A Natural Historym 123
Saab, V.A. and H.D.W. Powell. Fire and Avian Ecology in
North America Studies in Avian Biology Number
30, 121
Warren, A. and T. Allan. Guide to Deserts, 251
Miscellaneous
Burt, W. Marshes: The Disappearing Edens, 389
Callan, K. A Paddler’s Guide to Quetico and Beyond, 496
Czajkowski, C. Wildfire in the Wilderness, 498
Elder, J. Pilgrimage to Vallombrosa: From Vermont to Italy in
the Footsteps of George Perkins Marsh, 387
Hudson, R. A Field Guide to Gold, Gemstones & Mineral
Sites of British Columbia, Sites Within a Day’s Drive
of Vancouver, 128
Hudson, R. A Field Guide to Gold, Gemstones & Mineral
Sites of British Columbia, Sites Within a Day’s Drive
of Vancouver, 253
Koeppel, D. To See Every Bird on Earth, 498
Mabey, R. Gilbert White: A Biography of the Author of The
Natural History of Selborne, 496
Mayor, A. Fossil Legends of the First Americans, 127
Milne, P. Where to Watch Birds World Cities, 388
Mock. D.W. More than Kin and Less than Kind: The Evolu-
tion of Family Conflict, 497
THE CANADIAN FIELD-NATURALIST
Vol. 120
Yellow-cedar, 172
Yellowlegs, Lesser, 5
Yukon, First Records of the Southern Red-backed Vole, Myo-
des gapperi, in the, 331
Yukon Territory, Diversity and Range of Amphibians of the,
87
Yukon Territory, First Records of the Northern Long-eared
Bat, Myotis septentrionalis, in the, 39
Zannichellia palustris, 416
Zapus hudsonius, 332
Zea mays, 226
Zenaida macroura, 294
Quammen, D. The Reluctant Mr. Darwin: An Intimate Portrait
of Charles Darwin and the Making of His Theory of
Evolution, 252
Shtilmark, F. History of the Russian Zapovedniks 1985-1995,
254
Van der Flier-Keller, E. A Field Guide to the Identification
of Pebbles, 255
Zoology
Acorn, J. Deep Alberta. Fossil Facts and Dinosaur Digs, 485
Acorn, J. Ladybugs of Alberta: Finding the Spots and Con-
necting the Dots, 486
Adler, P., D. Currie, and D.M. Wood. The Black Flies (Simu-
liidae) of North America, 245
Arlott, N. Birds of Europe, Russia, China and Japan: Passer-
ines, Tyrant Flycatchers to Buntings, 484
Beadle, D. and J. Rising. Tanagers, Cardinals, and Finches of
the United States and Canada, 119
Bildstein, K.L. Migrating Raptors of the World: Their Ecology
and Conservation, 382
Bonin, F., B. Devaux and A. Dupre. Turtles of the World, 248
Chu, M. Songbird Journeys: Four Seasons in the Lives of
Migratory Birds, 385
Clements, J.F. The Clements Checklist of Birds of the World
6th Edition, 483
de Asua, M. and R. French. A New World of Animals: Early
Modern Europeans on the Creatures of Iberian
America, 487
del Hoyo, J., A. Elliott and D.A. Christie. Handbook of the
Birds of the World Volume IT Old World Flycatchers
to Old World Warbler, 381
Eisner, T., M. Eisner, M. Siegler. Secret Weapons Defenses of
Insects, Spiders, Scorpions, and Other Many-Legged
Creatures, 118
Elbroch, M. Animal Skulls, A Guide to North American Spe-
cies, 243
Elphick, J. Atlas of Bird Migration: Tracing the Great Jour-
neys of the World’s Birds, 380
Evans, H.E. and K.M. O’Neill. The Sand Wasps: Natural
History and Behavior, 489
Firouz, E. The Complete Fauna of Iran, 114
Forsyth, R.G. Land Snails of British Columbia, 383
Gibson, M. Gibson’s Guide to Bird Watching & Conservation,
485
Hoogland, J. Conservation of the Black-tailed Prairie Dog:
Saving North America’s Western Grasslands, 382
Jackson, D.C. Tracks, 490
Latta, S., C. Rimmer, A. Keith, J. Wiley, H. Raffaele, K.
McFarland, E. Fernandez. Birds of the Dominican
Republic and Haiti, 379
2007
Lynch, W. Penguins of the World, 488
Marshall, S.A. Insects: Their Natural History and Diversity:
With a photographic guide to insects of eastern North
America, 117
Marshall, S.A. Insects: Their Natural History and Diversity:
With a Photographic Guide to Insects of Eastern North
America, 246
Maybank, B. The Birding Sites of Nova Scotia, 114
Morrison, M.L. The Northern Goshawk: A technical Assess-
ment of its Status, Ecology and management, 247
Moss, S. The Birder’s Companion, 379
Nagorsen, D.W. Rodents and Lagomorphs of British Col-
umbia, 118
Nelson, J.S. Fishes of the World, 116
O’Connor, M. Why Don’t Woodpeckers Get Headaches? And
Other Bird Questions You Know You Want To Ask, 490
INDEX TO VOLUME 120
529
Panov, E.N. Wheatears of Palaearctic: Ecology, Behaviour
and Evolution of the Genus Oenanthe, 119
Proctor, N. and P. Lynch. A Field Guide to North Adantic
Wildlife, 244
Sale, R. A Complete Guide to Arctic Wildlife, 244
Stevenson, T. and J. Fanshawe. The Birds of East Africa:
Kenya, Tanzania, Uganda, Rwanda, Burundi, 113
Theberge, J.B. and M.T. Theberge. The Wolves of Algonquin
Park: A 12-year Ecological Study, 249
Wheeler, B.K. Raptors of Western North America and Raptors
of Eastern North America, 489
Young Naturalists
de Vries, M. Tale of a Great White Fish, 499
530
THE CANADIAN FIELD-NATURALIST
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TABLE OF CONTENTS (concluded) Volume 120 Number 4
300k Reviews
ZOOLOGY: The Clements Checklist of Birds of the World 6th Edition — Birds of Europe, Russia, China
and Japan: Passerines. Tyrant Flycatchers to Buntings — Deep Alberta: Fossil Facts and Dinosaur Digs
— Gibson’s Guide to Bird Watching & Conservation — Ladybugs of Alberta: Finding the Spots and
Connecting the Dots — A New World of Animals: Early Modern Europeans on the Creatures of Iberian
America — Penguins of the World — Raptors of Western North America and Raptors of Eastern North
| America — The Sand Wasps: Natural History and Behavior — Tracks — Why Don’t Woodpeckers Get
Headaches? And Other Bird Questions You Want to Ask
B30TANY: Alaska Trees and Shrubs Second edition
INVIRONMENT: The Atlas of Climate Change: Mapping the World’s Greatest Challenge — An Inconvenient
Truth: The Planetary Emergency of Global Warming and What We Can Do About It
ISCELLANEOUS: Gilbert White: A Biography of the Author of The Natural History of Selborne — A
Paddler’s Guide to Quetico and Beyond — More than Kin and Less than Kind: Evolution of Family Conflict
| — To See Every Bird on Earth — Wildfire in the Wilderness
‘OUNG NATURALISTS: Tale of a Great White Fish
arine Turtle Newsletter (114) — Canadian Association of Herpetologists/Association Canadienne des
Herpetologists Bulletin Fall 2006 — Wildlife Afield 3 (1 + Supplement to 1) — Errata The Canadian-Field
Naturalist 120(2) and 120(3)
| ttawa Field-Naturalists’ Club Annual Awards for 2005
‘dex to The Canadian Field-Naturalist Volume 120 Compiled by LEsLte Copy
| dvice to Contributors
i
{ailing date of the previous issue 120(3): 30 November 2007
2006
483
49]
493
497
499
500
502
530
THE CANADIAN FIELD-NATURALIST Volume 120 Number 4
Articles
A reevaluation of sexual dimorphism in the postcranium of the chasmosaurine ceratopsid
Chasmosaurus belli (Dinosauria: Ornithischia) | JORDAN C. MALLON and ROBERT B. HOLMES
Use of saltmarsh by dragonflies (Odonata) in the Baie des Chaleurs region of Quebec
and New Brunswick in late summer and autumn
PAUL M. CATLING, RAYMOND HUTCHINSON, and PAUL M. BRUNELLE
First records of the Yellow Bullhead, Ameiurus natalis, a loricariid catfish, Panaque suttonorum,
and a Silver Pacu, Piaractus cf P. brachypomus. in British Columbia
G. F. HANKE, M. C. E. MCNALL, and J. ROBERTS
Regional variation in Amelanchier in the Whitewood area of southeastern Saskatchewan and
the first Saskatchewan records of Amelanchier sanguinea PAUL CATLING and G. MITROW
Observations on the nesting of Northern Hawk Owl, Surnia ulula, near Timmins and Iroquois
Falls, northeastern Ontario, in 2001 MICHAEL PATRIKEEV
MtDNA analyses on hair samples confirm Cougar, Puma concolor, presence in southern
New Brunswick, eastern Canada
A.-S. BERTRAND, S. KENN, D. GALLANT, E. TREMBLAY, L. VASSEUR, and R. WISSINK
The leech Haemopis lateromaculata (Hirudinea; Haemopidae): Its North American distribution
and additional notes on species description PETER HOVINGH
Predation on artificial nests of Northern Bobwhites, Colinus virginianus, by mammalian
mesopredators: Does the problem-individual paradigm fit?
J. B. JENNINGS, M. L. KENNEDY, A. E. HOUSTON, and B. D. CARVER
Reactions of Narwhals, Monodon monocerus, to Killer Whale, Orcinus orca, attacks in eastern
Canadian Arctic KRISTIN L. LAIDRE, MADS PETER HEIDE-JORGENSEN, and JACK R. ORR
Urine-marking and ground-scratching by free-ranging Arctic Wolves, Canis lupus arctos,
in summer L. DAvID MECH
Notes
Double marking in Arctic Wolves, Canis lupus arctos: Influence of order on posture
FRED H. HARRINGTON
Use of “micro”-corridors by Eastern Coyotes, Canis latrans, in a heavily urbanized area:
Implications for ecosystem management JONATHAN G. Way and Davib L. EATOUGH
Bald Eagle, Haliaeetus leucocephalus, preying on a Maritime Garter Snake,
Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova Scotia STORRS L. OLSON
Rat poison kills a pack of eastern Coyotes, Canis latrans, in an urban area
JONATHAN G. Way, STEPHEN M. CIFUNI, DAviD L. EATOUGH, and Eric G. STRAUSS
A new record of Deepwater Sculpin, Myoxocephalus thompsonii, in northeastern Alberta
M. STEINHILBER and D. A. NEELY
(continued on inside back cover)
ISSN 0008-3550
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2006
LAN
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The CANADIAN
FIELD-NATURALIST
| Published by THE OTTAWA FIELD-NATURALISTS” CLUB, Ottawa, Canada
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Volume 121, Number 1 January—March 2007
The Ottawa Field-Naturalists’ Club
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Cover: “When Blackfeet and Sioux Meet” 1908 oil painting by Charles M. Russell (1864-1926). Courtesy of Sid Richardson
Museum where the original is housed. Born in St. Louis, Missouri, Charles M. Russell went west to Montana as a boy of
16 with a dream of becoming a real cowboy. Beginning in 1882, Charlie worked on various cattle drives wrangling horses
where he quickly established a reputation as a likeable cowboy who loved to draw and paint. During his 11 years on the
range, he witnessed the changing of the West as the days of free grass and unfenced range were ending. Charlie quit the
cowboy way of life in 1893 and married Nancy Cooper three years later. With Nancy’s keen business sense and belief in
her husband’s talent, Russell’s paintings began to be known nationally. Charlie Russell felt deeply the passing of the
West, the most evident theme of his art. The old ways of Indian and cowboy life had disappeared. His works reflected
the public’s demand for authentic pictures of the West, yet the soul of his art was romance. (Jan Scott, Director, Sid
Richardson Museum, 309 Main Street, Fort Worth, Texas 76102 USA; e-mail: jan @sidricharsonmuseum.org) See article
by Charles M. Kay “Were native people keystone predators: A continuous-time analysis of wildlife observations made by
Lewis and Clarke in 1804-1806” pages 1-16.
MCz
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FIELD-NATURALIST
Volume 121
2007
Volume 123
The Ottawa Field-Naturalists’ Club Transactions
Promoting the study and conservation
of northern biodiversity since 1880
THE OTTAWA FIELD-NATURALISTS’* CLUB
OTTAWA CANADA
The Canadian Field-Naturalist
January—March 2007
Volume 121, Number |
Were Native People Keystone Predators? A Continuous-Time Analysis
of Wildlife Observations Made by Lewis and Clark in 1804-1806
CHARLES E. KAY
Department of Political Science, Utah State University, Logan, Utah 84322-0725 USA
Kay, Charles E. 2007. Were native people keystone predators? A continuous-time analysis of wildlife observations made by
Lewis and Clark in 1804-1806. Canadian Field-Naturalist 121(1): 1-16.
It has long been claimed that native people were conservationists who had little or no impact on wildlife populations. More
recently, though, it has been suggested that native people were keystone predators, who lacked any effective conservation
strategies and instead routinely overexploited large mammal populations. To test these hypotheses, I performed a continuous-
time analysis of wildlife observations made by Lewis and Clark because their journals are often cited as an example of how
western North America teemed with wildlife before that area was despoiled by advancing European civilization. This
included Bison, Elk, Mule Deer, Whitetailed Deer, Blacktailed Deer, Moose, Pronghorn Antelope, Bighorn Sheep, Grizzly
Bears, Black Bears, and Grey Wolves. I also recorded all occasions on which Lewis and Clark met native peoples. Those
data show a strong inverse relationship between native people and wildlife. The only places Lewis and Clark reported an
abundance of game were in aboriginal buffer zones between tribes at war, but even there, wildlife populations were predator,
not food-limited. Bison, Grizzly Bears, Bighorn Sheep, Mule Deer, and Grey Wolves were seldom seen except in aboriginal
buffer zones. Moose were most susceptible to aboriginal hunting followed by Bison and then Elk, while Whitetailed Deer had
amore effective escape strategy. If it had not been for aboriginal buffer zones, Lewis and Clark would have found little wildlife
anywhere in the West. Moreover, prior to the 1780 smallpox and other earlier epidemics that decimated native populations
in advance of European contact, there were more aboriginal people and even less wildlife. The patterns observed by Lewis
and Clark are consistent with optimal foraging theory and other evolutionary ecology predictions.
Key Words: Native hunting, aboriginal buffer zones, Lewis and Clark, keystone predation, reference conditions, Elk, Bison,
Grizzly Bears, Mule Deer, Whitetailed Deer, western North America.
It has long been postulated that native people were
conservationists who had little or no impact on wildlife
populations (e.g.; Speck 1913, 1939a, 1939b). Studies
of modern hunter-gatherers, however, have found little
evidence that native people purposefully employ con-
servation strategies (Alvard 1993, 1994, 1995, 1998a,
1998b; Hill and Hurtado 1996), while archaeological
data suggest that prehistoric people routinely overex-
ploited large-mammal populations (Broughton 1994a,
1994b, 1997; Jones and Hilderbrant 1995: Janetski
1997; Butler 2000; Chatters 2004). Elsewhere, I have
proposed that native people were keystone predators,
who once structured entire ecosystems (Kay 1994,
1995, 1997a, 1997b, 1998, 2002).
To test these competing hypotheses, I performed a
continuous-time analysis of wildlife observations made
by Lewis and Clark on their expedition across North
America in 1804-1806 because their journals are often
cited as an example of how the West teemed with wild-
life before that area was despoiled by advancing Euro-
pean civilization (Botkin 1995, 2004; Patten 1998:
70; Wilkinson and Rauber 2002; Nie 2003: 1). Lewis
and Clark were the first Europeans to traverse what
eventually became the western United States, and many
of the native peoples they met had never before en-
countered Europeans. In addition, historians universal-
ly agree that Lewis and Clark’s journals are not only
among the earliest, but also the most detailed and ac-
curate, especially regarding natural history observations
(Burroughs 1961; Ronda 1984; Botkin 1995, 2004).
Thus, the descriptions left by Lewis and Clark are
thought by many to represent the “pristine” state of
western ecosystems (Craighead 1998: 597; Patten 1998:
70; Wilkinson and Rauber 2002; Botkin 2004). Botkin
(1995: 1), for instance, described Lewis and Clark’s
journey as “the greatest wilderness trip ever recorded.”
Methods and Study Area
Recently, Martin and Szuter (1999a, 1999b, 2002,
2004), Lyman and Wolverton (2002), and Laliberte and
Ripple (2003) presented contrasting interpretations
of western ecosystems based on Lewis and Clark’s jour-
nals, but those analyses are flawed, in part, because they
either did not separate ungulates by species or they did
DD THE CANADIAN FIELD-NATURALIST
not correlate wildlife sightings with the abundance of
native people on each day of the entire trip. Instead, I
developed three measures to quantify the wildlife ob-
servations recorded by Lewis and Clark in their orig-
inal journals, which have recently been re-edited and
republished (Moulton 1986, 1987a, 1987b, 1988, 1990,
1991, 1993 — hereafter cited only by volume and page).
First, game seen. If Lewis and Clark reported old sign
of a species, that was assigned a value of one, fresh sign
a two, and if they actually saw the animal, a three. This
included Bison (Bison bison), Elk (Cervus elaphus),
Whitetailed Deer (Odocoileus virginianus), Mule Deer
(O. hemionus hemionus), Blacktailed Deer (O. h. co-
lumbianus), Moose (Alces alces), Pronghorn Antelope
(Antilocapra americana), Bighorn Sheep (Ovis can-
adensis), Grizzly Bears (Ursus arctos), Black Bears
(U. americanus), and Grey Wolves (Canis lupus). This
was done each day for the entire 863 days of the expe-
dition.
Second, game killed. On each day, Lewis and Clark
recorded the exact number of animals that were killed
to provision their party. In three instances, though,
Lewis and Clark reported that “some” Whitetailed
Deer (day 78), Elk (day 365), or Bison (day 413) were
killed. In these cases, “some” was recorded as three
animals killed. In 12 instances, Lewis and Clark re-
ported that “several” Whitetailed Deer (days 46, 365,
367, 373, 408, and 811), Bison (days 354, 406, 408,
and 413), Mule Deer (day 404), or Blacktailed Deer
(day 602) were killed. In those cases, “several” was
recorded as seven animals killed. Similar to game seen,
the number of animals killed was recorded for all spe-
cies on all days.
Third, herd size. If Lewis and Clark reported sight-
ing large numbers of a particular animal, a value of ten
was assigned to that species on that day. A value of ten
was also assigned if Lewis and Clark reported killing
10 or more of one species on a single day. I then added
game seen, game killed, and herd size values for all
species on each day to obtain a daily measure of wild-
life abundance. Again, this was done for all 863 days
of the expedition.
I also developed a similar convention to quantify the
relative abundance of native people that Lewis and
Clark encountered each day of their journey. If Lewis
and Clark observed old sign, that was assigned a value
of one, fresh sign a two, and if Lewis and Clark actu-
ally saw native people, a three. If Lewis and Clark met
more than ten native people on a given day that was as-
signed a value of ten. On most days Lewis and Clark
traveled together but on a few occasions they took sep-
arate routes, most notably on the return trip. In those
cases, Lewis’ observations were recorded separately
from Clark’s. These conventions produced nearly 40 000
numerical data entries. To facilitate analysis, Lewis
and Clark’s route was divided into 55 trip segments
(Table 1), for which mean daily abundances of wildlife
and mean daily abundances of native people were cal-
Vol. 121
culated. It should be noted that Lewis and Clark gen-
erally sent their best hunters ahead of the main party
so that game would more readily be encountered.
Lewis and Clark left St. Louis, Missouri on 14 May
1804 and proceeded, via watercraft, up the Missouri
River through present-day Missouri, Kansas, Nebraska,
Iowa, South Dakota, and into North Dakota where they
built Fort Mandan in close proximity to the Mandan
and Hidatsa villages. Lewis and Clark over-wintered at
Fort Mandan, and then ascended the Missouri River
into present-day Montana during the spring of 1805.
After leaving their larger boats and portaging the Great
Falls, Lewis and Clark continued up the Missouri to
Three Forks before ascending the Jefferson and Beaver-
head Rivers, on whose upper reaches they met the Sho-
shone. After obtaining horses from the Shoshone, Lewis
and Clark cached their canoes where Clark Canyon
Reservoir is now situated and traveled over the Con-
tinental Divide into Idaho and down the Lemhi and
Salmon Rivers. From there, Lewis and Clark ascend-
ed the North Fork of the Salmon and crossed Lost
Trail Pass, re-entering Montana.
Next, Lewis and Clark traveled down the Bitterroot
Valley to Lolo Creek, which they traced to its source.
Lewis and Clark then followed the high ridges north
of Idaho’s Lochsa River and eventually descended to
the lower Lochsa, where the explorers met the Nez
Perce. At this point, Lewis and Clark left their horses
and proceeded via canoe down the Clearwater, Snake,
and Columbia Rivers through present-day Oregon and
Washington state. Finally, Lewis and Clark built Fort
Clatsop and overwintered on the south bank of the
Columbia near the Pacific Ocean.
During the spring of 1806, Lewis and Clark retraced
their route, with minor variations, until the expedition
reached present-day Lolo, Montana, where the party
divided. Lewis ascended the Blackfoot River, crossed
the Continental Divide, and proceeded to the Great
Falls on the Missouri River, where the party split a sec-
ond time. Lewis left most of his men to repair the boats
cached in 1805, while he and three companions trav-
eled by horseback to Cutbank Creek, where they met
the Blackfeet. After the only fatal encounter with native
people on the entire trip, Lewis retreated to the Mis-
souri, where he rejoined the rest of his men and togeth-
er they floated down that river until reunited with Clark
below the Yellowstone in present-day North Dakota.
Clark, on the other hand, left Lolo, Montana, and as-
cended the Bitterroot River to Chief Joseph Pass, where
he entered the Big Hole. From there, Clark crossed to
the Beaverhead and refloated the canoes cached in
1805. Clark’s party then proceeded by land and water
to Three Forks, where the group split a second time.
Clark sent some of his men and the canoes down the
Missouri to meet Lewis at Great Falls, while he trav-
eled overland via Bozeman Pass to the Yellowstone. At
this point, Clark fashioned canoes and floated down the
Yellowstone and Missouri Rivers until reunited with
2007
KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS?
TABLE |. Trip segments and itinerary of Lewis and Clark 1804-1806.
Trip
segment
CSmeAIDUNLWNH—
Segment
length (days)
30
o>)
DNOMMANWAO HW
Description
St. Louis to Grand River, Missouri
Grand River to Council Bluffs, Nebraska
Council Bluffs to Big Sioux River, Nebraska
Big Sioux River to above Vermillion River, Nebraska
Above Vermillion River
Above Vermillion River to Bad River, South Dakota
Bad River to below the Cannonball River, North Dakota
Cannonball River to Mandan Villages, North Dakota
Mandan Villages, North Dakota
Fort Mandan, North Dakota
Fort Mandan to Little Missouri River, North Dakota
Little Missouri River to Yellowstone River, North Dakota
Yellowstone River to Milk River, Montana
Milk River to Musselshell River, Montana
Musselshell River to Marias River, Montana
Marias River to Great Falls, Montana
Great Falls portage
Great Falls to Three Forks, Montana
Three Forks to Big Hole River, Montana
Big Hole River to meeting Shoshone in Idaho — Lewis’ account
Present Clark Canyon Reservoir across divide to Lemhi River,
Idaho — Lewis’ account
Above Three Forks to present Clark Canyon Reservoir, Montana
— Clark’s account
Clark Canyon Reservoir to Lemhi River to Salmon River and return
to Lemhi — Clark’s account
Lemhi River, Idaho, to present Lolo, Montana
Lolo, Montana, over Lolo Trail to lower Lochsa River, Idaho
Lower Lochsa River — canoe camp
Canoe Camp to the Dalles, Washington
Cascade Mountains and portage, Washington
Cascade Mountains to the mouth of Columbia River on the north bank,
Washington
Crossed to south bank of the Columbia River, Oregon
Ft. Clatsop, Oregon
Ft. Clatsop to the Cascade Mountains, Washington
Cascade Mountains to the Dalles, Washington
The Dalles to Camp Chopunnish, Idaho
Camp Chopunnish, Idaho
Attempted crossing of mountains
Crossed mountains on Lolo Trail
Present Lolo, Montana — here Lewis and Clark separated
Lewis — up Blackfoot River and across Continental Divide, Montana
Lewis — Divide to Great Falls (split party) Lewis to
Cutbank Creek, Montana
Lewis — Upper Cutbank Creek, met Piegan, fled back to Missouri River
Lewis — Marias River down Missouri to Yellowstone River,
North Dakota
Lewis — Down Missouri until reunited with Clark
Clark — Lolo, Montana, up Bitterroot into Big Hole Valley then to
present Clark Canyon Reservoir and down to Three Forks where
the party again split
Clark — Three Forks to Bozeman Pass and on to the Yellowstone
River, Montana
Clark — Down Yellowstone River to Missouri River, North Dakota
Clark — Down Missouri River until reunited with Lewis, North Dakota
Little Missouri River to Mandan Villages to Ft. Mandan, North Dakota
Heart River to Cannonball River, North Dakota
Aricara villages, South Dakota
Moreau River to White River, South Dakota
White River to Vermillion River, Nebraska
Big Sioux River to Council Bluffs, Nebraska
Platte River to Grand River, Missouri
Grand River to St. Louis, Missouri
Dates
5/14-6/12/1804
6/13-8/1/1804
8/2-8/21/1804
8/22-8/25/1804
8/26-9/2/1804
9/3-9/24/1804
9/25-10/16/1804
10/17-10/23/1804
10/24-11/1/1804
1 1/2/1804-4/6/1805
4/7-4/12/1805
4/13-4/25/1805
4/26-5/7/1805
5/8-5//19/1805
5/20-6/2/1805
6/3-6/15/1805
6/16-7/14/1805
7/15-7/24/1805
7/25-8/6/1805
8/7-8/12/1805
8/13-8/26/1805
8/1-8/16/1805
8/17-8/28/1805
8/29-9/10/1805
9/11-9/19/1805
9/20-10/6/1805
10/7-10/24/1805
10/25-1 1/2/1805
11/3-11/25/1805
11/26-12/6/1805
12/7-3/22/1806
3/23-4/6/1806
4/7-4/18/1806
4/19-5/13/1806
5/14-6/9/1806
6/10-6/24/1806
6/25-6/29/1806
6/30-7/2/1806
7/3-7/7/1806
7/8-7/21/1806
7/22-1/2711806
7/28-8/7/1806
8/8-8/11/1806
7/3-7/12/1806
7/13-7/15/1806
7/16-8/2/1806
8/3-8/11/1806
8/12-8/17/1806
8/18-8/20/1806
8/2 1-8/22/1806
8/23-8/29/1806
8/30-9/3/1806
9/4-9/8/1806
9/9-9/17/1806
9/18-9/23/1806
4 THE CANADIAN FIELD-NATURALIST
Lewis. Lewis and Clark then descended to St. Louis
(2: 64; 3: 6; 4: 6; 5: 6, 110, 176; 6: 6, 80; 7: 6; 8: 8-9,
49, 84).
Results
Lewis and Clark’s observations show an inverse rela-
tionship between wildlife and native people (Figure 1).
Wildlife was abundant only where native people were
absent, and if it had not been for the presence of abo-
riginal buffer zones between tribes at war (Hickerson
1965; Steffian 1991; Martin and Szuter 1999a, 1999b,
2002, 2004; Farr 2001; Laliberte and Ripple 2003),
there would have been little wildlife anywhere in the
West.
Yankton Sioux buffer zone
As Lewis and Clark ascended the Missouri River,
they met the Omahas and Ottes on day 97 and the
Yankton Sioux on day 108 (Figure 2). These two groups
were at war (2: 488), and wildlife was abundant only in
the buffer zone between the tribes. Bison, in particu-
lar, were found only in the center of the buffer zone.
Sioux-Mandan buffer zone
Lewis and Clark met the Teton Sioux on day 135,
the Arikaras on day 148, and the Mandan-Hidatsa on
day 164. Wildlife was not abundant in the area between
the Teton Sioux and the Arikaras, but was abundant
between the Arikaras and the Mandan-Hidatsa (Figure
3). This was because the Teton Sioux and Arikaras were
allied against the Mandan-Hidatsa (3: 156, 161, 195-
196, 207, 226, 233-234, 243-244, 251, 272-273, 295-
297, 304-305; Porsche and Loendorf 1987; Bouchet-
Bert 1999). That is, peace had a negative impact on
wildlife populations while war had a beneficial effect,
similar to conditions Hickerson (1965) reported in the
upper Mississippi Valley (Farr 2001).
Missouri- Yellowstone buffer zone
In 1804-1806 all of Montana between the Missouri
and Yellowstone Rivers was a six-sided buffer zone
between warring tribes (4: 21-22, 67, 108-109, 159-
160, 216, 222, 354, 379, 401, 426, 437; 5: 8-9, 45, 68-
71, 77-80, 85, 87-91, 96-97, 102-106, 123-124, 178,
197, 259, 318; 7: 242, 250; 8: 88, 93-94, 104, 113, 123,
143, 182, 195, 278, 321, 323). The north was con-
trolled by the Blackfeet Confederation, which consist-
ed of five tribes (Ewers 1958), while on the west were
the Flathead, Salish, Kootenay, and their allies. The
Shoshone occupied the southwest (Trenholm and Car-
ley 1964), the Crow the south-central, and the Sioux,
Cheyenne, and their allies the southeast. To the east
were the Mandan, Hidatsa, and their allies (Ahler et
al. 1991). Within this large buffer zone (Martin and
Szuter 1999a, 2002, 2004; Farr 2001), wildlife was rel-
atively more abundant (Figures 4-7) because the war-
ring factions did not hunt along the Yellowstone and
Missouri as frequently as they did more secure envi-
ronments closer to each tribe’s core area. As noted by
Lewis and Clark, tribes did venture into the buffer zone,
Vol. 121
MEAN DAILY ABUNDANCE OF ALL WILDLIFE
0
2 4 8 10
MEAN DAILY ABUNDANCE OF NATIVE PEOPLE
Ficure |. Relationship between the abundance of native peo-
ple and the abundance of wildlife as observed by Lewis
and Clark in 1804-1806. Plotted are the mean daily
abundance of all wildlife species and the mean daily
abundance of native people by trip segments — seg-
ments 1, 2, 54, and 55 were excluded because those
areas were near European settlements. Line fitted us-
ing a smoothing spline with cross validation (Math-
soft 1997: 158-167). X and Y axes are offset. Note that
there are no data points in the upper right as might
be expected if cultural beliefs fostered conservation.
Clearly, it made little difference what native people
believed, or said they believed. Instead, aboriginal
hunting followed predictions derived from optimal for-
aging theory and other evolutionary ecology models.
but only in force due to fear of attack. So the Missouri-
Yellowstone buffer zone was not unhunted (4: 232), in-
stead the area was just hunted less frequently (Farr
2001), which apparently was sufficient to permit great-
er numbers of wildlife.
Deer
Lewis and Clark killed more Whitetailed Deer than
all other large mammals combined. By comparison,
Mule Deer were rare and were found only in tribal
boundary zones, while blacktails were restricted to the
Cascade Mountains west to the Pacific (6: 328, 331,
403-404). Even along the lower Columbia, though,
Lewis and Clark encountered more whitetails than
blacktails. This was because whitetails had a more ef-
fective escape strategy than the other deer (Geist 1998;
Whittaker and Lindzey 2001; Lingle 2002; Robinson
et al. 2002) and thus were less affected by native hunt-
ing. Even where native people were abundant, some
whitetails were usually able to survive (Figure 8a) be-
cause, when discovered, whitetails quickly fled into
riparian thickets from which they could not easily be
dislodged (5: 87; 6: 403). Lewis and Clark noted that
Mule Deer and Elk when chased fled into the open
(4: 136-137; 6: 403), making those species easier to
hunt.
Elk
Lewis and Clark reported that Elk were easier to kill
than Whitetailed Deer (6: 85, 242), which 1s reflected
in the fact that native hunters had a greater impact on
2007 KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? 5
35
Yankton Sioux buffer zone
All
wildlife
30 +
All
wikslifen,
25
Bon!
& 20
£
2
ES
5 i
E Se eiane OMes 5 Yankton Sioux e
I\ /
B fay /
> 104 4 \ :
a Bison / \ ;
Y | / \ i B
\ ison
5 4 yy) / \ x
Y Lat | \ /
A) AGEN / \ /
VY) 7 N \ /
Z a Ts ete
5 - a 4 o4 ig ph tel
101 102 103 104 105 111 112 113
Day
FiGuRE 2. Yankton Sioux buffer zone along the Missouri River as reported by Lewis and Clark in 1804. Wildlife was abun-
dant only in the zone between warring tribes.
80
All
oe Sioux - Mandan buffer zone
70
All
wildlife
= 60 | wf
°o
a
a
As
cs
c
oo
g
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wi A Sioux yee Mandan - Hidatsa
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‘ . onffhtuvolsog oe EEEELSE CHU YE
130 135 140 145 165 170
are
FiGureE 3. Sioux-Mandan buffer zone along the Missouri River as reported by Lewis and Clark in 1804. The Teton Sioux
were allied with the Arikaras against the Mandan-Hidatsa and wildlife was abundant only in the zone between war-
ring factions. There was little wildlife and no Bison in the area between the allied tribes.
6 THE CANADIAN FIELD-NATURALIST Vol. 121
40 5
The abundance of wildlife in the Missouri River
buffer zone as reported by Lewis and Clark in 1805.
“6 Direction of travel was from east to west.
Ga All
Pua
25 |
20 4
154
Mean daily abundance of wildlife and native people
9 11 12 13 14 16 17 18 19 20 21
Trip Segment
FIGURE 4. The mean daily abundance of wildlife and native people along the Missouri River buffer zone as reported by
Lewis and Clark in 1805. On their trip across this section of Montana, Lewis and Clark did not see a single native
person from the time they left the Mandan (trip segments 9 and 11) until they met the Shoshone along the Montana-
Idaho border (trip segment 21). Bison were observed only where native people were absent.
40 5
The abundance of wildlife in the Missouri River
buffer zone as reported by Lewis in 1806.
Direction of travel was west to east.
Mean daily abundance of wildlife and native people
34 35 36 39 40 42 43 48
Trip segment
FIGURE 5. The mean daily abundance of wildlife and native people along the Missouri River buffer zone as reported by
Lewis in 1806. Trip segment 34 included tribes in central Washington, while Lewis and Clark spent trip segment 35
with the Nez Perce. Lewis and Clark then crossed the Bitterroot Mountains (trip segment 36) and separated at Lolo, |
Montana. By trip segment 40, Clark was back at Great Falls on the Missouri River, which he descended to the Man-
dan villages (trip segment 48). As on the upstream journey (Figure 4), Clark did not see a single native personon
this section of the Missouri and wildlife was abundant only where native people were absent. Bison, in particular,
occurred only in the center of the buffer zone.
2007 KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? 7
40
The abundance of wildlife in the Yellowstone
River buffer zone as reported by Clark in 1806
Direction of travel was from west to east
35
s
x
Nn
o
.
Mean daily abundance of wildlife and native people
8
g
GY
Trip Segment
FIGuRE 6. The mean daily abundance of wildlife and native people along the Yellowstone River buffer zone as reported by
Clark in 1806. Trip segments 34, 35, and 36 are the same as those in Figure 5. After Lewis and Clark separated,
Clark returned to Three Forks by trip segment 44 and was on the Yellowstone River by trip segment 46. Although the
Crow stole all of Clark’s horses on the Yellowstone, he did not actually see a single native person on his return trip
across Montana until he neared the Mandan villages (trip segment 48). The only place wildlife was abundant was
along the Yellowstone River and Bison were only seen in the center of that buffer zone.
35
Lewis -- Marias River buffer zone
30
= N nN
oa o uo
Daily abundance of wildlife and native people
r=
Ficure 7. Marias River buffer zone. After returning to Great Falls, Lewis ascended the Marias River and Cutbank Creek in
what is now the Blackfeet Indian Reservation. As Lewis traveled from the Missouri, wildlife became less and less
abundant, while Bison disappeared. Wildlife was reported on days 803 and 804 only because Lewis sent his hunters
downstream 30-40 km to kill Whitetailed Deer for food. On day 804 Lewis met seven Blackfeet who told him their
village was less than one-half day’s travel. Lewis camped with this small group of Blackfeet and next morning (day
805a) awoke to find the Blackfeet attempting to steal his guns and horses. An altercation followed and at least one
Blackfoot was killed, the only native person killed by Lewis and Clark on their entire journey. Fearing retaliation
and annihilation, Lewis fled back to the Missouri, and by his own account, traveled more than 160 km by nightfall
(day 805b). Thus within one day’s hard travel, Lewis went from an area with no game and native people to an area
with abundant game and no natives. Bison were found only where native people were absent (8: 112-140).
8 THE CANADIAN FIELD-NATURALIST
the abundance of Elk (Figure 8b) than they did deer
(Figure 8a). Lewis and Clark did kill a number of Elk
at Fort Clatsop, but only because they purposefully built
the fort where Elk were relatively more common and
native people infrequent (6: 92-93, 95-96, 105, 108,
112). That is, Lewis and Clark constructed Fort Clat-
sop in an intervillage buffer zone to take advantage of
the more abundant Elk. Nevertheless, Lewis and Clark
observed that most of the Elk they killed during the
winter of 1805-1806 had old arrow wounds (6: 208,
210), indicative of intense native hunting. “Many of the
Elk we have killed since we have been here, have been
wounded with these arrows, the short piece with the
barb remaining in the animal and grown up in the flesh”
(6: 208). Lewis and Clark also described how native
people used pit traps to kill Elk. “Then pits are emp-
loyed in taking the Elk, and of course are large and
deep, some of them a cube of 12 or 14 feet. These are
usually placed by the side of a large fallen tree, which
as well as the pit [lie] across the [trails] frequented by
the Elk. [The] pits are disguised with the slender boughs
of trees and moss; the unwary Elk in passing the tree
precipitates himself into the pit which is sufficiently
deep to prevent his escape” (6: 208). Thus, even in thick
coastal forests, Elk were intensely hunted by native
people.
Pronghorn Antelope
Native hunting had an even greater impact on the
abundance of Pronghorn Antelope (Figure 8c). Despite
their great speed, pronghorns were relatively easy for
native people to kill (3: 176; Frison 1991).
Bison
Native hunting controlled the distribution and num-
ber of Bison on the northern Great Plains (Figure 8d).
The only places Lewis and Clark saw Bison, and espe-
cially in large numbers, were in the center of aborigi-
nal buffer zones between warring tribes. This is similar
to what West (1995) documented on the central Great
Plains — if it had not been for warring tribes and buffer
zones, there would have been few Bison anywhere in
North America (Kay 2002).
Bighorn Sheep
Native hunting had an even greater effect on Bighorn
Sheep (Figure 8e). Lewis and Clark reported an abun-
dance of bighorns only in the center of buffer zones far
removed from native people.
Grizzly Bears
Native hunters also controlled the distribution and
abundance of Grizzly Bears (Figure 8f). This is simi-
lar to what Birkedal (1993) reported in Alaska. Aside
from a few grizzlies killed in the rugged Idaho moun-
tains, Lewis and Clark only observed grizzlies in abo-
riginal buffer zones.
Black Bears
Based on Lewis and Clark’s observations and kill
rates, Black Bears were less common than grizzlies,
even in forested areas.
Vol. 121
Moose
Despite spending substantial amounts of time in
what is currently prime Moose habitat, Lewis and Clark
recorded Moose only once (6: 313; 7: 326; 8: 95) and
that was in the center of the buffer zone between the
Blackfeet and the Flathead-Salish. As explained else-
where, native hunting controlled the distribution and
abundance of Moose throughout western North Ameri-
ca (Kay 1997b). Contrary to what is generally believed,
Moose are more abundant in western North America
today (Stevens 1971; Pierce and Peek 1984) than they
were in Lewis and Clark’s time, or any other point in
the past (Kay 1997b).
Grey Wolves
Lewis and Clark observed a direct relationship be-
tween the abundance of game and the abundance of
wolves. Wolves were common only where game was
relatively abundant (4: 85). Thus, wolves were largely
restricted to the same aboriginal buffer zones as were
Bison, Elk, and other ungulates.
Dogs and horses
I also recorded the number of dogs (Canis familaris)
Lewis and Clark purchased when game was in short
supply, and the number of horses (Equus caballus)
the explorers killed for food. Lewis and Clark killed
nine horses and bought (ate) 210 dogs, primarily in the
Columbia Basin, where native people were particu-
larly abundant and wildlife was virtually non-existent
(7: 49, 92). Lewis and Clark also bought large quan-
tities of other foodstuffs from various native peoples,
especially corn from the Mandan-Hidatsa and salmon
from tribes throughout the Columbia Basin.
Discussion
Optimal-foraging theory
According to optimal-foraging theory, high-ranked
diet items are more susceptible to overexploitation than
lower-ranked items (Smith 1983; Stephens and Krebs
1986; Smith and Winterhalder 1992; Butler 2000). The-
oretical considerations and studies of modern hunter-
gatherers both indicate that large mammals are the high-
est-ranked diet items, and that, in general, the larger the
animal, the higher its rank (Smith and Winterhalder
1992; Hill and Hurtado 1996). Moreover, if risk to the
hunter or travel distances are great, only the highest-
ranked diet items should be pursued (Smith and Win-
terhalder 1992). Thus, optimal-foraging theory would
predict that when native people entered aboriginal buf-
fer zones, they should have concentrated their hunt-
ing on the larger species, such as Bison and Elk, caus-
ing those species to decline accordingly. This would
also imply that native people lacked any effective con-
servation strategy regarding these prey items. This pat-
tern was, in fact, observed by Lewis and Clark as they
left various native peoples and entered buffer zones,
first Whitetailed Deer increased, then Elk and then
Bison. Conversely, as Lewis and Clark exited a buffer
zone, Bison disappeared first, followed by Elk, while
2007
=
no
=
co
MEAN DAILY ABUNDANCE OF DEER
r.)
2 4 6 8
MEAN DAILY ABUNDANCE OF NATIVE PEOPLE
MEAN DAILY ABUNDANCE OF PRONGHORN ANTELOPE
MEAN DAILY ABUNDANCE OF BIGHORN SHEEP
0 2 4 6 8 10
MEAN DAILY ABUNDANCE OF NATIVE PEOPLE
KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? Y
b
“x 10
ad |
w |
5 |
w 8
Z
< |
z 6
5
rs)
<
4
<
ra)
are ;
= — —
0 6 coud
0 2 4 6 8 10
MEAN DAILY ABUNDANCE OF NATIVE PEOPLE
11 d
MEAN DAILY ABUNDANCE OF BISON
2 “ 6 i) 10
MEAN DAILY ABUNDANCE OF NATIVE PEOPLE
wo
4
_
nN
=k
—t
MEAN DAILY ABUNDANCE OF GRIZZLY BEARS
°
0 2 a 6 8 10
MEAN DAILY ABUNDANCE OF NATIVE PEOPLE
Ficure 8. Relationship between the abundance of native people and the abundance of various wildlife species as observed
by Lewis and Clark in 1804-1806. Plotted are the mean daily abundance of species and the mean daily abundance of
native people by trip segments — segments 1, 2, 54, and 55 were excluded because those areas were near European
settlements. Lines fitted using a smoothing spline with cross validation (Mathsoft 1997: 158-167). X and Y axes are
offset. (a) Whitetailed Deer, (b) Elk, (c) Pronghorn Antelope, (d) Bison, (e) Bighorn Sheep, and (f) Grizzly Bears.
some Whitetailed Deer were usually able to escape
native hunters (Figure 9). Furthermore, Lewis and Clark
noted that native hunters preferred to kill female ungu-
lates (3: 61, 270) due to that sex’s higher fat content.
Now, killing females runs counter to any conservation
strategy (Kay 1994, 1997b, 1998; Kay and Simmons
2002).
Alvard (1998b, 2002) recently reviewed the condi-
tions under which evolution by natural selection might
favor resource conservation by humans. In short, con-
servation will be favored by evolution only if the
resource is economical to defend. For instance, if
1000 kcal are spent defending a resource, but less than
1000 keal are derived from that resource, evolution
will not favor conservation. For a variety of reasons,
including competition from carnivore predators, large
mammals were seldom, if ever, economical to defend
(Kay 1994, 1998, 2002). Instead the logical, rational
thing to do was to kill-out the large mammals as quickly
as possible and then move on to other resources,
10 THE CANADIAN FIELD-NATURALIST
which is exactly what aboriginal people did (Kay 1998,
2002). Counter-intuitively, once that was accomplished,
native populations actually increased because people
were forced to consume lower-ranked, but more abun-
dant diet items (Hawkes 1991, 1992, 1993). There is
also an evolved discount rate that acts to negate a wide
range of possible conservation practices (Rogers 1991,
1994).
Predator-limited
Even within buffer zones, though, wildlife was not
as abundant as one might think, because the animals
were predator, not food, limited (Kay 1998, 2002).
Food-limited ungulates invariably destroy berry-pro-
ducing shrubs and woody riparian vegetation due to
repeated browsing, and once willows (Salix spp.), cot-
tonwoods (Populus spp.), and aspen (Populus tremu-
loides) decline, so do associated species like Beaver
(Castor canadensis) (4: 189-190), which are dependent
upon those plants for food (Kay 1998 and references
therein; Nietvelt 2001). Lewis and Clark, however, re-
ported that riparian thickets were common in buffer
zones, as were Beaver and berry-producing shrubs
(e.g.; 4: 70, 145-146, 189-190, 247, 278, 332, 374, 391-
392, 399, 414, 419, 428, 435, 451; 5: 14, 42, 46, 59). In
addition, Lewis and Clark noted that Whitetailed Deer
often had twin fawns or triplets, and that even lactat-
ing deer were fat (4: 165), which would not have been
physiologically possible if ungulate populations had
been food-limited. Thus, carnivore predation and occa-
sional hunting by native people (4: 232) kept buffer
zone ungulate populations well below what the habi-
tat could otherwise support (White et al. 1998; Kay
2002).
Estimate of pre-Columbian wildlife populations
A number of investigators have cited Lewis and
Clark’s descriptions of abundant wildlife without real-
izing that those accounts only apply to the center of
buffer zones (Craighead 1998; Wilkinson and Rauber
2002). Botkin (1995: 49-86; 2004: 141-147), for in-
stance, used Lewis and Clark’s observations of griz-
zlies along the Missouri and Yellowstone Rivers to
estimate the number of bears in the western United
States prior to European contact, and arrived at a fig-
ure of 56 000, which others increased to 100 000 for
the entire continent (e.g.; Flores 1998: 61). Although
Botkin (1995: 165-169) acknowledged that native peo-
ple could be important ecological factors, he failed to
realize that native hunting controlled the distribution and
numbers of grizzlies throughout North America (Fig-
ure 8f; Birkedal 1993). During pre-Columbian times,
there may have been no more than 4000-5000 grizzlies
in all of North America because the bears were simply
large packages of fat meat that native hunters killed
at will (Hallowell 1926: 31-37; Birkedal 1993). Similar-
ly, there never were 60 million Bison on the Great Plains
(Seton 1929; Roe 1951), as is commonly believed
(Shaw 1995; Geist 1996; Kay 2002).
Vol. 121
Canadian buffer zones
Aboriginal buffer zones also occurred throughout
western Canada. Palliser, for instance, reported that
“As a general rule the more dangerous the country [due
to Indian attack] the greater the probability of finding
[an] abundance of game, showing in more ways than
one the truth of the old sportsman’s adage the more
danger the more sport. This part of the country is so
evidently the line of direction [demarcation] between
the three hostile tribes, that none of them dare venture
into it for hunting, except when driven to desperation
by hunger, they endeavor to snatch their game from
between the jaws of Scylla and Charybdis. Much there-
fore as I enjoyed [this] locality for a hunting camp, see-
ing buffalo on all sides, elk feeding in the distance, and
fresh deer tracks in every direction... Boucharville
[Palliser’s companion] did not relish it at all, and began
already to calculate how soon we were to go away”
(Palliser 1969: 266-267). “The abundance of game here
[and not anywhere else] is accounted for by its being
the neutral ground of the Crees, Assineboines, and
Blackfeet; none of these tribes are in the habit of re-
sorting to its neighborhood except in war parties.... We
are now in the heart of the buffalo country. This region
may be called a buffalo preserve, being the battle-ground
between the Crees and the Blackfeet...” (Spry 1968:
146). Like Lewis and Clark, Palliser observed grizzlies
primarily in aboriginal buffer zones.
Alexander Henry the Younger, describing conditions
on the Red River, also noted that grizzlies were found
almost exclusively in aboriginal buffer zones — “they
[grizzlies] are — very numerous, and seldom are molest-
ed by the hunters [Indians], it being the Frontier of the
Sieux [sic] and their enemy where none can hunt in
safety. Here they [the bears] breed and multiply in the
greatest security” (Gough 1988: 72). While Henry Hind
(1971: 28-29) reported the presence of five aboriginal
buffer zones on the northern Great Plains — four in
Canada and one in the United States. “The following
are celebrated “‘war-paths,’ where hunting is generally
disallowed although game from that circumstance is
usually more abundant. 1. “The War-path River’ and
war road of the Lac la Pluie, Ojibways and the Sioux,
from Rainy River to Red Lake River, thence across the
prairies in the Valley of Red Lake River to Miniwahen
or Devil’s Lake, in Dakotah [sic] Territory. 2. ‘War-
path River, from the southwest corners of Lake of the
Woods to Roseau River, thence to the prairies west of
Red River — same tribes. 3, “War-path River’, from Lake
Winnipeg to the Little Sashatchewan [sic], thence to
the prairies south of Manitobah [sic] Lake — the old
war-path of the Swampy Crees, the Assinaiboines and
Sioux, also of the Swampy Crees and the Lake Win-
nipeg Ojibways. 4. The ‘war-road’ near the Elbow of
the South Branch of the Sashatchewan [River], on the
flanks of the Grand Coteau, of the Blackfeet and Plain
Crees. 5. The ‘war-road’ of the Sioux, Blackfeet and
2007
10
Species buffer zone effects
Mean daily abundance of wildlife and native people
17 18
KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? 1]
Native
19 20 21
Trip segment
FiGurE 9. The effect of native hunting on Bison, Elk, and Whitetailed Deer. As Lewis and Clark ascended the Missouri
River from Great Falls (trip segment 17) and finally met the Shoshone (trip segment 21), wildlife became less and
less abundant. First, Bison disappeared, and then Elk, until only a few Whitetailed Deer remained. This is the pattern
that would be expected if native hunters foraged optimally without regard to conservation.
Crows, in the Valley of the Yellowstone [River]’. In
addition, both Alexander Mackenzie and Samuel Black
noted the presence of an aboriginal buffer zone along
the lower Peace River in northern British Columbia
(Rich 1955: 112; Lamb 1970: 279, 288-289, 404-407).
Similarly, Alexander Mackenzie reported a major In-
dian-Inuit buffer zone in the far north on his 1789 trip
to the Arctic Ocean (Lamb 1970: 181-233) as did Frank-
lin (1969) in 1821-1822.
As in the United States, most observations of wild-
life in western Canada recorded by early European
explorers, including virtually all sightings of Grizzly
Bears, occurred in aboriginal buffer zones. Heretofore
these buffer zone observations have been interpreted
to mean that all of western Canada once teemed with
game, which is simply not true. Instead, many areas
of western Canada were almost devoid of wildlife due
to intense native hunting (Kay et al. 2000).
Prey behavior
Lewis and Clark also reported a direct relationship
between prey behavior and native hunting. In the cen-
ter of buffer zones, where native people hunted only
infrequently, game was relatively tame and could eas-
ily be approached (e.g., 4: 67, 108). Elsewhere, how-
ever, game was exceedingly wary. “The country about
the mouth of this river [Little Missouri] had been
recently hunted by the Minetares, and the little game
which they had not killed and frightened away, was
so extreemly [sic] shy that ... [our] hunters could not
get in shoot [range] of them” (4: 26). “The Borders of
the river [Missouri] has so much hunted by those Indi-
ans ... [that] the game is scerce [sic] and veery [sic]
wild” (4: 39). This also applied to Grizzly Bears and
other animals. “[The bears] appear more shy here [near
the Shoshone] than on the Missouri below the moun-
tains” (4: 426). “These anamals [sic] [beaver] in conse-
quence of not being hunted [in a buffer zone] are ex-
treemly gentle, where they are hunted [though] they
[the beaver] never leave their lodges in the day” (4:
100). Similarly, in 1819 Long observed that Bison fled
in panic at the mere scent of humans. “The wind hap-
pening to blow fresh from the south, the scent of our
party was borne directly [to the Bison], and we could
distinctly note every step of [our scent’s] progress
through a distance of eight or ten miles, by the conster-
nation and terror it excited among the buffaloes. The
moment the tainted gale infected their atmosphere, [the
Bison] ran with as much violence as if pursued by a
party of mounted hunters” (Thwaites 1905: 255-256)
—not unexpectedly, these observations were made in an
aboriginal buffer zone along the Platte River in eastern
Colorado (West 1995). This is identical to what Dia-
mond (1984) reported in New Guinea, where even low-
intensity aboriginal hunting completely altered the be-
2 THE CANADIAN FIELD-NATURALIST
havior of prey species, as well as significantly reducing
prey number (Kay 2002).
Habitat
Over the years, I have retraced most of Lewis and
Clark’s route across North Dakota, Montana, Idaho,
Washington, and Oregon, and there are no habitat fea-
tures that could explain the distribution and abundances
of the various species observed by the explorers. Lewis
and Clark, for instance, did not find any “buffalo” in
the large, treeless valleys of southwest Montana. This
they attributed to the fact that Bison had been driven-
out and/or killed-out by Shoshone hunters, not habitat
characteristics (8: 182). At another point in their jour-
ney, Lewis and Clark commented on how they could
see no difference between the country west of the
mountains and the plains along the Missouri, except
that wildlife was common only on the latter. “I see
very little difference between the apparent face of the
country here [eastern Washington and western Idaho]
and that of the plains of the Missouri only that these
[the Columbia Basin grasslands] are not enlivened by
the vast herds of buffaloe [sic] Elk [etc] which orna-
ment the other” (7: 196). Bighorn Sheep are certainly
restricted to areas with precipitous escape terrain, but
Lewis and Clark found bighorns common only in the
center of aboriginal buffer zones. Other suitable habi-
tat was unoccupied because those areas were more
frequently used by native people.
Moreover, there is no evidence to support the asser-
tion by Mack and Thompson (1982) or Lyman and
Wolverton (2002) that Bison and other ungulates were
rare in the Columbia Basin because those grasslands
were nutritionally deficient. First as Daubenmire (1985)
and others have noted, the Columbia Basin supported
an open range livestock industry for nearly 40 years
during the late 1800s (Oliphant 1968; Galbraith and
Anderson 1971). Since cattle are less efficient herbi-
vores than Bison, there is no physiological reason Co-
lumbia Basin grasslands could have supported large
numbers of free-ranging cattle and not Bison (Van
Vuren 1987; Urness 1989). In addition, a modern Elk
herd on Columbia grasslands not only grew at near the
maximum rate of increase for that species, but pro-
duced huge record-book antlers, as well — all indica-
tive of excellent nutritional conditions (McCorquodale
et al. 1988, 1989; McCorquodale 1991, 1993). Free-
ranging Bison on other intermountain ranges have also
shown high rates of increase (Van Vuren and Bray
1986; Keiter 1997; Bjornlie and Garrott 2001). There
is also no evidence to support the notion that histori-
cally Columbia Basin Bison populations were kept at
low levels by severe winter weather, as proposed by
Daubenmire (1985). After all, Bison thrive in Yellow-
stone National Park (Keiter 1997; Bjornlie and Garrott
2001) and Wood Buffalo National Park (Carbyn et al.
1998) where winter climates are much more severe
than in the Columbia Basin (Urness 1989). Instead,
Bison and other ungulates were rare or absent from
Vol. 121
most of the Columbia Basin because large runs of sal-
mon, and other alternative resources, supported high
numbers of native people (Hunn and French 1981),
who took their preferred ungulate prey to low levels
or local extinction (Kay 1994, 1998, 2002; Chatters
2004).
Native populations and European diseases
It has long been known that native people in the
Americas had no immunological resistance to Euro-
pean diseases, but only recently has it been learned that
those diseases had a significant impact on native peo-
ple prior to direct European contact (Dobyns 1983), or
how this, in turn, caused abnormal increases in wildlife
populations (Neumann 1985; Preston 1996, 1997, 2002;
Kay 1998, 2002; Kay and Simmons 2002). European
diseases, for instance, preceded Lewis and Clark. The
smallpox epidemic of 1780 was especially devastating
(Boyd 1985; Trimble 1985), and its aftermath was
noted by Lewis and Clark (2: 478-482; 3: 285, 295,
311-312; 6: 81-82, 285, 308). In 1804-1806, Lewis
and Clark found four Mandan villages along the mid-
dle Missouri but observed that there had been 12 prior
to the 1780 epidemic. Similarly, Arikaras villages were
reduced from 32 to 2 (Ahler et al. 1991: 57). Thus, if
Lewis and Clark had journeyed west in 1775 instead of
1804-1806, they would have met more native people
and correspondingly there would have been even less
wildlife (Geist 1998: 4-5; Kay 1998, 2002). Further-
more, European diseases may have decimated native
populations throughout western North America as early
as 1550-1600 (Ramenofsky 1987; Campbell 1990;
Kornfeld 1994: 198; Preston 1996, 1997, 2002), which
suggests that pre-Columbian wildlife populations were
likely much lower than even what Lewis and Clark
experienced. Butler (2000), who studied resource de-
pression in the Columbia Basin, reported that high-
ranked diet items, such as ungulates, increased only
after epidemic diseases decimated native populations
ca. 1550. Similarly, Chatters (2004: 72-73) reported
that Bison numbers increased only when native pop-
ulations declined.
Conclusions
Contrary to prevailing paradigms (Lyman and Wol-
verton 2002; Moore 2002; Wilkinson and Rauber 2002),
native people controlled the distribution, abundance,
and behavior of wildlife, and large mammals were com-
mon only in boundary or buffer zones between warring
tribes (Martin and Szuter 1999a, 1999b, 2002, 2004;
Farr 2001). It is also clear that Lewis and Clark recog-
nized this phenomenon, for Clark (8: 328) “observed
that in the country between the [Indian] nations which
are at war with each other the greatest numbers of wild
animals are to be found.” This pattern can only be ex-
plained if native hunters pursued an optimal-foraging
strategy and did not employ any effective conservation
measures (Alvard 1998b, 2002). Only twice did Lewis
and Clark report high wildlife values and encounter
2007
large numbers of native people on the same day. In
both cases, native hunters were killing as many animals
as possible (3: 176, 253-255). Moreover, Lewis and
Clark were only able to complete their journey because
of the food, horses, and above all else, knowledge that
they received from native people. There were no un-
named streams, there were no unnamed mountains, and
there was no wilderness (Kay and Simmons 2002).
As noted by Lewis and Clark, the West was even more
densely populated prior to the smallpox pandemic that
decimated native people in 1780.
These data have important implications for anthro-
pology and archaeology, as well as other disciplines.
Most anthropological subsistence models, for instance,
incorporate the view that native people harvested un-
gulates at or near sustained yield levels, yet these and
other data do not support that assumption (Kay and
Simmons 2002). Similarly, cultural or religious beliefs
are often invoked to explain how aboriginal peoples
interacted with their environment (Krech 1999), yet
irrespective of what the 40 or so native groups encoun-
tered by Lewis and Clark believed, or said they be-
lieved, the ecological patterns were identical, at least
regarding large mammals (see Figure 1). This is simi-
lar to what Jerozolimski and Peres (2003) reported for
modern subsistence hunters in South America, where
neither ethnicity nor culture slowed the depletion of
game stocks. Finally, these data support the hypothe-
sis that native people were keystone predators, who
once structured entire ecosystems (Kay 1998, 2002;
Kay and Simmons 2002); i.e., ecologists need to aban-
don the myth of once abundant wildlife and instead
recognize that unhunted ungulate populations are out-
side the range of historical variability.
Acknowledgments
This research was supported by a grant from Utah
State University and a book contract from Oxford Uni-
versity Press. Susan Durham conducted the smoothing-
spline analyses. Fred Wagner, William Preston, Paul
Martin, Cliff White, Valerius Geist, Richard Keigley,
and Randy Simmons reviewed earlier drafts of this
manuscript, as did three anonymous reviewers.
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Received 4 July 2005
Accepted 27 June 2007
The Influence of Air Pollution on Corticolous Lichens near
the Strathcona Industrial Area, Alberta
M. EvsinGer!, E. BURRELL', N. DEBRUYN!, K. TANASICHUK!, and K. TIMONEY-
'Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E1 Canada
* Treeline Ecological Research, 21551 Township Road 520, Sherwood Park, Alberta T8E 1E3 Canada; email: ktimoney @
compusmart.ab.ca; corresponding author
Elsinger, M., E. Burrell, N. deBruyn, K. Tanasichuk, and K. Timoney. 2007. The influence of air pollution on corticolous lichens
near the Strathcona Industrial Area, Alberta. Canadian Field-Naturalist 121(1): 17-23.
Lichens that grow on the bark of mature trees were studied at 35 sites along an air pollution gradient east of Edmonton,
Alberta. Data on species composition, richness, and cover were recorded in October 1999 in a matrix of sites that extends
from a known source of pollutants (the Strathcona Industrial Area) east across Strathcona County. Air pollution is affecting
the corticolous lichen community. Lichen species richness and total cover increased with distance from the pollution source.
Species richness in areas distant from pollution was roughly twice that in areas near the Strathcona Industrial Area. Xantho-
ria fallax and Phaeophyscia orbicularis were the most pollution tolerant lichens. Xanthoria hasseana, Ochrolechia
arborea, Physcia adscendens, Parmelia sulcata, and Melanelia albertana were rare or absent near the pollution source and
common in more distant areas. Most of the 15 species assessed were sensitive to air quality to some degree. Some lichens
near the refineries and in Sherwood Park showed abnormal coloration and poor thallus integrity indicative of stress. We dis-
cuss implications for human health.
Key Words: corticolous lichens, air pollution, health, refineries, vehicle emissions, Alberta.
Lichens are well-known indicators of air quality.
While most lichen biomonitoring studies have been
conducted in Europe and the United States, there is a
growing body of such studies in Canada. In western
Canada, 49 lichen species have been used to study gen-
eral air pollution, while 39, 23, and 18 species have
been used to monitor sulphur dioxide, photochemicals,
and fluoride, respectively (Thormann 2006). Lichens
have proven useful to characterize the concentrations
and deposition patterns of air pollutants over large
areas, particularly for metals, sulphur, and nitrogen
compounds (Enns 2001*).
The fungal and algal partners of the lichen symbiosis
live in a delicate balance. Any pollutant that affects the
well-being of either partner affects the lichen as a whole
(Brodo et al. 2001). Sensitivity to air pollution is relat-
ed to several features of lichens. They obtain nutri-
ents only from air and precipitation; they accumulate
- compounds in their thalli and have inefficient mecha-
nisms of excluding or eliminating wastes, they have no
| protective epidermis or cuticle, and they are quick to
show symptoms of stress (Case 1981*; USGS 2005*).
The concentration and constituents of air pollution
vary in time and space. In Alberta, there are several
important constituents and sources of air pollution
(Alberta Environment 1996*; Alberta Health and Well-
ness 2003). Sulphur dioxide (SO) is released from na-
tural gas processing plants, oil sands facilities, power
plants, gas flaring, oil refineries, pulp and paper mills,
and fertilizer plants. In Alberta, 42% of SO, is emitted
from natural gas processing plants (Myrick and Hunt
1998*). Hydrogen sulphide is released from fugitive
emissions (leakages) at petroleum refineries, tank farms
for unrefined petroleum products, natural gas and petro-
chemical plants, oil sands plants, sewage treatment
plants, kraft pulp and paper plants, and animal feedlots.
Natural sources include sulphur hot springs and some
wetlands and lakes.
Nitrous oxides are produced from high temperature
combustion of natural gas, coal, oil, and gasoline. Nitro-
gen dioxide (NO,) is a pungent toxin. Ozone near
ground level comes from reaction of nitrous oxides and
volatile organic compounds in the presence of sunlight
and from transport of upper atmosphere ozone down
to ground level. Ozone levels are generally lower in
urban and polluted areas as ozone is destroyed by nitric
oxide (NO). Hydrocarbons come from a variety of in-
dustrial and natural sources. The most abundant hydro-
carbon in the Earth’s atmosphere is methane, a potent
greenhouse gas. Reactive hydrocarbons such as alkenes,
alkynes, and aromatics are toxic to humans and other
animals and vegetation. They react with nitrous oxides
in the presence of sunlight to form ozone.
Particulates come from a variety of sources (typi-
cally combustion) and can act as carriers for biologi-
cally active metals and combustion products. Particu-
late monitoring is based on the mass of particles filtered
per m° of air; PM,, and PM, . refer to particles <10 um
and <2.5 um respectively. In Alberta, the ratio of PM, .:
PM, in recent years has ranged from 0.52 to 0.59 (Pol-
lution Watch 2006*).
Alberta is the largest emitter of air pollutants in
Canada, with a 2005 release of 1.339 billion kg, 28.4%
of the national total, an increase of 62% over its 2002
pollutant release (Pollution Watch 2006*). Among the
criteria air contaminants that affect human and ecosys-
17
18 THE CANADIAN FIELD-NATURALIST
tem health, Alberta released 412.8 million kg of sulphur
dioxide and 403.9 million kg of oxides of nitrogen in
2005 (Pollution Watch 2006*).
Lichens can be harmed by a variety of air pollutants,
including sulphur dioxide, sulfuric and nitric acids,
ozone, hydrocarbons, fluorides, and metals such as
copper, lead, and zinc (Brodo et al. 2001). Sulphur
dioxide impacts on lichens have been noted in Europe
since the early 19" century (Simonson 1996*) and can
cause decreased species richness, diversity and vitality
(Case 1981*). Pollution-tolerant lichens may exhibit
increased cover in polluted areas because of decreased
competition with more sensitive species, such as Mel-
anelia albertana (Muir and McCune 1988; Forest
Health Highlights 1994*). Comparative study of the
lichens in an urban or industrial area with those dis-
tant from pollution can provide a good indicator of
air quality.
Our objective was to determine if there are spatially
detectable effects on the lichen communities of mature
tree bark within and downwind of pollution sources. We
focused on the corticolous lichen species that grow on
Balsam Poplar (Populus balsamifera) and Trembling
Aspen (Populus tremuloides) trunks.
Study Area
The study area and the Strathcona Industrial Area
(“Refinery Row”, Figure 1) are located east of Edmon-
ton, Alberta. Refinery Row contains a large group of
refineries and other industrial plants and is downwind
of Edmonton. Sherwood Park, Fort Saskatchewan, and
rural Strathcona County are downwind of Refinery
Row. Major sources of air pollution from Refinery Row
are industrial plants and vehicle emissions (Alberta
Environment 1997*).
Methods
Study sites spanned an area roughly 28 km east-west
and 14 km north-south downwind (east) of the primary
pollution source in the Strathcona Industrial Area.
Study sites were located in naturally forested areas both
public and private. Two crews of two people each sam-
pled a total 35 sites in mid-October 1999. Only mature
trees with diameters >20 cm at breast height were sam-
pled. Balsam Poplar was the preferred tree as its rough
bark afforded abundant microsites for lichens. Aspen
were also sampled. Prior to sampling, we learned to
recognize a subset of the lichen flora. We focused on
15 taxa readily identifiable in the field.
At each site, we chose the four trees with the high-
est lichen cover. These trees had rough or fissured bark
and were usually the largest and oldest trees present.
Our data indicate average maximum cover rather than
average cover. For each tree, we noted the tree species
and diameter at breast height and determined the per-
cent cover of each lichen species growing on the bark
in the zone 0.5-1.5 m above the ground. Locations were
recorded with a GPS unit. Lichen names follow Brodo
Vol. 121
et al. (2001).
For each site, we determined mean percent cover for
each species and all combined species and the total
number of species. Lichen cover and location data
were analysed statistically. While air pollutants origi-
nate from the entire industrial area, and from Edmon-
ton in general, we defined a single pollution source
point to quantify distance from source. That point was
defined as the junction of Baseline Road and 34" Street
on the west side of Refinery Row. Distances from each
site to the defined pollution source were determined
in a geographic information system.
Results
Of the 15 taxa of corticolous lichens assessed, six
tended to be common and dominant and nine uncom-
mon and non-dominant (Tables 1, 2). Lichen species
richness and cover increased with distance from the
pollution source (Figure 2). Relationships with dis-
tance were strongest for species richness and for non-
dominant species cover (Table 3). Both richness and
cover appeared to rise exponentially with distance.
Species richness and total cover increased gradually
from west to east over the first 15-20 km of the pollu-
tion gradient, then increased more rapidly (Figure 3).
Minimum richness and cover were found in the north-
western sites. Richness and cover continued to rise over
the gradient indicating that air quality influenced the
lichen community across the entire area. Some lichens
near the refineries and in Sherwood Park showed ab-
normal coloration (e.g., pale colors, or pinkish hues)
and poor thallus integrity (partly disintegrated thalli)
indicative of stress.
Of the six dominant species, all but one species
(Phaeophyscia orbicularis) increased in cover with dis-
TABLE 1. Corticolous lichen species observed on mature
Balsam Poplar and Trembling Aspen bark in the study.*
Dominant Species
Melanelia albertana
Ochrolechia arborea
Phaeophyscia orbicularis
Physcia adscendens
Xanthoria fallax
Xanthoria hasseana
Non-dominant Species
Arthonia patellulata
Bryoria spp.
Evernia mesomorpha
Flavopunctelia flaventior
Hypogymnia physodes
Parmelia sulcata
Parmeliopsis ambigua
Physcia aipolia
Usnea spp.
* non-dominant species occurred at <5% cover at most sites
2007 ELSINGER, BURRELL, DEBRUYN, TANASICHUK, and TIMONEY: AIR POLLUTION 19
Ficure 1. (a) Location of the 35 study sites. The area marked with an X, at the corner of Baseline Road and 34" Street
(Edmonton), was the reference point for measurement of distances from the pollution sources. (b) Part of the Strathcona
Industrial Area as seen from the corner of Baseline Road and Broadmoor Boulevard (west side of Sherwood Park.
9 June 2006).
20
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE 2. Cover (%) of six common, dominant lichen taxa, non-dominant species cover, total cover, species richness, distance
from pollution source, and location (UTM, Alberta 10TM projection) for the 35 study sites.
e S
aS} 3 a a eS
Ss s aS 5 aS
FOES SUS US) WUE Sis 5
SUS eS © 8 2 8 S
2 SiS) oS <8 © 3 5
a =3 O8 CaS) as >
| 1.53 1.28 9.50 10.50 9.25
2 1.53 WS) 6.25 27.50 15.50
3 1.25 So} 6.25 30.75 13.00
4 9.50. 6.25 5.50 18.75 9.25
5) 1.03 0.33 10.50 DBS) 27.00
6 1.03 1.05 17.00 9.50. 8.50
iT 4.03 5198 Te 9.50 13.50
8 0.08 SM Si20 25 8.25
9 0.28 18.25 15.00 DAS
10 0.55 12250 20:50 14.25
11 0.30 35) 9.53 ITS 34.50
12 0.28 3.53} 4.78 17.00 11.25
13 0.55 5.28 18.28 10.50
14 0.10 12.00 11.28 18.00
15 0.10 13.75 13.25 9.50
16 0.05 9.00 18.75 15.00
17 0.08 6.25 15.03 17.50
18 0.08 15.25 0.28 12.00
19 30.00 0.08 4.75
20 0.03 16.25 0.08 9.00
21 0.30 3.50 26.25 0.03 14.50
2D) 0.03 0.30 23.75 1.55 16.25
23 0.08 38.00 4.00 eS
24 10.00 18.78 1.03
25 0.55 14.25 0.28 20.00
26 0.05 14.50 155 15.00
27 0.03 MBAS 6.75 3.53
28 0.05 9.50 0.55 1.05
29 0.55 14.28 1.03 3.50
30 10.50 0.08 16.25
31 1.03 16.25 0.08 S25)
29) 3.03 26.25 6.30 4.78
33 0.53 35.00 155 DN DS
34 31.25 3.00 26.52
35 0.05 10.75 6.25 3.25
tance from the pollution source (Figure 4). Xanthoria
fallax and Phaeophyscia orbicularis were the most
hardy lichens, present in virtually all sample sites. Xan-
thoria hasseana and Ochrolechia arborea increased
appreciably in cover with distance from the refineries.
Physcia adscendens, Parmelia sulcata, and Melanelia
albertana were rare or absent near the pollution source
and common in more distant areas. Similarly, Flavop-
unctelia flaventior, Hypogymnia physodes, Usnea sp.,
Evernia mesomorpha, and Arthonia patellulata were
rare to absent near the refineries. Most species were
sensitive to air quality to some degree.
Discussion
Lichens are good bioindicators of air quality in the
region. There were clear trends of decreasing cover and
richness with increasing proximity to Refinery Row.
He Ee > = S bo
Ss 2G oe 2 es =
BS SS een = 23) Ss S =
S33 Sh 6 Be B ‘ e
1, reduced levels of
ARA and high EPA/ARA ratios decrease the inci-
dence of pseudoalbinism, incomplete eye migration
and hooked dorsal fin (Ness et al. 1995; McEvoy et al.
1998a; Neess and Lie 1998; Shields et al. 1999; Hamre
et al. 2002). Other nutritional differences between arte-
mia and copepods may contribute to pigmentation, such
as the greater proportion of polar lipids (McEvoy et al.
1998b), vitamin A (or its precursors, carotenoid com-
pounds) and iodine (Hamre et al. 2005) in copepods.
Less attention has been paid to the development of
ambicoloration. It can be voluntarily produced in the
Japanese Flounder, Paralichthys olivaceus, by immers-
ing larvae in water containing retinoic acid (Haga et
al. 2005). Ambicoloration is an undesirable trait in
hatcheries that is not normally induced voluntarily, but
rather occurs spontaneously at incidence rates that
depend on diet. Most diets that reduce the incidence of
CHABOT AND MILLER: AMBICOLOURED ATLANTIC HALIBUT 37
pseudoalbinism also increase the incidence of hyper
melanosis of the blind side (for a review, see Haga et al
2004). For instance, an excess of vitamin D, (Hasegawa
et al. 1998; Haga et al. 2004) or A (Miwa and Yamano
1999) can increase the rate of ambicoloration in the
Japanese Flounder. In Atlantic Halibut, copepod-based
diets sometimes resulted in ambicoloration for up to
30-40% of the fish (Shields et al. 1999; Hamre et al.
2002; Hamre et al. 2005), but in other studies it did not
(Ness et al. 1995; Ness and Lie 1998). This is likely
the result of variability in nutritional properties of the
copepods used, as the quantities and ratios of unsatu-
rated fatty acids, vitamins and other nutrients changed
between batches or size-classes of copepods within and
between studies (e.g., Ness et al. 1995; McEvoy et al.
1998a; Shields et al. 1999; Hamre et al. 2002). Diet-
induced ambicoloration is conceivable in wild halibut
as well: hatching time, egg condition and environmen-
tal conditions all influence when larvae begin to meta-
morphose and the availability and quality of prey items.
Laboratory studies have shown that the critical period
during which diet influences the development of post-
metamorphic pigmentation is short, making it possible
for wild fish to have a suboptimal diet near the begin-
ning of metamorphosis and become ambicoloured.
Malpigmentation can also occur in post-metamor-
phic flatfish. Many wild fingerlings of Speckled Floun-
der (Paralichthys woolmani) that were normally pig-
mented when captured developed white spots on their
ocular side within a few days. Most of them returned
to a normal pigmentation within one year, except for
a few that retained some pigmentation on their blind
side (Benetti 1997; Venizelos and Benetti 1999). Light
reaching the blind side where it is not in contact with
the bottom, and lack of sediment in which fish can bury
themselves can also result in some pigmentation on the
blind side of flatfish (Cunningham 1893; Ottesen and
Strand 1996). However these are examples of staining,
not ambicoloration. True ambicoloration appears to
be in place soon after metamorphosis.
It is difficult to assess whether factors leading to
malpigmentation in hatchery-reared halibut would do
so in the wild, or if these factors are even relevant in
the wild. Halibut larvae likely encounter appropriate
prey items in the wild. However, considering the high
incidence of malpigmentation when diet is inappro-
priate for only a few days in hatcheries, and the fact
that non-diet related causes of malpigmentation in cap-
tive Atlantic Halibut (light, sediment type, stocking
density, etc.) are even less likely to be problematic in
the wild, diet is the most likely of the known causes
of malpigmentation to occur in the wild. Other possi-
ble causes, such as a genetic causes, remain possible,
however. True albinism (i.e., presence of red eyes) is
inheritable in the Japanese Flounder (Shikano 2005),
but diet appears much more important than genetic fac-
tors in the development of pseudo-albinism and ambi-
colouration (Seikai and Matsumoto 1994).
38 THE CANADIAN FIELD-NATURALIST
Few wild ambicoloured Atlantic Halibut have been
reported, but reporting rate is unknown. Reporting
malpigmentation should be encouraged in scientific
surveys for this species as well as others to assess
incidence and mortality rate of ambicoloured fish and
improve our understanding of the causes of ambi-
colouration in Atlantic Halibut, and of taxonomically
related incidence rates in flatfish (e.g., Diaz de Astar-
loa et al. 2006).
The specimen presented in this study has been pre-
served and is part of the Maurice-Lamontagne Insti-
tute Collection (accession number 11633).
Acknowledgments
We thank G. Hogan and V. Puvanendran for their
helpful comments on an earlier version of the manu-
script.
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Accepted 22 February 2007
Micro-Propagation of White-top Aster, Sericocarpus rigidus, a Threatened
Species from the Garry Oak Ecosystem in British Columbia
BRENDA FREY, CHAIM KEMPLER, and DAvID L. EHRET!
Pacific Agri-Food Research Centre, Agriculture and Agri-Food Canada, PO Box 1000, Agassiz, British Columbia VOM 1A0
Canada
‘Author to whom correspondence should be addressed; e-mail: ehretd @agr.gc.ca
Frey, Brenda, Chaim Kempler, and David L. Ehret. 2007. Micro-propagation of White-top Aster, Sericocarpus rigidus, a
threatened species from the Garry Oak Ecosystem in British Columbia. Canadian Field-Naturalist 121(1): 40-45.
A study was undertaken to examine the feasibility of using micro-propagation techniques to regenerate White-top Aster (Seric-
ocarpus rigidus), a threatened species native to the Garry Oak ecosystem in coastal British Columbia. Shoot multiplication
from explants was compared in three different basal media formulations. The effects of varying [AA (indole-3-acetic acid)
and kinetin on shoot production were also investigated using one of the three basal media. Shoot production was most suc-
cessful on modified MS (Murashige and Skoog) media containing various concentrations of BAP (benzylaminopurine) and
GA, (gibberellic acid), followed by media containing IAA and kinetin at concentrations of 2 mg l'!. Root formation occurred
readily on modified MS media with IAA and reduced macronutrient and micronutrient concentrations. The study showed
that S. rigidus can be successfully propagated using in vitro shoot multiplication.
Key Words: Sericocarpus rigidus, Aster curtus, White-top Aster, micro-propagation, Garry Oak ecosystem, British Columbia.
The Garry Oak ecosystem has a geographic distri-
bution that spans the west coast of North America, from
southwestern British Columbia to southern Califor-
nia. In recent years, the survival of this ecosystem in
Canada has been threatened by fire suppression and
by residential, industrial and agricultural develop-
ment. These activities have resulted in loss of species
habitat, habitat degradation and the spread of inva-
sive exotic species (Garry Oak Ecosystem Recovery
Team 2002"). The Garry Oak ecosystem contains more
plant species than any other terrestrial ecosystem in
coastal British Columbia, and many plants that live in
the ecosystem do not occur elsewhere in Canada. It is
now estimated that less than 5% of all Garry Oak sites
remain in their natural state (Garry Oak Ecosystem
Recovery Team 2002’).
The White-top Aster, Sericocarpus rigidus (formerly
Aster curtus), is an herbaceous perennial species of
Asteraceae native to low elevation grasslands in the
Pacific Northwest in the United States and the associ-
ated Garry Oak ecosystem on Vancouver Island, British
Columbia (Clampitt 1987). Due to the serious loss of
habitat, the Committee on the Status of Endangered
Wildlife in Canada (COSEWIC) designated White-top
Aster as a threatened species in 1996, and the species
is now protected under the Species At Risk Act (SARA)
(Environment Canada 2006*). Sericocarpus rigidus pro-
duces underground rhizomes, and as a result, occurs in
patches which can vary tremendously in size. Shoots
emerge from rhizomes in April, flower in July and Au-
gust, then die back in the autumn. Seedlings are rarely,
if ever, observed, making rhizomes the major form of
propagation (Douglas and Illingworth 1997). Even
though S. rigidus produces an abundance of seeds, via-
bility is low and often uneven. In pollination studies,
Clampitt (1987) found that excluding pollinators by
bagging ramets significantly reduced the percentage
of viable seeds (measured with tetrazolium chloride)
from 20% in the unbagged treatment to 5% in the
bagged. Bigger (1999) found that exposed and hand-
pollinated flowers generally had the same number of
viable seed (as tested with tetrazolium chloride), but
differences in seed viability between hand-pollinated
and exposed plants varied greatly among sites, possi-
bly due to pollinator limitations in some patches. Patch
size was also found to influence the degree of preda-
tion by insects. Overall, seed viability varied from 0 to
83%. In a field comparison of open pollinated, between
patch pollinated, within patch pollinated and pollina-
tion excluded treatments, Giblin (1997) found that open
and between patch pollination treatments produced sig-
nificantly more filled seed (39% and 35%, respectively)
than within patch and pollination exclusion treatments
(18% and 9%, respectively). In terms of conditions re-
quired for germination, Clampitt (1987) showed that
stratification and exposure to light increased germina-
tion. Ehret et al. (2004) have shown that the seeds re-
quired an 8-12 week period of stratification for germi-
nation and seedlings are slow to establish. Even so,
Giblin and Hamilton (1999) suggest that the reproduc-
tive biology of the species does not contribute to its
rarity.
Sericocarpus rigidus was one of three species “at
risk” plants studied at the Pacific Agri-Food Research
Centre (PARC) with the objective of developing prop-
agation methods that would allow for the generation
of plant material that could be reintroduced into pro-
tected Garry Oak ecosystems, parks and ecological |
AO
2007
reserves or made available to native plant nurseries.
Because of the difficulties associated with seed germi-
nation, micro-propagation was studied with the objec-
tive of developing a method that could rapidly and reli-
ably produce plantlets.
Materials and Methods
Plant Material
Seeds were collected from two patches of S. rigidus
growing at an elevation of 220 m on Mt. Tzuhalem
(48°47'N, 123°38'W) on Vancouver Island in Septem-
ber 2001 and from another patch at the same location
in September 2002. The collection sites were 100 to
300 m apart. Given the close proximity, all plants may
have been derived from the same clone. Collected seed
count varied from 50 to 200 per plant. Seeds from 2001
were dried and stored at room temperature in the dark
for one year. In September 2002, all seeds were placed
in the dark in a cooler at 4°C. In October 2002, 15 to
30 seeds from each patch were placed on damp filter
paper in petri dishes and incubated in a growth cham-
ber maintained at 20°C and 14/10 h light/dark. After
three weeks, only four seeds had germinated and all
were collected from the same patch in 2001. These
were potted in 10 cm pots containing Sunshine Mix 4,
(Sun Gro Horticulture Inc., Vancouver, British Colum-
bia) and were over-wintered in a greenhouse. In May
2003 the seedlings were hardened off in a cold frame
for four weeks before being transplanted into a silt
loam soil in irrigated raised beds at PARC. After one
season of growth and flowering, seeds were collected
from those plants in October 2003, and after air drying,
were stored in a cooler at 4°C for 3 months. Surface
sterilization of 10 seeds was conducted by suspending
the seeds in a 5% solution of PPM (Plant Preservative
Mixture) (Plant Cell Technology Inc., Washington
D.C., USA) and deionized water (v/v) in a beaker on a
shaker for 24 h. Seeds were aseptically transferred to
a petri dish containing a 1.5% solution of sterile Bacto
agar in deionized water (w/v). The petri dish was sealed
with parafilm and placed in a plastic bag in a refriger-
ator at 3°C in the dark.
After nine weeks at 3°C, one seed germinated. This
was the only seed that germinated during the next five
months. The seedling was removed from the petri dish
under sterile conditions and placed in a 350 ml jar con-
taining half-strength MS (Murashige and Skoog 1962)
media containing 1.0 mg L' TAA and 0.1 mg L" kinetin.
The jar was placed in a growth room maintained at
22—25°C with a 16/8 h light/dark cycle using fluores-
cent light with a PPFD (photosynthetic photon flux
density) of ~50 uwE m? s"!. After five weeks, the fully
developed, healthy seedling was transferred to a mod-
ified MS medium containing half the concentration
of ammonium nitrate (825 mgeL), 2.0 mgeL IAA and
2.0 mg |! kinetin. After four weeks on this medium,
shoot and roots were excised and cultured on fresh
media. Cultures were maintained by bimonthly trans-
FREY, KEMPLER, AND EHRET: MICRO-PROPAGATION OF WHITE-TOP ASTER 4]
fer to fresh media until enough shoots were generated
to conduct shoot multiplication and rooting experi-
ments. The excised root tissue failed to grow after being
removed from the shoot and was discarded.
Shoot Induction
A replicated shoot multiplication experiment was
initiated to determine the effects of various concen-
trations of IAA plus kinetin or BAP plus GA, on the
growth of explant shoots produced on modified MS
media. The experimental design was a randomized
complete block design consisting of seven media treat-
ments with six replicate jars, each containing five ex-
plants. The experiment included three modified MS
basal medium (BM) formulations with each formula-
tion containing a different concentration of macronu-
trients, micronutrients and organic additives (Table 1).
BM1 was a formulation used in earlier micropropa-
gation media development using the cultivated violet
species, Viola praemorsa (Ehret et al. 2004). BM2 and
BM3 were modified MS media used routinely in the
PARC strawberry and raspberry breeding programs for
shoot proliferation. The ammonium nitrate concentra-
tion of BM1 was half that of the other media because
higher rates of ammonium nitrate have been found to
be toxic to cultures of some plant species because the
uptake of ammonium ions can acidify the medium and
increase the frequency of vitrification (Slater et al.
2003). BM1 was the basal media used in media treat-
ments | through 5 and it was supplemented with five
combinations of IAA and kinetin as follows: 0.5 mg*L
IAA + 1.0 mgeL" kinetin (M1), 0.5 mg*eL! IAA +
2.0 mg*L"! kinetin (M2), 2.0 mg*L! IAA + 1.0 mg*L'!
kinetin (M3), 2.0 mg*L IAA + 2.0 mg*L" kinetin (M4)
and 3.0 mgeL"! IAA +2.0 mg I! kinetin (M5). M4 was
the initial shoot multiplication media used to generate
plant material for the experiment. BM2 was the basal
media used in treatment 6 (M6) and it was supplement-
ed with 2.5 mg*L' BAP and 0.01 mg*L"! GA,. BM3
was the media used in treatment 7 (M7) and it was
supplemented with 1.0 mg*L' BAP and 0.01 mgeL'!
GA,,. The ferrous sulphate and Fe-EDTA supplied in
BMI and BM? was replaced by Sequestrene 138 (Beck-
er Underwood Inc., Ames, Iowa, USA) in BM3.
The pH of all media treatments was adjusted to 5.6—
5.7 and the media were then dispensed into 350 mL jars
which were covered with cellophane and autoclaved.
Explants which had a mean weight of 0.164 g + 0.008
(SE), were aseptically transferred into treatment jars
which were incubated in a growth room maintained at
22-25°C with a 16/8 h light/dark cycle using fluores-
cent light with a PPFD of ~50 uE m” s' for four weeks.
After four weeks, the number of explants that produced
shoots and roots, the number of shoots produced per ex-
plant, and shoot and explant fresh weights were recorded.
Root Induction
A separate rooting experiment was conducted using
BM4 (Table 1) which is used in the PARC berry breed-
42 THE CANADIAN FIELD-NATURALIST Vol. 121
TABLE |. Four modified Murashige and Skoog (MS) basal media (BM) formulations used for S. rigidus culture. Rates are
expressed in mgel! with the exception of Plant Preservation Mixture, which is expressed in ml I".
Induction Media
BM1
Macronutrients
Ammonium nitrate 825
Calcium chloride (anhydrous) 332
Magnesium sulphate (MgSO,-7H,O) 370
Potassium phosphate 170
Potassium nitrate 1900
Micronutrients
Boric acid 6.2
Cobalt chloride 0.025
Cupric sulphate 0.025
Ferrous sulphate 27.8
Manganese sulphate DS,
Potassium iodine 0.83
Sodium molybdate 0.25
Zinc sulphate 8.6
Na,EDTA SHk2
Sequestrene 138 -
Sodium phosphate (NaH,PO,-H,O) -
Organic Additives
Sucrose 20
myo-inositol 1000
Nicotinic acid 0.5
Pyridoxine-HCL 0.5
Thiamine 0.1
Glycine 20
Adenine —
Plant Preservation Mixture 2
Gerlite guar gum 2
ing program for root development. Fifteen shoot sam-
ples initiated on M4 were placed on the rooting medium
containing 1.0 mgeL"! IAA. After four weeks, rooted
plants were transferred to Sunshine Mix 4 potting mix
and moved to a greenhouse where they were acclima-
tized.
Statistical Analysis
Data were analysed using the general linear model
procedure in SAS (SAS Institute, Cary, North Carolina,
USA), for one-way analysis of variance (ANOVA). All
independent factors were fixed. Where ANOVA results
were significant, treatment means were separated by
Duncan’s multiple range tests. Data expressed in per-
centages were transformed by arcsin transformations
before statistical analysis.
Results and Discussion
In general, explants grew rapidly in culture, with
extensive shoot multiplication (Figure 1 a, b). After
four weeks of culture, explant survival ranged from
83.3 to 100 percent with no significant survival differ-
ences among media treatments (Table 2). The percent-
age of surviving explants that produced shoots ranged
from 61 to 100%. Just over 60% of M1 and M3 ex-
plants produced shoots compared to 90-100% of ex-
Shoot Root
BM2 BM3 BM4
1650 1237 464
332 B52) 332
370 370 370
170 170 64
1900 1425 534
6.2 6.2 3.1
0.025 0.025 0.013
0.025 0.025 0.013
27.8 = A
DOS 3} Lil
0.83 0.83 0.42
0.25 0.25 0.13
8.6 8.6 4.3
BD) — =
= 200 100
170 170 85
30 30 30
100. 100 100
0.8 0.8 0.4
80 80 —
D 2; 2
2 2 D
plants cultured on M4—M7. In addition, explants cul-
tured on M1, M2 or M3 produced roots while being
cultured on shoot multiplication media (data not pre-
sented). M1 and M3 which had the lowest concentra-
tions of kinetin produced the greatest number of roots,
13 and 20 % respectively. These results suggest that
aster media should contain kinetin at minimal rates of
2.0 mgeL to avoid root initiation during the shoot
multiplication stage. This supports previous recom-
mendations that kinetin be added to a medium for
cell division, shoot multiplication and axillary bud pro-
liferation while IAA should be added to promote cell
enlargement, root initiation and adventitious bud for-
mation (Kyte and Klein 1999).
Mean explant plus shoot weights ranged from 0.241
to 0.855 grams. No statistically significant differences
were observed among BMI treatments. M6 produced
a significantly greater biomass than all other media,
followed by M7 which had a significantly greater bio-
mass than M1 to M3 (Table 2, Figure 1c). Explants
and shoots cultured on M6 appeared slightly chlorot-
ic compared to those cultured on M7 which were the
darkest green of all treatments (data not presented).
Explants cultured on M2 and M3 were the least vigor-
ous; explant and shoot leaves appeared mottled and there
2007 FREY, KEMPLER, AND EHRET: MICRO-PROPAGATION OF WHITE-TOP ASTER 43
Treatment 7
bis)
a
— 2
—
— cw
— he
Ficure 1. A. Seriocarpus rigidus explant on culture media
after two weeks in culture. B. Explants and developed
shoots on media after 4 weeks. C. Representative sam-
ples of explants and shoots cultured on seven media
treatments after 4 weeks of culture. Treatment labels
1 through 7 represent treatments M1 through M7,
respectively. D. Four-month-old rooted aster plants
in the greenhouse.
ly fewer shoots than M4. Treatments M6 and M7, both
of which contained BAP and GA, and which were on
BM2 and BM3, respectively, produced a greater num-
ber of shoots per explant than most BM1 treatments
containing combinations of [AA and kinetin.
The mean weight of individual shoots cultured on
the different media ranged from 0.037 to 0.097 grams
per shoot. Few consistent differences were observed
were several dead leaves at the base of the explants. among the five BMI treatments. Compared to those
The number of shoots per surviving explant ranged _ treatments, M6 (on BM2 with BAP and GA,) pro-
from 1.72 to 6.19 shoots per explant (Table 2). Within duced shoots with the greatest weight. Despite some
the BMI treatments, M2 and M3 produced significant- differences among BMI treatments, the results show
44 THE CANADIAN FIELD-NATURALIST Vol. 121
that the ratio and the concentration of IAA and kinetin
in BM] did not have a consistent effect on overall shoot
multiplication. Shoots were small and difficult to handle
after four weeks of culture. Extending the culture time
from four to five weeks would likely have increased
shoots to a size that would have made them easier to
transfer and a more desirable size for rooting.
Root initiation was 100% on the BM4 medium (data
not shown). Roots appeared on the shoots after 7 days
of culture, and after 4 weeks a healthy root system had
fully developed. The rooted plants were easily acclima-
tized to a greenhouse environment and showed vigor-
ous growth (Figure 1d). Other methods to induce root
development were not attempted since this technique
was so highly successful.
Micro-propagation methods for commercially im-
portant species of Asteraceae such as sunflower (Hel-
ianthus annuus) employ a variety of techniques to re-
generate plants (Alibert et al. 1994). There are also
methods to micro-propagate other Asteraceae species
of commercial value, such as feverfew (Tanacetum
parthenium) (Simmons 1998), marigold (TJagetes erec-
ta) (Vanegas et al. 2002), Echinacea pallida (Koroch et
al. 2003) and safflower (Carthamus tinctorius) (Orli-
kowska and Dyer 1993). These techniques have in-
creasingly been used on medicinally important Aster-
aceae species such as goldenrod (Solidago virgaurea,
S. canadensis, S. gigantea and S. graminifolia) (Kelem-
ba and Thiem 2004) and those which are rare, such as
Saussurea obvallata (Joshi and Dhar 2003). In Canada,
micro-propagation of the Gulf of St. Lawrence Aster, a
rare species in Prince Edward Island, has been shown
to have potential in conservation efforts (Stewart and
Lacroix 2001).
Meristem culture has been extensively used for the
clonal propagation of horticultural plants. Since the
constituent cells of meristems are genetically stable,
plants regenerated by in vitro culture of shoot apices
without the callus-mediated process of organogenesis
should result in the recovery of genetically identical
progeny. Because the culture was started from a single
seed and mass regenerated in tissue culture, genetic var-
iability is limited. Sericocarpus rigidus propagates
clonally in the wild by underground rhizomes, so in
vitro mass clonal propagation could be a good strategy
for preservation and reintroduction of this threatened
species.
Our study is the first to show that the threatened
Asteraceae species, S. rigidus, can also be successfully
propagated using micro-propagation techniques. Only
one germinated seed was available for use in the study.
However, by its very nature, micro-propagation works
at the level of single seeds or explants, so our situa-
tion is not atypical. The genetic variability of the Mt.
Tzuhalem population of S. rigidus is not known, but
since the plant occurs in clonal patches, it is unlikely
that other seeds of S. rigidus would respond different-
ly to micro-propagation. The best results for shoot mul-
Shoot
weight
(g/shoot)
0.056 + .009 bed
0.056 + .007 bed
0.040 + .005 cd
0.037 + .004 d
0.070 + .013 be
0.097 + .009 a
0.079 + .005 ab
Number of
shoots per
surviving explant
2.68 + 0.29 cd
1.94+0.13 d
1.72+0.14d
4.13 + 0.50 be
2.97 + 0.58 cd
6.19+0.96 a
4.63 + 0.47 ab
oh oK
0.542 + 0.04 b
0.855 + 0.13 a
2K
Explant plus
shoot weight
(g)
0.262 + 0.02 c
0.278 + 0.02 c
0.241 + 0.04 c
0.439 + 0.07 be
0.379 + 0.08 be
Surviving explants
producing shoots
(%)
63.3 +6.1b
81.7 + 6.4 ab
61.1+17.1b
90.0 + 6.8 a
90.0+4.4a
96.7+3.3a
100.0+0a
2K
Explant
survival
(%)
100
86
83.3
100
93.0
100
96.7
NS
(mgeL!)
IAA + kinetin
0.5 + 1.0
0.5 + 2.0
2.0 + 1.0
regulators
2.0 + 2.0
Growth
BAP + GA,
2.5+0.01
1.04+ 0.01
3.0 + 2.0
Basal
medium
BM]
BM2
BM3
Means (+ SE) within columns followed by the same letter are not significantly different. NS: Non-significant or *significant at P <0.05 and **0.01, respectively.
TABLE 2. Growth and survival of Sericocarpus rigidus explants on basal medium amended with growth regulators after four weeks in culture.
Treatment
Significance
M1
M2
M3
M4
M5
M6
M7
2007
tiplication were obtained with combinations of BAP
and GA,, a regime which has not often been used in
other Asteraceae studies. Further work is required to
determine the optimum basal medium formulation to
use with the most successful combinations of BAP
plus GA, in order to generate the greatest number of
vigorous S. rigidus shoots. Additionally, research with
vegetative tissue as a source of explants, rather than
seeds, could be conducted in order to avoid the time and
difficulties associated with seed germination. Howev-
er, from a conservation point of view, harvesting veg-
etative tissue, particularly rhizomes, from a threatened
or endangered species may be less desirable than har-
vesting seeds because of damage to the plant and dis-
turbance of the site.
The costs of micro-propagation were not evaluated
against more conventional propagation techniques. Tis-
sue culture is cost effective and is used commercially
in the propagation of many plants. Although it requires
more knowledge and extra capital to start a micro-
propagation operation, the rate of propagation is much
faster than in conventional propagation. Given that
hundreds of propagules of an endangered or threatened
species may be generated using micro-propagation,
the effort seems reasonable.
Acknowledgments
The authors thank Georgia Kliever and Frederic
Bounaix for technical assistance, and Matt Fairbarns
for collecting seed and providing advice. This study
was partially funded by the Interdepartmental Recoy-
ery Fund of Environment Canada.
Documents Cited (marked * in text)
Environment Canada 2006. Species at Risk, White-top Aster.
http://www.speciesatrisk.gc.ca/search/speciesDetails_e.c
fm?SpeciesID=237.
Garry Oak Ecosystem Recovery Team. 2002. Recovery
Strategy for Garry Oak and Associated Ecosystems and
their Associated Species at Risk in Canada 2001 — 2006,
1 page.
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Received 5 September 2006
Accepted 26 March 2007
Pattern and Potential Causes of White-faced Ibis, Plegadis chihi,
Establishment in the Northern Prairie and Parkland Region of North
America
Jct A. SHAFFER!, GREGORY A. KNUTSEN?, RON E. MArtTIN?, and JoEL S. BRICE*
'U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 37" Street SE, Jamestown, North Dakota 58401
USA; e-mail: jshaffer@usgs.gov
°U.S. Fish and Wildlife Service, Long Lake National Wildlife Refuge Complex, 12000 353" Street SE, Moffit, North Dakota
58560 USA
316900 125" Street SE, Sawyer, North Dakota 58781 USA
‘Delta Waterfowl Foundation, 1305 East Central Avenue, Bismarck, North Dakota 58501 USA
Shaffer, Jill A., Gregory A. Knutsen, Ron E. Martin, and Joel S. Brice. 2007. Pattern and potential causes of White-faced
Ibis, Plegadis chihi, establishment in the northern prairie and parkland region of North America. Canadian Field-
Naturalist 121(1): 46-57.
The Northern Prairie and Parkland Waterbird Conservation Plan calls for renewed attention to determining the current status
of waterbird populations, their distributions, and conservation needs. It highlights the need for baseline information on the
White-faced Ibis (Plegadis chihi). In response, we examined the historical and current distribution of the ibis in North Dakota
and summarized first sightings and nest records for the provinces and other states composing the northern prairie and parkland
region. The establishment of breeding colonies of White-faced Ibis here may be due to climate and precipitation patterns,
invasion and spread of Narrowleaf Cattail (Typha angustifolia), changes in agricultural practices, habitat loss and range
expansion in the southern and western portions of the species’ range, and increases in ibis populations in the Intermountain
West. We placed special emphasis on North Dakota, a state for which there is scant published information concerning the
current status of this species. In recent decades, the ibis has become a regular breeding-season resident in North Dakota and
in other areas of the northern prairie and parkland region. From 1882 to 2002, there were 145 reports of one or more White-
faced Ibis in North Dakota, including 93 reports during the breeding season (15 May to 31 August), 49 during the non-
breeding season (1 September to 14 May), and three for which the season of occurrence was not reported. Prior to the 1960s,
there were only three records of the species in North Dakota. Observations of White-faced Ibises in North Dakota increased
dramatically between the 1960s and the early 21st century, and the species has been observed nearly annually since 1971.
The first White-faced Ibis nesting activity in the state was recorded in 1978, and to date, there have been 21 known records
of nesting activity in the state. The species nested in large (>300 ha) semipermanent or permanent wetlands within mixed-
species colonies ranging in areal extent from small (0.1 ha) to fairly large (27 ha), and colonies were located in patches of
emergent vegetation dominated by cattails (Typha) and bulrushes (Scirpus). We classify the White-faced Ibis as a fairly com-
mon migrant and a locally uncommon breeder east of the Missouri River and a casual migrant west of the Missouri River.
Key Words: White-faced Ibis, Plegadis chihi, colonial-nesting waterbird, distribution, status, northern prairie and parkland
region, North Dakota.
The White-faced Ibis (Plegadis chihi; hereafter ibis)
is a wading bird that nests in wetlands with emergent
vegetation and forages in shallow wetlands, flooded
hay meadows, or agricultural fields (Ryder and Manry
1994). Northernmost populations undertake regular
north-south migrations, whereas populations in Texas
and Louisiana are mainly year-round residents (Ryder
and Manry 1994). Extralimital wanderings seem to be
more prevalent before the nesting season than afterward
and may represent individuals seeking new nesting areas
when traditional sites are dry or have been destroyed by
humans (Ryder 1967; Ryder and Manry 1994). Dur-
ing the twentieth century, numerous sightings of the
species east and north of its historical range were report-
ed (Sabo 1992; Jorgenson and Dinsmore 2005).
The recently completed North American Waterbird
Conservation Plan (Kushlan et al. 2002) and its regional
offshoot, the Northern Prairie and Parkland Waterbird
Conservation Plan (NPPWCP) (Beyersbergen et al.
2004), called for renewed attention to determining the
current status of waterbird populations, their distribu-
tions, and conservation needs. The NPPWCP em-
phasized that information on the breeding distribution
of ibis in the northern prairie and parkland region (por-
tions of Alberta, Saskatchewan, Manitoba, Montana,
North Dakota, South Dakota, Minnesota, Iowa, and
Nebraska) is needed to serve as a baseline for future
monitoring needs (Beyersbergen et al. 2004). In re-
sponse to these information needs, we examined the
historical and current distribution of the ibis in North
Dakota and summarized first sightings and nesting
records for the provinces and states composing the
northern prairie and parkland region. We placed spe-
cial emphasis on North Dakota, a state for which there
is scant published information concerning the current
status of this species.
Lokemoen (1979) and Schmidt (1980a) summarized
the first occurrences and nest records of White-faced
46
2007
Ibis in North Dakota. Sabo (1992) described the status,
range, first occurrences, and first nesting observations
for some Great Plains states and Canadian provinces.
Goossen et al. (1995) provided a detailed list of obser-
vations and breeding records for the species through-
out Canada. Jorgenson and Dinsmore (2005) provided
an update of breeding records throughout the Great
Plains. In this paper, we summarize the reported occur-
rences of White-faced Ibis in North Dakota up to 2002,
and we report nest records through 2003, beyond which
time the U.S. Fish and Wildlife Service (USFWS) no
longer quantified colony and nest numbers. We discuss
factors influencing the species’ expansion into the state
and provide data on the species’ breeding habitat. We
also explain how the pattern of occurrence of the spe-
cies in North Dakota compares to the general pattern
throughout the rest of the northern prairie and park-
land region.
Methods
Records of ibis occurrences in North Dakota were
summarized from published reports, including those in
American Birds, Audubon Field Notes, National Audu-
bon Society Field Notes, and North American Birds,
and unpublished reports filed with the North Dakota
Birding Society (REM, unpublished data). We included
data from the North American Breeding Bird Survey
(BBS; 1966-2002; Sauer et al. 2005*), the National
Audubon Society’s Christmas Bird Count (NAS 2007*),
and banding records from the U.S. Geological Survey
Bird Banding Laboratory (USGS 2007*).
We defined a record as a museum specimen or a
reported occurrence of an individual, group of individ-
uals, or a colony. Records were summarized by date
(year, month, and day if known) and location (to coun-
ty level at a minimum). Nest records were defined as
the presence of nests, eggs, dependent young, or adults
exhibiting breeding behavior (e.g., carrying nesting
material). Observations from 15 May to 31 August
were considered breeding-season records, and obser-
vations from 1 September to 14 May were considered
nonbreeding-season records, based on authors’ obser-
vations of ibis behavior and phenological information
as described by Ryder and Manry (1994). Occurrences
of ibises at the same location over a range of days to
months were counted as a single observation for pur-
poses of annual summaries. To summarize data for the
breeding and nonbreeding seasons, reports of the spe-
cies at the same location within both seasons were
counted as separate records, unless information indi-
cated that individuals had nested at the location and
were still present during the nonbreeding season.
We summarized information on use of wetlands for
nesting from one published nest record and from three
studies that measured habitat attributes of nesting ibis
in North Dakota. Schmidt (1980a) provided a brief
description of the habitat attributes of a wetland that
supported several flightless young in 1978 and of one
SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 47
nest in southern North Dakota in 1979. During June and
July 2000, Brice (2003) documented a multi-species
colony that included ibis nests at Lake Alice National
Wildlife Refuge (NWR; 48°19'54"N, 99°06'38"W),
western Ramsey County, in northeastern North Dako-
ta. Colony size was defined as the area occupied by
nests of all colonially nesting species within the colony
boundary, not just by ibis nests. Within the colony
boundary, number of ibis nests, dominant vegetation,
and presence of other bird species nesting within the
colony were recorded. Two transects were placed
lengthwise through the colony. The percent of water
surface that was not covered or obstructed by stand-
ing or horizontal residual vegetation (1.e., percent open
water), residual stem density, and water depth were
measured within 86 1-m? sampling frames placed at
10-m intervals along the two transects (Table 1). Percent
open water within the colony was recorded by visual
estimation. Water depth was recorded using an approx-
imately 3-cm diameter closet rod marked in |-dm inter-
vals. In 2001, this same colony was monitored for the
presence of breeding ibises by USFWS personnel
(Phelps and Meeks 2001*). Percentages of colony area
composed of residual emergent vegetation and open
water were visually estimated. During June and July
2001 and 2002, the USFWS (2001*, 2002*) document-
ed ibis nesting activity on USFWS lands in Burleigh,
Emmons, and Kidder counties in south-central North
Dakota. The number of nests, the number of colonies,
the dominant vegetation within colonies, and the pres-
ence of other nesting bird species were recorded on one
NWR and one waterfowl! production area (WPA). The
ratio of live-to-residual emergent vegetation within
colonies was visually estimated. In 2001, water depth
at nests was measured using a meter stick. During June
2003, Knutsen (2004*) conducted a colonial-nesting
waterbird inventory on 183 wetlands located on NWRs
and WPAs in the Long Lake Wetland Management
District (Knutsen 2004*). Wetlands were divided into
three strata (high, moderate, and low) related to their
likelihood of supporting a waterbird colony, based on
habitat conditions in 2003, as well as the size and per-
manence of the wetlands (sensu Stewart and Kantrud
1971). Number of nests, number of colonies, dominant
vegetation, and the presence of other colonial bird
species nesting within colonies were recorded. The ratio
of live-to-residual emergent vegetation within colonies
and the percent of the colony’s area that was composed
of residual emergent vegetation and open water were
visually estimated. Water depth was not measured in
2003. Vegetation density was classified into one of
three categories: dense (heavy, thick stands of emer-
gent vegetation with few visible openings), moderate
(contiguous stands of emergent vegetation with numer-
ous small openings), or sparse (large “canoe-sized™
openings within the emergent vegetation of open-water
floating mat colonies).
THE CANADIAN FIELD-NATURALIST Vol. 121
Source
Schmidt (1980a)
Phelps and Meeks (2001*)
Knutsen (2004*)
Schmidt (1980a)
Brice (2003)
USFWS (2002*)
Knutsen (2004*)
USFWS (2001*)
Water Other
Yes
57-77 Yes
Yes
Yes
Yes
Yes
depth species
(cm) present
80
80
Ratio of
live-to-dead
vegetation
1:1 at 1 colony,
more live than
1:1 at the smaller
colony, more live than
dead at larger colony
9:1
dead at 3 colonies
% %
open emergent
water vegetation
20-50 30-50
73
Dominant
vegetation
Scirpus sp.
Typha spp.,
Schoenoplectus sp.
Typha spp.
Typha sp.
Typha spp.
Typha spp.
Phragmites australis,
Typha spp.,
Schoenoplectus sp.
Phragmites australis, 15
Typha spp.,
Schoenoplectus sp.
ibis nests
or pairs
Number of
2-4 nests in 1 colony
0.1-1.2 23 nests in 4 colonies
31 pairs in 3 colonies
11 nests in 2 colonies
7 nests in | colony
8 nests in | colony
1 nest
8 nests
Colony
area
(ha)!
10-27
2.8, 6.0
6.5
1]
Wetland
size (ha)
2
380
4913
4913
345
345
1978
2001
1979
2000
2001
2002
2003
Year
'Colony area refers to the area occupied. by nests of all colonially nesting species within the colony, not just to ibis nests.
The wetland size for the 1978 ibis observation includes the entire wetland basin size for Long Lake. The wetland size in 2001 and 2003 reflects the size of an impoundment that did
Long Lake NWR, Burleigh Co
not exist in 1978.
Kraft Slough, Sargent Co.
Lake Alice NWR, Ramsey Co.
location
TABLE |. White-faced Ibis colonies in North Dakota for which wetland and colony sizes, vegetation characteristics, and water depths were recorded.
Colony
(Dewald Slough), Kidder Co.
Kleppe Lang WPA
Results
Ibis in North Dakota
The only reported observation of ibis in North Dako-
ta in the nineteenth century was of a specimen that
was collected in 1882 and deposited in the University
of Michigan’s Museum of Zoology (Wood 1923).
Wood (1923) also reported the first two occurrences
in the twentieth century, including single individuals
in Morton County in 1912 and in Slope County in
May 1920. The fourth report of an ibis occurred over
40 years later, on 5 June 1961, in Stutsman County
(fide REM). This observation marked the beginning
of the species’ documented range expansion into North
Dakota (Figure 1). Observations of ibis increased mark-
edly between the 1960s and 1990s. Between 1961 and
2002, the species has been observed nearly annually,
occurring in all but nine of the 42 years.
From 1882 to 2002, there were 145 reports of one
or more White-faced Ibis in North Dakota, including
93 reports during the breeding season, 49 during the
nonbreeding season, and three for which the season of
occurrence was not reported. Of the 107 occurrences
of the species during the twentieth century, 71 occurred
during the breeding season, 34 during the nonbreeding
season, and the season of occurrence for two early
records was unknown (Figure 1). During the twenti-
eth century, no occurrences were reported from Janu-
ary through March, and two occurrences each were
reported in November and December (Figure 2). The
earliest observation within a year was 14 April, which
occurred in 2002 at Long Lake NWR (46°43'59"N,
100°05'43"W) in Burleigh County (P. C. Van Ningen,
U.S. Fish and Wildlife Service, Moffit, North Dakota,
personal communication). The latest observation with-
in a year was 17 December 1999 in Sargent County
(D. Kinzler, Oakes, North Dakota, personal commu-
nication). The largest group of birds observed during
the breeding season was about 400 individuals on 11
August 2001 in Sargent County; this group was un-
doubtedly a post-breeding flock (Martin 2002). The
largest group of birds observed during the nonbreeding
season was over 50 individuals in September 2000 in
Kidder County by H. F. Duebbert (Martin 2001).
From 1912 to 2002, ibises have been sighted in 28
(53%) of the 53 counties in North Dakota (Figure 3).
Observations first occurred west of the Missouri River.
The two observations in the early part of the twentieth
century both occurred in the western part of the state
(Wood 1923). Of the eight occurrences in the 1960s,
all were reported from southern counties, but only one
occurred west of the Missouri River, in Slope County
(G. B. Berkey, personal communication). Most obser-
vations after the 1960s were east and north of the
Missouri River. The first occurrences in the northern
half of the state occurred in 1973 at Upper Souris NWR
(48°30'11"N, 101°31'50"W), Renville County, by I. O.
Rostad (Houston 1974) and in 1974 at J. Clark Salyer
NWR (48°40'52"N, 100°47'23"W), McHenry County,
2007
70 ———
() Unknown Season
Nonbreeding Season
@ Breeding Season
Number of observations
1901-
1910
1911-
1920
1921-
1930
1931-
1940
194]-
1950
SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 49
1951-
1960
1961-
1970
1971-
1980
1981-
1990
1991-
2000
FicureE 1. Records of White-faced Ibis occurrences in North Dakota during the twentieth century, by decade.
by R. C. Fields (Houston and Houston 1974). The first
sightings in five counties (Bottineau, Foster, Mercer,
Pierce, and Ward) occurred recently, within 2001-2002.
To date, there are 21 records of nesting activity in
North Dakota (Table 2). The first nest was reported by
R. A. Schmidt and R. N. Randall in 1978 (Serr 1978:
Schmidt 1980a). From 1978 to 2003, nests have been
reported in ten of the 26 years, with the majority occur-
ring between 2000 and 2003. Nests or nesting activity
have been reported in each of the four recent years
from 2000 to 2003. The number of nests discovered
within an individual wetland varied from one to 31.
All 21 nesting observations were recorded in June, July,
or August. Observations in June indicated that nest-
ing had been initiated in May. By August, young or
fledglings were observed. Nesting ibises have been
observed in eight of the 53 counties (Table 2, Figure 3).
Most nesting activity was recorded in Burleigh and
Kidder counties (Table 2). Of the ten known wetlands
that have contained nesting ibises, six are federally
owned; two involve a mixture of federal, state, and/or
private ownerships; and two are privately owned.
The ibis has been recorded on a Breeding Bird Sur-
vey route in North Dakota in only one year since the
survey's inception in 1966. On 10 June 1999, three
individual ibis were reported by H. C. Talkington
(Mandan, North Dakota, personal communication) on
his BBS route south of Tappen in Kidder County.
According to banding records, three flightless young
were banded in late July 1979, in western Sargent Coun-
ty (K. Klimkiewicz, U.S. Geological Survey, Laurel,
Maryland, personal communication). No ibises have
been recorded in North Dakota on a Christmas Bird
Count, which have been conducted in the state since
the early 1900s.
In North Dakota, ibises have been found breeding
only in large (>300 ha) semipermanent or permanent
wetlands. Colonies, all of which were mixed-species
colonies, were located in patches of emergent vegeta-
tion dominated by cattails and bulrushes and ranged in
areal extent from 0.1 to 27 ha (Table 1). Ibises appeared
to prefer a fairly equal proportion of open water and
emergent vegetation within a wetland basin, with half
or more of the vegetation composed of live rather than
residual stems. Water depths varied from 60 to 80 cm.
Schmidt (1980a) reported that one ibis nest was built
just above the water level in residual cattails inter-
spersed with open water and was located 600 m from
shore and 20 m from the central open water area of the
wetland basin. Flightless ibises were located in emer-
gent vegetation about 700 m from shore and 35 m from
open water. Brice (2003) reported that the mean densi-
ty of residual cattail stems within an ibis colony was
44 stems/m?. Knutsen (2004*) found that two of 40
waterbird colonies contained ibis nests. Both of these
ibis colonies were located on wetlands categorized as
having a high probability of supporting a waterbird
colony. One of the colonies was located on an im-
poundment of Long Lake that contained ibises in 2001,
and the other was located on a WPA that contained
50
THE CANADIAN FIELD-NATURALIST
Vol. 121 |
nN oo
Sp
i) tN i) tO Ne
i) &
o
Number of observations
a
ie
Jan Feb Mar
18 +
14 +
12 +
10 +
3h
6
4
2
0 r T T T r 53 T r
Apr May Jun Jul Aug Sep
Oct Nov Dec
Month
FIGURE 2. Records of White-faced Ibis occurrences in North Dakota during the twentieth century, by month. An observation)
was counted once, regardless of whether the observation was of one or of multiple individuals.
ibises in 2002 (Table 1). Vegetation density within the
colony in the impoundment was categorized as mod-
erate, whereas vegetation density within the colony
in the WPA was categorized as dense. Several other
waterbird species occurred at colonies with ibises
(Table 1). These species included Ruddy Duck (Oxyura
Jamaicensis), Horned Grebe (Podiceps auritus), Eared
Grebe (Podiceps nigricollis), Western Grebe (Aech-
mophorus occidentalis), Clark’s Grebe (Aechmophorus
clarkii), Snowy Egret (Egretta thula), Cattle Egret
(Bubulcus ibis), Black-crowned Night-Heron (Nycti-
corax nycticorax), American Coot (Fulica americana),
Franklin’s Gull (Larus pipixcan), Forster’s Tern (Sterna
forsteri), and Black Tern (Chlidonias niger) (Schmidt
1980a; Phelps and Meeks 2001*; USFWS 2001*,
2002*; Brice 2003; Knutsen 2004*).
Ibis expansion in the northern prairie and
parkland region
There appears to be a consistent pattern throughout
the northern prairie and parkland region concerning the
chronological progression from the first occurrences
to the establishment of breeding colonies (Table 3).
From our examination of regional patterns of ibis dis-
tribution, we determined that the species rarely visited
the northern prairie and parkland region in the late
nineteenth and early twentieth centuries and did not
develop breeding populations until the late twentieth
century. North Dakota, Iowa, and Minnesota reported
ibis occurrences as early as the late nineteenth century
(Peabody 1896; Lokemoen 1979; Schmidt 1980a; Jans-
sen 1987; Kent and Dinsmore 1996), and Nebraska and
Alberta reported occurrences during the early twenti-
eth century (Kondla et al. 1973; Goossen et al. 1995;
Mollhoff 2001). Montana, South Dakota, Manitoba,
and Saskatchewan did not report ibis occurrences until
the mid-to-late twentieth century (Skaar 1969; Serr
1975; DuBois 1989; Goossen et al. 1995; Tallman et al.
2002). The first nests in the region were reported in’
1894 in Minnesota (Peabody 1896) and in 1916 in
Nebraska (Mollhoff 2001), but other states and prov-
inces did not report nesting ibis until the mid-to-late
twentieth century. Two nest records have been report-
ed for Saskatchewan, both of which occurred in 2000
(B. Hepworth, Ducks Unlimited Canada, Regina, Sas-
katchewan, personal communication; P. Taylor, Cana-
dian Wildlife Service, Environment Canada, Saska-
toon, Saskatchewan, personal communication). Bazin
and Artuso (2006) documented the first nest record for
Manitoba in 2005. Jorgenson and Dinsmore (2005)
summarized recent nesting records for Alberta and!
some of the states in the northern prairie and parkland
region. An additional nest record in Alberta was report-
ed by Flockhart (2001).
Discussion
The number of occurrences of ibises in North Dako-
ta reached the point that the North Dakota Birding
Society stopped recording new nonbreeding observa-
tions after about 2002 (REM, unpublished data). Nest-
ing ibises are becoming increasingly common, with
multiple reports of nests every year from 2000 to 2003
(Table 2). Spring arrival in North Dakota typically
begins in mid-to-late April, and birds begin departing
in September (Figure 2). Few birds remain in the state
in November and December, and no records exist for
January through March. Nesting occurs from mid-May |
to late August (Table 2). The ibis appears to nest colo- |
nially with other waterbird species as has been report-
2007
ii ae
Hettinger
SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 5]
FiGURE 3. Records of White-faced Ibis occurrences in North Dakota during the twentieth century and 2001-2002, by county.
Asterisks (*) denote counties in which nesting activity has been observed during the twentieth century and 2001-2003.
ed by other researchers in the northern prairie and park-
land region (Peabody 1896; Dinsmore and Dinsmore
1986; Mollhoff 2001). Based on our findings, we be-
lieve that the status of ibis in North Dakota is that of a
fairly common migrant and a locally uncommon breed-
er east of the Missouri River and a casual migrant west
of the Missouri River.
Of the ten known wetlands that have contained nest-
ing ibis, most are under federal management. These
wetlands are protected from drainage and tillage; there-
fore, thick stands of cattails and other tall wetland veg-
etation are able to become established in some years
(Table 1). These wetlands also may experience higher
levels of human visitation than privately owned wet-
lands. More human visits, either by federal employees
conducting biological surveys or by bird watchers and
nature enthusiasts visiting the NWRs, WPAs, and other
federal properties, could explain the greater number
of reports of nesting activity on federally managed
properties.
Because ibises were present as vagrants in some parts
of the northern prairie and parkland region in the late
nineteenth century, several authors have speculated that
the presence of breeding colonies in the twentieth cen-
tury was not so much an expansion of the species’ range
as it was the reclamation of portions of the species’ for-
mer range (Ryder and Manry 1994; Dinsmore et al.
1984). Goossen et al. (1995) advanced several explana-
tions for the expansion of breeding ibises into Canada,
which included population growth; loss of habitat to
draining, flooding, or drought in other portions of the
Species’ breeding range; dispersal; and an increase in
Suitable habitat due to the advent of managed or re-
stored wetlands and irrigation. The virtual absence of
sightings from the late nineteenth century through the
early twentieth century in the northern prairie and park-
land region (Table 3) coincided with the withdrawal of
ibises from inland areas in Texas, suggesting to Jor-
genson and Dinsmore (2005) that sightings in northern
regions in the late nineteenth century and first half of
the twentieth century were linked to fluctuations in the
Gulf Coast populations. They examined the historical
status of the species from Alberta to Texas and sug-
gested that the current range expansion appears to be
unprecedented. They contended that the idea that ibis-
es occupied the northern Great Plains in the nineteenth
century was based on only three nesting records that
occurred at a time when extralimital breeding of the
species was being documented elsewhere in the conti-
nent. They suggested that the development of breeding
populations of ibises was linked to population fluctu-
ations in the species’ core breeding areas of Texas,
Louisiana, and, more likely in modern times, to fluctu-
ations occurring in the Intermountain West, the region
of North America lying between the Rocky Moun-
tains to the east and the Cascades and Sierra Nevada
to the west.
Drawing upon knowledge of habitat, climate, and
socioeconomic conditions in the northern Great Plains,
using North Dakota as an example, we suggest fur-
ther explanations for the pattern of ibis observations.
Those explanations include habitat perturbations and
habitat loss in other parts of the ibis’ range, climate and
precipitation patterns in the northern prairie and park-
land region that impacted habitat conditions, and so-
cioeconomic conditions. We concur with Jorgenson
Vol. 121
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and Dinsmore’s (2005) suggestion that ibis occurrences
in the northern Great Plains during the late nineteenth
century and early twentieth century may have been due
to extralimital wandering as was occurring in other parts
of the U.S. at the same time, combined with the species’
predilection for post-breeding dispersal north of its
nesting grounds (Ryder 1967; Ryder and Manry 1994).
No ibises were reported in North Dakota between
1920 and 1961 (Table 3). Plausible explanations include
a scarcity of suitable nesting habitat and a low level of
human engagement in nature-based activities. Potential
suitable wetland habitat for ibises deteriorated with the
onset of drought that characterized the 1930s, the Dust
Bowl years (Robinson 1966). Given the duration and
intensity of that drought, it is highly likely that suitable
wetland habitat was largely nonexistent in the 1930s.
The widespread drought of the 1930s, combined
with economic collapse during the Great Depression,
resulted in a human exodus from North Dakota (Rob-
inson 1966). The human population in North Dakota
reached the highest levels to date, according to the 1930
census, and experienced a 9% decline between 1930
and 1950 (USCB 2005), with much of the decline occur-
ring in rural areas. The human population continues to
fluctuate and recovered somewhat from the 1950s, but
never has reached the levels experienced prior to the
1930s. During the 1940s, fewer people lived in North
Dakota than in the 1920s or 1930s (Kazeck 1956), and
some people that remained were called to war or were
engaged in war-related activities. During the war, am-
munition was rationed and hunting activities were sub-
sequently curtailed. Overall, then, there likely were
fewer people afield and thus there were fewer chances
of people encountering ibises during this time.
Also during the 1940s, moisture conditions, and thus
potential ibis habitat, began to improve, with the high-
est annual precipitation at Bismarck between 1900
and 2000 occurring in 1941 with 79.1 cm (NCDC
2004*). Concomitant with improved moisture condi-
tions was the invasion into North Dakota of an intro-
duced species of European cattail, the Narrowleaf Cat-
tail (Typha angustifolia), which was first collected in
North Dakota in 1942 at Long Lake NWR (Kantrud
1992). Prior to that time, there were few cattail-dom-
inated wetlands in North Dakota, and the few stands
that did exist were in boggy areas around freshwater
lakes (Kantrud 1992). Narrowleaf Cattail was restrict-
ed largely to the southeastern portion of North Dakota
in the early 1960s and spread to many wetlands in
central North Dakota by the 1970s (Kantrud 1992).
However, the invasive species hybridized with the native
Broadleaf Cattail (Typha latifolia), and the resulting
hybrid (7ypha xglauca) spread rapidly throughout
the prairie pothole region of North Dakota during the
1950s, becoming the most abundant large hydrophyte
in the state (Kantrud 1992). Wetlands dominated by the
hybrid cattail potentially provided prime breeding habi-
tat for ibises. Also, despite the decrease in the number
TABLE 3. Pattern for the northern prairie and parkland region, USA and Canada, that exhibits the lapse between time of first White-faced Ibis observations, time of consistent (>5
observations in a given decade) observations, and year of first nest record.
Consistent sightings
Early sightings
Mid-to-late
1900s
Mid-to-late Late Early
1900s
Early
Late
Sources
First nest
1900s
1800s
1900s
1912
1800s
1882
189]
Lokemoen 1979; Schmidt 1980a; this paper
SHAFFER ET AL.; WHITE-FACED IBIS IN NORTHERN PRAIRIE
Dinsmore and Dinsmore 1986; Kent and Dinsmore 1996
2 AND PARKI
_
/
Peabody 1896; Janssen 1987
Mollhotf 2001
Skaar 1969; DuBois 1989
+
Tallman et al. 200
Kondla et al. 1973; Goossen et al. 1995
SDOU 1991;
1960s
North Dakota
lowd
Min
1980s
=
1960s
1910
1894-1895
nesota
983-1984
1960s
1970s
1970s
AND
006
Goossen et al. 1995; personal communications: B. Hepworth’;
R. Luterbach?:
5
O05
O00
5
Serr 1975; Goossen et al. 1995; Bazin and Artuso
>
1970s
1916
=
h Dakota
=
Sou
194]
Manitoba
Saskatchewan
P. Taylor’.
*P. Taylor, Canadian
*R. Luterbach, Regina, Saskatchewan, personal communication
'B. Hepworth, Ducks Unlimited Canada, Regina, Saskatchewan, personal communication.
Wildlife Service, Saskatoon, Saskatchewan, personal communication.
53
54 THE CANADIAN FIELD-NATURALIST
of temporarily flooded wetlands due to drainage, the
number of permanent wetlands in the form of basin
wetlands and riverine impoundments increased in the
state (Lokemoen 1979).
Kantrud (1992) suggested that prairie wildfires and
grazing by large ungulates, two major ecological forces
during presettlement times, were responsible for the
restricted distribution of Broadleaf Cattail prior to the
invasion of Narrowleaf Cattail. Many semipermanent
wetlands in eastern North Dakota, eastern South Dako-
ta, and western Minnesota that were grazed and that
were dominated by semi-open stands of Hardstem Bul-
rush (Schoenoplectus acutus) presently are not grazed
and are dominated by dense stands of cattail. Gary
Krapu (U.S. Geological Survey, Jamestown, North Da-
kota, personal communication) recalls that, prior to
the mid-1950s in North Dakota, most wetlands expe-
rienced grazing pressure by livestock or were hayed.
In the late 1950s and 1960s, however, many farmers
began to focus exclusively on small-grain production,
phasing out livestock production. Livestock were ben-
eficial in stopping the growth of dense stands of cat-
tails by eating young, green cattail shoots. With the loss
of grazing pressure, Narrowleaf Cattail more easily
became established in wetlands. The decline in farm
operations that raised livestock was hastened by the Soil
Bank Program in the 1950s, which provided farmers
with annual payments under 5-to-10 year contracts to
idle their cropland by planting to perennial cover. This
program indirectly also encouraged the idling of wet-
lands as farmers reduced the number of livestock that
they produced.
During the 1950s and 1960s in southern portions of
the species’ breeding range, ibis breeding habitat was
being lost due to human encroachment or drought.
Breeding colonies of ibises disappeared or decreased
in size due to habitat loss or pesticide contamination in
Oregon, California, and Nevada (Ryder 1967; Ryder
and Manry 1994). This decline coincided with increased
sightings in Minnesota, North Dakota, South Dakota,
and Montana (Table 3). It is plausible that this highly
nomadic species sought and found better breeding con-
ditions in the northern prairie and parkland region than
in its more traditional breeding range.
Banding studies during the 1960s found that shoot-
ing was the major cause of reported deaths for this
species, with many bands recovered from Mexico
(DuBois 1989). The species was a gamebird until the
early twentieth century, when international treaties con-
ferred protection from hunting or harassment. Loke-
moen (1979) postulated that ibis population increases
might have been due in part to federal protection. Al-
though federal protection of migratory birds began in
1918 with the signing of the Migratory Bird Treaty
(Legal Information Institute 2005*), resources were
not available for widespread federal enforcement in
North Dakota until the 1940s (G. Krapu, personal com-
munication).
Vol. 121
After the late 1960s, pesticide contamination and
continued habitat loss occurred in southern states, coin-
ciding with more frequent ibis sightings and first nest
records in several northern states. King et al. (1980)
monitored eggshell thickness, pollutant residues, and
the population status of the species in Texas from 1969
through 1976. They found that nesting populations de-
clined by 42%, associated with low reproductive suc-
cess caused by DDE-induced shell thinning and diel-
drin mortality. Taylor et al. (1989) reported that the
ibis population doubled in the late 1970s in Utah and
also increased in Oregon. They attributed these increas-
es to flooding in the Great Basin during the early 1980s,
recovery from reduced productivity caused by DDT
in the 1970s, and/or natural fluctuations in ibis popu-
lations.
Other Cicontiformes species also increased in oc-
currences and expanded their distribution during the
same time period in North Dakota and other areas of
the northern prairie and parkland region (Jones and
Malcolm 1978; Lokemoen 1979; Schmidt 1979a,b,
1980b; Lambeth 1980). The first Cattle Egret in North
Dakota was reported in 1971 (Lokemoen 1979), and
the first nesting colony was discovered in 1976 among
nests of Black-crowned Night-Herons and Little Blue
Herons (Egretta caerulea) (Jones and Malcolm 1978).
More Cattle Egret colonies were discovered in the en-
suing years (Schmidt 1980b). It is possible that the
presence of the colonies of Cattle Egret and other colo-
nial-nesting birds provided the stimulus that prompt-
ed ibis to establish colonies.
More recently, the increase in White-faced Ibis ob-
servations in North Dakota has coincided with the wet
conditions experienced in the northern prairie and park-
land region from 1993 to the present (NCDC 2004*).
The current wet cycle may be the wettest period dur-
ing the past 130 years (Winter and Rosenberry 1998).
These wet conditions may be contributing to suitable
habitat conditions for the ibis. Johnson (1994) docu-
mented large-scale pioneering expansion of nesting dis-
tributions of several species of western North American
birds over the last three decades. Although Johnson
(1994) did invoke climatic warming as a possible expla-
nation, he believed that it was neither the sole expla-
nation nor even the primary cause. Rather, he hypothe-
sized that many of the species were responding to a
decades-long increase in summer rainfall in regions
beyond their former ranges, based on regional climatic
trends. Summer moisture, perhaps coupled with a high-
er mean temperature, encouraged avian range expan-
sions. Detailed correlations of local climatic data with
temperature and moisture requirements of individual
species would be necessary to establish causality (John-
son 1994),
We believe that the combination of adequate rain-
fall and invasion of Narrowleaf Cattail in the 1940s in
North Dakota, the rapid establishment of Narrowleaf
Cattail and its hybridization with the Broadleaf Cat-
2007
tail during the ensuing decades, the general expansion
of other cicontiform species, the conversion from ranch-
ing to small-grain production, wetland loss and poor
ibis reproduction in the southern portions of the ibis
breeding range, growing ibis populations in the Inter-
mountain West region, wet conditions suitable for colo-
nial waterbird establishment, and climatic changes have
contributed to the growth of ibis populations in North
Dakota. We suspect that the above factors also may
have played a role in population changes elsewhere
in the northern prairie and parkland region. Ibises are
considered rare, localized breeders in the northern
prairie and parkland region (Beyersbergen et al. 2004)
and, more specifically, are considered locally uncom-
mon in North Dakota (this paper), rare to locally un-
common in South Dakota (Sabo 1992), and a rare sum-
mer resident in Canada (Goossen et al. 1995).
The future status of ibis within the northern prairie
and parkland region likely will depend upon precipi-
tation patterns. With continuing years of adequate rain-
fall, wetlands that harbored colonies should continue
to provide suitable habitat. There are no indications that
the spread of monotypic stands of cattail within wet-
lands will abate, which should only aid in the spread of
ibis into new breeding areas. Already, breeding sites in
North Dakota that once contained only a few birds and
nests now contain larger groups of nesting birds. With
an onset of a prolonged drought, the quality of nest-
ing habitat could decline in some wetlands, but water
levels in other wetlands that had been inundated to
the degree that they no longer supported emergent veg-
etation would recede and again allow the growth of
deep-marsh vegetation that could harbor new nesting
colonies. Even in a prolonged drought, colonies could
persist at managed wetlands, such as those maintained
within certain NWRs. From these colonies, ibis could
disperse when precipitation levels again rise at previ-
ously dry wetlands.
Acknowledgments
We thank the many individuals who have con-
tributed records of White-faced Ibis in North Dakota.
We thank Richard Crawford, Todd Grant, Dave Grif-
fiths, Craig Hultburg, Lawrence Igl, Will Meeks, and
Mike Norton for providing detailed information on
unpublished sightings. Will Meeks of the U.S. Fish
and Wildlife Service procured funding and equipment
for Joel Brice’s graduate field research, and Richard
Crawford served as his graduate advisor. Kathy Klim-
kiewicz of the Bird Banding Laboratory provided infor-
mation on ibis banding records. Special thanks to Gary
Krapu for sharing his knowledge and experiences of
North Dakota birdlife and to him and Lawrence Igl for
prompting discussions on factors relating to ibis pop-
ulation increases. Thank you to Rachel Bush for con-
ducting library searches. Early drafts of the manuscript
were reviewed by Steve Dinsmore, Lawrence Ig], Dou-
glas Johnson, Gary Krapu, and Terry Shaffer.
SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 55
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Accepted 7 May 2007
Recent Distribution Records of the Little Brown Bat, Myotis lucifugus,
in Manitoba and Northwestern Ontario
JouN E. Dupois!? and KIMBERLY M. MONSON?
'Natural History Division, The Manitoba Museum, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2 Canada
*Present address: Wildlife and Ecosystem Protection Branch, Manitoba Conservation, Box 24, 200 Saulteaux Crescent, Winnipeg,
Manitoba R3J 3W3
3Geography Department, University of Winnipeg, 515 Portage Avenue, Winnipeg, Manitoba R3B 2E9 Canada
‘Dubois, John E., and Kimberly M. Monson. 2007. Recent distribution records of the Litthe Brown Bat, Myotis lucifueus, in
Manitoba and northwestern Ontario. Canadian Field-Naturalist 121(1): 57-61.
‘Until recently, the distribution of the Little Brown Bat (Myotis lucifugus) in Manitoba and northwestern Ontario was poorly
‘documented. Since 1988, we have been banding and recapturing little browns throughout Manitoba and adjacent Lake of the
Woods region in Ontario. All known hibernacula in the study area are recorded here for the first time, along with time of emer-
gence. Connections between some hibernacula and summer nurseries are verified by band returns, ranging from 37 to 540 km.
We began our study of the Little Brown Bat (Myotis
lucifugus) in 1988. The objectives of the study were:
(1) to map the distribution of Little Brown Bats in
‘Manitoba; (2) to find as many hibernacula and summer
nursery roosts as possible; (3) to learn where bats found
in hibernacula spend the summer; and (4) where bats
found in summer roosts spend the winter. An overriding
objective has been to encourage awareness and conser-
vation of bats among the public and responsible agen-
cies in Manitoba. All work was done under Manitoba
Conservation “Wildlife Scientific Research”, “Ecologi-
cal Reserves Research”, and Ontario Ministry of Natural
‘Resources (Kenora District) permits issued to Dubois.
_ Little previous research had been done on the six
“species of bats known to occur in Manitoba (Banfield
1974). Robert Barclay worked on some species and
made incidental observations of occurrences and migra-
tion dates of little browns at the University of Manitoba
| ar Marsh Field Station from 1981 to 1983 (Barclay
1984). Barclay (1993) stressed “Even for species that
are well studied elsewhere, such as the Little Brown
‘Bat, information about the biology of prairie popula-
‘tions is scarce. More than anything else, it is this lack
‘of information that hinders conservation efforts for
/prairie bats.” This study is meant to aid conservation
| efforts by filling gaps in our knowledge.
A. L. Rand (1948) reported sightings of the Little
Brown Bat in the Flin Flon district. J. Dewey Soper
(1961) reported the first specimens taken at Max Lake,
‘in Turtle Mountain Provincial Park, 20 August 1949.
Little browns were reported from the Delta area (Tam-
-sitt 1962). In 1978 the late Walter Cook of Grand
Rapids showed provincial biologist Bill Koonz a cou-
' ple of caves north of the town that over-wintered bats
_(Koonz, Manitoba Conservation, personal communica-
| tion). Since 1988 systematic searching for caves with
the Speleological Society of Manitoba (SSM) has pro-
‘Key Words: Little Brown Bat, Myotis lucifugus, distribution, Manitoba, northwestern Ontario.
duced more information on the Litthke Brown Bat in
Manitoba. Soliciting the public and various agencies
to report caves, summer bat roosts and sightings has
also been very productive.
Methods
We banded Little Brown Bats to monitor their distri-
bution and seasonal movements. Plastic coloured bands
were used in the first year of banding, 1988. Each cave
or location was identified by a different colour. For more
precise tracking of individual bats and their movements,
numbered aluminium bands (National Band & Tag Co.,
Kentucky), size 2 were used in 1989 and thereafter.
Hibernacula were located with the aid of the consid-
erable efforts of the members of the Speleological Soci-
ety of Manitoba and other volunteers. Starting from
reported hibernacula sites, the authors and other mem-
bers of the SSM used a combination of surficial geology,
topographic maps, and air photos to identify likely areas
for methodical ground searches. Karst features such as
caves, sinkholes and trenches were detected on air pho-
tos in areas where limestone, dolomite and gypsum bed-
rock are near the surface. These areas were then visited
and searched in a grid pattern, using map and compass, and
latterly, hand-held global positioning system (GPS) units.
Information provided by home and cottage owners,
Manitoba Department of Conservation, Ontario Min-
istry of Natural Resources, and other agencies led to
the banding of bats in residential and cottage summer
nursery colonies throughout the province, spilling over
into adjacent northwestern Ontario around the north
end of Lake of the Woods.
Greatest frequency of banding occurred in caves in the
Grand Rapids and Gypsumville areas, and in one large
cave (St. George Cave) north of Hodgson. [See map,
Figure 1] The karst features, where the caves are found
in gypsum and dolomite formations of these areas, are
SV/
58 THE CANADIAN FIELD-NATURALIST
described in detail in the field reports of the Manitoba
Mines Branch for 1988, 1989 and 1990 (e.g., Sweet
et al. 1988) and in McRitchie and Monson (2000).
We used standard techniques to capture and band
the bats (Kunz and Kurta 1988). We captured them in
caves by plucking the torpid bats carefully by hand
Vol. 121
from the cave wall, and placing them in a cloth bag. We
live-trapped Little Brown Bats with a harp trap (Tut-
tle 1974). Once captured, the bats were examined to
determine sex, reproductive status, age, injuries (both
healed and fresh), and, on a cursory basis, external par-
asites present. According to custom (Kunz and Kurta
TABLE |. Hibernacula banding locations in Manitoba and northwestern Ontario for Little Brown Bats.
Location
Cook’s Cave [GR]
Dale’s Cave [GR]
Firecamp Cave [GR]
Fold Cavern [Gyp]
Iguana Crypt [GR]
Longcrawl Cave [Gyp]
Microwave Cave [GR]
Squeaky Cave [GR]
St. George Cave [Hod]
Stormcloud Cave [Gyp]
Richard Lake, Ont. [LW]
Abyss Cave [GR]
Okaw Cave [GR]
[GR] = Grand Rapids area; [Gyp] = Gypsumville area; [Hod] = Hodgson area; [Wh] = Whiteshell area; [LW] = Lake of the Woods
TABLE 2. Summer nursery colony banding locations in Manitoba and northwestern Ontario for Little Brown Bats.
Location
Brereton Lake, Manitoba
Caliper Lake Provincial Park, Ontario
Deleau, Manitoba
French Portage Narrows, Ontario
Grand Rapids, Manitoba
Jackfish Lake, Manitoba
Keewatin, Ontario
Melita, Manitoba
Moosehorn, Manitoba
Pebble Beach, Manitoba
Rush Bay, Lake of the Woods, Ontario
Rushing River Provincial Park, Ontario
Schist Lake, Manitoba
Sherridon, Manitoba
Wasagaming, Manitoba
Wekusko Falls, Manitoba
Latitude Longitude
53°26'10"N 99°32'10" W
53°39'32"N 99°26'19" W
B29 25NN 99°20'50" W
51°51'02"N 98°29'30"W
53°26'14"N 99°30'58" W
51°47'10"N 98°37'00"W
53°34'05"N 99°26'25"W
53°37'42"N 99°20'3 1" W
51°36'07"N 97°24'52"W
51°50'28"N 98°29'40" W
49°50'54"N 94°41'30"W
53°42'55"N 99°25'40"W
53°51'53"N 100°07'12"W
Latitude Longitude
49°54'16"N 95°32'40"W
49°03'42"N 93°54'46"W
49°34'5S2"N 100°34'3 1" W.
49°26'58"N 94°40'14"W
53°12'30"N 99°18'00" W
50°18'05"N 100°02'10"W
49°45'33"N 94°33'20"W
49°16'05"N 100°59'45"W
SS W29EN 98°25'22"W
50°59'34"N 98°26'53"W
49°39'55"N 94°53'28"W
49°41'13"N 94°13'44"W
54°39'45"N 101°48'19"W
55°07'26"N 101°05'12"W
50°39'30"N 99°58'07" W
54°47'22"N 99°58'21"W
TABLE 3. Additional Little Brown Bat locations in Manitoba and northwestern Ontario verified by the authors.
Location
Souris, Manitoba
Reston, Manitoba
Minnedosa, Manitoba
Falcon Lake, Manitoba
Deception Lake, Ontario
Setting Lake, Manitoba
Woodridge, Manitoba
vicinity of Oak Lake, Manitoba
Kerr’s Lake, Manitoba
Pointe du Bois, Manitoba
Otter Falls, Manitoba
Taiga Biological Station, Wallace Lake, Manitoba
Winnipeg, Manitoba
Latitude Longitude
49°37'02"N 100°15'37"W
49°33'27"N 101°05'36"W
50°14'43"N 99°50'34" W
49°41'11"N 95°14'54"W
49°44'27"N 94°50'33" W
55°04'11"N 98°29'46"W
49°17'02"N 96°08'52" W
49°40'17"N 100°37'34"W
50°30'00"N 99°40'57" W
50°18'03"N 95°33'05"W
50°08'48"N 95°48'38"W
51°02'40"N 95°20'40""'W
49°53'04"N 97°08'47"W
DUBOIS AND MONSON: RECORDS OF LITTLE BROWN Bat 59
wr. FA .
a f Dale’s Cave
‘owave
Fire Camp
Cave
Lake
» Winnipeg
Dauphin :
Brandon’ \_~ Winnipeg
>
LY
Turtle Mountain \
Prov. PArk
MINNESOTA
aa NORTH DAKOTA 98° \ 96° Vine ees
Ficure 1. Map of Manitoba showing the portion of the Boreal Ecozone and major settlements. Circles are settlements, triangles
are other localities (see text).
60 THE CANADIAN FIELD-NATURALIST
1988), we banded males on the right forearm and fe-
males on the left. Those bats found to be previously
banded were duly recorded and injuries and/or para-
sites noted. Time in capture and handling was kept to
a minimum to avoid stressing the animals. Banding
locations are summarized in Tables | and 2, below.
Many of the Manitoba caves containing bats were
mapped, the annual maximum and minimum tempera-
tures recorded (average +5°C) and any other vertebrate
or invertebrate inhabitants noted, by us, other SSM
members, and by staff of the Manitoba Museum and
the Manitoba Geological Survey. These details and
more on the caves and surrounding landscapes are thor-
oughly described in McRitchie and Monson (2000).
Results
To date we have banded more than 9000 Little
Brown Bats. J. Dewey Soper (1961) speculated that
Little Brown Bats “probably extend almost to the Hud-
sonian Zone’, now called the Boreal Shield ecozone
(Ecological Stratification Working Group 1995), while
van Zyll de Jong (1985) stated: “It is found from the
Atlantic to the Pacific coast and occurs to the limits
of the boreal forest.”
In addition to those locations where banding has
taken place (Tables 1 and 2), we have verified the pres-
ence of this species throughout southern Manitoba and
at least to the latitude of Taiga Biological Station, Uni-
versity of Manitoba, Wallace Lake (51°01'50"N), east
of Lake Winnipeg (Table 3). On the west side of the
province, Little Brown Bats occur to the latitude of
Sherridon (55°07'26"N), and in north-central parts of
the province to Setting Lake (55°04'11"N) (Table 3)
— locations slightly north and just south of Herb Lake
(reported in van Zyll de Jong 1985, page 70).
Similar to many species across Canada, a combina-
tion of degree-days and night-length likely constrains
their range in the north — see discussions in Holroyd et
al. (1994), Nagorsen and Brigham (1993), Humphries et
al. (2002). A maternity roost of Little Brown Bats was
reported recently at 58°06'36"N on the Alaskan coast
(Parker and Cook 1996), as was one at Squanga Lake,
Yukon at 60°28'40"N (Jung and Slough 2005) .
Sherridon (55°07'26"N) is the site of the northern-
most nursery colony we have found in Manitoba to
date, while Okaw Cave (53°51'53"N) is the northern-
most hibernaculum. Individual Little Brown Bats have
been reported during summers from as far north as
Thompson (55°44'36"N). Little browns have been re-
ported to us as showing up at summer nursery colonies
in central and southern Manitoba as early as the last
week in April, and more often, the first week of May.
However, we routinely find large numbers of a given
hibernaculum still in torpor when entering the caves in
the Grand Rapids area on the Victoria Day long week-
end in May (normally the third weekend of the month,
range 17-28 May).
Band returns from this study have shown little
browns are very loyal to both their summer and winter
Vol. 121
roosts, with very low incidences of transference (>1%).
Recoveries from subsequent, different roosts ranged
from 5 km to 533 km distant from point of original
banding. Seasonal migration distance from hibernac-
ula to summer nursery colony for adult females ranged
from 37 km to 540 km. Details of inter-roost and sea-
sonal movements will be published separately.
We started banding Little Brown Bats around Lake
of the Woods (mainly in the Kenora area) in 1993 to
learn whether those populations use Manitoba caves to
hibernate. Band returns demonstrate that at least some
of those females forming nursery colonies in that region
definitely use central Manitoba caves to over-winter, as
well as one nearby abandoned mine. It has been spec-
ulated for some time that abandoned mines in the Keno-
ra area have served as hibernacula (Nagorsen 1980).
In July 1995, we were directed to an abandoned mine
at Richard Lake, 37 km east of Kenora and, upon re-
turning in late September, found it to contain at least
250 Little Brown Bats, none previously banded. In the
Richard Lake hibernaculum in subsequent years we
recaptured banded bats from nursery colonies on Lake
of the Woods (Keewatin, French Portage Narrows,
Rushing River Provincial Park), as well as bats previ-
ously banded in Manitoba hibernacula as far as 533 km
away (St. George, Iguana Crypt and Microwave caves).
Conservation
This study has verified that at least 11 caves in Mani-
toba and one abandoned mine in northwestern Ontario
serve as hibernacula for Little Brown Bats (Table 1).
Bats of many species are at their most vulnerable at
roost sites, particularly hibernacula (Brigham 1993).
The largest hibernaculum discovered to date in Man-
itoba, by several orders of magnitude, is St. George
Cave, 200 km north of Winnipeg. More than 20 000 lit-
tle browns spend each winter in that cave. This critical
cave and a small area around it was declared an ecolog-
ical reserve by the Province of Manitoba on 15 March
1997, after several years of effort by Willard Anderson,
the authors, and other members of the Speleological
Society of Manitoba. This is the most stringent legal
protection a natural feature can get in this province.
The Province of Manitoba, on 15 January 1996 official-
ly declared six species of bats to be “a wild animal”
under The Wildlife Act (by the Designation of Wild
Animals Regulation MR 3/96), as a first step in their
protection. As of December 1997, Manitoba Depart-
ment of Natural Resources (now Conservation) has
issued guidelines to direct all activities occurring on
Crown land around the known over-wintering caves
(Asmundson and Larche 1996*).
Acknowledgments
We thank our employers, the Manitoba Museum and
the University of Winnipeg, for their support during the
majority of the time period of this study. Additional
funding was granted by the Manitoba Museum Foun-
dation Fund. We were also substantially assisted by the
2007
staff of Manitoba Department of Conservation (Natural
Resource Officers, Operations Branch, particularly
of the Grand Rapids and Gypsumville offices, as well
as Parks and Natural Areas Branch), and Manitoba
Geological Survey, Department of Industry, Trade and
Mines. Thanks to Tolko Industries Ltd. and the hos-
pitable people of Grand Rapids and Gypsumville. Many
family members and friends have worked with us for
many hours. We would like to particularly thank Dale
Brown and David Wright. Other volunteer banders and
helpers over the years include Al Bisset, Melanie
Dubois, Michelle Bell, Liza McClintock, Marie Voth,
Lori Bilecki, Christy McDonald, Marg Rubin, Gloria
Goulet, Cecile Foster, Janis Klapecki, Dennis Peristy,
Kim Ottenbreit, Victoria Shemeliuk, Mike Claussen,
Justin Harrison, Donna Kurt, Rick Nash, Paul Buisson,
Robert Senkiw, Jocelyn Hildebrand, Marianne Setliff,
Chris Caslake, Bob Gill, Darren Keam, Amy Barker,
Lidi Kuiper, Lyndon Kivi, Kevin Campbell, Christine
Abraham, John Fitzmaurice, Lisa Lagerge and Trapper
John Christie. We thank Jim Duncan and Robert Nero
for reviewing an earlier draft of this paper.
Documents Cited (marked * in text)
Asmundson, J. D., and R. A. Larche. 1996. Bat Hibernacula
Management Guidelines. (unpublished manuscript) Wild-
life Branch, Manitoba Department of Natural Resources.
31 pages.
Literature Cited
Banfield, A. W. F. 1974. The mammals of Canada. National
Museum of Natural Sciences. University of Toronto Press.
438 pages.
Barclay, R. M. 1984. Observations on the migration, ecology
and behaviour of bats at Delta Marsh, Manitoba. Canadian
Field-Naturalist 98: 331-336.
Barclay, R. M. 1993. The biology of prairie bats in Proceed-
ings of the Third Prairie Conservation and Endangered
Species Workshop. Edited by G. L. Holroyd, H. L. Dick-
son, M. Regnier, and H. C. Smith. Provincial Museum of
Alberta, Natural History Occasional Paper Number 19.
Brigham, R. M. 1993. The implications of roost sites for the
conservation of Bats in Proceedings of the Third Prairie
Conservation and Endangered Species Workshop. Edited
by G. L. Holroyd, H. L. Dickson, M. Regnier, and H. C.
Smith. Provincial Museum of Alberta, Natural History
Occasional Paper Number 19.
Ecological Stratification Working Group. 1995. A National
Ecological Framework For Canada. Agriculture and Agri-
Food Canada, Research Branch, Centre for Land and Bio-
DUBOIS AND MONSON: RECORDS OF LITTLE BROWN BAT 6)
logical Resources Research and Environment Canada
State of the Environment Directorate, Ecozone Analysis
Branch, Ottawa/Hull.
Holroyd, S. L., R. M. R. Barclay, L. M. Merk, and R. M.
Brigham. 1994. A survey of the bat fauna of the dry interi-
or of British Columbia. Wildlife Working Report Number
WR-63, Wildlife Branch, Ministry of Environment, Lands
& Parks, B.C.
Humphries, M. M., D. W. Thomas, and J. R. Speakman.
2002. Climate-mediated energetic constraints on the dis-
tribution of hibernating mammals. Nature 418: 313-316
Jung, T. S., and B. G. Slough. 2005. Mortality of Little Brown
Bats, Myotis lucifugus, in a rodent trap in the boreal forest
Canadian Field-Naturalist 119: 589-590.
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holding devices. Pages 1-28 in Ecological and behavioral
methods for the study of bats. Edited by T. H. Kunz. Smith-
sonian Institution Press, Washington, D.C. 533 pages.
McRitchie, D. W., and K. M. Monson. 2000. Caves and
Karst of Manitoba’s Interlake Region. Speleological Soci-
ety of Manitoba, Second Edition. Winnipeg. 181 pages.
Nagorsen, D. W. 1980. Records of hibernating Big Brown
Bats (Eptesicus fuscus) and Little Brown Bats (Myotis
lucifugus) in northwestern Ontario. Canadian Field-Nat-
uralist 94: 83-85.
Nagorsen, D. W., and R. M. Brigham. 1993. The Bats of
British Columbia. Volume 1: The Mammals of British
Columbia. Royal British Columbia Museum Handbook.
UBC Press, Vancouver.
Parker, D. I., and J. A. Cook. 1996. Keen’s Long-eared Bat,
Myotis keenii, confirmed in southeast Alaska. Canadian
Field-Naturalist 10: 611-614.
Rand, A. L. 1948. Mr. W. H. Bryenton’s notes on Manitoba
mammals of the Herb Lake-Flin Flon area. Canadian Field-
Naturalist 62: 140-150.
Soper, J. D. 1961. The mammals of Manitoba. Canadian
Field-Naturalist 75: 171-219.
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Investigations in Palaeozoic Carbonates of the Grand
Rapids Uplands and Southern Interlake. Geological Paper
GP88-1, Province of Manitoba, Mines Branch.
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Lake Manitoba, Canada. Canadian Field-Naturalist 76: 71-
76.
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ences, Ottawa, Ontario.
Received 12 August 2003
Accepted 2 October 2007
Immobilization of Elk, Cervus elaphus, with Telezol and Xylazine and |
Reversal with Tolazine or Yohimbine
RICK ROSATTE
Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, DNA Building, 2140
East Bank Drive, Peterborough, Ontario K9J 7B8 Canada; e-mail: rick.rosatte @ ontario.ca.
Rosatte, Rick. 2007. Immobilization of Elk (Cervus elaphus) with telezol and xylazine and reversal with tolazine or yohim-
bine. Canadian Field-Naturalist 121(1): 62-66.
During January 2003 and March 2005, a Telazol®/xylazine mixture was used to immobilize 4 free-ranging Elk in Ontario,
Canada. A dosage of 3.3-3.6 mg/kg of Telazol® and 1.7-2.0 mg/kg of xylazine proved to be effective for the rapid immobi-
lization of Elk. Induction time for those dosages was as short as 3-4 minutes. The advantage of using Telazol® is that only
small volumes (3-4 ml) are needed to immobilize Elk-sized animals. In addition, Tolazine® and yohimbine both proved to
be effective antagonists for xylazine with recovery times of 8 to 15 minutes when administered at dosages of 3.3-3.6 and
0.08-0.14 mg/kg, respectively. The use of oxygen proved to be effective for treatment of hyoxemia in Elk immobilized with
a Telazol®/xylazine mixture. The immobilization procedures and the drug and antagonist dosage information will be useful
to researchers planning to capture free-ranging Elk for activities such as radio-collaring and blood sampling.
Key Words: Cervus elaphus, Elk, antagonist, immobilization, reversal agent, Telazol®, Tolazine, Yohimbine, Ontario.
During 1998 — 2001, 443 Elk (Cervus elaphus), also
known as American Elk, or Wapiti, were transported
from Elk Island National Park (EINP), Alberta, and
released in four areas of Ontario, Canada (Bellhouse
and Rosatte 2005; Rosatte et al. 2007). One of the
release sites, near Bancroft, Ontario (approximately
44°5'N, 77°30'W), received 120 Elk during 2000 and
2001. All Elk were aged and weighed at EINP during
processing, which included ear-tagging and fitting the
Elk with telemetry collars (VHF and GPS) (Rosatte
et al. 2002). The weights of Elk at EINP were used as
guidelines to estimate the weight of Elk in Ontario for
drug dosage calculations.
As ten of the Elk were fitted with GPS collars during
processing at EINP, some of the animals had to be
immobilized to retrieve the collars and download the
data. On 16 January 2003 (-20°C), an 8 year old, free-
ranging adult cow Elk (ear tag 341), previously fitted
with a GPS collar (148.188 Mhz) at EINP, Alberta, was
immobilized with Telazol® (tiletamine hydrochloride
and zolazepam hydrochloride) (Fort Dodge Animal
Health, Fort Dodge, Iowa) and AnaSed (xylazine
hydrochloride) (Vet-A-Mix, Shenandoah, Iowa) near
Bancroft, Ontario, for the purpose of collar removal.
The drug dosage (target dosage was about 3 mg/kg
Telazol® and approximately 1.5 mg/kg of xylazine)
was calculated using an estimated weight for the Elk
of 236 kg — her weight during processing at EINP in
January 2001. The immobilizing drug mixture was pre-
pared by injecting the desired volume and weight of
xylazine into a bottle containing lyophilized Telazol®.
The Telazol®/xylazine mixture was then drawn off with
a syringe and injected into the immobilization dart.
The dart was then plugged with vasoline to prevent the
leakage of drugs from the dart. A Model 193 Pneu-Dart
50 caliber rifle and 4x scope (Pneu-Dart Inc., Williams-
port, Pennsylvania) with a yellow CCI 22 caliber brass
power charge (Omaha Industries, Lewiston, Idaho) |
and a diffuser setting of 4, was used to project a 50 cal-
iber, 4 cc aluminum type C dart with a 3 cm long barbed
needle (Pneu-Dart Inc., Williamsport, Pennsylvania),
that contained the Telazol®/xylazine mixture, into the
hind leg musculature of the Elk. The distance of the
shot was about 35 m. As soon as the animal was in a
lateral recumbent position and there was no evidence ~
of leg or eye reflex, she was moved into a sternal re-
cumbent position and the head was elevated to mini-
mize the chance of bloat and regurgitation of rumen
contents. Ophthalmic ointment was applied to the
eyes of the immobilized Elk to prevent drying. The
respiration rate of the cow Elk was about 18/min at
30 min post-darting. At 36 min post-darting, the res-
piration rate increased to 54/min and breathing was
very shallow. At this point, bloat was evident and the
gum/ tongue area was blue in color (indicative of low
oxygen content in the blood). The animal was admin-
istered oxygen orally as a treatment for hypoxemia,
as well as rolled side to side in an effort to relieve the
bloat. The head was elevated and lowered as well to
stimulate belching. At 50 min post darting, respiration |
rate decreased to 42/min and the tongue/gum area
returned to a pinkish color following the administra- -
tion of oxygen. Hibitane antibacterial veterinary oint- -
ment (1% chlorhexidine acetate) (Ayerst, Guelph, On-
tario) was applied to the dart wound. After the animal
was processed an intramuscular injection of Tolazine
(tolazoline hydrochloride, Lloyd Laboratories, Shenan-
doah, Iowa) was administered in the right leg muscu-
lature to speed recovery. The concentration and dosage
of drugs as well as the induction, down and recovery
times are shown in Table 1. After the animal was ina
standing position, she orally expelled gas for several ©
62
2007 ROSATTE: IMMOBILIZATION OF ELK 63
seconds and the bloat decreased dramatically. The Elk
was monitored for the next few days and no adverse
co
effects were noted. She was still in good health when 3 5 we w.8
observed near Bancroft, Ontario in April 2005. Z Bs gZal4AR==
On 19 February 2004 (-6°C), a 10.5 year old cow £ Bagalezza
Elk (ear tag 360) was immobilized near Bancroft, On- es weak
tario, to remove a GPS collar (149.560 Mhz). The _ =
weight of the animal at EINP during January 2001 was i
234 kg. The animal was darted with a mixture of Tela- i ee ime
zol”/xylazine as described above using a Pneu-Dart 5 z E atl ee a eee
rifle, with a yellow 22 caliber power charge and a dif- = zg “Ss
fuser setting of 3, and a 4 cc dart. The distance of the 5
shot was about 25 m with the dart hitting the right upper z
hind leg musculature. As the drug took effect the Elk 2 ee. a
assumed a recumbent position beginning with the hind 3 5 E = $RHRF
end first (Figure 1). Some bloat was evident so oxygen A AT}
was administered for treatment of hypoxemia and the z
animal was rolled from side to side and the head elevat- @ a
ed to expel gases (Figure 2). Respiration rate increased g PIN Son ae ae Ps
from about 14/min to 70/min when bloating occurred. 2 | = =| ae i >
The gums and tongue remained pink colored through- S =) E =
out the handling period. When processing was com- fe S s
plete, Tolazine was injected intramuscularly to ® Cie ye 3
reverse the effects of xylazine and speed recovery = b Z eee = 5
from the drug. Induction, immobilization and recoy- 5 25a g/9NSS 3 =
ery times are noted in Table 1. || 2 = S
On 11 January 2005 (-17°C ambient temperature), = = = =
a yearling bull Elk, born in Ontario, suspected of being A oe & S = fc
infected with meningeal worm (Parelaphostrongylus 2 2 = fBalooa aes
tenuis), was immobilized with an intramuscular 5 22 El sor (Begs
injection of Telazol® and xylazine hydrochloride for the e er ERES%
purpose of collecting a blood sample. The dosage of z f Z7se
drug was calculated using mean weights of yearling 3 —q |eo wioe 8 oe
bull Elk at EINP. As the Elk had no fear of humans, = pag =l| spose = 2 oe a 25
possibly due to the effects of P. tenuis, the drugs were = Salss S S of Slt
administered intramuscularly in the right upper hind 3 Flare = 2885
limb area via a 5 cc sterile syringe (Burron Medical Z Oba 3 5
Products, Bethlehem, Pennsylvania, USA) and 22 g = eS ete oN 255
(4 cm) needle (Terumo, Belgium) attached to a jab = 26 SRTeonyss =a) =
stick made from a piece of copper tubing. When pro- > 2S Sa SSaalsus 225
cessing was complete, which included radio-collaring 8 AaX S| aaaalo 2 ass
(148.412 Mhz) and ear-tagging (#139) (as reported in 3 a5 252
McIntosh et al. 2007), the Elk was injected intramus- S Ham ig gpergelll eS 2 SUS ew
cularly in the hind limb musculature with Yobine (yo- 2 Sora aa cess = Mess ES
himbine hydrochloride) (Lloyd Laboratories, Shenan- S SeSE/SSssii<2e eee
doah, Iowa, USA) as a reversal agent for xylazine. Drug 5 ES re PaO fa Ee eo
dosages, induction, immobilization and recovery times : =o = = 2 = 5
are presented in Table 1. While immobilized, respira- aa r eSs5ce3
tion rate for the yearling bull Elk was 14-16/min, there i Helunooll SE Ese =
was no evidence of bloat, and the gums and tongue 8 2 S|SASS sR wee a's
remained pink in color. E s SE Ex
On 30 March 2005 (+5°C ambient temperature), a a v. Sgetecs
2% year old bull Elk (born in Ontario) was immobilized 22 Sel RR SS| PS sss gz
with Telazol® and xylazine for the purpose of de-antler- 2S Sosy te Ba otees
ing as the animal had broken the pedicle in the fall of oo S BY oe Be
2004. The drugs were administered as described above alee = = E i: > 3
using a Pneu-Dart rifle, with a green 22 caliber power = z oss S <= Sseacs
charge, a diffuser setting of 5, and a 4cc dart. The dis- agi 2 SSeS 2 53 2 ys
tance of the shot was about 30 m with the dart hitting a 3a Se She a ss
64
tne a be
THE CANADIAN FIELD-NATURALIST
Vol. 121
Ficure |. A GPS collared adult cow Elk (ear-tag 360) exhibiting hind limb paralysis 2 minutes after being darted with a
mixture of Telazol® and Xylazine. (Photo by R. Rosatte).
the right upper hind leg musculature. Respiration rate
was stable at about 12/min during processing (de-
antlered and ear-tagged — #141) and the gums and
tongue remained pink colored with no evidence of
hypoxemia or bloat. A 3 cc intramuscular injection of
Dystosel (Vitamin E — 136 IU/mL and selenium — 3
mg/mL) (Pfizer Canada Inc, London, Ontario) was
administered as a preventative measure against cap-
ture myopathy. When processing was complete, Yobine
was injected intramuscularly to speed recovery from
the effects of xylazine. Induction, immobilization and
recovery times are shown in Table 1.
Discussion
A dosage of 3.3-3.6 mg/kg (body weight) of Tela-
zol® and 1.7-2.0 mg/kg of xylazine proved to be an
effective drug combination for the rapid immobiliza-
tion of four free-ranging Elk in Ontario. Induction
time or the time from darting until the animal was
prone for those dosages was 3-4 minutes. This calcu-
lation did not include the Elk for which induction
time was noted as 10 minutes, as that animal was not
immediately located in the forest. Similarly, Mills-
paugh et al. (1995) reported a mean induction time of
4.6 minutes when Elk were immobilized with 2.5 mg/kg
Telazol® and 0.3 mg/kg xylazine. The advantage of
using a Telazol®/xylazine mixture over a drug combi-
nation such as ketamine hydrochloride (at a commer-
cially available concentration 100 mg/mL)/xylazine is
that a much lower volume of Telazol is required to at-
tain a state of immobilization. For example, only a 4 mL
(800 mg of Telazol® reconstituted in 3.5 mL of xyla-
zine) volume of drug was required to immobilize a
225 kg cow Elk. About 15 mL of a ketamine hydrochlo-
ride/xylazine hydrochloride (100 mg/mL ketamine)
mixture was required to immobilize a bull Elk in On-
tario during 2003. The advantage of a smaller volume
of Telazol®/xylazine mixture is that a much smaller
dart can be used (4 cc compared to a 10 or 15 cc) to
administer the drugs, which means less weight and less
damage to the animal when the dart is administered
via an immobilization gun. A smaller dart (4 cc) has
a better trajectory and is more accurate at longer dis-
tances (> 65 m) than a larger/heavier dart (10-15 cc).
In addition, as the induction time for ketamine in Elk
is generally longer than that for Telazol®, animals tend
to move farther before being completely immobilized
(Golightly and Hofstra 1989; Miller et al. 2004), which
could result in problems locating the animal in heavi-
ly forested areas.
2007
ROSATTE: IMMOBILIZATION OF ELK 65
FIGURE 2. Oxygen being administered to the immobilized cow Elk (ear tag 360) for treatment of hypoxemia (Photo by R
Rosatte).
One drawback with using a Telazol®/xylazine mix-
ture for the immobilization of Elk is the extended re-
covery time which could lead to decreased survival. In
one study, an Elk immobilized with a Telazol®/xylazine
mixture was not administered an antagonist and was
immobilized for 5 hours (Golightly and Hofstra 1989).
As well, there is no known antagonist for the tiletamine
component of Telazol® (Miller et al. 2004). Tolazine
and yohimbine both proved to be effective antagonists
for xylazine with recovery times of 8 to 15 minutes
when administered at dosages of 3.3-3.6 and 0.08-
0.14 mg/kg (body weight), respectively. However, the
key to rapid recovery was administering the antagonist
45-60 minutes (or longer) post -immobilization. This
allowed sufficient time for Telazol® to be metabolized
and diminish side-effects after injection of the rever-
sal agent. Animals may stagger for some time due to
the effects of Telazol® if the antagonist is adminis-
tered too soon after immobilization as the antagonist
only neutralizes the effects of xylazine. Millspaugh et
al. (1995) reported a mean recovery time of 14 min-
utes when yohimbine was administered intravenous-
ly to reverse xylazine administered to Elk at a dose of
0.3 mg/kg (in a mixture with Telazol® at a dose of
2.5 mg/kg). However, a recovery time in that study of
125 min was reported when yohimbine was given intra-
muscularly.
Hypoxemia or deficient oxygenation of the blood
may occur during the immobilization of wild rumi-
nants. This can lead to organ failure and capture myopa-
thy (Read et al. 2001). Bloating occurred in two of the
Elk immobilized in this study despite their being in a
sternal recumbent position. As well, respiration rate was
high in those two Elk {normal respiration rate for elk
is about 13 breaths/min (Hudson and Haigh 2002)}.
Bloat is a result of gas production in the rumen, which
may result in pressure being applied to the diaphragm
(due to the weight of the abdominal viscera on the
diaphragm) with resultant respiratory/circulatory prob-
lems and oxygen depletion in the blood. Bloating in
this study may have been aggravated due to the ani-
mals foraging at feeding stations (with alfalfa) prior
to darting. Oxygen quickly relieved the symptoms of
hypoxemia (blue tongue, lips), and the suggested rate
of oxygen administrated nasally for preventing hypox-
emia was 10 L/min for a period of about 5 minutes
(Read et al. 2001).
The four immobilized Elk were later (1 month to 2
years) located by radio-telemetry and had recovered
fully, with no observable side effects from immobiliza-
66 THE CANADIAN FIELD-NATURALIST
tion or handling. Capture procedures, drugs and antag-
onist data outlined above will be useful to researchers
planning to capture free-ranging Elk. For “off-label”
use of the above drugs on wild animals such as Elk,
an “Emergency Drug Release Authorization” must be
approved by the Veterinary Drugs Directorate, Health
Canada, and animals immobilized with agents such
as Telazol® are not to be used as food items.
Acknowledgments
The Ontario Elk Restoration program is supported
by the Provincial Elk Technical Team and the Ontario
Ministry of Natural Resources (OMNR), Wildlife Sec-
tion, and the Wildlife Research and Development Sec-
tion (WRDS). Thanks to Tom Simpson and Mike
Scafie, OMNR, Bancroft District; John O’ Donnell,
Barry Wannamaker, Ryan Cavanaugh, Jason Langis,
Bancroft Elk Program; Jim Young, Provincial Elk Pro-
gram Coordinator; and Terese McIntosh and Arthur
Dupuis, Trent University, for assisting with the immo-
bilization and processing of the elk. Special thanks to
Joe Neuhold for all of his assistance and dedication to
the Elk Program. Thanks to Norm Cool and Rob Kay,
Parks Canada, EINP, Alberta, and Bruce Chisholm,
Canadian Food Inspection Agency, Edmonton, Alberta,
for weighing and aging the elk during processing at
EINP. J. C. Davies and Mike Allan, OMNR, WRDS, re-
viewed the manuscript and provided helpful comments.
Literature Cited
Bellhouse T., and R. Rosatte. 2005. Assessment of the poten-
tial for negative interaction between re-introduced elk (Cer-
vus elaphus) and resident white-tailed deer (Odocoileus
virginianus) in their wintering areas in Ontario, Canada.
Mammalia 69 (1): 35-56.
Vol. 121
Golightly R., and T. Hofstra. 1989. Immobilization of elk
with a ketamine-xylazine mix and rapid reversal with yo-
himbine hydrochloride. Wildlife Society Bulletin 17: 53-
58.
Hudson R., and J. Haigh. 2002. Physical and physiological
adaptations. Pages 199-257 in North American elk, ecology
and management. Edited by D. Toweill and J. W. Thomas.
Smithsonian Institution Press, Washington, D.C.
McIntosh, T., R. Rosatte, D. Campbell, K. Welch, D.
Fornier, M. Spinato, and O. Ogunremi. 2007. Evidence
of Parelaphostrongylus tenuis infections in free-ranging
elk (Cervus elaphus) in southern Ontario. Canadian Veteri-
nary Journal 48: 1146-1154.
Miller B., L. Muller, T. Doherty, D. Osborn, K. Miller, and
R. Warren. 2004. Effectiveness of antagonists for tileta-
mine-zolazepam/xylazine immobilization in female white-
tailed deer. Journal of Wildlife Diseases 40: 533-537.
Millspaugh J., G. Brundige, J. Jenks, C. Tyner, and D.
Hustead. 1995. Immobilization of Rocky Mountain elk
with Telazol and xylazine hydrochloride, with antagonism
by yohimbine hydrochloride. Journal of Wildlife Diseases
31: 259-262.
Read M., N Caulkett, A. Symington, and T. Shury. 2001.
Treatment of hypoxemia during xylazine-tiletamine-
zolazepam immobilization of wapiti. Canadian Veterinary
Journal 42: 861-864.
Rosatte R., J. Hamr, B. Ranta, J. Young, and N. Cool.
2002. Elk restoration in Ontario, Canada: infectious dis-
ease management strategy, 1998-2001. Annals of the
New York Academy of Sciences 969: 358-363.
Rosatte, R., J. Hamr, J. Young, I. Filion, and H. Smith.
2007. The restoration of Elk (Cervus elaphus) in Ontario,
Canada: 1998-2005. Restoration Ecology 15: 34-43.
Received 19 January 2006
Accepted 11 January 2008
First Confirmed Breeding of the Marbled Godwit, Limosa fedoa,
in Québec
FRANCOIS MorNEAU!, BENOIT GAGNON,” and SIDNEY WHISKEYCHAN®
'63 rue Champagne, Saint-Basile-le-Grand, Québec J3N 1C2 Canada
Corresponding author. Hydro-Québec, Direction Environnement, 855 Sainte-Catherine Est, 9° étage, Montréal, Québec
H2L 4P5 Canada; e-mail: gagnon.benoit@hydro.qc.ca
> P.O. Box 167, Waskaganish, Québec JOM 1RO Canada
Morneau, Francois, Benoit Gagnon, and Sydney Whiskeychan. 2007. First confirmed breeding of the Marbled Godwit, Limosa
fedoa, in Québec. Canadian Field-Naturalist 121(1): 67—70
The objectives of this study were to confirm breeding of the Marbled Godwit (Limosa fedoa) in Rupert Bay, on the Québec
coast of James Bay, and to assess population abundance. Nest search was conducted 17—20 June 2003, using several tech-
niques. Two nests were found in the high marsh of Cabbage Willows Bay: these are the first confirmed breeding records of
this species in Québec. The Rupert Bay region probably holds a breeding population of no more than a few dozen pairs.
Cette étude visait a confirmer la reproduction de la Barge marbrée (Limosa fedoa) dans la baie de Rupert située sur la céte
est de la Baie James (Québec) et 4 estimer l’abondance de la population. Une recherche de nids a été réalisée entre le 17 et
le 20 juin 2003 a l’aide de différentes techniques. Deux nids ont été découverts dans le haut marais de la baie Cabbage
Willows. Ils confirment pour la premiére fois le statut nicheur de cette espéce au Québec. La région de la baie de Rupert
abrite probablement une population nicheuse d’au plus quelques douzaines de couples.
Key Words: Marbled Godwit, Limosa fedoa, nest search, breeding, abundance, James Bay, Rupert Bay, Cabbage Willows Bay,
Québec.
The Marbled Godwit (Limosa fedoa) breeds only in
North America, in three disjunct areas: the grassland
of the northern United States and southern Canada, the
Alaska Peninsula, and James Bay (Gratto-Trevor 2000).
The James Bay area probably holds only 1000-2000
birds (Gratto-Trevor 2000), but the species breeding
has been confirmed only in Ontario (Morrison et al.
1976; Peck and James 1983; Morrison 1987).
In Québec, the species was first recorded on the
James Bay coast by Spreadborough (in Macoun and
Macoun 1909), who stated that the species bred on
both coasts in 1904, although no direct evidence for
breeding was reported. The Marbled Godwit was not
recorded again on the east coast of James Bay before
1990. Since then it has been observed during the breed-
ing season in and near Rupert Bay (Létourneau and
Morrier 1996). Breeding behaviours were noted there
in 1990 and 1991, but nesting was not confirmed.
In 2003, as part of the Eastmain-1-A and Rupert
diversion hydroelectric project environmental studies,
we sought to clarify the breeding status of the Marbled
Godwit in Rupert Bay, on the Québec side of James
Bay. The objectives of this study were to confirm breed-
ing of the species in Québec and to assess population
abundance.
Study Area and Methods
Field work was carried out in the coastal wetlands of
Rupert Bay, Québec, Canada (Figure 1). These wet-
lands are particularly well developed owing to the
sandy deposits and flat landscape. They are especially
extensive in Cabbage Willows Bay, which opens into
Rupert Bay, and Boatswain Bay, which opens directly
into James Bay, but are narrower in Hall Cove and else-
where. They consist, in sequence from upland to open
water, of swamps, fens, high marshes and low marshes
(FORAMEC 2004*). The high marsh grounds are
somewhat spongy but not covered by water. Low
marshes are covered twice daily by the tide but the other
wetlands are flooded only by storm tides and equinoc-
tial tides.
Nest searches were conducted on 17—20 June 2003,
using several techniques. The rope-drag method (Klett
et al. 1986) was used on 0.3—1.4 km transects in parts
of the high coastal marsh of Cabbage Willows Bay
and Hall Cove. Nests were also searched for in the same
areas by watching birds flying from the low marsh to
potential nesting areas or vice versa. In addition, play-
backs of taped breeding calls were used regularly on
the high marsh and once in the low marsh. Finally, to
increase ground coverage, we conducted a helicopter
survey in Cabbage Willows Bay, in Hall Cove, over
some Tamarack (Larix laricina) fens on Ministikawatin
Peninsula, and on Jacob Island, north of Rupert Bay.
We searched along 50-m parallel transects, at speeds
that varied from 20 km/h to hover, and at altitudes
varying from 3 to 9 m.
Breeding chronology was estimated assuming a
mean egg-laying interval of 2 days and an incubation
period of 25 days (Gratto-Trevor 2000). Abundance
was estimated with counts of birds performing aerial
display and nests.
67
68 THE CANADIAN FIELD-NATURALIST
Vol. 121
—78° 45 *
] Boatswain Bay
Migratory Bird
La bs
‘Bay
lleTent'> £ ft 4 Sanctuary
i Di. 0
Island
Cabbage
Willows Bay
Biodiversity Reserve
==
—_ onli
Migratory Bird Sanctuary
Marbled Godwit nest
6675_hq_146_f1_080115.fh9
ag cd
’ Proposed Boatswain Bay
Biodiversity Reserve
Rupert
Waskagani
Ficure |. Location of Marbled Godwit study area and nests.
Results
The Marbled Godwit nest search was conducted on
slightly over 20 km of line transects using the rope-
drag method. It covered about 70 ha.
Two Marbled Godwit nests, 3.6 km apart, were dis-
covered in the high coastal marsh of Cabbage Willows
Bay in 2003 (Figure 1). The first nest (51°32.61'N,
79°16.13'W), containing four eggs (Figure 2) and
one adult incubating, was found on 17 June using the
rope-drag technique. The second nest (51°30.77'N,
79°15.13'W), discovered on 20 June following obser-
vation of courtship behaviour, contained only one egg,
but two adults were within 30 m of the nest. The first
nest was revisited again on 19 June and still contained
four eggs.
Nests were 5 cm in depth and 20 cm in diameter at
the rim. They were mostly, if not completely, made of
dry stems of Sweet Grass (Hierochloe odorata). Both
nests were found in the largest clumps of that species
in the vicinity. The first nest was partly covered by
2007
MORNEAU, GAGNON, and WHISKEYCHAN: MARBLED GOpDWIT 69
FIGURE 2. First Marbled Godwit nest discovered in Rupert Bay, Québec.
green stems. Throughout the high coastal marsh, grass
cover, about 10—15 cm tall, was relatively sparse, with
occasional clumps up to 15—25 cm.
In 2003, laying and potential incubation in Cab-
bage Willows Bay occurred at least between 11 June
and 19 July.
No Godwit responded to several playback trials in
the high coastal marsh. However, two individuals
answered spontaneously to the single trial on the low
coastal marsh, by flying and calling around the inves-
tigators. Over the same biotope, four different birds
were seen engaging in aerial courtship behaviour dur-
ing nest searches. Besides the birds associated with
the nests, only one Marbled Godwit was seen in the
high marsh: it was flying from the low marsh and
landed in the high marsh, on a potential nesting area.
Despite a long search, it was not seen again.
Partial aerial coverage of the Cabbage Willows Bay
low coastal marsh enabled us to identify four adults.
Another bird was spotted as it flew toward the low
marsh during the aerial survey; it seemed to be com-
ing from the second nest area. No birds were seen in
Hall Cove or in Ministikawatin Peninsula or Jacob
Island during aerial surveys.
Discussion
The two Marbled Godwit nests that we found in
Cabbage Willows Bay represent the first definitive
breeding evidences of this species in Québec. Some
previous observations had led us to believe the species
was nesting there. On 17 June 1991, two copulating
birds were observed in Cabbage Willows Bay. The
same year, three Marbled Godwits were seen on 21 July
in the same area, including two agitated birds giving
alarm calls, indicating the probable presence of young
(Létourneau and Morrier 1996). Finally, a similar ob-
servation was recorded on 27 July 2002, again in Cab-
bage Willows Bay, north of the Novide River (Hydro-
Québec, unpublished data).
Breeding habitat in Cabbage Willows Bay is simi-
lar to that described elsewhere. In North Dakota, the
Marbled Godwit shows a strong preference during
breeding season for short, sparse to moderately dense
cover along wetland shorelines and shorter vegetation
in upland habitats (Ryan et al. 1984). In James Bay,
Ontario, one nest was found in a raised, grassy ridge
between two ponds, in an open coastal marsh (Peck
and James 1983).
Estimated breeding chronology suggested that nest
search was carried out in the egg-laying and early in-
cubation period. This is consistent with the observed
copulation (17 June 1991) and other previous records
suggesting breeding (see above). The flight displays
observed in 2003 also support the breeding chronolo-
gy outline. Display behaviour is most frequent early
in the breeding season and is commonly performed by
70 THE CANADIAN FIELD-NATURALIST
unpaired or nest-scraping males until incubation duties
are initiated (Nowicki 1973). In Ontario, two breeding
records from James Bay indicate that egg-laying was
initiated approximately between 24 May and 31 May
(Morrison et al. 1976; Peck and James 1983).
In 2003, the Marbled Godwit breeding population
included at least four pairs in Cabbage Willows Bay.
In 2002, the species was also recorded in Boatswain
Bay, Jacob Island and Tent Island, between 23 July
and 1 September (Hydro-Québec, unpublished data).
However, considering the late dates and that no breed-
ing behaviour was observed, it is likely that these birds
were migrants rather than breeders. Although no Mar-
bled Godwit was recorded during the aerial survey
inland of Ministikawatin Peninsula, the species was
located there in June 1990 and 1991, in three areas, in-
cluding a bird on a shallow lake fen (Létourneau and
Morrier 1996). In Ontario, the Marbled Godwit is also
found up to 20 km inland (Gratto-Trevor 2000). Thus,
the species probably breeds inland of Ministikawatin
Peninsula. Its abundance there could exceed that of
Cabbage Willows Bay because of the vast expanse of
Tamarack fens. On 15 June 1991, 80 Marbled Godwits
were counted in Cabbage Willows Bay (Létourneau
and Morrier 1996). Their breeding status is question-
able. As noted earlier, that date probably coincides with
egg-laying and early incubation. Either these birds
were on the point of dispersal for local breeding or
were migrants on the way to northern James Bay
breeding grounds in Ontario. However, if Marbled
Godwits fly directly to breeding areas in spring from
wintering sites on southeastern U.S. coasts, as the re-
verse is presumed to occur in the fall (Morrison et al.
1976), then those birds may have been local or regional
breeders. Therefore, the Rupert Bay region probably
holds a breeding population of between four pairs and
a few dozens pairs.
Acknowledgments
This study was funded by Hydro-Québec. We are
grateful to Francoise Lebrun and Gilles Lefebvre, from
Hydro-Québec, for production of the map. We thank
Jeffrey Salt, from the Waskaganish Cree community,
for his contribution to this study. We are grateful to
helicopter pilot Laurie Gagnon for her field assistance.
Jean Doucet and Mary Antico provided helpful com-
Vol. 121
ments on an earlier draft of the manuscript. A. J. Ersk-
ine and an anonymous reviewer provided useful com-
ments to improve the manuscript.
Document Cited (marked * in text)
FORAMEC Inc. 2004. Centrale de l’Eastmain-1-A et déri-
vation Rupert. Etude de la végétation et des espéces floris-
tiques et fauniques menacées ou vulnérables. Préparé pour
la Société d’ énergie de la baie James. Québec.
Literature Cited
Gratto-Trevor, C. L. 2000. Marbled Godwit (Limosa fedoa).
The Birds of North America, No. 492. Edited by A. Poole
et F. Gill. The Birds of North America, Inc., Philadelphia.
Klett, A. T., H. F. Duebbert, C. A. Faanes, and K. F. Hig-
gins. 1986. Techniques for studying nest success of ducks
in upland habitats in the prairie pothole region. U.S. Fish
and Wildlife Service, Resource Publication (158). Wash-
ington, D.C.
Létourneau, V., and A. Morrier. 1996. Marbled Godwit.
Pages 1138-1139 in The breeding birds of Québec: atlas
of the breeding birds of Southern Québec. Edited by J.
Gauthier and Y. Aubry. Association québécoise des groupes
d’ornithologues, Montréal, Québec. Province of Québec
Society for the Protection of Birds, Canadian Wildlife
Service, Environment Canada, Québec Region, Montréal.
Xvill + 1295 pages.
Macoun, J., and J. Macoun. 1909. Catalogue of Canadian
birds, 24 edition. Geological Survey of Canada, Depart-
ment of Mines, Ottawa. Bulletin (973). 761 pages.
Morrison, R. I. G. 1987. Marbled Godwit. Page 528 in Atlas
of the breeding birds of Ontario. Edited by M. D. Cadman,
P. F. Eagles, and F. M. Helleiner. University of Waterloo
Press, Waterloo, Ontario.
Morrison, R. I. G., T. H. Manning, and J. A. Hagar. 1976.
Breeding of the Marbled Godwit, Limosa fedoa, in James
Bay. Canadian Field-Naturalist 90: 487-490.
Nowicki, T. 1973. A behavioral study of the Marbled Godwit
in North Dakota. MLS. thesis, Central Michigan University,
Mount Pleasant, Michigan.
Peck, G. K., and R. D. James. 1983. Breeding birds of On-
tario: nidiology and distribution, vol. 1: Non-passerines.
Life Sciences Miscellaneous Publications, Royal Ontario
Museum, Toronto. xii + 321 pages.
Ryan, M. R., R. B. Renken, and J. J. Dinsmore. 1984.
Marbled Godwit habitat selection in the northern prairie
region. Journal of Wildlife Management 48: 1206-1218.
Received 23 January 2006
Accepted 18 January 2008
Swift Fox, Vulpes velox, Den Use Patterns in Northwestern Texas
BRADY K. MCGEE, WARREN B. BALLARD, and Kerry L. NICHOLSON
Department of Natural Resources Management, Texas Tech University, Box 42125, Lubbock, Texas 79409 USA
McGee, Brady K., Warren B. Ballard, and Kerry L. Nicholson. 2007. Swift Fox, Vulpes velox, den use patterns in northwestern
Texas. Canadian Field-Naturalist 121(1): 71-75.
Predator avoidance may be a reason why Swift Foxes (Vulpes velox) are one of the most burrow-dependent canids in North
America. Typically Swift Foxes have multiple dens, which they frequently move among. As part of a larger study to reduce
Coyote (Canis latrans) related mortalities on Swift Foxes, we installed artificial escape dens in areas occupied by Swift
Foxes on Rita Blanca National Grassland, Dallam County, Texas. For this paper, our objective was to determine the effects of
artificial escape dens on Swift Fox den use patterns. From January 2002 to August 2004 we captured, radio-collared, and
monitored 55 Swift Foxes. We documented annual number of dens used, rate of den use (fidelity), distance between dens,
den area, and den sharing. We compared treated (artificial dens installed) and untreated (no artificial dens) areas but found
no differences in annual number of dens (P = 0.64; ¥ = 8), rate of den use (P = 0.96; ¥ = 35%), mean distance between dens
(P = 0.99; X = 2,311 m), den area (P = 0.55; X = 5.72 km’), or den sharing (P = 0.46; ¥ = 42% of time). We did not observe
an effect of artificial escape dens on Swift Fox den use patterns probably because artificial escape dens were designed for
.
|
|
|
if
|
1
\
temporary escape cover rather than diurnal den use.
Key Words: Swift fox, Vulpes velox, den range, den sharing, den use, fidelity, Texas.
Swift Foxes (Vulpes velox) were once abundant
throughout the short and mid-grass prairies of North
America but have rapidly declined with expansion of
human settlement (Egoscue 1979). Studies have shown
that the Swift Fox primarily inhabits areas of native
rangeland (Allardyce and Sovada 2003). Much of the
historical Swift Fox range has been fragmented into
patches of native rangeland, Conservation Reserve
Program, and agricultural fields (Allardyce and Sova-
da 2003). Habitat loss has been one of the reasons
Swift Foxes were temporarily a candidate for endan-
gered species listing with the United States Fish and
Wildlife Service (USFWS) from 1992 to 2001 (USFWS
1995; USFWS 2001; Allardyce and Sovada 2003).
Predation has been another limiting factor on Swift
Fox populations (Kamler et al. 2003a). Many studies
have shown that Coyotes (Canis latrans) are the pri-
mary source of Swift Fox mortality, with annual sur-
vival rates ranging from 43 to 53% (Sovada et al. 1998;
Kitchen et al. 1999; Matlack et al. 2000; Anderson et
al. 2003; Allardyce and Sovada 2003). Because Coy-
otes rarely consume Swift Foxes they kill (Sovada et
al. 1998; Kitchen et al. 1999; Allardyce and Sovada
2003), Coyote predation seems to be the result of both
interference and exploitative competition (Kamler et
al. 2003b).
Predator avoidance may be one reason why Swift
Foxes are one of the most burrow-dependent canids in
North America (Moehrenschlager et al. 2004). Dens
_ are an integral part of Swift Fox ecology. Swift Foxes
use dens year-round not only for protection from pred-
{
;
ators, but also for reproduction, resting, and avoidance
of extreme climatic conditions (Egoscue 1979). In
sparsely vegetated habitats occupied by Swift Foxes,
dens may constitute crucial escape cover. Arjo et al.
(2003) found the number of dens used by Kit Foxes
(Vulpes macrotis) was positively correlated with Coy-
ote numbers. In addition, White et al. (1994) suggest-
ed that Kit Foxes established a number of dens (220)
to facilitate escape.
It has also been suggested that Swift Foxes move
frequently among different den sites (Kilgore 1969;
Hines and Case 1991), but little detail on den use pat-
terns has been reported. Understanding Swift Fox den
behavior may be an important factor in sustaining
viable populations throughout their range.
Because Swift Foxes use dens year-round (Egoscue
1979), we installed artificial escape dens as part of a
larger study to determine if lack of den sites limited
Swift Fox populations in northwest Texas, USA
(McGee 2005). The objective of this paper was to doc-
ument the effects of artificial escape dens on Swift
Fox den use patterns. We determined annual number of
dens used, rate of den use (fidelity), distances between
dens, den area, and den sharing. We predicted that with
more dens available in treated areas Swift Foxes would
use more dens, have less fidelity for certain dens, have
greater distances between dens, and be less likely to
share a den. We made comparisons between treated
(artificial escape dens installed) and untreated (no arti-
ficial dens installed) areas.
Study Area
We collected data from a contiguous 100-km? area
on the Rita Blanca National Grassland (NG) in Dallam
County, Texas, approximately 43 km northwest of
Dalhart, Texas (Figure 1). The NG consisted of native
rangelands with short-grass prairie dominated by Blue
Grama (Bouteloua gracilis), Side-oats Grama (Boute-
loua curtipendula), Burrograss (Haplopappus tenui-
wal
YD THE CANADIAN FIELD-NATURALIST
sectus), and Buffalograss (Buchl6e dactyoides) that
were moderately to intensively grazed by cattle (Bos
taurus; Kamler et al. 2003a,b; Nicholson 2004;
McGee 2005).
Methods
We captured, handled, and radio-collared Swift Fox
using methods described by McGee (2005). We tracked
Swift Fox to their diurnal resting sites (dens) using a
hand-held antenna 1—2 times per week. We recorded
each den location with a Garmin global positioning
system receiver (Garmin International Inc., Olathe,
Kansas, USA). We only used Swift Fox that were
monitored =8 months of the year in data analysis. We
calculated annual estimates from September to August
of each year for all analyses. This was to allow Swift
Foxes an adjustment period after installation of artifi-
cial dens and to perform two full years of data analysis.
During April 2002, we placed 108 artificial escape
dens in three spatially separated areas (Figure 1). We
considered Swift Foxes belonging to a treated area if
their home ranges overlapped an artificial escape den
area by =50%. We considered untreated Swift Foxes
as those whose home ranges did not overlap artificial
escape den areas (McGee 2005). No Swift Foxes in un-
treated groups were ever located within an artificial
escape den treated area. Also, we considered foxes to
belong to the same family group if they used the same
area and dens concurrently (Kitchen et al. 1999; Kam-
ler et al. 2003a,b).
Escape dens consisted of corrugated plastic sewer
pipes 4.04 m long, 20.32 cm diameter with 20.32 cm
holes cut in the middle to allow foxes to modify and
expand subterranean dens ($6.41/m U.S.; Amarillo
Plumbing Supply, Inc., Amarillo, Texas, USA). The
diameter size of our artificial escape dens was based
on previous studies that reported a mean den opening
height of 20.0 cm for Swift Fox dens (Cutter 1958; Hill-
man and Sharps 1978; Pruss 1999; Jackson and Choate
2000). Coyote dens were reported to be 30-37 cm in
diameter (Bekoff 1977; Althoff 1980; Bekoff 1982;
Harrison and Gilbert 1985). We assumed that artifi-
cial escape den entrances, being the same diameter as
natural Swift Fox dens, were too narrow for Coyotes.
A John Deere 260 skid loader (Deere and Company
World Headquarters, Moline, Illinois, USA) was used
to install and cover the sewer pipe with only the two
open ends exposed. Escape cover was randomly ori-
ented and spaced approximately 322 m apart in a
2.59 km? grid pattern for a density of 36/2.59 km?
(McGee 2005).
We calculated rate of den use (fidelity) by dividing
number of dens by the number of times the fox was
located in dens (x 100%). Lower values represented
higher den fidelity. We calculated distances between
dens using Bearing and Distance Extension for
ArcView 3.2 (Environmental Systems Research Insti-
tute, Redlands, California, USA). We used den loca-
Vol. 121
FiGurE 1. Map of the 100-km? study area located on the
Rita Blanca National Grassland in northwest Dallam
County, Texas, USA. One hundred and eight artifi-
cial escape dens (black dots) where installed in three
separate grid locations.
tions to estimate annual den areas for Swift Foxes us-
ing 100% minimum convex polygon (MCP) method
(Mohr 1947) as calculated by Home Range extension
(Rodgers and Carr 1998) for ArcView 3.2 (Environ-
mental Systems Research Institute, Redlands, Cali-
fornia, USA). We calculated den sharing as percent-
age of time a radio-collared Swift Fox was found in a
den with another radio-collared fox.
We found no statistical differences among years so
we pooled data to increase power. We used 1-way
ANOVAs in SPSS 12.0 (SPSS 2003) to determine
differences between treated and untreated areas for
annual number of dens used, mean distances between
dens, and mean den areas. We compared average rate
of den use and sharing between treated and untreated
areas using Yates’ corrected chi-square tests (Zar
1999). Differences were deemed significant when
P <0.05.
Results
From January 2002 to August 2004, we captured
and radio-collared 55 Swift Foxes (31 males, 24 fe-
males). We documented a total of 104 Swift Fox dens
a
2007
McGEE, BALLARD, and NICHOLSON: Swift FOX DEN Usi 73
TABLE |. Average annual number of dens and den area of Swift Foxes on Rita Blanca National Grassland (NG) in northwest
Texas, 2002—2004.
Annual number of dens
Study area n % +t SE range
treated 8 7.88 + 0.48 6-10
untreated 4 7.50 + 0.50 7-9
during our study including four separate occasions when
we radio-tracked and observed Swift Foxes within arti-
ficial escape dens during the day. Due to the high turnover
rate in our Swift Fox population, only 12 Swift Foxes
(n = 8 treated, n = 4 untreated) were monitored for
28 months a year from September 2002 to August
2004.
There was no difference (P = 0.64) in annual num-
ber of dens used (mean + SE) by Swift Foxes between
treated (7.88 + 0.48, n = 8) and untreated (7.50 + 0.50,
n = 4) areas (Table |). Average rate of den use (fideli-
ty) was similar (Yates’ 7 = 0.003, P = 0.96) between
treated (35.9%) and untreated (35.4%) areas. Mean dis-
tance between dens (+ SE) was not different (F < 0.001,
P=0.99, 1-B = 0.05) between treated (2308 + 442 m,
n = 8) and untreated (2317 + 654 m, n = 4) areas.
There was no difference in den area (F = 0.38, P = 0.55,
1 — 8 = 0.09) between treated (6.34 + 1.86 km’, n = 8)
and untreated areas (3.81 + 1.91 km?, n = 4). Den shar-
ing only occurred between mated pairs. Average rate of
den sharing was similar (Yates’ x? = 0.56, P = 0.46)
between treated (44.9%) and untreated (39.0%) areas.
Discussion
We have shown that average number of dens used
by Swift Fox each year was eight in northwest Texas
(Table 1). Schauster et al. (2002) documented num-
ber of Swift Fox dens used was 2-8 for breeding and
gestation season, 5-10 for pup-rearing season, and
3-8 for the dispersal season. Similar multiple den use
has been documented with other small canids that share
a dependence on dens, such as Artic Fox (Alopex lago-
pus; Eberhardt et al. 1983) and Kit Fox (Tannerfeldt
et al. 2003; Moehrenschlager et al. 2004). Koopman et
al. (1998) reported an average of 11.8 dens per year
for Kit Foxes in California. Arjo et al. (2003) sug-
gested to an increase in the number of dens used by
Kit Foxes in western Utah could have been related to
the increase in Coyote presence.
We found that Swift Foxes had relatively high
fidelity to particular dens for both treated and un-
treated areas. Koopman et al. (1998) found that Kit
Foxes exhibited a strong affinity for particular dens
because Kit Foxes were located in their most frequent-
ly used den 32% of the time. Artic Foxes also preferred
certain dens while others were used infrequently (Eber-
hardt et al. 1983). We suspect that den fidelity may be
the result of den quality. Distinctions have been made
between good and bad dens for Artic Foxes (Tanner-
Den range (km*)
¥+SE range
6.34 + 1.86 0.52-13.21
4.50 + 1.91 0.81-9.20
feldt et al. 2003). On the other hand, Swift Foxes fre-
quently switch between dens (Egoscue 1979; Hines
and Case 1991), suggesting poor den quality. One of
the reasons for frequent changes between dens has
been attributed to the large numbers of fleas found in
Swift Foxes’ dens (Kilgore 1969). Other factors like
human disturbance, leaking, shifting towards food re-
sources, and predator avoidance may play a role in
Swift Foxes den switching and fidelity (Kilgore 1969;
Tannerfeldt et al. 2003).
In addition, we documented a mean distance be-
tween dens of 2311 m (range 729-3998 m). Similarly,
Moehrenschlager (2000) found that Swift Foxes in
Canada moved to dens up to 1900 m away. In contrast,
Cutter (1958) noted that Swift Fox dens in overgrazed
pastures of northern Texas were concentrated but did
not state the distance between dens. Cutter (1958) noted
that up to six dens were frequently observed within
65 hectares of pastureland. Greater distances between
dens would allow Swift Foxes access to more resources
within their environment if dens were used to escape
predators.
It has been suggested that carnivore home-range
size can be affected by habitat composition and food
distribution (Macdonald 1983). White and Ralls (1993)
observed that larger Kit Fox home ranges were asso-
ciated with low prey availability. We suspect that these
effects can be applied to den area of Swift Foxes as well.
Our results indicated a mean den area of 5.72 km”. It is
possible that Swift Foxes with larger den areas may
have had greater access to food resources while avoid-
ing predators. Ables (1969) recognized that Red Fox
(Vulpes vulpes) home range size was affected by food
abundance and availability. In addition, Hines and Case
(1991) speculated that carrion availability and prey
distribution probably affected Swift Fox home range
size and shape. Likewise, Olson and Lindzey (2002)
suggested that intensive hunting by Swift Foxes near
natal dens may have reduced prey availability and con-
sequently forced adults to expand the areas in which
they hunted. Thus, larger den areas may be the result
of fewer resources (Hines and Case 1991).
Although additional adult foxes have been observed
with mated pairs at Swift Fox (Egoscue 1979: Covell
1992; Kitchen et al. 1999; Lemons et al. 2003), Arctic
Fox (Eberhardt et al. 1983), and Kit Fox (Ralls et al.
2001; Tannerfeldt et al. 2003) den sites, we did not
document this occurrence. Of the Swift Foxes that we
monitored, only mated pairs were ever found occupy-
74 THE CANADIAN FIELD-NATURALIST
ing the same den. Adult radio-collared Swift Foxes in
our study denned with their mate 39-44% of the time.
Similarly, Koopman et al. (1998) found that mated
adult Kit Foxes denned together about 45% of the time.
Also, Ralls and White (2003) found that Kit Fox pair
members shared the same den 51% of the time. It is
possible that we underestimated the rate of den shar-
ing. We believe that on occasion a radio-collared fox
denned with adult foxes that were not radio-collared.
In conclusion, we were able to describe Swift Fox
den use patterns in northwest Texas even though we
did not observe an effect of artificial dens. One possi-
ble reason for not observing an effect may be the low
sample sizes. Even though we captured 55 Swift Foxes
during our study, residents were not abundant as data
analysis was only performed on eight Swift Foxes in
treated and four Swift Foxes in untreated areas that
were monitored =8 months per year. Also, no observed
effect of artificial dens may be due to the fact that arti-
ficial dens were designed for temporary escape from
predators, specifically Coyotes, while Swift Foxes were
away from their natural dens. As part of the larger study
mentioned before (McGee 2005), we found higher
annual Swift Fox survival (P = 0.07) in artificial den
treated areas (0.81) than in untreated areas (0.52) on
the same study site. Higher survival in treated areas
suggests that Swift Foxes were using artificial dens for
escape during their normal nocturnal activities. Diur-
nal use of artificial dens was limited. We only tracked
Swift Foxes to artificial escape dens on four occasions
during the day. Therefore, alternative artificial den
designs need further study to find a more suitable diur-
nal artificial den for Swift Foxes.
Acknowledgments
Funding was provided by the National Fish and
Wildlife Foundation and Texas Tech University. We
thank the United States Forest Service personnel for
letting us conduct research on the Rita Blanca Nation-
al Grasslands and helping install artificial escape dens.
We thank F. Pronger for allowing us to use his ranch
as a study site. We also thank E. and B. Hampton for
providing us with a place to stay while conducting
field research. Research protocols were approved by
the Animal Care and Use Committee at Texas Tech
University. This is Texas Tech University, College of
Agricultural Sciences and Natural Resources techni-
cal publication T-9-1091.
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Accepted 17 January 2008
Flowering During January in Antigonish County, Nova Scotia*
DAVID J. GARBARY and BARRY R. TAYLOR
Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia, B2G 2W5 Canada; e-mail: dgarbary @ gmail.com
*Contribution from the Herbarium of St. Francis Xavier University
Garbary, David J., and Barry R. Taylor. 2007. Flowering during January in Antigonish County, Nova Scotia. Canadian Field-
Naturalist 121(1): 76-80.
Over 85 records of 23 species of blooming, herbaceous angiosperms were made at 19 sites in Antigonish County between 7 and
21 January 2006, when daytime temperatures reached 15°C. These observations followed an unusually warm fall and early
winter. All species were observed on waste ground or in fields and garden plots, except for Epigaea repens L. which was part
of ground vegetation in a sparsely wooded site. The primary families represented were Asteraceae (six species), Brassicaceae
(six species) and Carophyllaceae (four species). The most commonly observed plants were Taraxacum officinale (L.) Weber
(11 sites), Capsella bursa-pastoris (L.) Medik. (nine sites), Thlaspi arvense L. (three sites) and Stellaria media (L.) Vill.
(four sites). Many plants and inflorescences were conspicuously frost-damaged, and flowers were rarely fully open. In
several species, e.g., TZ: arvense and Cerastium vulgatum L., many individual plants looked normal and there was no evidence
of frost damage. These observations are the latest flowering records for Nova Scotia.
Key Words: Asteraceae, Brassicaceae, Caryophyllaceae, winter flowering, phenology, Antigonish County, Nova Scotia.
In Nova Scotia most species of flowering plants
have ceased to flower by early November (Zinck
1998). Taylor and Garbary (2003) carried out an ex-
tensive survey of flowering during an unusually warm
late fall in Antigonish County and found 92 species in
bloom during November-December. These were the
latest flowering records of all but one of these species,
and extended the known flowering periods for plants in
Nova Scotia by an average of 45 days. Of particular
note were 15 species that were in bloom on 14-16 De-
cember 2001.
A detailed evaluation of the phenology of the entire
late fall flora was again undertaken in 2005 after anoth-
er warm fall in which abundant flowering was still oc-
curring in early November. This study will be described
elsewhere in the context of regional climate change
(Garbary et al.). Here, we provide records of 23 species
of herbaceous plants observed in flower during an ex-
traordinary warm period in January 2006. Our obser-
vations complement those on spring phenology by
Vasseur et al. (2001). They compared flowering times
in 12 species in the late 1990s with dates a century
earlier and showed significant changes in only two
species, of which only Syringa vulgaris L. had earlier
flowering.
Materials and Methods
Collection sites
Nineteen sites in Antigonish Town and County,
northern Nova Scotia, were explored between 7 and
21 January 2006 in three collection periods. Adjacent
sites were a minimum of 200 m apart, and the maxi-
mum distance between two sites was 23 km. Ten sites
were visited on 7 January, 16 sites during the period
11-16 January, and seven sites on 21 January. Five sites
were visited three times, and three other sites were
visited twice. Most of these sites had been previously
visited by Taylor and Garbary (2003). Sites were typi-
cally highly disturbed areas of waste ground or gravel
(Figure 1), or agricultural fields or garden plots. Each
site was surveyed for 15-45 minutes. Notes were taken
on the flowering status and plant condition of each
species in bloom. Representative plants were collected;
plants with cryptic flowers were dissected and exam-
ined under a stereomicroscope to ensure that flowers
were intact. Voucher specimens for most species were
prepared and deposited at St. Francis Xavier Univer-
sity. Nomenclature follows Zinck (1998).
Weather
Weather data were taken from the nearest active
Environment Canada meteorological station at Tra-
cadie, approximately 25 km from Antigonish (http://
climate.weatheroffice.ec.gc.ca/climateData/canada
_e.html). The first collecting period was on 7 January
when daytime temperatures reached 4.3°C. This fol-
lowed several days with daily maximum temperatures
above 0°C and daily minimal temperatures no lower
than -2.5°C. Each of the succeeding four nights had
frost, although daytime maxima were always above
freezing. The second collecting period, between 12 to
16 January, had minimum and maximum tempera-
tures of 0.4°C and 15.9°C, respectively. Temperatures
between 15 to 20 January were cooler and frosts oc-
curred most nights (minimum -6.8°C) although mean
daily temperatures were below freezing on only one
day (-4.6°C). The final collecting day was 21 January
when temperature ranged from 2.0 to 8.4°C. This was
followed by at least ten days when temperatures
dropped below freezing (-0.3 to -9.2°C) and condi-
tions were more typical of the season.
76
2007
r
GARBARY AND TAYLOR: FLOWERING DURING JANUARY
Ficure 1. Sample habitat. Waste ground on campus of St. Francis Xavier University adjacent to Trans-Canada Highway, 7 January
2006. Four species were found in flower at this site.
Results
ASTERACEAE
Taraxacum officinale (L.) Weber was the most wide-
ly distributed species, occurring at ten sites. One to
four plants were observed at each site during all three
collecting periods and blossoms occurred at two sites
on 21 January. Each plant had one or two ragged
inflorescences with short pedicels. Leaves had some
frost damage and were often flattened against the
substratum, although they had good colour on some
individuals.
Achillea millefolium L. occurred as a single plant
in a farmer’s field and as six plants on waste ground
(Figure 2). Plants had green leaves, but the flowering
shoots were often reclining, with several to dozens of
white florets. The flowering shoots observed 14 Jan-
uary were wilted and most of the flowers had turned
brown.
Two species of Matricaria were found: M. mariti-
ma L. and M. matricarioides (Less.) Porter. Plants of
M. maritima were found at two sites. A single, well-
developed inflorescence was found on 7 January as
well as several partially intact flower heads. By 15 Jan-
uary remaining ray florets were mostly lost; however,
the disc florets on many heads remained bright yellow.
The plants were often partly wilted but the leaves had
healthy colour. Up to three plants of M. matricari-
oides were found at each of three sites, and it was
observed during the first two collecting periods. In-
florescences typically had a mixture of dull yellow and
brownish florets. The plants appeared no less viable
than others observed in early December.
Leontodon autumnalis L. occurred at two sites on
7 January where single individuals were found with
only one or two wilted inflorescences on each plant.
Basal rosettes had good colour but were slightly wilt-
ed. Examination of both sites one week later revealed
no flowering individuals.
Two plants of Senecio vulgaris L. occurred at one
site on waste ground; a single individual was found
at a second site. Florets on most inflorescences had
turned brown but several had bright yellow florets.
Plants were a maximum of 10 cm in height and flow-
ering was observed only in the first two observation
periods.
78 THE CANADIAN FIELD-NATURALIST
ayy
i@
FIGURE 2. Achillea millefolium photographed in situ on 7
January 2006 at site shown in Figure 1. Flowering
shoot is about 30 cm tall.
BRASSICACEAE
Capsella bursa-pastoris (L.) Medik. was among
the most common plants flowering in January. It oc-
curred at nine sites with one to three plants per site.
Flowering individuals were up to 30 cm tall and car-
ried a few flowers at the apex of each plant. Leaves
were typically green and healthy, although larger plants
had generally become decumbent. During the second
week, plants were mostly very small (5-10 cm high)
with only a few florets. On 21 January flowers were
found at two sites.
Thlaspi arvense L. was among the most robustly
flowering species observed. The plants were erect, dark
green and with prominent white flowers. 7: arvense
in flower was as common in January as in any previ-
ous collecting period, and flowers were found at one
site on 21 January.
Raphanus raphanistrum L. occurred at two sites
where there were typically many plants. The stems and
leaves of these plants were badly wilted and reclin-
ing; however a few bright yellow florets and numer-
ous clusters of buds remained intact at many apices
on many plants. In the second week only a single, badly
damaged plant was found at one site.
Cardamine pensylvanica Muhl. ex Willd. was the
only species clearly protected by the heat shadow of a
large building. Dozens of small, prostrate plants each
with one to several flowers occurred in a flowerbed
along the side of a large brick building. No decline in
flowering was noted in the second week and flowers
persisted to 21 January.
Erysimum cheiranthoides L. was found at two sites
and only in the second week. Four large plants were
found along a bare concrete foundation for a house.
The surrounding lot supported numerous plants of
this species but all had gone to seed. A second site
had a few small individuals up to 5 cm tall. Both sites
were covered with snow on 7 January and so were not
examined until the second week.
Vol. 121
A single plant of Lepidium campestre (L.) R. Br.
was found 12 January on waste ground. The plant
was about 20 cm tall and had two clusters of flowers
with the most vigorous close to the ground and im-
mersed in a cluster of leaves.
CARYOPHYLLACEAE
Stellaria media (L.) Vill. was found at two or three
sites in all collecting periods (four sites overall), and
was among the most prominent of the plants in flower.
At two locations it was found as scattered individual
plants; however, at two other sites it formed large ex-
panses of continuous ground cover with numerous
inflorescences (Figure 3).
FiGure 3. Stellaria media photographed in situ on 7 January
2006 showing part of a large colony about 1 m in
diameter.
Flowering individuals of Cerastium vulgatum L.
were few and occurred at only three sites, although
many vegetative plants or plants with remains of inflo-
rescences were present at most locations. The repro-
ductive individuals appeared healthy and showed no
signs of frost damage. They grew on loose gravel where
there was no competition from surrounding vegeta-
tion. Flowering persisted at one site to 21 January.
A few flowers were present on several plants of
Spergula arvensis L. from a farmer’s field in the first
two weeks. The plants showed considerable frost dam-
age and dissection showed that very few bore intact
stamens and pistils. At a second site where vegetative
parts of the plants were less damaged, no intact flow-
ers occurred.
An extensive patch of Spergularia rubra (L.) J. &
C. Presl. occurred on waste ground. The prostrate plants
were 5-10 cm diameter and showed all reproductive
stages from buds to fruits. The site was only exam-
ined during the second week of observation.
OTHER FAMILIES
Euphorbia helioscopa L. (Euphorbiaceae) was pres-
ent at only one of the sites. There were hundreds of
individuals. Large plants (20-40 cm tall) were con-
2007
spicuously frost damaged, with the wilted stems caus-
ing the upper parts of the plants to topple over. The
terminal portions of these plants looked healthy, and
subsequent dissection of inflorescences showed intact
flowers. Small plants (5-15 cm tall) showed less col-
lapse of the stems and also had numerous intact flow-
ers. Flowering persisted until 21 January.
Individual plants of Fumaria officinalis L. (Fumari-
aceae) were found on each of the first two visits to one
site. The plants had two small inflorescences in which
a few flowers still retained their characteristic two-
toned colour. Inflorescences on other plants at this site
had been severely damaged and turned brown. On
most individuals the finely dissected leaves and deli-
cate stems showed no conspicuous wilting. At another
site dozens of plants grew in soil mounds, but there
were no inflorescences.
Viola tricolor L. (Violaceae) occurred at two sites.
Both were in flower beds and one was adjacent to a
building. Plants at both sites had leaves and stems in
excellent condition and several to dozens of fully open
flowers with blossoms persisting until 21 January.
There were numerous plants of Veronica persica
Poir. (Scrophulariaceae) at one site where they grew
as prostrate clumps or straggling plants over the bare
soil. The bilobed fruits were conspicuous, and a single
intact but only partially open flower was found on sin-
gle individuals returned to the laboratory at each of
the three visits.
Vegetative plants of Polygonum arenastrum Jord. ex
Boreau (Polygonaceae) were common at most sites.
Although most plants had conspicuous frost damage,
a few plants had two to three white-tipped buds and
open flowers that persisted until week two.
Epigaea repens L. (Ericaceae) was the only wood-
land plant found in the study. An extensive population
occurred at one site (14 January) among other ground
vegetation beneath an open, mixed forest growing on
sandy soil. Although buds on many plants appeared
swollen, only a single plant had open blossoms.
Discussion
That any plants should be flowering in January in
northern Nova Scotia, much less 23 species, is remark-
able. This late phenology may reflect the ability of
plants to have late flowering strains (e.g., Thlaspi ar-
vense, Warwick et al. 2002), or possibly reflect a west-
ern European origin where fall and early winter tem-
peratures would be much milder. Zinck (1998) does
not include December in the flowering periods of any
of these species; November is included in the flower-
ing periods only for Capsella bursa-pastoris, Matri-
caria matricarioides, Senecio vulgaris, Stellaria media
and Viola tricolor. The region falls within plant har-
diness zone 5B of Agriculture and Agri-Food Canada
(http://sis.agr.gc.ca/cansis/nsdb/climate/hardiness/intro.
html), in which harsh winters kill non-hardy species.
January is normally associated with severe winter
weather; the 30-year daily average temperature at Col-
GARBARY AND TAYLOR: FLOWERING DURING JANUARY 79
legeville (the nearest meteorological station for which
climate normals are available) is -6.6°C, and the aver-
age minimum is —11.6°C. Winters here are among the
coldest in the province (Davis and Browne 1997).
About 50 cm of snow falls on average in January, and
a heavy snowpack is common. Snowpack depths in a
wooded area of the St. Francis Xavier University cam-
pus were 40-50 cm in mid-January every year but one
in 1999-2005 (B. Taylor, unpublished data). In the
single low-snow year, the weather was very cold and
the ground was frozen.
The circumstances that led to plants blossoming in
mid-January were more complex than an extended peri-
od of warm weather. While fall and early winter of
2005-2006 were unusually mild, there had been reg-
ular nights of frost since November and two major
snowfalls (5 and 15 December) prior to the collection
period. The sites were visited over the course of three
weekends, and in the intervening periods there were
frosts on most night at Tracadie. These plant species
appear to be very resistant to short-term freezing, as
long as warm temperatures return quickly. In addition,
the most severe weather may have occurred when the
plants were covered with 20 cm of snow from the 15
December snowstorm. The snow would have insulat-
ed the plants from the deep frosts, but it was subse-
quently washed away by heavy rain.
None of the plants in bloom was restricted to the
first round of collecting; indeed, five species were found
in bloom only in the second round and ten species
had blossoms on 21 January. The two sites supporting
Erysimum cheiranthoides were snow-covered on 7 Jan-
uary and the single specimen of Lepidium campestre
found in week two may have been missed on the ear-
lier visit to the site. Only one site visited in this study
could be considered highly protected and influenced
by radiant heat from a building, and Cardamine pen-
sylvanica was the only species unique to this site. The
remaining sites were exposed to ambient air tempera-
tures and precipitation.
Nevertheless, even though plants in bloom were
found at 17 sites in mid-January, this was not a con-
spicuous phenomenon; surveys of additional sites re-
vealed no plants in blossom. The sites with plants in
flower comprised highly specialized habitats mostly of
limited extent: land cleared for development, margins
of abandoned or active agricultural fields, garden plots
and soil mounds. Even within these sites, most plants
of many species reported here had entirely succumbed
to the rigours of winter.
The plant species in bloom were not a cross-section
of the provincial flora. Most of the species reported
here are opportunistic, weedy species that thrive in
highly disturbed habitats, including agricultural set-
tings (e.g., Raphanus raphanistrum L., Warwick and
Francis 2005). Indeed many of our sites had less than
50% cover of vegetation. In addition, all species ob-
served in flower except Epigaea repens, Cardamine
pensylvanica and Cerastium vulgatum are non-native
80 THE CANADIAN FIELD-NATURALIST
members of the flora (Zinck 1998). Most of these
species are weeds that were introduced from Europe
with grain. It would appear that the ability to tolerate
harsh physical conditions (in contrast with competi-
tive ability) extends to tolerance of harsh weather as
well (Taylor and Garbary 2003).
Flowering in January in a cold climate would ap-
pear to be futile, and therefore likely of little evolution-
ary value. Excepting Epigaea repens, all of the species
reported here are annuals or short-lived perennials
(Zinck 1998). As early colonizers of disturbed ground,
they would be adapted to growing and reproducing
quickly in advance of competition from later species.
It seems likely that these species are adapted to grow,
flower and set seed as long as the weather permits (e.g.,
Senecio vulgaris, Robinson et al. 2003). This phenol-
ogy contrasts with that of longer-lived perennials in
which above-ground plant parts senesce in late autumn
in anticipation of winter. The ruderal species observed
here evidently lack this adaptation, presumably be-
cause either cold temperatures (annuals) or competi-
tion (perennials) normally curtails reproduction and
kills the plant. Consequently, when those events are
delayed, the plants have no physiological mechanism
to end flowering, even when seed production may be
impossible.
An important exception to the above is Pomquet
Beach Provincial Park, an undisturbed, coastal site
where Epigaea repens was part of a dense community
of ground vegetation beneath a diffuse tree layer. Unlike
the remaining fall-flowering species, Epigaea repens
normally flowers in April and May (Zinck 1998), when
it forms a conspicuous part of the spring-flowering
flora. The plants in flower in January may be consid-
ered either a second, late-fall flowering, or an excep-
tionally early spring flowering. This phenomenon con-
trasts with the situation of the remaining species, in
which normal late-fall flowering merely persisted for
a long time. It is doubtful that new flowers were initi-
ated among these species.
Epigaea repens was observed in flower in the An-
napolis Valley, about 250 km southwest of our study
area, in early November 2005 (N. Nickerson, Agricul-
ture and Agri-Food Canada, Kentville, personal com-
munication). Hence our flowering material from
Antigonish County probably should be considered a
further extension of a late fall return to flowering.
The same probably applies to Fumaria officinalis, for
which the latest flowering month given by Zinck (1998)
is August. It would be of interest to determine details
of plant phenology for E. repens and other species at
the extreme southern tip of the province, where Janu-
ary mean temperate is only just below freezing (plant
hardiness zone 6B; Davis and Browne 1997).
Vol. 121
Changes in plant phenology are indicators of climate
change and have been used as a proxy for tempera-
ture (Badeck et al. 2005; Menzel 2002; Sherry et al.
2007; Walther 2004). Consequently, the January flow-
ering records reported here might be considered as
evidence of regional climate warming. Thus, even if
regional climate warming has not been demonstrated,
the changes that we observed would be a natural con-
sequence of such a warming trend.
Acknowledgments
We thank Robert Garbary and Jonathan Ferrier who
assisted with field work and Nancy Nickerson for in-
formation on flowering of Epigaea repens. This work
was supported by grants from the Natural Sciences
and Engineering Research Council of Canada to both
authors.
Literature Cited
Badeck, F.-W., A. Bondeau, K. Boéttcher, D. Doktor, W.
Lucht, J. Schaber, and S. Sitch. 2005. Responses of
spring phenology to climate change. New Phytologist
162: 295-309.
Davis, D. S., and S. Browne (Editors). 1997. The natural
history of Nova Scotia, Volume 1. Topics and habitats.
Nimbus Publishing and Nova Scotia Provincial Museum,
Halifax, Nova Scotia. 518 pages.
Menzel, A. 2002. Phenology: its importance to the global
change community. Climate Change 54: 379-385.
Robinson, D. E., J. T. O’Donovan, M. P. Sharma, D. J.
Doohan, and R. Figueroa. 2003. The biology of Canadian
weeds. 123. Senecio vulgaris L. Canadian Journal of Plant
Science 83: 629-644.
Sherry, R. A., X. H. Zhou, S. L. Gu, J. A. Arnone, D. S.
Schimel, P. S. Verburg, L. L. Wallace, and Y. Q. Luo.
2007. Divergence of reproduction phenology under climate
warming. Proceedings of the National Academy of Sci-
ences of the United States of America 104: 198-202.
Taylor, B. R., and D. J. Garbary. 2003. Late-flowering plants
from northern Nova Scotia. Rhodora 105: 118-125.
Vasseur, L., R. L. Guscott, and P. J. Mudie. 2001. Moni-
toring of spring flower phenology in Nova Scotia: com-
parison over the last century. Northeastern Naturalist 8:
393-402.
Walther, G. R. 2004. Plants in a warmer world. Perspectives
in Plant Ecology Evolution and Systematics 6: 169-185.
Warwick, S. I., and A. Francis. 2005. The biology of Cana-
dian weeds. 132. Raphanus raphanistrum L. Canadian
Journal of Plant Science 85: 709-733.
Warwick, S. I., A. Francis, and D. J. Susko. 2002. The
biology of Canadian weeds. 9. Thlaspi arvense L. (up-
dated). Canadian Journal of Plant Science 82: 803-823.
Zinck, M., Editor. 1998. Roland’s flora of Nova Scotia, 3
edition. Nimbus Publishing Ltd. and Nova Scotia Provincial
Museum, Halifax, Nova Scotia. 1297 pages.
Received 24 January 2006
Accepted 15 January 2008
The Occurrence of Muskoxen, Ovibos moschatus in Labrador
| Tony E. Cuusss! and J. BRAZIL*
'Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and
Labrador AOP 1SO Canada; e-mail: techubbs @cablelab.net
*Endangered Species and Biodiversity Section, Wildlife Division, Department of Environment and Conservation, Govern-
ment of Newfoundland and Labrador, P.O. Box 2007, Corner Brook, Newfoundland and Labrador A2H 7S! Canada
Chubbs, Tony E., and J. Brazil. 2007. The occurrence of Muskoxen, Ovibos moschatus in Labrador. Canadian Field-Natu-
ralist 121(1): 81-84.
Muskoxen were first observed in Labrador in 1988 and have since been recorded at several locations in two separate river
valleys in northern Labrador. The occurrence of Muskoxen in Labrador is attributed to the dispersal of captive releases from
/ Québec in 1973 and 1983. Here we document observation records and examine the future management of the species in
The Muskox (Ovibos moschatus) is a relic from
‘the Pleistocene and is native to northern Canada,
‘northwestern Greenland, and the northern coast of
Alaska (Banfield 1974; Reynolds 1988). The elimi-
“nation of the species from Alaska and the reduction
~of Muskox numbers in Canada by the early 1900s re-
‘sulted in efforts to introduce and restore populations
‘in several arctic areas (Klein 1988; Reynolds 1988).
At present, Muskox populations are experiencing a
period of success, with high numbers throughout most
‘of their range (Groves 1997). Muskox apparently did
‘not colonize the Québec-Labrador peninsula after the
i last glaciation as no remains have been found at archae-
_ dlogical sites (Banfield 1975).
' In 1967, 15 Muskoxen (12 females, 3 males) were
‘captured near Eureka, Ellesmere Island (Centre D’E-
vudes Nordique 1981*), and brought to the Uming-
maqautik farm, near Kuujjuag in northern Québec
_ Figure 1) for Inuit to establish a Muskox farm, cre-
Ate employment, and build up a stock to introduce the
Bpecies into northern Québec (Le Hénaff and Créte
1989; Nault et al. 1993). A project of raising the ani-
“nals to collect their quviut (Muskox wool) was unsuc-
>essful. Subsequently, from 1973 to 1983, 54 animals
vere released at three different locations in northern
Québec (Le Hénaff and Créte 1989). The first release
sites were located to the north of Kuujjuaq and to the
Benuivest of Tasiujaq (Le Hénaff and Créte 1989). In
+983, Muskoxen were released east of Kuujjuag near
he outlet of the Whale River. The Umingmagqautik
arm was closed in 1983 and the remaining animals
vere given to the St-Félicien and the Orsainville zoos
-n Québec (Jean and Rivard 2005*).
__ The introductions in Québec proved successful. In
_ 983 a total of 148 Muskoxen were counted and by
986, with a finite rate of increase of 1.25, the popu-
_ ation had increased to 290 animals. By 1986, indi-
viduals and mixed groups of Muskoxen had dispersed
‘nall cardinal directions up to 650 km from the origi-
Labrador and speculate on its possible effect on this northern ecosystem.
/ Key Words: Muskox, Ovibos moschatus, range, distribution, management, Labrador, Québec.
nal release sites throughout the province of Québec
(Le Hénaff and Créte 1989).
In 2003, the Muskox population around Kuujjuaq
and Tasiujaq was estimated to be 1400 animals, includ-
ing calves (Jean and Rivard 2005*). In June 2005, aer-
ial surveys on Diana Island, near Quaqtaq, counted a
total of 112 adults and 25 calves. This new population
apparently originated from the Tasiujaq area (Jean and
Rivard 2005*).
Study Area
Over the past two decades, Muskoxen have been ob-
served on several occasions in the Low Arctic Alpine-
Torngat ecoregion in northern Labrador. This ecore-
gion has short, cool summers and long, cold winters
with mean average daily temperatures ranging from
-19 to -22°C in February and 7 to 9°C in July. Aver-
age annual precipitation is 500 to 700 mm with a total
average snowfall of 3 m (Meades 1990*). The vegeta-
tion is tundra (Alpine heath) with sedge meadows domi-
nating plateaus and Mountain Alder (Alnus crispa) and
willow (Salix spp.) thickets dominating on the lower
slopes and river valley slopes. Groves of White Birch
(Betula papyrifera) and Balsam Poplar (Populus bal-
samifera) occur sporadically in some river valleys.
Graminoids occur on the valley bottoms but are sparse
to absent on the upper slopes and at high elevations.
Dwarf Birch (B. glandulosa), Labrador Tea (Ledum
groenlandicum) and Black Crowberry (Empetrum
nigrum) dominate drier mountain ridges. Conifer trees
are absent from this region (Meades 1990*). Major
features of the terrain include braided east-flowing
rivers, foothills, elevated plateaus, rugged coastal cliffs
and inland mountains.
Methods
We recorded Muskox sightings during remote aeri-
al field surveys in northern Labrador. We queried res-
idents from remote communities throughout northern
81
82 THE CANADIAN FIELD-NATURALIST
Labrador and helicopter and fixed-wing aircraft pilots
for sightings of Muskoxen. We found only one record
of Muskoxen in other areas of Labrador apart from
our own observations. We classified individual Muskox
to age and sex from oblique aerial photographs. When
possible, animals were classified as adult male or fe-
male, based on body size, coat color and size and shape
of horns.
Observations
Five sightings of Muskoxen have been recorded in
Labrador over the past two decades in three different
river valleys, approximately 340 km apart. Four sight-
ings were made from a helicopter in different years
(1988, 1995, 2000, 2005) and included three sight-
ings of a single Muskox, likely different animals, and
one sighting of a group of three. The fifth sighting
(2006 — 2008) was reported by residents of Nain while
travelling by snowmobile to Voisey’s Bay.
Site 1. Komaktorvik River Valley
The first sighting of Muskoxen in Labrador was
recorded on 24 July 1988 at approximately 59°14'N,
63°56'W along the Komaktorvik River valley, which
flows into Seven Islands Bay (Figure 1). Three indi-
viduals were observed in a large patch of willows (Salix
spp.) in the river valley. All individuals had horns with
a slight separation in the butte and were all approxi-
mately the same size indicating that these animals were
likely adult males, although definite sex and age remain
uncertain (J. Brazil, personal observation).
On 28 July 1995, one Muskox was again observed in
the Komaktorvik River valley (59°14.65'N, 63°55.51'W)
very near the original 1988 sighting. From a photo-
graph, this animal, browsing in a large thicket of wil-
lows (Salix spp.) along the north slope of the river val-
ley, appeared to be an adult male.
On 30 July 2000, during aerial surveys for Harle-
quin Ducks (Histrionicus histrionicus) and Peregrine
Falcons (Falco peregrinus), we observed and pho-
tographed a lone Muskox in Seven Islands Bay at
(59°14.16'N, 63°57.66'W) 3 to 4 km west of the previ-
ous sightings in 1988 and 1995. This animal was large,
had well formed horns, and light coloration on the back
and saddle and was identified as a sub-adult male (Ole-
sen and Thing 1989).
The sightings along the Komaktorvik River valley
occurred approximately 250 km northeast of the re-
lease site at Whale River providing a straight-line rate
of dispersion of 50 km/year. The most probable dis-
persion route was along the Koroc River valley, which
traverses the Labrador Peninsula.
Site 2. Hebron Fiord
On 9 June 2004, a single Muskox was observed near
the outlet of Hebron Fiord (58°03.82'N, 63°12.71’ W)
(Figure 1, Site 2). This animal was feeding in a thick-
et of alders (Alnus spp.) and Willows (Salix spp.) at
the foot of a small plateau on the north side of Hebron
Brook where it flows into Hebron Fiord. A photo-
Vol. 121
graph identified this animal as an adult male > 4 years
old (D. Jean, personal communication). The straight-
line dispersal from the Whale River release site was a
distance of 220 km providing a rate of dispersion of
approximately 11 km/year. The most probable route
of dispersion was south along the George River val-
ley and then the Tasirlaq River to Hebron Fiord.
Site 3. Voisey’s Bay
In April 2006, the fifth Muskox reported in Lab-
rador was observed on a small unnamed island south
of Tabor Island in Voisey’s Bay at approximately
54°20'N, 61°43'W (Figure 1, Site 3). This animal was
readily identifiable as its right horn tip was broken at
the curl. This 4-5 year old male has remained on the
island and is in its second winter at this location. The
Straight-line dispersal from the Whale River release
site is a distance of 400 km providing a rate of dis-
persion of approximately 17 km/year. The most prob-
able route of dispersion was south along the George
River valley and then along either the Fraser or Kogaluk
river to Voisey’s Bay.
Discussion
Muskoxen have successfully dispersed and colo-
nized many remote regions of northern Québec. The
dispersal of Muskoxen from Québec into Labrador
may have been influenced by unoccupied habitat and
lack of dispersal barriers.
No mixed-aged groups (the reproductive segment of
the population) have been observed to date in Labrador.
It is thus likely that the separate observations at Kom-
aktorvik River valley, Hebron Fiord and Voisey’s Bay
are of different animals that immigrated to Labrador
through separate overland routes and at different times.
Based on the dispersion of Muskox south along the
George River in Québec through 1986 (Le Hénaff
and Créte 1989), it is possible that other individuals
or groups have traveled eastward into Labrador as far
south as Nain along the numerous river valleys ex-
tending inland.
Although Muskoxen and Caribou (Rangifer taran-
dus) are occasionally found together on the same veg-
etation type, caribou usually move through an area
rapidly, feeding on willows and flowering forbs where-
as Muskoxen are more sedentary and forage on sedges
and grasses (Parker 1978). Abundance and distribu-
tion of Muskoxen on many of the islands of the High
Arctic appear to be related to the abundance and dis-
tribution of sedge-producing meadows (Parker et al.
1975; Parker and Ross 1976).
Initial colonization by Muskox may be slow and
sporadic consisting mainly of adult males (Le Hénaff
and Créte 1989; Reynolds 1988). This may explain why
reproductive, mixed-aged groups have do not appear
to have been observed yet in Labrador. As Muskoxen
had populated many remote regions of Québec in all
cardinal directions north of 54°30'N latitude by 1986
(Le Hénaff and Créte 1989) and populations continue
2007
CHUBBS and BRAZIL: MUSKOXEN IN LABRADOR 83
Labrador
Legend
Sightings
Communities
Rivers
FiGuRE 1. Location of Muskox (Ovibos moschatus) sightings in northern Labrador. The two release sites were near Tasiujagq
and Kujjuaq in Québec.
to grow (lambda = 1.25) near their maximum finite
rate of increase (1.30), breeding groups may have be-
come established in Labrador but have gone undetect-
ed. Emigration of mixed-sexed groups out of regions
first occupied may result in shifts in population dis-
tribution and range expansion (Reynolds 1988). The
dispersal and colonization of Muskoxen to adjacent
ranges can be slow. However, with the present high rate
of population increase (Jean et al. 2004*) in northern
Québec, we believe that emigration may soon result in
a shift in an expansion of the distribution and range
of Muskoxen into Labrador. Reynolds (1988) identi-
fied three stages in the expansion of Muskox into new
range: (1) slow growth following release (2) the irrup-
tive phase and (3) decline and stabilization. It is likely
that in the next decade during the decline and stabi-
lization of Muskox numbers in Québec that the dis-
persal of mixed groups into Labrador will occur and
a breeding population be established.
In Alaska, where Muskoxen were extirpated and later
reintroduced, range expansion occurred only over 20%
of the former Muskox range and populations grew to
fewer than 1000 individuals over nearly 30 years
(Reynolds 1988).
Management Implications
Hunting of Muskox under a limited permit system
is conducted on Nunivak and Nelson islands and in
the Arctic National Wildlife Refuge of Alaska and in
Nunavut, Northwest Territories, and Québec in Can-
ada. Muskoxen are considered a unique and valuable
trophy by the outfitting industry and Muskox meat is
highly valued as an exported delicacy. This survivor
of the ice ages is an important attraction for tourists,
photographers, researchers and students of wildlife in
areas where the species is endemic or introduced.
The establishment of a Muskox population in Labra-
dor may provide an alternative to Caribou as a source
of protein for northern communities as well as a new
resource for developing the regional economy through
sport hunting similar to northern Québec and Alaska
(Le Hénaff and Créte 1989; Jean et al. 2004*; Reynolds
1988).
It appears that habitat for Muskoxen is widely dis-
tributed and largely unchanged in northern Labrador.
With public support and proper management, Muskox-
en may eventually become a more visible and famil-
iar wildlife species in Labrador. We recommend that
systematic aerial surveys be conducted to document the
current and future status and distribution of Muskoxen
in Labrador.
Acknowledgments
Thanks are extended to D. Jean, Ministére des Res-
sources naturelles et de la Faune, who provided liter-
ature on Muskoxen introduction and releases in Québec
84 THE CANADIAN FIELD-NATURALIST
as well as assistance in sexing and aging Muskoxen
from photographs. M. Créte, F. Phillips, G. Parker, the
late N. P. P. Simon and an anonymous reviewer pro-
vided comments on earlier drafts of the manuscript. L.
Elson drafted the figure. The Department of Environ-
ment and Conservation, Government of Newfoundland
and Labrador, provided funding for this publication.
Documents Cited (marked * in text)
Centre D’études nordiques. 1981. Rapport d’évaluation
du projet d’élevage de boeuf musqué, Umingmagautik,
Nouveau-Québec. (1967-1980). Centre d’ Etudes nordiques
et Faculté des sciences de l’agriculture et de l’alimentation,
Université Laval. Présenté au ministére de I’ Agriculture,
des Pécheries et de |’ Alimentation du Québec, Québec.
168 pages.
Jean, D., and S. Rivard. 2005. Inventaire de la population de
boeeufs musqués (Ovibos muschatus) sur Vile Diana, prés
de Quagtaq. Ministére des Ressources naturelles et de la
Faune — Secteur Faune Québec, Direction de l’aménage-
ment de la faune du Nord-du-Québec. 8 pages.
Jean, D., S. Rivard, and M. Bélanger. 2004. Inventaire et
structure de population du boeuf musqué (Ovibos mos-
chatus) au sud-ouest de la baie d’Ungava, aott 2003.
Ministére des Ressources naturelles, de la Faune et des
Parcs — Secteur Faune Québec, Chibougamau. 22 pages.
Meades, S. J. 1990. Natural regions of Newfoundland and
Labrador. Protected Areas Association, St. John’s, New-
foundland and Labrador. 103 pages.
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Banfield, A. W. F. 1975. Les mammiferes du Canada. Musées
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Groves, P. 1997. Intraspecific variation in mitochondrial
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Klein, D. R. 1988. The establishment of Muskox populations
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Le Heénaff, D., and M. Créte. 1989. Introduction of muskoxen
in northern Québec: the demographic explosion of a colo-
nizing herbivore. Canadian Journal of Zoology 67: 1102-
1105.
Nault, R., C. Mathieu, and M. Créte. 1993. Vegetation bio-
mass and habitat selection by a newly introduced population
of muskoxen in northern Québec. Rangifer 13: 71-77.
Olesen, C. R., and H. Thing. 1989. Guide to field classifi-
cation by sex and age of the Muskox. Canadian Journal
of Zoology 67: 1116-1119.
Parker G. R. 1978. The diets of muskoxen and Peary caribou
on some islands in the Canadian High Arctic. Canadian
Wildlife Service Occasional Paper (35). 21 pages.
Parker G. R., and R. K. Ross. 1976. Summer habitat use by
muskoxen (Ovibos moschatus) and Peary caribou (Rangi-
fer tarandus pearyi) in the Canadian High Arctic. Polar-
forschung 1976(1): 12-25.
Parker, G. R., D. C. Thomas, E. Broughton, and D. R.
Gray. 1975. Crashes of muskoxen and Peary caribou pop-
ulations in 1973-74 on the Parry Islands, Arctic Canada.
Canadian Wildlife Service Progress Note (56). 15 pages.
Reynolds, P. 1988. Dynamics and range expansion of a re-
established Muskox population. Journal of Wildlife Man-
agement 62: 734-744.
Received 15 March 2006
Accepted 8 March 2008
The Spindle Tree, Euonymus europaea L. (Celastraceae): A Newly
Naturalized Shrub in Nova Scotia
Davip J. GARBARY! and Amy E. DEVEAU
'Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2WS5 Canada; e-mail: dgarbary @ gmail.com
Garbary, David J., and Amy E. Deveau. 2007. The Spindle Tree, Euonymus europaea L. (Celastraceae); a newly naturalized
shrub in Nova Scotia. Canadian Field-Naturalist 121(1): 85-88.
Over 100 plants, including over 750 individual shoots, of Euonymus europaea L. are reported from the Town of Antigonish
in northern Nova Scotia. Plants were found in five discrete areas separated by 200-1000 m. A few individuals at one site
may represent human planting; however, over 90 % of plants occur on waste ground or in dense shrubbery where natural
colonization has occurred. Many individuals were observed in fruit in the fall of 2005, and over 95% of the plants had well
developed buds and flowers in May and early June 2006. Euonymus europaea is considered a newly naturalized species for
the flora of Nova Scotia.
Key Words: Spindle Tree, Euonymus europaea, introductions, naturalized, Nova Scotia.
Euonymus (Celastraceae) is a cosmopolitan genus
of over 200 species of vines, deciduous and evergreen
shrubs, and small trees (Gleason and Cronquist 1991).
Many species are ornamentals because of their col-
ourful fruits and fall foliage and their unusual winged
stems. At least six species are recorded from Canada,
of which one is regarded as doubtful (Scoggan 1978;
Catling 1997; Hinds 2000). Only E. occidentalis Nutt.
ex Torr. (Western Wahoo), E. atropurpurea Jacq. (East-
em Wahoo) and E. obovata Nutt. (Running Strawberry-
Bush) are native in Canada, with E. occidentalis res-
tricted to the west coast. Euonymus europaea L., E.
fortunei (Turcz.) Hand.-Mazz., E. alata (Thun.) Sie-
bold, and E. nana Bieb. are escapes from cultivation in
eastern Canada, although they have restricted distri-
butions in New Brunswick (Catling 1997; Hinds
2000). Other species occur in the northeastern United
States (i.e., E. alata and E. americana L., Gleason and
Cronquist 1991). No species is included in the flora
of Nova Scotia (Zinck 1998).
Euonymus europaea was first noted in Antigonish as
a single, non-cultivated plant in 2001 in Antigonish
County, Nova Scotia (DJG unpublished). In a sub-
sequent survey during fall and early winter of 2005-
2006 (Garbary and Taylor 2007), fruiting plants of E.
europaea were noted from the previously recognized
site as well as additional sites in Antigonish. The spe-
cies likely represents an escape from cultivation. Here
We report E. europaea as a newly naturalized plant in
Nova Scotia.
Materials and Methods
During the course of plant surveys in late fall and
early winter of 2001 and 2005-06 (Taylor and Garbary
2003; Garbary and Taylor 2007), fruiting specimens
of Euonymus europaea were noted. In May 2006, when
flower buds were well developed but unopened, a sur-
vey of all previously identified locations was under-
taken. Further surveys were conducted to determine
the size of the overall population in Antigonish and
the extent to which the species was reproducing. The
four primary sites are designated as University, Clay-
more, Hospital, and Brierly Brook (see Table 1). Two
additional sites with only 1-2 primary plants were
found but are not included in Table 1.
Sites were surveyed with hand-held GPS units (Gar-
min 12, Olathe, Kansas, USA). The largest stem in any
cluster was considered the primary plant, and adjacent
shoots were assumed to be derived from the same
base, or progeny from seed. The height of the largest
plant at each site was measured with a meter stick, and
the numbers of subsidiary shoots and nearby juveniles
were counted. These data were gathered to demonstrate
that plants were successfully reproducing in the area.
For each primary plant, the slope, exposure to full sun-
light, proximity to water (ditches, ponds or saturated
soil) and reproductive condition were recorded. The
number of flower buds on each peduncle was counted
on the population on St. Francis Xavier University.
Voucher specimens are deposited in the herbarium of
St. Francis Xavier University (STFX) with duplicates
deposited in herbariums of ACAD (Acadia University)
and NSPM (Nova Scotia Provincial Museum).
Results
With its opposite leaves on short petioles, the longi-
tudinal whitish markings on many young stems that
often gives them a four-angled outline, and the bril-
liant scarlet and orange fruits, Euonymus europaea 1s
a distinctive species during all seasons (Figure 1).
Most primary shoots (> 95%) and many adjacent
smaller plants had abundant clusters of flower buds in
mid- to late May 2006. The bud scales of branch buds
and developing inflorescences had red tips and a ciliate
85
86 THE CANADIAN FIELD-NATURALIST
Vol. 121
FIGURE 1. Euonymus europaea from Antigonish. A) Branch with leaves, developing inflorescences and bud scale (arrow).
Scale = 2 cm. B) Inflorescence with central bud partially open. Scale = 1 cm. C) Fully open flower. Scale = 5 mm.
D) Mature fruit after dehiscence showing single seed. Scale = 1 cm. E) Longitudinal view of stem showing leaf scar
and longitudinal ridge on bark (arrow). Scale = 2 cm. F) Transverse section of stem showing squarish outline and
position of longitudinal ridges. Scale = 5 mm.
margin. Flowers did not open until late May when a
single bud in each peduncle opened. The flowers are
not conspicuous, and plants are cryptic against a back-
ground of the larger and showier species of Prunus,
Crataegus and Amelanchier. The flowers are ca. 1 cm
diameter, and have diagnostic greenish-white petals.
The flowering peduncles had from 1-9 buds (5.0 + 1.8),
with most inflorescences having odd numbers of buds.
Over 100 primary plants over 50 cm high and over
650 secondary shoots and adjacent progeny were found
at six locations in Antigonish (Table 1). The primary
axes were 1.5 + 0.8 m (mean + s.d.) tall, and many
plants were over 3 m (Figure 2) with a trunk diameter
of up to 6 cm. There was wide variation (0 — >50) in
number of secondary shoots and progeny associated
with each primary plant, although about 30% had
2007
Frequency
o HE
Din EASES laiae Bian
ee os |
spr es sree ht ae Shy YO SI
NPI oR <= EN oe =e! |
Set eatin VA cA
Plant height (m)
FIGURE 2. Histogram showing sizes of 112 primary stems of
Euonymus europaea from Antigonish.
only 1-2 additional axes (Figure 3). The largest single
clump was at the Brierly Brook site. It occupied about
4 x 3 m and had about 50 separate stems that formed
a dense thicket.
Plants were typically found in the open where they
experienced full sun, or at the edges of dense shrub
thickets where they were either crowded or overtopped
by the surrounding plants. Most plants grew in moist
locations. Plants in the open tended to occur on bogg
ground, and plants were common close to ditches,
ponds or small pools or on the flood plain of a
stream. Some plants were present on steep slopes on
ground with obvious seeps. Only a few plants occurred
on drier soils; these were in grass on mowed areas
and may have been planted.
GARBARY and DEVEAU: SPINDLE TREE IN NOVA SCOTIA 87
50
40
30
20
10 y
Sy
rt
|
Number of shoots
FIGURE 3. Histogram showing number of secondary shoots and
possible progeny associated with each primary plant.
Discussion
All of the Euonymus plants we observed were E.
europaea. The plants are scattered over several square
kilometres in at least five separate sites. The abundance
of small plants within 3 m of larger individuals and
widely scattered small plants (1.e., less than | m) show
that successful seed germination is occurring. These
observations are consistent with the definition of na-
turalized of Pysek et al. (2004); i.e., that successful
reproduction has been occurring for at least ten years
in the vicinity of the parent plant.
The site where most of the plants were found is an
open field where EF. europaea is mixed with many other
shrubs. These shrubs are mostly widely scattered and
TABLE |. Populations of Euonymus europaea in Antigonish. Additional sites with only one or two primary plants not included
in the table.
Hospital
Latitude and longitude 45°37.743'N
61°59.033'W
Number of primary plants 12
Secondary shoots and juveniles 61
Number with flower buds 11/12
Habitats Steep slope with
extensive bed of lupins;
mowed and unmowed
grassy area
Shade Full sun
Claymore Brierly Brook
45°37.056'N 45°37.394'N
61°58.877'W 61°59.179'W
4 5
11 95
4/4 5/5
Shrub thicket Border of
beside hayfield at
small stream margin of thicket
adjoining stream
Partial to heavy
shade
Partial to high
shade
University
45°37.074'N
61°59.182'W
90
683
87/90
Open areas with
herbaceous plants
and shrubs: on flat
ground to steep
slopes
Full sun but some
heavily shaded
88 THE CANADIAN FIELD-NATURALIST
have an overall cover of less that 15%. The occasional
individuals that we observed in dense shrubbery had
few secondary shoots. Thus, once vegetation closes in,
E. europaea may not be able to establish or spread.
Species with fleshy fruits have low ability to recruit
under a canopy of E. europaea relative to nine other tall
shrub species (Kollmann and Grubb 1999). This has
not yet become important in Antigonish, since we have
observed only one continuous canopy of E. europaea.
Some Euonymus species, including E. alata, have
become invasive in several northeastern and mid-
western states (e.g., http://tncweeds.ucdavis.edu/alert/
alrteuon.html) following escape from ornamental
plantings. FE. alata is considered a threat in woodland
areas, fields and coastal scrubland, where it can out-
compete native species. There is no record of E.
europaea as a problematic invasive species, although
it has become widely naturalized in central Canada.
Montgomery (1957) refers to sites where it has be-
come “very common” in woods. The abundance of
mature and immature individuals in Antigonish sug-
gests that this is possible for E. europaea in Nova
Scotia. The extensive field survey carried out by DJG
in the fall and early winter of 2005-2006 identified four
of the sites reported here. Additional surveys in 2006-
2007 did not reveal further populations. While there
are no specimens based on wild collections of E.
europaea in other local herbaria (1.e., NSPM, ACAD,
NSAC [Nova Scotia Agriculture Canada]), other urban
areas may be expected to host wild plants (e.g., Wolf-
ville, Nova Scotia — Sam Vander Kloet, personal com-
munication). To satisfy the Pysek et al. (2004) defini-
tion of invasive, many offspring must be produced at
a distance from the parent individuals. The limited dis-
persal of E. europaeus in Antigonish has not yet satis-
fied this criterion.
Acknowledgments
We thank Greg Power, Kelly Clement and Meghan
Hines for assistance with fieldwork, Randy Lauff who
Vol. 121
pointed out the location of an additional Euonymus
site, and Barry Taylor for comments on the manuscript.
Curators of ACAD, NSPM and NSAC provided in-
formation and access to collections. This research was
supported by research grants from the Natural Sci-
ences and Engineering Research Council of Canada
to DJG.
Literature Cited
Catling, P. M. 1997. The problem of invading alien trees
and shrubs: some observations in Ontario and a Canadian
checklist. Canadian Field-Naturalist 111: 338-342.
Garbary, D. J., and B. R. Taylor. 2007. Plant flowering dur-
ing January in Antigonish County, Nova Scotia. Canadian
Field-Naturalist 121(1): 76-80.
Gleason, H. A., and A. Cronquist. 1991. Manual of vascular
plants of Northeastern United States and adjacent Canada,
2™4 edition. New York Botanical Garden, Bronx, New York.
Hinds, H. R. 2000. Flora of New Brunswick. University of
New Brunswick, Fredericton, New Brunswick. 695 pages.
Kollmann, J., and P. J. Grubb. 1999. Recruitment of fleshy-
fruited species under different shrub species: control by
under-canopy environment. Ecological Research 14: 9-21.
Montgomery, F. H. 1957. The introduced plants of Ontario
growing outside of cultivation. Transactions of the Royal
Canadian Institute 32: 3-35.
Pysek, P., D. M. Richardson, M. Rejmanek, G. L. Webster,
M. Williamson, and J. Kirschner. 2004. Alien plants in
checklists and floras: towards better communication be-
tween taxonomists and ecologists. Taxon 53: 131-143.
Scoggan, H. J. 1978. The Flora of Canada Part 3 — Dicotyle-
donae (Sauruaceae to Violaceae). National Museum of
Natural Sciences, National Museum of Canada, Ottawa.
568 pages
Taylor, B. R., and D. J. Garbary. 2003. Late-flowering plants
from northern Nova Scotia. Rhodora 105: 118-135.
Zinck, M. Editor. 1998. Roland’s Flora of Nova Scotia, 3"
edition Nimbus Publishing Ltd. and Nova Scotia Museum,
Halifax, Nova Scotia, Canada. 1297 pages.
Received 16 June 2006
Accepted 7 March 2008
Notes
Serviceberry, Amelanchier intermedia, Escaped from Cultivation in
Eastern Ontario
PAUL M. CATLING and GISELE MITROW
Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Saunders Building, Central Experimental Farm, Ottawa,
Ontario KIA 0C6 Canada; e-mail: catlingp @agr.ge.ca
Catling, Paul M., and Gisele Mitrow. 2007. Serviceberry, Amelanchier intermedia, escaped from cultivation in eastern Ontario
Canadian Field-Naturalist 121(1): 89-91.
Although not generally recognized as a cultivated plant, Amelanchier intermedia is widely cultivated in eastern Ontario
under the incorrect name, A. canadensis. Wild plants referable to A. intermedia in old fields south of Ottawa are likely to be
escaped from cultivation. Amelanchier intermedia likely originated as a hybrid involving A. canadensis and A. laevis. A key
to the Canadian species of Amelanchier with glabrous ovaries is provided.
Key Words: Saskatoon, Juneberry, Serviceberry, Shadbush, Amelanchier intermedia, status, cultivation, hybrid, Ontario.
Fernald (1950) and Kartesz and Meachum (1999)
reported Amelanchier intermedia Spach from New-
foundland to Minnesota and south to North Carolina.
It was not included in Michigan by Voss (1985), but
Cing-Mars (1971) showed a relatively extensive distri-
bution in southern Quebec. Although within the gen-
eral range outlined by Fernald (1950) and frequent in
adjacent Quebec (Cingq-Mars 1971), A. intermedia was
ig reported in Ontario by Soper and Heimburger
(1982) or McKay (1973). Although it was listed for
Ontario by Newmaster et al. (1998*), the listing was
without ranking indicating a lack of information on its
‘status. Over several years of examining populations of
Amelanchier in Ontario we have found plants that cor-
‘respond to the description of A. intermedia at only two
locations 1 in old field habitats south of Ottawa. Plants
similar to these wild plants are widely cultivated but
are never called A. intermedia.
' Most eastern Ontario nurseries offering shrubs for
sale offer Serviceberry (also called Juneberry, Shad-
bush and Saskatoon, Amelanchier spp.). The plants that
are most frequently available are either multi-stemmed
‘shrubs labelled as A. canadensis or they are named
cultivars referable to A. xgrandiflora Rehder, a hybrid
of A. arborea (Michx. f.) Fern. and A. laevis Wieg.
The plants referred to as Amelanchier canadensis (L.)
‘Medic. are particularly popular with companies pro-
‘viding landscaping for office buildings, whereas the
larger small trees are increasingly popular for use as
‘Street trees under utility wires (Gerhold 1999).
“>
Identification of wild and similar cultivated
' plants
__ With petals 9-13 mm long and reddish-tinged leaves
prominently acute at the tip, the plants growing in old
os
fields and woodland edges south of Ottawa (cited be-
low) and similar plants labelled as A. canadensis in
10 local nurseries as well as many plants in Ottawa gar-
dens could not be placed with A. canadensis, which has
petals 3-10 mm long and green young leaves with leaf
tips more or less rounded (Cruise 1964; Cing-Mars
1971). They are not A. /aevis because the leaves are
tomentose at flowering instead of glabrous. Nor are
they A. arborea because these are multi-branched shrubs
with reddish-tinged leaves and flowers with a rela-
tively broad hypanthium 4.5—5.5 mm across instead
of single-trunked trees with green leaves and flowers
with a relatively narrow hypanthium 2.5—3 mm across.
They also differ from A. arborea in having sepals that
are less reflexed and lanceolate with a concave outer
edge instead of strongly reflexed at the base and oblong-
lanceolate with the outer edge convex. In addition the
berries of these shrubs are pleasant-tasting rather than
insipid as in A. arborea. These plants do correspond to
A. intermedia Spach of Fernald (1950) and Cing-Mars
(1971).
While it is clear that A. intermedia occurs both as
a cultivated plant and outside of cultivation in eastern
Ontario, it is not clear whether or not the wild plants
are escapes from cultivation. It seems most likely that
they are escaped since one of the parents, A. canaden-
sis, does not occur in the local area and the similar
widely cultivated plants have become increasingly pop-
ular over the past few decades and are currently widely
available in nurseries and widely used by landscaping
companies. A key to the group with glabrous ovaries is
provided below.
Specimens referable to A. intermedia growing out-
side of cultivation in Ontario include: OTTAwA-CAR-
LETON: open, sandy ground 3 km N of Herbert’s Cor-
89
90 THE CANADIAN FIELD-NATURALIST
ners, Osgoode Twp., 45.2359°N, 75.5733°W, 17 May
2002, P. M. Catling s. n. (DAO); UNITED COUNTIES OF
STORMONT, DUNDAS AND GLENGARRY: open old field,
2 km WNW of Ormond and about 5 km N of Winches-
ter at 45.1552°N, 75.4173°W, 21 May 2005, P. M.
Catling 2005-41, 2005-42 (DAO). Cultivated speci-
mens referable to A. intermedia collected as part of this
study in Ottawa area gardens are also preserved in DAO
(see DAO 798819, 801264, 801265, 801272, 801275,
801280, 801328, 801329, 801330, 801332) and vouch-
ers obtained from nurseries are also in the collection
(see DAO 800923, 800924, 800925, 801262, 801267,
801269, 801270, 801271, 801333, 801335).
Taxonomic history and possible hybrid origin
of A. intermedia
Jones (1946) included the name intermedia with his
relatively narrowly defined A. canadensis showing a
largely eastern coastal plain distribution. Wiegand
(1920) used the name to represent a species related to
A. canadensis which he defined narrowly. Later Fer-
nald (1950) expanded the concept to include plants
clearly intermediate between A. canadensis and A. lae-
vis and this concept was maintained in the classic work
of Cing-Mars (1971).
Amelanchier intermedia may have arisen as a hybrid
of A. canadensis and A. laevis as noted by various
authors (e.g., Gleason and Cronquist 1991). Plants
referable to A. intermedia have leaves that are only
moderately pubescent and losing their pubescence at
flowering. This and their purplish tinge suggests that
one parent is a species with leaves purple-green and
glabrous and unfolded at flowering. The only species
with a glabrous ovary fitting this description is A. laevis.
The relatively short petal lengths and also the widely
open hypanthium of the putative hybrids suggest that
the other parent may have short petals, barely reflexed
sepals and a saucer-shaped hypanthium. This is true of
A. canadensis but not A. arborea which has relatively
long petals, prominently reflexed sepals and a rather
narrow, campanulate hypanthium. Furthermore a cross
of A. arborea and A. laevis would be expected to pro-
duce a more tree-like hybrid, one form of which, A.
xgrandiflora Rehd. is well known. Other species of
Amelanchier have pubescent ovary summits and/or
rounded and more coarsely toothed leaves and are thus
excluded from consideration as putative parents. Ame-
lanchier canadensis x laevis hybrids have been report-
ed to be common in some areas, whereas hybrids of A.
arborea and A. canadensis are less common (Cruise
1964).
The only material that can be confused with Ame-
lanchier interior that occurs in the lower Ottawa valley
are depauperate specimens of A. arborea which differ
in their early flowering with unexpanded and very hairy
leaves and apparent hybrids of A. /aevis and A.spicata
Vol. 121
which differ in their broad spatulate petals and partially
hairy ovaries. The typical habitat of these latter plants
is dried and disturbed heath bogs where A. spicata also
occurs. Such plants may have been the basis for reports
of A. intermedia from edges of bogs in Minnesota
(Nielsen 1939). Alternatively these reports may have
been based on fruiting material of A. interior of the
northwestern Great Lakes region which had lost some
of the pubescence on the ovary (as found to be the case
for reports of A. intermedia from northwestern Ontario).
The following key will readily separate flowering A.
intermedia from related Canadian species.
Key to Canadian flowering Amelanchier with
glabrous ovaries:
la. Leaves glabrous and purple at flowering;
petals 10-20 mm long; lowest pedicels
25-S 0 MMVON Sees eto Aesth eee A. laevis
1b. Leaves tardily pubescent and green or
purple-tinged at flowering; petals 3-15 mm long;
lowest pedicels 8-35 mm long
2a. Petals 3-10 mm long; leaves
FOUNG=(ippPed eee eee A. canadensis
2b. Petals 9-15 mm long; leaves pointed at tip ......... B)
3a. Leaves purple-tinged, more or less expanded and
becoming glabrous beneath during flowering;
SHEUDS 4c SHES as eT Ee, se A. intermedia
3b. Leaves green, small and folded at flowering,
pubescent beneath after flowering; trees ...A. arborea
Acknowledgments
Brenda Kostiuk assisted with field studies. W. J. Cody
provided useful comments on the manuscript.
Document Cited (marked * in text)
Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. McMurray,
and M. J. Oldham. 1998. Ontario plant list. Ontario Forest
Research Institute, Ontario Ministry of Natural Resources
(Sault Ste. Marie, Ontario), Forest Research Information
Paper 123. www.uoguelph.ca/foibis.
Literature Cited
Cing-Mars, L. 1971. Le genre Amélanchier au Québec. Le
Naturaliste canadien 98: 329-346.
Cruise, J. E. 1964. Studies of natural hybrids in Amelanchier.
Canadian Journal of Botany 42: 651-663.
Fernald, M. L. 1950. Gray’s manual of Botany. American
Book Company, New York. 1632 pages.
Gleason, H. A., and A. Cronquist. 1991. Manual of vascular
plants of northeastern United States and adjacent Canada.
2"4 edition New York Botanical Garden, Bronx, New York.
993 pages.
Gerhold, H. D. 1999. Serviceberry cultivars tested as street
trees: initial results. Journal of Arboriculture 25(4): 189-
192.
Jones, G. N. 1946. American species of Amelanchier. Illinois
Biological Monographs 20(2): 1-126.
Kartesz, J. T., and C. A. Meachum. 1999. Synthesis of the
North American flora. Version 1.0. Biota of North America
Program, University of North Carolina, Chapel Hill.
2007 CHUBBS and PHILLIPS: A TRIBUT!
rO NEAL PHILIP PERRY SIMON, 197
42006
FiGurE 1. Flowering inflorescence of Amelanchier intermedia showing relatively long petals
and pedicels and leaves pubescent below and more or less expanded at flowering time.
Photo of a cultivated plant in Ottawa by P. M. Catling.
McKay, S. M. 1973. A biosystematic study of the genus
Amelanchier in Ontario. M. Sc. thesis, University of Toron-
to. 241 pages.
Nielsen, E. L. 1939. A taxonomic study of the genus Ame-
lanchier in Minnesota. American Midland Naturalist 22:
160-205.
Soper, J. H., and M. L. Heimburger. 1982. Shrubs of Ontario.
Royal Ontario Museum Publications in Life Sciences. 495
pages.
Voss, E. G. 1985. Michigan flora, part I. Dicots (Saururaceae
— Cornaceae). Cranbrook Institute of Science Bulletin 59.
724 pages.
Wiegand, K. M. 1920. Additional notes on Amelanchier.
Rhodora 22: 146-151.
Received 15 October 2005
Accepted 23 February 2007
9]
92 THE CANADIAN FIELD-NATURALIST Vol. 121
A Freshwater Hydrobiid, cf. the Squat Duskysnail, Lyogyrus granum
(Mollusca), Widespread in the Hampton Marsh, New Brunswick
DONALD F. MCALPINE!, RACHEL H. MAILLET!, ANDREW J. ALBERT!, LAURA M. CROSSMAN!, REBECCA
R. SmitH!, and ANDRE L. MARTEL?
'New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada
2Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada
McAlpine, Donald F., Rachel H. Maillet, Andrew J. Albert, Laura M. Crossman, Rebecca R. Smith, and André Martel. 2007.
A freshwater hydrobiid, cf. the Squat Duskysnail, Lyogyrus granum (Mollusca), widespread in the Hampton Marsh,
New Brunswick. Canadian Field-Naturalist 121(1): 92-94.
In Canada, the Squat Duskysnail, Lyogyrus granum, is an apparently rare freshwater hydrobiid. We document putative L.
granum to be widespread throughout the Hampton Marsh, New Brunswick, with densities sometimes exceeding 150 snails/m?*
on substrates ranging from vegetated organic detritus over mud, to sparsely-vegetated cobble/sand lakeshore. That our identi-
fication of L. granum from Hampton Marsh remains tentative, in spite of large samples, emphasizes the need for detailed
taxonomic study of any putative L. granum populations in Canada. Such study will be necessary before the true conservation
status can be determined for this and any related taxa, as yet unrecognized in Canadian material.
Key Words: Squat Duskysnail, Lyogyrus granum, Hydrobiidae, conservation status, New Brunswick.
Of the two species of freshwater hydrobiid snails
recorded in Maritime Canada (Clarke 1981), the Ordi-
nary Spire Snail, Amnicola limosa, and the Squat
Duskysnail, Lyogyrus granum, the latter is very poorly
known. In part, this is due to difficulties in identifica-
tion; Smith (1991) notes that in Massachusetts L.
granum and L. pupoidea will rarely be confidently
distinguished from each other and Jokinen (1992) has
reported sexual dimorphism within the genus that is
poorly understood.
In Canada, L. granum has been reported from only
two locations in south-central New Brunswick (Jem-
seg and Baker Brook) and 11 sites in Nova Scotia
(COSEWIC 2003*). COSEWIC (2003*) suggested that
L. granum is rare in Canada and that the only known
Canadian populations in New Brunswick and Nova
Scotia should be considered nationally significant.
Nonetheless, a paucity of information on this species
has led to a designation of Data Deficient by the Com-
mittee on the Status of Endangered Wildlife in Canada
(COSEWIC). Here we document a L. granum-like
hydrobiid to be abundant and widespread in the Hamp-
ton Marsh, New Brunswick.
Considered one of the largest and most diverse fresh-
water marshes in the Saint John River floodplain, the
2000 hectare Hampton Marsh (45°33'N, 65°50'W) con-
sists of a series of eutrophic oxbows, tributaries and
backwaters of the Kennebecasis and Hammond Rivers.
The marsh has repeatedly been identified as a site of
ecological importance within the province (Stocek
1984*: Dionne et al. 1988; Austin-Smith 1994*) and
is currently designated as a wetland of significance by
the Wetlands and Coastal Habitat Program of the New
Brunswick Department of Natural Resources and Ener-
gy. Although about 40% of the land adjacent to the
marsh remains forested (Austin-Smith 1994*), shore-
line development, sedimentation, eutrophication, and
pesticide introduction from upstream agricultural areas
are issues of concern. The marsh is located largely with-
in the municipal boundaries of Hampton and Quispam-
sis, two of the fastest growing urban areas in New
Brunswick, and the Local Service District of Nauwige-
wauk (Austin-Smith 1994*).
Putative L. granum were discovered in the Hampton
Marsh during the course of ongoing mollusc surveys
on the lower Saint John River and tributaries (Sabine
et al. 2004; McAlpine et al. 2005). Data reported here
were collected from 20 June to 22 August 2005 from
25 survey sites scattered across the marsh. Survey sites
had been selected based on study of aerial photos,
topographic and nautical maps, and field inspection,
to represent a variety of habitat types (i.e., lakeshore,
river channel, backwaters, oxbows) within the marsh
complex. Molluscs were collected using a combina-
tion of techniques that included SCUBA, snorkeling,
dip netting, and sieving bottom sediments; voucher
material of all mollusc species reported here has been
placed in the New Brunswick Museum mollusc col-
lection. To provide abundance estimates for putative
L. granum in contrasting habitats within the marsh,
15 0.25-m? random quadrats were also sampled in
~ | m of water in August at three of the sites. Only live
snails were tabulated in quadrats following the com-
plete removal, sieving, and hand-sorting of the top 5 cm
of sediment along with all aquatic vegetation. Hampton
Marsh specimens of Lyogyrus were identified largely
on the basis of the structure of the verge in fresh mate-
rial as described and illustrated in Smith (1991) and
Jokinen (1992), but reference was also made to shell
and opercular morphology as noted by these authors,
and by Clarke (1981). As well, comparison was made
with Atlantic Canadian material in the collection of
the Canadian Museum of Nature. Although L. pupoidea
has not been recorded in Canada, our extensive series
of specimens agree with Smith (1991), in that inter-
mediates and extremes showing external morpholog-
2007
NOTES 93
TABLE |. Densities of putative Lyogyrus granum in contrasting habitats in the Hampton Marsh, New Brunswick
GPS
N 45° 30.025'
W 65° 52.50'
N 45° 31.358'
W 65° 52.867'
N 45° 31.856!
W 65° 51.297’
Habitat
rocky lakeshore,
clearwater
Ceratophyllum marsh
murky
backwater channel
Mean + sd(/m*)
96.8 + 52.6
76.0 + 63.9
17.6 + 12.5
Range (/m*)
Associated molluscs
48-184 Ai, Al, Cd, Ec, Gd, Ha, Lr, Pc,
Ph, Sc, Se, Vt,
4-160 Ai, Al, Ec, Gd, Ha, Pa, Ph,
Se, Ss, Vs, Vt
8-20 Al, Cd, Ec, Fp, Gd, Pe, Ph, Se,
Ss. Vs, Vt
' Ai=Anodonta implicata, Al\=Amnicola limosa, Cd=Campeloma decisum, Ec=Elliptio complanata, Fp=Ferrissia parallela,
Gd=Gyraulus deflectus, Ha=Helisoma anceps, Pa=Planorbella campanulata, Lr=Lampsilis radiata, Pe=Pyganodon
cataracta, Ph=Physa heterostropha, Ss=Sphaeriidae sp., Sc=Stagnicola catascopium, Se=Stagnicola elodes, Vs=Valvata
sincera, Vt=Valvata tricarinata
ical features of L. pupoidea are present. COSEWIC
(2003*) notes the possibility of L. pupoidea, as well as
closely related L. browni, overlapping the Canadian
range for L. granum. Clearly there is a need for detailed
taxonomic study of any putative L. granum populations
in Canada before the true distribution and conservation
status of this and related species in the country can
be determined.
Putative L. granum are widespread throughout the
Hampton Marsh, although they were noticeably ab-
sent from sites in the river channel, where currents of
0.1-0.3 m/sec were recorded, and in the lowest por-
tion of the marsh where there is periodic intrusion of
brackish water. Discounting two sites where brackish
water clearly intrudes on occasion, we recorded L.
granum at 17 of the 22 survey sites remaining. The
limited habitat data available for Lyogyrus indicates
that this species is generally found on organic debris
and vegetation in standing water (Clarke 1981; Smith
1987*: Jokinen 1983). Currents at sites where we re-
corded Lyogyrus did not exceed 0.1 m/sec. COSEWIC
(2003*) reports L. granum densities of up to 400/m”;
densities in the Hampton Marsh were lower but some-
times exceeded 150/m? (Table 1). We found the highest
densities along an open, sparsely-vegetated cobble/sand
lake shore on rocks and on the living shells of the
unionids Elliptio complanata, Anodonta implicata and
Lampsilis radiata. Lyogyrus granum was also abundant
in clear water on submersed vegetation where Cerato-
phyllum demersum dominated (lakeshore vs clear
water; Mann-Whitney U = 15, P > 0.05) but signifi-
cantly less abundant in a murky vegetation-clogged
backwater channel characterized by Vallisneria amer-
icana and Potamogeton perfoliatus than on the rocky
lake shore (lake shore vs backwater; Mann-Whitney
U=25, P < 0.05) (Table 1). Both of the more heavily
vegetated sites were characterized by deep bottom
layers of organic detritus over mud.
COSEWIC (2003*) suggests that the few sites from
which the species has been recorded in Canada indi-
cate either that L. granum is a habitat specialist, that
human impacts are reducing the availability of suit-
able habitat, that L. granum is a truly rare species, or
some combination of these factors. That the rarity of
this species may only be apparent, due to a lack of col-
lecting, is not mentioned. Jokinen (1983, 1992) found
this snail to be uncommon in Connecticut, and both
Smith (1987*) and Jokinen (1992) found the species
to be rare in Massachusetts and New York, respectively.
While L. granum may indeed prove to be naturally rare,
this species is small and easily overlooked and there
have been few detailed surveys of the freshwater gas-
tropods of Atlantic Canada. That L. granum has been
recorded from 11 widely distributed sites across Nova
Scotia also suggests that further field investigation may
show this species, in association with related taxa as
yet unrecognized in Canada, to be more prevalent in
the region than current records indicate.
Acknowledgments
Financial support for this project was provided
through grants to DFM from the Community-Univer-
sity Research Alliance program of the Social Sciences
and Humanities Research Council of Canada, The Sala-
mander Foundation, the New Brunswick Wildlife Trust
Fund, and the Province of New Brunswick Summer
Mentorship Program for Female Students. Laurie Mills
and James Wilson of the Hampton Area Environmen-
tal Group assisted with field arrangements while Archie
and Connie Downey were unfailingly gracious in pro-
viding access to the marsh through their property and
providing use of a wharf and other amenities; Gart
Bishop and Geoffry McBriarty assisted with the field
sampling of molluscs.
Documents Cited (marked * in text)
Austin-Smith, P. 1994. Hampton-Kennebecasis Marsh Com-
plex — Status Report. Town of Hampton, New Brunswick,
99 pages.
COSEWIC. 2003. Assessment and update status report on
the Squat Duskysnail, Lyogyrus granum, in Canada. Com-
mittee on the Status of Endangered Wildlife in Canada.
Ottawa. [unpublished report]. vi + 27 pages.
Smith, D. G. 1987. Keys to the freshwater invertebrates of
Massachusetts. Number 2. Mollusca Mesogastropoda (oper-
culate snails). Massachusetts Department of Environmental
Quality Engineering, Division of Water Pollution Control
(Westborough). 34 pages.
94 THE CANADIAN FIELD-NATURALIST
Stocek, R. 1984. Environmentally significant areas in the
Saint John Planning Region. Environmental Services
Branch, Environment New Brunswick, Fredericton, New
Brunswick. 116 pages.
Literature Cited
Clarke, A. H. 1981. The freshwater molluscs of Canada.
National Museum of Natural Sciences, National Museums
of Canada, Ottawa. 446 pages.
Dionne, L. A., D. M. Farmer, and M. C. Young. (Compilers).
1988. Critical natural areas in New Brunswick. New Hori-
zons Critical Natural Areas Committee, Fredericton, New
Brunswick. 313 pages.
Jokinen, E. H. 1983. The freshwater snails of Connecticut.
State Geological and Natural History Survey of Connecti-
cut. Department of Environmental Protection Bulletin 109:
1-83.
Vol. 121
Jokinen, E. H. 1992. The freshwater snails (Mollusca: Gas-
tropoda) of New York State. New York State Museum Bul-
letin 482. 112 pages.
McAlpine, D. F., D. L. Bateman, and C. A. Davis. 2005.
Spurwinkia salsa (Pilsbry 1905) (Gastropoda: Hydrobi-
idae) in the Kennebecasis River estuary, New Brunswick:
A brackish water snail new to Canada. Journal of Conchol-
ogy 38: 602-604.
Sabine, D. L., S. Makepeace, and D. F. McAlpine. 2004.
The Yellow lampmussel (Lampsilis cariosa) in New Bruns-
wick: A population of significant conservation value. North-
eastern Naturalist 11: 407-420.
Smith, D. G. 1991. Keys to the freshwater macroinverte-
brates of Massachusetts. Department of Zoology, Univer-
sity of Massachusetts, Amherst, Massachusetts. 236 pages.
Received 27 February 2006
Accepted 5 November 2007
Northern Range Extension of the Pygmy Shrew, Sorex hoyi, in the Yukon
THOoMaAs S. JuNG!, TRoy D. PRETZLAW2, and Davip W. NAGORSEN?
'Fish and Wildlife Branch, Yukon Department of Environment, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada; e-mail:
thomas.jung @ gov.yk.ca
"Department of Natural Resource Sciences, McGill University, 21111 Lakeshore Blvd., Montreal, Quebec H9X 3V9 Canada
3Mammalia Biological Consulting, 4268 Mechosin Road, Victoria, British Columbia V9C 3Z4 Canada
Jung, Thomas S., Troy D. Pretzlaw, and David W. Nagorsen. 2007. Northern range extension of the Pygmy Shrew, Sorex
hoyi, in the Yukon. Canadian Field-Naturalist 121(1): 94-95.
A Pygmy Shrew, Sorex hoyi, was captured in a pitfall trap on the Blackstone River (65°04.6'N, 138°10.8'W) in central Yukon.
This represents a northern range extension of about 110 km for S. hoyi in the Yukon.
Key Words: Distribution, Pygmy Shrew, Sorex hoyi, Yukon.
Shrews, Sorex, have been undersampled in north-
western North America because they are often not the
focus of surveys (Jarrell 1986; Nagorsen 1996) and
they are not readily captured in traps designed for small
rodents (Nagorsen 1996). As such, our knowledge of
the distributions of shrews in northwestern North Amer-
ica (i.e., Alaska, Yukon, northern British Columbia,
western Northwest Territories) is limited. In recent
years, however, directed surveys of shrews in this vast
region have resulted in discovery of a new species, the
Alaskan Tiny Shrew, Sorex yukonicus (Dokuchaev
1997), and range extensions for a number of species
(e.g., American Water Shrew, S. palustris, Jarrell 1986;
Cook et al. 1997; Tundra Shrew, S. tundrensis, Nagors-
en and Jones 1981; Alaskan Tiny Shrew, S. yukonicus,
and Pygmy Shrew, S. hoyi, Peirce and Peirce 2000).
Nevertheless, much work is needed to better understand
the distributions of shrews in northwestern North Amer-
ica. Herein, we describe a northern range extension
for S. hoyi in the Yukon.
During an extensive survey of the small mammal
fauna along the Dempster Highway in central Yukon, a
gmy Shrew was captured on 12 July 2005, 2 km west
of the Blackstone River (65°04.6'N, 138°10.8'W),
129 km northeast of Dawson City. The shrew was cap-
tured in an unbaited pitfall trap installed flush with the
substrate. Pitfall traps have been found to be particu-
larly effective for Pygmy Shrews (Prince 1941). No
other shrews were captured at this site. The specimen
was identified using dental characteristics and keys
in Nagorsen (2002) and van Zyll de Jong (1983). The
specimen (Field ID: DEMP-0015) is held by the Yukon
Department of Environment, but will be deposited at
the Museum of Southwestern Biology.
Our record represents a northern range extension
of approximately 110 km for S. hoyi in the Yukon.
Youngman (1975) mapped the hypothetical range as
being as far north as the Old Crow Flats (approximate-
ly 68°N). Prior to our capture, however, the northern-
most specimen records of S. hoyi in the Yukon were
from 22 km east of Dawson City (Youngman 1975; van
Zyll de Jong 1983) and the abandoned village of Forty-
mile (64°25'N, 140°32'W), 67 km northwest of Dawson
City (B. G. Slough and T. S. Jung, unpublished data).
This Yukon range extension of S. hoyi is not surpris-
ing as there are more than 125 specimens held at the
University of Alaska Museum of the North that were
collected north of 65°N in Alaska, with the most
northerly taken near the Dalton Highway at the Diet-
rich River (67°37'N, 149°46.8'W; UAM 23050). In
2007
addition, van Zyll de Jong (1983) reported specimens
from Chick Lake and Fort Franklin, two locations
north of 65°N in the Northwest Territories. It is likely
that S. hoyi ranges further north in the Yukon; this is
a species of the boreal forest and it likely extends north
to the treeline. Further observations are needed to doc-
ument the range of S. hoyi and other soricids in the
Yukon and elsewhere in northwestern North America
(Cook et al. 1997).
Acknowledgments
We thank Kevin Lake, Elise Bolduc, Kyle Russell,
Kieran O’Donovan, Shannon Barker, Lea Randall,
Brian Bell, Sara Nielsen, Michael Smith, Becky Cad-
sand and Matthias Clyde for able assistance in the field
or lab. Funding for this work was provided by McGill
University, the Yukon Department of Environment,
and NatureServe Yukon. We thank Murray Humphries,
Julie Frisch and Gordon McRae for logistical support
while on the Dempster Highway.
Literature Cited
Cook, J. A., C. J. Conroy, and J. D. Herriges, Jr. 1997.
Northern record of the water shrew, Sorex palustris, in
Alaska. Canadian Field-Naturalist 111: 638-639.
Dokuchaey, N. E. 1997. A new species of shrew (Soricidae,
Insectivora) from Alaska. Journal of Mammalogy 78: 811-
817.
NOTES 95
Jarrell, G. H. 1986. A northern record of the water shrew,
Sorex palustris, from the Klondike River, Yukon Territory
Canadian Field-Naturalist 100: 39)
Nagorsen, D. W. 1996. The mammals of British Columbia
Volume 2: Opossums, shrews and moles of British Colum-
bia. Royal BC Museum Handbook. UBC Press, Vancouver
169 pages.
Nagorsen, D. W. 2002. An identification guide to the small
mammals of British Columbia. Ministry of Sustainable
Resource Management, Ministry of Water, Land, and Air
Protection, Biodiversity Branch, and Royal BC Museum
153 pages.
Nagorsen, D. W., and D. M. Jones. 1981. First records of
the tundra shrew (Sorex tundrensis) in British Columbia.
Canadian Field-Naturalist 95: 93-94.
Peirce, K. N., and J. M. Peirce. 2000. Range extensions for
the Alaska tiny shrew and pygmy shrew in southwestern
Alaska. Northwestern Naturalist 81: 67-68.
Prince, L. P. 1941. Water traps capture the Pygmy Shrew
(Microsorex hoyi) in abundance. Canadian Field-Naturalist
Se:
van Zyll de Jong, C. G. 1983. Handbook of Canadian mam-
mals. 1: Marsupials and insectivores. National Museums
of Canada, Ottawa. 210 pages.
Youngman, P. M. 1975. Mammals of the Yukon Territory.
National Museums of Canada, Ottawa, Ontario. Publica-
tions in Zoology 10: 1-192.
Received 25 January 2006
Accepted 15 January 2008
A Tribute to Neal Philip Perry Simon 1973-2006
Tony E. Cuupss! and FRANK R. PHILLIPS?
'Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and
Labrador AOP 1S0 Canada; e-mail: techubbs @cablelab.net
*Department of Natural Resources, Government of Newfoundland and Labrador, Box 175, Station C, Happy Valley-Goose
Bay, Newfoundland and Labrador, Newfoundland AOP 1CO Canada
Chubbs, Tony E., and Frank R. Phillips. 2006. A tribute to Neal Philip Perry Simon, 1973-2006. Canadian Field-Naturalist
121(1): 96-99.
Neal Philip Perry Simon, our friend, hunting com-
panion, and colleague, was born in Labrador City,
Newfoundland and Labrador, on Sunday 30 December
1973, to Dave and Doreen Simon. He passed away sud-
denly and unexpectedly in a tragic accident on 23 Sep-
tember 2006, at the age of 32 near Happy Valley —
Goose Bay where he resided since 1988. His parents,
Dave and Doreen Simon, sister Nicole and fiancée,
Leanne Elson, survive him. Neal had a mutual adora-
tion for his grandparents, Carrie and Gordon Temple-
man, who will miss him dearly.
Neal’s early education was in Catholic schools: Notre
Dame Academy (primary and elementary) and Labra-
dor City Collegiate (high school) both of which were
in Labrador City and have since closed. Neal, along
with his grade school pal Colin Carroll, developed a
keen interest in biology at an early age, trying his hand
at taxidermy in grade seven. Francis Schwab, Neal’s first
year university professor, was instrumental in introduc-
ing him to scientific research, especially the effects of
post-fire succession on songbird abundance. His post-
secondary education began at Memorial University of
Newfoundland in St. John’s, Newfoundland and Lab-
rador, where in 1995 he obtained a B.Sc.(Hons) with a
major in ecology and evolution and a minor in statis-
tics. It was there, under the instruction of Eric Baggs
and the late Dr. Gary Cowen, he was instilled with an
appreciation of the importance of small mammals in
understanding forest structure dynamics.
In 1996, Neal began his Master of Science in For-
estry at the University of New Brunswick under the
supervision of Dr. A. W. Diamond. As part of the
Atlantic Cooperative Wildlife Ecology Research Net-
work (ACWERN), Neal regularly contributed presen-
tations at the annual workshop from 1998 to 2003. In
2000, Neal continued under the supervision of Dr.
Diamond to pursue his Ph.D. on the effects of site
productivity and heterogeneity on bird habitat quality
and species richness. His research ranged from mod-
elling species distributions to studying the effects of
landscape features on habitat use by passerines, with a
focus on their demography. Much of Neal’s work had
an overlying conservation theme linked quite strongly
to ecological theory. Neal completed his Ph.D. in 2005
and was about to embark on his post-doctoral research
in the spring of 2007.
Neal taught in various capacities at Memorial Uni-
versity of Newfoundland, the University of New Bruns-
wick and St. Mary’s University. From 1996 to 1998,
Neal worked as a contract biologist for the College of
|
'
the North Atlantic in Labrador City and Happy Valley —
— Goose Bay where he also offered his services as a
university tutor in various disciplines, including chem- —
istry and biology. Neal was always there to help out
whenever or wherever he could and would always with-
out hesitation say “Yes, just provide me the materials.”
Students found him friendly and informative and they
usually had few problems grasping concepts from him.
He will be remembered for his willingness to help, his
ability to teach a difficult content area and his friendly —
personality.
From November 1998, Neal was employed by the
Provincial Government of Newfoundland and Labra-
dor, Department of Natural Resources, occupying the
position of Regional Ecologist for the Labrador Region.
Neal’s work concentrated on experimental design, data
collection, and analysis of research projects relating to
forest management plans in Labrador. Neal’s research
interests included effects of forest management and
changing forest structures and specifically how these
changes may affect the spatial and temporal distribu-
tion of threatened Woodland Caribou (Rangifer taran-
dus caribou) in central Labrador. Neal’s most recent
interests involved sustainable forest management in)
collaboration with the Government of Newfoundland:
and Labrador, Department of Natural Resources, and)
the Université du Québec 4 Montréal. Neal was about
to embark on his post-doctorate research using inno-
vative simulation models to ensure the sustainable man-
agement of Labrador’s forests while his fiancée and co-;
worker Leanne would be completing her Master of)
Science in a supporting area of research. Neal was one¢
of those rare ecologists who not only recognize thes
power of quantitative methods, but also actually under-
stand how to appropriately apply these methods. Hav-
ing just barely embarked upon his research career,
Neal, as attested by his already growing list of publi-
cations, was destined for greatness in the world of
research and statistics.
Since the summer of 2000, Neal had been an active
participant in the North American Breeding Bird Sur- |
vey (BBS) program and ran the Newfoundland anc:
96
2007 CHUBBS and PHILLIPS: A TRIBUTI
rO NEAL PHILIP PERRY SIMON, 1973-2006 g
75
Ficure |. Neal P. P. Simon holding his first seal darted off Mulligan Point, 2 May 2003. (Photo
courtesy Sarah Townley).
Labrador, Happy Valley — Goose Bay survey route for
Environment Canada. Neal was also a committed par-
ticipant in the Christmas Bird Count (CBC).
Neal was a dedicated member of both the local
Squash League and Goose Bay Judo Club. He exem-
plified the philosophy of Judo in all aspects of his life,
striving to maximize his physical, intellectual and per-
sonal character by doing his best; and being his best,
Neal truly reflected this teaching through his physical
strength and fitness, his academic accomplishments
and desire to learn. As a competitor, Neal was intense
and spirited while always maintaining his youthful grin
and sense of humour. Neal’s enthusiasm for research
in ecology and his jubilant attitude radiated from him,
rapidly instilling in his colleagues a source of continual
positive motivation. With an acute and intense sense of
humour, Neal added a unique and welcome aspect to
his interaction with both friends and fellow researchers.
As an editor and reviewer of prospective manuscripts,
his dedication and support were immeasurable and
will be greatly missed.
In 1999, Neal was an original member of the Lab-
rador Wolverine Working Group, and became one of
the founding members of the Labrador Woodland Cari-
bou Recovery Team in 2001. Neal was a member of
The Society for Conservation Biology, the Wildlife
Society, the Atlantic Region of the Canadian Climate
Impacts and Adaptation Research Network, the Atlan-
tic Society of Fish and Wildlife Biologists, Atlantic
Co-operative Wildlife Ecology Research Network
98 THE CANADIAN FIELD-NATURALIST
(ACWERN), Newfoundland and Labrador Biodiver-
sity working group, Forest Management Planning
Team Member for Labrador Districts 19, 20, 21 and
22 and a member of the Institute for Environmental
Monitoring and Research — Osprey Technical Com-
mittee. Neal was about to join the Groupe de recherche
en écologie forestiére interuniversitaire at 1’ Université
du Québec 4 Montréal to complete his post-doctoral
research developing habitat and timber harvesting-
conservation trade-off models for Woodland Caribou.
Neal’s interests were varied and included, but were
not limited to, modelling species movements, predict-
ing species distributions and understanding the effects
of forest management on species occurrence and demo-
graphy. He spent most summers in Labrador, along with
his students and co-workers; in the field conducting
songbird point count samples and vegetation plot counts.
We came to know Neal and build a strong friendship
when he moved to Happy Valley — Goose Bay in 1998
and shortly thereafer began work as the Ecosystem
Ecologist for Labrador. Neal was friendly, energetic and
an avid waterfowl and big game hunter. We anxiously
awaited mid spring each year when we would travel by
snowmobile over the bay ice to Portage Island, Mul-
ligan Point. Here we would participate in the eons old
traditional Inuit hunting practice of waiting patiently
at breathing holes for Ringed Seals (Pusa hispida).
Evenings would be filled with laughter as we swapped
humorous exaggerated stories of past good times. Such
wonderful times were set against the backdrop of a
blazing driftwood fire and the assurance of deep slum-
ber later in our traditional Labrador canvas tents heat-
ed with the warmth of woodstoves. Neal often swapped
related stories of how he had enjoyed the outdoors
with his best hunting buddy, his father Dave, in West-
ern Labrador. It was more important to Neal to enjoy
the outdoors than to be successful on a hunt, though
more often than not he was. As he so often mentioned
to his hunting partner Colin Carroll, “If we don’t get
anything today, it doesn’t matter. Just being here doing
this is making my day.”
His friends will best remember Neal as a shining,
though comedic intellectual, with a love of life and a
passion for the outdoors. Through discussions with
some of Neal’s colleagues and friends in the prepara-
tion of this tribute, we have found that Neal existed in
several parallel universes, having the same mannerisms,
characteristics and sharp wit despite the varied per-
sonalities and context of his friends. He will be remem-
bered for his comic additions to his work and conver-
sation slipping easily between extreme seriousness and
laughter.
Acknowledgments
We thank Neal’s parents (Dave and Doreen), his
sister Nicole, fiancée Leanne, and others for personal
information. Colin Carroll, Neal’s schoolmate and col-
league provided information on Neal’s early schooling
in Labrador City. We thank Tony Diamond for provid-
Vol. 121
ing a compilation of Neal’s academic achievements.
Matt Betts and Joe Nocera provided insight on Neal’s
life that could only be shared between close colleagues
and best friends. Bob Simms shared personal interac-
tions with Neal through their judo club and Bernice
Tracy provided details of Neal’s association with the
College of the North Atlantic. Finally, we would like
to thank all of Neal’s friends, family, colleagues and
acquaintances that contributed to the Neal Simon
Memorial Trust to fund an academic scholarship in
Neal’s honour and memory. Neal appreciated the out-
doors and enjoyed hunting and fishing and spending
time in the Labrador wilds with Leanne and his dog,
Boss. He will be greatly missed by his family, his
many friends and his academic colleagues.
Bibliography of Neal P. P. Simon
Compiled by Tony E. CHUBBS
Peer Reviewed Publications:
Elson, L. T., N. P. P. Simon, and D. Kneeshaw. /n press.
Regeneration differences between fire and clearcut logging
in southeastern Labrador: a multiple spatial scale analysis.
Canadian Journal of Forest Research: 000-000.
Elson, L. T., and N. P. P. Simon. /n press. Plant abundances
following clearcutting and stripcutting in central Labrador.
Northern Journal of Applied Forestry, 00: 000-000.
Elson, L. T., F. E. Schwab, and N. P. P. Simon. Jn press
Winter food habits of willow ptarmigan (Lagopus lagopus)
as a mechanism to explain winter sexual segregation. North-
eastern Naturalist 00: 000-000.
Roberts, B. A., N. P. P. Simon, and K. W. Deering. Jn press.
The forests and woodlands of Labrador, Canada: Ecology,
distribution, and future management. Ecological Research
00: 000-000.
Schwab, F. E., N. P. P. Simon, S. Stryde, and G. Forbes. /n
press. Effect of post-fire snag removal on breeding birds of
western Labrador. Journal of Wildlife Management 00:
000-000.
Schwab, F. E., N. P. P. Simon, and A. R. E. Sinclair. 2006.
Breeding birds related to vegetation structure in southeast
British Columbia. Journal of Wildlife Management 70:
189-197.
Schwab, F. E., N. P. P. Simon, and S. Nash. 2005. Sex and
age segregation of wintering willow ptarmigan in Labrador.
Northeastern Naturalist 13: 113-118.
Simon, N. P. P., and F. E. Schwab. 2005. Plant community
structure following wildfire in the subarctic forests of Lab-
rador. Northern Journal of Applied Forestry 22: 229-235.
Newbury, T. L., and N. P. P. Simon. 2005. The effects of
clearcutting on snowshoe hare (Lepus americanus) rela-
tive abundance in central Labrador. Forest Ecology and
Management 210: 131-142.
Betts, M., N. P. P. Simon, and J. J. Nocera. 2005. Point count
summary statistics differentially predict reproductive activ-
ity in bird-habitat relationship studies. Journal of Ornithol-
ogy 146: 151-159.
Simon, N. P. P., and F. E. Schwab. 2005. The response of
conifer and broad-leaved trees and shrubs to wildfire and
clearcut logging in the boreal forests of central Labrador.
Northern Journal of Applied Forestry 22: 35-41.
Otto, R. D., N. P. P. Simon, S. Couturier, and I. Schmelzer.
2003. Evaluation of satellite collar sample size require-
ments for mitigation of low-level military jet disturbance
2007
of the George River caribou herd. Rangifer special issue
number 14; 297-302.
Simon, N. P. P., A. W. Diamond, and F. E. Schwab. 2003.
Do northern forest bird communities show more ecological
plasticity than southern forest bird communities in eastern
Canada? Ecoscience 10: 298-296.
Simon, N. P. P., F. E. Schwab, and R. D. Otto. 2002. Song-
bird abundance in clear-cut and burned stands: a compari-
son of natural disturbance and forest management. Cana-
dian Journal of Forest Research 32: 1343-1350.
Simon, N. P. P., C. B. Stratton, G. J. Forbes, and F. E.
Schwab. 2002. Similarity of small mammal abundance in
post-fire and clearcut forests. Forest Ecology and Manage-
ment 165: 163-172.
Schwab, F. E., N. P. P. Simon, and C. G. Carroll. 2001.
Breeding songbird abundance in the subarctic forests of
western Labrador. Ecoscience 8: 1-7.
Schwab, F. E., F. G. Pitoello, and N. P. P. Simon. 2001.
Relative palatability of green manure crops and carrots to
white-tailed deer. Wildlife Society Bulletin 29: 317-321.
LeCoure, M. I., F. E. Schwab, N. P. P. Simon, and A. W.
Diamond. 2000. Effects of post-fire salvage logging on
breeding birds in western Labrador. Northeast Wildlife 55:
39-46.
Simon, N. P. P., F. E. Schwab, and A. W. Diamond. 2000.
Patterns of bird abundance in relation to logging in west-
ern Labrador. Canadian Journal of Forest Research 30:
257-263.
Simon, N. P. P., F. E. Schwab, M. I. LeCoure, F. R. Phillips,
and P. G. Trimper. 1999. Effects of trapper access on mar-
ten population in central Labrador. Northeast Wildlife 54:
73-76.
Simon, N. P. P., F. E. Schwab, M. I. LeCoure, and F. R.
Phillips. 1999. Fall and winter diet of Martens, Martes
americana, in central Labrador related to small mammal
densities. Canadian Field-Naturalist 113: 678-680.
Simon, N. P. P., F. E., Schwab, E. M. Baggs, and G. I. McT.
Cowan. 1998. Distribution of small mammals among suc-
cessional and mature forest types in western Labrador.
Canadian Field-Naturalist 112: 441-445.
Peer Reviewed Manuscripts in Preparation:
Newbury, T. L., N. P. P. Simon. Moose (Alces alces) browse
use in central Labrador Submitted to Canadian Field-Nat-
uralist.
Simon, N. P. P., and A. W. Diamond. Songbird habitat quality
across a timber productivity gradient within an unfragment-
ed northern boreal forest: local and landscape effects. Sub-
mitted to Landscape Ecology.
Simon, N. P. P., and A. W. Diamond. Evaluation of environ-
mental factors influencing vegetation structure in mature
Picea mariana forests using constrained ordination and
constrained classification. Submitted to Canadian Journal
of Forest Research.
Simon, N. P. P., F. E. Schwab, and J. K. Colbert. Vegetation
response to summer and winter logging of Black Spruce
Picea mariana forests in central Labrador. Submitted to
Northern Journal of Applied Forestry.
Theses:
Simon, N. P. P. 1996. The effects of secondary succession on
vegetation structure and its effects on small mammal bio-
diversity in western Labrador. B.Sc. Honours thesis.
Memorial University of Newfoundland, St. John’s, New-
foundland and Labrador.
CHUBBS and PHILLIPS: A TRIBUTE TO NEAL PHILIP PERRY SIMON, 1973-2006 YY
Simon, N. P. P. 1998. Patterns of bird abundance in relation
to logging in western Labrador. M.Sc. University of New
Brunswick, Fredericton, New Brunswick. 83 pages
Simon, N. P. P. 2006. The effects of site productivity and
heterogeneity on bird habitat quality and species richness
Ph.D. thesis. University of New Brunswick, Fredericton,
New Brunswick. 184 pages.
Reports:
Schmelzer, L., J. Brazil, T. Chubbs, S. French, S. B. Hearn,
R. Jeffery, L. LeDrew, H. Martin, A. McNeill, R. Otto,
F. Phillips, G. Mitchell, G. Pittman, N. Simon, and G.
Yetman. 2004. Recovery Strategy for three Woodland cari-
bou herds (Rangifer tarandus caribou; Boreal population)
in Labrador, Canada. Department of Environment and Con-
servation, Government of Newfoundland and Labrador,
Corner Brook. 51 pages.
Simon, N. P. P. 2003. A summary of some effects of full tree
logging — literature review. Internal report — Department of
Forest Resources and Agrifoods, Wildlife Division.
Simon, N. P. P., E. Baggs, and G. I. McT. Cowan. 1998.
the effect of forest fire on small mammals in western Lab-
rador. Internal report — Department of Forest Resources
and Agrifoods, Wildlife Division. 23 pages.
Simon, N. P. P., F. E. Schwab, and M. I. LeCoure. 1998.
Labrador marten demographics as determined by winter
diet and trapper access. Interim report submitted to the
Newfoundland and Labrador Department of Forest Re-
sources and Agrifoods, Wildlife Division.
Simon, N. P. P., F. E. Schwab, and A. W. Diamond. 1998.
Patterns of bird abundance in relation to logging in western
Labrador. Interim report submitted to: Wildlife Habitat
Canada, Newfoundland and Labrador Department of For-
est Resources and Agrifoods, Wildlife Division.
Simon, N. P. P., F. E. Schwab, and A. W. Diamond. 1997.
The effects of logging on breeding birds in western Lab-
rador. Interim report submitted to: Wildlife Habitat Cana-
da, Newfoundland and Labrador Department of Natural
Resources and Wildlife Division.
Workshop and Conference Presentations:
Simon, N. P. P. 2003. Evaluating the effects of landscape
change on abundance, productivity, and survival of forest
birds (presented with M. Betts). ACWERN, Wolfville,
Nova Scotia.
Simon, N. P. P. 2003. Natural disturbance and forest man-
agement. Workshop on Natural Disturbance Management.
Sustainable Forest Management Network, Corner Brook,
Newfoundland and Labrador.
Simon, N. P. P. 2002. The relationship between forest pro-
ductivity and songbird habitat quality. ACWERN, Rocky
Harbour, Newfoundland and Labrador.
Simon, N. P. P. 2000. Songbird abundance and fecundity in
relation to forest structure and productivity. ACWERN,
St. Andrews, New Brunswick.
Simon, N. P. P. 1998. Patterns of bird abundance in relation
to logging in western Labrador/Bird plasticity in relation to
latitude. Institute of Environmental Monitoring and Re-
search Seminar Series — Happy Valley — Goose Bay, New-
foundland and Labrador.
Simon, N. P. P. 1997. The effects of logging on birds in west-
erm Labrador. ASWFB/ACWERN conference — Alma. New
Brunswick.
Simon, N. P. P. 1996. The effects of logging on birds in westem
Labrador. ACWERN, Bon Portage Island, Nova Scotia.
Received 14 February 2007
Book Reviews
ZOOLOGY
Amphibians and Reptiles of British Columbia
By Brent M. Matsuda, David M. Green, and Patrick T. Gre-
gory. 2006. Handbook, Royal British Columbia Museum,
Victoria, British Columbia. 266 pages. $17.33.
The British Columbia Provincial Museum (now the
Royal British Columbia Museum) was second only to
the Royal Ontario Museum to issue guidebooks to their
provincial amphibian and reptile faunas for the public
(by E. B. S. Logier in 1937 and 1939 for Ontario, and
Clifford Carl in 1943 and 1944 for British Columbia).
The Carl guides (Handbooks 2 and 3) subsequently
went through reprintings with some revisionary materi-
al added in 1951 and 1968. In 1984 completely rewrit-
ten guides were published on reptiles (Handbook 44
by Gregory) and amphibians (Handbook 45 by Green),
both coauthored with R. Wayne Campbell.
Now a new handbook (unnumbered) is a combined
revised and partially rewritten text from the earlier
Green and Gregory texts with updated distribution
maps and the addition of a new coauthor. The previ-
ous authors are long-established, still active research
herpetologists with extensive publications, Green pri-
marily on frogs (in the broad sense including, as well
as typical “frogs”, toads, treefrogs, spadefoots, tailed
frogs) and Gregory primarily on snakes but also includ-
ing lizards, turtles and some amphibians. Dr. Gregory
has spent his entire post-graduate career at the Univer-
sity of Victoria; Dr. Green was born British Columbian
but his post-graduate career has been at the Redpath
Museum at McGill in Quebec. Both have extensive
field experience in British Columbia. The new addition,
Matsuda, is characterised in the tongue-in-cheek auth-
ors’ profiles (page 258) as an “overachieving ecto-
therm” (although this is surely not a unique charac-
terization within this triumvirate) with a UBC M.Sc.
who is now settled in British Columbia as an envi-
ronmental consultant.
This handbook, like its predecessors, follows a stan-
dard guide layout. Preface, introduction (covering a
wide range of aspects of amphibians and reptiles in
general), checklist and conservation status, individual
species accounts (for 39 species of which 4 are defi-
nitely and 2 possibly introduced) and a further section
which includes additional introduced species which
apparently have not persisted or whose occurrence is
only speculative, an appendix which gives contact in-
formation for regional authorities, glossary of terms,
bibliography (divided into additional reading and ref-
erences), acknowledgements (about authors with illus-
trator credits) and an index. The black-and-white text
drawings are mostly repeats from the earlier guides
except for newly included wall lizard and all of the
amphibians. A few of the latter are superior to those
of the earlier amphibian guide but many are not. The
new salamander drawings are particularly disappoint-
ing with the number of costal grooves shown rarely
agreeing with the counts given in the text. However,
all are sufficient for identification. New are 60 small
colour photographs inserted between pages 186 and —
187.
Additions since the last guide new to British Colum- |
bia and Canada are the secretive salamander Plethodon _
idahoensis and the introduced European wall lizard
Podacris muralis. Also new is the conclusion from
molecular evidence by T. R. Jackson in 1998 that the
Aneides salamander on Vancouver Island is an import
with bark from California, A. vagrans, and not the
geographically closer A. ferreus to which it had long
been assigned. Other nomenclature updates from recent
revisionary studies are Dicamptodon tenebrosus, Asca-
phus montanus, Spea intermontanus, Actinemys mar-
morata, Pseudacris maculata, Rana luteiventris, Pitu-
ophis catenifer, and Crotalus oreganus. One accepted
elsewhere that was not made is Pseudacris for Hyla
regilla, and the more controversial generic changes
Lithobates for Rana sylvatica and Rana pipiens (but
not for R. aurora, R. pretiosa, and R. luteiventris)
and Anaxyrus for Bufo boreas, even though one of
the authors (David Green) is a coauthor on the paper
detailing the latter changes (American Museum of
Natural History Bulletin 297).
Many incidental facts liven the text (such as the
observation that frogs always close their eyes when)
they leap (page 14)), that frogs can “hear” ground-
borne vibrations by transmissions through their front!
legs (pages 21-22) and that females of many turtles:
species can store sperm from a single male for up to
four years (page 32).
The marine Cadborosaurus and the Okanagan Lakee
“Ogopogo” are only mentioned under folklore (pagee
49) with the (arguable by some) statement “Sporadic¢
sightings of these beasts continue, but evidence of!
their existence is lacking.”
Few slips were missed in proofing such as the fail-
ure to italicise Hyla versicolor and H. chrysoscelis on
page 29, and the only British Columbia specimens ol
the horned lizard Phrynosoma given as “two neal
Osoyos in 1898” on page 182 but as “two records from
Osoyos area in 1910” on the facing page 183 (the firs
is correct).
100
2007
This guide brings British Columbia herpetology
effectively into the 21" century as to be expected from
the distinguished authors. It is a must for any west coast
naturalist’s bookshelf and/or field jacket with an out-
sized pocket.
BooK REVIEWS 10]
FRANCIS R. Cook
Canadian Museum of Nature, Box 3443, Station D, Ottawa,
Ontario K1P 6P4 Canada
Blue Grouse: Their Biology and Natural History
By Fred C. Zwickel, and James F. Bendell. 2004. National
Research Council of Canada, Ottawa, Ontario. 284 pages.
$69.95 Paper.
This monograph is the culmination of two life-
times devoted to studying Blue Grouse population
biology and behaviour, both in the field and in aviaries,
predominantly in coastal British Columbia. One of the
advantages of a lengthy monograph is that the authors
were able to synthesize much unpublished data and
“gray” literature in addition to published references,
making it the most comprehensive reference on this
species group that is currently available. As such, it is
not a book for the general reader of natural history (the
colour and black-and-white photographs scattered
throughout are quite small), but a scientific reference
full of tables and graphs and lots of detail (there are
12 pages each of references and results of statistical
tests).
Unfortunately, only two years after Zwickel and Ben-
dell’s monograph was published, the American Ornith-
ologists’ Union (Banks et al. 2006) re-split Blue Grouse
into two taxa, Dendragapus obscurus in the Pacific
Coast Range and the Sierra Nevada, and D. fuliginosus
in the Rocky Mountains. These two taxa had origi-
nally been considered two species (Brooks 1929), but
were conspecific for most of the 20" century. The re-
split resulted from the recent DNA-based work of
Barrowclough et al. (2004), who also found that the
New Mexico populations of Dusky Grouse were per-
plexingly different, but not enough to consider them
separate species. Populations in the northern parts of
the range (i.e., north of the Chilcotin in central B.C.,
through to Alaska) were not included in the Barrow-
clough et al. study, but the separation into coastal and
inland species is assumed to hold true. Unfortunately,
the AOU split makes the monograph more difficult to
use, but in their favour, Zwickel and Bendell use the
scientific names, often with full trinomial reference
Birds of Peru
By Thomas S. Schulenberg, Douglas F. Stotz, Daniel F. Lane,
John P. O'Neill, and Theodore A. Parker III. 2007 Prince-
ton University Press. 656 pages. U.S. $49.50 Cloth.
In 2001 I was delighted to find Birds of Peru, by J.
Clements and N. Shany. It was a good modern field
guide and proved its worth in Peru. However, there
were a few times when I needed to borrow a more com-
prehensive text to be certain of my identification. Now
to the eight accepted races at the time of publication,
throughout, as well as referring to “coastal” and
“interior” subspecies analogous to the new split. I
recommend that the serious reader really study the
chapter on taxonomy and distribution with Barrow-
clough’s paper (available on the internet) in hand.
As a serious naturalist, | found the chapters on histor-
ical review, physical environment, integument (plumage
especially), behaviour, habitat use and movement, pop-
ulation parameters, predators and disease the most
interesting. The seven chapters on form and function
— integument, morphology, reproduction, growth and
development, food and nutrition, energetics and genet-
ics — will be of interest mostly to serious students of
Blue Grouse.
Although Zwickel and Bendell refer to studies from
other parts of Blue Grouse range, they admit that the
strong focus on the coastal species (D. fuliginosus) is
unfortunate because there are a number of character-
istics, such as vocalizations, that are clearly different
between the two species. The authors hint at a future
publication that will explore the population ecology of
Blue Grouse, and it is to be hoped that they are able
to overcome this geographical disparity.
Literature Cited
Banks, R. C., C. Cicero, J. L. Dunn, A. W. Kratter, P. C. Ras-
mussen, J. V. Remsen, J. D. Rising, and D. F. Stotz. 2006.
Forty-seventh supplement to the American Ornithologists’ Union
Check-list of North American Birds. Auk 123(3): 926-936.
Barrowclough, G. F., J. G. Groth, L. A. Mertz, and R. J. Gutierrez.
2004. Phylogeographic structure, gene flow and species status in
Blue Grouse (Dendragapus obscurus). Molecular Ecology 13:
1911-1922.
Brooks, A. 1929. On Dendragapus obscurus obscurus. Auk 46: 111-
113.
CyYNDI M. SMITH
Box 5, Waterton Park, Alberta TOK 2MO Canada
we have a new guide by Schulenberg et al. and the
obvious question is whether it is an improvement.
The first difference is that the new guide has range
maps. This means I do not have to struggle with phras-
es such as “on the east bank of the Rio Utacamba at the
south end of Cordillera de Colon.” [Marvelous Spatule-
tail] Now I can look at the map and have an immediate
understanding. The range maps are conveniently set
102
next to the text in what is normally the margin, making
great use of space. This alone is a great improvement.
Peru has about 1800 species or one fifth of the
world’s birds. Most sources cite this vague number.
The few that are more precise give numbers that vary
by up to 100 species. It is not surprising that a state
of flux exists in such a vast and complex area. This
book covers 1792 species which is probably as good
as you do under such fluctuating circumstances. Try-
ing to reconcile the list of species covered by both
books or given in other sources is very difficult. It is
a bit like trying to compile a list of “honest” politi-
cians. The new book drops some questionable records
like Plumbeous Ibis [one old, suspect, badly labeled
specimen] but adds Bogota Rail [based on a current
valid sighting.] Some birds are treated by Shulenburg
as sub-species; Andean Ibis is listed as a sub-species
of Black-faced Ibis whereas other sources give it full
species status. There are complications due to name
changes. For example, there are four toucans listed in
both books. However, the names do not correlate well.
This includes the scientific binomials, normally a very
stable source of information. The old book has Toco
Toucan Ramphastos toco, Black-mandibled Toucan
Ramphastos ambigus Yellow-ridged Toucan Ramphas-
tos culminates, and Cuvier’s Toucan Ramphastos cu-
vieri. The new book has Toco Toucan Ramphastos toco,
Black-mandibled Toucan Ramphastos ambigus plus
White-throated Toucan Ramphastos tucanus, and
Channel-billed Toucan Ramphastos vitellinus. The last
two are new names for Cuvier’s and Yellow-ridged
Toucans. [Speciation in the whole toucan complex is
confused.] I also noted that Andean Flamingo and
James’s [Puna] Flamingo have been updated to Phoeni-
coparrus andinus and Phoenicoparrus jamesi, respec-
tively, but the Guanay and Red-legged Cormorants
The Freshwater Fishes of British Columbia
By J. D. McPhail, illustrated by D. L. McPhail, foreword by
Joseph S. Nelson. 2007. University of Alberta Press,
Edmonton. lxxiv + 620 pages. $90.00.
This book is the latest treatment covering the fresh-
water fishes of British Columbia. Earlier books ap-
peared in 1948 (132 pages) and 1967 (192 pages).
They were published by the British Columbia Provin-
cial Museum (now Royal British Columbia Museum)
while the latest book is published, strangely, in Alberta.
The present volume is 694 pages long with a larger
format and is indicative of both the growth in knowl-
edge and the need for a comprehensive work on this
provincial fauna.
The book is composed of introductory sections coy-
ering purpose, use of keys, names of fishes, layout of
the book, origins of the fauna, present distributions,
and conservation. The sections on glacial events and
THE CANADIAN FIELD-NATURALIST
Vol. 121
have not been changed to Leucocarbo bougainvillii
and Leucocarbo gaimardi. So those with a penchant
for lists may need to do some research. That being
said, this book is more than adequate as a field guide
for the resident and visiting birder alike.
The color plates are opposite the descriptions and
distribution maps. There are 13 illustrators, so there is
some difference in style. The most disparate are Lau-
rence McQueen and John O’ Neill. O’ Neill’s work has
the precise, almost photographic rendering we see in
most modern field guides. McQueen’s paintings are
more flowing and look like paintings. My junior school
art teacher would have said they were more “painterly.”
Daniel Lane’s work is somewhat in between these two
styles. Which style you like is a matter of taste. I have
looked at many of the illustrations for accuracy and can
find no problems. In fact, I particularly like the tou-
canets by O’ Neill and the jaegers by McQueen. There
are 4000 color illustrations at about six to a plate, mean-
ing the illustrations are quite large [again the publish-
er has used the margins].
In addition to the species accounts there are well-
written sections on Habitats, Molts-and-Plumages, and
Conservation. In particular the terminology used for
habitat in the species accounts is clearly documented.
The text in the species accounts covers the key iden-
tification features, status, distribution, and song.
This book will please birders and biologists living
in or visiting Peru. It is also useful beyond the borders
of Peru into adjacent areas of South America. It is a lit-
tle on the heavy side for dragging through a hot jungle,
but what else can you expect with such an enormous
bird list.
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada
current distributions are a very useful synthesis as is
the discussion on what a species is and why the vari-
ous forms of sticklebacks and others have not been
named taxonomically.
The bulk of the text is the descriptions of species.
Family accounts give general information on the in-
cluded species and the identification keys are found
there. The species accounts are preceded by a pictori-
al key to families that works for the more distinctive
body shapes but would have benefited from some anno-
tations for those fish that are more similar in shape to
enable the naive reader to correctly identify the family.
It is always a debate as to whether keys should be with
the family or should form a separate section. The for-
mer works when reading through the book but the latter
is much easier for the actual identification process in a
laboratory or the field. The keys are well illustrated al-
2007
though both illustrations and text are too small. Partic-
ularly useful are keys to fry, parr, and adult salmons and
trouts as these life history stages differ in appearance.
Each species account has a scientific name (but no
date with the author), English common name, a black-
and-white illustration, sections on distinguishing fea-
tures, taxonomy, sexual dimorphism, hybridization, dis-
tribution with a spot map, life history, reproduction,
age, growth and maturity, food habits, habitat and con-
servation comments.
A total of 81 species are covered, including 10 exot-
ics. The main families of the 17 known from British
Columbia are salmons, trouts and whitefishes with
23 species, minnows (20), and sculpins (8). Minor er-
rors are inevitable in any work; e.g., Lindsey is mis-
spelled in the dedication, Qadri as Quadri in the Ref-
erences, but some can be confusing; e.g., on page 498
Pomoxis nigromaculatus is said to have 5 anal spines
in the sunfish identification key but most fish have 6-
7 spines; the stickleback key separates two species by
non-overlapping dorsal fin spine counts (which do,
however, overlap occasionally) where in one species
(Pungitius) the spines are staggered left to right, an
easily seen and unique character (mentioned in Dis-
tinguishing Characters).
The book ends with an extensive Bibliography, an
Appendix I (a checklist of the fishes, with no other
appendices), a Glossary and an index to scientific and
common names. The Glossary explains a variety of
terms, even that the plural of dentary is dentaries which
is fairly standard English, but gives axillae without
explaining it is the plural of axilla (a knowledge of
Latin is ever more dead). One obscure word is snye,
defined as a dead-end side channel. This is listed in
dictionaries as an Ontario word and can also mean a
channel joining two rivers.
Cost is increasingly a factor in production of books
and this may explain the absence of any colour plates.
Owls of the World
By J. Duncan. 2003. Key Porter Books, Toronto, Ontario.
319 pages. Hardcover, $60.
This beautiful book actually fulfills the two roles that
many natural history books aspire to but usually fall
short of ... being both a detailed scientific reference
and easy to read. The first two-thirds of the book cover
the biology and life history of owls, stories by people
who have uncovered these amazing details and the
methods they used, the substantial role that owls have
in mythology around the world, and threats to their sur-
vival (but also the passion and hope that people hold
out for owls). The U.S. edition, published by Firefly
Books, carries the apt subtitle “their lives, behaviour
and survival”.
The last third of the book is a detailed look at 205
species of owls: physical description, habitat, natural
history, general distribution and their global conserva-
Book REVIEWS
103
In my mind this argues for including a CD/DVD with
the book to carry photographs of habitats and coloured
depictions of fishes. The text can also be put on the disk,
making the book searchable. I still prefer a book in the
hand to a computer screen but suspect this does not
apply to a younger generation. Even the CD in a pock-
et at the back of the book is a transitional stage and
“books” should now appear on the internet, thereby
saving trees, allowing a plethora of colour plates and
other illustrations, enabling the author to receive and
incorporate feedback and new data, and facilitating
correction of errors of fact and of omission. A “book”
on a website can be built up incrementally although
some will inevitably fail to reach a completed form.
Each stage allows for criticism and correction, from
the initial checklist of species, the identification keys,
distribution maps (allowing zoom-in for finer details
than part of a book page allows), and finally text
descriptions.
For those worried about vanishing websites, EM
pulses and cyber trashing, a printout of say 20 copies
of the digital “book” can be deposited in various
libraries. This trend can be seen in some scientific
works such as the “Annotated Checklists of Fishes”
produced by the California Academy of Sciences which
are on-line at that institution but have copies deposited
in a minimum of six natural history institutions world-
wide, admittedly as CDs.
This is the definitive book on this fish fauna and an
essential tome for any student of the aquatic environ-
ment in British Columbia. Future versions will require
multiple authors to match and improve on the experi-
ence and knowledge exhibited here by Don McPhail.
BRIAN W. CoAD
Canadian Museum of Nature, P.O. Box 3443, Station D,
Ottawa, Ontario K1P 6P4 Canada
tion status. Even a quick look highlights how difficult
it is to study some of these species, as many do not have
photographs.
Duncan brings not only his own wealth of knowl-
edge and field experience (particularly with Great Gray
Owls) to this task, but also those of a “whoo’s whoo”
in the world of owls, such as Robert W. Nero, Irina
Menyushina, C. Stuart Houston, and Aki Higuchi.
Duncan’s use of anecdotes helps to de-mystify sci-
ence and raises the readability level, such as when he
relates being dared to taste a freshly regurgitated owl
pellet in front of a crowd. He said it tasted “sweet,”
but his incredulous darer (R. W. Nero) bit into it and
quickly spat it out, declaring that it tasted “bitter.” They
were both right ... the pellet is coated with a pH-neutral
film of mucous that protects the owl’s throat and mouth
from the low-pH acid-soaked mass of fur and bones,
which Nero had bitten into!
104
As with many species, owl populations are often in
trouble around the world for the usual reasons: habitat
destruction, direct (vehicles, shooting, trapping) and
indirect (hitting barbed wire fences) mortality, and in-
vasive alien species such as West Nile virus. But Dun-
can argues that an awareness and understanding of
how people have perceived owls in the past may help
support efforts to conserve them today.
The one small section of the book that many people
will struggle to read describes the DNA sequencing
and classification of owls. It would have been more
understandable if Duncan had linked this highly tech-
nical description, especially the Maximum Likelihood
Tree showing ow] relationships, to some of the species
accounts, where it is obvious from a glance at range
maps for many of the Scops Owls in the Malaysian
THE CANADIAN FIELD-NATURALIST
Vols 121
archipelago that these species probably had a com-
mon ancestor, and allopatric speciation occurred due
to island isolation.
The photos in this book are luscious, with rich colour.
Today, I randomly opened it and there was a full-page
picture of Verraux’s Eagle Owl — the best photo that
I’ve seen of this owl that adequately shows its pinkish-
purple eyelids. I recall driving along a sandy track in
Chobe National Park in Botswana when one of my
companions excitedly sputtered “purple eyelids, pur-
ple eyelids,” as she pointed at the bird that had caught
our attention in the guidebook!
Excuse me while I read up on the Tucuman Pygmy
Owl before heading off to Argentina ....
Cynpbi M. SMITH
Box 5, Waterton Park, Alberta TOK 2MO Canada
Ecology, Conservation, and Status of Reptiles in Canada
Edited by C. N. L. Seburn, and C. A. Bishop. 2007. Herpeto-
logical Conservation volume 2. Society for the Study of Am-
phibians and Reptiles, Salt Lake City, USA. x + 246 pages.
illus., hardbound. Available from the Publications secretary
of SSAR, ssar@herplit.com/U.S. $40 plus shipping.
This book is an edited compilation of peer-reviewed
contributions on several aspects of conservation and
more specifically, on the ecology of the reptiles that
occur in Canada. It contains chapters by many (figu-
rative) heavyweights of Canadian herpetology, who
bring combined centuries of experience to the proj-
ect. The book is a very good application of accumu-
lated biological knowledge to the question of conser-
vation. It is also a good example of how a professional
community can collaborate in a common cause.
It begins with a short introduction, then a histori-
cal review of work done on reptiles in Canada. Next
are four chapters summarizing the life history of
lizards, snakes, freshwater turtles, and marine turtles.
Contributions on the applications of genetics to con-
servation, health and disease in reptile populations,
and traffic mortality follow. The future of Canadian
reptiles and a strategy for conservation are discussed.
The book ends with an illustrated checklist of species,
with the official conservation status of each. There is
also a comprehensive summary of literature, both his-
toric and recent, with few omissions.
The book succeeds in its aim of providing a sum-
mary of the biology and conservation status of Cana-
dian reptiles. The chapters are of a consistently high
calibre — a testimony to the knowledge of the authors
and the thoroughness of the editors. The book is use-
ful on more than one level — as a summary of reptile
ecology in Canada and as a conservation guide; the
extensive literature is also a valuable resource.
Several themes are common to the contributions.
Reptiles living in Canada face two problems — the cli-
mate and humans. Because climate affects life history,
reptiles are concentrated in southern Canada, where
it’s warmer. Unfortunately for the reptiles, humans are
concentrated in the same areas. Human activities that
result in habitat modification are usually in direct
conflict with the best interests of reptiles.
The book makes the point that although the Cana-
dian climate may not be ideal for reptiles, they have
adapted to life here. A reader soon realizes that many,
if not all, declines in reptile numbers are due to human
action. For example, road mortality is a common theme
in several chapters. Human alterations to the landscape
can have direct effects such as habitat destruction, and
also secondary effects such as increases in reptile pred-
ators like cats and raccoons. Several authors make the
point that declines in reptile, especially turtle, popula-
tions can occur slowly, drawing little public attention.
Although I was familiar with much of the informa-
tion presented, there were some surprises, one being
the high number of marine turtles off the Atlantic and
Pacific coasts. This is but one example of how much
is still not known about our reptile fauna.
Quibbles are few and minor. Colour variations and
aberrant scutellation in turtles should not be included
among diseases and parasites; the authors themselves
admit that these “..are not detrimental...” (page 149).
What is the prognosis for reptile conservation? Al-
though knowledge of the need for conservation is high
among the public, and although the number of conser-
vation programs is high, the future still looks bleak,
because of the juxtaposition of human and reptile pop-
ulations. The point is made repeatedly that while atlases
and monitoring programs are good, they are ultimate-
ly ineffective without concrete action; “...ecological
knowledge alone will not save species when the real
problems facing them are social and political.” (page
47).
Ross D. MACCULLOCH
Department of Natural History, Royal Ontario Museum,
100 Queen’s Park, Toronto, Ontario M5S 2C6 Canada; e-
mail: rossm@rom.on.ca
2007
BOOK REVIEWS
105
Rodent Societies: An Ecological & Evolutionary Perspective
Edited by J. O. Wolff and P. W. Sherman. 2007. The University
of Chicago Press, Chicago, 610 pages. U.S. $49. Paper,
$125 Cloth.
This opus on rodent behaviour brings together 61 re-
searchers in a single edited volume. There are 42 chap-
ters grouped into various topics including systematics,
sexual behaviour, life histories, behavioural develop-
ment, social behaviour, antipredator behaviour, com-
parative socioecology, conservation and disease. The
main objectives are to present the latest research on
the behaviour of rodents and to place it in the context
of ecology and evolution. The resultant compilation of
papers is firmly based on comparative biology, which
goes beyond merely describing phenomena and ex-
plains it in terms of the larger ecosystem and within a
phylogenetic framework. There is an emphasis on
hypothesis testing and explicit experimentation that
results in robust and scientifically-sound studies. This
approach fostered by the editors gives more than just
summaries of particular topics but highlights the on-
going nature of our understanding of behaviour in not
only rodents but also as related to other organisms.
At over 2200 species, Rodentia is the most speciose
order of mammals and accounts for approximately 40%
of the diversity. In terms of breadth of coverage, almost
half of the chapters summarize behavioural data across
all major lineages of rodents or within a higher-level
classification. In addition, many different groups or spe-
cies of rodents are directly studied as case examples in
this compendium, with taxa examined ranging from
squirrels to beavers to rats to capybara.
After an introductory chapter on rodents as model
systems, the evolutionary background is set with a
paper discussing phylogenetics and biogeography. This
is followed by several chapters on sexual behaviour
dealing with mating and reproductive strategies. Life
histories of rodents are covered by topics on dimor-
phism, sex ratios, stress, dispersal and philopatry, gene
dynamics, and self-regulation. There is discussion of
behavioural development such as neural regulation,
ontogeny, learning, and kin recognition. Social behay-
iour is examined with papers on parental care, ecology
of sociality, scent marking, non-parental infanticide,
monogamy, and pacifism. There is a small section with
three chapters on antipredator behaviour including
Seashells: Jewels from the Ocean
By Budd Titlow. 2007. Voyageur Press, 729 Prospect Avenue,
P.O. Box 1, Osceola, Wisconsin 54020. 112 pages. US $20.
Cloth.
Sea shells have fascinated people since the dawn of
time. Intrinsically beautiful, they have inspired artists
and architects, been used to fashion tools, and have
even served as money. They are also the prize of col-
lectors the world over. All of these themes feature in
alarm communication. Comparative socioecology has
several papers on social organization and structure of a
number of different species or higher-level groups. The
book ends with a section on conservation and disease
followed by a chapter summarizing conclusions and
future directions on the research of rodent behaviour.
One weakness in an otherwise comprehensive anthol-
ogy was the lack of molecular study on the genetic basis
for much of the evolution of behaviour. In particular,
DNA sequence data nowadays seems to be churning
out by the genome. Was this an editorial oversight, or
is there just nobody looking into this field of research
for rodents? If so, this highlights an area that needs
attention, especially since there are several candidate
model-system species in rodents to choose from. Like-
wise, I found it odd that only four of the chapters have
figures of phylogenies, considering that an evolutionary
perspective is supposed to be one of the major underly-
ing themes of the volume. Perhaps this is an indication
that there is still an obvious gap that requires bridg-
ing between micro- and macroevolution.
Another observation is that over 80% of the authors
are based at institutions in the Americas. Has this biased
the presentation of the state of knowledge of rodent
behaviour? A more subjective criticism is the com-
bined literature cited section for the whole book. For
edited volumes, I prefer references listed separately
at the end of each chapter, especially when topics are
as varied as this monograph. However, there are some
advantages such as the elimination of repeated cita-
tions for general publications on rodents and a single
compiled source useful for searching purposes. None-
theless, these minor detractions by no means diminish
from the scientific merits of the book.
This book is definitely aimed at the specialist re-
searching the behaviour of rodents, but will also be of
interest to biologists working on other organisms and
studying similar social systems. At over 600 pages,
the price is reasonable for purchase by a university
library, or would be a valuable addition to any ecolo-
gist’s bookshelf because it is packed with the latest
information on ethology.
BURTON K. LIM
Department of Natural History, Royal Ontario Museum,
100 Queen’s Park, Toronto, Ontario M5S 2C6 Canada
Budd Titlow’s handsome book Seashells: Jewels from
the Ocean.
This relatively slender volume, copiously illustrated
with photographs by the author or from stock footage,
provides a good introduction to the world of mollusks
and the shells which adorn so many of them. Succes-
sive chapters deal with the physiology of mollusks, their
classification, their habitats, their cultural and econom-
106
ic importance to humans, and the threats that mollusks
face. The latter include marine and land-based sources
of pollution and over-exploitation of commercially
valuable species.
There is also a primer on shell collecting, and also
short notes on good collecting sites in North America.
At its heart, this is a book aimed at shell collectors, be
they casual beachcombers or fanatical conchologists.
The former will learn much, the latter will no doubt
admire the stunning photographs. There is a distinct
American bias, both in the selection of photographs
and in the shell collecting sites profiled; indeed at times
the book reads as an ode to Sanibel Island, Florida,
the Mecca of American shell collectors.
The book oscillates between straightforward expla-
nations in layman’s terms of the science underpinning
malacology, and a rather folksy, and at times highly
personal, appreciation of the cultural aspects of sea
shells and shelling. While occasionally rather affected,
on the whole it is highly readable. Many of the pho-
tographs selected sacrifice ecological verisimilitude
for art; for example, there are several very beautiful
assemblages that would be highly improbable in nature.
And, belaying the book’s American bias, it would be a
fortunate day indeed where one found a Spider Conch
(from the tropical Indo-Pacific) on a North American
beach. For a field naturalist, the book is faintly disap-
pointing in that it tends not to identify in specific terms
THE CANADIAN FIELD-NATURALIST
Vol. 121
most of the shells portrayed. Shelling, unlike birding,
is One pastime where getting a grip on the Latin ter-
minology and the taxonomy is essential; for this the
reader would be well advised to turn to any one of the
several excellent works cited in the short but useful
reference section. By the same token, a few sketches
illustrating such things as mollusk morphology would
have been helpful.
There are a few errors in the text; for example the
swans that winter at Chesapeake Bay are Tundra Swan,
not Trumpeter Swan, but these errors do not detract
from the overall value of the book. One somewhat dis-
appointing aspect of the book, and one in counterpoint
to the otherwise conservationist approach, is a section
dealing with the collection of live specimens. While
pale in comparison to other threats, collectors can pose
a considerable threat to vulnerable species and it would
have been preferable if the author had concentrated
the reader’s attention on the collection of non-living
material. That said, the author does emphasize the need
to respect local regulations. In conclusion, this book
is not a scientific reference; however, it does provide
a useful primer for anyone keen to explore the world
of shelling, and a handsome addition to any collection
of “coffee table” books.
MARK GAWN
Mission of Canada, 5 Avenue de |’ Ariana, Geneva 1202
Switzerland
Turtles: An Extraordinary Natural History 245 Million Years in the Making
By Carl J. Franklin. 2007. Voyageur Press, 729 Prospect
Avenue, PO Box 1, Osceola, Wisconsin 54020. 160
pages. U.S. $35.
Turtles provides a brief overview of the turtles of
the world along with abundant colour photographs of
many of the species. The book is divided into two parts.
Part 1, the life and lifestyle of turtles, consists of three
chapters. The first chapter covers the taxonomy and
internal biology of turtles in roughly 10 pages. Chap-
ter two, the ecology of turtles, covers topics such as
thermoregulation, feeding, reproduction, and preda-
tors. The third chapter, covering the origin and fossil
history of turtles, is only 3 pages long. Part 2, the diver-
sity of modern chelonians, consists of two chapters.
One chapter covers the families of Pleurodira, or side-
necked turtles, and the other discusses the families of
Cryptodira, or hidden-necked turtles. For each family
account there is a map illustrating the global distribu-
tion of the family, then text discussing shared traits
among members of the family and then information
on many of the species within the family.
Roughly two-thirds of the books is allocated to Part
2, the family accounts. I found this an unsatisfactory
balance. Part 1, which should provide a coherent pic-
ture of the various different turtle life history strategies,
is just too short and superficial. Likewise, the family
accounts remain superficial because they do not pro-
vide systematic species accounts. The information sel-
ected for individual species should highlight the unique
qualities of that species, but all too often the species
profiles rely on just some basic numbers (size, number
of eggs laid), distribution (which countries it occurs
in) and some general habitat info.
The text also has a number of glaring mistakes. In a
text box on Linnaean taxonomy of turtles, the author
states that turtles belong to the class Anapsida (page
14). In Linnaean terms, Anapsida is the subclass,
while Reptilia (or Chelonia, for splitters) would be the
class. The author claims that both McCord’s Box Tur-
tle (Cuora mccordi) and Zhou’s Box Turtle C. zhoui)
are believed to be extinct (pages 39-40), when really
he should state they are likely extinct in the wild, as
both species still occur in captivity. The author makes
use of the most recent phylogenetic work on the genus
Clemmys, by transfering the Wood Turtle (Glyptemys
insculpta) to its new genus, but still leaves the Bog Tur-
tle (G. muhlenbergii) in the genus Clemmys. Other
careless errors include stating that there is more than
one species within the genus Actinemys (page 98), and
claiming that there are four species of Painted Turtle
(Chrysemys picta; page 101), when really there are four
subspecies. The Indian Flapshell Turtle (Lissemys
2007
punctata) receives two different and somewhat con-
tradictory species accounts, one brief account under
its correct subfamily (Cyclanorbinae; page 134), and
a longer account under an incorrect subfamily (Tri-
onychinae; page 139). Despite these errors, there are
some good things about this book. Part | does pro-
vide a very brief and readable introduction to the world
Whales and Seals Biology and Ecology
By Pierre-Henry Fontaine. 2007. Schiffer Publishing, 4880
Lower Valley Road, Atglen, Pennsylvania 19310 USA.
448 pages. U.S. $35. Paper.
During my career as a marine mammal research
scientist I was reluctantly dragged in on several occa-
sions to necropsy and dispose of large dead mammal
carcasses found along the beaches of Canada’s east
or west coast. Almost always these mammoth moun-
tains of mammalian flesh were in an advanced stage
of putrefaction. Upon opening the body cavity one
would confront what has been termed a “gaseous pud-
ding” and it was challenging to properly identify even
the most conspicuous organs. It is with this in mind
that I say that I am impressed with the work in hand,
which has yielded for the author a large amount of
useful, detailed information, gathered with fortitude
and persistence, from a source that many would have
been wont to avoid.
The illustrations, particularly the excellent photo-
graphs of the organs and musculature, will serve as a
valuable resource for field biologists learning their way
around marine mammal carcasses. The author’s expert-
ise in marine mammals comes primarily from his exten-
sive experience in performing necropsies of beach cast
cetaceans and to a lesser extent pinnipeds. This book
uses this as the point of departure relating the specific
adaptations, the senses, nutrition, and locomotion, to
the anatomical structures which he has studied and
illustrated during his years of studies.
The remainder of the book attempts to give a gener-
al account of whales and seals, particularly from the
North Atlantic, where the author has worked. The chap-
ter on strandings is devoted to the question of why live
cetaceans beach themselves. As the author points out,
almost every imaginable reason has been offered from
geomagnetic anomalies to suicide. The chapter leaves
one hanging as do most articles on this subject. One
very important recent work on this subject is the mono-
graph Marine Mammals Ashore (J. R. Geraci and V.
Loundsbury. 1993. Texas A&M Sea Grant Publica-
tion). This not mentioned by the author and would be
a valuable source of additional information.
The chapter on mounting marine mammal skele-
tons will be found especially useful by museums and
institutions wishing to do such displays. Little detailed
BoOoK REVIEWS
107
of turtles. The photographs are generally quite good.
Ultimately, however, this book could have been great-
ly improved by a thorough review from an expert on
turtles.
DAVID SEBURN
Seburn Ecological Services, 2710 Clarenda Street, Ottawa,
Ontario K2B 785 Canada
information exists on this subject and the author has
learned valuable lessons in avoiding errors in skeletal
reconstruction from his numerous and careful field
dissections.
The remaining chapters, while not based on the au-
thor’s direct experience, are well researched and illus-
trated. The long association of people with whales and
seals is presented in an interesting manner. Some spe-
cific examples of marine mammal hunting in the St.
Lawrence are little known. The technique on Walrus
hunting in the Magdalene Islands, which led to the
early extinction of the southern-most Atlantic Walrus
stock, is one such example.
The chapter dealing with the complicated subject
of marine mammal paleontology gives a good intro-
duction to a discipline which is both difficult to assimi-
late and where the information is not well organized in
any single comprehensive source. This will serve as a
good point of departure for students who want to learn
more about the origins of marine mammals.
The last chapters, the fact sheets on Cetaceans and
Pinnipeds, might be considered useful additions to this
general work on marine mammals only because they
include the species found in the North Atlantic, which
is where most of the author’s work has been done. One
wonders why the author has included the Northern
Elephant Seal (only found in the Pacific) and why the
Pacific Walrus (Odobenus divergens) is mentioned and
shown in one of the illustrations. The Arctic Ringed
Seal, which is found as far south as the Labrador coast
(Lake Melville) is omitted. A number of up-to-date
field guides exist, which give a comprehensive account
of the seals and whales in the North Atlantic and would
serve their readers more effectively.
I recommend this book to the non-specialist who
has an interest in whales in particular. It will also be
a useful source of information for field biologists and
naturalists who might deal with beach cast remains of
marine mammals. It succeeds in what the author set
out to accomplish by sharing his considerable know]-
edge of a subject, in which he has thoroughly immersed
himself.
THOMAS G. SMITH
EMC Eco Marine Corp, 5694 Camp Comfort Road, Beaulac-
Garthby, Quebec GOY 1B0 Canada
108
ENVIRONMENT
China Ecosystems
Sun Honglie, Editor-in-chief: 2005. The Science Press of China.
1823 pages. 350 Yuan RMB Cloth.
The publication of such a magnum opus can be re-
garded as a landmark achievement accomplished by
this group of Chinese ecologists. The book summed
up the main results of the long-term research on the
ecosystem structure and function at more than 80 eco-
logical research stations, extensively distributed over
China, of the Chinese Academy of Science. The re-
search comprehensively covered the various fields such
as ecosystem classification, ecosystem structure and
dynamics, ecosystem energy flow, water and nutrient
cycling in ecosystems, ecosystem management and
sustainable use and so on.
China is a vast territory, richly endowed by nature
and abounding in diverse types of ecosystems. Tradi-
tional Chinese integrative philosophy, an abundance
of resources and the fast increasing economic power
confer on China satisfactory conditions and capacity for
ecosystem research. However, in China, the initiation
of ecosystem research was rather late compared with
many other countries. At the beginning stage in the
1950s, ecosystem research in China was very rare and
scattered. This was nearly stopped during the ten-year
period of the Cultural Revolution. China began to par-
ticipate in the large international ecosystem research
plans, such as IBP [International Biological Program],
MAB [Man and Biosphere Program], IGBP [Interna-
tional Geosphere-Biosphere Program], GTOS [Glob-
al Terrestrial Observation System], EHA [Ecosystem
Health Assessment], PAGE [Pilot Analysis of Global
Ecosystem] [Pesticide Assessment Guidelines?] and
MA [Millennium Ecosystem Assessment] from the
late 1970s. Driven by these international programs,
Chinese ecosystem research developed quickly, espe-
cially in recent years.
Long-term observational studies play an irreplace-
able role in understanding and assessing the ecosys-
tem structure, function and their health status. In this
course, the ecological research stations made great
contributions. The Chinese ecological stations, which
are distributed throughout the typical ecological zones
of China, have accumulated long-term continuous and
dynamic data, not only on the changes of the same
ecosystem types under different driving forces, and on
the integrated features of different types of ecosystems
at the regional scale, but also on the long-term ecologi-
cal experiments and management practices conducted
in these stations.
Most of Chinese ecosystems were generally healthy
in the past; however, in recent years, at least some of
them have undergone serious degradation, and some
are even close to irreversible damage. Not long ago,
China Environmental Protection Administration an-
THE CANADIAN FIELD-NATURALIST
Vol. 121
nounced the first Report on the Ecological Protection in
China, which admits that the trend of environmental
deterioration in China has not been effectively curbed
yet, and more than 60% of lands are becoming envi-
ronmentally fragile. Obviously, to achieve sustainable
development, China must continually improve its dete-
riorating environment, maintain and continuously up-
grade the capacity of its ecosystem service, which needs
a sound understanding of the effective management
of various types of ecosystems in China.
The results of the long-term research on the Chinese
ecosystems were achieved with a background of large
temporal and spatial environmental changes. There-
fore, the data accumulated by the Chinese ecological
research stations (data used in this book), include the
information not only on the progressive but also de-
grading succession of Chinese ecosystems, and the
summary of some restoration practices. These results
are particularly unique and valuable, and could be used
as references for some other regions and countries.
The objective of the Millennium Ecosystem Assess-
ment of the UN, which involves a large number of
ecologists from different regions and countries, is to
systematically and comprehensively reveal the trends
of changes of various ecosystems, and accordingly pro-
pose effective countermeasures for the future. China
ecosystems is certainly helpful for such an assessment
and its publication is timely.
The book is composed of eight parts. The first part
is General review, including Chapter one: the history,
evolution and perspective of the ecosystem research;
Chapter two: the natural conditions for the formation
of the Chinese ecosystems; Chapter three: the evolu-
tion of the Chinese ecosystems; Chapter four: the im-
pacts of human activity on the ecosystems; Chapter
five: the principles and the systems of the Chinese eco-
system classification.
The second part is Forest ecosystems, including
Chapter one: introduction; Chapter two: cold-temper-
ate zone northern conifer forest ecosystem; Chapter
three: middle-temperate zone conifer-broad-leaved for-
est ecosystem; Chapter four: warm-temperate zone
deciduous forest ecosystem; Chapter five: subtropical
zone evergreen and deciduous broad-leaved mixed for-
est ecosystem; Chapter six: subtropical zone evergreen
broad-leaved mixed forest ecosystem; Chapter seven:
subtropical zone western mountainous region ever-
green hard-and-broad-leaved forest ecosystem; Chap-
ter eight: subtropical zone alpine conifer forest and
mountainous region conifer-broad-leaved mixed for-
est ecosystem; Chapter nine: tropical zone seasonal
rain forest ecosystem; Chapter ten: tropical zone rain
forest ecosystem; Chapter eleven: sustainable devel-
opment of China forest ecosystems.
2007
The third part is Grassland ecosystems, including
Chapter one: the formation and classification of the
grassland ecosystems; Chapter two; temperate zone
grassland ecosystems; Chapter three: high and cold
region grassland ecosystems; Chapter four: warm-
temperate zone grassland ecosystems; Chapter five:
tropical zone grassland ecosystems.
The fourth part is China’s water ecosystems, in-
cluding Chapter one: introduction; Chapter two: fresh
water ecosystems; Chapter three: lake ecosystems;
Chapter four: the succession and primary productivity
of the lake ecosystems; Chapter five: ocean ecosystems.
The fifth part is Farmland ecosystems, including
Chapter one: introduction; Chapter two: northeastern
Song-nen Plain farmland ecosystems; Chapter three:
lower field of Liao River Plain farmland ecosystems;
Chapter four: north China Taihang Piedmont Plain
farmland ecosystems; Chapter five: Huang-huai Plain
farmland ecosystems; Chapter six: Tai Lake Plain farm-
land ecosystems; Chapter seven: South China hilly region
farmland ecosystems; Chapter eight: Dongting Lake
Plain farmland ecosystems; Chapter nine: Central Sichuan
hilly region farmland ecosystems; Chapter ten: Loess
Plateau farmland ecosystems; Chapter eleven: oasis
farmland ecosystems in desert regions.
The sixth part is nutrient cycling in terrestrial eco-
systems, including Chapter one: a review on the re-
search progress of biogeochemical cycling; Chapter
two: cycling of carbon and main nutrient elements in
farmland ecosystems; Chapter three: cycling of main
New Zealand — A Natural History
By Tui De Roy and Mark Jones. 2006. Firefly Books Ltd.,
Richmond Hill, Ontario. 160 pages. $40. Hardcover.
The authors, experienced professional photogra-
phers Tui De Roy and Mark Jones, moved to New
Zealand about 16 years prior to the publication of the
book, and, during this time, they explored many of
New Zealand’s distinct and often rugged and remote
natural regions. The result of these explorations and
hard work is this beautifully illustrated coffee table
book containing more than 450 magnificent photo-
graphs which offer a wonderful introduction to many
of the unique species and habitats of New Zealand.
In keeping with one of the main areas of expertise of
the authors — nature photography, the book is largely
made up of beautiful pictures. Early in the book,
there is a helpful map, which includes many of the
locations and national parks referred to later on. The
book also contains a reasonably detailed and useful
index. As a biologist, I appreciated the inclusion — in
an appendix just before the index — of a list of scien-
tific and common names for many of the species
named in the previous pages.
The text sections are fairly brief overall. I counted
39 pages covered mainly by text (not including the
acknowledgments and index), out of the total 160 pages
Book REVIEWS
109
nutrient elements in forest ecosystem; Chapter four:
cycling of main nutrient elements in grassland ecosys-
tems.
The seventh part is water cycling in terrestrial eco-
systems, including Chapter one: introduction; Chapter
two: water cycling in farmland ecosystems; Chapter
three: water cycling in forest ecosystems; Chapter four:
water cycling in grassland ecosystems.
The eighth part is the theories and methods for the
ecosystem network research, including Chapter one:
the purposes and tasks for the ecosystem network re-
search; Chapter two: the establishment and develop-
ment of the ecosystem network; Chapter three: the
methods for the ecosystem network research; Chapter
four: the scales for the ecosystem network research.
The book was well written with few errors. Abun-
dant illustrations are helpful for readers to easily under-
stand the research results. The book is suitable for the
professionals who engage in ecology or environmental
science, or other persons who are interested in these
fields.
Li Dezut', Qin AIL’, AND QIN HonG?
'Lab of Urbanization and Ecological Restoration of Shang-
hai; National Field Observation and Research Station in
Tiantong Forest Ecosystem of Zhejiang; Department of
Environmental Science, East China Normal! University,
3663, Zhongshan Rd (N). Shanghai, China 200062)
*Jilin Forestry Staff School, Jilin, China
Zhejiang University, Hangzhou, China
of the book. Thus, if one is planning a visit to New
Zealand, reading the book prior to the departure may
be easier than putting such a weighty, hardcover tome
in the luggage.
Although brief, the text is not without merit. The
authors have done their best to include information
on a wide variety of animals and plants, as well as a
few details about New Zealand’s geological past and
present. There are some interesting, although usually
very short, stories about several species, including fist-
sized giant carnivorous land snails — some of them
endemic (restricted) to a single hill — that hunt large
earthworms at night (some of the local earthworms
can reach | meter in length) or the inquisitive kea, the
very active mountain parrots of New Zealand, which
are apparently capable of sometimes killing sheep and
dismantling cars. There is a strong conservation theme
throughout the book, and the main purpose of the text
is clearly to highlight the plight of the many endan-
gered species unique to New Zealand, and to discuss
some of the conservation programs undertaken to save
these rare species. This is a worthy goal, of course, and
the authors’ commitment to conservation is evident
throughout.
However, as a scientist, I would have liked to see a
more balanced and in-depth discussion of certain envi-
110
ronmental and conservation issues. The authors seem
to take an uncritical approach towards some controver-
sial control programs undertaken by the local Depart-
ment of Conservation against species introduced to
New Zealand by people, over the centuries. It is some-
what unsettling, for example, to read about the “Her-
culean aerial poisoning operation” — “the largest rodent
control program ever undertaken anywhere in the
world”, on the same page where the authors refer to
the “humane integrity” associated with the conserva-
tion initiatives such massive poisoning operations are
a part of. It should perhaps have been mentioned, for
example, that such massive poisoning operations can
occasionally backfire, and affect rare native species as
well. For instance, poisons used to eradicate mice also
killed the North Island saddleback, a rare New Zealand
native forest bird (Davidson and Armstrong 2002).
Also, the magnitude of the effects of introduced herbi-
vores on New Zealand’s native plants can sometimes
be difficult to determine (Veblen and Stewart 1982).
In addition, it is worth noting that some species, such
as the swallows and silvereyes mentioned by De Roy
and Jones, expanded their ranges naturally into New
Zealand in recent times. When it comes to the many
interactions among introduced species and their new
environments, the story is often considerably more
complex than this book suggests. Thus, the repeated
demonization of introduced species found in the text
is often overly simplistic.
The authors often mention the negative impact of
introduced wild mammals such as rats or Australian
brush-tailed possums in New Zealand, but, curiously,
seem to largely ignore the huge impact of New Zea-
land’s domestic animals, particularly the country’s
enormous sheep population. According to Ponting
(1993), there are more than 70 million sheep and 8
million cattle in New Zealand, and many of the habi-
tats of this country have been changed drastically and
MISCELLANEOUS
THE CANADIAN FIELD-NATURALIST
Vol. 121
irreversibly as a result. Also largely ignored in the
book is a discussion of the early, pre-European, human-
caused species extinctions. New Zealand’s fauna used
to be dominated by very large (the largest ones reached
3 meters in height) flightless birds called moa. These
birds became extinct mainly due to overhunting by
the Maori — the country’s first human colonizers (Ellis
2004). The disappearance of the moas had important
effects on some of the native plant species these giant
birds were browsing on and had coevolved with. Thus,
although conservation is, of course, a very important
goal, it is also essential to recognize that nature is
dynamic, and that the initial, pre-human colonization
web of life found in New Zealand long ago can no
longer be re-created due to the many irreversible
changes that have taken place since then. A much more
thorough and detailed discussion of New Zealand’s
prehistoric life and ecology can be found in the book
by Worthy and Holdaway (2002).
Despite the shortcomings, the current work by De
Roy and Jones represents a nice introduction to the
unique natural world of this fascinating and remote
country.
Literature Cited
Davidson, R. S., and D. P. Armstrong. 2002. Estimating impacts of poison
operations on non-target species using mark-recapture analysis and simula-
tion modelling: an example with saddlebacks. Biological Conservation 105:
375-381.
Ellis, R. 2004. No turning back: The life and death of animal species. Harper
Collins Publishers, New York. 428 pages.
Ponting, C. 1993. A green history of the world: The environment and the
collapse of great civilizations. Penguin Books, New York. 430 pages.
Veblen, T. T., and G. H. Stewart. 1982. The effects of introduced wild animals
on New Zealand forests. Annals of the Association of American Geographers
72: 372-397.
Worthy, T. H., and R. N. Holdaway. 2002. The lost world of the moa:
Prehistoric life of New Zealand. Indiana University Press, Bloomington and
Indianapolis. 575 pages.
RADU CORNEL GUIASU
Environmental and Health Studies Programme, Glendon
College, York University, 2275 Bayview Avenue, Toronto,
Ontario M4N 3M6 Canada
Beneath My Feet: The Memoirs of George Mercer Dawson
By Phil Jenkins. McClelland & Stewart, Toronto. 2007 Hard-
cover. 350 pages. 18 photographs, 6 sketches. $34.99.
Phil Jenkins set out to write the biography of George
M. Dawson, the son of Sir John William Dawson, the
first principal and then first president of McGill Uni-
versity. Jenkins has instead taken nearly 5000 of George
Dawson’s letters, most of them previously unpublished,
from the McGill University archives, and has wisely
allowed Dawson to write his own “autobiography.”
Jenkins has created an introduction in Dawson’s style,
has abridged an article published by Dawson in Harp-
er’s Magazine, and has written over 5% of the text (we
don’t know which segments) to fill in gaps not avail-
able in Dawson’s own words. The result is a personal,
frank, well-written, entertaining life of George M. Daw-
son, CMG, LLD.
A designer, “Mr. Richardson” (unaccountably, Jenk-
ins does not provide his first name) has produced a
charming book cover that fits perfectly with what would
have been expected in the 1890s.
George M. Dawson was crippled in late childhood
by spinal tuberculosis which left him a hunchback
and a good foot shorter than he should have been. Nev-
ertheless as an adult he struggled manfully through
arduous hiking, canoeing and horseback riding that
would have deterred many men of normal stature. He
obtained superb training at the Royal School of Mines
in London, England, where he was taught by Professor
2007
Thomas Henry Huxley, a leading proponent of Dar-
win’s theory of evolution. Young George was fav-
ourably inclined to Huxley’s views and had spirited
discussions on the subject with his father.
George M. Dawson was a geologist, author, teacher,
civil servant, anthropologist, palaeontologist, surveyor,
mapper, photographer, artist and poet. A number of his
photographs, sketches, and poems are interspersed
throughout this book. Although he was named geolo-
gist and naturalist to the North American Boundary
Commission in 1873-74, his skills in natural history
were weak (Jenkins fails to share this fact). In north-
ern Montana, just south of the 49" parallel, he observed
the last great herds of Bison and came across skele-
tons resulting from the slaughter of 21 plains Indians.
Dawson named a number of geographic features in
the Queen Charlotte Islands and the interior of British
Columbia that are still in use today. He suffered the
pain of unrequited love and remained a bachelor until
his premature death from an acute chest infection at
age 51.
I agree with Jenkins (page 2), that Dawson was “a
writer ... of endearing wit, evocative description, and
illuminating fact.” He described an Orkneyman in the
employ of the Hudson’s Bay Company as “an amiable
Hudson Bay Fossil.” He used the term “liquid excis-
able articles” to describe spirituous liquors.
This book has not achieved its full potential. I spot-
ted the following misspellings and factual errors, pre-
sumably made by Dawson and not by Jenkins: alumni
(when alumnus was indicated), latitude (when longi-
tude was meant), astrological (astronomical), penin-
sular (peninsula), tit bits (tid bits), 100 meridian (110"
meridian), prairie chickens (Sharp-tailed Grouse), Dr
Helckin (Dr Helmcken), F.R.C. (E.R.S.), silver load
(silver lode), and Bearing Sea (Bering Sea). There is
The Wolf Totem
By Jiang Rong. 2004. Changjiang Literary Art Press. Pages:
478; $18. Yuan RMB.
[Editors note: The Penguin Group, under the British
Pearson Education Company, bought the copyright and
will publish an English version in 2007. This researched-
based novel has topped China’s bestseller list for 16
straight months, selling about | million copies. ]
The Chinese press has not published a significant
book in many years. However, in April of 2004, the
Changjiang Literary Art Press of China published the
novel-style book The Wolf Totem, which was written
by an author bearing the pseudonym Jiang Rong. The
response from the media and the public was rapid and
it became a best seller.
The book is composed of various relatively inde-
pendent stories. The narration of each story is full of
craftsmanship, firing one’s imagination and stirring
one’s emotion. The novel-style book is a marvellous,
BOOK REVIEWS 11]
little doubt that a University press would have en-
couraged Jenkins to insert the correct spellings in square
brackets, as well as to include an index, explanatory
footnotes (e.g., to add the initials and explain the im-
portance of Dr. W. F. Tolmie, surgeon to the Hudson's
Bay Company), and a bibliography. The latter would
have included Life and Letters of George Mercer Daw-
son, 1849-1901, by Lois Winslow-Spragge, Dawson's
niece, and would have listed the available photocopies
of Dawson’s detailed field notebooks which Jenkins
seems to have ignored. Similarly, we are not told that
Sapper Duckworth was Dawson’s collector and taxi-
dermist in 1873-74, that white cranes were Whoop-
ing Cranes, that the blue jay was probably a Steller’s
Jay, that the hummingbird nest was likely that of a
Rufous Hummingbird, and that the name of the des-
tructive but now extinct Rocky Mountain Locust has
since changed from Caloptenus spretus to Melanoplus
spretus.
A longer introductory essay or a postscript could
have allowed a more complete assessment of Dawson’s
great achievements which Dawson modestly did not
mention. We learn, for example, that Dawson received
an honorary doctorate from the University of Toronto,
but those from Princeton, Queen’s and McGill, and
other high honors, are not mentioned.
In spite of the shortcomings mentioned, Dawson's
skills make this book a treasure, a lasting contribution
to Canadian history. I recommend it highly to the gen-
eral public. It deserves a place in every university and
high school library.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N OJ8
Canada
unprecedented way to present many stories of the
enigmatic animal of the steppe, the wolf. The stories
are not only charming in their sense of humanity, but
also colourful and valuable in their understanding of
the ecological role played by the Wolf. After reading
the book, readers will be enlightened, moved and even
shocked by the vivid ecological ethics conveyed in the
book. The publication of the book can be regarded as
an important event for both literature and ecologists.
From the viewpoint of ecology, a steppe may be re-
garded as a special type of ecosystem seemingly sim-
ple in structure and function, but essentially diverse,
complicated and unpredictable. From the perspective
of humanity, a boundless steppe may be mysterious,
thought inspiring and soul stirring, sometimes touching
and sometimes plaintive. In the cultural evolutionary
history of the world, the acted steppe as an endless well-
spring of artistic creation and a boundless field for sci-
112
entific exploration. The anecdotes and the legends, the
music and the paintings related to the steppe are un-
countable; the study cases concerning the steppe eco-
system are also voluminous.
Turning aside from the fascinating visions and thrill-
ing scenes the author describes in each story, and turn-
ing aside from the totem of wolf at the level of politics
and humanities the author elucidates in his narration,
the book is of important ecological values. The descrip-
tions of the behaviours of the Wolf populations in team-
work, reconnaissance, ambuscade, and surprise attack,
and the unusual wisdom exhibited in making use of
weather and topography, as well as the discussions on
the relationships among the populations of Wolf and
human, sheep and grass can be regarded as very vivid
and enlightening material. However, it is a pity that in
an otherwise valuable book that the author does not
give the basic scientific and quantitative relationships
among the component populations of the ecosystem.
This may be beyond the scope of this style of book.
All these mysterious stories took place in the steppes
of northern China during a special period more than
nearly 40 years ago when the author and his classmates
as urban school graduates laboured in the countryside
in response to the political call of Chairman Mao. At
that time, the vast area of steppe of northern China
had plenty of water and lush grass, creating a beauti-
ful landscape. But afterward, due to overgrazing and
farming, the steppe ecosystem degenerated rapidly.
China is the second largest country in the world in
terms of the steppe area. However, due to the increase
in large scale agriculture in the half century from
1949, 5% of the natural steppe area was turned into
farmland [This constituted one fifth of the total farm-
land of China]. This has resulted in more than a quarter
of Chinese land being threatened by desertification.
More than 90% of the steppe has been degenerated
and such degeneration is expanding at the rate of 2
million km?/yr. Since steppe restoration is becoming
harder and harder in China, all the stories about the
steppe in the past are becoming almost unrepeatable.
This further implicates the extreme rarity and the
important value of the material in this book.
Ecosystems with multiple-trophic levels are com-
plicated and maintaining the stability is critical. In a
typical steppe ecosystem with three trophic levels:
carnivore, herbivore and grass, the carnivore usually
plays a top-down effect on the community structure,
and by this means grass tends to be protected to some
extent. The theory of trophic cascade and HSS hypoth-
esis is useful in explaining, “Why the terrestrial world
tended to be green.’ McLaren and Peterson (1994)
used tree ring data to show how a top-down effect of
Wolf populations on Isle Royale reduced Moose abun-
dance resulting in an increase in the growth increment
of forage trees. The Wolf Totem shows that in the north-
em China about one century ago, harmonious rela-
tionships among the component populations of the
THE CANADIAN FIELD-NATURALIST
Vol. 121
ecosystem did exist, and the relative stability of the
pasture was naturally maintained. This supports the
rationality of the theories as mentioned above. How-
ever, considering the present situations of the rapidly
degenerated steppe ecosystem in northern China, the
above-mentioned theories encounter limitations,
because they do not deal with how and why “the ter-
restrial world turned from green to yellow or black”
due to over-use and human disturbance. If a distur-
bance dimension was added to the theories and each
dimension were given a threshold, the theories would
become more general, compatible and extendable.
The Mongolians, the native people in the northern
China, fully understood the natural laws of mutual
restriction and dynamic balance among the popula-
tion of wolf, sheep and grass in the steppe ecosystem
in their long history dealing with nature, and knew
how to efficiently live within these laws. They never
killed all of the Wolves, never bred the sheep exces-
sively and never over-grazed the pasture. The accu-
mulation of such ecological wisdom imitating nature,
acclimating nature and making use of nature, moulded
the Mongolian nationality. On the contrary, by disobey-
ing the laws of nature, they have unavoidably learnt a
painful lesson about the degeneration of the steppe.
When human or social-economic dimensions are
added into a natural ecosystem, it would become a
social-economic-natural-complex-ecosystem (SENCE).
The present steppe ecosystem in northern China is
such a SENCE. Keeping such a SENCE developing
in a harmonious way is a new challenge to China with
ever-increasing population. The classical Chinese
philosophy, such as The Doctrine of the Mean,
emphasizes human’s active roles such as participation,
promotion, coordination and cultivation rather than
domination, so as to reach harmony between human
and nature. Combining the classical Chinese philoso-
phy with the modern ecological theories and using the
actual experiences and lessons for references, steppe-
like SENCE would be managed more effectively. The
Wolf Totem with its stories of a rural youth living
among Inner Mongolian nomads and wolves tells of
this grassland environment and nomadic culture.
Reading The Wolf Totem will bring the reader a better
understanding of the ecology of the Chinese-Mongo-
lian steppe.
Literature Cited
McLaren, B. E., and R. O. Peterson. 1994. Wolves, moose and tree rings on
Isle Royale. Science 266: 1555-1558.
Li Dezut', QIN AILP, AND QIN HONG?
'Lab of Urbanization and Ecological Restoration of Shang-
hai; National Field Observation and Research Station in
Tiantong Forest Ecosystem of Zhejiang; Department of
Environmental Science, East China Normal University,
3663, Zhongshan Rd (N). Shanghai, China 200062)
“Jilin Forestry Staff School, Jilin, China
Zhejiang University, Hangzhou, China
2007
YOUNG NATURALISTS
Gracie. The Public Gardens Duck
By Judith Meyrick. Illustrations by Richard Rudnicki, 2007.
Nimbus Publishing Limited. Halifax, Nova Scotia B3K
SMB Canada. 32 pages. $16.95. Cloth.
This is a delightful children’s book, told from the
perspective of Gracie the duck. The story is about the
duck’s quest for food crumbs in a public park near the
CBC building in Halifax. Gracie spends most of her
time, quite successfully, getting the attention of differ-
ent people she sees in the park so that they will share
their food with her.
However, one day, people are not giving food to her.
“Mr. Black Suit’, the person who usually feeds her,
does not proffer up any of his muffin; nor does a pair
of school girls, who actually apologize to Gracie. A
young girl almost shares a piece of her peanut butter
sandwich, until the girl’s mother reminds her of the
sign near the park entrance. What sign? What is going
on? Gracie asks herself, puzzled about this change in
her daily feeding regimen.
NEw TITLES
Prepared by Roy John
+ Available for review * Assigned
ZOOLOGY
Book REVIEWS
113
Finally, hungry Gracie reads some of the words on
the sign and discovers that park policy has changed
and feeding the ducks is no longer allowed. Gracie
takes action, but even this doesn’t work. The hungry
duck is then forced to try the insects in the grass and
then some plants in the pond, and finds that these are
very satisfying. Pleased with herself, she now realizes
that there is plenty of food in the park without rely-
ing on people for crumbs such as Mr. Black Suit —
who comments to Gracie that she has adapted nicely.
The story is very compelling and suitable for ages
3 to 8 (at least) —- my 12-year-old niece picked up the
book and enjoyed it as much as her younger sisters
did. The illustrations are dynamic and it is almost as if
the people and ducks in this story are moving as you
read it. Truly a delight — with a good lesson learned
along the way.
ELLEN WOODLEY
Liana Environmental Consulting Ltd., RR 4, Fergus, Ontario
N1M 2W5 Canada
[Editor’s note: The following publications are available from China Scientific Book Services, Jian Nei Da Jie Post
Office, 100001-88, Beijing, China or e-mail order: order-csbs@94book.com.cn, Online order: http://www.-hceis.
The publication dates and costs vary and not all have English text.
Fauna Sinica Invertebrata Volume 1: Crustacea Freshwater
Cladocera
Fauna Sinica Invertebrata Volume 2: Arthropoda Crustacea
Freshwater Copepoda
Fauna Sinica Invertebrata Volume 3: Platyhelminthes:
Trematoda: Digenea (1)
Fauna Sinica Invertebrata Volume 4: Cephalopode
Fauna Sinica Invertebrata Volume 5: Hirudinea
Fauna Sinica Invertebrata Volume 6: Holothuroidea
Fauna Sinica Invertebrata Volume 7: Gastropoda: Meso-
gastropoda: Cypraeacea
Fauna Sinica Invertebrata Volume 8: Arachnida: Ara-
neae:Thomisidae and Philodromidae
Fauna Sinica Invertebrata Volume 9: Polychaeta: Phyllo-
docimorpha
Fauna Sinica Invertebrata Volume 10: Arachnida: Araneae:
Araneida
Fauna Sinica Invertebrata Volume 11: Gastropoda: Opis-
thobranchia: Cephalaspidea
Fauna Sinica Invertebrata Volume 12: Bivalvia: Mytiloida
Fauna Sinica Invertebrata Volume 13: Arachnida Araneae
Theridiidae
Fauna Sinica Invertebrata Volume |4: Sacodina: Acan-
tharia and Spumellaria
Fauna Sinica Invertebrata Volume 15: Myxosporea
Fauna Sinica Invertebrata Volume 16: Anthozoa: Acti-
niaria, Ceriantharia and Zoanthidea
Fauna Sinica Invertebrata Volume 17: Crustacea:
Decapoda: Parathelphusidae and Pota midae
Fauna Sinica Invertebrata Volume 18: Protura
Fauna Sinica Invertebrata Volume 19: Gastropoda: Pul-
monata: Stylommatophora: Clausiliidae)
Fauna Sinica Invertebrata Volume 20: Bivalvia: Proto-
branchia Anomalodesmata
Fauna Sinica Invertebrata Volume 21: Crustacea: Mysidacea
Fauna Sinica Invertebrata Volume 22: Monogenea
Fauna Sinica Invertebrata Volume 23: Anthozoa: Sclerac-
tinia: Hermatypic coral
114
Fauna Sinica Invertebrata Volume 24: Bivalvia Veneridae
Fauna Sinica Invertebrata Volume 25: Nematoda: Rhab-
ditida: Strongylata
Fauna Sinica Invertebrata Volume 26: Phylum Granulore-
ticulosa Class Foraminiferea Agglutinated
Fauna Sinica Invertebrata Volume 27: Hydrozoa and Scy-
phomedusae
Fauna Sinica Invertebrata Volume 28: Crustacea: Amphi-
poda:Hyperiidea
Fauna Sinica Invertebrata Volume 29: Gastropoda:
Archaeogas-tropoda: Trochacea
Fauna Sinica Invertebrata Volume 30: Crustacea: Brachy-
ura: Marine Primitive Crabs
Fauna Sinica Invertebrata Volume 31 : Bivalvia: Pteriina
Fauna Sinica Invertebrata Volume 32: Polycystinea: Nasel-
laria: Phaeodarea: Phaeodaria
Fauna Sinica Invertebrata Volume 33: Annelida Polychaeta
II Nereidida
Fauna Sinica Invertebrata Volume 34: Mollusca Gastropoda
Tonnace
Fauna Sinica Invertebrata Volume 35: Arachnida: Araneae:
Tetragnathidae
Fauna Sinica Invertebrata Volume 36: Crustace Decapoda
Atyidae
Fauna Sinica Invertebrata Volume 37: Mollusca Gastropoda
Fauna Sinica Invertebrata Volume 38: Chaetognatha:
Sagittoidea
Fauna Sinica Invertebrata Volume 39: Arachnida Araneae
Gnaphosidae
Fauna Sinica Invertebrata Volume 40: Echinodermata
Ophiuroidea
Fauna Sinica Invertebrata Volume 41: Crustacea Amphipoda
Gammaridea (1)
Fauna Sinica Invertebrata Volume 44: Crustacea Decapoda
Palaemonoidea
Fauna Sinica Invertebrata Volume 46: Sipuncula Echiura
Seashells of China
Illustrated Animals of China-Mollusca (Volume 3)
Economic Mollusca of China
Bivalve Mollusca of China Seas
Pearl Science
Biology of the Economic Species of Cephalopods in the
World Oceans
Fossil Bivalves of Yunnan
Marine Molluscs and Their Collection
Rotifera of China Latin-Chinese Names of Invertebrate
Animal
THE CANADIAN FIELD-NATURALIST
Vol. 121
Invertebrates of Wuling Mountains Area, Southwestern
China
Vermiculture & vermiprotein
Polychaetous Annelids Commonly Seen From the Chinese
Waters
The Parasitic Helminth Fauna of Shaanxi
A List of Parasites for Livestock and Poultry in China
Nematology Advances and Perspective (2 volume set)
1: Nematode Morphology, Physiology and Ecology
2: Nematode Management and Utilization
Parasitic Nematodes from Equus spp.
The Studies on the cestodes in China
Monogeneans of Chinese Marine Fishes
Parasites and Parasitic Disease of Fishes
Fauna of Zhejiang-Trematoda
Offshore Radiolaria in China
Pathogenic Protozoa in Mariculture
Foraminifera and Ostracoda in Bottom Sediments of the
East China Sea
The Echinoderms of Southern China
Crabs of China Seas
Illustrated Animals of China-Class Crustacea (Volume 1)
Illustrated Animals of China-Class Crustacea (Volume 2)
Illustrated Animals of China-Class Crustacea (Volume 3)
Fauna of Zhejiang-Crustacea
Pictorial Keys To Soil Animals Of China
Atlas of shrimps in Taiwan straight
Penaeoid Shrimps of the South China Sea
Economic Shrimp and Crab of the East China Sea
Pelagic Siphonophora in China Seas
The Sichuan Farmland Spiders
Spiders in Qinghai-Tibet Plateau of China
The fauna of Hebei, China-Araneae
Economic Fauna of China: Terrestrial Mollusks
Economic Fauna of China: Freshwater Mollusks
Economic Fauna of China: Marine Mollusks
The Ferocious Summer — Adelie Penguins and the Warm-
ing of Antarctica. By M. Hooper. 2008. Greystone Books,
#201 — 2323 Quebec Street, Vancouver, British Columbia
VST 4S7. 336 pages. $29.95 Cloth.
* The Ecology and Behavior of Amphibians. By K. Wells.
2007. University of Chicago Press, 1427 E. 60" Street,
Chicago, Illinois 60637.1400 pages. US$75. Cloth.
2007
Threatened Amphibians of the World. Edited by S. N.
Stuart, M. Hoffmann, J. S. Chanson, N. A. Cox, R. J. Ber-
ridge, P. Ramani and B. E. Young. 2008. Lynx Edicions,
Montseny, 8, 08193 Bellaterra, Barcelona, Spain. 850 pages.
US$24.
The Atlas of Breeding Birds of Alberta: A Second Look.
The Federation of Alberta Naturalists fan@fanweb.ca
* Birds in Love. By Jean Levelle. 2007. Voyageur Press,
729 Prospect Avenue, PO Box 1, Osceola, Wisconsin 54020.
160 pages. $25.
Birder’s Conservation Handbook — 100 North American
Birds at Risk. By J. Wells. 2007. Princeton University Press,
41 William Street, Princeton, New Jersey, USA, 08540-5237.
464 pages. US$35. Paper.
A Climatic Atlas of European Breeding Birds. By Brian
Huntley, Rhys E. Green, Yvonne C. Collingham & Stephen
G. Willis. Hardback. Lynx Edicions, Montseny, 8, 08193
Bellaterra, Barcelona, Spain. U.S. $65 2007.
A Photographic Guide to the Birds of Japan and North-
East Asia. By Tadao Shimba. 2007. Yale University Press
P.O. Box 209040, New Haven, Connecticut 06520-9040 US
$40. Paper.
* Birds of Peru. By T. Schulenburg, D. Stotz, D. Lane, John
O’Neill and T Parker III. 2007. Princeton University Press,
41 William Street, Princeton, New Jersey, USA, 08540-5237
656 pages. US $49.50.
The Birds of Scotland. Edited by Ron Forrester and Ian
Andrews. Scottish Ornithologists’ Club (SOC), Waterston
House, Aberlady EH32 OPY, Scotland. 2 Volumes 1600
pages. GBP 75 [Excluding P&P] Cloth.
Roberts Bird Guide Southern Africa. Edited by Hugh
Chittenden. 2007 Africa Geographic, 1‘* Floor Devonshire
Court, 20 Devonshire Road, Wynberg 7800 Cape Town,
South Africa. GBP 17.99 448 pages. Cloth.
Speciation in Birds. By Trevor Price. 2007. Roberts &
Company Publishers, 4950 S. Yosemite Street, F2 #197,
Greenwood Village, Colorado 80111. U.S. $71 Paper
* Handbook of the Birds of the World. Volume 12: Pica-
thartes to Tits and Chickadees. Edited by Josep Del Hoyo,
Andrew Elliot and David A Christie. 2007. Lynx Edicions,
Montseny, 8, 08193 Bellaterra, Barcelona, Spain. 798 pages
US$294, Euros 205,00 Cloth.
+ What Bugged the Dinosaurs. By G. and R. Poinar. 2008.
Princeton University Press 41 William Street, Princeton,
New Jersey, USA 08540-5237. 312 pages. US$29.95 Cloth.
* The Return of Caribou to Ungava. By A. T. Bergerud,
Stuart N. Luttich and Lodewijk Camps. 2007. McGill-
Queen’s University Press, 3430 McTavish Street, Montreal,
Québec. Cloth.
* The Freshwater Fishes of British Columbia. By J.
McPhail. 2007. University of Alberta Press, Ring House 2 ,
University of Alberta, Edmonton, Alberta T6G 2E1 Canada.
696 pages. $90. Cloth.
How and Why Species Multiply — The Radiation of Dar-
win’s Finches. P. and R. Grant. 2008. Princeton University
Book REVIEWS
115
Press 41 William Street, Princeton, New Jersey USA 08540-
5237. 272 pages. US$35. Cloth.
Handbook of European Freshwater Fishes. By Maurice
Kottelat and J. Freyhof. 2007. Steven Simpson Books, 5
Hardingham Road, Norwich, Norfolk NR9 4LX, United
Kingdom. US$122. Cloth.
The Natural History and Management of the Gopher
Tortoise. By R. and P Ashton. Krieger Publishing, P.O. Box
9542, Melbourne, Florida USA 32902-9542. 288 pages.
US$60. Cloth.
The Wild Mammals of Wisconsin. By Charles A. Long.
Pensoft Publishers, Sofia-Moscow, Pensoft Series Faunistica
68. In English. 524 pages. EURO 45 (Paper), EURO 78
(Cloth).
+ Ecology and Management of the North American
Moose, Second Edition. By A. Franzmann and C. Schwartz.
2007. University Press of Colorado, 5589 Arapahoe Avenue,
Suite 206C, Boulder, Colorado 80303. 776 pages. US$75.
Cloth.
+ Operation Orca. By D. Francis and G. Hewlett. 2007.
Harbour Publishing, Box 219, Madeira Park, British Columbia
VON 2HO. 280 pages. $34.95. Cloth.
Owls of the United States and Canada: A Complete Guide
to Their Biology and Behavior. Wayne Lynch. 2007. The
Johns Hopkins University Press, 2715 North Charles Street,
Baltimore, Maryland 21218-4363 USA. 264 pages. US$39.95
Cloth.
A Field Guide to Seashells and Shellfish of the Pacific
Northwest. By R. Harbo. 2008. Harbour Publishing, Box
219, Madeira Park, British Columbia, VON 2HO. Laminated
pamphlet, $7.95.
*Biology of the Snapping Turtle (Chelydra serpentina).
Edited by A.C. Steyermark, M. S. Finkler, and R. J. Brooks.
2008. The Johns Hopkins University Press, 2715 North Charles
Street, Baltimore, Maryland 21218-4363. 240 pages. US$75.
Cloth.
Choosing Wildness — My Life Among the Ospreys. By C.
Arbour. 2008. Greystone Books, #201 — 2323 Quebec
Street, Vancouver, British Columbia, VST 487. 256 pages.
$24.95. Cloth.
Behavioral Ecology of Insect Parasitoids. Edited by E.
Wajnberg, C Bernstein and Jacques van Alphen. 2007.
Blackwell Publishing Inc., Commerce Place, 350 Main Street,
Malden, Massachusetts 02148, USA. 464 pages. $89.99
Cloth.
* Seashells — Jewels from the Ocean. By B. Titlow. 2007.
Voyageur Press. Galtier Plaza, Suite 200, 3800 Jackson Street,
St. Paul, Minnesota 55101. 112 pages. US$20. Cloth.
* The Smaller Majority. By P. Naskrecki. 2007. Harvard
University Press, 79 Garden Street, Cambridge, Massachusetts
02138, USA. 288 pages. US$24.95. Paper.
The Snakes of Europe. By Guido Kreiner. 2007. Chimaira.
Publishing, Groenesteinlaan 22, 9722 BX Groningen. The
Netherlands [distributed in the US by Eco Publishing] GBP
44.
116
Orangutans. By J. Payne and C. Prudente. 2008. The MIT
Press, Five Cambridge Center, 4th Floor, Cambridge,
Massachusetts 02142-1493. 160 pages. $29.95 Cloth.
* Turtles. By C. Franklin. 2007. Voyageur Press, 729 Prospect
Avenue, PO Box 1, Osceola, Wisconsin 54020. 160 pages.
US$45.
BOTANY
The Curious World of Carnivorous Plants. By Wilhelm
Barthlott, Stefan Porembski, Riidiger Seine, and Inge Theisen
[Translated by Michael Ashdown]. Timber Press, 133 SW
2™4 Avenue, Suite 450 Portland, Oregon 97204 USA. 224
pages. US$39.95 Cloth.
Figs, Dates, Laurel, and Myrrh — Plants of the Bible and
the Quran. By Lytton John Musselman. Timber Press, 133
SW 2nd Avenue, Suite 450, Portland, Oregon 97204, USA
336 pages. US$24.95. Cloth.
*Grasses of Colorado. By R. Shaw. 2008. University Press
of Colorado. 5589 Arapahoe Avenue, Suite 206C, Boulder,
Colorado 80303. 662 pages. US$75. Cloth.
Bringing Nature Home — How Native Plants Sustain
Wildlife in Our Gardens. By Douglas W. Tallamy. 2007.
Timber Press, 133 SW 2nd Avenue, Suite 450, Portland,
Oregon 97204 USA. 288 pages. US$27.95. Cloth.
Wild Plants of Eastern Canada. By M. Walker. 2008. Nim-
bus Publishing Ltd., P.O. Box 9166, Halifax, Nova Scotia
B3K 5MB8. 240 pages. $29.95 Paper.
OTHER
+ The Explorer’s Guide to Algonquin Park. By M. Runtz.
2008. The Boston Mills Press, 132 Main Street, Erin,
Ontario NOB 1TO. 222 pages. $29.95 Paper.
Antarctica. By Jonathan Scott and Angela Scott. 2007. Har-
per Collins Publishers Ltd. 2 Bloor Street East, 20" Floor,
Toronto, Ontario M4W 1A8. 256 pages. $49.00. Cloth.
* Arc of the Medicine Line: Mapping the World’s Longest
Undefended Border across the Western Plains. 2007.
Duncan and McIntyre, # 201 — 2323 Quebec Street, Vancouver,
British Columbia VST 4S7. 384 pages. $36.95 Cloth.
* Arctic Hell Ship. By W. Barr. 2007. University of Alberta
Press, Ring House 2, Edmonton, Alberta T6G 2E1. 318 pages.
$34.95 Paper.
The Big Earth Book. By James Bruges. 2007. Alistair Sawday
Publishing, The Old Farmyard Yanley Lane, Long Ashton,
Bristol BS41 9LR U.K. £19.99. Cloth.
Birds and People. Nigel J. Collar, Adrian J Long, Patricio
Robles Gil and Jamie Rojo. CEMEX-Agrupaci6én Sierra
Madre-BirdLife International, Mexico. 360 pages. GBP
$29.99. Cloth.
Go Wild with your Camera — DVD. 2007. Chris Gomer-
sall, Peter Cairns and Mark Hamblin. http://www.gowildtv.
com/index.htm. GBP 19.95.
THE CANADIAN FIELD-NATURALIST
Vol. 121
The Ornithologist’s Dictionary. By Johannes Erritzoe, Kaj
Kampp, Kevin Winker and Clifford B. Frith. 2007. Lynx
Edicions, Montseny, 8, 08193 Bellaterra, Barcelona, Spain.
290 pages. 19 Euros Paper.
Earth. By Iain Stewart and John Lynch. 2007. The Random
House Group Limited, Random House, 20 Vauxhall Bridge
Road, London SW1V 2SA. US$31 Cloth.
Culturing Wilderness in Jasper National Park — Studies
in Two Centuries of Human History in the Upper Atha-
basca River Watershed. By I. S. MacLaren, Michael Payne,
Peter J. Murphy, PearlAnn Reichwein, Lisa McDermott, C.
J. Taylor, Gabrielle Zezulka-Mailloux, Zac Robinson, and Eric
Higgs. 2007. University of Alberta Press, Ring House 2, Ed-
monton, Alberta T6G 2E1. 400 pages. $45. Paper.
Natural Sable Island. By Z. Lucas. 2008. Nimbus Publishing
Ltd., P.O. Box 9166, Halifax, Nova Scotia B3K 5M8. 128
pages. $18.95. Paper.
+ Nikkei Fishermen of the BC Coast. By M. Fukawa.
Harbour Publishing, PO Box 219, Madeira Park, British
Columbia VON 2HO. 208 pages. $ 39.95. Cloth.
Ornithology, Evolution, and Philosphy: The Life and
Science of Ernst Mayr 1904-2005. By Jiirgen Haffer. 2007.
Springer Medizin Verlag, Heidelberger Platz 3, 14197 Berlin,
Germany. 473 pages. US$119 Cloth.
Reefs Revealed. By Alex Mustard. 2007. Constable &
Robinson Ltd., 3 The Lanchester, 162 Fulham Palace Road,
London W6 9ER U.K. 192 pages. GBP 30. Cloth.
Beneath the Surface. By David Miller. 2007. Langford Press
10 New Road, Langtoft, Peterborough PE6 9LE UK. £37.99.
Cloth.
Encyclopedia of Tidepools and Rocky Shores. Edited by
Mark W. Denny and Steven D. Gaines. 2007. University of
California Press, 120 Berkeley Way, Berkeley, California
94704-1012. GBP 56. Cloth.
* Hunters at the Margin: Native People and Wildlife Con-
servation in the Northwest Territories. By John Sandlos.
UBC Press, 2029 West Mall, Vancouver British Columbia
V6T 1Z2 . 360 pages. $85 Cloth, 39.95 Paper.
Paddling and Hiking the Georgian Bay Coast. By Kas
Stone. 2008. The Boston Mills Press, 132 Main Street, Erin,
Ontario, NOB 1TO. 256 pages. $29.95 Paper.
Science and Conservation of Vernal Pools. By A. Calhoun
and P. Maynadier. CRC press LLC Headquarters 2000 NW
Corporate Boulevard, Boca Raton, Florida 33431, USA.
363 pages. US$99.95. Cloth.
How to be Wild. By Simon Barnes. 2007. Short Books, 3A
Exmouth House, Pine Street, Exmouth Market, London,
ECIR OJH. 282 pages. GBP 14.99 Cloth.
2007
BooK REVIEWS
117
News and Comment
Marine Turtle Newsletter (115)
January 2007. 32 pages: EpiroriALs: Guest Edi-
torial (P C. H. Pritchard) — Beach Driving Manage-
ment Cape Hatteras and Cape Lookout National
Seashores, USA (L. R. Nester and N. B. Frazer) —
ARTICLES: Interactions Between Marine Mammals
and Turtles (D. Fertl and G. L. Fulling) — Identifica-
tion of Bacterial Isolates from Unhatched Loggerhead
Sea Turtle Eggs in Georgia, USA (K. S. Cravem. J.
Awong-Taylor, L. Griffiths, C. Bass, and M. Musca-
rella — Nores: Captive-raised Loggerhead Turtle
Found Nesting Eight Years After Release (A. de Padua
Almeida, C. Baptistotte, A. S’A dos Santos, and T. Z.
Serafinai) — Tarballs and Early Life Stages of Sea
Turtles in Paraiba, Brazil (R. G. Santos and E. F.
Mariano) — Rapid Survey of Marine Turtles in Agalega,
Western Indian Ocean (O. Griffiths and V. Tatayah)
— Kemp’s Ridley Shell Damage (W. N. Witzell) —
Live Loggerhead Observed in Newfoundland, Canada
in Late Autumn (W. Ledwell) — IUCN-MTSG Quar-
TERLY REPORT — MEETING REPORTS — OBITUARY: Boyd
Nathaniel Lyon 1969-2006 (Peter C. H. Pritchard) —
ANNOUNCEMENTS — News & LEGAL BRIEFS — RECENT
PUBLICATIONS.
The Marine Turtle Newsletter is edited by Lisa M.
Campbell, Nicholas School of Environment and Earth
Sciences, Duke University, 135 Duke Marine Lab
Road, Beaufort, North Carolina 28516 USA; and
Matthew H. Godfrey, North Carolina Wildlife Re-
sources Commission, 1507 Ann Street, Beaufort, North
Carolina 28516 USA. Subscriptions and donations
towards the production of the MTN can be made online
at or postal mail to
Michael Coyne (Managing Editor) Marine Turtle
Newsletter, 1 Southampton Place, Durham, North
Carolina 27705 USA; e-mail: mcoyne @seaturtle.org.
118
THE CANADIAN FIELD-NATURALIST
Vol. 121
Advice for Contributors to The Canadian Field-Naturalist
Content
The Canadian Field-Naturalist is a medium for the publi-
cation of scientific papers by amateur and professional natu-
ralists or field biologists reporting observations and results
of investigations in any field of natural history provided that
they are original, significant, and relevant to Canada. All read-
ers and other potential contributors are invited to submit for
consideration their manuscripts meeting these criteria. The
journal also publishes natural history news and comment items
if judged by the Editor to be of interest to readers and sub-
scribers, and book reviews. Please correspond with the Book
Review Editor concerning suitability of manuscripts for this
section. For further information consult: A Publication Policy
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian
Field-Naturalist 97(2): 231-234. Potential contributors who
are neither members of The Ottawa Field-Naturalists’ Club
nor subscribers to The Canadian Field-Naturalist are encour-
aged to support the journal by becoming either members or
subscribers.
Manuscripts
Please submit by post to the Editor, in either English or
French, three complete manuscripts written in the journal
style. Manuscripts may also be submitted (one copy) by e-
mail. The research reported should be original. It is recom-
mended that authors ask qualified persons to appraise the
paper before it is submitted. All authors should have read and
approved it. Institutional or contract approval for the publica-
tion of the data must have been obtained by the authors. Also
authors are expected to have complied with all pertinent leg-
islation regarding the study, disturbance, or collection of ani-
mals, plants or minerals. The place where voucher specimens
have been deposited, and their catalogue numbers, should be
given. Latitude and longitude should be included for all indi-
vidual localities where collections or observations have been
made.
Manuscripts should be printed on standard-size paper, dou-
blespaced throughout, generous margins to allow for copy
marking, and each page numbered. For Articles and Notes
provide a bibliographic (citation) strip, an abstract, and a list
of key words. Generally, words should not be abbreviated but
use SI symbols for units of measure. The names of authors of
scientific names may be omitted except in taxonomic manu-
scripts or other papers involving nomenclatural problems.
“Standard” common names (with initial letters capitalized)
should be used at least once for all species of higher animals
and plants; all should also be identified by scientific name.
The names of journals in the Literature Cited should be
written out in full. Unpublished reports and web documents
should not be cited here but placed in a separate Documents
Cited section. List the captions for figures numbered in arabic
numerals and typed together on a separate page. Present the
tables each titled, numbered consecutively in arabic numerals,
and placed on a separate page. Mark in the margin of the text
the places for the figures and tables.
Check recent issues (particularly Literature Cited) for
journal format. Either “Canadian” or “American” spellings
are acceptable in English but should be consistent within one
manuscript. The Oxford English Dictionary, Webster’s
New International Dictionary and le Grand Larousse
Encyclopédique are the authorities for spelling.
Illustrations
Photographs should have a glossy finish and show sharp
contrasts. Electronic versions should be high resolution. Pre-
pare line drawings with India ink on good quality paper and
letter (don’t type) descriptive matter. Photographic reproduc-
tion of line drawings should be no larger than a standard
page. Write author’s name, title of paper, and figure number
on the lower left corner or on the back of each illustration.
Reviewing Policy
Manuscripts submitted to The Canadian Field-Naturalist
are normally sent for evaluation to an Associate Editor (who
reviews it or asks another qualified person to do so), and at
least one other reviewer, who is a specialist in the field, cho-
sen by the Editor. Authors are encouraged to suggest names
of suitable referees. Reviewers are asked to give a general
appraisal of the manuscript followed by specific comments
and constructive recommendations. Almost all manuscripts
accepted for publication have undergone revision—some-
times extensive revision and reappraisal. The Editor makes
the final decision on whether a manuscript is acceptable for
publication, and in so doing aims to maintain the scientific
quality, content, overall high standards and consistency of
style, of the joumal.
Special Charges — Please take note
Authors must share in the cost of publication by pay-
ing $90 for each page, plus $30 for each illustration (any size
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ing on size). Authors may also be charged for their changes in
proofs. Reproduction of color photos is extremely expensive;
price quotations may be obtained from the Editor.
Limited joumal funds are available to help offset publi-
cation charges to authors without grants or institutional sup-
port. Requests for financial assistance should be made to the
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Reprints
An order form for the purchase of reprints or pdf will ac-
company the galley proofs sent to the authors. Invoices for
publication costs will be sent when the submission is pub-
lished.
FRANCIS R. Cook, Editor
RR 3 North Augusta, Ontario KOG IRO Canada
TABLE OF CONTENTS (concluded) Volume 121 Number 1
Tributes
A tribute to Neal Philip Simon 1973-2006 TONY E. CHUBBS and FRANK R. PHILLIPS
Book Reviews
Zoo.oGcy: Amphibians and Reptiles of British Columbia — Blue Grouse: Their Biology and Natural History
— Birds of Peru — The Freshwater Fishes of British Columbia — Owls of the World — Ecology,
Conservation, and Status of Reptiles in Canada — Rodent Societies: An Ecological & Evolutionary
Perspective — Seashells; Jewels from the Ocean — Turtles: An Extraordinary Natural History 245 Million
Years in the Making — Whales and Seals Biology and Ecology
ENVIRONMENT: China Ecosystems — New Zealand — A Natural History
MISCELLANEOUS: Beneath My Feet: The Memoirs of George Mercer Dawson — The Wolf Totem
YOUNG NATURALISTS Gracie, The Public Gardens Duck
New TITLES
News and Comment
‘Marine Turtle Newsletter (115)
Advice to Contributors
Mailing date of the previous issue 120(4): 12 February 2008
2007
100
108
110
113
113
117
118
THE CANADIAN FIELD-NATURALIST Volume 121 Number 1 2007
Articles
Were native people keystone predators? A continuous-time analysis of wildlife
observations made by Lewis and Clark in 1804-1806 CHARLES E. KAy 1
The influence of air pollution on corticolous lichens near Strathcona
industrial area, Alberta M. ELSINGER, E. BURRELL, N. DEBRUYN,
K. TANASICHUK, and K. TIMONEY Ay
The hepatic flora and floristic affinity of hepatics around Takakia Lake,
Queen Charlotte Islands, British Columbia WON SHIC HONG 24
Comparison of scales, pectoral fin rays and opercles for age estimation of Ontario
Redhorse, Moxostoma species S. M. REID 29
A rare case of completely ambicoloured Atlantic Halibut, Hippoglossus hippoglossus,
from the lower St. Lawrence estuary, Quebec DENIS CHABOT and ROBERT MILLER 35
Micro-propagation of White-top Aster, Seriocarpus rigidus, a threatened species
from the Garry Oak ecosystem in British Columbia
BRENDA FREY, CHAIM KEMPLER, and DAviD L. EHRET 40
Pattern and potential causes of White-faced Ibis, Plegadis chihi, establishment in the
northern prairie and parkland region of North America
JILL A. SHAFFER, GREGORY A. KNUTSEN, RON E. MartIN, and JOEL S. BRICE 46
Recent distribution of the Little Brown Bat, Myotis lucifugus, in Manitoba and
northwestern Ontario JOHN E. DUBOIS and KIMBERLY M. MONSON 57
Immobilization of Elk, Cervus elaphus, with Telezol and Xylazine and reversal
with Tolazine or Yohimbine RICK ROSATTE 62
First confirmed breeding of the Marbled Godwit, Limosa fedoa. in Quebec
FRANCOIS MORNEAU, BENOIT GAGNON, and SIDNEY WHISKEYCHAN 67
Swift Fox, Vulpes velox, den use patterns in northwestern Texas
BRADY K. MCGEE, WARREN K. BALLARD, and KERRY L. NICHOLSON 71
Flowering during January in Antigonish County, Nova Scotia
DaviD J. GARBARY and BARRY R. TAYLOR 76
The occurrence of Muskoxen, Ovibos moshatus, in Labrador
Tony E. CHUBBS, and J. BRAZIL 81
The Spindle Tree, Euonymus europaea L. (Celastraceae): A newly naturalized shrub
in Nova Scotia Davib J. GARBARY and Amy E. DEVEAU 85
Notes
Serviceberry, Amelanchier intermedia, escaped from cultivation in eastern Ontario PAuL M. CATLING 89
A freshwater hydrobiid, cf. the Squat Duskysnail, Lyogyrus granum (Mollusca), widespread
and abundant in the Hampton Marsh, New Brunswick |
DONALD F. MCALPINE, RACHEL H. MAILLET, ANDREW J. ALBERT,
LAURA M. CROSSMAN, REBECCA R. SMITH, and ANDRE MARTEL 92
Northern range extension of the Pygmy Shrew, Sorex hoyi, in the Yukon |
THOMAS S. JUNG, TRoy D. PRETZLAW, and Davip W. NAGORSEN 94
(continued on inside back cover)
ISSN 0008-3550
CAN
27 The CANADIAN
FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
Volume 121, Number 2 April-June 2007
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Her Excellency The Right Honourable Michaille Jean
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage: to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields |
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members
Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope
Charles D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr.
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson
2007 Council
President: Mike Murphy Annie Belair Diane Kitching Dan Millar
Vice-President: Ken Allison Ronald E. Bedford Karen McLachlan Hamilton Stanley Rosenbaum
Recording Secretary: Susan Laurie-Bourque Fenja Brodo David Hobden Henry Steger
Treasurer: Frank Pope Julia Cipriani Diane Lepage Chris Traynor
Past President: Gary McNulty William J. Cody Ann Mackenzie Eleanor Zurbrigg
Francis R. Cook Gillian Marston
To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate
P.O. Ottawa, Canada K1Z 1A2; or e-mail: ofnc @ofne.ca.
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca.
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.
We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward
our mailing costs. PAP Registration Number 9477. Canada
Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KO G IRO; (613) 269-3211; e-mail: cfn@ofne.ca
Copy Editor: Elizabeth Morton
Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5; e-mail: r.john@rogers.com
Associate Editors:
Robert R. Anderson Charles D. Bird Paul M. Catling Anthony J. Erskine Donald F. McAlpine
Warren B. Ballard Robert R. Campbell Brian W. Coad David Nagorsen William O. Pruitt, Jr.
Chairman, Publications Committee: Ronald E. Bedford
All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail
or e-mail.
Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor.
Subscriptions and Membership
Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 pe
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) ant
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) mus!
add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Loca
Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries a
$33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be maile:
to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publication
Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: April-June 2007 (December 2007).
Cover: Four Maritime Garter Snakes, Thamnophis sirtalis pallidulus, observed in courtship behavior on the branches C’
Rhododendron shrub at 325 Prince Albert Road, Dartmouth, Nova Scotia, on 5 September 2006. See note by Gilhen an
Strum pages 210-211.
WIZ
LIBRARY
DEC 04 2008
HARVARD
PHY ER SEY
April-June 2007
The Canadian Field-Naturalist
Volume 121, Number 2
Diversity and Distribution of the Terrestrial Mammals of the Yukon
Territory: A Review
BRIAN G. SLOUGH! and THoMas S. JUNG?
'35 Cronkhite Road, Whitehorse, Yukon Territory Y! A 589 Canada; e-mail: slough@northwestel.net
Yukon Department of Environment, Box 2703, Whitehorse, Yukon Territory Y 1A 2C6 Canada; e-mail: thomas.jung @ goy.yk.ca
Slough, Brian G., and Thomas S. Jung. 2007. Diversity and distribution of the terrestrial mammals of the Yukon Territory: A
review. Canadian Field-Naturalist 121(2): 119-127.
The diversity and distribution of the terrestrial mammals of the Yukon has not been systematically reviewed since 1975, a time
when the occurrences of many species were not well known. Since then, single species and community studies, biological col-
lections and expert observations have increased our knowledge of the land mammals of the Yukon. Taxonomic studies have
resulted in some additional species. We provide an update on the diversity and distribution of recent land mammals of the
Yukon, including previously unreported range extensions, and include a revised checklist. Research in adjoining jurisdictions
has contributed more hypothetical species for the Yukon. The mammalian fauna of the Yukon is in a relatively dynamic state,
and human-induced changes, particularly climate warming, will undoubtedly influence mammalian diversity and distributions
in the coming decades.
Key Words: biological diversity, Beringia, checklist, distribution, Mammalia, Yukon.
Philip Youngman (1975) provided the most com-
prehensive survey of the recent mammals of the Yukon.
That information was based on a survey of observa-
tions and biological collections from the late 1800s
to mid-1970s, and is now over 30 years old. A recent
increase in interest in mammals, particularly non-har-
vested species, rare species, and species at risk, has
resulted in new information and corrections to previ-
ously reported distributions. The evolution of the ter-
ritorial wildlife management agency, the establishment
of a conservation data centre, several university-based
research projects (most notably near Kluane Lake; see
Krebs et al. 2000), documentation of local and tradi-
tional knowledge, and a number of resident and visit-
ing naturalists have all contributed much new infor-
mation. Changes in taxonomy and nomenclature have
also occurred since Youngman (1975), with the former
particularly advanced with the use of phylogeographic
analyses to differentiate taxa. DNA analyses have also
helped detect the presence of cryptic species (e.g.,
Myotis, Myodes, and Peromyscus spp.) that are difficult
to identify to the species level in the field. Herein, we
provide an update of the terrestrial mammals of the
Yukon Territory; an interesting zoogeographic area of
Canada with both Beringian and post-glacial immi-
grant species from southern refugia.
Methods
We reviewed information from published and un-
published work, collections, and sightings made since
Youngman (1975), primarily chronicling the period
from the mid-1970s to the present. We identified and
report on changes in species known to be present and
update their distributions since Youngman (1975) and
others (e.g., Banfield 1974; van Zyll de Jong 1983,
1985). Species without noteworthy changes are not dis-
cussed in our review; however, all documented species
occurring in the Yukon are provided in a checklist
(Table 1). Scientific names and English common names
follow Wilson and Reeder (2005) except Caribou which
replaces Reindeer. In addition, we have also extrapo-
lated ranges of species found in neighboring jurisdic-
tions to compile a list of hypothetical species. We did
not consider the marine mammals of the Beaufort Sea.
Order Soricomorhpa (Insectivores)
Seven species of soricomorphs, representing one
family (Soricidae), are recorded from the Yukon (van
Zyll de Jong 1983; Table 1). Few surveys specifically
for shrews have been undertaken in the Yukon and
because they are most efficiently captured in pitfall
traps (Prince 1941; MacLeod and Lethiecq 1963), they
are likely undersampled using standard methods for
small rodents (i.e., live-traps or snap-traps).
My)
120
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Checklist of the terrestrial mammals of Yukon. Scientific and English common names follow Wilson and Reeder
(2005).
Order / Family Scientific Name English Common Name
SORICOMORPHA
Soricidae Sorex arcticus Arctic Shrew
Sorex cinereus Cinereus Shrew
Sorex hoyi American Pygmy Shrew
Sorex monticolus Dusky Shrew
Sorex palustris American Water Shrew
Sorex tundrensis Tundra Shrew
Sorex ugyunak Barren Ground Shrew
CHIROPTERA
Vespertilionidae Eptesicus fuscus Big Brown Bat
Myotis lucifugus Little Brown Myotis
Myotis septentrionalis Northern Myotis
LAGOMORPHA
Ochotonidae Ochotona collaris Collared Pika
Leporidae Lepus americanus Snowshoe Hare
RODENTIA
Sciuridae Glaucomys sabrinus Northern Flying Squirrel
Marmota caligata Hoary Marmot
Marmota monax Woodchuck
Spermophilus parryii Arctic Ground Squirrel
Tamias minimus Least Chipmunk
Tamiasciurus hudsonicus Red Squirrel
Castoridae Castor canadensis American Beaver
Cricetidae Dicrostonyx groenlandicus Nearctic Collared Lemming
Dicrostonyx nunatakensis Ogilvie Mountains Collared Lemming
Lemmus trimucronatus Nearctic Brown Lemming
Microtus longicaudus Long-tailed Vole
Microtus miurus Singing Vole
Microtus oeconomus Root Vole
Microtus pennsylvanicus Meadow Vole
Microtus xanthognathus Taiga Vole
Myodes gapperi Southern Red-backed Vole
Myodes rutilus Northern Red-backed Vole
Neotoma cinerea Bushy-tailed Woodrat
Ondatra zibethicus Common Muskrat
Peromyscus keeni Northwestern Deermmouse
Peromyscus maniculatus North American Deermouse
Phenacomys ungava Eastern Heather Vole
Synaptomys borealis Northern Bog Lemming
Dipodidae Zapus hudsonius Meadow Jumping Mouse
Zapus princeps Western Jumping Mouse
Erethizontidae Erethizon dorsatum North American Porcupine
CARNIVORA
Canidae Canis latrans Coyote
Canis lupus Wolf
Vulpes lagopus Arctic Fox
Vulpes vulpes Red Fox
Ursidae Ursus americanus American Black Bear
Ursus arctos Grizzly Bear
Ursus maritimus Polar Bear
Mustelidae Gulo gulo Wolverine ,
Lontra canadensis North American River Otter
Martes americana American Marten
Martes pennanti Fisher
Mustela erminea Ermine
Mustela nivalis Least Weasel
Neovison vison American Mink
Felidae Lynx canadensis Canadian Lynx
Puma concolor
Cougar
2007
TABLE |. (continued)
Order / Family Scientific Name
ARTIODACTYLA
Cervidae Alces americanus
Cervus elaphus
Odocoileus hemionus
Odocoileus virginianus
Rangifer tarandus
Bison bison
Oreamnos americanus
Ovibus moschatus
Ovis dalli
Bovidae
Stewart et al. (2003) reported a specimen of Pygmy
Shrew, Sorex hoyi, from the Ogilvie Mountains which
would be a northern range extension; however the co-
ordinates given for the specimen align with Dawson
City (64°03'N, 139°25'W). Subsequently, a specimen
was retained in 2005 from the Blackstone River in
the Ogilvie Mountains (65°05'N, 138°11'W; Jung et
al. 2007), representing a northern range extension of
about 110 km.
The range of the American Water Shrew, S. palus-
tris, has been extended northward since Youngman
(1975) and van Zyll de Jong (1983) and it is now
known to as far north as the Tombstone Range of the
southern Ogilvie Mountains (Jarrell 1986; Cook et
al. 1997). It has been observed near the tree line on
Hamilton Creek, 30 km southeast of the Tombstone
Campground (64°19'N, 137°38'W; J. Schou, person-
al communication).
The Tundra Shrew, S. tundrensis, is known from the
northern Yukon south to Fortymile (64°26'N, 140°32'W
Youngman 1975), where three additional specimens
were obtained in 2003 (B. Slough and T. Jung, unpub-
lished data). In the late 1970s five Tundra Shrews
were trapped in alpine and subalpine habitats on the
Haines Road in British Columbia within 20 km of the
southwest Yukon, and about 500 km south of its pre-
viously known limits (Nagorsen and Jones 1981). It was
thought to be limited to Beringia, a region unglaciat-
ed during the Wisconsin glaciation; however, the
British Columbia locations are 500 km from the lim-
its of Beringia and confound the expected distribution
pattern (Nagorsen and Jones 1981). It may yet be
found in the intervening area in the Yukon.
The Arctic Shrew, S. arcticus, remains known only
from a single skull collected in 1965 (Youngman 1975),
despite substantial subsequent survey effort for shrews
(e.g., T. Jung and colleagues, unpublished data; J.
Cook and colleagues, unpublished data). Further col-
lections are required to assess its distribution in the
Yukon.
Order Chiroptera (Bats)
Three species of bats, representing one family (Ves-
pertilionidae), are reported from the Yukon (Table 1).
SLOUGH AND JUNG: TERRESTRIAL MAMMALS OI
rHE YUKON 12]
English Common Name
Moose
Elk
Mule Deer
White-tailed Deer
Caribou
American Bison
Mountain Goat
Muskox
Dall’s Sheep
Youngman (1975) reported only the Litthe Brown
Myotis, Myotis lucifugus, from the Yukon, where it is
widespread and common south of 64°N. Since Young-
man (1975), the Northern Myotis, M. septentrionalis,
has been confirmed (Jung et al. 2006a) and another
species, either the Big Brown Bat (Eptesicus fuscus)
or Silver-haired Bat (Lasionycteris noctivagans), has
been recorded via acoustic sampling (B. Slough, un-
published data). Until recently, few mist-net or echolo-
cation monitoring surveys for bats had been under-
taken in the Yukon, and it is likely that the Order
remains undersampled. Based on surveys in adjacent
jurisdictions, there may be other species of bats pres-
ent in the Yukon that are not yet documented.
The Northern Myotis, M. septentrionalis, was first
found in the Yukon in 2004 when three individuals
were captured at the La Biche River (60°08'N, 124°04'W;
Jung et al. 2006a). Five additional specimens were cap-
tured in 2007 near Watson Lake (60°07'N, 128°49'W;
Lausen et al. 2008). This species has also been cap-
tured at several locations in the Liard watershed of
northern British Columbia (Wilkinson et al. 1995*;
Bradbury et al. 1997*; Crampton et al. 1997*; Vonhof
et al. 1997*; Vonhof and Wilkinson 1999*) and in the
southern Northwest Territories as far west as Nahanni
National Park (van Zyll de Jong 1985; Lausen 2006*).
Youngman (1975) had listed the Big Brown Bat,
E. fuscus, as a hypothetical species for the Yukon based
on a record from interior Alaska (Reeder 1965). In
September 1999, B. Slough (unpublished data) made
a recording of a bat echolocation call sequence with
an AnaBat II bat detector (Titley Electronics, Ballina,
New South Wales, Australia) at Morris Lake (60°25'N,
131°38'W) that was most likely E. fuscus (C. Corben,
personal communication; Keinath 2004*). This is pos-
sibly the first record of the species from the Yukon,
although it may be difficult to distinguish the echolo-
cation calls of E. fuscus from those of Silver-haired
Bats, as recorded with an AnaBat detector (Betts 1998).
There is a single record from central Alaska (Reeder
1965) and unsubstantiated reports exist for southeast
Alaska (MacDonald and Cook 1996). E. fuscus had
been recorded on bat detectors in the Liard watershed
in northern British Columbia (Wilkinson et al. 1995*:
122
Bradbury et al. 1997*; Crampton et al. 1997*) and
southwestern Northwest Territories (Lausen 2006*).
Order Lagomorpha (Pikas and Hares)
As noted by Youngman (1975), two species of lago-
morphs, representing two families (Ochotonidae and
Leporidae) are recorded from the Yukon (Table 1).
The southeastern limit for the Collared Pika, Ocho-
tona collaris, shown by Youngman (1975) was the
Logan Mountains. It has since been sighted approxi-
mately 200 km east-southeast in the Kotaneelee (2004;
60°27'N, 124°12'W) and La Biche (1998; 60°14'N,
124°32'W) ranges (B. Bennett, personal communica-
tion).
Order Rodentia (Rodents)
There are 26 known species of rodents, represent-
ing five families (Sciuridae, 6 species; Castoridae, |
species; Cricetidae, 16 species; Dipodidae, 2 species;
Erithizontidae, 1 species), recorded from the Yukon
(Table 1).
The Northern Flying Squirrel, Glaucomys sabrinus,
occurs throughout the forests of the southern Yukon
(Youngman 1975). It has been observed near the tree-
line on Hamilton Creek in the Ogilvie Mountains
(64°20'N, 137°39'W; J. Schou, personal communica-
tion), confirming the northern range limit projected
by Youngman (1975).
The Hoary Marmot, Marmota caligata, has been
confirmed in the La Biche Range in the extreme south-
east Yukon (60°14'N, 124°32'W; B. Bennett, personal
communication).
The Woodchuck, Marmota monax, apparently has
a broad range through the southern Yukon; however,
sightings and specimens indicate a geographic sepa-
ration of at least 370 km between the southern (Ross
River; 61°59'N, 132°27'W) and central (Klondike;
63°39'N, 138°40'W) populations which Youngman
(1975) shows as contiguous.
The Least Chipmunk, Neotamias minimus, was re-
ported by Youngman (1975) at Bonnet Plume Lake. It
has since been sighted approximately 100 km north-
west on the Bonnet Plume River (seven locations from
Bonnet Plume Lake to Fairchild Creek (64°58'N,
133°46'W; B. Slough, unpublished data) and approx-
imately 80 km northwest on the Wind River (65°03'N,
134°54'W; S. Gilbert, personal communication). The
species appears well distributed in the Wernecke Moun-
tains.
The distribution of the Ogilvie Mountains Lemming,
Dicrostonyx nunatakensis, is poorly understood, since
it is known from only 13 specimens collected at Angel-
comb Peak in the Ogilvie Mountains (64°36'N,
138°16'W; Youngman 1964, 1967, 1975), two of which
were recorded recently (T. Jung and B. Slough, un-
published data).
Krebs and Wingate (1976, 1985) captured Nearctic
Brown Lemmings, Lemmus trimucronatus, near the
Alaska Highway south of Kluane Lake and from the
THE CANADIAN FIELD-NATURALIST
Vol. 121
Chilkat Pass, near the Haines Road, in British Colum-
bia, extending its known range about 170 km southwest
of that reported by Youngman (1975).
The Singing Vole, Microtus miurus, is known at
present only from the British Mountains, the southern
Ogilvie Mountains, and from the Kluane region of the
southwestern Yukon (Youngman 1975; Krebs and Win-
gate 1976, 1985). Krebs and Wingate (1976, 1985)
trapped extensively in the Kluane region and found
the southern limit of distribution to be the Slims River
area at the south end of Kluane Lake (61°N, 138°31'W).
Nagorsen (2002) reported that it has not been found
south to the British Columbia border as suggested by
Banfield (1974) and Youngman (1975).
Jung et al. (2006b) reported the Southern Red-
backed Vole, Myodes gapperi, from the Yukon at the
La Biche River (60°08'N, 124°04'W). Youngman (1975)
did not report M. gapperi from the Yukon; however, he
considered M. gapperi and the Northern Red-backed
Vole (M. rutilus) to be conspecific. Based on the dis-
tribution of M. gapperi in adjacent northeastern British
Columbia and southwestern Northwest Territories
(Banfield 1974; Nagorsen 2005) we suspect that M.
gapperi may be more widespread in the Liard River
watershed.
Youngman (1975) gave the northwestern limit of
the Bushy-tailed Woodrat, Neotoma cinerea, as Klu-
ane Lake. It has since been been observed on Ogilvie
Island, Yukon River, near Youngman’s projected north-
ern limit (63°34'N, 139°44'W; B. Bennett, personal
communication). In 2003, a nest of this species was
observed at Bear Creek (64°02'N, 139°15'W; B.
Slough, unpublished data), representing a 160 km
northward range extension.
Muskrat, Ondatra zibethicus, inhabit wetlands
across the territory (Youngman 1975). The northern-
most observations were made in the Arctic wetlands of
the Mackenzie Delta in the Yukon (68°46'N, 136°32'W;
B. Slough, unpublished data).
The Northwestern Deermouse, Peromyscus keeni,
had been confirmed through DNA analysis as being
present in the Coast Mountains and adjacent areas of
the southwestern Yukon (Wike 1998; Lucid and Cook
2004; M. Lucid, personal communication) within the
range of the former P. maniculatus algidus.
In July 2005, a North American Deermouse, P. man-
iculatus, was captured in the Ogilvie Mountains
(64°30'N, 138°13'W; T. Jung et al. unpublished data);
representing a 70 km northward range extension, but
within the range projected by Youngman (1975).
Interestingly, Lucid and Cook (2007) provide DNA
evidence of an undescribed species of Peromyscus in
southwestern Yukon. Wike (1998), working in central
Yukon, also reported the possibility of an undescribed
species of Peromyscus in the Yukon.
Banfield (1974) and Youngman (1975) show the
Northern Bog Lemming, Synaptomys borealis, ranging
throughout the Yukon north to the Porcupine River
drainage. Smits et al. (1989) reported S. borealis teeth
2007
and bone fragments in the scat of an Arctic Fox, Vulpes
lagopus, collected in 1985, on Herschel Island, extend-
ing the range of S. borealis about 150 km north to the
Beaufort Sea (69°35'N, 139°05S'W).
The Western Jumping Mouse, Zapus princeps, is
known from only two specimens collected in 1944
(Rand 1945; Youngman 1975) and another collected
in 1980 (S. Beare, unpublished data); all three speci-
mens are from the South Canol Road, near the Rose
and Nisutlin rivers.
Order Carnivora (Carnivores)
Sixteen species of carnivores, representing four fam-
ilies (Canidae, 4 species; Ursidae, 3 species; Mustel-
idae, 7 species; Felidae, 2 species), are recorded from
the Yukon (Table 1).
Coyotes, Canis latrans, expanded northward from
55°N to southern Alaska between 1829 and 1907
(Banfield 1974). The earliest reported sighting for the
area is from “near Whitehorse, on the Alsek River
Alaska, in February 1907” (J. A. Allen 1908, report-
ed in Hall 1981). Cairnes (1909) reports the presence
of Coyotes in the southern Yukon in 1908. The range
has been represented as the entire Yukon (Banfield
1974; Youngman 1975); however, trapper and hunter
returns and observations suggest that Coyotes are
absent or scarce north of 65°N (Yukon Department of
Environment, unpublished data). Extraordinary obser-
vations, such as a sighting at Old Crow (67°N, 135°W,
C. P. Charlie reported in Youngman 1975) should be
considered as extra-limital and not evidence of range
expansion.
The Arctic Fox, occurs on the Yukon Coastal Plain
and Herschel Island, roughly northward of the 150 m
ASL contour (Smits and Slough 1993). There is no
evidence of denning or other range use further inland,
therefore extraordinary specimens from the Old Crow
Flats and the Porcupine River (reported in Youngman
1975) should be considered as extra-limital and not
evidence of range expansion. Similarly, extraordinary
observations of Polar Bear, Ursus maritimus, near Old
Crow (reported in Youngman 1975) should be consid-
ered as extra-limital and not evidence of range expan-
sion. The Yukon range of Polar Bears is also restricted
to the North Coastal Plain and Herschel Island.
An analysis of fur harvest data from registered trap-
lines to 2006 shows that the Fisher, Martes pennanti,
is largely confined to the Liard River watershed in
the southeast Yukon, ranging west to the Teslin River
(60°28'N, 133°18'W) as projected by Youngman (1975).
Youngman’s northern limit was approximately 61°N;
however, the species ranges 80 km north to 61°40'N
(upper Hyland River and Finlayson Lake; B. Slough
and T. Jung, unpublished data). Several observations
have been made west of the Teslin River, and Fishers
have been trapped north of Whitehorse (Klusha Creek;
61°13'N, 135°58'W) and near Haines Junction (60°51'N,
137°36'W). We believe that the species is expanding
its range northwards and westwards.
SLOUGH AND JUNG: TERRESTRIAL MAMMALS OF THE YUKON 123
The Least Weasel, Mustela nivalis, occurs through-
out the Yukon (Banfield 1974) and, contrary to Young
man (1975), it is present in the Liard watershed. In-
dividuals have been captured by trappers in the Liard
watershed (B. Slough, unpublished data; Yukon Depart-
ment of Environment, unpublished data).
American Mink, Mustela vison, occurs throughout
the Yukon, as far north as the Old Crow Flats (Young-
man 1975) where it has been captured by trappers
(Yukon Department of Environment, unpublished data)
but does not appear to be a regular inhabitant north of
the treeline (68°25'N; Bee and Hall 1956; Banfield
1974) as suggested by Youngman (1975).
Similarly, the North American River Otter, Lontra
canadensis, occurs throughout the Yukon, as far north
as the British and Richardson mountains (Youngman
1975), but does not likely inhabit the Arctic Coastal
Plain (north of the 150 m ASL contour, Bee and Hall
1956; Banfield 1974) as suggested by Youngman
(1975). We suggest that an observation from Herschel
Island (Wildlife Management Advisory Council (North
Slope) and the Aklavik Hunters and Trappers Commit-
tee 2003*) was an extra-limital movement. A similar
occurrence was reported for the Beaufort Sea of north-
eastern Alaska by Haskell (2006). Additionally, we
suggest that the Arctic range for Canadian Lynx, Lynx
canadensis, shown by Youngman (1975) is extra-limital,
and occurs only following years during a population
crash.
The range of Cougar, Puma concolor, in the Yukon
remains somewhat obscure, given the species’ propen-
sity for long-distance extra-limital movements. Spo-
radic sightings have occurred throughout the southern
half of the territory. Although most sightings are unsub-
stantiated, they occasionally occur in concurrent geo-
graphic clusters and are often associated with the
occurrence of Mule Deer, Odocoileus hemionus, (Yukon
Department of Environment, unpublished data), pro-
viding some credibility to the sighting data. The first
Yukon specimen was found dead at Watson Lake
(60°04'N, 128°43'W) in November 2000 (Jung and
Merchant 2005). Successful breeding by P. concolor
in the Yukon has not been documented.
Order Artiodactyla (Even-toed Ungulates)
Ungulates have long been the management priority
in the Yukon, and consequently the best known Order.
Nine species of ungulates, representing two families
(Cervidae, 5 species; Bovidae, 4 species), are recorded
from the Yukon (Table 1). Since Youngman’s (1975)
review, White-tailed Deer, Odocoileus virginianus,
have been recorded in the Yukon (Hoefs 2001).
Since Youngman (1975), introduced populations of
Elk, Cervus elaphus, north and west of Whitehorse
have grown substantially and now may number over
200 animals, although no reliable estimates are avail-
able. In addition to introduced populations, Elk have
been reported regularly since 1977 in southeastern
Yukon, primarily in the Hyland, Beaver, Coal, La-
124
Biche, and Liard river valleys (Hoefs et al. unpub-
lished data*). The most recent reported location in
southeast Yukon is from October 2004 at Beaver River,
where the resident trapper has seen Elk with some reg-
ularity over the past several years (60°'N, 124°25'W;
D. MacDonald, personal communication). Whether
these animals in the southeast represent populations
expanding their range northward from British Colum-
bia (Shackleton 1999), or are extra-limital movements,
is unknown.
Mule Deer, Odocoileus hemionus, appear to have
extended their range in the Yukon since the review by
Youngman (1975). They are now found throughout the
Yukon, south of the Ogilvie Mountains (64°05'W) and
including westward to Alaska (Hoefs 2001). Hoefs
(2001) reported a 1998 observation north of the Ogilvie
Mountains at Chapman Lake (64°50'N, 138°25'W).
We regard this observation, however, as an extra-limital
movement and not evidence of Mule Deer being dis-
tributed north of the Ogilvie Mountains.
Reliable observations of White-tailed Deer, Odo-
coileus virginianus, some including photographs, have
been made in the Yukon (Hoefs 2001). The first obser-
vation was reported in 1975 near Tagish Lake (60°10'N,
134°20'W, Hoefs 2001). Mayo (63°05'N, 135°53'W)
appears to be the most northern location that White-
tailed Deer have been observed in the Yukon (M.
O’ Donoghue, personal communication). Hoefs (2001)
reported an antlered skull found near Ross River (62°N,
132°25'W) as further evidence of White-tailed Deer
in the Yukon.
American Bison (Bison bison) (Wood Bison sub-
species, B. b. athabascae) have been reintroduced to
the Yukon since the review by Youngman (1975). An
attempt in 1951 to establish a small herd of Plains
Bison, B. b. bison, in the Yukon, by releasing five
animals from Alaska, had failed (Youngman 1975).
Three Wood Bison herds may now be found in the
Yukon; all are reintroduced. The Aishihik Herd ranges
in southwestern Yukon, near Aishihik Lake (61°36'N,
137°30'W). Two other, smaller herds — the Nahanni
and Nordquist herds — mainly range in the Northwest
Territories and British Columbia; Shackleton 1999),
respectively, but a portion of their annual range is in
southeastern Yukon. The Nahanni Herd ranges into
the LaBiche River valley (60°07'N, 124°03'W); the
Nordquist Herd ranges along the Alaska Highway,
north of Liard Hot Springs in British Columbia, and
into the Yukon at Contact Creek (60°01'N, 127°37'W).
Both of the southeastern herds were also reintroduced
to their native range in the 1990s (Gates et al. 2000).
Muskox, Ovibos moschatus, were reintroduced to
Barter Island (70°08'N, 143°38'W) and the adjacent
mainland in northern Alaska in 1969 and 1970. Muskox
were sighted almost yearly in the Yukon since the rein-
troduction but they did not appear to establish a year-
round breeding population at the time of Youngman
(1975). The first mixed sex groups including calves
were reported in the Yukon in 1986 (D. Cooley, per-
THE CANADIAN FIELD-NATURALIST
Vol. 121
sonal communication). They currently range as far east
as the Blow River (ca. 137°W; D. Cooley, personal
communication). Summer movements to the Old Crow
Flats (ca. 68°N, 140°W) have been somewhat com-
mon since 1995. A new resident breeding population
appears to have formed in the Richardson Mountains
(ca. 68°N, 137°W) that remains all year in the moun-
tains (D. Cooley, personal communication).
Hypothetical Species
The following species are found in neighbouring
Alaska, British Columbia or the Northwest Territories,
and their ranges may extend into the Yukon (Table 2).
Some taxa, such as shrews and bats, are undersampled,
as are some of the more remote geographic areas.
The Alaskan Tiny Shrew, Sorex yukonicus, likely
occurs in the Yukon (Table 2), but has yet to be col-
lected outside Alaska. This species has been recently
described from seven specimens from Alaska in 1993
(Dokuchaev 1997). Subsequently, other specimens
(ca. 30) have been obtained from Alaska as a result
of directed pitfall trapping efforts (Peirce and Peirce
2000; MacDonald and Cook 2001*; Cook and Mac-
Donald 2003*), indicating that the species is wide-
spread, but uncommon, in Alaska. Specimens were
obtained from Yukon-Charley Rivers National Pre-
serve and Wrangell-St. Elias National Park and Pre-
serve only 50 km and 12 km, respectively, from the
western Yukon border (MacDonald and Cook 2001*:;
Cook and MacDonald 2003*). Directed sampling
effort in 2003 near the abandoned village of Fortymile
(64°25'N, 140°32'W) failed to secure a specimen (B.
Slough and J. Jung, unpublished data).
Youngman (1975) noted the Long-legged Bat,
Myotis volans, as a hypothetical species for the Yukon,
based on a record near Atlin, British Columbia, re-
ported by Swarth (1936). The species was recently
captured in the southwestern Northwest Territories
(Lausen 2006*). In addition to M. volans, we add the
Western Long-eared Bat, M. evotis, and Silver-haired
Bat, Lasionycteris noctivagans, to the list of hypotheti-
cal bat species in the Yukon. All three of these hypo-
thetical species have recently been captured within
150 km of the Yukon border in northeastern British
Columbia (see: Wilkinson et al. 1995*; Bradbury et al.
1997*; Crampton et al. 1997*; Vonhof et al. 1997*;
Vonhof and Wilkinson 1999*; Lausen 2006*) and occur
in southeast Alaska (Parker et al. 1997). These captures,
however, have not been substantiated with a specimen,
echolocation recording or a genetic sample (D. Nagors-
en, personal communication). Acoustic records also
exist for the Hoary Bat, Lasiurus cinereus, and the
Eastern Red Bat, L. borealis, from the southwestern
Northwest Territories (Lausen 2006*), suggestive that
these species may also occur in southeastern Yukon.
Parker and Cook (1996) reported Keen’s Myotis,
Myotis keenii, from southeastern Alaska (north to ca.
58°N), and it may yet be reported from in or near Klu-
ane National Park Reserve in southwestern Yukon.
2007
SLOUGH AND JUNG: TERRESTRIAL MAMMALS OF THE YUKON
125
TABLE 2. Hypothetical terrestrial mammals that may occur in the Yukon.
Order / Family Scientific Name
SORICOMORPHA
Soricidae Sorex yukonicus
CHIROPTERA
Vespertilionidae Lasionycteris noctivagans
Lasiurus borealis
Lasiurus cinereus
Myotis evotis
Myotis keenti
Myotis volans
LAGOMORPHA
Leporidae Lepus arcticus
Lepus othus
RODENTIA
Sciuridae Marmota broweri
Muridae Mus musculus
CARNIVORA
Memphitidae Mephitis mephitis
In addition to the resident Snowshoe Hare, Lepus
americanus, Youngman noted the possibility of Arc-
tic Hare, L. arcticus, or Alaskan Hare, L. othus, being
found in northern Yukon. There is an unsubstantiated
sight record of L. arcticus from the eastern Richard-
son Mountains of the Yukon in the 1970s (P. Frank-
ish, personal communication).
The Alaska Marmot, Marmota broweri, may occur
in the Yukon, but it has not yet been recorded from
there. M. broweri is found as far east in Alaska as the
Brooks Range in the Arctic National Wildlife Refuge
(Bee and Hall 1956); it may also range in the British
Mountains in Ivvavik National Park, Yukon. The Wild-
life Management Advisory Council (North Slope) and
the Aklavik Hunters and Trappers Committee (2003*)
reports “two people said they remembered seeing (mar-
mots) in 1960s and 1970s in the Babbage and Firth
River areas”.
The House Mouse, Mus musculus, occurred in the
vicinities of Dawson and Whitehorse in the 1970s
(Youngman 1975); however, no further specimens have
been obtained. This species may be periodically intro-
duced, but does not appear capable of sustaining com-
mensal populations in the Yukon.
Banfield (1974) shows the Striped Skunk, Mephitis
mephitis, entering the Yukon in the extreme southeast
(La Biche River region) and coming very close in the
Teslin area. It has not been documented from the
Yukon with the nearest record being two skins taken
at the junction of the Fort Nelson and Liard rivers in
British Columbia (Rand 1944; Hatler et al. 2008),
about 52 km SE of the Yukon. In the Northwest Terri-
tories, it is present at Fort Smith, and is expected at
Fort Liard (S. Carriére, personal communication), the
latter being only 30 km from the Yukon border.
English Common Name
Alaskan Tiny Shrew
Silver-haired Bat
Eastern Red Bat
Hoary Bat
Long-eared Myotis
Keen's Myotis
Long-legged Myotis
Arctic Hare
Alaskan Hare
Alaska Marmot
House Mouse
Striped Skunk
Conclusions
Knowledge of the diversity and distribution of the
terrestrial mammals of the Yukon has improved steadi-
ly since the last review was published in 1975 (Young-
man 1975). Excluding Humans, there are now 63 spe-
cies of terrestrial mammals that are believed to occur
in the Yukon, representing 40 genera, 15 families and
6 orders (Table 1). There are an additional 11 hypo-
thetical species (Table 2). All species, except Mus mus-
culus, are native. Only one species, Dicrostonyx nuna-
takensis, is endemic. Populations of some species native
to the Yukon have been introduced (Cervus elaphus)
or reintroduced (Martes americana, Oreamnos amer-
icanus, Bison bison).
The mammalian fauna of the Yukon is in a relatively
dynamic state. Several species have colonized the Yukon
in the past 125 years (e.g., Canis latrans, Odocoileus
hemionus, and Puma concolor). The history of mam-
mal colonization and evolution in Yukon and neigh-
bouring Alaska have been discussed by Youngman
(1975) and MacDonald and Cook (1996). Approxi-
mately one-third of the recent terrestrial mammals of
the Yukon originated in Beringia; the remainder are
southern immigrants (Youngman 1975). Many of the
southern immigrants have limited distributions in the
Yukon today. For example, the newly recorded bat
species are likely southern immigrants, and Chiropter-
ans may still be colonizing northwestern North Amer-
ica (Jung et al. 2006a). Myodes gapperi and Peromyscus
keeni are similarly southern immigrants. Further cryp-
tic species may be present in the Yukon, for example,
in the genus Peromyscus (Lucid and Cook 2007) and
Myotis (Jung et al. 2006a). Human-induced changes,
particularly climate change, will undoubtedly affect
mammalian diversity and distributions, particularly in
126
northern regions (e.g., Humphries et al. 2002, 2004).
Ranges of some species in the Yukon will likely grow
with anthropogenic habitat changes and climate warm-
ing (e.g., Mule Deer), while others may contract (e.g.,
Caribou). Detailed mapping of distribution in the Yukon
can assist in monitoring any change in distributional
patterns.
Appropriate sampling techniques for species typi-
cally undersampled, such as echolocation monitoring
and mist-netting for bats and pitfall trapping of shrews,
has rapidly increased our knowledge of these secre-
tive and cryptic species in the Yukon. Sampling in
remote areas not well surveyed for mammals will also
likely yield interesting records. Further mammal sur-
veys, particularly for the smaller species (<1 kg) and in
geographic areas with inadequate sampling, will con-
tinue to improve our understanding of Yukon mam-
mals. Such knowledge is imperative for the conserva-
tion of biodiversity and predicting the impact of climate
change and landscape change on the diversity, distri-
bution and conservation status of Yukon’s mammals.
Acknowledgments
We thank the numerous individuals and agencies
that contributed species-specific information, includ-
ing collections and observations, or funding, that sub-
stantially improved our knowledge of the diversity,
distribution and status of terrestrial mammals of the
Yukon. Financial support was provided by the Yukon
Department of Environment and the Northern Re-
search Institute, Yukon College. David Nagorsen and
an anonymous referee reviewed an earlier version of
the manuscript.
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Received 21 March 2006
Accepted 9 May 2008
State Records and Habitat of Clam Shrimp, Caenestheriella gynecia
(Crustacea: Conchostraca), in New York and New Jersey
ROBERT E. ScHMipT! and ERIK KIVIAT
Hudsonia Limited, P.O. Box 5000, Annandale, New York 12504 USA
'Present address and correspondence to: Bard College at Simon’s Rock, 84 Alford Road, Great Barrington, Massachusetts
01230 USA; e-mail: schmidt @simons-rock.edu
Schmidt, Robert E., and Erik Kiviat. 2007. State records and habitat of Clam Shrimp, Caenestheriella gynecia (Crustacea:
Conchostraca), in New York and New Jersey. Canadian Field-Naturalist 121(2): 128-132.
We report new records of the Clam Shrimp, Caenestheriella gynecia Mattox 1950, from three localities in the Hudson Valley
of New York and one locality in northeastern New Jersey. All of our specimens were collected in puddles on dirt roads. The
New Jersey specimens have meristics that are well within the range of Caenestheriella Daday 1915; however, the eastern
New York specimens resemble those reported from Massachusetts with meristics that are closer to Cyzicus Audouin 1837.
We hypothesize that C. gynecia was established as a parthenogenetic species due to an unlikely dispersal event in the west-
ern part of its range and has migrated eastward since the last glacial maximum. Dispersal of this species may occur by wind,
in the gut of birds, or stuck to animal feet or fur, or to vehicles. We recommend that C. gynecia be treated as rare and vulner-
able to extinction throughout its range unless demonstrated otherwise.
Key Words: Clam Shrimp, Caenestheriella gynecia, temporary pond, biogeography, dirt road, New York, New Jersey.
Clam Shrimps are small freshwater bivalvular arthro-
pods in the order Conchostraca. Clam Shrimps inhabit
isolated ephemeral pools, temporary ponds, and some-
times lakes; the taxon is found nearly worldwide (Wil-
liams 1987). Caenestheriella gynecia Mattox 1950 is
a parthenogenetic species with no records of males,
although males can be uncommon in related species
and may be discovered (see Eder et al. 2000). Until
Smith and Gola (2001) reported two populations of this
species from western Massachusetts, the known range
extended east from Ohio (Smith 2001) to northeastern
Pennsylvania. Here we report four localities of C.
gynecia in the eastern United States, representing range
extensions to New York and New Jersey.
Methods
Specimens were captured from rain puddles and pre-
served in 50% isopropanol. Preserved specimens were
measured with digital calipers (greatest length in mm).
Spines on the telson and second antennal segments were
counted at 320x with a stereomicroscope.
Results
Here we document four collections of Caenestheriel-
la gynecia, representing the first records of this species
from New Jersey and New York.
New JERSEY: rain puddles on unimproved Paterson Lateral
Gas Pipeline road between Empire Avenue and the New Jersey
Turnpike, Borough of Carlstadt and Township of South Hack-
ensack, Bergen County, 74°02'26"W longitude, 40°49'37"N
latitude, Hackensack Meadowlands. [Weehawken, New Jersey—
New York. 1967, U.S. Geological Survey (USGS) 7.5 minute
topographic map sheet], E. Kiviat, 2 September 2001. 6 spec-
imens (Kiviat and MacDonald 2004).
New York: rain puddle on abandoned farm road, 3.5 km south
of Rhinebeck, Dutchess County, 73°55’15”W longitude,
41°53’ 19”N latitude, Town of Rhinebeck, [Kingston East, New
York. 1963, photorevised 1980 USGS 7.5 minute topograph-
ic map sheet], S. Nyman, 27 May 1994. 1 specimen, 7.6 mm,
dried (Site C-3 in Kiviat et al. 1994*).
NEw York: rain puddles on dirt road, ca. 1.4 km NNW of East
Park, Dutchess County, 73°55'20"W longitude, 41°48'19"N
latitude, Town of Hyde Park, [Hyde Park, New York. 1963,
USGS 7.5 minute topographic map sheet] E. Kiviat & J.
Bridges, 26 June 2002. 5 specimens.
NEw YorRK: rain puddles on ATV trail, ca. 3.4 km NNE of
Saugerties, Greene County, 73°55'58"W longitude, 42°06'33"N
latitude, Bristol Beach State Park [Saugerties, New York. 1963,
USGS 7.5 minute topographic map sheet], E. Kiviat, 25 July
2007. 1 specimen.
Clam Shrimp habitats (Figure 1) at the New York
and New Jersey localities were all rain puddles on dirt
roads. Clam Shrimp were observed swimming slowly
on the underside of the surface film in New Jersey, and
swimming in the water column and along the bottom
in New York. The puddles were ca. 1.5-30 m? and ca.
5-15 cm deep at the dates of collection. The rain puddles
essentially lacked vascular plants. We have visited the
puddles at the New Jersey locality several times at dif-
ferent seasons 2002-2007; some of the puddles main-
tained their depths even during dry periods, but did dry
during the summer 2005 drought. The New Jersey pud-
dles were consistently very turbid and Clam Shrimp
were not visible below the surface; the Hyde Park pools
were clear enough to see Clam Shrimp swimming along
the bottom. The roads at the New Jersey and Saugerties,
New York localities are regularly used by all-terrain
128
2007
ts Ee, ans - Sas
SCHMIDT AND KIVIAT: CLAM SHRIMP IN NEW YORK AND NEW JERSEY
129
FiGurE |. Habitat of Caenestheriella gynecia in New Jersey. Photograph by Erik Kiviat.
vehicles (including “‘four-wheelers” and motorcycles),
and road vehicles (pickup trucks and sport-utility vehi-
cles) were observed using the road at the Hyde Park
locality. The puddles at the Bristol Beach State Park
site were created and maintained by all-terrain vehicle
use. We believe the puddle habitats at these four sites
were created and maintained by wheeled vehicles; the
puddles at the Rhinebeck locality appeared less recent-
ly disturbed and may have been created by farm vehi-
cles. The dirt road at the Hyde Park locality was bor-
dered by hardwood forest and woodland pools. The dirt
road at the Rhinebeck locality was fringed by tall shrubs
and small trees in the midst of a wet meadow and old
field. In New Jersey, the dirt road was fringed by Com-
mon Reed (Phragmites australis), tall shrubs, and
small trees, and was raised ca. 1.5 m above the level of
extensive, formerly tidal, marsh now dominated by
common reed. The ATV trail at the Bristol Beach site
was bordered by second-growth hardwood forest.
The Hyde Park and Rhinebeck roads appear to com-
prise in situ soil materials which are mapped as Nas-
sau-Cardigan complex (Lithic and Typic Dystrochrepts)
and Sun silt loam on glacial till, and Hudson-Vergennes
soil (Glossaquic Hapludalf) on glaciolacustrine silty
clay, respectively (Faber 2002*). The Saugerties site
has soils mapped as Hudson silt loam, a Glassaquic
Hapludalf and Madalin silty clay loam, a Mollic Ochra-
qualf according to Tormes (1979*).
The pH of these soils varies from strongly or very
strongly acidic at the surface to moderately akaline in
the substratum (Faber 2002*). Wetlands adjoining the
New Jersey site were mapped as Sulfhemists and Sul-
faquents (Goodman 1995*); however, the dirt road ap-
pears to have been constructed from imported materi-
al with a light reddish color that may have originated
from nearby red Triassic shale and sandstone (Schu-
berth 1968). We have not found Clam Shrimp in wet-
lands adjoining the dirt road habitats.
Smith and Gola (2001) discussed generic placement
of this species and showed that the meristics on their
Massachusetts specimens were not typical of Caenes-
theriella but were intermediate or closer to species of
Cyzicus Andouin 1837. The most significant difference
in the genera is the shape of the male rostrum which
cannot be determined in our specimens. There are dif-
ferences in the number of spines on the telson, but the
counts on the Massachusetts specimens overlapped the
genera and Smith and Gola (2001) questioned whether
the two genera were distinct. The size and meristics of
our New Jersey specimens (Table 1) fall clearly within
Caenestheriella Daday 1915 (all specimens contained
eggs) as did the single specimen from Saugerties (west
of the Hudson River). Our New York specimens from
east of the Hudson River, however, are similar to those
in western Massachusetts. It is possible that the eastern
New York and Massachusetts populations are Cyzicus.
Discussion
Caenestheriella gynecia inhabits temporary fresh-
water pools and was considered a “warm water” species
130
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Size and meristics of Caenestheriella gynecia (Clam Shrimp) specimens from New Jersey and New York.
State Number Mean Number of Number of
of specimens (range) size, mm telson spines segments, 2nd antennae
New Jersey 6 4.75 (4.2-5.2) 18-22 15-19
New York 5 9.10 (8.6-10.2) 27-32 15-20
by Weeks and Marcus (1997). This is typical habitat for
a suite of clam shrimp and relatives (Anostraca, Noto-
straca) and for certain distantly related species of Cla-
docera and Ostracoda (Williams 1987). The diversity
of aquatic animals in a given pool is low, however,
probably due to the habitat’s inherent instability and
simplicity. Caenestheriella gynecia appears to be dif-
ferent from its close and distant relatives in that it is
known mostly from rain puddles on dirt roads; the type
locality (Mattox 1950), a second location in Ohio (Em-
berton 1980), one of the populations in Massachusetts
(Smith and Gola 2001), and this paper. Although the
puddles in which we found C. gynecia are more-or-
less permanent, their small size (and high turbidity in
the case of the New Jersey pools) may exclude some
aquatic predators much as does impermanence.
Wiggins et al. (1980) and Williams (1985) character-
ized the resident biota of ephemeral aquatic habitats as
having high fecundity, short life spans, and rapid mat-
uration. Additionally Clam Shrimp produce “resting
eggs” (Hann 1996*) which can tolerate heat, cold, and
dessication and may require freezing or drying to hatch
(Mattox and Velardo 1950; Belk and Belk 1975).
Hann (1996*) observed that the small scale distribu-
tion of Caenestheriella setosa was patchy, with some
puddles inhabited while nearby puddles were empty.
This suggests that, despite the Clam Shrimp’s invest-
ment in large numbers of resting eggs, egg survival is
low, even short-distance dispersal is difficult, or local
extinction (in a pool) is frequent. Boileau and Taylor
(1994) showed that related organisms had poor disper-
sal over even very short distances and calculated that
there were 0.6-4.5 dispersers per generation in small
pools in South Carolina. We presume that dispersal of
Clam Shrimp is primarily in the resting egg stage and we
can envision several modes of dispersal of these eggs.
Eggs of C. gynecia from desiccated puddles could
be blown along with soil particles from one puddle or
locality to another. Animals passing through puddles,
especially species that spend time in muddy habitats,
could pick up eggs in mud adhering to their feet, shells,
or fur. Potential vectors of dispersal include Raccoons
(Procyon lotor, see Maguire 1963), Mallards (Anas
platyrhynchos), Snapping Turtles (Chelydra serpenti-
na), and humans. More recently, human traffic in the
form of wheeled or tracked vehicles could act as a
dispersal agent.
Although our few specimens of C. gynecia do little
to establish a pattern that would resolve the taxonomic
question posed by Smith and Gola (2001), there is a
geographic pattern that deserves discussion. All known
localities for this species are within the maximum ex-
tent of the Wisconsinan glaciation (Figure 2) except
one location in central Ohio (Emberton 1980). It is, of
course, possible that not all populations of this species
have been documented (especially since we are here
reporting several) and that there may be populations
to the south of the known distribution. This is impor-
tant because robust biogeographic hypotheses do
require accurate knowledge of an animal’s distribu-
tion. Although this species is rare, there are a number
of researchers actively working with Clam Shrimp in
the midwestern and southern United States and there
have been no other reports of this species.
This distribution suggests the following hypotheses.
Caenestheriella gynecia is derived from a sexually re-
producing species, probably C. setosa, whose range is
closest to C. gynecia. Dispersal into the glaciated area
occurred in the Ohio Valley and was an unlikely event
given the dispersal mechanisms of the species. Unlikely
dispersal would favor establishment of a parthenoge-
netic individual over at least two sexually reproducing
ones. This scenario suggests that C. gynecia is relative-
ly young, less than 12 500 years old (the approximate
age of the last glacial retreat).
Dispersal eastward from the Ohio Valley would be
equally unlikely except that seasonally muddy dirt roads
were created soon after European colonization of the
northeastern states. Caenestheriella gynecia could have
spread as eggs in mud splattered on wagons or horses’
hoofs and its distribution would follow traffic flow. Re-
cently, many of these colonial roads have been paved,
leaving C. gynecia to persist in the remaining dirt road
habitats, including farm roads, logging roads, and serv-
ice roads for powerlines and pipelines.
The above scenario could be corroborated by phy-
logenetic and genetic analyses. The closest living rel-
ative of C. gynecia should be geographically very close
to C. gynecia but we are unaware of any phylogenetic
analyses of this genus. Additionally, the genetic vari-
ation in C. gynecia should be minimal because it is
parthenogenetic, has been derived from a very small
founder population, and is geologically young.
Caenestheriella gynecia or its ancestors could have
evolved in association with American Bison (Bison
bison), living in “buffalo wallows” and dispersing in
mud caked on Bison fur or hoofs. Butler (1995) noted
buffalo wallows somewhat larger than the puddles
where we collected C. gynecia and alluded to observa-
tions of mud stuck to fur. Dispersal of resting stages in
the guts of birds is also possible for C. gynecia. Proctor
et al. (1967) found that Artemia eggs and the eggs and
adults of Ostracoda could remain viable after passing
through the gut of Killdeer (Charadrius vociferus).
2007
SCHMIDT AND KIVIAT: CLAM SHRIMP IN NEW YORK AND NEW JERSEY 13]
Lae
FIGURE 2. Map of the eastern United States showing collection localities of Caenestheriella gynecia in relation to the south-
ern extent of the most recent glaciation.
Long distance (1400 km) dispersal of Caenestheriel-
la belfragei was presumed due to eggs being carried
on or in waterfowl (Donald 1989). Proctor (1954)
showed that viable eggs of some phyllopods could pass
through the gut of ducks. If C. gynecia had been car-
ried to the glaciated areas by waterfowl, its close rel-
atives could be anywhere along the waterfowl flyways.
Two of the New York and the New Jersey Clam
Shrimp localities are privately owned. The Hyde Park
road is part of a residential subdivision and reportedly
has been excavated for installation of a water pipeline.
The New Jersey road is subject to gas pipeline main-
tenance activities and adjoins The Richard P. Kane
Natural Area, possibly subject to habitat restoration.
Because field surveys have not been conducted to
determine the distribution of, and degree of threat to,
C. gynecia populations in the northeastern states, we
recommend this species be treated as rare and vulner-
able to extinction unless demonstrated otherwise. Only
11 localities (Figure 2) have been reported for C. gyne-
cia, including our localities, although a recent Ohio
survey listed just one site (Weeks and Marcus 1997).
If this Clam Shrimp were reviewed by state Natural
Heritage Programs in New York and New Jersey it
would be ranked as G2, S1 assuming all the reported
sites are extant. We recommend that dirt roads one year
or more old with long-lasting rain puddles > 1 m* and
>5 cm deep be checked for Clam Shrimp during June-
September as part of land management or environmen-
tal review processes. Clam Shrimp may sometimes be
observed swimming in the pools and may be captured
by sweeping a fine-mesh dipnet through the pools.
Research, conservation, management, and restoration
of biodiversity are most often done in wildlands and
natural areas. Yet urban, industrial, and other altered
landscapes are expanding rapidly in North America.
Certain rare or declining native species survive or even
thrive in human-altered landscapes. Examples of rare
species occupying human-disturbed soils are: Knies-
kern’s Beaked Rush (Rhynchospora knieskernii) in a
tire rut and on a waste dump in the New Jersey Pine
Barrens (Schuyler 1999) and larvae of the Regal Fritil-
lary (Speyeria idalia) overwintering in soils disturbed
by tanks in Pennsylvania (Rosenzweig 2003). These
ecological distributions can be the result of human
activities creating a surrogate habitat type or physi-
cal-chemical environment not otherwise available, an
abundance of food, reduced predation or competition,
or persistence during landscape change (see, e.g., Adams
1994). It can be said that “Biodiversity is where you
find it.” Therefore conservation biologists and environ-
mental planners need to pay more attention to organ-
isms such as Caenestheriella gynecia and the artificial
or altered habitats that support them. Altered or arti-
ficial habitats of types known to support rare organ-
isms should be surveyed for those species routinely,
much as natural and semi-natural habitats are surveyed
for rare species. Those sites discovered to support a rare
132
species, such as the Clam Shrimp sites reported here,
should be considered for conservation management. In
the case of Caenestheriella gynecia, management may
need to include a degree of controlled motor vehicle
activity or its equivalent to maintain the puddle habitat
and potentially disperse Clam Shrimp among puddles.
We recommend that this species be considered rare and
vulnerable to extinction throughout its range unless
demonstrated otherwise.
Acknowledgments
Joseph T. Bridges and Stephen Nyman collaborated
in the field. Kathleen Schmidt drew Figure 2. Clam
Shrimp were collected and the manuscript prepared dur-
ing studies funded by the Geraldine R. Dodge Foun-
dation, H2O Fund (Highlands to Ocean Fund), Geof-
frey C. Hughes Foundation, Mary Jean and Frank P.
Smeal Foundation, Hackensack Meadowlands Part-
nership, Matthew D. Rudikoff Associates, and U.S.
Environmental Protection Agency. This paper has not
been reviewed by the funders; opinions herein are sole-
ly those of the authors. This is Bard College Field Sta-
tion — Hudsonia Contribution 87.
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Received 30 January 2006
Accepted 1 May 2008
Colonization of Non-Traditional Range in Dispersing Elk, Cervus
elaphus nelsoni, Populations
FRED VAN DYKE
Montana Department of Fish, Wildlife and Parks, Red Lodge, Montana 59068, USA
Present address: Department of Biology, Wheaton College, Wheaton, Illinois 60187 USA; e-mail: fred.g. vandyke @ wheaton.edu
Van Dyke, Fred. 2007. Colonization of non-traditional range in dispersing Elk, Cervus elaphus nelsoni, populations. Canadian
Field-Naturalist 121(2): 133-141.
As ungulate populations increase in density on traditional range, resulting increases in intraspecific competition can encour-
age dispersal of some individuals to new areas. Such areas, although lower in density of conspecifics, might present unfamil-
iar arrays and types of habitat that could require altered patterns of home range and habitat use by dispersers. However, the
specific adaptations employed by dispersers in such circumstances are not well documented or understood. I investigated
three cases of range expansion by Elk (Cervus elaphus nelsoni) populations experiencing population growth on traditional
ranges in south-central Montana, USA. Each source population produced a group that dispersed to non-traditional areas.
Compared to source populations, dispersing groups increased average size of home ranges, changed patterns of use in core
areas of home ranges, and used habitats differently than Elk on traditional range. Dispersing groups demonstrated fidelity to
new ranges equal to that of source populations, but their seasonal tenure on non-traditional range was strongly linked to envi-
ronmental conditions, especially rainfall. Dispersal of groups increased the overall range of the population and its range of
habitat use. In growing populations of Elk, managers should determine if dispersing groups exist and whether they should be
protected to establish new populations in marginal areas or be reduced to limit potential Elk-landowner conflicts.
Key Words: Elk, Cervus elaphus, dispersal, habitat use, population, Montana.
On traditional ranges, animals typically concentrate
use in the most productive patches of habitat (Charnov
1976; Parker and Stuart 1976). However, intraspecific
population pressure, often associated with population
growth, can cause a species to increase its range of
habitat use in less predictable ways (Rosenzweig 1991).
For non-territorial species, the tendency to increase
range of habitat use with increasing population densi-
ty has been formalized as the Ideal Free Distribution
(IFD) Theorem. The Ideal Free Distribution Theorem
asserts that individuals choose habitats at densities asso-
ciated with equal levels of fitness (Fretwell and Lucas
1970). Thus, IFD predicts that when populations occu-
py new ranges at low densities, individuals will occu-
py only optimal habitat. As density increases, so does
interference with conspecifics, causing individual fit-
ness to decline. Individuals leave optimal habitat when
fitness drops below what they could achieve in a sub-
optimal habitat at a lower population density.
Among ungulates, the behavior of Elk (Cervus ela-
phus nelsoni) in populations experiencing long-term
population growth corresponds to predictions of IFD.
Dispersive behavior associated with population growth
and an increasing range of habitat use might contribute
to the initial stages of colonization and the establishment
of a permanent population in a previously unoccupied
area (Safriel and Ritte 1983). The colonization of vacant
habitat without decreased fitness of colonists is an alter-
native to increased intraspecific competition on tradi-
tional range. Although the IFD Theorem makes den-
sity-dependent dispersal theoretically plausible, such
dispersal has been difficult to demonstrate in ungulates
(Clutton-Brock et al. 1985; Boyce 1989), and determi-
nation of the mechanisms of colonization by individu-
als dispersing from established ungulate populations,
or their relation to population growth, has been prob-
lematic.
In three increasing populations in south-central Mon-
tana, USA, Elk demonstrated changes in habitat use and
home-range characteristics, increasing spatial separa-
tion of adjacent populations, and fissioning of individ-
ual populations into multiple populations (Van Dyke
and Klein 1996; Van Dyke et al. 1998), all of which can
lead to an increasing range of habitat use. The last
response, population fissioning, was initiated by the
formation of a dispersing group that made seasonal
movements to new range, often containing a distribu-
tion of vegetation communities different from the tradi-
tional range of the established population. In investigat-
ing the role of such groups in expanding range and
habitat use of established populations, I sought to deter-
mine what changes dispersing Elk make in their use
of home range and habitats compared to their popula-
tions of origin that might enable them to persist on new
ranges that the established population did not use.
Study Area
Three populations of Elk in south-central Montana
(USA) were investigated, locally known as the Line
Creek (LC), Picket Pin (PP), and Silver Run (SR)
populations (Van Dyke and Klein 1996: Van Dyke et
al. 1998).
133
134 THE CANADIAN FIELD-NATURALIST
These populations summer in the Absaroka-Bear-
tooth Wilderness Area and winter in lower elevation
foothills and prairies on the Custer National Forest
and adjoining private land (Figure 1).
Climate in this area was characterized by long, cold
winters and short, cool summers during the study peri-
od. Mean January and July temperatures at the Mys-
tic Lake, Montana, reporting station (elevation 2339 m)
near the approximate center of the study area were -4.1
and 17.1°C, respectively. Average annual rainfall and
snowfall reported at the same location were 64.7 and
497.9 cm, respectively (NOAA 1992).
Forests were dominated, from lowest to highest ele-
vations, by Limberpine (Pinus flexilis), Lodgepole Pine
(P. contorta), Douglas Fir (Pseudotsuga menziesit),
Engelmann Spruce (Picea engelmannii), and White-
barked Pine (Pinus albicaulis). Non-forested commu-
nities at lower elevations were dominated by Bunch-
grass (Pseudoroegneria spicata)-forb and sagebrush
(Artemisia spp.) associations. Alpine areas were dom-
inated by sedges (Carex spp.), cottongrass (Eriopho-
rum spp.), Alpine Bluegrass (Poa alpina), and vari-
ous alpine forbs. Detailed descriptions of these plant
communities are provided in Van Dyke et al. (1991)
and Van Dyke et al. (1994).
Methods
Sampling Procedures
Thirty-five female Elk were captured and radio-
collared in January-March 1988 (19), December and
February 1989 (11), March 1990 (1), and March 1991
(4). All but three were captured by darting from a
helicopter (Nielson and Shaw 1967) with Carfentanil
(dosage 0.0082 mg/kg). Two Elk were captured in bait-
ed traps and one Elk by a net-gun fired from a helicop-
ter. All were fitted with telemetry collars (AVM Instru-
ment Co., Telonics, Inc.). Procedures associated with
locating individual animals have been described (Van
Dyke and Klein 1996; Van Dyke et al. 1998).
Detection of Dispersing Groups
I documented movements of small groups (<40 in-
dividuals) away from source populations (120-160 indi-
viduals) through relocations of =1 radio-collared Elk
in each group. Such groups used different ranges and
remained isolated from individuals in the source pop-
ulations for 6-9 months each year. One group separat-
ed from each source population. These were the Trout
Creek (TC) group from the PP population, the Hell-
roaring (HR) group from the SR population, and the
Wolf Creek (WC) group from the LC population.
The relationship of dispersing groups to source pop-
ulations, as well as the boundaries of seasonal ranges
of the dispersing groups, was unknown prior to the
study. As a result, individuals in dispersing groups were
unintentionally collared when they were near source
populations on winter range. Although only a small
number of Elk were radio-collared in each group, the
percentages of radio-collared Elk in such groups were
Vol. 121
= those collared in source populations (5-12% vs. 4-
6%, respectively). Given the sociability of Elk, previ-
ous studies have been successful in tracking large
groups even when only one radio-collared animal was
present (e.g., Craighead et al. 1973), and that also was
the case in this study. Locations of radio-collared
individuals were considered representative of the entire
group because: (1) dispersing groups contained few
(7-40) individuals; (2) dispersing groups remained con-
stant in size during occupancy on non-traditional range,
indicating that groups possessed a high degree of cohe-
siveness; and (3) radio-collared individuals always
were present in the group when they were located.
Estimation and Analysis of Home Range Characteristics
Home range of each population or group was defined
as the 95% contour minimum convex polygon (MCP)
area and associated activity centers of source popula-
tions and dispersing groups as computed by HOME
RANGE, a home range analysis program (Ackerman
et al. 1990). To estimate seasonal home ranges, sea-
sons were defined as winter-spring (1 December — 31
May) and summer-autumn (1 June — 30 November).
Seasons were combined in this manner because source
populations had substantial overlap between winter and
spring ranges and between summer and autumn ranges
(Van Dyke and Klein 1996; Van Dyke et al. 1998).
Because MCPs are sensitive to sample size, I eval-
uated only populations with = 21 locations/season for
calculation of seasonal range and =45 total locations
for calculation of annual range. All samples were
equal to or larger (seasonal x = 97, range 21-438;
annual x = 189, range 45-787) than these minimums.
Previous tests of the relation between numbers of loca-
tions and home area estimates indicate that such sam-
ple sizes estimated = 81% of population home ranges
for ungulates in this area (Van Dyke et al. 1995; Van
Dyke et al. 1998). Core areas within home ranges
(Kaufman 1962) were identified by comparing actual
range use distributions generated from harmonic mean
estimates of home range (Dixon and Chapman 1980)
to hypothetical uniform use distributions (Samuel et
al. 1985). I also determined the percent contribution
of core areas to total home areas and core area per-
cent contribution to total use, and then determined
the ratio of use contribution to area contribution as an
index of the concentration of use in core areas.
Evaluation of Range Use
Home areas of dispersing groups and source popu-
lations were compared in four ways. First, as a first
approximation of potential movement differences in
landscape pattern and scale, I compared distances Elk
in dispersing groups moved from winter to spring or
summer ranges with the same movements in their
associated traditional populations using linear, straight-
line distances between seasonal geometric activity
centers in each group as determined by pooled loca-
tions from 1988-1992. I compared distances associ-
2007
30 km
STONE RNG
Ry
x
~ GALLATIN
NATIONAL
FOREST
VAN DYKE: COLONIZATION OF NON-TRADITIONAL RANGE BY ELK
Ny NORTH
ABS
Sd Sebi
PICKET PIN Nair
POPULATION
x ewarer ae
SILVER RUN
POPULATION
LINE CREEK
POPULATION
te
SHOSHONE NATIONAL FOREST
Ficure 1. Study area and approximate ranges of the Line Creek, Picket Pin, and Silver Run populations of Elk in south-central
Montana, USA, 1988-1991.
ated with such movements to the radius of the shared
winter home area as an index of relative movement
scale. Second, I compared home range size between
source populations and dispersing groups using 2-
sample t-tests. Third, I evaluated Euclidean distances
between points in associated dispersing group and
source populations through a multiple range permuta-
tion procedures test, which evaluates whether distri-
butions of distances in different groups come from a
common probability distribution (Mielke et al. 1976).
Fourth, I compared use-area ratios of core areas be-
tween dispersing groups and associated source popu-
lations to determine if differences existed in degree of
concentration of core area use using a 2-sample rtest.
The ratio of core to non-core use is important because
core areas would be likely to include the most opti-
mal habitat within a home range. Therefore I treated
the concentration of use in core areas as an index of
relative differences between optimal and suboptimal
habitats within a home range.
Use of Vegetation Communities and Production of
Forage
The types of vegetation communities present on
all seasonal ranges of all populations varied, but were
simplified for analysis to categories of alpine, grass-
forb (meadows), Douglas Fir, Limberpine, Lodgepole
Pine, sagebrush, Spruce (subalpine forest), and White-
bark Pine. Together, these vegetation communities cov-
ered the range of elevations used by all populations
in all seasons as well as broadly covering the array of
vegetation communities encountered and used by EIk.
Comparisons of use of vegetation communities by Elk
in source populations, based on locations of animals, to
availability of such communities visually identified at
568 randomly selected points within home areas of such
populations were evaluated using the Design I version
of Manly et al. (2002) resource selection function (RSF)
model. This version of the RSF model, which uses
independently sampled counts of available and used
resources, assumes that the availability of vegetation
communities does not change over the course of the
study period, that vegetation communities were cor-
rectly identified, and that Elk had free and equal access
to all vegetation communities within their designated
ranges, assumptions that were met in this study. The
model’s null hypothesis that Elk select vegetation com-
munities in proportion to their occurrence was tested by
comparing use to availability via a Chi-square good-
ness of fit test (Manly et al. 2002). The application of
this procedure to similar tests for random selection of
136 THE CANADIAN FIELD-NATURALIST
each vegetation community was adjusted using Bon-
ferroni corrections to prevent error accumulation. I
used Manly’s standardized selection ratio to represent
relative strength of selection for a given vegetation
community. Vegetation communities with 0 availability
in a given range were omitted from analysis.
Locations of Elk during 1988-1991 were used to
determine core foraging areas of dispersing groups
on summer ranges. Using radio telemetry locations
and visual observation, I established five 404-m?
(20.1 m x 20.1 m) macroplots selected within core
foraging areas determined in each dispersing group
and in the core foraging area of the PP population in
1991. Ten macroplots already had been established in
core foraging areas of LC and SR populations (five
in each area) as part of another study (Van Dyke et al.
1994). I compared volumes of forbs and graminoids
produced from each core area associated with dispers-
ing groups to core areas of respective source popula-
tions through 2-sample t-tests. Sampling methods have
been described (Van Dyke et al. 1991, 1994). Shrubs
were not included in the analysis because shrub cov-
erage was low on most plots and summer diets of Elk
in all populations averaged <10% shrubs (Van Dyke
et al. 1994). Although species-specific forage selec-
tion was evident (Van Dyke et al. 1994), all available
graminoids and forbs were consumed. Thus, estimates
of forb and graminoid volume were considered to rep-
resent volume of edible browse.
Results
Seasonal dispersal and home range characteristics
All dispersing groups moved greater absolute and
relative distances from winter to spring or summer
ranges than traditional populations, and, in two of
three cases, traveled in entirely different directions and
followed different elevational gradients (Table 1). Rela-
tive to the size of winter home areas, traditional pop-
ulations migrated distances 2.3-7.5x the radius of their
winter home areas to reach spring and summer ranges,
but dispersing groups moved 5.6-11.0x the radius of
their winter home areas to reach new, non-traditional
ranges in the same seasons.
Elk in dispersing groups used annual or seasonal
ranges differently than Elk in associated source pop-
ulations (Table 2). Differences also existed in home
range sizes of dispersing groups and source populations
(Figure 2). The annual and seasonal home ranges of
Elk in the TC and WC groups were larger than home
ranges of Elk in associated source populations. In con-
trast, seasonal home ranges of Elk in the HR group
were similar to those estimated for individuals in the
associated source population. Differences between
source and dispersing group populations also were evi-
dent in use of core areas (Figure 3). Hellroaring Elk
made greater use of smaller areas (higher use/area
ratios) in summer and autumn than did Elk in their
source (SR) population. A similar trend was evident
in the more concentrated use of winter and spring
Vol. 121
core areas by the WC group. In contrast, the TC
group demonstrated more dispersed range use than
its source (PP) population, and had no identifiable
core area in winter-spring (use/area ratio = 1).
Patterns of habitat and elevation use in source
populations: Comparisons of source populations and
dispersing groups
No population showed random proportional selec-
tion of vegetation communities (P < 0.001, all cases),
but selection patterns differed in different populations,
and, in two of three cases, patterns of selection in source
populations differed from selection patterns in asso-
ciated dispersing groups. The SR source population
and HR dispersing group were the only source popu-
lation-dispersing group pair in which selection of vege-
tation communities did not differ. Both used alpine
areas less frequently than expected and used Lodge-
pole Pine communities more than expected (Table 3).
The Line Creek source population and its associated
WC dispersing group differed in selection patterns in
four of seven vegetation communities (Table 4), and
the PP source population and its TC dispersing group
in four of eight (Table 5), although patterns were dif-
ferent in each case. Wolf Creek Elk used sagebrush
communities most commonly in the spring, followed
by an intensive shift to alpine communities in the sum-
mer, resulting in nearly equal rates of use in these com-
munities over the combined period. However, WC Elk
did not use any one vegetation community differently
than expected, although their overall pattern of use
was different from availability. In contrast, LC Elk
made more equitable use of all available vegetation
types, although they were most commonly located in
stands of Limberpine (33%). Line Creek Elk used
grass-forb and Limberpine communities more than
expected, and sagebrush and alpine communities less
than expected. Actual proportional use in grassland-
forb communities was identical in the two populations.
Observed differences in significance in this case were
attributable to differences in the number of observa-
tions in the two groups.
The majority of PP Elk locations were in grass-forb
vegetation (56%), and this population used this vegeta-
tion type and Limberpine at greater than expected fre-
quencies. Picket Pin Elk used Lodgepole Pine less than
expected. In contrast, TC Elk were most commonly
located in stands of Douglas Fir (42%). They used this
vegetation community more than expected, but used
spruce-dominated vegetation less than expected.
Core foraging areas of dispersing groups and source
populations differed in volumes of forbs and gram-
inoids produced (Table 6). Trout Creek core areas had
higher volumes of graminoids and forbs compared to
PP core areas, but differences in other comparisons
displayed no consistent pattern.
Two dispersing groups, TC and WC, showed dif-
ferent patterns in seasonal elevational use compared
to their source populations (Figure 4). Trout Creek
2007
VAN DYKE: COLONIZATION OF NON-TRADITIONAL RANGE BY ELK
137
TABLE 1. Scale considerations in distance, direction, and elevational differences in pairs of traditional source populations
and dispersing groups of Elk in south-central Montana, U.S.A.,
1988-1991. In each pair, the traditional source population is
listed first. Migration distance (km) reflects distance from shared winter range activity center to traditional source popula-
tion or dispersing group spring or summer range. Home range radii estimated as square root of home range area ( km*)
divided by st.
Migration Migration Elevation Population home
Population distance direction change range radius — winter
Line Creek 4.9 W Higher 2.12
Wolf Creek 11.8 NE Lower
Picket Pin 11.6 W Higher 1.54
Trout Creek 17.0 NE Same
Silver Run 11.8 SW Higher 1.70
Hellroaring 14.6 SW Higher
TABLE 2. Results of Multiple Range Permutation Procedures analysis comparing range utilization between dispersing groups
and associated source populations in south-central Montana 1988-1991. P < 0.05 interpreted as evidence of significantly
different distributions between dispersing group and associated source population. n = number of Elk locations used in analysis
(source population and dispersing group, respectively). Seasons selected represent periods of movement to new ranges by
dispersing groups.
Population Season n Standardized test statistic 12
Picket Pin-Trout Creek Annual 348/71 -102.89 <0.0001
Silver Run-Hellroaring Summer-Autumn 291/58 - 46.24 <0.0001
Line Creek-Wolf Creek Spring-Summer 263/73 - 10.99 <0.0001
2 Se ae ees ee ee
Seasonal Home-range Size Core Area Concentration
300 54 @ Trout Creek
O Picket Pin
@ Trout Creek els
O Picket Pin v Heliroaning
v_ Hellroaring v Silver Run
Vv Silver Run 44 @ Wolf Creek
Lf @ Wolf Creek 0 Line Creek
Y= 200 0 Line Creek e =? ==+
‘6 e co =
> s a o T ~- |
2 & 34 |
< ® ro)
on
= = y | a
T = We it
100 i> ml)
5 1 2 4 j I a Mg I]
! a & 5 | :
a = |
: A
| 8 |
Winter/Spring ‘Summer/Autumn Annual |
Season 0 =
Winter/Spring Summer/Autumn Annual
FiGuRE 2. Seasonal and annual home area sizes of source
populations and associated dispersing groups of Elk
in south-central Montana, USA, 1988-1991. Bars
indicate 95% CI.
Elk used lower elevations in summer (f,,, = 3.62,
P < 0.0001), consistent with their tendency to move
to traditional, high elevation summer range later than
PP Elk. Similarly, WC Elk used lower elevations in
summer (t,,, = 2.03, P = 0.05) and autumn (,,, = 2.46,
P = (0.02), reflecting their tendency to spend at least
part of these periods off the Line Creek Plateau and
away from the main LC population. Hellroaring Elk
used elevations similar to SR Elk in all seasons, had
non-overlapping ranges in summer and autumn, and
Season
FiGuRE 3. Core area concentration indices (% use of core
area/% contribution of core area to home area) of
source populations and associated dispersing groups
of Elk in south-central Montana, USA, 1988-1991.
Bars indicate 95% Cl.
persisted on their new ranges throughout both seasons
in all years. In contrast, TC and WC Elk always aban-
doned the non-traditional ranges at some point during
the summer.
Over time, it became apparent that dispersing groups
abandoned non-traditional ranges more quickly in drier
years. I examined the strength of this relationship at
the conclusion of the study through a logistic regres-
sion analysis relating June and July rainfall (estimated
138 THE CANADIAN FIELD-NATURALIST Vol. 121
TABLE 3. Proportional availability and use of vegetation communities on summer-autumn range by the Silver Run Elk popu-
lation and an associated population (Hellroaring) colonizing non-traditional range in disjunct areas in south-central Mon-
tana, USA, 1988-1991. P values (Bonferroni corrected) equal the probability that proportional use equals availability.
Vegetation Proportion Proportion used Standardized selection ratio P
community available Silver Run Hellroaring SilverRun Hellroaring Silver Run Hellroaring
Alpine 0.70 0.48 0.48 0.08 0.07 <0.001 0.02
Lodgepole Pine 0.04 0.22 0.29 0.60 0.70 <0.001 <0.001
Sagebrush 0.08 0.14 0.12 0.21 0.16 0.27 1.00
Whitebark Pine 0.18 0.16 0.11 0.11 0.07 1.00 1.00
TABLE 4. Proportional availability and use of vegetation communities on spring-summer range by the Line Creek Elk popu-
lation and an associated population (Wolf Creek) colonizing non-traditional range in disjunct areas in south-central Mon-
tana, USA, 1988-1991. P values (Bonferroni corrected) equal the probability that proportional use equals availability.
Vegetation Proportion Proportion used Standardized selection ratio P
community available Line Creek Wolf Creek Line Creek Wolf Creek Line Creek Wolf Creek
Alpine 0.33 0.17 0.39 0.05 0.16 <0.001 1.00
Douglas Fir 0.08 0.14 0.08 0.15 0.13 0.11 1.00
Grass-forb 0.02 0.06 0.06 0.31 0.44 0.001 0.17
Limberpine 0.10 0.33 0.07 0.30 0.09 <0.001 1.00
Sagebrush 0.30 0.18 0.40 0.05 0.18 <0.001 0.60
Spruce 0.08 0.09 0.00 0.04 0.00 1.00 0.17
Whitebark Pine 0.09 0.04 0.00 0.10 0.00 0.21 0.13
TABLE 5. Proportional availability and use of vegetation communities on annual range by the Picket Pin Elk population and
an associated population (Trout Creek) colonizing non-traditional range in disjunct areas in south-central Montana, USA,
1988-1991. P values (Bonferroni corrected) equal the probability that proportional use equals availability.
Vegetation Proportion Proportion used Standardized selection ratio JB
community available Picket Pin Trout Creek Picket Pin Trout Creek Picket Pin Trout Creek
Alpine 0.07 0.02 0.12 0.04 0.25 0.57 1.00
Douglas Fir 0.15 0.13 0.42 0.11 0.39 1.00 <0.001
Grass-forb 0.25 0.56 0.33 0.28 0.18 <0.001 1.00
Limberpine 0.04 0.10 0.01 0.35 0.05 0.01 1.00
Lodgepole Pine 0.26 0.05 0.09 0.02 0.05 <0.001 0.06
Sagebrush 0.05 0.02 0.03 0.06 0.08 1.00 1.00
Spruce 0.16 0.11 0.00 0.09 0.00 1.00 0.03
Whitebark Pine 0.02 0.01 0.00 0.05 0.00 1.00 1.00
TABLE 6. Volume (m*/ha) of forbs and graminoids in core foraging areas of dispersing population groups and associated
source populations in south-central Montana 1988-1991. n (number of sampled macroplots) = 5 except for Line Creek (n = 4)
and Silver Run (n = 3). Means of plant volume in areas used by dispersing groups and areas used by source populations
compared by t-test, same categories.
Volume (m?/ha)
Area Forbs Graminoids
m3/ha SE m?/ha SE
Trout Creek SIS 62 1218 230
Picket Pin 228 32 307 2D
Hellroaring 3184 11 326° 84
Silver Run 215 15 1002 376
Wolf Creek 698 18 B24 65
Line Creek 415 108 210 i
* Different from source population. P < 0.0001.
2007
3000 Seasonal Elevation Use
2800 —@®- Trout Creek ff
Oo Picket Pin /
—¥— Hellroaring | ff o v
2600 9 SilverRun | .
—f— Wolf Creek | /
© LineCreek | yi
Elevation (m)
~
s
~ Sy
| ~
Ny aa Pe | / °
2000 ~ 7 oY ae
a
Ol pad en OZ
1800 ie
Winter Spring Summer Autumn
Season
FIGURE 4. Seasonal elevational use (m) of source populations
and associated dispersing groups of Elk in south-cen-
tral Montana, USA, 1988-1991.
from the Red Lodge, Montana recording station for
WC Elk and the Nye, Montana recording station for
TC Elk, NOAA 1992) to the total estimated number
of days in June, July, and August on new range. Vari-
ation in combined June and July rainfall explained
two-thirds (77 = 0.67, P < 0.0001) of the variation in
days on non-traditional summer range in the TC and
WC groups (Figure 5).
Discussion
Dispersal and density of populations
Historical expansion and contraction of ranges of
ungulate species has been influenced by many fac-
tors, both natural and anthropogenic (Laliberte and
Ripple 2004), but range expansions at local levels are
often achieved by dispersal, which in turn may be
mediated by density (Andersen et al. 2004). An eco-
logical correlate of a population’s capacity to colonize
new range is its ability to yield a large colonizing group
(Safriel and Ritte 1983), something which can be best
achieved during periods of population growth. In this
area, annual aerial counts indicated that all source
populations experienced population growth on tradi-
tional range in the decade prior to this study. From
1979 to 1988, the PP population increased from 44 to
190 individuals (355%), the SR population from 72 to
139 individuals (93%), and the LC population from
85 to 127 individuals (49%) (Van Dyke et al. 1998),
and all populations continued to increase through 1991.
These increases are small relative to fluctuations in
historically large populations of Elk like the nearby
Northern Yellowstone Population which has ranged
from 10000 to 20 000 individuals in recent years
(Committee on Ungulate Management in Yellowstone
National Park US National Research Council 2002).
Nevertheless, such increases were proportionately
large relative to past population size and occurred on
very small annual ranges (84 — 166 km?) (Van Dyke
VAN DYKE: COLONIZATION OF NON-TRADITIONAL RANGE BY ELK
139
60
° 7
e
.
50 P
|
|
e 40 ,
2 y = 36 44L n(x) - 17 707
f = 06724
30 /
Sa
20 e
10 + r =" —— —r
2 a 6 )
Rainfall (cm)
FiGuRE 5. Relation between June and July rainfall (combined)
and estimated number of days on non-traditional sum-
mer range in the Trout Creek and Wolf Creek dispers-
ing groups in south-central Montana, USA, 1988-1991.
et al. 1998) compared to those of the Northern Yel-
lowstone Population, whose annual range encompass-
es thousands of km? (Committee on Ungulate Man-
agement in Yellowstone National Park U.S. National
Research Council 2002). Thus, it is reasonable to
believe that the numerical increases observed in these
herds on these restricted ranges would increase den-
sities and competition for space and other resources,
contributing to the formation of dispersing groups,
although the dispersive behavior seen in these small
populations might be very different from that observed
in larger populations.
Size of home range and use and productivity of
vegetation communities
Variability in use and selection patterns of vegeta-
tion communities in different populations is, in part,
evidence of the behavioral plasticity of Elk. As these
populations experienced relatively large proportional
increases, the movement and fidelity of colonists to new
ranges appeared similar to observed seasonal move-
ments of White-tailed Deer (Odocoileus virginianus)
in areas fragmented by agriculture (Nixon et al. 1991).
However, whereas deer occupied traditional vegetation
communities in the new areas to which they dispersed,
Elk in these dispersing groups demonstrated their plas-
ticity in habitat selection by changing their patterns of
use of vegetation communities as they moved to new
ranges.
It is not only the proportion of a vegetation commu-
nity in the landscape that determines its use by Elk,
but also its juxtaposition and interspersion with other
such communities (Porter and Church 1987), a dimen-
sion of availability that was beyond the scope of this
study to measure. Thus, use of vegetation communities
by specific populations would be a poor guide for
landscape scale habitat management. In these popu-
lations Elk adjusted use of vegetation communities
according to local and seasonal availabilities.
140
Use of marginal habitats is associated with increased
rates of range expansion in some ungulate species
(Andersen et al. 2004). The larger home ranges of TC
and WC groups using non-traditional lower elevation
ranges in summer might indicate the same trend. Fur-
ther, larger home ranges in these groups also suggest-
ed that forage on non-traditional ranges might have
been more dispersed. However, comparisons of forage
volume in core foraging areas of group and source pop-
ulations did not support the premise that productivity
was lower in ranges of dispersing groups. On the TC
range, forage volume was higher in both graminoids
and forbs. However, foraging areas for TC Elk were
concentrated in riparian areas and probably could not
support higher densities or more extended use by Elk
in most years. Nutritional quality of vegetation is often
better at higher elevations (Johnston et al. 1968; Boyce
1989), a trend confirmed on these ranges in measure-
ments of protein and other nutrients in low vs. high-
elevation plants (Van Dyke et al. 1991; Van Dyke et
al. 1994). However, McCorquodale (1991) found that,
in sagebrush habitats used by Elk in central Washing-
ton similar to those used by the WC group, low inter-
community variability in forage production, lack of a
nonforage overstory, and the relative abundance of
foraging areas all mitigated against lower primary
production.
The low-elevation, mostly treeless habitat of the WC
group appeared to provide the greatest potential for
heat stress of Elk during summer months because of its
absence of thermal cover. However, Peek et al. (1982)
have argued that thermal cover is not a requirement
for Elk. Elk historically occurred on the Great Plains
and in other areas that lacked thermal cover. Loss of
Elk in these areas was due to human influence, not
lack of adaptability to the habitat (Laliberte and Rip-
ple 2004). In recent years Elk have recolonized areas
in treeless, sagebrush-steppe habitats in central Wash-
ington (Rickard et al. 1977; McCorquodale et al. 1986;
McCorquodale 1991) similar to those used by the
WC Elk.
Both source populations and dispersing groups
demonstrated fidelity to their selected home ranges
from year to year. Such fidelity suggested that dis-
persing groups had already established some degree
of tradition in their use of new ranges and were not
dispersing randomly. However, early abandonment
of non-traditional range at low elevations under drier
conditions suggested that such new ranges might have
been sub-optimal environments for Elk compared to
traditional range. Dispersing groups responded to peri-
ods of environmental stress by rejoining source pop-
ulations on traditional areas. By doing so, individuals
in dispersing groups might have benefited from the
source population’s collective knowledge of available
resources on traditional range and its successful, long-
term strategies of range use (Edge et al. 1985).
THE CANADIAN FIELD-NATURALIST
Vol. 121
Management Implications
Elk in dispersing groups demonstrated an ability to
locate and adapt to non-traditional range through var-
ious combinations of altered habitat use, changes in
patterns of seasonal elevational migration and adjust-
ments in home range size and use. Given this reper-
toire of adaptive strategies, managers should assume
that areas adjacent to existing Elk populations but
presently without Elk could be colonized, and could
be of importance to future growth of such populations
and expansion of their range. For example, in restora-
tions of Elk in Kentucky (USA), translocated Elk
demonstrated fidelity to a variety of habitat types, in-
cluding some not previously encountered on their orig-
inal ranges (Larkin et al. 2004). Such considerations
are of importance to managers as efforts to restore Elk
populations in several U. S. states (Didier and Porter
1999; McClafferty and Parkhurst 2001; Larkin et al.
2004) are being considered or have commenced.
As noted previously, dispersing groups from these
smaller populations might not exhibit the same pat-
terns or behaviors associated with dispersal as has been
observed in historically large herds. Thus, it would
be advisable to consider and evaluate dispersive behav-
ior associated with populations of different sizes before
making broad generalizations about patterns of dis-
persive behavior that could apply to populations of all
sizes. However, even with this caveat in hand, it will
be true that expanding ranges and growing populations
of Elk, regardless of size, might create added opportu-
nity to view and hunt Elk, but such expansion could
contribute to increasing conflict between Elk and other
land uses such as agriculture and must be examined in
a regional context. Where colonization of new range
by Elk enhances the persistence of marginal popula-
tions and provides desired recreational opportunities
for hunting and viewing Elk, managers should identify,
provide access to, and, if possible, acquire adjacent
range to which individuals might disperse, especially
within the broader context of regional strategies that
follow the principle of retaining large contiguous or
connected areas that contain critical habitats (Dale et
al. 2000). In these cases, dispersing groups should be
protected from disturbance until they have established
fidelity to new ranges (Larkin et al. 2004). Where
dispersion to adjacent, but non-traditional ranges and
habitats is likely to create or exacerbate land use con-
flicts between Elk and humans, managers should con-
sider lowering densities of Elk on traditional range to
reduce the likelihood of such dispersal.
Acknowledgments
R. J. Mackie and E. H. Merrill offered comments on
earlier versions of this manuscript. S. H. Yoo assisted
in preparation of text, tables, and figures. C. E. Eustace,
Montana Department of Fish, Wildlife and Parks, pro-
vided logistical support for fieldwork.
2007
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The Rumsfeld Paradigm: Knowns and Unknowns in Characterizing
Habitats Used by the Endangered Sharp-tailed Snake, Contia tenuis, in
Southwestern British Columbia
SHANNON F. WILKINSON!*, PATRICK T. GREGORY! >, CHRISTIAN ENGELSTOFT?, and KARI J. NELSON?
'Department of Biology, University of Victoria, P.O. Box 3020 STN CSC, Victoria, British Columbia V8W 3N5 Canada
*Alula Biological Consulting, 1967 Nicholas Road, Saanichton, British Columbia V8M 1X8 Canada
Ecosystems Branch, British Columbia Ministry of Environment, P.O. Box 9338 STN PROV GOVT, Victoria, British
Columbia V8T 5J9 Canada
4Present Address: 824 Leslie Drive, Victoria, British Columbia V8X 2Y4 Canada
Author for correspondence: viper @uvic.ca
Wilkinson, Shannon F., Patrick T. Gregory, Christian Engelstoft, and Kari J. Nelson. 2007. The Rumsfeld paradigm: knowns
and unknowns in characterizing habitats used by the endangered Sharp-tailed Snake, Contia tenuis, in southwestern
British Columbia. Canadian Field-Naturalist 121(2): 142-149.
The Sharp-tailed Snake, Contia tenuis, has a small and highly fragmented range in British Columbia, where it is considered | _
endangered. Known sites are few in number and generally small in spatial extent; numbers of snakes apparently are correspond-
ingly low. Furthermore, most known sites for the species are on private lands in areas that are fairly heavily developed or being
developed. Thus, the species is under serious threat of habitat alteration or loss. Although land stewardship has been a valuable
conservation tool in this case, we also need to identify the key habitat requirements of Sharp-tailed Snakes to identify potential
new sites, modify former or potential ones, or even create new ones. In this study, we compared sites known to harbour Sharp-
tailed Snakes with those that seemed subjectively similar and therefore potentially suitable. We also compared these known and |
potential sites with randomly chosen nearby locations. Variability of most measured features was high, both within and among
site/location categories. Nonetheless, we found significant differences between known and potential sites and between those
locations and random ones. Overall, locations known to be used by snakes had a more southerly aspect, more rock cover, shal-
lower soil and litter, and less shrub cover than other sites. This study was constrained by the small number of known sites for
Sharp-tailed Snakes in southwestern British Columbia, making our conclusions suggestive rather than definitive. Future work
should incorporate additional variables. It also might be useful to undertake comparative habitat studies elsewhere in the range
of the Sharp-tailed Snake where it is more common.
Key Words: Sharp-tailed Snake, Contia tenuis, habitat, British Columbia.
On 12 February 2002, U.S. Secretary of Defense
Donald Rumsfeld made the following widely cited
observation:
“As we know, there are known knowns; there are things we
know we know. We also know there are known unknowns; that
is to say, we know there are some things we do not know. But
there are also unknown unknowns, the ones we don’t know we
don’t know. And if one looks throughout the history of our
country and other free countries, it is the latter category that
tend to be the difficult ones.”
Although Rumsfeld was not discussing science, his
circumlocutional comments nonetheless may well be
apt in science (Shermer 2005). In conservation biology,
the unknowns, whether known or unknown, usually
exceed the known knowns and we often have to make
educated guesses in, for example, trying to determine
where there might be additional suitable habitats for a
threatened or endangered species.
To manage and conserve any species, it is crucial to
understand its habitat requirements. The fitness of any
species should be greatest in a habitat for which it is
best suited and that it presumably prefers (Garshelis
2000). Determining critical habitats is an important
component of the background knowledge that allows
researchers and managers to develop strategies to either
recover or maintain an endangered or threatened species
and to choose areas in which to explore such strategies.
The Sharp-tailed Snake (Contia tenius) is a small
secretive snake, from 20 to 45 cm in total length when
adult (Leonard and Ovaska 1998), that ranges from
southern California to southern British Columbia (Cook
1960). Although the southern part of its range, to north-
ern Oregon, is more-or-less continuous, the distribution
of this species is highly fragmented in Washington
and British Columbia (Leonard and Ovaska 1998). In ~
British Columbia, Sharp-tailed Snakes are known only |
from a small number of scattered locations on southern
Vancouver Island and some of the adjacent Gulf Is--
lands. A previous report from Chase in the interior of |
British Columbia is of doubtful origin (Matsuda et al.
2006). Because of the low numbers of snakes found in)
Canada and few known sites, the snake was consid-
ered an endangered species by COSEWIC in 1999
(COSEWIC 2002*). Although formal study of Sharp-
tailed Snakes has been limited, recent studies done in
British Columbia and adjacent Washington cover as-
pects of their ecology such as defensive behaviour |
(Ovaska and Engelstoft 1999; Leonard and Stebbins
1999), aggregative behaviour (Leonard et al. 1996),
142
2007
movements (Engelstoft et al. 1999), habitat (Leonard
and Leonard 1998), and cover use (Engelstoft and
Ovaska 2000).
Most of the areas where the Sharp-tailed Snake is
found in British Columbia have fairly dense human
populations and highly disturbed habitat. Consequent-
ly, many of the sites at which snakes are found are on
private land. Although the Sharp-tailed Snake seems
somewhat tolerant of anthropogenic habitat distur-
bance, habitat loss is still considered the primary threat
to its persistence in Canada. Therefore, management
and stewardship by landowners are important factors
in conservation programs for this species.
The aim of this study was to determine the key char-
acteristics that define habitats used by the Sharp-tailed
Snake, in order to guide future habitat management.
We did this by comparing sites known to be used by
Sharp-tailed Snakes with sites that were subjectively
similar but not known to be used (potential sites), and
with locations chosen at random with respect to both
known and potential site types. The first comparison
objectively tested whether potential sites actually are
similar to those sites known to be used by these snakes
and the second tested whether Sharp-tailed Snakes use
a subset of those habitats that are available. Initial
observations suggested that south-facing rocky slopes
and forest clearings appeared to provide the most
suitable habitat for this species (see also Leonard and
Leonard 1998), but no quantitative assessment of habi-
tat characteristics has previously been done. Thus, we
measured a wide range of possibly influential habitat
variables at each sample location and compared them
statistically among locations.
Although our comparisons among sites and loca-
tions were objective, selection of potential sites obvi-
ously was biased by our general knowledge of snake
ecology and our previous experience with Sharp-tailed
Snakes. This mainly reflected an aim of the recovery
program for this species to find as many occupied sites
as possible.
Study Area
In the past few years, federal lands in the Capital
Regional District (including Gulf Islands National Park,
Department of National Defence lands, and Coast
Guard properties) have been assessed and rated for
suitability as habitat for the Sharp-tailed Snake (Engel-
stoft and Ovaska 1998*). These sites are in various
locations on North and South Pender Islands, Salt-
spring Island and southern Vancouver Island. In addi-
tion, artificial cover objects (ACOs) have been installed
at all sites at which Sharp-tailed Snakes are known to
occur, including private lands, and at sites rated as hay-
ing high potential as habitat for these snakes. ACOs
provide easily sampled, attractive cover and, combined
with naturally occurring cover objects, are an effective
means of determining presence of snakes (Engelstoft
and Ovaska 2000). Sharp-tailed Snakes are secretive
WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKE
143
Potential 2004
Known and potential 2004
New since 2004
FiGurE |. Map of study area with sites known to be occupied
by Sharp-tailed Snakes up to 2004 and since, and
potential sites identified in 2004. Squares labeled
“known and potential 2004” indicate known and
potential sites that are too close together to represent
separately on this map. For reference, latitude and lon-
gitude for Victoria are 48°25'N and 123°19'W, respec-
tively, and for Ganges 48°51'N and 123°31'W, respec-
tively. Modified from a map obtained from the Natural
Areas Atlas on the Capital Regional District (2008*)
website.
and rarely seen in the open, and ACOs provide the main
means by which they are discovered in the field. Poten-
tial sites in this study were identified by CE based on
his extensive field experience with Sharp-tailed Snakes.
The sites where ACOs have been placed can be
classified into two general types. Unrestricted sites are
those with relatively large extents of contiguous habi-
tat under the same ownership surrounding the location
of the ACO. These areas typically have experienced
relatively little disturbance in the last century (e.g..
Department of National Defence lands or Canada
Parks lands). Restricted sites are those on relatively
144
small, privately owned properties surrounded by private
lands in relatively developed areas. These locations
have generally experienced a greater degree of habitat
modification (buildings, roads, gardens, etc.).
Methods
Fieldwork for this study was undertaken in summer
and autumn of 2004. Our general sampling scheme
was straightforward. The 31 sites were divided into
two site types, those known to harbour Sharp-tailed
Snakes (known knowns) and those that appeared to
have the potential to harbour them (known unknowns).
Hereafter, we call these known and potential sites,
respectively (Figure 1). At each of these sites, in addi-
tion to the known or potential assigned location, we
also chose a corresponding random location (unknown
unknowns) for sampling. Random locations were 50 m
from an assigned location, but in a randomly chosen
direction. Because 50 m was well within the known
movement capabilities of Contia (Engelstoft et al.
1999), yet far enough away for the environment to vary
significantly, this distance provided a test of non-ran-
dom use of the local habitat. However, because most
known sites are restricted, we were able to obtain a
random location for only seven of the 16 known sites
(compared to 14/15 for potential sites). Our sampling
protocol was thus dictated to a large degree by access
to private property, and by the amount of contiguous
habitat under the same ownership surrounding the site.
The total number of assigned and random locations at
all sites combined was 52. At each of these locations,
we took samples along three randomly chosen 10-m-
long transects.
The first transect at each assigned location was
placed in a random direction from a randomly chosen
ACO, so that the O0-m mark was at the centre of the
ACO. The other two transects were placed at two dif-
ferent randomly chosen locations (and in random
directions) within a 25-m radius of the initial ACO.
We measured canopy cover, aspect, and slope of loca-
tion at the 0-m mark of the first transect; that is, these
variables were measured once per location. All other
variables were measured along each of the three tran-
sects (i.e., three replicates for each location). This
same protocol was repeated for random locations, with
a point on the ground standing in for an ACO. We
ensured that transects around an assigned location
and its associated random location did not overlap or
abut each other.
We measured the following variables at each of the
52 locations:
1. Percentage canopy cover — Canopy cover was measured
once for each location at the starting point of the first
transect, directly overhead, and then at a 45° angle in each
of the cardinal directions (south, north, east and west —
surrounding canopy cover). The tree species making up
the canopy cover also were recorded.
. Aspect — The aspect of each location was measured in
degrees using a compass at the starting point of the first
i)
THE CANADIAN FIELD-NATURALIST
Vol. 121
transect. A measurement of 180° indicates a southerly
aspect.
3. Slope — The general slope of the location was recorded
in degrees as well as the slope for each separate transect.
Both were measured using a compass equipped with a
clinometer.
4. Substrate — At the l1-m mark along each transect, we
determined the depth of both soil and litter layer by
measuring the distance a thick wire penetrated into the
ground. The composition of the litter (needles, leaves,
etc.) also was recorded.
5. Rock coverage — Rock coverage was determined at
each transect by measuring the total distance intercepted
by rocks along the length of the transect. The rocks were
rated by size as follows: loose gravel (pebbles <7.5 cm),
cobbles (7.5-19.9 cm), rocks (20-50 cm), boulders (>50
cm), and bedrock.
6. Coarse woody debris — This was determined by
measuring the total distance intercepted by coarse woody
debris along the transect. Each piece of coarse woody
debris was identified to species (if possible), and assigned
a decay class (1-5) as per Table 5.13 in Meidinger (1998*).
7. Vegetation — Shrub cover and ground cover were
determined by measuring the total distance intercepted
by vegetation along the transect. For logistical reasons,
coverage by each of lichens, bryophytes, grass and forbs
was expressed as the total distance intercepted by these
plants between the 2-m and 4-m marks of each transect.
Statistical Analyses
We used SAS 8.0 to analyze the data, except for
aspect, which we also analyzed using ORIANA soft-
ware. We compared locations and site types using both
univariate and multivariate statistics. Although one
level of the factor “location” was a randomly selected
partner to a known or potential assigned location for
Sharp-tailed Snakes, all factors (site, site type, location)
were treated as fixed factors because all sites initially
were chosen deliberately as particular site types and
locations. In cases of non-orthogonal data, we used
Type III sums of squares for F-tests. As this was essen-
tially an exploratory study, we used a liberal criterion
for tests of significance, highlighting all results for
which P < 0.10.
Most variables showed no clear pattern of hetero-
geneity among categories of sites, so we combined
them in various ways, which also reduced the number
of variables to manageable levels, as follows: total
shrub coverage (all shrubs — e.g., broom, Oregon grape,
salal); tall, non-shrub coverage (ferns and asters); thick
ground cover (e.g., ivy, periwinkle, trailing blackberry);
light ground cover (lichens, bryophytes, grasses, forbs);
rock cover (sum of all rock-size classes); coarse woody
debris (sum of all CWD decay classes); and surround-
ing canopy cover (average of directional values around
sample location). All other variables (e.g. overhead
canopy cover, soil depth, slope) were left unaltered.
Because assigned and random locations were paired
at unrestricted sites, we compared them via a 2-way
factorial analysis of variance (ANOVA), with the three
transects treated as replicates within each combination
2007
of site and location. We did this analysis separately for
known and potential site types, as they were in differ-
ent places. In each case, we had to drop restricted sites
from the analysis because they lacked a random loca-
tion. In all cases in which the interaction between fac-
tors was non-significant, we dropped it from the analy-
sis to test main factors only. In cases in which the
interaction was significant, compromising interpreta-
tion of main effects, we then did a comparison of loca-
tions for each site separately.
To compare known and potential site types, we used
a nested ANOVA (sites nested within site type), ana-
lyzing assigned and random locations separately, with
transects serving as replicates within locations. F-tests
were adjusted to those appropriate for nested ANOVA
(Sokal and Rohlf 1981).
Because aspect is an angular variable, with the low-
est possible value (0°) identical to the highest possible
(360°), we treated it differently from the other variables.
We categorized aspect at each location as one of four
quadrants representing the four cardinal directions and
then compared frequency of directions among site
type/location categories (known/assigned, known/ran-
dom, potential/assigned, potential/random) by contin-
gency-table analysis. We also used circular (angular)
Statistics, as described by Zar (1984), to compare mean
aspect between pairs of site type/location categories.
In addition to aspect, a few other variables (e.g. can-
opy cover) were recorded only once for each location,
rather than separately for each transect. We therefore
reduced the data set to average values for each loca-
tion and then analyzed those new variables using two-
way ANOVA (site x type x location, sites serving as
replicates within each combination of factor levels).
Again, we dropped the interaction term when it was
non-significant, but did a separate comparison of
locations at each site type when the interaction was
significant. We also tested for correlations among
these variables across all sites/locations.
Studies of habitat relationships are necessarily mul-
tivariate (Reinert 1993) and typically involve methods
such as Principal Components Analysis (PCA) and
Discriminant Function Analysis (DFA). In comparing
categories of habitats, such as we have in this study,
two approaches are possible (Quinn and Keough 2002).
One is the multivariate analysis of variance (MANO-
VA) approach embodied by Discriminant Function
Analysis (DFA). A second is to do a Principal Com-
ponents Analysis (PCA) of all groups combined and
then compare the derived components among groups
using univariate ANOVA. Although the two approach-
es do not produce identical results, they generally result
in broadly similar patterns of group differences (Quinn
and Keough 2002). Advantages of the PCA approach
include ease of analysis of multiple derived compo-
nents and straightforward post hoc comparisons of
particular groups (Quinn and Keough 2002), so we
used it here.
WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKI
145
To avoid pseudo-replication, especially for variables
for which only one measurement was made per loca-
tion (see above), we used the reduced data set (i.e.,
means of the three transects treated as data) for mul-
tivariate analysis. Using the same variables as in the
ANOVAs above, we subjected the data to PCA, ex-
amined each of the first three principal components
for influence of the original variables, and then treat-
ed each of the principal components as variables in
separate two-way ANOVAs (as above). We also used
the first principal component (PC 1) as the independ-
ent variable in a logistic regression comparing sites/
locations known to be used by Contia ys. all other
sites.
Results
Two-way ANOVAs using individual transect data
as values yielded few differences attributable to loca-
tion, although sites differed significantly for several
variables. In comparisons of assigned vs. random loca-
tions for known site types, we found an effect only of
rock cover, which was higher at assigned than at ran-
dom locations (F, ,, = 3.42, P = 0.07). In cases of
variables for which there was a significant interaction
between site and location (extent of litter, slope of
transect, coarse woody debris), differences between
locations were significant at some sites, but not in any
consistent direction.
For potential site types, assigned and random loca-
tions differed only in thick ground cover (F, ,.. = 38.8.
P = 0.053), which was higher in assigned locations.
Sites again differed significantly for several variables
and the interaction between site and location was sig-
nificant for four variables (extent of litter, rock cover,
light ground cover, shrub cover). Patterns from inter-
actions again generally were indistinct, but for the three
sites that differed significantly in rock cover, the value
was higher at assigned than random locations; for
three sites differing significantly in shrub cover, the
value was lower at assigned locations.
Nested ANOVAs comparing assigned locations be-
tween known and potential site types showed only that
known site types had lower soil depth (F, ,. = 3.20.
P = 0.08) and less coarse woody debris (F, ,. = 7.45.
P = (0.01) than potential site types. Again, there were
significant differences among sites for some variables.
Similar comparison of random locations between
known and potential site types revealed only that rock
cover was higher at known site types (F, ,, = 3.15.
P = 0.09), with sites again varying significantly for
some variables.
Two-way ANOVAs, using mean values (per 3 tran-
sects) as data (and all site type x location combinations,
n = 52), revealed more differences (Table 1). Overall
(i.e., assigned and random locations considered togeth-
er), known site types had shallower soil (F, ,, = 2.92.
P = 0.09), shallower litter (F, ,, = 2.95, P = 0.09),
1.49
and higher rock cover (F, 4 = 4.67. P = 0.04) than
146 THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Statistical summary of main habitat variables by site type (known vs. potential) and location (assigned vs. ran-
dom). Only variables that were significantly different in at least one analysis (see text) are shown. Individual data are means
for 3 transects at each site type/location combination; shown in the table are mean (top row) and + standard deviation, mini-
mum value — maximum value (bottom row) of these transect means.
Site type Known Potential
Location Assigned Random Assigned Random
(n= 16) (n=7) (n= 15) (n= 14)
Rock Cover (m) Dell 2.1 1.6 0.9
2.45, 0 — 7.46 2.24, 0.02 — 6.03 1.78, 0.02 — 6.40 1.04, 0.01 — 3.11
Coarse Woody Debris (m) 0.1 0.6 0.4 0.5
0.18, 0 —0.71 1.37, 0 — 3.70 0.43, 0- 1.71 0.72, 0 — 2.07
Thick Ground Cover (m) 0.6 0 0.3 0.03
0.96, 0 — 3.50 0,0-0 0.56, 0 — 1.61 0.05, 0 — 0.14
Shrub Cover (m) 1.8 2.3 1.8 3.5
1.83.0-—5.29 1.42, 0.41 — 4.00 1.85, 0- 6.95 3.04, 0 — 9.95
Soil Depth (cm) 6.6 8.4 9.0 9.4
2.44, 3.5 -— 10.9 3.58, 3.6 — 13.2 4.85, 4.3 — 22.2 4.30, 2.7 — 17.0
Litter Depth (cm) Dal De) 2.7 Bs
1.26, 0- 4.3 1.46, 1.0—5.3 1.14, 0.7-4.2 2.23, 0.7 — 8.7
Overhead Canopy (%) 28.8 17.1 eS 28.2
21.17, 0 — 60.0 22.15, 0 — 50.0 14.25, 0 — 50.0 21.6, 0 — 50.0
Surrounding Canopy (%) 25.4 21.1 19.1 Spl
1239825) 52-5 16.51, 0 — 42.5 14.74, 0 - 46.3 18.66, 0 — 52.5
Transect Slope (°) 13.0 15.4 11.2 Holl
7.76, 0 — 28.33 15.39, 0 — 45.33 9.56, 0 — 27.67 7.24, 0.67 — 27.33
potential site types, on average. Assigned locations had F, ,. = 3.02, P = 0.1; known random vs. potential ran-
more thick ground cover (F, ,, = 4.15, P = 0.047), dom —F, ,, = 3.55, P = 0.07, although this last result
less shrub cover (F, ,, = 3.59, P = 0.06), less overhead
canopy cover (potential site types only, F, ,, = 9.55,
P =0.005) , less surrounding canopy cover (potential
site types only, F,,, = 6.62, P = 0.02), and steeper
transect slopes (potential site types only, F, ,, = 5.90,
P =0.02) than random locations, on average.
As expected, many of the variables above were sig-
nificantly correlated over all site types/locations (n = 52
in all cases). These included mean soil depth vs. mean
litter depth (r = 0.40, P = 0.004), mean rock cover vs.
mean litter depth (r = -0.29, P = 0.04), mean soil
depth vs. mean rock cover (r = -0.38, P = 0.005) and
overhead canopy cover vs. surrounding canopy cover
(r = 0.58, P < 0.0001).
Direction (aspect) differed significantly among site
type/location categories (Fisher’s exact test, P = 0.004).
Thirteen known/assigned locations (81%) faced south
and the rest (3) all faced west, whereas only six (40%)
of potential/assigned locations faced south and another
six faced east. Potential/random locations were partic-
ularly deviant from known/assigned, with eight (57%)
facing either north or east. These results were consis-
tent with those from analysis via circular statistics
(mean vectors: known assigned — 203.99°; known ran-
dom — 215.15°; potential assigned — 155.73°; potential
random — 125.86°). Significant differences between
site type/location categories were as follows: known
assigned vs. potential assigned — F, ,. = 4.27, P = 0.05;
known assigned vs. potential random — F, ,, = 7.21,
P = 0.01; known random vs. potential assigned —
may not be reliable because of a uniform distribution
of points.
We ran PCA using various combinations of vari-
ables, dropping those that had low weights on PC 1.
Results were broadly similar in all cases, but they were
most clearly defined when we used mean rock cover,
mean soil depth, mean surrounding canopy cover, and
mean shrub cover as variables in the analysis. In that
case, PC 1 accounted for 44% of the total variance and
PC 2 and PC 3 23% and 20%, respectively. This high
cumulative percentage variance for the first three prin-
cipal components suggests that they summarized the
data efficiently (McGarigal et al. 2000). PC 1 was pos-
itively weighted by soil depth, surrounding canopy
cover, and shrub cover, but negatively weighted by
rock cover (Table 2).
Two-way ANOVAs on the PCs showed that PC 1
differed overall between both site types (F, 4. = 4.19,
P = 0.046) and locations (F, ,, = 4.84, P = 0.03).
Known site types had lower average values of PC 1
than potential site types and assigned locations had
lower values than random, consistent with the two-way
ANOVAs performed above on the individual variables
and with the signs of coefficients in PC 1. Random
locations had higher values of PC 2 than assigned loca-
tions, but only for potential site types (F, ,, = 3.86,
P = 0.06). Neither site types nor locations differed
with respect to PC 3.
Despite statistically interesting differences in PC 1
between site types and locations, differences overall
2007
WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKE
147
TABLE 2. Summary of weights (coefficients) of four variables on first three principal components in analysis of habitat vari
ables potentially influencing occurrence of Sharp-tailed Snakes. PC | accounts for 44% of variance and PC 2 and PC 3 23%
and 20%, respectively.
Variable PC | Coefficients
Soil Depth 0.497
Surrounding Canopy Cover 0.440
Shrub Cover ().489
Rock Cover -0.566
Site — type
Known Potential
tS
5 2
S
1
£
Suio
Penal:
3°
= -2
—3
A R A R
Location
FIGURE 2. Box-and-whisker plots of values of principal
component | by site type (known vs. potential) and
location (actual, A, vs. random, R). Horizontal line
in each box is median. Upper and lower boundaries
of box (hinges) indicate quartiles (25" and 75" per-
centiles). Whiskers from box reach to the most ex-
treme value that does not exceed 1.5 times the dis-
tance between the quartiles; n = 52.
among the four site type/location combinations were
not large (Figure 2). In fact, differences among these
categories were graded rather than discrete. We there-
fore divided site type/location combinations into two
categories, those known to be occupied by Sharp-tailed
Snakes and those not. The resulting binomial variable
was then regressed (logistic regression) against PC 1.
The logistic model was a satisfactory fit to the data
(Homer and Lemeshow test, x *, = 4.46, P = 0.81)
and the relationship was significantly negative (Wald’s
xX z = 4.25, P = 0.04; Figure 3). We also compared PC |
of known/assigned locations successively, via logis-
tic regression, to known/random, potential/ assigned
and potential/random locations. The trend also was
negative in each case, but significant (and strongly
so) only in the last one. Thus, sites known to be occu-
pied by Sharp-tailed Snakes differed from those not
known to be occupied.
Discussion
The study of how and why animals occur in partic-
ular habitats has a long history in ecology and has been
the subject of numerous reviews and syntheses (e.g.
PC 2 Coefficients PC 3 Coefficients
-0.59] 0.346
0.549 0.670
0.490 -(). 562
0.331 0.339
P (Presence of Contia)
ee aT
=) 0 3
Principal Component 1
FIGURE 3. Logistic regression of presence(1)/absence(0) of
Sharp-tailed Snakes vs. principal component | for
all site type/location combinations. Absence implies
“not recorded” rather than confirmed absence. Verti-
cal lines are data points, solid line is predicted
regression, and dashed lines are upper and lower
95% confidence limits on regression; n = 52.
Morse 1980; Gray and Craig 1991; Reinert 1993;
Garshelis 2000). Habitat use by a species is a function
not only of habitat selection based on resource require-
ments and physical factors such as temperature, but of
other factors such as population density and the pres-
ence of competitors and predators (Huston 2002: Mor-
ris 2003). Thus, the habitats used by animals may not
be those that are optimal for them in terms of fitness.
In fact, it is even possible that some occupied habi-
tats have negative implications for fitness (Van Horne
1983), for example if those habitats are sinks in a
source-sink system (Pulliam 1988). Conversely, opti-
mal habitats that could be used might not be occu-
pied simply because the species’ ability to disperse
there is limited (Huston 2002). In short, the analysis
and interpretation of habitat-use studies is fraught with
difficulties (Garshelis 2000). In that light, this study
148
is but a small first step towards understanding the
habitat relationships of the Sharp-tailed Snake. It is
further limited by its reliance on presence/absence
(really presence/not recorded) data rather than data
on abundance, or better, demographic performance
(Pulliam 1988; Garshelis 2000). In terms of Garshelis’
(2000) classification of methods, our approach is a
variant of the site-attribute design, but with the added
inclusion of predetermined potential sites for the
species’ occurrence.
Animals typically do not use their environments at
random, but non-randomly, selecting some sites over
others because of particular habitat and micro-habitat
features. Thus, the mere demonstration that places used
by a particular species are different from those not used
is unremarkable by itself. For the conservationist or
manager, however, determining which habitat charac-
teristics are important is fundamental to preservation,
restoration, or even creation of habitats.
What are the key features that describe suitable habi-
tats for the Sharp-tailed Snake? The data collected in
this study were highly variable and can only hint at
differences between site types and locations that appear
to be important. More detailed, longer-term work with
larger samples would allow us to reach stronger con-
clusions, but this study was limited by the small num-
ber of sites at which the Sharp-tailed Snake is known
to occur in southwestern British Columbia. It also
was limited by the subjectivity with which potential
sites were identified. For example, south-facing slopes
are often favoured by snakes and this is reflected in
the distribution of known and potential sites in this
study. However, a Sharp-tailed Snake recently was
found on a north-facing slope on Pender Island (D.
Spalding, personal communication), suggesting that
the species may occur in a wider range of conditions
than previously thought. Currently, a more randomized
approach to sampling sites for Sharp-tailed Snakes is
under way to reduce this kind of bias, but we still have
some way to go before we can convincingly move
known and unknown unknowns to the fully known
side of the ledger.
Aside from the necessary constraints of small num-
bers of known sites for Sharp-tailed Snakes, this study
also might have been limited by failure to measure
key environmental variables. For example, as is the
case for other ectotherms, the ecology of Sharp-tailed
Snakes is no doubt strongly influenced by temperature,
which will affect the kinds of habitats and microhabi-
tats they use at different times (Huey 1991; Reinert
1993; Blouin-Demers and Weatherhead 2001). Simi-
larly, given the potential for desiccation in this small
species, as well as a requirement for incubation sites
for eggs, moisture also is likely to be an important
physical factor. Therefore, cracks and openings for
underground access and suitable sites for egg-laying
or hibernation are potentially critical habitat features
for these snakes. However, measuring subterranean
access would be very difficult and we know nothing
THE CANADIAN FIELD-NATURALIST
Vol. 121
about the kinds of sites favoured by Sharp-tailed
Snakes for egg deposition or for spending the winter.
To deal adequately with temperature and moisture,
we would need to measure them across all seasons
and in multiple microhabitats per location simultane-
ously.
Despite these deficiencies, we found fairly clear in-
dications that sites used by Sharp-tailed Snakes tend
to be south-facing and to have relatively shallow soil
and litter, relatively high rock cover, and relatively low
shrub cover. Differences from randomly chosen sites
emphasize this species’ non-random use of habitats, but
differences between locations actually used by snakes
and those deemed to be potential habitat underline
the subjectivity of visual habitat assessment. That said,
this subjective approach based on field experience
has proven effective. Although none of the potential
sites used in this study have since yielded evidence
of Sharp-tailed Snakes, two of the known sites were
initially identified by CE as potential sites and anoth-
er site designated as potential by CE after this study
was completed has since turned out to be occupied by
Sharp-tailed Snakes. Even so, quantitative measures
for objective assessment of potential sites also would
be valuable. Future work could focus on variables
found to be important in this study and on others that
we were unable to measure, such as those mentioned
above as well as shrub height, tree density, soil com-
position, drainage, and type of bedrock.
Given the rarity of the Sharp-tailed Snake in British
Columbia and the obvious difficulties in studying
rare species in general, perhaps an alternative (or sup-
plementary) approach would be to undertake compar-
ative studies of this species in more southerly parts of
its range where it is more widespread and more abun-
dant. Extrapolating from a species’ ecology in one part
of its range to another distant one requires caution as
there may be significant adaptive differences in differ-
ent areas. For example, Sharp-tailed Snakes in Cali-
fornia occupy a wide diversity of habitats (D. Wake,
personal communication), most of which do not occur
in British Columbia. Nonetheless, this approach might
reveal fundamental species’ characteristics that can
be used to delineate more clearly the critical habitats
required for conservation of the Sharp-tailed Snake
in British Columbia.
Acknowledgments
We thank Laura Matthias and Michelle Masselink
for showing us sites known to be occupied by Sharp-
tailed Snakes and Andrea Pilski for assisting in the
field. The Sharp-tailed Recovery Team offered useful
help and advice and both David Wake and David
Spalding provided us with important personal obser-
vations. We also thank various landowners for allow-
ing access to their land: Pender Island — Don and
Theresa Williams, Sylvia O’Reilly, Robert Dill, Jon
Crawford, Pat and Ian Haugh; Saltspring Island —
Frances Hill and Dave Kerman, Sandra and Vince
2007
Knight, Leslie Northey, Mark and Brenda Haughley.
The British Columbia government provided staff time.
The views presented are those of the authors, not their
respective agencies. The Endangered Species Recovery
Fund (World Wildlife Fund, Canada) and the Habitat
Conservation Trust Fund (BC) provided financial
support.
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Received 11 April 2006
Accepted 20 May 2008
Do Repugnant Scents Increase Survival of Ground Nests? A Test with
Artificial and Natural Duck Nests
VANESSA B. HARRIMAN! 4, JusTIN A. Pitt?, and SERGE LARIVIERE?
'Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan S7N SE2 Canada
"Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada
3Cree Hunters and Trappers Income Security Board, Quebec, Quebec G1 V 4K5 Canada
4Corresponding author: vanessa.harriman@ec.gc.ca
Harriman, Vanessa B., Justin A. Pitt, and Serge Lariviere. 2007. Do repugnant scents increase survival of ground nests? A test
with artificial and natural duck nests. Canadian Field-Naturalist 121(2): 150-154.
Ground-nesting birds typically experience high predation rates on their nests, often by mammalian predators. As such, researchers
and wildlife managers have employed numerous techniques to mitigate nest predation. We investigated the use of scents as repel-
lents to deter predators from both artificial and natural ground nests. Survival rates of artificial nests did not differ among six
groups of substances (Wald y?,,_ ;= 4.53, P < 0.48); however the chronology of predation among groups differed. A commercial
Coyote urine based deterrent (DEER-D-TER™), human hair, and Worcestershire sauce were depredated faster than the control
(F, ; = 40.3, P < 0.001). Nest survival of natural nests differed among those groups tested (Wald ren — SSP 01005);
the eight mothball treatment decreased survival (Wald x? ,,_ , = 11.5, P < 0.005), which indicated that novel smells may
attract predators or result in duck nest abandonment when coupled with natural duck scent. Chronologies of predation events
among treatment groups were not different for natural nests (F,, = 1.9, P = 0.22). These findings indicate an interaction
between novel scents and predator olfactory cues.
Key Words: Coyote, Canis latrans, urine, human hair, mothballs, napthaldehyde, nest predation, olfactory cues, waterfowl,
Manitoba.
In the prairies of North America, predation on nests
of ground-nesting ducks is considered the most impor-
tant factor limiting waterfowl populations (Chafloun
et al. 2002). As such, wildlife managers have devel-
oped and employed various techniques in an attempt to
mitigate nest predation with varying degrees of success.
Typically, a predation event by a mammalian predator
such as the Striped Skunk (Mephitis mephitis), Red Fox
(Vulpes vulpes), or Raccoon (Procyon lotor) is oppor-
tunistic (Lariviere and Messier 1997). These predators
are nocturnal and locate nests using audition (when
the hen flushes) and olfaction (Lariviére and Messier
1997). Although substances that may attract predators
to nests have been investigated (Whelan et al. 1994;
Clark and Wobeser 1997), scents that may repel pred-
ators from nests have yet to be tested.
Chemical compounds have been used successfully
to repel animals, including mammals. Some animals
have been effectively repelled by means of conditioned
taste aversion (e.g., Hoover and Conover 1998) and
through the use of noxious scents such as toxic com-
pounds or odor from predatory animals (Sullivan et
al. 1985; Rosell 2001).
We investigated the effectiveness of various scents to
increase survival of duck nests in the Canadian prairies.
Specifically, we tested whether applying DEER-D-
TER™ (a commercial product containing Coyote
(Canis latrans) urine), human hair, Worcestershire
sauce, and mothball treatments could increase survival
of eggs in artificial nests and the capability of DEER-
D-TER and mothball treatments to increase the sur-
vival of eggs in natural duck nests. We also tested for
differences in speed of predation events to determine
whether specific scents may temporarily deter preda-
tors from a nest. Finally, we compared survival of
eggs in artificial and natural nests.
Study Area
This study was conducted southeast of Minnedosa,
Manitoba (50°14'54"N, 99°50'30"W), in an area sup-
porting numerous small (<3 ha) seasonal to permanent
wetlands (ca. 14% of study site) and characterized by
intensive cultivation (63%) of small grains (wheat, bar-
ley, and oats) and oil crops (canola and flax). Inter-
mixed among agricultural fields are small aspen bluffs
composed of poplar species (largely Quaking Aspen
(Populus tremuloides) and Bur Oak (Quercus macro-
carpa). The predator community is diverse and includes
Raccoons, Striped Skunks, Coyotes (Canis latrans), Red
Foxes, American Badgers (Taxidea taxus), and Ameri-
can Mink (Neovison vison). Predation of bird nests in
the area is largely attributed to mammals (Pasitschni-
ak-Arts and Messier 1995) but avian predation also
occurs, primarily by birds such as the Common Raven
(Corvus corax), American Crow (Corvus brachyrhyn-
chos), and Black-billed Magpie (Pica pica).
Methods
The effectiveness of five groups of substances was
compared to a control using artificial nests. The repel-
lent substances were chosen based on unsubstantiated
reports of their efficacy against mammals (human hair
150
2007 HARRIMAN, PITT, AND LARIVIERE: DO REPUGNANT SCENTS REPEL PREDATORS? 15]
10 ie
re 08
S 06 $ 06
g §
3 a
- D
8 0.4 ZS 04
0.2 02
0.0 ue r T T T 7 0.0
cots? ow 5554 ee ow? worn wor Control Coyote urine 8 Mothballs
quo" Treatment
Treatment
FiGurE |. Daily nest survival (all eggs remaining in nest) of
artificial duck nests treated with scents in Manitoba,
2002.
Control
1.2 Koyrcae Coyote urine
———*-—— Human hair
— —-y—-- Worcestershire sauce
1.0 — -#— 4 Mothbalis
8 Mothballs
08
2
© 06 se) oa’ ee <<
S "OR asl
© ~ Oo---0
no ~S
2 0.4 ~- Vy
o
Zz
0.2
0.0
a0 SS ree fe
0 5 10 15 20 25 30
Exposure days
FIGURE 2. Percent of surviving artificial duck nests (all eggs
remaining in nest) treated with scents at X exposure
day in Manitoba, 2002.
and Worcestershire sauce) or from their commercial
claims (DEER-D-TER™ and mothballs). Nests in the
control group were each treated with 0.1 L of marsh
water, which was poured along the outside edge of the
nest. A packet of DEER-D-TER™ was placed on the
outside edge of each nest of the second group. DEER-
D-TER™ consisted of a green plastic packet contain-
ing a mixture of sand and Coyote urine and was adver-
tised to repel deer (Odocoileus spp.) and skunks.
According to the manufacturer, this product was effec-
tive in the field for ca. 90 days. Group three received
one cup of human hair which was collected from three
different salons and was combined to create a homoge-
neous mixture. Group four received 0.15 L of Worces-
tershire sauce applied evenly to each egg and was re-
applied on every subsequent visit. The fifth group
FIGuRE 3. Daily nest survival (all eggs remaining in nest) of
natural duck nests treated with scents in Manitoba,
2002.
125
—®— _ Control
° Coyote urine |
104 lms 8 Mothbalis |
084
2
g 064
oS
oO
2 044
o
z
024 re) 02000
[5 es
0.0 1 ee hea
L er ee. =
0 5 10 15 20 25 30
Exposure days
FIGURE 4. Percent of surviving natural duck nests (all eggs
remaining in nest) treated with scents at X exposure
day in Manitoba, 2002.
received four mothballs double wrapped in cheesecloth
to prevent accidental ingestion by ducks or predators.
The sixth group received eight mothballs, to determine
if a greater amount yielded a different or stronger res-
ponse, with two cheesecloth-wrapped groups of four
mothballs placed on opposite sides of the outside edge
of the nest. Three groups of substances were applied to
natural nests: marsh water, DEER-D-TER™, and eight
mothballs. Methodology and application of treatments
for natural nests followed the same protocol as those
of artificial nests.
Data were collected from 12 June to 7 July 2002.
Artificial nests were created by depositing six medium-
sized chicken eggs in a depression which was then cov-
ered with grass. Nests were marked by a thin welding
rod at a distance of | m. To avoid biases associated with
2
habitat, all artificial nests were deployed in rights-of-
way. Each nest was randomly placed within a 0.8 km
stretch along four different, but intersecting, second-
ary unpaved grid roads. Artificial nests were located
300-1000 m apart on alternating sides of the road. Nests
were placed in the densest vegetation as close as possi-
ble to the randomly selected location to reduce possi-
ble effects of concealment on survival. Artificial nests
were visited every five days for 25 days and were ap-
proached from a different angle each visit to avoid the
creation of a pathway to the nest. Rubber boots and
latex gloves were worn at all times in hopes of con-
cealing human scent. Each nest was randomly assigned
one test substance until an equal number of nests per
treatment was obtained.
Natural nests were located via the drag chain method.
Only upland areas of dense nesting cover were searched
and nests were marked with stakes placed 10 m away
in arandom direction. Nests were located on four sep-
arate quarter sections within 1.6-8.0 kilometers of each
other. Each quarter section consisted of dense nesting
cover and was bordered by either a crop or fallow land
and a secondary gravel road.
Treatments of natural nests were assigned systemat-
ically because we were unaware of how many nests we
would eventually locate and we sought to have a sim-
ilar number of nests per treatment. If, on return to nests
treated with mothballs, any of the mothballs had fallen
out of the cheesecloth, the entire packet was replaced
with a new one. Hens often incorporated the wrapped
mothballs into their clutch and occasionally moved the
packet(s) directly (< 0.5 m) outside of the nest bowl.
In these cases, the wrapped mothballs were left where
the hen placed them. The first re-visit to the nest oc-
curred between five and eight days after initial loca-
tion and treatment application. Subsequent visits were
made every seven days for up to a total of five visits
depending on the fate of the nest. Natural nests were
excluded from the analyses if they were abandoned but
not depredated. However, we were unable to determine
whether a nest had been abandoned and subsequently
depredated.
All nests were deemed depredated (failed) after at
least one of the eggs was removed from or destroyed
in the nest. Mayfield logistic regression (Hazler 2004)
was used to determine whether the nest survival of each
treatment group differed from the nest survival of con-
trol nests within each nest type (artificial or natural).
Because nests were not visited daily the exact date of
failure was unknown. The number of days a nest was
active (“exposure days”) was calculated differently
depending on the fate of the nest. Exposure days of
nests that hatched (or survived to the last day of the
study in the case of artificial nests) was the number of
days that nest was under observation. Exposure days
of failed nests was the number of days the nest was
known to be active plus the midpoint between the last
THE CANADIAN FIELD-NATURALIST
Vol. 121
check and the last known active day. Nests that fail
early are less likely to be detected and the Mayfield
logistic regression reduces this bias by incorporating
exposure days. Also, because this is a logistic-regres-
sion analysis, covariates can be incorporated, allowing
for easy analysis of experimental data (Hazler 2004).
All artificial nests were deployed on the same day and
nest dragging was completed within three days which
minimized the potential impact of seasonality on the
data. An analysis of variance with repeated measures
was used with respect to nest survival as a function of
exposure days to determine whether there was a differ-
ence in the chronology of predation events between
treatments as time progressed. All post-hoc tests were
conducted using an all-pair-wise comparison Tukey
test. Survival of artificial and natural nests was com-
pared using Mayfield logistic regression.
Results
Survival of eggs in artificial nests was higher than
that of natural nests (Wald y? ,,_ , = 43.4, P < 0.001).
Artificial nest daily survival was 0.54 (SE + 0.003)
for all substances combined, while daily nest survival
for natural nests was 0.04 (SE + 0.01) for all substances
combined.
We deployed 120 artificial nests, with each treatment
group containing 20 nests. Due to flooding, seven arti-
ficial nests were lost at various phases of the experi-
ment and were excluded from further analyses. We
found no significant difference in artificial nest survival
rates among treatment groups (Wald 7 Aes =e
P = 0.48; Figure 1).
For all treatments combined, there was a positive
correlation between survival of artificial nests and
exposure days, with survivorship of nests increasing
with time (F, ,,= 6.77, P < 0.001) and the first five
exposure days being the most fatal (Figure 2). Addi-
tionally, the chronology of predation differed among
groups (F, , = 40.3, P < 0.001). Artificial nests con-
taining both four and eight mothball treatments were
depredated similarly to control nests (q = 1.63, p = 6,
P > 0.05;.q = 1.40, p = 6, P > 0.05). However, nests
treated with DEER-D-TER™ (q = 9.14, p= 6, P < 0.05),
human hair (q = 4.84, p = 6, P < 0.05), and Worcester-
shire sauce (q = 13.9, p= 6, P < 0.05) were depredated
more quickly than control nests.
In total, 70 natural duck nests were located, with 24
receiving the control, 23 treated with DEER-D-TER™,
and 23 treated with eight mothballs. Natural nest sur-
vival of the treatment groups differed from that of the
control (Wald x? ,,_5= 11.82, P < 0.005; Figure 3)
with survivorship of nests treated with eight moth-
balls being significantly lower (Wald x? ,,_ , = 11.47,
P < 0.005).
Nest survival did not vary as a function of exposure
days for all natural duck nests combined (F, , = 2.14,
P =().17) despite a notably rapid decrease in nest sur-
2007
vival during the first five exposure days (Figure 4). Nest
survival did not vary as a function of exposure days
among treatments for natural duck nests (F, , = 1.99,
P= 22):
Discussion
We assessed the effect of various scent treatments on
the survivorship of artificial and natural duck nests in
the prairie pothole region of Canada. The application
of scents did not increase nest survival of artificial or
natural nests. Addition of mothballs decreased survival
of natural nests. These results not only indicate that
DEER-D-TER™, human hair, Worcestershire sauce,
and mothballs were ineffective at repelling predators,
but that certain scents may actually attract predators
and/or increase duck nest abandonment rates.
The chronology of predation differed among treat-
ments for artificial nests but not for natural nests. Con-
trary to expectations, artificial nests treated with DEER-
D-TER™, human hair, and Worcestershire sauce were
depredated more quickly than control nests. Again these
results may indicate a neophilic predator response. In
particular, human-associated scents (i.e., hair and food
items) may elicit predator attraction depending on the
study site and the animal’s experience with people. Al-
though this study took place in a rural location, it is
plausible that local predators may have positive asso-
ciations with human dwellings and food, especially
opportunistic species such as Striped Skunks and Rac-
coons (Pitt et al. 2008). Raccoons in particular often
rely on anthropogenic foods at the northern edge of
their range distribution (Lariviére 2004; Pitt et al.
2008). Additionally, DEER-D-TER™ may have also
attracted predators to nests, especially con-specifics.
Mothballs and hair regularly went undisturbed at de-
predated nest sites whereas urine packets were often
chewed on and/or ripped open.
The result of predators exhibiting neophilic tenden-
cies may have important implications for waterfowl
due to the propensity for which they are studied. The
concealment of human scent should be an important
consideration for researchers. Neophilic predators may
increase their search radius when a particular scent cue
is present (Whelan et al. 1994), possibly rendering
human-visited nests more susceptible to predation.
Our study indicates that the survival rates of artificial
and natural duck nests in the prairie pothole region are
different and this may indicate that these nests are
viewed differently from the perspective of a predator.
Artificial nests realized higher survival rates compared
to natural nests, which has been previously document-
ed (Guyn and Clark 1997). The disproportionate dif-
ference in survival between artificial and natural nests
could also in part be attributed to the presence of scents
associated with natural nests, as well as the presence
of a hen and the ability to cue in on the nest after a flush
event (Lariviere and Messier 1997). Alternatively, the
HARRIMAN, PITT, AND LARIVIERE: DO REPUGNANT SCENTS REPEL PREDATORS?
153
observed difference may have been mediated by uncon-
trolled factors affecting natural nest survival, such as
nest abandonment, hen species and behavior, nest con-
cealment, and variation in nest spacing. Specifically,
the eight mothball treatment may have provoked aban-
donment by nesting female ducks as some ejected
mothballs from nests.
Conditioned taste aversion has been effective in
deterring predators from nests (Nicolaus 1986; Con-
over 1990). Napthaldehyde (the noxious substance in
mothballs) has been found to be effective in causing
adverse responses by Coyotes when ingested (Hoover
and Conover 1998). After repeated exposure, Coyotes
avoided eggs injected with napthaldehyde. Although
highly successful, it is expensive and time consum-
ing to set up nests with injected chemicals. For future
studies, it may be equally effective and less time con-
suming to combine the use of conditioned taste aver-
sion with scents. Our results indicate that applied by
themselves, these scents do not increase survival of
either artificial or natural duck nests.
Acknowledgments
Delta Waterfowl Foundation provided funding and
assistance for this study. Ducks Unlimited Canada kind-
ly granted permission to search nesting cover. D. Coul-
ton, H. Remenda, J. Estrella provided assistance in the
field. Two reviews contributed to the clarity of the man-
uscript. This research was approved and conducted under
Manitoba Wildlife Animal Care Committee #2002-10.
SL was supported through a National Science and
Engineering Research Council discovery grant.
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THE CANADIAN FIELD-NATURALIST
Vol. 121
Sullivan, T. P., L. O. Nordstrom, and D. S. Sullivan. 1985.
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Received 8 May 2006
Accepted 11 April 2008
Translocation and Recovery Efforts for the Telkwa Caribou, Rangifer
tarandus caribou, Herd in Westcentral British Columbia, 1997-2005
ASTRID VIK STRONEN!?, PAUL PAQUET!, STEPHEN HERRERO!, SEAN SHARPE®, and NIGEL WATERS*
'Faculty of Environmental Design, University of Calgary, 2500 University Drive NW, Calgary, Alberta T2N 1N4 Canada
*Present Address: Department of Biology, University of New Brunswick, Fredericton, New Brunswick E3B 6C2 Canada:
Mailing Address: 6533 rue de Normanville, Montreal, Quebec H2S 2B8 Canada; e-mail: astrid.stronen@unb.ca
‘Ministry of Environment, #325, 1011 Fourth Avenue, Prince George, British Columbia V2L 3H9 Canada
‘Department of Geography, George Mason University, 4400 University Drive MS 1E2, Fairfax, Virginia 22030 USA
Stronen, Astrid Vik, Paul Paquet, Stephen Herrero, Sean Sharpe, and Nigel Waters. 2007. Translocation and recovery efforts for
the Telkwa Caribou, Rangifer tarandus caribou, herd in westcentral British Columbia, 1997-2005. Canadian Field-
Naturalist 121(2): 155-163.
During 1997-1999, 32 Woodland Caribou (Rangifer tarandus caribou) were translocated from the Sustut Herd to the Telkwa
Mountains in westcentral British Columbia to augment recovery of the Telkwa Caribou Herd. The animals were fitted with
radiocollars and located during 1997-2000 to determine selection of habitat features and terrain variables. Six Caribou calves
were also collared to determine causes and timing of calf mortality during summer 1999. Defining available habitat for newly
translocated animals is often arbitrary and subjective, and we based the analyses on ranks for habitat use and availability as
this is less sensitive to the inclusion or exclusion of a questionable resource. This method represents some loss of information
but provides indications of the relative importance of various habitat types without classifying any as avoided. High elevation
habitat (> 1700 masl) on moderate slopes (16 — 45°) received the highest ranks, as did “warm” (136 — 315°) aspects and forests
> 250 years old. Three calves died shortly after birth. One calf appeared to have been killed by predation, likely by a Golden
Eagle (Aquila chrysaetos), and one calf was abandoned by the cow. Cause of death for the third calf is unknown. To assess
habitat use associated with calving we compared summer locations with data obtained throughout the rest of 1999 for eight
cows with calves and eight without calves. We found significant difference in use of elevation during calving time, when cows
with calves remained at high elevations and barren cows generally descended to lower elevation habitat. Surveys conducted
in 2005, five years after the completion of the initial study, produced a count of approximately 90 Caribou. This suggests
that in the short term, the translocation was successful in re-establishing a self-sustaining Caribou population in the Telkwa
- Mountains.
Key Words: Woodland Caribou, Rangifer tarandus caribou, calf mortality, calving habitat, habitat selection, rank analysis,
wildlife management, British Columbia.
Woodland Caribou (Rangifer tarandus caribou), pop-
ulations have generally experienced decline and range
reduction over the last century in British Columbia
(Heard and Vagt 1998), across Canada (COSEWIC
2000*) and throughout the circumpolar region (Mal-
lory and Hillis 1996). Causes of decline include a com-
bination of habitat loss, habitat fragmentation, increased
| human access, hunting and predation (Bloomfield 1980;
Edmonds and Bloomfield 1984; Heard and Vagt 1998),
_ poaching (Edmonds 1988), and possible interactions
_ between disturbance and predation (Mallory and Hillis
1996). Woodland Caribou in the Southern Mountain
_ National Ecological Area of British Columbia (includ-
| ing the Telkwa Caribou Herd) were designated as
| threatened in 2000 (COSEWIC 2000*). Caribou were
once widely distributed throughout the mountainous
areas surrounding Telkwa and Smithers, British Col-
umbia, and might have formed part of a larger herd
_ whose range extended south into Tweedsmuir Provin-
cial Park (TCHRP 1998*). Few data are available on
_ the Telkwa Caribou Herd before 1997; however, sur-
yveys indicated fluctuations in herd size from less than
100 in 1949 to a high of 271 in 1965. A hunting clo-
sure was implemented for the herd in 1973 but only
68 animals were counted in 1984 (van Drimmelen
1986*) and monitoring flights in the 1990s resulted in
counts of less than 20 Caribou (TCHRP 1998*). During
1997-1999, 32 Caribou were captured from the Sustut
Herd about 150 km north of Smithers and translocated
to the Telkwa Mountains to help the recovery of the
Telkwa Caribou Herd.
Translocation is now a common tool in the recovery
of populations at risk (Griffith et al. 1989) and the need
for documenting habitat use and availability is critical
in efforts to preserve endangered species and manage
small populations (Manly et al. 1993). We studied
habitat use and calf survival of the Telkwa Caribou
Herd from 1997 to 2000 to determine preliminary habi-
tat selection of translocated animals and assess timing
and causes of calf mortality. With small populations
following a translocation, there are usually limited data
available to investigate habitat selection, and therefore
little or no information for land managers. Although
defining available habitat for newly translocated ani-
mals is often arbitrary and subjective, a preliminary
evaluation of habitat selection can provide some infor-
155
156
mation on priority areas for protection when assess-
ing other land uses such as forestry and recreation.
Translocation success can be evaluated in three phases:
(1) initial occupation and reproduction in the new eco-
system; (2) presence of a short-term self-sustaining
population, and (3) ultimate population recovery
(Compton et al. 1995). We outline findings on habitat
selection and calf mortality between 1997 and 2000
and report for phases | and 2 of the Telkwa Caribou
Herd Recovery Project. We also discuss challenges
involved in providing habitat management priorities
based on preliminary data from a small reintroduced
population.
Study Area
The study area comprises approximately 25 000 km?
in westcentral British Columbia, with elevations rang-
ing from 300-3000 meters above sea level (Figure 1).
It is situated between 53.40 and 55.20 degrees of lati-
tude north and 125.10 to 128.30 degrees of longitude
west. Glaciers, rugged peaks, high alpine plateaus,
and deep glaciated U-shaped valleys with extensive
wetlands characterize the alpine areas. Major tree spe-
cies in the study area include Subalpine Fir (Abies
lasiocarpa), Engelmann Spruce (Picea engelmannii),
and Hybrid White Spruce (Picea glauca x engelman-
nii). The climate has both coastal and inland influ-
ences, with cool summers and moderately cold winters.
Precipitation averages 500 mm per year, of which
about half falls as snow. The Telkwa Mountains have
a population of approximately 250 Mountain Goats
(Oreamnus americanus), and lower elevations have
populations of Moose (Alces alces), Mule Deer (Odo-
coileus hemionus hemionus), and some Elk (Cervus
elaphus). Large predators within the range of the Telk-
wa Caribou Herd include: Grizzly Bears (Ursus arc-
tos), Black Bears (Ursus americanus), Wolverine (Gulo
gulo), Wolves (Canis lupus), Coyotes (Canis latrans),
and Golden Eagles (Aquila chryseatos). To account
for the difficulty in defining available resources for
the herd and thus delineating our study area, where
most animals have had < 2 years of experience, we
selected an area of available habitat based on three
factors: (1) locations of collared caribou 1997-1998
(G. Schultze, British Columbia Ministry of Environ-
ment, personal communication); (2) Traditional Eco-
logical Knowledge of the Telkwa Caribou Herd (B.
Holland, Wet’suwet’en First Nation, personal commu-
nication in Stronen 2000) and (3) availability of forest
cover data from the British Columbia Ministry of
Forests (Inventory Branch, Victoria, scale 1: 20 000).
Methods
We located radiocollared animals throughout the
year from fixed wing aircraft (Cessna 206, 185, or 172)
using standard telemetry equipment. Tracking intervals
varied from every four days to every three weeks, de-
pending on funding, season, and weather conditions.
THE CANADIAN FIELD-NATURALIST
Vol. 121
We determined geographic location using yoke-mount-
ed, handheld, or panel-mounted Global Positioning
System (GPS) units and noted habitat characteristics,
activity, group size, and presence of young. The accu-
racy of relocation data was tested and averaged + 300
meters. Most locations were determined during morn-
ing or mid-day. For the purpose and scale of this study,
the data were assumed representative of Caribou habi-
tat use for day and night. Six calves were captured
shortly after birth and equipped with radiocollars dur-
ing the summer of 1999. Caribou calves were captured
as close to within 24 hours of birth as possible using
a Bell 206 Jet Ranger and a Long Ranger helicopter and
a capture crew of three people. Calves were sexed,
weighed, and fitted with VHF radiocollars (Lotek En-
gineering, Newmarket, Ontario) with motion sensitive
mortality sensors. We estimated age by assessing the
presence/condition of the umbilical cord, condition of
hooves and pelage, posture, and general appearance
(Haugen and Speake 1958; Miller et al. 1988), and
calculated weight by subtracting 0.6 kg for each day of
estimated age (Adams et al. 1995). Handling time was
approximately five minutes from capture to release.
Calves were monitored daily, weather permitting, from
date of capture until the end of August. During Sep-
tember and October, we monitored calves weekly and
after October they were monitored as a part of regular
telemetry flights for the herd. We investigated mortal-
ity signals via helicopter the day of discovery.
Data for elevation (meters above mean sea level),
slope (%), and aspect (in azimuth measured from north)
were derived from a Digital Elevation Model using
Arc/INFO v. 7 software. Tiles from the Digital Eleva-
tion Model were merged to form a seamless grid cov-
ering the study area. Floating point grids were gener-
ated for slope and aspect and we created a variable
combining elevation and slope. Data on habitat type
(leading tree species or non-productive timber area),
crown closure (%), and tree age (using leading tree spe-
cies) were derived from Forest Cover Inventory Poly-
gons (British Columbia Ministry of Forests, Inventory
Branch, Victoria, scale 1: 20 000). Values for terrain
and habitat variables were summarized (km?) for the
entire study area to determine percent availability of the
various classes of each variable. Habitat availability
was analyzed by the British Columbia Ministry of En-
vironment’s regional GIS data analyst, who also pro-
duced cartographic output. We used data collected
between November 1997 and April 2000 to analyse
habitat use and determined terrain and forest cover
values for all telemetry locations. To assess habitat sel-
ection, we followed sampling design 2 of Manly et al.
(1993), where individual animals were identified and
the use of resources was measured for each individual,
but availability was measured at the population level.
To assess habitat use associated with calving, we
used locations for eight cows with calves (n = 123 loca-
tions) and eight without calves (n = 128 locations)
2007
200 0
STRONEN, PAQUET, HERRERO, SHARPE, and WATERS: TELKWA CARIBOL 157
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200 Kilometers
FiGuRE |. The study area in British Columbia, 1997 — 2000.
collected during the summer of 1999. We compared
summer data (n = 150 locations) with locations ob-
tained during the rest of the year (n = 145 locations)
to account for differences not attributed to calving.
Locations were divided into two groups:
(1) calving (25 May — 6 July)
(2) all-year (1 January — 24 May, and 7 July — 31
December)
We analyzed habitat selection using Resource Sel-
ection for Windows (RSW), a 32-bit Windows program
for analyzing preference (Leban 1999). Using Johnson’s
(1980) method, which orders all variable components
by rank for usage and availability, we measured habitat
preference for each animal as the difference between
the rank of usage and the rank of availability. We aver-
aged these differences across animals to obtain a mean
value for each habitat component and then compared
the values to determine a ranking from least preferred
habitat component to most preferred. We tested for
habitat selection using alpha = 0.05. We compared
habitat use of reproductive cows and cows without
calves using analysis of variance (repeated measures,
alpha = 0.05) within each group and within each sea-
son using SPSS 10.0 (SPSS Inc. 1999). The herd con-
sisted primarily of translocated Caribou during the
study and we considered the radio-collared animals
to be a random sample of the Telkwa Herd.
Results
High elevation habitat on moderate slopes (> 1700
meters above sea level, 16 — 45%) received the highest
ranks (Table 1). Caribou also made extensive use of
the steep rugged mountains in the Telkwa range
(>1700 m, > 45%). Aspects 136 — 315° received the
highest rank (Table 2). The rank values for aspect ap-
pear to contradict the information on use versus avail-
ability. However, data on use report only overall percen-
tage of use, and taking behavior of individual animals
into account; the 136 — 315° aspects receive the highest
rank. Less than 4 of locations (405 of 1424) were in
forested habitats (Table 3), and of these locations,
forests > 250 years old received the highest rank.
For analysis within groups, we found a significant
difference in mean elevation between calving time and
the rest of the year for cows without calves, but not
for cows with calves. For analysis within season, we
found a significant difference in mean elevation for
calving time but not for the rest of the year. Although
sample sizes were smaller for calving time, both groups
had a narrower range in elevation during calving and
individual differences in elevation use were also high-
er during calving than the remainder of the year (Fig-
ures 2 — 5). Figure boxes represent the interquartile
range containing 50% of the values, with a line show-
ing the median value. Error bars represent the highest
and lowest values, excluding outliers (extreme values).
Most cows gave birth in alpine locations, in exposed
areas free of snow. Of the 16 cows sampled, only cow
#151 made extensive use of elevations below 1400
meters throughout the year (Figure 4).
Calving time for Telkwa Caribou ranged from 26
May until 10 June, and six Caribou calves were col-
lared between 3 June and 10 June 1999. Five were
males born to transplanted Caribou cows and one was
a female calf born to the only resident collared cow.
Average birth weight was 8.4 kg (range 6.9 — 9.4 kg).
Three of the six calves died shortly after birth. One
calf likely died from abandonment, whereas cause of
158
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Use, availability and rank of classes for elevation and slope. Telkwa Mountains, British Columbia, Canada
1997-2000. N = 1424
Elevation (m) Slope (%) Class Locations
<1100 0-15 1 116
<1100 16-45 2 38
<1100 >45 3 4
1100 — 1700 0-15 4 166
1100 — 1700 16-45 5) 303
1100 — 1700 >45 6 168
>1700 0-15 7 144
>1700 16-45 8 313
>1700 >45 9 172
% use % available Rank
8.15 38.43 9
2.67 20.75 8
0.28 6.40. 7
11.66 17.98 6
21.28 9.71 4
11.80 3.00 5
10.11 2.18 3
21.98 1.18 1
12.08 0.36 2
TABLE 2. Use, availability and rank of classes for aspect. Telkwa Mountains, British Columbia, Canada 1997—2000. N =
136 — 315 1 627
14162
Aspect (azimuth) Class Locations
316 — 135 D) 789
% use % available Rank
44.03 47.24 1
55.41 48.23 2
4 Areas of zero aspect (usually where relocation error positioned animals in water) are excluded.
TABLE 3. Use, availability and rank of classes for forest age. Telkwa Mountains, British Columbia, Canada 1997-2000. N = 405°
8.47 4
Forest age (years) Class Locations
1-40 I 18
41 — 100 D 84
101 — 140 3 105
141 — 250 4 161
>250 5 Sy)
> Including only locations below tree line (1700 m).
death for the second calf appeared to be avian preda-
tion, most likely Golden Eagle. Cause of death for the
third calf was unknown. The calf had remains of what
appeared to be milk curds in the stomach and small
amounts of vegetation. However, membranes inside the
hide appeared dry and indicated dehydration. No signs
of predation or scavenging were evident. We found
adult tracks, likely made by the mother, circling the
carcass. During the first year after the transplant (1998),
we observed five calves in the herd. The following year,
we saw one yearling in the herd, as well as 12 new
calves and one cow with an extended udder, indicat-
ing she recently had given birth and lost her calf. We
observed eight calves during the 1999 October calf
count. Recent surveys of the herd found 16 cows with
calves in October 2003 and 23 the following year. In
October 2005 ten cows with calves were counted. How-
ever, the decrease in cow/calf numbers from 2004 to
2005 may reflect fewer functioning radio collars in the
herd during 2005, thus increasing the probability that
some Caribou groups went undetected (G. Schultze,
personal communication).
% use % available Rank
1.26
5.90 17.82 3
7.37 14.85 Dp,
11.31 24.60 5
2.53 7.08 1
Discussion
The Telkwa Caribou Herd appears to select high
elevation habitat with moderate slopes, which corre-
sponds with earlier observations (van Drimmelen
1986*; Bill Holland, personal communication in Stro-
nen 2000). The herd uses gentle to moderate slopes
throughout much of the year, which is common for
Woodland Caribou (Edmonds and Bloomfield 1984;
Seip 1990*; Terry et al. 1996; Wood and Terry 1999).
However, use of steep slopes is also reported (Hatler
1987*; Terry et al. 1996). Woodland Caribou have been
found using southern/warm aspects (Edmonds and
Bloomfield 1984; Warren et al. 1996), northern/cool
aspects (Warren et al. 1996) or to show no apparent
selection (Seip 1990*). Use also varies between years
and seasons (Terry et al. 1996; Wood and Terry 1999).
When in forested habitat the Telkwa Caribou Herd
appears to select old forests, which corresponds with
other studies (Cichowski 1993; Terry et al. 1996; Wood
and Terry 1999; Apps and Kinley 2000*). Woodland
Caribou in British Columbia have been classified into
two ecotypes (Stevenson and Hatler 1985). The north-
:
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CARIBOU COWS
STRONEN, PAQUET, HERRERO, SHARPE, and WATERS: TELKWA CaArisot
159
610
FiGuRE 2. Barren cows’ use of elevation all-year, 1999. Telkwa Mountains, British Columbia.
‘
4s
DERIVED ELEVATION
Ke 1¢ 1 10
004 o71 272
CARIBOU COWS
i.
.
i"
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283 291 177 610 801
FiGure 3. Barren cows’ use of elevation at calving time, 1999. Telkwa Mountains, British Columbia.
er ecotype (including the Telkwa Caribou Herd)
depends primarily on terrestrial lichens during winter
and the mountain ecotype depends more on arboreal
lichens. Arboreal lichens are most abundant in old for-
est (Apps and Kinley 2000*), often on trees > 120 years
old (Stevenson and Hatler 1985; Wood and Terry 1999).
We found the Telkwa Caribou Herd in forested habitat
only one-third of the time. However, divisions between
arboreal- and terrestrial lichen-dependent groups are
not absolute. The temporal impacts of icing, deep snow,
and other environmental conditions may at times pre-
vent Caribou from obtaining terrestrial lichens, making
areas of arboreal lichens provisionally more impor-
tant (van Drimmelen 1986*; Edmonds 1988: Cichow-
ski 1993). Forests 141—250 years old received the high-
est use but the lowest rank. This is the most abundant
forest age class (Table 3). Whereas the small number
of Caribou during this study did not use this age class
160
in proportion to availability, the low rank does not equal
avoidance or lack of importance. Future monitoring
could also help clarify whether certain areas within
this forest age class are locally important.
The difference in elevation use between barren and
reproductive cows during calving season might be due
to reproductive status. There are, however, wide dif-
ferences within each group. As four of eight barren
cows remained at high elevations (which presumably
meant forgoing nutritional opportunities, at least in
spring/early summer), this could be an antipredator
strategy employed regardless of reproductive status.
Based on this study, possible antipredator tactics for
protecting a calf can not be distinguished from those
that cows seem to use for their own protection. Ed-
monds and Smith (1991) reported that cow/calf pairs
localised movement to relatively small geographic
areas in early June (1-15) and did not increase the size
of their ranges until after July. The Telkwa Caribou
displayed similar behaviour, as cow/calf pairs in the
Telkwa Herd remained relatively stationary until the
middle of July. Subsequently, some pairs made long
distance movements (one pair moved 50 km over a
few days). The calving time for Telkwa Caribou cor-
responded to that reported by Edmonds (1988) for
westcentral Alberta. As only one birth from a resident
Caribou was recorded (10 June), it was not possible to
examine any differences between transplant and resi-
dent animals.
Some cows in the Telkwa Caribou Herd appeared
more sensitive to fixed-wing aircraft during the first
couple of weeks post-calving than during the rest of
the year, but we noted a wide range in behaviour.
Cichowski (1993) reported that most adult females
were found alone on ridge tops during calving. Two
Telkwa cows calved on the top of a rocky ridge and
were joined by two other cows with calves in the mid-
dle of June. The ridge had few access points but bio-
physical information suggests low potential for lichen
production. All four calves were found alive in Octo-
ber. The use of ridges may be a trade-off between pred-
ator avoidance and the availability of abundant high
protein food (Bergerud and Miller 1984). Cichowski
(1993) found that most calving sites were within 10 km
of previous calving sites and Seip (1990*) reported
many Caribou use the same calving locations each year.
We could not analyse fidelity to calving areas within
the Telkwa Herd and it would take several years before
this could be assessed. Caribou calving areas could
change from year to year as an antipredator strategy
(Bergerud and Miller 1984), and Valkenburg et al.
(1988) warned against making management decisions
based on Caribou calving areas located during narrow
time intervals. However, previously known calving
areas identified by the Traditional Ecological Knowl-
edge of the Wet’suwet’en First Nation (B. Holland,
in Stronen 2000) should be considered when making
management decisions, as this information is based
THE CANADIAN FIELD-NATURALIST
Vol. 121
on years of observation and would indicate fidelity to
specific calving sites.
Small sample size precluded analysis of calf mortal-
ity for the Telkwa herd but suggested that avian preda-
tion was a factor. Recruitment data from November
1998 to April 2000 seemed to indicate that low pro-
duction of calves was a factor of equal or higher im-
portance than the loss of calves. If the low recruitment
of calves was due to calf predation, a higher frequen-
cy of Caribou cows with extended udders but without
calves would be expected. Females without calves
commonly shed their antlers in March and April and
exhibit fresh antler growth during the calving period,
whereas pregnant females shed their antlers near calv-
ing in May and June (Bergerud 1980). During our sur-
veys at calving time, many cows exhibited fresh antler
growth, indicating that they were not expecting a calf
that year. The Caribou transplanted from the Sustut Cari-
bou Herd would be expected to show pregnancy rates
comparable to those of other populations when they
arrived, and reported pregnancy rates usually range
from 80% to 90% (Bergerud 1980; Seip 1990*; Ed-
monds and Smith 1991). It is possible that stress relat-
ed to capture, immobilization, and transplant may have
caused abortions or fetal absorptions, or that a large
number of the cows were by chance not pregnant. In
addition, there may not have been enough bulls to
achieve normal pregnancy rates for the Telkwa herd
during the first years. At least one resident mature bull
was present in the herd during 1998 but due to difficul-
ties associated with movement and collaring of adult
bulls only younger males were translocated to the Tel-
kwa Mountains.
Human activities such as recreation and resource
extraction may also influence Caribou habitat use. Van
Drimmelen (1986*) expressed trepidation over snow-
mobiling on the winter ranges of the Telkwa Herd
and this has been a concern for over 25 years (Bustard
1977*). The presence of dogs may also cause Caribou
to avoid otherwise useful habitat. This is particularly
important during calving time, when cows may be
extra sensitive to disturbance (Bergerud and Miller
1984). Need for information related to species translo-
cations and/or small remnant populations will increase
in the future. Where habitat availability is poorly esti-
mated it may be particularly important not to label
habitats and resources, which may be important in the
long term, as avoided. We chose Johnson’s (1980)
method because it is relatively insensitive to decisions
on what constitutes available habitat for the Telkwa
Caribou Herd. If habitat components are used in pro-
portion to availability, the rank ordering of use and
availability will be the same. However, the converse is
not necessarily true; the use and availability propor-
tions could be different across habitat components,
while still having the same rank, thus resulting in high
Type 2 errors (where a difference in proportional
selection is not detected) (Alldredge and Ratti 1986,
:
:
|
2007
DERIVED ELEVATION
co
o
3
CARIBOU COWS
STRONEN, PAQUET, HERRERO, SHARPE, and WATERS: TELKWA CARIBOI 16]
FIGURE 4. Reproductive cows’ use of elevation all-year, 1999. Telkwa Mountains, British Columbia.
2200
2000
1800
1600
1400
1200
1000
800
DERIVED ELEVATION
024 O32 034
CARIBOU Cows
151 251 770 256 $70
FiGuRE 5. Reproductive cows’ use of elevation during calving time, 1999. Telkwa Mountains, British Columbia.
1992). A bias may also occur against small habitat
patches when the size of error area for locations [here
+ 300 m] is independent of habitat patch size (Rettie
and McLoughlin 1999). Many wetlands in our study
area were too small to be captured in Forest Cover data
and Houwers (1996*) reported that distribution of ar-
boreal lichens in the Telkwa Mountains was patchy.
These two habitat units, too small to be noted by our
analysis, may be vital for long-term survival of the
Telkwa Caribou Herd. The size of habitat required by
Caribou to avoid predation may be significantly greater
than the amounts required to obtain sufficient forage
(Stevenson et al. 1994*: Seip and Cichowski 1996),
and requirements for predation avoidance could be
10 times larger (Bergerud 1980). It is theorized that
the Telkwa Mountains could support approximately
162
300 caribou (van Drimmelen 1986*). Given sufficient
habitat and calf survival the herd could recover to
historic numbers. At present, the translocation appears
to have been successful in establishing a self-sustaining
Caribou herd in the Telkwa Mountains.
Acknowledgments
The Habitat Conservation Trust Fund, Forest Ren-
ewal BC, the British Columbia Ministry of Environ-
ment, and a Graduate Research Scholarship from the
Faculty of Environmental Design, University of Cal-
gary, provided funding for this study. Staff at the Bri-
tish Columbia Ministry of Environment in Smithers
assisted with fieldwork, and G. Schultze provided
survey data from 2000 to 2005. We thank J. Warren
and other staff at the GlIS-section for help with all
GIS-related work. Northern Lights Air piloted fixed-
wing telemetry flights, and Canadian Helicopters pilot-
ed flights to capture caribou calves and investigate
mortalities. An anonymous reviewer provided helpful
comments on the manuscript.
Documents Cited (marked * in text)
Apps, C., and T. Kinley. 2000. Multi-scale habitat associa-
tions of Mountain Caribou in the Southern Purcell Moun-
tains, British Columbia. East Kimberley Environmental
Society, Kimberley British Columbia and Crestbrook
Forest Industries Ltd., Cranbrook, British Columbia.
Bustard, D. 1977. Caribou observations — Hunter’s Basin —
Camel Humps Area. Fish & Wildlife Branch memorandum,
Ministry of Environment, Lands and Parks, 14 March,
LOTT:
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An Inventory of the Aquatic and Subaquatic Plants in SASK Water
Canals in Central Saskatchewan, Canada, Before and After the
Application of the Herbicide Magnacide
J. HUGo CoTa-SANCHEZ! and KIRSTEN REMARCHUK
Department of Biology and Herbarium of the University of Saskatchewan, University of Saskatchewan, 112 Science Place,
Saskatoon, Saskatchewan S7N 5E2 Canada
' Corresponding author: hugo.cota@usask.ca
Cota-Sanchez, J. Hugo, and Kirsten Remarchuk. 2007. An inventory of the aquatic and subaquatic plants in SASK Water canals in
central Saskatchewan, Canada, before and after the application of the herbicide Magnacide. Canadian Field-Naturalist
121(2): 164-167.
This study focuses on the floristic composition of aquatic and semi-aquatic plants in the SASK Water canal system and their
potential effect on irrigation systems. A checklist, evaluation, and synthesis of the species identified in this survey before and
after the application of the herbicide Magnacide are provided, in addition to a brief discussion of the environmental effects of
Magnacide. Thirty-three species in 26 genera within 20 plant families were identified. Two unidentified green algae were also
collected. Common aquatics (i.e., green algae, Potamogeton spp., Alisma gramineum, A. plantago-aquatica, Ceratophyllum
demersum, and Myriophyllum sibiricum) combined with debris from terrestrial plants were the primary contributors to block-
age of irrigation drains. In general, the concentration of Magnacide used in this study had a minor effect on aquatic plant
diversity, but effectively reduced plant density. However, the long-term effects of pesticides on the surrounding aquatic and
terrestrial environments of the SASK Water irrigation system are unknown.
Key Words: aquatic plant inventory, environmental effect, SASK Water canals, Magnacide, acrolein, Saskatchewan.
Plants are vital to the function of aquatic ecosystems
for their role in providing food, oxygen, and habitat for
other organisms. However, floating and submerged
vegetation along with debris from terrestrial plants
may be problematic in different types of water bodies.
For example, in shallow lakes, profuse plant growth
creates dense mats that prevent the regular movement
of watercraft. In addition, aquatic plants, including
algae, can significantly reduce the aesthetic value of
water bodies. Furthermore, rapid growth of vegetation
and deposition of debris in irrigation canals can result
in a decline in water flow rates and a subsequent in-
ability to supply water to crops. Hence, the removal of
selected aquatic species via physical or chemical meth-
ods may be required to improve ecosystem function,
aesthetic value, and availability for human consump-
tion.
Aquatic plants can be placed into four broad cate-
gories: (1) algae, (2) floating-leaved, (3) submerged, and
(4) emergent plants (Shelton and Murphy 1989). Of the
four categories listed above, algae are of the greatest
concern in canal systems because of the likelihood of
interfering with water flow. Algae grow profusely in
slow moving, shallow water. Floating-leaved and sub-
merged plants are rooted, with the foliage at or below
the water surface. Flowing water or disturbance, such
as wind, can uproot plants leading to subsequent ob-
struction of water flow. Finally, emergent species typ-
ically grow in shallow, fluctuating water, such as drain-
age ditches, canals, rivers, periphery of water bodies,
and ponds. Plants in this category are of minor concern
in canal systems and in some cases may prevent non-
aquatic plant debris from entering the system.
Aquatic species impede water flow by clogging
gates, intake screens, valves, and pumps in irrigation
and drainage channels (Lancar and Krake 2002). As a
consequence, localized flooding may result under high
water conditions. In addition, Holm et al. (1969) indi-
cate that abundant vegetation in canals can lead to an
excessive loss of water through evapotranspiration.
pondweed (Potamogeton L.), Milfoil (Myriophyllum
sibiricum), Canada Waterweed (Elodea canadensis),
wild celery (Vallisneria L.), and American Eel-grass
(Zostera marina L.) are a few of the problematic coarse
or large species in shallow aquatic environments (Lan-
car and Krake 2002).
Of particular interest to this study are aquatic taxa
that interfere with canal irrigation systems in central
Saskatchewan. Accelerated growth of plants in SASK-
Water canals restricts the amount of water available for
irrigation. Given the clogging effect of aquatic plant
growth on water flow, agricultural canals were treated
with Magnacide, a volatile algaecide and aquatic her-
bicide, in which the active ingredient is acrolein (Score-
card 2006*). Acrolein contains two functional groups,
a reactive double bond and an aldehyde group (Nor-
done et al. 1996) and is, in turn, toxic to some organ-
isms (Albarifio et al. 2007). However, it is widely used
in agricultural canals and water bodies as an herbicide
to control aquatic weeds (Burland et al. 1984; San-
godoyin and Smith 1996). The use of high concentra-
tions of acrolein could pose a significant risk to aquat- |
ic environments (Nordone et al. 1996); therefore, the
controlled application and use of Magnacide and other
164
2007
herbicides is highly recommended. Previous studies
have shown that toxic residues deposited in soils and
water bodies have detrimental effects in the life cycles
of local flora and fauna (Sangodoyin and Smith 1996).
Thus, even though several sources, e.g., Sangodoyin
and Smith (1996); Nordone et al. (1996); Albarino et
al. (2007), claim that the transient use of acrolein in
agricultural waters at minimal amounts has no negative
impact in natural aquatic environments, rigid control
must be enforced to prevent ecological damage.
In this study, we investigated the floristic composi-
tion of aquatic and semi-aquatic plants in the SASK-
Water canal system and their potential effect on ir-
rigation systems. Our study is a contribution to the
knowledge of Saskatchewan aquatic and sub-aquatic
flora and provides a preliminary assessment and syn-
thesis of the species identified prior to and after the
application of Magnacide. The potential effect of
Magnacide on aquatic flora is also discussed.
Methods
Compared to most wetland inventories, this survey
covers a limited geographic area and a restricted range
of wetland flora. The site selection and sampling strat-
egy were designed to survey all major areas of the
SASK Water canal system to be evaluated in terms of
aquatic vegetation within the Saskatoon Southeast Water
Supply Project Location Plan (Brightwater-Blackstrap-
Zelma Reservoir areas; 51°25' to 52°55'N, 106°15' to
107°00'W). This system irrigates an estimated area of
20 120 acres. The survey sites include locations of
Magnacide application by SASKWater personnel at
intake screens. Acrolein was directly applied to water
in strategic sites to control submersed, floating and
emergent aquatic weeds. The herbicide was applied at
a concentration of 2.5-5.0 ppm/30 minutes every two
weeks from June through August 2005 in an attempt to
eradicate aquatic species with excessive growth rates
(J. Mander, SASK Water, personal communication).
Surveys were conducted before and after the chemi-
cal treatment to determine the effect of Magnacide
on the taxonomic richness and occurrence of aquatic
plants in the canal system. Plant collections were con-
ducted in 25-50 m areas before and after the point of
application. Plants were pressed and identified in the
herbarium of the University of Saskatchewan (SASK)
using several local and regional references (Hotchkiss
1972: Larson 1993: Johnson et al. 1995; Moss 1996:
Vance et al. 1984; Royer and Dickinson 1999). The
plants collected were prepared as voucher specimens
to serve as a permanent reference of this study in the
SASK collection.
Results and Discussion
The survey indicates that the taxonomic composi-
tion of the SASK Water canals includes a total of 33
species in 26 genera and 20 plant families (Table 1).
In addition, two different, unidentified, algae were col-
lected (Table 1). Our survey also indicates that no plant
CotaA-SANCHEZ and REMARCHUK: AQUATIC AND SUBAQUATIC PLANTS
165
species were eradicated from the sites as a result of
Magnacide application; that is, the same species were
identified before and after the treatment. However,
the plant density was noticeably lower after the treat-
ment with Magnacide. Various submerged species in-
cluding Potamogeton richardsonii, P. pectinatus, and
the unknown green algae were present in most of the
sites surveyed. Other aquatics, e.g., Elodea canadensis
and Lemna trisulca, were less frequent and account-
ed for a smaller proportion of plant biomass in the
canal system. Emergent species in the Cyperaceae,
Equisetaceae, and Asteraceae do not appear to play a
role in impeding water flow, but floating debris from
non-aquatic taxa, such as Melilotus albus and Hordeum
jubatum, contributes to the clogging of intake screens
throughout the canal system.
Based on this survey, it is not possible to determine
a single species that adversely affects water flow by
clogging intake screens in the SASK Water canal sys-
tems. A combination of common aquatic plants ( Pota-
mogeton spp., Alisma gramineum, A. plantago-aquat-
ica, Ceratophyllum demersum, and Myriophyllum
sibiricum), which are frequently dislodged by flowing
water or wind, are the primary contributors to blockage
of drains and intake screens. The presence of large,
coarse water pondweeds such as Potamogeton spp. and
other filamentous species in SASK Water canals is un-
desirable because their accumulation affects the reg-
ular water flow. A similar situation has been reported
in the Alberta irrigation systems (Burland and Catling
1986). In addition, green algae are common and are
presumably the greatest problem in clogging the in-
take screen systems because of their filamentous nature
and massive growth. Furthermore, some plants or frag-
ments from non-aquatic plants, particularly Melilotus
albus and Hordeum jubatum, are involved in reducing
water flow because the species tend to concentrate near
the uptake areas, especially in the vicinities of mowed
ditches along the canals.
Since this survey is mainly concerned with the flo-
ristic composition of the SASK Water canal system and
the probable effect of some species on irrigation canals,
only general comments can be made regarding the
chemical treatment and subsequent effect in the aquat-
ic and subaquatic plants. In general, the use of Magna-
cide at the concentration previously specified appears
to have a minor effect on aquatic plants because spe-
cies diversity before and after the application remained
similar. This suggests that the amount of chemical
applied is too low to notably reduce the growth and
development of aquatic plants and possibly other non-
target organisms. Acrolein is a contact herbicide; there-
fore, the root system is not damaged and plants quickly
resume growth. This leads to the frequent re-applica-
tion of acrolein (J. Mander, SASK Water, personal com-
munication). Other studies, [e.g., Bowmer and Sainty
(1977)], have shown that acrolein is effective in con-
trolling aquatic plant growth, but no concentration rates
are given. On the other hand, Sangodoyin and Smith
166
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE 1. Taxonomic list of the inventory of plants in the SASK Water canal system in south central Saskatchewan. The
arrangement is alphabetical by family, then by genus and species.
Family Species
Characeae Unknown green algae
Chara L.
Alismataceae Alisma gramineum Le}.
Alisma plantago-aquatica L.
Apiaceae/Umbelliferae Sium suave Walter
Asteraceae/Compositae
Symphiotrichum lanceolatum (Willd.) G.L. Nesom
subsp. hesperium (A. Gray) G.L. Nesom
Ceratophyllaceae Ceratophyllum demersum L.
Cyperaceae Carex rostrata Stokes
Eleocharis acicularis (L.) Roem. & Schult.
Eleocharis palustris (L.) Roem. & Schult.
Schoenoplectus acutus (Muhl. ex Bigelow)
A. Love & D. Love
Schoenoplectus tabernaemontani (C.C. Gmel.) Palla
Scirpus microcarpus J. & K. Presl.
Equisetaceae Equisetum palustre L.
Fabaceae/Leguminosae Melilotus albus Medik.
Halogaridaceae Myriophyllum sibiricum Kom.
Hydrocharitaceae Elodea canadensis Michx.
Lamiaceae/Labiatae Lycopus asper Greene
Lemnaceae Lemna trisulca L.
Onagraceae Epilobium ciliatum Raf.
Oenothera biennis L.
Poaceae/Graminae Glyceria grandis S. Watson
Hordeum jubatum L.
Phalaris arundinacea L.
Potamogetonaceae Potamogeton filiformis Pers.
Potamogeton pectinatus L.
Potamogeton richardsonii (Benn.) Rydb.
Potamogeton vaginatus Turcz.
Potamogeton zosteriformis Fern.
Ranunculaceae Ranunculus aquatilis L.
Salicaceae Salix exigua Nutt.
Sparganiaceae Sparganium sp. L.
Typhaceae Typha latifolia L.
Zannichelliaceae Zannichellia palustris L.
Common Name
Narrow-leaved Pondweed
Broad-leaved Water Plantain
Water Parsnip
Western Willow Aster
Hornwort
Beaked Sedge
Needle-Spike Rush
Common or Pale Spike-Rush
Great Bulrush
Small-fruited Bulrush
Marsh Horsetail
White Sweet Clover
Water Milfoil
Canada Waterweed
Water Horehound
Duckweed
Northern Willow Herb
Yellow-Evening Primrose
Manna Grass
Wild Barley
Reed Canary Grass
Leafy Pondweed
Sago Pondweed
Clasping Leaf Pondweed
Large-sheath Pondweed
Flat-stemmed Pondweed
Thread-leaved Buttercup
Narrow-leaved Willow
Bur-Reed
Common Cattail
Horned Pondweed
(1996) caution that low concentrations of pesticides
and agricultural chemical on surface waters have poten-
tial effects on fisheries and aquatic environments, but
conclude that Canadian water sources in general exhib-
it concentrations of pesticides below the guideline lim-
its, which suggests the proper use and handling of pes-
ticides and herbicides in Canada. Nonetheless, more
detailed research is required to draw conclusions on
water quality issues.
Considering the losses and adverse effects to farm-
ing activities caused by obstructed canal systems, the
control of aquatic plants is necessary in water bodies,
particularly in areas with intake screens, so that irri-
gation systems can be most efficiently utilized. In addi-
tion to the chemical methods currently used for the
eradication of aquatic plants, new research to imple-
ment a biological control program (involving perhaps
an herbivorous species) of aquatic plants in irrigation
canals should be conducted. Since numerous non-
aquatic species clog irrigation systems, in our view,
the most effective, environmentally friendly, and in-
expensive strategy for maintaining levels of water flow
for irrigation is to periodically manually clean intake
screens and surrounding areas. Nonetheless, cautious
and planned applications of chemicals such as acrolein
(Bowmer and Sainty 1977), Endothall (Sisneros et al.
1998), and Hydrothall (Corbus 1982) may be effective
to control excessive growth levels of species such as
Potamogeton pectinatus and filamentous algae species.
Since several native, relatively poorly represented
species are characteristic of these ecosystems, consis-
tent monitoring is recommended to preserve wild nat-
ural resources. Furthermore, the long-term effects of
pesticides on the surrounding aquatic and terrestrial
environments of the SASK Water irrigation system are
unknown. The current challenge is to assess the use of
herbicides, in particular the long-term effect and impact
of Magnacide residues, in the water quality, environ-
ment, and diversity of the wetland flora and fauna in
Saskatchewan and other Canadian ecosystems.
2007 CoTa-SANCHEZ and REMARCHUK:
Acknowledgements
We thank SASK Water personnel, in particular K.
Berkers for help with surveys, J. Mander for comments
and for showing the canal system, and two anonymous
reviewers for their comments on the manuscript. We
also thank the personnel of the Herbarium of the Uni-
versity of Saskatchewan (SASK) for their assistance.
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Accepted 23 May 2008
Life History of the Marine Isopod Cyathura polita in the Saint John
River Estuary, New Brunswick: a Species at the Northern Extent of
its Range
SARAH C. MERCER!, GLENYS D. G1BSON2, and MICHAEL J. DADSWELL?
Department of Biology, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada
'Sarah_mercer9 @hotmail.com
* glenys.gibson @acadiau.ca
>Corresponding author. Email: mdadswell @bwr.eastlink.ca Phone: (902) 585-1161. Fax: (902) 585-1059
Mercer, Sarah C., Glenys D. Gibson, and Michael J. Dadswell. 2007. Life history of the Marine Isopod Cyathura polita in the Saint
John River estuary, New Brunswick: A species at the northern extent of its range. Canadian Field-Naturalist 121(2): 168-
WIGS
The marine isopod, Cyathura polita, inhabits estuaries on the east coast of North America from the Gulf of Mexico to the Bay
of Fundy, Canada. We studied C. polita in the Saint John estuary to test for potential differences in life history that might occur
because of the northern location of the population. In the Saint John, based on our interpretation from a six-month sampling
program (May-October), the population exhibits a three-year life cycle, one year longer than more southern populations, and
stretching over four summers. Our study supported the occurrence of protogynic hermaphroditism. After two summers as
juveniles, individuals matured as females during their third summer, then displayed sex reversal by becoming males that fall,
and finally reproducing as males in their fourth summer of life before death. Mean length of C. polita from the Saint John was
greater than individuals from more southern populations (females, 13.8 + 2.14 mm; males, 16.3 + 2.41 mm). Annual brood
release occurred in late July-early August. Mean fecundity of females was 53.2 + 18.9 embryos per brood, which was greater
than found in southern populations. Cyathura polita is rare in Canada and is known only from the Saint John and along the
northern shore of the Bay of Fundy to the border of the United States.
Key Words: Isopod, Cythura politica, growth, reproductive cycle, fecundity, protogynic hermaphroditism, status, St. John
River, New Brunswick.
Our knowledge of life history evolution depends on
understanding intraspecific variation in life history
traits, such as those correlated with latitude. Cyathura
polita (Stimpson, 1855) is a benthic isopod that occurs
in estuaries along the Atlantic coast of North America
from the Gulf of Mexico to the Bay of Fundy, Canada
(Burbanck et al. 1979), and is an appropriate model for
such a study. Latitudinal variations in life history traits
have been demonstrated for C. polita populations from
Florida to Cape Cod, and include adult size and life
span (Burbanck and Burbanck 1979). Latitudinal dif-
ferences in life history traits have been described in the
common cyathuran of the eastern Atlantic, Cyathura
carinata, among populations from Portugal to Swe-
den (Bamber 1985; Olafsson and Persson 1986; Mar-
ques et al. 1994; Cruz et al. 2003). In the present study,
we investigated life history characteristics of C. polita
for its northernmost known population in the Saint John
River estuary, New Brunswick.
Cyathurans are interstitial isopods that live in bur-
rows and feed on detritus and algae in the sediment
(Burbanck 1959). They are eurytopic animals, tolerat-
ing wide fluctuations in conditions such as temperature
and salinity, an adaptation suitable to estuaries. They
are also an important indicator organism for healthy
estuarine ecotones because they are usually present
only in unpolluted waters (Burbanck and Burbanck
1979). Cyathurans in general do not swim as adults nor
do they have a pelagic larval stage, resulting in popula-
tions that are relatively immobile, sessile and disjunct.
According to Burbanck (1959, 1962a, b), C. polita
inhabits estuaries where fresh and salt water mix over
stable sediment containing gravel, silt and detritus.
They are obligatory aquatic animals and cannot with-
stand long periods of dryness (Burbanck and Bur-
banck 1979). Often C. polita are located in transition-
al zones where cattails (Typha) and salt marsh grasses
(Spartina) meet along the shoreline (Burbanck 1962b).
They are an important food for ducks and fishes (Bur-
banck and Burbanck 1979; Dadswell 1979).
The life cycle of C. polita varies with latitude (Bur-
banck and Burbanck 1979). In southern populations, .
the life cycle duration is generally one year. In Florida,
young are released from May to October, and grow
over the winter until they mature and produce young }
the following spring and summer. Most southern cya- -
thurans die at the end of their second summer (Kruczyn- -
ski and Subrahmanyam 1978). In more northern loca-
tions, such as Cape Cod, the young are released in early |
summer (late June or early July), grow rapidly in the®
fall, and produce young the following year. In these
populations some cyathurans die at the end of their
second summer while others may survive a second win-
ter and reproduce the following year, often exhibiting
protogynic hermaphroditism (Burbanck and Burbanck
ID),
168
2007
Cyathuran females carry their embryos in a ventral
marsupium made of overlapping pairs of oostegites
(Stromberg 1972). The embryos develop in the mar-
supium until they hatch from a vitelline membrane and
are released into the environment as juveniles. This
method of reproduction permits detailed study of fec-
undity and development in cyathurans, making the
species an excellent model for life history studies.
Cyathura polita has been studied extensively in the
United States, particularly in the southern states (Bur-
banck and Burbanck 1979). The Saint John River estu-
ary and along the north Bay of Fundy shore to the
United States border are so far the only known occur-
rences of C. polita in Canada, and these populations are
located at the northern extent of its range (Burbanck
et al. 1979, Atlantic Reference Center, St. Andrews,
New Brunswick). We investigated the life history of
C. polita in the Saint John River estuary, and compared
these data to published observations to explore the
hypothesis that northern cyathurans would exhibit
differences in life history relative to those at southern
latitudes. We suspected that, with probable slower
growth in this population, body size of adults would
be greater because of delayed maturity and the larger
females would carry more embryos. Also, since this
is one of the few Canadian populations of an organ-
ism which has poor powers of dispersal caused by its
non-swimming life style and a brooding reproductive
method, it may be important to characterize its life
history for future reference under the Species at Risk
Act. Finally, the Saint John River estuary is a unique
habitat in Canada containing elements of Virginian,
coastal marine ecosystems in the summer-warm shal-
lows (Bousfield 1962; Burbanck et al. 1979; Dadswell
1979), boreal, marine species in its deep, fjord-like
basins (Trites 1960; Carter and Dadswell 1983) along
with the wide range of freshwater, diadromous, and
marine species typically found in Atlantic Canada
(Rogers 1936; Bousfield 1962; Gorham 1965; Gillis
1974).
Study Area and Methods
Samples of Cyathura polita were collected from the
subtidal shore of the lower Saint John River estuary
from four stations: Bell Point, Westfield Beach, Sum-
merville and Millidgeville (Figure 1). This region is
the junction between the Bay of Fundy and the Saint
John River and both estuary outflow and tidal inflow
are inhibited by a sill at the Reversing Falls. As a result,
there are large fluctuations in salinity because of fresh-
water discharge, and inside the estuary tidal range is
reduced to 0.5 m and summer surface temperatures are
higher than in the Bay of Fundy (Metcalfe et al. 1976).
The Saint John River has a salt wedge estuary com-
bined with deep, fjord-lake basins, and surface salini-
ties at the seaward end fluctuate throughout the year,
falling to 0% during spring freshet and increasing to
20% during low freshwater flow (Metcalfe et al. 1976).
MERCER, GIBSON, and DADSWELL:
LIFE HISTORY OF CYATHURA POLITA 169
Samples were collected approximately bi-weekly,
usually at low tide, from 29 May to 27 August 2004,
with an additional collection on 10 October 2004
Samples of C. polita were collected randomly by re-
moving the surface substrate from subtidal sites in
water depths of 20 to 150 cm. Substrate was collected
with a shovel, and sieved through | mm or 0.25 mm
mesh. The smaller mesh size was used to capture juve-
niles too small to be retained by the larger mesh. Ap-
proximately 30 adults and 30 juveniles, if possible,
were collected from each location during a sampling
effort. Individuals were fixed on the day collected in
10% formalin buffered in sea water and transferred to
70% isopropyl! alcohol within three days. On each col-
lection date at each station, the water temperature at
0.5 m depth was measured using a mercury thermome-
ter, and a IL sample of surface water was collected for
salinity measurements later in the laboratory. Sedi-
ment samples from each station (1L) were collected
on 17 July 2004.
Salinity was measured using an YSI 30 Salinity
meter, after calibration with a mercury thermometer.
Sediment samples were analyzed for proportions of
gravel, sand, and silt, using a sieve series where gravel
was <38.]1 mm, sand <2.0 mm, and silt <63 um (Miall
2000).
Within a few days of capture, the total length of all
individuals was measured to the nearest mm from the
base of the antennae to the tip of the telson. Measur-
ing the total body length was preferred to estimating
size from prosomal length (Bamber 1985), for com-
parison with other literature on C. polita. Males were
distinguishable by the presence of an appendix mas-
culina on the second pleopod; females could be accu-
rately distinguished only when with oostegites forming
a brood pouch or marsupium (Figure 2). Individuals
without these sexual dimorphisms were considered
to be juveniles.
All gravid females with intact marsupia were ana-
lyzed for number and size of embryos. Embryos were
removed from marsupia and classified into four stages
of development based on electron microscopy (Mer-
cer et al., in press): Class 1 embryos were round and
contained within two membranes, the chorion and
vitelline membranes; Class 2 embryos were elongated
and showed differentiation of the three body regions
(head, thorax, abdomen), but lacked differentiation of
the appendages; Class 3 embryos were still housed
within the vitelline membrane, but had considerable
differentiation of appendages and head: and Class 4
embryos were hatched from the vitelline membrane
and had well-differentiated appendages. Embryos were
measured with a compound microscope to the nearest
17 um.
Adults and embryos were prepared for scanning
electron microscopy (SEM) with primary fixation in
2.5% glutaraldehyde (1 hr), followed by a post-fixation
in 1% osmium tetraoxide (4 hrs, both fixes in 0.1M
170
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Ficure |. Map of the Saint John River estuary, with the four study locations indicated. BP = Bell Point; WF = Westfield;
SV = Summerville; MV = Millidgeville.
cacodylate buffer in seawater). Tissues were dehydrat-
ed in an ascending ethanol series, critical point dried
using a Bio-Rad E3000 critical point drier with liq-
uid CO, and sputter coated with a Technics Hummer
II sputter coater. Samples were viewed with a JEOL
5600LV Scanning Electron Microscope at 10KV.
Statistics were estimated with MINITAB 14, Mic-
rosoft Office Excel 2003, and SPSS 12. Graphs were
plotted in Excel and SigmaPlot 9.0. Images were
manipulated for size in Corel PhotoPaint 9 and labelled
with CorelDraw 9.
Results
Water temperature, salinity and substrate were com-
pared among all stations. For most of the study period
water temperature ranged between 15 and 20°C (mean
19°C), with the highest temperature of 28°C on 17
July 2004 (Mercer 2005). Water temperatures at the
four stations were similar on each collection date
(one-way ANOVA, P = 0.993). Surface salinity ranged
from 0.2 ppt in spring to a maximum of 6.6 ppt dur-
ing mid-summer (Mercer 2005). However, mean salin-
ity differed among the four stations (one-way ANOVA,
2007
MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA 17]
FIGURE 2. Scanning electron micrograph of a female Cyathura polita with marsupium, collected from the Saint John River
during the summer of 2004. Symbols are: g = gnathopod, m = marsupium, pll= |“ pleopod.
4.5 5
4.0 |
3.9
201
2.9 5
2.0
1.5
1.0
05
0.0
(3)
Sex ratio (males:females)
(37)
29 May 4 Jun 10 Jun 18 Jun
(41)
1 Jul
Date
(61) (7) ~~ (13)
| x =~ |
17 Jul 314 Jul 17 Aug 27 Aug 10 Oct
FiGuRE 3. Male to female sex ratio of Cyathura polita during the study period in the Saint John River estuary. Data were
pooled from four stations, and sample size (7) is given in brackets above each column. On 10 October, most adult
isopods collected were male, resulting in a different scale, thus the sex ratio is given above the bar.
P < 0.001). Mean salinities at Bell Point (0.69 + 0.31
ppt) and Westfield (1.5 + 0.86 ppt) were similar
(Fisher’s 95% CI of -0.35, 0.78), while Millidgeville
(4.7 + 1.3 ppt) and Summerville (4.9 + 1.5 ppt) were
comparable (Fisher’s 95% CI of -0.94, 0.19).
Sediment composition varied among the four study
sites but all generally consisted of easily drained sub-
strates (no anoxia; Mercer 2005). Bell Point had the
highest gravel content of the four (84.5%), with little
sand (15.1%) and hardly any silt (0.4%). Westfield
had the highest percentages of sand (66.4%) and silt
(7.3%) and the least amount of gravel (26.3%). Mil-
lidgeville contained more gravel (55.9%) than sand
(43.5%) and very little silt (0.6%). Summerville was
similar to Millidgeville with abundant gravel (47.1%)
and sand (51.5%) and little silt (1.4%).
The sex ratio of C. polita varied during the study
period, from a male:female ratio of 2:1 in May, declin-
ing to 0.5:1 in July, and increasing to 21:0 in October
(Figure 3). In late spring, numbers of males exceeded
females and large juveniles (probably maturing fe-
males) were present (Figure 4). From early June to the
end of August, there were more females than males.
Small, free-living juveniles, 2 to 3 mm in length, were
first observed on 31 July 2004 numbers of presum-
ably recently released from female marsupia. By the
end of August, few distinguishable adults were col-
lected at any station (Figure 4), but there were many
172
large juveniles (probably adults without secondary
sexual characteristics). In October, despite the rarity
of males during late August, there was an abundance
of relatively large males (Figure 3).
Over the entire study, C. polita in the Saint John
River estuary were an average of 10.1 + 4.02 mm
long, although the mean length varied among stations
(Table 1). The mean length of all females collected
was 13.8 + 2.14 mm (n = 163) while males averaged
16.3 + 2.41 mm (n = 68). Males were similar in
length at all stations, as were females (Table 1; one-
way ANOVA). Mean length of individuals collected
from Bell Point and Millidgeville over the study peri-
od were similar, but isopods from both stations were
significantly longer than those from Westfield and
Summerville largely because the size of juveniles
varied among stations (Table 1; one-way ANOVA).
For three stations, mean juvenile length averaged
10.82 + 2.50 mm while juvenile cyathurans from
Summerville were smaller (8.60 + 2.49 mm; Table 1).
Most C. polita collected at Summerville throughout
the study period were juveniles. This fact was high-
lighted when comparing the size of juveniles at Sum-
merville (8.60 + 2.49 mm in length) to the size of all
individuals collected at Summerville throughout the
study (8.71 + 2.92 mm).
Seasonal changes in body length were estimated
by comparing lengths of C. polita from each collec-
tion date, after sorting individuals by sex and repro-
ductive stage (Figure 4). Since the physical charac-
teristics were relatively similar among all stations
growth data were pooled to assess life history. Females
(with oostegites) were observed from 29 May to 27
August. Mature males were observed throughout the
study, with peak abundance in late spring and early
fall. Newly-released juveniles were first observed on
31 July and were abundant into the fall.
Cohort growth was estimated by comparing the
mean length of each group of cyathurans at each col-
lection date over the entire study period (Figure 5).
Growth was estimated by extrapolating the mean length
of each cohort in October back to the mean length of
the next cohort in May. These estimates suggest that
cyathurans from Saint John have a three-year life cycle,
which spans four summer seasons. Young juveniles
(Age 0) released from the marsupia at the end of July
had a mean body length of 2.7 + 0.25 mm and aver-
aged 7.5 mm by October. Young juveniles grew ap-
proximately 0.07 mm/day, the fastest of all groups,
where growth was represented by: y = 0.068x -11.9
(R* = 0.99). The second cohort of juveniles (Age 1)
had a mean length of 8.8 + 3.4 mm in May and grew
approximately 0.02mm/day were growth was repre-
sented by: y = 0.018x + 6.44 (R?= 0.22), resulting in
a mean length in October of 12.4 + 1.90 mm. Mature
females (Age 2) were 11.0 mm mean length in May,
and grew slowly at a rate of 0.01 mm/day where growth
was represented by: y = 0.009x + 11.3 (R* = 0.05)
THE CANADIAN FIELD-NATURALIST
Vol. 121
15 July 17
=
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S 15 August 17
— 10
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B 5
2
® 0
ial 20 4
15 August 27
10 n=139
5
0
20
15 October 10
10 n=78
5
O A
0 5 10 15 20 25
Body length (mm)
FIGURE 4. Size distribution of Cyathura polita in the Saint
John River estuary for selected sampling dates during
the summer of 2004, pooled from the four stations
and grouped by sex and reproductive stage. New
juveniles, horizontal bars; juveniles, cross-hatching:
females, white; gravid female, stippled; males, black.
until they reached a mean length of 12.3 + 1.53 mm
by the end of August. Females grew little during the
summer; however, it was the reproductive period, sug-
gesting resources were partitioned into reproduction,
not growth. The size distribution of male cyathurans
2007 MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA 173
TABLE |. Body lengths of Cyathura polita in the Saint John River estuary collected over the entire study period (29 May to
10 October 2004). Differences analyzed with a one-way ANOVA and a Scheffé post-hoc comparison among sites
Location N Mean length + SD (mm) Minimum (mm) Maximum (mm)
All isopods : nitghrdrs ar:
Bell Point 247 11.2 + 3.06 ) He) 19.0
Westfield 231 9.2 + 5.00 2.0 21.0
Summerville 297 8.7 + 2.90 2 Mos 19.0
Millidgeville 272 11.5 + 4.20 2.5 24.0
one-way ANOVA df = 3, 1046, f = 35.90, p<0.000 (sites BP=>MV>WF=SV)
Males
Bell Point 26 15.1 + 1.77 12.0 18.0
Westfield 11 17.3 + 2.57 14.0 21.0
Summerville 3 16.3 + 2.51 14.0 19.0
Millidgeville 28 16.9 + 2.54 13.0 24.0
one-way ANOVA df = 3, 67, f = 3.633 ns
Females
Bell Point 50 13.4 + 1.87 9.0 19.0
Westfield 34 14.2 + 2.38 8.0 20.0
Summerville 16 12.7 + 2.79 9.0 18.0
Millidgeville 63 14.1+ 1.9] 9.0 17.0
one-way ANOVA df = 3, 162, f = 2.75 ns
Juveniles
Bell Point 136 10.6 + 2.01 6.0 15.0
Westfield 96 11.4 + 2.69 6.0 18.0
Summerville 262 8.6 + 2.49 4.0 16.0
Millidgeville 149 LOM 2515 7.0 17.0
one-way ANOVA df = 3, 642, f = 43.75, P < 0.000 (sites BP = MV = WF>SYV)
25.0
20.0
@ Males
15.0 O Females
= Juveniles
10.0 O New Juveniles
320
Mean body length (mm)
0.0
4 May 23 Jun 12 Aug Oat 20 Nov
Date
FicurE 5. Mean length of Cyathura polita from the Saint John River estuary pooled from all stations at each sampling interval
during the summer of 2004 and grouped by sex and reproductive stage. Vertical bars are standard deviations.
174
was very different from that of juveniles and females.
Males (Age 3) were largest at the end of May (mean
length 18.5 + 3.54 mm), but smaller in October (mean
length 15.1 + 1.95 mm). A bi-modal size distribution
during the growing season suggests the die-off of larg-
er and older males in early summer, and the addition
of new, smaller males in the fall as females underwent
protogynic sex change or some large juveniles matured
as males.
Other studies noted latitudinal variation in body
length of C. polita, with individuals from more
northern populations being generally larger than in
southern populations. However, environmental con-
ditions, such as the type of substrate or depth of
water, may also strongly affect body size (Burbanck
and Burbanck 1979). To assess whether cyathurans
from Saint John followed this latitudinal trend, their
mean body lengths were compared to values reported
in the literature (Table 2). Females pooled from all
stations had a mean length of 13.8 + 2.14 mm (maxi-
mum: 20.0 mm), and males had a mean length of
16.3 + 2.41 mm (maximum: 24.0 mm). These lengths
were similar to cyathurans populations studied north
of North Carolina, although none collected at Saint
John were as large as the maximum length (27 mm)
of the individuals collected from Chewonki Creek,
Maine (Burbanck 1959). The individuals collected in
the Saint John estuary differed markedly from south-
ern populations such as Georgia and Florida: the
largest male collected at St. Mark’s, Florida, was only
1.8 mm larger than the smallest male collected in the
Saint John (Kruczynski and Subrahmanyam 1978).
Broods were examined from 72 females, pooled
from all four stations. Fecundity ranged from 16 to
98 embryos per brood, with a mean of 53.2 + 18.9
embryos (Table 3). Abundant gravid females with
embryos were first observed on | July. All of these
broods contained early embryos (Class 1), with a
mean length of 0.45 + 0.06 mm. Two weeks later,
more than half of the broods contained elongated
Class 2 and 3 embryos, while the remainder contained
Class | embryos. By the end of July, 50% of broods
contained Class 4 young that had hatched out of their
vitelline membranes but were still developing within
the marsupia. By mid-August, only six gravid females
were found: two had very large Class 1 broods and two
had late-stage Class 4 broods. Within the marsupium,
embryos grew from a mean length of 0.49 + 0.08 mm as
Class 1 embryos to a mean length of 2.07 + 0.16 mm
when they were ready to be released as juveniles. Class
2 and 3 embryos had a mean length of 1.05 + 0.15 mm
and 1.12 + 0.19 mm, respectively (Table 3).
There was no correlation between female length
and fecundity (R*= 0.01). There was also no effect of
embryos/brood on embryo size (R? = 0.01) and little
mortality within broods during development (Mercer
2005). Within-brood survivorship was 96.4% with
only a slight decrease observed in mean embryos/brood
THE CANADIAN FIELD-NATURALIST
Vol. 121
when compared between Class | (57.9 + 18.77 embry-
os/brood) and Class 4 broods (55.8 + 19.64 embryos/
brood).
Discussion
Cyathura polita from the Saint John River estuary
exhibits several modifications in life history associat-
ed with its northern distribution. In particular, this
population exhibits a longer life span, a shorter and
later reproductive season, and produces larger adults
with higher fecundity, in comparison to southern
populations. Additionally, our data suggest that pro-
togynic hermaphroditism occurs, as found in some
other populations (Burbanck and Burbanck 1974).
In the Saint John River estuary, C. polita exhibits a
three-year life cycle. Juveniles are released during
July and August (Age 0) and, after little growth dur-
ing the winter, remain immature the following sum-
mer (Age 1). Over the course of their second winter,
juveniles develop into mature females (Age 2), which
reproduce the following summer. Then, based on our
length and sex ratio data, females either die or become
males (Age 3) during the late summer-fall, and repro-
duce as males during their final summer of life. The
result is a three-year life cycle, which spans four sum-
mers. In contrast, C. polita from southern populations
have shorter life cycles. For example, in St. Mark’s,
Florida, C. polita are mature one year after hatching
and reproduce throughout their second summer, sug-
gesting a life cycle of less than two years (Kruczynski
and Subrahmanyam 1978), while on southern Cape
Cod, C. polita reproduce in their second, and some-
times third, summer, suggesting a two-year life cycle
(Burbanck 1962a; Burbanck and Burbanck 1979).
Latitudinal changes in the duration and timing of
reproduction are also evident. Cyathura polita repro-
duce from April to August in St. Mark’s, Florida
(Kruczynski and Subrahmanyam 1978), from June to
July at Cape Cod, Massachusetts (Burbanck 1962a)
and July and August in the Saint John estuary (pre-
sent study). Although the breeding season is short
and relatively simultaneous in the Saint John popula-
tion, females were found with broods at different
stages of embryonic development throughout the
brooding period, especially in mid- to late-July.
The longer life cycle and delayed reproduction in
northern cyathurans probably occur because of a
shorter growing season. The population in the Saint
John lives in water that averages 19.0°C during sum-
mer, but shallow water temperatures decline to near
O°C from December until April (Metcalfe et al. 1976).
The result is limited opportunity for growth and
reproduction in warmer waters, resulting in delayed
maturation. In contrast, cyathurans from Florida live
at temperatures ranging between 13.8° and 33°C, and
above 20°C for ten months of the year (Kruczynski and
Subrahmanyam 1978). Previous studies suggest that
rapid growth in cyathurans occurs only during periods
2007
MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA
TABLE 2. Summary of body lengths for C. polita reported in the literature, as means + S.D. or range, depending on the
source,
Location of study
Saint John, New Brunswick SO 16.3+0,29
9 13.8+0.17
Marsh River, Maine GO 14-21
9 14-19
Marshfield, Massachusetts 13.3 + 0.33
Sapelo Island, Georgia 9.3 + 0.64
St. Mark’s, Florida GO 7.3 - 13.8
9 8.3 - 13.0
Body length (mm)
Author(s)
present study
Haefner et al. 1969
Frankenberg 1962
Frankenberg 1962
Krucsynski and Subrahmanyam 1978
TABLE 3. Fecundity and mean embryo length of Cyathura polita in the Saint John River estuary. Embryos were grouped by
class of development where: Class | embryos were round and contained in two membranes, the chorion and vitelline, Class
2 embryos were elongated and differentiated into three body regions (head, thorax and abdomen), Class 3 embryos were
within vitelline membrane but had differentiated appendages, Class 4 embryos were hatched from the vitelline membrane
(Mercer et al. in press).
Date Class of Number of
collected development gravid females
1 July 15
17 July 13
2 14
3 7)
31 July 1 3
2, l
3 7!
4 11
17 August I 2
2 1
3 |
4 2
of higher water temperature, and that growth ceases
when waters cool (Olafsson and Persson 1986). The
two-year life cycle described for Cape Cod may be
more similar to the southern populations, perhaps
because the southern side of Cape Cod, where the
study was conducted, is more Virginian in nature
than Boreal (Gosner 1971).
Cyathura carinata, common along the coast of
Europe, exhibits a similar cline in life history traits.
Studies of C. carinata in Portugal have suggested rel-
atively short life cycles of 19 (Cruz et al. 2003) or 18
to 22 months (Marques et al. 1994). However, three-
year life cycles are reported for C. carinata populations
in more northern locations such as Poland (Jazdzewski
1969), Sweden (Olafsson and Persson 1986) and Eng-
land (Bamber 1985). Olafsson and Persson (1986) sug-
gest that C. carinata populations in Sweden require
two years before juveniles are able to mature and repro-
duce, in contrast to southern populations, which are
able to reproduce after one year. Bamber (1985) sug-
gests that a longer life cycle is necessary in northern
populations as the isopods cannot grow during the win-
ter; whereas southern populations are able to grow
continuously and attain mature size sooner.
Fecundity Total number Embryo length (mm)
Mean + SD of embryos Mean + SD
52.8 + 22.48 956 0.45 + 0.06
59.4 + 11.08 658 0.52 + 0.09
47.0 + 18.88 630 1.05 + 0.15
38.3 + 14.21 207 1.10+0.11
52.0 + 22.61 181 0.52 + 0.01
23 1.03 + 0.11
50.5 + 15.34 34] 1.14+0.24
51.3 + 24.18 540 2.06 + 0.16
90.0 + 11.31 180 0.59 + 0.07
4 1.07 + 0.15
37) 1.13 + 0.07
SS Ee ee 67 2.15 +0.18
Cyathura polita in the Saint John River estuary ap-
pears to exhibit protogynic hermaphroditism. Sequen-
tial hermaphroditism is rare in marine peracaridans,
although relatively common in terrestrial and parasitic
species (Johnson et al. 2001). Brooks et al. (1994)
report that of the approximately 60 species of crus-
taceans known to be sequential hermaphrodites, only a
few are protogynous, including three species of Cya-
thura, three other isopods and five species of tanaids.
Evidence for protogyny from the present study includes
sex ratio and adult size. Males were abundant in the
population in May and again in October, but rare dur-
ing summer when the majority of adults were females.
Also, males in the fall were similar in length to spent
females at the end of the summer (approximately
15 and 12 mm in length, respectively), while most
large juveniles were smaller (approximately 10 mm
in length). Males continue to grow during the fall and
reach a length of approximately 18 mm in the spring
of their final year. These observations support a model
of protogynic hermaphroditism, whereby mature fe-
males become reproductive in early summer of their
second year (i.e., third summer) and release juveniles
during late July and August. After the release of young,
176
females molt, change sex, and function as mature males
in their fourth spring and summer, before death.
The hypothesis that C. polita in the Saint John River
are protogynic hermaphrodites is consistent with evi-
dence of sex change reported from other populations
under both laboratory (Burbanck and Burbanck 1974)
and field conditions (Kruczynski and Subrahmanyam
1978). While it is possible that males migrate to other
areas of the estuary after reproducing and return to
shallow waters in the fall, previous work suggests that
mostly immature cyathurans occur in the deeper waters
of this estuary, while reproductive adults are in shal-
lower water (Gillis 1974; Burbanck et al. 1979). Bur-
banck and Burbanck (1974, 1979) have reported that
some primary males can develop directly from juve-
niles, which may account for some of the variation in
sex ratios and body lengths observed here.
As hypothesized by Bamber (1985), it would be
advantageous for male cyathurans to be larger than
females to reduce their chances of mortality. Males
are able to mate with more than one female, but need
to leave their burrows in order to mate, thus they are
at a greater risk of predation. Females do not need to
expose themselves to that risk, and thus may mature
at a smaller size. Warner (1988) additionally suggest-
ed that it would be advantageous to become male at a
larger size if there is competition among males for
females.
With a few exceptions, the development of embryos
collected at Saint John was synchronized within a
brood, which agrees with observations on cyathurans
by Stromberg (1972) and Bamber (1985). Our obser-
vations suggest that the brooding period for females in
the Saint John River estuary lasts from 30 to 45 days.
Class 1 embryos were first observed in marsupia on |
July. The first hatched embryos (Class 4) were observed
on 31 July, which corresponds with the first appear-
ance of new juveniles. While comparable observa-
tions are not available for more southern populations
of this species, a 30 to 45 day brooding period may be
typical for this genus. Olafsson and Persson (1986)
found that a C. carinata population on the southern
coast of Sweden had a three to four week brooding
period, while others studying C. carinata in Portugal
determined embryonic development to also last 30 to
45 days (Cruz et al. 2003).
Brood size is not widely reported for C. polita, how-
ever broods collected from Saint John were much larg-
er (53.2 + 18.9 embryos/brood) than broods from Flori-
da (mean: 14 embryos/ brood, range 2-32; Kruczynski
and Subrahmanyam 1978). A similar south-north cline
is known in the European C. carinata. Bamber (1985)
found that C. carinata females in England carry a
mean of 29.8 embryos/brood (range 14-45), while C.
carinata females in Sweden averaged 42 + 12 embryos/
brood (Olafsson and Persson 1986). The large brood
size in Saint John C. polita is likely a function of
increased female size because of greater age (3 vs. 2
THE CANADIAN FIELD-NATURALIST
Vol. 121
years). Larger females would likely have larger marsu-
pia and therefore may be able to carry more embryos.
Cyathura polita at the northernmost extent of its
range exhibits several modifications in life history
compared to more southern North American popula-
tions. These modifications are similar to those observed
among European C. carinata populations. Our results
support the existence of a latitudinal cline in life his-
tory traits in Cyathura, with implications for life span,
size and fecundity. Plasticity in reproduction, coupled
with protogynic hermaphroditism, makes C. polita an
excellent model for studies of the effects of the envi-
ronment on life history traits.
Cyathura polita is an important indicator of estu-
arine health (Burbanck and Burbanck 1979) and since
the Saint John River estuary has one of the few popu-
lations in Canada there are compelling reasons to
understand its life history and population size. We sug-
gest a more comprehensive knowledge of the biology
of this isopod in the Saint John could assist both with
pollution monitoring and with further the distinction
of the Saint John estuary as a distinct ecosystem in
Canada. Future study of the Saint John cyathurans
should be undertaken to verify the 3-year life cycle
with sampling throughout the year and to define its
estuarine distribution and abundance. Cyathura poli-
ta occurs as far inland in the Saint John estuary as
Washademoak Lake (Figure 1; Burbanck et al. 1979)
and it is a common prey of the Shortnose Sturgeon
(Acipenser brevirostrum) another southern species
that is endemic to this estuary in Canada (Dadswell
O79):
Acknowledgments
This project was supported by NSERC (Natural
Sciences and Engineering Research Council of Canada)
through a grant to G. Gibson. The authors are grateful
to Acadia University for use of equipment; D. Mac-
Donald and G. Cheeseman of the Acadia Centre for
Microstructural Analysis; T. Avery for reviews and
corrections, Y. Zhang for statistics assistance (Acadia
University Statistical Consulting Centre), and R. Mer-
cer and C. Mercer for assistance with field work.
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Received 8 August 2006
Accepted 30 May 2008
Post-Emergence Movements and Overwintering of Snapping Turtle,
Chelydra serpentina, Hatchlings in New York and New Hampshire
GorRDON R. Uttscu!”, Matt DRAub?, and BARRY WICKLOW*
‘Corresponding author: Courtesy Professor, Department of Zoology, University of Florida, Gainesville, Florida 32611 USA
*Mailing address: 4324 NW 36th St., Cape Coral, Florida 33993 USA; e-mail: gultsch@bama.ua.edu
*Department of Biology, C. W. Post — Long Island University, 720 Northern Boulevard, Brookville, New York 11547 USA
*Department of Biology, St. Anselm College, 100 Saint Anselm Drive, Manchester, New Hampshire 03102 USA
Ultsch, Gordon R., Matt Draud, and Barry Wicklow. 2007. Post-emergence movements and overwintering of Snapping Turtle,
Chelydra serpentina, hatchlings in New York and New Hampshire. Canadian Field-Naturalist 121(2): 178-181.
Hatchling Common Snapping Turtles (Chelydra serpentina) were captured within, or as they emerged from, their nest cavities
in Long Island, New York, and in southeastern New Hampshire. They were fitted with radiotransmitters and released at their nest
sites. Their movements were monitored for as long as possible, which for some included tracking them to their overwintering
sites and relocating them the following spring. On Long Island, all hatchlings initially moved to water. Later movements were
both aquatic and terrestrial, and those that could be located while overwintering had left the water and hibernated in spring
seeps, where they were recovered alive the following April. In New Hampshire, hatchlings moved directly to nearby aquatic
habitats after emergence, where they spent the winter submerged in shallow water in root masses near banks.
Key Words: Common Snapping Turtle, Chelydra serpentina, hatchling turtles, hibernation in turtles, New York, New Hampshire.
In northern climates, the behavior of hatchling
aquatic turtles, once they have pipped and left the
eggshell, is variable both among and within species.
In some species the majority of hatchlings remain in
the nest until the following spring, although some fall
emergence may occur. These include Painted Turtles
(Chrysemys picta — Zwiefel 1989; Lindeman 1991;
Pappas et al. 2000), Common Map Turtles (Grap-
temys geographica — Pappas et al. 2000; Baker et al.
2003; Nagle et al. 2004), and Diamondback Terra-
pins (Malaclemys terrapin; Baker et al. 2006). For the
two former species, it is assumed that if the hatchlings
leave the nest, they overwinter aquatically, while for
the latter species fall emergence coupled with aquatic
or terrestrial overwintering has been reported (Draud
et al. 2004), as well as overwintering in the nest cou-
pled with fall emergence (Baker et al. 2006). Other
species have been consistently reported to be over-
whelmingly fall emergers, including Snapping Turtles
(Chelydra serpentina — Obbard and Brooks 1981; Car-
roll and Ultsch 2007), Spiny Softshell Turtles (Apalone
spinifera — Christiansen and Gallaway 1984; Costanzo
et al. 1995), Musk Turtles (Sternotherus odoratus —
Cagle 1942; Mitchell 1988), Wood Turtles (Glyptemys
insculpta — Harding 1991; Carroll and Ultsch 2007),
Spotted Turtles (Clemmys guttata — Ernst 1976; Car-
roll and Ultsch 2007), and Blanding’s Turtles (Emy-
doidea blandingii — Standing et al. 1997, 1999; Pap-
pas et al. 2000). Hatchlings of most of these species
are assumed to overwinter aquatically, even though
some may remain on land for extended periods before
entering the water; e.g., Wood Turtles (Tuttle and
Carroll 2005; Castellano, Behler, and Ultsch, unpub-
lished data) and Blanding’s Turtles (Standing et al.
1997, 1999). However, there are some reports of sus-
pected terrestrial overwintering in the hatchlings of
some of these species, based largely on the capture of
one or a few individuals in spring in microhabitats that
are suggestive of non-aquatic overwintering (Ernst
1966, 1976; Standing et al. 1997; Pappas et al. 2000,
Parren and Rice 2004). While such behavior does not |
appear common, its degree of occurrence is uncertain.
There are no reports of following individual hatch-
lings from the time of emergence from the nest until
the resumption of activity the next spring. The typical
approach of a study of overwintering behavior, a num-
ber of which have been done with adult turtles (re-
viewed by Ultsch 2006), has been the use of radiotele-
metry, permitting tracking of the turtles throughout
the hibernation period, including entrance and exit.
Recently, transmitters small enough to attach to hatch-
lings have become available, which allows tracking of
the animals from the nest to their wintering grounds,
although battery life is not yet long enough to track
the turtles until the following spring. However, once
the turtles go into hibernation, they move little, which 1 —
potentially allows recovery during the early spring:
before movement is resumed.
Here we report on the post-emergence and over- -
wintering behavior of Snapping Turtle (Chelydra ser- -
pentina) hatchlings in Long Island, New York, and in|
southeastern New Hampshire. In both localities the
hatchlings moved to water rather directly, but their
eventual overwintering sites differed considerably at’
the two locations.
178
2007
Methods
On Long Island, 15 Snapping Turtle hatchlings were
collected in the nest on 11 September 2004 from two
nest sites (seven from one and eight from the other)
in the vicinity of Cold Spring Harbor, Nassau County.
The nests were located along an earthen dam separat-
ing a lake from the stream formed by the dam over-
flow. They were brought to the laboratory, fitted with
0.9 g BD-2 transmitters (Holohil Systems Ltd., Carp,
Ontario, Canada) attached to the carapace with cyano-
acrylate glue (Krazy Glue”), and released into their
respective nest cavities on 13 September 2004. All
hatchlings left the nest cavity within 10 min. Telemetry
commenced on 14 September, using a Communication
Specialists model R1000 receiver. The expected life of
the transmitters was 30 days, and some were replaced
on day 30. Whenever possible, a visual location was
paired with the telemetric location, which was record-
ed with a hand-held Magellan GPS 315. Hatchlings
were located on nine days from 14-30 September, 10
days from 2-28 October, and on 5 and 12 November.
Five hatchlings were lost during the study. Hatchlings
that entered final hibernation sites were marked by
surveyor flags for recovery, which occurred on 5 April
2005.
In New Hampshire, hatchlings were collected from
both riverine and palustrine habitats. Six hatchlings
were collected from a nest site 20 m from a third-order
stream (riverine site) and 18 hatchlings were collected
from two adjacent nests 3 m from a first-order stream
(palustrine site) within a wetland complex. On 17 Sep-
tember 2005 six hatchlings from the riverine site were
brought to the laboratory and fitted with 0.5 g A2415
transmitters (Advanced Telemetry Systems [ATS], Inc.,
Isanti, Minnesota) attached to the carapace with epoxy
resin. They were released at the nest site on 19 Sep-
tember 2005. Hatchlings were located using an ATS
FM-100 receiver and their GPS coordinates recorded
using a Trimble Geoexplorer 3. At the palustrine site,
nine hatchlings emerging from a nest (22 September
2005) were fitted on site with Holohil transmitters (see
above) attached to the carapace using epoxy resin.
These hatchlings were then returned to the nest. After
a flash flood event during the first week of October
2004, 12 hatchlings were lost. At that time transmitters
were attached on site to eight additional hatchlings
captured at the site in shallow water. Hibernation sites
were marked using surveyor flags. These sites were
searched in March 2006.
Results
On Long Island, 10 hatchlings were followed. The
transmitter became detached from three after their
movement to water. Two others were found dead. One
moved to the water and was found dead on a beach
after 26 days. One exhibited considerable movement
activity (34-260 m between location periods), includ-
ing some terrestrial movements, and by 5 October had
ULTSCH, DRAUD, and WICKLOW: OVERWINTERING SNAPPING TURTLE HATCHLINGS
179
moved to a seep, buried in leaves near the surface; it
was found dead there on 3 April at a depth of 5 cm.
One moved into and out of water and then back into
the water, and was located on 12 November dug into
sand under a tree root in about 8 cm of water. This
animal could not be located the following April.
Four hatchlings were tracked to their hibernation
quarters and located the following spring. One moved
immediately to water, then was found on 2 October
under a log 8 m from water. By 10 October it had
moved uphill to a spring seep, and two days later had
dug to about 20 cm depth and did not move through
12 November. It was found alive and responsive at
the same site on the following 3 April in 8 cm of sat-
urated mud. A second moved directly to water and then
emerged onto land on 30 September about 1.5 m from
the water. By 2 October it had moved to a site about
20 m from the water, where it buried under leaves
and remained until 8 October. By 10 October it had
moved 15 m and was at the surface under cat briar.
On 12 October it had moved to a second spring seep,
where it buried, moved, and reburied over a 6-day
period to depths of 10-13 cm. By 21 October it had
relocated in the seep and buried to about 15 cm and
had not moved by the last observation on 12 Novem-
ber. It was found alive and alert near the surface at
the same location on 3 April. A third moved to water,
and moved aquatically until 30 September, when it
moved into the same seep area as the second, where
it moved intermittently, being found buried at depths
of 8-13 cm until 18 October, when it settled into the
wet organic detritus along the side of a fallen tree,
and had not moved by 12 November. It overwintered
there and was found alive and well at the surface of
the mud at the same site on 3 April. A fourth moved to
and about in shallow water for five days. By the sev-
enth day it had left the water and buried into moist
sphagnum moss about 0.25 m from shore for 3-5 days
without moving. It then moved back to water for
2-3 days and then back onto land on 30 September,
where it buried for three weeks about 8 cm into the
same seep used by the second and third hatchlings. It
then moved downward to about 15 cm by 21 October
and did not move through 12 November. It was recoy-
ered alive at this site on 3 April.
In New Hampshire, at the riverine site, 5 of 6 hatch-
lings moved first up a gentle slope and then down a
steep slope to the river 6 m below. Three of these
entered the river within 3 h of release, one after 24 h,
and one after 36 h. The sixth moved 8 m on land, down
a slope, and burrowed under leaves and then entered
the water the next day, all within 24 h of release. The
hatchlings remained within 20 m of their entrance
points into the river, remaining in shallow (2-3 cm)
water, and often buried in sand or detritus with only
the head exposed. After extreme flooding caused by
heavy rains (25-30 cm in 3 days) in early October, the
hatchlings were lost. Three detached transmitters were
180
found, suggesting significant buffeting of the hatch-
lings.
At the palustrine New Hampshire site, 15 hatchlings
(five with and 10 without transmitters) traveled 3-5 m
to enter a nearby wetland stream within 4 h of release.
Of the four additional hatchlings with transmitters,
one burrowed back into the sand of the nest, where it
remained for 2 days until dug out by a predator. Two
more entered the water within 24 h, and the fourth trav-
eled 5 m, remained under wetland vegetation for one
day, burrowed into sand one day, then traveled 20 m
to shallow water (8 cm) at the edge of the stream. Six
hatchlings were lost after a flood during the first week
of October. On 11 October 2005, eight additional
hatchlings were located along the stream within 5 m
of the nest. Each was fitted with a transmitter on site
and then returned to the stream. Of these, one moved
15 m downstream into the flooded wetland within 24 h,
two moved 12 and 17 m downstream into the wet-
land by 16 October 2005. A fourth moved 15 m into
the wetland by 2 November 2005. Four hatchlings re-
mained tucked into roots of sedges and silky dogwood
in 2-3 cm water along the edge of the wetland stream,
3-5 m from the nest. A fifth had remained in this
location since 22 September 2004 (visually located on
2 November 2005). Three hatchlings were relocated
in March 2006, and they were all at the locations at
which they were last observed in November 2005.
One was located on 8 March, 6 cm below the surface
and 2.5 m from the nest site, under sedge roots at the
edge of the stream in 1°C water. A second was found
on 8 March, 6 cm below the surface in 1°C water under
leaves at the base of a submerged log in the wetland
15 m from its nest. A third was found on 10 March,
7 cm below the surface in 2°C water under silky dog-
wood roots at the edge of the stream 2.5 m from its
nest. The three hatchlings increased in mass by 12.9 to
19.5% from autumn release to spring capture, but we
cannot partition mass gain between possible autumn
feeding and potential winter water uptake.
Discussion
Although the transmitters were small, they still were
up to ~10% of the body mass of the hatchlings, and
potentially could snag on vegetation during locomo-
tion. While we cannot discount effects on behavior,
since the hatchlings within a given group behaved sim-
ilarly, we do not think that transmitter attachment had
a major effect on behavior relative to that of an unfet-
tered hatchling.
On Long Island, the four hatchlings that were recov-
ered from their hibernacula in April all overwintered
in spring seeps away from standing water, in contrast
to the behavior of the New Hampshire hatchlings,
which all moved to shallow water and remained there
to overwinter. The four Long Island hatchlings also
initially moved to open water from their release in the
nest cavity (two from each nest), but they then followed
THE CANADIAN FIELD-NATURALIST
Vol. 121
different paths, sometimes including both aquatic and
terrestrial movements, before eventually settling into
the spring seeps for overwintering, as did the fifth,
although we could not find it in April. In the sense
that Long Island hibernacula were not under standing
water, the hibernacula were terrestrial and differed from
those expected by assuming that hatchlings move to
aquatic lentic or lotic habitats and overwinter there, as
was the case with the New Hampshire hatchlings.
However, the Long Island hatchlings were surround-
ed by a constantly saturated medium that appeared to
have an underground flow, so desiccation was not a
threat. We do not know the oxygen content of the
interstitial water, but assume that it was appreciable
or the turtles would have died if they had remained
submerged throughout the winter. Reese et al. (2004)
found that Snapping Turtle hatchlings will die in 30
days if submerged in anoxic water at 3°C, compared
to survival of adults for at least 125 days under simi-
lar conditions (Reese at al. 2002). The limited survival
of anoxic hatchlings is attributable to their reduced
whole-body buffering capacity, and therefore reduced
tolerance of the metabolic acidosis that accompanies
the accumulation of lactic acid during anaerobiosis.
The reduced buffering capacity is due to the reduced
relative amount of bone, especially in the shell, of the
hatchlings compared to adults, and is a general char-
acteristic of hatchling turtles (Reese et al. 2004). This
inability to tolerate anoxia would preclude the turtles
from hibernating for extended periods in anoxic con-
ditions, such as mud under standing water. Thus the
seep water that surrounded the hatchlings must have
had a significant amount of dissolved oxygen, perhaps
maintained by a constant percolation of oxygenated
spring water that exceeded the biochemical oxygen
demand of the detritus at the low wintertime temper-
atures. Alternatively, it is possible that the hatchlings
may have occasionally air-breathed, since the soil sur-
face of the seeps did not freeze during the winter. Air-
breathing would require that the animals move to the
surface to breathe (on the assumption that the water
saturation prevents any underground air pockets from
forming); since turtles are not completely immobilized
by cold, air-breathing cannot be ruled out. Why the
hatchlings moved out of open water to hibernate in the
seeps, or how they oriented toward them, is uncertain.
Thus the overwintering habitats of hatchling Snap-
ping Turtles are varied, ranging from the expected
shallow-water retreats typical of the New Hampshire
hatchlings, to the use of wet but not aquatic sites (e.g.,
seeps), even though aquatic sites are nearby and easily
accessible, as seen with the Long Island hatchlings.
The use of seeps as hibernacula for turtles, especially
young ones, may be more common than has been ap-
preciated. V. Lamoureaux (personal communication),
while studying the use of seeps as hibernacula for
Green Frogs (Rana clamitans) in New York (Lam-
oureux and Madison 1999), found one adult and three
2007
juvenile (9.5 — 11 cm) Snapping Turtles leaving a seep
area in late April, where they apparently overwintered,
and moving toward a nearby pond. Dunson (1986)
radiotracked four snapping turtles in a tidal marsh area
in Virginia, and they all moved to seep/spring areas on
higher ground, which remained unfrozen throughout
the winter, suggesting that the use of seeps as hibernac-
ula may not be limited to immature turtles, although
almost all reports of hibernacula of adult Snapping
Turtles are of those covered by standing or flowing
water (Ultsch 2006).
Acknowledgments
We thank Sara Zimnavoda for help with radiotrack-
ing in Long Island and students of the Aquatic Ecology
course at St. Anselm College for similar aid in New
Hampshire. This study was supported by a National
Science Foundation grant to GRU (IBN 0076592).
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Received 27 October 2006
Accepted 29 January 2008
Notes on Succession in Old Fields in Southeastern Ontario: the Herbs
A. CROWDER!, R. HARMSEN!, and S. E. BLATT?
'Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6 Canada
26 Wallace Place, Wolfville, Nova Scotia B4P 2R6 Canada
3Corresponding author
Crowder, A., R. Harmsen, and S. E. Blatt. 2007. Notes on succession in old fields in southeastern Ontario: the herbs. Canadian
Field-Naturalist 121(2): 182-190.
Vegetation in abandoned hayfields was monitored during 1976-1998. An earlier successional stage followed ploughing.
Changes in tree, shrub and vine populations have been reported earlier and showed expected increases in species richness
and cover. Highest species richness of herbs occurred three years after ploughing. Non-woody species richness trended
irregularly downward, while non-woody cover was variable, peaking in 1987. Within the herbaceous community, year-to-
year changes in cover and frequency of species in the following selected groups are reported here: 18 grasses including sown
and adventive species; 13 legumes including two sown species; 14 macroforbs of the Compositae, including a goldenrod,
Solidago canadensis, which dominated parts of the fields; a rosette weed, Taraxacum officinalis; sedges, horsetails and some
other minor components. Grasses and goldenrods were grazed, sometimes intensively and repeatedly, by insects; grasses
were impacted by skipper larvae (Thymelicus lineola), and goldenrods by beetle larvae (Trirhabda spp.). Effects of repeated
outbreaks on host plant cover are shown for two plots (100 m2) matching the scale of outbreaks.
Key Words: old field herbs, phytophagous insects, plant succession, goldenrods, grasses, legumes, Ontario.
Two abandoned hayfields in southeastern Ontario
were selected as sites in which to investigate possible
effects of insect herbivory on early stages of secondary
succession. The location, geology, history of land use,
and soils have been described earlier (Crowder and
Harmsen 1998). Their nutrient status was discussed by
Epp and Aarssen (1988). Estimates of increasing cover
and abundance of trees, shrubs and vines (51 species)
during 1976-1998 were shown to follow classic succes-
sional models, although with unusual pioneer species.
After two decades the structure of the vegetation had
become heterogeneous, with groves of small trees,
shrub thickets, patches dominated by macroforbs and
patches still dominated by grasses. Cover varied with
fluctuations of weather from year to year, while soil
differences influenced cover and species richness.
Most successional studies on old fields use synoptic
records from fields of different ages, a method which
obscures minor fluctuations and differences between
fields. In this paper, year-to-year observations are used
to follow temporal changes in the herbaceous com-
munity and to relate its cover and species richness to
those of the woody layers. Herbaceous species found
in 27 experimental plots are listed to allow comparisons
with other areas; lichens and bryophytes are omitted.
Differences in soil texture (Crowder and Harmsen
1998) influenced cover and richness.
Within the herbaceous community, details of tem-
poral change in selected groups of plants and some
specific interactions are reported here. Grasses were
the main constituents of the hayfields, including both
agricultural and adventives species. Legumes also in-
cluded sown and invasive species. Invasive macroforbs,
182
mainly Solidago and Aster species, dominated parts of
the fields; population changes of 15 species are reported.
Taraxacum officinalis, Dandelion, was selected to rep-
resent rosette plants. Sedges, horsetails and a few other
minor components are described.
During the decades when vegetation was monitored,
arthropods were also sampled (McBrien et al. 1983).
Two insect populations had repetitive outbreaks which
caused significant decreases of plant cover, and are
therefore the only arthropods reported here. Larvae of ©
the hesperiid European Skipper (Thymelicus lineola)
grazed on grasses, particularly on Timothy, Phleum |
pratense (Henderson 1986). Larvae of beetles of the
genus Trirhabda Le Conte (Coleoptera, Chrysomeli- |
dae), mostly Trirhabda virgata, grazed primarily on
leaves of Solidago canadensis, causing intense defo-
liation. The strong effect of one or more outbreaks of
beetles on plant vigour was replicated experimentally
(McBrien and Harmsen 1987). These defoliations can
cause changes in successional pathways at the grass-
macroforb stage by affecting the relative dominance *
of grass or goldenrods. Blatt et al. (2001) have used |
these data to develop a successional model including :
herbivore effects, showing possible outcomes for the °
herbs.
Methods
A hayfield abandoned in 1970 and one mown for
hay until 1975 were selected at Queen’s University |
Biology Station at Opinicon, Frontenac County, On-
tario (44°03'N, 76°0'W). Nine permanent plots, each |
100 m?, were demarcated in 1975 in the older field,
which was called S field, for Solidago. Eighteen similar \
fi
2007
plots were placed in the younger field, and nine of
these were ploughed once; these ploughed plots were
called P and the undisturbed plots G, for grass. All
plant species present in the 27 plots were listed in June
and September of sampling years: 1976-1988, 1995,
1998. Cover was estimated using five randomly placed
quadrats within each plot; cover values used in this
paper were from September (McBrien et al. 1983;
Henderson 1986). Each quadrat was 50 cm x 50 cm
in area. Values were compared using Scheffe’s mean
separation test (Zar 1984; Blatt et al. 2003). Percentage
frequency, the number of quadrats in which a species
was recorded at a particular time (Mellinger 1972;
Eriksson and Jakobssen 1998), was used as a measure
of relative abundance (Maycock and Guzikowa 1984).
Nomenclature follows Newmaster et al. (1998);
voucher specimens were lodged at the Fowler Herba-
rium, Queen’s University. Lichens, mosses and liver-
worts have been omitted from this paper.
Results and Discussion
Species richness of woody species and of herbs per
100 m? plot, through 1976-1998, is shown in Figure 1.
Note gaps in sampling after 1988. Herb richness trended
downward with high annual variance while woody spe-
cies richness increased over time (see Crowder and
Harmsen 1998).
The range of species richness was 7-65 in ploughed
(P) plots, 6-42 in grass-dominated (G) plots, and 6-33
in Solidago (S) plots. Species richness of herbs and of
woody plants in a grass plot (G1) and a ploughed plot
(P2) are shown in Figures 2 and 3. Because soil texture
was found to be correlated with species richness, the
selected plots were matched for texture; both are sandy,
with relatively low richness (Crowder and Harmsen
1998). Note the difference in scale between Figures
2B and 3B. Species richness peaked in P2 three years
after ploughing when several weedy species were pre-
sent (cf. Armesto and Pickett 1985).
Herbaceous taxa observed in 27 plots during 1976-
1998 are listed in Table 1, in alphabetical order, with
English names added (Newmaster et al. 1998). The
closest comparable study site is a 50-year-old field, at
Erindale (Toronto area), described by Maycock and
Guzikowa (1984), in which 93% of the species listed
at Opinicon were found. Other old fields in Ontario
have been described by Lamoureux (1970), Reader
(1990) and Reader and Buck (1991). Fewer of the
Opinicon species (37%) were found in abandoned
hayfields in New York State described by Mellinger
(1972), and only 20% in old fields in Quebec (Dan-
sereau and Gille 1949). The proportion of introduced
species at Opinicon was 37%, which is higher than the
average for the Kingston region (Beschel et al. 1970).
They included both crop plants and weeds.
During 1976-1998 the cover value of woody plants
for all plots combined increased significantly (Crowder
and Harmsen 1998): herbaceous cover, however, did
CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS
183
not decrease significantly over time (Figure 1). Mean
percent cover of non-woody species in 1998 was not
significantly different from that in 1988, 1986, 1984,
1978, 1977 and 1976. Years with high cover were 1979,
1980, 1981, 1985, or 1987 (Figure 1). These values
suggest that herbaceous cover was related to year-to-
year changes in weather or herbivory rather than to a
steady increase in shading. No correlations of cover
values with available climatic data were found, al-
though the years with low cover had partial droughts
that caused soil cracking.
Herb cover in selected plots is shown in Figures 2A
and 3A, with relatively low variance between years.
Plot G1 consistently had higher values than P2. The
main components of herb cover are discussed in the
following section. The hayfields had been sown with
Phleum pratense (Timothy) and Poa pratensis (Ken-
tucky Blue Grass), which maintained their frequency
after the cessation of mowing (see Table 2). Mellinger
(1972) also reported their high frequency in old hay-
fields in New York. In the Opinicon fields the two spe-
cies are associated randomly (Epp and Aarssen 1988).
Table 2 lists annual percentage frequency of nine
grass species. The most frequent invasive grasses have
been Elymus repens, Poa compressa and Calamagrostis
canadensis. Other less common invasive grasses, not
shown on Table 2, were Bromus inermis ssp. pum-
pellianus, Echinochloa crusgalli, Elymus trachycaulon
ssp. trachycaulon, Muhlenbergia mexicana, Panicum
acuminatum, Poa capillare, Poa annua, Setaria pumila
and Setaria viridis. Annual cover of all grass species
in all plots varied from a mean of 35% in 1998 to a
mean of 87% in 1987. Cover in the G (grass) plots ex-
ceeded 100% in some years, because layers of two or
more species developed. Differences between the three
sets of plots were not consistent.
Cover of grasses did not show synchronous trends
within sets of plots. Changes in grass cover were not
correlated with soil texture; the two plots shown in
Figures 2 and 3 were matched for soil texture but the
undisturbed plot G1 in Figure 2 had high grass cover
and the ploughed plot in Figure 3 had not. A temporal
pattern in the grass-dominated plot showed successive
peaks three to five years apart followed by declines
in cover values. This pattern was typical of grass-
dominated plots. The disturbed ploughed plot shown
in Figure 3 had two less marked peaks and declines.
Herbivory was demonstrated by Henderson (1986) to
reduce grass cover significantly; he counted larvae of
a hesperiid European Skipper (Thymelicus lineola)
and found that peaks of larval numbers coincided with
low cover of Elymus repens, Phleum pratense and Poa
pratensis, and low concentrations of skipper larvae with
high grass cover. The grass species differed in their
capacity to recover after an insect outbreak, with
Phleum pratense being most negatively affected. Out-
break behaviour by the European Skipper has been
documented by Pengelly (1961) elsewhere in Ontario.
184 THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Alphabetic list of herbs recorded within 27 plots during 1975-1998 at two old fields at Opinicon, with English
names. Nomenclature follows Newmaster et al. (1998).
Acalypha virginica. Copperleaf
Achillea millefolium ssp. millefolium. Yarrow
Agrimonia gryposepala. Hooked Agrimony
Agrostis gigantea. Redtop
Ambrosia artemisiifolia. Common Ragweed
Anaphalis margaritacea. Pearly Everlasting
Anemone canadensis. Canada Anemone
Antennaria neglecta. Field Pussytoes
Antennaria parlinii. Plantain-leaved Pussytoes
Arctium minus ssp. minus. Common Burdock
Artemisia absinthium. Absinth
Asclepias syriaca. Common Milkweed
Aster cordifolius. Heart-leaved Aster
Aster ericoides. Heath Aster
Aster lanceolatus. Panicled Aster (includes Aster simplex)
Aster lateriflorus. Calico Aster
Aster umbellatus. Flat-topped White Aster
Aster urophyllus. Arrow-leaved Aster
Barbarea vulgaris. Wintercress
Bromus inermis ssp. pumpellianus. Smooth Brome
Calamagrostis canadensis. Canada Blue Joint
Capsella bursa-pastoris. Shepherd’s Purse
Carex aurea. Sedge
Carex communis. Common Sedge
Carex debilis. Sedge
Carex foenea. Sedge
Carex gracillima. Filiform Sedge
Carex granularis. Granular Sedge
Carex laxiflora. Slender Sedge
Carex normalis. Normal Sedge
Carex scoparia. Sedge
Carex vulpinoidea. Fox Sedge
Cerastium fontanum ssp. triviale. Common Mouse-ear
Chickweed
Chenopodium simplex. Maple-leaved Goosefoot
Chrysanthemum leucanthemum. Ox-eye Daisy
Cichorium intybus. Chicory
Cirsium arvense. Canada Thistle
Clinopodium vulgare. Wild Basil
Convolvulus arvensis. Field Bindweed
Conyza canadensis. Canada Fleabane
Dactylis glomerata. Orchard Grass
Danthonia spicata. Poverty Oat Grass
Daucus carota. Wild Carrot
Dianthus armeria. Deptford Pink
Echinochloa crusgalli. Echinochloa
Echium vulgare. Blueweed
Elymus repens. Couchgrass
Elymus trachycaulus ssp. trachycaulus. Slender Wheat
Grass
Epilobium hirsutum. Hairy Willow Herb
Equisetum arvense. Field Horsetail
Equisetum hyemale. Scouring Rush
Erigeron annuus. Daisy Fleabane
Erigeron philadelphicus ssp. philadelphicus. Common
Fleabane
Erigeron strigosus. Lesser Fleabane
Eupatorium maculatum. Joe Pye Weed
Eupatorium perfoliatum. Boneset
Eupatorium rugosum. White Snakeroot
Euthamia graminifolia. Grass-leaved Goldenrod
Fragaria vesca ssp. americana. Wood Strawberry
Fragaria virginiana. Wild Strawberry
Galinsoga quadriradiata. Quickweed
Galium obtusum. Wild Madder
Galium trifidum ssp. trifidum. Small Bedstraw
Gentiana andrewsii. Bottle Gentian
Geranium bicknellii. Bicknell’s Crane’s Bill
Geum aleppicum. Yellow Avens
Geum canadense. White Avens
Hieracium aurantiacum. Orange Hawkweed
Hieracium piloselloides. King Devil
Hypericum perforatum. Common St. John’s Wort
Juncus dudleyi. Dudley’s Rush
Lactuca canadensis. Wild Lettuce
Lepidium campestre. Field Peppergrass
Lithospermum officinale. Gromwell
Lobelia inflata. Indian Tobacco
Lycopus americanus. Cut-leaved Water Horehound
Lycopus uniflorus. Water Horehound
Lysimachia terrestris. Yellow Loosestrife
Medicago lupulina. Black Medick
Medicago sativa ssp. sativa. Alfalfa
Medicago sativa ssp. falcata. Medick
Melilotus alba. White Sweet Clover
Melilotus officinalis. Yellow Sweet Clover
Mentha arvensis ssp. borealis. Wild Mint
Muhlenbergia mexicana. Dropseed
Oenothera perennis. Sundrops
Onoclea sensibilis. Sensitive Fern
Osmunda regalis. Royal Fern
Oxalis stricta. Yellow Wood Sorrel
Panicum acuminatum. Woolly Panic Grass
Panicum capillare. Witch Grass
Panicum depauperatum. Starved Panic Grass
Phleum pratense. Timothy
Physalis heterophylla. Clammy Ground Cherry
Plantago lanceolata. Narrow-leaved Plantain
Plantago major. Common Plantain
Plantago rugelii. Rugel’s Plantain
Poa annua. Annual Blue Grass
Poa compressa. Canada Blue Grass
Poa pratensis ssp. pratensis. Kentucky Blue Grass
Polygonum convolvulus. Black Bindweed
Polygonum persicaria. Lady’s Thumb
Polygonum sagittatum Arrowleaved Tear-Thumb
Potentilla argentea. Silvery Cinquefoil
Potentilla canadensis. Old Field Cinquefoil
Potentilla inclinata. Downy Cinquefoil
Potentilla norvegica. Rough Cinquefoil
Potentilla recta. Erect Cinquefoil
Prunella vulgaris ssp. vulgaris. Selfheal
Ranunculus abortivus. Kidney-leaf Buttercup
Ranunculus acris. Field Buttercup
Rudbeckia hirta. Black-eyed Susan
Rumex acetosella ssp. acetosella. Sheep Sorrel
Rumex crispus. Curly Dock
Sanicula marilandica. Black Snakeroot
Setaria pumila. Yellow Foxtail
Setaria. viridis. Green Foxtail
Silene vulgaris. Bladder Campion
Sisymbrium officinale. Hedge-Mustard
Sisyrinchium angustifolium. Blue-eyed Grass
Solidago caesia. Blue-stemmed Goldenrod
2007
TABLE |. (continued)
Solidago canadensis. Canada Goldenrod
Solidago gigantea. Yall Goldenrod
Solidago juncea. Early Goldenrod
Solidago nemoralis ssp. nemoralis. Gray Goldenrod
Solidago rugosa. Rough-stemmed Goldenrod
Sonchus asper ssp. asper. Spiny Sow-thistle
Spiranthes romanzoffiana. Hooded Ladies’ Tresses
Stellaria borealis ssp. borealis. Northern Stitchwort
Stellaria graminea. Stitchwort
Stellaria media, Chickweed
Taraxacum officinale. Dandelion
Teucrium canadense ssp. canadense. Germander
Thlaspi arvense. Field Pennycress
Larval outbreaks are considered to affect the succes-
sional pathway in the Opinicon fields in a model
developed from our data by Blatt et al. (2001). When
insect herbivory on grasses in old fields was studied by
Brown et al. (1988), Elymus repens was the grass
species most affected by Homoptera/Auchenorrhycha;
these authors considered that both texture and struc-
ture of the successional vegetation were affected by
the insects.
The initial seed mix contained Medicago sativa ssp.
sativa (Alfalfa) and possibly Trifolium pratense (Red
Clover) which have persisted in the fields. Medicago
sativa frequency was 3-5% in 1976-1980, rose to 12%
in 1983 and fell again to 2% in the 1990s. Trifolium
pratense frequency was 35% in 1976, fell to less than
8% during 1980-1995 and rose again to 22% in 1998.
Trifolium aureum, Trifolium campestre and Trifolium
hybridum were recorded occasionally.
Vetches (Vicia sativa ssp. nigra, Vicia cracca, and
Vicia tetrasperma) became more frequent in later years
with a maximal frequency of 69% for Vicia cracca in
1987. Possibly these straggling plants benefited from
the cessation of mowing. Other legumes recorded were
Melilotus alba, Melilotus officinalis and Medicago
lupulinus, randomly distributed except for Medicago
lupulina, which was most frequent in 1976-1979 in
the ploughed plots.
Macroforb areas of the fields have been and are
dominated by Solidago canadensis (Canada Golden-
rod), which we observed to spread by both seeds and
rhizomes. Its importance in old field succession in
North America has been widely reported (e.g., Dale
et al. 1966; Goldberg and Gross 1988). Percentage fre-
quency of Solidago canadensis in all plots during
1976-1998 is shown as mean values in Table 2, start-
ing with minimal values in 1976 and reaching a maxi-
mum in 1987-1988. Frequencies of Solidago nemoralis
(Old Field Goldenrod), Solidago caesia (Blue-stemmed
Goldenrod), and Euthamia graminifolia (Grass-leaved
Goldenrod), also given in Table 2, were higher in the
1980s than in the 1970s or 1990s. When the three sets
of plots were compared, Solidago canadensis and Soli-
CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 185
Tragopogon dubius. Goatsbeard
Tragopogon pratensis ssp. pratensis. Showy Goatsbeard
Trifolium aureum. Yellow Clover
Trifolium campestre. Smaller Hop Clover
Trifolium hybridum. Alsike Clover
Trifolium pratense. Red Clover
Verbascum thapsus. Common Mullein
Verbena hastata. Blue Vervain
Veronica serpyllifolia ssp. serpyllifolia. Thyme-leaved
Speedwell
Vicia cracca. Cow Vetch
Vicia sativa ssp. nigra. Common Vetch
Vicia tetrasperma. Slender Vetch
Viola sp. Violet
dago caesia were found to be most frequent in the S$
plots (during 1976-1998 the mean frequency of Solli-
dago canadensis in S plots was 47%, in the P plots
37% and in the G plots 26%). Solidago nemoralis and
Solidago juncea were generally more frequent in sandy
dry plots. This distribution may be explained by dif-
ferent soil moisture tolerances among the congeners
(Werner and Platt 1976).
Cover values for Solidago canadensis ranged from
1 to 100%. Patterns were not synchronous between
plots. Two examples are shown. Figure 3 shows Plot
P2, with three irregular peaks of goldenrod cover, and
Figure 2 shows Plot G1 with only three years of spor-
adic goldenrod cover. The plot with three peaks and
lows (Figure 3) had a periodicity also observed in other
plots, while others had one, two, or three-and-a-half
episodes. Some sandy plots similar to that shown in
Figure 2 failed to develop dense Solidago cover, and
did not have a series of highs and lows.
We have found that the cover of Solidago canadensis
was negatively and significantly correlated with larval
density of three species of Trirhabda beetles in the field
(McBrien et al. 1983; Blatt et al. 2001). Trirhabda vir-
gata was most abundant. Field experiments showed that
effects on biomass and cover similar to those caused by
larval grazing could be produced by manual defoliation
(McBrien and Harmsen 1987).
Within a field, beetle outbreaks were not synchro-
nous between plots. Outbreaks in the two fields also
occurred at different times. In the P/G field, years with
several plot peaks of Solidago cover were 1980, 1981,
and 1985, and years with most lows were 1976, 1985
and 1998. That both peaks and lows occurred in 1985
shows the small scale of the beetle outbreaks, some-
times confined to a plot 100 m?. Outbreaks on a much
larger scale have been reported, for example, on a scale
of several hectares (Werner et al. 1980), and over two
hundred insect species have been reported on Solidago
canadensis.
Macroforbs growing to the same size as some gold-
enrods included the asters Aster umbellatus (Flat-
topped White Aster) and Aster novae-angliae (New
186
THE CANADIAN FIELD-NATURALIST
Vol. 121
Mean non-woody species nchness
30
25 finan By
pie a a
\ es v4 \
ve
20
15
10
AD
abca be
ia i
defg
cto bedef'” A”
> fb 5 ao &
es “st S&S Ss PSH SF SF PS & SS S
Year
[___] Mean percent cover of non-woody species, all plots
ber A...... Mean percent cover of woody species, all plots
Ficure |. Comparison of woody and non-woody vegetation. Mean species richness of herbaceous plants in 27 plots (species
number/100 m7) for the years 1976-1988, 1995 and 1998 is shown by dots. Mean percent cover of non-woody species
in all plots in September is shown in the histogram; means which do not share the same letter are significantly different.
Mean percent cover of woody species in all plots is indicated by a dotted line with triangles.
England Aster). During the years of observation the
percentage frequency of these two species increased
(Table 2). Three smaller species, Aster lanceolatus (in-
cluded in Aster simplex in Table 2), Aster lateriflorus
and Aster ontarionis, have been lumped together be-
cause they were generally intermingled in straggling
clumps which also increased in frequency. Aster
cordifolius (Heart-leaved Aster) and Aster urophyllus
(Arrow-leaved Aster), both of which are shade-tolerant
(Semple et al. 1996), had low frequency. Five of the
aster species found in the Opinicon fields were re-
corded by Maycock and Guzikowa (1984).
When all aster species are lumped together, mean
annual cover ranged from | to 45%. Comparing dif-
2007 CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 187
ferent plots, patterns of peaks and lows like those ob-
a bh ns ty my fe a served in Solidago canadensis occurred in some plots,
= MANM=OOVVSTOMOVONATO but with a longer periodicity ranging up to ten years.
ats Since these fluctuations in cover were not synchronous
ES o> ee ce a, oe a, i a between plots or fields, it is possible that they were due
to herbivory, like the changes in Solidago and grasses.
Two species of horsetail, Equisetum arvense and
Equisetum hyemale, were present in the P/G field
throughout 1976-1998 with a combined mean per-
1988
4
0
6
]
5
5
4
0
% ie, alt ES bo os a centage frequency of 68%. Both species were most
Bre SS EN ot eovwrer N'| frequent in’ satidy plots and) are locally. common in
disturbed habitats such as sandpits (Crowder et al.
% mons_s = > ~ 1997). Cover did not exceed 29% for the two species
ee OVS VN VIN O ONO Om ST combined in any yearnand was,eenerally low:
ss Mean frequency of Jaraxacum officinale ranged
oe etd tase er ve es os from 13% in 1981 to 37% in 1988. Dandelions were
mE en VV VO VONNANANS™ HAOTAOSH resent in all three sets of plots every year, with highest
frequency in the S and lowest in the G plots. The
x rosette growth form of Taraxacum officinale is similar
FIFA ROR CVV OMS +n CoH ammAc]| to that of many genera in the plots, such as Hieraciwn
spp. and Antennaria parlinii, and probably facilitates
eS early spring growth.
Rloann = oA _ ~ Sedges, Carex spp., were observed in all years and
AS WOarIANNOCSOVOOCOTAONODOO VANMOMNTNS : > m
in all sets of plots, but were not analysed because of
Q difficulty in distinguishing non-flowering stages. Carex
DPT oon voted Vornono|| granularis, Carex laxiflora and Carex normalis were
relatively abundant. Mean annual frequency of all spe-
a Bil, es S cies was 39% and most plants were found in plots with
Nemo ms ootast =n vyenee*e™2 high moisture content (Crowder and Harmsen 1998).
Mean cover was less than 2% until 1979, rose to 7%
SSBmooVVOFate+Voemnac]| in 1987, and then declined again.
Species which increased in both frequency and
Lis Seal bs a cover during 1976-1998 included Asclepias syriaca
SEAS SUG SO OO OS SOO ANd Sil!’ (Milkweed) and Ranunculus acns (Field Buttercup).
Species which declined in both frequency and cover
Sea Of MAO VOkmm unl aecluded (Rumex acetosella ssp. acetosella (Sheep’s
Sorrel), Ambrosia artemisiifolia (Ragweed), Oxalis
stricta (Yellow Wood Sorrel), and Prunella vulgaris
Bae ealtosg Veotoauaacoaaw sal ssp. lanceolata (Self-Heal). Rianex acetosella and
Ambrosia artemisiifolia, which are weedy species,
were mostly in the ploughed plots, whereas Prunella
1981
ay
52
1979 1980
1978
1977
TABLE 2. Mean percentage frequency of selected herbs in all plots during 1976-1988, 1995 and 1998 (n = 162)
S Recien a = a vulgaris was initially concentrated in grass plots. A
~ PPrepoomomomona = Vom ~~2 1 minor component with low cover value but high fre-
quency (40-50%) was Potentilla recta (Erect Cin-
8 quefoil).
s When grass cover was lowered by grazing of
= Thymelicus lineola, feeding preferentially on Phleum
S pratense, cover of Solidago and Aster spp. in both
aS fields increased rapidly; they then maintained their
Be dominance at levels equal to that of Poa pratensis,
2 S§ preventing the recovery of Phlewm pratense (Hen-
S§s s a derson 1986; Blatt et al. 2001; Blatt et al. 2003). In
SSs s See 8 = aS addition to interactions between grasses and golden-
2 S as SSSEs . Ses $< _ || rods, other layers were affected. For example, when
28 S oes S Ss Sos ue SS Sse S|} the cover of Solidago canadensis was decreased by
. SS LSS = S < : ge SSS SPSS the feeding of beetle larvae a chamaephyte. Fragaria
Sis Sg E Sas Bes YS &s £3 S|] virginiana (Wild Strawberry). showed significantly
a ls8 = SSEE S ss = = = = =S 3 $2 $]] increased cover (McBrien et al. 1983). This finding
FIRQAHQTOCAACHAAANHAHAAXTIICK
188 THE CANADIAN FIELD-NATURALIST Vol. 121
30 A. Non-woody species
25
20
15 /
~ | a ~
a 7 ° ie :
Non-woody
species nchness
oOo WA
a B. Woody species
50 C. Grass species
3
2
D. Solidago species
oO
Oo
Mean percent (+ SE) cover or species richness
i=)
WD SS
OO A
S
>
,
&
“te,
9p
N
FIGURE 2. Conditions in grass plot G1 during 1976-1988, 1995 and 1998. 2A shows mean percent cover of non-woody species,
and non-woody species richness (species number/100 m7). 2B shows cover and species richness of woody plants. 2C shows
mean cover of all grass species. 2D shows Solidago species cover, which was only present in 1986, 1988 and 1998.
2007 CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 189
A. Non-woody species
35 a
abe
30 Yani
Non-woody species nchness
23 ~ ae. 7 e ~
B. Woody species
Woody species nchness
70 — C- Grass species
Mean percent (+ SE) cover or species richness
60 — D. Solidago species
wo dA oe OD DOD GD &
SP OOM NL
N
ro So Dd FP SF
Or OTe ane een ase
Year
FIGuRE 3. Conditions in ploughed plot P2 during 1976-1988, 1995 and 1998. 3A shows non-woody species richness (species/
100 m2) and mean annual percent cover. 3B shows woody species richness and cover. 3C shows cover of all grass species
with peaks and lows. 3D shows cover of Solidago species with peaks and lows.
190
has been confirmed elsewhere by Carson and Pickett
(1990) who attributed it to decreased shading by the
macroforb.
Shade-tolerant herbs in the plots included Sanicula
marilandica (Black Snakeroot), Teucrium canadense
ssp. canadense (Germander), Lactuca canadensis (Wild
Lettuce), Solidago caesia and Aster cordifolius. The
increase in woody plants documented by Crowder and
Harmsen (1998) could have been expected to create
conditions favouring increases of such shade-tolerant
species. They did not, however, increase in frequency
or cover over time. Presumably shaded microniches
under plants such as goldenrods were present in early
years and were later replaced by similar conditions
under shrubs or trees.
Acknowledgments
Partial funding was received from the Ontario
Ministry of Natural Resources; we particularly thank
R. Wagner, S. Strobl and P. Neave. Preliminary analysis
of field results was done by C. Vardy, and we thank her
and the many others who helped with field surveys.
Our thanks are due to J. Pringle and P. Catling who
identified some of the composites and sedges. We are
grateful for the continued support of Queen’s University
Biology Station.
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Received 16 October 2006
Accepted 28 April 2008
Growth of White Spruce, Picea glauca, Seedlings in Relation
to Microenvironmental Conditions in a Forest-Prairie Ecotone
of Southwestern Manitoba
SOPHAN CHHIN! and G. GEOFF WANG?
' Corresponding author: Department of Forestry, Michigan State University, East Lansing, Michigan 48824-1222 USA
* Department of Forestry and Natural Resources, Clemson University, 261 Lehotsky Hall, Clemson, South Carolina 29634-0317
USA; e-mail: gwang @clemson.edu
| Chhin, Sophan, and G. Geoff Wang. 2007. Growth of White Spruce, Picea glauca, seedlings in relation to microenvironmental!
conditions in a forest-prairie ecotone of southwestern Manitoba. Canadian Field-Naturalist 121(2): 191-200.
The influence of microenvironmental conditions on the growth performance (1.e., diameter and height growth) of White Spruce
[Picea glauca (Moench) Voss] seedlings was examined within three contrasting habitats: White Spruce tree islands, open
prairies and Trembling Aspen (Populus tremuloides Michx.) groves. These habitats exist within a disjunct occurrence of White
Spruce at its southern limit of distribution in three mixed-grass prairie preserves in the Spruce Woods Provincial Park within
the forest-prairie ecotone of southwestern Manitoba. Microenvironmental measurements (i.e., light, temperature, relative
humidity, soil moisture) were obtained on 10 sites in each of the three habitats and growth characteristics of 60 White Spruce
seedlings were measured in each of three habitats. Higher light and soil temperature conditions occurred within the open
prairie. In contrast, lower light and soil temperature conditions occurred under the tree canopy of aspen groves and the north-
ern aspect of spruce islands, which moderated the effect of the dry regional climate. Height growth did not vary significantly
among the three habitats. The greater diameter growth and decreased slenderness of White Spruce seedlings in the open ver-
sus the shaded habitats appears to be a result of increased photosynthesis at higher light intensity and may also represent a
morphological adaptation to withstand the effect of increased wind exposure. The increased slenderness of White Spruce in
the shaded habitats appears to be a morphological adaptation of increasing carbon allocation towards height growth and thus
maximizing effective competition for light.
Key Words: White Spruce, Picea glauca, prairie-forest boundary, aspen parkland, growth, microenvironment, seed-seedling
conflicts, Manitoba.
In the prairie provinces of Canada, the southern
limit of conifer species occurs in the aspen parkland.
Vegetation in the aspen parkland is dominated by
Trembling Aspen (Populus tremuloides Michx.) which
exists as continuous forest or as groves intermixed
with prairie (Bird 1961). The aspen parkland is a tran-
sitional vegetation zone (ecotone) between prairie to
the south and boreal forest to the north (Bird 1961;
Zoltai 1975). The prevailing regional climate of the
aspen parkland and mixed-grass prairie in southwestern
_ Manitoba is characterized by low precipitation, high
summer temperatures, and dry winds, any of which may
be limiting factors to plant distribution and growth
- (Coupland 1950). For instance, previous studies which
examined the radial growth-climate association of White
| Spruce [Picea glauca (Moench) Voss] in the aspen
parkland region demonstrated that growth was restrict-
ed by moisture deficiency exacerbated by tempera-
ture-induced drought stress (Chhin et al. 2004).
Tree invasion into prairies under the dry climate of
_ the aspen parkland must be assisted by favorable micro-
environments or “safe sites” (Harper 1977; De Steven
» 1991a, 1991b; Kellman and Kading 1992; Chhin and
- Wang 2002). In many prairie environments, shrubs are
_ known to facilitate tree seedling establishment (Call-
away 1995; Chambers 2001) by suppressing grass com-
petition (Bird 1961; Werner and Harbeck 1982) and by
providing protective conditions from abiotic forces such
as fire (Coupland 1950; Bird 1961). The presence of
previously established trees in open prairie not only
reduces evaporative water loss from plants (Kellman
and Kading 1992) but also increases soil moisture and
nutrients (Wilson and Kleb 1996; Li and Wilson 1998:
Wilson 1998). Furthermore, woody canopies modify
the understory microclimate, including shading, inter-
ception of precipitation, litter fall (Vetaas 1992), reduced
wind speed (Archibold et al. 1996), lower soil temper-
ature, and higher humidity (Carlson and Groot 1997).
Within the aspen parkland of southwestern Manitoba,
White Spruce typically establishes under a Trembling
Aspen canopy and will eventually replace aspen to form
White Spruce forests in the absence of disturbances
(Bird 1961). Abundant White Spruce seedlings and
saplings are commonly observed in many Trembling
Aspen stands, indicating a favorable environment for
regeneration. However, White Spruce can occasional-
ly invade open prairies within previously established
Creeping Juniper (Juniperus horizontalis Moench)
patches (Bird 1961; Chhin and Wang 2002). Creeping
Juniper is believed to reduce mortality of White Spruce
seedlings from prairie ground fires (Bird 1961). The
progressive recruitment and establishment of White
19]
Spruce in the open prairie results in the development
of White Spruce tree islands which are surrounded
by mixed-grass prairie. The islands have an asym-
metric spatial structure since White Spruce regenera-
tion is concentrated on the northern aspect of the tree
islands. Chhin and Wang (2002) determined that the
best environment for White Spruce regeneration with-
in mixed-grass prairie was under the influence of estab-
lished White Spruce trees (i.e., on the north vs. south
aspect and between 4 to 12 m from the spruce island
centre, and close association with Creeping Juniper).
Other studies have also observed growth of White
Spruce in the aspen parkland region on favourable
microenvironments such as the north side of trees, north
side of river valleys, and other north-facing slopes
and microsites (Zoltai 1975; Hogg 1994; Hogg and
Schwartz 1997). Together, these previous studies
suggested that the suitability for White Spruce estab-
lishment decreased from aspen groves (1.e., under a
Trembling Aspen canopy), to White Spruce tree islands
(i.e., under the influence of established White Spruce
trees) and to Creeping Juniper patches in open prairies.
However, environmental conditions associated with
these habitats have not been documented and how
White Spruce growth differs among these habitats has
not been quantified.
Ecotones are expected to be sensitive to climatic
change, and the environment of the aspen parkland is
believed to characterize the future boreal forest under
a warmer climate (Hogg 1994). Moderated microen-
vironmental conditions may potentially serve as micro-
scale refugia or “safe sites” (Harper 1977) for tree
seedlings in the context of global warming during the
21* century (IPCC 2007) within a climatically sensi-
tive ecotonal region of the aspen parkland. The first
objective of this study was to examine the microenvi-
ronmental conditions of five habitats within the aspen
parkland of the Spruce Woods Provincial Park (SWPP)
in southwestern Manitoba, Canada. These habitats
include one habitat in Trembling Aspen groves (AG)
[interior (AG-I)], two habitats in White Spruce tree
islands (SI) [northern (SI-N) vs. southern (SI-S) aspect],
and adjacent to the islands, two habitats in open prairies
(OP) [ground vegetation dominated by grass (OP-G)
vs. juniper (OP-J)]. The second objective of the study
was to compare White Spruce seedling growth (i.e.,
height and diameter) among three habitats (OP-J, SI-
N, and AG-I; seedlings were not sampled in OP-G and
SI-S since it was extremely rare to find them in these
habitats) and relate growth patterns to the microenvi-
ronmental patterns.
Methods
Study area
The study was conducted in the SWPP which is
located in southwestern Manitoba (49°40'N, 99°15'W)
(Figure 1). The nearest meteorological station with a
long-term climatic record is the Brandon Agriculture
THE CANADIAN FIELD-NATURALIST
Vol. 121
Station (49°52'N, 99°59'W), which is located about
88.5 km northwest of the SWPP. The area experienced,
for the reference period of 1971-2000, an average annu-
al temperature of 2.4°C (Environment Canada 2002).
Average annual precipitation amounts to 474.0 mm,
with 78.3% as rainfall and the remainder as snowfall.
Precipitation peaks in June (75.7 mm), and tempera-
ture is highest in July, reaching a daily mean of 18.9°C
(Environment Canada 2002). Vegetation in the park
is characterized as the aspen-oak grove of the boreal
forest region (Rowe 1972). A sandhill, mixed-grass
prairie community unique to the aspen parkland is
located in the SWPP and consists of Trembling Aspen
intermixed with a disjunct population of White Spruce
at its southern limit of distribution (Schykulski and
Moore 1997). The SWPP resides over an extensive belt
of deltaic sands (6500 km”) created 12 000 years B.P.
when a predecessor of the Assiniboine River flowed
into glacial Lake Agassiz. The SWPP is located on
this delta, and this area has been designated as the
Assiniboine Delta Natural Region (ADNR). Within the
ADNR, the SWPP contains some of the best exam-
ples of the few remaining remnants of native mixed-
grass prairie, since most of the native prairie in the
ADNKR has been lost to agricultural cultivation. The
mixed-grass prairie preserves of the SWPP are pro-
tected under the mandate of the Prairie Management
Plan of Manitoba Conservation in order to preserve
its unique and rich biodiversity of plants, insects, and
wildlife (Schykulski and Moore 1997).
Natural and anthropogenic disturbances have con-
tributed to the landscape mosaic of the aspen park-
land (Bird 1961). In pre-European settlement times,
disturbances included fire which occurred naturally
or started by Native people in order to maintain the
prairie environment for the Bison (Bison bison L.).
Grazing by herbivores such as Bison and Elk (Cervus
canadensis Erxl.) prevented the encroachment of the
forest onto the prairie. European settlement contributed
to the decline of the Bison population due to hunting,
livestock grazing, intensive agriculture, and the policy
of fire suppression (Bird 1961). The reduction of graz-
ing and fire pressure has resulted in the encroachment
of the forest onto the prairie. A policy of fire suppres-
sion continues within the SWPP, with intermittent
prescribed burning permitted within the mixed-grass
prairie preserves (Schykulski and Moore 1997).
Site selection
The study was located in three mixed-grass prairie
preserves north of the Assiniboine river: Picnic, Aspen
Bluff, and Jackfish prairie (Figure 1). Ten White Spruce
islands (SI) with adjacent open prairie (OP), and ten
aspen groves (AG) were selected using aerial photo-
graphs and field reconnaissance. Four sites of each
habitat were chosen in Picnic prairie and Aspen Bluff
prairie, whereas two sites of each habitat were selected
in Jackfish prairie. Although the occurrence of habitat
types could be confounded with environmental con-
2007
99°14'W
Spruce Woods :
Provincial Park |
Epinette
mag Prairie Sites
—— Park Boundary
— Roads
Township Line
1:75000
500 0 500 1000 Meters
Le ae
CHHIN AND WANG: GROWTH OF WHITE SPRUCE SEEDLINGS
193
99°06'W MANITOBA /
L«
of’ ¢ .
Lo
SYS
z
<
=
WwW
x
O
|
0.05). Light intensity in the aspen
grove interior (AG-I, 18.3%) was significantly less
than any of the other habitats (all P < 0.05).
Soil temperature differed among some of the habi-
tats (Table |). Soil temperature within spruce islands
was lower in the northern (SI-N, 13.5°C) versus south-
ern (SI-S, 16.4°C) aspect (P < 0.05). The spruce island
habitats (SI-N, SI-S) in turn were not significantly
different from soil temperature in the open prairie
grass habitat (OP-G, 15.7°C) (all P > 0.05). Both the
SI-S and the OP-G habitats had significantly greater
soil temperature than the open prairie juniper habitat
(OP-J, 13.4°C) (all P < 0.05). Soil temperature of the
aspen grove habitat (AG-I, 10.2°C) was lower than
any of the other habitats (all P < 0.05). While surface
and air temperature did not differ significantly among
the habitats they followed similar relative patterns
among the habitats as described for soil temperature.
Although relative air humidity was greatest in aspen
groves (AG-I, 36.7%), lowest in the open prairie (OP-
G, 30.4%; OP-J, 31.8%), and intermediate in spruce
islands (SI-N, 32.7%; SI-S, 33.7%) there were no
significant differences in relative air humidity among
these habitats (all P > 0.05) (Table 1).
Surface soil moisture in the northern island aspect
(SI-N, 7.1%) was not significantly different from that
in the southern aspect (SI-S, 4.7%) (P > 0.05) (Table
1). Soil moisture in the southern island aspect in turn
was significantly less than in either the open prairie
grass (OP-G, 10.9%) or aspen grove (AG-I, 8.9%) (all
P < 0.05). Surface soil moisture of the open prairie
juniper (OP-J, 8.8%) habitat was not significantly dif-
ferent from the other habitats (all P > 0.05). Average
soil moisture of all profile depths in the SI-S (1.8%)
habitat was less than that in the AG-I (8.0%) habitat
(all P < 0.05) (Table 1).
Seedling growth
White Spruce seedlings in the open prairie and
spruce islands established exclusively amongst Creep-
ing Juniper. Seedlings in aspen groves established ex-
clusively on forest floor litter (i.e., Trembling Aspen
leaves). General seedling attributes are shown in Table
2. Mean annual height growth of seedlings did not dif-
fer significantly among the three habitats (all P > 0.05)
(Figure 2). Mean annual diameter growth differed
significantly amongst the three habitats (all P < 0.05)
(Figure 3). Open prairie seedlings had the greatest diam-
eter followed by seedlings from spruce islands and
then aspen groves. The slenderness coefficient also dif-
fered significantly amongst the three habitats (all
P <0.05) (Figure 4). Aspen grove seedlings were most
slender, followed by seedlings from spruce islands and
then open prairies.
Discussion
Less exposure to solar radiation was expected under
the canopy of aspen groves and spruce islands because
of shading. Light conditions under aspen canopy
196 THE CANADIAN FIELD-NATURALIST Vol. 121
TABLE |. Microenvironmental conditions of Trembling Aspen groves [interior (AG-I) habitat], and White Spruce tree islands
{north (SI-N) and south (SI-S) aspect habitat] with adjacent open prairies [grass (OP-G) and juniper (OP-J) habitat]. The
lower and upper limits of a 95% confidence interval of the mean of ten replicates per habitat are shown in parentheses. For
each microenvironmental variable, habitats with different letters are significantly different (P < 0.05). Except for the tem-
perature data, comparisons were made based on log-transformed data. Microenvironmental variables with no letters for each
habitat were not significantly different (P > 0.05).
Habitat
Variable! OP-G OP-J SLS SIN AG-I
Light (% PAR) 100.0 c 100.0 c 79.1 ¢ 54.5 b 18.3 a
Temp. (°C) (100.0, 100.0) (100.0, 100.0) (62.6, 100.0) (44.1, 67.2) (15.9, 21.1)
Soil lSeade 13.4 be 16.4e 13.5 cd 10.2 a
(14.1, 17.3) (11.8, 14.9) (IS.7/, 7eib)) (12.1, 14.9) (9.7, 10.7)
Surface 31.3 30.8 30.1 29.1 28.0
(28.1, 34.5) (27.9, 33.8) (27.3, 32.9) (27.4, 30.7) (25.6, 30.3)
Air 31.7 31.6 29.3 30.5 29.1
(28.7, 34.6) (28.7, 34.4) (26.8, 31.9) (28.9, 32.1) (26.7, 31.6)
RH (%) 30.4 31.8 33)57/ 32.7 36.7
(25.0, 37.1) (26.0, 38.8) (26.7, 42.6) (28.0, 38.1) (30.5, 44.1)
SM (% vol)
Surface 10.9b 8.8 ab 47a 7.1 ab 8.9b
(8.7, 13.6) (6.5, 11.9) (2.6, 8.5) Gell, 97) (7.4, 10.8)
Profile 5.3 ab 2.4 ab 1.8a 2.0 ab 8.0 b
(2.0, 12.0) (0.4, 7.2) (0.4, 4.5) (0.7, 4.4) (6.2, 10.1)
'PAR, photosynthetically active radiation; Temp., temperature; RH, relative air humidity; SM, soil moisture
TABLE 2. Attributes of White Spruce seedlings from White Spruce tree islands, open prairies, and Trembling Aspen groves.
Standard error of the mean in parentheses.
Attribute Open Prairie Spruce Island Aspen Grove
Sample size (7) 60 60 60
Height (cm) 50.4 (4.9) 49.8 (4.6) 69.6 (6.7)
Diameter (cm) 1.66 (0.21) 1.24 (0.13) 1.16 (0.12)
Age (year) 11.4 (0.6) 11.9 (0.7) 15.1 (0.8)
(18.3%) occurred at levels similar to other studies of
aspen stands (Archibold et al. 1996: 23%; Carlson
and Groot 1997: 18%). The dominant overstory White
Spruce trees cast shade onto the understory, with more
shade on the northern versus southern aspect. The
reduced soil temperature of the northern versus south-
ern island aspect as well as the lowest soil temperature
in aspen groves indicated that areas with increased
shade were also cooler. This was expected since tem-
perature is generally a function of solar radiation
(Archibold et al. 1996; Morecroft et al. 1998).
Decreased soil temperature may lead to decreased
evapotranspirative demands in the aforementioned
shaded areas. In our study, soil temperature in aspen
groves was 3.2-5.5°C less than in the open prairie.
Other studies have also reported similar levels of
reduced soil temperature in aspen (Archibold et al.
1996: 5°C) and other deciduous stands (Morecroft et
al. 1998; 2.5°C) compared to adjacent open conditions.
Reduced solar radiation has the potential to moderate
the microclimate in forest/grassland ecosystems and
thus provide a less hostile environment for tree regen-
eration and growth (Zoltai 1975; Hogg 1994; Bres-
hears et al. 1997; Hogg and Schwartz 1997).
While light and soil temperature varied significant-
ly among different habitats, surface and air tempera-
ture as well as relative air humidity did not. Morecroft
et al. (1998) also found that a deciduous forest canopy
had a greater effect on reducing soil temperature than
air temperature. Although air temperature and humid-
ity did not differ significantly among the habitats, the
lower level of air temperature in aspen groves (2.5-
2.6°C lower) versus the open prairie was comparable
to that observed in other aspen (Archibold et al. 1996:
1.5°C lower; Carlson and Groot 1997: 0.7°C) and
deciduous stands (Morecroft et al. 1998: 0.6-0.9°C).
Relative humidity was 4.9-6.3% higher in aspen groves
compared to the open prairie. This is comparable to a
study in the aspen parkland of Saskatchewan by Archi-
bold et al. (1996). They observed that average relative
humidity in the summer was generally 4% higher in-
side than outside an aspen grove. Higher air humidity
2007
i a) Open Prairie (4.1 + 0.2)
o 3
©
= 4
®
ic 1
0
3 + 5
Mean annual height growth (cm y-")
b) Spruce Island (3.7 + 0.1)
Frequency
OF — GN) 16> =|
3 4 5
Mean annual height growth (cm yr-")
c) Aspen Grove (4.2 + 0.2)
- NYO wo Ff
Frequency
3 4 5
Mean annual height growth (cm yr-")
FIGURE 2. Frequency histogram of height/age ratio (i.e.,
mean annual height growth) of White Spruce seedlings
from open prairies (a), White Spruce tree islands (b),
and Trembling Aspen groves (c). Mean + | standard
error of the mean of 10 replicates per habitat shown
in parentheses. Mean annual height growth did not
differ significantly among habitats (P > 0.05).
is usually linked to the effects of reduced wind speed
and reduced air temperature (Archibold et al. 1996)
Some significant differences in soil moisture were
observed among the five habitats. Better moisture re-
tention due to increased shading may explain signifi-
cantly higher surface and profile soil moisture in aspen
groves when compared to SI-S. Due to different degrees
of shading, both soil surface and profile moisture were
also slightly higher on the northern versus southern
aspect of spruce islands. These results support the
CHHIN AND WANG: GROWTH OF WHITE SPRUCE SEEDLINGS
197
a) Open Prairie (0.126 + 0.010 C)
4
o 3
Cc
52
oO
®
iy i
0
0.05 0.10 0.15 0.20 0.25
Mean annual diameter growth (cm y-")
A b) Spruce Island (0.091 + 0.003 B)
o 3
S
iD
ion
oO
ic 1
0
0.05 0.10 0.15 0.20 0.25
Mean annual diameter growth (cm y-")
4 c) Aspen Grove (0.069 + 0.004 A)
an)
©
ae
oO
Liter
0
0.05 0.10 0.15 0.20 0.25
Mean annual diameter growth (cm yr-")
FIGURE 3. Frequency histogram of RCD/age ratio (i.e., mean
annual radial growth) of White Spruce seedlings from
open prairies (a), White Spruce tree islands (b), and
Trembling Aspen groves (c). Mean + | standard
error of the mean of 10 replicates per habitat shown
in parentheses, and habitats with different capital
letters are significantly different (P < 0.05).
hypothesis of soil moisture retention which asserts
that soil moisture is expected to be greater in areas of
increased shading (Wilson and Kleb 1996; Li and
Wilson 1998; Wilson 1998). The significantly higher
soil surface moisture found in OP-G, and the lack of
significant difference between open prairies and spruce
islands or aspen groves in soil profile moisture are,
however, difficult to explain. A possible explanation is
that tree or shrub canopies, especially those of conifers,
intercept rainfall and thus limit the amount of through-
198
fall (Vetaas 1992). Although water may also enter soil
via stemflow, it has been shown that such water is more
likely to infiltrate to deep soil layers to be used by tree
species (Parker 1983; Vetaas 1992; Breshears et al.
IGS).
Although open prairies represent the worst habitat
for White Spruce establishment (Chhin and Wang
2002), they support the best seedling growth. These
results support the hypothesis of seed-seedling con-
flicts which asserts that the habitat requirements for
optimal seedling establishment are discordant with that
for seedling growth (Schupp 1995). White Spruce did
not show any significant differences in height growth
amongst the three habitats. In contrast, White Spruce
significantly had the greatest diameter growth within
the open prairie followed by tree islands and then aspen
groves. These results suggest that diameter growth was
more responsive to the environmental factors associ-
ated with each habitat than height growth. These results
are consistent with that of Groot (1999) who reported
that diameter growth of young, planted White Spruce
was primarily influenced by light availability, whereas
height growth exhibited complex relationships with
multiple environmental factors.
The larger diameter growth of White Spruce seedlings
in the open prairie was not expected given the harsher
abiotic microenvironment (i.e., heat stress) of the open
prairie compared to the moderated microclimatic con-
ditions within the spruce islands and aspen groves.
There are a number of factors which may have con-
tributed to the better diameter growth of seedlings in
the open prairie and the lower than expected diameter
growth within spruce islands and aspen groves. These
factors include the effect of substrate growth condi-
tions, light availability, and morphological adaptations
to wind. First, surveying White Spruce seedlings in the
open prairie was difficult given the rarity of finding
open prairie spruce. Furthermore, those White Spruce
individuals that were found grew exclusively in asso-
ciation with Creeping Juniper. Therefore, the better
than expected growth in the open prairie may be due
to the moderating influence of a Creeping Juniper sub-
strate, and Creeping Juniper is likely providing White
Spruce seedlings protective cover from prairie ground
fires (Bird 1961). However, seedlings on the northern
aspect of spruce islands also established exclusively on
juniper. If substrate was a dominant factor influencing
diameter growth it is unclear why seedlings from the
open prairie and spruce islands would have different
diameter growth rates given the same growth substrate.
Corresponding to the diameter growth reduction
from OP-J to SI-N to AG-I, the degree of shade in-
creased. The reduced supply of solar radiation could
have outweighed the benefits of a moderated micro-
climate. In other words, while shade is an important
moderating microclimatic factor that is conducive to
successful White Spruce establishment (Chhin and
Wang 2002), shade becomes a limiting factor to sub-
sequent seedling growth. Although ecophysiological
THE CANADIAN FIELD-NATURALIST
Vol. 121
a) Open Prairie (38.3 + 1.9 A)
Frequency
Oo - NY WwW FA
20), 650) 4055 450)) (GOV e70mco
Slendemess coefficient
b) Spruce Island (45.9 + 1.3 B)
Frequency
Oo =] NY WwW A
20° 230) 240 350) 260 e700
Slendemess coefficient
c) Aspen Grove (64.2 + 2.3 C)
w WwW F#&
Frequency
20K. E50) ee AOe 50 a0 eae Oe oO.
Slendemess coefficient
FiGuRE 4. Frequency histogram of height/RCD ratio (ie.,
slenderness coefficient) of White Spruce seedlings
from White Spruce tree islands (a), open prairies
(b), and Trembling Aspen groves (c). Mean + | stan-
dard error of the mean of 10 replicates per habitat
shown in parentheses, and habitats with different
capital letters are significantly different (P < 0.05).
studies indicate that White Spruce seedlings general-
ly attain maximal height growth at 40% full sunlight
(Lieffers and Stadt 1994; Lieffers et al. 1996), seedling
diameter growth is greatest at full light intensity (Logan
1969). This may explain the reduced diameter growth
of seedlings from the northern aspect of spruce islands
which was characterized by light levels of 54.5% of
full sunlight. Light levels below 8% generally results
in mortality of White Spruce (Lieffers and Stadt 1994).
While light conditions under the aspen canopy (18.3%)
were greater than this minimal light level, they are
2007
well below that required for maximal diameter growth,
Man and Lieffers (1997) also confirmed that light
intensity under an aspen canopy was not sufficient to
reach the saturation point although light levels were
above the compensation point for White Spruce pho-
tosynthesis.
Another likely factor which may explain the dis-
crepancy in diameter growth between the three habi-
tats may be morphological adaptations to the effect of
wind. Although wind speed was not examined here,
it has been shown in another study in Saskatchewan
that wind speed in an aspen grove was reduced to 7%
of that in adjacent prairie (Archibold et al. 1996).
Forest openings are generally characterized by greater
wind speeds than forest interiors (Carlson and Groot
1997). Wind causes trees to sway and this can lead to
morphological adaptations to resist the effect of wind
via increased diameter growth particularly at the base
of trees (Kozlowski et al. 1991; Telewski 1995).
The effect of shade and wind is further supported
by White Spruce being most slender in aspen groves
than in tree islands and the open prairie. These results
confirmed our morphological observations in the field.
White Spruce seedlings from aspen groves were gen-
erally tall and thin, while seedlings from the open
prairie were short and thick. White Spruce seedlings
from spruce islands were intermediate along this mor-
phological continuum. The morphology of White
Spruce under aspen canopy is thus suited for concen-
trating growth in height to counter the effects of over-
topping vegetation (Lieffers and Stadt 1994; Lieffers
et al. 1996). The morphology of White Spruce in the
open prairie is suited for a more robust stature to with-
stand the effect of winds during summer (Archibold
et al. 1996) and wind and snow abrasion during winter.
Wind and snow abrasion have been shown to influence
the growth form of open grown spruce at its northern
limit of distribution at the tree-line in Churchill, Man-
itoba (Scott et al. 1993).
The possible discordance in habitat requirements of
different developmental stages of White Spruce seed-
lings may potentially influence successional dynam-
ics in a forest-prairie ecotone in the Spruce Woods
Provincial Park. For instance, projecting the initial
slow degree of White Spruce seedling establishment
in the open prairie onto all developmental stages of
White Spruce may underestimate the rate of succes-
sional expansion of forest onto prairie, since subse-
quent seedling diameter growth in the open prairie
appears to benefit from high light availability. Nonethe-
less, further research is required to separate the rela-
tive effects of high light availability and increased wind
exposure on increased diameter growth of White Spruce
seedlings in the open prairie. Projecting the initial
establishment success of White Spruce under a Trem-
bling Aspen canopy onto all developmental stages of
White Spruce may overestimate the rate of succession-
al replacement of Trembling Aspen by White Spruce,
CHHIN AND WANG: GROWTH OF WHITE SPRUCE SEEDLINGS
199
since shade appears to outweigh the benefits of a mod-
erated microclimate for diameter growth of White
Spruce seedlings under a Trembling Aspen canopy
In conclusion, higher light and soil temperature con
ditions occurred within the open versus shaded habi-
tats. The greater diameter growth and decreased slen-
derness of White Spruce seedlings in the open versus
shaded habitats appears to be a result of increased
photosynthesis at higher light intensity and may also
represent a morphological adaptation to withstand the
effect of increased wind exposure. The increased
slenderness of White Spruce in the shaded habitats
appears to be a morphological adaptation of increas-
ing carbon allocation towards height growth and thus
maximizing effective competition for light. While shade
is an important moderating microclimatic factor that is
conducive to successful White Spruce establishment
(Chhin and Wang 2002), our study has shown that at
a later developmental stage, shade is a contributing
factor towards limiting diameter growth. Our results
support the hypothesis that there are conflicting habi-
tat requirements for trees at different developmental
stages (Schupp 1995). Thus, in the context of future
global warming (IPCC 2007) in a climatically sensitive
ecotonal region of the aspen parkland, although mod-
erated microclimates may act as micro-scale refugia
or “safe sites” (Harper 1977) for tree seedling estab-
lishment, this may be at the expense of reduced growth
at a later developmental stage. Models of forest growth
in response to climate change should therefore take into
account the role of conflicting habitat requirements
for trees at different developmental stages.
Acknowledgments
This study was funded through a Natural Sciences
and Engineering Research Council of Canada (NSERC)
Postgraduate Scholarship to S. Chhin, and a research
grant from Global Forest to G. G. Wang. We thank
Drs. R. Staniforth and J. Tardif for their contributions
to the initial research proposal. Thanks also go to K.
Ryan, R. Klos, and M. Kreiner for their assistance in
the field data collection; and K. Schykulski and H.
Hernandez of Manitoba Conservation for their logis-
tical support.
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Received 2 January 2007
Accepted 9 May 2008
|
Natural Recolonization of Cultivated Land by Native Prairie Plants and
its Enhancement by Removal of Scots Pine, Pinus sylvestris
PAUL M. CATLING! and BARRY KING?
' Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Wm. Saunders Building, Central Experimental Farm,
Ottawa, Ontario K1 A 0C6 Canada; e-mail: catlingp @agr.ge.ca
2 217 King St. West, Apartment 175, Cobourg, Ontario K9A 2N2 Canada
Catling, Paul M., and Barry King. 2007. Natural recolonization of cultivated land by native prairie plants and its enhancement
by removal of Scots Pine, Pinus sylvestris. Canadian Field-Naturalist 121(2): 201-205.
A combination of native and introduced plants colonized abandoned cultivated land with adjacent relict prairie and savanna in
the Rice Lake region of southern Ontario. After 71 years, the native colonizers included 86 species found in regional prairie
relicts, but much of the area was also colonized by introduced Scots Pine, Pinus sylvestris, which apparently spread from an
adjacent planting. The pines formed expanding patches of dense growth that excluded other species. Removal of the invasive
Scots Pine from a 200 m? plot within the abandoned land led to colonization 17 years later by 36 native species characteristic
of the prairie, savanna and sand barrens of the region. Included in this group of native prairie colonizers were keystone species
such as Andropogon gerardii, Carex siccata, Ceanothus americanus, Comptonia peregrina and Quercus velutina. Areas where
patches of Scots Pine had been allowed to continue growing were either devoid of vegetation or had a sparse understory of
introduced species and Poison Ivy (Rhus radicans). These observations support the concepts of (1) protecting islands of
native dry ground flora which can serve as sources for recolonization and (2) protection of old field and particularly recently
cultivated land adjacent to protected native grassland to allow natural restoration. The fact that keystone native species were
able to colonize the area from which the introduced Scots Pines had been removed suggests that the pines are aggressive
competitors that occupy space to the exclusion of the native species. Scots Pine is thus a driver of ecological change in degraded
ecosystems. Management including removal of Scots Pine to support native plant biodiversity is strongly supported.
Key Words: Scots Pine, Pinus sylvestris, prairie, sand barren, savanna, recolonization, driver, Andropogon gerardii, Carex siccata,
Ceanothus americanus, Comptonia peregrina, Quercus velutina, Rice Lake, Ontario.
Although information on restoration of prairie veg-
etation is abundant (e.g., Schramm 1992; Packard and
Mutel 1997), information on natural recolonization of
prairie flora following human impacts (e.g., Inouye et
al. 1987; Rabinowitz and Rapp 1985) is relatively
scarce. Despite the lack of information it is often con-
_ sidered a good idea to preserve degraded habitats that
have islands of pristine habitat or have adjacent pris-
_ tine habitats that can serve as sources for natural re-
colonization. It appears in some cases that invasive
alien plants can limit the success of natural recoloniza-
tion in these circumstances but a lack of information is
apparent in this area as well. While some have expressed
concern that invasive plant species are one of the lead-
ing direct causes of biodiversity loss (e.g., Catling
2005), others have questioned the extent to which in-
troduced species threaten biodiversity (e.g., Davis
2003) and several recent articles have recommended
changes in language and attitude in dealing with inva-
Sive species problems based to some extent on the need
for more accurate information (Gobster 2005; Larson
2005). Recently it has been suggested that invasive
plants are the passengers rather than the drivers of eco-
logical change in degraded ecosystems (MacDougal
and Turkington 2005). As “drivers” the invasives would
compete with native species and may also change the
environment to reduce the capability of native species
to survive. As “passengers” they are not the limiting
factor for native species but just along for the ride.
MacDougal and Turkington (2005) suggested that if
interactive processes were limiting, then removal of in-
vasive plants should result in a direct increase in native
species richness (the “driver” model). On the other
hand, if invasive plants are not the limiting factor for
native plants, then removal of the invasive plant(s)
should have little effect (the “passenger” model).
Although common sense suggests that (1) natural
recolonization could occur from pristine nuclei and
(2) that invasive alien plants may be either drivers or
passengers, the cost of managing natural areas requires
stronger evidence. A simple field experiment conduct-
ed in the Rice Lake region of eastern Ontario provides
information relating to these assumptions. It involves
invasive alien Scots Pine (Pinus sylvestris) and a poten-
tially diverse savanna, sand barren and prairie flora.
In situations where a native sand barren, prairie or
savanna has been replaced by a dense stand of Scots
Pine with a completely barren and dark understory (e.g.,
Catling and Carbyn 2005), it is compelling to think of
this invasive as a “driver of ecological change” (Did-
ham et al. 2005). Indeed, Scots Pine is listed as the fifth
most significant invasive alien plant of natural habitats
in Canada (Catling and Mitrow 2005). However, the
present question is whether or not Scots Pine is a driv-
er of ecological change in a degraded ecosystem where
the native prairie vegetation has been reduced by other
201
202
factors. If following perturbation the native prairie
species respond by colonizing an area of previously
cultivated ground and they also respond by colonizing
a portion of that same ground from which invading
Scots Pine is removed, then native prairie plants do
naturally recolonize and the introduced pine is a driver
of ecological change that is clearly in the way since it
is preventing the natural restoration of native flora fol-
lowing the cessation of degrading processes.
Methods
Natural Recolonization
The degraded ecosystem in this experiment origi-
nated from an area of sandy soil that was completely
utilized for the cultivation of cereal crops and espe-
cially potatoes until 1935 when it was abandoned. It
is located at 44.1268°N, 77.9839°W in Lot 4, Conces-
sion 7, originally Haldimand Twp. but now Alnwick/
Haldimand township in Northumberland County, On-
tario. The eradication of native flora is confirmed by
BK whose family occupied the land, and further by
aerial photographs taken in 1929. Cultivation was es-
sentially continuous following settlement in the 1850s,
but prior to this the region had been a rolling prairie
dominated by prairie grasses with localized Black Oak
(Querus relutina) Red Pine (Pinus resinosa) savanna
and sand barren flora (Catling et al. 1992). By 2006 a
combination of native species and alien species, in-
cluding alien Scots Pine, had colonized the previously
cultivated field. It had become a semi-open forest with
50-year-old pines, including the Scots Pine, along with
the native White Pine (Pinus strobus) and both Red
(Querus rubra) and Black oaks, but throughout the
forest were openings of various sizes with a combina-
tion of native prairie species (see Table 1) and inva-
sive alien species, including Hypericum perforatum,
Poa compressa, Hieracium pilloselloides, Rumex ace-
tosella, Potentilla norvegica, and especially Pinus syl-
vestris. Native species that have colonized the aban-
doned potato field are believed to have spread from
the adjacent Russ Creek pioneer cemetery and from
the adjacent Barr property preserve, which was never
cultivated. The early pioneer graveyard dates from
1849 (earliest marker, although it had a schoolhouse
beside it in 1845 and was likely used much earlier)
with few burials after 1900. The prairie plant commu-
nity in pioneer graveyards in prairie regions was usu-
ally not cultivated or grazed unlike much of the sur-
rounding landscape because it was sacred ground.
Since it was only controlled occasionally with a scythe,
the prairie vegetation survived and this seems to be the
case with the Russ Creek Cemetery, which is a nucle-
us Of rare and restricted native prairie species. In 2006
an open (treeless) portion of the recolonized field that
was 0.8 hectares in extent was selected as a sample
plot to document recolonization of native species. All
native plant species present in this sample plot were
recorded during spring, summer and fall visits in 2006.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Effect of removing Scots Pine
The invading Scots Pines appeared to be preventing
the re-colonization of native species in the old field
since they grew in dense stands that contained no other
species of vascular plants. In 1991 a dense stand of
pines was removed from an area of 200 m? adjacent to
the sample plot to allow colonization of native species.
Beneath this stand, and beneath others within 100 m,
the only plants at the time of tree removal were occa-
sional depauperate specimens of Poison Ivy (Rhus rad-
icans) which does well in the decomposing leaf litter
of Scots Pine needles. In 2006 the area from which the
trees were removed was surveyed and all native plant
species present were recorded during spring, summer
and fall visits. Native flora was also recorded from
adjacent stands of Scots Pine (where the trees had not
been removed).
Scientific and common names used in the text gen-
erally follow Kartesz and Meacham (1999). Repre-
sentative specimens are at the AAFC (Agriculture and
Agri-Food Canada) vascular plant herbarium in Ottawa
(acronym — DAO).
Results and Discussion
Natural Colonization
In 2006, 71 years after the old potato field was aban-
doned as cultivated land, a total of 86 species charac-
teristic of natural prairie, savanna and sand barrens in
the region was recorded in the 0.8 hectare sample plot
of an open (treeless) area (Table 1). It is clear that na-
tive species can recolonize seriously degraded habitat
in this region but it is evidently a slow process since
the occurrence of native species was still very patchy
after 71 years despite a relatively uniform substrate and
elevation throughout the area. The return of prairie
flora to the previously cultivated field is to be expect-
ed on the basis of observations elsewhere where dis-
persal and colonization from adjacent protected sites
have proven to be the major factors in old field succes-
sion (e.g., Inouye et al. 1987). The results of this study
support the concept of protecting islands of native dry
ground flora which can serve as sources for recolo-
nization. The results also suggest that even in the north-
eastern-most prairie and savanna habitats, protection
of previously cultivated land adjacent to protected
native grassland is likely to result in an old field suc-
cession that includes many native prairie species. Fur-
thermore, some studies have shown that the native
prairie species increase with time (over 60 years) and
with increasing soil nitrogen at the expense of both
the introduced plants and native non-prairie plants
(Inouye et al. 1987: Figure 8). Thus protection of aban-
doned farmlands adjacent to protected sites is likely
to result in a degree of natural restoration that can be
supported through plantings and management.
Effect of Removing Scots Pine
It is likely that the colonization by native species in
the old potato field would have been more extensive
|
2007 CATLING and KING: RECOLONIZATION OF CULTIVATED LAND 203
TABLE |, Native species of prairie, savanna and sand barren habitats recorded in 2006 that naturally colonized a 0.8 hectare plot
of previously cultivated land abandoned in 1935, The native species recorded in the 200 m* subplot in 2006, Le., 15 years after
removal of invading Scots Pines, are indicated with an asterisk (*).
Amelanchier arborea (Michx.) Fern. var. arborea, DOWNY SERVICE-BERRY
* Amelanchier laevis Wieg., ALLEGHENY SERVICE-BERRY
Amelanchier stolonifera Wieg., RUNNING SERVICE-BERRY
* Andropogon gerardii Vitman, BIG BLUESTEM
Anemone cylindrica Gray, LONG-HEAD THIMBLEWEED
Antennaria howellii ssp. petaloidea (Fern.) Bayer, SMALL PUSSYTOES
* Antennaria parlinii ssp. fallax (Greene) Bayer & Stebbins, PARLIN’S PUSSYTOES
* Apocynum androsaemifolium L., SPREADING DOGBANE
Aquilegia canadensis L., RED COLUMBINE
Arabis divaricarpa A. Nels., ROCKCRESS
Arabis hirsuta var. pycnocarpa (M. Hopkins) Rollins, HAIRY ROCKCRESS
Arabis holboellii var. retrofracta (Graham) Rydb., HOLBOELL’S ROCKCRESS
Arctostaphylos uva-ursi (L.) Spreng., RED BEARBERRY
Artemisia campestris ssp. borealis var. scouleriana (Hook.) Crong., PACIFIC WORMWOOD
Asclepias exaltata L., POKE MILKWEED
* Asclepias syriaca L., COMMON MILKWEED
* Asclepias tuberosa ssp. tuberosa, BUTTERFLY MILKWEED
* Calystegia spithamaea ssp. spithamaea, LOW FALSE BINDWEED
* Carex backii Boott, BACK’S SEDGE
Carex gracillima Schwein., GRACEFUL SEDGE
* Carex molesta Mackenzie ex Bright, TROUBLESOME SEDGE
* Carex muehlenbergii var. muehlenbergii, MUHLENBERG’S SEDGE
Carex pensylvanica Lam., PENNSYLVANIA SEDGE
Carex richardsonii R. Br., RICHARDSON’S SEDGE
* Carex siccata Dewey, DRY-SPIKE SEDGE
* Carex tonsa var. rugosperma (Mackenzie) Crins
* Ceanothus americanus L., NEW JERSEY-TEA
Comandra umbellata ssp. umbellata, BASTARD TOADFLAX
* Comptonia peregrina (L.) Coult., SWEET-FERN
Cornus racemosa Lam., GRAY DOGwWooD
Cyperus houghtonii Torr., HOUGHTON’S FLAT SEDGE
Cyperus lupulinus (Spreng.) Marcks ssp. lupulinus, GREAT PLAINS FLAT SEDGE
* Danthonia spicata (L.) Beauv. ex Roemer & J.A. Schultes, POVERTY WILD Oat GRASS
* Dichanthelium sabulorum (Lam.) Gould & C.A. Clark var: thinium (A.S. Hitche. & Chase) Gould and C.A Clark (D.
columbianum Scribner), HEMLOCK ROSETTE GRASS
Dichanthelium depauperatum (Muhl.) Gould, STARVED ROSETTE GRASS
Dichanthelium linearifolium (Scribn. ex Nash) Gould, SLIM LEAF ROSETTE GRASS
Dichanthelium perlongum (Nash) Freckmann, LONG SLIM LEAF ROSETTE GRASS
Elymus trachycaulus ssp. subsecundus (Link) A.& D. Léve (includes unilaterale), SLENDER WILD RYE
* Fragaria virginiana Duchesne ssp. virginiana, VIRGINIA STRAWBERRY
Galium boreale L., NORTHERN BEDSTRAW
Galium circaezans Michx. var. circaezans, LICORICE BEDSTRAW
Helianthus divaricatus L., WOODLAND SUNFLOWER
Lechea intermedia var. intermedia, ROUND FRUIT PINWEED
Liatris cylindracea Michx., ONTARIO GAYFEATHER
Lilium philadelphicum L. var. philadelphicum, Woop LILy
* Maianthemum canadense Desf., FALSE LILY-OF-THE-VALLEY
* Maianthemum stellatum (L.) Link (Smilacina stellata), STARRY FALSE SOLOMON’S SEAL
Melampyrum lineare Dest. var. lineare, AMERICAN COW WHEAT
Monarda fistulosa L. ssp. fistulosa var. fistulosa, OSWEGO-TEA
* Physalis heterophylla Nees, CLAMMY GROUND CHERRY
Penstemon hirsutus (L.) Willd., HAIRY BEARDTONGUE
Pinus strobus L., EASTERN WHITE PINE
* Poa pratensis L. ssp. pratensis, KENTUCKY BLUE GRASS
* Polygala polygama Walt., RACEMED MILKWORT
Polygonatum biflorum (Walt.) Ell. var. commutatum (J.A. and J.H. Schultes) Morong, KING SOLOMON’ S-SEAL
* Potentilla arguta Pursh ssp. arguta, TALL CINQUEFOIL
Potentilla simplex Michx., OLDFIELD CINQUEFOIL
Prunus pumila L. var. susquehanae (hort. ex Willd.) Jaeger, SUSQUEHANNA SAND CHERRY
* Prunus serotina Ehrh., BLACK CHERRY
Prunus virginiana L. var. virginiana, CHOKE CHERRY
204
TABLE |. (cont.)
THE CANADIAN FIELD-NATURALIST
Vol. 121
Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underwood ex Heller, NORTHERN BRACKEN FERN
Quercus alba L., NORTHERN WHITE OAK
Quercus rubra L., NORTHERN RED OAK
* Quercus velutina Lam., BLACK OAK
Ranunculus rhomboideus Goldie, LABRADOR BUTTERCUP
* Rhus typhina L., SLIM LEAF ROSETTE GRASS
* Rosa acicularis Lindl. ssp. acicularis, PRICKLY ROSE
* Rosa blanda Ait. var. blanda, SMOOTH ROSE
* Rubus idaeus L. ssp. idaeus, COMMON RED RASPBERRY
Rudbeckia hirta L. var. hirta, BLACK-EYED SUSAN
Salix humilis Marsh var. humilis, PRAIRIE WILLOW
Schizachyrium scoparium (Michx.) Nash var. scoparium, LITTLE FALSE BLUESTEM
Shepherdia canadensis (L.) Nutt., RUSSET BUFFALO-BERRY
Solidago canadensis L. var. canadensis, CANADA GOLDENROD
Solidago gigantea Ait., LATE GOLDENROD
* Solidago juncea Ait., EARLY GOLDENROD
* Solidago nemoralis Ait. var. nemoralis, GRAY GOLDENROD
Sorghastrum nutans (L.) Nash, YELLOW INDIAN GRASS
* Symphyotrichum ericoides (L.) Nesom vat. ericoides, WHITE HEATH AMERICAN-ASTER
* Symphyotrichum novae-angliae (L.) Nesom, NEW ENGLAND ASTER
* Symphyotrichum oolentangiense (Riddell) Nesom var. oolentangiense, SKY-BLUE ASTER
* Toxicodendron rydbergii (Small ex Rydb.) Greene, WESTERN POISON Ivy
Viola adunca Sm. var. adunca, HOOK-SPUR VIOLET
* Viola sagittata Ait. var. ovata (Nutt.) Torr. & Gray, ARROW-LEAF VIOLET
Vitis riparia Michx., RIVER-BANK GRAPE
and less patchy if the Scots Pines had not invaded.
Scattered dense stands of Scots Pine were without other
species in 1991 and in 2006. However, within the recol-
onized area of the old field where the Scots Pines were
removed in 1991, 36 native species of prairie, savanna
and sand barren affinity were found in 2006, and had
thus colonized this space within 15 years (Table 1). The
absence of native species under the dense Scots Pines
that remained suggested that this colonization would
not have occurred if the pines had not been removed.
Although introduced species had also colonized this
area from which the Scots Pines were removed, includ-
ing particularly Poa compressa, Hieracium pilosel-
loides and young Scots Pines, the native species were
at least 50% of the cover. Therefore, the pines occupy
space to the exclusion of the native species and restrict
recolonization of disturbed sites. Since only more pines
and Poison Ivy and raspberries Rubus spp. occur in the
older (more than 50 years old) Scots Pine stands in
the region, there is very good evidence that the Scots
Pines interfere with the recolonization of native prairie
species following cessation of degrading processes and
apparently also change the environment to reduce the
capability of native species to survive. In adjacent
stands of Scots Pines the understory was bare or cov-
ered only by a scant growth of Poison Ivy, raspberries,
and introduced plants. The native species that respond-
ed to the removal of the pines included the character-
istic and keystone native species of prairie, savanna
and sand barren such as Andropogon gerardii, Carex
siccata, Ceanothus americanus, Comptonia peregrina
and Quercus velutina. The fact that these native species
were able to colonize the area from which the pines
were removed, but did not where the pines remained,
suggests that the pines are aggressive competitors that
occupy space to the exclusion of the native species and
change conditions by competing for nutrients, reduc-
ing light and changing soil characteristics with leaf lit-
ter accumulation, so that most of the native species of
prairie, savanna and sand barren cannot exist. Scots
Pine in this situation and evidently in many other situa-
tions (e.g., Catling and Carbyn 2005) is thus a driver of
ecological change. Management including removal of
Scots Pine to support native plant biodiversity is strong-
ly supported. Since the trees are relatively slow grow-
ing, slow to mature, reproduce only by seed, are con-
spicuous at all times, and easily eliminated by pulling
and cutting and/or burning, control of Scots Pine is
much less problematic than the control of some other
invasive plant species.
Conclusion
These observations suggest that natural recoloniza-
tion by native prairie species can occur, but that it is
hindered by invasive alien Scots Pine, which drives
ecological change in a degraded prairie ecosystem.
Acknowledgments
Ms. B. Kostiuk provided extensive assistance in
the field survey. Earlier surveys in the area were con-
ducted by D. White and E. Heuvel.
2007
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Catling, P. M., and S. Carbyn. 2005. Invasive Scots Pine,
Pinus sylvestris, replacing Corema, Corema conradii,
heathland in the Annapolis valley, Nova Scotia, Canadian
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Catling, P. M., V. R. Catling, and S. M. McKay-Kuja.
1992. The extent and floristic composition of the Rice
Lake Plains based on historical records. Canadian Field-
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Catling, P. M., and G. Mitrow. 2005. A prioritized list of
the invasive alien plants of natural habitats in Canada.
Canadian Botanical Association Bulletin 38: 55-57.
Didham, R. K., J. M. Tylianakis, M. A. Hutchison, R. M.
Ewers, and N. J. Gemmell. 2005. Are invasive species the
drivers of ecological change? Trends in Ecology and
Evolution 20: 470-474.
Davis, M. A. 2003. Biotic globalization: does competition
from introduced species threaten biodiversity? Bioscience
53: 481-489.
Gobster, P. H. 2005. Invasive species as ecological threat:
is restoration an alternative to fear-based resource manage-
ment? Ecological Restoration 23: 261-270.
Inouye R. S., N. J. Huntly, D. Tilman, J. R. Teste, M. Still-
well, and K. Zinnel.1987. Old-field succession on a Min-
nesota sand plain. Ecology 68: 12-26
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invasion biology. Frontiers in Ecology and Environment
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MacDougal, A. S., and R. Turkington. 2005. Are invasive
species the drivers or passengers of change in degraded
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Packard, S., and C. F. Mutel. Editors. 1997. The tallgrass
restoration handbook: for prairies, savannas, and wood-
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Rabinowitz, D., and J. K. Rapp. 1985. Colonization and
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Received 19 September 2006
Accepted 22 May 2008
Notes
A High Elevation Record of the Star-nosed Mole (Condylura cristata)
in Northeastern Vermont
RYAN W. Norris and C. WILLIAM KILPATRICK
Department of Biology, University of Vermont, Burlington, Vermont 05405 USA;
e-mail: rnorris@zoo.uvm.edu and wkilpatr@ zoo.uvm.edu
Norris, Ryan W., and C. William Kilpatrick. 2007. A high elevation record of the Star-nosed Mole (Condylura cristata) in
northeastern Vermont. Canadian Field-Naturalist 121(2): 206-207.
A single male Star-nosed Mole, Condylura cristata, was captured in a rock pile at the top of East Mountain (elevation 1042
m; 3240 ft) in northeastern Vermont. Although known from high elevations (up to 1676 m; 5500 ft) in southern U.S. states,
this species was not known to occur above 573 m (1880 ft) in the northern part of its range. This record is also important in
confirming that C. cristata has some climbing ability when travelling above ground.
Key Words: Star-nosed Mole, Condylura cristata, elevation, talus, Vermont.
The Star-nosed Mole, Condylura cristata, has the
largest and most northerly geographic range of all
North American moles (Peterson 1966; Hall 1981).
The species ranges from southwest Manitoba to
Labrador and Nova Scotia, down the eastern United
States to South Carolina, and to the Great Lakes region
(Petersen and Yates 1980). The species is known to be
associated with moist soils near marshes and streams
and is an excellent swimmer (Hamilton 1931; Petersen
and Yates 1980). Habitat preference for the Star-nosed
Mole is usually characterized as low wet ground near
bodies of water, swamps, wet meadows, wet spots in
fields or low-lying woods, and dry soils within mixed
hardwoods near water (Godin 1977; DeGraaf and
Yamasaki 2001).
Beane (1995), Laerm et al. (1997), and McCay et al.
(1999) have suggested that C. cristata is quite com-
mon in high elevations in states at the southern edge
of its distribution such as North Carolina and South
Carolina. Elevational records reported for southern
U.S. states include a record at 760 m (2494 ft) in South
Carolina (Laerm et al. 1997) and several individuals
from the Smoky Mountain National Park at elevations
ranging from 488 to 1676 m (1600-5500 ft; Linzey
and Linzey 1971). Literature records from Canada and
northern U.S. states rarely report elevation, but all avail-
able records are lower. The highest elevation reported
at northern latitudes that we have been able to find is
Saunders’s (1988) suggestion that C. cristata ranges
up to 573 m (1880 ft) in elevation in the Adirondacks
of New York.
Study Area and Methods
The summit of East Mountain in northeastern Ver-
mont was modified in 1955 with the construction of the
206
North Concord Air Force Station. The ridgeline was
levelled creating an area of approximately 17 acres on
the summit for this radar station. The rubble from this
modification was pushed to the edges, levelling the
surface and creating substantial areas of talus. The
station was deactivated in 1963; the buildings have
slowly deteriorated, and have been partially disman-
tled. The native fauna and flora have since recolo-
nized the modified area. Dense regenerative spruce-
fir forest now covers much of the summit and small
mammals associated with talus habitat, such as the
Yellow-nosed Vole (Microtus chrotorrhinus), have been
found in this human-made talus habitat.
During a small mammal survey, 30 Sherman live
traps, baited with rolled oats, were set in and adjacent |
to one of several human-made talus piles at the top of
East Mountain (elevation 1042 m or 3420 ft), East |
Haven township, Essex County, Vermont, 44°39.5'N
71°46.1'W over three nights (23-25 August 2004).
Traps were checked daily and rebaited as needed. The
talus pile was approximately 20 m in diameter and
contained bare, flattish rocks ranging from ~ 0.5 m to
3 m in diameter. It was located on a gentle slope and
had small pools of standing water. No larger streams
or bodies of water were observed in the vicinity. Traps ;
in the talus were placed on rocks, not soil, but a small |
quantity of humus had collected in some areas.
Results
A single male C. cristata was discovered dead in a
Sherman trap placed among rocks approximately 5 m
from the edge of the talus pile on the first night of
trapping (23 August). This night saw moderate rain
with nighttime low temperatures at or approaching’
O°C. The measurements of this specimen are: total
2007
length 174 mm, tail length 80 mm, and hind foot length
27 mm. Skin, skull, and tissue are housed in the Zadock
Thompson Natural History Museum at the University
of Vermont (catalog number 5273). Other species cap-
tured over the course of these three nights of trapping
were Short-tailed Shrew, Blarina brevicauda (1);
Smokey Shrew, Sorex fumeus (1), Southern Red-
backed Vole, Myodes gapperi (5), and Deer Mouse,
Peromyscus maniculatus (8).
Discussion
The Star-nosed Mole, Condylura cristata, is clearly
a more northern-adapted species than other moles in
eastern North America. The relatively high number of
reports of this species in mountainous areas in south-
ern states is not surprising considering its distribution
as far north as Labrador. Our capture of the species at
a high elevation site in Vermont was unexpected, par-
ticularly considering that it was taken in a talus pile and
not associated with soil or long-term standing water.
The single specimen reported here is evidence that
the Star-nosed Mole also occurs at high elevations (at
least 1042 m; 3420 ft) even in the northeastern USA.
This capture also suggests that they can live, or travel
through, rocky areas away from soil. Peterson (1966)
suggested that Star-nosed Moles spend more time
above ground than other mole species and, as a result,
are occasionally found dead on the surface or captured
on the surface with snap or live traps.
The capture of the mole amongst rocks and not on
bare soil suggests that it must have been maneuvering
within this rock pile. Hickman (1982) reported observ-
ing a vertical climb of 60 cm by a Star-nosed Mole in
the corner of a concrete column, and Schmidt (1931)
reported a Star-nosed Mole climbing two feet out of
a swimming tank. Our capture provides further indica-
tion that C. cristata is capable of some degree of climb-
ing, a feature not often attributed to fossorially adapted
mammals. Hickman (1982) hypothesized that the
climbing ability of C. cristata may serve to aid in mo-
bility across rocky areas. Our capture is also consis-
tent with this idea.
NOTES
207
Acknowledgments
The authors thank Charles A. Woods for important
discussion leading to the preparation of this manu-
script. This specimen was collected in association with
a small mammal survey funded by the Kingdom Com-
mons Group.
Literature Cited
Beane, J. C. 1995. New distributional record for the star-
nosed mole, Condylura cristata (Insectivora: Talpidae),
in North Carolina, with comments on its occurrence in the
Piedmont Region. Brimleyana 22: 77-86.
DeGraaf, R. M., and M. Yamasaki. 2001. New England
Wildlife: Habitat, Natural History, and Distribution. Uni-
versity Press of New England, Hanover, New Hampshire
482 pages.
Godin, A. J. 1977. Wild Mammals of New England. Johns
Hopkins University Press, Baltimore, Maryland. 304 pages.
Hall, E. R. 1981. The mammals of North America. Volume
1. John Wiley and Sons, New York. 600 pages.
Hamilton, W. J., Jr. 1931. Habits of the star-nosed mole,
Condylura cristata. Journal of Mammalogy 12: 345-355.
Hickman, G. C. 1982. Climbing ability of the star-nosed
mole, Condylura cristata (Talpidae). Saugetierkundliche
Mitteilungen 30: 296-297.
Laerm, J., G., Livingston, C. Spencer, and B. Stuart. 1997.
Condylura cristata (Insectivora: Talpidae) in the Blue Ridge
Province of western South Carolina. Brimleyana 24: 46-49.
Linzey, A. V., and D. W. Linzey. 1971. Mammals of Great
Smoky Mountains National Park. University of Tennessee
Press, Knoxville. 114 pages.
McCay, T. S., M. J. Komoroski, and W. M. Ford. 1999. Use
of an upland pine forest by the star-nosed mole, Condylura
cristata. The Journal of the Elisha Mitchell Scientific Soci-
ety 115: 316-318.
Petersen, K. E., and T. L. Yates. 1980. Condylura cristata.
Mammalian Species 129: 1-4.
Peterson, R. L. 1966. The mammals of eastern Canada. Ox-
ford University Press, Toronto. 465 pages.
Saunders, D. A. 1988. Adirondack Mammals. State Univer-
sity of New York. 216 pages.
Schmidt, F. K. W. 1931. Mammals of Western Clarke County,
Wisconsin. Journal of Mammalogy 12: 99-117.
Received 22 February 2006
Accepted 8 February 2008
208 THE CANADIAN FIELD-NATURALIST Vol. 121
Northern Long-eared Bat, Myotis septentrionalis (Chiroptera:
Vespertilionidae), on Prince Edward Island: First Records of
Occurrence and Over-Wintering
JENNIFER A. BRowN!, DONALD F. MCALPINE“, and ROSEMARY CURLEY?
University of Guelph, Department of Animal and Poultry Science, Guelph, Ontario N1G 2W1 Canada
’New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada (author to whom correspondence
should be addressed; e-mail: donald.mcalpine @nbm-mnb.ca)
Forests, Fish and Wildlife Division, Prince Edward Island Department of Environment, Energy, and Forestry, P.O. Box 2000,
Charlottetown, Prince Edward Island C1A 7N8 Canada
Brown, Jennifer. A., Donald F. McAlpine, and Rosemary Curley. 2007. Northern Long-eared Bat, Myotis septentrionalis
(Chiroptera: Vespertilionidae), on Prince Edward Island: first records of occurrence and over-wintering. Canadian
Field-Naturalist 121(2): 208-209.
First occurrence of the Northern Long-eared Bat, Myotis septentrionalis, on Prince Edward Island is reported. A mixed-
species hibernaculum of M. septentrionalis and the Little Brown Bat, M. lucifugus, is also described from southeastern
Prince Edward Island. This is the first record of bats over-wintering in the province and the first time either species has been
reported making use of a building as a hibernaculum.
Key Words: Little Brown Bat, Myotis lucifugus, Northern Long-eared Bat, Myotis septentrionalis, Prince Edward Island,
distribution, hibernaculum.
The bats of Prince Edward Island are poorly known.
Although seven species have been recorded from the
adjacent mainland of New Brunswick and Nova Sco-
tia (van Zyll de Jong 1985), only the Little Brown Bat
(Myotis lucifugus) and Hoary Bat (Lasiurus cinereus)
are known from the Island (van Zyll de Jong 1985;
McApine et al. 2002). Cameron (1958) reported a sight-
ing which he believed to be a Red Bat (Lasiurus bore-
alis) near coastal Fortune Bridge, Kings County, Prince
Edward Island (46°21'23"N 62°29'57"W) on 7 June
1954, but this species remains unconfirmed on the Is-
land. Neither have any bat species been documented
to hibernate on Prince Edward Island. Here we docu-
ment the first occurrence of the Northern Long-eared
Bat (Myotis septentrionalis) from the province and
report a mixed-species hibernaculum of M. lucifugus
and M. septentrionalis from southeastern Prince Ed-
ward Island.
On 24 September 1988, two bats retrieved by a
house cat (Felis domesticus) were collected in
Breadalbane, Queens County, Prince Edward Island
(46°21'23"N 63°29'57"W) by JAB and subsequently
identified as M. septentrionalis. These specimens,
deposited in the Canadian Museum of Nature (CMN),
include a male (CMN 52612) and a female (CMN
52613). Measurements for both bats (M (male), TL
(total length) 85 mm, T (tail) 37 mm, HF (hind foot)
10 mm; F (female), TL 90 mm, T 36 mm, HF 10 mm)
fall within the range for the species presented by van
Zyll de Jong (1985).
On or about 10 January 1989 a commercial pest con-
trol officer removed about 800 bats, both Little Brown
and Northern Long-eared bats, from the basement of a
home at Culloden, Queens County, Prince Edward
Island (46°0'58"N 62°42'8"W; reported in the media as
Murray River, the larger nearby community). All bats
were destined to be incinerated. However, DFM was
able to secure 781 of these bats alive and an unsuc-
cessful attempt was made to maintain them in a state
of torpor over the winter with the intention of spring
release. Various media reports suggest that the hiber-
naculum contained 1500 bats (Anonymous 1989*;
Cody 1989 a*, b*; White 1989*). However, communi-
cation with the pest control officer (personal commu-
nication to DFM) confirmed that all hibernating bats
observed were removed. Information presented here is
based on an examination of all 781 bats, apparently
the contents of virtually the entire hibernaculum. We
visited the hibernaculum site on 29 November 1990
and found only two torpid female M. septentrionalis,
clinging to wooden floor joists. A voucher series of
both species from the site have been deposited in the
mammal collection of the New Brunswick Museum.
The hibernation site was the unheated basement of
an occupied home, built about 1980. The house had
been recently re-occupied following a 3-5 year vacancy
(resident personal communication to DFM). The base-
ment had an earthen floor and island stone and mortar
foundation walls with inside dimensions about 11 x 8 m
and a ceiling height of 1-2 m. Entry to the basement
was via a trap door on the first floor. Air temperature
in the hibernaculum recorded on the night of 29 No-
vember 1990 was 11.3°C. This hibernation tempera-
ture, while above the mode of 6°C for vespertilionids,
falls within the range of -4° to 13°C and 0.6°C to
13.9°C reported for Little Brown and Northern Long-
eared bats; respectively (Webb et al. 1996). The hiber-
naculum was dominated by the Little Brown Bat
(82.6%); with smaller numbers of the Northern Long-
eared (17.4%). In both species sex ratios were slightly
]
4
at
ees
2007
female biased (62.1% and 57.1% in M. lucifugus and
M. ‘septentrionalis, respectively). We also assessed
reproductive status by dissection of all Northern
Long-eared Bats following the criteria and terminol-
ogy of Sluiter and Bouman (1951), Sluiter (1954,
1961), and Racey (1974), with the following results:
males; 5.9% sexually immature, 94.1% sexually
mature; females; 17.5% nulliparous, 14.2% non-
parous, 68.3% parous.
The Northern Long-eared Bat is widespread across
Atlantic Canada (van Zyll de Jong 1985). Although
less abundant, apparently, in the region than the Little
Brown Bat, the presence of the Northern Long-eared
Bat on Prince Edward Island is not unexpected. Cullo-
den and Breadalbane are 67 km straight-line distance
apart, suggesting M. septentrionalis may be widespread
in the province. Conversely, the discovery of large num-
bers of hibernating bats on the Island was unantici-
pated. The province is underlain largely by Permian
sandstones, siltstone and glystone (van de Poll 1983)
and we are aware of no natural caves or abandoned
mines on the Island that might provide suitable hiber-
nacula. The closest natural caves serving as bat hiber-
nacula occur in adjacent New Brunswick and Nova
Scotia (McAlpine 1983; Mosely 2007). Fenton and
Barclay (1980) were unable to locate any records of
Little Brown Bats hibernating in buildings. Likewise,
the Northern Long-eared has previously only been
recorded hibernating in caves and mines (van Zyll de
Jong 1985). Both these species of Myotis may be more
flexible in their hibernation requirements than the lit-
erature suggests, being able to rapidly exploit suit-
able artificial hibernacula where present.
Acknowledgments
We are grateful to the late C. G. van Zyll de Jong,
both for his encouragement and for confirming the
identity of the two bats collected at Breadalbane. We
also thank the resident of Culloden who so graciously
allowed us access to her home, and Earl Peterson, Mari-
time Varmint and Rodent Control, for agreeing to make
the bats removed from this home available to DFM.
Documents Cited [marked* in text]
Anonymous. 1989. Going batty (photo with caption). The
Telegraph Journal, Saint John, New Brunswick, 14 January
1989, Page 1.
NOTES
209
Cody, J. 1989a. P.E.1. bats likely to become “ex-bats”: people
curious about their fate. The Guardian, Charlottetown,
Prince Edward Island, 19 January 1989, Page 6
Cody, J. 1989b. Bat removal poses question of animal's
right to live. The Evening Patriot, Charlottetown, Prince
Edward Island, 18 January 1989, Page 3
White, D. 1989. Bats take up residence in Culloden basement.
The Times-Transcript, Moncton, New Brunswick
Literature Cited
Cameron, A. W. 1958. Mammals of the islands in the Gulf of
St. Lawrence. Bulletin 154, National Museum of Canada,
Ottawa.
Fenton, M. B., and M. R. Barclay. 1980. Myotis lucifugus.
Mammalian Species 142: 1-8.
McAlpine, D. F. 1983. Status and conservation of solution
caves in New Brunswick. New Brunswick Museum. Pub-
lications in Natural Science (1). 28 pages.
McAlpine, D. F, F. Muldoon, and A. I. Wandeler. 2002.
First record of the Hoary Bat, Lasiurus cinereus (Chi-
roptera: Vespertilionidae), from Prince Edward Island.
Canadian Field-Naturalist 116: 124-125.
Mosely, M. 2007. Acadian biospeleology: composition and
ecology of cave fauna of Nova Scotia and southern New
Brunswick, Canada. International Journal of Speleology
36: 1-21.
Racey, P. A. 1974. Ageing and assessment of reproductive
status of Pipistrellus bats, Pipistrellus pipistrellus. Jour-
nal of Zoology 173: 264-271.
Sluiter, J. W. 1954. Sexual maturity in bats of the genus
Myotis I. Females of M. mystacinus and supplementary
data on female M. myotis and M. emarginatus. Koninklijke
Nederlandsche Academie van Wetenschappen 57: 696-700.
Sluiter, J. W. 1961. Sexual maturity in males of the bat Myoris
myotis. Koninklijke Nederlandsche Academie van Weten-
schappen 64: 243-249.
Sluiter, J. W., and M. Bouman. 1951. Sexual maturity in bats
of the genus Myotis I. Size and histology of the repro-
ductive organs during hibernation in connection with age
and wear of the teeth in female Myotis myotis and Myotis
emarginatus. Koninklijke Nederlandsche Academie van
Wetenschappen 54: 595-603.
van de Poll, H. W. 1983. Geology of Prince Edward Island.
Province of Prince Edward Island, Department of Energy
and Forestry. Report $3-1.
van Zyll de Jong, C. G. 1985. Handbook of Canadian Mam-
mals 2: Bats. National Museum of Natural Sciences,
Ottawa.
Webb, P. I, J. R. Speakman, and P. A. Racey. 1996. How hot
is a hibernaculum? A review of the temperature at which
bats hibernate. Canadian Journal of Zoology 74: 761-765.
Received 9 May 2007
Accepted 14 January 2008
210
THE CANADIAN FIELD-NATURALIST
Vol. 121
Arboreal Late Summer Courtship Behaviour of Maritime Garter Snake,
Thamnophis sirtalis pallidulus, in Dartmouth, Nova Scotia, Canada
JOHN GILHEN! and ROBERT STRUM2
' Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada; e-mail: gilhenja
@gov.ns.ca
2 325 Prince Albert Road, Dartmouth, Halifax County, Nova Scotia B2Y 1N5 Canada
Gilhen, John, and Robert Strum. 2007. Arboreal late summer courtship behaviour of Maritime Garter Snake, Thamnophis sirtalis
pallidulus, in Dartmouth, Nova Scotia, Canada. Canadian Field-Naturalist 121(2): 210-211.
We report, for the first time, Maritime Garter Snake, Thamnophis sirtalis pallidulus, arboreal, late summer, courtship behaviour in the
wild observed 4-5 September 2006.
Key Words: Maritime Garter Snake, Thamnophis sirtalis pallidulus, arboreal, late summer, courtship behaviour, Dartmouth,
Nova Scotia, Canada.
On the morning of 5 September 2006 Robert Strum
reported to John Gilhen that a ball of snakes had
gathered on a Rhododendron shrub, Rhododendron sp..,
in his yard at 325 Prince Albert Road, Dartmouth, Nova
Scotia. In the afternoon of the previous day, 4 Sep-
tember 2006, Robert Strum could see, from the kitchen
window, that a number of brown snakes had gathered
inside a gazebo style bird feeder on the Rhododendron
shrub and he was successful in moving them out. The
following morning, 5 September 2006, he noticed the
snakes had gathered, again, on a branch near the bird
feeder, about two meters above the ground (Cover).
Upon investigation it was determined that there were
four (three males and one female) Maritime Garter
Snakes, Thamnophis sirtalis pallidulus, involved in
courtship behaviour: male number 1, the largest male;
males 2 and 3 about equal total length; and 1 female
(the largest of the four snakes). The weather at 1200
hours on 5 September 2006 was cloudy and overcast
with ambient air temperature of 20°C and temperature
near the ground was also 20°C.
At 1215 hours the males were observed moving
alongside and over the back of the female. At the same
time the body of male number | was pulsating rhyth-
mically. This male rubbed his chin along the sides of
the female. At this time the tail and about one half of
the female’s posterior trunk (about 20 cm) drooped
downward from one of the supporting branches, with
the tails of all three males wrapped around her tail.
While the female’s posterior body and tail were dang-
ling down, the most active male (male number 1) ap-
peared to maintain ventral apposition to copulate with
her (Figure 1).
All four snakes remained on the Rhododendron shrub
from the morning to early afternoon but dispersed at
some time late in the afternoon, as they were not seen
there again that evening or in the following days. It is
not known if copulation occurred over the two-day
period these snakes occupied the Rhododendron shrub.
One of the two subordinate males was collected as a
voucher specimen (NSM55388).
Courtship behaviour of the Maritime Garter Snake
in captivity has been previously observed by John
Gilhen in late August (unpublished). On two separate
occasions (August 2001 and August 2004), days after
individual females had given birth, two males were
observed following alongside the female and trying to
attain cloacal apposition. Although other adult females
were present in the enclosure, the males were only in-
terested in courting with a recently spent female. These
observations of captives and the present observation in
the wild suggests that the Maritime Garter Snake mates
in late summer, sometime after females give birth, but
prior to hibernation.
Galbraith (2001) reports similar arboreal late sum-
mer courtship by Eastern Garter Snakes, Thamnophis
sirtalis sirtalis. Cook (1984) mentions snakes in the
genus Thamnophis most frequently copulate in spring
but have occasionally been noted to copulate in late fall,
immediately prior to entering hibernacula. Arboreal
spring breeding activity of the Red-sided Garter Snake,
Thamnophis sirtalis parietalis, was observed on 8 May
1972 near Inwood, Manitoba (Gregory 1975).
Rossman et al. (1996) and Ernst and Ernst (2003)
agree mating of Thamnophis sirtalis usually occurs in
spring after emergence from hibernation but may also
take place in autumn. Rossman et al. (1996) also states
that, until recently, most of the studies of mating be-
havior and the factors that regulate the timing of re-
production in T. sirtalis have focused primarily if not
exclusively on spring activities. Previous accounts of
the species in Nova Scotia (Barnes et al. 2006; Gilhen
1984, 2000), have not noted fall mating.
The urbanization of both Halifax and Dartmouth
cities over the past two decades has been extensive
leaving many isolated pockets of amphibians and rep-
tiles. The Maritime Garter Snakes at the observation
site in Dartmouth continue to survive in small numbers
in a back yard landscape, surrounded by apartment
buildings on the south side, a major city road on the
north side and other homes on the east and west sides.
|
Galbraith’s (2001) observation of the Eastern Garter |
|
2007 NOTES 2]
Ficure |. An adult male Maritime Garter Snake, Thamnophis
sirtalis pallidulus, seeking ventral apposition with
adult female in courtship behavior on the branches
of a Rhododendron shrub at 325 Prince Albert Road,
Dartmouth, Nova Scotia, on 5 September 2006.
Snake also took place on private land in a cottage
community on the coast sand dune area of the eastert
shore of Lake Huron in Ontario
In Nova Scotia, the Maritime Garter Snake is most
common along the shores of lakes, ponds and stream
but has readily adapted to a host of urban habitat
Robert Strum has commonly observed Maritime
Garter Snakes in his yard and has noted they move
down along the base of the swimming pool and prob
ably hibernate under the pool.
Acknowledgment
The authors appreciate the advice offered by Andrew
Hebda, Curator of Zoology, Nova Scotia Museum of
Natural History, during the preparation of the manu-
script.
Literature Cited
Barnes, S., C. M. Dubesky, and T. B. Herman. 2006
Ecology and morphology of Thamnophis sirtalis pallidulus
(Maritime Garter Snake) on Georges Island, Nova Scotia
Northeastern Naturalist 13(1): 73-82.
Cook, F. R. 1984. Introduction to Canadian amphibians and
reptiles. National Museum of Natural Sciences, Ottawa,
Ontario. 200 pages.
Ernst, C. H., and E. M. Ernst. 2003. Snakes of the United
States and Canada. Smithsonian Books, Washington and
London. 668 pages.
Galbraith, D. A. 2001. Arboreal courtship behaviour by
Eastern Garter Snakes, Thamnophis sirtalis sirtalis, in Sep-
tember in Bruce County, Ontario. Canadian Field-Naturalist
115: 347-348.
Gilhen, J. 1984. Amphibians and reptiles of Nova Scotia
Nova Scotia Museum. Halifax. 162 pages.
Gilhen, J. 2000. Amphibians and reptiles of Nova Scotia
Species recorded in the accession books of Harry Piers
from 1899 to 1939. Manuscript Report. Nova Scotia Mu-
seum of Natural History.
Gregory, P. T. 1975. Arboreal mating behaviour in the Red-
sided Garter Snake. Canadian Field-Naturalist 89: 461-462.
Rossman, D. A., N. B. Ford, and R. A. Seigel. 1996. The
garter snakes: Evolution and ecology. University of Okla-
homa Press, Norman, Oklahoma.
Received 2 November 2006
Accepted 11 March 2008
PWD THE CANADIAN FIELD-NATURALIST Vol. 121
Human-assisted Movements of Raccoons, Procyon lotor, and Opossums,
Didelphis virginiana, between the United States and Canada
Rick RosaTTE!, DENNIS DONOVAN!, MIKE ALLAN!, LAURA Bruce!, and CHRIS Davies!
‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, 2140 East Bank Dr.,
Peterborough, Ontario K9J 7B8, Canada
*Corresponding author (e-mail: rick.rosatte @ ontario.ca) Tel. (705) 755-2280, Fax (705) 755-1559
Rosatte, Rick, Dennis Donovan, Mike Allan, Laura Bruce, and Chris Davies. 2007. Human-assisted Movements of Raccoons,
Procyon lotor, and Opossums, Didelphis virginiana, between the United States and Canada. Canadian Field-Naturalist
121(2): 212-213.
Movements of Raccoons (Procyon lotor) (x = 479 km) and Opossums (Didelphis virginiana) (xX = 688 km) by means of transport
trailers and a train from the USA and Quebec into Ontario were significantly greater than Raccoon movements (x = 15 km)
from Ontario to New York State determined by mark-recapture. Human-assisted movements of wildlife could have significant
impacts with respect to cross-border movements of diseases such as rabies. Proactive communication programs, especially at
International border crossing areas, should be encouraged to decrease the occurrence of the unintentional movement of wildlife
and associated diseases.
Key Words: Didelphis virginiana, Opossum, Procyon lotor, Raccoon, movement, disease, rabies, Ontario.
The movement of wildlife and domestic animals by
humans has resulted in the national, international, and
sometimes global movement of diseases such as rabies,
West Nile virus, bovine tuberculosis, parvovirus, chron-
ic wasting disease (CWD), and bovine spongiform en-
cephalopathy (BSE) (Jenkins and Winkler 1987; Niel-
son and Brown 1988; Woodford and Rossiter 1993;
Williams and Barker 2001). The international move-
ment of Raccoons (Procyon lotor) from the United
States to Ontario, Canada, has been documented previ-
ously (Rosatte et al. 1997, 2001); however, those pub-
lications did not document the distances moved or
geographic areas involved.
During January 1997 and December 2005, a total of
14 Raccoons were documented as having moved across
the St. Lawrence and Niagara Rivers from Ontario,
Canada, to New York State. Those Raccoons were ear-
tagged and released in Ontario as part of a wildlife
rabies control program during 1997-2005 (Rosatte et
al. 1997, 2001). They were later recaptured or found
dead in the vicinity of Buffalo or Ogdensburg, New
York (Figure 1). Exact locations were known for 13 of
those. Mean movement of the 13 Raccoons was 15 km
(SD=9.0) (range = 3-30 km). The sex and age were
known for 11 of the 13 Raccoons. More adult males
(9) than adult females (2) moved from Ontario to New
York (Chi Square = 8.9; P<0.0028). Mean movement
(18 km, SD=8.4) (range = 6-30 km) of adult male
Raccoons was greater than mean movement of adult
females (7.5 km, SD=6.4) (range=3-12 km) (t=1.65;
e—(at))
In addition, during 1998-2005, a total of 14 Rac-
coons and two Opossums (Didelphis virginiana) were
found in transport trailers and in a box car on a train
in the greater Toronto area. The point of departure of
those vehicles included Ohio, Ilinois, Tennessee, New
York, Michigan, Quebec, and Ontario (Figure 1). In
most cases the trailer or container was locked and sealed
at the point of departure and was not opened until it
arrived in Ontario. Mean movement of Raccoons by
vehicle from the USA and Quebec (n=11) to Ontario,
Canada, was 479 km (SD=250) (range=250-890 km).
This was significantly greater (t=6.9, P<0.00002) than
movements of Raccoons noted above that did not in-
volve transport via vehicles. The two Opossums moved
1000 km from Tennessee and 376 km from Michigan,
respectively.
In Ontario, rabid Raccoons have been documented
as having moved 0.7-4.1 km (Rosatte et al. 2005, 2006).
Median movements of non-rabid Raccoons in Ontario
were generally less than 1 km, with movements rarely
exceeding 50 km in distance (Rosatte 2000; Rosatte
et al. 2006). However, those movements were assumed
not to have been due to human involvement (intentional
or otherwise) (i.e. by vehicles). The exceptional move-
ments for Raccoons, as well as Opossums, documented
in this paper, indicate that those species have the poten-
tial to move infectious diseases such as rabies signifi-
cant distances. In view of this, people should be vigi-
lant for wildlife such as Raccoons riding on vehicles.
Transport trailer drivers (as well as operators of other
vehicles such as ferries, trains, boats) should be encour-
aged to check their vehicles and trailers for wildlife
prior to departure to minimize the unintentional move-
ment of animals. This will assist in decreasing the
movement of wildlife diseases. Signage at internation-
al border crossing areas advising drivers to check their
vehicles for wildlife should also be posted. These tac-
tics were used during the 1990s in Ontario in an attempt
to prevent the movement of Raccoon rabies from the
United States to Canada (Rosatte et al. 1997).
Acknowledgments
Special thanks to Brad Gates, AAA Wildlife Con-
trol, Scarborough, Ontario; Scarborough Animal Con-
trol; Mississauga Animal Control staff; Dan Lawrence,
|
|
)
|
|
|
|
q
2007
Animal Control, Markham; Brampton Animal Control;
Art King, Canadian Food Inspection Agency, Ft. Erie,
Ontario; Rick Stevens and Ted Smith New York State
Department of Environmental Conservation; James
Lucas, Lockport, New York; Laura Bigler, Cornell Uni-
versity; Mark Carrera, United States Department of
Agriculture, Potsdam, New York; and Jim Masters,
Niagara County Health Department, New York. The
manuscript was reviewed by Mark Gibson, Tore
Buchanan, and Kirk Sobey, Ontario Ministry of Nat-
ural Resources, Peterborough, Ontario.
Literature Cited
Jenkins, S., and W. Winkler. 1987. Descriptive epidemiol-
ogy from an epizootic of raccoon rabies in the Middle
Atlantic States, 1982-1983. American Journal of Epidemi-
olgy 126: 429-437.
Nielson, L., and R. Brown. 1988. Translocation of wild ani-
mals. Wisconsin Humane Society and the Caesar Kleberg
Wildlife Research Institute publishers. 333 pages.
Rosatte, R. 2000. Management of raccoons (Procyon lotor)
in Ontario, Canada: Do human intervention and disease
have significant impact on raccoon populations? Mam-
malia 64 : 369-390.
Rosatte, R., C. MacInnes, R. Taylor Williams, and O.
Williams. 1997. A proactive prevention strategy for rac-
coon rabies in Ontario, Canada. Wildlife Society Bulletin
25: 110-116.
Rosatte, R., D. Donovan, M. Allan, L. Howes, A. Silver,
K. Bennett, C. MacInnes, C. Davies, A. Wandeler, and
B. Radford. 2001. Emergency response to raccoon rabies
introduction into Ontario. Journal of Wildlife Diseases
37: 265-279.
Rosatte, R. M. Allan, R. Warren, P. Neave, T. Babin, L.
Buchanan, D. Donovan, K. Sobey, C. Davies, F. Mul-
doon, and A. Wandeler. 2005. Movements of two rabid
Raccoons, Procyon lotor, in eastern Ontario, Canada.
Canadian Field-Naturalist 119: 453-454.
Rosatte, R., K. Sobey, D. Donovan, L. Bruce, M. Allan, A.
Silver, K. Bennett, M. Gibson, H. Simpson, C. Davies,
A. Wandeler, and F. Muldoon. 2006. Behavior, move-
ments and demographics of rabid raccoons in Ontario,
Canada: management implications. Journal of Wildlife Dis-
eases 42: 589-605.
Williams, E. S., and I. K. Barker. 2001. Infectious diseases
of wild mammals. Iowa State Press. 558 pages.
NOTES 213
FIGURE |. Map depicting Raccoon movement to and from
Ontario, Canada. Direction of arrows represents direc-
tion of Raccoon movements. Arrows depicting move-
ments of > 30 km represent Raccoon movement via
transport trailers. One arrow in Michigan represents
Raccoon movement on a train. Arrows depicting move-
ments of < 30 km probably represent natural Rac-
coon movements (i.e. not human assisted). Locations
of arrows are approximate.
Woodford, M., and P. Rossiter. 1993. Disease risks associ-
ated with wildlife translocation projects. Revue Scientific
et Technique de L’ Office International des Epizootics. 12:
115-135.
Received 10 April 2006
Accepted 21 May 2008
214 THE CANADIAN FIELD-NATURALIST Vol. 121
Correlation Between Age Estimates for Elk, Cervus Elaphus, Using
Tooth Wear/Eruption Patterns and Counts of Annuli in Tooth Cementum
Rick RosAtTE!“*, ANDREW SILVER!, MARK GIBSON!, BRUCE CHISHOLM, and NorRM CooL?
‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, DNA Building,
2140 East Bank Drive, Peterborough, Ontario K9J 7B8 Canada
*Canadian Food Inspection Agency, Edmonton, Alberta, T6H 5T6 Canada
3Parks Canada, Elk Island National Park, Fort Saskatchewan, Alberta T8L 2N7 Canada
4Corresponding author e-mail: rick.rosatte @ ontario.ca
Rosatte, Rick, Andrew Silver, Mark Gibson, Bruce Chisholm, and Norm Cool. 2007. Correlation between age estimates for Elk,
Cervus elaphus, using tooth wear/eruption patterns and annuli in tooth cementum. Canadian Field-Naturalist 121(2):
214-215.
Two different techniques, (1) tooth wear, tooth eruption patterns and other attributes and (2) counting annuli and seasonal growth
zones in the cementum of incisor teeth, were used to estimate the age of Elk, Cervus elaphus, that were relocated from Alberta
to Ontario, Canada. Age estimates for Elk ranged from 2 to 20 years, and a significant relationship was found between the ages
acquired by the two methods. Critical to acquiring accurate age estimates for Elk were staff with extensive skill with respect to
the aging technique utilized.
Key Words: Elk, Cervus elaphus, age estimates, annuli, cementum, tooth wear.
During 1998-2001, 443 Elk (Cervus elaphus) were
acquired from Elk Island National Park (EINP), Al-
berta, Canada, and released in four different areas of
Ontario, Canada (Rosatte et al. 2002; Rosatte et al.
2007). Elk were aged according to tooth wear, eruption
patterns (deciduous and permanent teeth), colouring
and staining of teeth, as well as by comparison to a
reference sample of known-age Elk jaws, during pro-
cessing at EINP (Hudson et al. 2002) (Figure 1). By
about two years of age the upper pair of permanent in-
cisiform canines erupt (the ivory) and by about 3 years
of age, the 4 pairs of permanent incisors have erupted.
The weight of the animal was also considered when
estimating the age of Elk at EINP; e.g., the weight of
7-month-old calves, 1¥2-year-old, and = 22-year-old
Elk was approximately 100-120 kg, 130-170 kg, and
> 180 kg (+ 10 kg), respectively. The weights of the
Elk were checked following age estimation to ensure
that the age estimate fell into the above age/weight
categories.
During 1999-2004, incisor teeth were collected op-
portunistically from Elk that died in Ontario post-
release. These teeth were aged via counting annuli and
seasonal growth zones in tooth cementum (Figure 2).
The annuli or rings in the cementum represent yearly
growth. By counting the number of annuli in the
cementum an estimate of the animals age can be det-
ermined (Johnston et al. 1999). The age estimates were
compared with those obtained during processing at
EINP after the tooth wear age estimates were extrap-
olated to coincide with the time of death of the Elk in
Ficure 1. Photo of an Elk (Cervus elaphus) mandible showing
incisor, premolar, and molar teeth.
Outer edge
of tooth
FiGurE 2. Longitudinal section (decalcified, Harris modified
hematoxylin stain using bright field illumination) of an
incisor tooth (100x magnification) from an Elk (Cervus
elaphus) depicting annual growth lines (annuli) in the
cementum. Estimated age is 20 years.
sue and soaked in 70% ethanol for 5 minutes to prevent
Ontario. Staff involved in aging had many years of ex-
perience with the aging technique.
Incisor teeth were extracted (via tooth extractors)
from a sample (7 = 10) of Elk that died in Ontario
post-release. Two yearling bulls, for which the age was
known, served as controls. Teeth were cleaned of tis-
the transmission of infectious pathogens. They were
then preserved in 10% formalin or frozen at -20°C.
Teeth were sectioned using an Isomet double-bladed
saw (Buehler Isomet, Techmet, Scarborough, Ontario)
according to Johnston et al. (1999). Each section (60-
100u) was mounted on a microscope slide with Per-
2007
mount and covered with a cover slip. Two microscopic
techniques were used to determine an age estimate for
each incisor. To distinguish juveniles from adults, the
undecalcified, unstained sections were viewed under
a compound light microscope (SOx and 100x magni-
fication) using polarized light to view seasonal grow
zones. In the majority of sections (especially older ani-
mals), the annuli were not distinct and not easily dis-
cerned under polarized light. These were decalcified
(according to Johnston et al. 1999), stained (Harris
modified hematoxylin), and placed in 70% ethanol for
2-3 minutes. Slides were then air-dried, covered with
Permount and a cover slip, and examined under the
microscope (unpolarized light) to count the number
of annuli to obtain an estimate of age (Johnston et al.
1999) (Figure 2).
Age estimates for Elk acquired by tooth wear (and
other indicators as noted above) and annuli or seasonal
growth zone counts in incisor teeth, ranged between
2 and 20 years of age (Figure 3). The first week of June
was used as the birth date for Elk (Hudson et al. 2002).
A simple linear correlation and a f statistic (Zar 1974)
were used (Statistica 6.0 software, StatSoft, Tulsa,
Oklahoma, USA) to measure the relationship and level
of significance between tooth wear and counting annuli
for age determination in Elk. The correlation was found
to be significant (P < 0.0001; t = 20.1; r = 0.9903;
7 = 0.98). There was a positive slope of 1.09 between
the ages acquired by the two methods (Y = 1.0918
x = 1.261) (Figure 3).
This study examined the relationship between two
methods used to estimate the ages of wild Elk. Tooth
wear estimates of age may be inexact as tooth wear
could be affected by diet, soil type and substrate (Hud-
son et al. 2002). However, as the Elk at EINP were in an
enclosed (fenced) situation, these variables were fairly
constant as their geographical location remained un-
changed among years. Therefore, we believe the age
estimates acquired by tooth wear (and other attributes
that were examined) were fairly accurate. Some authors
(Keiss 1969; Hamlin et al. 2000; Hudson et al. 2002)
suggest that staining sections of teeth and counting an-
nuli provides the most reliable estimate of age for Elk.
Furthermore, Keiss (1969) and Hamlin et al. (2000)
only found about a 50% agreement between tooth erup-
tion/wear and cementum annuli counts and concluded
that wear/eruption estimates are unreliable for aging
Elk. On the contrary, in our study, although the sam-
ple size was small, age estimates for Elk obtained by
tooth wear correlated well with those acquired by
counting annuli in stained and unstained sections of
incisor teeth. We believe that the different result in this
study was due to staff expertise with both aging tech-
niques and that this is critical to obtaining accurate
age estimates for Elk.
Acknowledgments
The Ontario Elk restoration and research program is
supported by the Ontario Ministry of Natural Resources
NOTES
215
ge i
© 20 4 y = 1.0918x - 1.261
=e 7 = 0.9807
a3
c
c
oO
a
c
7
a
o
io)
<
0 5 10 15 20
Age using tooth wear (years)
FIGURE 3. Relationship between age of Elk (Cervus elaphus)
determined by tooth wear and annual growth lines
(annuli) in tooth cementum (n= 10).
(OMNR), Wildlife Research and Development Section,
Dr. J. Chris Davies, manager, and the OMNR, Wildlife
Section, Deb Stetson, manager. Special acknowledg-
ment goes to the staff of the OMNR, Rabies Research
and Development Unit, Peterborough, Ontario, to Trent
University graduate students, Peterborough, Ontario,
to the Canadian Food Inspection Agency, Edmonton,
Alberta, and to Parks Canada, EINP, Alberta (especial-
ly Rob Kaye), who assisted with the acquisition of
teeth for aging and/or the processing of elk at EINP.
The manuscript was reviewed by Dr. J. Chris Davies,
B. Stevenson, and P. Bachmann, OMNR.
Literature Cited
Hamlin, K., D. Pac, C. Sime, R. DeSimone, and G. Dusek.
2000. Evaluating the accuracy of ages obtained by two
methods for Montana ungulates. Journal of Wildlife Man-
agement 64: 441-449.
Hudson, R., J. Haigh, and A. Bubenik. 2002. Physical and
physiological adaptations. Pages 199-257 in North Amer-
ican elk: ecology and management. Edited by D. Toweill
and J. W. Thomas. Smithsonian Institution Press, Washing-
ton, D.C.
Johnston, D. H., D. G. Joachim, P. Bachmann, K. Y. Kar-
dong, R. E. A. Stewart, L. M. Dix, M. A. Strickland,
and I. D. Watt. 1999. Aging furbearers using tooth struc-
ture and biomarkers. Pages 228—243 in Wild Furbearer
Management and Conservation in North America. Edited
by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch.
[CD Edition] Ontario Fur Managers Federation, Sault Ste.
Marie, Ontario.
Keiss, R. 1969. Comparison of eruption-wear patterns and
cementum annuli as age criteria in elk. Journal of Wildlife
Management 33: 175-180.
Rosatte, R., J. Hamr, B. Ranta, J. Young, and N. Cool.
2002. Elk restoration in Ontario, Canada: Infectious dis-
ease Management strategy, 1998-2001. Annals of the New
York Academy of Sciences 969: 358-363.
Rosatte, R., J. Hamr, J. Young, I. Filion, and H. Smith.
2007. The restoration of Elk (Cervus elaphus) in Ontario,
Canada: 1998-2005. Restoration Ecology 15: 34-43.
Zar, J. 1974. Biostatistical analysis. Prentice Hall Inc.,
Engelwood Cliffs, New Jersey, 620 pages.
Received 6 March 2006
Accepted 14 March 2008
216 THE CANADIAN FIELD-NATURALIST Vol. 121
Predator-Prey Interaction Between an American Robin, Turdus
migratorius, and a Five-lined Skink, Eumeces fasciatus
E. NATASHA VANDERHOFE!: 2
'Department of Biology, University of Louisville, Louisville, Kentucky 40292 USA
*Current address: Department of Biology, Francis Marion University, Florence, South Carolina 29501 USA; e-mail: evanderhoff@
fmarion.edu
Vanderhoff, E. Natasha. 2007. Predator-prey Interaction between an American Robin, Turdus migratorius, and a Five-lined
Skink, Eumeces fasciatus. Canadian Field-Naturalist 121(2): 216-218.
I observed a predator-prey interaction between a juvenile American Robin (Turdus migratorius L.) and a juvenile Five-lined
Skink (Eumeces fasciatus L.). Although Robins are considered omnivorous, there are no previous reports of a robin eating
lizards although they have been recorded as occasionally taking snakes. I discuss the age of the individuals involved as it
relates to prey capture and escape.
Key Words: American Robin, Turdus migratorius, Five-lined Skink, Eumeces fasciatus, predation.
The American Robin (Turdus migratorius Linnaeus)
and Five-lined Skink (Eumeces fasciatus Linnaeus) are
commonly found in deciduous forests, as well as rural
and urban areas and parks, in eastern North America.
However, although they inhabit the same areas, there
have been no accounts of predator-prey interactions
between these two species. The American Robin, the
most common thrush species in North America, is a
versatile forager, feeding largely on invertebrates dur-
ing the spring and summer and shifting to a diet with
more fruits in the fall and winter (Sallabanks and James
1999; Wheelwright 1986). Robins also occasionally
consume vertebrate prey, including snakes (Davis 1969;
Netting 1969; Richmond 1975; Erickson 1978), shrews
(Powers 1973), and even fish (Bayer 1980; Kimball
1944). However, there are no published records of
Robins pursuing or consuming any species of lizard
(Sallabanks and James 1999).
The Five-lined Skink is a small lizard associated
with forest openings throughout the eastern United
States and southern Ontario (Fitch 1954; Behler and
King 1997). Juveniles and adults differ in their appear-
ance; adults develop a solid brownish color, whereas
juveniles have five light-colored stripes on a black body
and a bright blue tail (Behler and King 1997). Juveniles
can lose their blue tail (autotomy) during interactions
with predators. This, along with their striped pattern,
is thought to confuse predators and allow the skink to
escape (Fitch 1954; Clark and Hall 1970; Vitt and
Cooper 1986; Goodman 2006). Numerous mammalian
and reptilian predators of the Five-lined Skink have
been documented (Fitch 1954). Although several birds,
including mimids, icterids, corvids, and accipiters, prey
upon lizards (Cooper and Vitt 1985), the American
Robin has not been known to prey upon lizards. Addi-
tionally, there have only been a few confirmed avian
predators of the Five-lined Skink, including the Amer-
ican Kestrel (Falco sparverius) (Heintzelman 1964),
Broad-winged Hawk (Buteo platypterus) (Fitch 1974),
Cooper’s Hawk (Accipiter cooperii) (Toland 1985),
Greater Roadrunner (Geococcyx californianus) (Brown
1963) and Red-shouldered Hawk (Buteo lineatus) (Fitch
et al. 1946).
I observed a predator-prey interaction between an
American Robin and a Five-lined Skink on 24 August
2006 at Joe Creason Park in Jefferson County, Kentucky
(38°12'N 85°42'W). The interaction took place in a
field at the edge of a forested area and I used 10 x 43
binoculars to aid my observations. The initial observa-
tions of two juvenile robins (J1 and J2), and the juve-
nile skink were made from a distance of over 20 m.
Subsequent observations of J2 were made from about
10 m. At 18:18 I observed J1 with a skink in its beak.
At 18:19 J2 approached J1, at which point the skink
escaped by jumping out of J1’s beak. The robins pur-
sued the skink for approximately 30 cm; J2 captured
the skink and carried it approximately 5 m away while
J1 flew off. I moved closer and observed the robin han-
dle and consume the skink over the next six minutes.
The robin, holding the skink in its beak, beat the skink
on the ground multiple times and then took several bites.
The robin repeated this four times, stopping periodi-
cally to stand alert and scan the area. The robin also
took one large bite and swallowed a large portion of the
skink. After the robin flew off, I examined the spot and
found only the skink’s entrails. Closer examination of
the site where the skink was initially captured revealed
the skink’s blue tail. Although I was unable to measure
the skink, based on the tail remnant and entrails the
skink was approximately 90 mm.
This is the first account of a predator-prey interac-
tion between the American Robin and the Five-lined
Skink. The Five-lined Skink spends most of its time
under cover such as woody debris and thus may large-
ly avoid predation from avian foragers (Fitch 1954).
However, the ranges and habitat of both species over-
lap, and it is likely that skinks may occasionally be
pursued and successfully captured by robins. At least
one other thrush species is known to consume skinks;
Turdus caleanops of Japan has been recorded to prey
upon juvenile Eumeces okadae (Hasegawa 1990). It is
interesting that the interaction I observed took place
2007
between juveniles of both species. Previous studies
have shown that juvenile robins are less skilled when
foraging for invertebrates and fruits (Vanderhoff and
Eason 2007). Therefore, one might not expect that a
juvenile robin would attempt to capture a skink. How-
ever, juveniles are often more likely than adults to pur-
sue and consume novel food items (Wunderle 1991).
Juvenile Five-Lined Skinks may also be less skilled
at escaping predators than are adults. I did not witness
the initial capture of the skink by JI so | am unsure of
any anti-predator behaviors that the skink may have
used at that time. However, many species of Eumeces
flicker their vividly colored tails to distract predators
(Cooper 1998), and studies have shown that juveniles
that display this behavior are better able to escape from
snakes (Cooper and Vitt 1985).
Five-lined Skink populations may be more intermit-
tent and less common in Ontario than in Kentucky
where this interaction was observed. In recent years
Five-lined Skink populations have been in decline in
Canada, and under the Species at Risk Act the Five-
lined Skink is listed as a species of special concern
(Quirt et al. 2006). Human removal of woody debris
has eliminated an important microhabitat for the
skink, and the decline in appropriate microhabitat is
thought to be one of the main causes for the skink’s
decline in some areas of Canada (Hecnar and M’Clos-
key 1998). Woody debris is important for nest sites, but
also acts as an important refuge from predators (Hec-
nar 1994), and removal of these refuges may increase
the Five-lined Skinks’ encounter rate with avian pred-
ators, like the American Robin, as well as increase the
skinks’ likelihood of being captured.
Birds and lizards are commonly studied organisms,
but documentation of predator-prey interactions be-
tween these two groups is rare (Blomberg and Shine
2000), although avian predation is thought to be the
selective pressure responsible for the vivid blue tail of
juvenile skinks (Cooper and Vitt 1985), little informa-
tion exists on interactions between birds and skinks.
Both quantitative and qualitative information on pred-
ator-prey interactions between birds and lizards is need-
ed to add to knowledge of both groups.
Acknowledgments
I thank Catherine Byers, Chadwick Hanna, Lina
Rifai, my advisor Perri Eason, and two reviewers for
helpful comments on the manuscript. I also thank
Louisville Metro Parks for allowing me to conduct
my research at Joe Creason Park.
Literature Cited
Bayer, R. D. 1980. Novel use of an unusual food: American
robin eating parts of fish. Journal of Field Ornithology 51:
74-75.
Behler, J. L., and F. W. King. 1997. National Audubon Society
Field Guide to North American Reptiles and Amphibians.
Alfred F. Knopf, New York. 743 pages.
NOTES
217
Blomberg, S. P., and R. Shine. 2000. Size-based predation
by kookaburras (Dacelo novaeguineae) on lizards (Eulam
prus tympanum: Scincidae): What determines prey vul
nerability? Behavioral Ecology and Sociobiology 48
484-489.
Brown, L. N. 1963. Status of the Roadrunner in Missouri
The Condor 65: 242-243.
Clark, D. R., and R. J. Hall. 1970. Function of the blue tail-
coloration of the Five-lined Skink (Eumeces fasciatus)
Herpetologica 26: 271-274.
Cooper, W. E. 1998. Reactive and anticipatory display to de-
flect predatory attack to an autotomous lizard tail. Cana-
dian Journal of Zoology 76: 1507-1510.
Cooper, W. E., and L. J. Vitt. 1985. Blue tail and autotomy
Enhancement of predation avoidance in juvenile skinks.
Zeitschrift fiir Tierpsychologie 70: 265-276.
Davis, W. F. 1969. Robin kills snake. The Wilson Bulletin
81: 470-471.
Erickson, D. B. 1978. Robin feeding upon snake. The Mur-
relet 59: 26.
Fitch H. 1954. Life history and ecology of the five-lined
skink, Eumeces fasciatus. Kansas University Museum of
Natural History Publications 8(1): 1-156. University of
Kansas Publications, Lawrence.
Fitch H. 1974. Observations on the food and nesting of the
Broad-winged Hawk (Buteo platypterus) in northeastern
Kansas. The Condor 76: 331-333.
Fitch, H., F. Swenson, and D. F. Tillotson. 1946. Behavior
and food habits of the Red-tailed Hawk. The Condor 48:
205-237.
Goodman, R. M. 2006. Effects of tail loss on growth and
sprint speed of juvenile Ewmeces fasciatus (Scincidae).
Journal of Herpetology 40: 99-102.
Hasegawa, M. 1990. The thrush 7iurdus celaenops as an avian
predator of juvenile Eumeces okadae on Mayake-Jima, Izu
Islands. Japanese Journal of Herpetology 13: 65-69.
Hecnar, S. J. 1994. Nest distribution, site selection, and brood-
ing in the five-lined skink (Eumeces fasciatus). Canadian
Journal of Zoology 72: 1510-1516.
Hecnar, S. J., and R. T. M’Closkey. 1998. Effects of distur-
bance on five-lined skink, Eumeces fasciatus, abundance
and distribution. Biological Conservation 85: 213-222.
Heintzelman, D. S. 1964. Spring and summer Sparrow
Hawk food habits. The Wilson Bulletin 76: 323-330.
Kimball, J. W. 1944. A fishy bird story. The Auk 61: 646-
647.
Netting, M. G. 1969. Does the robin eat DeKay’s snake?
The Wilson Bulletin 81: 471.
Powers, L. R. 1973. Record of a robin feeding shrews to its
nestlings. The Condor 75: 248.
Quirt, K. C., G. Blouin-Demers, B. J. Howes, and S. C.
Lougheed. 2006. Microhabitat selection of five-lined
skinks in northern peripheral populations. Journal of
Herpetology 40: 335-342.
Richmond, M. L. 1975. American robin feeds garter snake
to its nestlings. The Wilson Bulletin 87: 552.
Sallabanks, R., and F. C. James. 1999. American Robin
(Turdus migratorius). Birds of North America (462). Ameri-
can Ornithologists’ Union, Philadelphia, Pennsylvania. 27
pages.
Toland, B. 1985. Food habits and hunting success of
Cooper’s Hawk in Missouri. Journal of Field Ornithology
56: 419-422.
218
Vanderhoff, E. N., and P. K. Eason. 2007. Disparity be-
tween adult and juvenile American Robins Turdus migra-
torius foraging for ground invertebrates and cherry fruits.
Ethology 113: 1212-1218.
Vitt, L. J., and W. E. Cooper. 1986. Tail loss, tail color, and
predator escape in Eumeces (Lacertilla: Scincidae): Age-
specific differences in costs and benefits. Canadian Journal
of Zoology 64: 583-592.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Wheelwright, N. T. 1986. The diet of American robins: An
analysis of U.S. Biological Survey Records. The Auk 103:
710-725.
Wunderle, J. 1991. Age-specific foraging proficiency in birds.
Pages 273-324 in Current Ornithology. Edited by Dennis
Power. Plenum Press, New York.
Received 15 December 2006
Accepted 20 March 2008
Book Reviews
Book Review Editor’s Note: From this issue forward we will use the current currency codes. Thus Canadian dollars are
CAD, U.S. dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars are AUD, and so on. You will
find these are the codes now used by financial institutions and internet currency converters. I will include an updated note
for the next few issues as a reminder.
ZOOLOGY
Atlas of Breeding Birds of Ontario 2001 — 2005
By M. D. Cadman, D. A. Sutherland, G. G. Beck, D. Lepage,
A. R. Couturier. 2008. Published by Bird Studies Canada,
Environment Canada, Ontario Field Ornithologists, Ontario
Ministry of Natural Resources, Ontario Nature. xxii +
706 pages. 96.00 CAD Cloth.
There’s nothing like a good, hefty book to make
reading in bed difficult; there’s also nothing like a
good, hefty book that’s full of all the information it’s
supposed to have. This second atlas of Ontario birds
is both. Like all recent atlases, this book has enough
coffee-table appeal to be of interest to non-birders, and
could in fact, entice them into the fold. This is not at
the expense of the data, since the book typically pres-
ents the data thoroughly and well. I am very happy
that Ontario (as most jurisdictions) chose to publish a
book, and not a CD-ROM, as did Oregon...people will
always be able to read books...CDs may already be
on their way out.
Given that this is a second atlas for Ontario, the
opportunity to compare results with the first atlas was
there, and was used. The reader can see instantaneous-
ly whether 10 x 10 km squares (in southern Ontario)
or 100 x 100 km blocks (in northern Ontario) have had
a change in breeding status. This segues me to describe
the regrettable, but unfortunately practical practice of
giving less detail to the north than the south; being a
native born-and-raised northern Ontario boy...this has
always bothered me, but as stated, I see the practicality.
What I don’t understand, though, is the situation where
the distribution of a bird like the Palm Warbler is
shown as six squares on a large, and large scale south-
er Ontario map, even though the distribution of 99%
of the population is shown on a smaller, and smaller
scale, northern Ontario map. At least northern Ontario
is covered, unlike the otherwise-fantastic Quebec atlas,
which didn’t attempt to describe birds in the northern
two-thirds of that province.
This atlas has several important chapters of pream-
ble, though, like the Alberta atlas, these are essential-
ly limited to what is necessary to best understand the
Species accounts. Other jurisdictions have provided a
greater diversity of introductory chapters to create an
even more useful atlas — these include chapters on the
local history of birding and ornithology, aboriginals
and their relationships with birds, conservation, and
others. Although not necessary, chapters like these
would have enhanced the Ontario atlas.
Somewhere in the fleet of introductory chapters or
appendices, one would have expected to come across
a very basic piece of information — the number of
squares in Ontario. There are tables in the introducto-
ry chapters and appendices which tell you the num-
ber of squares in which a species was found...obvi-
ously, without knowing the total number of squares, the
former information is less valuable than it could be.
As with the Alberta atlas, each species account is
given one double-page spread. This is convenient for
the reader and allows for enough text, graphs and maps
in most cases. It must have been an editorial decision
to do this — surely some species could have done with
three pages, without getting verbose. However, a two-
page allotment does give a decent treatment to all. The
accounts are divvied into appropriate subheadings in
Distribution and Population Status, Breeding Biolo-
gy and Abundance. A multitude of authors wrote these
species accounts, and even so, I didn’t find a single one
lacking. All were well, to me extremely well, written —
congratulations go to the authors, reviewers and editors
of the species accounts. By using the multi-author
approach, the workload is spread out and the project
takes advantage of the areas in which these folks spe-
cialize.
This atlas incorporated point counts into its method-
ology as an important way to quantify many species.
Very intuitive maps show the distribution of different
population densities. By doing this, the third atlas will
be better able to quantify population changes on the
whole, as well as shifts in population centres and bor-
ders within Ontario.
In contrast, there are many other surveys from
which information could have been incorporated into
the atlas; not all species are effectively sampled using
point counts. For example, the Ontario Nocturnal Ow!
Survey data could have been summarized for each of
Ontario’s owls; this would not have been a daunting
task. The Breeding Bird Survey data, along with the
19
220
data from other sources, were incorporated into the
Oregon BBA, to the overall benefit of atlas users.
There are some technical points to the data presen-
tation that I did not find appealing or useful, though I
will only mention the bigger ones here. For each spe-
cies, there is a histogram which illustrates the probabil-
ity that a person spending 20 h birding will encounter
a species in a square from one of the five biogeograph-
ic regions of Ontario (and another bar illustrating the
data for all of Ontario). On the y-axis are the labels
for the five regions and the whole of Ontario; the x-
axis shows the probability. There are two bars for each
region, one for the first atlas, one for the second. Hav-
ing both bars does give the reader a good idea of the
change in abundance of the species. However, there
are two issues with these histograms. The first is that
the exact value of the probability is put at the end of
each of the twelve bars...isn’t that what the x-axis is
for? For those very few people who need to know an
exact value (e.g., a 59.9% chance of finding a House
Finch in Lake Simcoe-Rideau) instead of the ballpark
x-axis value of 60%, those data can be retrieved from
the atlas project. For the rest of us, the data labels
(made popular by many software packages) are simply
so much clutter.
Secondly, regardless of the data, the x-axis is always
calibrated in 20% increments, from 0-100%. That
means, for species like the Ruddy Duck and Wilson’s
Phalarope, where eight of the twelve bars are at 1% or
less, the reader barely sees the bars...why not scale
~ the axis from 0-10% to show the data more effectively?
This becomes ridiculous with birds like the Worm-eat-
ing Warbler and Northern Wheatear, which have some
of the six categories blank, and all of the others with
non-existent bars labelled at 0.0%.
Each of the species accounts features one photo-
graph of the bird and sometimes a habitat and nest
shot as well. The quality was from good to great...
there were a few shots that I would have replaced, but
nothing serious here. To increase the visual appeal, I
would have included more habitat shots; both The
Birds of British Columbia and Birds of the Yukon Ter-
ritory had more of this, and I think that added quite a
bit to those books [Neither of those books are atlases
per se, but both do show distribution and breeding
records]. A few photographers contributed many of
The Return of Caribou to Ungava
A. T. Bergerud, Stuart N. Luttich, and Lodewiih Camps. 2007.
McGill-Queen’s University Press, Montreal, Quebec. 2007.
586 pages. 49.95 CAD Cloth.
This is the most comprehensive book on Caribou
ecology and predator-prey relationships that has ap-
peared in many years, perhaps ever. Not only is the
research seminal, but the authors systematically dis-
mantle paradigms that have been in vogue for years.
According to the authors, Caribou biologists have
THE CANADIAN FIELD-NATURALIST
Vol. 121
the pictures, though overall there was a good diversity
of photographers; that many peoples’ works get fea-
tured is always good to see in a volunteer effort...so
this is definitely a plus in my mind.
Finally, the cover photo. A Prairie Warbler? Really?
Why? This bird was recorded in only 45 squares in
Ontario — that’s less than 0.5% of the total. If I were
to ask you to name the best-known bird of the Yukon,
what would it be? And yes, it’s on the cover of their
book. The widely-dispersed Red-tailed Hawk was a
fine choice for the cover of the first atlas of Maritime
birds. Surely something much more widespread and
charismatic like their provincial bird, or one known
to almost anyone who feeds birds in Ontario, like the
Dark-eyed Junco, would have been more appropriate.
The decision to have the very local Prairie Warbler as
the coverbird just boggles my mind.
Overall my impression is that this is quite a good
book that could have so easily become a great book.
Ontario atlassers should be very satisfied with their
second atlas — it largely presents the efforts of their
long hours well; users of this book will be faced with
a lot of information that is well-organized, and pleas-
ing to read.
Literature Cited
Adamus, P. R., K. Larsen, G. Gillson, and C. R. Miller. 2001.
Oregon Breeding Bird Atlas. Oregon Field Ornithologists, Eugene.
CD ROM.
Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M.
Cooper, G. W. Kaiser, M. C. E. McNall and G. E. J. Smith.
The Birds of British Columbia Volume 3. University of British
Columbia Press, Vancouver. 693 pages.
Erskine, A. J. 1992. Atlas of Breeding Birds of the Maritime
Provinces. Nimbus Publishing Company and Nova Scotia Museum,
Halifax, Nova Scotia 270 pages.
Gauthier, J., and Y. Aubry. 1996. The Breeding Birds of Québec:
Atlas of the Breeding Birds of Southern Québec. The Province of
Québec Society for the Protection of Birds and the Canadian
Wildlife Service. Montréal. 1302 pages.
Hess, G. K., R. L. West, M. V. Barnhill and L. M. Fleming. Birds of
Delaware. University of Pittsburgh Press. Pittsburgh. 635 pages.
Sinclair, P. H., W. A. Nixon, C. D. Eckert, and N. L. Hughes.
2003. Birds of the Yukon Territory. UBC Press, Toronto. 595 pages.
The Federation of Alberta Naturalists. 2007. The Atlas of Breeding
Birds of Alberta: A Second Look. FAN, Altona. 626 pages.
RANDY F. LAUFF
St. Francis Xavier University, Antigonish, Nova Scotia
B2G 2W5 Canada
wasted the last 50 years measuring lichens on winter
ranges, when they should have been documenting plant
production on summer ranges. Wolves, along with
human hunters, both limit and regulate caribou popu-
lations, not habitat. Food on the summer range only
regulates at high densities and only after the range has
been overgrazed. Wolves are driving Woodland and
Mountain Caribou to extinction. Caribou populations
where Wolves are absent maintain densities 100 times
2007
greater than predated herds. The reason arctic Caribou
migrate to barren ground calving areas is to avoid
Wolves tied to den sites at treeline. Even so, if it were
not for periodic rabies epidemics, migratory Caribou
populations would be severely limited by Wolf pre-
dation. Volcanic eruptions half a world away trigger
population declines in arctic Caribou at high densi-
ties. And this is just for starters.
The book chronicles the history of the George River
Caribou in Labrador and Quebec from near extinc-
tion during the early 1900s to an estimated 600 000
animals before the herd declined. The authors explore
the various hypotheses that have been proposed to
explain these fluctuations and present data set after
data set to separate between competing explanations.
In addition, the authors discuss virtually every other
Caribou population that has been studied in North
America, Scandinavia, and beyond, including the dif-
ference between migratory and sedentary herds, which
is key to understanding this species’ ecology.
To the south of Ungava are small non-migratory
populations of Woodland Caribou that are being driven
to extinction by Wolf predation. But in reality, Moose
and White-tail Deer are to blame. Historically, these
areas sustained low-density, widely-spaced Caribou
that in and of themselves could support few or no
Wolves. Moose and whitetails were absent. But since
the early 1900’s, Moose and whitetails have extended
their range providing alternative prey for Wolves,
where none existed before. The Wolves then drive the
more vulnerable Caribou ever downward. That is to
say, the addition of alternative prey did not buffer
predation on Caribou, but instead increased predation
pressure, contrary to what many people would expect.
But that is not the most intriguing part.
Why were Moose and whitetails absent historically
and prehistorically? The authors contend that logging
changed coniferous forests to secondary deciduous
species favored by Moose and whitetails. In this I
believe they erred because fire history data indicate
there was always a strong deciduous component in
those forests. Besides, Moose and Whitetails can sur-
vive on a winter diet of Balsam Fir, as they do on Isle
Royale and Anticosti Island. Instead, I believe that
native hunters once kept eastern Moose populations
in check, as I know native hunters did in western North
America, where there are more Moose today than at
any time in the last 12 000 years (see Alces 33:141-
164). Historically and prehistorically, native hunters
extirpated Moose over large areas because, like the
Wolves discussed above, humans had a multitude of
alternative prey, including vegetal resources and fish
unavailable to carnivores. As aboriginal hunting pres-
sure declines, prey populations increase. In fact, the
Book REVIEWS
221
authors note that the influenza epidemic of 1918 dec-
imated native populations on Ungava, which in turn
allowed Caribou to increase.
I certainly commend the authors for presenting data
on aboriginal peoples since the time Ungava was first
inhabited and for describing how human hunting im-
pacts Caribou. Most other studies of ungulate ecology
begin with the premise that native people are irrele-
vant because everything was a “wilderness” untouched
by the hand of man prior to the arrival of Europeans:
e.g., see The Kruger Experience. As | have explained
elsewhere, however, this is a fatal error. The authors
did not make that mistake, but I would suggest they
need to look deeper into human evolutionary ecology.
Take the seemingly random movements of Caribou, a
subject covered at length in this book.
Unfortunately, the authors neglected to consult Bin-
ford’s data on Inuit Caribou hunters — see Numamint
Ethnoarchaeology. One of the questions Binford asked
was how do caribou hunters select a direction to hunt
when they have no prior knowledge of where the Cari-
bou are? The Inuit base their decisions on what we in
the West would call mysticism. By careful observa-
tion, however, Binford determined that Inuit pre-hunt
behavior was simply a random number generator. That
is to say, in these cases, the Inuit hunted randomly,
which makes perfect ecological sense, odd though it
may seem.
If the Caribou moved in a predictable pattern, they
would be easy prey for aboriginal hunters, as the au-
thors note when the Ungava herd is forced by topog-
raphy to cross the George River at Indian House Lake.
If the hunters hunted in a predictable pattern, the Cari-
bou would quickly learn to avoid the hunters, and the
people would starve. The solution to the Caribou’s
problem is to move as randomly as possible, while the
solution to the hunter’s predicament is to hunt random-
ly. This co-evolution occurred over thousands of years
and probably is the only evolutionary stable strategy
available to both Caribou and humans and then only
because the Caribou’s range was vast and diverse. The
authors note that even when Ungava Caribou num-
bered only 15 000 animals, spread over an immense
area, aboriginal hunting alone kept the herd from in-
creasing. Using dog sleds, native hunters would follow
caribou tracks for days, until the animals were killed
or the trail lost.
The Return of Caribou to Ungava should be read
by everyone with even a passing interest in northern
ecology, caribou management, or predator-prey rela-
tionships. It should also be read by historians, anthro-
pologists, and archaeologists.
CHARLES E. KAy
Utah State University, Logan, Utah 84322-0725 USA
Dpape
A Guide to the Mammals of China
Edited by A. Smith and Y. Xie. 2008. Princeton University
Press 41, William Street, Princeton, New Jersey USA
08540-5237. 576 pages. 60.00 USD Cloth.
China, like many other areas outside North America
and Europe, has had good field guides to the birds for
some years. Finally it now has a decent guide to the
mammals. Unlike the mammal guides for Africa and
South America, this new Chinese book includes a more
thorough coverage of the small stuff, such as rodents,
pikas, shrews and the like.
It was produced with contributions from nine people
from the United States, the United Kingdom, Italy, and
China. It is refreshing to see such a cooperative effort
by a multi-national team in a region where politics
often clash. The book covers all of China, including
Taiwan and Xizang [Tibet].
The artwork is by Federico Gemma of Italy and is
top quality. I am particularly impressed by the way he
has occasionally moved away from the “classic” guide
poses. For example, he portrays Tibetan Gazelle leap-
ing through the air. This accentuates the gracefulness
of this lovely species. At first it may appear that Gemma
has an easy job because so little is known of Chinese
mammals in the west. This is not true. We are familiar
with the cute poster mammals — the Giant and Red
pandas and the Tiger. Many animals are familiar zoo
and park species — Muntjac [Red], Sika and Sambar
Deer. Others are common species in more accessible
places — Ermine, Caribou, Moose, Red Fox. Using
these as a guide, the reader will soon see that the illus-
trations are accurate. If you want to examine the accu-
racy further, then you can verify details like the facial
features of the Greater Horshoe Bat. However, not all
of the species are covered by this fine artwork. I esti-
mate that a little over 67% of the 556 species are depict-
ed [although some of the non-illustrated species are
in other, readily available, books].
The book has a great introductory section covering
geography, the history of mammalogy in China, and
conservation. This includes some useful maps and pho-
tographs. The text consists of the typical sections on
characteristics, distribution, natural history, additional
comments, and status. I was pleased to find this infor-
mation is current and honest. Chinese literature meant
for the West often glosses over key problems. This is not
the case here, and the status of the Tiger, for example,
is clearly stated. The reasons for its decline are given
frankly. As this book covers 81 species more than I
have on my existing Chinese mammal list, I tried com-
paring the two. I gave up after about 150 species
because there were considerable differences. Many
additional species, primarily from the fringe areas of
Xinjiang, Xizang and Yunnan, were included and many
others were omitted. I realised many of the missing
ones had been assigned a new genus name making
direct comparison more difficult. I did wonder why
THE CANADIAN FIELD-NATURALIST
Vol. 121
the Eastern Black Crested Gibbon (Nomascus nasutus)
was not mentioned except in a list of subspecies. This
is generally regarded as a full species and is consid-
ered the most endangered primate. There are a few [in
the low tens] on Huinan [plus a similar number in Viet-
nam]. Other species like the Manchurian Zokor /Myo-
spalax epsilanus] and the Chinese Scrub Vole [Micro-
tus irene] are not even mentioned, yet are listed in Duff
and Lawson’s Mammals of the World and other ref-
erences.
In addition, there are some changed common names
— Onager is called a Kulan, for example. Also only
five coastal species of marine whales out of over two
dozen pelagic species are included. [However an Ap-
pendix lists an additional 29 cetaceans]. Yet all five
possible seals recorded, mostly as vagrants, on China’s
coast are in the book. Another problem is character-
ized by the distribution of the Northern Birch Mouse,
Sicista betulina, given by some authorities as being in
China, but others as being absent [Including this book].
I think that all of these differences reveal the status of
the taxonomy of Chinese mammals and the need to in-
crease research. I will now use this book as the author-
itative list, but will likely still consider fringe species.
The issue I have with the text is in the “characteris-
tics” sections. The descriptions will be difficult to use
in the field as there is very little comparison with like
species. The descriptions are generic enough to over-
lap and the illustrations are similar. For example, Jer-
boas are very difficult to separate. I know how hard it is
to see these small fast creatures [the Great Jerboa goes
over 40 km/hr!] in the field. There is a key to each
genus, but this is more useful for museum specimens.
[In fact there were a few places where I felt strong
museum overtones as opposed to field experience.
However, the book is not called a field guide.]
In addition to re-assigning genera, the authors have
split the Eurasian Moose or Elk from the North Amer-
ican Moose. In China there are two widely separated
populations: one in Xinjiang [north west] and one in
Heilongjiang [north east]. They have made this split
based on the larger size, redder fur and bigger dewlap
of the “American” Moose. No other reference makes
this difference.
The range maps plot actual museum and other rec-
ords rather than shaded estimations of current distribu-
tion. This is more precise and factual, but may not rep-
resent today’s status and may be more of a historical
record. This will mean the reader will have to interpret
the data carefully so as not to get a wrong impression.
I did spot a couple of errors. Savi’s Pipistrelle’s
range is given as Xinjiang ... to Northern Europe. It
actually is found in Greece, Italy and eastern Spain,
not what I consider northern Europe. The index indi-
cates that the Tibetan Antelope is on Plate 60 when it
is depicted on Plate 57. I also question the range map
of the Spotted Seal, shown with many specimens going
2007
far inland. | would be surprised if this pack-ice loving
beast would go up fresh water rivers.
Birds of the Saskatoon Area
Edited by Anna L. Leighton, Jim Hay, C. Stuart Houston, J.
Frank Roy, and Stan Shadick. 2002. Nature Saskatchewan
Special Publication Number 23, Regina and Saskatoon
Nature Society, Saskatoon. 345 pages. 40.00 CAD [30.00
CAD to Saskatoon Nature Society members] Paper.
During my second encounter with the late J. Bernard
Gollop in Regina some time between 1968 and 1970,
the late Margaret Belcher and her sister, Mary Hous-
ton, mentioned that his planned book on the birds of
the Saskatoon area would probably be the next of the
esteemed Saskatchewan Natural History Society spe-
cial publications to appear. Unfortunately, Bernie did
not live to see it in print, but 77 authors and six edi-
tors finally brought his 1965-1988 efforts to fruition.
The long gestation period produced an outstanding
addition to Saskatchewan’s ornithological literature.
This largest book in the series to date is also the first
to be co-published [by the Saskatoon Nature Society]
and is the fifth in the Manley Callin series on birds of
specific areas of the province. It is jammed with infor-
mation, with even the covers and fold-outs from the
covers containing information [an area map, a Landsat
image of the study area and a few tables]. Introductory
material covers the first 68 pages of text, including pub-
lication blurbs, tables of contents, drawings, maps and
photographs, a dedication to Bernie Gollop, a preface,
a foreword by Farley Mowat, a poem, acknowledg-
ments, chapters on birding aids, histories of birding in
the area and of the project, a detailed chapter on habi-
tats and other ecological influences on the local avi-
fauna, a detailed chapter on “hot spots” [essentially a
bird-finding guide], a list of “unpublished” sources [in-
cluding published seasonal compilations] consulted, a
list of abbreviations, a topography and a glossary. These
are well illustrated with figures, maps and drawings.
The bulk of the text consists of 340 species accounts
in taxonomic order except for one [Prairie Warbler]
that was added while the book was in late stages of
production. Eight appendices, a list of references cited
and indices to English and French bird names close
the book.
The species accounts are preceded by a two-page
introduction, explaining their contents and definitions
used for seasonal occurrence, status in the area, abun-
dance, tables of arrival and departure dates and other
general information. Species accounts range in length
from two paragraphs to two and a quarter pages, de-
pending on regularity of occurrence in the area, amount
of knowledge of the species’ life history features in the
Book REVIEWS
493
Notwithstanding my comments, this is a much-
needed guide and it will be indispensable for natural-
ists and mammalogists for years to come.
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario. K1J 6K5 Canada
area, numbers banded in the area, and other factors,
such as change in status, locally, provincially, and/or
generally. If migratory, earliest and median spring ar-
rival and latest and median fall departure dates are
included. If banded, numbers of the species banded in
the area are listed, as are numbers (if any) recovered in-
side and/or outside the Saskatoon bird area. If known
to nest, data are included on whatever details of nest-
ing biology, chronology, habitat and/or success are
available. Remarks on population changes and their
believed causes are also included if appropriate. Draw-
ings accompany several species accounts. Current En-
glish, French and scientific names and one or more
four-letter species code(s) are indicated for each spe-
cies. Former English names are also indicated for sev-
eral. Nineteen nesting species are listed as hosts of
Brown-headed Cowbird documented in the Saskatoon
area in both the cowbird and the host species accounts.
Several miscellaneous observations of biological, his-
torical or population trend interest are also mentioned,
often in a “side bar” or box.
The book ends with a series of eight appendices (on
annual Christmas, May Day and fall counts, local
Breeding Bird Surveys, numbers of each of 79 species
banded in the area, numbers of birds banded by each
bander who has banded in the area, nest-box dimen-
sions and requirements for 12 species and a list of 12
theses based on birds studied within the area), a list
of references cited and indices in English and French
to the first page of each species account.
This volume continues the trend of improving on the
already high quality of earlier contributions in this
series, using as many sources of published and unpub-
lished sources as possible. Although some variability
in style is inevitable in a book with so many authors
and editors, the quality of writing and accuracy of
information are high throughout. I noted a few minor
grammatical deviations, but few errors or omissions of
substance. The statement (page 284) that Dickcissels
“continue to breed” in southern Manitoba implies reg-
ular occurrence there. While they occasionally nest
there, rarely in fairly large numbers, their occurrence is
irregular, often with several years between sightings
(Davis et al. 2003). Minor errors include the omission
of the hyphen from screech-owl (page 68), reference
to Townsend’s Solitaires as “thrush-sized™ (page 232).
implying that they are not thrushes and a few instances
of species names in lower case. The vocalization of
Baltimore Oriole referred as a “call” (page 291) ap-
1)
tO
TS
pears to be the song. Cross-referencing between species
accounts is quite thorough, but the list of 19 known
local cowbird host species (page 290) omits Hermit
Thrush (page 235) and Common Yellowthroat records
(page 259). Also, a record of House Sparrows nesting
in Bank Swallow holes is mentioned in the swallow
account, but not in that of the sparrow and a case of
hybridization between Eastern and Mountain bluebirds
is mentioned only in the Eastern species account. Odd
omissions from the glossary are the terms “albinism”
and “leucism,’ especially since the less common “mel-
anism” is included. An odd feature, though not an error,
is the use of U.S. spelling for behaviour, centered,
colour, favoured, harbour, moulting, neighbourhood
and variants of these. The difference in slope of the
bill between Tundra and Trumpeter swans, a feature
much easier to see than the visual differences noted,
would have been a useful addition to the identification
tips in the swan accounts (pages 89 and 90). The head-
er for Wilson’s Snipe indicates that two nesting records
have been documented in the area, whereas three are
mentioned in the text. The record of a robin feeding a
young cowbird (page 237), while interesting, is not
definitive evidence of successful parasitism by cow-
birds on robins, as non-parental birds sometimes feed
offspring of birds of other species (for example, Salt
1997). Most references cited are included in the ref-
erence list, but one by Allen (1952) is cited as both
Allan [incorrectly] and Allen on page 138, one by
Dunn et al. (2000) cited on page 189 is not listed un-
less 2002 is intended, the title is missing from a paper
by Greenberg and Droege, the pages of the 1996 owl
paper by C. S. Houston should be 125-133 [not 25-
132] and the second author of the book by Robbins et
al. should be B. Bruun, not B. Brown.
Although such regional works serve primarily to
document occurrence, dynamics and changes in local
THE CANADIAN FIELD-NATURALIST
Vol. 121
bird populations, they also serve to document various
life history observations that the author(s) may other-
wise never find time to write up. Examples of such
tid-bits in this book include fondness of Snow Geese
for peas, television tower casualties of Green-winged
Teal, albino and/or “partial albino” Ruddy Duck, mag-
pies, Mountain Bluebird, Yellow-rumped Warbler and
Yellow-headed Blackbird, a pair of Red-necked Grebes
incubating golf balls, predation by American Crows
on Eared Grebe eggs, cormorants perching on power
lines, a dry land nest of an American Bittern, Willet
courtship flight, magpies chasing and catching bats
and eating berries, a Bank Swallow colony in a hole in
a wheat field, House Sparrows nesting in Bank Swal-
low holes, robins nesting on antlers and large numbers
of starlings flycatching.
Another measure of success in such regional works
is the degree to which they stimulate publication to fill
in gaps and document changes. Judging by the num-
ber of notes and papers on bird observations and re-
search in the Saskatoon area that have appeared in
ornithological and natural history journals since “Birds
of the Saskatoon Area” appeared in print, this volume
has succeeded in that respect as well. It is an impor-
tant contribution towards a forthcoming two-volume
account of the birds of Saskatchewan.
Literature Cited
Davis, S. K., G. Walz, and P. Taylor. 2003. Dickcissel Dickcissel
d’amerique Spiza americana (Gmelin). Pages 372-373 in The
birds of Manitoba. Edited by P. Taylor. Manitoba Naturalists
Society, Winnipeg.
Salt, J. R. 1997. MacGillivray’s and Black-throated Gray warblers
compete to feed fledgling cowbird. British Columbia Birds 7: 15.
MARTIN Mc NICHOLL
Apartment 105, 8752 Centaurus Circle, Burnaby, British
Columbia V3J 7E7 Canada
Book Review Editor’s note: In a brainstorming session between the Editor, Francis Cook and myself, on future directions, we
agreed I would investigate suitable websites and, if appropriate, include their reviews. I proposed we use Brian Coad’s site as
a test case, hence the following review. If others know of similar suitable sites [Cornel University, USDA and USGS come to
mind] please e-mail me [r.john@ rogers.com] with your suggestions.
Brian Coad’s Ichthyology Site
http://www.briancoad.com/main.asp?page=fishdictionary.htm
By Brian Coad. 2008.
Unlike books, websites are not static. This is the
case with Coad’s site, which is defined as a work in
progress. What it currently contains is sections on
Projects, Fishes of Canada: Annotated Checklist,
Dictionary of Ichthyology, Freshwater Fishes of Iran,
Fishes of Canada’s National Capital Region and Fresh-
water Fishes of Iran.
The first section lists the author’s eight current proj-
ects, with a few sentences of explanation. The checklist
of Canadian Fish is a straightforward list of scientific
and common names in both languages, plus occasion-
ally a few comments. The dictionary, co-authored by
the late Don E. McAllister, defines terms specifically
used in the study of fish and includes some generally
used common words.
The Freshwater Fishes of Iran has individual species
accounts arranged by genera. These start with an intro-
duction to the genus. Each detailed account has sec-
tions on the Common name in English and Farsi with
the phonetic pronunciation, Systematics, Key charac-
ters, Morphology, Sexual dimorphism, Colour Size,
Distribution, Zoogeography, Habitat, Age and growth,
Food, Reproduction, Parasites and predators, Econom-
ic importance, Conservation, Further work and Sources.
2007
These accounts are illustrated [black-and-white line
drawings] and give the reader a very exhaustive under-
standing of the individual species.
Fishes of Canada’s National Capital Region [NCR]
has similar coverage to the above. Also included are
photographs of out-of-water fish where these are avail-
able and range maps within the National Capital
Region. I decided to try out the site to identify small
fish I saw in Mud Lake in Ottawa. After comparing
descriptions and other information I decided it might
be a Central Mudminnow. To be certain I would need
the fish in hand. I might raise eyebrows, or worse, if I
started netting fish in a public park.
The Freshwater Fishes of Iraq contains only a
checklist and bibliography at this date.
Some of the advantage that this web site [and all
others] have is that material [such as photos] can be
included as required, poor quality photos can be re-
placed if better become available, multiple photo can
shows all features (not the case in a book where cost
and space are limitations), text and other errors can be
corrected, feedback from readers is easily incorporated,
new information can be added as science progresses,
in some cases differing view points can be presented
at length, saving on costs of paper and distribution
BOTANY
BOOK REVIEWS
995
over the book format. The site can be accessed from
anywhere in the world [with Internet access] without
needing to carry around hard copy, it is copied by stu-
dents, researchers and naturalist for their own projects
This site is a wonderful source of information in a
subject area that is often difficult to access. It is very
easy to navigate and loads quickly [but it is not as
user-friendly as a book]. | am not one to load up my
favourites folder unless it is a site I will use frequently.
I suggest this is one of those sites, especially if you
live in the Ottawa-Gatineau area (National Capital
Region). I also need to trust the information supplied.
The generic question remains in that sites like this have
not been reviewed as a book would be: Readers on the
internet will need to verify the quality of a website
before trusting the data. As Brian Coad is a respected
research scientist (at the Canadian Museum of Nature)
with extensive field experience in the Ottawa District
and Iran, this is not an issue, despite its being a person-
al, rather than an institutional site. At least you should
take a look at it as | am sure you will learn as I did.
Roy JOHN
2193 Emard Cresent, Ottawa, Ontario K1J 6K5 Canada
Exploitation and Utilization of Chinese Spice Plant Resources
By Zhang Weiming and Xiao Zhengchun et al. 2007. South-
east University Press. 722 pages. Price 186.00 CNY.
Spice plants are in a special category in the plant
kingdom, and have been popularly used for various
purposes in different countries or regions of the world.
The history of the use of spice has even exceeded re-
corded history of human beings. Early humans might
have used spice plants to preserve meat, fish or other
food, reduce the annoying odors of various foods, or
increase a special flavor when cooking. Gradually,
these began to expand the range of use of spice plants.
Spice plants are closely related to the daily lives of
human beings. Systematic research and summary of
them is necessary, however, as this has not been well
done before, at least in China. Now, this situation has
been changed by the publication of the book Exploita-
tion and Utilization of Chinese Spice Plant Resources
by Zhang Weiming and Xiao Zhengchun et al. The
authors have been engaged in research work on spice
plant resources for a long time. They have conducted
several research projects relating to Chinese spice
plants in the last few decades, and obtained valuable
and abundant data in this field. Meanwhile, they also
collected other abundant information on spice plant
resources at home and abroad. Based on these data
and information, they produced this massive book.
There are a number of climate zones in China. Dif-
ferent environmental conditions allow for the growth
of an abundance of plant species, including spice plants.
Except for a few tropical species, most of the spice
plants grow in China. China was one of the earliest
countries to use spices. There are some records on the
aromatic and spicy plants in the Book of Songs, the
Book of Mountains and Seas, and other book that are
more than 2000 years old. Over a long time, due to the
various usage of different types of spice plants in cook-
ing, the people living in various areas of China formed
a variety of eating habits, reflected in the so-called
eight genres of cuisine of Chinese dishes. Hence, we
might say that one of the most important reasons that
Chinese food (or any other regional food) enjoys a
world-wide reputation is its unique use of spices in
cooking.
The indigeneous spice plants in China are abundant.
Furthermore, over the years, China introduced many
foreign-origin species. The book not only is a system-
atic and comprehensive monograph on the spice plant
resources of China, but also has its value as a refer-
ence for other related countries. The book is divided
into two parts. The first part is a general introduction
to spice plants, including the history of spice plants,
the general situation of Chinese spice plant resources,
the main ingredients and the application of spice plants,
the processing technology and the development of spice
plants, and the strategy of the development of China
spice plant resources. The second part is the special
issues, including a total of 134 kinds of commonly
used spices, and more than 200 spice plant species.
For each spice plant species, the ecological environ-
ment, history of use, chemical ingredients, compre-
hensive methods of use, exploitation of related prod-
ucts, standardization and cultivation techniques, and
sO on were comprehensively introduced. The typically
introduced spice plant species in this part include
Capsicum frutescens, Allium sativum, Aglaia adorata,
Allium cepa, Allium fistulosum, Allium porrum, Allium
sativum, Artemisia vulgaris, Illicium verum, Foenicu-
lum vulgare, Levisticum officinale, Mentha haploca-
MISCELLANEOUS
Arctic Hell Ship
By W. Barr. 2007. University of Alberta Press, Ring House
2, Edmonton, Alberta T6G 2E1. 318 pages, 34.95 CAD
Cloth.
More of a human history than a natural history
book, this tale covers part of one of the great sagas of
Canada’s north. The Franklin Expedition left Britain
with high hopes and the best resources available —
well almost. It vanished in 1845 and spawned at least
two dozen searches to solve the mystery of the disap-
pearance. These efforts in turn have led to a flood of
books and scholarly articles.
So do we need another treatise? The answer is yes
in this case. Barr’s contribution covers the 12" search
by Captain R. Collinson aboard the HMS Enterprise
with Robert John Le Mesurier M’Clure, as second in
command, on the HMS Investigator. Barr gives us an
extremely detailed description of life on one of Her
Majesty’s ships in the 1800s. So often the other books
concentrate on the leaders and their accomplishments,
so we learn little of how the crew fares. Barr remedies
this by telling us about many of the individual sailors.
The “story” is very simple. This was a voyage that pro-
duced limited results and suffered a lot of bickering
between the officers. As a result, the ship’s, logs and
letters contain irritating daily details that show what
life was really like.
Collinson was a good seaman and surveyor, but he
was Clearly not a leader. He picked on very minor
transgressions and turned them into major discipli-
nary actions. He sowed unnecessary seeds of discord,
interspersed by bouts of humanity. In hindsight, his
biggest failure was that he did not do enough. He was
charged with rescuing Franklin’s men and given the
resources to do it, and yet he spent a lot of time sailing
back and forth, unable to make a clear decision. His
sledging trips were too little and too late. The other
ships in his command (which got “separated” early
in the expedition) went farther and did more. When
THE CANADIAN FIELD-NATURALIST
Vol. 121
lyx, Pimpinella anisum, Sesamum indicum, Zanthoxy-
lum bungeanum, Zanthoxylum simulans, and Zingiber
officinale, among others.
The book was well written with few errors, in a
style of writing that can be easily understood by non-
scientists. As a comprehensive book, it is suitable for
anyone who is engaged in the various areas of research
and application of spice plants.
Li DEZHI
Lab of Urbanization and Ecological Restoration of Shanghai;
National Field Observation and Research Station in
Tiantong Forest Ecosystem of Zhejiang; Department of
Environmental Science, East China Normal University,
3663, Zhongshan Road (N). Shanghai, China 200062
Collinson did make progress he found notes from
either M’Clure or Dr. John Rae showing that they had
searched the area earlier. In the tough conditions on
board ship during an Arctic winter this debilitating
caution and indecisiveness coupled with petty deci-
sions on discipline must have made an unhappy ship
full of exasperated people.
Collinson returned to Britain, having failed to solve
the Franklin issue, demanding courts-martial for his
officers. The lords of the Admiralty wisely declined to
take action. Public exposure of such silly shenanigans
would do the service no good.
The most notorious British naval captain of this era
is Bligh. In spite of the fact that his reputation owes
more to the movies than reality, Bligh was a superb
navigator, excellent seaman and a relatively humane
captain. He personally intervened in the trials of the
captured mutineers, got some set free and obtained
reduced sentences for others. He died as a Vice-Admi-
ral. Collinson’s dithering and his stubborn need for
absolute discipline would have made him a better can-
didate for Hollywood’s spite, yet even this would be
unfair. How would we all fare under such harsh con-
ditions?
Collinson did make Vice-Admiral, after serving at
Trinity House, the establishment that maintained the
aids to navigation, where he rose to deputy master. He
remained at Trinity House until retirement, clearly a
better bureaucrat than expedition leader. Perhaps the
error lies with the lords of the Admiralty. They fell into
that awful trap. They promoted a very technically com-
petent person to a supervisor, a move that does not al-
ways work. In particular, voyages to the Arctic, even
today, need competent leaders. Collinson did not come
close to Nansen, Shackleton, Amundsen, or even Scott
in his ability to deal with people.
While Barr has produced a minutely detailed doc-
ument, using all available evidence, rather than a dis-
2007
tillation of the key events, it is those details that give
the book its appeal. I found I wanted to read just one
more paragraph each time before I put the book down.
I was surprised when I realized I was over half way
through the book and it seemed I had just started. As |
stated initially this is really a history text, but it covers
a key era in the story of Canada. The only important
Kighteenth-century Naturalists of Hudson Bay
By Stuart Houston, Tim Ball and Mary Houston. 2003.
McGill-Queen’s University Press, 3430 McTavish Street,
Montreal, Quebec, H3A 1X9. xxiv + 333 pages 49.95
CAD Cloth.
In 1959, C. Stuart Houston and Maurice G. Street
began to introduce prairie naturalists to the pioneering
contributions of employees and other associates of the
Hudson’s Bay Company and to expand our under-
standing of the natural history of Canada’s Arctic and
prairie regions (Houston and Street 1959) in the first
of a long series of articles, books, notes and papers by
Houston and numerous colleagues on biographies of
specific contributors and the history of various aspects
of these contributions. In this volume, Stuart and his
chief collaborator (Mary I. Houston) bring many of the
details of their earlier publications together, expand
on the European collaborators of these naturalists and
compare the Hudson’s Bay contributions with approx-
imately contemporary efforts in South Carolina. They
also collaborate with Timothy E. Ball in outlining the
contributions of H.B.C. officials in documenting cli-
matological details at various sites and their changes
over a long time period and with Arok Wolvengrey
and Jean Okimasis in documenting Cree names for
numerous natural history species in the lands of the
vast territory formerly controlled by H.B.C.
The book opens with a series of introductory sec-
tions, including a list of McGill-Queen’s “Native and
Northern Series” (of which this is the 34"), a dedica-
tion to William B. Ewart (whose efforts led to the mov-
ing of the extensive Hudson’s Bay Company archives
from England to Winnipeg), lists of contents, illustra-
tions and tables, eight colour plates of birds originally
published in two of four 1743-1751 volumes by George
Edwards, acknowledgments, a list of supplementary
documents available through the internet, a list of ab-
breviations and a foreword by the Keeper of the Hud-
son’s Bay Archives, Judith Hudson Beattie. The main
text of the book (pages 1-142) consists of an unnum-
bered introductory chapter, followed by 12 numbered
chapters. Seven appendices occupy almost as many
pages (pages 143-251). About 40 pages of footnotes,
25 pages of references and a 14-page index close the
book.
The main text starts with a short outline of the his-
tory of Hudson’s Bay Company activities in North
America and the role of HBC officials in documening
natural history features of much of the prairie prov-
Book REVIEWS
227
note of natural history concerns the surgeon, Edward
Adams, for whom the Yellow-billed Loon was named
Gavia adamsii. It is well worth the read if you love
the Arctic, enjoy history or are a Franklin buff
Roy JOHN
2193 Emard Crescent Ottawa, Ontario K1J 6K5 Canada
inces, Ontario, Quebec, Nunavut and adjacent areas in
collaboration with native North Americans and Euro-
pean scientists. The first numbered chapter consists
of brief biographies of seven European scientists who
received and/or examined animal specimens collected
by HBC personnel, described and often named them,
and wrote the first accounts of their behaviour and
other natural history features. The role of these speci-
mens in developing Linnaeus’s system of classification
of plants and animals is outlined. Eight chapters fol-
low on the contributions and efforts by eight specific
HBC officials (Peter Fidler, Andrew Graham, Samuel
Hearne, Thomas Hutchins, James Isham, Alexander
Light, Humphrey Marten and Moses Norton). The tenth
numbered chapter summarizes these contributions, lists
Hudson Bay area bird species mentioned in ten early
manuscripts, provides several background details and
notes several corrections and omissions in earlier pub-
lications. The eleventh chapter, based primarily on four
years of archival research by Ball, outlines weather,
climate and climatic changes in the area, as document-
ed through records of 30 or more years at 30 HBC
posts. The final chapter outlines early contributions
to North American natural history by early settlers of
Charles Town, South Carolina, when that area was
still a British colony. Brief biographies of two contrib-
utors of meteorological data and two natural history
collectors are included, as well as a list of species
named as new in Mark Catesby’s 1729-1747 two-vol-
ume treatise on the birds of South Carolina (58 from
“South Carolina,” seven from other parts of eastern
U.S. states, seven from the Caribbean, and one from
“Quebec.”
Seven appendices cover about 100 pages. The
longest (about 50 pages) by the Houstons, Jean Oki-
masis and Arok Wolvengrey lists words used in the
eighteenth century in four Cree dialects for various taxa
that live in the territory controlled by the Hudson’s Bay
Company. Another lists departure and arrival dates and
the captains of ships sailing between York Factory
and Europe from 1716 to 1892.
Smiles will cross the faces of Saskatchewan natural-
ists, long aware that two of their most prominent mem-
bers (Margaret Belcher and Mary Houston) were sis-
ters, when they read that one of two ships captained by
a Belcher (twice) was “Mary.” A two-page appendix
by Deirdre Simmons outlines the history of the Hud-
son’s Bay Company Archives. Others concern ten man-
228
uscripts that have variously been attributed to Andrew
Graham and Thomas Hutchins, and plagiarism of Gra-
ham by Hutchins. The Houstons provide an interesting
account of their research in the archives, in London and
at a symposium in the Orkney Islands that brought
them to the conclusion that these manuscripts resulted
from collaboration. Other appendices concern the ten-
year cycles of various northern birds and mammals,
the effects of nineteenth century trade on Trumpeter
Swans, and the naming of Canada Goose prior to the
official existence of Canada. I assumed that the latter
was a combined reprint of two earlier publications in
Blue Jay (Houston 1994, 1995), but some of it presents
additional findings of historical research that was stim-
ulated by the earlier notes.
Naturalists interested in historical aspects of nature
study will find plenty of interest in this volume. It is
written well and researched thoroughly. I found no sub-
stantial errors, although a few references cited were
either not included in the literature lists or the publica-
tion dates and/or author list in the text differed slightly
from those in the literature list. One reference (Ross
1834) is listed in a footnote (page 256), but not in the
references. Three references cited are not included in
the literature list (Gmelin [1788 on page 140], McIn-
tyre and Houston [1999] on page 269) and Rousseau
[1969] on page 279, although enough information on
the latter is included to enable a reader to track it
down). The 1983 book by Peck and James should be
listed as Volume 1. Footnote 39 on page 255 on
William Wales should refer to Chapter 11, not 12.
References to Appendix E and Figure 3.13 on page 52
apply to Appendix F and Figure 3.14; respectively.
THE CANADIAN FIELD-NATURALIST
Vol. 121
The abbreviated forms of the authors’ names from the
more usual versions that they use in publications (Tim-
othy E. Ball, C. Stuart Houston and Mary I. Houston)
provides potential confusion for bibliographers who
may think that other, presumably related, authors are
involved, especially as their usual publication versions
are used in some appendices. As noted by Meyer
(2004), some of the appendices would be more appro-
priate within the main text. Although the literature
list is substantial, readers new to the subject who
wished to follow up with further reading would have
benefited from inclusion of more of Houston’s previ-
ous biographies, book chapters, notes and papers on
some of the subjects covered. Similarly, although bio-
graphical references are included for many people,
none are included for a few (for example, William
Rowan) for whom several are available. None of these
criticisms detract substantially from the overall high
quality of this excellent contribution to the history of
nature study in North America.
Literature Cited
Houston, C. S. 1994. How did the Canada Goose get its name before
there was a Canada? Blue Jay 52: 141-143.
Houston, C. S. 1995. Earlier use of the name Canada Goose. Blue
Jay 53: 116-117.
Houston, C. S. and M. G. Street. 1959. The birds of the Saskatchewan
River Carlton to Cumberland. Saskatchewan Natural History Soci-
ety Special Publication Number 2, Regina.
Meyer, D. 2004. [Review of] Eighteenth-century naturalists of Hudson
Bay. Blue Jay 62: 120-122.
- MARTIN K. Mc NICHOLL
Apartment 105, 8752 Centaurus Circle, Burnaby, British
Columbia V3J 7E7 Canada
Hunters at the Margin: Native People and Wildlife Conservation in the Northwest Territories
By John Sandlos. UBC Press, 2029 West Mall, Vancouver,
British Columbia V6T 1Z2. 360 pages, 85.00 CAD
Cloth, 39.95 CAD Paper
Professor Sandlos has written a very well-researched
text on three major conservation issues. He examines
the plight of the Bison, Muskox and Caribou in Cana-
da’s north. The author has examined the questions
raised about hunting of these three key species and
the development and enforcement of laws relating to
them. He has done this with a very sympathetic atti-
tude towards Canada’s original inhabitants. He com-
pares the lifestyle and requirements of aboriginals to
the efforts of Ottawa’s bureaucrats to control hunting
and to conserve these mammals. These two philoso-
phies are rarely compatible, and this leads to conflict.
I had a hard time reading this book. While I under-
stand the author’s sympathy with Canada’s aboriginal
people I found his constant innuendo very galling.
When discussing the native viewpoint he writes with-
out emphasis. When writing about the non-native peo-
ple’s actions he frequently uses parentheses, causing
the reader to doubt the adjective’s veracity. When
defending the native people Sandlos ignores much per-
tinent information. He proposes the natives are not the
wanton killers portrayed by white policemen, wardens
and travelers, but long before the arrival of the white
man the natives hunted Mammoths, Giant Sloths and
others to extinction. This was also before climate
change pushed these creatures over the edge [as in
Europe]. The latest book on Caribou (Bergerud et al.
2007) notes that after an influenza epidemic decimated
native populations, Caribou increased [see review this
issue]. In my own experience, two years ago at Cam-
bridge Bay, I was informed the hunting ban on Muskox
had been lifted a few months earlier and the town’s
residents had shot the entire local herd. I had to travel
out of town for an hour by truck and an hour and a
half on foot to see distant Muskox on this island.
Before the arrival of whites, there were few natives
in the north; it was too hostile a climate. Living was
easier on the plains with its abundant Bison. In fact
the author provides evidence for this concept when he
quotes a convicted poacher, Joseph Wakwan, as pre-
ferring to hunt the larger Bison over the legal Moose.
2007
The natives moved north after they found they could
‘make money for guns and supplies by trapping Beaver.
The demand for Beaver hats was purely European.
I do not know if the author has been involved in
creating laws, but I suspect not. This a long and com-
plex process where nobody wins everything they want.
At the conclusion you achieve compromise regulations
that all can accept, more or less. To constantly criti-
cize the Ottawa law-making bureaucrats, who were try-
ing their best with the information and attitudes of the
day, is a futile process. Trying to judge past events with
today’s understanding is fun, but useless. National
committees have learned long ago not to react too
quickly to the latest reports and studies. It takes time
and careful study to be certain any new data are valid.
There have been many reports that claimed some-
thing was bad, only to be followed by another that said
it was good. We do not want our laws flip-flopping
with every new announcement.
If we applied Sandlos’s logic that white man’s laws
do not match aboriginal situations and therefore real-
ly do not apply, then I could ignore the laws of Eng-
land as I am Welsh, a true English aboriginal. [The
Welsh inhabited all of Britain before they were con-
quered by the Angles and Saxons. These in turn were
reduced to serfdom by the Norman French.] I, like all
those in England, have to obey today’s laws. Such is
history. The Canadian native people actually fared rea-
sonably well for a conquered community. Compare
their fate to that of the Inca after the arrival of the Span-
ish or the original inhabitants of the Amazon after the
Portuguese took over. The arrival of the white man
brought white-man’s laws, as well as welfare, modern
The Ornithologist’s Dictionary
By Johannes Erritzoe, Kaj Kampp, Kevin Winker and Clifford
B. Frith. 2007. Lynx Edicions, Montseny, 8, 08193 Bella-
terra, Barcelona, Spain. 290 pages, 19 EUR Paper.
This is a great little book that will be handy for all
who study birds. I can easily slip out of my depth when
reading a book or article because I do not understand
some terms. Typically I stick a mark in the page and
later go to my computer for help. Google searches are
normally wonderful ways to fill in your knowledge
gaps. But this also disrupts the continuity of your read-
ing and lowers the value you can pull from the text.
Having a resource that can sit by your side is a big
advantage.
So how accurate and comprehensive is the text?
First, let me say the authors use the English spellings
[colour etc.] This does not mean a European bias as
there are plenty of references to purely North American
terms [National Audubon, AOU etc.] and the U.S.
spellings are included [spishing vs pishing!]. I looked
up several definitions of words which are frequently
misused and could only find one significant error.
Parameter is not “any variable” but a constant in an
BooK REVIEWS
299
medicine and current technologies. Whether these are
good or bad is a moot question. When | wrote my last
report on people of the north, the data sets I accessed
on diet used by northern residents showed that a high
proportion of junk food had replaced food gathered
from the wild. In my view, this represents a poor
choice; on the other hand, I do not recall any native
village starving or freezing to death [as happened in
the past, before the arrival of the white man]. We
need to remember we all live in today’s conditions,
which includes a plethora of laws which we must all
obey [Whether we agree with them or not. Ask me
about CITES. A well-meaning idea, whose concept I
support. In practice it is often counterproductive to
conservation and frequently punishes those who are
promoting conservation. |
Do not let my criticisms put you off reading this
book. There is a lot of useful and thought provoking
material embedded in the text. The questions and
issues surrounding the native people and wildlife in
the north are not easy to deal with and | doubt there
will ever be “an” answer. This book makes a significant
contribution to the continuing research and debate
required to make rational decisions. I just wish it was
written in a more balanced and critical style.
Literature Cited
Bergerud, A. T., Stuart N. Luttich, and Lodewiih Camps. 2007.
The Return of Caribou to Ungava. McGill-Queen’s University
Press, Montreal, Quebec.
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario K1J 6K5
equation that varies in other equations of the same gen-
eral form [the classic example is the force of gravity
— always constant in Ottawa but different yet con-
stant in Banff.] This is an incredibly common error,
even among scientists who should know better. I have
a few other less significant comments, such as the
needless use of interrelationship [relationship is fine,
particularly in a dictionary] and I would not say the
use of “mirrors” for the white tip of a gull’s wing is un-
common as it is used in most of my books. I thought,
too, that the authors could have included newer terms
like sahel [note the lower case s] — A region having
characteristics of a savanna or a steppe and bordering
on a desert — as well as the classic definition of Sahel
— a transition zone between the Sahara and the trop-
ical forests to the south.
Each letter of the alphabet starts with a neat black-
and-white drawing of a bird. These are not identified,
but I could easily determine the species in all but X
and Y. Also I could not find the identity of the artist.
Several pages in my original copy were greyed. As
this is totally unlike the publisher, Lynx Edicions, who
230
normally produce beautiful, crystal clear copies, I con-
tacted them. It appears this was an isolated incident;
however, you should check the particular copy you
intend to buy. Lynx sent me another, perfect copy.
This will be of greatest use to non-professional avid
birders who lack formal training. You can quickly
resolve the occurrence of melanin, the nature of re-
miges, the difference between granivorous and grami-
nivorous, mimesis and mimicry and other sources of
Pioneering Women in Plant Pathology
Edited by Jean Beagle Ristaino. 2008. The American Phyto-
pathological Society, 3340 Pilot Knob Road, St. Paul,
Minnesota, U.S.A. 339 pages. 89.00 USD Cloth.
As seen by one woman, an M.Sc. graduate in plant
pathology from the mid-1970s, it would appear that
women scientists in this field are rapidly proving their
abilities to shine with accomplishment in what was
once an all male profession; moving from being seen
as pretty women hobbyists picking wild flowers to seri-
ous taxonomists of vascular and non-vascular plants,
many of which are plant pathogens. Agriculture has
itself moved from its primitive beginnings of slash and
burn to a precise science of food production in the 21*
century.
As the possibility that a woman may progress from
a mere technician and housekeeper of the laboratory
to the more stellar role of research scientist has im-
proved, their research results have proved their abili-
ties. The first woman plant pathologist, hired by the
United States Department of Agriculture, was Effie
A. Southworth, in 1887. Her most significant contri-
bution was the description of the pathogen Colleto-
trichum gossypii, the cause of anthrocnose on cotton,
and the recommendation of measures of control of
this disease. In 1895, Flora W. Patterson became the
first woman mycologist at United States Department
of Agriculture. She is remembered for the develop-
ment of the U.S. National Fungus Collections, which
are still of importance to mycologists and plant pathol-
ogists today. She also was very involved with the in-
spection of imported materials for invasive fungal
pathogens, and intercepted the dangerous potato wart
disease before it was imported into the United States
for the first time. Another woman of importance was
Edna Marie Buhrer, who brought forward the impor-
tance of nematodes in plant diseases; up till 1920 the
role of nematodes in crop production was considered
unimportant. This led to various treatments of soils with
Birdwatcher: The Life of Roger Tory Peterson
By Elizabeth J. Rosenthal. 2008. The Lyons Press, Guilford,
Connecticut. 437 pages. 34.95 CAD.
Roger Tory Peterson’s parents could not afford to
send him to college and he had no formal training in
science. He moved to New York City, where he paint-
THE CANADIAN FIELD-NATURALIST
Vol. 121
confusion. This book would have rapidly ended my
last bird-term argument — the meaning of leucism —
if I had had it handy. It may not be as valuable to pro-
fessionals, but I think they will likely find it useful
too. Buy this one for yourself!
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada
nematicides which results in improved crop produc-
tion.
In England also the importance of women in this
field was recognized at Rothamsted Research Center
and among those employed there and making a valu-
able contribution was Mary Gwynne. She started her
career in 1917, worked mainly on diseases of cereal
crops, and in 1960 was awarded the Order of the British
Empire for her contribution. Margaret Newton, at Mac-
Donald College, McGill University, was a Canadian
woman who made a considerable contribution to the
knowledge of stem rusts of wheat, so important to the
agricultural economy of Canada. At this period of time
in the early twentieth century, women from European
countries were also making their contributions.
It is recommended that this book be on the shelves
of every plant pathology department library, not only
to show the valuable contributions made by these sci-
entists, but also to show the courage and dedication
of these pioneering women in the face of the prejudices
of those times. This book also highlights the impor-
tance of this field of science to the agriculture of the
American continent and the safety of the world’s food
trade. We have here a book about pioneering women
plant pathologists who have overcome prejudice to
make considerable contributions to the economy of
North America.
Pioneering Women Plant Pathologists is edited by
Jean Beagle Ristaino; and the careers of 26 women
scientists are outlined by 37 authors from the United
Kingdom, United States of America, Canada, Portu-
gal, Italy and the Netherlands. It is most interesting to
read, and gives one a very good indication of what
these plant pathologists have contributed to this field.
ELIZABETH PARNIS
12-1063 Valewood Trail, Victoria, British Columbia V8X 5G5
Canada
ed designs on furniture in the mornings to earn enough
to attend art classes in the afternoon. He joined the
Bronx County Bird Club, following the new concept
of identifying birds in the field without the need to
shoot any specimens for confirmation. The club’s pres-
2007
ident, Ludlow Griscom, was an early advocate of this
method and RTP soon rivaled him in auditory and
visual skills.
RTP’s first book, A Field Guide to the Birds, pub-
lished 27 April 1934 at the height of the depression,
made bird identification easy for the first time. His
wide-ranging combination of skills and knowledge
made possible the succeeding Peterson field guides,
all published by Houghton Mifflin in Boston. Each
used the “Peterson system of field identification,’ with
an arrow pointing to important field marks for each
species. Millions of copies were sold. His first bird
field guide went through four more revisions, the last
posthumously; it was 85% complete when he died a
month before his 88" birthday. Through his multifac-
eted achievements: books, articles, photographs,
movies and paintings, Peterson became the best-known
and arguably the best-loved naturalist in the world.
RTP’s first teaching experiences were at a boy’s
camp in Maine, and in writing pamphlets for members
of Junior Audubon Societies. He remained a teacher
all his life. His dedicated second wife, Barbara, man-
aged his home and business affairs with great distinc-
tion, freeing him for extensive world travel.
Liz Rosenthal’s informative book is based largely
on 147 interviews with 117 people, including many of
the “who’s who” of the birding world. Their words and
memories provide inside stories about their interactions
with Roger. The inevitable result of this methodology
is a few minor omissions. As one example, Rosenthal’s
book omits one of RTP’s early pupils at the Rivers
School near Boston: Eliot Richardson, who much later
became U.S. Attorney-General and successfully nom-
inated RTP as the teacher who had influenced him
most, resulting in the nationwide “Teacher of the Year”
Award. Nor we do we learn that RTP’s final total of
honorary doctorate degrees reached 23.
Conservation themes are interwoven throughout this
book. These began with Peterson’s long involvement
with the National Audubon Society, as a director, sec-
retary, columnist in their magazine, and presenter of
some of their most popular screen tours. He helped
found the World Wildlife Fund, helped save fragile
areas such as the fabulous Coto Donana in Spain, Mid-
Book REVIEWS 23]
way Atoll in the Pacific and Aldabra Atoll in the Indi-
an Ocean, and helped Kenya's Lake Nakuru become
a national park, where a million flamingos form “the
world’s greatest ornithological spectacle.” He pho-
tographed all 17 of the world’s species of penguin.
We learn of Peterson’s close friendships with two
other world-famous ornithologists, James Fisher and
Sir Peter Scott. As evidence of his stature among pro-
fessional ornithologists, Peterson served as president of
the Wilson Ornithological Society and as Vice-Presi-
dent of the American Ornithologists’ Union (AOU). An
icon, he received the William Brewster Award from
the AOU in 1944 at the early age of 36; Presidential
Medal of Honor, presented by Jimmy Carter; Audubon
Medal from the National Audubon Society; Gold
Medal of the World Wildlife Fund, presented by Prince
Bernhard of the Netherlands; Order of the Golden Ark
from the Netherlands; Linnaeus Gold Medal from the
Swedish Academy of Sciences, and the Smithson
Medal from the Smithsonian Institution. He attended
the dedication of the Charles Darwin Research Station
in the Galapagos. RTP was happiest when serving as
naturalist-lecturer aboard Lars-Eric Lindblad’s Explor-
er on cruises to the Antarctic, Galapagos, Amazon
River and Aldabra Atoll.
I detected two errors. Iceland is NOT north of the
Arctic Circle (page 133). The Peregrine nest on the
Sun Life building was in Montreal, not Toronto (page
229). I also regret that, in a book about people, the peo-
ple index is incomplete, omitting some names entirely,
and incompletely listing mentions of others.
Elizabeth J. Rosenthal, a graduate in both journalism
and law, has previously written His Song: the Musical
Journey of Elton John. Her Peterson life deals with a
man of very different but not lesser talents, who was
unusually helpful to beginner and expert alike, and
whose personality shines through in the interview
process. The potential audience includes the millions
of people who began nature study with a Peterson field
guide.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N OJ8
Canada
Jee
New TITLES
Prepared by Roy John
+ Available for review * Assigned
ZOOLOGY
Audubon: Early Drawings. By S. Edwards [Annotater].
2008. Harvard University Press, 79 Garden Street, Cambridge,
Massachusetts 02138 USA. 272 pages. 125.00 USD Cloth.
* Backyard Birdwatch. By S. Kress. 2008. Dorling Kin-
dersley/Tourmaline Editions Inc., 662 King Street West, Suite
504, Toronto, Ontario MSV 1M7 Canada. 72 pages. 11.95
CAD Paper.
* Beautiful Minds: The Parallel Lives of Great Apes and
Dolphins. By M. Bearzi and C. Stanford. 2008. Harvard
University Press, 79 Garden Street, Cambridge, Massachu-
setts 02138 USA. 322 pages. 24.95 USD Cloth.
Butterfly. By T. Marent. 2008. Dorling Kindersley/Tour-
maline Editions Inc., 662 King Street West, Suite 504, Toronto,
Ontario MSV 1M7 Canada. 280 pages. 35.00 USD Paper.
Life in old Blood. By Sir D. Attenborough. 2008. Princeton
University Press, 41 William Street, Princeton, New Jersey,
08540-5237 USA. 288 pages. 29.95 USD Cloth.
* Ecology. By M. Cain, W. Bowman and S. Hacker. 2008.
Sinauer Associates Inc., Box 407, Sunderland, Massachusetts
01375-0407 USA. 621 pages. 107.95 USD.
Egg & Nest. By R. Purcell, L. Hall and R. Corado. 2008.
Harvard University Press, 79 Garden Street Cambridge,
Massachusetts 02138 USA. 224 pages. 39.95 USD Cloth.
Evolution. Edited by M. Ruse and J. Travis. 2009. Harvard
University Press, 79 Garden Street, Cambridge, Massachusetts
02138 USA.
* The Ferocious Summer. By M. Hooper. 2008. Greystone
Books, #201-2323 Quebec Street, Vancouver, British Col-
umbia V5T 4S7 Canada. 336 pages. 29.95 USD.
+ Fossil Ecosystems of North America. By J. Nudds and P.
Selden. 2008. University of Chicago Press, 5801 Ellis Avenue,
Chicago, Illinois 60637 USA. 288 pages. 39.00 USD.
* A Guide to the Mammals of China. Edited by A. Smith
and Y. Xie. 2008. Princeton University Press, 41 William
Street, Princeton, New Jersey 08540-5237 USA. 576 pages.
60.00 USD.
* A Guide to the Mammals of Southeast Asia. By C.
Francis. 2008. Princeton University Press, 41 William Street,
Princeton, New Jersey 08540-5237 USA. 392 pages. 55.00
USD.
Seashore Life in the North Atlantic. By J. D. Duane. 2008.
Princeton University Press, 41 William Street, Princeton,
New Jersey 08540-5237 USA. 240 pages. 19.95 USD.
BOTANY
* The Macrolichens of New England. By James W. Hinds
and Patricia L. Hinds. 2008. The New York Botanical Garden
Press, 200th Street & Kazimiroff Boulevard, Bronx, New
York 10458 USA. 600 pages. 65 USD Cloth.
THE CANADIAN FIELD-NATURALIST
Vol. 121
MISCELLANEOUS
Culturing Wilderness in Jasper National Park. By LS.
MacLaren. 2008. University of Alberta Press, Ring House 2,
Edmonton, Alberta T6G 2E1 Canada. 312 pages. 39.95 CAD
Paper.
Nahanni Journals — R.M. Patterson’s 1927-1929 Journals.
Edited by R. Davis. 2008. University of Alberta Press, Ring
House 2, Edmonton, Alberta T6G 2E1 Canada. 316 pages.
29.95 CAD Paper.
Ocean. 2008. By Dorling Kindersley. Tourmaline Editions
Inc., 662 King Street West, Suite 504, Toronto, Ontario
M5V 1M7 Canada. 512 pages. 27.95 USD Paper.
*Birdwatcher: The Life of Roger Tory Peterson. By
Elizabeth J. Rosenthal. 2008. The Lyons Press/Globe Pequot
Press, Guilford, Connecticutt USA. 437 pages.
Smithsonian Rock and Gem. 2008. By Dorling Kindersley,
Tourmaline Editions Inc., 662 King Street West, Suite 504,
Toronto, Ontario M5V 1M7 Canada. 360 pages. 22.95 USD
Paper.
Shelter at the Shore — The Boathouses of Muskoka. By J.
De Visser and J. Ross. 2008. Tourmaline Editions Inc., 662
King Street West, Suite 504, Toronto, Ontario M5V 1M7
Canada. 72 pages. 11.95 USD Paper.
YOUNG NATURALISTS
Dangerous Dinosaurs Q & A. 2008. By Dorling Kindersley.
Tourmaline Editions Inc., 662 King Street West, Suite 504,
Toronto, Ontario MSV 1M7 Canada. 64 pages. 14.99 USD
Paper.
Encyclopedia of Dinosaurs & Prehistoric Life. 2008. By
Dorling Kindersley. Tourmaline Editions Inc., 662 King Street
West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 376
pages. 22.99 USD Paper.
My Terrific Dinosaur Book. 2008. By Dorling Kindersley.
Tourmaline Editions Inc., 662 King Street West, Suite 504,
Toronto, Ontario M5V 1M7 Canada. 14 pages. 14.99 USD
Cloth.
Earth Matters. 2008. By Dorling Kindersley. Tourmaline
Editions Inc., 662 King Street West, Suite 504, Toronto, On-
tario MSV 1M7 Canada. 256 pages. 27.99 USD Cloth.
Eyewitness Climate Change. 2008. By Dorling Kindersley.
Tourmaline Editions Inc., 662 King Street West, Suite 504,
Toronto, Ontario MSV 1M7 Canada. 72 pages. 18.99 USD
Cloth.
Sharpedia. 2008. By Dorling Kindersley. Tourmaline Editions
Inc., 662 King Street West, Suite 504, Toronto, Ontario
MSV 1M7 Canada. 128 pages. 14.99 USD Cloth.
News and Comment
Bill Cody Receives Honorary Doctorate of Science from McMaster University, June 5, 2007
William James (Bill) Cody was born in Hamilton, Ontario
in 1922. He grew up in Hamilton during the Depression.
During summers, he worked on his uncle Bruce’s fruit farm
near St. David’s in the Niagara Peninsula. He always enjoyed
the outdoors, but it was with the help of McMaster University’s
Professor Lulu Gaiser that his love for plants grew. Bill re-
ceived his bachelor’s degree from McMaster University in
1946,
That same year, Bill joined the Canada Department of
Agriculture at the Central Experimental Farm, in Ottawa, as
Junior Agricultural Assistant. He was subsequently promoted
to Senior Agricultural Assistant, Technical Officer and then
Research Assistant. His enthusiasm for the outdoors led him
to travel extensively during the summer months across nine
provinces and two territories. During these excursions, he
collected over 35,000 plant specimens. He has published over
343 articles, including books, scientific papers, and popular
articles. He is Canada’s leading expert on the flora of the Yukon
and is an Honorary Research Associate with Agriculture and
Agri-Food Canada.
Bill was appointed Curator of the Vascular Plant Herbarium
in 1959. He held this position for 28 years. Under his super-
vision it became the largest vascular plant herbarium in Can-
ada with more than one million specimens, including the thou-
sands collected by him personally. The herbarium provides a
basis for research and identification of economically impor-
tant plants in Canada.
Officially retiring in 1987, after 41 years of service, Bill
continues to go to the office. His expertise in identifying spec-
imens and his knowledge is sought after by the younger gen-
eration of botanists. He remains an internationally recognized
Marine Turtle Newsletter (116)
April 2007. 36 pages: EDITORIALS: Proceeding to the
future (M. S. Coyne) — The oldest place where there is
always something new (J. G. Frazier) — ARTICLES: An up-
date on marine turtles in Cote d’Ivoire, West Aftrica (J. G.
Penate, M. Karamoko, S. Bamba, and G. Djadji) — An
update on sea turtle conservation activities in the Republic of
Congo (G. Bal, N. Brehert, and H. Vanleeuwe) — Conflicts
and social dilemmas associated with the incidental capture
of marine turtles by artisanal fishers in Benin (J. S. Dossa,
B. A. Somsom and G. A. Mensah) — Marine turtle mortali-
ty in southern Gabon and northern Congo (R. Parnell, B.
Verhage, S. L. Deem, H. Van Leeuwe, T. Nishihara, C. Mou-
koula, and A. Gibudi) — Artificial lights as a significant
cause of morbidity of Leatherbacks in Pongara National
Park, Gabon (S. L. Deem. F. Boussamba, A. Z. Nguema,
G-P Sounguet, S. Bourgeois, J. Cianciolo, and A. Formia)
— Observations of marine turtles in relation to seismic air-
gun sond off Angola (C. R. Weir) — Fibropapillomatosis
confirmed in Chelonia mydas in the Gulf of Guinea, West
Africa (A. Formia, S. Deem, A. Billes, S. Ngouessono, R.
Parnell, T. Collins, G-P Sounguet, A. Gibundi, A. Villarubia,
authority in Canada and continues to contribute to the devel-
opment of the collection.
He joined the Ottawa Field-Naturalist’s Club when he
arrived in Ottawa in 1946. The position of Business Manager
of a scientific journal, The Canadian Field-Naturalist was his
for over 50 years. His duties included managing finances,
monitoring stock, billing for reprints and public relations.
Bill was married for 47 years to Lois and together they
raised five children: David, Margaret, Leslie, Douglas and
Gordon. Having sung in a barber shop quartet and church
choir, his joy of music was very evident. Bill’s baritone voice
could be heard resonating throughout the halls of his office
and apartment building. He often had a pun or quip on the tip
of his tongue.
In recent years, Bill had the opportunity to return to the
Yukon four times. Each time, he took one of his children to
help him collect and log specimens in order to determine
range extensions and also to let them experience a place that
had been a large part of his life.
Bill is very proud of his family heritage. His father, William
MacPherson Cody, was the first appointed anesthetist to the
Hamilton General Hospital. Also, his second cousin, 3 times
removed, was William Frederick Cody, known to most as
“Buffalo Bill” Cody, a famous hero of the American Wild
West.
Among the tributes awarded to Bill are the following: the
Lawson Medal, most prestigious award of the Canadian Bo-
tanical Association (1997); the Distinguished Technical Comm-
unication Award of the Society for Technical Communica-
tions (1997); induction into the McMaster University Alumni
Gallery (2002); The Queen’s Golden Jubilee Medal (2002):
and the Yukon Biodiversity Awareness Award (2006).
G. H. Balazs, and T. R. Spraker) — Presence of sea turtles
in traditional pharmacopoeia and beliefs of West Africa (J.
Fretey, G. H. Segniagbeto, and M’M. Soumah) — Discovery
of a Gabonese Leatherback in South Africa (J. Fretey, A.
Billes, B. Baxter, and Caterine Hughes) — Second TAMAR-
tagged Hawksbill recaptured in Corisco Bay, West Africa
(A. Grossman, C. Bellini, A. Fallabrino, A. Formia, J. M. Mba,
J. N. Mba, and C. Obama) — IUCN-MTSG QUARTERLY
UPDATE — MEETING REPORTS — NEWS & LEGAL BRIEFS
— RECENT PUBLICATIONS — ACKNOWLEDGEMENTS
The Marine Turtle Newsletter is edited by Lisa M. Camp-
bell, Nicholas School of Environment and Earth Sciences,
Duke University, 135 Duke Marine Lab Road, Beaufort, North
Carolina 28516 USA; and Matthew H. Godfrey, North
Carolina Wildlife Resources Commission, 1507 Ann Street,
Beaufort, North Carolina 28516 USA. Subscriptions and
donations towards the production of the MTN can be made
online at or postal mail to
Michael Coyne (Managing Editor), Marine Turtle Newsletter,
1 Southampton Place, Durham, North Carolina 27705 USA:
e-mail: mcoyne @seaturtle.org.
234
THE CANADIAN FIELD-NATURALIST
Vol. 121
Atlas of amphibians and reptiles of Quebec, 2007 inventory
Instructions for participants in the 20" herpetological
survey for the Québec Atlas of amphibians and reptiles,
coordinated by the St. Lawrence Natural History Society in
collaboration with the Ministere des Ressources naturelles de
la Faune du Québec. The data from this annual inventory pro-
gram are incorporated into the Centre de donnees sur le pat-
rimone natural du Québec database; to date this database con-
tains more than 60 000 date entries on Québec species. This
provincial inventory is done every year with the important co-
operation of volunteers. Contributors are encouraged to send
their e-mail address and request for instruction on trasmitting
data, Observation cards in paper format are available on re-
quest.
Mathieu Oulette, Research Officer, Research and Conser-
vation, Ecomuseum — St. Lawrence Valley Natural History
Society, 21 125 chemin Saint-Marie, Saint-Anne-de-Bellevue,
Quebec H9X 3Y7; phone 514-457-9449; fax 514-457-0769;
e-mail: info @herpetofaune.org.
The Boreal Dip Net/L’epousette boreale 11(1) Spring 2007
Newsletter of the Canadian Amphibian and Reptile Con-
servation Network/Réseau Canadien de Conservation des
Amphibians et Reptiles
Contents: Editor’s Note (Sara Ashpole) — Notes from the
chairperson (David Galbraith) — Presentaions at CARC-
NET/RECCAR 2006 Annual Meeting, October, Victoria,
British Columbia: Student Presentation Awards: Best
Platform in 2006 Julie A. Lee-Yaw — Best Poster in 2006
Isabelle Deguise — 2006 Silver Salamander Award Mike
Rankin (posthumously) — 2006 Silver Salamander Award:
Fish & Wildlife Compensation Program: Jean Wai Jang —
2006 Silver Salamander Award: Gord McAdams and the West
Kootenay Ecosociety: They stuck their necks out to save the
turtles — 2006 Scholarship for Amphibian and Reptile Re-
search: Gregory Bulte — Keynote Speakers with Extended
Abstracts : Captive Breeding of Amphibians - Conservation
or Cosmetic Surgery? (Richard A. Griffiths); The Way of a
Gecko on a Rock — Why the Animal’s Real Environment
Matters — How the Student Presentations are Judged —
Presentations Made (titles and authors of 50 presentations)
— Book notice: Amphibians and Reptiles of British Columbia
— Towards a Cooperative Strategy for Dealing with Disease
Issues in Amphibian and Reptile Conservation — Blanding’s
Turtle Management Symposium and Roads and Ecopassages
Forum — Announcement: CARRCNET/RECCAR Annual
General Meeting 21 to 24 September 2007 Kingston,
Ontario — CARCNET/RECCAR Board of Directors.
Canadian Association of Herpetologists/Association Canadienne des Herpetologists Bulletin
14(2) Spring 2007
Contents: Instructions for Authors — EDITORIAL NOTES:
Membership and Association News — MEETINGS: CARCNet
11" Annual Meeting 27 to 29 October in Victoria, British
Columbia; Joint Meeting of Ichthylogists and Herpetologists
in St. Louis, Missouri, 11 to 16 July 2007 — FEATURE
ARTICLE: Accuracy and precision of site occupancy estimates
of anuran acoustic surveys when incorporating detection prob-
abilities (Shane de Solla) — FIELD NOTES: Use of digital
images obtained from volunteers to increase sample size and
accuracy of morphometric ratios (Wayne Weller) — BOOK
REVIEWS: Herpetological Conservation and Biology [an
online journal] (assessed by Mare J. Mazerolle, Jim Bogart,
Kate Jackson) — Diseases of Amphibians and Reptiles (re-
viewed by Dale A. Smith) — THESIS ABSTRACTS IN CAN-
ADIAN HERPETOLOGY: Sara M. Delannoy MSc 2006 Univer-
sity of Calgary, Alberta (Supervisor Anthony P. Russell)
Subdigital setae of the Tokay Gecko: Variation in form and im-
plication for adhesion; Patrick S. Druckenmiller PhD 2006
University of Calgary, Alberta (Supervisor Anthony P. Russell)
Early Cretaceous Plesiosaurs (Sauroppterygia: Plesiosaura)
from northern Alberta: Palaeoenvironmental and systematic
implications; Virgil C. Hawkes MSc 2007 University of Vic-
toria (Supervisor Patrick Gregory) Riparian management and
ampibians: Does buffer width matter?; Geoffrey Hughes
Honours BSc 2007 Laurentian University, Sudbury, Ontario
(Supervisor Jacqueline Litzgus) Nest site selection by Wood
Turtles (Glyptemys insculpta) in a thermally limited environ-
ment; Heather A. Jamniezky PhD 2006 University of Calgary,
Alberta (Supervisor Anthony P. Russell) Turtle cranial arterial
circulation: Intergrative analysis of a systematically influen-
tial character complex; Sheila Smith BSc 2006 Laurentian
University, Sudbury, Ontario (Supervisor Jacqueline Litzgus)
Geographic variation of sexual size dimorphism in Painted
Turtles, Chrysemys picta; Eric D. Snively PhD 2006 Univer-
sity of Calgary, Alberta (Supervisor Anthony P. Russell) Neck
musculoskeletal function in the Tyrannosauridae (Theropoda,
Coelurosauria): Implications for feeding dynamics — THESES
IN HERPETOLOGY AT THE UNIVERSITY OF VICTORIA (Patrick
Gregory) a list by student and title for 33 theses (1974-2005):
9 BSc, 18 MSc, 6 PhD — RECENT PUBLICATIONS IN
CANADIAN HERPETOLOGY: 17: 2006 (15), 2007(1), in
press (1) — ANNOUNCEMENTS New Fiction Book about Sea
Turtles: Sand Turtles.
Editor’s Report for Volume 120 (2006)
Mailing dates for issues in volume 120(1) 15 July TABLE 2. Number of articles and notes published in The Can
2007, (2) 1 November 2007; (3) 30 November 2007; (4) @4/an Field-Naturalist Volume 120 (2006) by major field of
31 December 2007. study.
A summary of circulation to members and sub- Subject. ~~~~~-~Arficiles”~—~—sONotes”~—=~=S*«*STowal
scribers (institutional and individual) totals for 2006
is given in Table 1. The number of articles and notes in ne ae = : =
volume 121 is summarized in Table 2 by topic; totals for eae: :
; d ; : Amphibians + reptiles | I 2
Book Reviews and New Titles are given in Table 3, and 5, 5 4 7
the distribution of content by page totals per issue in Iycects 3 0 3
Table 4. ' : Other invertebrates | | 2
Council continued to authorize 40% of membership Plants 13 0 13
dues for publication of the journal and all of sub- Paleontology | 0
scriptions (both individual and institutional). Council — Tributes 2 0 2
also has allocated 80% of the annual interest from the ea ,
Totals 56 16 72
Manning Fund and other capital funds to The Canadian
Field-Naturalist. Use of the Manning fund portion was
broadened on an issue-by-issue basis to offset the pub-
lication costs where authors lacked grant or institutional
funds to support publication. tents and abstracts are now posted on the Ottawa Field-
The journal was printed at Gilmore Printers, Ottawa, Naturalists’ Club website by webmaster Sandra Gar-
and thanks are due business representative Tom Smith; land, and PDFs may be ordered by authors as well as
customer representative Chuck Graham for overseeing _ traditional print reprints.
production, and Wendy Cotie for typesetting and cor- Manuscripts (excluding book reviews, notices, and
rections. Leslie Cody prepared the Index for volume reports) submitted to The Canadian Field-Naturalist
120; Elizabeth Morton proofed the galleys. When Bill _ totalled 67 in 2006, down by 8 from the total for 2005.
Cody, Businesss Manager for 60 years, was unable to The following reviewed for papers submitted in 2006
carry on, most of his duties were shouldered by Trea- (with number of manuscripts reviewed in parentheses
surer Frank Pope and Assistant Treasurer Jim Ward _ if more than one):
with invoice preparation assumed by the editor. Roy Associate Editors: R. Anderson, Canadian Museum of
John arranged and edited book reviews and prepared the Nature, Ottawa, Ontario; C. D. Bird, Erskine, Alberta (8); R.
New Titles listings. Special thanks are due to Jenifer i woe Pa aCe ace elie pe ae
aed eee aie iste, a ae and 2 Canadian Museum of Nature, Ottawa, Ontario (6); A. UE Er-
‘ ; u : : : skine, Sackville, New Brunswick (10); D. F. McAlpine, New
ing for 120(1) the eight papers on conservation evalu- Brunswick Museum, Saint John, New Brunswick (4); D. W.
ation of some British Columbia plants originated by the — Nagorsen, Mammalia Biological Consulting, Victoria, British
late George Douglas and also for preparing the tribute — Columbia (8); W. O. Pruitt, Jr., University of Manitoba, Win-
and updated bibliography for Douglas. The journal con- _nipeg, Manitoba (6).
TABLE 1. The 2006 circulation of The Canadian Field-Naturalist (2005 in parentheses). Prepared by Robin Murphy, bmr
group, Ottawa. Totals are copies mailed. As some members opt not to receive the journal, membership total here is only those
receiving the journal. For total club membership see the Annual Report of the membership committee to the 2007 Annual Business
Meeting.
Canada USA Other Totals
Memberships
Family & individual 556 (624) 24 (26) 5 (6) 585 (656)
Subscriptions
Individuals 131 (130) 57 (63) 5 (4) 193 (197)
Institutions 158 (156) 232 (229) 32); 64) 422 (419)
Ags 845 (910) 313 (318) 42 (44) 1200 (1272)
Note: 20 countries (outside Canada and United States) are included under “Other” (number of copies in parentheses):
Australia, Austria, Belgium, Brazil, Denmark, United Kingdom (9: including | to Scotland), Finland (2), France (2),
Germany (2), Iceland, Ireland, Japan, Netherlands (2), New Zealand, Norway (6), Poland, Russia, South Africa, Spain (2),
) Switzerland, Taiwan (2).
235
236
TABLE 3. Number of reviews and new titles published in Book
Review section of The Canadian Field-Naturalist Volume 120
(2006) by topic.
Reviews New Titles
Zoology 37 126
Botany 4 1S)
Environment 9 58
Miscellaneous 10 4
Young Naturalists 1 5
Totals 61 208
Other Reviewers: P. Achuff, Waterton Lakes, Alberta; W.
B. Ballard, Texas Tech University, Lubbock, Texas (4); J. R.
Bider, Baie d’Urfe, Quebec (2); G. Blouin-Demers, University
of Ottawa, Ontario; E. L. Bousfield, Victoria, British Columbia
(2); K. Conlan, Canadian Museum of Nature, Ottawa (2); D.
Boyd, Helena, Montana; R. J. Brooks, University of Guelph,
Ontario; S. Carriere, Environment and Natural Resources,
Yellowknife, Northwest Territories; R. W. Campbell, Bio-
diversity Centre for Wildlife Studies, Victoria, British Colum-
bia; J. A. Cook, Museum of Southwestern Biology, University
of New Mexico, Albuquerque; S. Cooke, Carleton University,
Ottawa, Ontario (2); W. J. Crins, Ontario Ministry of Natural
Resources, Peterborough, Ontario; D. B. Donald, Environment
Canada, Regina, Saskatchewan; J. Dubois, Manitoba Natural
Resources, Winnipeg, Manitoba; J. Ferron, Universite du
Quebec a Rimouski, Quebec; D. R. Galbraith, Royal Botanical
Gardens, Hamilton, Ontario; A. J. Gaston, Environment Can-
ada, Gatineau, Quebec; D. Giblin, Burke Museum, University
of Washington, Seattle, Washington; J. Gilhen, Nova Scotia
Museum of Natural History, Halifax, Nova Scotia (2); P.
Goossen, Environment Canada, Edmonton, Alberta; P. T.
Gregory, University of Victoria, British Columbia; F. H. Har-
rington, Mt. St. Vincent University, Halifax, Nova Scotia; S.
J. Hecnar, Lakehead University, Thunder Bay, Ontario; E.
Hendrycks, Canadian Museum of Nature, Ottawa, Ontario;
P. Hicklin, Environment Canada, Sackville, New Brunswick;
L. V. Hills, Cochrane, Alberta; S. Hooker, University of St.
Andrews, Fife, United Kingdom; B. D. Johnson, Herbarium,
Northern Forest Research Centre, Canadian Forest Service,
Edmonton, Alberta (2); C. S. Houston, Saskatoon, Saskat-
chewan; J. Kamler, Polish Academy of Sciences, Bialowieza,
Poland; C. R. Lacroix, University of Prince Edward Island,
Charlottetown; J. Lanteigne, Travaux publics et services,
gouvernementaux Canada, Gatineau, Quebec; G. Lee, Ottawa,
Ontario; R. MacCulloch, Royal Ontario Museum, Toronto,
Ontario; J. Madill, Canadian Museum of Nature, Ottawa,
Ontario; F. F. Mallory, Laurentian University, Sudbury, Ontario
(2); M. Mallory, Environment Canada, Iqaluit, Nunavut;
André Martel, Canadian Museum of Nature, Ottawa, Ontario;
L. D. Mech, U.S. Geological Survey, The Raptor Center,
University of Minnesota, St. Paul (3); J. Metcalfe-Smith,
Aquatic Ecosystem Protection Branch, National Water Re-
search Institute, Burlington, Ontario; A. Moehrenschlager,
Calgary Zoological Society, Alberta; E. L. Mills, Dalhousie
University, Halifax, Nova Scotia; G. Morrison, Environment
Canada, Gatineau. Quebec; G. Parker, Environment Canada,
Sackville, New Brunswick; E. Pip, University of Winnipeg,
Manitoba; G. Pohle, Huntsman Marine Science Centre, St.
Andrews, New Brunswick; W. Preston, Winnipeg, Manitoba;
G. Proulx, Alpha Research & Management Ltd., Sherwood
THE CANADIAN FIELD-NATURALIST
Vol. 120
TABLE 4. Number of pages per section published in The Cana-
dian Field-Naturalist Volume 120 (2006) by issue.
(1) (2) (3) (4) Total
Articles 105 85 104 68 362
Notes 7 11 3 12 33
Tributes 0 12 9 0 21
Book Reviews 19 15 13 19 66
CEN/OFNC Reports” °0 2 9 5 16
News and Comment 2 2 1 D) 7
Index 0 0 0 21 21
Advice to Contributors | 1 1 1 4
Totals ey ARS IX) 128 530
* Total pages for book review section include both reviews
and new titles listings.
™ Includes CEN Editor’s report in (2), OFNC Annual Business
Meeting (3) and OFNC Awards (4).
Park, Alberta (2); R. Reeves, Okapi Wildlife Associates,
Hudson, Quebec (2); T. E. Reimchen, University of British
Columbia, Victoria, British Columbia; G. J. Robertson, Envi-
ronment Canada, Mount Pearl, Newfoundland; L. Rogers, .
Ely, Minnesota; R. Rosatte, Ministry of Natural Resources,
Trent University, Peterborough, Ontario; A. Russell, University
of Calgary, Calgary, Alberta; F. Scott, Acadia University,
Wolfville, Nova Scotia; F. W. Schueler, Bishops Mills Natural
History Center, Oxford Mills, Ontario; J. W. Sheard, University
of Saskatchewan, Saskatoon; Brian Slough, Whitehorse,
Yukon Territory; K. W. Stewart, University of Manitoba,
Winnipeg, Manitoba (3); P. Weatherhead, Queen’s Biological
Station, Elgin, Ontario; M. J. Wetzell, Illinois, Natural History
Survey, Champaign, Illinois; N. Yan, York University, North
York, Ontario.
I am also indebted once again to the President of
the Ottawa Field-Naturalists’ Club, Gary McNulty, and
the Club Council for continuing support of the journal:
Chairman Ron Bedford and the Publications Com-
mittee of the OFNC for editorial encouragement and
support, the Finance Committee for input and encour-
agement to speed things up, the Canadian Museum of '
Nature for access to its library and the facilities at the «
Natural Heritage Building, 1740 Pink Road, Aylmer, |
Quebec, and to Joyce for everything else.
I must add my special thanks due William J. (Bill)
Cody who served the club as council member for more «
than 60 years, vitually all ot this time as Business!
Manager of The Canadian Field-Naturalist. Those of)
us exposed to Bill’s unfailing good humour, determi-'
nation, efficient focus on the job on hand, and some-:
times his inevitable serenades, have lasting memories’
of the time shared. Bill served with, and kept each on)
track, five editors (one twice) — Harold Senn to 1955),
Bob Hamilton (1956-1961), myself (1962-1966), Ted
Mosquin (1967-1972), Lorraine Smith (1972-1981,
and me again (1981 — present). Efficient management
and financial stability through good times and bad_
were due in large measure to Bill’s initiative which
assured, in some periods, the journals very survival.
2007
A decade ago, to mark 50 years of service, the club
paid tribute to Bill’s contribution to that point with a
reception at the Central Experimental Farm, Depart-
ment of Agriculture, Ottawa, where Bill had spent his
whole career, including “retirement”, as a research
botanist. Stephen Darbyshire for Awards Committee
stated in part (1998. Canadian Field-Naturalist 1 12(4):
719-720):
“Such long-term service and dedication is unprece-
dented in the history of the Club and is not likely to ever
be repeated. The titles of office do not reveal the extent
to which Bill has laboured for the Club. One cannot
completely list the jobs that he has taken on (even Bill
cannot remember them all), many of which are among
the most distasteful and time consuming (imagine the
years of dealing with government forms for postage
rates, charitable status, etc.). Executive officers have
7
EpIror’s REPORT
237
come and gone, Canadian Field-Naturalist editors have
come and gone, Council is re-elected every year, but
Bill Cody continues as a major constituent of the glue
that keeps the Club together and keeps it, and its scien-
tific publication, running smoothly.”
Bill continued to perform these duties for close to
another decade before failing health forced him pass
them on to others. The full truth in the earlier words
of tribute has become increasingly realized by those
of us Bill left to share his multitude of tasks. Most of
the burden has fallen on Frank Pope, already contri-
buting tremendous service as Treasurer of the Ottawa
Field-Naturalist’s Club, and Jim Ward as Treasurer's
Assistant. They have made the transition work.
FRANCIS R. COOK
Editor
238
THE CANADIAN FIELD-NATURALIST
Vol. 120
Advice for Contributors to The Canadian Field-Naturalist
Content
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of investigations in any field of natural history provided that
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Review Editor concerning suitability of manuscripts for this
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for the Ottawa Field-Naturalists’ Club, 1983. The Canadian
Field-Naturalist 97(2): 231-234. Potential contributors who
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FRANCIS R. CooK, Editor
RR 3 North Augusta, Ontario KOG IRO Canada
TABLE OF CONTENTS (concluded) Volume 121 Number 2 2007
Notes (continued)
Human-assisted movements of Raccoons, Procyon lotor, and Opossums, Didelphis virginiana, between
the United States and Canada Rick C. ROSATTE, DENNIS DONOVAN,
MIKE ALLAN, LAURA BRUCE, and CHRIS DAVIES PAV
Correlation between age estimates for Elk, Cervus elaphus, using tooth wear/eruption patterns and
counts of annuli in tooth cementum
RICK ROSATTE, ANDREW SILVER, MARK GIBSON, BRUCE CHISHOLM, and NORM CooL 214
Predator-prey interaction between and American Robin, Turdus migratorius, and a Five-lined Skink,
Eumeces fasciatus E. NATASHA V ANDERHOFF 216
Book Reviews
Zoo.ocy: Atlas of Breeding Birds of Ontario 2001-2005 — The Return of Caribou to Ungava — A Guide
to the Mammals of China — Birds of the Saskatoon Area — Brian Coad’s Ichthyology Site 219
Borany: Exploration and Utilization of Chinese Spice Plant Resources pipe)
MISCELLANEOUS: Arctic Hell Ship — Eighteenth-century Naturalists of Hudson Bay — Hunters at the
Margin: Native People and Wildlife Conservation in the Northwest Territories — The Ornithologist’s
Dictionary — Pioneering Women in Plant Pathology — Birdwatcher: The Life of Roger Tory Peterson
226
New TITLES DBD
News and Comment
Bill Cody receives Honorary Doctorate of Science from McMaster University, June 2007 — Marine Turtle
Newsletter 116 April 2007 — Atlas of amphibians and reptiles of Quebec, 2007 inventory — The Boreal
Dip Net/L'epuosette boreale 11(1) Spring 2007 — Canadian Association of Herpetologists/Association
Canadien des Herpetologists Bulletin 14(2) Spring 2007 233
Editor’s Report for Volume 120 (2006) FRANCIS R. COOK 235
Advice to Contributors 238
Mailing date of the previous issue 121(1): 6 June 2008
THE CANADIAN FIELD-NATURALIST Volume 121 Number 2
Articles
Diversity and distribution of the terrestrial mammals of the Yukon Territory: A review
BRIAN G. SLOUGH and THOMAS S. JUNG
State records and habitat of Clam Shrimp, Caenestheriella gynecia (Crustacea: Conchostraca),
in New York and New Jersey ROBERT E. SCHMIDT and ERIK KIVIAT
Colonization of non-traditional range in dispersing Elk, Cervus elaphus nelsoni,
populations FRED VAN DYKE
The Rumsfeld Paradigm: Knowns and Unknowns in characterizing habitats used by the
endangered Sharp-tailed Snake, Contia tenuis, in southwestern British Columbia
SHANNON F. WILKINSON, PATRICK T. GREGORY, CHRISTIAN ENGELSTOFT, and KARL J. NELSON
Do repugnant scents increase survival of ground nests?
VANESSA B. HARRIMAN, JUSTIN A. PITT, and SERGE LARIVIERE
Translocation and recovery efforts for the Telkwa Caribou, Rangifer tarandus caribou,
herd in westcentral British Columbia, 1997-2005
ASTRID VIK STRONEN, PAUL PAQUET, STEPHEN HERRERO,
SEAN SHARPE, and NIGEL WATERS
An inventory of aquatic and subaquatic plants in SASK Water Canals in Central Saskatchewan,
Canada J. HuGo Cota-SANCHEZ and KIRSTEN REMARCHUK
Life history of the marine isopod Cyathura polita in the Saint John River Estuary, New
Brunswick: a species at the northern extent of its range
SARAH C. MERCER, GLENYS D. GIBSON, and MICHAEL J. DADSWELL
Post-emergence movements and overwintering of Snapping Turtles, Chelydra serpentina,
hatchlings in New York and New Hampshire
GORDON R. ULTScH, MATT DRAuD, and BARRY WICKLOW
Notes on succession in old fields in southeastern Ontario: Il The herbs
A. CROWDER, R. HARMSEN, and S. E. BLATT
Growth of White Spruce, Picea glauca, seedlings in relation to microenvironmental
conditions in a forest-prairie ecotone of southwestern Manitoba
SOPHAN CHHIN and G. GEOFF WANG
Natural recolonization of cultivated land by native prairie plants and its enhancement by
removal of Scots Pine, Pinus sylvestris PAUL CATLING and BARRY KING
Notes
A high elevation record of the Star-nosed Mole, Concylura cristata, in northeastern Vermont
RYAN W. Norris and C. WILLIAM KILPATRICK
Northern Long-eared Bat, Myotis septentrionalis, (Chiroptera: Vespertilionidae) on Price Edward Island:
First records of occurrence and over-wintering
JENNIFER A. BROWN, DONALD F. MCALPINE, and ROSEMARY CURLEY
Arboreal, late summer, courtship of Maritime Garter Snakes, Thamnophis sirtalis pallidulus,
in Dartmouth, Nova Scotia JOHN GILHEN and ROBERT STRUM 210
i MUCUS N (continued on inside back cover)
ier erhegmwccrand” SEN ISSN 0008-3550
FSC www.fscorg Cert no. SGS-COC-003284
© 1996 Forest Stewardship Council
2007
1]
12
133
14
15
206
The CANADIAN
FIELD-NATURALIST
Canada
ASTS’ CLUB, Ottawa,
Published by THE OTTAWA FIELD-NATURAI
July-September 2007
Volume 121, Number 3
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Her Excellency The Right Honourable Michaille Jean
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to)
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields|
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-)
ments of high quality for living things.
Honorary Members
Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope
Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr.
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson
2007 Council
President: Mike Murphy Annie Belair Diane Kitching Dan Millar
Vice-President: Ken Allison Ronald E. Bedford |= Karen McLachlan Hamilton Stanley Rosenbaum
Recording Secretary: Susan Laurie-Bourque Fenja Brodo David Hobden Henry Steger
Treasurer: Frank Pope Julia Cipriani Diane Lepage Chris Traynor
Past President: Gary McNulty William J. Cody Ann Mackenzie Eleanor Zurbrigg
Francis R. Cook Gillian Marston
To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069.
Westgate P.O. Ottawa, Canada K1Z 1A2
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed 1 ir
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. |
We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) towarc
our mailing costs. PAP Registration Number 9477. Canada
Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca
Copy Editor: Elizabeth Morton
Honorary Business Manager: William J. Cody
Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5; e-mail: r.john @rogers.com
Associate Editors: | Robert R. Anderson Robert R. Campbell Anthony J. Erskine Donald F. McAlpine
Warren B. Ballard Paul M. Catling David Nagorsen William O. Pruitt, Jr.
Charles D. Bird Brian W. Coad
Chairman, Publications Committee: Ronald E. Bedford
All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail
or e-mail.
Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor.
Subscriptions and Membership
Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 pe;
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) ani
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) mus
add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Locz
Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries <
$33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be maile
to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publication
Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: July-October 2007 (December 2008).
Cover: A Red-winged Blackbird nest with three blackbird eggs and five Brown-headed Cowbird eggs. The nest was found in
Conservation Reserve Program grassland in Day County, South Dakota. Photographed on 16 June 2005 by Lawrence I
Igl. See article on cowbird parasitism by Ig] and Johnson, pages 239-255.
]
|
Volume 121, Number 3 April
June 2007
Brown-headed Cowbird, Molothrus ater, Parasitism and Abundance in
the Northern Great Plains
LAWRENCE D. IGL! and DouGLAS H. JOHNSON?
'U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 37" Street SE, Jamestown, North Dakota 5840]
USA; e-mail: larry_ig] @usgs.gov
2 U.S. Geological Survey, Northern Prairie Wildlife Research Center, 204 Hodson Hall, 1980 Folwell Avenue, University of
Minnesota, St. Paul, Minnesota 55108 USA
Igl, Lawrence D., and Douglas H. Johnson. 2007. Brown-headed Cowbird, Molothrus ater, parasitism and abundance in the
northern Great Plains. Canadian Field-Naturalist 121(3): 239-255.
The Brown-headed Cowbird (Molothrus ater) reaches its highest abundance in the northern Great Plains, but much of our
understanding of cowbird ecology and host-parasite interactions comes from areas outside of this region. We examine cow-
bird brood parasitism and densities during two studies of breeding birds in the northern Great Plains during 1990-2006. We
found 2649 active nests of 75 species, including 746 nonpasserine nests and 1902 passerine nests. Overall, <1% of non-
passerine nests and 25% of passerine nests were parasitized by Brown-headed Cowbirds. Although the overall frequency of
cowbird parasitism in passerine nests in these two studies is considered moderate, the frequency of multiple parasitism among
parasitized nests was heavy (nearly 50%). The mean number of cowbird eggs per parasitized passerine nest was 1.9 + 1.2 (SD;
range = 1-8 cowbird eggs). The parasitism rates were 9.5% for passerines that typically nest in habitats characterized by
woody vegetation, 16.4% for grassland-nesting passerines, 4.7% for passerines known to consistently eject cowbird eggs,
and 28.2% for passerines that usually accept cowbird eggs. The Red-winged Blackbird (Agelaius phoeniceus) was the most
commonly parasitized species (43.1% parasitism, 49.6% multiple parasitism, 71.2% of all cases of parasitism). Passerine nests
found within areas of higher female cowbird abundance experienced higher frequencies of cowbird parasitism than those
found in areas of lower female cowbird abundance. Densities of female cowbirds were positively related to densities and rich-
ness of other birds in the breeding bird community.
Key Words: Brown-headed Cowbird, Molothrus ater. brood parasitism, grasslands, multiple parasitism, spatial variation, temporal
variation, northern Great Plains.
The Brown-headed Cowbird (Molothrus ater; here-
after cowbird) is an obligate brood parasite that lays its
eggs in nests of a variety of host species. Cowbird dis-
tribution, abundance, and brood parasitism have long
been major topics in avian ecology in North America
(Nice 1937; McGeen 1972; May and Robinson 1985;
Ortega 1998; Herkert et al. 2003). Cowbird abundance
and rates of brood parasitism vary geographically
(Robinson et al. 1995; Ortega 1998; Rothstein and
Robinson 1998). The cowbird reaches its highest abun-
dance in the northern Great Plains (Sauer et al. 2005*),
yet much of our understanding of cowbird parasitism
and the dynamics of host and parasite populations
comes from areas outside of this region. Nonetheless,
some studies have reported high rates of cowbird par-
asitism and multiple parasitism in the northern Great
Plains (e.g., Linz and Bolin 1982; Romig and Craw-
ford 1995; Davis and Sealy 2000; Koford et al. 2000).
Robinson and Smith (2000), however, suggested that
some of these oft-cited studies may not be representa-
tive of the northern Great Plains. There are few pub-
lished examples of community-wide studies of cow-
bird parasitism in the northern Great Plains (e.g., Gran-
fors et al. 2001; Woolfenden et al. 2004). Rather, most
reports of cowbird parasitism from this region focus
on a single species (e.g., Linz and Bolin 1982), a sub-
set of species (e.g., Winter et al. 2004), or species that
have evolved defenses against cowbird parasitism
(Sealy 1996).
At the continental level, cowbird abundance dec-
lines with distance from the center of the species’
breeding range in the northern Great Plains (Thomp-
son et al. 2000). The frequencies of cowbird parasitism
appear to follow a similar biogeographical pattern (e.g.
Smith and Myers-Smith 1998), especially for grass-
land birds (Jensen and Cully 2005a, 2005b), although
this relationship may not be generalized across habitats
(Robinson and Smith 2000) and may not be indicative
of parasitism frequencies at the local level (Chace et
al. 2005). Nonetheless, Chace et al. (2005) acknowl-
239
240
edged that, at the continental scale, cowbird abundance
appears to be a reasonable predictor of cowbird para-
sitism levels. Because of the difficulty in obtaining data
on both cowbird abundance and brood parasitism across
the continent (or even over a large region), there are
few examples of studies that have evaluated this rela-
tionship using cowbird and nest data collected concur-
rently at the same study sites over a large region (e.g.,
Robinson et al. 2000; Jensen and Cully 2005a, 2005b).
Only a few evaluations of this biogeographical rela-
tionship have included data from study sites in the
northern Great Plains (e.g., Smith and Myers-Smith
1998; Herkert et al. 2003).
At local and regional scales, geographic differences
in cowbird distribution and abundance also may result
from differences in avian communities (e.g., Farmer
1999; Thompson et al. 2000). That is, given that cow-
birds are host generalists, measures of avian abundance
or richness might serve as proximate cues for cowbirds
in determining where to settle (McGeen 1972; Farmer
1999). Chace et al. (2005) listed four nested levels of
an avian community that cowbirds could use as cues
for settlement: (1) all breeding birds in the avian com-
munity, (2) all passerine species, (3) all host species,
and (4) a single host species. Very few studies have
evaluated this relationship at all four levels (Farmer
1999). Using BBS (Breeding Bird Survey) data, Hahn
and O’Connor (2002) concluded that cowbird distri-
bution was not simply the result of shared habitat
preferences with their hosts, but rather host abun-
dance was an important predictor of cowbird occur-
rence in recently colonized regions of the United
States (1.e., eastern and western states) and to a lesser
extent in the cowbird’s ancestral range (i.e., central
Great Plains). Except for Hahn and O’Connor’s (2002)
evaluation using BBS data, the influences of avian
abundance and richness on cowbird distribution and
abundance have not been evaluated using regional
data from the northern Great Plains (reviewed in
Chace et al. 2005).
In this paper, we report cowbird densities and par-
asitism frequencies during two breeding-bird studies
in the northern Great Plains: a statewide study in
North Dakota (1992 and 1993: Igl and Johnson 1997;
Igl et al. 1999) and a Conservation Reserve Program
(CRP) grassland study in nine counties of four states
(1990-2006: Johnson and Schwartz 1993a, 1993b; Igl
and Johnson 1995, 1999; Johnson and Igl 1995, 2001).
We use data from nests found incidentally during both
studies to examine weekly and annual variation in
parasitism rates in the northern Great Plains, and nest
data from the CRP grassland study to examine region-
al variation in parasitism rates. We use data from the
long-term CRP grassland study to evaluate the rela-
tionships between female cowbird densities and cow-
bird parasitism and between female cowbird densities
and avian densities and richness in different regions
of the northern Great Plains. CRP grasslands are a suit-
THE CANADIAN FIELD-NATURALIST
Vol. 121
able habitat to evaluate these relationships because this
habitat has been shown to be a dominant landscape
predictor of cowbird distribution in the United States
(Hahn and O’Connor 2002).
Study Areas
North Dakota statewide study
The study area for the North Dakota statewide
study was described in detail by Stewart and Kantrud
(1972) and Ig] and Johnson (1997) and is only briefly
described here. North Dakota was divided into eight
major strata based on biogeographical, physiographi-
cal, and ecological characteristics. From these eight
strata, 130 quarter-sections (about 64.75 ha each) were
selected randomly (Figure 1). The number of sample
units allocated to each stratum was proportional to the
area of the stratum. Landowners denied access to one
quarter-section in 1992 and a different quarter-section
ims 19935
CRP grassland study
The CRP study is an ongoing investigation (1990
to present) that is examining breeding-bird communi-
ties annually in nearly 350 CRP grassland fields in
nine counties of four states (Figure 2), including Fal-
lon and Sheridan counties in Montana; Hettinger,
Kidder, and Eddy counties in North Dakota; Butte,
McPherson, and Day counties in South Dakota; and
Grant County in Minnesota (Johnson and Schwartz
1993a, 1993b). Field size varied from <1.0 to 111.7
ha. In the northern Great Plains, most CRP land was
planted to a mixture of native and/or non-native grass-
es and legumes (Johnson and Schwartz 1993a).
Woody vegetation has encroached into some idle
CRP fields, and some CRP fields contain wetlands or
were partially inundated by water during a recent wet
period (Ig] 2001; Ig] and Johnson, unpublished data).
Methods
Breeding bird and cowbird surveys
In both studies, we conducted total area counts of
breeding birds using the strip-transect procedure em-
ployed by Stewart and Kantrud (1972: also see Ig] and
Johnson 1997). During the North Dakota statewide
study, bird surveys were conducted by two observers
on foot, and each observer surveyed cowbirds and other
breeding birds on a rectangular half (805 * 402 m;:
32.37 ha) of a quarter-section by following a standard-
ized survey route. This route was 100 m inside of, and
parallel to, the boundary of the rectangle. The rectan-
gular halves were usually surveyed simultaneously, and
an interval of 400 m was maintained between observers.
Deviations of up to 100 m from the route were often
necessary to adequately survey all habitats.
Bird species were identified by sight or sound. Counts
during precipitation and strong winds (> 24 km/h) were
avoided. Surveys of open country birds were con-
ducted between 0.5 h after sunrise and 15:00 CST.
Although some surveys occurred outside the time of
2007
FiGurRE |. Distribution of 130 quarter-sections (64.75 ha
each) in North Dakota in which bird surveys were
conducted during the breeding seasons in 1992 and
1993,
most active bird vocalizations (i.e., early morning or
late evening), Stewart and Kantrud (1972) concluded
that singing and other activities of open-country birds
were not appreciably affected by time of day (also
see Vickery 1995*). Quarter-sections containing exten-
sive woodland habitats were usually covered on rela-
tively calm (<8 km/h), sunny days between 0.5 h after
sunrise and 10:00 CST. These limitations were neces-
sary because song frequencies and other activities of
most woodland birds are reduced on cloudy days, in
moderate or high winds, and at mid-day.
Counts of breeding birds were based primarily on
the number of indicated breeding pairs on territories
or home ranges during peak breeding periods. All
sample units were surveyed for early-nesting species
between 24 April and 7 June, for mid-nesting species
between 14 May and 10 July, and for late-nesting
species between 22 May and 21 July. When a survey
was conducted during an overlapping portion of the
peak breeding periods, counts of early-, mid-, and late-
nesting species coincided. Thus, quarter-sections that
were visited between 22 May and 7 June were only
surveyed once, and those that were surveyed before
22 May were surveyed again after 7 June so as to in-
clude species from all three breeding periods. For most
species, nearly all indicated pairs were observed as
segregated pairs or as territorial males. For the sexually
dimorphic Brown-headed Cowbird, we separately
tallied the males and females seen per sample unit;
herein, we report only female cowbird densities. We did
not consider certain birds observed during the census-
es to be breeding and excluded them from our results.
These included late-migrating flocks and other birds
passing overhead in high, direct flight. By counting
birds only during their peak breeding periods, we max-
imized the potential for recording breeding birds and,
at the same time, minimized the likelihood of con-
founding breeding birds with migrants.
In the CRP study, total area counts of breeding
birds were conducted by using a minor modification of
the strip-transect procedures used by Stewart and Kan-
IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS
24)
Sheridan e
NDISaya Ne
es {Ys eA J
Hetiinger u Kidder
a
McPherson
zB
Day
SD
FIGURE 2. Counties in which Conservation Reserve Program
grassland fields were surveyed for breeding birds in
North Dakota, South Dakota, Minnesota, and Mon-
tana, 1990-2006.
trud (1972) and Ig] and Johnson (1997). Small fields
usually were surveyed by a single observer; large fields
typically were surveyed by two observers on foot, each
simultaneously covering about one-half of the field.
The number and configuration of transects were con-
sistent among years but varied depending upon the field
size and shape to achieve total coverage of a field. We
surveyed each CRP grassland field once each year
between 21 May and 9 July in 1990-2006.
Nests and cowbird parasitism
Prior to entering the field in each year, the observers
were trained in bird, nest, and egg identification. Nests
were found serendipitously during breeding bird sur-
veys and related research activities. Most nests were
found by unintentionally flushing an adult host (or
rarely a female cowbird) from the nest or nest vicinity.
The remainder of the nests were found by observers
as they were simply walking through a habitat. When
a nest was located, we inspected it and recorded the
species (based on the identification of the flushing
adult, the nest, or the nest contents) and the number of
eggs or nestlings of the host and of cowbirds. If the
nest species could not be identified at the time of the
initial observation, the nest location was temporarily
marked and then revisited and identified at the end of
the survey by an experienced observer. We classified
a nest as parasitized if it contained at least one cow-
bird egg or nestling, regardless of the stage of the nest-
ing cycle. Cowbird and host nestlings were identified
based on size, gape and flange colors, and down or
contour feather characteristics and development. We
were unable to return to nests to monitor their out-
come or to document additional brood parasitism. We
rarely located or checked the contents of nests of up-
per-canopy nesting species. Most of the nests that we
located were found in the egg-laying stage or the in-
cubation stage (see below); cowbirds lay most of their
eggs before or during the hosts’ egg-laying stage (Johns-
gard 1997). Although we found many nests in the
nest-building stage (i.e., before the onset of egg lay-
ing), we summarize here only active nests, which we
defined as nests attended by adults that contained at
least one host or cowbird egg and/or nestling. We also
242
found a few abandoned nests that contained cowbird
eggs, but these are not reported because cowbirds are
known to parasitize inactive nests (Freeman et al. 1990).
The presence of cowbird fledglings that were fed by
adult hosts also was noted during the surveys of breed-
ing birds, although an observation of a potential host
feeding a cowbird fledgling does not establish conclu-
sively the true foster-parentage of the fledgling cow-
bird (sensu Klein and Rosenberg 1986).
We use the frequency (%) of nests parasitized as an
index to the levels of parasitism. We define the frequen-
cy (%) of cowbird parasitism as the total number of
parasitized nests (*100) divided by the total number of
active nests found for a given species, group of species
(e.g., grassland birds), county, or time period (week
or year). We define the frequency (%) of multiple par-
asitism as the total number of nests containing more
than one cowbird egg (*100) divided by the total num-
ber of parasitized nests found for a given species, group
of species, county, or time period. The average (+ SD)
number of cowbird eggs per parasitized nest (1.e., par-
asitism intensity) was calculated by summing the num-
ber of cowbird eggs found in all parasitized nests in
both studies and dividing by the total number of par-
asitized nests. Although cowbirds may avoid parasitiz-
ing nests of ejector species (Sealy and Bazin 1995), we
include nests of all passerine species, because ejector
species are sometimes parasitized (e.g., Scott 1977;
Peer et al. 2000), because parasitism of ejector species
may vary geographically (e.g., Haas and Haas 1998)
or with time of sympatry with cowbirds (Briskie et al.
1992), and because it is difficult to estimate how often
ejector species are parasitized (e.g., Scott 1977). To
evaluate changes in cowbird parasitism within the
breeding season, we divided each month into four,
roughly weekly, periods (7, 8, 7[8] and 8 days, respec-
tively) (sensu Orians et al. 1989). For discussion pur-
poses, we grouped rates of parasitism and multiple par-
asitism into five categories: low (<10%), low-to-
moderate (10-20%), moderate (20-40%), moderate-
to-heavy (40-50%), and heavy (>50%) (sensu Ortega
1998: 184). Scientific names of potential host species
are included in Table 2.
To determine whether cowbirds avoid laying eggs
in previously parasitized nests (Mayfield 1965a; Ori-
ans et al. 1989), we evaluated the distribution pattern
of cowbird eggs (i.e., random or non-random) using a
zero-truncated Poisson distribution model for only para-
sitized nests (Lindsey 1997). A random distribution of
cowbirds eggs suggests that female cowbirds did not
discriminate among parasitized nests and that the pro-
portion of nests with 1, 2, 3, 4, ..., i cowbird eggs ap-
proximated a Poisson distribution (Orians et al. 1989).
A non-random distribution of cowbird eggs indicated
that the random distribution model was rejected
(P <0.01).
For both studies, we calculated overall parasitism
frequencies, mean densities of female cowbirds, mean
avian densities, and mean avian richness. For the CRP
THE CANADIAN FIELD-NATURALIST
Vol. 121
grassland study, we calculated mean avian density and
richness among five groups (all of which excluded cow-
birds): (1) all avian species, (2) all passerine species,
(3) all passerines known to have raised cowbird young
(Ortega 1998; Davis and Sealy 2000), (4) all passer-
ines excluding ejectors (Peer and Sealy 2004), and
(5) a single, preferred host (i.e., Red-winged Black-
bird; see below). We were interested in the regional
avian influences that affect cowbird distribution and
abundance rather than short-term influences, and thus
we averaged across the 17 years within a county and
did not analyze the survey data separately for individ-
ual years. For the CRP grassland study, we used female
cowbird densities both as an explanatory variable in
relation to rates of cowbird parasitism and as a res-
ponse variable in relation to avian community variables
(i.e., density and richness). Linear regression analyses
(PROC REG; SAS Institute, Inc. 2004) were performed
(1) to determine the relationship between overall cow-
bird parasitism and female cowbird densities, and (2)
to examine the relationship between female cowbird
densities and the five levels of the avian community.
Results
In the North Dakota study, we observed 160 breed-
ing bird species, including 78 nonpasserine species and
82 passerine species (Igl and Johnson 1997). Given that
grasslands, croplands, and wetlands are the three most
common habitats in North Dakota (Ig] and Johnson
1997), most of the common breeding bird species in
North Dakota are associated with such open habitats.
The five most abundant species in the North Dakota
study, in decreasing order of abundance, were the
Horned Lark, Chestnut-collared Longspur, Red-winged
Blackbird, Western Meadowlark, and Lark Bunting
(scientific names given in Table 2). Average breeding
bird densities in a quarter-section, excluding cow-
birds, were 126 breeding pairs per 100 ha in 1992 and
143.9 breeding pairs per 100 ha in 1993 (Table 1).
The average numbers of species observed in a quar-
ter-section (64.75 ha) were 18.5 species in 1992 and
21.6 species in 1993. Average cowbird densities were
3.0 female cowbirds per 100 ha in 1992 and 4.3
female cowbirds per 100 ha in 1993. Female Brown-
headed Cowbirds were found in 66.7 and 76.7% of the
quarter-sections in 1992 and 1993, respectively.
In the CRP grassland study, we observed 143 breed-
ing bird species, including 78 nonpasserine species and
65 passerine species. The five most abundant species,
in decreasing order of abundance, were Savannah Spar-
row, Grasshopper Sparrow, Red-winged Blackbird,
Clay-colored Sparrow, and Western Meadowlark. Av-
erage breeding-bird densities within a county (aver-
aged across years and excluding cowbirds) ranged
from 104.1 to 208.1 breeding pairs per 100 ha, and the
average number of species observed within a county
ranged from 26.8 to 52.7 species (Table 1). Average
cowbird densities within a county ranged from 0.6 to
8.0 female cowbirds per 100 ha, and the average annual
i
2007
IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS
243
TABLE |. Summary of Brown-headed Cowbird densities (females/100 ha), cowbird occurrence, breeding bird densities (all
species excluding cowbirds; breeding pairs/100 ha), and number of breeding bird species per quarter-section in two studies
in the northern Great Plains, 1990-2006,
Female
cowbird
Year or Study Location density
Mean SE
North Dakota statewide study
1992 3.0 0.3
1993 AS» (0/5
CRP grassland study (1990-2006)
Butte County, South Dakota 0.6 0.1
Day County, South Dakota 8.0 0.7
Eddy County, North Dakota 6.3 0.4
Fallon County, Montana 0:9" 10:2
Grant County, Minnesota 3h 0.4
Hettinger County, North Dakota 7) 0.5
Kidder County, North Dakota 519, 1033
McPherson County, South Dakota fell 0.5
Sheridan County, Montana 2.8 9 o 30. ,
frequency of female cowbird occurrence ranged from
12.4 to 54.1%. Average cowbird densities and frequen-
cies were lowest in the counties on the eastern and
western edges of our study area (Table 1, Figure 2).
This pattern mirrors cowbird distribution maps from
the North American BBS (Price et al. 1995; Sauer et
al. 2005*), which show cowbird abundance declining
both east and west of central North Dakota and South
Dakota.
We found 351 active nests of 51 species (18 non-
passerine and 33 passerine species) during the North
Dakota statewide study and 2298 active nests of 62
species (23 nonpasserine and 39 passerine species)
during the CRP grassland study, for a total of 2649
nests of 75 species (Table 2). Most of the nests of
nonpasserine species were found during the egg-
laying or incubation stages; 89.9% of the 746 nests
of nonpasserine species contained only eggs, and
10.1% contained only nestlings or both eggs and
nestlings. Only one nonpasserine nest was parasitized
by a cowbird (Table 2). An Upland Sandpiper nest
with four sandpiper eggs and one cowbird egg was
found in a CRP grassland field in Sheridan County,
Montana, on 28 June 1993. Hereafter, we summarize
data only for passerine nests.
Most of the active nests of passerines were found
during the egg-laying or incubation stages; 85.8% of the
1902 passerine nests contained only eggs, and 14.2%
of the nests contained only nestlings or both eggs and
nestlings. Overall, the parasitism rate of passerine nests
was moderate; 476 of the 1902 (25%) passerine nests
that we found during the two studies were parasitized
by cowbirds (Table 2).
Twenty-four of the 45 passerine species were found
parasitized (Table 2). Among species with 25 or more
nests, parasitism levels were low for Western Kingbird
(0%), Eastern Kingbird (1.7%), Brown Thrasher (3.8%),
Savannah Sparrow (9.8%), Chestnut-collared Longspur
Female Breeding Number
cowbird bird of breeding bird
occurrence density Species
Mean % SE Mean SE Mean SE
0.67 - 126.0 7.8 18.5 0.8
0.77 ~ 143.9 7,3 216 09
0.14 0.02 105.8 5.9 28.5 1.4
0.54 0.03 208.1 14.5 48.8 2.7
0.49 0.03 188.7 15.1 45.5 2.5
0.12 0.02 104.1 9.| 26.8 1.2
0.25 0.02 181.8 13.7 39.6 2.2
0.52 0.03 160.0 9.3 35/74 1.3
0.50 0.01 160.2. 14.1 Sse LS
0.54 0.02 164.5 9.8 43.8 2.4
0.29 0.05 122.3) 30.4 1.1
(7.7%), and Common Grackle (1.1%); low-to-moder-
ate for Horned Lark (15.6%), Clay-colored Sparrow
(11.7%), Lark Bunting (19.4%), Grasshopper Sparrow
(19.7%), Bobolink (19.2%), and Western Meadowlark
(16.1%); moderate for Brewer's Blackbird (33.3%);
and moderate-to-heavy for Red-winged Blackbird
(43.1%). The Red-winged Blackbird was the most
commonly parasitized species in the two studies (71.2%
of all cases of parasitism), and we classified it as a
preferred host in subsequent analyses.
Parasitized passerine nests contained from zero to
six host eggs and/or nestlings and from one to eight
cowbird eggs and/or nestlings (Table 3). Of the 476
parasitized passerine nests, 50.6% contained one cow-
bird egg, 28.6% contained two, 12.0% contained three,
and 8.8% contained four or more cowbird eggs. The
maximum number of eggs or nestlings in a parasitized
nest (host and cowbird combined) was nine. The aver-
age number of cowbird eggs per parasitized nest was
1.9 + 1.2 (Tables 2 and 3). Multiple parasitism occurred
at 12.4% of all passerine nests and at 49.4% of all par-
asitized passerine nests. Parasitized nests of 18 passer-
ine species contained multiple cowbird eggs and/or
nestlings (Table 2). Among parasitized species with
25 or more nests, multiple parasitism rates were low
for Eastern Kingbird (0%), Brown Thrasher (0%), and
Common Grackle (0%); moderate-to-heavy for Clay-
colored Sparrow (43.5%), Lark Bunting (38.9%), Grass-
hopper Sparrow (44.4%), Chestnut-collared Long-
spur (50.0%), and Red-winged Blackbird (49.6%):
and heavy for Horned Lark (80.0%), Savannah Spar-
row (62.5%), Bobolink (53.3%), Western Meadowlark
(100%), and Brewer’s Blackbird (58.3%). Among these
species, the average number of cowbird eggs per par-
asitized nest was highest for the Western Meadowlark
(3.4 + 1.6); all nine parasitized nests of the Western
Meadowlark contained two or more cowbird eggs.
Vol. 121
THE CANADIAN FIELD-NATURALIST
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246
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE 3. Contents of parasitized passerine nests found incidentally during breeding bird surveys in two studies in the northern
Great Plains, 1990—2006.
Number of
host eggs or
nestlings per nest J 2 3
0) 37 2) 8
J 40 23 9
2 50 4| 14
3 58 37 18
4 45 13 zy
5 10 | 5
6 |
Total number
of nests 24\ 136 57
Number of Brown-headed Cowbird eggs or nestlings per nest
Total number
of nests
1 1 77
83
114
4 5 6 7 8
5
zy
2
|
SW BON WwW
NN We
22 10 8 ] 1 476
TABLE 4. Yearly variation of Brown-headed Cowbird parasitism in passerine nests (all species combined) found incidentally
during breeding bird surveys in two studies in the northern Great Plains, 1990-2006.
Frequency
Total number of cowbird
Year of nests parasitism (%)
1990, 57 29.8
199] 52 15.4
1992! 18] 12.7
1993' 90 20.0
1994 4] ID
1995 32 18.8
1996 20 0)
1997 42 16.7
199% 96 24.0
1999 96 25.0
2000 75 25.3
2001 129 Sbeyei
2002 184 34,8
2003 189 24.9
2004 234 29.9
2005 231 23,8
2006 153 26.8
Mean + SE UMA ete Uhre PLUG) a0) Pel
Frequency
of multiple
parasitism (%)
Distribution of
cowbird eggs in
parasitized nests>
Parasitism
intensity
(mean + SD)
5B) 1.6+0.9 random
50.0 1.8+0.9 random
PJP M phe \\ 5) random
55.6 Dd w= MN L6) random
62.5 1.9+0.8 random
Bos lgs) (05) random
0 ~ -
42.9 16+0.8 random
60.9 2.0 + 1.4 random
62.5 24+1.6 random
57.9 EO FENRZ random
34.8 1.5+£1.0 random
50.0 f yse ell random
46.8 LS + 1.2 random
Sif 1.8+0.8 random
36.4 17+ 1.4 non-random
56.1 19+ 1.2 random
46.9 + 3.8
"Mean number of cowbird eggs or nestlings per parasitized nest.
"Random distribution of cowbird eggs in parasitized nests was evaluated using a zero-truncated Poisson distribution model
(Lindsey 1997), Random distribution of cowbirds eggs indicated that female cowbirds did not discriminate among parasitized
hests and that the proportion of nests with 1, 2, 3,4, .../ cowbird eggs approximated a Poisson distribution, Non-random
distibution indicated that the random distribution model was rejected (P< 0.01).
"Data for the North Dakota statewide study and the CRP study were combined for 1992 and 1993,
Not including nests of the Red-winged Blackbird,
our sample of passerine nests from the two studies in
cluded mostly nests of grassland-nesting (e.g, West
ern Meadowlark) and edge-nesting (¢.¢., Yellow War
bler) species and very few nests of cavity-nesting (c.2.,
House Wren) or woodland-interior (¢.g., Ovenbird)
species (Table 2), We found 482 nests of 15 passerine
species that typically nest in grasslands, Overall, the
parasitism rate Of grassland bird nests was low-to
moderate (16.4%), We found 602 nests of 23 passerine
species that typically nest in habitats characterized by
woody vegetation (Le,, woodland, shrubland, open
habitat with scattered trees or shrubs, open or semi
open deciduous woodland, and edge), The parasitism
rate of bird species associated with woody vegetation
was low (9.5%).
Of the 45 passerine species for which nests were
found, nine species are known to eject cowbird eggs
(Table 2; Peer and Sealy 2004), Of the 257 nests of
these nine ejector species that we found, t2 (4.7%)
contained cowbird eggs. The overall parasitism rate tor
nests of all non-ejector (.c,, acceptor) passerine species
combined was 28.2% (1 = 1645), In addition, Yellow
Warblers often bury the contents of parasitized nests
under a new nest floor (Clark and Robertson 198t;
Burgham and Picman 1989), Eight of 24 (44.4%) Yel
2007 IGL and JOHNSON; COWBIRDS IN THE NORTHERN GREAT PLAINS 247
TABLE 5. Weekly variation of Brown-headed Cowbird parasitism in passerine nests (all species combined) found incidentally
during breeding bird surveys in two studies in the northern Great Plains, 1990-2006
Frequency Frequency Parasitism Distribution of
Weekly Total number of cowbird of multiple intensity cowbird eggs in
period of nests parasitism (%) parasitism (%) (mean + SD)' parasitized nests’
April IV 5 0 0
May I 0 0 0
May II 6 0 0
May III 1] 9.1 100.0 4.0 + 0.0
May IV 9] 11.0 60.0 2.6+2.0 random
June | 274 19.7 50.0 J Vee Wis} random
June II 4020 S53 Sill See non-random
June Il 421 27.8 53.0 19+ 1.1 random
June [V 454 20.9 41.1 1.6+0.9 non-random
July I 162 21.0 50.0 2.0 + 1.4 random
July U 39 23.1 33.3 1.4+0.7 random
July 1 37 16.2 50.0 2,2 E16 random
Mean + SE 158.5 +51.9 20.7 + 2.8° 54.3 + 6.2° - -
“To evaluate changes in cowbird parasitism within the breeding season, each month was divided into four, roughly weekly,
periods (week I = 7 days, week II = 8 days, week III = 7[8] days, and week IV = 8 days) (sensu Orians et al. 1989).
See Table 4 for explanation.
°Mean includes third week in May to third week in July (i.e., weeks with cowbird parasitism).
low Warbler nests each contained a single cowbird
egg. During the CRP study, buried (but visible) cow-
bird eggs were observed in two Yellow Warbler nests
(one in Kidder County, North Dakota, on 27 June 2003
and one in Eddy County, North Dakota, on 25 June
2005) and one Clay-colored Sparrow nest (McPher-
son County, South Dakota, on 13 June 2006).
Parasitized nests of passerines were found in 16 of
the 17 years (Table 4). No parasitized nests were found
in 1996, which also is the year that we found the fewest
number of active passerine nests. Excluding 1996, an-
nual parasitism rates ranged from 12.7 to 35.7% and
averaged 22.6% (Table 4). The average annual rate of
multiple parasitism was 46.9%. The zero-truncated
Poisson distribution model for parasitized nests was
rejected in one (i.e., 2005) of the 16 years in which par-
asitized nests were found.
In both studies combined, passerine nests were
found in 11 of 12 weekly periods from the last week of
April through the third week of July (Table 5). Eleven
passerine nests were found in two of the three weeks
before the first cowbird egg was found (i.e., before the
third week in May), although cowbirds were recorded
in each of those three weeks (Igl and Johnson, unpub-
lished data). In the North Dakota statewide study, we
found active nests between 27 April and 20 July, par-
asitized nests between 20 May and 18 July, and mul-
tiply parasitized nests between 20 May and 18 July.
In the CRP grassland study, we found active nests be-
tween 22 May and 9 July, parasitized nests between
29 May and 9 July, and multiply parasitized nests be-
tween 29 May and 9 July. During both studies, non-
parasitized and parasitized nests were found largely
between late May and early July, which corresponds
with the peak breeding periods of most passerine
species in the northern Great Plains, including the
Brown-headed Cowbird (Stewart 1975; Ig] and Johnson
1997; Tallman et al. 2002). After the third week in May,
weekly parasitism rates ranged from 9.1 to 37.3% and
averaged 20.7%; the highest parasitism rates were
found in the second and third weeks of June (Table 5).
The average weekly rate of multiple parasitism was
54.3%. The zero-truncated Poisson distribution model
for parasitized nests was rejected in two (i.e., second
and fourth week of June) of eight weekly periods.
In the CRP grassland study, parasitized and multi-
ply parasitized nests of passerines were found in all nine
counties (Table 6). Cowbird parasitism of passerine
nests was low in Fallon County in southeastern Mon-
tana (5.8%) and Grant County in western Minnesota
(9.3%). Parasitism rates were low-to-moderate in Butte
County in western South Dakota (12.8%) and Sheridan
County in northeastern Montana (11.6%), and moder-
ate in Hettinger (29.9%), Kidder (25.3%), and Eddy
(23.8%) counties in North Dakota and Day County in
northeastern South Dakota (28.9%). Heavy parasitism
was found in McPherson County in north-central South
Dakota (51.5%). The frequency of multiple parasitism
was moderate in Butte (21.4%) and Day counties in
South Dakota (39.8%) and Fallon County, Montana
(25%), and moderate-to-heavy to heavy in the other
six counties (range: 42.9-58.7%). The zero-truncated
Poisson distribution model for parasitized nests was
rejected in one (i.e., Day County, South Dakota) of the
nine counties (Table 6). Nests within the counties of
highest female cowbird abundance experienced high-
er frequencies of cowbird parasitism (R* = 0.64,
F = 12.65, P = 0.0093) than those found in counties
with the lowest female cowbird abundance (Figure 3).
The quadratic term for cowbird densities was non-
significant (P = 0.62) and was not included in the final
model.
248
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE 6. Regional variation in Brown-headed Cowbird parasitism in passerine nests (all species combined) found incidentally during
breeding bird surveys in Conservation Reserve Program grassland fields in nine counties in the northern Great Plains, 1990-2006.
Frequency
of cowbird
Total number
County of nests parasitism (%)
Butte, South Dakota 109 12.8
Day, South Dakota BD 28.9
Eddy, North Dakota We 23.8
Fallon, Montana 69 5.8
Grant, Minnesota 151 9.3
Hettinger, North Dakota 211 29.9
Kidder, North Dakota 332 25.3
McPherson, South Dakota 233) 51.5
Sheridan, Montana 69 11.6
“See Table 4 for explanation.
In the CRP grassland study, cowbird densities, total
breeding bird densities, and total breeding bird rich-
ness were lowest in the southwestern counties (Butte
County, South Dakota, and Fallon County, Montana)
and highest in counties near the center of the cowbird’s
breeding bird range. There was a positive relationship
between female cowbird densities and all measures of
avian abundance and richness. That is, female cowbird
densities increased with increasing density and rich-
ness at all levels of the avian community (Table 7).
Breeding bird density for all bird species (R* = 0.72)
was a better predictor of cowbird abundance than the
other four levels of avian densities (Table 7, Figure 4a).
Breeding bird richness for all bird species (R* = 0.69)
was a better predictor of cowbird abundance than the
other three levels of avian richness (Table 7, Figure 4b).
There was only a weak suggestion that cowbirds use,
as a settlement cue, the number of passerines species
known to have raised cowbird young (P = 0.115).
During breeding bird surveys in both studies, ob-
servers noted 586 instances of adult passerines provi-
sioning food to individual fledglings, of which only
26 (4.4%) instances involved hosts feeding cowbird
fledglings. These 26 observations occurred between
4 June and 7 July and involved eight host species: Clay-
colored Sparrow (5 cases), Grasshopper Sparrow (1
case), Le Conte’s Sparrow (1 case), Savannah Sparrow
(6 cases), Song Sparrow (4 cases), Brewer’s Blackbird
(2 cases), Red-winged Blackbird (5 cases), and Orchard
Oriole (2 cases). These eight species accounted for
83.6% of the total cowbird parasitism in these two stud-
ies (Table 2). Although we cannot establish conclu-
sively the true foster-parentage of these fledgling cow-
birds, we did not observe any non-parasitized species
(i.€., species whose nests were not parasitized during
the two studies) feeding cowbird fledglings during the
two studies.
Discussion
Brown-headed Cowbirds parasitize nests of a wide
variety of host species (Friedmann 1963; Lowther
1993), including some inappropriate hosts. As in other
Distribution of
cowbird eggs in
parasitized nests*
Parasitism
intensity
(mean + SD)
Frequency
of multiple
parasitism (%)
21.4 1.2+0.4 random
39.8 ilove IL non-random
46.3 1.8+ 1.1 random
25.0 1.3+0.5 random
42.9 1.5+0.7 random
58.7 Dp), = M5) random
45.2 1.7+0.9 random
59.2 AAdes IIail random
50.0 Allee i} random
studies (Ortega 1998), we found almost no cowbird
parasitism of nonpasserine nests. Most nonpasserines
are considered unsuitable hosts, and their nests usu-
ally are avoided by Brown-headed Cowbirds (Roth-
stein and Robinson 1998). Although the only parasi-
tized nonpasserine, an Upland Sandpiper, was clearly
an unsuitable host (1.e., a large-bodied host with pre-
cocial, nidifugous young), several other studies have
documented cowbird eggs in Upland Sandpiper nests
(Higgins and Kirsch 1975; Faanes and Lingle 1995*;
Davis and Duncan 1999).
Our overall frequency of cowbird parasitism (25%)
for passerine species was within the range that Ortega
(1998) considered moderate (20-40%). This overall
rate seems low given that cowbird densities are highest
in the northern Great Plains (Sauer et al. 2005*) and
that cowbirds are more abundant than many of their
common hosts in this region (Ig] and Johnson 1997;
Ig] et al. 1999). However, our combined parasitism rate
is based on nests from two studies, an extensive region,
17 years, and numerous passerine species, including
several species that ostensibly have developed behav-
ioral defenses against cowbird parasitism (see below).
After we excluded nests of ejector species, our over-
all parasitism rate remained moderate (28.2%). Some
studies in this region have found higher combined
rates of cowbird parasitism than those reported here-
in. For example, Koford et al. (2000) found an over-
all parasitism rate of 40.3% for passerines nesting in
seeded and native grasslands and cropland in North
Dakota and Minnesota, and Davis and Sealy (2000)
found an overall rate of 32.6% for species nesting in
grasslands in southwestern Manitoba. In both of these
studies, some individual species experienced moder-
ate-to-heavy levels of cowbird parasitism (e.g., >40%
parasitism for Western Meadowlarks in both studies).
Robinson and Smith (2000) suggested that some of
these commonly cited reports of high parasitism rates
from the northern Great Plains might not be represen-
tative of this region. Indeed, some recent reviews of
geographic patterns of cowbird parasitism (e.g., Peer et
al. 2000; Chace et al. 2005) focused on the extreme
2007
>
oO
y = 4.2169x + 3.1958, R? = 0.6437
ho ta abe Ch
Cee it eo
Parasitism rate (%)
DR inidise oii ode Bion iarn8 o 9
Cowbird Densities (99 / 100 ha)
FIGURE 3. Relationship between female cowbird densities and
the percentage of the total number of passerine nests
that were parasitized in Conservation Reserve Program
grasslands in North Dakota, South Dakota, Minnesota,
and Montana, 1990-2006. The data from the North
Dakota (N) statewide study were included for compar-
ison (1992 and 1993 were averaged).
cases of parasitism in the northern Great Plains (e.g.,
Linz and Bolin 1982; Davis and Sealy 2000; Koford et
al. 2000). There are many less-cited studies of cowbird
parasitism (see Shaffer et al. 2003) with low-to-mod-
erate rates of cowbird parasitism in this region (e.g.,
Hill and Sealy 1994; Granfors et al. 2001; Davis 2003:
Woolfenden et al. 2004). For example, in Manitoba,
Davis (2003) found an overall parasitism rate of 19.9%
for six grassland species in southern Saskatchewan over
a 5-year period.
Although our overall rate of cowbird parasitism
was 25%, we did find higher rates of parasitism in
some years (e.g., 35.7% in 2001), weeks (e.g., 37.3%
in the second week of June), counties (e.g., 51.5% in
McPherson County, South Dakota), and individual
species (e.g., 43.1% for the Red-winged Blackbird).
The results from our studies can serve as a reminder
that cowbird parasitism rates are not constant across
space or time (i.e., among years or within a breeding
season) and that cowbirds differentially parasitize hosts
within the same habitats. Differences in parasitism lev-
els among studies in the northern Great Plains may
simply reflect variation among species, regions, habi-
tats, or years. For example, Davis and Sealy’s (2000)
overall parasitism rate of 32.6% was based on data col-
lected at three study sites over two years; parasitism
rates were low-to-moderate at two of the study sites (18
and 20%) and heavy at the third study site (67%). We
echo Hahn and Hatfield’s (1995) caveat that patterns
of cowbird parasitism are complex and variable, and
that field studies of cowbird parasitism would be more
effective if they were long-term, community-wide stud-
ies set in strategically chosen communities in different
regions.
Our nest data were slightly unorthodox compared
to nest data from many other studies. We located all of
IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS
249
en
v—
Oo
Oo
ot
Ba
z
®
(=)
a] With Grant: y = 0.0616x - 5.0673, R? = 0.715
ri Without Grant: y = 0.0707x - 6.0706, R? = 0.908
=
°o
° 100 120 140 160 180 200 22
Breeding Bird Densities (pairs / 100 ha)
= 10
f=
oOo
aug
= 6
a=
y = 0.2357x - 4.6603, R? = 0.687
oO
Cowbird Densities (4
ho
ine)
or
30 35 40 45 50 55
Species Richness (al! species)
FiGuRE 4. Relationship between (a) mean female cowbird
densities and average densities of all other breeding
bird species, and (b) mean female cowbird densities
and mean species richness within the breeding bird
community (excluding cowbirds) in Conservation
Reserve Program (CRP) grasslands in North Dakota,
South Dakota, Minnesota, and Montana, 1990-2006.
The circle in Figure 4a indicates a potential outlier,
Grant County in Minnesota, which is discussed in the
text.
our nests fortuitously and observed each nest only once.
Any cowbird eggs ejected before our surveys, any nests
abandoned before our surveys, any cowbird eggs buried
completely in the nest lining, and any cowbird eggs
added or removed after our brief observations were not
recorded in our studies, and thus our frequencies of
cowbird parasitism should be considered minimal esti-
mates. These limitations, however, are not unique to
our data but rather are common features of all studies
of cowbird parasitism that do not involve cameras,
because most nests are not monitored continuously
throughout the nesting cycle but rather are visited at
relatively fixed intervals (e.g., every 24 days) for rela-
tively brief periods of time (e.g., less than a few min-
utes). Even systematic nest searches are neither inces-
sant nor comprehensive, but rather occur at fixed
intervals (e.g., Koford et al. 2000; Davis 2003; Winter
250 THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE 7. Five levels of avian abundance (density) and richness (number of species) that female Brown-headed Cowbirds could
use as proximate cues for settlement in Conservation Reserve Program grassland fields in nine counties in the northern Great
Plains, 1990-2006. Linear regression analyses were performed to examine the relationship between female cowbird densities
and each different subset of avian abundance and richness at the five levels of the avian community (also see Figure 4).
Level of avian community* Breeding bird density Number of breeding species
R? F P R? F IP
All breeding birds 0.72 17.53 0.004 0.69 15.56 0.006
All passerines 0.63 12.02 0.011 0.40 4.65 0.068
All passerines that have raised cowbirds 0.66 13.64 0.008 0.32 325) 0.115
All passerines that are non-ejectors 0.62 11.39 0.012 0.38 4.32 0.076
Preferred host (Red-winged Blackbird) 0.54 8.35 0.023 - = =
4 Excluding cowbirds.
et al. 2004). In addition, many studies that use sys-
tematic nest searches also include a fortuitous com-
ponent to their nest searching methodology (e.g.,
Koford et al. 2000; Davis 2003). Thus, nearly all esti-
mates of cowbird parasitism in the literature could be
considered minimal estimates.
In our two studies, female cowbirds did not avoid
laying eggs in previously parasitized nests. Nearly one-
half of the parasitized nests in our two studies con-
tained more than one cowbird egg, with the average
parasitized nest containing about two cowbird eggs.
The assumption that female cowbirds laid eggs ran-
domly in parasitized nests could only be rejected in
two of eight weekly periods, one of 16 years, and one
of nine counties in the CRP grassland study. Deviations
from a random Poisson distribution were likely relat-
ed to a greater number of nests with multiple cowbird
eggs or nestlings than would be expected by chance.
Multiple parasitism has been widely documented in
nests parasitized by the Brown-headed Cowbird (Mc-
Laren et al. 2003). Although multiple parasitism can
vary by region or host species, it typically occurs in
about one-third of all reported cases of cowbird para-
sitism (Friedmann 1963; Lowther 1993; Johnsgard
1997; Ortega 1998) and seems to be more common in
nests of larger hosts (Lorenzana and Sealy 1999; Trine
2000). Little is known about the factors that might
influence multiple parasitism, but some mechanisms
have been proposed or investigated. For example, mul-
tiple parasitism might occur if (1) a female cowbird
exhibits host preference (Smith and Myers-Smith
1998), (2) there is a high ratio of female cowbirds to
host nests (McGeen 1972), (3) there is a shortage of
alternative host nests (Smith and Myers-Smith 1998),
or (4) an inexperienced and nonselective female makes
a poor host choice (McLaren et al. 2003). Cowbird
hosts in the Great Plains seem to be particularly vul-
nerable to multiple parasitism (e.g., Elliott 1978; Davis
and Sealy 2000; this study), which may reflect higher
densities of cowbirds (Elliott 1977; Ortega 1998) or
lower densities of available or suitable hosts (Robin-
son and Smith 2000).
We likely underestimated the frequency of cowbird
parasitism of those species that eject or bury cowbird
eggs or that abandon parasitized nests (Ortega 1998).
Less than 5% of the nests of ejector species in the two
studies contained cowbird eggs, which was much lower
than the overall parasitism rate (28.2%) for acceptor
species. Ejection of cowbird eggs by a host is an effec-
tive anti-parasite strategy among some North America
passerines (Ortega 1998; Peer and Sealy 2004), espe-
cially among edge-nesting passerines that are common
in the northern Great Plains (Ig] and Johnson 1997;
Table 1). Even within acceptor species, however, some
individuals are prone to abandon parasitized nests and
then re-nest (Lorenzana and Sealy 1999). Hosoi and
Rothstein (2000) showed that the frequency of deser-
tion of parasitized nests was higher in non-forest than
forest species and suggested that increased nest deser-
tion was an evolved response to cowbird parasitism.
Among small-sized hosts, nest desertion seems to be
a common response to parasitism or defense against
parasitism (Ortega 1998). Many species that breed in
the northern Great Plains are known to abandon their
nests occasionally as a result of parasitism or cowbird
removal of host eggs (e.g., Ortega 1991; Hosoi and
Rothstein 2000), including some grassland and shrub-
grassland species, although the frequency of abandon-
ment may be quite low (Sealy 1999; Davis and Sealy
2000).
The Red-winged Blackbird was the third most
abundant species observed in both of our studies.
Despite its aggressive nest-defense behaviors against
cowbirds and its colonial-nesting tendencies (Free-
man et al. 1990), the Red-winged Blackbird was the
most commonly parasitized species in the two studies
(43.1% parasitism, 49.6% multiple parasitism, 71.2%
of all cases of parasitism) and appears to be a pre-
ferred cowbird host in the northern Great Plains. Our
data support Hanka’s (1979) contention that cowbirds
may show a phylogenetic preference for Red-winged
Blackbirds and other Icterine species (e.g., Brewer’s
Blackbird, Orchard Oriole; Table 2) in the cowbird’s
range. Linz and Bolin (1982) and Koford et al.
(2000) reported similar parasitism rates (42 and 43%,
respectively) for Red-winged Blackbirds in_ this
region. In the eastern portion of the cowbird’s range,
Red-winged Blackbirds are rarely parasitized (e.g.,
2007
Hahn and Hatfield 1995). Although we categorized
the Red-winged Blackbird as a wetland species
(Table 2; Igl and Johnson 1997), many of the red-
wing nests that we found during the CRP grassland
study were scattered in upland habitats (grassland,
shrub-grassland), where group defense is generally
lower (Robertson and Norman 1977; Ortega 1991)
and parasitism rates are generally higher (Robertson
and Norman 1976; Krapu 1978). Several host species
are known to avoid or reduce parasitism by acting
aggressively toward cowbirds (Robertson and Nor-
man 1976, 1977), but aggression is not always an
effective host defense against cowbird parasitism, espe-
cially in habitats in which host densities are low be-
cause cowbirds may use host aggression (Robertson
and Norman 1977) or vocalizations (Clotfelter 1998)
to locate nests. Nonetheless, the Red-winged Blackbird
is one of a few species reported to reduce parasitism
by mobbing (Freeman et al. 1990; Chace et al. 2005).
The presence of Red-winged Blackbirds may be a
double-edged sword for other passerines nesting in the
same habitats, such as CRP grasslands. Red-winged
Blackbirds are conspicuous breeding birds in CRP
grasslands and thus might increase parasitism in nests
of other species. Barber and Martin (1997) found that
a conspicuous, co-occurring species in a breeding bird
community can increase the risk of cowbird parasitism
on alternative host species. Alternatively, other species
may benefit from nesting near Red-winged Blackbirds
if (1) the defensive behavior of blackbirds toward cow-
birds reduces cowbird activity in the area, or (2) higher
cowbird parasitism of blackbird nests reduces para-
sitism pressure on other species. Fretwell (1972)
noted higher rates of cowbird parasitism in Dickcissel
nests that were built near Red-winged Blackbird nests,
whereas Clark and Robertson (1979) found the oppo-
site for Yellow Warblers. Woolfenden et al. (2004)
speculated that Red-winged Blackbirds might have in-
creased the risk of parasitism on Yellow Warblers in
their Manitoba study sites. The influence of Red-winged
Blackbird presence and abundance on cowbird para-
sitism rates of other species requires further study.
Mayfield (1965b) suggested that grassland birds
may have evolved defenses against brood parasitism
because of their long evolutionary history of co-
occurrence with the Brown-headed Cowbird in the
Great Plains. To date, however, the species that have
shown the strongest anti-parasite behaviors (i.e., egg
ejection) have not been the grassland species but rather
have been the species associated with woody vegetation
(Peer and Sealy 2004, Table 2). Our overall rate of cow-
bird parasitism for grassland bird nests (16.4%) was
much lower than those reported in some studies (e.g.
Elliott 1978 [57.7%]; Davis and Sealy 2000 [32.6%];
Jensen and Finck 2004 [42.9%]) but comparable to or
higher than those reported in other studies (e.g., Straus-
berger and Ashley 1997 [0%]; Kershner and Bollinger
1998 [1.7%]; Robinson et al. 2000 [7.8%]; Granfors
et al. 2001 [14%]; Davis 2003 [19.9%]; Winter et al.
IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS 251
2004 [6.7%]). Nonetheless, we found a higher fre-
quency of parasitism in nests of grassland species than
in nests of species that nest in edge and woodland habi-
tats (9.5%). In contrast to our results, in Illinois, Rob-
inson et al. (1999, 2000) and Strausberger and Ashley
(1997) found a higher frequency of parasitism in spe-
cies nesting in edge and woodland habitats than in nests
of grassland species. Although the difference between
the two regions for grassland birds is consistent with
the pattern that cowbird parasitism declines with dis-
tance from the center of the cowbird’s range in the
northern Great Plains, the difference for species asso-
ciated with woody vegetation is contrary to this gen-
eralization. Again, these patterns emphasize the com-
plexity and variability of cowbird parasitism in North
America (Hahn and Hatfield 1995). Robinson and
Smith (2000) and others (Peer et al. 2000; Jensen and
Finck 2004) have suggested that the higher levels of
parasitism in grasslands in the Great Plains than those
in the Midwest might reflect the near absence of large
forested areas, where more tolerant hosts can absorb
cowbirds and cowbird parasitism from grasslands.
Seemingly, our data support this contention. In our
most-forested county in the CRP grassland study (Grant
County in west-central Minnesota; Ig] and Johnson,
unpublished data), the density of female cowbirds (and
cowbird parasitism) was much lower than would be
expected given the density of breeding birds in the avian
community (Figure 4a). Moreover, Minnesota typical-
ly is lumped with midwestern states rather than with
Great Plains states. In the other eight counties, there
are few, if any, large forested areas that could absorb
cowbird parasitism from the open grasslands. Trees and
shrubs in these eight counties occur largely in small,
often linear, patches, where cowbird egg ejectors (e.g.,
kingbirds, thrashers, catbirds, robins) dominate the
breeding bird community. The low parasitism rates of
edge and woodland species in our study may reflect the
predominance of ejector species in woody habitats in
the northern Great Plains (Igl and Johnson 1997) and
the avoidance by cowbirds of a habitat that may con-
tain many less-tolerant hosts (Sealy and Bazin 1995).
In Illinois, Peer et al. (2000) recently found that
Western Meadowlarks rejected (i.e., ejected or dam-
aged) 78% of artificial and real cowbird eggs experi-
mentally added to nests. In contrast, in the northern
Great Plains, some Western Meadowlark populations
are moderately to heavily parasitized (23-67%: Davis
and Sealy 2000; 19-47%: Koford et al. 2000), and the
species often experiences extreme levels of multiple
parasitism in this region (Davis and Sealy 2000; Davis
2003; this study). It is unclear why some meadowlark
individuals or populations experience heavy or intense
parasitism (e.g., average of 3.4 and 3.1 cowbird eggs
per parasitized nest in this study and Davis and Sealy
2000, respectively), whereas others apparently expe-
rience little or none. Davis and Sealy (2000) suggest-
ed that the apparent geographic variation in para-
sitism levels in Western Meadowlark populations in
252
the northern Great Plains and elsewhere might reflect
differences in the size of suitable grassland patches
and the availability of other host species in the com-
munity. Peer et al. (2000) suggested that geographic
differences in parasitism levels in the northern Great
Plains and the Midwest might reflect geographic vari-
ation in egg rejection by Western Meadowlarks. Egg
rejection behavior in the Western Meadowlark should
be tested in more geographic locations or landscapes
to resolve these issues. Moreover, the Western Mead-
owlark and its congener, the Eastern Meadowlark
(Sturnella magna), use a similar tactic to destroy eggs
or clutches of other birds (Creighton and Porter 1974;
Schaef and Picman 1988; Picman 1992). The function
of this egg-destroying behavior has been speculative
(e.g., predation, interference competition for limited
resources), but in view of the similarities (e.g., egg
removal, puncture) between this egg-destroying behav-
ior at nests of other birds and cowbird egg ejection at
their own nests, experiments are needed to evaluate
the relationship between general egg destruction and
cowbird egg rejection by meadowlarks.
In our CRP grassland study, female cowbird den-
sities varied greatly among the nine counties in the
northern Great Plains, and rates of cowbird parasitism
were higher in the counties in which densities of fe-
male cowbirds were higher. Our results support the
interpretation of Miles and Buehler (2000) and Chace
et al. (2005) that female cowbird densities are a rea-
sonable predictor of the frequencies of cowbird para-
sitism. Our results also support the contention that cow-
bird parasitism declines with distance from the center
of the cowbird’s range, although on a smaller scale
(Hoover and Brittingham 1993; Smith and Myers-
Smith 1998; Thompson et al. 2000).
Few studies have evaluated the relationship be-
tween cowbird abundance and parasitism levels using
cowbird and nest data collected concurrently at the
same study sites over a large region (e.g., Robinson
et al. 2000; Jensen and Cully 2005a, 2005b). Large-
scale evaluations of this relationship typically have
relied on data from unconnected sources, such as abun-
dance data from the North American Breeding Bird
Survey (BBS) and nest data from the Cornell Labora-
tory of Ornithology or from multiple studies (e.g.,
Hoover and Brittingham 1993; Basili 1997; Smith
and Myers-Smith 1998; Herkert et al. 2003). Most
evaluations of this relationship have focused on single
species (Hoover and Brittingham 1993; Basili 1997;
Smith and Myers-Smith 1998; Herkert et al. 2003).
Nonetheless, many studies have found similar positive
relationships between cowbird densities and cowbird
parasitism levels. For example, Basili (1997) and
Herkert et al. (2003) showed that cowbird densities on
BBS routes were positively related to frequencies of
cowbird parasitism of Dickcissel nests from multiple
studies across several states. Jensen and Cully (2005a,b)
found a similar, positive relationship between female
cowbird densities and parasitism levels of Dickcissel
THE CANADIAN FIELD-NATURALIST
Vol. 121
nests at eight study sites in Kansas and Oklahoma. In
a meta-analysis of nest and cowbird data from six stud-
ies from four midwestern states, Thompson et al. (2000)
similarly found that cowbird abundance and para-
sitism levels of multiple species were positively cor-
related across study areas. In contrast, Woolfenden et
al. (2004) found that female cowbird abundance was
not related to parasitism frequency among three species
in an avian community in Manitoba, and Robinson et
al. (2000) found that the percentage of nests para-
sitized was not related to cowbird abundance in mul-
tiple forest sites across Illinois.
In CRP grasslands, we also found a positive rela-
tionship between cowbird abundance and all five lev-
els of the avian community, including all avian species,
all passerine species, all passerines that are non-ejec-
tors, all passerines known to raise cowbird young, and
a single, preferred host (i1.e., Red-winged Blackbird).
The abundance and richness of all avian species were
the strongest predictors of cowbird abundance, which
suggests that cowbirds might use the overall breeding
bird community as a settlement cue for optimal areas
to breed rather than more specific passerine or host
categories (e.g., all passerines that are known to have
raised cowbird young). The number of individuals and
the number of species represent different components
of a cowbird’s breeding resource. The presence of more
individuals and more species with different breeding
chronologies ensures a larger number of potential hosts
within the breeding bird community throughout the
cowbird’s breeding season (Farmer 1999). Although
we evaluated these relationships using only data from
the CRP grassland study, cowbirds also might use other
cues (e.g., vegetation or landscape factors) as an indi-
cator of the richness or abundance of the avian com-
munity (Chace et al. 2005).
Chace et al. (2005) tabulated 16 published studies
and one unpublished study that evaluated the relation-
ship between different measures of the avian commu-
nity and cowbird abundance or occurrence. As with
our results, many studies found positive relationships
between cowbirds and different measures of avian den-
sities and richness. To our knowledge, Farmer (1999) is
the only other published study to have evaluated mul-
tiple host and avian community measures using both
abundance and richness data. Farmer (1999) found
positive relationships for nearly every level of the avian
community, including abundance and richness cate-
gories. Because host species are necessary for cow-
birds to be reproductively successful, most studies have
evaluated these relationships at the host level. Results
of such evaluations have been inconsistent. Moreover,
comparisons among studies using host categories are
difficult because a variety of definitions of host com-
munities were used (Chace et al. 2005). For example,
Thompson et al. (2000) found that cowbird abundance
was significantly correlated with host abundance (i.e.,
hosts included species that bred on the study area and
accepted cowbird eggs), whereas Robinson et al. (2000)
2007
found no relationship between cowbird abundance
and host abundance (i.e., hosts included species that
accept cowbird eggs regularly but excluding cavity
nesters, cowbird egg ejectors, and large species). Evans
and Gates (1997) found that cowbird abundance was
positively related to the abundance of all avian species
combined, but the relationship was non-significant for
richness of all avian species and richness and abun-
dance of host species (1.e., known cowbird hosts).
In summary, our data show that, even within the
northern Great Plains, there is considerable variation in
parasitism rates among species and across space and
time. Our results demonstrate that parasitism rates over-
all are moderate in the northern Great Plains, although
some individual species, counties, years, and weeks ex-
perience heavier rates of parasitism or multiple para-
sitism than others. Our study is one of the first to show
a strong, positive relationship between regional cow-
bird abundance and nest parasitism rates using both
cowbird abundance and nest data collected at the same
study sites. We also found a clear, positive relation-
ship between female cowbird densities and the over-
all breeding bird communities in CRP grasslands in
the northern Great Plains. More studies are needed to
determine if these relationships hold elsewhere in the
northern Great Plains with different host communi-
ties or within other habitats.
Acknowledgments
We thank K. L. Andersson, I. Balodis, K. A. Dal-
ton, J. E. Doster, D. A. Hobbick, C. J. Johnson, K. F.
Kuehnl, J. M. Legge, R. L. Manson, J. W. Marlow,
M. C. Marlow, L. A. Murphy, K. L. Richardson, T. R.
Runia, M. D. Schwartz, C. M. Shoemaker, M. L.
Sondreal, J. M. Steiner, and K. A. Ward for their assis-
tance in the field. We are grateful to the numerous land
owners and operators who allowed us access to their
property. J. E. Austin, D. A. Buhl, A. J. Erskine, D. A.
Granfors, C. S. Houston, R. R. Koford, W. E. Newton,
P. J. Pietz, and S. G. Sealy provided helpful comments
on earlier versions of this paper.
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Received 27 August 2007
Accepted 12 March 2008
Wolf, Canis lupus, Behavior in Areas of Frequent Human Activity
ELLEN HEILHECKER!, RICHARD P. THIEL’, and WAYNE HALL, JR.?
University of Wisconsin-Stevens Point, College of Natural Resources, Stevens Point, Wisconsin 54481 USA
*Wisconsin Department of Natural Resources, Sandhill Wildlife Area, Box 156, Babcock, Wisconsin 54413 USA
Heilhecker, Ellen, Richard P. Thiel, and Wayne Hall, Jr. Wolf, Canis lupus, behavior in areas of frequent human activity.
Canadian Field-Naturalist 121(3): 256-260.
We report incidental observations of Wolves (Canis lupus) tolerating human activity in central Wisconsin. Three monitored
packs raised pups in close proximity to varying levels of human activity. Wolf pups were raised <350m from rearing pens of
the endangered Whooping Crane (Grus americana), which saw daily human activity. One pack used cornfields as ren-
dezvous sites within 175 m of a maintenance shed visited regularly by workers. Another pack centered their activities along a
well-traveled state highway using both the verge and the road center for activity. Aerial locations of 10 yearling and adult
dispersing Wolves were plotted to evalute human densities in natal territories relative to dispersal and post-dispersal territo-
ries. Township densities (¥ = 9.02 humans/km?, SE = 4.015) and residential densities (¥ = 5.59 housing units/km?, SE =2.12 )
in natal pack territories were significantly greater (P <.01) for dispersal and post-dispersal township densities (¥ = 43.98
humans/km?, SE =7.37) and residence densities (¥ = 23.12 housing units/km*, SE =3.49). Furthermore, a pup negotiated the
densely populated region of northern Illinois and dispersed from central Wisconsin to east-central Indiana, a distance of at
least 690 km. As Wolves live 1n closer proximity to humans, living in areas of higher township and residential densities, they
can be expected to be more habituated to people, increasing the probability of human/Wolf conflicts.
Key Words: Wolf, Canis lupus, pups, humans, habituation, behavior, Wisconsin.
Historically, Gray Wolves (Canis lupus) were one of
the most widely distributed land mammals in North
America inhabiting all regions except arid deserts
(Young 1944; Mech 1970; Mech and Boitani 2003).
However, human settlement and persecution restrict-
ed Wolves to northern forested regions and eventually
the only substantial population within the lower 48
states occurred in northern Minnesota (Mech 1970).
Under the protection of the federal Endangered
Species Act of 1973, Wolves have returned to some
areas within their historical range (Mech 1995). They
began recolonizing Wisconsin in the mid-1970s (Mech
and Nowak 1981; Thiel and Welch 1981; Wydeven et
al. 1995). Wolves have since repopulated much of north-
ern Wisconsin and an isolated area in the central part
of the state referred to as the Central Forest Region
(CFR). In 1999, the Wisconsin Department of Natural
Resources (WDNR) downlisted the Wolf from an
endangered to a threatened species (WDNR 1999*).
Wolves continued to surpass target recovery levels. In
April 2003, the U.S. Fish and Wildlife Service pro-
posed to delist the Eastern Distinct Population Seg-
ment of Gray Wolf, which includes Wisconsin (Fed-
eral Register 2003*).
Wolves initially recolonized areas in Wisconsin with
significantly lower road densities (y = 0.23 km/km/?)
and human densities (y = 1.52 people/ km*) (Mlade-
noff et al. 1995). As recovery progresses, Wolves are
demonstrating an increasing tolerance to human activ-
ity throughout the upper Great Lakes region, including
Wisconsin (Thiel et al. 1998). In recent years, Wolves
have denned near active sphagnum moss drying sites,
occupying homesites inside artillery impact zones on
military bases, crossed four lane highways and large
expanses of non-forested areas, and bypassed major
U.S. cities (Licht and Fritts 1994; Mech et al. 1995;
Thiel et al. 1998). While such behavior is not unique to |
the species, it is unique to Wolves living in the conti-
nental United States (McNay 2002; Mech and Boitani
2003). Observations of Wolves displaying increased
tolerance towards humans are of value insofar as it af-
fects management of Wolf recovery. This paper reports
on the success of Wolves attempting to colonize areas
of higher human densities than previously reported in
the continental United States.
Study Area and Methods
Wisconsin’s Central Forest Region (CFR) encom-
passes 7767 km? of deciduous forest (Acer saccharum,
Tilia americana, Quercus alba, Q. borealis, Q. veluti-
na), coniferous swamps (Picea mariana, Larix larici-
na), aspen stands (Populus tremuloides, P. grandiden-
tata), pine barrens (Pinus banksiana, P. resinosa, P.
strobus), sphagnum bogs, and marshes (Curtis 1959;
Finley 1976*). A 12 141-hectare swamp, the largest in
Wisconsin, lies within the CFR (Martin 1965). The»)
CER includes portions of Eau Claire, Chippewa, Clark, ||
Jackson, Wood, Monroe, Juneau, Adams, and Mar-
quette counties. The majority of the CFR consists of |
private industrial, county, state and federal forestland.
Major economic industries include logging and cran-
berry agriculture as well as dairy operations (WDNR
2003*). Road density within the CFR is 1.23 km/km?
(U. S. Census Bureau 2000*; WDNR 2003%*).
Wolves were captured on or near rendezvous sites, |
located by howling and track surveys, using offset,
256
2007
modified #14 Newhouse traps during summers 2002
and 2003, following guidelines established by Ani-
mal Care and Use committees within the Wisconsin
Department of Natural Resources and University of
Wisconsin-Stevens Point. With the exception of setting
traps, researchers minimized time spent in pack ren-
dezvous sites to avoid Wolf-human interactions with
study animals. Wolves were sedated with a 5:1 mL
mixture of ketamine hydrochloride and xylazine and
were weighed, sexed, and examined to assess general
condition (Wydeven et al. 1995).
Radio collars were placed on adults. Pups received
ear tag transmitters developed for a Wolf pup mortal-
ity study (Advanced Telemetry Systems, Isanti, Min-
nesota) programmed to transmit from 0700 — 1900.
Pups and adults were located from the ground using a
receiver with either a 5-element hand held Yagi anten-
na or a 9-element vehicle mounted antenna once daily
from time of capture to 15 January each study sea-
son. All Wolves were located from fixed-wing air-
craft 1-2 times/week.
Human activity observed near pack homesites was
documented. Additionally, we interviewed people and
collected reports from people who encountered study
pack Wolves to assess the relative level of tolerant be-
havior of study packs towards humans.
Aerial locations of 10 dispersing Wolves were plot-
ted in Universal Transverse Mercator (UTM) coordi-
nates utilizing ArcView 3.X (ESRI, Inc., Redlands,
California). Township (human) and residential (struc-
tural) densities for each county in which Wolf loca-
tions were plotted were queried (U. S. Census Bureau
2005*). Measurements obtained included township
densities and residential densities in the county for
(1) natal territories from which Wolves dispersed, (2)
locations while Wolves were dispersing, and (3) after
dispersers settled and either became territorial or be-
came breeders.
Results
We monitored 17 Wolves from three study packs
in 2002 and nine Wolves in five study packs in 2003.
Seven Wolves and three packs were new in 2003. In
three of the six study packs we were able to detail
repeated interactions with humans.
Suk Cerney pack: Three male pups, one female pup,
a yearling female, and an adult male Wolf were radioed
near their den site between 3-6 July 2002 on Necedah
National Wildlife Refuge (NNWR) in Juneau County
(44°04.957'N, 90°10.399'W). The den site was locat-
ed approximately 350 m from an observation bunker
and several rearing pens used as part of a Whooping
Crane (Grus americana) reintroduction program locat-
ed on the NNWR. Between July and October employ-
ees and volunteers of the crane project, NNWR staff.
along with visiting news media and guests, visited the
cranes at least twice per day. Crane staff and Wolf re-
searchers routinely observed pups before and during
HEILHECKER, THIEL, and HALL: WOLF BEHAVIOR
tN
A
~~
. LY.
from inet
Of
f Wisconsin YA Ai \
Uy
. fee J
\
( Madison Q
j
f
y |Miwauked
ey ) |} Michigan ya
1, Chicago / (
jf -
oe SS _/ Gary i ew
) |
|
, Minis | * Jet
Ne } Indiana |
“be ’ ~ 2 é
t 4 ¥
i; ) ma ARAN
( s we =)
\ a
x ae ra
bela
of ~
W E
FIGURE |. Capture site of Wolf 409 in central Wisconsin on
10 August 2002 and death site in eastcentral Indiana.
June 2003.
our study period. One pup watched Wolf researchers
from the den, located in oak savannah dominated by
Hill’s oak (Quercus ellipsoidalis) for approximately
10 minutes as traps were set on 3 July 2002, approxi-
mately 10.5 m away. Adult Wolves were rarely seen.
However, radio signals confirmed they were often in
the immediate vicinity. The 4 radioed pups stayed in
the area despite daily human activity, until their deaths
between July 15 and 5 September 2002 (Heilhecker
2003*).
Bear Bluff pack: Six Wolves were captured in and
adjacent to cornfields on private land in Jackson Coun-
ty between 20 July and 3 August 2002 (44°15.355'N,
90°20.582'W). Two male pups and two female pups, a
yearling female, and a yearling male were radio-tagged.
A partially planted, sandy soil cornfield with stunted
corn stalks < 0.9 m tall, used as a daily rendezvous site
from July to early November in 2002 and 2003, was
approximately 175 m from the maintenance building
used by employees of the cranberry farm at least twice
a week. Distance from the rendezvous site to the main-
tenance building and cranberry beds were approximate-
ly 450 m and 1300 m, respectively. White-tailed Deer
(Odocoileus virginianus), the Wolves’ primary prey in
the region, were abundant and cause $35000/year dam-
age to local commercial cranberry beds (K. Rice,
Flying Dollar Cranberry Company, personal commu-
nication). Deer flies (Chrysops sp.) and mosquitoes
(Culicidae) were generally less abundant in the corn-
258
fields compared to the surrounding forested land. In
both years, and for years prior to our study, employees
of the cranberry company regularly observed Wolves.
Once during our study they witnessed a Wolf in their
parking lot sniffing vehicle tires. The property man-
ager also reported his dog interacting with Wolves in
what looked like play behavior.
Noch Hanai pack: Wolf 429, a yearling female, was
radio-collared on 3 June 2002. In summer 2002, she
was observed multiple times acting indifferently to
vehicles along State Highway 54 on the northern border
of her natal pack territory (44°20.15'N, 90°35.51'W).
In the autumn she dispersed and created a new pack
that occupied a 102-km/? territory centered along the
same state highway. The pack’s den site was discov-
ered approximately 230 m from State Highway 54 ina
Red Pine (Pinus resinosa) plantation with no under-
story. Vehicles passing on the state highway were visi-
ble from the den. The wolves’ rendezvous sites, as
determined by matted grass, trails, human garbage with
teeth marks, and deer legs, were on embankments along
the shoulder of this moderately traveled state highway,
adjacent side roads, and recently logged forests span-
ning a distance of 7.7 km by 60 m along the state high-
way.
In summer 2003, over 60 citizen phone calls and e-
mails reported a radio-collared Wolf with pups walk-
ing, playing, and lying in the middle of the state high-
way. Initially four pups and a radio-collared adult were
observed. On 19 July 2003, a dead male pup was col-
lected from the shoulder of the highway. The necropsy
revealed injuries consistent with a vehicle collision.
None of the Wolves appeared to be afraid of vehi-
cles. On 20 July 2003, a citizen reported watching
three pups and a collared female in the middle of the
state highway. They moved off to the shoulder of the
road to let vehicles pass and then returned to the middle
of the road. E. Heilhecker and W. Hall, Jr., observed
Wolf 429 looking both directions before stepping onto
the highway. Pups were observed sitting on the side of
the road chewing on deer legs as vehicles passed. Pups
once walked next to Heilhecker and Hall’s vehicle
parked on the shoulder of the road. In all instances the
pups appeared oblivious to the presence of vehicles.
It was later learned that the Wolves had been fed by
loggers working in this area during the late spring/early
summer.
On 13 January 2004, WDNR received a report of an
uncollared Wolf on State Highway 54 jumping into
the bed of a truck containing deer hides and carcass-
es after its occupant had stopped to take pictures of
the Wolf (M. Windsor and T. Babros, WDNR, person-
al communication). On 29 March 2004, an uncollared
Wolf was observed walking back and forth across State
Highway 54 stopping traffic. At one point the Wolf
picked up a yellow plastic oil container, crossed the
road, and lay down within 3 m of a vehicle. Several
vehicles slowed down to watch the Wolf. While no one
exited their vehicles, people rolled down their win-
THE CANADIAN FIELD-NATURALIST
Vol. 121
dows and continued to talk within hearing range of
the Wolf. The Wolf did not appear to be affected by
the commotion. On 27 April 2004, the radio-collared
female, Wolf 429, was humanely dispatched along
Hwy 54 because she was unable to ambulate on her
hind legs. A preliminary necropsy was inconclusive,
but her paralysis was not caused by a vehicle collision.
This Wolf did have a healed injury to the pelvic girdle
that was caused by blunt trauma suggestive of an ear-
lier collision with a vehicle (A. Dassow, University of
Wisconsin, Madison, personal communication).
Dispersal: We compared aerial locations for 10
Wolves, as yearlings and adults, dispersing between
1998 and 2004. Seven Wolves (W002, W269, W309,
W338, W341, W426, W429) established new territo-
ries; five of which (W309, W338, W341, W426, W429)
were known to have successfully bred. Three Wolves
(W337, W427, W480) continued to disperse until
their deaths and never set up known territories. Town-
ship densities (x = 9.02 humans/km?, SE = 4.015) and
residential densities (y = 5.59 housing units/km?,
SE =2.12 ) in natal pack territories were significantly
lower (P < 0.01) than township densities (z = 43.98
humans/km?, SE = 7.37) and residential densities
(x = 23.12 housing units/km?, SE = 3.49) in dispersal
and post-dispersal territories (U. S. Census Bureau
2005*).
A male pup, W409, dispersed to eastern Indiana in
less than 156 days. The pup was last monitored on 15
January 2003 and found dead on 20 June 2003. If the
pup dispersed in a straight line, it would have passed
through relatively densely settled areas of northern
Illinois and the greater metropolitan Chicago area to
reach eastern Indiana (40°06.4'N, 85°04.3'W) (Figure
1). The human density of Jackson County, its birth-
place, was 7.8 people/km?. The least populated coun-
ties along the Wisconsin/Illinois state border had 22.9
people/km? (Green County, Wisconsin) and 14.5 peo-
ple/km? (Jo Daviess County, Illinois). The human den-
sity in Randolph County, Indiana, where the Wolf was
found shot was 22.84 people/km? (U. S. Census
Bureau 2005*). This was the first documented Wolf
in Indiana since their extirpation in 1908 (Mumford
and Whitaker 1982), and the southern-most movement
of a radio-collared Wisconsin Wolf to date (A. Wyde-
ven, WDNR, personal communication).
Discussion
Wolves’ reactions to humans are a reflection of their
experiences with people and the relative tolerance of
humans to Wolves (McNay 2002; Fritts et al. 2003).
By the mid-1900s Wolves were largely restricted to
wilderness areas in North America due to intense
human persecution (Young 1944; Mech 1995). With
legal protection from the Endangered Species Act of
1973, a gradual increase in acceptance of Wolves
(Fritts et al. 2003), and an expanding population,
Wolves have inevitably come in closer contact with
humans (Mech 1995; Thiel et al. 1998; Mech and
2007
Boitani 2003). By the 1990s an increasing number of
Wolves began dispersing into and colonizing more
settled regions and agricultural areas in the northern
United States (Licht and Fritts 1994; Mech 1995; Mech
and Boitani 2003). Thiel et al. (1998) and Merrill and
Mech (2000) reported tolerance of humans close to den
and rendezvous sites in Minnesota and Wisconsin.
As Wolves colonize more fragmented habitat dom-
inated by humans they will come into more frequent
contact with humans, as our findings demonstrate. Dis-
persing Wolves from the numerous packs presently
inhabiting the forest-farmland zones in Wisconsin and
Minnesota are capable of penetrating great distances
into landscapes highly dominated by humans (Licht
and Fritts 1994; Merrill and Mech 2000; Fritts et al.
2003; Mech and Boitani 2003).
The male Wolf pup, 409, provides an example of
this progression of tolerance to humans, a behavior
that can be expected to be exhibited by Wolves in
succeeding generations. Although movements of
= 670 km from their homesites have been well docu-
mented in Wolves (Van Camp and Gluckie 1979; Bal-
lard et al. 1983; Fritts 1983; Ream et al. 1991; Licht
and Fritts 1994; Merrill and Mech 2000; Mech and
Boitani 2003), this is the first documentation of a wild
North American Wolf moving through areas of such
high human densities. While Wolf 409 was not known
to have any contact with nor was known to have been
conditioned to humans as a pup, it was subjected to
increasing levels of human activity as it dispersed south
and east of Lake Michigan. Interestingly, Wolf 429, the
founder of what became the human-habituated Noch
Hanai pack, came from the very same pack in which
Wolf pup 409 was born.
We observed two forms of habituation of Wolves in
our study: (1) pups that became accustomed to the pres-
ence of humans through repeated, benign association,
and (2) association of humans with a food reward. In
the two cases where litters were raised in close proxim-
ity to human activity and were frequently seen, the pups
fled at the close presence or approach of a human(s).
These Wolves created no known threats to public safe-
ty or caused any damage to personal property. Wolves
displaying avoidance or fear behavior during direct
encounters with humans are considered to be showing
“normal” behavior (Smith and Stahler 2003*).
In the remaining case in our study, pups received
food awards from humans. Wolves are not known to
attack people during their first human encounter but
require repeated exposure to humans before attack-
ing (McNay 2002; Smith and Stahler 2003*). A pre-
requisite for aggressive Wolf behavior towards humans
is habituation, with food being the most influential
factor (McNay 2002; Smith and Stahler 2003*). Our
human-habituated Wolves’ whose bold behavior rap-
idly evolved into a public nuisance issue.
We expect more Wolf-human encounters and human-
habituated Wolves due to the recovering Wolf popula-
HEILHECKER, THIEL, and HALL: WOLF BEHAVIOR
259
tion in the Great Lakes area. Wisconsin's Wolf popu-
lation has increased from 83 Wolves in 1995 to 425
Wolves in 2005 (Wydeven and Wiedenhoeft 2005*)
The growing Wolf population and increased number
of Wolf sightings have raised safety concerns from
the general public. A continuous educational program
that focuses on how human actions affect Wolf behav-
ior may minimize the establishing of human-habituated
Wolves. Periodic news releases and public education
programs at the federal and state forests, posters at road
side kiosks emphasizing the reasons why feeding of
Wolves is improper, writing stipulation in public con-
tracts on public land prohibiting feeding of Wolves
especially in areas where Wolf habituation is occur-
ring, or fining individuals who feed Wolves may be
useful approaches. When habituation occurs, attempts
should be made to deter Wolves displaying bold behav-
ior with aversion techniques such as rubber bullets
and cracker shells before Wolf aggression develops.
If aversion conditioning is unsuccessful, for public
safety reasons, aggressively bold Wolves should be
humanely euthanized.
Acknowledgments
This study was funded by the Zoological Society of
Milwaukee County, National Fish and Wildlife Foun-
dation’s Budweiser Conservation Scholarship, Univer-
sity of Wisconsin-Stevens Point, and the WDNR. We
thank Angela Dassow (UW-Madison), Adrian Wyde-
ven (WDNR), Paul Anderson (WDNR), Peggy Calla-
han and the Wildlife Science Center, Kirby Rice and
the Flying Dollar Cranberry Company, Advanced
Telemetry Systems, Inc., Neal Paisley and the employ-
ees of Sandhill Wildlife Area (WDNR), Michele
Windsor (WDNR), Tim Babros (WDNR), Tim Beyer
(WDNR), the staff at the NNWR, the Bloomington
Field Office of the Indiana Department of Natural
Resources (IDNR), Lori Pruitt, Scott Pruitt, Larry Har-
ris (USFWS), Kent Hanauer (IDNR), and Jayne Bel-
sky (NNWR volunteer).
Documents Cited (marked * in text)
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Report | July-15 January 2003. University of Wisconsin
— Stevens Point. Stevens Point, Wisconsin.
Smith D. W., and D. E. Stahler. 2003. Management of
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United States Census Bureau. 2000. Wisconsin 2000 Roads:
2000 TIGER line files, Office of Land Information Serv-
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United States Census Bureau. 2005. State and county quick
facts. http://quickfacts.census.gov
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Received 11 April 2006
Accepted 5 June 2008
Influence of Gender and Den Type on Home Range Shape for Striped
Skunks, Mephitis mephitis, in Saskatchewan
SERGE LARIVIERE!, DAVID HOWERTER?, and FRANCOIS MESSIER®
'Delta Waterfowl Foundation, R.R. 1, Box 1, Site 1, Portage La Prairie, Manitoba RIN 3A1 Canada. Present address: Cree Hunters
and Trappers Income Security Board, 2700 boulevard Laurier, Champlain #1110, Québec (Québec) G1 V 4K5 Canada
“Institute for Wetland and Waterfowl Research, Ducks Unlimited Canada, P.O. Box 1160, Stonewall, Manitoba ROC 270 Canada
3 Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada
Lariviere, Serge, David Howerter, and Francois Messier. 2007. Influence of gender and den type on home range shape for
Striped Skunks, Mephitis mephitis, in Saskatchewan. Canadian Field-Naturalist 121(3): 261-264.
We investigated gender differences in shape of home ranges for Striped Skunks, Mephitis mephitis, in southcentral Saskatchewan,
Canada, during 1993-1994. Data collected on free-ranging Striped Skunks indicated that shape of female home ranges was
unaffected by den type (building versus burrow) and was similar to shape of male home ranges. Moreover, type of maternity
den did not influence the position of the den within the home range (inside or outside of core area). We conclude that the dis-
tribution of Striped Skunks in summer, and the distribution of their foraging activity, are not affected by the availability of
anthropogenic den structures.
Key Words: Striped Skunk, Mephitis mephitis, home range, denning site, spatial distribution, Canadian prairies.
Fragmentation of habitats affects not only how ani-
mals are distributed in the landscape, but also their use
of space. In fragmented landscapes, suitable habitats
for foraging may not occur in proximity to other criti-
cal habitats for denning, parturition, or rearing of
young. Therefore, mammalian females that live in frag-
mented landscapes face two problems: establishing a
home range that includes suitable foraging habitat and
finding a site for parturition and rearing of young.
For males that do not provide parental care, obtain-
ing food is a primary drive affecting behavior during
summer. In contrast, females give birth and raise a lit-
ter during the same period, and habitat preferences
for raising young may not coincide with good forag-
ing habitats. In fragmented landscapes, the compro-
mise between optimal foraging and optimal rearing
habitats may affect the establishment of home ranges,
and consequently their shape. Thus, the location of
maternity dens may have several implications for the
distribution of foraging activity of females.
First, den location may affect the shape of the home
range or the position of the den site within the home
range. For example, species that rely on a burrow and
central place for refuge and storage of food may benefit
from a centrally located den and circular home range
| (Bowers 1995). However, in fragmented landscapes,
females may have to prioritize either establishment of
home range (find a good foraging area first, then find a
suitable place to rear young within the home range), or
_ the establishment of maternity den (find a good den
_ first, then find a good place to forage).
We investigated the influence of gender and den
_ types on home range shape of Striped Skunks (Mephi-
_ tis mephitis) in the parklands of Saskatchewan, Cana-
ae!
da. Striped Skunks are distributed across most of Cana-
da and the United States (Rosatte and Lariviére 2003).
Females give birth in mid-May, and because males pro-
vide no parental care, the progeny are dependent on
the mother until mid-July, when the young disperse
(Lariviére and Messier 1997). In Saskatchewan, female
Striped Skunks use two main types of shelter for mater-
nity dens, either anthropogenic structures (e.g., build-
ings, ca. 60% of maternity dens) or underground burrows
(ca. 40% of maternity dens) (Lariviere and Messier
1998c). The distribution and availability of both den
types differ, and thus, we hypothesized that den type
could influence home range shape. For example, build-
ings may provide good denning structure, but their
position may not be in proximity to good foraging areas,
a scenario that would yield an elongated home range.
In contrast, a burrow may be dug presumably closer to
good foraging areas, and thus home range of females
using burrows could be more circular in shape. Thus,
Striped Skunks provide a good model species to test
predictions of den site positioning and home range
shape by animals occupying fragmented landscapes.
Study Area and Methods
We conducted this study in the Prairie Pothole Region
of southcentral Saskatchewan (52°45'N, 107°08'W).
Small grain (1.e., wheat, barley, oats) and oil crops
(mostly canola but also flax) occupy 60% of the land-
scape. Numerous wetlands and stands of Trembling
Aspen (Populus tremuloides) occur throughout the
area. Topography is gently rolling, and an extensive
network of roads divides the land. Areas managed for
nesting waterfowl are common throughout the area, and
consist mostly of dense nesting cover, a mixture of
native and tame grasses seeded specifically to attract
upland nesting waterfowl. General characteristics of
the Prairie Pothole Region are detailed elsewhere
(Greenwood et al. 1995).
261
262
From April to August, 1993-1994, Striped Skunks
were captured in opaque-sided livetraps (Larivieére and
Messier 1999) and anesthetized using halothane and
Telazol® (Lariviére and Messier 1996a). All individuals
were equipped with a 5-sec delay motion-sensitive
radio-collar (150-152 Mhz, Telonics Inc., Mesa, Ari-
zona, USA) to monitor activity during tracking (Lari-
viere and Messier 1997). Skunks were released at the
site of capture.
Radio-collared animals were located by an observer
using hand-held receiving equipment from 18:00 to
06:00, during the period of greatest activity (Lariviere
and Messier 1997). Individuals were located every 15
min by direct observation, auditive location, or short-
range (<50 m) triangulation (Lariviere and Messier
1998b). Light amplifying, night-vision goggles (AN-
PVS 5, Bill’s Electronics Ltd., Mildmay, Ontario)
enabled visual observations at night. Tracking effort
was systematically distributed among all skunks, and
no animals were tracked during two consecutive nights.
Striped Skunks exhibit aposematic behavior, and
defensive postures are obvious (Lariviére and Messier
1996b). Locations immediately following an observer-
induced defensive posture were discarded from analy-
ses due to possible observer disturbance (Lariviére
and Messier 1998b).
Universal Tranverse Mercator coordinates were ob-
tained from the exact location of the skunk following
its departure from the area using a portable Global
Positioning System (Ensign GPS, Trimble Naviga-
tion, Cansel Survey Equipment, Burnaby, British
Columbia). To increase the precision of the coordinates,
we only considered locations for which =3 satellites
were available (D’ Eon 1995).
Our radio-tracking schedule was intensive (1.e., loca-
tions every 15 min), and designed to maximize direct
behavioral information instead of statistical independ-
ence within an animal’s home range. Using Shoener’s
ratio, we estimated that statistical independence be-
tween locations occurred at intervals of 270 min. How-
ever, Striped Skunks are highly mobile, and are capable
of moving >1 km in 15 min, even in dense vegetation
(Lariviére and Messier 1998b). Thus, we assumed that
short movements reflected behavioral decisions by
the skunk, not physical constraints.
We defined maternity dens as any sites for which
at least two of the following criteria were satisfied:
repeated use for >3 consecutive days during the par-
turition/rearing period (15 May - 30 June), collection of
grass for preparation of nest chamber, or presence of
young during the rearing period (Lariviére and Messier
1998a). Because some females may relocate their prog-
eny during the rearing period, and because we could
not assess why females changed maternity dens (e.g.,
to reduce parasite loads, disturbance, relocation to a
better foraging area, etc.), we only used data on females
with a single maternity den for spatial analysis of den
locations within home ranges.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Our entire study area was digitized into a Geograph-
ic Information System (SPANS GIS, PCI Geomatics,
Richmond Hill, Ontario, Canada). Locations of the
maternity dens were plotted for all female skunks, as
well as all telemetry locations to determine home range
size (Lariviere and Messier 1998c) and shape. Home
range shape was quantified using a shape index
(McGarigal and Marks 1994) that was calculated by
the following formula:
shape index = perimeter / (2* Vv (pi * area)).
Thus, a smaller shape index indicates a more cir-
cular range. We compared average shape indices for
males and females with a Student t-test for unpaired
samples. Spatial analyses of activity centres were
performed using software package TRACKER and
RANGES IV, and core areas (50% minimum convex
polygon; Lariviere and Messier 1998c) were consid-
ered activity centers (Samuel et al. 1985).
Results
In 1993 and 1994, we radiotracked Striped Skunks
during 1858 hours, and obtained home range informa-
tion for 20 females and 5 males. Of the 20 females,
11 used only | maternity den during the entire period.
First, we predicted that because of the fixed position
of buildings in the landscapes, females using buildings
as maternity dens would have home ranges less circular
(larger shape index) than those using burrows. This pre-
diction was not supported by our data: the shape index
of females using buildings (mean = 1.53, SE = 0.08,
n=6) did not differ (one-tailed t= 0.45, df =9, P=0.33)
from the shape index of females using burrows as
maternity dens (mean = 1.48, SE = 0.07, n =5).
Second, we hypothesized that if females try to min-
imize the travel distance to any point within their res-
pective home ranges due to lactation constraints, then
home range of females should be more circular (small-
er shape index) than home ranges of males. Our pre-
diction that home ranges of females are more circular
than those of males was not supported: the average
shape index for females (mean = 1.46, SE = 0.03,
n = 20) did not differ (two-tailed t = 0.59, df = 23,
P =0.56) from the average shape index of male home
ranges (mean = 1.50, SE = 0.06, n = 5).
Third, we hypothesized that because buildings are
preferred denning sites (Lariviére and Messier 1998a),
some buildings may be used as maternity dens but not
be located within the core area of the home range. We
predicted that underground burrows should occur in
the core areas more often than dens in buildings. Our
data also did not support this prediction: there were
no associations between den type and whether or not
the den was located inside the core area (n = 12 fe-
males; Fisher exact test, P = 0.52). Overall, three
underground burrows were inside versus two outside
the core area, whereas six buildings were inside versus
one outside the core area. In total, 75% of dens were
inside core areas.
2007
In this study, we did not detect any effect of den type
(anthropogenic or not) on shape of female home ranges
or position of the den. Further, the shape of Striped
Skunk home ranges was similar among males and fe-
males. These findings suggest that female skunks estab-
lish home ranges irrespective of den type and with
rules similar to males.
Examination of the natural history of Striped Skunks
may help explain the patterns observed. First, Striped
Skunks in northern environments undergo long periods
of inactivity during winter, and to survive such period
requires sufficient body reserves (Gunson and Bjorge
1979; Hwang et al. 2007; see also Rogers 1987). Thus,
establishing a home range that includes good foraging
areas may be more critical than choice of maternity
den, but whether choice of maternity den influences
juvenile survival remains unknown. In our study area,
many buildings that were deemed suitable for occupa-
tion by skunks were not utilized (Lariviere et al. 1999).
As well, numerous females that used underground bur-
rows to rear young had suitable unoccupied buildings
within the boundaries of their home ranges. Thus, al-
though buildings may be convenient denning sites, they
may not be necessary or provide reproductive advan-
tages to Striped Skunks. If juvenile mortality in skunks
mostly occurs after emergence, then anthropogenic den
types such as buildings may be used for other reasons
such as thermoregulation, protection from the elements,
or simply convenience.
During the parturition/rearing period of Striped
Skunks, females make nocturnal foraging trips but
return to the maternity den by morning to nurse the
young (Lariviére and Messier 1997). Foraging move-
ments are highly variable (0-20 km per night; Lariv-
iére et al. 1998b) and highly opportunistic, although
mostly focused on insects and small mammals (Green-
wood et al. 1999) and targeted to specific habitats (Bix-
ler and Gittleman 2000; Lariviere and Messier 2000).
Moreover, habitat availability around used and unused
buildings in our study area was similar, suggesting that
distribution of resources may not affect location of
maternity dens, and conversely that choice of maternity
den does not affect access to resources. Suitable den
sites likely were not limiting for Striped Skunks (La-
riviére et al. 1999). The preference of Striped Skunks
for buildings may be most obvious where buildings
are abundant, but absence of buildings as maternity
dens probably does not affect distribution of skunks
in the landscape during summer. Hence, our study
reinforces the previous suggestion that food, instead
of denning structures, probably affects distribution of
Striped Skunks and their foraging activity in the land-
scape.
Acknowledgments
This study was financed by Ducks Unlimited Cana-
da (Institute for Wetland and Waterfowl Research),
the Canadian Wildlife Service, and logistic support
from Delta Waterfowl Foundation. SL is grateful to
LARIVIERE, HOWERTER, and MESSIER: HOME RANGE SHAPE FOR SKUNKS
263
Fonds pour la Formation de Chercheurs et I’ Aide a la
Recherche (FCAR), Québec, for postgraduate and
postdoctoral support, as well as to the National Sci-
ence and Engineering Research Council for an oper-
ating research grant. Technical assistance was provid-
ed by B. Dinter, E. Howard, G. Poon, L. R. Walton,
B. Fry, and D. Anderson. W. A. Gorsuch reviewed an
earlier draft of this manuscript. Methods used in this
study were approved by the University of Saskatche-
wan Animal Care Committee (Approval #920091).
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Received 18 April 2006
Accepted 7 July 2008
Road-kill of Mammals in Nova Scotia
DAvID FupGE!*, BILL FREEDMAN!, MICHAEL CROWELL”, TONY NetrTe?, and VINCE POWER
' Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada
? Jacques-Whitford Environment Limited, 3 Spectacle Lake Road, Dartmouth, Nova Scotia B3B 1Q9 Canada
4 Nova Scotia Department of Natural Resources, Wildlife Branch, Kentville, Nova Scotia B4N 4E5 Canada
Fudge, David, Bill Freedman, Michael Crowell, Tony Nette, and Vince Power. 2007. Road-kill of mammals in Nova Scotia
Canadian Field-Naturalist 121(3): 265-273.
We examined road mortality of wild mammals in Nova Scotia using data from (a) five years of province-wide data on
wildlife collisions involving larger mammals, and (b) 20 months of observations of smaller mammals along a 160-km route
near Halifax. An average of 2079 White-tailed Deer was reported killed annually on highways during 1999 through 2003,
along with 14 Moose and 33 Black Bear. Female White-tailed Deer were more likely to be road-killed than males (by a fac-
tor of 1.8), yearlings more so than older animals, and there were peaks of mortality in the late spring and late autumn. The
road-kill of smaller mammals was highest in the summer and least in the winter, and the most frequent mortalities were of
Raccoon (28% of observations), Porcupine (27%), Skunk (17%), Snowshoe Hare (6%), Coyote (4%), and Groundhog (3%).
The standardized kill-rate (number/100 km) was highest on a 100-series highway compared to trunk highways or urban
streets, but the rate standardized to vehicle density was highest on trunk highways.
Key Words: White-tailed Deer, Odocoileus virginianus, Moose, Alces alces, Black Bear, Ursus americanus, Raccoon, Procyon
lotor, Porcupine, Erethizon dorsatum, Skunk, Mephitus mephitis, Snowshoe Hare, Lepus americanus, Coyote, Canis
latrans, Groundhog, Marmota monax, road-kill, mortality, wild mammals, Nova Scotia.
Roads are a defining feature of modern transportation
systems. Networks of public roads connect population
centers, allowing the movement of people and goods in
increasingly larger numbers of fast-moving vehicles.
Because much highway infrastructure traverses wildlife
habitat, an unfortunate consequence is accidental col-
_ lisions that threaten the lives of both people and ani-
mals, while also causing damage to property (Romin
and Bissonette 1996). In Nova Scotia, for example,
503 collisions between vehicles and large wild mam-
mals were reported in 2003, with 68 of the incidents
resulting in personal injury requiring emergency serv-
ices (NSDTPW 2006*). Data relating to the economic
_ costs of such accidents have been quantified in Maine,
where they result in US$32 million/year in property
damage and medical costs (MEDOT 2001*). Overall
in the United States, vehicle accidents involving only
White-tailed Deer (Odocoileus virginianus) annually
-tesult in about 200 human fatalities, 30 thousand
injuries, and more than US$1 billion in property dam-
age (Conover 1997). In Europe, collisions with ungu-
| lates result in 300 human fatalities and US$1 billion
in property damage annually (Bruinderink and Haze-
broek 1996). Clearly, a large socio-economic cost is
associated with collisions with large animals on roads.
Roads also have important ecological impacts, in-
cluding the following: (a) damage to populations of
some species through mortality; (b) acting as barriers to
movement or disrupting behaviour in other ways; (Cc)
destroying and fragmenting natural habitat; (d) caus-
» ing pollution by de-icing salt, metals, and pesticides;
_and (e) enhancing access for hunters (Bennet 1991;
Evink 1999). The direct mortality of larger animals due
to vehicle collisions is relatively apparent and well-
quantified (Clarke et al. 1998; Cain et al. 2003; Cle-
venger et al. 2003; Saeki and MacDonald 2004). Road-
kill may affect the conservation status of species —
about 65% of the mortality of the endangered Florida
Panther (Puma concolor coryi) is caused by vehicular
collisions (Harris and Gallagher 1989). Other at-risk
populations threatened by road-kill mortality include
Ocelot (Leopardus pardalis) in the U.S. southwest
(Tewes and Blanton 1998), the European Badger
(Meles meles) in Great Britain (Clarke et al. 1998),
and the endangered mainland Moose of Nova Scotia
(Alces alces americana) (NSDNR 2008*).
Many jurisdictions have implemented mitigations
to reduce the frequency of collisions between vehicles
and wildlife. Often, this is limited to warning signs in-
stalled along highway sections known to have high
collision rates (AMEC 2004*). In some other places,
highways may have perimeter fencing or even wildlife
under- and over-passes to keep animals off the roads,
particularly in national parks and wilderness areas
(Clevenger 1997; AMEC 2004*; NBDOT 2008*). Such
mitigations help prevent animals from entering the
right-of-way and are successful in reducing mortality,
particularly of mammals (Foster and Humphrey 1995).
Interestingly, the effectiveness of animal-crossing signs
in reducing collision rates has not yet been quantified
(MEDOT 2001*).
Despite the fact that collisions with vehicles are an
important source of mortality of wildlife, there has
been little research into the phenomenon. Putman
(1997) argued that such work is necessary, because
understanding the biological and ecological influences
265
266
on road-kill rates is integral to the design and imple-
mentation of effective mitigations to deal with this
problem. Within this context, the great majority of
research into road-kill has dealt with larger mammals.
This focus is due to several socio-economic factors,
including the danger to people associated with colli-
sions with large mammals, their economic importance
in hunting, and their high public profile (Cain et al.
2003; Clevenger et al. 2003). A few studies have also
examined factors influencing road-kills of smaller
mammals (Garland and Bradley 1984; Clevenger et
al. 2003; Saeki and Macdonald 2004).
In the present study the road-kill mortality of larger-
and medium-sized mammals is examined for a variety
of roads in Nova Scotia. Two data sets were used: (a)
province-wide incident data for larger mammals over
a five-year period, and (b) data for medium-sized mam-
mals over a 20-month period along a 160-km work-day
commute. We examine biological and environmental
factors possibly influencing the road-kill rates, includ-
ing gender, age, and proximity to distinctive habitats.
Methods
This study was done in Nova Scotia, an eastern
province with a population of about 934 thousand peo-
ple and 23 thousand km of public roads (NSDTPW
2008*). Data for larger mammals were obtained from
incident reports of the Nova Scotia Department of
Natural Resources (NSDNR), which records road-kill
of White-tailed Deer, Moose (Alces alces), and Black
Bear (Ursus americanus), including the location, date,
species, sex, and age of the dead animals. Although
the reporting of kills is not mandatory, NSDNR esti-
mates that 60-90% of bear and deer collisions and all
involving Moose are reported to them (see Conover
et al. 1997). The original NSDNR data were taken from
a FoxPro database and converted to Excel spread-
sheets. The original location coordinates were con-
verted to UTM using ArcView software to allow for
analysis by a geographical information system (GIS).
The number of road-kills for each species was
summed monthly and annually from 1999 through
2003. The data for White-tailed Deer and Moose were
also analyzed by gender and by yearling and multi-
year animals. A GIS analysis was used to plot the
distribution of road-kills on digital maps, and spatial
relationships with road type and traffic volume were
examined. In cases where the carcass was not found
immediately beside a road, because the wounded ani-
mal had moved before it died, the nearest road was
inferred as the place of impact. The traffic volume of
highway segments was identified for reaches between
successive exits and interchanges, using archived data
from NSDTPW (2008*). The 100-series highways of
Nova Scotia have relatively high traffic volumes and
are often four lanes with controlled access, while the
trunk highways are relatively low-volume, two-laned,
and have free access. The annual road-kill data were
standardized on a per-km and per-10* vehicles basis,
THE CANADIAN FIELD-NATURALIST
Vol. 121
and were compared across highway types and on the
basis of adjacent habitat (categories were: mature soft-
wood forest (>60% conifer trees), mature hardwood
(>60% angiosperm trees), mature mixedwood, imma-
ture forest, agricultural, and urban land-use; a sepa-
rate analysis involved surface water within 100 m.
The relationship of road-kill density was also com-
pared with indicators of local species abundance for
White-tailed Deer and Moose. For deer, density was
indicated by county-level Pellet Group Inventory (PGI)
data (obtained by NSDNR in the spring, after snowmelt
but before green-up, along 460 random | km x 2 m
transects throughout the province). For Moose, the
abundance polygons of Snaith (2001) were used.
Data for smaller mammals were collected over a
20-month period during 1993 through 1994 along a
160-km weekday commuting route (by M. Crowell)
from the rural village of Urbania to the urban Burn-
side Industrial Park in Dartmouth. The route included
106 km of 100-series highway (part of Highway 102),
38 km of trunk highway (Highway 215), and 16 km of
urban road. Animals observed dead beside the road
were identified to species, as juvenile or adult, and the
location was mapped. The annual road-kill data were
standardized per km of road, per km? of road-bed, and
per 10? vehicles of traffic density (using archived data
from NSDTPW (2008*)). The total-kill data and that
for Raccoon (Procyon lotor), Porcupine (Erethizon
dorsatum), and Striped Skunk (Mephitis mephitis) were
analyzed with respect to adjacent habitat along the
surveyed reaches of 100-series highway, including:
(a) roadbed being lower than, raised, or level with the
adjacent terrain; (b) adjacent habitat being mature
softwood forest, mature hardwood, mature mixed-
wood, immature forest, agricultural, or urban land-use;
and (c) surface water within 100 m. A coefficient of
similarity was used to make pair-wise comparisons of
the relative dominance of species among road-types:
S, = 2C/(A+B), where C is the number of species
killed on both road-types, while A and B are the num-
bers on particular road-types.
Observations and Discussion
White-tailed Deer
A total of 10 397 White-tailed Deer were reported
killed in vehicle collisions from 1999 through 2003,
or an annual average of 2 079. This mortality is equiv-
alent to about 6% of the estimated deer population,
and is a 28% increment on top of the reported hunt (on
average, hunters take 20% of the Nova Scotia herd
during the fall hunt; NSDNR 2007*).. The road-kill
rate was similar during 1999 through 2001 (average
+ SD: 2 137 + 102), but greater than during 2002 and
2003 (1 449 + 44: P = 0.003 for difference between
the groups of years). The smaller road-kill of 2002-
2003 is likely related to a reduced deer herd during
that period (NSDNR 2007*).
There was a bimodal pattern of peak mortality of
multi-year individuals (of both sexes) during June-
2007 FUDGE, FREEDMAN, CROWELL, NETTE, AND POWER: ROAD-KILL OF MAMMALS IN NOVA SCOTIA 267
July and October-November (Figure 1a). Other authors
have reported a fall peak of deer road-kill, and have
attributed this to breeding activities (Puglisi et al. 1974;
Case 1978). During the rut, bucks move about in search
of receptive females, and does are also relatively vagile
during that period (Banfield 1974). The fall hunting
season may also increase road-kill rates (Puglisi et al.
1974; Allen and McCullough 1976), although Kilgo
et al. (1998) found that deer in Florida generally avoid
roads and other open areas when under hunting pres-
sure. The increase of deer road-kill in June-July could
be related to such factors as increased rural traffic asso-
ciated with tourists and cottagers, the availability of
high-quality forage and residual de-icing salt along
verges, harassment by biting flies and partial refuge
from them in open areas, and increased mobility of
does with fawns during this time. Puglisi et al. (1974)
commented on the attractiveness of early-greening road
verges to deer in the springtime. We observed a par-
ticularly large peak of road-kill in the spring of 2001
following an extremely harsh winter, during which
about half of the provincial herd starved to death
because of deep snow cover (NSDNR 2007*).
Adult females were 1.8-times more likely to be killed
in vehicle collisions than adult males. This is likely due
to the Nova Scotian deer herd being disproportionate-
ly female because of the selective removal of males by
hunters (Feldhamer et al. 1986; Bruinderink and Haze-
broek 1996). Yearling mortality was 11% of the total
road-kill mortality, and the kill rate was similar to that
of adults (when standardized by estimated population
size), although there was little seasonality of the year-
ling mortality.
The road-kill rate was higher on two-lane secondary
highways (0.38 kills/km.yr) than on four-lane 100-series
highways (0.13/km.yr). Bashore et al. (1985) made a
similar observation in Pennsylvania, where there were
more deer road-kills along two-lane roads than along
a larger interstate highway. Putman (1997) found that
deer would routinely cross secondary roads within their
territory, but would avoid larger interstate highways,
likely because of their greater width and faster and more
plentiful traffic.
The road-kill rate per 10* vehicles was higher on
secondary (0.092/10+ vehicles) than on 100-series high-
ways (0.012/10*). Overall, however, we did not find a
clear relationship between road-kill rates and traffic
volume at most of the tested spatial scales, suggesting
that more cars do not directly relate to more acci-
dents. Carbaugh et al. (1975) also did not find a linear
relationship between traffic density and deer kills, and
Armstrong (2005*) reported a slight inverse relation-
ship. Although we did not examine the phenomenon, it
is likely that high traffic rates during crepuscular hours
would be particularly lethal to deer trying to cross roads
(Allen and McCullough 1976).
We observed seven clusters of deer road-kills, each
located near a human population centre with a high-
density road network. A linear regression using average
county data found a positive correlation between the
estimated density of deer and that of road-kill (R* = 0.71;
P<0.01).
In some areas, road-kill may exert a controlling in-
fluence on local deer populations and slow their recov-
ery from high winter mortality. There are economic
implications of such effects because White-tailed Deer
are the most valuable wildlife resource in North Amer-
ica in terms of hunting (Conover 1997; DuWors et al.
1999*). Romin and Bissonette (1996) estimated that
the inclusive value of deer in North America is about
$2 200 per animal (Canadian dollars adjusted for infla-
tion to 2005).
Moose
Seventy Moose were reported killed in vehicle colli-
sions from 1999 through 2003 (average = 14/yr). About
two-thirds of the mortality involved the relatively abun-
dant population on the highlands of Cape Breton, which
is derived from 18 released individuals (Alces alces
andersoni) imported from Alberta in the late 1940s
(Pulsifier and Nette 1997). The annual road-kill mor-
tality is equivalent to about 0.23% of the Cape Breton
population, and 3.7% of the reported harvest by hunters
(in 2003; this does not include the Mi’ kmaq harvest,
which is estimated to be similar to or exceeding the
licenced hunter harvest). However, the native mainland
population of Moose (Alces alces americana) is much
less abundant, likely fewer than a thousand individuals,
and since 2003 it has been listed as endangered under
the Nova Scotia Endangered Species Act. The annual
road-kill mortality of mainland Moose is equivalent
to about 0.5% of the population (there is no legal hunt-
ing of this population).
Moose road-kills occurred in all months except April,
with a peak of male mortality in October, and of fe-
males during June-July (Figure 1b). The autumn peak
of male mortality occurs during the rut, when they are
competing and moving about in search of receptive
females (Banfield 1974). Males also become more
aggressive during the rut and may be less averse to
roads and traffic (Joyce and Mahoney 2001). The
female peak of road-kill in early summer is related to
the high volume of tourism-related traffic that occurs
then on Cape Breton highways, particularly on the
Cabot Trail.
Similar numbers of males and females were killed
on the mainland (10 vs. 12), but 1.6-times more fe-
males were killed on Cape Breton roads. The latter
observation is related to the female-skewed sex dis-
tribution of Moose on Cape Breton (the gender ratio
of kills by hunters is about three males per two fe-
males). Yearling mortality was 11% of the total, but
their kill rate was similar to that of adults (when stan-
dardized by their estimated population size). There
was no obvious seasonality of the yearling mortality.
Of the road-kills on Cape Breton, 85% occurred
along eastern sections of the Cabot Trail, where traf-
fic volumes are relatively high. The highest kill rate
(0.4/10 km. yr) occurred along a stretch from French
268
River Bridge to Wreck Cove along the Cabot Trail,
within Cape Breton Highlands National Park. On
mainland Nova Scotia, the highest road-kill rate was
on Highway 104 on the upland Cobequid Pass Toll
highway, where 38% of the mainland kills occurred
(0.2/10 km.yr). These high-kill areas coincide with
regions of Nova Scotia that have the highest density
of Moose (Snaith 2001).
Black Bear
One hundred and sixty-four Black Bears were report-
ed killed in vehicle collisions from 1999 through 2003
(average = 33/yr). The annual mortality is equivalent
to about 0.5% of the estimated population of Black
Bears in Nova Scotia, and 7.2% of the reported har-
vest by hunters (in 2003). The lowest kill-rate was in
1999 (10 individuals) and the highest in 2003 (57). An
unusually large number of 27 bears were road-killed
in June 2003, compared to the 1999-2002 June average
of 5. This may have been due to an increasing bear
population, or the springtime of 2003 being unusually
cool and wet, so that animals emerging from hiberna-
tion had trouble finding adequate natural food and were
attracted to areas near human settlement.
There was a bimodal pattern of road-kill mortality,
with peaks in May-June and in October-November
(Figure Ic). The first peak is likely due to activity as-
sociated with the breeding season, when males wander
in search of females (Klenney 1987). The early sum-
mer is also when sub-adult animals, at about 15 months
of age, leave their mothers to find their own territory
(Matson 1967). The fall peak of mortality is likely relat-
ed to activity associated with pre-hibernation feeding
and movement to denning sites (Matson 1967; Davis
and Browne 1996). No road-kills were recorded from
mid-November to early April, when bears are in hiber-
nation. It was not possible to factor the influence of age
or sex on bear road-kill, as these data were not con-
sistently recorded.
Bear road-kills occurred throughout the province,
except in southeastern Cape Breton, where the species
does not have an established population (Macmichael
2007). The 100-series highways had a higher linear
mortality rate (1.58 kills/100 km.yr) than secondary
highways (0.032/100 km.yr). According to traffic den-
sity, however, the 100-series and secondary highways
had similar mortality rates (both 0.02 kills/10° vehi-
cles.year). Relative to their total estimated population
size in Nova Scotia, Black Bear road-kill rates are
lower than those of White-tailed Deer. This may be
due to bears being relatively wary of humans and
roads, except for habituated animals (Herrero 1985).
In addition, deer tend to be relatively abundant in
areas of Nova Scotia with high populations of peo-
ple, especially in agricultural and coastal regions.
Other Mammals
Three hundred and ninety-two smaller mammals of
sixteen species were recorded as road-kill on a 160-km
THE CANADIAN FIELD-NATURALIST
Vol. 121
study route during a 20-month survey (Figure Id). The
most frequent mortalities were of Raccoon (28% of
kills), Porcupine (27%), and Striped Skunk (18%).
These are relatively abundant in the study area, often
occurring near roads and other habitat edges, and are
at inherent risk because they are relatively deliberate
in their movements and have a body too large to pass
beneath most moving vehicles without impact (Gar-
land and Bradley 1984; Rolley and Lehmen 1992; Cle-
venger et al. 2003). All species recorded as road-kill are
indigenous to the study area, except for Coyote, Canis
latrans, domestic cat, Felis domesticus, and domestic
dog, Canis familarus (Banfield 1974; Davis and
Browne 1996). Additionally, a Red-necked Wallaby
(Macropus rufogriseus) that had been a 6-month
escapee from a wildlife park was found by one of us
as road-kill in the Annapolis Valley of Nova Scotia.
Discoveries of small-mammal road-kills (rodents
and shrews) were uncommon, but this is likely due to
the difficulty of detecting these animals when scan-
ning from a moving vehicle, plus the likelihood that
they are quickly scavenged. In addition, small mam-
mals are reluctant to cross multi-lane highways (Cle-
venger et al. 2003). An innate reluctance to cross open
spaces may also explain why Snowshoe Hare, Lepus
americanus (the most abundant medium-sized mam-
mal in the study area), was killed less frequently than
Raccoon, Porcupine, or Striped Skunk, and why their
kill rate was higher on the two-lane Highway 215
than on the four-lane Highway 102. Clevenger et al.
(2003) also found that Snowshoe Hare was more
likely to be killed on narrow roads than on wider ones
in Banff National Park, Alberta.
The road-kill rate, when standardized by distance
surveyed, was highest on the four-lane Highway 102
(6.0 kills/km) than on the two-lane Highway 215
(4.0/km) or city streets (1.3/km). However, when stan-
dardized to traffic density, Highway 215 had a greater
rate of mortality (6.3 kills/10* vehicles) than Highway
102 (1.9/10* vehicles). The kill rate per 10* vehicles
approached zero on urban streets because of the large
traffic volumes, lower vehicle speeds, and likely a rel-
atively small abundance of wild mammals. The lower
rate on Highway 102 is likely due to its much greater
width and larger volume of faster-moving vehicles,
which likely discouraged animals from crossing. Cle-
venger et al. (2003) also found that relatively wide and
high-traffic roads discouraged small- and medium-sized
mammals from crossing in Alberta, as did Clarke et
al. (1998) for badgers in England.
Similar numbers of species were observed killed on
the four-lane Highway 102 (n=12) and on the second-
ary Highway 215 (n=11), and both exceeded urban
streets (only Red Squirrel, Zamiasciurus hudsonicus,
were observed road-killed in the urban area studied).
The most frequent mortalities on the secondary High-
way 215 were Raccoon (19% of the total), Striped
Skunk (18%), Red Squirrel (16%), Snowshoe Hare
2007 FUDGE, FREEDMAN, CROWELL, NETTE, AND POWER: ROAD-KILL OF MAMMALS IN Nova SCOTIA 269
(a) White-tailed Deer
14 — 7 —— =
12 —_
— 10
Ble i
re M
A
S&S 6 Tot:
4
2
oO
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
(b) Moose
30 —
25
| 20 a
3 oF
epee te) M
6
Tota
S 10 a
mE
g _ Dai. ig we eo
1 zZ 3 t 5 6 P 8 9 10 11 12
(c) Black Bear
35
;
eee es
£ 20
ee ey as EX =ans
SS
I CE ee a ae ae |
ee a SP EN
ee ee Ses eee
ie)
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
(d) Smaller Mammals
S
=
qy
is)
SS
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Ficure |. Average monthly road-kill mortality of large mammals from 1999 through 2003: (a) White-tailed Deer (m = 10 397):
(b) Moose (n = 70); (c) Black Bear (n = 164); (d) smaller mammals (7m = 392). Sum of genders may not add to totals,
because the sex of some animals was not identified.
270
(14%), and Porcupine (11%). The most frequent on the
four-lane Highway 102 were Porcupine (31%), Raccoon
(30%), and Striped Skunk (18%). The species spec-
trum of mortality of Highways 102 and 215 had a
similarity coefficient of 0.78, but their similarity with
urban streets was very low — only 0.00 and 0.08, respec-
tively.
The greatest rate of road-kill occurred in the summer
months of June (35 mortalities), July (34) and August
(22) (Figure 1d). Smaller numbers were recorded in the
winter months of November through February, when
animals are hibernating or generally less active. The
road-kill of Raccoons was highest in May and June,
when adult males search widely for females with which
to mate (Banfield 1974). In addition, 43% of the Rac-
coon mortality in May and June was yearlings, which
disperse during that time. The mortality of Striped
Skunks was highest in September, when juveniles dis-
perse from their natal home range. The mortality of
Porcupines was relatively even throughout the spring,
summer, and fall.
Partial-deviance tests indicate that roadside topogra-
phy and habitat were significant predictors of Raccoon
road-kill. The likelihood of a Raccoon road-kill was
much greater at level roadside topography (x? = 11.3,
df = 3, P <0.001) and along segments of highway
bisecting mature forest (y? = 9.4, df = 3, P <0.05).
Proximity to water did not have a significant effect
(x° = 0.50, df = 1, P >0.05). The likelihood of Rac-
coon road-kill at different roadside topographies was
assessed using log-likelihood ratios, and was much
greater at level crossings (T-value = 3.1; P <0.05) and
along segments bordering mature forest (T-value = 5.9;
P <0.0025). For Porcupines, topography (x2 = 29.4,
df = 4, P <0.005) and proximity to water (7 = 5.9,
df = 1, P <0.025) were significant predictors of road-
kill. Log-likelihood ratios indicate that road-kill was
most frequent at level roadside topography (T-value
4.1; P <0.025) and adjacent to mature softwood forest
(T-value 4.0; P <0.025). For Striped Skunks, topogra-
phy (x? = 27.4, df = 2, P <0.005) was the only signifi-
cant predictor of road-kill, and the log-likelihood of
mortality was much higher at level roadside topogra-
phy (T-value = 4.1; P <0.025).
Overall, no road-kills of any species were observed
at sites where the roadbed was raised (higher than the
adjacent terrain), and the likelihood of road-kill was
higher on roads bisecting mature forest (mixed, soft-
wood, or hardwood) than adjacent to immature for-
est, agricultural fields, or urban habitat (T-value 3.8;
P <0.05). Proximity to water did not have an overall
significant predictive effect.
The results suggest that far fewer road-kills occurred
on raised than on level or buried roadbeds. Raised
roadbeds could serve as a visual barrier to habitat on
the other side, and the banked incline may act as a
physical deterrent to crossing. Clevenger et al. (2003)
made similar observations for smaller-animal road-
THE CANADIAN FIELD-NATURALIST
Vol. 121
kills in Banff National Park, where raised roadbeds had
a lower risk of mortality than other topographies.
Another factor potentially affecting road-kill rates
is the presence of vegetated center medians on divid-
ed highways, such as the grassy ones occurring on
Highway 102 in the present study. Bellis and Graves
(1971) suggest that these habitats function as temporary
refuge areas for animals attempting to cross major
highways.
Our study does not allow for an assessment of the
impact of road-kill mortality on the local abundances
of smaller mammals, as population estimates are not
available. If our data were extrapolated to the total
length of 100-series (3 x 10° km) and secondary
(4 x 10° km) highways in Nova Scotia, the estimated
annual mortality within this size range would be
20 x 10°? mammals. It can be concluded that the num-
bers of animals killed on Nova Scotian highways is
large, and that this is a significant anthropogenic stres-
sor. Our field data for smaller mammals were collect-
ed in 1993-1994, and since then traffic volumes on
Nova Scotian roads have increased by about 24% since
then (NSDTPW 2008*), so it is likely that road-kill
rates have also increased. Moreover, some new roads
are being built and the 100-series highways are being
expanded, which may also be increasing the rates of
road-kill.
Mitigation
Clearly, large numbers of mammals are being killed
on Nova Scotian roads, and the further creation and
upgrading of highways and increasing traffic may wors-
en this situation. Moreover, there are important socio-
economic consequences of collisions with wildlife.
These include injuries to people involved in the colli-
sions, the associated medical costs and time off work,
damages to property, and decreases in the popula-
tions of animals available to be hunted or trapped.
For example, during our study period of 1999
through 2003, White-tailed Deer were involved in
204 vehicle collisions in which people suffered
injuries requiring hospitalization, and seven incidents
involved a human fatality (NSDTPW 2006%*). In fact,
White-tailed Deer cause more collision-related prop-
erty damage and human mortality than any other
wild animal in North America (Conover 1997).
According to Putman (1996), an average collision with
a deer costs about $3 400 in vehicle damage (Canadi-
an dollars, adjusted for inflation to 2005). Using this
cost estimate, deer collisions in Nova Scotia during
our study period resulted in about $6.8 million in
annual property damage, not including costs associ-
ated with health care and emergency services.
Fortunately, there are ways to reduce road-kill along
highways (Woods 1990*; Clevenger et al. 2001;
USDOT 2002*; AMEC 2004*). Some mitigations are
intended to change the behavior of animals in ways
that make them less likely to enter the right-of-way.
For example, fencing is used to prevent animals from
2007 FUDGE, FREEDMAN, CROWELL, NETTE, AND POWER: ROAD-KILL OF MAMMALS IN Nova ScoTIA 27]
TABLE |. Road-kill of smaller mammals over a 20-month study period.
Highway 101
i
Species n
Raccoon (Procyon lotor) 97
Porcupine (Erethizon dorsatum) 10]
Striped Skunk (Mephitis mephitis) 60
Snowshoe Hare (Lepus americanus) 14
Domestic Cat (Felis domesticus) 17
Coyote (Canis latrans) 13
Groundhog (Marmota monax) 10
Red Squirrel (Jamiasciurus hudsonicus) 0
Domestic Dog (Canis familiaris) 4
Red Fox (Vulpes vulpes) 4
Eastern Chipmunk (7amias striatus) 0
Beaver (Castor canadensis) |
Mink (Mustela vison) ]
Muskrat (Ondatra zibethicus) |
Red-backed Vole (Clethrionomys gapperi) 0
Woodland Jumping Mouse (Zapus hudsonicus) 0
Unknown mammal |
Total Species = 16 324
crossing major roadways in at least eleven U.S. states
(MEDOT 2001*), New Brunswick, and in the Cana-
dian Rockies. In Banff National Park, for example,
fencing resulted in a 96% reduction in large-mammal
road-kill (Clevenger 1997). It is generally recommend-
ed that fencing be at least 2.6 m in height, including
above winter snow levels (USDOT 2002*). However,
fencing also has drawbacks, because it can fragment
populations and may trap animals within a right-of-
way that they may enter via a break or at a fence-end
(Phillips 1999*). Phillips (1999*) estimated that fenc-
ing along a major highway in New Brunswick typi-
cally costs $40 000 to $80 000 (Canadian) per kilome-
ter. At present, the only areas in Nova Scotia with
wildlife fencing along road verges are on the Cabot
Trail on Cape Breton Island to prevent devastating col-
lisions with Moose in segments where there are steep
drop-offs along the highway, and an area near Maga-
zine Hill in Dartmouth that had experienced a high
number of accidents with deer.
Although extremely expensive, structural crossings
can be installed to allow for the movement of animals
across a right-of-way. This mitigation can help pre-
vent local and even regional populations of certain
animals from becoming fragmented, with potentially
deleterious consequences, while also avoiding risks to
people and property from collisions. Crossing structures
include underpasses, overpasses, and modified water-
course structures (a kind of underpass). However, not
all animals will use structural crossings. Reed (1981)
observed that Mule Deer (Odocoileus hemionus) were
reluctant to use enclosed underpasses, and they did not
acclimatize to them over a study period of 10 years.
He suggested that deer were far more willing to use
open “bridge” style overpasses than underpasses. A
review by USDOT (2002*) also concluded that ungu-
Highway 215 Urban lotal
c ‘
n c n n a
29.9 1 19.3 0 108 27.5
Ships 6 10.5 0 107 27.3
18.5 10 17.5 0 70 17.9
4.3 8 14.0 0 22 35
5.2 4 7.0 0) 21 5.4
4.0 9) 0.0 0 13 3.3
2 BiG 0 12 3.)
0.0 9 15.8 3 9 2.3
1.2 2 35 5) 6 ics
1.2 0 0.0 6) 4 1.0
0.0 3 5.3 (0) 3 0.8
0.3 0 0.0 0 ] 0.3
0.3 0 0.0 0 ] 0.3
0.3 0 0.0 0 l 0.3
0.0 | 1.8 0 l 0.3
0.0 | 1.8 0 | 0.3
0.3 0 0.0 0 l 0.3
57 3 392
lates were most likely to use overpass structures. The
installation of a single overpass crossing structure
can cost Canadian $1.4-2.6 million, while underpass-
es and modified watercourse structures are generally
$92 000 to $420 000 (MEDOT 2001*).
Another useful mitigation is to make drivers more
aware of potential dangers and to reduce vehicular
speeds (AMEC 2004*). These driver-orientated ap-
proaches include signage and educational programs.
Passive warning signs (sometimes with flashing lights)
of White-tailed Deer and Moose are used in many
places in Nova Scotia and elsewhere to alert motorists
to increased risks of collision with wildlife, generally
in places where repeated collisions have occurred
(AMEC 2004*). Although there are no data supporting
the efficacy of signs, they likely have limited effective-
ness because they rely on driver attentiveness, yet peo-
ple quickly become desensitized to them (USDOT,
2002*). Passive signs are, however, relatively inexpen-
sive to install and maintain.
Driver education related to road-kill may make
motorists more aware of the potential for wildlife en-
counters, and therefore better prepared to avoid them.
For example, a program called “Watch out for Wild-
life” is delivered in the public-school system and by
driver-education messaging in Florida (Evink 1999).
The effectiveness of education programs in reducing
road-kill is difficult to quantify, although people skilled
at spotting animals and predicting their behaviour tend
to be involved in fewer collisions (Joyce and Mahoney
2001).
In Nova Scotia, enhanced public awareness of wild-
life-collision risks could be promoted by an increased
educational program. In addition, fencing verges in
stretches of all major roads known to have high kill
rates of larger mammals would reduce hazards, partic-
272
ularly for conservation of the at-risk mainland Moose
population along the Cobequid Pass of Highway 104
(which also has high rates of bear and deer road-kills).
However, roadside fencing could also detrimentally
fragment the habitat of the already-small populations
of mainland Moose in these areas.
Of course, mitigations have significant costs, but they
provide large benefits by reducing property damage
and risks of injury to people and wildlife.
Acknowledgments
This work was supported by a Discovery Grant to
BF from the Natural Sciences and Engineering Research
Council of Canada. Lester Hanley of the Nova Scotia
Department of Transportation provided traffic statis-
tics for roads, Jennifer Milne and James Boxall of
Dalhousie University assisted with the GIS analysis,
and Cathy Wambolt with statistical analysis.
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DuWors, E., M. Villeneuve, F. L. Filion, R. Reid, P.
Bouchard, D. Legg, P. Boxall, T. Williamson, A. Bath,
and S. Meiss. 1999. The Importance of wildlife to Cana-
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MEDOT (Maine Department of Transportation). 2001.
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2006. The Furbearers and Upland Game Program.
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Received 13 September 2006
Accepted 15 April 2008
Third Census of Seabird Populations of the Gaspé Peninsula, Québec,
2002
RICHARD COTTER and JEAN-FRANCOIS RAIL
Canadian Wildlife Service, P. O. Box 10100, Ste-Foy, Québec G1V 4H5 Canada
Cotter, Richard, and Jean-Francois Rail. 2007. Third census of seabird populations of the Gaspé Peninsula, Québec, 2002.
Canadian Field-Naturalist 121(3): 274-286.
In the tradition of the quinquennial census of seabirds in the Migratory Bird Sanctuaries along Québec’s North Shore, which
began in 1925, this paper presents the results of the first three seabird censuses of the Gaspé Peninsula conducted in 1979,
1989, and 2002, with an emphasis on the third census and on changes in seabird populations that occurred between the sec-
ond and third censuses. In 1979 the population of seabirds was estimated at 134 163 birds, and this increased 72% to 231 186
birds in 1989. The upward trend in population continued in the 1990s but less sharply. The 2002 census recorded 279 276 birds,
a 21% increase over the second census. The seabird community of the Gaspé Peninsula comprises 14 species. In each of the
three censuses, three species accounted for over 75% of the total seabird population: Northern Gannet, Black-legged Kittiwake,
and Common Murre. Between 1989 and 2002, Common Eider, Northern Gannet, Great Cormorant, Ring-billed Gull, Razorbill,
and Black Guillemot increased in numbers, whereas Herring Gull, Black-legged Kittiwake, and Common Tern all suffered
declines. Populations for Double-crested Cormorant, Great Black-backed Gull, and Common Murre all remained stable. Very
small populations (<25 birds) of Leach’s Storm-Petrel and Atlantic Puffin have been recorded during the censuses at only
one location, Bonaventure Island. In the 2002 census there were no petrels observed, neither of adults nor of active burrows.
A Vinstar des inventaires quinquennaux des oiseaux marins dans les refuges d’ oiseaux migrateurs de la Céte-Nord, entrepris
en 1925, cet article présente les résultats des trois premiers inventaires complets des oiseaux de mer nicheurs en Gaspésie, menés:
en 1979, 1989 et 2002. Une attention particuliére est mise sur le statut des espéces d’aprés le troisiéme inventaire, et sur les
tendances des populations entre les deux derniers inventaires. En 1979, le nombre d’oiseaux marins nicheur était évalué a
134 163 oiseaux, puis ce total a augmenté de 72% pour atteindre 231 186 oiseaux en 1989. Cette tendance a la hausse a con-
tinué dans les années 90, mais moins rapidement. En 2002 l’estimation était de 279 276 oiseaux, soit une hausse de 21 % par
rapport au deuxieme inventaire. La communauté d’ oiseaux de mer de la Gaspésie est formée de 14 especes nicheuses. Dans
chacun des trois inventaires, trois espéces représentaient plus de 75 % de la population totale des oiseaux marins: le Fou de
Bassan, la Mouette tridactyle, et le Guillemot marmette. Entre 1989 et 2002, I’ Eider a duvet, le Fou de Bassan, le Grand Cor-
moran, le Goéland a bec cerclé, le Petit Pingouin et le Guillemot a miroir ont vu leurs nombres croitre, tandis que le Goéland.
argenté, la Mouette tridactyle et la Sterne pierregarin subissaient des déclins. Les populations de Cormoran a aigrettes, Goéland
marin et Guillemot marmette sont demeurées stables. Les petites populations (<25 oiseaux) d’Océanite cul-blanc et de
Macareux moine n’ont été vues durant ces inventaires qu’a l’ile Bonaventure. Cependant, aucun nid actif d’océanite ne fut
localisé a cet endroit en 2002.
Key Words: Seabirds, populations, larids, alcids, Gaspé Peninsula, Bonaventure Island, Forillon National Park, Québec.
Seabird populations of the Gaspé Peninsula have
long been well known due to two very popular sum-
mer tourist destinations: Forillon National Park of
Canada and the town of Percé. At Percé boat tours are
offered around Rocher Percé and Bonaventure Island,
which together form a provincial park as well as a fed-
eral Migratory Bird Sanctuary. The most striking fea-
tures of these sites are the high vertical cliffs holding
large Black-legged Kittiwake colonies and the spectac-
ular Northern Gannet colony on Bonaventure Island.
(Scientific names are in Table 1.) With eight breeding
seabird species numbering more than 100 000 pairs in
2002, Bonaventure Island is undoubtedly the most
important seabird colony in the area; nevertheless, the
smaller mainland cliffs, together with a few islands
and sand bars distributed around the peninsula, are also
home to numerous colonies of cormorants, gulls,
guillemots and Razorbills.
Chapdelaine and Brousseau (1992) published the
first account the distribution and abundance of all
274
seabird species breeding along the entire Gaspé Penin-
sula in eastern Québec. They presented the results of the
first systematic censuses of seabirds for the region—in
1979 and 10 years later in 1989—as well as changes in’
populations and distribution for each species. After ani
interval of 13 years, in 2002 the Canadian Wildlife
Service conducted a third seabird census. The objec-
tive of this paper is to present the results of all three
censuses, but with an emphasis on describing thei
changes in seabird populations and distribution that
occurred between 1989 and 2002. This paper thereforet
complements and updates Chapdelaine and Brousseau
(1992) who discussed the changes that occurred:
between 1979 and 1989.
Methods
On the Gaspé Peninsula, seabird species (Table 1)
were censused at 59 breeding sites (Table 2, Figure 1)
during June and early July in 1979, 1989, and 2002 by
the Canadian Wildlife Service-Québec Region. There,
2007
COTTER AND RAIL: SEABIRD POPULATIONS OF GASPE PENINSULA
275
TABLE 1. Species names and summary of census methods used for each species in the 1979, 1989, and 2002 seabird census-
es of the Gaspé Peninsula.
English name Scientific name
Common Eider Somateria mollissima
Oceanodroma leucorhoa
Morus bassanus
Phalacrocorax auritus
Leach’s Storm-Petrel
Northern Gannet
Double-crested Cormorant
Phalacrocorax carbo
Larus delawarensis
Larus argentatus
Great Cormorant
Ring-billed Gull
Herring Gull
Great Black-backed Gull Larus marinus
Black-legged Kittiwake Rissa tridactyla
Sterna hirundo
Uria aalge
Common Tern
Common Murre
Razorbill
Black Guillemot
Alca torda
Cepphus grylle
Atlantic Puffin Fratercula arctica
were three exceptions with respect to the year of the
census, and these were: Black Guillemots at Clori-
dorme to Marsoui were counted in 1980 instead of in
1979, Common Eiders at Pointe de Sandy Beach were
counted in 2003 instead of in 2002, and Northern
Gannets at Bonaventure Island were counted in 2004
instead of in 2002. The census techniques employed
varied among species and colonies but were consis-
tent for each colony among the three censuses (i.e.,
1979, 1989, and 2002). A detailed explanation of the
techniques used in 1979 and 1989 can be found in
Chapdelaine and Brousseau (1992); with few excep-
tions the same methods were used in 2002. A brief
description of the techniques used in 2002 for each
species is presented in Table 1. Certain species, such
as Black Guillemots and Herring and Great Black-
backed gulls, nest in colonies as well as solitary pairs
or in very small groups. As such, for these species the
distribution and colony size data may pertain to a sec-
tion of shoreline rather than a discrete colony. At
colonies where a species was believed to have been
present but there are no population estimate available,
a “+” is indicated in the corresponding table. To exam-
ine changes in population sizes between the three cen-
suses, that is from 1979 to 1989 and from 1989 to
2002, the compound annual growth rate (CAGR) was
calculated for each species using the formula: CAGR
= (((N(t)-N(0))/N(0)+1)!)-l where M(t) = population
size at time t, N(0) = population size at time 0 and t =
number of years between two censuses.
Methodology
systematic counts of all nests, except for llot Mahy nord
where flushed females were counted
systematic counts of active burrows
population analysis from aerial photography
nests counted from a boat, except for the Upper Plateau
of Rocher Percé where nests were counted from a look
out point using a spotting scope
nests counted from a boat
systematic counts of all nests
systematic counts of nests; ratio calculated where nested
with Great Black-backed Gulls; counts of incubating
adults on cliffs made from a boat
systematic counts of nests (sometimes from boat); ratio
calculated where nested with Herring Gull
Bonaventure Island, Cap d’Espoir, Forillon Peninsula:
counts derived from photographs (taken from boat);
elsewhere nests were counted from a boat
systematic counts of all nests
counts of birds at colonies on cliffs; used a conversion
factor of 0.76 pair/bird observed
counts of birds at colonies on cliffs
counts of birds at colonies on cliffs and on the water;
used Cairns (1979) conversion factor to estimate the
number of breeding pairs
counts of adult birds
Results
During the surveys in 1979, 1989, and 2002, a total
of 14 species of seabirds was recorded at 59 breeding
sites (colonies and/or shoreline sectors) around the
Gaspé Peninsula (Tables 1, 2). Twelve species were
observed in all three censuses, with the remaining two
absent in one of the census years (Tables 3-8). In 2002,
13 species were observed, with only the Leach’s Storm-
Petrel not recorded. In terms of total abundance (num-
ber of individuals) and breeding sites, the censuses
recorded 134 163 birds at 44 sites in 1979, 231 186
birds at 58 sites in 1989, and 279 276 birds at 57 sites
in 2002 (Table 8). In each of the three survey years,
three species accounted for over 75% of the total sea-
bird population—Northern Gannet, Black-legged Kit-
tiwake, and Common Murre. Individual species ac-
counts are as follows:
Common Eider: In 2001, the total population of
breeding eiders (subspecies dresseri) in the St.
Lawrence Estuary and Gulf was estimated at 42 000
pairs (The Joint Working Group on the Management
of the Common Eider 2004). The 2002 seabird cen-
sus of the Gaspé Peninsula recorded only 235 pairs
of this eider, which represents less than 1% of the
total population in Québec. Nevertheless, the popula-
tion is steadily increasing, from 41 pairs in 1979 to
128 pairs in 1989 to 235 pairs in 2002 (Table 8). In
2002 this seabird bred at [lot Mahy nord, Ile Plate,
and Pointe de Sandy Beach (Table 6). A few pairs are
also believed to nest each year at the foot of the cliff
on the Forillon Peninsula.
276
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE 2. Number, name, and centroid coordinates of sites on the Gaspé Peninsula surveyed for nesting seabirds in 1979,
1989, and 2002. (Colony numbers correspond to locations in Figure 1.)
Miguasha to [lot Mahy Sud sector
Colony Coordinates!
1. Miguasha to Saint-Omer 48°04'N, 66°18'W
2. Saint-Omer 48°07'N, 66°15'W
3. Banc de Carleton 48°06'N, 66°08'W
4. Ile au Pique-Nique 48°06'N, 66°07'W
5. Ile Taylor 48°09'N, 65°51'W
6. New Richmond to Bonaventure 48°09'N, 65°50'W
7. Pointe Howatson 48°08'N, 65°49'W
8. Caps Noirs 48°07'N, 65°48'W
9. Caplan 48°06'N, 65°39'W
48°0S'N, 65°37'W
48°04'N, 65°33'W
48°02'N, 65°29"'W
48°03'N, 65°28'W
10. Ruisseau Leblanc
11. Marais de Saint-Siméon
12. Bonaventure to Paspébiac
13. [lots de la riviére Bonaventure
14. Paspébiac Ouest 48°02'N, 65°17'W
15. Marais de Paspébiac 48°02'N, 65°15'W
16. Paspébiac to Shigawake 48°O1'N, 65°15'W
48°02'N, 65°13'W
48°03'N, 65°11"'W
48°03'N, 65°09'W
48°0S'N, 65°06'W
48°06'N, 65°04'W
48°06'N, 65°03'W
48°08'N, 64°59'W
48°11'N, 64°56'W
48°11'N, 64°53'W
48°11'N, 64°52'W
48°12'N, 64°49'W
48°15'N, 64°45'W
17. Pointe aux Corbeaux
18. Pointe Huntington
19. Pointe a Ritchie
20. Saint-Godefroi
21. Shigawake to Chandler
22. Shigawake
23. Colline de Port-Daniel
24. La Vieille (Pointe Pillar)
25. Gascons-Ouest
26. Pointe Reddish
27. Anse a Pierre Loiselle
28. Tlot Mahy sud
llot Mahy Nord to Forillon sector
Coordinates!
48°15'N, 64°45'W
48°20'N, 64°42'W
48°21'N, 64°41'W
48°21'N, 64°40'W
48°21'N, 64°39'W
48°23'N, 64°35'W
48°24'N, 64°29'W
48°25'N, 64°26'W
48°25'N, 64°25'W
48°25'N, 64°19"'W
48°26'N, 64°19'W
48°27'N, 64°19'W
48°29'N, 64°17'W
48°30'N, 64°13'W
48°30'N, 64°10'W
48°32'N, 64°12'W
48°32'N, 64°12'W
48°32'N, 64°13'W
48°34'N, 64°18'W
48°36'N, 64°18'W
48°38'N, 64°09'W
48°37'N, 64°10'W
Colony
29. lot Mahy nord
30. Banc de la baie du Grand Pabos
31. Barachois de Chandler
32. Ile Dupuis
33. Chandler to Grande-Riviére
34. Barachois de Pabos
35. Grande-Riviere to Cap d’ Espoir
36. Grande-Riviere
37. Sainte-Thérese-de-Gaspé
38. Cap d’Espoir to Percé
39. Cap d’Espoir
40. Cap Malin
41. Weygand
42. Cap Blanc
43. Bonaventure Island
44. Rocher Percé
45. Percé to Cannes de Roches
46. Les Trois Soeurs
47. Coin-du-Banc to Pte St-Pierre
48. Barachois de Malbaie
49. Ile Plate
50. Pointe St-Pierre to Douglastown
51. La Grande Anse 48°41'N, 64°14"'W
52. Bois-Brilé 48°46'N, 64°22'W
53. Douglastown to Sandy Beach 48°46'N, 64°22'W
54. Cap Haldimand 48°48'N, 64°23'W
55. Pointe de Sandy Beach 48°50'N, 64°24'W
56. Pointe Jacques Cartier (Gaspé) 48°51'N, 64°29'W
57. Forillon Peninsula 48°47'N, 64°12'W
58. Cloridorme to Marsoui 49°11'N, 64°50'W
59. Marsoui to Baie des Capucins 49°13'N, 66°04'W
' For shoreline sections, the coordinates of the starting locations are given.
Leach’s Storm-Petrel: On the Gaspé Peninsula,
this petrel has only ever been known to nest at two
locations, Forillon Peninsula and Bonaventure Island.
By the 1930s, the Forillon colony no longer existed
(Ball 1938), leaving only the Bonaventure Island
colony. This colony was probably never large—in the
1979 and 1989 censuses only 10 and 21 pairs were
counted, respectively. Unfortunately, no petrels (nor
active burrows) were observed in the 2002 census
(Table 7). In fact, this species’ breeding status in
Québec is currently uncertain. Historically, it nested
at several locations along the North Shore of the Gulf
of St. Lawrence and on the Magdalen Islands. Howev-
er, the only active colony found in the 1998-1999
census of North Shore Migratory Bird Sanctuaries
(Corossol Island with 718 active burrows, Rail and
Chapdelaine 2004) was deserted in the 2005 census
(Rail and Cotter 2007), and no active burrow has been
recorded on the Magdalen Islands in the last 20 years,
although observations in 2007 of adults in the [le Brion
Ecological Reserve during a survey for this species sug-
gest a small colony still exists there (CWS, unpublished
data). In some cases, including Bonaventure Island,
the presence of Red Foxes (Vulpes vulpes) might be
responsible of the abandonment of the colony.
Northern Gannet: On the Gaspé Peninsula, this
species nests only on Bonaventure Island. During the
last census the Northern Gannet was the most abundant
of all seabirds with an estimated 53 635 pairs (Table 8).
Since 1979 this gannet colony has grown at a fairly
consistent rate; the compound annual growth rate
(CAGR) was 2.8% between 1979 and 1989 and 5.5%
between 1989 and 2004 (Table 8). This colony was
also censused in 1999 and was estimated at 36 936
pairs; the CAGR for 1999 to 2004 was 7.7%. In 1999
and 2004 the colony showed no sign of density-depen-
dant effects, and because breeding success estimates
obtained in 1999 and 2004 were high and stable (CWS,
unpublished data), we think that the colony should con-
tinue to increase in the next few years.
Double-crested Cormorant: After a doubling in
population size between 1979 and 1989, the size of this
cormorant’s population on the Gaspé Peninsula did not
change substantially between 1989 and 2002 (CAGR
of only 0.4%), reaching 4 889 pairs in 2002 (Table 8).
A stabilization in populations between the mid-1980s
2007
GULF OF ST. LAWRENCE
GASPE PENINSULA
>
47° 45'N
67° 00' W
COTTER AND RAIL: SEABIRD POPULATIONS OF GASPI
77
Py NINSULA
49° 25'N
_- 28-29 30-
A 24.26 ~27
FIGURE 1. Location of colonies and shoreline sectors on the Gaspé Peninsula surveyed for breeding seabirds in 2002. See
Table 2 for location listings and coordinates.
and the early 2000s has also been recorded in Nova
Scotia (Boyne and Beukens 2004) and New Brunswick
(Boyne and Hudson 2002). In Québec, the population
on the Magdalen Islands also remained fairly stable
between 1990 and 2002 (CWS, unpublished data), as
did the overall population in breeding colonies within
the Migratory Bird Sanctuaries along the St. Lawrence
North Shore between 1993 and 2005 (Rail and Cotter
2007). Interestingly, the increase in the size of the
Gaspé population between 1979 and 1989 was accom-
panied with an additional nine new colonies in 1989,
bringing to a total of 29 active colonies that year. In
2002, however, there was a contraction in the number
of sites where it nested, as it was recorded at only 25
sites (Tables 4, 6, 7). Of the nine new colonies in
1989, three were not active in 2002. Additionally, six
colonies that were active in both 1979 and 1989 har-
boured no cormorants in 2002. The loss of these nine
colonies was offset by five new colonies, including
one comprising 393 pairs in the New Richmond-
Bonaventure region and, interestingly, a small colony
on Bonaventure Island. As Chapdelaine and Brousseau
(1992) stated, the majority of Double-crested Cor-
morant colonies on the Gaspé Peninsula are found on
sea-cliffs that in some areas are unstable and subject
to erosion. Thus it is not surprising that between cen-
suses with an interval of =10 years there are colonies
that become inactive and new ones are created.
Great Cormorant: World-wide this species has the
largest breeding distribution of all cormorant species,
but only a small fraction of its population nests in
North America (Hatch et al. 2000). In Québec there
are approximately 20 small colonies (total population
<1500 pairs) (CWS, unpublished data), with histori-
cally only one colony on the Gaspé Peninsula at
Rocher Percé. In 1979 this colony had 16 nesting pairs,
and in the 1989 census this number had doubled to
35 pairs. This upward trend continued in the 1990s as
the total Gaspé population reached 96 pairs in 2002
(Table 7). In that census, the Rocher Percé colony had
changed little in size since 1989, but a new and fairly
large colony of 50 pairs was found at Cap d’Espoir
(Table 7). Small numbers (1-3 pairs) were also observed
at four other sites in 2002 (Tables 4, 6), which along
with the new colony at Cap d’Espoir may indicate the
population is expanding not only in size but also in its
distribution. Rail and Chapdelaine (2004) also report-
ed on the recent appearance of small Great Cormorant
colonies in North Shore Migratory Bird Sanctuaries,
although the most recent survey of these sanctuaries,
in 2005, recorded only 48 Great Cormorants, a sharp
decrease from the previous survey in 1998-1999 when
342 adults were counted (Rail and Cotter 2007).
Ring-billed Gull: This gull, whose first breeding
record in the Maritime Provinces was in 1965 (Lock
1988), is also a newcomer to the Gaspé Peninsula. It
278
was absent during the 1979 census, and in 1989 only
21 pairs were counted at two sites, Barachois de
Chandler and Pointe de Sandy Beach (Table 5). In the
2002 census, however, there was only one colony and
it was not at either of those two sites. Instead it was
at Saint-Omer, and the colony was surprisingly quite
large with 1663 pairs (Table 3). There were more
Ring-bills at this site than the traditional species that
nest there, that is Herring and Great Black-backed gulls
and Common Terns. With numbers of Ring-billed Gulls
still rising in the Maritimes (Boyne and Hudson 2002;
Boyne et al. 2006) and apparently also on the North
Shore (Roberge 2004), it is not surprising to observe
the same trend around the Gaspé Peninsula.
Herring Gull: Among all seabirds on the Gaspé
Peninsula, the Herring Gull is the only species whose
population declined in size between 1979 and 1989 and
again between 1989 and 2002 (Table 8). From 1979
to 1989 its CAGR was -1.8%, and this rate of decline
doubled to -3.5% between 1989 and 2002. Between
1979 and 2002, the total population declined from
9796 to 5134 pairs. This decline is attributable to
colonies decreasing in size as opposed to a reduction
in the number of breeding sites (24 in 1979 vs 29 in
2002; Table 8). In 2002, 22 of the 29 colonies had
fewer gulls than in 1989. There were, however, four
colonies/sectors that increased in size by more than
50% between 1989 and 2002: Saint-Omer, Ile Taylor,
Marais de Paspébiac, and the New Richmond to Bona-
venture sector (Tables 3, 5). These four areas along
with [lot Mahy sud and Pointe de Sandy Beach com-
prised 64% of the total Herring Gull population on
the Gaspé Peninsula in 2002. The decline in colony
size is probably not a result of competition, such as
for nest or foraging areas, with other larids. Among
the 22 sites where the population declined between
1989 and 2002, there were only two-Ilot Mahy sud
and Ile Plate-that had a sizable increase in the popu-
lation of another seabird, and at both sites it was of
Great Black-backed Gulls (three other sites had small-
er increases in their Great Black-backed Gull popula-
tions; Tables 3, 5). Along coastal New England, how-
ever, the more aggressive Great Black-backed Gulls
have displaced Herring Gulls from certain good breed-
ing sites, and Rome and Ellis (2004) found evidence
there that Great Black-backed Gulls may suppress
foraging by Herring Gulls. Elsewhere in Canada, pop-
ulations of Herring Gulls have also declined in New-
foundland (Robertson et al. 2004), New Brunswick
(Boyne and Hudson 2002), Nova Scotia (Boyne and
Beukens 2004), and Prince Edward Island (Boyne
and McKnight 2005). Chapdelaine and Rail (1997)
showed that there was a positive relationship between
Herring Gull populations in the North Shore sanctu-
aries and cod fishery activities in the Gulf of St.
Lawrence, the latter providing substantial amounts of
fish offal and discards to breeding gulls. After a thriv-
ing period that lasted up to the end of the 1980s, the
cod fishery collapsed, a situation which probably trig-
THE CANADIAN FIELD-NATURALIST
Vol. 121
gered the decline of the Herring Gull populations on
the North Shore (-70% between 1988 and 1993; Chap-
delaine 1995) and around the Gaspé Peninsula as well
(-37% between 1989 and 2002). We think that the
disappearance of this abundant anthropogenic food
source brought Herring Gull populations back to a
more “natural” level, which will stabilize in the future,
as has been observed in North Shore sanctuaries since
the early 1990s (Rail and Cotter 2007).
Great Black-backed Gull: This large gull often
nests in mixed colonies with its more abundant rela-
tive, the Herring Gull. Unlike that species, however,
Great Black-backs increased dramatically in num-
bers on the Gaspé Peninsula between 1979 and 1989,
with a CAGR of 23.7%. After reaching a population
of 1 337 pairs in 1989, the population in 2002 was rel-
atively unchanged with 1 322 pairs. The large increase
from 1979 to 1989 was due in part to colonies increas-
ing in size but also to a doubling in the number of
colonies (Table 8). Comparing 2002 with 1989, there
were a few changes in colony locations. In 2002 this
gull did not nest at Barachois de Chandler but colo-
nized the Banc de la baie du Grand Pabos in the adja-
cent bay, less than 1 km away. It also deserted the cliffs
from Paspébiac to Shigawake, but became established
between New Richmond and Bonaventure. Elsewhere,
in each of the neighbouring Maritime Provinces Great
Black-backed Gull numbers declined between the late
1980s and early 2000s (Boyne and Beukens 2004;
Boyne and Hudson 2002; Boyne and McKnight 2005).
In the Migratory Bird Sanctuaries (excluding Gros
Mécatina and Saint-Augustin) along the St. Lawrence
North Shore, however, the population has been rela-
tively stable, with | 883 individuals counted in 1988
and | 835 individuals counted in 2005 (Rail and
Chapdelaine 2004; Rail and Cotter 2007).
Black-legged Kittiwake: This small larid was the
most abundant seabird species in both 1979 and 1989
(Table 8). The size of the kittiwake population on the
Gaspé Peninsula doubled in size in the 1980s, from
21 827 pairs in 1979 to 43 423 pairs in 1989. This
species’ distribution and abundance between 1974
and 1985 have been described in detail for the entire
Gulf of St. Lawrence by Chapdelaine and Brousseau
(1989). In the 1990s, the population stopped grow-
ing, and in 2002 there were 13% fewer pairs (37 649)
than there were in 1989. Recent downward trends (1.e.,
negative annual growth rates) have been observed in
other regions of eastern Canada: Magdalen Islands
(Québec), -1.0%/year, 1989-2000; Anticosti Island
(Québec), -2.3%/year, 1985-2004; North Shore Migra-
tory Bird Sanctuaries (Québec), -8.0%, 1988 to 1998-
1999 but followed by a stabilization with a rate of
0.64%, 1998-1999 to 2005; Gull Island (Newfound-
land), -2.2%, 1971-2001; Great Island (Newfound-
land), -9.9%, 1994-2003; and Baccalieu Island (New-
foundland), -3.6%, 1984-2003 (CWS, unpublished
data; Rail and Cotter 2007; Robertson et al. 2004; G.
Robertson, personal communication). In each of the
SEABIRD POPULATIONS OF GASPE PENINSULA
OTTER AND RAIL:
.
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Vol. 121
THE CANADIAN FIELD-NATURALIST
282
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2007 COTTER AND RAIL: SEABIRD POPULATIONS OF GASPE PENINSULA 283
three Gaspé Peninsula censuses, five colonies com-
prised 295% of the total kittiwake population: Cap
al onocoeonn > enn
a S € peed a é 0 o d’ Espoir, Bonaventure Island, Les Trois Soeurs, Rocher
E =| ve aA Percé, and Forillon Peninsula (Tables 5, 7). Compar-
3 2s) Rete ale a nak * ing colony sizes in 2002 with 1979, the former three
o e 2 is rm i colonies had changed by less than +35%, whereas
& aline ms too aa S +r the latter two had increased substantially in size, in par-
Re Bel de Ean Gh mS ticular Rocher Percé, which increased from 16 pairs
ro “ a a = in 1979 to | 094 pairs in 2002 (Table 7). In each of
2 the three censuses, the Bonaventure Island colony
2 harboured the most kittiwakes, with an estimated 18
cE” ||| ee aa 2 alent Sale whe Serr ee ee 550 pairs nesting in 2002. After the 1989 census, at
AE a + = - 3 “= which time there were 23 650 pairs at this colony,
o — ~ ay Chapdelaine and Brousseau (1992) postulated that
bb ia alle a, ME A the Bonaventure Island colony may have reached its
=| ZR oF as zl TR peak due to a lack of available nesting habitat.
é S “ = Common Tern: Of all species censused in 1979,
‘@ ee es CcuMccve®) as a 1989, and 2002, the Common Tern had the greatest
s ies a o z aS change in CAGR over the 1989 to 2002 period as
2 .. e compared with 1979 to 1989 (Table 8). In that first
5 interval, the population increased at an annual rate of
S a Terr ws =ee 18.0%, from 459 pairs in 1979 to 2 407 in 1989. From
s |3s aS 2 z SSS 1989 to 2002, however, the tern population declined
S 2 7. at a CAGR of -13.1%. The 2002 population was esti-
a | So A) Sere wey mated at only 388 pairs, which is fewer birds than
Ei BG SC AS a Sue} there were in 1979 (Table 8); only one other species,
ie) ae im the Leach’s Storm-Petrel, had a smaller population in
2 + D Beas ont 2002 than in 1979. Terns have never nested at a large
= 3 Se number of sites on the Gaspé Peninsula. In 1979 the
ef Common Tern nested only at Bane de Carleton and
% Bs Pointe de Sandy Beach, and both of these colonies had
z BQ wena & ; ON 0 Fe tn increased considerably in size by 1989 and were the
ra aa Ro) | Sane largest of all tern colonies, with each containing over
e : ve eas 800 pairs (Tables 3, 5). In addition to these two
= 5 % Ss ci om SS g iB m S ; colonies, there were four new colonies in 1989, and
6 Sai a as among these six colonies four had 200 or more nest-
2 a EN) Sree sty SA aI) | NOES == ing pairs. In 2002, however, there were only four
3S ee e g ae 3 colonies, of which three consisted of 20 or fewer pairs.
& oh Pr yee = During this census there were no terns at Banc de
= 30%, this was not the sit-
uation between 1989 and 2002. Over this period, six
of 14 species increased in population by =>70%, three
species decreased in numbers by >35%, and the pop-
ulations of five remained relatively stable with changes
of <15%. The latter group includes the Atlantic Puf-
fin, for whom numbers are too low to assess changes
in population size (Table 8).
COTTER AND RAIL: SEABIRD POPULATIONS OF GASPE PENINSULA
285
In each of the 1979, 1989, and 2002 censuses, three
Northern Gannet, Black-legged Kittiwake,
and Common Murre—accounted for over 75% of the
total population, and therefore changes in the popula-
tions of these species had the main impact on overall
seabird population trends. Interestingly, between 1989
and 2002 these three species each exhibited a differ-
ent trend; the population of the Northern Gannet in-
creased by 122%, that of the Black-legged Kittiwake
decreased by 13%, while the population of the Com-
mon Murre remained stable (increase of only 3%).
Of the different seabird groups, between 1989 and
2002 the total population of larids declined whereas
the populations of alcids and cormorants increased.
Three of the five larid species that breed on the Gaspé
Peninsula declined sharply (>35%), whereas popula-
tions of all alcid and cormorant species either remained
stable or increased. Of the remaining three individual
species, Common Eiders and Northern Gannets in-
creased in numbers, whereas the small population of
Leach’s Storm-Petrels declined (there were no active
burrows found in 2002). Rail and Chapdelaine (2004)
reported a similar pattern in the overall trends of sea-
birds along the St. Lawrence North Shore between
1993 and 1998-1999. Over that interval, they report-
ed increases in population for four species, decreases
for four species, and stable or moderate trends for the
remaining species. As we observed on the Gaspé
Peninsula in the 1990s, larids along the North Shore
declined in numbers whereas populations of alcids and
cormorants increased. It should be noted that among
larids, the Ring-billed Gull was an exception to the
downward trend; it increased considerably in numbers
in both regions, with annual growth of 31.0% (1993 to
1998-1999) and 40.0% (1989-2002) for the North
Shore sanctuaries (Rail and Chapdelaine 2004) and
the Gaspé Peninsula, respectively. The coastlines of
both regions are fronted by the Estuary and Gulf of
St. Lawrence, which breeding seabirds of both regions
use almost exclusively as their foraging grounds. It is
not surprising, therefore, that the seabird populations
of both regions show similar trends. Strong growth of
Ring-bill populations also occurred in the Maritime
Provinces, where the population grew at an annual rate
of 20.9% between 1972 and 1986 (Lock 1988), and
recent censuses in Prince Edward Island (Boyne and
McKnight 2005) and New Brunswick (Boyne et al.
2006) have shown that populations in these two prov-
inces have continued to grow.
An interesting observation from our results is the
1989-2002 trend for species that are “surface feeders”
as compared with those that are “divers”. The pro-
portion of the total seabird population made up of
divers increased from 50% in 1989 to 70% in 2002,
and conversely the proportion for surface feeders fell
from 50% to 30%. A similar pattern has occurred with
seabird populations on the Magdalen Islands. In this
archipelago, between 1989-1990 and 2000-2002 divers
increased by 20% while surface feeders declined by
20% (CWS, unpublished data). This trend seems gen-
species
286
eralized throughout the Gulf of St. Lawrence, as with-
in Migratory Bird Sanctuaries of the North Shore the
same 20% increase in the proportion of diving species
(and 20% decrease of surface feeders) can be noted
between 1988 and 2005 (CWS, unpublished data).
For the Gaspé Peninsula, the recent declines in lar-
ids, in particular among kittiwakes and terns, as well
as the possible extirpation of the Leach’s Storm-Petrel,
are major concerns for seabird biologists and mana-
gers. Factors that may be responsible for the declines,
specifically possible changes in productivity, breed-
ing conditions, predators, and/or survival, should be
studied. Despite concern for these species, the over-
all population trends of seabirds along this peninsula
are encouraging. The total number of seabirds has dou-
bled between the first comprehensive survey in 1979
and the most recent survey in 2002, with nine of the
14 breeding seabirds showing strong population growth
over that period.
Acknowledgments
We extend special thanks to our recently retired
colleague, Gilles Chapdelaine, who in addition to hav-
ing organized and carried out the field work for this
third seabird census of the Gaspé Peninsula is largely
responsible for the development in Québec of a mod-
ern seabird monitoring program. We also thank Jean-
Frangois Cloutier, personnel from the Parc national de
I’ Ile-Bonaventure-et-du-Rocher-Percé and Forillon
National Park of Canada, and Serge Brodeur for their
assistance in the surveys. John Chardine was of great
assistance with the gannet survey.
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of the Gulf of St. Lawrence, 1998-1999. Canadian Field-
Naturalist 118: 256-263.
Rail, J.-F., and R. Cotter. 2007. Sixteenth census of seabird
populations in the sanctuaries of the North Shore of the
Gulf of St. Lawrence, 2005. Canadian Field-Naturalist
121: 287-294.
Roberge, B. 2004. Plan de gestion de l’avifaune marine et
aquatique. Réserve de parc national du Canada de |’ Archi-
pel-de-Mingan. Volume 2: Atlas de l’avifaune marine et
aquatique. Parks Canada, Mingan Field Unit. 106 pages.
Robertson, G. J., S. I. Wilhelm, and P. A. Taylor. 2004.
Population size and trends of seabirds nesting on Gull and
Great Islands, Witless Bay Ecological Reserve, Newfound-
land, up to 2003. Canadian Wildlife Service Technical
Report Series Number 418, Atlantic Region. 45 pages.
Rome, M. S., and J. C. Ellis. 2004. Foraging ecology and
interactions between Herring Gulls and Great Black-
backed Gulls in New England. Waterbirds 27: 200-210.
Received 12 January 2007
Accepted 22 May 2008
Sixteenth Census of Seabird Populations in the Sanctuaries of the
North Shore of the Gulf of St. Lawrence, 2005
JEAN-FRANCOIS RAIL and RICHARD COTTER
Canadian Wildlife Service, P.O. Box 10100, Ste-Foy, Québee G1V 4HS Canada
Rail, Jean-Francois, and Richard Cotter. 2007. Sixteenth census of seabird populations in the sanctuaries of the North Shore
of the Gulf of St. Lawrence, 2005. Canadian Field-Naturalist 121(3): 287-294.
The Migratory Bird Sanctuaries of the North Shore of the Gulf of St. Lawrence are important breeding grounds for many
seabird species. The quinquennial censuses of seabirds in these sanctuaries have been carried out for 80 years (since 1925),
in order to detect changes in the distribution and population levels of seabirds breeding on the North Shore. Between 1998-
1999 and 2005, the most striking trends observed were the near doubling in numbers of Common Eiders in most sanctuaries,
and the strong population growth rate of Razorbill. Unexplainable declines of Common Murre and Atlantic Puffin were
observed at some of the larger colonies of these species. The last active colonies of Leach’s Storm-Petrel (Corossol Island)
and Caspian Tern (ile 4 la Brume) on the North Shore seem on the verge of disappearing, with zero and three birds observed
in 2005, respectively. Seabird populations in the larger, more accessible and disturbed sanctuaries are still markedly smaller
than in historical times.
Les refuges d’oiseaux migrateurs de la Céte-Nord du Golfe du Saint-Laurent constituent d’importants sites de nidification
pour plusieurs espéces d’oiseaux marins. Des inventaires quinquennaux des oiseaux marins dans ces refuges sont effectués
depuis 80 ans (depuis 1925), pour détecter les changements dans la répartition et les niveaux de population des oiseaux
marins nichant sur la Cote-Nord. Entre 1998-99 et 2005, les tendances les plus marquantes étaient chez |’ Eider 4 duvet, dont
les effectifs ont pratiquement doublé dans la plupart des refuges, et chez le Petit Pingouin, dont les populations ont égale-
ment connu une croissance rapide. Des déclins énigmatiques chez le Guillemot marmette et le Macareux moine ont été
observés a plusieurs des colonies les plus importantes de ces espéces. Les derniéres colonies actives d’ Océanite cul-blane (a
Vile du Corossol) et de Sterne caspienne (a I’ile 4 la Brume) sur la Cote-Nord semblent sur le point de disparaitre, avec
respectivement zéro et trois oiseaux observés en 2005. Les populations d’ oiseaux marins demeurent a des niveaux bien en-
de¢a de ceux rapportés historiquement dans les refuges de grande superficie, davantage accessibles et sujets au dérangement.
Key Words: Seabirds, populations, sanctuaries, Atlantic Puffin, larids, alcids, Gulf of St. Lawrence.
The Migratory Bird Sanctuaries of the North Shore
of the Gulf of St. Lawrence not only contribute to sea-
bird conservation by protecting important breeding
grounds, but through the quinquennial censuses they
serve as monitoring areas to detect changes in the dis-
tribution and population levels of 15 species of seabirds
(scientific names given in Table 1), which in turn ori-
ent future research and management.
Since the creation of the sanctuaries on the North
Shore in 1925, seabirds breeding in these sanctuaries
_have been censused approximately every five years.
The results of previous surveys have been consistently
_ published (Lewis 1925, 1931, 1937, 1942; Hewitt 1950;
’
|
Tener 1951; Lemieux 1956; Moisan 1962; Moisan and
Fyfe 1967; Nettleship and Lock 1973; Chapdelaine
1980, 1995; Chapdelaine and Brousseau 1984, 1991;
| Rail and Chapdelaine 2004). Thus the sixteenth census,
| conducted in 2005, perpetuated an 80 year-old tradition
that constitutes, over time, an invaluable database for the
_ management and conservation of seabird populations in
Québec. For some species, a very significant propor-
_ tion of their total breeding population in the province
is found in North Shore Migratory Bird Sanctuaries.
’ Furthermore, many species are represented well enough
in these sanctuaries to assume that the observed pop-
ulation trends reflect the demographic changes for the
entire North Shore. These censuses of seabird colonies
are of even greater importance as for most of our sea-
birds they are the only means to effectively monitor
population trends, because most seabird species are not
currently covered by other existing monitoring pro-
grams. Ultimately, the integrity of the whole seabird
community in the sanctuaries can also be used as a sen-
tinel to track environmental changes and to evaluate the
health of the Gulf of St. Lawrence marine ecosystem.
The goals of this article are to update population
estimates and trends for seabirds breeding in the ten
Migratory Bird Sanctuaries of the North Shore of the
Gulf of St. Lawrence, to evaluate the situation of each
species, and to identify priority issues for their con-
servation.
Methods
We used basically the same survey techniques as in
1998-1999 (see Rail and Chapdelaine 2004). The
methodology, described in detail for each species and
site in Rail and Chapdelaine (2002), is summarized
below for each family of birds.
ANATIDS — On Corossol Island, the average nest den-
sity (nests/ha) as calculated from six 60 m x 60 m
(0.36 ha) quadrats, was extrapolated to the entire area
287
288
on the island deemed suitable for Common Eiders
(Chapdelaine 1978*). In the Betchouane Bird Sanctu-
ary, we used a system of line-transects of varying length
on Innu Island and conducted a systematic nest count
elsewhere. In Watshishou and Ile 4 la Brume, we count-
ed all eider nests on the same sample of islands used
in previous censuses, which covered 12% and 27% of
the sanctuaries’ land area, respectively, and we then
extrapolated the average density over the entire area
encompassing all the islands. At the Baie des Loups
Sanctuary, a combination of methods was used: aver-
age nest density was extrapolated from two quadrats
on Ile Les Blacklands, from a sample of islands at [les
Factory, and from the closest island at Tle no 4; the
remaining islands in the sanctuary were systematically
censused. Total nest counts were conducted on most
large islands at the [les aux Perroquets and Iles Sainte-
Marie Sanctuaries, representing 57% and 96% of the
total land area, respectively; average nest densities
were extrapolated to the rest of the sanctuaries. At the
Gros Mécatina and Saint-Augustin Sanctuaries we
also searched for nests on the largest islands, but for the
smaller islands, on which we did not land, we counted
from a boat all females that flushed and assumed one
nest per female observed.
Gaviips — Red-throated Loon nests were systemati-
cally counted by walking around the ponds on islands
presenting suitable open habitat in each sanctuary. In
one instance, instead of landing on the island, we
noted the presence of a territorial adult.
HyYDROBATIDS — We systematically searched for active
burrows where colonies had been found in previous
censuses. Traditionally, a burrow was considered active
if we detected signs, such as soil freshly excavated
and/or oily odour characteristic of petrels, or when we
were able to reach an adult inside the burrow. In
2005 we also used in some instances a burrow probe
(Peep-A-Roo, Sandpiper Technologies Inc.), which
consisted of a 1.0-inch diameter video imager with
infrared illumination source, on a 3-meter long flexi-
ble cord.
PHALACROCORACIDS — Great and Double-crested
Cormorant nests were systematically either from the
ground, from a lookout point, or from a boat.
Larips — On Corossol Island, we counted the num-
ber of nests (Np) and the number of adults (Ni) in sub-
colonies, then we estimated the number of pairs in
sectors where we counted only the number of adults
present by using the factor k (k = Np/Ni). In these sec-
tors we applied the observed species ratio (Great
Black-backed / Herring Gulls) to the calculated num-
ber of nests. All gull nests were counted systemati-
cally at the Betchouane Sanctuary, and attributed to
Herring and Great Black-backed Gulls afterwards
according to the observed proportions of adults of both
species. A combination of nest counts, adult counts
and species ratios was used elsewhere for gulls and
THE CANADIAN FIELD-NATURALIST
Vol. 121
terns, except that results have been combined for Arc-
tic and Common Terns in Table 1. For the Black-legged
Kittiwake, we counted apparently occupied nests.
A.Lcips — Adult bird counts were used to estimate
Black Guillemot populations in all sanctuaries. When
colonies of Common Murres, Razorbills and Atlantic
Puffins could be visited with minimal disturbance, we
systematically counted the eggs and apparently active
burrows. For puffins, we occasionally used the Peep-
A-Roo burrow probe (described in the Hydrobatids
survey method) to determine the real occupancy rate
of apparently active burrows. At many colonies, for
example, burrows that were inaccessible in cliffs or
featuring large numbers of birds (especially murres),
we avoided disturbance by counting from a distance
all individuals at the colony and on nearby waters, us-
ing binoculars or a spotting scope. At Corossol Island
and Gros Mécatina Sanctuaries, only bird counts were
used, whereas all eggs and active burrows were noted
at the Betchouane Sanctuary. Population estimates
for the large concentrations of alcids at the Baie des
Loups, Iles aux Perroquets and [les Sainte-Marie
Sanctuaries were obtained mostly from adult counts.
For Atlantic Puffins nesting in the Baie des Loups
Sanctuary, we used the method described above for
larids, that is we calculated the factor k (number of
active burrows per individual observed at the colony)
for sectors on Ile Les Blacklands and Ile du Loup and
used it to estimate the number of pairs for puffins ob-
served elsewhere. Evenly spaced grids and line tran-
sects were used on Ile aux Perroquets in the Baie de
Brador Sanctuary, in order to (1) delimit the area of
both the puffin and Razorbill colonies, (2) evaluate
mean nest densities, and (3) extrapolate the calculat-
ed density to total colony area (see Nettleship 1973* ;
Chapdelaine 1978*). Incubating murres were count-
ed on this same island. Also, in the Baie de Brador
Sanctuary, numbers of puffins around L’Ile-Verte
were counted.
Results
The number of seabird species breeding in North
Shore sanctuaries remained at 14 between 1998-1999
and 2005, with the reappearance of the Caspian Tern
(3 adults) at the Ile 4 la Brume Sanctuary, and the ap-
parent disappearance of the Leach’s Storm-Petrel. If
we exclude results from the Saint-Augustin Sanctu-
ary (not surveyed in 1998-1999), the estimated total
number of breeding seabirds censused in 2005
(n = 107 O11 individuals) was very similar to 1998-
1999 estimate (n = 103 579; increase of only 3.3%).
However, some species showed marked population
changes (Tables | and 2). In particular, Common Eider
numbers doubled in nearly every sanctuary. The
Razorbill population also increased greatly in most
sanctuaries (overall annual growth rate of +7.3%).
On the other hand, the population of Common Mur-
res dropped by half between 1998-1999 and 2005.
289
IXTEENTH CENSUS NORTH SHORE GULF OF ST. LAWRENCE
.
.
RAIL AND COTTER
2007
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290 THE CANADIAN FIELD-NATURALIST Vol. 121
TABLE 2. Changes in the numbers of seabirds in sanctuaries on the North Shore of the Gulf of St Lawrence, Québec, 1993 to
2005.
Years of survey rate by period* Compound annual growth
Species 1993 1998-1999 2005 1993-1999** 1999-2005**
Common Eider 14548 13072 25570 -2.10% 9.98%
Red-throated Loon 66 82 92 3.21% 1.92%
Leach’s Storm-Petrel 1840 718 0) -18.38% -100.00%
Double-crested Cormorant 3472 2830 3346 -3.36% 2.82%
Great Cormorant 78 342 48 24.54% -32.713%
Ring-billed Gull 104 484 1344 28.43% 21.54%
Herring Gull 4879 4988 4812 0.58% -0.27%
Great Black-backed Gull 2284 2427 1870 1.27% -3.79%
Black-legged Kittiwake 6294 3856 3994 -9.98% 0.64%
Caspian Tern 0 0 3 - 100.00%
Common and Arctic Terns 545 394 2324 -10.99% 29.01%
Common Murre 30829 30124 14877 -0.39% -11.78%
Razorbill 8389 14341 22472 8.58% 7.32%
Black Guillemot 411 788 924 8.90% 2.45 %o
Atlantic Puffin 46684 29133 25335 -7.91% -2.34%
loge N (t) - loge N (0)
r= 100%
t
vee Data from the Saint-Augustin Sanctuary (censused only in 2005) are excluded from this table, as well as data from the Gros
Mécatina Sanctuary (not surveyed in 1993) in the 1993-1999 growth rate calculations. Populations in sanctuaries censused in
1998 were extrapolated to 1999 to derive the compound annual growth rate for all North Shore sanctuaries after 6 years.
Although smaller murre colonies grew substantially
in size, the bulk of the population, concentrated at
Iles Sainte-Marie and Iles aux Perroquets, suffered
an unexplainable and severe decline. In other res-
pects, the sharp decreases in Black-legged Kittiwake
numbers that were noted in 1993 and 1998-1999 had
finally ceased by 2005 at Corossol Island, the largest
colony on the North Shore. The 2005 census also
showed that Herring Gull populations were quite sta-
ble, whereas Great Black-backed Gulls suffered a
23% overall reduction in numbers compared with
1998-1999. All major Atlantic Puffin colonies showed
declines except for the Baie de Brador colony which
increased by 27%, and this colony thereby represents
the only stronghold of the species in our province.
Corossol Island was visited from 28 May to 30 May.
In 1998, the island held the last known colony of
Leach’s Storm-Petrel on the North Shore (359 active
burrows). Unfortunately, no nests were found in 2005.
The Double-crested Cormorant colony had 156 nests
in 2005, which is only a third of its size in 1998. Con-
versely, the estimated 980 breeding pairs of Common
Eiders in 2005 represent a 69% increase compared to
the 1998 count. We found that the rapid decline of
Black-legged Kittiwake numbers, noted during the
two preceding censuses, had finally stopped between
1998 and 2005. This species is still the most abundant
seabird on the island. Alcids in general showed clear
upward trends, in particular the Razorbill population
which increased by 170% during the 1998-2005 peri-
od. Finally, Herring and Great Black-backed Gulls
numbers were somewhat reduced in 2005, shrinking
back to the levels observed in 1993. The total number
of breeding seabirds was similar in 1998 and 2005 at
Corossol Island.
On 2 June 2005 the Betchouane Sanctuary was
surveyed and Common Eider numbers were found to
have had increased by 81% compared to 1998. The
observed density of nests was impressive as over
one-fifth (21.8%) of the eiders breeding in all North
Shore sanctuaries were concentrated in Betchouane,
the smallest sanctuary. Alcids also did well between
1998 and 2005; numbers of Razorbills and Atlantic
Puffins had risen significantly, and the 64 Common
Murre pairs discovered in 2005 were the first breed-
ing record of this species in the sanctuary. Population
levels of gulls and kittiwakes were quite stable between
1998 and 2005. The overall number of breeding sea-
birds increased by more than 60%.
When we visited the Watshishou Bird Sanctuary
on 4 and 5 June 2005, we found the Common Eider
population had literally exploded since 1998 (+132%).
The terns had bounced back with an estimated 331
individuals breeding on ten islands, after very few of
these birds were seen in 1998 (n = 38). Three of the
four Double-crested Cormorant colonies found in 1998
had grown in numbers in 2005 (+56.6% total increase).
Numbers of Herring and Great Black-backed Gulls
stayed remarkably stable during the same period. On
the other hand, Black Guillemots were less abundant
in 2005 than in 1998, and no Razorbills were seen in
2005.Population trends between 1999 and 2005 were
2007 RAIL AND COTTER: SIXTEENTH CENSUS NORTH SHORE GULF OF ST, LAWRENCE 29]
LABRADOR STOO W
f 57°00 N
y } QUEBEC
°
@ Baie de Brador
Saint-Augustin
Gros Mécatina
«
% irs : eed lles Sainte-Marie
3s hi
ile du Corossol Retchouane lles aux Perroquets
Gaspé Peninsula .
Baie des Loups
ile a la Brume
Gulf of St. Lawrence
NEWFOUNDLAND
Magdalen Islands
NEW BRUNSWICK
50 km
FIGURE |. Location of the sanctuaries of the North Shore of the Gulf of St. Lawrence.
minor for most species breeding at the [le 4 la Brume
Sanctuary. Even Common Eiders were stable here,
whereas spectacular increases were noted for this spe-
cies in other sanctuaries. The Caspian Tern did a timid
comeback with three (presumably) breeding individ-
uals seen in 2005. This species had not been recorded
on the two previous quinquennial censuses. Converse-
ly, for the first time since the beginning of the quinquen-
nial censuses in 1925, no Razorbills were seen in this
sanctuary in 2005. Only Herring and Great Black-
backed gulls exhibited marked positive trends; their
small populations grew more than twofold between
1999 and 2005. The Ile 4 la Brume Sanctuary was vis-
ited on 9 and 10 June.
The most striking result for the Baie des Loups Sanc-
tuary (visited on 11, 14 and 19 June) was the extreme-
ly low number of Atlantic Puffins observed at Ile Les
Blacklands, an island which has always been well
known for its large puffin colony. Data analysis con-
firmed the catastrophic puffin decline (-79%) between
1999 and 2005 in this sanctuary. Since 1960, the Baie
des Loups Sanctuary had never held less than 25% of
the total puffin population breeding in North Shore
sanctuaries, until this proportion dropped to 7% in
2005. Black Guillemots also had suffered a sharp dec-
line (-75%). Razorbill numbers were similar in 1999
and 2005, in contrast with the general upward trend
observed elsewhere for this species during the same
period. On the bright side, the small Common Murre
population had become firmly established in the sanc-
tuary in 2005 with a near fivefold increase since 1999,
and with the appearance of a second colony. Mean-
while, the population of Common Eiders had also risen
substantially (+87%).
At the Iles aux Perroquets Sanctuary, the total num-
ber of breeding alcids stayed relatively stable between
1999 and 2005, but this is because the Razorbills’ large
population increase (+172%) compensated for the
declines in Common Murres (-56%), Atlantic Puffins
(-61%) and Black Guillemots (-39%). The Common
Eider population grew by +80%. Meanwhile, Herring
and Great Black-backed Gull numbers were nearly
stable. Only one of the seven Great Cormorant nests
found in 1999 was still active in 2005. Black-legged
Kittiwakes seemed also on the verge of disappearing
from Iles aux Perroquets, with seven nests left in 2005.
The sanctuary was visited on 13 and 22 June.
The Iles Sainte-Marie Sanctuary comprises large is-
lands that hold important concentrations of seabirds,
and six days were devoted to their census (15, 17, 18,
and 20 to 22 June) in 2005. Alcids showed moderate
population trends except for the major drop in Com-
mon Murre numbers (-54%), the most abundant sea-
bird in the sanctuary. There were more than twice as
many eider nests as in 1999. Between 1999 and 2005,
the Black-legged Kittiwake colony at Ile Cliff grew un-
expectedly (+184%), but other gull populations con-
tinued to dwindle in the sanctuary. Such low numbers
of breeding Great Black-backed and Herring Gulls had
not been seen there since 1950 and 1925, respectively.
The two species of cormorants showed opposite trends.
With an 86% drop between 1999 and 2005, the size of
the Great Cormorant colony in 2005 (23 nests) was the
smallest ever recorded since its establishment in the
early 1930s, whereas the Double-crested Cormorant
colony increased to 650 nests (+62%).
The Gros Mécatina Sanctuary was surveyed in two
stages: the [les aux Marmettes and Ile Plate were vis-
ited on 23 June, whereas Ile aux Trois Collines could
not be accessed before 7 July. This was only the second
quinquennial survey of the Gros Mécatina Sanctuary,
which was created in 1996. In 2005, two species had
disappeared (Great Cormorant, Black-legged Kittiwake)
but were replaced by two others (Double-crested Cor-
morant, Common Eider). The most significant change
was in the tern colony at Ile Plate which grew from
50 to nearly 900 pairs between 1999 and 2005. Terns
now comprise two-thirds of all breeding seabirds in the
sanctuary. Also, the small number of Common Mur-
res observed had increased fivefold. Considering the
small size of the sanctuary, the growth (+75%
between 1999 and 2005) and size (n = 240 in 2005)
of the Black Guillemot population were impressive.
Our visit to the Saint-Augustin Sanctuary on 25 June
2005 was the first in 17 years. The last time was at the
end of the 1980s, when Herring Gull populations
were thriving, and just before a widespread decline
was observed on the North Shore. The 2005 census
confirmed that this trend also occurred at the Saint-
Augustin Sanctuary, as Herring Gull numbers were
down 82% compared with 1988. The Great Black-
backed Gull population also suffered a 73% decline
during the same period. Alcids, represented by a mere
four (4) Black Guillemots in 2005, did not show any
signs of recovery. Common Eiders, on the other hand,
made a noticeable comeback as they were more than
ten times more abundant than in 1988. However, con-
sidering that the Saint-Augustin Sanctuary has the
largest land surface area of all North Shore sanctuar-
ies and the fact that between 1935 and 1960 a mini-
mum of five hundred pairs of eiders were recorded in
any one census, the estimate of 73 pairs for 2005
looks far from the potential of these islands. The
terns, though, were well represented in 2005, with
numerous (n = 13) colonies and a total population of
nearly a thousand birds, the largest ever recorded in
this sanctuary.
The largest concentration of breeding Atlantic
Puffins in the province of Québec has always been
located in the Baie de Brador Sanctuary. Fortunately,
our results show a 27% increase of that colony between
1999 and 2005, in contrast with the declines observed
at other major puffin colonies on the North Shore. Baie
de Brador now shelters 79% of all puffins breeding in
North Shore sanctuaries. During the same period, the
Razorbill population mean annual growth rate was
high (6.9%) but similar to elsewhere, and numbers of
Common Murres went up by an amazing 50% annual-
THE CANADIAN FIELD-NATURALIST
Vol. 121
ly. In other respects, the colony of Black-legged Kit-
tiwakes discovered in 1999 had already disappeared
by 2005. We landed on Ile aux Perroquets on 30 June
and 1 July, whereas L’Ile-Verte could not be reached
before 12 August.
Discussion
Here we present a short analysis of the latest popu-
lation trend and current status of each species in the
North Shore sanctuaries, in order of species in best
health (population) to worst. The Common Eider has
to come first on the list. Its population has increased
dramatically since 1982, and the near-doubling in num-
bers between 1998-1999 and 2005 represents a very
strong population growth rate. The Common Eider is
now the most abundant breeding seabird in North Shore
sanctuaries. The Razorbill is another species that has
been increasing in numbers for more than 20 years in
the sanctuaries (as well as everywhere in Eastern
North America, Chapdelaine et al. 2001), and there is
no indication that this upward trend will slow. Fur-
thermore, many new colony sites were discovered
during this period. The size of the Red-throated Loon
population was larger than ever in 2005, particularly
due to the high numbers reported in the Iles Sainte-
Marie and Iles aux Perroquets sanctuaries. The addi-
tion of the latter sanctuary in 1982 helped to protect a
significant proportion of this species’ small popula-
tion of only a few hundred individuals spread along
the northern coasts of the Gulf of St. Lawrence
(DesGranges and Laporte 1979*; Barr et al. 2000;
Chapdelaine and Rail 2004). The Black Guillemot has
also fared well in the sanctuaries, an exception being
at Ile 4 la Brume and Baie des Loups, where breeding
numbers have been much higher in the past. Important
for the species is the recently created Gros Mécatina
Sanctuary (1996); this sanctuary encompasses less
than 3% of the total land surface area of North Shore
sanctuaries yet it harboured one-fourth of the Black
Guillemot population in 2005. The Double-crested
Cormorant, the Common and Arctic Tern, and the
Ring-billed Gull were species very well represented
in North Shore sanctuaries in 2005. However, the
trends observed with these species in the sanctuaries
may not be representative for the whole North Shore.
In fact, the few colonies of Double-crested Cormorants
in the sanctuaries show variable and sometimes con-
trasting trends. And colonies of terns and Ring-billed
Gulls are highly unpredictable, too variable in size and
localization (in and out of the sanctuaries from one
year to another) to derive meaningful trends.
Herring Gull population trends also tended to vary
between sanctuaries, but as a whole the North Shore
population has been quite stable since the crash that
occurred between 1988 and 1993 (which was associ-
ated with the collapse of the cod fishery, see Chapde-
laine and Rail 1997). In 2005 the species was well
established in all sanctuaries. Great Black-backed
2007
Gulls were also well represented in every sanctuary,
and leaving aside the 43% decline at Corossol Island,
elsewhere the population was stable between 1998-
1999 and 2005. Another species showing relative sta-
bility in numbers between the last two quinquennial
censuses is the Black-legged Kittiwake. At Corossol
Island, the only large colony in North Shore Sanctuar-
ies, the sharp declining trend (-58%) observed between
1988 and 1998 had ceased by 2005. Because of this
colony’s importance for the species on the North Shore
and evidence of very poor reproductive success in 2005
and 2006 (CWS, unpublished data), it is imperative that
careful monitoring of the colony occurs in the future.
Examination of their diet suggests that when staple food
(capelin, sandlance) becomes scarce in their diet, the
larger gulls (Herring and Great Black-backed gulls)
are more likely to prey on kittiwake chicks.
The Common Murre population had expanded so
much between 1972 and the 1990s that it was disap-
pointing to see this recent setback (51% decline
between 1998-1999 and 2005) which brought the
population back to the level observed in 1982. The
Common Murre is still the fourth most abundant bird
in the sanctuaries, but the underlying factors responsi-
ble for the substantial population decline at the
largest colonies are not known and will need to be
investigated if the species declines in number any
further. The Iles Sainte-Marie holds the only well-
established Great Cormorant colony in North Shore
sanctuaries. The number of nests, which once peaked
at 339 (in 1955), has been highly variable since 1993.
We can only hope that numbers will bounce back
after the lowest number of nests ever recorded there
was in 2005, otherwise the species may well disappear
from the sanctuaries in the near future. The Atlantic
Puffin is not on the verge of vanishing from the sanc-
tuaries; however, its situation is worrying, as the larg-
er colonies located at Baie des Loups, Iles aux Perro-
quets, Iles Sainte-Marie and Baie de Brador declined
by 47% between 1993 and 2005. The 79% decline in
six years (1999 to 2005) at Baie des Loups is particu-
larly mysterious, and the colony at the Baie de
Brador sanctuary, currently the species’ stronghold in
the province of Québec, appears vulnerable to distur-
bance, poaching and bycatch in fishing nets. The fact
that three Caspian Terns were observed in 2005 in the
Ile a la Brume Sanctuary, the only active breeding site
in the province, confirms that a small number of indi-
viduals of this species continues to frequent the area.
We suspect that a few pairs may still breed there spo-
radically, but it is doubtful that such a tiny population
can persist for long if no recovery plan is implement-
ed (but see Shaffer et al. 2004). The first active bur-
rows of Leach’s Storm-Petrels were discovered in
1972 in the Baie des Loups Sanctuary. In the census-
es that followed, the species was found in three other
sanctuaries, with a total of close to 900 burrows in
1982 and 1988. Inexplicably, storm-petrels vanished
RAIL AND COTTER: SIXTEENTH CENSUS NORTH SHORE GULF OF ST. LAWRENCI
293
as fast as they appeared and in 2005 not a single nest
ing site was found at the last active colony, Ile du
Corossol. We have no reason to believe that the
(unknown) factors responsible for the loss of this
species will be reversed in the future.
We identified sanctuaries that seemed in worse
shape by comparing the 2005 seabird numbers with
historical data in each sanctuary. Some current sea-
bird populations estimates are markedly lower than
in historical times. For example, the Ile 4 la Brume
Sanctuary once protected a fair number of alcids, in-
cluding up to 2000 pairs of Common Murrtes (in 1935).
These species declined and never recovered, and in
2005 only a few pairs of Black Guillemot were still
breeding in the sanctuary. Alcid populations had also
been much larger in the past at the Baie des Loups
Sanctuary. At the Saint-Augustin Sanctuary, popula-
tion levels of Common Eider, Black Guillemot and
Red-throated Loon seem far from the potential showed
in the 1940s. Moreover, while the Ile 4 la Brume and
Saint-Augustin Sanctuaries together represent 40% of
the total land area of the ten North Shore sanctuaries,
only 4.1% of the seabirds were found there. Of course,
seabird populations also reflect habitat features, and
these two sanctuaries may not have habitats such as
cliffs and deep crevices that are most suitable for kitti-
wakes and murres. But even the Common Eider, which
in 2005 was abundant in similar habitat in Watshishou,
was rather rare in the Ile a la Brume and Saint-Augustin
sanctuaries as they harboured only 2.2% of the total
North Shore sanctuaries population. Besides, we saw
evidence of eider egg collection in the Saint-Augustin
sanctuary in 2005. In these sanctuaries that are com-
prised of a large number of islands and are easily
accessed by boat from nearby villages, the poor den-
sity of breeding birds is likely the result of a lack of
law enforcement and public awareness the vulnera-
bility of seabird breeding colonies. In addition to im-
provements in law enforcement and educational pro-
grams in local communities, priorities for seabird
conservation in North Shore sanctuaries should in-
clude population studies to identify the underlying
causes of the unexplained declines in Atlantic Puffins
and Common Murres and a restoration project for the
Caspian Tern colony at Ile 4 la Brume.
Acknowledgments
Many thanks to everyone who contributed to this
census. First, our special thanks go to the Gallienne
family for their hospitality on Corossol Island. Field-
work was carried out with the help of our colleagues
Pierre Brousseau, Renaude Bender, Jocelyn Thibault,
Raphaél Lavoie and Olivier Meyer, as well as Cana-
dian Wildlife Service game wardens Wilson Evans and
Mathieu Nadeau-Monger. Members of Parks Canada
in Havre Saint-Pierre, Louis Lalo and many volunteers
also gave us a big hand at the Betchouane and Wat-
shishou sanctuaries. Dean and Peggy Martin helped us
294
to renew the traditional survey of the Saint-Augustin
Sanctuary, which their late father, Canadian Wildlife
Service game warden Londus Martin, had patrolled for
so many years. We would also like to acknowledge
Trish Nash and Tess Cecil-Cockwell from the Quebec-
Labrador Foundation for their efforts at Baie de Brador
and Saint-Augustin sanctuaries, as well as boatmen
Lawrence Jones and his son Edward Jones at Blanc-
Sablon.
Documents Cited (marked * in text)
Chapdelaine, G. 1978. Onziéme inventaire des oiseaux colo-
niaux des refuges de la cote nord du Golfe Saint-Laurent
et révision globale de la fluctuation des populations depuis
1925 jusqu’a 1977. Service canadien de la faune, région
du Québec, Environnement Canada, Sainte-Foy. 166 pages.
DesGranges, J.-L., and P. Laporte. 1979. Preliminary con-
siderations on the status of loons (Gaviidae) in Quebec.
Unpublished report prepared for the waterfowl working
group of the Joint Committee of the James Bay and North-
ern Quebec Agreement, Canada.
Nettleship, D. N. 1973. Census of seabirds in the sanctuaries
of the North Shore of the Gulf of St. Lawrence, summer
1972. Studies on northern seabirds No 20. Canadian
Wildlife Service Report, Ottawa. 160 pages.
Literature Cited
Barr, J. F., C. Eberl, and J. W. McIntyre. 2000. Red-
throated Loon (Gavia stellata) in The Birds of North
America, No. 513. Edited by A. Poole and F. Gill. The
Birds of North America, Inc., Philadelphia, Pennsylvania.
Chapdelaine, G. 1980. Onziéme inventaire et analyse des
fluctuations des populations d’oiseaux marins dans les
refuges de la Cote Nord du Golfe Saint-Laurent. Canadian
Field-Naturalist 94: 34-42.
Chapdelaine, G. 1995. Fourteenth census of seabird popu-
lations in the sanctuaries of the North Shore of the Gulf
of St. Lawrence, 1993. Canadian Field-Naturalist 109: 220-
226.
Chapdelaine, G., and P. Brousseau. 1984. Douziéme inven-
taire des populations d’ oiseaux marins dans les refuges de
la Céte-Nord du golfe du Saint-Laurent. Canadian Field-
Naturalist 98: 178-183.
Chapdelaine, G., and P. Brousseau. 1991. Thirteenth cen-
sus of seabird populations in the sanctuaries of the North
Shore of the Gulf of St. Lawrence, 1982-1988. Canadian
Field-Naturalist 105: 60-66.
Chapdelaine, G., A. W. Diamond, R. Elliot, and G. J.
Robertson. 2001. Status and population trends of the
Razorbill in eastern North America. Canadian Wildlife
Service Occasional Paper 105. Ottawa.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Chapdelaine, G., and J.-F. Rail. 1997. Relationship between
cod fishery activities and the population of herring gulls
on the North Shore of the Gulf of St Lawrence, Québec,
Canada. ICES Journal of Marine Science 54: 708-713.
Chapdelaine, G., and J.-F. Rail. 2004. Québec’s Waterbird
Conservation Plan. Migratory Bird Division, Canadian
Wildlife Service, Québec Region, Environment Canada,
Sainte-Foy, Québec. 99 pages.
Hewitt, O. H. 1950. Fifth census of non-passerine birds in
the bird sanctuaries of the North Shore of the Gulf of St.
Lawrence. Canadian Field-Naturalist 64: 73-76.
Lemieux, L. 1956. Seventh census of nonpasserine birds in
the bird sanctuaries of the North Shore of the Gulf of St.
Lawrence. Canadian Field-Naturalist 70: 183-185.
Lewis, H. F. 1925. The new bird sanctuaries in the Gulf of
St. Lawrence. Canadian Field-Naturalist 39: 177-179.
Lewis, H. F. 1931. Five years’ progress in the bird sanctuaries
of the North Shore of the Gulf of St. Lawrence. Canadian
Field-Naturalist 45: 73-78.
Lewis, H. F. 1937. A decade of progress in the bird sanctu-
aries of the North Shore of the Gulf of St. Lawrence.
Canadian Field-Naturalist 51: 51-55.
Lewis, H. F. 1942. Fourth census of non-passerine birds in
the bird sanctuaries of the North Shore of the Gulf of St.
Lawrence. Canadian Field-Naturalist 56: 5-8.
Moisan, G. 1962. Eighth census of non-passerine birds in
the bird sanctuaries of the North Shore of the Gulf of St.
Lawrence. Canadian Field-Naturalist 76: 78-82.
Moisan, G., and R. W. Fyfe. 1967. Ninth census of non-
passerine birds in the sanctuaries of the North Shore of
the Gulf of St. Lawrence. Canadian Field-Naturalist 81:
67-70.
Nettleship, D. N., and A. R. Lock. 1973. Tenth census of
seabirds in the sanctuaries of the North Shore of the Gulf
of St. Lawrence. Canadian Field-Naturalist 87: 395-402.
Rail, J.-F., and G. Chapdelaine. 2002. Quinziéme inventaire
des oiseaux marins dans les refuges de la C6te-Nord : tech-
niques et résultats détaillés. Série de rapports techniques
No. 392. Service canadien de la faune, région du Québec,
Environnement Canada, Sainte-Foy. xvi + 307 pages.
Rail, J.-F., and G. Chapdelaine. 2004. Fifteenth census of
seabird populations in the sanctuaries of the North Shore
of the Gulf of St. Lawrence, 1998-99. Canadian Field-
Naturalist 118: 256-263.
Shaffer, F., M. Robert, J.-F. Rail, and V. Létourneau. 2004.
La Sterne caspienne (Sterna caspia) au Québec: bilan des
connaissances. Série de rapports techniques No. 415.
Service canadien de la faune, région du Québec, Environ-
nement Canada, Sainte-Foy, x + 65 pages.
Tener, J. S. 1951. Sixth census of non-passerine birds in the
bird sanctuaries of the North Shore of the Gulf of St.
Lawrence. Canadian Field-Naturalist 65: 65-68.
Received 12 January 2007
Accepted 22 May 2008
Discovery of a Possibly Relict Outbreeding Morphotype of
Sparrow’s-egg Lady’s-slipper Orchid, Cypripedium passerinum,
in Southwestern Yukon
Pau M. CATLING! and BRUCE A. BENNETT?
' Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Wm. Saunders Building, Central Experimental Farm,
Ottawa, Ontario, Canada K1A 0C6 e-mail: catlingp @agr.ge.ca
*Department of Environment, Government of the Yukon, Box 2703, Whitehorse, Yukon, Canada Y1A 2C6 e-mail: Bruce
Bennett @ gov.yk.ca
Catling, Paul M., and Bruce A. Bennett. 2007. Discovery a possibly relict outbreeding morphotype of Sparrow’s-egg Lady’s-
slipper Orchid, Cypripedium passerinum, in southwestern Yukon. Canadian Field-Naturalist 121(3); 295-298.
An outbreeding morphotype of Cypripedium passerinum with separate anthers and stigma was discovered in the Beringian
region of southwestern Yukon. The occurrence of this breeding system in Beringia is attributed to the persistent advantage of
outbreeding over hundreds of thousands of years in this unglaciated area, whereas the widespread occurrence of exclusively
self-pollinating races elsewhere in Canada is attributed to advantages in colonization of recently deglaciated territory which
first became available less than 10 000 years ago, and/or to a changing and more forested habitat.
Key Words: Sparrow’s-egg Lady’s-slipper Orchid, Cypripedium passerinum, Orchidaceae, autogamy, self-pollination, unglaciat-
ed, Beringia, Kluane, Yukon.
In fresh flowers of Sparrow-egg Lady’s-slipper
Orchid (Cypripedium passerinum Richards.), the lat-
eral parts of the stigma are in contact with the anther
allowing self-pollination (Figure 1A). Self-pollina-
tion is evident by a mass of pollen tubes connecting
the expanded anther tightly to the stigmatic surface. It
occurs when the flowers are still in bud or before they
are fully open and germination of pollen prevents sub-
sequent removal by pollinators resulting in little chance
for cross-pollination. This obligate self-pollination is
widespread (Catling 1983; Catling, personal observa-
tion) in the geographic range, which extends from
‘Quebec west across the boreal forest and montane
region of Canada to Alaska (Sheviak 2002; Figure 2).
‘Plants that do not self-pollinate have not been
observed prior to the observations reported here.
An examination of long-faded and shrivelled flow-
ers, clearly past peak flowering, collected in alcohol
on 13 July 2006 from the Alsek River trail (60.7748°N,
-137.6939°W, approximately 10 km west of Haines
Junction) in the Kluane Range, St. Elias Mountains of
southwestern Yukon, revealed that self-pollination had
“not occurred in 7 of 8 flowers removed from a clump of
30. In most of these flowers self-pollination was pre-
‘vented because the anthers were clearly separated from
the stigmatic surface (Figures 1B-C) but in one flower
they were touching but without self-pollination (Figure
1D) and the anthers were bowed so that most of the
length of the suture was not in contact with the stig-
matic surface. In one of these flowers self-pollination
“had occurred apparently due to contact of anther and
‘stigma (Figure 1E). All eight of the flowers from this
clump were presumably from one individual and sug-
gest variation in self-pollination at the individual! level.
A voucher specimen (Catling & Bennett, 13 July 2006)
from the Alsek River location has been placed in the
National Collection (acronym — DAO) of Agriculture
and Agri-Food Canada on the Central Experimental
Farm in Ottawa.
Ten long-faded flowers collected from several indi-
vidual plants south of Whitehorse at Cowley Creek
(60.5908°N, 134.8953°W) were self-pollinated (Fig-
ures 1F-H) and so were three flowers collected from a
location near Mayo (not illustrated). In one of the flow-
ers from Whitehorse an anther was sufficiently sepa-
rated that self-pollination had not occurred (Figure
LF). Although only the distal tip of the anther suture
appeared to contact the stigmatic surface in most speci-
mens, in one flower (Figure 1G) the proximal part had
also come in contact, with the result that pollen tubes
entered the stigma from both anther tips. Self-pollina-
tion in orchids is frequently partial or facultative (e.g.,
Catling 1990). In western Yukon there may be substan-
tial variation in column morphology leading to varia-
tion in amounts of self-pollination, both within indi-
viduals and between plants. This variation may have
provided the basis for the selection of obligate self-
pollinating morphotypes that are now widespread.
Selection for self-pollination may have favoured a
number of characteristics including: (1) more extensive
development of the lateral parts of the stigmatic sur-
face; (2) less divergence of the anther stalks; (3) less
elongation and curvature of the column resulting in less
displacement of the anthers above the stigmatic sur-
face; (4) increased bowing of the anthers to bring the
tips of the suture into contact with the stigmatic sur-
295
296
THE CANADIAN FIELD-NATURALIST
Vol. 121
5mm
FicureE |. Columns of Cypripedium passerinum viewed from below showing anthers (a), pollinia (p), stigmatic surface (s)
and suture of anther (su). A, camera lucida drawing of flower from Hattie Cove, Lake Superior, Ontario (modified
from Catling 1983); B-E flowers from Alsek trail, Kluane Range, Yukon; F-H, flowers from south of Whitehorse,
Yukon. Columns A and E-H demonstrate self-pollination with one or both anthers connected to the stigmatic surface
by a mass of pollen tubes. Columns B-D have not self-pollinated.
face; and (5) increased secretion of stigmatic fluid
which would connect the anther and the stigma.
Regardless of how the flowers have changed to pro-
mote self-pollination, both the separation of anther
and stigma, and the lack of self-pollination in old
flowers are novel observations suggesting outbreeding
in a species that was hitherto known only to obligately
self-pollinate.
The structure related to breeding system in C. pas-
serinum is difficult to discern in reconstituted pressed
herbarium specimens and softening the few flowers
on a specimen to evaluate it results in damage to the
specimen. As a consequence, an investigation of
herbarium material was not conducted. The examina-
tion of fresh plants from at least 20 localities (Catling,
personal observation) across Canada, including Ontario
(3 locations), Manitoba (1 location), British Colum-
bia (8 locations), Alberta (2 locations) and Northwest
Territories (6 locations) is believed to reliably indicate
the predominance of the obligately self-pollinating
morphotype outside of the Beringian region (Figure 2).
Nevertheless, more research on the geographical occur-
rence and related morphological and anatomical as-
pects is needed to better understand the evolution of
self-pollination in this species, and to provide a basis
for consideration of formal taxonomic recognition.
Self-pollination originates through evolution by nat-
ural selection from cross-pollinated plants (e.g., Catling
1990). In general, self-pollination in orchids is more
frequent in cooler and more extreme environments
such as recently glaciated territory (Catling 1983,
1990). This is believed to be a consequence of strong
selection for colonizing ability, the selfing plants being
able to set seed in the absence of others of their kind
and/or in the absence of pollinators and being able to
mass produce adapted genotypes. This would have
been advantageous in colonizing huge areas of land
(most of northern North America) exposed following
melting of the Wisconsin ice sheets less than 10 000
years ago (Dyke 2005). It may also be advantageous
within Beringia as a consequence of the change from
pollinator-rich willow (Salix spp.) scrub and tundra to
2007
Cypripedium passerinum
in northwestern
North America
CATLING AND BENNETT: LADY’ S-SLIPPER ORCHID IN YUKON
297
1000 K veler
FiGure 2. Distribution of Cypripedium passerinum in northwestern North America (dots) and location of plants with non-auto-
pollinating column structure on Alsek River (circle). The unglaciated region of Beringia is darkly shaded (redrawn from
Dyke 2005). The dot distribution of C. passerinum was copied from Cody (1996), Hultén (1968), Packer (1959),
Porsild and Cody (1980), and Szezawinski (1959).
a pollinator-depauperate boreal forest that occurred
between 9000 and 5000 ka kiloannum BP (Dyke 2005).
As a result of the strong selection, self-pollinating mor-
photypes are presumed to have evolved from cross-pol-
linated ancestors in unglaciated territory, and these new
selfers spread over the newly available or recently
changed landscape. Many self-pollinating orchids have
a non-self-pollinating relative that has a more restricted
range in a region of more temperate climate and/or
longer established habitat. Examples in North Ameri-
ca include self-pollinating Liparis loeselii which is
widespread north of the range of the non-self-pollinat-
ing Liparis liliifolia, and self-pollinating Spiranthes
ovalis var. erostellata which is widespread north of the
more restricted distribution of Spiranthes ovalis vat.
ovalis (see Catling 1990 for other examples).
The occurrence of a non-self-pollinating race of
Cypripedium passerinum in the northwestern part of
the range rather than the warmer southern or south-
eastern part of its range is at first contrary to expecta-
tion. However, in this example a relict outbreeding
morphotype may have existed for hundreds of thou-
sands of years in the unglaciated Beringian area of
Alaska and western Yukon (Figure 2 and see Scudder
1997). It may have been from this source that mor-
photypes more committed to self-pollination spread
southward and eastward following deglaciation 10 000
years ago. The present distribution of Cypripedium
passerinum fits what Hultén refers to as “North Amer-
ican boreal plants radiating from Beringia.” (Hultén
1937, see also Cody 1971 and Ritchie 2003). It is
possible that the outbreeding ancestral morphotype
of Cypripedium passerinum is a Beringian relict. In
Beringian habitats that have existed for hundreds of
thousands of years, and presumably include pollina-
tors, cross-pollination was likely advantageous. Cypri-
pedium passerinum presently occurs in Alnus crispa
or Betula glandulosa shrubland habitats that are be-
lieved to have existed in Beringia during full glacial
times (e.g., Hopkins 1982; Ritchie and Cwynar 1982:
Cwynar 1990; Cwynar and Spear 1991; Lacourse and
Gajewski 2000) when the rest of Canada was covered
by ice up to a few km deep. Cypripedium passerinum
also occurs in open aspen (Populus) and spruce
(Picea) woodland. Both of these species evidently
existed in isolated valley-bottom habitats within the
Beringian steppe plateaus, although Populus pollen
does not preserve well making its widespread but
isolated occurrences more difficult to detect (Zazula
et al. 2006; Zazula, personal communication). Cypri-
pedium passerinum also occurs in parts of the north,
to 68°N, where Beringian species are predominant.
As it spread from Beringia across the rest of northern
Canada into new, recently deglaciated territory, there
298
was likely a strong selection for self-pollination, this
explaining the apparent predominance of self-polli-
nating morphotypes.
Literature Cited
Catling, P. M. 1983. Autogamy in eastern Canadian Orchi-
daceae: a review of current knowledge and some new
observations. Le Naturaliste canadien 110: 37-53.
Catling, P. M. 1990. Auto-pollination in the Orchidaceae.
Pages 121-158 in Orchid Biology, reviews and perspec-
tives V. Edited by J. Arditti. Timber Press, Oregon. 451
pages.
Cody, W. J. 1971. A phytogeographic study of the floras of
the Continental Northwest Territories and Yukon. Le Natu-
raliste canadien 98: 145-158.
Cody, W. J. 1996. Flora of the Yukon Territory. NRC Research
Press, Ottawa, Ontario. 643 pages.
Cwynar, L. C. 1990. A late quarternary vegetation history
from Lily Lake, Chilkat Peninsula, southeast Alaska. Cana-
dian Journal of Botany 68: 1106-1112.
Cwynar, L. C., and R. W. Spear. 1991. Reversion of forest
to tundra in the central Yukon. Ecology 72: 202-212.
Dyke, A. S. 2005. Late quaternary vegetation history of north-
ern North America based on pollen, macrofossil, and fau-
nal remains. Géographie physique et Quaternaire 59: 211-
162.
Hopkins, D. M. 1982. Aspects of the paleogeography of
Beringia during the late Pleistocene. Pages 3-28 in Paleo-
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thews Jr., C. E. Schweger, and S. B. Young. Academic
Press, New York, New York.
Hultén, E. 1937. Outline of the history of Arctic and Boreal
biota during the Quartenary Period: their evolution during
and after the glacial period as indicated by the Equiformal
Progressive Areas of Present Plant Species. Bokforlags
Aktiebolaget Thule, Stockholm. 167 pages.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Hultén, E. 1968. Flora of Alaska and neighbouring districts.
Stanford University Press, Stanford, California. 1008 pages.
Lacourse, T., and K. Gajewski. 2000. Late Quaternary vege-
tation history of Sulphur Lake, southwest Yukon Territory,
Canada. Arctic 53: 27-35.
Packer, J. G. 1959. Flora of Alberta, 2nd edition. University
of Toronto Press, Toronto, Ontario. 687 pages.
Porsild, A. E., and W. J. Cody. 1980. Vascular plants of con-
tinental Northwest Territories. National Museum of Nat-
ural Sciences, Ottawa, Canada. 667 pages.
Ritchie, J. C. 2003. Postglacial vegetation of Canada. Cam-
bridge University Press, Cambridge, UK. 178 pages.
Ritchie, J. C., and L. C. Cwynar. 1982. The late Quaternary
vegetation of the North Yukon. Pages 113-126 in Paleo-
ecology of Beringia. Edited by D. M. Hopkins, J. V.
Matthews Jr., C. E. Schweger, and S. B. Young. Academ-
ic Press, New York, New York.
Scudder, G. G. E. 1997. Environment of the Yukon. Pages
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J. A. Downes. Biological Survey of Canada Monograph
Series (2): 1034 pages.
Sheviak, C. J. 2002. 1. Cypripedium Linnaeus. Pages 499-
507 in Flora of North America volume 26. Edited by Flora
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object_id=8891 &flora_id=1.
Szczawinski, A. 1959. Orchids of British Columbia. British
Columbia Provincial Museum Handbook (16): 1-124.
Zazula, G. D., A. M. Telka, C. R. Harington, C. E.
Schweger, and R. W. Mathewes. 2006. New spruce (Picea
spp.) macrofossils from Yukon Territory: implications for
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391-400.
Received 22 January 2007
Accepted 14 July 2008
Twenty-Four-Hour Activity Budgets of Mule Deer, Odocoileus
hemionus, in the Aspen Parkland of Eastcentral Alberta
GERALD W. Kuzyk!? and Ropert J. HUDSON!
'Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G 2H! Canada
*Present address: Ministry of Environment, Wildlife Science Section, P.O. Box 9338, Victoria, British Columbia V8W 9M]
Canada; e-mail: Gerald. Kuzyk @ gov.be.ca
Kuzyk, Gerald W., and Robert J. Hudson. 2007. Twenty-four-hour activity budgets of Mule Deer, Odocoileus hemionus, in the
aspen parkland of eastcentral Alberta. Canadian Field-Naturalist 121(3): 299-302.
We documented seven 24-h activity budgets of two adult Mule Deer (Odocoileus hemionus) does without fawns in the Aspen
Parkland of Alberta when the vegetation was green (June, July), maturing (September) and cured (October). They spent about
40% (mean = 42.3%) of each day foraging, somewhat more when vegetation was green (40-47%) and maturing (45%), and
less time in October, when the vegetation was cured (38-39%). The percentage of time Mule Deer were bedded each day
generally equaled or slightly exceeded (33-53%) that of foraging. Walking was a prevalent activity (3-24%), especially in
autumn and little time was invested in other activities such as social interactions and grooming (1-4%). There were generally
more feeding bouts (10-16/day) than bedding bouts (4-10/day) per day, and feeding bouts were shorter (37-68 minutes) than
bedding bouts (69-133 minutes). Mule Deer displayed polycyclic feeding cycles with slight increases in feeding activity near
dawn and dusk.
Key Words: Mule Deer, Odocoileus hemionus, activity, foraging, Alberta.
Mule Deer (Odocoileus hemionus) range over much
of western North America, yet little is known about their
foraging ecology on northern ranges (Kuzyk 2008),
especially regarding their daily activities. Document-
ing how much time Mule Deer spend feeding during
different stages of vegetation phenology provides in-
sight into behavioral adaptations in seasonal northern
environments. Browsing ruminants normally spend
~ about 50% each day foraging, with feeding times vary-
ing with forage phenology, morphological and physio-
logical characteristics such as mouth shape, gut capac-
ity and food passage rate (du Toit and Yetman 2005).
_ Activity budgets can be used to quantify daily foraging
times and other major activities. One difficulty in con-
_ ducting activity budgets is remaining close enough to
a focal animal to accurately quantify activity patterns,
especially during darkness. Using tame deer is one way
to maintain consistent visual contact with a focal ani-
' mal for a 24-h period (Collins and Urness 1983) and
} provides reliable results, as food intake in tame deer
_ is deemed innate (Spalinger et al. 1997). Maintaining
, visual contact with tame deer also allows daily feeding
[ cycles and duration of feeding bouts to be quantified.
_ The purpose of this study was to quantify activity
t budgets of Mule Deer on northern ranges when the veg-
| etation was green (June and July), maturing (Septem-
_ ber) and cured (October). From allometric considera-
‘ tions (Mysterud 1998; du Toit and Yetman 2005), we
; predicted that Mule Deer would spend 50% of their
- daily activity budget foraging and foraging time would
_ change with forage phenology. We predicted that Mule
Deer would spend least time foraging in summer when
vegetation was green and the most time in autumn when
the vegetation had cured because required selectivity
increases search time.
Methods
Our study animals were two unbred adult Mule Deer
does which were hand-reared as fawns. The animals
were four and six years old and weighed 59 and 60 kg,
respectively, at the beginning of the study. These ani-
mals would allow researchers to remain within five
meters of them, apparently without disturbing their nor-
mal activities. The deer were released into 3-7 hectare
fenced enclosures (2.2 m high) in an Aspen Parkland
habitat (Strong 1992) at the University of Alberta
Ranch (53°N, 111°31'W) approximately 150 kilome-
ters southeast of Edmonton, Alberta. Each deer was
instrumented with a radiocollar (Telonics TGW-3570)
and placed in a paddock with two other Mule Deer as
part of a larger study (Kuzyk 2008). This study was
approved by the University of Alberta Faculty Animal
Policy and Welfare Committee (Protocol HUDS
2004—33B), following guidelines of the Canadian Coun-
cil on Animal Care.
Scan sampling (Altmann 1974) was conducted on
focal deer throughout one complete 24-h period and
repeated seven times during the summer and autumn
of 2004 and 2005 (Table 1). One Mule Deer (501H)
was used for six of the sampling periods and was sub-
stituted for another Mule Deer (515K) to compare be-
haviors at the same season but different years (Table 1).
Only one 24-h activity budget was conducted per sam-
pling period. The activity budget was classed into pre-
dominant activities and recorded every 10 minutes (Col-
lins and Urness 1983). One observer closely followed
299
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Activity budgets (24-h) for one Mule Deer doe (501H) at the University of Alberta Ranch, Kinsella, Alberta.
Date Feeding Bedding Walking Other
% day hrs % day hrs % day hrs % day hrs
2005 June 9 40.8 9.8 50.0 12 6.3 il-5) 229 0.7
2004 = July 13 41.7 10 47.9 11.5 6.3 1S) 4.2 1
2005 July 7 47.1 11.3 47.1 Hil 3.3} 0.8 2.1 0.5
2004 September 11 45 10.8 32.5 7.8 DCT 52 0.8 0.2
2005 September 9 44.6 10.7 38.3 92 15.8 3.8 1.3) 0.3
2004 October 8 38.8 9.3 36.7 8.8 23.8 S)o// 0.8 0.2
2005* October 12 38.3 9.2 52.9 Wa Tol c9/ Doll 0.5
* Note: Different Mule Deer (515K).
the deer and used a stopwatch to time predominant
activities within a 10-minute bout. An activity had to
occur for 5 minutes or longer within a 10-minute time
interval to be classed as predominant. Predominant
activities were feeding, bedding, walking and other
(social interactions, grooming). Feeding was defined
as ingesting, chewing or intently searching (smelling
plants) for food, while walking was defined as a deer
moving without biting vegetation. Ruminating was
associated with bedding. Each observer spent between
6 and 10 hours with the Mule Deer and overlapped
shifts by approximately 15 minutes to minimize dis-
turbance to the deer. At night, a flashlight was used to
monitor the Mule Deer by concentrating the light beam
on the posterior portion of the body to minimize dis-
turbance.
Daily patterns of feeding and bedding were deter-
mined by the number and duration of bouts that were
separated by at least 10 minutes (Pépin et al. 2006).
Daily feeding cycles were evaluated by the proportion
of time spent feeding each hour over a 24-hr period.
The proportion of time spent feeding at dawn and dusk
was determined by the mean amount of time spent
feeding 1-3 hours before, | hour before and | hour after
and 1-3 hours after sunrise and sunset (Colman et al.
2001). Descriptive analyses of activities are presented
due to insufficient sample sizes for statistical compar-
isons.
Results
Mule Deer spent about 40% of each day foraging
with slight increases when the vegetation was green
and maturing (40-47%) and less time when the vege-
tation was cured (38-39%). Mule Deer spent between
33-53% of each day bedded (Table 1). Feeding bouts
were generally shorter (37-68 min) than bedding bouts
(69-133 min) and there were more feeding bouts (10-
16/day) than bedding bouts (4-10/day) per day (Table
2). Feeding cycles were polycyclic (Figure 1) with in-
creased feeding activity near sunrise and sunset (Fig-
ure 2).
Discussion
Our research documented and compared daily for-
aging activities of Mule Deer on a northern range when
the vegetation was at three phenological states, and
deer were at successive stages of their annual growth
cycle. Our findings did not meet our prediction that
Mule Deer would spend less time foraging in summer
than in autumn. We assumed abundant forage quantity
and quality would account for reduced foraging activ-
ity in summer, but early summer is the period when
weight gain and appetite are greatest, despite the ab-
sence of milk production for fawns. Possibly Mule
Deer were being very selective in forage quality in the
summer period and spent more time consuming the
TABLE 2. Number and duration (mean min/bout + SE) of feeding and bedding bouts for one Mule Deer doe (501H) during 24-h
periods at the University of Alberta Ranch, Kinsella, Alberta.
Date Feeding
Bouts/24-hr Min/bout
2005 9 June 16 BOOED ES
2004 13 July 15 40.0 + 8.7
2005 7 July 10 68.0 + 10.6
2004 11 September 12 54.2+11.5
2005 9 September 1] S32) s8 ILI ,3)
2004 8 October 13 43.1 +9.0
2005* 12 October 12 45.8 +9.4
Bedding
Bouts/24-hr Min/bout
10 72.0 + 16.5
10 69 + 11.6
6 SES
4 117.5 + 29.3
5 110 + 27.6
4 132.5 + 41.7
7 108.6 + 11.0
* Note: Different Mule Deer (515K).
2007 KUZYK and HUDSON: ACTIVITY BUDGETS OF MULE DEER )]
o- = a Po "Ssq
N= Oe Zor oy = ce
QO. @G>, 'O@ t639 Go = 2c
4
>
>
-
o
.
. 4
2
aa
e
Proportion of Time Feeding
4
Sa
2
=
~)
LS
SP OPA Pt PHP H
OK a iw: ©: d° © Ps
oy Denilss
OS™ AS” av
hr
Osu
XV Ah r ©
FicureE 1. Feeding cycles of adult Mule Deer does in the Aspen Parkland of east-central Alberta. Data are averaged and
pooled for season (June, July, September, October) and years (2004-2005).
0.90
0.80
Oo
~
ro)
—¢e— June/July
—«— September
—@#— October
o 9°
es uw (op)
(o) oO (@)
o 9°
W
io)
Proportion of Time Feeding
Oo
in
oO
0.10
0.00
Before Sunrise After Before Sunset After
Sunrise Sunrise Sunset Sunset
FIGURE 2. Proportion of time a Mule Deer doe spent feeding 1-3 hours before. | hour before and after, and 1-3 hours after
sunrise and sunset in the Aspen Parkland of east-central Alberta. Data are averaged and pooled for season and years
(2004-2005).
302
easily digestible forbs when they were readily avail-
able. In September, Mule Deer may have increased for-
aging times as the vegetation started to senesce and
deer spent more time searching for green vegetation,
especially forbs beneath the grass layer.
Mule Deer spent the least time foraging in October
when the vegetation had cured. This may be due to
Mule Deer adjusting their bite sizes (Kuzyk and Hud-
son 2006) to compensate for the lack of green vegeta-
tion. Mule Deer on northern ranges have high con-
sumption rates (g/min) in October and can forage at
rates that are near theoretical maximum by feeding on
Canadian thistle (Cirsium arvense) (Kuzyk and Hud-
son 2006). Increased bite sizes in October could lead
to a quicker gut fill compared to other seasons.
The percentage of time Mule Deer bedded each
day equaled or exceeded that of the time spent forag-
ing. High diurnal temperatures can reduce Mule Deer
activity (Hayes and Krausman 1993), especially during
midday (Ager et al. 2003), and could account for the
difference. Beir and McCullough (1990) found that
White-tailed Deer (Odocoileus virginianus) were rela-
tively inactive in summer when feeding on abundant
forage. Long bedding bouts may reflect requirements
for rumination of coarse forage.
There was variation in behaviors between the two
Mule Deer in October in different years. Time spent
foraging was almost identical, whereas the time spent
bedding contrasted greatly as the individual used in
multiple trials (501H) spent 36.7% of the day bedding
compared to 52.9% for the other individual (515K).
Also, 501H spent substantially more time walking
(24%) than did 515K (7%), which may be due to indi-
vidual variations in behaviors.
Mule Deer had polycyclic feeding cycles which has
been determined for some other northern ungulates
(Colman et al. 2001). The cycles may be driven by
rumen fill and time required for digestion (Pérez-Bar-
berfa and Gordon 1999), as these Mule Deer exper-
ienced little disturbance and few intra-specific interac-
tions (Kuzyk and Hudson 2007). Mule Deer increased
feeding activity near sunrise and sunset which is sim-
ilar to the behavior of White-tailed Deer (Beir and
McCullough 1990), as crepuscular activity is a com-
mon behavior for small ruminants to reduce predation
risk (Pérez-Barberia and Gordon 1999). There appeared
to be a more pronounced feeding peak at sunrise, pos-
sibly due to rapid daytime heating in summer and fall
or increased insect harassment. Additional studies quan-
tifying activity budgets of Mule Deer would help clar-
ify determining factors of daily feeding cycles.
Acknowledgments
This study was supported by a strategic research
grant through the Natural Sciences and Engineering
Research Council of Canada and a University of Alber-
THE CANADIAN FIELD-NATURALIST
Vol. 121
ta Graduate Research Assistantship. We appreciate the
field assistance of P. DeWitt, K. Kuzyk, A. Lockwood,
N. Maleki, and S. Vinge and thank B. Irving and staff
at the University of Alberta Ranch (Kinsella) for ani-
mal handling.
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Altmann, J. 1974. Observational study of behaviour — sam-
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Beir, P., and D. R. McCullough. 1990. Factors influencing
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Monographs 109: 1-51.
Collins, W. B., and P. J. Urness. 1983. Feeding behavior and
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Colman, J. E., C. Pedersen, D. @. Hjermann, @. Holand,
S. R. Moe, and E. Reimers. 2001. Twenty-four-hour feed-
ing and lying patterns of wild reindeer Rangifer tarandus
tarandus in summer. Canadian Journal of Zoology 79:
2168-2175.
du Toit, J. T., and C. A. Yetman. 2005. Effects of body size
on the diurnal activity budgets of African browsing rumi-
nants. Oecologia 143: 317-325.
Hayes, C. L., and P. R. Krausman. 1993. Nocturnal activity
of female desert mule deer. Journal of Wildlife Manage-
ment 57: 897-904.
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lates in central Alberta. PhD thesis. University of Alber-
ta, Edmonton, Alberta.
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ers to estimate forage intake of mule deer. Canadian Jour-
nal of Zoology 84: 1576-1583.
Kuzyk, G. W., and R. J. Hudson. 2007. Animal-unit equiv-
alence of bison, wapiti and mule deer in the aspen parkland
of Alberta. Canadian Journal of Zoology 85: 767-773.
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gia 113: 442-446.
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logia 120: 193-197
Spalinger, D. E., S. M. Cooper, D. J. Martin, and L. A.
Shipley. 1997. Is social learning an important influence on
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Received 5 February 2007
Accepted 16 July 2008
Cavity Nest Materials of Northern Flying Squirrels, Glaucomys
sabrinus, and North American Red
Squirrels, Tamiasciurus hudsonicus,
in a Secondary Hardwood Forest of Southern Ontario
Jesse E. H. PATTERSON!?, STEPHEN J. PATTERSON?2, and Jay R. MALCOLM!
'Faculty of Forestry, University of Toronto, 33 Willcocks Street, Toronto, Ontario, MSS 3B3 Canada
*Hummingbird Services, 251 Queen Street South, Suite 523, Streetsville, Ontario, LSM 1L7 Canada
‘Present address: Department of Biological Sciences, University of Calgary, 2500 University Drive, Calgary, Alberta T2N IN4
Canada
Patterson, Jesse E. H., Stephen J. Patterson, and Jay R. Malcolm. 2007. Cavity nest materials of Northern Flying Squirrels,
Glaucomys sabrinus, and North American Red Squirrels, Tamiasciurus hudsonicus, in a secondary hardwood forest of
southern Ontario. Canadian Field-Naturalist 121(3): 303-307.
Through deployment of artificial nest boxes, we examined the composition of cavity nest materials used by Northern Flying
Squirrels (Glaucomys sabrinus) and North American Red Squirrels (Tamiasciurus hudsonicus) in a secondary hardwood forest of
southern Ontario, Canada. We collected 32 nests of known species association and found that 85.7% of G. sabrinus nests and
77.8% of T: hudsonicus nests were constructed almost entirely of shredded bark from Eastern White Cedar (Thuja occidentalis).
Mean nest depth across all samples was 12.2 cm and showed no significant difference between species or between spring and sum-
mer nests. We review the antiparasitic properties of 7: occidentalis and suggest that the use of shredded cedar bark by G. sabrinus
and 7: hudsonicus to line nest cavities may be a behavioural adaptation, which reduces ectoparasite loads in the nest environment.
Key Words: Northern Flying Squirrel, Glaucomys sabrinus, North American Red Squirrel, Tamiasciurus hudsonicus, ectoparasites,
nest box, nest material, nest-protection hypothesis, thermoregulation, Eastern White Cedar, Thuja occidentalis, Ontario.
Tree cavities are a critical resource for many tree
squirrels in forest ecosystems, providing sites for rais-
ing young, avoiding predators, and improving thermo-
regulation (Collias 1964; Weigl 1978; Wiebe 2001).
Tree cavities are generally most abundant in old growth
forests where large diameter trees and well-decayed
snags are commonplace (Holloway and Malcolm 2006;
Holloway and Malcolm 2007). Cavities also may form
in live trees where the entry of rot has been facilitated
_ by disease, deformities, broken limbs or woodpecker
excavations. Where tree cavities are not readily avail-
able, many tree squirrels use external nests (dreys) or
_ subterranean nests, although evidence is emerging that
tree cavities are preferentially selected when available
(Bakker and Hastings 2002). Nest materials used in
Cavity nest construction may not only act to increase
ambient temperature in the nest environment and pro-
vide comfort for nest occupants, but may also reduce
ectoparasite loads and repel moisture in these dark,
_ damp microclimates (Stapp et al. 1991; Hemmes et al.
2002.) However, despite their importance, tree cavities
‘and cavity nests are rarely incorporated into studies of
' tree squirrels (Fokidis and Risch 2005).
| The nocturnal Northern Flying Squirrel (Glaucomys
_ sabrinus) and the diurnal North American Red Squirrel
- (Tamiasciurus hudsonicus) are both occasional cavity
nesting, arboreal rodents common in temperate and
_ boreal forests of North America. Both species are pri-
' marily associated with conifer-dominated forests, al-
_ though in many areas of eastern North America they
inhabit hardwood and mixed hardwood-conifer forests
(Flyger and Gates 1982; Holloway and Malcolm 2006;
Patterson 2008). Both G. sabrinus and T. hudsonicus
have been found to use cavities in trees more inten-
sively than external or subterranean shelters, especially
for natal nests (Layne 1954; Bakker and Hastings
2002; Holloway and Malcolm 2007).
Nest materials used by arboreal squirrels have been
found to vary geographically (Muul 1974). However,
most reports describing nest materials employed by
squirrels have been qualitative in nature and have fo-
cused on dreys. Tamiasciurus hudsonicus dreys have
been noted to include grape bark, deciduous leaves,
dried grasses, moss, feathers, fur and soft inner bark
(Layne 1954). Similarly, dreys of G. sabrinus have been
found to consist of dried grasses, shredded bark, moss-
es, twigs and lichens (Rust 1946; Mowrey and Zasada
1984). Cavity nests have been described quantitatively
only once for G. sabrinus (Hayward and Rosentreter
1994). Using nest boxes, these authors found that G.
sabrinus in the northern Rocky Mountains of central
Idaho and western Montana primarily constructed cay-
ity nests from arboreal lichens, especially of the gen-
era Bryoria and Letharia. We are not aware of studies
that have quantified cavity nest materials of 7. hud-
sonicus; however, Hayward and Rosentreter (1994)
suggest a preference for grasses and shredded bark.
Published data on G. sabrinus and T: hudsonicus nest
materials, and associated seasonal differences and
depths, in the Great Lakes-St. Lawrence forest region
of eastern North America do not exist. In general, the
study of nest material use and function in mammals has
303
received very little attention by researchers and thus
warrants investigation.
In order to gain a more complete understanding of
the habitat requirements for G. sabrinus and T: hud-
sonicus in northern secondary hardwood forests, we
characterize cavity nest materials used by these two
species in a large network of nest boxes in southern
Ontario, Canada. We hypothesized that G. sabrinus
and T. hudsonicus would preferentially select nest mate-
rials that favoured cavity nest occupation (1.e., parasite
reduction, moisture repellency). Nest depth measure-
ments were investigated as a function of the type of
nest material and season (spring and summer); we also
assessed seasonal variation in nest material composi-
tion. We predicted that nest depths would exhibit inter-
seasonal variability if nest construction was primarily
to enhance thermoregulation, such that deeper nests
would be constructed under colder temperature re-
gimes. Given the prevalence of shredded cedar bark in
the nests of both G. sabrinus and T: hudsonicus, we
discuss the potential antiparasitic and thermoregulatory
benefits of Eastern White Cedar (Thuja occidentalis)
bark.
Study Area
The study was conducted in Bruce and Grey coun-
ties, Ontario, Canada (43°59' to 45°12'N, 80°22' to
81°39'W), which together encompass an area of about
22 000 km?. Nest boxes were placed in trees growing at
various elevations, ranging from approximately 200 m
to 450 m above sea level. Approximately 22% of the
landscape is currently composed of closed-canopy sec-
ondary forests, which are dominated by Sugar Maple
(Acer saccharum), White Ash (Fraxinus americana)
and American Beech (Fagus grandifolia). Other com-
mon tree species include Eastern White Cedar, East-
ern Hemlock (Tsuga canadensis), aspen (Populus tre-
muloides, P. grandidentata) and White Birch (Betula
papyrifera) (Patterson 2008). Small stands of Eastern
White Pine (Pinus strobus) and spruce (Picea glauca,
P. mariana, P. rubens) occur throughout the area, often
in plantations.
Materials and Methods
A total of 266 nest boxes were installed between
2002 and 2004 as part of a larger study being conduct-
ed by SJP on G. sabrinus. Sampling effort varied among
years due to logistical constraints. In 2002, 154 boxes
were installed; an additional 36 boxes were installed
in 2003 and the final 76 boxes were installed in 2004.
All nest boxes were placed at heights of 3-4.5 m on
trees that had a diameter at breast height >17 cm.
Following Carey (2002), nest boxes were established
at a density of three nest boxes/ha to allow for a bal-
ance between occupancy rates and monitoring effort.
Three or more boxes were placed at each of 70 sites
across the study region.
Boxes were constructed following Sonenshine et
al. (1973), except for the door which was top mounted
THE CANADIAN FIELD-NATURALIST
Vol. 121
and could be flipped up to remove any animals and
nest material. Our entrance hole was also of larger
diameter (4 cm) to permit use by G. sabrinus and T.
hudsonicus, but exclude Eastern Grey Squirrels (Sci-
urus carolinensis). All boxes were mounted to the tree
trunk and any tree branches were removed from below
the nest box.
All boxes were intensively checked twice per year
during the day: once in the spring and once again in
autumn. Spring checks were conducted between 12
May and 12 June following the onset of parturition,
whereas autumn checks were conducted between 12
October and 12 November. Cursory visual nest box
inspections were conducted monthly to monitor the
condition of the boxes, but no sampling was done on
these occasions. These visual inspections indicated that
neither G. sabrinus nor T. hudsonicus used the nest
boxes in late autumn and winter (early December —
late February), presumably because individuals aban-
doned the nest boxes for larger, better insulated natu-
ral cavities or subnivean nests. Therefore, nests found
at the spring inspection had been constructed between
early March and the date of inspection in spring (here-
after termed “spring” nests), whereas those found at
the autumn inspection had been constructed between
the spring and autumn inspection dates (hereafter
termed “summer” nests). Where nests were present and
occupied at time of inspection, we recorded the loca-
tion, number and reproductive condition of occupants
and nest depth. Samples of nest material were col-
lected from all nests for identification of materials. In
cases where the nest box was not occupied at the time
of inspection, the entire nest was removed. In cases
where the nest box was occupied at the time of inspec-
tion (14% of the nests), a small sample of nest material
was initially removed and the entire nest was subse-
quently removed during targeted monthly nest box
examinations following the departure of nest occu-
pants. We removed nest material not only to examine
nest materials, but also because Hayward and Rosen-
treter (1994) observed that nest boxes were typically
not occupied in successive years if used nest material
was present. Nest depth was always determined during
the initial investigation and measured in situ along the
vertical dimension from the internal base of the nest
box to the top of the nest. Nest volume was deter-
mined by multiplying the nest depth by the internal
width and length dimensions of the nest box.
Results
Nest Box Occupation
Between 2002 and 2005, we found 224 occupied
and unoccupied nests. Of these, 32 were occupied when
checked (14 G. sabrinus and 18 T. hudsonicus). Fe-
males were found occupying the nest boxes in 12 cases
for G. sabrinus and 14 cases for T: hudsonicus. Lone
males were occasionally found occupying the nest
boxes (two G. sabrinus and four T. hudsonicus).
2007
Nest Materials of G. sabrinus and T. hudsonicus
Twelve of the 14 boxes occupied by G. sabrinus and
14 of the 18 boxes occupied by 7. hudsonicus con-
tained >90% shredded cedar bark by volume. One G.
sabrinus nest was composed entirely of fibreglass insu-
lation and another was composed of >80% shredded
deciduous bark. Trace amounts of peat moss, dried
grasses, cedar leaves and twigs also were found in
occupied G. sabrinus nests. Two T. hudsonicus nests
were composed entirely of dried grasses and two nests
contained >80% deciduous leaves. Trace amounts of
shredded plastic, bird feathers, animal fur, dried grass-
es and deciduous leaves also were found in occupied
T. hudsonicus nests.
Nest materials did not appear to differ between sea-
sons (t = -1.892, P = 0.06), with shredded cedar bark
appearing as the most prominent nest material in both
spring (84.5%) and summer (79.3%). Small sample
sizes for male nests did not allow for robust compar-
isons between male and female nest materials.
Nest Depth
Mean depth of nests occupied by G. sabrinus was
12.1 cm (SD = 2.4, range = 6.7 to 15.3 cm) and for T.
hudsonicus was 12.3 cm (SD = 2.1, range = 9.1 to
17.8 cm), although the difference was not statistically
different (t = 0.31, P = 0.76). We found no statistical dif-
ference between spring (G. sabrinus mean = 11.9 cm;
T. hudsonicus mean = 12.6 cm) and summer (G. sabri-
nus mean = 11.1 cm; T: hudsonicus mean = 12.7 cm)
nest depths for either species for occupied nests (G.
sabrinus: t = 0.55, P = 0.30; T. hudsonicus: t = 0.17,
P = 0.43). Small sample sizes for male nests did not
allow for robust comparisons between male and female
nest depths.
Discussion
The prominence of cedar bark in G. sabrinus and
T: hudsonicus nests may be explained by at least two,
nonmutually exclusive, hypotheses: nest-protection and
thermoregulation. The nest-protection hypothesis has
received much attention from ornithologists to explain
the use of green vegetation in avian nests. Initially sug-
gested by Wimberger (1984) and tested by Clark and
Mason (1985), the nest-protection hypothesis posits
that animals exploit the antiparasitic properties of cer-
tain plant species that emit specific volatile compounds
(Dawson 2004). All plants contain secondary metabo-
lites that are used as a defense against disease and
herbivory (Clark and Mason 1988) and, when these
plants are used as nesting materials, certain com-
pounds may reduce ectoparasite loads in the nest envi-
ronment. Numerous studies have supported the nest-
protection hypothesis in birds (Wimberger 1984:
Rodgers et al. 1988; Fauth et al. 1991; Lafuma et al.
2001). The nest protection hypothesis has found sup-
port on at least one occasion for small mammals:
Hemmes et al. (2002) reported that Dusky-footed
Wood Rats (Neotoma fuscipes) placed California Bay
PATTERSON, PATTERSON, and MALCOLM: CAVITY NEST MATERIALS OF SQUIRRELS 305
(Umbellularia californica) \eaves around their nest
sites. Laboratory tests revealed that when incubated
with torn U. californica leaves for 72 h, survival of flea
larvae was reduced by 74% compared to controls. By
using shredded Eastern White Cedar bark as a primary
nest material, G. sabrinus and T. hudsonicus also may
be limiting exposure to nest-borne ectoparasites. Phy-
tochemical analysis of 7 occidentalis bark has revealed
22 volatile compounds, including monoterpenes, fen-
chene, camphene, camphor, carvacol and paracymene
(Shaw 1953; Witte et al. 1983; Yatagai et al. 1985;
Keita et al. 2001), several of which have noted anti-
parasitic or insecticidal properties (Adams et al. 1988;
Adams 1993; Keita et al. 2001).
Support for the nest protection hypothesis is bol-
stered by the use of lichens in nests of G. sabrinus
found in western North America. Antiparasitic effects
have been described for several species of lichen. Hay-
ward and Rosentreter (1994) found that G. sabrinus
selected lichens containing the secondary compounds
norstictic acid, fumarprotocetraric acid, vulpinic acid
and atranorin, all of which have been attributed with
varying degrees of antiparasitic or antimicrobial effects
(Giez et al. 1994; Tay et al. 2004; Yilmaz et al. 2004).
These various studies support the possibility that the
use of 7: occidentalis, as well as several lichen species,
by G. sabrinus and T. hudsonicus may be a behavy-
ioural adaptation for ectoparasite control in the nest
environment. Further testing of the antiparasitic effects
of 7: occidentalis volatile oils and secondary com-
pounds on known G. sabrinus and T. hudsonicus ecto-
parasites is required.
An alternative hypothesis is that shredded cedar bark
may offer greater insulative properties than other avail-
able materials. Hayward and Rosentreter (1994) sug-
gested the same hypothesis when characterizing the
primary use of lichens as nest material by G. sabri-
nus. This hypothesis is based on the finding by Stapp
et al. (1991) that nests composed of plant fibres allowed
Southern Flying Squirrels (Glaucomys volans) to
reduce their energy expenditure when experimentally
subjected to cold temperatures. We are not aware of
studies on the thermal properties of cedar bark, lichens
or other nest materials found in this study. A common
assumption is that nest materials with greater thermal
properties are favoured, and/or that nest depth would
peak, during colder periods. Contrastingly, we found
that shredded cedar bark was used equally in spring
and summer and that nest depth did not vary between
the two seasons. This suggests that cedar bark may not
be chosen solely for thermoregulatory purposes: how-
ever, we are unable to reject the thermoregulation
hypothesis outright without further empirical testing.
We dismiss a third potential hypothesis that squirrels
choose the most widely and readily available materi-
als for use as nest substrate. Patterson (2008) found
that deciduous trees accounted for approximately 80%
of the tree composition in our study area by basal area,
306
while Eastern White Cedar accounted for less than
10% of the total forest composition on average, by
basal area. Deciduous leaves and deciduous tree bark
are therefore more greatly available than 7: occiden-
talis bark in our study area. At some sites where cedar
bark nest material was found to occur, Eastern White
Cedar trees accounted for only 2% or less of the total
tree composition (Patterson 2008). On one occasion
during this study we observed a single female G.
sabrinus travel over 500 m to the nearest 7: occiden-
talis tree to acquire cedar bark for nest construction.
Mosses, lichens, and grasses are also widely available
throughout the study area (J. Patterson, unpublished
data) and yet were rarely found as nest materials in
this study.
Acknowledgments
We thank the Grey Sauble Conservation Authority,
the Saugeen Valley Conservation Authority and the
various private landowners who graciously made their
land available for this study. We also would like to
thank Thistlewood Timber Frame Homes in Markdale,
Ontario, for generously donating the lumber used for
the nest boxes. J. Bowman, D. Nagorsen and two anony-
mous reviewers provided helpful suggestions for the
improvement of this manuscript. We appreciate the
support and field assistance provided by H. Maddin
and M. Patterson. This project was funded, in part, by
the Natural Sciences and Engineering Research Coun-
cil of Canada Undergraduate Student Research Award
to JEHP.
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Received 12 February 2007
Accepted 10 April 2008
Depredation of Common Eider, Somateria mollissima, Nests on a
Central Beaufort Sea Barrier Island: A Case Where No One Wins
JoHN A. REED!, DEBORAH L. LAcRorx2, and PAUL L. FLINT!
'U.S. Geological Survey, Alaska Science Center, 1011 East Tudor Road, Anchorage, Alaska 99503 USA. Corresponding author
e-mail: paul_flint@usgs.gov
Centre for Wildlife Ecology, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia VOR 1B0 Canada
Reed, John A., Deborah L. Lacroix, and Paul L. Flint. 2007. Depredation of Common Eider, Somateria mollissima, nests on a
central Beaufort Sea barrier island: a case where no one wins. Canadian Field-Naturalist 121(3): 308-312.
Along the central Beaufort Sea, Pacific Common Eiders (Somateria mollissima v-nigra) nest on unvegetated, barrier islands;
often near nesting Glaucous Gulls (Larus hyperboreus). Nest-site choice likely reflects a strategy of predator avoidance: nest-
ing on islands to avoid mammalian predators and near territorial gulls to avoid other avian predators. We observed a nesting
colony of Common Eiders from first nest initiation through nesting termination on Egg Island near Prudhoe Bay, Alaska (2002
— 2003). Resident gulls depredated many eider nests, mostly during initiation. All nests failed when an Arctic Fox (Alopex
lagopus) visited the island and flushed hens from their nests, exposing the eggs to depredation by the fox and gulls (resident
and non-resident). Common Eiders actively defended nests from gulls, but not from foxes. Likely all three species (1.e., eiders,
gulls, and foxes) ultimately achieved negligible benefit from their nest-site selection or predatory activity: (a) island nesting
provided no safety from mammalian predators for eiders or gulls, (b) for Common Eiders, nesting near gulls increased egg
loss, (c) for Glaucous Gulls, nesting near colonial eiders may have reduced nest success by attracting the fox, and (d) for
Arctic Foxes, the depredation was of questionable value, as most eggs were cached and probably not recoverable (due to
damage from fall storms). Thus, the predator-prey interactions we observed appear to be a case where little or no fitness
advantage was realized by any of the species involved.
Key Words: Arctic Fox, Alopex lagopus, Common Eider, Somateria mollissima, Glaucous Gull, Larus hyperboreus, nest
depredation, predator-prey interaction, Beaufort Sea, Alaska.
Along the central Beaufort Sea, nest success of Pacif-
ic Common Eiders (Somateria mollissima v-nigra) 1s
highly variable, with egg depredation by Glaucous Gulls
(Larus hyperboreus) and Arctic Foxes (Alopex lagopus)
being the primary cause of nest failure (Johnson and
Herter 1989; Johnson 2000). Common Eider nest-site
choice on coastal barrier islands is thought to reflect a
strategy of predation avoidance. Typically, Common
Eiders nest in colonies, often within the territory of nest-
ing gulls (Noel et al. 2005). By nesting on islands,
Common Eiders may avoid mammalian predators such
as foxes (Johnson 2000; Noel et al. 2005). By nesting
near gulls, eiders may benefit from territorial defense
by gulls (Bourget 1973; Schamel 1977, Gotmark and
Ahlund 1988; Fournier and Hines 2001), but may incur
additional risk if the resident gulls depredate their nests.
If the depredation rate exceeds the gulls’ protective
benefit, then Common Eiders could be experiencing
an unsustainable situation similar to the “ecological
trap” described by Dwernychuk and Boag (1972).
As part of a larger study of Common Eider breed-
ing ecology, we conducted an intensive observational
study of nesting biology on Egg Island, near Prudhoe
Bay, Alaska. Our primary objective was to replicate
the study by Schamel (1974, 1977) and document the
behavioral interactions among Common Eiders and
nest predators with minimal researcher disturbance.
Methods
We selected Egg Island for this study because data
from annual surveys showed relatively consistent con-
centrations of Common Eider nests (Noel et al. 2005)
and a similar observational study of Common Eider
reproduction had been conducted there in the 1970s
(Schamel 1974, 1977). Egg Island is a small unvege-
tated island that is part of the chain of barrier islands
that form Simpson Lagoon (Noel et al. 2005). Prior to
Common Eider nest initiation in 2002 and 2003, we
erected elevated observation blinds near the main
colony. In 2002, we used two blinds with 30 m
above-ground plywood access tunnels, and in 2003 we
used one blind with a 25 m access tunnel. The blinds
were 1.2 m x 1.2 m x 1.5 m plywood boxes raised 2.5
m above the ground. Observers accessed the island in
small boats, using the tunnels to hide our approach.
Boating was limited to a single landing and departure
per day and there was no observer activity on Egg
Island outside of the tunnels and blinds. Common
Eiders were never observed to flush while we
accessed the blinds. However, the entrance of one tun-
nel in 2002 was near three Glaucous Gull nests. These
gulls often flushed as we approached, but remained
near their nest-site and returned to the nest quickly
after we entered the blind.
With few exceptions due to poor weather, we rec-
orded daily nest observations during blocks of 4-6 x 1 h
308
2007
observation sessions. We staggered our observation
blocks to cover the entire 24-hour period, because, at
the latitude of our study site, the sun does not set dur-
ing the entire nesting period. We conducted obser-
vations from initiation of the first eider nest until
incubation at all nests terminated each year. During
observation sessions, we mapped all nests within view
of the blinds and recorded the amount of time each
nest was occupied. We recorded all interactions with
predators and, each hour, we performed 15 min pred-
ator watches during which we counted all aerial and
ground patrols by avian predators. We tallied these
counts to create an index of avian predation pressure.
We considered a nest to be a site in which at least
one egg was seen or a site that was occupied by a hen
during observations for > 24 h. We considered incu-
bation to have begun when a hen was seen on a nest
full-time after at least three consecutive days of atten-
dance. Nests were deemed failed when no eider was
seen using the site on two consecutive days. Nests occu-
pied > 24 h after an observed depredation event were
counted as partial depredation, whereas nests aban-
doned shortly after depredation were considered to be
completely depredated (i.e., entire clutch loss). To esti-
mate inter-nest distances, we visited each nest site after
all nesting birds left the island and recorded coordi-
nates with a GPS unit. All means are provided + SE.
Results
We conducted 126 h of observations on 16 days (15
June — 3 July) in 2002, and 140.5 h of observation on
23 days (19 June — 13 July) in 2003. We observed 45
Common Eider and 10 Glaucous Gull nests in 2002,
and 37 Common Eider and 9 Glaucous Gull nests in
2003. Gull nest initiation preceded eider nest initiation
by several days. The majority of Glaucous Gull nests
were already in the incubation stage when observations
began each year, whereas the median initiation dates
for Common Eiders were 23 June and 25 June, in
2002 and 2003, respectively. In both years, eider and
gull nesting periods were terminated prior to hatch
when a single Arctic Fox accessed the island and
destroyed all nests during the incubation stage.
Gull Predation
We conducted 328 avian predator patrol watches.
Glaucous Gulls were the most common predator; other
avian predators (jaegers, Stercorarius spp., and Ravens,
Corvus corax) were seen on only seven occasions.
The search rate by gulls appeared similar each year
(X patroisyh = 17.2 + 1.6 in 2002, and 14.1 + 1.0 in 2003),
but the search technique changed between years. In
2002, 68.2% + 3.0 of patrols were aerial, while in
2003 aerial patrols accounted for only 32.9% + 2.7 of
the total. Concurrently, the mean distance between
eider nests and the nearest Glaucous Gull nest de-
creased significantly (¢,, = 3.58, P < 0.001) from
75.8 m + 4.3 in 2002 to 35.2 m + 9.2 in 2003. This
REED, LACROIX, and FLINT: DEPREDATION OF EIDER NESTS
309
shift in Glaucous Gull patrols may reflect variation in
the distance between Common Eider and Glaucous
Gull nests between years.
Prior to the fox arrival in 2002, 42% (19 of 45) of
attempted Common Eider nests had failed and, in 2003,
70% (26 of 37) had failed but the observation period
lasted 10 days longer in 2003. The daily failure rates
of 1.2 nests/day in 2002 and 1.3 nests/day in 2003 were
similar. Glaucous Gull depredation was the likely cause
of all of these failed nests. We directly observed Glau-
cous Gull depredation in 7 and 11 cases, in 2002 and
2003, respectively. Most nests were lost during the
egg laying stage; however, some nests were also
destroyed during incubation (4 in 2002, 10 in 2003).
Proximity to gull nests appears to have been impor-
tant in nest failure. Fifty percent and 75% of Common
Eider nests < 30 m from gull nests failed in 2002 and
2003, respectively. In 2003, more nests (n = 20) were
within 30 m of gull nests than in 2002 (n = 4). From
the blinds, we were unable to see into nests, and thus
could not quantify clutch size or proportional egg
loss. Yet, in 2003 we did see 38 individual eider eggs
eaten by gulls; of these, 14 were taken by Glaucous
Gulls known to be nesting in the midst of the Common
Eider colony (10 eggs were taken by a single pair of
nesting Glaucous Gulls).
Partial predation was uncommon, as Glaucous Gulls
typically removed all eggs once they gained access to a
nest. We observed six cases of partial predation (4
Common Eider eggs, 2 Glaucous Gull eggs) across
years. In one case, a Glaucous Gull pressured a pair
of Common Eiders for 41 min before finally stealing
an egg from beneath the hen. During this encounter,
the hen never moved from the nest and did not indi-
cate awareness of the egg loss. The attending drake
actively attempted to defend the nest. In another in-
stance, a Glaucous Gull pulled nest material from under
an incubating hen, but was unable to remove any eggs.
We never observed a Glaucous Gull displace a Com-
mon Eider from a nest.
Common Eiders took attendance breaks through-
out the nesting period, averaging 8.2 + 1.3 min/break
based on 57 breaks of known length. Across years,
Glaucous Gulls attempted to depredate nests during
nearly one quarter (i.e., 21) of the 86 observed atten-
dance breaks, and were successful in half of their at-
tempts (11 of 21). Regularly, we observed Glaucous
Gulls. standing within 1 m of occupied Common
Eider nests. On five of these occasions the incubating
hens left their nests unattended; only two returned to
successfully defend their nest.
Common Eiders showed three defense strategies
against Glaucous Gulls in this study: (1) passively ig-
noring harassment and sitting tight on nests, (2) active-
ly nipping/lunging at nearby gulls from occupied nests
(35% of active behaviors), and (3) actively chasing
gulls from their nest after leaving it unoccupied for an
310
attendance break (65% of active behaviors). The male
Common Eider nest defense described earlier was not
a singular event; rather, while attending females at nest
sites, males often played an active role in defense. Male
attendance of females during nest prospecting and
initiation was high (for the three days prior to peak
initiation in 2002 and 2003, 24% and 71% of active
nests were attended by males) then tapered through
early incubation. Across years, males were involved in
13 of 15 observed nest defense events prior to 29 June
and 0 of 8 defenses thereafter.
Fox Predation
In both years, a single Arctic Fox gained access to
Egg Island during the incubation period. On 3 July
2002 it arrived and departed across ice floes and on
13 July 2003 it swam at least one km to Egg Island
from a nearby island and left by swimming toward the
mainland (> 2 km away). In both cases, the fox was
responsible (directly or indirectly) for the destruction
of all nests that were active at the time of arrival. The
entire fox visit to Egg Island was observed in 2002
(duration = 3 h 11 min) but not in 2003. When ob-
servers arrived in 2003 (after a 37 h absence due to
weather conditions) the fox was still on the island, but
all nests had been destroyed. There were 32 active
nests (26 Common Eider, 6 Glaucous Gull) on the is-
land when the fox arrived in 2002, and in 2003 there
were 18 active nests (11 Common Eider, 7 Glaucous
Gull) during the last observation prior to the fox visit.
When the fox arrived in 2002, there were approxi-
mately 30 non-resident Glaucous Gulls loafing on Egg
Island in addition to the resident Glaucous Gulls from
the six active nests. During observations in 2002,
Glaucous Gulls with nearby nests attacked the fox,
whereas the other gulls searched through the unpro-
tected nests (both eider and gull) and ate the contents
of all eggs encountered. Some gulls ate egg contents
at the nests, but others carried eggs up to 20 m away
before consuming them.
The fox observed in 2002 cached the vast majority
of eggs that it encountered, eating only one gull egg,
one eider egg, and the contents of a Snow Bunting
(Plectrophenax nivalis) nest. It cached at least 34 eider
and 4 gull eggs, taking on average 2.3 min to find an
egg, cache it, and find another egg. Eggs were carried
20-100 m from the nest and buried 2-5 cm deep in
areas of open sand or gravel. We do not have clutch
size data from Egg Island; however, the mean Com-
mon Eider clutch size from neighboring islands in
2002 was 2.6 eggs/nest. Given this estimate, the fox
took approximately 56% of the Common Eider eggs
on the island, while the remaining eggs were eaten by
Glaucous Gulls.
Resident Glaucous Gulls actively attacked and har-
assed the fox, but the fox was never seen to leave a site
or egg as a result of an attack. Common Eiders never
defended their nests from the fox; rather, they flushed
from their nest when the fox approached to within 20 m.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Discussion
None of the three species observed in this study
appeared to have gained fitness from the use of Egg
Island or from interactions on the island. In fact, it ap-
pears that they all may have fallen into a situation sim-
ilar to an “ecological trap”. That is, each was enticed by
initial conditions (or expectation of conditions) which
could have confered a fitness benefit, but due to changes
in those conditions, the final result was the loss of fit-
ness.
Glaucous Gulls nested earlier than Common Eiders
and may have chosen nest sites on Egg Island in antic-
ipation of Common Eider nests as a potential source
of prey or in an attempt to avoid mammalian predators.
Regardless, Common Eiders, ultimately, established
their nests near existing Glaucous Gull nests. Our
results suggest that proximity to Glaucous Gull nests
increased avian depredation risk for Common Eiders.
Similar to the results of Schamel (1977), the majority
(71%) of eider nests were depredated when located
within 30 m of gull nests.
On the barrier islands of the central Beaufort Sea
there is little vegetation, and the sparse driftwood pro-
vides minimal concealment for incubating birds (Noel
et al. 2005). Female Common Eiders taking incubation
breaks seem to be obvious to nearby gulls, as one-third
of our observations of a Glaucous Gull predation event
were apparently cued by the eider leaving for an incu-
bation break. Although Common Eider incubation
breaks were generally short and probably infrequent
(Bolduc and Guillemette 2003; Swennen et al. 1993),
our observations indicate that the detection of any
break by a Glaucous Gull could lead to the loss of an
entire clutch. Female Common Eiders generally fast
through the entire nesting period (Korschgen 1977;
Bolduc and Guillemette 2003; Swennen et al. 1993),
but several authors have noted the physiological impor-
tance of drinking periodically throughout incubation
(Bolduc and Guillemette 2003; Criscuolo et al. 2000).
Thus, it appears that eider hens were forced to trade
current reproductive effort for survival and future
reproductive potential. It is notable that attending male
Common Eiders probably did not increase nest detec-
tion substantially, as Common Eider hens were already
visible to Glaucous Gulls nesting nearby, and males
were, in fact, able to provide additional nest defense in
several cases.
Predators, too, failed to gain a fitness benefit during
this study. There was no successful Glaucous Gull
reproduction in either year. As the fox and non-resi-
dent gulls were, most likely, attracted to Egg Island by
the concentration of Common Eiders, it appears that
Glaucous Gulls, too, fell into an “ecological trap.”
Common Eider nests were highly vulnerable to depre-
dation by locally nesting Glaucous Gulls (a single
nesting pair of Glaucous Gulls consumed over one
quarter of all eggs we saw eaten in 2003), so it follows
that Common Eider eggs were an important food
2007
source for breeding Glaucous Gulls on Egg Island.
Yet, when unprotected eggs were most available (i.e.,
during the fox visit), non-resident gulls consumed the
majority of eggs (including gull eggs), while breeding
gulls were occupied with defense of their own nests.
Ultimately, these locally breeding Glaucous Gulls
failed reproductively, in spite of the apparent benefit
of nesting in proximity to Common Eider nests.
Ultimately, Arctic Foxes caused the complete nest
loss for all birds in this study, yet the fitness gained by
the foxes was likely minimal. The foxes cached almost
all the eggs and departed the island shortly after all
nests were destroyed. It is doubtful that the cached
eggs were ever recovered. To return to the island later
in the season to retrieve cached eggs, foxes would face
a longer, more arduous, swim due to continued sea
ice retreat. Further, the barrier islands of the central
Beaufort Sea experience substantial wave-battering
and redistribution of sand and gravel from storms dur-
ing the ice-free period. Thus, it is unlikely that intact
cached eggs could be recovered even if the foxes re-
turned to the island later in the season or after freeze-up.
Our results demonstrate that successful Common
Eider and Glaucous Gull reproduction in the central
Beaufort Sea can occur only in years when foxes do
not have access to the barrier islands and even then
nesting success of eiders is likely to be low (Quinlan
and Lehnhausen 1982). Island access by foxes is
unpredictable and can occur during any stage of nest-
ing. These particular foxes were probably non-breed-
ers as they were roaming the islands in July when
breeders would be attending mainland dens. The
number of non-breeding foxes fluctuates year-to-
year; however, in the oilfields around Prudhoe Bay
the degree of fluctuation has been dampened
(Burgess 2000) and the number of fox dens and the
average litter size have increased (Ballard et al. 2000;
Burgess 2000). For this eider-gull nesting association
to have evolved, the historical access of foxes to
island nesting colonies must have been relatively rare
(Ahlén and Andersson 1970). These associations can-
not persist on the barrier islands of the central Beau-
fort Sea if nesting success is consistently driven to
near zero by a changed system in which the frequen-
cy of Arctic Foxes access to the islands is increased.
Despite the total nest failure caused by foxes, our
observations provide information regarding rates of
nest depredation by gulls in the absence of human
disturbance. Gulls destroyed a fairly high proportion
of nests during the egg-laying period, and some of
these failed-early nests (often characterized by mini-
mal down or nest bow] delineation) would probably go
undetected by researchers searching for nests weeks
later during the incubation stage. This has important
implications for studies of nesting effort and success
involving incubation-stage nest searches, as a bias
against the detection of failed nests would lead to
biased estimates of these parameters. Overall, even in
REED, LACROIX, and FLINT: DEPREDATION OF EIDER NESTS 311
the absence of human disturbance, the nesting produc
tivity of Common Eiders on the barrier islands of the
central Beaufort Sea is low due to the impact of avian
and mammalian predators.
Acknowledgments
We thank J. Arnold, C. Crews, A. Farris, S. Iverson,
D. Koontz, T. Pearson, L. Skerratt, S. Sonsthagen, and
S. Wallace for field assistance. Particular thanks go to
R. Lanctot, K. Cairns, W. Cullor, J. Sarvis, W. Streev-
er, and ERA Helicopters, Inc., for logistical support
of this project. S. Are, A. Erskine, T. Fondell, H. Wil-
son, and two anomynous reviews provided helpful
comments on earlier versions of the manuscript. Fund-
ing was provided by the Minerals Management Serv-
ice and the U.S. Geological Survey — Alaska Science
Center.
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Received 7 March 2007
Accepted 23 May 2008
Management Implications of Molt Migration by the Atlantic Flyway
Resident Population of Canada Geese, Branta canadensis
SUSAN E. SHEAFFER!®, RICHARD A. MALECK?*”, BRYAN L. Swirt?, JOHN DUNN*, and Kim SCRIBNER*
'Department of Natural Resources, Cornell University, Ithaca, New York 14853 USA
U.S. Geological Survey, New York Cooperative Fish and Wildlife Research Unit, Department of Natural Resources, Cornell
University, Ithaca, New York 14853 USA
‘New York State Department of Environmental Conservation, Bureau of Wildlife, Game Bird Unit, 625 Broadway, Albany.
New York 12233 USA
‘Pennsylvania Game Commission, 911 Big Spring Road, Shippensburg, Pennsylvania 17257 USA
‘Department of Fisheries and Wildlife, Michigan State University, East Lansing, Michigan 48824 USA
Present address: Livingston Ripley Waterfowl Conservancy, P.O. Box 210, Litchfield, Connecticut 06759 USA
Sheaffer, Susan E., Richard A. Malecki, Bryan L. Swift, John Dunn, and Kim Scribner. 2007. Management implications of molt
migration by the Atlantic Flyway resident population of Canada Geese, Branta canadensis. Canadian Field-Naturalist
121(3): 313-320.
We used satellite-tracked transmitters in 2001 and 2003 to document the timing, location, and extent of molt migrations by
female Canada Geese (Branta canadensis) affiliated with the Atlantic Flyway Resident Population (AFRP) of Canada Geese
that breed in the temperate region of eastern North America. Twenty-seven adult females were captured during the nesting
period in late May and fitted with a satellite transmitter mounted either on a plastic neck collar or backpack harness. Nests of
24 birds were destroyed late in incubation to prevent renesting and ensure nest failure; three females did not have nests.
Twelve of the 27 birds (44%) made a northward migration to molt in northern Quebec, Canada: seven to the eastern coast of
Hudson Bay (58°12'N, 76°60'W), three to lowland areas east of James Bay (53°30'N, 79°02'W), and two to interior locations
south of Ungava Bay (55°54'N, 68°24'W). Molt migrants were present in northern Quebec from June to September, a period
that coincides with breeding ground aerial surveys and banding operations conducted for Atlantic Population (AP) Canada
Geese that breed in this same region of northern Quebec. With >1 million AFRP geese estimated in the Atlantic Flyway, the
potential exists for substantial numbers of yearling, sub-adult, and nest-failed or non-breeding adults to molt migrate to
northern breeding areas and bias efforts to survey and mark AP geese. Within AFRP breeding areas, many local flocks have
reached nuisance levels. We hypothesized that by inducing molt migration in breeding adults, through destruction of nests
late in incubation, we would lessen recruitment, reduce numbers of summer resident adults with young, and increase adult
mortality from hunting. However, molt migration behavior was not uniform throughout our study area. Molt migrants were
from rural areas in New York, Pennsylvania, and Vermont, whereas marked birds that did not make molt migrations were
from more coastal regions of the flyway. The 14 birds that did not make a molt migration remained within 60 km of their
banding site. A genetic comparison of these two groups revealed no detectable differences. We conclude that failure to
undergo a molt migration is likely attributed to the historical origin of captive-reared birds of mixed subspecies that comprise
AFRP flocks in the eastern regions of the flyway and the availability of quality local habitat, distinct from brood-rearing
areas, for molting.
Key Words: Branta canadensis, Canada Geese, molt migration, temperate-nesting, Resident Population, Atlantic Flyway.
_ Canada Geese (Branta canadensis) often undergo a as the Atlantic Flyway Resident Population (AFRP:
late spring migration, flying long distances from their Atlantic Flyway Waterfowl Council 1999*). Because
breeding locations to more northerly areas, where they these populations are mostly indistinguishable when
undergo an annual molt. A form of molt migration, they mix in the field on migration routes and wintering
these movements have been documented for Canada locations, their population and harvest management
Geese from both subarctic-nesting populations, which depends on monitoring efforts directed at their popu-
typically breed in remote areas of northern Canada __lation-specific breeding areas. For example, annual
. (Sterling and Dzubin 1967; Abraham et al. 1999), and —_ assessments of population size are dependent on spring
_temperate-nesting populations, that breed in southern aerial surveys of breeding areas. However, movement
Canada and the United States (Zicus 1981; Davis et al. of molting groups of temperate-nesting geese from
1985; Lawrence et al. 1998). Birds that undergo a molt Michigan to subarctic breeding areas in northern
migration are primarily nonbreeding subadults and un- Canada has been confirmed through monitoring with
successful breeding adults (Salomonsen 1968; Law- _ satellite-tracked transmitters (Mykut et al. 2004). If
tence et al. 1998). molt migrations by temperate-nesting geese occur while
There are two subarctic-nesting populations (Atlantic spring surveys are being conducted on northern breed-
Population [AP] and North Atlantic Population [NAP]) ing grounds, then population estimates for subarctic-
associated with the states of the Atlantic Flyway, where- nesting populations can be positively biased by the
as temperate-nesting geese are collectively recognized —_ inclusion of molt migrants (Abraham et al. 1999).
o>)
13
314
Similarly, population-specific assessments of harvest
rely on band recoveries obtained from birds marked on
their breeding grounds as flightless adults and young.
In recent years, recaptures on subarctic breeding
grounds of previously banded adults and young from
temperate-nesting areas (Abraham et al. 1999; Nichols
et al. 2004) have raised concern over the potential bias
in the banded samples of subarctic-nesting geese.
Molt migrations of temperate-nesting geese to north-
ern breeding areas can also increase competition for
resources and potentially degrade brood-rearing habi-
tat for subarctic-nesting geese. Previous studies have
demonstrated that a large proportion (50% — 60%) of
temperate-nesting flocks potentially undergo a molt
migration (Zicus 1981; Lawrence et al. 1998). Given
that estimated numbers of temperate-nesting geese cur-
rently exceed | million birds in each of the Mississippi
and Atlantic flyways (United States [U. S.] Fish and
Wildlife Service 2007*), the potential problems with
survey efforts and impacts to habitat could be substan-
tial.
Temperate-nesting Canada Geese also are of con-
cern within their breeding range, where they often
occur in areas with few natural predators. Because of
their relatively high survival and stable reproductive
rates, local flocks can rapidly reach nuisance levels in
urban areas and rural areas with low harvest pressure.
Given that some unsuccessful breeders undergo a molt
migration, we reasoned that deliberate nest destruction
late in the incubation period, to ensure nest failure,
would induce these birds to molt migrate. This proce-
dure has the potential to reduce annual recruitment of
young into the population, lessen the number of adults
and young residing 1n local areas during the summer,
and expose this component of the population to hunt-
ing pressure in other regions when they return in the
fall. Our objective was to assess both the efficacy of
this technique and to monitor the resultant timing, loca-
tions, and extent of molt-migration movements as they
relate to management of AFRP Canada Geese in the
Atlantic Flyway.
Methods
In 2001, eight female Canada Geese were captured
at four locations in New York during late May (Table
|). Nests of five birds were destroyed during the last
7-10 days of incubation; no nests were detected for
three of the eight (Table 1). Each goose was fitted with
a 30 g satellite-tracked transmitter (PTT Model 100,
Microwave Telemetry, Inc., Columbia, Maryland; men-
tion of trade names or commercial products does not
constitute endorsement or recommendation for use by
the U.S. Government) attached dorsally with a harness
constructed of Teflon ribbon. Transmitters had speci-
fications for ~480 hours of battery life and were pro-
grammed to transmit for an eight-hour period every four
days. Expected battery life was about eight months.
We expanded the study in 2002 to include 19 nesting
females captured under similar nesting conditions in
THE CANADIAN FIELD-NATURALIST
Vol. 121
New York (n = 4), Pennsylvania (n = 6), New Jersey
(n = 4), Massachusetts (n = 2), Connecticut (n = 1),
Vermont (n = 1), and Maine (n = 1). To simplify attach-
ment procedures and reduce costs, we switched to trans-
mitters mounted on plastic neck collars produced by
Telonics, Inc. of Mesa, Arizona. Program features were
similar to those used in 2001. Unfortunately, most of
the transmitters deployed in 2002 did not provide
sufficient signals to the satellites to produce usable
locations. The 2002 transmitters were replaced by the
manufacturer and 19 nesting females were marked in
2003: New York (n = 4), Pennsylvania (n = 4), New
Jersey (n = 4), Massachusetts (n = 2), Connecticut
(n = 2), Vermont (n = 1), Maine (n = 1), and Maryland
(n = 1) (Table 1).
Data from the radio transmitters were obtained from
the Argos satellite system of the French Space Agency
via a preferential tariff agreement with the U. S. Depart-
ment of Commerce’s National Oceanic and Atmos-
pheric Administration. Location estimates were classi-
fied by Argos based on their estimated accuracy and
the number of transmissions received from a transmit-
ter during a satellite overpass. Location classes 3, 2, and
1 had accuracy ratings within 1000 m. Accuracy for
location class 0 was >1000 m, and location classes A
and B did not receive enough transmissions during an
overpass for accuracy to be estimated (Service Argos
1996*). However, Britton et al. (1999) demonstrated
that poor locations (classes 0, A, and B) received from
satellite transmitters (30 g) averaged 35 km from the
true location of the transmitter. We therefore included
all locations with a classification of 3, 2, 1, 0, A, or B,
because the accuracy of these classes was sufficient to
describe large-scale movements.
Argos estimated locations by measuring the Doppler
shift of the received signals, which produces two pairs
of latitude and longitude coordinates from an individ-
ual satellite overpass. Argos designates the location
with the better frequency continuity as the most prob-
able location (location 1), and the alternate location is
designated as the image (location 2). Examination of
our data occasionally indicated that sometimes loca-
tion 2 was a more probable fix than location | based on
the flight dynamics of Canada Geese. We developed a
sorting routine similar to that of Britton et al. (1999)
that sequentially examined the location pairs to iden-
tify locations that appeared most probable. The initial
location for each bird was the site of banding. For each
subsequent pair of locations, flight speed (distance/hr)
was calculated from location | in the previous location
pair to location 1 of the next location pair. If the bird
had flown <65 km/hr to reach both locations | and 2,
we selected location 1. If the bird had flown >65 km/hr
to reach location 1, but <65 km/hr to reach location 2,
we selected location 2. If both locations violated our
65 km/hr rule, we deleted both. We retained all loca-
tions with an accuracy classification of 3, 2, and 1.
When an 8-hour transmission period contained only
locations with a classification of 0, A, and B, we esti-
2007
mated one location for that period as the centroid of
the most probable latitude and longitude coordinates.
During the course of our study, we hypothesized that
the differential propensity to undergo a molt migration
could be related to genetic differences among geese
from interior and coastal areas comprising our sample.
In 2004, we attempted to test this by collecting blood
samples from molting adult geese with young in rural
areas of central New York (n = 15) and Pennsylvania
(n = 15) and compared these with similarly collected
samples from more coastal and urban/suburban areas
of Long Island, New York (n = 5), New Jersey (n = 15),
Massachusetts (m = 5), and Connecticut (nm = 5). Genet-
ic testing of microsatellite markers and mitochondrial
DNA was done using methods described by Scribner
et al. (2003 a).
We used seven microsatellite loci that have previous-
ly been described for Canada Geese for other popula-
tions across North America (Scribner et al. 2003a,
2003b). Loci included TTU-CG-1, TTU-CG-5 (Cathey
et al. 1998) and Bcau7, Bcau9, Beaull, and Hhil,
(Buchhollz et al. 1998), and CRG (Baker, unpublished
data). After electrophoresis on denaturing 6% acry-
lamide gels, PCR products were visualized using an
FMBIO II laser scanner (Hitachi Software Engineering
Co., Alameda, California). Genotypes were scored
based on 20 base-pair standards and reference samples
of known allelic size.
We obtained sequence information from a ~ 385 base-
pair (bp) fragment of the 5' end of the Canada Goose
mitochondrial DNA control region using primers and
conditions described in Pearce et al. (2000). Sequenc-
ing was performed using a SequiTherm Excel DNA
sequencing kit (Epicentre, Inc.), following product
protocols for use of fluorescently labeled primers.
Sequences were aligned manually.
We estimated degree of differentiation in allele and
haplotype frequency between Canada Geese sampled
from coastal and interior regions using a Fisher’s exact
test in program GENEPOP and using F-statistics (Weir
1996) using the program F-STAT (Goudet 1995). Mea-
sures of inter-population variance in allele frequency
were summarized as pair-wise estimates of population
F,. Significance of mean F,, values (across the seven
loci) were determined by jack-knifing procedures (Weir
1996). Analyses of population differences in mtDNA
haplotype frequency were conducted using an analy-
sis of molecular variance (Excoffier et al. 1992).
Results
Of the eight birds marked in New York in 2001,
seven made molt migrations to areas in Quebec, Cana-
da: four traveled to the eastern coast of Hudson Bay
(58°12'N, 76°60'W), two went to lowland areas east of
James Bay (53°30'N, 79°02'W), and one moved to an
interior location ~350 km southwest of Ungava Bay
(56°06'N, 70°04'W) (Figure 1). Locations from four
transmitters deployed in 2002 (two from Pennsylvania,
SHEAFFER, MALECKI, SWIFT, DUNN, and SCRIBNER: CANADA GEES 315
TABLE |. Bird identification numbers, state and year of band
ing (cohort), latitude and longitude of banding location, and
approximate distance between banding and molting location
(km). Females with no nest are identified with an (*)
Banding location
ID Cohort latitude longitude km
33115 NY-2001 43°06N 78°30 W 1122
33116* NY-2001 43°06'N = 78°24'W 897
33117 NY-2001 42°12'N 78°54'W 1830
33118 NY-2001 42°06'N 78°36'W 1295
33119* NY-2001 41°36'N = 74°06"W 1641
33120 NY-2001 41°36'N 74°06'W <60
33121* NY-2001 42°42'N 75°30'W 194]
33122 NY-2001 42°42'N 75°30'W 1725
19873 CT-2003 41°20'N 71°32'W <60
19874 PA-2003 40°39'N 77°45'W 1830
19875 MD-2003 38°35'N 76°07'W <60
19876 CT-2003 41°32'N 73°04'W <60
19909 ME-2003 44°43'N 68°54'W <60
1991] MA-2003 41°37'N 71°06'W <60
19916 MA-2003 41°17'N = 71°27'W <60
19922 NJ-2003 40°42'N = 74°30'W <60
19923 NJ-2003 40°25'N 74°29'W <60
19930 NJ-2003 39°35'N 75°29'W <60
19932 NJ-2003 41°00'N 74°09'W <60
19934 NY-2003 41°0S'N = _73°51'W <60
19940 N Y-2003 42°50'N 73°50'W 1573
19955 NY-2003 40°45'N = 73°07'W <60
19957 NY-2003 40°37'N 3°15'W <60
19962 PA-2003 39°48'N = 80°03"'W 1396
19973 PA-2003 40°07'N 72°24'W 564
19978 PA-2003 40°32'N 75°95'W <60
20024 PA-2003 40°36'N = 80°00'W <60
20036 PA-2003 42°09'N —- 80°02"W <60
20043 PA-2003 41°1S'N = 75°18'W <60
20059 VT-2003 44°05'N = 73°20'W 1521
NY = New York, CT = Connecticut, PA = Pennsylvania,
MD = Maryland, ME = Maine, MA = Massachusetts,
NJ = New Jersey, VT = Vermont
one from New York, and one from Vermont) suggested
that these birds also made a molt migration to northern
Quebec; however, the data were not sufficient to iden-
tify migration timing or provide reliable locations.
Only five of the 19 birds marked in 2003 made molt
migrations to northern Quebec (three from Pennsylva-
nia, one from New York, and one from Vermont). Three
traveled to the eastern coast of Hudson Bay (58°12'N,
76°60'W), one went to eastern James Bay (53°54'N,
79°00'W), and one moved to an interior location south
of Ungava Bay (57°00'N, 74°36'W) (Figure 1). The
14 females that did not make a molt migration exhibit-
ed little movement during June through early Novem-
ber; movement of these geese was limited to within
60 km of the banding site. All of the birds that did not
make a molt migration were from more coastal regions
of the flyway (Figure 2).
Molt migrations in both 2001 and 2003 occurred in
June. Marked geese that molt migrated (7 = 12) were
located in the U.S. as late as 30 May — 20 June. They
arrived at terminal locations in northern Quebec (Fig-
316
ure 1) between 9 and 26 June. Fall migration of these
molt migrants occurred primarily in September. Birds
were located in northern Quebec as late as 28 August
— 29 September. They arrived in southern Canada or
the U.S. between 6 September and 3 October (Figure
3). There was no distinguishable difference in timing
of molt and fall migrations between marked geese in
2001 and 2003. Molt migrations ranged in distance
from 564 — 1941 km (Table 1).
Genetic analysis of the two groups of geese showed
no evidence across loci of significant differences in
microsatellite allele frequency (F = 0.005, P > 0.05).
Likewise, analysis of variance results for mtDNA hap-
lotype frequencies also revealed no evidence of sig-
nificant differences between the two groups (® = 0.00,
P > 0.40). When analyses were extended to include
reference samples from the two subarctic-nesting Cana-
da Goose populations (AP and NAP), we observed dif-
ferences in allele frequencies among all three popula-
tions for both microsatellite loci and mtDNA haplotype
(P < 0.05 for both markers).
Discussion
Our results provide convincing evidence of the
movement of some molting AFRP geese to the north-
ern breeding ground of AP geese at a critical time dur-
ing the spring population monitoring and summer pre-
hunting season banding periods. Zicus (1981) and
Lawrence et al. (1998), working with temperate-breed-
ing Canada Geese in Wisconsin and Illinois, each re-
ported 50-60% of their spring population departing on
molt migrations. Mykut (2002) satellite-tracked female
Canada Geese in Michigan, whose nests were des-
troyed, as in this study, late in incubation. He found
62% (n = 37) in 2000 and 73% (n = 52) in 2001 of
these birds molt-migrating to James and Hudson Bays
in northern Canada. The literature is replete with other
reports of long distance molt migrations in temperate-
nesting geese.
The propensity to molt migrate, observed in nest-
failed females from rural New York and Pennsylvania,
but not for birds nesting in more coastal areas of the
flyway, prompted us to suspect a possible genetic dif-
ference resulting from regional differences in the ori-
gin of these birds. Blandin and Heusmann (1974) re-
ported that geese breeding in eastern Massachusetts
were of mixed racial stock, nonmigratory in nature, and
the progeny of decoy birds released in the late 1930s.
Morphometric measurements of these birds (Pottie and
Heusmann 1979) indicated a probable mixing of sev-
eral sub-species, in which B. c. maxima (Giant Canada
Goose) predominated. In contrast, geese in rural New
York and Pennsylvania are primarily large Canada
Geese of the B. c. maxima and B. c. moffitti (Great
Basin Canada Goose) stock introduced after the 1950s
to areas previously devoid of breeding geese. These
birds may be more similar to mid-continent temperate-
nesting populations of restored or reintroduced giant
THE CANADIAN FIELD-NATURALIST
Vol. 121
Canada Geese. Molt migration in giant Canada Geese
is considered a common behavior (Abraham et al. 1999).
We were able to detect genetic differences of sufficient
magnitude to discern the AFRP geese we sampled from
subarctic-nesting populations of geese occurring in the
flyway (AP and NAP), but we did not detect genetic
differences between the two regional cohorts sampled
within the AFRP.
Two factors favoring movement of waterfowl to re-
mote regions to molt are the reduced risk to survival
and greater food availability (Hohman et al. 1992;
Baldassarre and Bolen 2006). In Canada Geese, suc-
cessful breeders also are often behaviorally dominant
over nonbreeders (Raveling 1970), which may influ-
ence movement of the nonbreeding segment away
from nesting areas. However, in New Jersey, Nichols
et al. (2004) estimated that only 22-31% of resident
Canada Geese migrated out of the state to molt. They
suggested that birds in and near urban-suburban dom-
inated landscapes may have many suitable molting sites
consisting of large expanses of lush, fertilized lawns
interspersed with water bodies and few predators. The
gathering of nonbreeders into larger flocks to molt on
or near breeding areas may also lessen stress associat-
ed with behavioral interactions of adults with young.
Lawrence et al. (1998) also noted that in more recent
times, with greater emphasis on hunting of temperate-
nesting geese to manage their numbers, birds leaving
the sanctuary of local breeding areas are more likely
to be harvested.
The coastal region of our study, from which molt
migration did not occur (Connecticut, Massachusetts,
Maryland, and New Jersey), contains ~21% (N =
245351) of the AFRP spring population estimate (Unit-
ed States Fish and Wildlife Service 2007*) in the U.S.
Geese in New York, Pennsylvania, and Vermont make
up ~45% (N = 514181) and the coastal states of Dela-
ware and Virginia, where geese were not sampled as
part of this study, comprise ~14% (N = 163655). If we
assume that 50% of the geese in New York and Penn-
sylvania are molt migrant non-breeders or unsuccess-
ful breeders (Zicus 1981; Lawrence et al. 1998), then
the potential exists for a substantial movement of geese
(=250 O00) to subarctic breeding areas in northern
Canada. This number is further increased by temper-
ate-nesting birds moving into subarctic regions from
the Mississippi Flyway.
The population estimate for AP geese in northern
Quebec, where we suspect most AFRP birds molt mi-
grate to, is ~1.2 million (Harvey and Rodrique 2007*).
However, the authors caution that differences in survey
timing and the abundance of molt migrants can clearly
introduce substantial variability in the total population
estimate. Similar conflicts are reported by Abraham et
al. (1999) for Southern James Bay Population Canada
Geese in northern Ontario. Spring surveys of AP geese
are timed to cover the mid to late incubation period;
generally the last two weeks in June. Arrival of AFRP
2007
60°
95°
90°
Ontario
45°
40°
SHEAFFER, MALECKI, SWIFT, DUNN, and SCRIBNER: CANADA GEES! 317
Legend
@ Release location
A Telemetry location
— Banded in 2001
__ Banded in 2003
1000 Kilometers
Ficure 1. Locations of Atlantic Flyway Resident Population Canada Geese during spring molt migration in 2001 and 2003.
Locations were identified using birds marked with satellite-tracked transmitters. Birds arrived at terminal molting
areas in northern Quebec between 9 June and 16 June.
molt-migrants coincides very closely with the begin-
ning of these surveys. Similarly, summer banding of
AP geese, which usually takes place in late July —
early August, also occurs when molt migrants are
still on the AP breeding grounds. Given that molting
large Canada Geese can require up to 40 days to
obtain the 85% of primary feather development
required to regain flight (Hanson 1965), geese arriving
in mid-June have the potential still to be flightless
when subarctic-breeding adults with young are being
banded. This mixing of populations re-enforces the
need to (1) monitor the status of subarctic-breeding
populations using numbers of breeding pairs surveyed,
rather than total counts involving groups of geese of
unknown origin, and (2) band only flightless adults
with young during the summer pre-season banding
period to ensure a representative sample of subarctic-
nesting populations of Canada Geese.
In the late 1980s, as an aid in the control of increas-
ing numbers of temperate-nesting geese, the U.S. Fish
and Wildlife Service endorsed the implementation of
special hunting seasons in September and late-winter,
when subarctic-nesting goose populations would be
less likely affected (Heusmann et al. 1998). While
effective in many rural areas, geese in urban and sub-
urban environments often receive little exposure to
hunter harvest (Smith et al. 1999). Efforts to control
nuisance geese generally rely on non-lethal methods
involving addling and oiling of eggs in nests (Chris-
tens et al. 1995) and harassment techniques to move
birds from heavily used residential and industrial areas
(Holevinski et al. 2007). Both methods provide only
a moderate degree of relief, at best.
318
Ontario
3°55
“SY Of
100 Kilometers
THE CANADIAN FIELD-NATURALIST
Vol. 121
Quebec
a
ve
Vermont | ‘Sil
Arar -
Ae!
ew
Hampshire
an a ie A
“Rhode Island
_)
ao
Connecticut
New Jersey
Delaware
Legend
@ Underwent a molt migration
O Did not undergo a molt migration
FIGURE 2. Banding locations of Atlantic Flyway Resident Population Canada Geese marked with satellite-
tracked transmitters during May 2001 and 2003.
Many studies have demonstrated that altering sur-
vival of adult geese is much more effective in changing
population size than altering recruitment rates (Trost
et al. 1986; Schmutz et al. 1997). However, direct
culling of molting birds continues to remain a socially
unacceptable option for population control in many
areas. One alternative, transplanting flightless adults
with young from nuisance areas to rural areas prior to
the fall hunting season, successfully reduced survival
of adults in comparison to non-transported birds, but
was both time consuming and costly (Holevinski et al.
2006). Inducing molt migration of adult breeding geese
by destroying nests late in incubation, as demonstrated
in this study, showed potential for both removal of birds
from their breeding area during the spring and sum-
mer period and exposing them to hunting outside the
region. However, our failure to demonstrate this in
coastal areas, which have more prominent nuisance
Canada goose issues, was problematic. We can only
assume that the reason for this was attributed to the
historical origin of captive-reared birds of mixed sub-
species making up AFRP flocks in this part of the fly-
way and the availability of quality local molting
habitat distinct from brood-rearing areas.
Acknowledgments
We acknowledge the valuable assistance and fund-
ing received from the following individuals and their
respective agencies: P. Castelli and T. Nichols, New
Jersey Division of Fish and Wildlife; H. W. Heusmann,
Massachusetts Division of Fisheries and Wildlife; M.
Huang, Connecticut Department of Environmental
Protection; W. Crenshaw, Vermont Fish and Wildlife
Department; B. Allen, Maine Department of Inland
Fisheries and Wildlife; L. Hindman, Maryland Wildlife
and Heritage Service; and the agencies and university
affiliates of the authors. Valuable editorial comments
were provided by G. Baldassarre, College of Environ-
mental Science and Forestry at Syracuse University, L.
Hindman, and the anonymous referees for this journal.
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SHEAFFER, MALECKI, SWIFT, DUNN, and SCRIBNER: CANADA GEESI 319
Legend
@ Molt location
A Telemetry location
— Banded in 2001
_. Banded in 2003
1000 Kilometers
Ficure 3. Locations of Atlantic Flyway Resident Population Canada Geese returning from molting locations in the fall 2001
and 2003. Locations were identified using birds marked with satellite-tracked transmitters. Birds initiated fall migra-
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Received 8 August 2006
Accepted 7 May 2008
Recovery of DNA from Footprints in the Snow
Love DALEN!®, ANDERS GOTHERSTROM!2, TOMAS MEUER®, and BETH SHAPIRO"
' Centro Mixto UCM-ISCIII de Evolucién y Comportamiento de Humanos, 28029, Madrid, Spain
* Evolutionary Biology Centre, Uppsala University, 752 36, Uppsala, Sweden
+ Department of Zoology, Stockholm University, S-10691, Stockholm, Sweden.
* Department of Zoology, Oxford University, South Parks Road, Oxford OX1 3PS United Kingdom
*Present address: School of Biological Sciences, Royal Holloway, University of London, Egham TW20 OEX, United Kingdom:
corresponding author email: Love.Dalen@rhul.ac.uk.
Dalén, Love, Anders Gotherstr6m, Tomas Meijer, and Beth Shapiro. 2007. Recovery of DNA from footprints in the snow
Canadian Field-Naturalist 121(3): 321-324.
The recovery of trace amounts of DNA has been demonstrated to be a reliable tool in conservation genetics and has become a
key component of modern forensic casework. To date, genetic data have been successfully recovered from a variety of sources,
including biological fluids, faeces, clothing, and even directly from fingerprints. However, to our knowledge and despite their
widespread occurrence and clear potential as a source of DNA, genetic information has not previously been recovered direct-
ly from footprints. Here, we extract and amplify mitochondrial DNA from a snow footprint, <48-hours old, made by a
Swedish Arctic Fox (Alopex lagopus). Our results demonstrate that it is possible to recover sufficient DNA from recent foot-
prints to accurately type the source of the print, with implications for conservation biology and forensic science.
Key Words: Arctic Fox, Alopex lagopus, Red Fox, Vulpes vulpes, trace DNA, conservation biology, forensics.
Both conservation and forensic genetics rely on the _ fluids. However, previous attempts to isolate DNA from
ability to retrieve data from material containing only _ randomly selected patches of ice and snow have shown
minute amounts of DNA. In forensic studies, key data’ that DNA is rare in the general environment (Hansen
are often generated from small amounts of biological and Willerslev 2002). Nonetheless, if it were possible
tissue or liquids recovered from crime scenes (Bond _ to identify patches in which DNA is concentrated, the
2007). Conservation geneticists are often interested in combination of cold preservation and the purity of the
applying non-invasive sampling techniques in their sample should make these patches an excellent source
research, many of which also target low copy-number _ of genetic information.
biological sources such as hair or faeces (DeSalle and The ability to amplify DNA from fingerprints has
Amato 2004). Methods for recovering trace amounts shown that a few shed skin cells are sufficient for
of DNA have been fine-tuned over the last decade, genetic typing. Footprints made by people or animals
resulting in the successful amplification of DNA from should therefore be equally viable sources of DNA;
a variety of sources. For example, previous work has _ however, the nature of the location of footprints (on the
shown that it is possible to recover nuclear DNA from ground, mixed with myriad other genetic data) has
items of worn fabric, such as clothing and the insoles made this hypothesis difficult to test. Footprints made
of shoes (Wickenheiser 2002; Bright and Petricevic in snow, however, are unlikely to be contaminated by
2004), and from other fabrics such as bedding (Petri- other sources of DNA or chemical inhibitors. Here,
cevic, Bright, and Cockerton 2006). Hair shafts and we investigate whether it is possible to recover DNA
faeces (Kohn and Wayne 1997) from humans, animals, directly from animal footprints made in snow, and
and even ancient specimens (e.g., Higuchi et al. 1988; discuss the potential of such an approach for forensic
Gilbert et al. 2004; Pfeiffer et al. 2004; McNevin et al. | science and conservation biology.
2005), fingernails and fingernail scrapings (e.g., Oz and
Zamir 2000), and latent fingerprints (e.g., van Oorschot Materials and Methods
and Jones 1997; Alessandrini et al. 2003; Balogh et al. Six samples from fresh (<48h old) fox tracks were
2003) have also been shown to be good sources of both —_ collected in Jamtland County, Sweden, during the win-
mitochondrial DNA (mtDNA) and nuclear DNA. ter in 2006. Samples were taken from different tracks,
It is well known that DNA survival is affected by _ so as to minimise the risk of sampling the same indi-
ambient temperature, with the greatest probability of | vidual twice. Each imprint was carefully excavated
survival occurring in locations with constant low tem- —_ from the snow using sterile equipment and put into 50
peratures (Smith et al. 2003). It is therefore conceivable ml Falcon tubes. Three additional samples were taken
that should biological material be deposited in ice or from undisturbed snow, for use as negative controls.
snow this frozen environment would provide ideal con- Due to the possibility of low DNA yields from foot-
ditions for DNA survival. Potential sources of DNA prints, and ensuing risk of contamination, the DNA
In ice or snow include shed cells, hair, and biological extractions were done in a dedicated ancient DNA
;
ws)
i)
—
Alopex_NEA1
Alopex_ICE2
Alopex_NEA2
Alopex_HOL1
Alopex_HOL2
Alopex_HOL4
Alopex_SIB3
AF338795_Vulpes
AF338794_ Vulpes
AF338792_Vulpes
AF338791_ Vulpes
AF338801_ Vulpes
AF338800_Vulpes
AF338799_Vulpes
AF338796_Vulpes
THE CANADIAN FIELD-NATURALIST
Volt
B
Alopex_HOL9
Alopex_GRE3
Alopex_CAN6
Alopex_GRE1
Alopex_GRE2 P Pr
Alopex_HOL7 r) 6
AF338798_Vulpes wt
FiGuRE 1. (A) Maximum liklihood tree describing the relationships between mitochondrial DNA sequences from Arctic
Foxes, Red Foxes, and that recovered from the footprint in central Sweden. Details of the phylogenetic analysis and
Statistical support are given in the text. (B) Map showing the Holarctic distribution of the two species of fox. Lighter
and dark grey shading indicates the distribution of the Red Fox and Arctic Fox, respectively, and black shading indi-
cates where the two ranges overlap. The line indicates the location in Sweden from which the footprints were recov-
ered.
facility in Madrid, Spain. The samples were concen-
trated using 15 ml Amicon filters (Millipore Massa-
chusetts, USA) and extracted from the resulting fil-
trate following the protocol by Yang et al. (1998), as
modified by Svensson et al. (2007). An approximately
150 base pair (bp) fragment of the mitochondrial con-
trol region was amplified using primers Pex3F and
H3R (Dalén et al. 2007). Resulting PCR products
were sequenced using the Big Dye Terminator v3.1
Cycle Sequencing Kit (Applied Biosystems, Califor-
nia, USA) and analysed on an Applied Biosystems
3730xl sequencer following the manufacturer’s
instructions.
Previously published Arctic Fox (Alopex lagopus)
and Red Fox (Vulpes vulpes) mitochondrial DNA
sequences covering the length of the 150-bp amplifid
fragment were collected from GenBank (accession
numbers AF338791-AF338792, AF338794-AF338796,
AF338 798-AF338801, AY321120-AY321148) and
aligned by eye. A Maximum Likelihood (ML) phylo-
genetic analysis was performed using the program
PAUP v4.10b (Swofford 1999). An HKY model of
nucleotide substitution was assumed, so as to allow
for different rates of transitions and transversions along
the sequences. Starting trees were generated by neigh-
bor-joining (NJ), from which subsitution parameters
were estimated and fixed. Heuristic searches were then
performed using the estimated substitution parameters
and SPR branch swapping. Substitution parameters
were then re-estimated from the most likely trees and
the analysis was repeated. Figure | shows the resulting
ML tree describing the relationship of the sequence
amplified from the footprint with Arctic and Red foxes.
The two clades are separated by bootstrap (100 full
heuristic runs, with starting trees generated by NJ and
NNI branch swapping) support values of 96%.
Results
Mitochondrial DNA was recovered from one of the
six footprints from which snow was collected. While
it has been shown recently that DNA can be recovered
from faecal and hair samples collected along animal
2007
tracks (Ulizio et al. 2006), this is the first report of
DNA being recovered directly from the tracks them-
selves. Identification of species from tracks presents
an important resource for conservation and manage-
ment of rare species. In this case, both Arctic and Red
foxes inhabit the Swedish mountain tundra, and the
footprint could thus have derived from either of the
two species. However, the resulting mtDNA sequence
showed that the footprint originated from an Arctic
Fox (Figure |) and was identical to one of the three
mtDNA haplotypes that exist today in Sweden (Dalén
et al. 2005).
Discussion
While the overall success of this initial study was
small, it is worthwhile to note that, in this case, only
one footprint was sampled from each track. If this
method were to be applied in a forensic or conservation
genetics context, it would be reasonable to sample
several footprints from the same track, significantly
increasing the potential for DNA recovery. While this
approach would increase the volume of snow or other
substrate from which the biological material must be
recovered, modern filtration devices such as those em-
ployed in this study make it feasible to concentrate
large volumes in relatively little time.
The ability to recover DNA from footprints has
implications for many areas of research that rely on
the ability to amplify low copy number DNA. For ex-
ample, DNA from footprints could be used to identify
individual carnivores that prey on livestock, or to link
animal prints found at a crime scene to a specific in-
dividual. In conservation genetics, this method makes
it feasible to follow individual movements across a
landscape, and to directly establish the dimensions of
individual home ranges. Perhaps most importantly, the
amplification of DNA from footprints could serve as an
additional source of non-invasive sampling, comple-
menting conventional materials such as faeces (H6ss
et al. 1992), hair (Taberlet et al. 1993) and urine (Valiere
and Taberlet 2000), and potentially avoiding many of
the problems with inhibition that are associated with
these sources of DNA.
Acknowledgments
LD was funded by a Marie Curie Intra-European
Fellowship; BS was funded by the Royal Society. The
authors thank the Ministerio de Ciencia y Tecnologia in
Spain and EU-Life SEFALO+ for financial and logis-
tical support.
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as a source of DNA for species and individual identifi-
cation. Molecular Ecology 9: 2150-2152.
THE CANADIAN FIELD-NATURALIST
Vol. 121
van Oorschot, R. A. H., and M. K. Jones. 1997. DNA fin-
gerprints from fingerprints. Nature 387: 767-767.
Wickenheiser, R. A. 2002. Trace DNA: A review, discus-
sion of theory, and application of the transfer of trace quan-
tities of DNA through skin contact. Journal of Forensic
Sciences 47: 442-450.
Yang, D. Y., B. Eng, J. S. Waye, J. C. Dudar, and S. R.
Saunders. 1998. Technical note: Improved DNA extraction
from ancient bones using silica-based spin columns. Amer-
ican Journal of Physical Anthropology 105: 539-543.
Received 8 August 2007
Accepted 15 July 2008
Notes
A River Otter’s, Lontra canadensis, Capture of a Double-crested
Cormorant, Phalacrocorax auritus, in British Columbia’s Gulf Island
Waters
MICHAEL H. H. Price! and CLARE E. ARIES2
'Raincoast Conservation Foundation. PO Box 2429, Sidney, British Columbia V8L 3Y3 Canada
’*Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8W 9W2 Canada
Price, Michael H. H., and Clare E. Aries. 2007. A River Otter’s, Lontra canadensis, capture of a Double-crested Cormorant
Phalacrocorax auritus, in British Columbia’s Gulf Island Waters. Canadian Field-Naturalist 121(3): 325-326.
Direct and apparent predation events by River Otters (Lontra canadensis) on birds have been recorded on marine islands and
freshwater lakes. We add to this the first known observation of a River Otter capturing a marine bird on the ocean.
Key Words: River Otter, Lontra canadensis, diet, birds, foraging, conservation, coastal, islands, British Columbia.
The River Otter (Lontra canadensis) is a largely pis-
civorous carnivore inhabiting riparian areas, coastal
shorelines, and archipelagos of North America (Cowan
and Guiguet 1965). Their diets have been well des-
cribed along inland river systems and lakes (Sheldon
and Toll 1964; Knudsen and Hale 1968; Reid et al.
1994), yet little is known about their ecology in marine
communities. Nevertheless, fishes are their primary
food in all habitats (Lagler and Ostenson 1942; Sten-
son et al. 1984; Bowyer et al. 1994). Birds are consid-
ered rare and sporadic food items (Toweill 1974; Sten-
son et al. 1984).
Early reports of bird remains in the stomachs or fae-
ces of River Otters suggested that they scavenged car-
casses (Lagler and Ostenson 1942; Sheldon and Toll
_ 1964). Years later, it was postulated that River Otters
_ probably consume wounded or incidentally killed birds
hunted by humans (Toweill 1974). In coastal British
Columbia, Washington, and Alaska, however, research-
ers have recently described predation events by River
Otters on nesting marine birds (Hayward et al. 1975;
_ Foottit and Butler 1977; Verbeek and Morgan 1978;
Quinlan 1983; Speich and Pitman 1984; Duffy 1995).
We herein describe a River Otter’s successful aquatic
| capture of a Double-crested Cormorant (Phalacroco-
rax auritus) in marine waters.
' At approximately 1130 hours on 18 February 2005,
a family of four River Otters [one adult — three juve-
| niles] swam in an unnamed cove adjacent to Swartz
1 Bay, Vancouver Island (48°42'N, 123°25'W). The sky
| Was overcast, and the sea and winds were calm. At this
i
=
Sa
time, we also observed a mixed raft of approximately
120 Double-crested and Pelagic (P. pelagicus) cor-
morants and Pacific Loons (Gavia pacifica) situated
.
|
}
400 meters from the River Otters to the north. The
birds drifted east with the current, roughly 100 meters
south of Piers Island. They consistently dived and sur-
faced, presumably feeding on small fishes [Pacific
Sandlance (Ammodytes hexapterus) or Pacific Herring
(Clupea pallasii)).
The four River Otters swam directly towards the raft
with their heads above water. When within 50 meters,
all Otters dived in succession; apparently the foraging
birds failed to detect their approach, as they continued
to dive and surface. Less than two minutes later, a lone
River Otter began swimming south to the unnamed
cove with its head above water, towing a large object in
its mouth. It dived, and re-surfaced less than 3 meters
from our viewing platform with a gasping Double-
crested Cormorant held by its throat. The River Otter
then swam beneath a shoreline cabin, directly adjacent
to a known den-site, and presumably fed upon its prey.
We failed to observe the juvenile River Otters from that
point on, and saw no evidence of them joining the adult
to consume the cormorant. We also saw no physical
evidence [i.e., carcass or feathers] after the event.
Observations of River Otters preying on birds are
rare and almost exclusively observed on land or fresh-
water. For example, River Otters have ambushed nest-
ing Glaucous-winged Gulls (Larus glaucescens), Aleu-
tian Terns (Sterna aleutica) (Hayward et al. 1975:
Foottit and Butler 1977; Verbeek and Morgan 1978:
Duffy 1995), and Slavonian Grebes (Podiceps auritus)
(Perkins et al. 2005), and preyed on incubating Storm-
petrels [((Oceanodroma furcata and O. leucorhoa)] by
digging them out of nesting burrows (Quinlan 1983).
River Otters have also been observed preying on moult-
ing American Widgeon (Anas americana) and Green-
326
winged Teal (A. crecca) on a freshwater lake in Alberta
(Reid et al. 1994), and a Common Gallinule (Gallinu-
la chloropus) on a slurry pond in Florida (Meyer-
riecks 1963). To our knowledge, this is the first obser-
vation of a River Otter preying on a bird on the ocean.
Acknowledgments
We thank the Coast Salish peoples for the use of their
traditional territory, the Price family for use of their
research cabin, and C.T. Darimont and two anonymous
reviewers for improving the manuscript.
Literature Cited
Bowyer, R. T., J. W. Testa, J. B. Faro, C. C. Schwartz, and
J. B. Browning. 1994. Changes in the diet of river otters in
Prince William Sound, Alaska: effects of the Exxon Valdez
oil spill. Canadian Journal of Zoology 72: 970-976.
Cowan, I. M., and C. J. Guiguet. 1965. The mammals of
British Columbia. British Columbia Provincial Museum.
Handbook (11).
Duffy, D. C. 1995. Apparent river otter predation at an Aleu-
tian tern colony. Colonial Waterbirds 18: 91-92.
Foottit, R. G., and R. W Butler. 1977. Predation on nesting
glaucous-winged gulls by river otter. Canadian Field-
Naturalist 91: 189-190.
Hayward, J. L., Jr., C. J. Amlaner Jr., W. H. Gillet, and
J. F. Stout. 1975. Predation on nesting gulls by a river
otter in Washington State. Murrelet 56: 9-10.
Knudsen, G. J., and J. B. Hale. 1968. Food habits of otters
in the Great Lakes region. Journal of Wildlife Management
32: 89-93.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Lagler, K. F., and B. T. Ostenson. 1942. Early spring food
of the otter in Michigan. Journal of Wildlife Management
6: 245-254.
Meyerriecks, A. J. 1963. Florida otter preys on common
gallinule. Journal of Mammalogy 44: 425-426.
Perkins, A. J., M. H. Hancock, N. Butcher, and R. W.
Summers. 2005. Use of time-lapse video cameras to
determine causes of nest failures of Slavonian grebes. Bird
Study 52: 159-165.
Quinlan, S. E. 1983. Avian and river otter predation in a
storm-petrel colony. Journal of Wildlife Management:47:
1036-1043.
Reid, D. G., T. E. Code, A. C. H. Reid, and S. M. Herrero.
1994. Food habits of the river otter in a boreal ecosystem.
Canadian Journal of Zoology 72: 1306-1313.
Sheldon W. G., and W. G. Toll. 1964. Feeding habits of the
river Otter in a reservoir in central Massachusetts. Journal
of Mammalogy 45: 449-455.
Speich, S. M., and R. L. Pitman. 1984. River otter occur-
rence and predation on nesting marine birds in the Wash-
ington Islands Wilderness. Murrelet 65: 25-27.
Stenson, G. B., G. A. Badgero, and H. D. Fisher. 1984.
Food habits of the river otter Lontra canadensis in the
marine environment of British Columbia. Canadian Journal
of Zoology 62: 88-91.
Toweill, D. E. 1974. Winter food habits of river otters in
western Oregon. Journal of Wildlife Management 38: 107-
111.
Verbeek, N. A. M., and J. L. Morgan. 1978. River otter
predation on glaucous-winged gulls on Mandarte Island,
British Columbia. The Murrelet 59: 92-95.
Received 13 June 2006
Accepted 3 July 2008
2007
NOTES
327
Winter Occurrences of Ivory Gulls, Pagophila eburnea, in Inland
Labrador
Tony E. Cuupss! and FRANK R. PHILLIPS?
'Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and
Labrador AOP 1SO Canada; e-mail: techubbs @cablelab.net
*Department of Natural Resources, Government of Newfoundland and Labrador, Box 175, Station C, Happy Valley-Goose
Bay, Newfoundland and Labrador, Newfoundland AOP 1CO Canada
Chubbs, Tony E., and Frank R. Phillips. 2007. Winter occurrences of Ivory Gulls, Pagophila eburnea, in inland Labrador
Canadian Field-Naturalist 121(3); 327-328.
Ivory Gulls were observed during winter on three occasions up to 325 km inland from open coastal water, feeding on Caribou
carcasses in inland Labrador. Other recent observations in central Labrador are also noted.
Key Words: Ivory Gulls, Pagophila eburnea, range, distribution, Labrador.
The Ivory Gull (Pagophila eburnea) is a plump,
medium-sized gull (40-43 cm long; 108-120 cm wing
span), about 10% larger and longer-winged than the
Black-legged Kittiwake (Rissa tridactyla), with sexes
alike and plumage entirely white in adults (Haney and
MacDonald 1995). Ivory Gulls are High Arctic birds
that have a circumpolar distribution ranging across
northern Canada, Greenland, and arctic western Europe
(Haney and MacDonald 1995; Gilchrist et al. 2005).
The known Canadian population breeds in remote
locations on Ellesmere, Devon, Seymour, northwest-
ern Baffin and Perley islands, Nunavut (Todd 1963;
Haney and MacDonald 1995; Thomas and MacDonald
1987; Stenhouse et al. 2004; Gilchrist and Mallory
2005).
Ivory Gulls, locally referred to as “Ice Partridges”
(Peters and Burleigh 1951) or “Ice-Gulls” (Tuck 1971),
are uncommon winter visitors in coastal Labrador and
northern Newfoundland (Todd 1963). They are rarely
found far from drifting pack ice offshore in winter near
the southern pack ice between Canada and Greenland
(Haney and MacDonald 1995; Stenhouse et al. 2004).
Ivory Gulls appear along the northeast coast of New-
foundland and Labrador in December, usually well
offshore, but they are most abundant in March when
they concentrate on Harp Seal (Pagophilus groenlandi-
cus) Whelping areas in the pack Ice (Tuck 1967, 1971).
Historic inshore observations of Ivory Gulls in New-
foundland and Labrador have occurred along the north-
east coast of insular Newfoundland between December
and March (Tuck 1971) with fewer records along the
Labrador coast (Todd 1963). The largest numbers of
Ivory Gulls recently recorded in our region have been
in southeastern Labrador and northern Newfoundland
following strong northeast gales (P. Linegar, personal
communication). Peters and Burleigh (1951) indicat-
ed that there were a number of reports of the “Ice Par-
tridge” from interior localities in Newfoundland and
one specifically from Badger, but those sightings were
not confirmed. Several adult Ivory Gulls have been
banded at sea off Cape Chidley, Labrador (Thomas and
MacDonald 1987), and a band was recovered in 1940
at Port Hope Simpson, Labrador (52°30'N, 56°10'W)
that was banded in Franz-Joseph Archipelago (North
of Russia) six years previously (Tuck 1967, 1971).
Occasionally, there have been anecdotal observations
of Ivory Gulls at North West River, Labrador, during
winter, where the water remains open (P. Linegar per-
sonal communication). A recent observation at North
West River was of a single immature Ivory Gull on
16 March 2006 (J. Thomas, Department of Natural
Resources, personal communication). An adult and an
immature Ivory Gull were also observed on 10 and 17
February 2007 at Dry Island (53°42.00'N, 59°59.50'W)
near North West River (S. Russell, personal commu-
nication).
The Ivory Gull is designated as Endangered in New-
foundland and Labrador under the province’s Endan-
gered Species Act (NL ESA Regulation 57/2, 2002*)
and in Canada under the federal Species at Risk Act
(SARA Schedule 1) (COSEWIC, April 2006*). Recent
aerial breeding colony surveys in the eastern Canadian
Arctic revealed an 85% decline in numbers of breeding
Ivory Gulls since the early 1980s (Gilchrist and Mal-
lory 2005; Gilchrist et al. 2005).
Our observations were made while travelling along
the Trans-Labrador Highway between Happy Valley-
Goose Bay and Churchill Falls in central Labrador.
This area lies within the Lake Melville-High Boreal
Forest ecoregion and is dominated by Black Spruce
(Picea mariana) with lesser amounts of Trembling
Aspen (Populus tremuloides), White Birch (Betula
papyrifera), Balsam Fir (Abies balsamea) and White
Spruce (P. glauca) in river valleys (Meades 1990*).
This region has the most moderate climate in Labrador
with mean average daily temperatures ranging from
—14 to —18°C in February and +13°C in July. Average
annual precipitation is 1100 mm with an average snow-
fall accumulation of 4 m (Meades 1990*).
On 8 March 2003, F. R. Phillips and F. Taylor ob-
served an adult Ivory Gull along the Trans-Labrador
Highway approximately 5 km west of Metchin River
(53°26'N, 63°18'W). The Ivory Gull was feeding on
Caribou (Rangifer tarandus) remains left by hunters of
328
the migratory George River Caribou herd. The obser-
vation was located approximately 325 km west of the
nearest permanently open coastal water at Rigolet
(54°11'N, 58°27'W), and nearly 400 km inland from
the edge of the coastal ice pack.
On 23 March 2006 we observed an adult Ivory Gull
near the same location as the sighting recorded in
2003. This Ivory Gull was at a campsite adjacent to
the Trans-Labrador Highway, 10 km west of Metchin
River (53°26'N, 63°21'W) where approximately 15
Innu hunters had recently harvested 58 caribou. The
Caribou carcasses, laid on the snow, were intact with
the hide, head and legs attached with only the entrails
removed. The Ivory Gull was feeding on the exposed
flesh and offal of the Caribou adjacent to the road.
The hunters indicated that the Ivory Gull had been
present and feeding on scraps of Caribou meat for sev-
eral days, alternating feeding sites between the road
and the Churchill River, where the animals were har-
vested. On 26 March 2006, F. R. P. returned to the
Innu camp but did not observe the gull. The hunters
indicated that they had last observed the Ivory Gull
on 24 March 2006.
On the morning of 26 March 2006, T. E. C. ob-
served an adult Ivory Gull adjacent to his home in the
community of Happy Valley-Goose Bay (53°18.92'N,
60°22.95'W) feeding on scraps of Caribou meat from
a hunt the previous week. This location is approxi-
mately 195 km east of our sighting at Metchin River,
three days earlier, and was possibly the same bird. We
placed several large pieces of Caribou meat on the
snow to feed the Ivory Gull and to encourage it to stay
for further observation and photographs. At approxi-
mately 1500 hrs on 27 March 2006, an adult Glaucous
Gull (Larus hyperboreus) joined the Ivory Gull and
both fed on the Caribou meat. Both Gulls appeared to
have departed on the evening of 27 March 2006 and
were not observed again.
To our knowledge, our sightings are the first con-
firmed inland observations of Ivory Gulls recorded
for insular Newfoundland or Labrador. These obser-
vations of Ivory Gulls in central Labrador may have
been influenced by the recent southern extension of
the winter range of the George River Caribou herd,
which has occurred during the last two decades
(Chubbs and Phillips 2005). Caribou from the George
River herd have been recorded as far south as 52°10'N
in central regions and may have facilitated the inland
range expansion of other scavengers such as Coyotes
(Canis latrans) into Labrador (Chubbs and Phillips
2005). With the Canadian Arctic breeding population
of Ivory Gulls possibly in peril, and little information
on their life history outside their breeding grounds,
our inland observations add to the knowledge of this
species’ feeding habits and migration stopovers in
Labrador that may provide some insight into the recent
decline of this species in Canada or aid in the recoy-
ery of the population.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Acknowledgments
P. Linegar, G. J. Robertson and the late N. P. P.
Simon commented on earlier drafts of the manuscript.
We thank A. J. Erskine and an anonymous reviewer
for their helpful comments. J. Thomas and S. Russell
provided us with details of their observations. Jonathan
and Kayla Chubbs correctly identified and alerted us
to the presence of the Glaucous Gull. Funding for this
publication was provided through the Department of
Natural Resources.
Documents Cited (marked * in text)
COSEWIC (Committee on the status of Endangered
Wildlife in Canada). 2006. Canadian Species at Risk,
April 2006, Ottawa, Ontario, Canada. http://www.news
wire.ca/en/releases/archive/May2006/01/c9773.html.
Accessed online August 2006.
Meades, S. J. 1990. Natural regions of Newfoundland and
Labrador. Protected Areas Association, St. John’s, New-
foundland and Labrador. 103 pages.
NL ESA (Newfoundland and Labrador, Endangered
Species Act) E-10.1. 2002. http://www.gov.nl.ca/env/wild
life/wildlife_at_risk.htm. Accessed online August 2006.
Literature Cited
Chubbs, T. E., and F. Phillips. 2005. Evidence of range
expansion of eastern Coyotes, Canis latrans, in Labrador.
Canadian Field-Naturalist 119: 381-381.
Gilchrist, H. G., and M. J. Mallory. 2005. Declines in
abundance and distribution of the Ivory Gull, Pagophila
eburnea, in Arctic Canada. Biological Conservation 121:
303-309.
Gilchrist, H. G., M. J. Mallory, and F. Merkel. 2005. Can
local ecological knowledge contribute to wildlife man-
agement? Case studies of migratory birds. Ecology and
Society 10(1) 20[online] URL: http:/www.ecologyand
society.org/vol1O/iss 1/art20/
Haney, J. C., and S. D. MacDonald. 1995. Ivory Gull,
Pagophila eburnea. Number 175 in The Birds of North
America. Edited by A. Poole and F. Gill. The Birds of
North America, Inc., Philadelphia, Pennsylvania.
Peters, H. S., and T. D. Burleigh. 1951. The birds of New-
foundland. Department of Natural Resources, Province of
Newfoundland and Labrador, St. John’s, Newfoundland,
Canada.
Stenhouse, I. J., G. J. Robertson, and H. G. Gilchrist.
2004. Recoveries and survival rate of Ivory Gulls banded
in Nunavut, Canada, 1971-1999. Waterbirds 27: 486-492.
Thomas, V. G., and S. D. MacDonald. 1987. The breeding
distribution and current population status of the Ivory
Gull in Canada. Arctic 40: 211-218.
Todd, W. E. C. 1963. The birds of the Labrador Peninsula
and adjacent areas. University Toronto Press, Toronto,
Ontario, Canada.
Tuck, L. M. 1967. The birds of Newfoundland. Pages 265-
316 in The Book of Newfoundland, Volume III. Edited by
J. R. Smallwood. Newfoundland Book Publishers (1967)
Ltd., St. John’s, Newfoundland, Canada.
Tuck, L. M. 1971. The occurrence of Greenland and Euro-
pean birds in Newfoundland. Bird Banding 42: 184-209.
Received 20 September 2006
Accepted 27 June 2008
2007
NOTES
329
Northern Goshawk, Accipiter gentilis, Exploits a Beagle Hound,
Canis familiaris, as a “Beater” to Catch a Snowshoe Hare, Lepus
americanus
JOHN T. NEVILLE
Wildlife Division, Department of Environment and Conservation, P.O. Box 2007, Corner Brook, Newfoundland and
Labrador A2H 7S1 Canada
Neville, John T. 2007. Northern Goshawk, Accipiter gentilis, exploits a beagle hound, Canis familiaris, as a “beater” to catch
a Snowshoe Hare, Lepus americanus. Canadian Field Naturalist 121(3): 329-330.
During a winter Snowshoe Hare (Lepus americanus) hunt with a beagle hound (Canis familiaris), | observed an adult North-
ern Goshawk (Accipiter gentilis) exploiting the running, barking hound as a “beater” and use strategic locations for attacks
The hawk made three attacks on a hare being pursued by the hound, and subsequently caught and killed it. To my knowledge.
this report represents the first record of a Northern Goshawk using this behavior to capture prey.
Key Words: Northern Goshawk, Accipiter gentilis, Snowshoe Hare, Lepus americanus, beagle, Canis familiaris, beater,
hunting, Newfoundland.
In eastern boreal forests, Spruce Grouse (Dendra-
gapus canadensis), Ruffed Grouse (Bonasa umbel-
lus), Red Squirrel (Tamiasciurus hudsonicus), and
Snowshoe Hare (Lepus americanus) comprise major
components of Northern Goshawk (Accipiter gentilis)
diets (Johnsgard 1990: pages 177-182). Goshawks
winter throughout their breeding range when these
prey species are abundant; however Goshawks in the
northern parts of their range may move further south
when populations of Ruffed Grouse and especially
Snowshoe Hares crash (Palmer 1988; Johnsgard 1990).
In my described observation area, hares were abundant,
squirrel numbers low, and both grouse species, as well
as alternative prey items, very low in abundance (per-
sonal observation).
The observations described occurred in western
Newfoundland (49°05'N, 57°31'W) on 4 December
2004. The area consists of approximately 70% mixed
Balsam Fir (Abies balsamea) Black Spruce (Picea
marina) in various-aged stands and 30% forest open-
ings consisting of small bogs, blowdowns and alder
(Alnus sp.) swales. Approximately 10 cm of snow had
accumulated on bare ground over the previous 10 days.
During a Snowshoe Hare hunt with a beagle hound
I observed unusual hunting behavior by an adult female
Northern Goshawk (sex and age judged by size and
plumage; Palmer 1988). The hawk exploited the run-
ning, barking dog as a “beater”, and positioned itself
into opportunistic attack locations where prey would
likely be flushed ahead of the oncoming hound. The
hawk regularly repositioned itself and selected tree-top
perches (Kenward 1982) ahead of the dog in locations
where it could scan the snow surface. Over approxi-
mately 20 minutes I witnessed three attacks on a hare
actively pursued by my hound, the last resulting in the
hare being caught and killed. With the aid of binocu-
lars, I was able to observe the hawk’s head and its fix-
ation on snow surfaces ahead of the barking hound. I
noted the hawk relocate itself 10 times during the
described hunt.
The first attack occurred after the hawk had posi-
tioned itself on a tree-top perch above a 10 m-wide
alder swale which offered enhanced viewing of the
snow surface from above. As the barking hound pro-
gressed towards the perched hawk, the hare suddenly
appeared in the alder swale and the hawk swooped
down and attempted to catch it. However, dense alders
hindered the bird, and the hare evaded capture. A sec-
ond capture attempt occurred in similar fashion as
the hare ran along the edge of a small bog. The hare
re-entered dense cover before the attacking hawk ap-
proached to within about 8 m. Both of these capture
attempts occurred approximately 40 m from my loca-
tion. The final attack occurred roughly 60 m from my
location in a 10 m by 15 m opening in the canopy. I did
not see the attack but heard the hare emit a distress cry
about 50 m ahead of the barking dog indicating that
it had been caught. After approximately | minute the
hound reached the attack location where its steady
barking changed to intense growling indicating that the
dog was then in the presence of the hawk. I advanced
toward the kill site when from a distance of less than
5 m an adult Goshawk flew off the snow surface car-
rying an intact limp hare in its talons. The bird strug-
gled in flight due to the mass of the hare and it was
assumed that my sudden appearance had caused the
bird to flee with its prey. The snow revealed indications
of the attack: hare tracks, wing tip impressions, hare fur,
blood and an obvious struggle site while backtracking
verified that it was the hare under pursuit of my hound.
Due to the quality of trailing by the hound, the hare’s
running ability and the apparent normal size of the hare.
it is believed that it was in good condition. In addition,
hares that I harvested that day and on following hunts
in the same area were in excellent condition. These
scenes occurred in fairly dense spruce/fir forest with
the likelihood that additional attacks, without my de-
tection, had occurred.
On 11 and 18 December 2004 during hare hunts in
the described area, an adult Goshawk was observed
330
several times in the vicinity of the barking hound which
was trailing a hare; one unsuccessful attack on a hare
trailed by the hound was noted. At no time during these
events did I observe the hawk attempt to strike the dog.
It was also noted that Goshawks had walked along and
followed hare trails on the snow surface (also cited in
Palmer 1988), indicating a strong reliance upon hares
in that area.
The effectiveness of exploiting beaters has been
demonstrated in several species. Cattle Egrets (Bubul-
cus ibis) exploit cattle as “beaters” to flush insects and
small mammals (Welty and Baptista 1988: page
422), and by doing so were more efficient than when
foraging alone (Dinsmore 1973; Grubb 1976). Both
Merlins (Falco columbarius) and Peregrine Falcons
(Falco peregrinus) have followed Northern Harriers
(Circus cyaneus) (Dickson 1984 in Palmer 1988);
and Rough-legged Hawks (Buteo lagopus) have been
observed following Arctic Fox (Alopex lagopus) to
catch disturbed voles and lemmings (Welty and Bap-
tista 1988). Reynolds (1965) suggested that beater
associations may be regular, as with the Cattle Egret,
or may represent an individual capacity to take advan-
tage of a favorable situation. The latter better describes
my observations, which likely originated out of curios-
ity as the Goshawk investigated the dog’s barking and
was rewarded with prey capture opportunities. As far
as I can determine, this is the first report of a Northern
THE CANADIAN FIELD-NATURALIST
Vol. 121
Goshawk or any Accipiter sp. utilizing “beating” behay-
ior to aid in prey capture.
Acknowledgments
I thank R. Perry and R. Otto for helpful suggestions
or an earlier draft, and the helpful comments of two
anonymous reviewers.
Literature Cited
Dinsmore, J. J. 1973. Foraging success of Cattle Egrets,
Bubulcus ibis. American Midland Naturalist 89: 242-246.
Grubb, T. C, Jr. 1976. Adaptiveness of foraging in the Cattle
Egret. Wilson Bulletin 88: 145-148.
Johnsgard, P. A. 1990. Hawks, eagles and falcons of North
America: biology and natural history. Pages 177-182. Edit-
ed by Smithsonian Institution Press, Washington U.K. and
London, U.S.A.
Kenward, R. E. 1982. Goshawk hunting behavior, and range
size as a function of food and habitat availability. Journal
of Animal Ecology 51: 69-80.
Palmer, R. S. 1988. Handbook of North American birds. Vol-
ume 4: Pages 355-378, Northern Goshawk. Yale University
Press, New Haven, USA.
Reynolds, J. 1965. Association between the Little Egret and
African Spoonbill. British Birds 58: 468.
Welty, J. C., and L. F. Baptista 1988. The life of birds, Fourth
Edition. Harcourt Brace College Publishing, Fort Worth,
USA.
Received 9 August 2006
Accepted 3 June 2008
Giant Beaver, Castoroides ohioensis, Remains in Canada and an
Overlooked Report from Ontario
C. RICHARD HARINGTON
Canadian Museum of Nature (Paleobiology), P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada
Harington, C. Richard. 2007. Giant Beaver, Castoroides ohioensis, remains in Canada and an overlooked report from Ontario.
Canadian Field-Naturalist 121(3): 330-333.
The Giant Beaver (Castoroides ohioensis) was the largest ice age rodent in North America, reaching about the size of a Black
Bear (Ursus americanus). In Canada, fossils of this species are commonly found in the Old Crow Basin, Yukon, and single
specimens are known from Toronto, Ontario and Indian Island, New Brunswick. A hitherto overlooked 1891 record of a Giant
Beaver skull from near Highgate, Ontario is the earliest for Canada.
Key Words: Giant Beaver, Castoroides ohioensis, Pleistocene, Highgate, Ontario, earliest Canadian record, paleoenvironment.
The Giant Beaver (Castoroides ohioensis) was the
largest rodent in North America during the ice age
(Quaternary — approximately the last 2 million years)
(Kurtén and Anderson 1980). The great morphological
similarity between Giant Beaver and modern Beaver
(Castor canadensis) leaves no doubt that the two ani-
mals were much alike in appearance and were adapted
to similar surroundings (Figure 1).
But there was one remarkable difference — size!
Adults reached a length of nearly 2.5 m, the size of a
Black Bear (Ursus americanus), and may have weighed
as much as 200 kg (compared to a 1 m-long modern
Beaver weighing about 30 kg) (Kurtén and Anderson
1980). However, Reynolds (2002), treating statistically
a comprehensive sample, estimates Giant Beavers had
a body mass of only 60 to 100 kg while assuming a
length of 1.5 m [this ignores the nearly 2.5 m length of
the skeleton displayed in Chicago’s Field Museum of
Natural History and the 2.2 m length for the Earlham
College skeleton, which represents an immature indi-
vidual (Barbour 1931, Figure 109)]. Using a regression
she established for all rodents, Hopkins (2008) calculat-
ed a mass of 67 kg for Castoroides, which corresponds
to Reynolds’ estimate. Average weights of adult female
Black Bears range from 40 to 70 kg and for adult
males from 60 to 140 kg with body lengths of 1 to 2m
2007
Ficure 1. Locations of Giant Beaver (Castoroides ohioensis) fossil sites in Canada: 1. Old Crow Basin, Yukon Territory: 2.
Highgate, Ontario; 3. Toronto, Ontario; 4. Indian Island, New Brunswick. Insert shows two Giant Beavers (with fox
for scale) in the foreground and Woolly Mammoths (Mammuthus primigenius) in the background. Reduced black
and white image of a painting by George Teichmann (C.R. Harington and N. Rybezynski, scientific advisors).
(Pelton 1982), so Reynolds’ estimate is still within the
range for Black Bears. Other differences include a
relatively narrow tail, cheek teeth with S-shaped
enamel patterns on the grinding surfaces, and cutting
teeth (incisors) about 15 cm long with prominently
ridged outer surfaces (Harington 1996).
A primitive beaver (smaller than the modern beaver)
called Dipoides that occupied Eurasia and North Amer-
ica during the late Tertiary (some 5 million years ago
— indeed members of this genus occupied beaver
ponds in the vicinity of Strathcona Fiord, Ellesmere
Island, about 4 million years ago, Tedford and Haring-
ton (2003), and the Hand Hills, Alberta perhaps even
earlier (Owen and Burns 2006) evidently gave rise to
Procastoroides, a large beaver about two-thirds the
size of the Giant Beaver. That beaver probably
NOTES 33)
+
evolved into the Giant Beaver some 3 million years
ago.
Castoroides ranged from Florida to the Yukon and
from New Brunswick to Nebraska, but it has not
been found outside of North America. Giant Beavers
seem to have flourished in the region south of the
Great Lakes toward the close of the last glaciation,
about 10 000 years ago, and became extinct about
that time (Harington 1996).
In Canada, Giant Beaver fossils have been found in
great abundance in the Old Crow Basin of the north-
ern Yukon (Harington 1977, 1978), and a single inci-
sor was collected from the Don Formation in Toronto
(Coleman 1933; Harington 1978) — fossils from both
places probably extending back to at least last (Sang-
amonian) interglacial time some 130 000 years ago.
An isolated Giant Beaver incisor tooth was recovered
from Indian Island, New Brunswick, near the mouth
of the Bay of Fundy. It may have been deposited on a
storm beach from near-shore deposits (Miller et al.
2000) (Figure 1).
It has recently come to my attention (personal com-
munication, P. F. Karrow 2006) that a much earlier
report from near Highgate, Ontario (approximately
42°30'N, 81°49'W) had been overlooked (e.g. Hay
1923; Harington 1996, 2003). J. H. Panton (Professor
of Natural History and Geology, Ontario Agricultural
College, Guelph, Ontario) on page 4 of an 1891
booklet titled The Mastodon and Mammoth in
Ontario..., noted: “Associated with the remains of
the [Highgate] Mastodon, a skull was found which
the writer has identified to be that of an animal allied
to the beaver, but this animal must have been fully six
feet [about 2 m] in length, instead of two or three
[about 0.61 or 0.91 m] like the Beaver of the present.
One of its gnawing [incisor] teeth in the lower jaw is
eight inches [20.3 cm].” I think that this description is
sufficient to provide a veritable record of the Giant
Beaver. Presumably this report was overlooked
because of the relative rarity of Panton’s (1891) pub-
lished report — the only original copy known to me is
in the rare book room of the Dufferin County Muse-
um at Guelph, and because the specimen has not
appeared in a major public collection. Unfortunately, I
have yet to track down the actual specimen. It is not
with the remains of the Highgate Mastodon (personal
communication, J. Hoganson 2006).
The Highgate report is the earliest from Canada. The
first recorded Giant Beaver remains were found in a
peat swamp near Nashport, Ohio, and were described,
but not named, by S.P. Hildreth (1837). The geologist
J. W. Foster called the specimen Castoroides ohioen-
sis in a publication a year later (Cahn 1932).
Giant Beavers, being well-adapted to swimming,
seem to have preferred lakes and ponds bordered by
swamps as their habitat because their remains have
been found in ancient swamp deposits so often (Har-
ington 1996). American Mastodons (Mammut ameri-
canum) evidently occupied similar habitat in open
spruce forests. Indeed, at Boney Spring, Missouri,
eight mastodon cheek teeth show tooth marks proba-
bly made by the lower incisors of Castoroides (Saun-
ders 1977). It is worth noting that pollen analysis of
sediment recovered from a hole in the sternum of the
Highgate Mastodon (with which the Giant Beaver
skull was associated) indicate that both mastodon and
Giant Beaver occupied a boreal forest dominated by
spruce probably between about 12 000 and 10 300
years ago (Hoganson 2006). Presumably this pollen
analysis was carried out by McAndrews (1994, page
183) who indicates that his “recent analysis of mud”
from the Highgate Mastodon shows that it belongs to
pollen subzone 1b, which in his Figure 10.2 dates be-
tween about 11 800 and 10 000 BP [radiocarbon years
before present (taken as 1950)].
THE CANADIAN FIELD-NATURALIST
Vol. 121
Acknowledgments
I am indebted to the vigilance of my colleague Paul
F. Karrow for noting Panton’s (1891) record of the
Giant Beaver skull from the vicinity of Highgate, On-
tario, and thank John Hoganson for searching for the
Highgate Giant Beaver skull in collections of the North
Dakota Heritage Centre, Bismarck, where the High-
gate Mastodon 1s kept. I thank two anonymous review-
ers for valuable comments that helped to improve this
paper.
Literature Cited
Barbour, E. H. 1931. The giant beaver, Castoroides, and
the common beaver, Castor, in Nebraska. Nebraska State
Museum Bulletin 1(20):171-186.
Cahn, A. R. 1932. Records and distribution of the fossil
beaver, Castoroides ohioensis. Journal of Mammalogy
13: 229-241.
Coleman, A. P. 1933. The Pleistocene of the Toronto region
(including the Toronto interglacial formations). 41‘ Annual
Report of the Ontario Department of Mines 41(7) [1932]:
1-69.
Harington, C. R. 1977. Pleistocene mammals of the Yukon
Territory. Ph.D. thesis, University of Alberta, Edmonton.
Harington, C. R. 1978. Quaternary vertebrate faunas of
Canada and Alaska and their suggested chronological
sequence. Syllogeus 15: 1-105.
Harington, C. R. 1996. Giant beaver. Beringian Research
Notes (6): 1-4. (Published by the Yukon Beringia Interpre-
tive Centre, Whitehorse).
Harington, C. R. 2003. Annotated bibliography of Quater-
nary vertebrates of northern North America — with radio-
carbon dates. University of Toronto Press, Toronto. 539
pages.
Hay, O. P. 1923. The Pleistocene of North America and its
vertebrated animals from states East of the Mississippi and
from Canadian provinces East of Longitude 95°. Carnegie
Institute of Washington Publication (322). 499 pages.
Hildreth, S. P. 1837. Cabinet of the Atheneum of Zanesville.
American Journal of Science 31:79.
Hoganson, J. 2006. The strange and incredible journey of
the Highgate Mastodon. 66" Annual Meeting, Society of
Vertebrate Paleontology (Ottawa, 18-21 October 2006),
Abstracts of Papers, Journal of Vertebrate Paleontology 26,
Supplement to Number 3: 76A-77A.
Hopkins, S. S. B. 2008. Reassessing the mass of exceptional-
ly large rodents using toothrow length and area as proxies
for body mass. Journal of Mammalogy 89: 232-243.
Kurtén, B., and E. Anderson. 1980. Pleistocene mammals
of North America. Columbia University Press, New York.
442 pages.
McAndrews, J. H. 1994. Pollen diagrams for southern
Ontario applied to archaeology. Pages 171-195 in Pro-
ceedings of Great Lakes Archaeology and Paleoecology.
Edited by R. 1. MacDonald. Symposium presented by the
Quaternary Sciences Institute at the University of Water-
loo, 21-23 September, 1991.
Miller, R. F., C. R. Harington, and R. Welch. 2000. A
giant beaver (Castoroides ohioensis Foster) fossil from
New Brunswick, Canada. Atlantic Geology 36: 1-5.
Owen, P., and J. Burns. 2006. New record of Taxidea (Mus-
telidae: Carnivora) from a Miocene deposit in the Hand
Hills, Alberta, Canada. 66" Annual Meeting, Society of
Vertebrate Paleontology (Ottawa, 18-21 October 2006).
2007
Abstracts of Papers, Journal of Vertebrate Paleontology
26, Supplement to Number 3: 108A.
Panton, J. H. 1891. The mastodon and mammoth in Ontario:
Being a scientific description of these gigantic extinct
quadrupeds, and the fossil remains of them discovered by
John Jelly, Esq., Shelburne. Thomas Moore and Company,
Printers, Toronto. 24 pages.
Pelton, M. R. 1982. Black bear (Ursus americanus). Pages
NOTES
333
Reynolds, P. 8S. 2002. How big is a giant? The importance of
method in estimating body size of extinct mammals. Jour
nal of Mammalogy 83: 321-332
Saunders, J. J. 1977. Late Pleistocene vertebrates of the
western Ozark Highland, Missouri. Illinois State Museum
Reports of Investigations 33: 1-118
Tedford, R. H., and C. R. Harington. 2003. An arctic mam
mal fauna from the Early Pliocene of North America
504-514 in Wild mammals of North America: Biology,
management, economics. Edited by J. A. Chapman and G.
A. Feldhamer, Johns Hopkins University Press, Baltimore,
Maryland, USA.
Nature 425: 388-390.
Received 9 January 2007
Accepted 30 June 2008
A Late Born White-tailed Deer, Odocoileus virginianus, Fawn in
Southcentral Wisconsin
CHRISTOPHER N. JACQUES!**, WILLIAM E. ISHMAEL”, TIMOTHY R. VAN DEELEN®, and ROBERT E. ROLLEY'
' Wisconsin Department of Natural Resources, 2801 Progress Road, Madison, Wisconsin 53716 USA
+ Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, Wisconsin, 53706 USA
‘Corresponding author: email: christopher.jacques @ wisconsin.gov
Jacques, Christopher N., William E. Ishmael, Timothy R. Van Deelen, and Robert E. Rolley. 2007. A late born White-
tailed Deer, Odocoileus virginianus, fawn in southcentral Wisconsin. Canadian Field-Naturalist 121(3): 333-335.
Published reports of peak breeding and parturition dates for White-tailed Deer (Odocoileus virginianus) indicate that deer in
northern regions typically breed during November and give birth during late May and early June. However, we report a late born
White-tailed Deer fawn killed by a vehicle between 12-13 March 2007 in south central Wisconsin. Morphology measurements
and body weight indicated the individual was 63-76 days old, was born between 26 December 2006 and 8 January 2007.
and was conceived between 14-27 June 2006. To our knowledge, this observation represents the latest documented breeding
activity in northern deer populations.
Key Words: White-tailed Deer, Odocoileus virginianus, parturition, breeding, conception, Wisconsin.
Seasonally polyestrous breeding activities of north-
ern White-tailed Deer (Odocoileus virginianus) are
clearly synchronized by photoperiod (Verme and UlI-
rey 1984; Verme et al. 1987). However, timing of peak
breeding activities and parturition can be affected by
nutrition and deer herd demographics (Demarais et al.
2000). Low quality nutrition can prolong breeding and
fawning activities, while high quality nutrition can shift
peak breeding and fawning periods to earlier dates
(Verme and Ullrey 1984).
Breeding dates vary throughout the range of the
White-tailed Deer and may occur throughout the year
in Central America or within short (2-3 weeks) time
periods in northern deer populations (Dahlberg and
Guettinger 1956; Moore and White 1971; Verme 1977;
Clark 1981; Webb and Nellis 1981; Ozoga and Verme
1984). Similarly, gestation for deer varies from 196 to
213 days (Cheatum and Morton 1946; Haugen and
Davenport 1950; Verme 1965, 1969), with peak par-
turition occurring in late May to early June in most
northern deer populations (Verme et al. 1987; Brink-
man 2003; Burris 2005).
Although rarely documented, occurrences of late
breeding activity have been reported in northern deer
populations. Throughout southcentral and northern
Minnesota, previous investigations have reported con-
ception dates in captive and free-ranging deer popula-
tions between 3 March and 9 April (Erickson 1952;
Kerr and Peterson 1988; DePerno and Anderson 2000).
However, we were unable to find documentation in the
scientific literature of breeding activity occurring dur-
ing June in northern deer populations. Our purpose was
to report an occurrence of a late-born fawn and subse-
quent evidence of late breeding activity by White-tailed
Deer in southcentral Wisconsin.
On 14 March 2007 a vehicle-killed spotted male
fawn was retrieved from State Highway 23 near the
village of Plain in Franklin Township, Sauk County,
Wisconsin (43°16'30"N, 90°03'00"W). Because High-
way 23 is frequently traveled by WEI, we are certain
this fawn was killed between 12-13 March 2007. At
the time of carcass retrieval, recorded body weight
was 17.7 kg (39.2 Ibs); however, the hindquarters
were slightly scavenged. Because the amount of tissue
removed by scavengers was minimal (s | kg), the esti-
mated body weight of this fawn was 18.7 kg (41 Ibs).
Normally, fawns from well-fed adult females aver-
age 3-4 kg (6.6 to 8.8 lbs) at birth (Verme and Ullrey
1984; Nelson and Woolf 1985: Verme 1989) and typi-
cally gain 0.20 to 0.24 kg (0.44 to 0.53 Ibs) per day
(Robbins and Moen 1975; Verme and Ullrey 1984).
Thus, we assumed the fawn weighed approximately
334
3.5 kg (7.7 Ibs) at birth and gained approximately
15.2 kg (33.5 Ibs) between its birth and death. Using
body measurements and estimated weight gains, we
back-dated from the date of collection, suggesting
the fawn was 63-76 days old and was born between
26 December 2006 and 8 January 2007. Assuming a
gestation period of between 196 and 213 days (Hau-
gen and Davenport 1950; Verme 1965, 1969), concep-
tion likely occurred between 14 and 27 June 2006.
Also, physical examination indicated the fawn was in
good physical condition with obvious white spotting
on a relatively coarse hair coat. Admittedly, we used
typical daily weight gains previously reported for fawns
born during late spring or early summer (i.e., when
nutritious forage is abundant) to estimate age and breed-
ing and conception dates. Given the estimated birth date,
it is possible that energy acquisition was limited and
impeded growth in this individual due to reduced avail-
ability of nutritious food resources and limited milk
production by the lactating mother during winter
months. Consequently, breeding activity and the subse-
quent birth date may have occurred earlier than our
estimates.
Reasons for late conception may be associated with
nutritional limitations or winter weather conditions
(DePerno and Anderson 2000). Although unlikely in
the farmland region of southern Wisconsin, local nutri-
tional limitations may exist. Verme (1965) noted that
adult female deer on low quality diets initiated breed-
ing activities several weeks later than individuals on
higher quality diets. Van Deelen et al. (2007) reported
a female deer from southern Wisconsin carrying five
fetuses and McCaffery et al. (1998) noted that fetal
rates among yearling and adult female deer in south-
ern Wisconsin varied from 1.7 to 2.0 fetuses per preg-
nancy, indicating that high quality food resources are
available throughout the farmland region of southern
Wisconsin.
Late conception may be associated with mild win-
ter weather conditions and deer breeding age. Because
age of the breeding female was unknown, we were un-
able to determine the influence of breeding age. How-
ever, Raedeke et al. (2002) noted lower reproductive
success in older-aged (i.e., = 7 years of age) Elk (Cer-
vus elaphus) while Rosatte and Neuhold (2006) sug-
gested that age of a cow Elk (i.e., 11 years) may have
contributed to her inability to meet annual energy re-
quirements for normal reproduction and subsequent
late conception and parturition dates during their study.
Alternatively, DePerno and Anderson (2000) suggest-
ed that mild winter conditions may have extended es-
trous in a female deer fawn in southern Minnesota,
partly explaining their observation of a late-born fawn.
Verme and Ullrey (1984) suggested that yearling fe-
male deer achieved estrous later than older aged adults
while Cheatum and Morton (1946) and Clark (1981)
noted that fawns were bred one month later than adults.
Also, female deer evidently experience recurrent es-
THE CANADIAN FIELD-NATURALIST
Vol. 121
trous cycles (Plotka et al. 1977). For example, Chea-
tum and Morton (1946) suggested that females were
capable of cycling into January, while Knox et al.
(1988) reported two to seven estrous cycles in adult
female deer and a potential breeding season of 172
days. It is possible that an older-aged female deer
may have been unable to meet annual energy require-
ments for normal reproduction, which contributed to
late conception and parturition observed in the pres-
ent study. However, Woodford and McCaffery (2006)
concluded that mild winter conditions during 2005
did not contribute to excessive mortality or depressed
spring fawn production, and that deep snow conditions
were limited throughout northern Wisconsin and absent
by April when deer were most vulnerable to winter
effects. To this end, it is also possible that a young
White-tailed Deer reached sexual maturity during De-
cember 2005 and experienced multiple (6-8) estrous
cycles before successfully breeding in mid to late June
2006 and consequently giving birth in late December
2006 or early January 2007.
Despite this uncertainty, we suggest that favorable
winter conditions and distribution of farmlands
throughout southern Wisconsin likely enable deer to
maximize growth and reproductive effort at younger
age classes, thus contributing to atypical breeding and
parturition activities. Moreover, Rosatte and Neuhold
(2006) suggested supplemental feeding may have en-
hanced survival of a late-born Elk calf by providing
sufficient energy for the lactating cow to continue milk
production during winter months. We further suggest
that abundant food resources throughout the southern
farmlands minimized potential growth impediments of
this fawn by providing sufficient energy for the lactat-
ing mother to continue milk production during the win-
ter season, thereby minimizing potential biases in our
estimates of age, birth date, and conception dates.
Nonetheless, the sporadic observation of late-born
fawns throughout the farmland regions of Wisconsin
highlights the need for a greater understanding of the
underlying factors that potentially contribute to late
breeding activities by White-tailed Deer throughout the
Midwest.and Great Lakes states. It may be that by
monitoring occurrence of late breeding activities that
wildlife biologists may be able to identify additional
factors affecting deer population dynamics.
Acknowledgments
We thank A. Kraemer and W. Wehrwein for assis-
tance with specimen collection, and L. Ishmael for
assistance with skeletal measurements. We appreciate
helpful comments from T. W. Grovenburg, J. A. Jenks,
K. R. McCaffery, and two anonymous reviewers on
earlier drafts of our manuscript.
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2007
Burris, B. M. 2005. Seasonal movements of white-tailed deer
in eastern South Dakota and southwestern Minnesota rel-
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Cheatum, E. L., and G. H. Morton. 1946. Breeding season
of white-tailed deer in New York. Journal of Wildlife Man-
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tral Wisconsin. Thesis, University of Wisconsin, Stevens
Point, USA. 29 pages.
Dahlberg, B. L., and R. C. Guettinger. 1956. The white-
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DePerno, C. S., and J. R. Anderson. 2000. Documentation
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Erickson, A. B. 1952. A late breeding record for the white-
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Kerr, K. D., and W. J. Peterson. 1988. Late breeding in the
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Knox, W. M., K. V. Miller, and R. L. Marchinton. 1988.
Recurrent estrous cycles in white-tailed deer. Journal of
Mammalogy 69: 384-386.
McCaffery, K. R., J. E. Ashbrenner, and R. E. Rolley.
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261.
Moore, S. A., and C. M. White. 1971. Intrauterine distribu-
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NOTES
335
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and K. D. Keenlyne. 1977. Reproductive steroids in the
white-tailed deer (Odocoileus virginianus borealis). |
Seasonal changes in the female. Biological Reproduction
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Raedeke, K., J. Millspaugh, and P. Clark. 2002. Population
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Robbins, C. T., and A. N. Moen. 1975. Milk consumption
and weight gain of white-tailed deer. Journal of Wildlife
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Rosatte, R. C., and J. Neuhold. 2006. Late-born Elk, Cervus
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Field-Naturalist 120: 188-191.
Van Deelen, T. R., P. Kaiser, and S. R. Craven. 2007.
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Verme, L. J. 1965. Reproductive studies on penned white-
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Monona, Wisconsin, USA.
Received 17 October 2007
Accepted 21 May 2008
Book Reviews
Editor’s Note: We are continuing to use the current currency codes. Thus Canadian dollars are CAD, U.S. dollars are USD,
Euros are EUR, China Yuan Remimbi are CNY, Australian dollars a AUD and so on. You will find these are the codes now
used by financial institutions and internet currency converters. I will include an updated note for the next few issues as a
reminder.
ZOOLOGY
Evolution and Biogeography of Australasian Vertebrates
Edited by J. R. Merrick, M. Archer, G. M. Hickey, and M.
S. Y. Lee. 2006. Australian Scientific Publishing Ltd.,
P. O. Box 68, Oatlands, NSW 2117, Australia. xxviii +
942 pages. 230.00 AUD Cloth, 170.00 AUD. Paper.
This massive tome, covering the current state of
knowledge on how and why the Australasian verte-
brate fauna came to be what and where it is today, is
absolutely absorbing. There is a massive amount of
content, supplemented by numerous illustrations of
continental movements, possible movement corridors,
fossil and extant organisms, phylogenetic trees, and
reconstructions of extinct organisms and ecosystems.
Furthermore, much of the writing, although detailed,
is in an accessible and sometimes humorous form.
The book is subdivided into 38 chapters, each writ-
ten by experts on their particular subject matter. These
chapters are grouped into seven broad categories, five
of them covering the major traditional vertebrate tax-
onomic subdivisions: fishes, amphibians, reptiles,
birds, and mammals. The other two subdivisions pro-
vide introductory context (an introduction to system-
atics, the geological history of the Australasian region,
historical and current climatic influences on evolution
and dispersal, and the links between the biotas of New
Guinea, Indonesia, New Zealand, and Australia) and
a final section entitled “Accelerated Change — the
Regional Future”, dealing with modern techniques in
molecular systematics, tracking of marine movements
of Australasian pinnipeds, and the importance of con-
servation planning for the preservation of the unique
biota of the region. The introductory set of chapters
provides critically important background information
for understanding the relationships of the various land
masses that make up Australasia, how these land mass-
es were arranged within the southern supercontinent
Gondwana prior to its fragmentation in the Jurassic, the
influence of rising and lowering sea levels during geo-
logical history, ancient climates and glaciation, and a
plethora of other environmental attributes that have
been instrumental drivers in the evolution and move-
ments of vertebrates in this part of the world.
Because the focus of the book is on evolution and
dispersal, there is heavy emphasis on the fossil record
and on phylogenetic reconstruction. In virtually all
chapters, there is a review of the fossil record, includ-
ing early hypotheses of origins of the given taxonom-
ic group, as well as the influence of recent discoveries
of fossils, as well as of modern molecular and other sys-
tematic techniques, on revising those earlier hypotheses.
Thus, these chapters provide not only a summary of
what is now known or hypothesized, but also a histori-
cal perspective on how the concepts developed and
changed. There is interesting information in these
chapters for both the specialist and the generalist, and
also for those interested in the history of science. I did
find the introductory chapter on systematics to be some-
what dogmatic, downplaying older approaches and
advocating phylogenetic taxonomy, when I believe that
most systematists now realize that even the older
approaches have something to offer and their results
can be integrated into the more modern approaches.
Nevertheless, on balance, all of the chapters dealing
with the systematics and evolution of the various ver-
tebrate taxa are enjoyable to read and packed with
information.
The authors clearly had fun writing these chapters,
with many of them having a very engaging style. To
give you an idea of some of the interesting chapter
titles and passages in the book, here is a small sam-
ple: “Eons of Fishy Fossils” [Chapter 7], “Furry Egg-
layers: Monotreme Relationships and Radiations”
[Chapter 26], “Humans among Primates” [Chapter
34], referring to Deinosuchus, the largest crocodilian
ever to exist, at 15 m in length “This generalist croco-
dilian could have taken on its contemporary, Tyran-
nosaurus rex, and won” [Chapter 18], referring to a
fossil lizard “Huehuecuetzpalli ... from the early Cre-
taceous of Mexico, surely one of the most frighten-
ing names in all of palaeontology ...” [Chapter 19],
and regarding marsupial origins “A far more com-
plete fossil record will be required before we can
confidently finger any particular taxon as the “origi-
nal” marsupial. Whether North America or Asia is
ultimately honoured as the marsupial “homeland”, a
curious twist to the story is that Gondwana, specifi-
cally Australia, might be the point of origin of placen-
336
2005
tals.” Scattered throughout these chapters, there also are
important messages regarding conservation approaches
and philosophy. In Chapter 23, dealing with island bio-
geography and the patterns of distribution of birds in
Australasia, the author makes the point that conserva-
tion-oriented management must be directed at the entire
landscape, not just the reserves. In various chapters,
there are interesting thoughts on the selection of con-
servation reserves through the use of genetic charac-
teristics of target populations, and the value of corridors
to the movement of organisms across the landscape.
A Wildlife Guide to Chile
By S. Chester. 2008. Princeton University Press 41 William
Street, Princeton, New Jersey, 08540-5237 USA. 392
pages, 19.95 USD Paper.
This is a compact guide with a wide coverage. Not
only does it cover birds and mammals, but includes
reptiles, amphibians, butterflies, moths, marine life and
flora. In addition there are descriptions of the Chilean
landscape and a where-to-watch section. While it does
not cover all wildlife, it does portray those you are like-
ly to see in a typical trip. You get all this for less than 20
USD - a real bargain.
I was puzzled by the illustrations until I read they
were a composite of scanned photos using Adobe
Photoshop and a Cintigq monitor. The result is a pho-
tograph-like illustration arranged in the same format
as a painted version. This works very well as the depic-
tions are clear without the spurious shadows and odd
shading that often plague pure photographs. This man-
ipulation has allowed the author to show creatures at
different angles, highlighting special field marks.
The sections covering birds, mammals, reptiles and
amphibians follow the usual field guide format. The
names are given in English, Spanish and Latin, with
a listing of other names where appropriate. There fol-
lows a description of the key characteristics. This is
essential as not all plumages are illustrated. So you
have to read to know that a female Vermillion Fly-
catcher is radically different from the male. There are
no range maps, but ranges are written and you need
to refer to the country maps at the front of the book.
The plants are presented somewhat differently. The
author has them organised into eight zones and she
describes and illustrates a dozen or so key species in
each zone. Some, like Opuntia, are presented at the
genus level only. In total the book covers less than 100
species or about 2% of Chile’s plants. For example,
there are three orchids out of the fifty species listed
for Chile.
Similarly, about one third of the frogs and only four
snakes are illustrated. This is not as Spartan as it
Book REVIEWS
337
There is so much of interest and value in this book
that it 1s difficult to summarize all of it. Suffice to say
that this book contains a great deal of detail, present
ed in an interesting way, and that anyone interested in
the evolution and biogeography of vertebrates in Aus
tralasia, or in evolution generally, will find something
of value to them in this book. I recommend it highly.
WILLIAM J. CRINS
Ontario Ministry of Natural Resources, Ontario Parks, 300
Water Street, Peterborough, Ontario K9J 8M5 Canada
sounds. On a typical trip I usually see a few of the
commonest species of frogs and I average seeing one
snake for every 10 days in the field. Similar com-
ments can be made about the butterflies and moths.
The author has added numerous sections on basic
natural history [e.g. Life of a frog or butterfly] through-
out the text. While these are informative and fun, I am
not sure they should be in a guide. Typically you want
to compromise between expanding coverage and reduc-
ing weight. Also they tend to be rather curt and overly
definitive. For example, the short explanation on the
characteristics of butterflies and moths says moths fly
by night. Most, but not all do.
The regional coverage includes mainland Chile and
Easter Island, Desventurades Islands, Juan Fernandez
Archipelago and the Chilean Antarctic Territory. The
text includes location, a little history, a physical descrip-
tion and coverage of the key species [especially endem-
ics]. In particular there is a short, but interesting,
coverage of Antarctica.
I believe this would be very useful as a guide in the
field. The coverage is good for mammals and birds
and is adequate for plants and the other animals. The
illustrations show the key characteristics as well as
any other method. Take, for example, the three stocky
skuas, Chilean, Brown and South Polar; the text and
plates do as good a job as can be expected on these
difficult species. While it does not contain as many
juvenile, female and flight illustrations for the small
birds as Jaramillo’s book [Birds of Chile. By A.
Jamarillo. 2003. Princeton University Press, New
Jersey], for the other sections it is really good. The
large birds, such as albatrosses, hawks, and shorebirds
are typically shown in flight. I would prefer to have
depictions of, say, female siskins so I would take this
and Jamarillo’s book on any trip to Chile. For the
Chilean Territories, however, this guide is a must.
Roy JOHN
2193 Emard Cresent, Ottawa, Ontario K1J 6K5 Canada
338
THE CANADIAN FIELD-NATURALIST
Vol. 119
Ecology & Evolution in the Tropics: A Herpetological Perspective
Edited by Maureen A. Donnelly, Brian I. Crother, Craig
Guyer, Marvalee H. Wake, and Mary E. White. 2005.
University of Chicago Press, Chicago, Illinois. x + 675
pages. 45 USD.
This volume contains most of the papers presented
in a symposium held as part of the 2000 joint meet-
ing in La Paz, Baja California, Mexico, of the three
major herpetological professional groups in North
America: The American Society of Ichthyologists and
Herpetologists, the Society for the Study of Amphib-
ians and Reptiles, and the Herpetologist’s League. The
symposium was organized as a tribute to the submis-
sion of the manuscript The Amphibians and Reptiles
of Costa Rica: A Herpetofauna between Two Conti-
nents, between Two Seas (ultimately published in 2002,
University of Chicago Press) by Jay Mathers Savage,
and the latter’s lifetime accomplishments in promot-
ing long-term study of tropical diversity.
The contents are divided somewhat awkwardly but
equally (9 chapters each) between (I) Evolution and
Biogeography and (II) Ecology, Biogeography and
Faunal Studies. The content is heavily weighted (13
of the 18 contributions) on tropical American species,
not surprising, perhaps, considering Central and South
America’s richness in species and diversity. These
American studies range from molecular phylogeny of
caecilians, diversity of Costa Rican salamanders, an
endemic Honduran frog, chromosomal variation in a
group of southern Central American frogs, secondary
sexual characters in tropical frogs, selection and male
reproductive success in a neotropical frog, co-occur-
rence of hylid frogs in a temporary wetland, frog-eat-
ing anurans in the Paraguayan Chaco, biogeography
of the anole genus Norops and of bothropoid pitvipers,
and herpetofaunal analyses of the Rincon Area in
Costa Rica, a Guyanan rainforest, and the Guayana
highlands. The one Old World contribution is a long-
term frog monitoring in Papua New Guinea, The four
primarily overview contributions on a wider geograph-
ic scale are on phylogenetic taxonomy as a replace-
ment for the Linnean system, on snake phylogeny
based on Ribosomal DNA and morphology, elapid
relationships, and theories of snake mimicry.
There are 27 contributors, many well known in the
herpetological literature, others less so (in order of
appearance): Arnold G. Kluge, Marvelee H. Wake,
Gabriela Para-Olea, Judy P. Y. Sheen, David B. Wake,
W. Ronald Heyer, Rafael O. de Sa, Sarah Muller,
Shyh-Hwang Chen, Sharon B. Emerson, Mary E.
White, Maria Kelly-Smith, Brian I. Crother, Joseph
B. Slowinski, Robin Lawson, Harry W. Green, Roy
W. McDiarmid, Karen R. Lips, Craig Guyer, Maureen
A. Donnelly, Norman J. Scott Jr., A. Luz Acquino,
David P. Bickford, Kirsten E. Nicholson, Steven D.
Werman, Roy W. McDiarmid (again), Jay M. Savage,
Maureen A. Donnelly (again), Magan H. Chen, Gra-
ham G. Watkins, Roy W. McDiarmid (again) and
Maureen A. Donnelly (again).
This is not a comprehensive anthology of tropical
diversity of amphibians and reptiles but rather selective
glimpses from a random assortment of studies. Even
as such it samples effectively the range of research
into these in the southern climes and their importance
in perspective of the variation of these groups in the
world. The baseline studies presented are of particu-
lar importance in this era of unprecedented habitat
destruction and climatic change after a period of rela-
tive stability against the background of well publi-
cised observations of amphibian declines and the
more poorly studied condition in reptiles.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P O. Box 3443, Station D, Ottawa, Ontario
Canada
Fossil Ecosystems of North America: A Guide to the Sites and Their Extraordinary Biotas
By John R. Nudds and Paul A. Selden. 2008. The Universi-
ty of Chicago Press, Chicago. 288 pages, 39 USD Paper.
Fossil Ecosystems of North America is a science
book, travel guide, mystery story, and historical saga
in one package — a must-read for any naturalist with
an interest in fossils, evolution, and geological time.
Written by paleontologists John R. Nudds, University
of Manchester, and Paul A. Selden, University of
Kansas, it is a North American sequel to the 2004
Evolution of Fossil Ecosystems, in which the authors
focused on 14 renowned fossil sites around the world.
Their new book features just as many sites, all
located in North America, including three Canadian
locations, one site which spans the Canada-United
States border, and another in the Dominican Republic.
Each site is covered by one chapter, with sub-sections
devoted to its evolutionary context, its discovery and
subsequent research, its stratigraphic setting, biota,
and paleoecolgy, and comparisons with other similar
sites. An appendix, written in true travel guide fash-
ion, provides details on exploring the fossil locations
and visiting related museums.
The Canadian sites, interestingly, contain evidence
of the oldest biota. They include northern Ontario’s
Gunflint Chert, Newfoundland’s Mistaken Point, and,
probably Canada’s best known fossil site, the Burgess
Shale in British Columbia, also a UNESCO World
Heritage Site.
The Gunflint Chert in northern Ontario, discovered
in 1953 by geologist Stanley Tyler, has recently been
dated at a staggering 1878 million years, which places
it in the Paleoproterozoic era. It helps tell the story of
2005
life without oxygen on the planet, of a time when only
bacteria lived here, in rich and unusual diversity.
The evidence of early microbial ecosystems found
at the Gunflint Chert includes filamentous, spherical,
and star-shaped bacteria, as well as the curious and
graceful umbrella-shaped Kakabekia bacteria, with
its spheroidal bulb, slender stipe, and umbrella-like
crown. All the fossils in the book are depicted in a
combination of photographs and drawings.
The fossils at Mistaken Point on Newfoundland’s
Avalon Peninsula were discovered in 1967 by Shiva
Balak Misra, an Indian graduate student studying at
Memorial University. They help fill a critical gap in
the history of life on Earth: the span between the
microbial ecosystems of the Precambrian era, and the
animal ecosystems of the Phanerozoic.
Mistaken Point biota range from disc-like Aspidella,
to frond-shaped Charnia and bush-shaped Bradgatia.
One of the most unusual shapes is Triforillonia, with
its three-lobed body and rounded lobes radiating from
a central rosette, possibly a holdfast or polyp-like
Book REVIEWS
339
organism. The particularly rich array of fossils pre
served at Mistaken Point
thousands of individuals
a census population of
offers valuable snapshots
of living communities at the moment they were
smothered by volcanic ash.
Fossil Ecosystems of North America is a fascinating
book, easy-to-read, with highly comprehensible scien-
tific explanations, extensive details, and helpful maps,
photographs and drawings. It is, essentially, a rivet-
ing mystery story about life on this planet, filled with
cataclysmic events, extinctions, takeovers, and human
quests for clues and explanations, often involving con-
flict and heated debate. At another level, Fossil Ecosys-
tems of North America 1s a science book for students
and an interested lay readership. At yet another level,
it is an unusual and informative “time travel” guide.
Three books in one — a valuable addition to the
bookshelves of any naturalist curious about deep
ne RENATE SANDER-REGIER
3, 11" Line, Bristol, Quebec JOX 1GO Canada
Headless Males Make Great Lovers and Other Unusual Natural Histories
By Marty Crump. 2005. University of Chicago Press, Chicago,
Illinois, USA. 199 pages. 25.00 USD Cloth, 14.00 USD
Paper.
Marty [Martha] Crump, herpetologist, author of Jn
Search of the Golden Frog (University of Chicago
Press) and Amphibians, Reptiles, and Their Con-
servation, has compiled a remarkable collection of
capsule glimpses of diverse natural history behaviour.
Five section headings set the text style: Ain't Love
Grand, The Mamas and the Papas, Eat to Live and
Live to Eat, Don’t Tread on Me, Ya Don’t Say, but the
serious reader should not be put off by this flippant
approach — its aim is merely to grab the attention of
readers who would not normally pick up a natural his-
tory book. The science is authoritative and reliable.
The detailed documentation of the scientific litera-
ture, on which the text is based, is contained in a 10-
page concluding section arranged by topic with simi-
larly innovative headings: Rampant Machismo, Come
Up and See My Etchings, Sneakers and Deceivers,
Survival of the Pampered, Nests Aren’t Just for the
Birds, Babies on Board, A Pouch Full of Miracles,
Stomping for Worms, A Team Effort: How (Some)
Ants Get Food, Chameleons of the Sea, Tears of Blood,
Casting the Insides Outside, Spit ‘n’ Spray, Living
State of North America’s Birds of Prey
By Keith L. Bildstein, Jeff P. Smith, Ernesto Ruelas Inzunza,
and Richard R. Veit. Nuttall Ornithological Club and The
American Ornithologists’ Union, Series in Ornithology
(3) 2008. Softcover. 466 pages, 98 figures, 42 tables, 6-
page glossary.
This is the third volume in a promising and com-
mendable new series of ornithological publications,
Flashlights, Rapturous and Rapacious Reptiles, Smell-
ing is Understanding, Love Potion Number Nine, It
Pays to be Neighborly. This is followed by a seven-
page index of organisms in the text.
This is a book ideally suited for idle moments or
bedtime reading, the entries are as short and punchy
as the section headings. The multitude of observations
are gleaned and summarized from an amazingly vast
diversity of research by ethologists and naturalists
around the world. The title comes from the generally
well-known mating of certain spiders where the female
decapitates the male while he is still in the process of
copulation. But the text covers a vast array of other
behaviours, such as the stomping of Wood Turtles to
bring the earthworms cherished as food to the sur-
face. It is a remarkable testament to the seemingly
endless variety of life styles and unique behavioural
innovations present in the animal world, and is highly
recommended to all naturalists for both education and
amusement. The text is enlivened by sketches by Alan
Crump.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4
Canada
each appreciably larger than the AOU Monographs
of long standing.
The book begins with a succinct history of raptor
conservation by senior author Keith Bildstein. He tells
the history of bounties as a method of raptor control.
Between 1917 and 1952, Alaska paid bounties rang-
ing from 50 cents to two dollars on over 128 000 Bald
340
Eagles, in spite of the bird being the American national
emblem. Bounties were an expensive method of con-
trolling predators; for example, farmers in Pennsyl-
vania may have saved only one dollar’s worth of chick-
ens for each $1205 in bounties paid by the state.
Most members of the public viewed hawks and
owls as enemies to be shot on sight. “The only good
hawk is a dead hawk,” people would rationalize. In
Pennsylvania, so-called sportsmen thronged to spe-
cific sites along the margin of Hawk Mountain to
engage in wholesale killing of most species of raptor
that flew by.
Those who believe that a single person can do little
to change conservation practices will be impressed
by the story of Rosalie Edge, who purchased a portion
of Hawk Mountain that included the shooting sites.
She then hired a warden to police the land. Against
intense local opposition, the land was posted and
hunters were deprived of the best place in all Penns-
ylvania to shoot the state-bountied Northern
Goshawk. Because of Rosalie’s vision and determina-
tion, Hawk Mountain has become a major educational
and conservation center, increasing in importance each year.
This book lists each of the growing number of rap-
tor monitoring sites, more than one hundred, where
keen volunteers flock to help. The greatest effort is
given to the fall migration. The “Veracruz River of
Raptors” has posted numbers as high as 5.26 million
in a single fall migration. There, the top four species
are the Turkey Vulture, just shy of 2 million, Broad-
winged Hawk at 1.92 million, Swainson’s Hawks up
to 915 000 and Mississippi Kites an impressive
155 000. A million Turkey Vultures have also been
counted at Kekoldi in Costa Rica.
This book is replete with maps, graphs and tables.
Seven excellent maps, for example, together demon-
strate the impressive band recoveries for Sharp-
shinned Hawks. Movements of many other raptors,
however, are less well known and knowledge of their
population levels is limited.
Raptor monitoring in Canada receives appropriate
attention. Since hawks only come together in increas-
ing numbers as they move south, they do not coa-
lesce into large flocks in some parts of Canada. An
exception are the 107 000 Broad-winged Hawks
counted during the 1996 fall migration at Holiday
Beach near Windsor, Ontario. There are three sites in
Quebec where over 1 000 of one raptor species may be
THE CANADIAN FIELD-NATURALIST
Vol. 119
counted in a single day. Ontario has ten sites, includ-
ing Hawk Cliff, and there are two each in Manitoba
and British Columbia. In Alberta, at Windy Point,
manned since 1967, over one thousand each of
Sharp-shinned Hawks and Golden Eagles may be
counted passing by. At nearby Mount Lorette, the
Golden Eagle count can reach the neighbourhood of
4700. Canadian contributors to the book include
David Hussell, a pioneer in developing the Raptor
Population Index, a new counting method that aims
for more consistency in raptor counting. Erica Dunn
is the lead author for the Watchsite Methods chapter.
While insect-eating and ground-nesting birds con-
tinue to decline across North America, there is more
good news than bad from the raptor front. While
Northern Harrier and American Kestrel are diminish-
ing at many sites, Swainson’s Hawks, Bald Eagles,
Peregrine Falcons and Turkey Vultures are generally
increasing. The ground-nesting harrier is at risk from
many predators, but it makes less sense that the
kestrel, with relatively safer nest sites in boxes and
cavities, should also be in decline. The impact of
habitat changes, weather and food availability is being
studied for many species.
There are minor defects in this compendium. There
is a misleading absence of “Diurnal” before “Birds of
Prey” in the title. Owls are not included, although the
four-year cyclical irruptions of the Great Gray Owl
and Northern Hawk-Owl can be dramatic. We are told
in one sentence that irruptive migrations of the
Northern Goshawk “occur periodically in years of
low prey availability” but we are not told that this is
part of a general 10-year cycle involving hare, lynx
and Great Horned Owls. In the section on types of
migration, “irruptive migration” should have been
added to “loop,” “altitudinal” and “leap-frog” migra-
tions. The book lacks a detailed index.
Despite these minor flaws, this book belongs in the
library of every raptor enthusiast and in every univer-
sity and college library. Some chapters warrant reading
in their entirety and others are excellent reference
material. Anyone with the slightest interest in the rap-
idly growing field of hawk monitoring should read
the salient chapters, and become aware of the con-
tents of the others.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N OJ8
Canada
Owls of the United States and Canada: A Complete Guide to Their Biology and Behavior
By Wayne Lynch. 2008. University of British Columbia
Press, 6344 Memorial Road, Vancouver, British Colum-
bia V6T 1Z2. 242 pages 39.95 CAD Cloth.
Wayne Lynch is an accomplished photographer
and writer, I had no expectations that his newest book
would represent anything less than the standard for
which he is known.
The list of “things I would do differently” is quite
small, but worth mentioning. The title, A complete
guide... strikes me as being somewhat pompous...
we know so much about owls that offering up a sin-
gle tome as being somehow complete is a bit much.
Then there’s the tank of information yet to be discoy-
ered! Secondly, I'd be willing to bet that the vast
2005
majority of research that was done by the original
biologists was reported in metric. However, all of that
was converted to imperial, rounded off and reported
as such; parenthetically, the imperial values were
reconverted and oft-reported as inappropriately-accu-
rate metric values again, which muddles the original
values and makes the reading stumbly. Both the bird-
ing and ornithological literature have long used a con-
vention of capitalizing bird names; this avoids some
potential confusion when, for instance, referring to
“the Northern Hawk Owl” (the species) versus “the
northern hawk owl” (the hawk owl found more norther-
ly than others). I find newspapers reticent to adopt this
standard, but there’s no reason for this book to have
done so. Those well-read in bird literature will do men-
tal hiccups frequently while reading this book.
Chapters often start with a journal entry or descrip-
tion of an individual encounter with owls that beauti-
fully sets up the chapter’s theme. By doing this, Lynch
maintains a personal connection with readers, which
then encourages them to continue reading the more-
matter-of-fact, but still very well-written material later.
There is a nice 10-page identification guide to the
owls, with range maps and a synopsis of information
such as habitat preferences, life span, diet and status.
Although the text is good, the maps are deceptive.
Unlike most field guides, the maps here use only one
colour to report the species’ ranges. And if you didn’t
know any better, you’d think the Snowy Owl bred
everywhere in Canada, and into the northern tier of the
United States (which it doesn’t). Johnsgard’s book on
North American Owls does more justice to the ranges.
The rest of the book is written topic-by-topic, much
like Duncan’s Owls of the World. Other owl books,
like Mikkola’s Owls of Europe, take a more species-
by-species account, with a few introductory chapters
Owls of North America
By Frances Backhouse. 2008. Firefly Books Ltd, 66 Leek
Crescent, Richmond Hill Ontario L4B 1H1. 215 pages,
34.95 CAD Cloth.
Owls of North America, by Canadian writer Frances
Backhouse, is a beautiful book — literally. The first
thing I did when it came into my possession was to
look at the photographs. A Great Horned Ow! silhouet-
ted against a sunset. A close-up of the same species’
primary flight feathers. An Elf Owl delivering a cricket
to its cactus nest. The large eyespots on the back of a
Northern Pygmy-Owl’s head. Great Gray Owls
engaging in mutual preening. A young Long-eared
Owl hanging upside down from a twig. A Short-Eared
Owl stretching its leg as a warm-up to hunting.
I would have been thrilled with the book for the
photographs alone. When I started reading, it got even
better. Eight chapters discuss topics covering a wide
range: owl and human relations, owl family groupings.
life trajectories, feeding habits, communication, adap-
tations for nocturnal life, and more. Almost 80 pages
Book REVIEWS 34]
on topics like food, anatomy, movements, etc. I don't
think one format can be said to be better than the
other in general, each excels for different reasons
The approach adopted by Lynch makes for a more
coffee-table friendly book; different species can be
depicted in any one chapter, making comparisons
among, for instance, insectivores to carnivores, readi-
ly apparent.
Throughout the book are sidebars containing syn-
opses of topics related to the chapters’ contents. I’ve
always liked this approach as it allows for informa-
tion that is otherwise not readily incorporable to be
presented. In this book, the topics range from the pH
scale (used to discuss digestion) to brooding behav-
iour.
Finally...the photographs. Dr. Lynch is perhaps best-
known for his stunning photography. There are some
awe-inspiring habitat shots, and double-page spreads
that leave the reader with a sense that “I must visit that
place.” The shots of the owls themselves rate with the
best I’ve ever seen and include depictions of some
behaviours that are not frequently seen. Not to dimin-
ish the writing at all, but the photographs really make
this book what it is.
Literature Cited
Duncan, J. Owls of the World: Their Lives, Behavior, and Survival
2003. Firefly Books, 66 Leek Crescent, Richmond Hill Ontario
L4B 1H1 319 pages
Johnsgard, P. 1988. North American Owls: Biology and Natural
History. Smithsonian Institution Press, Washington. 298 pages
Mikkola, H. 1983. Owls of Europe. Buteo Books, Vermillion. 397
pages
RANDY LAUFF
St. Francis Xavier University, Antigonish, Nova Scotia B2G
2W5 Canada
are devoted to relatively comprehensive species pro-
files that provide photographs, maps, and details on
appearance, voice, activity, roosting, distribution, habi-
tat, feeding, breeding, migration, and conservation. A
glossary and bibliography complete the information
offerings.
The book brought alive owl species I was only
vaguely aware of. The Elf Owl of the southwest, the
world’s smallest owl, which feeds mainly on insects.
The Flammulated Ow! of the west, also an insect-eater,
with ventriloqual abilities that make singers difficult
to locate. The Vermiculated Screech-Owl of Mexico.
Central and South America, which sings like a toad.
The book also deepened my understanding of owls
which are more familiar to me. The Great Horned Owl.
that consumes a greater diversity of prey — from
grasshoppers to great blue herons — than any other
North American owl. The Great Gray Owl, which is
benefiting from nesting platforms; one study revealed
that nesting success was higher on platforms than natu-
ral nest sites.
342
The Eastern Screech Owl which, in Texas, brings
live Texas blind snakes to its nest to eat the soft-bod-
ied larvae of insects attracted to pellets, fecal matter,
uneaten food, and occasional dead nestling accumu-
lated in the cavity. The Burrowing Owl, which produces
a sound remarkably like a rattlesnake’s rattling buzz
— a sound that may have evolved to deter predators
such as badgers and weasels from entering nest holes.
It has also, the author writes with a twist of humour,
“definitely proven effective on occasion in discourag-
The Tree of Life: A Phylogenetic Classification
By Guillaume Lecointre and Herve Le Guyader. 2006. The
Belknap Press of Harvard University Press, Cambridge,
Massachusetts. 560 pages. 39.95.USD.
Here is a textbook to usher in the 21st Century
based on the use of cladistics and the steadily increasing
data from molecular biology. These have been gradual-
ly changing systematic biology over the past 30
years. It may not be the last word in this dynamic field
but it certainly is in the current mainstream. It presents
a genealogical approach to classification of the diver-
sity of all life on earth through the phylogenetic analy-
sis increasingly promoted by many researchers as the
successor to the Linnean System. The latter was out-
lined for plants in 1735 and for all organisms in 1758
and thereafter long universally adopted and taught
throughout the world. The Linnean system accepted
the concepts of its time — divine creation and the
Great Chain of Being, each individually created
group of organisms, from single celled to human, con-
ceived of as progressively “higher” with Homo sapi-
ens as the culmination. After a hundred years, in 1858,
Charles Darwin outlined descent with modification by
natural selection which showed how evolution could
occur. Shortly after, in 1866, Ernst Haeckel coined
the term phylogeny but it was nearly a hundred years
later before Willi Hennig outlined the comprehensive
application of an objective cladistics approach (phylo-
genetic systematics) published in German in 1950
and in English in 1966.
The Tree of Life outlines major groupings for the
present estimated | 749 577 described and currently
recognized organisms. After a preface and introduc-
tion there are 15 chapters. The initial one on “Life” is
followed by Eubacteria, Archaea, Eukaryotes, Chloro-
bionta, Embryophyta, Metazoa, Protostomia, Moll-
usca, Eurarthopoda, Deuterostomia, Sarcopterygii,
Mammalia, Primates, Actinopterygii. Some group
names are familiar from traditional classifications,
some will be new and strange to most naturalists.
Abolished are familiar classification categories (King-
dom, Class, Order, Family) to be replaced by dia-
THE CANADIAN FIELD-NATURALIST
Vol. 119
ing ornithologists from reaching blindly into bur-
rOWS.”
Owls, the book clearly communicates, are fascinating
earth residents with unique skills and adaptations that
equip them for important ecological roles. As such, they
deserve admiration, respect, and help in reversing the
habitat loss from which virtually all owl species suffer.
RENATE SANDER-REGIER
3, 11" Line, Bristol, Quebec JOX 1GO Canada
grams (dendrograms) depicting relationships. Also
vanished are what are now regarded as mixed group-
ings such as reptiles and fish.
Each chapter is subdivided and a chart presented
of the relationships of the included forms discussed.
Each grouping has line drawings of example organ-
isms and features, a brief general description of includ-
ed forms and sections on ecology, some unique
derived features, and examples of included forms. A
sidebar gives number of species, oldest known fos-
sils, and current distribution.
The book concludes with 13 Appendices of “trees”
showing the cladistic view of relatedness, and includ-
ing “Where are they” giving the new positions of
major traditional groups of both one-celled and multi-
celled life, Sequenced Genomes (listing organisms for
which the genome is entirely sequenced or almost so
as of January 2006), a one-page General Bibliography,
a Glossary of 70 entries from Alignment to UPGMA
(Unweighted Pair Group Method Using Arithmetic
Average), Index of Common Names and Index of
Latin Names.
The intent of the volume is to move this phyloge-
netic approach to classification out of its currently
restricted academic position and bring it into the
mainstream to replace the traditional teaching now
rendered archaic, at least in the view of the presenters.
The authors are well suited to the task, throughly
immersed in the modern concepts and teaching.
Lecointre is Professor and Research Scientist at the
Museum National d’Histoire Naturelle and Le
Guyader is Professor of Evolutionary Biology at the
University of Pierre and Marie Curie, both in Paris,
France. The book was originally published as Classi-
fication phylogenetique du vivant (third edition, Edi-
tions Belin, Paris, 2006) and translated by Karen
McCoy. It is illustrated by Dominique Visset.
FRANCIS R. CooK
Emeritus Curator and Researcher, Canadian Museum of Na-
ture, PO. Box 3443, Station D, Ottawa, Ontario K1P 6P4
Canada
2005
Book REVIEWS 343
Biology of the Snapping Turtle (Chelydra serpentina)
Edited by A. C. Steyermark, M. S. Finkler, and R. J. Brooks.
2008. The Johns Hopkins University Press, 2715 North
Charles Street, Baltimore, Maryland 21218-4363. x +
225 pages 75 USD.
The Biology of the Snapping Turtle consists of 17
review papers on various aspects of the life history of
one of the most recognizable turtles of North America.
Part 1, Taxonomy and Systematics, consists of four
chapters covering the systematics of the family Chely-
dridae, the fossil history of the family, the anatomy of
the skull, and molecular insights into the systematics of
the family. Part 2, Physiology, Energetics, and Growth,
consists of seven chapters covering such disparate top-
ics as respiration, reproductive physiology, thermal
ecology, energetics, embryology, overwintering adapta-
tions and growth patterns. The final section of the book,
Life History and Ecology, consists of six chapters on
nesting ecology, water relations of the eggs, sex deter-
mination, physiology of hatchlings, population biology
and geographic variation in life history traits.
Although many of the chapters are strictly review
papers, a number of the authors have included previ-
ously unpublished data from their own work or the
work of others. For example, H. Bradley Shaffer and
his co-authors include their analysis of the genetic
variation across the currently recognized four sub-
species of Snapping Turtle. Their results are consis-
tent with earlier less extensive work: first of all, there
is no molecular evidence to support the recognition
of the Florida Snapping Turtle C. s. osceola) as a dis-
tinct taxa; second of all, the Central American Snap-
ping Turtle C. s. rossignonii) and the South American
Snapping Turtle C. s. acutirostris) both appear to be
separate evolutionary lineages and should be viewed
as separate species.
While many of the papers in this volume are tightly
focussed on the Snapping Turtle, a number of the
papers could just as easily have appeared in a volume
on the biology of turtles in general. For example,
Gordon Ultsch and Scott Reese's paper on overwin-
tering is a thorough review of the literature on turtle
physiology with regards to hibernation.
Overall, the quality of the papers is high. One of the
highlights of the book is the chapter on nesting ecolo-
gy written by researchers involved in three long-term
studies of Snapping Turtles at Algonquin Park,
Ontario; the Edwin S. George Reserve, Michigan; and
the Savannah River Site, North Carolina. There are
also a few careless mistakes in the book that more
careful editing could have caught. The Wood Turtle
(Glyptemys insculpta) and the Bog Turtle (G. muh-
lenbergii) are still placed in the genus Clemmys in the
text and in the index. There is also more overlap in
material than is required. Growth of the embryo is dis-
cussed in both the chapter on the embryo and the chap-
ter on growth. The relationship between clutch size
and body size is thoroughly charted in the chapter on
nesting ecology, discussed in the chapter on popula-
tion biology and re-visited in the chapter on geo-
graphic variation. The editors are also strangely silent.
There is no introduction from them explaining the
origin or rationale for the volume or any kind of con-
cluding chapter synthesizing the themes of the vol-
ume. Nonetheless, anyone with a technical interest in
the biology of turtles (not just Snapping Turtles) will
find this an extremely valuable reference work.
DAVID SEBURN
Seburn Ecological Services, 2710 Clarenda Street, Ottawa,
Ontario K2B 7S5 Canada
Wildlife of North America: A Naturalist’s Lifelist
By Whit Bronaugh. 2006. University Press of Florida,
Gainesville, Florida, USA. 565 pages. 29.95 USD.
This is a bold and daring compilation that few
would have the courage to undertake and present. It is
a contribution to the attempt to promote systematic
record keeping by the vast number of casual or non-
academic naturalists. The volume is essentially a
checklist of all native and introduced mammals, birds,
reptiles, amphibians, freshwater fishes, butterflies,
dragonflies and damselflies recorded to date for
North America. These groups include relatively large
and often conspicuous animals for which a naturalist
can identify a significant number of species on sight.
It is based on the latest (at the time of publication)
authoritative listing for each group with the full real-
ization that work is ongoing for all group names and
species recognition is constantly changing with each
new study that produces new knowledge that necessi-
tates revisions. Despite this, probably enough stability
has been reached in each included group that names
used here will still be able to at least be equated with
those that will be current in another decade or two
hence.
The contents include a Preface (the author's pur-
pose and how he came to compile the book and his
acknowledgments), a Checklist of North American
Orders and Families for the included groups, A List
of Symbols (indicating species extinct and since
when), an Introduction to the main text, How to Use
this Book, Biodiversity and Zoogeography of North
America (with maps of number of species in each
province and state) and Extinct Species. The bulk of
the book follows (pages 83 to 430) listing English
and scientific name for each species by family and
blank spaces for user remarks such as first observa-
tion date, place or other data considered relevant. The
344
are additional blank entry spaces at the end of each
group section for addition of species defined after the
publication of this book. The book concludes with an
Appendix of recent taxonomic and nomenclatural
changes which depart from standard field guides
(pages 431-478), Works Cited (pages 469-478), Index
to Scientific Names (479-528), and Index to Common
Names (pages 529-565). The last two are particularly
valuable for equating a name used in older field
guides to the name recognized currently.
What this is not is a field book. It is far too large and
bulky to be carried on outings. It is a book for post-
expedition entries. It is devoid of keys or text on iden-
tifications, these are left to consultation of the many
guides listed in the references. But even naturalists
who may purchase the book with the best of, but later
unrealised, intentions, will find it a valued quick refer-
ence for their library for its analysis of North America
distribution patterns, comparative species totals, and
post-Pleistocene zoogeography, among many other con-
BOTANY
Grasses of Colorado
By Robert B. Shaw. 2008. University Press of Colorado,
5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado
80303 USA. xi + 647 pages. 75 USD Cloth.
The grasses constitute one of the most important
families of flowering plants in the world, both eco-
nomically and ecologically. They are extremely
diverse and generally comprise about one-tenth of the
vascular flora in many parts of North America. They
are dominant in many ecosystems, including several
of the most extensive ecosystems in Colorado.
Grasses of Colorado provides a thorough, up-to-date
treatment of the 335 species of grasses known to
occur outside of cultivation in the state. The book is
arranged in several sections. The introductory section
deals mostly with utilitarian aspects of grasses,
including food and forage, soil conservation, turf and
ornamental uses, and harmful grasses. The ecological
significance of grasses receives rather limited discus-
sion, despite the prominent role of the family in the
state’s major ecosystems. Following the introduction
is a section on the physiography and ecoregions of
Colorado which is informative and useful in under-
standing the distribution patterns of the grasses in the
state. Maps outlining the major landforms, river sys-
tems, mountain ranges, and ecological subdivisions
assist considerably in understanding these patterns.
This section is followed by a chapter on grass mor-
phology which contains good illustrations that cover
most of the important features that are used in grass
identification. These illustrations are particularly
helpful when coupled with the illustrations of each
species that are interspersed in the main taxonomic,
descriptive part of the book.
THE CANADIAN FIELD-NATURALIST
Vol. 119
siderations, all carefully compiled from recent author-
itative sources.
Whit Bronaugh now lives in Oregon and makes a
living as a nature writer and photographer, but he grew
up among the carefully manicured horse farms of cen-
tral Kentucky and did not really discover nature until
a university field course in Africa. He has remained in
awe of the diversity of animal life ever since and, as an
initial lifelist of birds expanded to include other groups,
the concept to this book was generated initially to fill
his own needs. His other work has appeared in popu-
lar periodicals such as Wildlife Conservation, National
Geographic, Natural History, and Smithsonian. His
photography presented here is restricted to a few small
black-and-white reproductions in the book itself and
colour ones on the attractive dust jacket.
FRANCIS R. CooK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario
K1P 6P4 Canada
The bulk of the book is comprised of the taxonom-
ic treatments, including keys, descriptions of subfami-
lies, tribes, genera, and species, illustrations, and range
maps depicting counties of occurrence. The species
descriptions are quite detailed, including the impor-
tant floral and vegetative characteristics. A concerted
effort has been made to ensure that all descriptions are
parallel, not just between species within a genus, but
across all genera. Each species treatment also con-
tains information on major synonyms, common
names, origin (native vs. alien), a general habitat
description, and a section for comments on status in
the state, taxonomic issues, and similar species. The
accompanying illustrations are reproduced with per-
mission from the two grass volumes of the Flora of
North America (Barkworth et al. 2003, 2007). These
generally are excellent and, when used in conjunction
with the keys and descriptions, should enable success-
ful identification. On occasion, I found the comment
subsection in the species treatments to be redundant
(same information repeated for several species within
a genus), but more often than not, there was useful
supplementary information there.
In general, Shaw follows the taxonomic concepts
espoused by other grass experts, as reflected in the
Flora of North America. However, there are a few
departures, such as the segregation of Melica into two
genera, Melica and Bromelica, the segregation of
Bromus into four genera, Anisantha, Bromopsis, Bro-
mus, and Ceratochloa, corresponding to the sections
of Bromus in most other North American treatments,
and the separation of Critesion from Hordeum. Most
of the rest of the taxonomic treatment will be familiar
2005
to those who follow grass taxonomy, including the
segregations and realignments of species within genera
in the tribe Stipeae.
The content of the book generally is of high quali-
ty, and, as much as is possible with a group of plants
that has its own set of descriptive terminology, the
text and keys are readable with jargon minimized.
There are occasional inconsistencies in the text; for
example, in the treatment of Digitaria sanguinalis,
under Origin, the species is said to be native, but in
the Comments section, it is stated to be “a European
species now established as a global weed.” I detected
relatively few proof-reading errors (e.g., synonym not
italicized, punctuation misplaced, rare spelling
errors), and these do not detract substantively from
the book.
This book provides a welcome updated treatment
of the grasses of Colorado. In spite of the fact that
there is a recent North American taxonomic treatment
The Macrolichens of New England
By James W. Hinds and Patricia L. Hinds. 2007. Memoirs
of the New York Botanical Garden Volume 96. New York
Botanical Garden Press, Bronx, New York. 608 pages,
65.00 USD Cloth.
The most recent publication on lichens from the
New York Botanical Garden is the first detailed treat-
ment of New England macrolichens. Before the reader
even opens The Macrolichens of New England the 14
beautiful photos on the front and back covers give the
sense that it is something special. This much-awaited
guide, drawing on 35 years of collected photos and
knowledge of the lichenologist team of James and
Patricia Hinds, describes and illustrates the lichen flora
from a very unique region rich in lichen flora. The
Hinds share with the reader a plethora of information
ranging from specific species treatments to general
lichen biology so it appeals to the serious professional
and novice alike.
A quick key index inside the front cover provides
the reader with direct access to a group or classifica-
tion of their choice; a useful tool allowing this book to
be used strictly as a field guide. The front sections
should not to be overlooked as they provide, in much
the same way as I. M. Brodo, S. D. Sharnoff, and S.
Sharnoff (2001) Lichens of North America, 70 pages
of useful information that will inform and help the
reader to explore the world of lichenology. These early
chapters provide essays on anatomy, ecology, lichen
conservation and population trends, and identifica-
tion. The general key section follows the background
(approximately 30 pages) which will direct you into
the genus descriptions (some 400 pages). The exten-
sive references cited, glossary, two appendixes, and
index, organized by Latin name, round out the book.
A total of 502 species is covered, including 308 with
colour photos. Some of these species are extremely
Book REVIEWS 345
of the family, there will always be a need for regional
and local treatments, especially for large and diverse
families such as this. The book should prove to be
useful for the identification of grasses in several adja
cent states, including much of the upper Midwest,
from Montana to North Dakota and south to Kansas. It
should also be useful in the southern portions of the
Prairie Provinces.
Literature Cited
Barkworth, M. E., K. M. Capels, S. Long, L. K. Anderton, and M.
B. Piep, Editors. 2007. Flora of North America North of Mexico
Volume 24. Magnoliophyta: Commelinidae (in part): Poaceae
part 1, Oxford University Press, New York. xxviii + 911 pages
Barkworth, M. E., K. M. Capels, S. Long, and M. B. Piep, Editors
2003. Flora of North America North of Mexico. Volume 25
~
Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford
University Press, New York. xxv + 783 pages
WILLIAM J. CRINS
Ontario Ministry of Natural Resources, Ontario Parks, 300
Water St., Peterborough, Ontario K9J 8M5 Canada
rare, not typically described in literature of this type.
Included are species found in adjacent localities, which
could occur in the region of study. Microlichens are
briefly discussed in the introductory chapters but are not
treated with the same detail as the macrolichens. Many
of the species discussed have coastal affinities, with
many others found inland, making this book a great
resource for readers interested in species across the
entire continent.
For the most part the photos are stunning, and
beautiful, capturing key identification features need-
ed for the reader to compare species observed or col-
lected with the photos in the book and make accurate
and correct identifications. Capturing the entire depth
of field in macro photography is a common problem
and generally the authors do a great job. Some of the
images, however, suffer from poor depth of field sharp-
ness and have blurred or fuzzy portions. (i.e., figures
111, 165, 272). There are many more stunning and
beautiful images with great depth of field than with
out. Species described without an image include a
reference, when available, to another source where a
photo was provided.
Two omissions of occurrence records should be
noted. One was on page 51. When describing the rar-
ity of Leptogium rivulare the authors suggest that the
most recently known collections occurred in the early
1970s (Southern Ontario); however, collections of
this species occurred in the early 2000s (Southern
Ontario 2002 and Manitoba in 2003) and more recently
in Southern Ontario in 2007 (not published). The sec-
ond was a reference to the distribution of Xanrhoria
parietina, page 525, which is described as restricted
to oceanic coastal areas. In general this is true but
recent inland discoveries, including records in South-
ern Ontario (2002 and 2006): have been documented.
346
Using photographs only for identification can
result in a misidentified specimen. In The Macro-
lichens of New England the blend of text and images
will enable the user to avoid this error. There are in-
credible photos of very rare lichens not typically seen
by the average lichenologist. The paired photo exam-
ples of various species showing the upper and lower
surface (Peltigera sp.), wet thallus versus dry thallus
(1.e., Placidium arboretum, Physconia detersa, and
Pyxine sorediata), and macro versus micro features
(1.e., Leptogium, Sterocaulon, Usnea) make this book
incredibly valuable because some of these species
can only be separated by comparing these features.
The book is 7 % inches wide 10 % inches long and
1 % inches deep. It weighs approximately 2.0 kg. Al-
though the book is by no means light, it is a more
manageable field guide than Brodo et al, which is
Wild Plants of Eastern Canada
By Marilyn Walker. 2008. Nimbus Publishing Ltd., P.O. Box
9166, Halifax, Nova Scotia B3K 5MB8. 202 pages, 24.95
CAD.
Wild Plants of Eastern Canada was not what I ex-
pected. I was looking forward to another wild plant
field guide, a cross-referencing source, a book offer-
ing additional details and visuals to aid in identifying
those hard-to-pin-down native asters and goldenrods.
That’s not what Wild Plants of Eastern Canada is
about.
My first reaction to the book was, consequently, mild
annoyance — which developed into growing curiosity,
then pleasant surprise the further I read. Wild Plants
of Eastern Canada turns out to be more than a wild
plant identification guide. Divided into three parts —
(1) “Plants and People of the Northeast’, (2) “Field
Guide’, (3) “Recipes” — the book is a human and nat-
ural history, an identification tool (of sorts), and a
recipe book.
Part 2, the “Field Guide”, takes up the most pages.
It contains extensive information on each plant pro-
filed, including scientific and “other” names, general
information about the plant family, habitat details, a
physical description of the plant, and, depending on the
species, sections on usable parts, common uses, medic-
inal properties, and garden-related particulars. The
accompanying black-and-white photographs are help-
ful, while the leaf prints add a subtle artistic touch.
The recipes covering almost 40 pages in Part 3 of
Wild Plants of Eastern Canada are intriguing. They
range from “classics” such as teas and bannock, to
preparations using petals, buds, roots, and berries. I
have not yet had the opportunity to test any of them,
but I am looking forward to trying out goldenrod
crepes, dandelion leaf sandwiches, and clover butter.
Part 1 of the book I found the most interesting and
most trying. It was there, not far into the section, that
I discovered the book’s most frustrating limitation:
its geographical scope. I had assumed the “Eastern
THE CANADIAN FIELD-NATURALIST
Vol. 119
roughly twice the size and weight, making this book
a more welcome field companion. I have toted both
into backwoods locations and appreciated the size and
weight difference. It has a hard durable cover with a
small ruler drawn on the last page that could be used
for field measurements. If taken into the field, how-
ever, the glossy covers will inevitably get scratched as
there is no slip cover to protect it.
The authors tout this book as the “the most compre-
hensive work of its kind” and they deliver. This book,
like Brodo et al., will be an instant classic. Those who
read this book will have no choice but to be drawn
into the world of these fascinating organisms.
Curis LEwIs
16 Prospect Street, P.O. Box 1384, Lakefield, Ontario
KOL 2HO Canada
Canada” part of the book’s title to refer to the usual
territory stretching from the Atlantic Ocean to the
western end of Ontario. Yet Wild Plants of Eastern
Canada ends up covering the Maritime provinces
only — not a bad thing if that is what the reader is
expecting. But as a Quebec resident, I was hoping my
region was included in the book.
The rest of the information in Part 1 comes as a
pleasant surprise. Rich in historical and anthropolog-
ical detail, it provides a succinct and comprehensive
overview of Atlantic Canada’s paleoenvironment,
European settlement, and Aboriginal history and cul-
ture, including First Nations traditional plant knowl-
edge.
I found the first sub-section, “Like Plants, Like
People” the most intriguing. There, the author writes:
“Like people, plants are great colonizers. They wan-
der about, singly or in “herds”, trying out new condi-
tions until they find somewhere to settle in. If the
environment is not suitable, they go elsewhere out of
necessity, or perhaps out of the same sense of adven-
ture that inspired human migration” — an intriguing,
and for me appealing, perspective.
My main gripe with the book is its main title. Wild
Plants of Eastern Canada is misleading, and the extra
text on the front and back covers does not provide any
clues regarding the geographical scope of the book.
Yet for individuals able to get their hands on it, the
smaller typeface on the front cover — “Identifying,
Harvesting and Using”, “Includes recipes & medici-
nal uses” — provides helpful hints regarding the
book’s content.
Wild Plants of Eastern Canada is a welcome addi-
tion to the wild plants section of my bookshelves,
despite its limitations and frustrations.
RENATE SANDER-REGIER
3, 11" Line, Bristol, Quebec JOX 1G0O Canada
2005
MISCELLANEOUS
Book REVIEWS
347
Sods, Soil, and Spades: The Acadians at Grand Pre and their Dykeland Legacy
By J. Sherman Bleakney. 2005. McGill-Queens Press,
Montreal, Quebec, and Kingston, Ontario. xxvii + 221
pages. 14.95 CAD.
Sherman Bleakney is widely known in Canada as
a zoologist of remarkeably diverse interests and pub-
lications. As curator of amphibians, reptiles and fish
at the National Museum of Canada from 1952 to 1958
his expeditions from the Maritimes to Alberta dou-
bled the national herpetology collection to 14 000
specimens. He published a ground-breaking analysis
of the distribution of amphibians and reptiles in East-
ern Canada (Bleakney 1958) based on his 1956 Ph.D.
thesis from McGill University supervised by leg-
endary Max Dunbar, the first Ph.D. in herpetology at
a Canadian university (Cook 2007). In 1958 he
returned to Nova Scotia where he had been raised, and
to Acadia University where he had obtained his B.S.c
and M.Sc., to teach zoology and where he has remained
until and past retirement. Although initially his her-
petological contributions continued there, particular-
ly on both freshwater and marine turtles, he became
increasingly immersed in publishing on fishes and
invertebrates of the marine ecosytem of the Minas
Basin, culminating in a book on_ nudibranchs
(Bleakney 1996).
Since 1994 he has further broadened his environ-
mental perspective and turned his critical trained eye
NEw TITLES
Prepared by Roy John
+ Available for review * Assigned
and literary skills and enthusium to the dykes origi-
nally built by the French settlers (Acadians) along the
Minas Basin at the head of the Bay of Fundy in Nova
Scotia. The result is a fascinatingly detailed and
throughly documented history and construction
analysis, with sections on the geology, post-glacial
events, and fauna of an area which boasts the world’s
highest tides and the land reclaimed and protected
from them. For this he has been able to draw on writ-
ten and oral accounts and has paid particular tribute
not only to the builders but also to those who left an
accurate record of techniques of dyking brought from
the low countries of Europe and adapted to the local
conditions.
Literature Cited
Bleakney, J. S. 1958. A zoogeographical analysis of the amphibians
and reptiles of Eastern Canada. National Museum of Canada Bul-
letin 155, Ottawa, Ontario.
Bleakney, J. S. 1996. Sea slugs of Atlantic Canada and the Gulf of
Maine. Nimbus Publishing, Halifax, Nova Scotia.
Cook, F. R. 2007. Early Canadian reptile studies: anecdote to inven-
tory. Chapter 2 in Ecology, Conservation and Status of Reptiles
in Canada. Edited by Carolyn Seburn and Christine Bishop. Socti-
ety for the Study of Amphibians and Reptiles Conservation
Monograph 2: 5-22.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario K!P
6P4 Canada
Currency Codes — CAD Canadian Dollars, USD U.S. Dollars, EUR Euros, AUD Australian Dollars.
ZOOLOGY
Albatross — Their World, Their Ways. By T. De Roy, M.
Jones and J. Fitter. 2008. Firefly Books Ltd., 66 Leek Crescent,
Richmond Hill Ontario L4B 1H1. 240 pages, 49.95 CAD,
Cloth.
* Bees. By C. Savage. 2008. Greystone Books, #201 — 2323
Quebec Street, Vancouver, British Columbia VST 4S7. 136
pages, 28 CAD Cloth.
* Extreme Birds — The World’s Most Extraordinary and
Bizarre Birds. By D. Couzens. 2008. Firefly Books Ltd., 66
Leek Crescent, Richmond Hill Ontario L4B 1H1. 288 pages,
45.00 CAD, Cloth.
Smithsonian Field Guide to the Birds of North America.
Edited by Ted Floyd. Harper Collins Publishers Ltd., 2 Bloor
Street East, 20th Floor, Toronto, Ontario M4W 1A8. 528
pages, 26.95 CAD.
* A Wildlife Guide to Chile. By S. Chester. 2008. Princeton
University Press 41 William Street, Princeton, New Jersey,
USA, 08540-5237. 392 pages, 19.95 USD, Paper.
Carnivores of British Columbia. By D. Hatler, W. Nagors-
en, and A Beal. 2008. UBC Press, 20219 West Mall, Vancou-
ver, British Columbia V6T 1Z2. 416 pages, 85 CAD Cloth.
Coral Reef. By S. Wells. 2008. Firefly Books Ltd., 66 Leek
Crescent, Richmond Hill, Ontario L4B 1H1. 288 pages, 39.95
CAD, Cloth
* Cranes — A Natural History of a Bird in Crisis. By J.
Hughes. 2008. Firefly Books Ltd., 66 Leek Crescent, Rich-
mond Hill, Ontario L4B 1H1. 256 pages, 45.00 CAD, Cloth.
Deer World. By D. Taylor. 2008. Firefly Books Ltd., 66
Leek Crescent, Richmond Hill, Ontario L4B 1H1. 400 pages.
39.95 CAD, Cloth.
Fifty Years of Flukes and Flippers: A Little History and
Personal Adventures with Dolphins, Whales and Sea
Lions (1958-2007). By William E. Evans. Pensoft Publishers.
Sofia-Moscow. In English 147 pages EUR 20.00
Elephant Man. By N. Baumgartl and C. Galluci. 2008. Fire-
fly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1. 160 pages, 39.95 CAD, Cloth.
348
Arctic Fox — Life at the Top of the World. By G. Hamil-
ton. 2008. Firefly Books Ltd., 66 Leek Crescent, Richmond
Hill, Ontario L4B 1H1. 232 pages, 39.95 CAD, Cloth.
Lost Worlds of the Guiana Highlands. By Stewart McPher-
son. 2008. Redfern Natural History Productions Limited, 61
Lake Drive, Hamworthy, Poole, Dorset BHI5 4LR Eng-
land, UK. 385 pages, 29.99 GBP.
Dictionary of Herpetology. By H. Lillywhite. 2008. Krieger
Publishing, P.O. Box 9542, Melbourne, Florida 32902-9542
USA. 384 pages, 112.50 USD Cloth.
500 Insects — A Visual Reference. By S. Marshall. 2008. Fire-
fly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1. 528 pages, 29.95 CAD, Cloth.
* Owls of North America. By F. Backhouse. 2008. Firefly
Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1. 216 pages, 34.95 CAD, Cloth.
* Choosing Wildness — My life among the Ospreys. 2008.
By C. Arbour. Greystone Books, #201 — 2323 Quebec Street,
Vancouver, British Columbia VST 487.
*Central Park in the Dark. By M. Winn. 2008. Farrar,
Straus and Giroux [Douglas & McIntyre Publishing Group],
Suite 201, 2323 Quebec Street, Vancouver, British Colum-
bia VST 4S7. 263 pages, 27.50 CAD Coth.
Firefly Encyclopedia of Reptiles and Amphibians. [Sec-
ond Edition] By C. Mattison. 2008. Firefly Books Ltd., 66
Leek Crescent, Richmond Hill, Ontario L4B 1H1. 240 pages,
49.95 CAD, Cloth.
The Encyclopedia of Sharks. By S. Parker. 2008. Firefly
Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B
1H1. 224 pages, 24.95 CAD, Paper.
Spiders — The Ultimate Predators. By S. Dalton. 2008. Fire-
fly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario
L4B 1H1. 208 pages, 34.95 CAD, Cloth.
* Evolution and Biogeography of Australasian Verte-
brates. By John R. Merrick, M. Archer, G. Hickey, and M.
Lee. 2008, Australian Scientific Publishing, P.O. Box 68,
Oatlands, New South Wales 2117, Australia. 942 pages, 230
AUD Cloth, 170 AUD Paper.
BOTANY
Rain Forest. By J. Burton. 2008. Firefly Books Ltd., 66 Leek
Crescent, Richmond Hill, Ontario L4B 1H1. 288 pages,
39.95 CAD, Cloth.
* Wild Plants of Eastern Canada. By M. Walker. 2008
Nimbus Publishing Ltd., P.O. Box 9166, Halifax, Nova Sco-
tia B3K 5MB8. 240 pages, 24.95 CAD.
Between Earth and Sky — Our Intimate Connections to
Trees. By N. Nadkarni. 2008. University of California Press,
2120 Berkeley Way, Berkeley, California 94704 USA. 336
pages, 24.95 USD Cloth.
OTHER
Wilderness America — A Visual Journey. By T. Fitzharris.
2008. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill,
Ontario L4B 1H1. 160 pages, 39.95 CAD, Cloth.
THE CANADIAN FIELD-NATURALIST
Vol. 119
Antarctica — Secrets of The Southern Continent. By D.
McGonigal. 2008. Firefly Books Ltd., 66 Leek Crescent,
Richmond Hill, Ontario L4B 1H1. 400 pages, 59.95 CAD,
Cloth.
A Primer of Conservation Biology [Fourth Edition]. By R.
Primack. 2008. Sinauer Associates, Inc., 23 Plumtree Road,
P.O. Box 407, Sunderland, Massachusetts 01375-0407. 292
pages, 49.95 USD, Paper.
Costa Rica — A Journey through Nature. By A. Hepworth.
2008. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill,
Ontario L4B 1H1. 204 pages, 39.95 CAD, Cloth.
Wild Costa Rica — The Wildlife and Landscapes of Costa
Rica. By A. Hepworth. 2008. The MIT Press, Five Cam-
bridge Center, 4th Floor, Cambridge, Massachusetts 02142-
1493. 176 pages, 29.95 USD, Cloth.
Setting the Standard — Certification, Governance, and
the Forest Stewardship Council. By C. Tollefson, F. Gale
and D. Haley. 2008. UBC Press, 20219 West Mall, Vancou-
ver, British Columbia V6T 1Z2. 352 pages, 85 CAD Cloth.
David Suzuki’s Green Guide. By D. Suzuki and D. Boyd.
2008. Greystone Books, #201 — 2323 Quebec Street, Van-
couver, British Columbia V5T 487. 192 pages, 19.95 CAD.
Snakebit: Confessions of a Herpetologist. By L. Anthony.
2008. Greystone Books, #201 — 2323 Quebec Street, Van-
couver, British Columbia V5T 4S7. 288 pages, 29.95 CAD.
The Lizard King: The True Crimes and Passions of the
World’s Greatest Reptile Smugglers. By Bryan Christy.
2008. Twelve. http://www.twelvebooks.com/content/index.
asp. 241 pages. 24.99 CAD.
The Wisdom of Birds — An illustrated history of ornithol-
ogy. By T. Birkhead. 2008. Greystone Books, #201 — 2323
Quebec Street, Vancouver, British Columbia V5T 4S7. 448
pages, 42.95 CAD.
National Audubon Society Guide to Nature Photography
— Digital Edition. By T. Fitzharris. 2008. Firefly Books
Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1.
208 pages, 24.95 CAD, Cloth.
* Wilderness Prophet — Arthur Carhart. By T. Wolf. 2008.
University Press of Colorado, 5589 Arapahoe Avenue, Suite
206C, Boulder, Colorado 80303. 294 pages, 34.95 USD Cloth.
* The Riverscape and the River. By S. Haslam. 2008.
Cambridge University Press, 32 Avenue of the Americas,
New York, New York 10013-2473. 404 pages, 130 USD
Cloth.
* Serengeti III - Human Impacts on Ecosystem Dynamics.
By A. Sinclair, C. Packer, S. Mduma and J. Fryxell. 2008.
The University of Chicago Press, 1427 East 60th Street
Chicago, Illinois 60637 USA. 464 pages, 45 USD Paper.
* On the Wings of Cranes: Larry Walkinshaw’s Life Story.
By Lowell Schake and James Walkinshaw. 2008. iUniverse,
Inc. 24.95 USD Paper, 35 USD Cloth.
News and Comment
Marine Turtle Newsletter (117) July 2007
25 pages: ARTICLES: Marine turtles in Mozambique:
Towards an effective conservation and management program
(A. Costa, H. Motta, M. A. M. Pereira, E. J. S. Videira, C. M.
M. Louro, and J. Joao) — Predation on the zoanthid Palythoa
caribaeorum (Anthozoa, Cnidaria) by a Hawksbill Turtle (rer-
mochelys imbricata) in southeastern Brazil (S. N. Stammpar,
P. F. da Sola, and O. J. Luiz Jr.) — Rat eradication as part of a
Hawksbill Turtle (Eretmochelys imbricat) conservation pro-
gram in an urban area in Cebedelo, Paraibaaa State, Brazil (D.
Zeppelini, R. Mascarenhas, and G. G. Meier) — Morphody-
namics of an Olive Ridley nesting beach in the Baja Peninsula
(V. M.Gomez-Munoz, and L. Godinez-Orta) — NOTES: First
records of Olive Ridley Turtles (Lepidochelys olivacea) in Sey-
chelles (S. Remie and J. A. Mortimer) — Sexual harassment by
a male Green Turtle (Chelonia mydas) (B. W. Bowen) — Inci-
dental capture of a Leatherback along the coas of Ceara, Brazil
(E. H. S. M. Lima, M. T. D. Melo, and M. H. Godfrey
BOOK REVIEW IUCN-MTSG QUARTERLY REPORT
ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — RECENT
PUBLICATIONS.
The Marine Turtle Newsletter is edited by Lisa M
Campbell, Nicholas School of Environment and Earth Sci
ences, Duke University, 135 Duke Marine Lab Road, Beaufort,
North Carolina 28516 USA; and Matthew H. Godfrey, North
Carolina Wildlife Resources Commission, 1507 Ann Street,
Beaufort, North Carolina 28516 USA. Subscriptions and dona-
tions towards the production of the MTN can be made online
at or postal mail to Michael
Coyne (Managing Editor) Marine Turtle Newsletter, | South-
ampton Place, Durham, North Carolina 27705 USA; e-mail:
mcoyne @seaturtle.org.
The Boreal Dip Net/L’epuosette boreal 11(2) Summer 2007
Newsletter of the Canadian Amphibian and Reptile Con-
servation Network/ Reseau Canadien de Conservation des Am-
phibians et Reptiles
Contents: Editor’s Note (Sara Ashpole) — Summary of
2006 Annual Meeting 27 to 29 October, Victoria, British
Columbia (Jean Wai Jung) — Prairie Rattlesnake movement
patterns though the City of Lethbridge (Wonnita Andrus) —
Hibernation sites of Western Toads in Alberta (Constance
Browne) — Prairie Skinks in the Manitoba sandhills: Deter-
mining a habitat suitability model (Nichlas Cairns and Pamela
I. Rutherford) — The effects of de-icing salt (NaCl) on am-
phibian community structure in Nova Scotia (Sara Collins
and Ron Russell) — Arguments, counter arguments and geneal
mystification over new scientific names for amphibians (David
M. Green) — Reptiles at risk on the road again! (Jeff Hath-
away) — Fertilizers and enrichment of nitrogen isotopic sig-
nature in larval Green Frogs (Dale Jefferson amd Ron Russell)
— Amphibian tropic interactions (Dale Jefferson and Ron
Russell) — Seabird Island Oregan Spotted Frog Project (Keena
McNeil) — Are Bullfrogs being spread by vehicles (Elke
Wind) — Annual General Meeting 2007 Queens University,
Kingston, Ontario 21 to 24 September — Critical habitat work-
shop — Fieldtrip — Member publications.
Alliance of Natural History Museums of Canada Annual Meeting and Awards
_ On 24 and 25 September 2007 the Alliance of Natural
History Museums held its annual meeting in Ottawa. The
~ANNHMC is a network of Natural History Museums created
in 2003 to enhance collaborative work in the areas of research,
collections development, and education about the natural envi-
ronment. The 12 founding members (from west to east) are
the Royal British Columbia Museum, Royal Alberta Museum,
i Royal Tyrell Museum of Palaeontology, Royal Saskatchewan
Museum, Manitoba Museum, Canadian Museum of Nature,
_ Montreal’s Nature Museums (Biodome, Insectarium, Botanical
Gardens and Planetarium), New Brunswick Museum, Nova
j Scotia Museum of Natural History, and the Rooms, New-
i foundland and Labrador. The Royal Ontrario Museum joined
_ in the summer of 2007.
_ “By sharing knowledge and resources among member
museums, we can impove our efforts in opening up the world
of Canada’s rich natural history to Canadians and educating
them about important environmental issues” explained Johanne
Landry, President of the ANHMc and Director of the Insect-
arium (part of Montreal Nature Museums). We have a stragegic
plan in place and have completed a survey of our combined
collections. This meeting was an important opportunity for
our representatives coast to coast to come together and iden-
tify our next steps.” Together, the member museums safe-
guard more than 19 million catalogued specimens of plants,
animals, minerals and fossils collected over 150 years that
are the record of Canada’s natural history through time.
At a reception 25 September on Parliament Hill in Ottawa
the Alliance welcomed the Royal Ontario Musuem as a new
member and presented a new award, the Gold Leaf Award
for significant contribution to the study of natural history in
Canada to Dr. Francis R. Cook, researcher emeritus at the
Canadian Museum of Nature and editor of the Canadian Field-
Naturalist, and posthuously to the late Dr. Bruce Naylor, former
director of the Royal Tyrell Museum of Paleontology and
founding ANHMC president. The award has been renamed
for Dr. Naylor.
349
Minutes of the 128" Annual Business Meeting of
The Ottawa Field-Naturalists’ Club 9 January 2007
Place and time: Canadian Museum of Nature, Ottawa, Ontario, 7:30 pm
Chairperson:
Attendance:
Mike Murphy, President
Twenty-six persons attended the meeting.
Attendees spent the first half-hour reviewing the minutes of the previous meeting, the Treasurer’s report and the
Report of Council. The meeting was called to order at 7:35 pm with some opening remarks from the President.
1. Minutes of the Previous Meeting
1. Under Report of the Nominating Committee; D.
Kitching should be listed under “Members at
Large”
2. Under Presentation of Awards for the 2005 OFNC
Photo Contest, “The Grand Prize winner received
a digital camera and photo printer donated by
Galaxy Camera”
3. Grand Prize winner: Dave Sangster
It was moved by Henry Steger and seconded by Ann
MacKenzie that the minutes be accepted.
(Motion Carried)
2. Business Arising from the Minutes
There was no business arising from the Minutes.
3. Communications Relating to the Annual
Business Meeting
There were no communications relating to the Annual
Business Meeting.
4. Treasurer’s Report Frank Pope
Frank Pope reviewed the financial report for the year
ending 30 September 2006, noting that the Club’s net
assets have increased by approximately $2000. Listed
under “Other revenue” was the balance of a bequest
from the estate of Mildred Groh ($23 717).
Frank answered questions from the audience with
regards to the increased publishing costs of the CEN.
He explained that this was due to the publishing of 4
issues in 2006 versus the 2 issues published in 2005.
Several audience members spoke about electronic pub-
lishing as being the way of the future for scientific jour-
nals, and one which the CEN should actively consider.
Moved by Frank Pope and seconded by Ken Allison
that the Financial Report be accepted as amended.
(Motion Carried)
5. Committee Reports
Mike Murphy asked for questions and comments
on the Committee reports which had been distributed
to the attendees. He thanked the committee chairs and
committee members for their work over the past year.
Moved by Gillian Marston and seconded by Diane
Lepage, that the reports be accepted.
(Motion Carried)
ie)
6. Nomination of the Auditor
Moved by Frank Pope and seconded by Ann Mac-
Kenzie, that Janet Gehr continue as Auditor for another
year.
(Motion Carried)
7. Report of the Nominating Committee
Eleanor Zurbrigg for Fenja Brodo
President
Vice President
Secretary
Treasurer
Past President
Business Manager
Editor CFN
Editor T&L
Committee Chairs
Birds
Computers
Conservation
E&P
E&L
Finance
FWG
Membership
Publications
ON Rep
Members at large
Julia Cipriani
Diane Kitching
Annie Belair
Diane Lepage
Chairs not on Council
Awards
Macoun
Nominations
Retiring from the council: Susan Howell, Justin Peter
Mike Murphy
Ken Allison
Susan Laurie-Bourque
Frank Pope
Gary McNulty
Bill Cody
Francis Cook
Karen McLachlan-Hamilton
Chris Traynor
Dan Millar
Stan Rosenbaum
Gillian Marston
Fenja Brodo
Ann MacKenzie
David Hobden
Henry Steger
Ron Bedford
Eleanor Zurbrigg
Ernie Brodo
Rob Lee
Fenja Brodo
New on the council: Annie Belair
Moved by Eleanor Zurbrigg and seconded by Frank
Pope, that the slate of nominations for the 2007 Council
be accepted.
(Motion Carried)
2007
8. New Business
There was no new business.
There was a comment from a Club member on the
need to involve the younger generation of Club mem-
bers in Club activities. Ways of bringing these members
together at the monthly Club meetings such as the use
of name tags and formal introductions were discussed.
9.The OFNC Website; a virtual tour by our
Webmaster Sandra Garland
The OFNC has a wonderful, informative website
maintained by Sandra Garland. Sandra gave us a tour
of the Website with aid of a laptop and digital projec-
tor. She informed us that the website was started 10
years ago by the E&P committee. Since then the site
MINUTES OF THE 128' ANNUAL BUSINESS MEETING
351
has grown to become the information rich resource for
Club members and the public at large that it is today
Web statistics reveal that the site is actively used and
accessed regularly.
Mike Murphy thanked Sandra for her excellent work
and for her entertaining and informative presentation
10. Adjournment
Moved by David Hobden and seconded by Diane
Lepage that the meeting be adjourned at 9:35 pm.
(Motion Carried)
SUSAN LAURIE-BOURQUE
Recording Secretary
The Ottawa Field-Naturalists’ Club Committee Reports for 2006
Birds Committee
The Birds Committee, in conjunction with the Club des
Ornithologues de ’ Outaouais, compiled the 2005 Christmas
Bird Count. The fall bird count was held in October 2006
and was a successful event. This year marked the 10" anniver-
sary of the Peregrine Falcon Watch. Commemorative hats were
produced to mark the occasion. One male was successfully
fledged. The Bird Records Sub-committee continues to review
records of rare birds and has made Rare Bird Report forms
available on the Club’s web site. This year’s Seed-a-Thon was
successful in raising over $600.00 for Club feeders. A new
feeder was built for the OFNC by the National Capital Com-
mission and was installed at the Mer Bleu. The Committee
continues to operate the Ottawa Bird Status Line and the
Rare Bird Alert.
CHRIS TRAYNOR
Chair, Birds Committee
OFNC Computer Management Committee
The Computer Management Committee did not meet dur-
ing 2006.
MICHAEL MURPHY
OFNC President
Education and Publicity
Kiosks, Displays and Other Events
The committee created another set of display panels for the
large display using the Watershed as the theme. Committee
members participated with displays at the Ottawa-Carleton
Elementary Teachers’ Federation Publishers’ Display, National
Wildlife Week’s Wildlife Festival, several Environment Week
events (National Resources Canada, Agriculture and Agri-food
Canada, Citizenship & Immigration and National Defense),
the Sierra Club “People & the Planet’ conference, and the
OFNC Soiree.
Once again the OFNC participated in the Ottawa Science
Fair sponsoring 3 prizes for projects co-judged by Jeff Sleev-
ington and Ed & Pub committee member Kathy Conlan.
The committee also arranged speakers for outside groups.
Several digital photo presentations were also made by com-
mittee members.
OFNC Membership Drive
The portable kiosks were used for approximately 8 outings
for a successful fall club membership drive.
Digital Library
The committee continued to enhance the OFNC’s ‘digital
photo library and catalogue’ scanning many of the slides from
the slide library.
OFNC Nature Photography Contest
The Committee initiated a 2 Annual Photography Contest
(to run until September 2007) to encourage discovery of nature
and further enhance the club’s digital library. This contest is
open to the general public.
Brochures
The committee continued to distribute the club brochure to
area libraries and nature shops.
Club T-Shirt
Committee member Laura Penney designed a beautiful new
OFNC T-Shirt and 300 shirts of various sizes were printed for
sale.
GILLIAN MARSTON
Chair, Education & Publicity Committee
Excursions and Lectures Committee
This Committee took great pleasure in arranging (and at-
tending) 36 trips, ten monthly meetings (including the ABM)
and the Soirée. We had one bus trip to Montreal (Biodome,
etc.); the rest were by car pool leaving from either Lincoln
Fields or Elmvale Acres. The monthly meetings from Sep-
tember to December were held at the Neatby Building, Agri-
culture and Agri-food Canada because the CMN was under-
going renovations. These facilities were quite satisfactory.
We had 11 birding trips, 2 insect trips, | on plants, | on fun-
gi, lichens and ferns, | on glaciation and the landscape, and
17 general natural history outings. Trips a little further afield
(wetlands near Spencerville and snowshoeing in Algonquin
Park) were very popular as were two mid-week events (early
morning birding and a museum visit).
The Soirée was well-attended because many
Macoun Club members were specially invited.
former
FENJA BRODO
Chair, Excursions & Lectures Committee
352
OFNC Executive Committee
The Executive Committee met once to review issues and
consider strategic alternatives during 2006.
MICHAEL MURPHY
Chair, Executive Committee
Finance Committee
The Finance Committee met three times during the year.
Highlights include:
The Committee recommended, effective for the 2006-07
year, an increase in individual membership from $28.00 to
$33.00 and an increase in family membership from $30.00 to
$36.00. A subscription to CFN will be similarly increased.
Finance Committee considered a request from the Ottawa
Riverkeepers to fund the printing of the first River Report for
the Ottawa River. The report supports the objectives of the Club.
Council agreed with the recommendation to provide $10,000
from the Groh bequest for this purpose.
A proposed budget for 2006-07 was developed and sub-
mitted to Council. A deficit of $13,400 was forecast for the
Club and a deficit of $8,000 was forecast for the Canadian
Field-Naturalist. Finance Committee emphasized the need to
raise additional funds since these deficits cannot continue.
ANN MACKENZIE
Chair, Finance Committee
Fletcher Wildlife Garden
The Fletcher Wildlife Garden has completed another
successful year. Wildlife continues to make good use of it and
several new species have been added to our lists. The Man-
agement Committee met eleven times during the year. The
12 meeting was an end of season volunteer appreciation at-
tended by about 30 people.
On March 24" in Toronto FWG was presented with the On-
tario Association of Landscape Architects’ 2006 Certificate
of Merit for Service to the Environment. This award was later
featured at Agriculture and Agri-food Canada’s volunteer
appreciation event.
Volunteers contributed over 3000 hours, mainly on Friday
and Sunday mornings or Wednesday evenings. This year in
addition to work in the Backyard Garden there was extensive
work to extend the Butterfly Meadow, with plantings of native
perennials, trees and shrubs. Sections of split rail fencing and
rock piles were also added. Flowers were planted in the Ash
Woods by a Pathfinder (Girl Guides) group.
Our struggle with invasive species continues. Garlic mus-
tard and buckthorn are fairly well under control, but not elim-
inated. Pale swallow-wort is more difficult to remove with-
out damage to other plants. We are reduced to measures to
prevent overgrowth and seed production in critical areas while
we wait for biocontrol methods. A group of Ontario Stew-
ardship Rangers and a group from Nepean High School vis-
ited to help with invasive control.
About 30 people attended our International Migratory Bird
Day event in May and a similar number visited during the
Central Experimental Farm Open House. Our display was at
the Wildlife Festival and at Agriculture and Agri-food Canada
for Environment Week. FWG was used for 4 OFNC excur-
sions, and by one school class and a field trip for the Monarch
Teachers’ Network. Talks were presented to three outside
groups.
Our fund-raising this year proved quite successful. The
annual Plant Sale raised almost $3000. A grant from Human
THE CANADIAN FIELD-NATURALIST
Vol. 121
Resources and Skills Development Canada allowed us to
employ a summer student for 10 weeks. She operated the
Interpretation Centre, worked with visitors, prepared mate-
rials for displays and the Backyard Garden and made and re-
corded wildlife observations. Once a week she provided activ-
ities for a children’s day camp from Carleton University. The
TD Friends of the Environment Foundation funded the print-
ing of a brochure and contributed to remodeling of the map
sign by the parking area. At third grant from the Evergreen
Foundation. in the form of a gift card for Home Depot, also
helped with the sign and the Butterfly Meadow work.
Our web site (http://www.ofnc.ca/fletcher.php) has been
improved with news of recent events and observations and a
plant database.
Davib HOBDEN
Chair, Fletcher Wildlife Garden Committee
Macoun Club Committee
The Committee met once, and otherwise coordinated the
weekly and monthly planning by telephone and e-mail. Com-
mittee members supervised or gave presentations at 17 indoor
meetings, and led 16 field trips.
For the second year running, the Macoun Club was with-
out a high-school aged group (our Seniors). Almost all recruit-
ing efforts now are focused on building a strong Junior group
that, growing up in Macoun Club, should eventually re-estab-
lish the Seniors.
As the year drew to a close, we were advised by our co-
sponsor, the Canadian Museum of Nature, that it would not be
able to provide any meeting space during the remainder of the
renovation period at the Victoria Memorial Museum Building
(to 2009). The Museum has, however, warehoused the Macoun
Club’s collections and file cabinets, and expressed a willing-
ness to assist in other ways.
Issue No. 60 of the Macoun Club’s annual publication, The
Little Bear, was put together by a young member and his fam-
ily. It was distributed to members at the annual party in June.
ROBERT E. LEE
Chair, Macoun Field Club
Membership Committee
The distribution of the membership for 2006 is shown in
the table (on next page), with the corresponding numbers for
2005 in brackets. “Others” represent, for the greatest part,
affiliate organizations that receive complimentary copies of
the Club’s publications. Local membership (within 50 km of
Parliament Hill) was 684 and 680 in 2005 and 2006, respec-
tively. The reduction of 11 in total membership is a continu-
ation of this trend in recent years and presents a challenge
to the Club.
This year, the Club lost a long time member, John Cameron,
who joined the Club in 1970. In particular, his contribution to
the Education and Publicity Committee must be acknowledged.
HENRY STEGER
Chair, Membership Committee
Publications Committee
The Publications Committee met three times in 2006, one
of these in conjunction with the Finance Committee.
Four issues of The Canadian Field-Naturalist were pub-
lished in 2006: Volume 118(4) and Volume 119(1,2,3). These
Ea -
NY
2007
CANADIAN USA
Individual 401 (408) 16 (18)
Family 332 (331) | (2)
Sustaining 9... (1) 05 .4(0)
T and L 6 (6) 0 (QO)
Honorary Ze (26) Omni)
Life 42 (41) G6)
Other ae’) (31) 2 eG)
Total 845
four issues contained 666 pages; 60 articles; 28 notes; 56 book
reviews; 115 new titles; 2 commemorative tributes; 20 pages
of News and Comments; 4 pages of miscellany; and a 28 page
index. No funds were drawn from the Manning Memorial
Fund. Re: electronic publishing, the OFNC web-site now
carries the table of contents; all of the abstracts of articles
and notes; and the News and Comments from each issue, as
well as the annual indices. The committee feels that this is as
far as we should go in this regard for the foreseeable future.
MINUTES OF THE 128" ANNUAL BUSINESS MEETING
~
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OTHER [Oral
5 (5) 422 (431)
2 (1) 335 (334)
0 (O) 9 (11)
0 (QO) 6 (6)
0 (O) 25 (26)
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9 (8) 878 (889)
(854) 24 (27)
Publishing costs are still a matter of concern.
Volume 40 of Trail & Landscape was published in four
issues containing 226 pages. Issue 4 contains the 5-year index
for Volumes 36 — 40. Preparation of the second 20-year index
is underway. Costs have risen because of the loss of the
Canadian Heritage subsidy for mailing.
Respectfully submitted,
RONALD E. BEDFORD
Chairman, Publications Committee
354 THE CANADIAN FIELD-NATURALIST Vol. 121
Auditor’s Report
To The Members of THE OTTAWA FIELD NATURALISTS’ CLUB
I have audited the statement of financial position The Ottawa Field-Naturalists’ Club
of THE OTTAWA FIELD-NATURALISTS’ CLUB as at Balance Sheet
September 30, 2006, the statement of changes in net September 30, 2006
assets, the statement of operations, and the statement
of cash flows for the year then ended. These financial ‘Agauie 2006 me!
statements are the responsibility of the organization's Ginn
management. My responsibility is to express an opin- Cher $26,480 $27,819
ion on these financial statements based on my audit. Investment certificates (Note 1) 15,145 29.401
Except as explained in the following paragraph, I Marketable securities (Note 2) 31,187 0
conducted my audit in accordance with Canadian gen- Accounts receivable 31,491 21,837
erally accepted auditing standards. Those standards Prepaid expenses ae OP es ECOOm
require that I plan and perform an audit to obtain rea- $104,303 80,057
sonable assurance whether the financial statements
are free of material misstatement. An audit includes © LAND — ALFRED BoG 3,348 3,348
examining, on a test basis, evidence supporting the Marketable securities (Note 2) 315,501 338,834
amounts and disclosures in the financial statements. $423,152 $422,239
An audit also includes assessing the accounting prin-
ciples used and significant estimates made by man- LIABILITIES AND FUND BALANCES
agement, as well as evaluating the overall financial
statement presentation. CURRENT
In common with many non-profit organizations, THE Accounts payable and
OTTAWA FIELD-NATURALISTS’ CLUB derives some of eoomed Lelolhues $2500 Beh
P j as Jini Deferred revenue 11,688 12,049
its revenue from donations and fund-raising activi- oo ——
ties. These revenues are not readily susceptible to 14,188 15,549
complete audit verification. Accordingly, my verifi- | LIFE MEMBERSHIPS 14,079 14,079
cation of these revenues was limited to the amounts
recorded in the records of THE OrrawA FieLp- NET ASSETS
NATURALISTS’ CLUB, and I was not able to determine Unrestricted 145,238 147,823
whether any adjustments to the recorded amounts might CSOT Ue ONEDD MOD eae
be necessary. Manning principal 100,000 100,000
oe f Manning interest - OFNC 1,948 1,014
In my opinion, except for the effects of adjustments, _CFN 24,225 20,490
if any, which I might have determined to be necessary Seedathon 329 810
had I been able to satisfy myself concerning the com- Anne Hanes memorial 788 870
pleteness of the revenue referred to in the preceding de Kiriline-Lawrence 17,174 16,619
paragraph, these financial statements present fairly, in Macoun Baillie Birdathon 1,253 1,180
all material respects, the financial position of the orga- Alfred Bog __ 3,930 __3,805_
nization as at September 30, 2006, and the results of NEGUS: BSSesSee! Veale IE BICD 394,885 392,611
its Operations and cash flows for the year then ended in $423,152 $422,239
accordance with Canadian generally accepted account-
ing principles.
JANET M. GEHR
C.A., Licensed Public Accountant
North Gower, Ontario
6 January 2007
2007
The Ottawa Field-Naturalists’ Club
Statement of Operations
For the Year Ended September 30, 2006
2006
REVENUE
Memberships $ 13,490
Trail and Landscape 146
Interest 2,730
GST rebate 1,005
Other — 625:
17,994
OPERATING EXPENSES
Affiliation fees $625
Computer 841
Membership 836
Office Assistant 1,275
Telephone 1,782
Insurance 725
Audit 2,000
GST 1,460
Other 946
10,490
C.LuB ACTIVITY EXPENSES
Awards $232
Birds 385
Education and Publicity 2,627
Excursions and Lectures 832
Macoun Field Club ESS
Origins and History of OFNC 0
Soiree 46
Trail and Landscape 10,014
Fletcher Wildlife Garden (Note 4) 2,321
Other 323
17,919
SURPLUS (DEFICIT) OF REVENUE
OVER EXPENSES $ (10,415)
MINUTES OF THE
2005
$ 13,635
220
2,129
720
622
17,326
652
1,846
1,363
1,000
II 7/313
725
2,500
1,755
1,072
12,646
390
618
1,149
(1,805)
466
128™ ANNUAL BUSINESS MEETING
REVENUE
Memberships
Subscriptions
Reprints
Publication charges
Interest and exchange
GST rebate
Other
EXPENSES
Publishing
Reprints
Circulation
Editing
Office Assistant
Honoraria
GST
Other
EXCESS EXPENSES OVER
REVENUE
The Ottawa Field-Naturalists’ Club
Canadian Field-Naturalist — Statement of Operations
For the Year Ended September 30, 2006
2006
$9,311
24,314
5,267
27,470
10,829
1,542
1,299
80,032
57,735
3,832
9,678
1,856
5,000
9,000
4,870
24
91,995
$ (11,963)
355
2005
$ 9,090
25,135
6,578
23,270
9 $24
2,503
1,23]
77.631
20,777
4.658
8,532
2,394
5,000
9.000
2,693
19]
53,245
$ 24,386
356 THE CANADIAN FIELD-NATURALIST
The Ottawa Field-Naturalists’ Club Notes to the Financial Statements
September 30, 2006
1. Cash
2006
Chequing $ 1,945
Savings 20,630
Nesbitt Burns 919
Fletcher Wildlife Garden 2,986
$ 26,480
Investment Certificates: Maturity Maturity Yield
Value Date
$ 15,145 05/01/06 3.25%
2. Marketable Securities
(Market Value $343,241) Maturity Maturity Yield
Value Date
Province of Newfoundland Coupon $ 44,782 10/17/11 4.525%
Province of Ontario Coupon 15,376 12/02/12 4.591%
Province of Manitoba Coupon 45,740 09/05/13 4.694%
Res CIBC Int BB6 70,827 10/31/14 4.144%
CMHC Global Debs 52,000 12/01/06 5.250%
Province of Ontario Bond 30,000 09/12/07 6.125%
Province of Newfoundland Bond 20,000 10/07/08 6.263%
Government of Canada Coupon 30,167 12/01/09 5.605%
Province of New Brunswick Bond 60,000 12/03/15 3.965%
Province of New Brunswick Bond 20,000 06/15/10 6.231%
3. Capital Assets
Equipment at a cost of $16,748 is fully amortized.
4. Fletcher Wildlife Garden
2006
REVENUE
Human Resources and Skills Dev. Canada $ 2,500
TD Friends of the Environment Fund 1,500
Taverner Cup 67
Sales 3,083
GST 221
Donations 480
7,851
EXPENSES
Program 3,235
Backyard 2,167
Habitats 2,584
Interpretation centre By
Administration 409
Publications 1,295
GST 356
Library 94
10,172
$(2,321)
Vol. 121
2005
$ 16,144
6,963
459
4,253
$ 27,819
Book
Value
$ 15,145
Book
Value
$ 36,400
11,703
33,612
51,188
53,538
31,187
20,538
24,944
62,510
21,068
$346,688
7,877
$(3,667)
2007
5. Statement of Changes in Net Assets
a) Unrestricted other revenue includes the balance of a
bequest of $23,717 from the Estate of Mildred Groh.
b) Unristricted other expenses: a donation to enable the
publication of the River Keepers’ River Report.
c) Seedathon Expenses: birdseed for club feeders, $1314.
6. Publication Liability
An annual membership fee or subscription entitles the
member/subscriber to four issues of The Canadian Field-
Naturalist based on a calander year. As the year end of the
MINUTES OF THE 128" ANNUAL BUSINESS MEETING 357
Club is September 30, the Club incurs a liability for publishing
the fourth issue of each publication
At this time, however, the publication of The Canadian
Field-Naturalist is running late. At September 30, 2006, the
Club owes members/subscribers issue number 3 and 4 of
2005 and three issues of 2006. Although most of the work
preparing these publications is done by volunteers, the Club
must pay for printing and mailing. Based upon recent costs,
it is estimated that the Club has a liability of $68,000 for the
outstanding issues. This amount will be reduced by page charg-
es to the authors in the amount of 40% of the printing costs
The Ottawa Field-Naturalists’ Club Statement of Changes in Net Assets
For the Year Ended September 30, 2006 (see note 5 for footnotes)
$(10,000) b) $145,238
Net Beginning Excess
Assets Balance Expenses CFN
Unrestricted $147,823 $(11,963)
Club reserve 100,000 0
Manning Principal 100,000 0
Manning - CFN 20,490 0
Manning - OFNC 1,014 0
Seedathon 810 0
Anne Hanes Memorial 870 0
de Kiriline-Lawrence 16,619 0
Macoun Baillie Birdathon 1,180 0
Alfred Bog 3,805 0
$392,611
Excess Other Other Ending
Expenses OFNC Revenue Expenses Balance
$(10,415) $29,793 a)
0 0) 0 100,000
0 0) 0 100,000
0 3,735 0 24,225
0 934 0 1,948
0 833 (1,314) c) 329
0 0 (82) 788
0 555 0 17,174
0 146 (73) 1,253
0 125 0 3,930
$(10,415) $36,121 $(11.469) $394,885
$(11,963)
The Ottawa Field-Naturalists’ Club Summary of Significant Accounting Policies
September 30, 2006
1 Nature of Business
The organization is non-profit and incorporated under the
laws of Ontario (1884). The organization promotes the appre-
ciation, preservation, and conservation of Canada’s natural
heritage. It encourages investigation and publishes the results
of the research in all fields of natural history and diffuses infor-
mation on these fields as widely as possible. It also supports
and cooperates with other organizations engaging in pre-
serving, maintaining or restoring environments of high quality
for living things.
2. Financial Instruments
The organization’s financial instruments consist of cash,
accounts receivable, marketable securities, and accounts pay-
able. Unless otherwise noted, it is the management’s opinion
that the organization is not exposed to significant interest, cur-
rency, or credit risks arising from these financial instruments.
The fair value of these instruments approximate their carry-
ing values, unless otherwise noted.
3. Capital Assets
Capital assets in excess of $4,000 cost are recorded as assets
at cost and amortized on a straight-line basis. These assets have
been fully amortized.
4. Revenue Recognition
Revenue is recognized when earned for donations and
fund-raising when received.
5. Foreign Currency
Transactions during the year in U.S. dollars have been con-
verted in the accounts to Canadian dollars at the exchange
rate effective at the date of the transaction. All monetary assets
in U.S. dollars at year end have been converted to Canadian
dollars at the rate effective on Sept. 30, 2006.
Gains or losses resulting therefrom are included in revenue
or expenses.
358
THE CANADIAN FIELD-NATURALIST
Vol. 121
Advice for Contributors to The Canadian Field-Naturalist
Content
The Canadian Field-Naturalist is a medium for the publi-
cation of scientific papers by amateur and professional natu-
ralists or field biologists reporting observations and results
of investigations in any field of natural history provided that
they are original, significant, and relevant to Canada. All read-
ers and other potential contributors are invited to submit for
consideration their manuscripts meeting these criteria. The
journal also publishes natural history news and comment items
if judged by the Editor to be of interest to readers and sub-
scribers, and book reviews. Please correspond with the Book
Review Editor concerning suitability of manuscripts for this
section. For further information consult: A Publication Policy
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian
Field-Naturalist 97(2): 231-234. Potential contributors who
are neither members of The Ottawa Field-Naturalists’ Club
nor subscribers to The Canadian Field-Naturalist are encour-
aged to support the journal by becoming either members or
subscribers.
Manuscripts
Please submit by post to the Editor, in either English or
French, three complete manuscripts written in the journal
style. Manuscripts may also be submitted (one copy) by e-
mail. The research reported should be original. It is recom-
mended that authors ask qualified persons to appraise the
paper before it is submitted. All authors should have read and
approved it. Institutional or contract approval for the publica-
tion of the data must have been obtained by the authors. Also
authors are expected to have complied with all pertinent leg-
islation regarding the study, disturbance, or collection of ani-
mals, plants or minerals. The place where voucher specimens
have been deposited, and their catalogue numbers, should be
given. Latitude and longitude should be included for all indi-
vidual localities where collections or observations have been
made.
Manuscripts should be printed on standard-size paper, dou-
blespaced throughout, generous margins to allow for copy
marking, and each page numbered. For Articles and Notes
provide a bibliographic (citation) strip, an abstract, and a list
of key words. Generally, words should not be abbreviated but
use SI symbols for units of measure. The names of authors of
scientific names may be omitted except in taxonomic manu-
scripts or other papers involving nomenclatural problems.
“Standard” common names (with initial letters capitalized)
should be used at least once for all species of higher animals
and plants; all should also be identified by scientific name.
The names of journals in the Literature Cited should be
written out in full. Unpublished reports and web documents
should not be cited here but placed in a separate Documents
Cited section. List the captions for figures numbered in arabic
numerals and typed together on a separate page. Present the
tables each titled, numbered consecutively in arabic numerals,
and placed on a separate page. Mark in the margin of the text
the places for the figures and tables.
Check recent issues (particularly Literature Cited) for
journal format. Either “Canadian” or “American” spellings
are acceptable in English but should be consistent within one
manuscript. The Oxford English Dictionary, Webster’s
New International Dictionary and le Grand Larousse
Encyclopédique are the authorities for spelling.
Illustrations
Photographs should have a glossy finish and show sharp
contrasts. Electronic versions should be high resolution.
Prepare line drawings with India ink on good quality paper
and letter (don’t type) descriptive matter. Photographic
reproduction of line drawings should be no larger than a
standard page. Write author’s name, title of paper, and fig-
ure number on the lower left corner or on the back of each
illustration.
Reviewing Policy
Manuscripts submitted to The Canadian Field-Naturalist
are normally sent for evaluation to an Associate Editor (who
reviews it or asks another qualified person to do so), and at
least one other reviewer, who is a specialist in the field, cho-
sen by the Editor. Authors are encouraged to suggest names
of suitable referees. Reviewers are asked to give a general
appraisal of the manuscript followed by specific comments
and constructive recommendations. Almost all manuscripts
accepted for publication have undergone revision—some-
times extensive revision and reappraisal. The Editor makes
the final decision on whether a manuscript is acceptable for
publication, and in so doing aims to maintain the scientific
quality, content, overall high standards and consistency of
style, of the joumal.
Special Charges — Please take note
Authors must share in the cost of publication by pay-
ing $90 for each page, plus $30 for each illustration (any size
up to a full page), and up to $90 per page for tables (depend-
ing on size). Authors may also be charged for their changes in
proofs. Reproduction of color photos is extremely expensive;
price quotations may be obtained from the Editor.
Limited joumal funds are available to help offset publi-
cation charges to authors without grants or institutional sup-
port. Requests for financial assistance should be made to the
Editor when the manuscript is submitted.
Reprints
An order form for the purchase of reprints or pdf will ac-
company the galley proofs sent to the authors. Invoices for
publication costs will be sent when the submission is pub-
lished.
FRANCIS R. Cook, Editor
RR 3 North Augusta, Ontario KOG IRO Canada
TABLE OF CONTENTS (concluded) Volume 121 Number 3
Notes (continued)
Northern Goshawk, Accipiter gentilis, exploits a Beagle Hound, Canis familiaris, as a “beater” to
catch a Snowshoe Hare, Lepus americanus JOHN T. NEVILLE
Giant Beaver, Castoroides ohioensis, remains in Canada and an overlooked report from Ontario
C. RICHARD HARINGTON
A late born White-tailed Deer, Odocoileus virginianus, in southcentral Wisconsin
CHRISTOPHER N. JACQUES, WILLIAM E. ISHMAEL, TIMOTHY R. VAN DEELEN, and ROBERT E. ROLLEY
Book Reviews
Zoo.LoGy: Evolution and Biogeography of Australian Vertebrates — A Wildkife Guide to Chile —
Ecology & Evolution in the Tropics: A Herpetological Perspective — Fossil Ecosystens of North
America: A Guide to the Sites and Their Extraordinary Biotas — Headless Males Make Great Lovers
— State of North America’s Birds of Prey — Owls of the United States and Canada: A Complete Guide
and Other Unusual Natural Histories to Their Biology and Behavior — Owls of North America — The
Tree of Life: A Phylogenetic Classification — Biology of the Snapping Turtle (Chelydra serpentina) —
Wildlife of North America: A Naturalist’s Lifelist
Botany: Grasses of Colorado — The Macrolichens of New England — Wild Plants of Eastern Canada
MISCELLANEOUS: Sods, Soil, and Spades: The Acadians at Grand Pre and their Dykeland Legacy
New TITLES
News and Comment
Marine Turtle Newsletter (117) July 2007 — The Boreal Dip Net/L’epuosette boreal 11(2) Summer 2007
— Alliance of Natural History Museums of Canada Annual Meeting and Awards
Minutes of the 128th Annual Business Meeting of The Ottawa Field-Naturalists’ Club Tuesday
9 January 2007
Advice to Contributors
Mailing date of the previous issue 121(2): 6 November 2008
2007
350
358
THE CANADIAN FIELD-NATURALIST Volume 121 Number 3 2007
Articles
Brown-headed Cowbird, Molothrus ater, parasitism and abundance in the Northern Great Plains |
LAWRENCE D. IGL and DOUGLAS H. JOHNSON 239
Wolf, Canis lupus, behavior in areas of frequent human activity
ELLEN HEILHECKER, RICHARD P. THIEL, and WAYNE HALL, JR. 256
Influence of gender and den type on home range shape for Striped Skunks, Mephitus mephitis,
in Saskatchewan SERGE LARIVIERE, DAVID HOWERTER, and FRANCOIS MESSIER 261
Road-kill of mammals in Nova Scotia DAvID FUDGE, BILL FREEDMAN, MICHAEL CROWELL
Tony NETTE, and VINCE POWER 265
Third census of seabird populations of the Gaspé Peninsula, Québec, 2002
RICHARD COTER and JEAN-FRANCOIS RAIL 274
Sixteenth census of seabird populations in the sancturaries of the North Shore of the Gulf of the
St. Lawrence, 2005 JEAN-FRANCOIS RAIL and RICHARD COTER 287
Discovery of a possibly relict outbreeding morphotype of Sparrow’s-egg Lady’s-slipper Orchid,
Cypripedium passerinum, in southwestern Yukon
PAUL M. CATLING and BRUCE A. BENNETT 295
Twenty-four hour activity budgets of Mule Deer, Odocoileus hemionus, in the aspen parkland
of eastcentral Alberta G. W. KuZyK and R. J. HUDSON 299
Cavity nest materials of Northern Flying Squirrels, Glaucomys sabrinus, and North American
Red Squirrels, Tamiascius hudsonicus, in a secondary harwood forest of southern Ontario
JESE E. H. PATTERSON, STEPHEN J. PATTERSON, and JAY R. MALCOLM 303
Depredation of Common Eider, Somateria mollissima, nests on a central Beaufort Sea barrier
island: A case where no one wins JOHN A. REED, DEBORAH L. LACROIX, and PAUL L. FLINT 308
Management implications of molt migration by Atlantic Flyway resident population Canada
Geese, Branta canadensis SUSAN E. SHEAFFER, RICHARD A. MALECKI,
BRYAN L. SwIFT, JOHN DUNN, and KIM SCRIBNER 313
Recovery of DNA from footprints in the snow LOVE DALEN, ANDERS GOTHERSTROM,
THOMAS MEIJER, and BETH SHAPIRO 32h
Notes
A River Otter’s, Lontra cadadensis, capture of a Double-crested Cormorant, Phalacrocorax auritus,
in British Columbia’s Gulf Island waters MICHAEL H. H. PRICE and CLARE E. ARIES 325
Winter occurrences of Ivory Gulls, Pagophila eburnea, in inland Labrador
Tony E. CHUBBS and FRANK R. PHILLIPS 327
Product group from well-managed JP <0,
Rae eee ISSN 0008-3550
recycled wood or fibre
FSC www.fsc.org Cert no. SGS-COC-003284
© 1996 Forest Stewar ‘dship Council I
i Mixed Sources “& (continued on inside back cover)
The CANADIAN
FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
Volume 121, Number 4 October—December 2007
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Her Excellency The Right Honourable Michaille Jean
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields |
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members
Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope
Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr.
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson
2007 Council
President: Mike Murphy Annie Belair Diane Kitching Dan Millar
Vice-President: Ken Allison Ronald E. Bedford |= Karen McLachlan Hamilton Stanley Rosenbaum
Recording Secretary: Susan Laurie-Bourque Fenja Brodo David Hobden Henry Steger
Treasurer: Frank Pope Julia Cipriani Diane Lepage Chris Traynor
Past President: Gary McNulty William J. Cody Ann Mackenzie Eleanor Zurbrigg
Francis R. Cook Gillian Marston
To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate i
P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofcn@ofnc.ca.
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.
We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward
our mailing costs. PAP Registration Number 9477. Canada
Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: ofen@ofne.ca
Copy Editor: Elizabeth Morton
Honorary Business Manager: William J. Cody
Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwegsc.gc.ca
Associate Editors: | Robert R. Anderson Robert R. Campbell Anthony J. Erskine Donald F. McAlpine
Warren B. Ballard Paul M. Catling David Nagorsen William O. Pruitt, Jr.
Charles D. Bird Brian W. Coad
Chairman, Publications Committee: Ronald E. Bedford
All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail
or e-mail.
Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor.
Subscriptions and Membership
Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 per
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) and
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must
add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local
Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at
$33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed |
to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications
Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: October-December 2007 (March 2009).
Cover: Bufflehead, Bucephala albeola. Photographed at Shoal Harbour Migratory Bird sanctuary, southeastern Vancouver Island
by Suzanne Huot. See articles by James K. Findley pages 370-374 and 375-378.
The Canadian Field-Naturalist Ky
ARV
== <5 aL D
Volume 121, Number 4 October-December ‘2 SIT y
Moose, Alces alces, Winter Browse Use in Central Labrador
TINA L. NEwsury!4, NEAL P. P. SIMon?, and Tony E. CHUBBS?
'Western Newfoundland Model Forest, P.O. Box 68, Corner Brook, Newfoundland and Labrador A2H 6C3 Canada
*Newfoundland and Labrador Department of Natural Resources, P.O. Box 3014, Station B, Happy Valley-Goose Bay, New-
foundland and Labrador AOP 1E0 Canada (Deceased: see Canadian Field-Naturalist 121(1): 96-98).
3Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and
Labrador AOP 1S0 Canada; e-mail: techubbs @cablelab.net; corresponding author
4Present address: Jacques Whitfield Limited, 19 Union Street, P.O. Box 772, Corner Brook, Newfoundland and Labrador
A2H 6G7 Canada
Newbury, Tina L., Neal P. P. Simon, and Tony E. Chubbs. 2007. Moose, Alces alces, winter browse use in central Labrador.
Canadian Field-Naturalist 121(4): 359-363.
To determine the effect of forestry practices on the availability of winter Moose forage, we recorded Moose browse along
four 250 m transects in each of five forest regeneration ages. Browse use was greater on 20- and 30-year-old regenerating
stands as compared with recently clearcut stands (5 and 10 years old) or mature forest (> 150 years old). Willow (Salix sp.)
followed by White Birch (Betula papyrifera) had the highest proportion of browsing by Moose.
Key Words: Moose, Alces alces, browse, logging, Labrador.
Since the early 1950s, Moose (Alces alces) have ex-
panded into Labrador, independent of introductions on
the Labrador coast (Mercer and Kitchen 1969; Dalton
1986*:; Chubbs and Schaefer 1997). Conversions of
mature forests into early and mid-successional seres
are partly responsible for Moose increases in most of
North America (Bergerud and Manuel 1968; Collins
and Helm 1997; Rempel et al. 1997) since Moose tend
to be associated with mid-successional forests (Ber-
gerud and Manuel 1968; Telfer 1974; Pierce and Peek
1984; Collins and Helm 1997; McCracken et al. 1997;
McLaren et al. 2000). It is unlikely that the first expan-
sion of Moose into Labrador resulted from forest cut-
ting; most cutting is limited to discrete areas isolated
from the historical distribution of Moose in eastern
Canada and logging largely began in the late 1960s
_ (FMDPT 2003*) after Moose had become established
in Labrador (Chubbs and Schaefer 1997). However,
anticipated increases in logging in Labrador (FMDPT
2003*) will likely increase the amount of forest in suc-
- cessional stages favorable to Moose, possibly increas-
_ ing Moose densities.
Moose favor areas of highest forest productivity,
preferring 5- to 15-year-old regenerating stands where
vegetation reaches heights of 3 m and is thus available
; above snow (Dodds 1960; Bergerud and Manuel 1968;
Telfer 1974; McLaren et al. 2000). The mosaic of food
and cover produced by logging can benefit Moose.
High quality food is important for storing winter fat
-and provides females with the nutrients required for
rearing young (Leptich and Gilbert 1989), while ther-
mal cover is important for energy conservation (Schwab
and Pitt 1991). Early and mid-aged clearcuts are favor-
able to Moose relative to forested stands in part due to
more total browse (Telfer 1974; Schwab et al. 1987;
Collins and Helm 1997) and because wind exposure
reduces snow depths (Schwab et al. 1987). Early regen-
erating and pre-commercially thinned Balsam Fir (Abies
balsamea) stands may also attract Moose, depending on
site type (Thompson et al. 1992; McLaren et al. 2000).
There is considerable geographical and seasonal
variation in Moose diets, yet coarse patterns exist (Peek
1974). In boreal forests during winter, Moose make
high use of White Birch (Betula papyrifera), Mountain
Maple (Acer spicatum), and Balsam Fir saplings, wil-
low (Salix sp.), and with lesser amounts of Red-osier
Dogwood (Cornus stolonifera), Pin Cherry (Prunus
pennsylvanica), Mountain Ash (Sorbus americana), and
Viburnum spp. (Dodds 1960; Bergerud and Manuel
1968; Peek 1974; Proulx and Joyal 1981; McLaren et
al. 2000). Leaves and annual growth stems of the above
species are dominant spring and summer forage. Addi-
tional summer foods include plants such as Yellow
Water Lily (Nuphar microphyllum), horsetail (Equise-
tum sp.), fireweed (Epilobium spp), and sedges (Carex
spp.) (Dodds 1960; Irwin 1985).
To determine the effects of forest harvesting on
Moose browse, we documented Moose browsing across
regenerating clearcuts of four ages (5 to 30 years old)
and mature, uncut forests.
359
360
Study Area
The study was conducted from 28 June to 10 Au-
gust 2004 within 40 km of Happy Valley-Goose Bay
(53°19'N, 60°25'W), Newfoundland and Labrador,
Canada (Figure 1). Sites were located in the High
Boreal Forest Ecoregion of Labrador (Meades 1990*);
this region contains the most productive forests for
commercial timber in Labrador (Wilton1959; Lopouk-
hine et al. 1975). The moderately rolling terrain is dom-
inated by Black Spruce (Picea mariana) and Feather
Moss (Pleurozium schreberi) forest at the higher ele-
vations and Balsam Fir/Black Spruce/White Birch
forest at slightly lower elevations (Lopoukhine et al.
1975). This area experiences a mean annual tempera-
ture of -0.5°C (mean monthly range: -18.1 to 15.4°C)
and precipitation amounts of 949 mm, half of which
falls as snow (Environment Canada Climate Normals:
http://www.msc-smc.ec.gc.ca; viewed 27 September
2004). Snow remains on the ground from October
through June. Approximately 12% of this study area
was commercially harvested from the late 1960s to
2004 (FMDPT 2003"). The area is accessible to hunters
and the most recent density estimates reported 0.168
Moose/km? in 1994 (Chubbs and Schaefer 1997).
Methods
We established twenty 250 m-long transects in dif-
ferent stands, with four transects representing four
clearcut ages (approximately 30, 20, 10, and <5 years
following cutting) and uncut mature (>150 years old)
forest. We selected stands ranging from 30 to 700 ha
and those within the same age group were chosen as
far apart as possible (= 700 m) while still being acces-
sible. Prior to harvest, stands were dominated by
Black Spruce and classified as commercial, i.e., sup-
porting > 100 m of timber per ha with canopy height
ranging from 9.5 to 18.5 m tall and crown closure
ranging from 50 to 75%. Our mature forest transects
reflected this variation and all sites had regenerated.
Transects in harvested stands started at commercial
mature forest edges and were oriented to avoid all
other stand edges, roads and patches of remnant forest.
Transects consisted of evenly spaced plots at 50 m
intervals. Each plot consisted of five 4.5 m?* circular
subplots: one central and the remaining four in the
cardinal directions 10 m from the centre. Within each
subplot, plant species, number of stems and occur-
rence of Moose browsing were recorded.
Results
Most browsing occurred in the 20- and 30-year-old
stands, with willow being the most proportionately,
0.50 and 0.80, respectively, browsed species. Willow
accounted for only 0.8% of total stems but represented
16.5% of the total stems browsed across 20- and 30-
year-old stands. The second most proportionately
browsed species was White Birch (0.33 and 0.48 in
20- and 30-year-old stands, respectively). Considerably
THE CANADIAN FIELD-NATURALIST
Vol. 121
smaller proportions of Mountain Alder (Alnus crispa),
Balsam Fir and Black Spruce were browsed. There was
virtually no browsing in clearcuts < 15 years old and
forests > 150 years old.
Discussion
Throughout their range and between seasons, spe-
cies browsed by Moose varies, but tends to be domi-
nated by willow when available (Peek 1974; Mc-
Cracken et al. 1997; Collins 1999). Similarly, we found
that willow was the most proportionately browsed
species in Labrador. Our second most proportionately
browsed species, White Birch, was also a prominent
Moose browse throughout Canada (Dodds 1960; Peek
1974). However, in contrast to other regions (Dodds
1960; Bergerud and Manuel 1968; Thompson et al.
1992) proportionately little Balsam Fir was browsed
in our study. Balsam Fir is generally a winter food
(Dodds 1960) and may be selected only when decid-
uous species are unavailable or where Moose densities
are high (McLaren et al. 2000). Moose are reported
absent from apparently suitable habitat in Labrador
and are possibly limited by Wolf (Canus lupus) preda-
tion, illegal hunting, and snow depths (Trimper et al.
1996). Although snow depth could limit food supply
(Schwab et al. 1987), it also increases expended ener-
gy (Schwab and Pitt 1991). Illegal hunting, Wolf pre-
dation and energy costs of snow depth may depress
Moose populations enough that they can forage on the
preferred willow and White Birch rather than resort-
ing to Balsam Fir.
Our finding of more Moose browsing in 20- and 30-
year-old stands is similar to other studies that found
greater amounts of browse and Moose densities in re-
generating clearcuts (Telfer 1974; Schwab et al. 1987;
Leptich and Gilbert 1989; Collins and Helm 1997;
Thompson et al. 1999; McLaren et al. 2000). However,
our peaks in Moose browsing occurred 10 — 15 years
later than suggested by Dodds (1960) and Telfer (1974),
likely due to the slower regeneration rate in our study
area than in more southerly Moose ranges. Our results
suggest that increases in forest cutting may increase
Moose densities by enhancing browse production.
However, the lack of browsing on less preferred
species (e.g., Balsam Fir) indicates Moose are proba-
bly not limited by winter browse and therefore Moose
may not increase as rapidly as in other areas. Although
we found few browsed stems outside 20 — 30 year old
clearcuts, Moose may use these areas to graze on herbs
during summer and fall. Browse surveys alone may not
be enough to quantify Moose diets — fecal and rumen
analyses and foraging observations would help to
better determine seasonal Moose forage preferences
(McCracken et al. 1997).
Increased Moose densities resulting from logging
may enhance recreational and sustenance activities;
e.g., viewing and hunting. However, increased Moose
densities are believed to increase Woodland Caribou
2007 NEWBURY, SIMON, and CHUBBS: MOOSE BROWSE USE IN CENTRAL LABRADOR 36]
Legend
b Saad Roads
1500 ——-_ Transects
150D
30C
150C
20C
ie yy
10D —
———— 16
ee
150B wie 8D
204
150A
20B
AS
20D
30D
5B = 10A
30A
30B
Ficure 1. Study area showing the locations of each of the four (A, B, C and D) transect locations for each clearcut age (5, 10.
20, 30, and 150 years).
362
THE CANADIAN FIELD-NATURALIST
Vol. 121
TABLE |. Proportion of plant stems browsed by Moose and standard error according to clearcut age (n = 100 per age).
Plant species Number of stems
5 years
Abies balsamea 302
Picea mariana 623
Alnus crispa 0
Betula papyrifera 3
Salix spp. 0
10 years
Abies balsamea 165
Picea mariana 657
Alnus crispa 0)
Betula papyrifera i
Salix spp. 8
20 years
Abies balsamea 912
Picea mariana 1429
Alnus crispa 118
Betula papyrifera 45
Salix spp. 26
30 years
Abies balsamea 627
Picea mariana 1111
Alnus crispa 0
Viburnum edule 0
Betula papyrifera 129
Salix spp. 10
> 150 years
Abies balsamea 345
Picea mariana 930
Alnus crispa 0
Betula papyrifera 1
Salix spp. 7
(Rangifer tarandus) mortality possibly thorough an
influx of large predators; e.g., Wolves (Bergerud and
Elliot 1986; Klein 1991; Seip 1992; Schaefer et al.
1999) and Black Bears, Ursus americanus (Mahoney
and Virgl 2003). Boreal populations of Woodland Cari-
bou are threatened in Labrador (Schmelzer et al.
2004*), so it may be desirable to reduce Moose for-
age, notably White Birch and willow, on regenerating
clearcuts. Our results indicate that accelerating the pas-
sage of clearcuts through earlier successional stages by
aggressive Black Spruce planting and targeting Moose
forage in pre-commercial thins would have the great-
est negative impact on Moose winter forage.
Acknowledgments
We thank S. Barr, R. Dove, M. Hynes, D. Jacque,
and K. Osmond for their assistance with data collec-
tion and entry. We thank F. Schwab and an anony-
mous reviewer for the helpful comments on the man-
uscript. D. Jennings and J. Thomas provided maps and
forest stand information. This study was supported
through funding and in-kind support from Human
Resources and Skills Development Canada, Innu
Nation, Newfoundland and Labrador Department of
Browsed stems
lon
Proportion Standard error
0.006 0.006
0.007 0.003
0.001 0.001
0.093 0.013
0.333 0.071
0.500 0.100
0.006 0.003
0.002 0.001
0.488 0.044
0.800 0.133
Natural Resources, and the Western Newfoundland
Model Forest.
Documents Cited (marked * in text)
Dalton, W. J. 1986. Moose census in Labrador on Manage-
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1986. A report prepared for the Government of Newfound-
land and Labrador Wildlife Division, Project Number 4403,
53 pages.
(FMDPT) Forest Management District 19A Planning
Team. 2003. Five Year Operating Plan for Forest Manage-
ment District 19A (Goose Bay). 75 pages. Government of
Newfoundland and Labrador, Goose Bay, Newfoundland
and Labrador, Canada
Meades, S. J. 1990. Natural regions of Newfoundland and
Labrador. Protected Areas Association, St. John’s, New-
foundland. 103 pages.
Schmelzer, I., J. Brazil, T. Chubbs, S. French, B. Hearn, R.
Jeffrey, L. LeDrew, H. Martin, A. McNeill, R. Nuna,
F. Phillips, G. Mitchell, G. Pittman, N. Simon, and G.
Yetman. 2004. Recovery strategy for three woodland
caribou herds (Rangifer tarandus caribou; boreal popu-
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and Wildlife Division, Department of Environment and
Conservation, Government of Newfoundland and Labra-
dor, and the Nationally Endangered Wildlife Committee
(RENEW).
2007
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Received 12 September 2005
Accepted 21 July 2008
Movements of Transient Coyotes, Canis latrans, in Urbanized Eastern
Massachusetts
JONATHAN G. Way!”
'Science Department, Barnstable High School, 744 West Main Street, Hyannis Massachusetts 02601 USA; e-mail: jw9802@
yahoo.com.
"Eastern Coyote Research, 89 Ebenezer Road, Osterville, Massachusetts 02655 USA
Way, Jonathan G. 2007. Movements of transient Coyotes, Canis latrans, in urbanized eastern Massachusetts. Canadian Field-
Naturalist 121(4): 364-369.
I document the movements of five transient (or nomadic) eastern Coyotes (Canis latrans) in heavily urbanized eastern Massa-
chusetts. Linear movements from capture location to end location varied from 23.0 to 100.5 km and averaged 63.8 + 52.0 km
for two females and 38.7 + 17.2 km for three males (t = 0.657, df = 1.15, P = 0.618). Transients ranged in age between 1-2 yr
old. There was no relationship between Coyote body weight and dispersal distances (r = 0.389, P = 0.518). Coyotes travel long
distances even in human-dominated areas, allowing transients to find vacant territories. Because of the ability of Coyotes to
colonize and recolonize areas, I recommend that Coyote management efforts focus more on educating the public about actual |
Coyote behavior and their life history needs than on killing them.
Key Words: Eastern Coyote, Canis latrans, dispersal, human-dominated landscapes, movements, suburban, transient, urban-
ized Massachusetts.
Coyotes (Canis latrans) typically live in packs con-
sisting of a breeding pair, resident associates (helpers),
and pups of the year (Gese et al. 1996; Way et al.
2002a). Coyote packs defend a territory while nomadic
or transient Coyotes, usually young, but sometimes old
individuals (see Way 2007a), travel among a matrix of
territories as they disperse from their natal range (Har-
rison 1992; Way et al. 2002a). Individual Wolves
(Canis lupus) and Coyotes are known to disperse sev-
eral hundred kilometers from their natal range (Fritts
1983; Carbyn and Paquet 1986; Gese and Mech 1991;
Harrison 1992; Mech and Boitani 2003a), which facil-
itates recolonization of areas where control actions
limit their numbers or where they are expanding their
range (Parker 1995; Mech and Boitani 2003a).
Movements of dispersing transient Coyotes have
been documented in rural/forested areas (Harrison
1992), agricultural landscapes (Person 1988), and in
southern Canada (Carbyn and Paquet 1986). Coyotes
have also been documented to travel across seemingly
disparate areas, such as wide canals (Way 2002),
islands (Thomas and Dibblee 1986), and on drifting
pack ice (Chubbs and Phillips 2002). However, aside
from one documented Coyote in urban/agricultural
southern Canada (Rosatte 2002), there are no data on
transient/ dispersing Coyote movements in urbanized
areas. Documenting the movement of transient Coyotes
in urbanized areas will give managers data on how
Coyotes move in these landscapes compared to more
rural environs (e.g., Harrison 1992; Gese et al. 1996).
This could have practical implications. For example, if
transient Coyotes do not move far in urbanized locales
(because of the high amount of roads) then localized
control efforts may be more successful in reducing
364
Coyote numbers in those regions; conversely, if the op-
posite is true (1.e., Coyotes move similar distances in |
urban and rural areas), then control efforts would likely
be less successful, unless targeting a specific individ-
ual(s). This paper, part of a larger ongoing study on
Coyote ecology (Way et al. 2001; Way et al. 2002a;
Way et al. 2004) in eastern Massachusetts, documents
the movement of transient Coyotes in a heavily urban-
ized region.
Study Areas
This research took place in two urbanized locations:
Cape Cod and the towns and cities north of Boston
(Figure 1). Most research conducted on the heavily
urban north edge of Boston (~100-150 km?; 42.43°N,
71.06°W) took place in the cities of Revere (3089 peo-
ple/km2, housing density = 1318/km?), Everett (4345 |
people/km?, housing density = 1817/km7), and Malden |
(4291 people/km?, housing density = 1800/km7) (U.S. |
Census Bureau 2000 estimates). The area is charac- |
terized by high-density housing with small woodland |)
areas (including cemeteries) non-strategically situated |
in towns and cities. Coyotes were captured and spent |
most of their time in these wooded, green areas as the ;
high-density housing areas were often fenced and pro-~
vided nowhere for Coyotes to travel, except for maintl
roads. Railroad tracks and holes in some of the fences »
provided small corridors between some of the green'
areas (Way and Eatough 2006).
Cape Cod research was conducted within Barnstable
County, Cape Cod, Massachusetts (approximate study
area 250 km?), with a concentration in the town/city of
Barnstable (although called a town, Barnstable is techni-
cally a city; 41.67°N, 70.28°W; land area = 155.5 km”).
|
2007
Human population density in the town/ city of Barn-
stable was 308 people/km’ and housing density was
161/km’, while the entire Barnstable County (3382
km’) averaged 217 people/km? and 144 houses/km?*
(U.S. Census Bureau 2000 estimates). The town/ city
of Barnstable has a distinct rural-urban gradient with-
in its borders; the highest and lowest densities of peo-
ple were found in urban Hyannis (556 people/km’,
housing units = 328/km?) and rural West Barnstable
(89/km’, housing units = 39/km*) (Cape Cod Com-
mission 1998*). Road density, defined as centerline km
of roadway per km’, was 4.7 for the town of Barnsta-
ble and 4.0 for Barnstable County (Cape Cod Commis-
sion 1998*). Cape Cod is characterized by being resi-
dential as well as having numerous small (5-10 ha)
and a few large (~1000 ha) conservation areas inter-
spersed throughout. Most of the neighborhoods are
not fenced, however, and Coyotes were readily able to
travel through these areas to access various portions
of their home range (Way et al. 2004). Coyote pack
territories were roughly 30 km? and were non-over-
lapping, similar to more rural areas (Gese et al. 1996;
Way et al. 2002a).
Methods
Coyotes were captured by box trap (Way et al.
2002b) then radio-collared or radio-implanted (juve-
niles — i.e., pups of the year) using Telonics, Inc.
(Mesa, Arizona) transmitters, aged based on tooth wear
(Bowen 1982; Landon et al. 1998), weighed, blood
drawn (ca. 4 cc), then released. Transient (or nomadic)
Coyotes were classified as Coyotes who had no dis-
cernable territory and nomadically moved throughout
the study areas, including within resident collared
Coyote home ranges. These Coyotes are typically clas-
sified as young Coyotes that are in the process of dis-
persing from their natal pack (Way et al. 2002a).
Tracking protocols were described by Way et al.
(2002a) and Way et al. (2004). Portable receivers (Cus-
tom Electronics, Urbana, Illinois, USA) and hand-held
3-element Yagi antennas were used to radio-track Coy-
otes both on foot and from a vehicle. Due to the highly
developed landscape with many roads, I mostly radio-
tracked in a vehicle as Coyotes did not react nega-
tively to them as much as they did to people (e.g., by
running away; Way 2007a; J. Way, unpublished data);
occasionally I approached radio-collared Coyotes as
closely as possible on foot without disturbing them. I
used binoculars and video-cameras when observing
Coyotes, and city street lights, nightscopes and occa-
sionally headlights when following Coyotes at night
with a vehicle (Way et al. 2002a; Way et al. 2004).
Due to funding constraints, I did not use airplanes to
search for Coyotes that left our study areas; extended
trips were made in vehicles to locate missing Coy-
otes but this was often unsuccessful. Instead, I relied
on recovering Coyotes from sightings by the public
(n = 1), opportunistically receiving radio-locations in
Way: MOVEMENT OF TRANSIENT COYOTES 365
new areas (n = 1), and recovering carcasses from
human-related kills (i.¢., road-kill or gunshot; n = 3)
Transient movement distances were calculated from
where they were first captured (if not on natal territo-
ry) or from their natal territories (if known) to their
final location when they either settled (i.¢., estab-
lished a resident home range in a new area) or died.
I used an independent sample two-tailed t-test to
detect differences between male and female dispersal.
Levene’s test was used to detect for equal variance
between samples; a significant Levene’s result indi-
cated that equal variances were not assumed. I corre-
lated Coyote dispersal distances with body weight
using two-tailed bivariate Pearson Correlation Coef-
ficients (SPSS Inc., Chicago, Illinois) tests. I set sig-
nificance at < 0.05.
Results and Discussion
I documented the movement of five transient
Coyotes: one from north Boston and four from Cape
Cod (Figure 2). Upon release, all Coyotes appeared
to be in the process of dispersal as their movements
were nomadic. Movements from capture to final
location varied from 23.0 to 100.5 km and averaged
63.8 + 52.0 km for two females and 38.7 + 17.2 km
for three males (Levene’s Test = 0.030, t = 0.657,
df = 1.15, P = 0.618). Although I had a low sample
size, these movements were within the range of dis-
persal by transient Coyotes in more rural environ-
ments, with an average distance of 98 km for Coyotes
in forested Maine (Harrison 1992), 16 — 152 km in
rural Ontario (Kolenosky et al. 1978), 20 — 140 km in
rural, agricultural Vermont (Person 1988), 80%) of the shrub particles were classified as
“Other” or “Unknown”, so we were not able to deter-
mine if they should be classified as evergreen or decid-
uous shrubs. Therefore we were unable to apply Rus-
sell et al.’s (1993) correction factor for digestibility.
Paired t-tests were employed to identify significant
changes in percent cover and diet between 1995/1996
and 2005. We calculated Ivlev’s index of selectivity
(Ivlev 1961) using the formula: (U-A) / (U+A), where
U (utilization) was the amount of a vegetation type
found in the fecal matter and A (availability) was the
percent cover of that type found on the paired transect.
P T df
0.001 3.73 23
0.002 3.53 23
0.351 0.95 23
0.169 1.42 23
0.155 1.47 73
Results
The permanent vegetation transects indicated
changes in vegetation from 1995/1996 to 2005 in the
winter range of the WAH. Significant declines in
lichen abundance were noted, while both graminoids
and shrubs increased during this time period (Table
1; Joly et al. 2007b).
The changes in vegetative cover were mirrored by
changes in the paired fecal composition of WAH Cari-
bou between 1995/1996 and 2005 (Table 2). The pro-
portion of lichen found in the fecal pellets declined
significantly (T = 3.73, P < 0.001, df = 23), while
the proportion of graminoids increased (T = 3.53,
P < 0.0021, df = 23). The mean proportion of Cladina
spp. declined from 21.8% to 15.2% from 1995/1996
to 2005 (T = 4.71, P < 0.001, df = 23). This decline in
Cladina spp. represented 84% of the decline in over-
all lichen in fecal matter over this time period. There
were no significant differences in the proportion of
Cetraria spp., Cladonia spp., or non-preferred lichens
(e.g., Alectoria spp., Bryoria spp., Peltigera spp., etc.)
between 1995/1996 and 2005. There were no signifi-
cant differences in the other vegetative classes (Table 2).
Lichens and mosses were found in the fecal matter
of WAH Caribou more than they were present in the
environment in both 1995/1996 and 2005 (Figure 1).
Ivlev’s (1961) index of selectivity increased for lichens
between 1995/1996 and 2005, despite declines in
lichen availability. The index decreased for mosses as
its availability declined (Figure 1). Graminoids and
shrubs were more common on the landscape than in
the fecal matter of Caribou in both 1995/1996 and
2007
0.8
0.6
Ivlev's Index of Selectivity
i)
N o
Lichens Graminoids
JOLY, COLE, AND JANDT: DIETS OF OVERWINTERING CARIBOI 38]
Moss Shrubs Forbs
FiGurE 1. Ivlev’s index of selectivity for Caribou utilizing different vegetative categories on the winter range of the Western
Arctic Herd, northwestern Alaska, USA. Light-colored bars represent data from 1995/1996 and the dark bars from
2005. An index of 0 would occur when the amount of that category was found in equal percentages in the fecal mat-
ter and along paired permanent vegetation transects. Ratios > 0 reveal vegetation types that were greater in the diet
than found on the landscape. Ratios < 0 reveal vegetation types that were less prevalent in fecal matter than on the
landscape.
2005 (Figure 1). Ivlev’s (1961) index of selectivity
declined between 1995/1996 and 2005 in the face of
increasing availability of these vegetation types.
Discussion
Lichen content in the winter diet of Caribou has
been reported to be from 62 — 69% (Thompson and
McCourt 1981; Boertje 1984; Boertje et al. 1985;
Russell et al. 1993). While our results show somewhat
lower lichen content (51 — 59%), we were not able to
apply correction factors to account for differential
rate of digestibility among vegetation types. Lichen
content is typically under-represented without cor-
rection factors (Russell et al. 1993), so our results are
likely similar to these previous studies (though see
Boertje et al. 1985).
We know of no other studies that have tracked the
diets of Caribou over a decade and directly related
detailed vegetative cover surveys to dietary analyses.
Our results show that the recent winter diet of WAH
Caribou contains less lichen and more graminoids
than in the past, tracking changes in vegetative cover
from 1995/1996 to 2005 reported by Joly et al.
(2007b). The percent of lichen in the fecal matter was
3.5 — 4 times greater than the percent of lichen cover
in the environment, suggestive of selection. Caribou
used areas of high lichen cover more than other areas
(Joly et al. 2007b). The significant reduction of Clad-
ina spp. in the fecal matter between time periods was
also reflected in declines of primary forage lichen on
the landscape (Joly et al. 2007b). Grazing was likely
a factor that drove the decline in lichen in northwest
Alaska (Joly et al. 2007b). It may take a couple of
decades for lichens to recover from intense grazing,
barring further disturbance (Henry and Gunn 1990).
Shrubs are significantly increasing in the Arctic
(Sturm et al. 2001; Joly et al. 2007b); however, we
did not detect a corresponding increase in shrubs in
the diet of WAH Caribou. Shrub expansion could have
negative impacts on individual Caribou energetics and
multiplier effects on the herd (see Lawler and White
2006). Other researchers have reported increased moss
cover where Caribou range has been heavily grazed
(Klein 1987; van der Wal 2006), but changes in moss
cover were not detected within the range of the WAH
during this time period (Joly et al. 2007b). Mosses were
present in the fecal matter in greater percentages than
were found on the landscape; however we believe this
is due to its low digestibility rather than Caribou select-
ing for this forage type (Boertje et al. 1985).
Climate warming is another factor that has been
implicated in declining lichen cover within the range
of the WAH (Joly et al. 2007b). Fire activity is strong-
ly correlated with warm summer temperatures in
Alaska and Canada (Duffy et al. 2005; Kasischke and
Turetsky 2006). The frequency and extent of fires is
predicted to increase due to global warming (Rupp et
al. 2000; McCoy and Burn 2005). Lichens within the
range of the WAH are slow to recover from wildfire,
often taking many decades to recover to initial levels
(Arseneault et al. 1997; Jandt et al. 2008). Overwin-
tering Caribou are known to avoid burned areas in
both tundra and boreal forest, in northwest Alaska,
for up to 55 years (Joly et al. 2007a). Lack of lichens
382
in these areas has been hypothesized to be a reason
for this avoidance though other factors have been
suggested (see review by Klein 1982).
Lichen cover has been declining in the study area
since 1981 (Joly et al. 2007b). Potential outcomes of
continued declines in lichen cover in winter range in-
clude range shifts and population-level effects (Fer-
guson et al. 2001; Joly et al. 2007b). Caribou nutri-
tional status should not be assessed based on lichen
abundance alone (Boertje 1990). Studies in the high
Arctic studies have shown that Caribou can be less
reliant on lichens (Thomas and Edmonds, 1983;
Adamezewski et al. 1988). However, these Caribou do
not face the predatory pressure and migratory expense
that WAH Caribou do and also differ phenotypically.
Additional differences between these types include
greater ability of the high Arctic Caribou to store fat
and digest graminoids and mosses (Tyler 1987). Large,
migratory herds of barren-ground Caribou, like the
WAH, are thought to be reliant on abundant terricolous
lichens (White et al. 1981; Klein 1991; Heggeberget
et al. 2002). Nevertheless, it has been suggested that
Caribou may not be negatively affected by a change
from a lichen-rich diet to a graminoid-rich diet as
ecosystems transition to graminoid-dominated states
(van der Wal 2006).
Adult survival rates would be the least sensitive pop-
ulation parameter to potential declines in nutritional
status, so biologists and managers should monitor
pregnancy rates and calf to cow ratios to detect early
signs of nutritional stress. Declines in these parame-
ters may imply that the nutritional status of the herd
is declining due to poor range conditions and that the
transition to a graminoid diet may impact the herd.
Recruitment in the WAH has been slowly declining
(Dau 2005a) and the most recent photo-census (2007)
revealed a 20% decline in the herd (J. Dau, personal
communication), which may be incipient indications
of the importance of lichens to this herd. Predation is
not likely a major factor in these declines with the
herd size being so large and predator densities rela-
tively low (Ballard et al. 1997; Haskell and Ballard
2007), though density-independent icing events may
be a factor (Dau 2005b). Although the debate over the
relative importance of lichens in the winter diet of
Caribou remains unresolved, our results reveal that
WAH Caribou use lichens extensively in the winter
despite their declining abundance on the landscape in
northwestern Alaska.
Acknowledgments
Fecal analysis was conducted by the Wildlife Habi-
tat Nutrition Analysis Laboratory at Washington State
University, Pullman, Washington, USA. R. Meyers,
(retired) Bureau of Land Management, was an integral
member of the research group during both study peri-
ods. We thank J. Lawler, D. Spalinger, J. Welker, S.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Wilson and two anonymous CEFN reviewers for pro-
viding recommendations for substantially improving
previous drafts of this manuscript.
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Climatic
Received 4 July 2007
Accepted 20 August 2008
An Analysis of the Historical Records for the Native Mammalian
Fauna of Prince Edward Island
DouGLAS G. SOBEY
Research Associate, Institute of Island Studies, University of Prince Edward Island, Charlottetown, Prince Edward Islanq)
CIA 4P3 Canada
Summer address: P.O. Box 4004, Bedeque, Prince Edward Island COB 1CO Canada
Formerly: School of Applied Biological Sciences, University of Ulster, Jordanstown, Northern Ireland, BT37 OQB Unitec
Kingdom
Sobey, D. G. 2007. An analysis of the historical records for the native mammalian fauna of Prince Edward Island. Canadiat
Field-Naturalist 121(4): 384-396.
A search was carried out for historical records, both published and unpublished, that make reference to the native mammaliar’
fauna of Prince Edward Island. Based on documents dating from 1721 to 1890, a comprehensive list of the records for the.
native mammals of the island has been compiled. Among the new information found is evidence for the presence of the Grey)
Wolf (as well as the Woodland Caribou) at the time of the first French settlement in 1720, and for the absence of the Beaver anc
Moose. Historical information has been assembled on the abundance and food-chain relationships of each of the mammaliat,
species, as well as on their interactions with the European population, including the attitudes of the new settlers towards eacl,
species. The records indicate that seven of the mammals were extirpated: the Grey Wolf, American Black Bear, Americar’
Marten, River Otter, Canada Lynx, Atlantic Walrus and Woodland Caribou. All of these extirpations were due to the activities!
of the European population, with the attitudes of the settlers contributing to four of them: an indifference to the survival o
the otter and Marten, and a direct hostility to the bear and lynx (due to their predation on livestock), leading to the paymeni
of bounties.
Key Words: historical records of mammals, mammalian fauna, Prince Edward Island, Grey Wolf, American Marten, Atlantic:
Walrus, Woodland Caribou, Moose, Beaver.
At the time of the first European settlement on Prince
Edward Island in 1720, the island supported a varied
mammalian fauna. In the course of the next 180 years,
the native fauna was to become greatly reduced, with
six terrestrial species (five of them predators) and one
marine mammal being eliminated from the system.
Despite an awareness of this general picture (e.g., Clark
1959), the many historical accounts relating to the is-
land’s native mammalian fauna have never been sub-
jected to comprehensive analysis. It was hoped that a
detailed examination of the records would not only
reveal what species were present when European set-
tlement began, but also yield information on the inter-
actions between the native fauna and the European
population, including the extent to which the fauna
contributed to the food supply of the human popula-
tion and to any fur-trade economy that developed, as
well as an understanding of the factors that led to the
extirpation of most of the fur-bearing predators. Ini-
tially, my aim was to examine only the records for the
terrestrial fauna, though later the pinnipeds occurring in
island waters (i.e., the seals and the walrus) were also
included.
The historical context
Prince Edward Island (known as Ile Saint-Jean to the
French) was “discovered” by Jacques Cartier in 1534.
However, historical accounts in any significant num-
bers only begin to occur from 1720, the year of the
384
|
arrival of some 250 colonists from France. (See Clark
(1959) for an in-depth study of the island’s historica
geography.) Prior to the arrival of the French, aborigina
use by the Mi’kmaq appears to have been confined tc.
the summer period, as part of their seasonal migratior |
to the coastal areas of the Maritimes, though with the:
establishment of the first French settlement some of the
Mi’ kmaq began to spend the whole year. The Frenct |
population grew slowly, so that by 1748 there were
only 735 persons. However, numbers then rose sharply
reaching almost 5000 by 1758, due to the arrival oi)
Acadian refugees from the Bay of Fundy area on ac-
count of the political unrest and deportations occurring;
in Nova Scotia. After the fall of Louisbourg in 175&
almost all of the Acadian population was deported tc)
France by the British military authorities, or fled to they
mainland, leaving only a few hundred refugees. From
the 1770s British settlement proceeded slowly, with by)
1805 the population reaching only 6957. However, i
then began to grow exponentially, reaching 71 000 by)
1855, when immigration from the British Isles began
to end. Even so, natural growth continued, with num)
bers peaking at 109 000 in 1891 (Clark 1959). This in-
crease in the human population was associated with é
decrease in the area forested, and by 1935 only 32% ol
the land area still had a forest cover (Glen 1997).
Prior to European settlement virtually all of the is-
land’s land area of some 5750 km? had been covered by
old-growth forest, with the predominant forest-type
/
|
2007
TABLE |. Twenty-one lists of the mammals of Prince Edward Island
The lists are in chronological order, with the names of the recorders given below the table
end of the paper. See the main text for the identification of the species. (E
occurring on the island, or the evidence ts for the past occurrence only; A
SOBEY: HISTORICAL RECORDS FOR NATIVE MAMMALS OF PEI
385
made by different recorders between 1721 and 1890
these are fully referenced at the
exurpated — 1.¢., reported as either no longer
reported as absent from the island.) *
THE Lists o%n
MAMMAL Le Sa ae Sar 6 Tee 8 LO Tv ies aea TS eT 7 Peo Zoot aor
‘Shrew’ . . 2
‘Bat’ © . . . 4
‘Mole’ A . . 2
‘Wolf’ ° A ie 3
‘Fox’ . . ° . . . ° . . . . . . . . . . . . . . 21
‘Bear’ . . . 2 . . . . ° . . . . . 7 . . . . 19
‘Marten’ ° ° ° ° . ° ° . . ° . . . . . . . . . 19
‘Weasel’ or ‘ermine’ ° . . <— . 6
‘Mink’ e e ° othe. ce met © tefl tle e ° ° eit, 14
‘Otter’ sth ve av ie ol 's ou te 6 et! Ne ° st ‘cs ° owas 17
“Wild cat’ or lynx orire ome: Wie? CCH cine Se Te aire as BC . UE 18
‘Seals’ e e ° ° ° ° ° ° ° ° . . . . . 15
‘Sea cow’ or walrus ° ° OO” OREC JE E B. WBE 1]
‘Caribou’ Can ° BE ACE 6
‘Moose’ A EB Au A. aie sre oe 5
‘Deer’ A A E l
‘Squirrels’ Om FC ae os 6
Red squirrel ov Prony ke on Rinre . 6
Flying squirrel ° ohihaels hte Siaine ° 7
Chipmunk ow ieniite oh) is . 6
‘Beaver’ A A A E l
‘Muskrat’ ° e e ° ° ° ° e ° °° . : ° . © l 5
‘The plague mouse’ Oo OO ° ° ° ° ° 25 ae 10
Other mice and voles ° ° ° ° ° . 6
‘Hare’ or ‘rabbit’ e e ° e e e e e e e e e . ° ° ° . . . 19
TOTAL Con eOn Oe sleno cit 2 or 12" 16a tO. Lae S es OF ere tye
The recorders: 1 — La Ronde 1721; 2 — Roma 1750; 3 — Franquet 1751; 4 — Pichon 1760; 5 — Holland 1765; 6 — Patterson
1770; 7 — Patterson 1774; 8 — Shuttleworth 1793; 9 — [Cambridge] c.1796; 10 — Walsh 1803; 11 — Stewart 1806; 12 — John-
stone 1822; 13 — MacGregor 1828; 14 — Hill 1839; 15 — Lawson 1851; 16 — Bagster 1861; 17 — Sutherland 1861; 18 -
Rowan 1876; 19 — Anonymous 1876; 20 — [Lawson] 1877-1878; 21 — Bain 1890.
* List 19 is a composite list assembled from the responses of eighteen men to several questions on the mammals occurring
on the island in their “young days”, contained in a questionnaire sent out in 1876 to the “oldest inhabitants” by a Charlotte-
town historical committee; list 20 has been put together from anecdotes on the island’s native fauna contained in a series of
nineteen articles on pioneer life published in 1877 and 1878 in the Presbyterian and Evangelical Protestant Union.
particularly in the central part, consisting of hardwood
forest in which American Beech (Fagus grandifolia)
especially, but also Sugar Maple (Acer saccharinum)
and Yellow Birch (Betula alleghaniensis) predominat-
ed (Sobey and Glen 2002, 2004; Sobey 2006). In the
eastern and western parts, where there was a higher
proportion of poorly drained soils, other forest-types
occurred, especially a type of “swamp” woodland,
containing Red Maple (Acer rubrum), Eastern White-
cedar (Thuja occidentalis), Black Ash (Fraxinus nigra)
and American Elm (U/mus americana), as well as a
boreal forest-type in which Black Spruce (Picea mari-
ana) predominated along with a minority element of
Tamarack (Larix laricina). The upland hardwood for-
est in particular was subjected to large-scale clearance
for agriculture, with any remnants (often in the form
+ See List-makers cited in Table 1 after Literature Cited.
of farm woodlots), undergoing considerable change
due to the continual harvesting of selected species for
timber and firewood. As well, in all forest-types there
were the damaging effects of the ubiquitous forest-
fires associated with forest clearance. The removal of
this old-growth forest and the alteration of that which
remained would have had considerable consequences
for the island’s mammalian fauna.
Results
From the beginning of European settlement in 1720
there is a regular flow of relevant documents from the
French (1720-1758), British (1758-1873) and post-
Confederation periods (1873 to c.1900). These com-
prise both published books, such as immigrants’
handbooks and travel accounts (most of which were
386
available in the P.E.I. Collection of the Robertson
Library of the University of Prince Edward Island),
and unpublished documents, including government
records and private journals, many of which were
housed in the Provincial Archives and Public Records
Office in Charlottetown (PEI PARO). Some 58 docu-
ments were found that make reference to one or more
of the native mammals of the island — these are fully
reported in Sobey (2002, 2006). Especially informa-
tive are those records where the writer constructed a
list of the mammals of the island: twenty-one such
lists have been assembled in Table 1. Four come from
the French period (all based on observations made be-
tween 1720 and 1752) and seventeen from the British
and post-Confederation periods. Many of these list-
makers also made short comments on the abundance
and/or ecology of some or all of the mammals, as well
as on their relations with the human population.
The lists range from four recorders who mention
only six species, to one listing twenty-one. All but four
of the list-makers appear to have been resident on the
island for at least a year, and most for many years.
The exceptions are Franquet (17517), Pichon (17607),
Walsh (18037) and Rowan (18767), who spent from
seventeen days to a few months. However, given the
elusive nature of most of the mammals, it is unlikely
that even the resident recorders would have seen every
animal they listed, and it is especially likely that ani-
mals that were known to have occurred in the past
would have continued to have been listed for some
time after their extirpation. Also, there is a likelihood
that some of what they record may derive not from
their own observations, or even from the anecdotal ex-
perience of other island residents, but from general
written sources from elsewhere.
Table | indicates that, collectively, there is a bias in
the selection of the animals recorded. The smaller
mammals (the shrews, bats and rodents) were either
ignored, or else were treated generically, except by a
few of the late nineteenth century “scientific” record-
ers, such as Sutherland (1861+) and Bain (18907),
whereas a greater coverage was given to those mam-
mals considered “useful” to the human population; i.e.,
as a source of food or products such as furs. Also, some
of the animals received greater attention because of
their predation on the settlers’ livestock and poultry.
The records for each mammal are presented separately
below, with the nomenclature and order following that
of Scott and Hebda (2004).
Shrews
The Atlantic Canada Conservation Data Centre
(ACCDC 2008*) lists five species of shrew as cur-
rently occurring on Prince Edward Island: the Masked
Shrew (Sorex cinereus), Water Shrew (Sorex palus-
tris), Smoky Shrew (Sorex fumeus), Pigmy Shrew
(Sorex hoyi), and Northern Short-tailed Shrew (Blari-
na brevicauda). However, the only historical records
THE CANADIAN FIELD-NATURALIST
Vol. 121
for the presence of shrews are those of Sutherland
(18617) and Bain (18907), who recorded the presence
of what they called the “shrew mouse” and “shrew
mole’, respectively (Table 1), both mentioning that it
fed on insects.
Bats
ACCDC (2008*) lists four species of bat for Prince
Edward Island: the Little Brown Bat (Myotis lucifugus),
Northern Long-eared Bat (Myotis septentrionalis),
Hoary Bat (Lasiurus cinereus), and Eastern Red Bat
(Lasiurus borealis). For the Eastern Red Bat there is a
sight record only. Not surprisingly, the four recorders
who noted the presence of bats (Table 1) listed them as
if only one species were present. All four recorders
indicated that bats were common during the summer,
and Bain (18907) said that they “[hid] away in some
secure cranny in buildings or in a hollow tree”.
Grey Wolf
The only certain record for the Grey Wolf (Canis
lupus) is that of La Ronde (17217), who said that there
were wolves “of a prodigious size” on the island. That
he also added that he was sending a wolf pelt back to
France from the island, both supports his record and
indicates they were vulnerable to the presence of the
newly arrived Europeans, though it is not impossible
that it was the Mi’kmaq who were responsible for the
kill. Since there is no further mention of the presence
of wolves in eighteenth century records, it is possible
that the Grey Wolf was extirpated shortly after settle-
ment began, or else moved across the ice to the main-
land. It is thus surprising that there are a couple of late
post-Confederation mentions of its presence ([Lawson]
1877-18787; Bain 1890+), both of which report, on the
basis of anecdotal and second-hand information of un-
known date, wolves as transient visitors. What Bain
(18907) says is that “wolves have been known to cross
the Northumberland Strait on the ice and visit the
Island”. If this ever occurred, it was likely to have been
long before the 1890s when the island was at the peak
of European settlement and forest clearance, and when
according to Lohr and Ballard (1996) the Grey Wolf
was very scarce on the mainland.
Red Fox
The Red Fox (Vulpes vulpes) occurs in all twenty-
one lists (Table 1), with several recorders noting it
as common (MacGregor 18287; Sutherland 18617;
Anonymous 1876+; Bain 18907). Others noted the
presence of different colour phases (e.g., Franquet
1751+; [Cambridge] c.1796+; Stewart 18067): red,
black, and silver-grey are the most frequently listed,
though all noted that the red phase was by far the pre-
dominant. A fox fur of any colour seems to have had
some value, though it was the blacks and silver-greys
that were especially sought after. Stewart (18067) said
that the number of foxes caught on the island was
2007
“very considerable”, while the means of catching them
included steel traps and shooting (Stewart 18067;
MacGregor 18287; Hill 1839+). There are varying
opinions on the fox’s pest status, several persons re-
cording that it occasionally took poultry, though they
differ as to whether it attacked sheep (Stewart 18067;
MacGregor 18287; Johnstone 18227). As for their nat-
ural food, according to Bain (1890%), it was “young
birds and quadrupeds”, while MacGregor (18287)
more vaguely mentions that they fed “in the woods or
along the shore”.
Black Bear
The American Black Bear (Ursus americanus) was
recorded by nineteen of the twenty-one list-makers
(Table 1) and during the British period several re-
corders devoted a paragraph or more to it, and espe-
cially to its interactions with the human population.
It would seem that in the early years bears were abun-
dant on the island: for example, in February 1780 the
ensign of a visiting regiment ({Ritter] 178077) record-
ed that his soldiers were shooting the bears that came
near their huts at Charlottetown, while the proprietor
Shuttleworth (17937) reported that where he lived
(near St. Peters Bay) bears “swarmed to a degree that
[he] could not have believed”, and the visiting bishop
Plessis (181277) said “they are seen in greater num-
bers than one would expect’. However, just ten years
later Johnstone (18227) noted that they were “decreas-
ing in numbers greatly”, with the “few of them yet in
the woods” being only “seen occasionally by the in-
habitants”. There are similar comments on its decline
by MacGregor (18287), Bagster (18617) and Anony-
mous (18767), with by the end of the century, Bain
(18907) saying that bears were restricted to “the large
wooded tracts” in the east and west of the island.
Since, beginning as early as Stewart in 1806, several
writers were predicting the Black Bear’s impending
extirpation, it comes as a surprise that the last record-
ed bear on Prince Edward Island was shot in 1927
(Hornby 1987).
There was an almost total antipathy to the Bear,
largely due to its habit of killing livestock, a practice
not discouraged by the fact that in new settlements,
farm animals were let loose in the woods (e.g., Stew-
art 18067). Sheep are the most frequently mentioned
as being taken (by fourteen recorders), followed by
pigs and cattle (by eight and nine recorders respec-
tively). Not surprisingly, these attacks seem to have
been greater in the early years, Stewart (18067) record-
ing that “the quantity of black cattle, sheep and hogs
destroyed by them annually is considerable”. Howev-
er, later reports (Johnstone 1822+; MacGregor 18287)
suggest that such predation was only occasional, pre-
sumably due to the decline in the number of bears.
Bain (18907) also reported that bears could do “great
damage” by feeding on oats in the field.
SOBEY: HISTORICAL RECORDS FOR NATIVE MAMMALS OF PEI
387
In terms of their danger to people, the general con-
sensus was that bears avoided humans and only
attacked when either threatened or with their cubs
(Stewart 18067; MacGregor 18287; Bagster 18617;
Sutherland 1861+; Anonymous 18767; Bain 18907).
All the same, it was the perception of the Black Bear
as a potential danger, based on perhaps a few incidents,
which greatly added to the human antipathy towards
them. In fact, even in the early years the list of their
“evils” was considered so great that an official extermi-
nation program was instituted in the 1790s (Vass 1987).
This involved the payment of a bounty of 15 shillings
per bear (a large sum at the time), and remained in
effect until the 1860s. Vass (1987) estimated that be-
tween 1820 and 1861 bounties were paid on over a
thousand bears. Apart from the bounty, the Black Bear
provided other resources to some of the island’s human
population: its skin was especially valuable (MacGre-
gor 18287; Sutherland 18617), and it was also a source
of food for some, especially the Mi’ kmaq (Johnstone
18227), and the Acadians during their refugee years
(Holland 17657; Patterson 17707), and seemingly for
others as well (Sutherland 18617; Bagster 18617).
We are told a little about the Black Bear’s natural
diet: several writers noted that it fed primarily on
“berries”, “wild fruits” and “small shrubs”, as well as
on insects (ant hills and grubs in old fallen trees were
favourite foods), and on smelts in the spring (Stewart
18067; MacGregor 18287; Sutherland 18617; [Law-
son] 1877-18787; Bain 18907). Curiously, none of the
recorders mention them feeding on the island’s abun-
dant beech mast resource, for which there is evidence
from elsewhere (e.g., Vass 1987; Telfer 2004).
Marten
The American Marten (Martes americana) occurs
in nineteen of the twenty-one lists (Table 1). The only
comments on its abundance are Stewart's (18067) that
it was “a very shy little animal, seldom seen in the
woods, though some years in great abundance”, and
Sutherland’s (18617) inclusion of it among those
mustelids that were “much more numerous” than the
“comparatively scarce” otter. Bain (18907) noted that
it was “a woodland animal nesting in hollow trees and
feeding on birds and small animals”, and they also ap-
pear to have fed on voles during the “mouse plagues”
(MacLeod 187677). None of the recorders seemed
aware of its approaching extirpation, for at some peri-
od, perhaps even in the late nineteenth century, the
Marten became extinct on the island. As for its fur,
several recorders commented on its value (e.g., Mac-
Gregor 18287; Hill 18397; Sutherland 18617), while
Holland (17657) reported that martens were one of
those animals that the refugee Acadians were com-
pelled to use as food after the deportation of most of
the French population in 1758.
388
Weasel
The first record for the presence of the Short-tailed
Weasel or Ermine (Mustela erminea) on the island is
that of Stewart (18067) (Table 1). In fact, he treated it
as two separate species, as did several later recorders.
One of these he called the “weasel”, which he said was
“common” and “often destructive amongst poultry”,
while the other, the “ermine”, was “uncommon” and
made its nest inside fallen trees. MacGregor (18287),
continuing this distinction, said that “weasels and
ermines were not common’, as did Bagster (18617),
who, however, considered only one species to be pres-
ent, which he called the ermine or “white weasel”. In
the same year Sutherland (18617), still listing it as two
separate species, said they were “much more numer-
ous” than the otter, and that their fur “brings a fair
price’, which is the only comment on its value as a fur-
bearing animal. Apart from Stewart (18067) above,
no other author implies that it was a pest, and Bain’s
(18907) comment that it was “frequently seen about
farmsteads where it comes in search of mice’, indi-
cates that it was then not uncommon, and at the same
viewed as an assistant in mouse control.
Mink
The American Mink (Mustela vison) is first record-
ed in the French period (Franquet 17517) and there-
after appears in thirteen of the British period lists
(Table 1). The only comment on its abundance is
Sutherland’s (18617) that it was much more numer-
ous than the “comparatively scarce” otter. Its habitat
associations are encompassed in Stewart’s (18067)
description of it as “amphibious”, while Bain (18907)
used the term “semi-aquatic”’, saying that it fed on fish
and frogs, as well as on “small land animals”. Bagster
(18617) also noted fish as its main food item. In terms
of its pest status, both Stewart (18067) and Bain
(18907) recorded that it took poultry and eggs, while
Bagster (18617) said that it also caused damage by
burrowing into mill-dams. Several recorders reported
its fur to be of value (Stewart 1806+; MacGregor
18287; Hill 1839+; Sutherland 1861+; Bain 18907),
though Bagster (18617) said it varied in quality.
Otter
The River Otter (Lontra canadensis) makes an ap-
pearance in seventeen of the twenty-one lists, begin-
ning with that of La Ronde in 1721 (Table 1). Due to
the high quality and monetary value of its fur (Stew-
art 18067; MacGregor 1828+; Hill 1839+; Bagster
18617; Sutherland 1861+; Bain 18907), in the British
period the otter was under considerable pressure from
trapping, and such trapping had also been recorded in
the French period (Roma 1750+; Pichon 17607). As
early as 1806 Stewart’s use of the past tense: “they
have been very plenty on the island” implies they
were in decline, despite his adding that “they are still
caught in considerable numbers”. By 1861 both Bag-
ster and Sutherland could note that otters were “‘get-
THE CANADIAN FIELD-NATURALIST
Vol. 121
ting very scarce” and “comparatively scarce”, respec-
tively, Sutherland attributing this to habitat loss, or as
he put it, “the increasing occupation of the country”.
Bain in 1890, by which time it must have been even
scarcer, if not already extirpated, makes no comment
at all on its abundance, instead noting that it lived in
ponds and streams feeding on fish, and that when these
failed, “it will attack lambs and poultry”. Holland
(1765+) noted that it was among those animals hunt-
ed by the refugee Acadians for food.
Lynx
The Canada Lynx (Lynx lynx) was recorded by
eighteen of the twenty-one list-makers (Table 1).
Though it was sometimes trapped for its fur (Stewart
18067; MacGregor 18287; Bagster 18617), and for
food (Holland 1765+; Patterson 1774+; Stewart
18067), it was more generally viewed as a harmful
predator of farm livestock. Johnstone (18227) said
that ‘wild cats’ sometimes preyed on cattle and sheep
in the woods, while MacGregor (18287), Bagster
(18617) and ({Lawson] 1877-18787) noted that it
specifically attacked sheep, killing several at a time
without eating them. This, however, is contradicted
by Stewart (18067), who said that “very few of them
have been known to attack sheep or lambs”. Given
this reputation, it is not surprising that in the 1820s a
bounty (of five shillings) was instituted in an effort to
exterminate the lynx, and from government records,
Vass (1987) estimates that 357 bounties were collect-
ed between 1820 and 1861. Given the ease with which
it could be killed with dogs, or caught in snares or
traps (e.g., Stewart 18067), combined with the loss of
its habitat (Lawson 18517), it is not surprising that an
animal that MacGregor (18287) had said was “rather
numerous”, and Bain noted had once been “not un-
common”, was by 1890 “unknown” (Bain 18907). As
for its natural foods, Stewart (18067) said that it lived
on “hares and partridges”; Sutherland (1861+) men-
tions “small mammals, mice, birds, rabbits etc.”; Rowan
(1876+) noted its predation on hares; while Bain
(18907) said it ate “small quadrupeds and birds”.
Seals
The presence of seals around the shores of the island
was noted by many recorders throughout the French
and British periods (Table 1), but without any other
information being given than that they provided a
source of meat and oil to the inhabitants, as well oil
and skins for export (the export of some 860 seal skins
was recorded in the island’s customs records between
1802 and 1807). All of the recorders write as if only
a single species were present except Stewart (18067)
who distinguished between two different “kinds” of
seal, one of which, he said, was “very common” in all
of the rivers and harbours of the island — this presum-
ably was the Harbour Seal (Phoca vitulina). The other,
he said, was “a larger kind brought on the coast annu-
ally in the month of April by the floating ice from the
2007
northward; ... sometimes vast quantities come, other
years little or none”. This must be the Harp Seal (Phoca
groenlandica), and Stewart goes on to describe the
hunting of them from schooners. MacGregor (18287*)
and Sutherland (1861+) combine together details appli-
cable to each of these species, without realising they
were describing two separate species, Sutherland add-
ing that by his day seals were less common than for-
merly.
Walrus
It is odd that French period recorders make no ref-
erence to the presence of the Atlantic Walrus (Odobe-
nus rosmarus) along the shores of the island, though
La Ronde (17217) noted the presence of the ‘vache
marine’, as he called it, in the Magdalen Islands, to
which he said they were sending a ship from the island
to participate in the summer hunt. The first record of
the walrus directly on the island itself is thus that of
the British surveyor Samuel Holland (1765*) (Table
1), who, however, stated that they “seldom or never
come on shore but are now and then killed in the spring
by the inhabitants in their canoes, when they happen
to float too near the shore on pieces of ice”. This, how-
ever, conflicts with some of the later British records,
especially that of Stewart (18067), who claimed that
in the 1770s (he himself having arrived on the island in
1775) “sea cows” had been “found in great numbers
on the north coast” (he especially mentions the area
near North Cape), and he added that between 1770
and 1775 they were caught in considerable numbers
there, the method being to drive the herds inland and
then slaughter them using long spears. However, at the
time that he was writing in 1806 he said that they “are
now become scarce and seldom seen ashore”. All of
the later historical records for the presence of the wal-
rus are thus retrospective (e.g., MacGregor 18287;
Hill 18397; Sutherland 18617), and all state that it was
no longer observed around the island. The animal was
killed especially for its oil, but there is also mention
of the value of its hide for making ropes, harness and
shoes (e.g., Curtis 1775++; Stewart 1806+; MacGre-
gor 18287; Hill 18397; [Lawson] 1877-18787), as well
as of its tusks for ivory (e.g., Stewart 18067; Mac-
Gregor 18287).
Caribou
The first record of the presence of the Woodland
Caribou (Rangifer tarandus) on the island is that of
Nicholas Denys (167277): he said that there were
only a few on account of the Mi’ kmaq being too fond
of them for food to allow them to increase. Somewhat
in conflict with this statement is that of La Ronde
(17217), who said that the Mi’kmag did not find the
Caribou easy to kill. The last French period record of
their presence is that of Roma (17507), who had left
the island in 1746. The only record of their possible
survival into the British period is that of Holland
SOBEY: HISTORICAL RECORDS FOR NATIVE MAMMALS OF PEI
389
(1765+) who noted that there were “some, but very
few Carriboux”. If the species was still present in
1765, then it would seem to have become extinct short-
ly after, for ten years later, in 1775, Thomas Curtis
recorded that none of the inhabitants whom he had
met (and one of these had been resident since 1768)
knew of any “deer” on the island (Curtis 177577). I
also say “if” for Holland’s record because, as has often
been demonstrated elsewhere, there is a tendency for
people to believe that an extirpated animal is still pres-
ent long after its disappearance, so that Holland, per-
haps having been told by the Acadians of its past pres-
ence in small numbers, recorded it as still present. It
is also significant that he did not include them as one
of the animals that the refugee Acadians hunted for
food, which they surely would have done, had it been
present. Thereafter in the nineteenth century, the past
presence of the Caribou was witnessed by the occa-
sional finding of a “time-worn” antler in the woods
(Lawson 1851+; Bagster 18617; Rowan 18767), though
such antlers were also sometimes erroneously cited
as evidence for the past occurrence of the Moose on
the island (see below).
Squirrels and Chipmunk
That squirrels occur in only thirteen of the twenty-
one lists (Table 1) is an indication not of their scarci-
ty, but that, being of little value as either food or for
their fur, they tended to be overlooked, except by those
who set out to list all of the island’s mammals. In fact
during the French period the only recorder to note the
presence of any of the squirrel species was La Ronde
(17217), who recorded the elusive flying squirrel, pre-
sumably because of its unusual mode of movement. In
the early decades of British settlement, although sev-
eral recorders noted the presence of different “kinds”
of squirrel, they did not name them. Thus it was not
until 1806 that the three species currently occurring
are specifically recorded: the American Red Squirrel
(Tamiasciuris hudsonicus), the Eastern Chipmunk
(Tamias striatus), and the Northern flying squirrel
(Glaucomys sabrinus) (Stewart 18067), and thereafter,
all three are listed by another five recorders (Table 1).
The red squirrel and the chipmunk were consid-
ered very plentiful (Stewart 18067; Johnstone 18227:
Sutherland 18617), but due to its nocturnal and elusive
habits there was some uncertainty as to the abundance
of the flying squirrel: Stewart (18067) considered it
not as common as the other two species, though Bain
(18907) said it was “not rare”. Stewart (18067) said
that all three species, and particularly the chipmunk,
increased “vastly” the year after a crop of beech
mast, while Bain (18907) said that the red squirrel
was “plentiful in every wood where beech nuts were
found”. Bain (18907) said that the chipmunk dug its
burrow “under the roots of the great trees in a spruce
wood”, while the flying squirrel nested in hollow trees.
390
Muskrat
The Muskrat (Ondatra zibethicus) was listed by
fifteen of the twenty-one list makers (Table 1), most
of whom make no further comment about it. Though
Bagster (18617) said it was “by no means plentiful’,
Rowan (18767) considered it to be “very plentiful’,
and Stewart (1806+), Sutherland (18617), and Bain
(18907) said it was frequently seen about streams and
ponds. There were opposing opinions about the value
of its pelt: Bagster (18617) said the fur was “useful
and saleable”, while Sutherland (18617) said it was
“thin and poor and little esteemed”. Holland (17657)
included Muskrats as one of the animals that the
refugee Acadians used for food, while Bain (18907)
noted that Muskrats, due to their habit of burrowing
into the banks, were “troublesome about mill dams”.
As for its natural foods, Sutherland (18617) and Bain
(18907) noted that it ate roots, shellfish and seeds.
“Plague Mouse”
During the French period, as well as in the first
eighty years of British settlement, virtually the only
species of mouse or vole that entered the records was
the one responsible in occasional years for the destruc-
tion of the settlers’ crops. This “plague mouse” dam-
aged the crops several times during the French regime
(e.g., Roma 17507; Franquet 17517; La Roque
175277), while during the British period there were
at least eight further outbreaks between 1770 and
1815 (Sobey 2006). However, by 1828 MacGregor
noted that within the previous twenty years little injury
had been done to the crops by “these mischievous
animals”.
Identifying the species responsible for the outbreaks
has not been easy. Roma’s (17507) detailed description
of the habits of the vole that caused the fléau [plague]
of 1738, appears to better fit the Southern Red-backed
Vole (Clethrionomys gapperi), rather than the Meadow
Vole (Microtus pennsylvanicus), which has been more
generally viewed as the culprit (e.g., MacQuarrie 1987):
Roma’s mulot, as he called it, was a woodland species
(this, confirmed by all of the other French and British
recorders, fits only the Southern Red-backed Vole); it
made substantial underground food caches for the win-
ter; and it was able to climb (although Roma said not as
well as the souris (probably the Deer Mouse, Peromys-
cus maniculatus). These last two properties are char-
acteristic of the Southern Red-backed Vole as described
by Hamilton and Whitaker (1979) and Godin (1977),
and appear not to apply to the Meadow Vole. Howey-
er, I note that Banfield (1974) states that the Southern
Red-backed Vole does not store up food for winter,
whereas he says the Meadow Vole does. Unfortunate-
ly, the subsequent British period recorders do not sup-
ply any additional information that would enable us to
settle the matter: what records there are, indicate the
animal was a largish “mouse” with a short tail (e.g.,
Patterson 17707; Shuttleworth 17937). Even so, the fur-
ther history of the outbreaks — the fact that they
THE CANADIAN FIELD-NATURALIST
Vol. 121
occurred only during the pioneer phase when new
clearings were being opened up in the forest, and
ceased as the forests were cleared — points also to the
Southern Red-backed Vole.
Two alternative theories were proposed for the cause
of the outbreaks by contemporary observers. Roma
(17507) and Patterson (17707) considered that win-
ters with heavy snow gave the voles protection from
their predators, at the same time allowing them to feed
from their stores under the snow; whereas Franquet
(17517), La Roque (175277), Stewart (18067) and
Plessis (181277) considered that the high production
of beech mast in certain years was the cause. It is
impossible retrospectively to determine which if either
of these explanations is correct.
Other mice and voles
The first mention of a mouse other than the “plague
mouse” is, Roma’s (1750+) reference above to the
souris [mouse] that he said was a better climber than
the plague vole — this can only be the Deer Mouse.
The next mention is over a century later: Sutherland
(18617) refers to two species of “field mice”: the “bur-
rowing field mouse”, which appears from his descrip-
tion to be the Meadow Vole, and the “leaping field
mouse’, which, given its “long tail and strong hind
legs”, is likely to be the Meadow Jumping Mouse
(Zapus hudsonius). Sutherland did not realize that
there was another species of jumping mouse on the
island, the Woodland Jumping Mouse (Napaeozapus
insignis), first recorded by Cameron (1958).
The only other recorder to distinguish between the
native species of mice and voles was Bain (18907) who
listed three native species: his “short-tailed Meadow
Mouse” can only be the Meadow Vole since his brief
description fits this species: “it lives on grain and grass-
es, and builds a nest of dry grass and makes long gal-
leries under the snow in winter when it causes much
destruction by barking young orchard trees”. His two
other species were “much less common”: his “White-
Footed Mouse” can only be the Deer Mouse, while his
‘Hamster Mouse” must be the Southern Red-backed
Vole. He thus did not record either species of jump-
ing mouse.
Snowshoe Hare
The Snowshoe Hare (Lepus americanus) is listed
by all but two of the list-makers (Table 1), with many
of the recorders noting that hares were abundant on
the island. The only recorders to give any information
on its habitat are Rowan (18767), who said “their
favourite resort is the second growth of young forest”,
and Bain (18907), who noted they liked “dry ground
in summer but frequeni the swamps in winter”. This
was presumably related to the food they fed on in
winter, which Bain (18907) said was “young shoots
of trees” — these were more specifically identified by
Rowan (18767) as “the tender twigs of maple, moose-
wood, birch, willow, alder etc.”. Both recorders said
2007
that in summer they fed on “grass”. Rowan said that
on the island their natural enemies were the Lynx,
Marten, Weasel and Fox, as well as birds of prey, and
man. Almost all of the recorders refer to the Snow-
shoe Hare in the context of a game animal, and in the
early years of settlement it seems to have been an im-
portant supplementary food source for the human pop-
ulation (e.g., Curtis 1775*+; Chappell 1775-18187*).
Hares continued to be eaten throughout the nine-
teenth century and were considered a delicacy by
many. However, Johnstone (1822+) and Sutherland
(1861%) considered their fur to be of little value.
Absences in the records
Moles
Although Johnstone (18227) stated categorically
that “there is not a mole on the island”, two later writ-
ers of school textbooks for the island (Sutherland
1861+; Bain 18907), included it in their lists of the
island’s mammals, both using similar phrasing: “it bur-
rows in the ground and lives on worms”. Despite their
assertions, there is no evidence that a species of mole
has ever occurred on Prince Edward Island, and nei-
ther Cameron (1958) nor ACCDC (2008*) have in-
cluded it in their lists of the native mammals.
Fisher
The Fisher (Martes pennanti), which currently
occurs in Nova Scotia and New Brunswick, is entirely
absent from the historical records for the island. Thus
the inclusion of Prince Edward Island in the animal’s
historical range by Cameron (1958), Peterson (1966),
Banfield (1974) and ACCDC (2008*) is not support-
ed by the early records.
Moose
There is no historical evidence that the Moose
(Alces alces) ever occurred on Prince Edward Island,
at least during the settlement period, and in fact there
is strong evidence to the contrary: during the French
period Denys (167277) and La Ronde (17217) had
made a point of recording that the Moose was absent
from the island, as did Sutherland (18617) during the
nineteenth century. The five British period records of
its occurrence are late and retrospective (Table 1), and
are all based on the anecdotal finding of “moose horns”
in the woods — it was this evidence that led Cameron
(1958) and Banfield (1974) to list the Moose as once
occurring on the island. These “horns”, however, are
far more likely to have belonged to the Caribou, for
which there is irrefutable evidence of its past occur-
rence.
Beaver
The only historical record for the occurrence of the
American Beaver (Castor canadensis) on Prince
Edward Island (that is, prior to its deliberate intro-
duction in the twentieth century) is that of Bain
(18907) who said that it was “once common here and
SOBEY: HISTORICAL RECORDS FOR NATIVE MAMMALS OF PEI 39]
the remains of its dams are still to be seen in many
parts of the country”. This is 4 curious statement since
fourteen years earlier Rowan (18767) had said that
he had not seen or heard of any “beaver works” on
the island, and so he was “inclined to think” they had
“never lived on the island”, as “their traces endure long
after they have become extinct’. So also had Suther-
land (18617) included it in a list of the mammals found
on the mainland that did not occur on the island. The
validity of Rowan’s and Sutherland’s observations is
strongly supported by the fact that its absence from
the island had been noted at the beginning of French
settlement by La Ronde (17217), as well as by the fact
that none of the other list-makers had included the
beaver in their lists of the island’s mammals, including
Stewart (18067), Johnstone (18227) and MacGregor
(1828+) who had made fairly comprehensive lists.
Given the commercial importance of its fur and its rel-
atively large size, it was a mammal that, had it been
present, would not have been overlooked. Apart from
Bain’s comment, the only other mentions of the beaver
that [ have come across in the historical records are its
inclusion in a list of the retail prices of pelts, printed in
a pamphlet for immigrants to the island (Anonymous
1808+7+), and the presence of beaver pelts among a
consignment of animal skins shipped to Halifax in
1802 (Sobey 2006). Since pelts were easily trans-
portable from the mainland, where the species did
occur, neither record indicates its presence on the is-
land. It should be noted that the beaver’s absence
from the island from 1720 onward does not exclude the
possibility that it had been trapped-out prior to the
eighteenth century. The evidence in support of such a
view, however, is limited to the finding of a beaver
incisor tooth in an archaeological dig at a prehistoric
aboriginal site (probably Mi’kmaq) near East Point
that was occupied from about 800 to 1000 AD (Keen-
lyside 1983) (though such a small item could have
been easily carried from the mainland), and the unref-
erenced statement of Cameron (1958) that beaver
tooth-marks had been found on sticks from island
peat-bogs.
Other absences
Sutherland (18617) noted the absence of the Rac-
coon (Procyon lotor) from the island, as well as the
American Porcupine (Erethizon dorsatum) (as had
La Ronde IN 17217), both of which he noted as occur-
ring in Nova Scotia and New Brunswick; while Rowan
(18767) noted the absence of the “Virginian deer” (i.e.,
the White-tailed Deer, Odocoileus virginianus). The
validity of these statements is supported by the fact
that none of these animals was noted as present by
any other of the historical recorders.
Discussion
The island’s native mammalian fauna
Thirteen of the island’s native terrestrial mammals
were recorded by French period observers (Table 1).
392
with British period recorders taking the total to twenty.
On the basis of the current ACCDC (2008*) mammal
list for the island, we may surmise that the number of
terrestrial species at the time of settlement might
have been 28. This includes my deletion of the
beaver and Fisher from the ACCDC list, and ignores
the post-settlement arrivals of the Brown Rat (Rattus
norvegicus), House Mouse (Mus domesticus), Striped
Skunk (Mephitis mephitis), Racoon and Coyote (Canis
latrans), the latter three in the twentieth century. The
eight omissions from the historical record are all small
mammals: the island’s five species of shrew (assum-
ing those currently in the ACCDC list were also pres-
ent in the eighteenth century) were treated by the early
recorders as one species, as were the four species of
bat; while one of the five species of native mice and
voles went unrecorded (the Woodland Jumping Mouse
Napaeozapus insignis, first recorded by Cameron
1958). As for the marine fauna, three species are dis-
tinguished in the historical records: the Walrus, and
seemingly, the Harbour Seal and Harp Seal. Thus the
two other seal species listed by ACCDC (2008*) as
currently occurring in the Gulf of St. Lawrence (the
Grey Seal, Halichoerus grypus and Hooded Seal,
Cystophora cristata) are not discernable in the his-
torical records for the island.
Despite the absence of three key boreal elements
from the records (the Moose, porcupine and beaver),
the terrestrial fauna recorded at the beginning of Euro-
pean settlement is a boreal fauna, signalled in the
records by the presence of the Snowshoe Hare, Red
Squirrel, Northern Flying Squirrel, Southern Red-
backed Vole, American Marten, Short-tailed Weasel,
Canada Lynx, and Woodland Caribou. All of the other
mammals recorded (e.g., the American Black Bear,
Grey Wolf, Red Fox, American Otter, Mink and
Muskrat) are also quite at home in the boreal forest
zone, though occurring widely in more southerly areas
as well (e.g., Banfield 1974; Hall 1981). Concomitant-
ly, there is a total absence of mammals characteristic
of the more southern deciduous forest zone (e.g., the
Striped Skunk, Racoon, Eastern Grey Squirrel, Sciurus
carolinensis, and White-tailed Deer); and if Denys
(167277) is to be relied upon, none of these were pres-
ent in the rest of Acadia (including eastern Maine) in
the seventeenth century — though in the last two cen-
turies such southern species have reached the adjacent
mainland and some have even been introduced onto
the island (i.e., Striped Skunk and Racoon).
The presence of the Woodland Caribou, along with
its predator, the Grey Wolf, is one of the more interest-
ing pieces of information to emerge from the records
of the period, and is a corrective to Lohr and Ballard’s
(1996) statement that wolves never occurred on Prince
Edward Island. As I have noted, it seems that neither
species was abundant, and La Ronde (17217) said that
the reason the Mi’kmaq did not spend the winter on
the island was that they found the Caribou difficult to
THE CANADIAN FIELD-NATURALIST
Vol. 121
kill (he actually called them cerfs or “deer’), the impli-
cation being that there was little else for them to eat
in winter — which would indeed have been so, given
the absence of the beaver and the Moose, two of their
traditional winter food animals (e.g., Denys 167277).
We thus have here a clear instance of the natural fauna
(both presences and absences) having important effects
on the aboriginal population.
The importance of the beech forests
The historical evidence indicates that American
Beech was a very important tree in the upland hard-
wood forests of the island, with large areas of the land
surface being covered by beech forest (Sobey 2006).
Beech produces a nut (traditionally called mast), which
provides an important food for many different ani-
mals (e.g., Tubbs and Houston 1990). In the histori-
cal records for the island we are told, as we have
seen, that the Red and Flying squirrels, the chipmunk
and the “plague mouse” fed on the mast, as also, we
may presume, did all of the other woodland mice and
voles. Evidence from elsewhere (e.g., Vass 1987;
Telfer 2004) indicates that the American Black Bear
was also likely to have done so.
The North American Beech has long been known to
be irregular in its seed production, with “mast years”
occurring every two to eight years (Tubbs and Houston
1990). That the beech forests on the island also exhib-
ited such mast years is recorded by Franquet (17517),
La Roque (175277), Stewart (18067), Plessis (181277)
and Bain (1868-188477). As we have seen, such mast
years were considered by some to be the cause of the
vole irruptions, while Stewart (18067) said that the
three squirrel species (and of these, especially the chip-
munk) also showed a “great increase” in numbers
“after an abundant crop of beech mast”. And though
they did not enter the written record, we may presume
that all of the other feeders on beech mast would also
have been affected. As well, the distribution of beech
forests on the island might have affected the spatial dis-
tribution of the mast-feeders, and this may not have
been only in mast years. However, the only evidence
that we have for this is Bain’s (1890+) comment that
the red squirrel was “so plentiful in every wood where
beech nuts are found”.
Given this recorded increase in the mast-eaters, we
might expect that the island’s predator populations
might have concentrated on them when they were
abundant — Roma (17507) had certainly noted that the
fox and the marten fed on the voles during a plague
year. We have a hint that this abundance of prey may
have also led to a increase in the numbers of the pred-
ator species, in Franquet’s (17517) statement that foxes,
martens, mink and lynx were more abundant in some
years than in others, as well as Stewart’s (18067) that
the marten was in “great abundance” in some years.
However, confounding factors here are the so-called
“Jong” or “nine to ten year cycle” in the numbers of
the Snowshoe Hare and its predators (especially the
2007
lynx) that has long been recorded throughout its
North American range and the “short” cycle (of two
to three years) that commonly occurs among rodents
and their predators in northern environments (e.g.,
Finerty 1980).
We can only speculate on the effect that the great
decline in the area of beech forest has had on the abil-
ity of the forests to sustain the food-webs that they did
before European settlement. Prior to forest clearance,
beech was a predominant tree, Stewart (18067%), for
example, saying that “one-half of the island is cov-
ered with it, in some districts it forms nine-tenths of
the forest”. By 1991, however, beech contributed only
4.1% of the woody biomass in the remaining upland
hardwood forest (Sobey and Glen 2002), and far less
when all of the forest-types are considered. Thus
those mammals still occurring on the island that were
once substantial mast-eaters (the chipmunk, squirrels,
voles and mice) have had to turn to other types of food.
The utilization of the mammalian fauna for food and
fur
All of the French period mentions of the Caribou
refer to it in the context of game, either for the abo-
riginal Mi’kmaq (Denys 1672++; La Ronde 17217) or
the new French settlers (Gotteville de Belile 172077;
Roma 17507). Then, after the Caribou’s extirpation
in the mid-eighteenth century, the animal most men-
tioned as a source of food is the Snowshoe Hare. As
well, other animals that most people nowadays would
not consider edible are also recorded as being eaten
by the European population, notably the bear and the
lynx, while Holland (17657) reported that the refugee
Acadians were eating even martens, otters and
Muskrats. The island’s seals also served as food for
the Acadian population in the early years (La Ronde
1721+; Roma 1750+; Patterson 17707), as did oil from
the walrus (Stewart 18067). However, once settlement
was established, the native fauna seems to have become
unimportant as a food source, though the hare contin-
ued to be mentioned as a game animal throughout the
nineteenth century.
Would the hunting of any of these animals for food
have had any effect on their numbers? It is probably
not a coincidence that the Caribou disappeared from
the island within a few decades of the establishment
of the first permanent settlement in 1720: the small
numbers reported to occur (e.g., Denys 1672+7+; Hol-
land 1765+) would have been vulnerable to even low
levels of hunting. Also, the hunting of the walrus for
its oil may have contributed to its disappearance from
island waters. By contrast, the Snowshoe Hare con-
tinues to thrive on the island, and any hunting, even
in the unregulated early days, seems to have had little
effect.
As for those animals that served as food only under
extreme conditions, it is clear that hunting was anoth-
er pressure that contributed to their extirpation, name-
ly, their being hunted for their furs. However, though
SoBEY: HISTORICAL RECORDS FOR NATIVE MAMMALS OF PEI 393
many recorders, as we have seen, stress the value of
the furs of the otter, marten, mink, Red Fox, and
black bear, and to a less extent the lynx and the
Short-tailed Weasel, concrete data on any fur trade
that developed on the island is sparse in the historical
record. For the French period, the only specific refer-
ences are Roma’s (17507) comment that “/a chasse
aux martres |1.e., marten-hunting}| should be reserved
to the Mi’kmagq to enable them to “procure their
needs”, and the entry in the 1757 journal of an officer
aboard a French warship at Louisbourg that “the Indi-
ans of [le Saint-Jean usually come to Louisbourg in
August and September to sell their animal pelts
which brings them a very good profit” (Sobey 2006).
In the British period, the only recorder to give the
fur trade more than a passing mention is Samuel Hill
(18397), who wrote that “there was once a consider-
able export of furs from the island”. The island's cus-
toms records contain evidence of two such early fur
shipments: the Betsy, bound for Halifax in June 1802,
had amongst its cargo “4 Trunks of furs, containing
Rabbit, Fox, Martin, & Beaver skins, a few seal skins
& 30 Dozen loose rabbit skins”; while in May 1819
the Nelly, also bound for Halifax, had 1200 “rabbit”
skins, 162 martin, 14 otter, | lynx, and | bear. (The
presence of beaver skins in the 1802 shipment indi-
cates that at least some of the furs had come from the
mainland.) Also, the inventory of the estate of one
John Rennie in 1791 recorded that at his death he was
in possession of “six dozen dressed marten skins”
(Nicholson 2004).
When Hill goes on to comment on the fur trade of
the 1830s he highlights the role of the Mi’kmagq;: “foxes
are chiefly shot or trapped by the Indians, and their
skins exported to England by the resident merchants”,
and other writers also refer to the skills of the Mi’k-
maq in the trapping of the native fauna (Johnstone
1822+; MacGregor 18287). And there is also an 1830s
record of the export of furs in the customs records:
the Catherine Jane bound for Liverpool in May 1834
carried 493 fox pelts (of which 19 were cross and 4
silver), 440 marten, 60 “rabbit”, 40 lynx, 31 otter, 29
mink, 2 Raccoon, and | Muskrat. As was so for the
beaver above, the presence of the two Raccoon pelts
indicates that some of the skins must have come from
the mainland, since the Raccoon is otherwise absent
from the historical records for the island.
Extirpations among the fauna
Four of the fur-bearers were eliminated from the
island: the marten, otter, lynx and black bear, as were
also the caribou and the wolf — as well as the walrus
from the waters around the island. We may question
whether all of these extirpations were inevitable.
Granted that the large-scale destruction of forest to
make way for farmland would have resulted in a con-
siderable loss of habitat and a resulting decline in the
total numbers of all of the terrestrial mammals: even
394
so, with the exception of the caribou and the wolf,
which are both likely to have required a greater area
of forested habitat than what remained, the other four
land mammals should have been able to survive in
the remaining areas of forest and other habitat. In the
British Isles the Pine Marten (Martes martes) and
European Otters (Lutra lutra) have been able to sur-
vive in landscapes greatly altered by human activity,
and in association with as high or even higher human
population densities than on Prince Edward Island
(e.g., Corbett and Southern 1977; Jefferies 1989). It
thus seems that had there not been the uncontrolled
trapping of the marten — an animal notoriously easy
to trap (e.g., Banfield 1974; Hamilton and Whitaker
1979) — it very likely could have survived. As for the
Otter, though habitat loss may have played a role, the
rivers and bays that were its habitat being directly adja-
cent to the areas first selected for settlement, unregu-
lated trapping appears also to have been the main rea-
son for its extirpation. In fact, it was not until 1879
that the island’s House of Assembly passed its first
act “for the protection of Game and Fur-bearing Ani-
mals”, which, among other measures, restricted the
season for taking Muskrat, marten, and otter to be-
tween | November and 1 May (Glen1995).
As for the lynx and the black bear, had there not
been active hostility towards them, including the insti-
tution of bounties, it is possible that they could also
have survived. According to Anonymous (2003*), “as
long as they are not disturbed”, lynx are “remarkably
tolerant of human settlement” (see also Poole
(2003)), while the last bear on the island was shot as
late as the 1920s — some forty years after the peak of
human population density. As for the Walrus, though
the island’s first British governor was aware of the
need for controls on the “sea cow fishery” and even
had his council pass an ordinance in 1770 to regulate
it, he was aware that the hunt, being largely carried
out by “vessels from New England”, was beyond the
control of his infant government (Patterson 17747).
We may thus conclude that though the activities of
the settler population led to the seven extirpations, an
important factor contributing to four of them was the
attitude of the settlers: a positive hostility to the lynx
and the bear (understandable in the light of their pre-
dation on livestock), and an indifference to the sur-
vival of the otter and the marten so that the necessary
protections were not instituted until it was too late.
Acknowledgments
I thank Professor Gerry McKenna, then Dean of
Science at the University of Ulster, for enabling me
to take six months sabbatical in 1996 so that I could
initiate the study, and Harry Baglole of the Institute
of Island Studies at the University of Prince Edward
Island for supporting a research associateship at the
university. I also thank the staff responsible for the
THE CANADIAN FIELD-NATURALIST
Vol. 121
P.E.I. Collection at the Library of the University of
P.E.I. and those at the PEI. Public Archives and
Record Office for assisting me with locating and copy-
ing documents, as well as the PE.I. Department of
Environment, Energy and Forestry, and in particular
William Glen and Kate MacQuarrie, for supporting
the research. I also thank William Glen for directing
me to some of the fur data in the customs records.
Documents Cited (marked * in text)
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Sackville, New Brunswick. At website: www.accdc.com
Anonymous. 2003. The Canada Lynx. Hinterland Who’s
Who. Canadian Wildlife Service, Environment Canada.
At website: www.hww.ca.
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sion 2702, Series 20, 296-317.
[Lawson, Stephen G.] 1877-1878. [A series of articles on
pioneer days.] The Presbyterian and Evangelical Protes-
tant Union, Charlottetown. February 1877 to March 1878.
[The relevant extracts are reprinted in Sobey (2006).|
Bain, Francis. 1890. The Natural History of Prince Edward
Island. G. H. Hazard, Charlottetown, P.E.I.
Additional Historical Documents Cited (listed
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Denys, Nicolas. 1672. Description geographique et his-
torique des costes de |’Amerique Septentrionale, avec
Histoire naturelle du Pais. [Re-published 1908 by the
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by W. F. Ganong — this was reprinted in 1968 as a facsimi-
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Gotteville de Belile, Robert-David de. 1720. Letter to the
Council of the Marine. PAC, AC, C''A, Volume 43, fols.
134-36.
La Roque, Joseph de. 1752. Voyage d’inspection du Sieur
de la Roque, Recensement 1752. [Published 1906 in Rap-
port concernant les Archives canadiennes pour l’année
1905, Volume II, pages 1-168. Imprimerie du Roi, Ottawa.
English translation in Report concerning Canadian
Archives for the year 1905, Volume I, Appendix A, Part
1, pages 1-172].
Curtis, Thomas. 1775. A Narative of the Voyage of Tho*.
Curtis to the Island of St. John’s in the Gulf of St. Law-
rence in North America, in the year 1775. Published
1955 in Journeys to the Island of St. John or Prince Edward
Island. Edited by D. C. Harvey. MacMillan, Toronto, pages
9-69]
396
Chappell, Benjamin. 1775-1818. Daybooks. P.E.I. PARO:
2227. [Published 2003 by the P.E.I. Genealogical Society,
Charlottetown, as: The Daybooks of Benjamin Chappell
(Edited by L. Chappell)].
[Ritter, Ensign]. 1780. The regimental log of the von Kny-
phausen Fusilier Regiment (8 September 1779 — 30 June
1780). [An English translation by L. Zimmermann of the
original German was printed 1987 in Abegweit Review
5: 20-28].
Anonymous. 1808. A True Guide to Prince Edward Island,
formerly Saint John’s in the Gulph of St. Lawrence, North
America. G. F. Harris, Liverpool.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Plessis, Joseph-Octave. 1812. Le journal des visites pasto-
relles de Mgr. J.-O. Plessis (Evéque de Québec) en Acadie
1811, 1812, 1815. [Published 1980 in Les cahiers de la
société historique acadienne 11: 1, 2, 3.]
Bain, Francis. 1868-1884. Notes in Natural History, 1868-
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MacLeod, D. 1876. Memoir of Norman MacLeod, D.D. by
his Brother. Belford Brothers, Toronto.
Received 13 July 2007
Accepted 25 July 2008
Social and Play Behavior in a Wild Eastern Coyote, Canis latrans, Pack
JONATHAN G. Way!”
'Science Department, Barnstable High School, 744 West Main Street, Hyannis Massachusetts 02601 USA; e-mail: jon@ eastern
coyoteresearch.com
*Mailing address: Eastern Coyote Research, 89 Ebenezer Rd, Osterville Massachusetts 02655 USA
Way, Jonathan G. 2007. Social and play behavior in a wild eastern Coyote, Canis latrans, pack. Canadian Field-Naturalist
121(4): 397-401.
I had close and consistent observations of a wild eastern Coyote pack (Canis latrans) from January 2000 to August 2007.
During this time, I obtained 3156 radio-locations on a specific radio-collared breeding male (“Sill”) and observed him and/or
members of his pack on 375 occasions. The average group size = 3.0 + 2.3 (SD) Coyotes with 1.9 + 1.2 (SD) being adults
and 1.1 + 1.9 being pups. Maximal group size involved 12 Coyotes (9 pups, 3 adults). During these observations, Coyotes
most often behaved in a friendly manner toward each other as indicated by 80 of my observations involving play between
pups, and 15 involving play among adult Coyotes. On the evening of 6 July 2007 I observed the breeding male (>8 yr old),
his mate (>5 yr old), one of their full-sized probable yearlings, and five pups playing intensely for 33 minutes. This paper
details social and play behavior from this pack, especially from the 6 July 2007 observation.
Key Words: Canis latrans, eastern Coyote, amiable/affiliative interactions, play behavior, Massachusetts.
Although not exclusively a mammalian trait, play
is an important social activity for mammals, and espe-
cially for carnivores (Bekoff 1972a, b, 1989; Ewer
1973; Rennicke 2007). It is characterized by (1) friend-
ly, non-aggressive (or amiable) intentions; (2) more
energy than is strictly necessary; and (3) a typical air
of eagerness, excitement, and enjoyment during inter-
actions (Ewer 1973; Bekoff 1974a, b, 1989). Although
play is not unknown in adults, it is displayed more com-
monly in young animals (Bekoff 1978), and provides
the means of doing all the things the central nervous
_ system is prepared for without any of the risks inher-
ent in a “real” (e.g., hunting, fighting) situation (Ewer
1973). Hypotheses about the function of play in canids
_have been reviewed by Bekoff (1974a) and Packard
(2003) and include (1) the ability to learn counter-tac-
tics during play that may provide a basis for learning
| complex social relationships later in life; (2) physical
exercise related to aerobic conditioning; (3) develop-
_ ment of muscular routines; (4) practice of instincts
useful for hunting later in life; (5) a demonstration that
“animals that play together, tend to stay together” i-e..,
social cohesion; and (6) an indication that it may sim-
1 ply be fun and a pleasurable experience.
In the vast majority of studies on Coyotes (Canis
latrans) (and on carnivores in general), play behavior
has occurred with juveniles in captivity where
researchers can easily observe their study subjects
fend quantify behavioral observations in a controlled
) setting (Bekoff 1972a, 1974b, 1978, 1989; Way et al.
2000), Packard (2003) noted that our understanding
of Wolf (Canis lupus) social behavior will advance
| more rapidly when we integrate the information from
both captive and field populations, as long as we take
great care to recognize the limitations of each per-
Spective and to understand the whole as the sum of the
’
3
parts. However, few studies have examined the play of
Coyotes and Wolves in the wild (Ortega 1988; Mech
1997; Packard 2003). Of these studies, all detailed play
behavior was observed only in pups.
For instance, Mech (1997: 75) watched wild Wolf
pups in the Arctic play “for about 45 minutes as the
pups scrambled back and forth across the snowfield,
chasing one another, tackling, sliding, rolling, skidding,
and carrying on to a degree I have never seen nor heard
of before for any species.” Observations of pups play-
ing is acommon theme in studies on play behavior in
canids, and despite some authors claiming that adult
canids play (e.g., Bekoff 1989; Lehner 1978; Packard
2003), there is scant evidence of this in the wild, pos-
sibly due to their need for conservation of energy,
certainly in harsh climates. In this paper I describe a
social, amiable, wild Coyote pack, including observa-
tions of the older adults in the pack playing.
Methods
Research was conducted in an urbanized area (town
of Barnstable on Cape Cod) in eastern Massachusetts
(see Way et al. 2001, 2002a, 2004). Average human
density on the study site was 290 people/km? and road
density was 4.66 km of roadway per km? (Cape Cod
Commission 1998*). Observations began on 18 Janu-
ary 2000 when the Coyote “Sill” (ID # 0003), a 16.8 kg
9-month old male Coyote was captured in a box trap
(model 610B, 152.4 cm x 50.8 cm x 66.0 cm, Toma-
hawk Live Trap Co., Tomahawk, Wisconsin, USA:
Way et al. 2002b) in Cotuit (within the town of Barn-
stable), Massachusetts, and radio-collared as part of
an ongoing study of eastern Coyote ecology on Cape
Cod (Way et al. 2002a, 2004). Sill was tracked on his
natal range (his father, “Kett.” ID # 9805, was also
radio-collared) until January 2001 when he dispersed
397
398
and settled into a vacant territory to the immediate east
of his natal range in February/March 2001 (see Way
2007* for more details). He was the dominant male of
that new pack (Newtown/Bog Pack) in Marstons Mills
(within the town of Barnstable) from 2002 to 2007,
when the study ended.
Tracking protocols were fully described by Way et
al. (2002a, 2004). Portable receivers (Custom Elec-
tronics, Urbana, Illinois, USA) and hand-held 3-ele-
ment Yagi antennas were used to radio-track Coyotes
both on foot and from a vehicle. Animals were typi-
cally homed in on using the loudest-signal method of
radio-tracking (Springer 1979). I approached radio-
collared Coyotes as closely as possible without disturb-
ing them. I used binoculars, 15-45 x spotting scopes
(Bushnell, Overland Park, Kansas, USA), and video-
cameras when observing Coyotes at den and rendez-
vous sites (Way et al. 2001, 2003), and spotlighting
and headlights when following Coyotes at night with
a vehicle (Way et al. 2002a, 2004).
Collared Coyotes were often seen with untagged
companion(s), especially when at rendezvous sites
(Way 2003). A detailed description (e.g., size, col-
oration, distinguishing markings, and behavior) of the
uncollared animals was made during every direct obser-
vation. In this manner, the unmarked Coyotes were
identified based on appearance, as described by Way
et al. (2002a) and Way (2004). Overall, I identified as
many Coyotes as possible from Sill’s pack, as well as
from other groups within the study area. I classified
Coyotes as adults or pups/juveniles. Adults were clas-
sified as all full-sized Coyotes and often included year-
lings that remained on their natal territory. Pups were
born in April (Way et al. 2001) and were classified as
such until October, when they approached full body
size and became indistinguishable from adults when
observed in the field. Due to the logistic difficulties
of keeping track of large groups (25) of Coyotes
observed at the same time (especially when they tem-
porarily separated and were all visible at the same
time but only one observer was there to monitor them
in separate locations), it was necessary to began with
this simple coding scheme (pups or adults).
Most of the lengthy sightings were made at two sets
of cranberry bog complexes (2-3 bogs at each location;
complexes were | km apart) where distant (~400 m)
viewing was possible (Way 2003, 2007*). Sill’s pack
began using these bogs in summer 2003 following the
death of an old female Coyote (“Mole” #0110) in the
pack to their immediate northeast (Way and Strauss
2004; Way 2007*). To avoid autocorrelation (see Way
et al. 2004), sightings were separated by > 4 h from
each other. Typically this meant that Coyotes were
observed during dawn and/or dusk on a given day
with occasional daytime observations (n ~25).
Play was defined as the behavior that was performed
during social interactions in which there was a decrease
in social distance between the interactants, and no
THE CANADIAN FIELD-NATURALIST
Vol. 121
evidence of social investigation or of agonistic or pas-
sive-submissive behaviors on the part of the members
playing, although these actions may occur as derived
acts during play (e.g., passive submission during a play
bout — Bekoff 1972a, 1974a; Way et al. 2006). Bekoff
(1974a) classified play as (1) incorporating various
contexts into unpredictable temporal sequences; (2)
preceded by a metacommunicative signal (e.g., play
initiation); (3) repeated and performed actions in an
exaggerated manner; and (4) appearance of being
pleasurable to the participants, e.g. a play face (wide
open jaws and eyes) is apparent. Included in play were
numerous role reversals where more dominant Coy-
otes allowed lower ranked Coyotes to pin them to the
ground; this was never documented during agonistic
displays (Way et al. 2006).
I used continuous animal sampling (Martin and Bate-
son 1986) to record Coyote activity and recorded the
behavior (via scan sampling) of as many Coyotes con-
currently as I could during my observations. I record-
ed data into a field notebook and video-camera for
later analysis.
Results
I obtained 3156 radio-locations on Sill and observed
him and/or members of his pack 375 times (>4 hours
apart) from January 2000 to August 2007. The aver-
age Coyote group size during the 375 sightings was
3.0 + 2.3 (SD) Coyotes with 1.9 + 1.2 being adults
and 1.1 + 1.9 being pups. Maximal group size ob-
served during an observation involved 12 Coyotes
(9 pups, 3 adults), while two sightings involved 10
Coyotes (3 adults and 7 pups) and eight sightings in-
volved 9 Coyotes (ranging from 2-7 adults and 2-7
pups). This pack was typical of other eastern Coyote
social groups (see Way et al. 2002a, 2003) in that Sill
had a mate (“Mange-back”, positively identified
from 2004/2005 to 2007, having a distinct missing
patch of fur on the top of her neck and a large soft-
ball-sized bleached-like looking mark on her left rib
which looked like a distinct blonde splotch on an
otherwise brown body) and those two comprised the
breeding pair while the remaining adults appeared to
be yearlings, or pack associates, most likely their off-
spring from the previous year. On any given year there
were 3 to 5 adults (including associates/yearlings)
and four or more pups. Sill’s group was especially
visible for the four and a half years from 2003 to 2007
when they adjusted their territory to include two com-
plexes of several cranberry bogs each, formerly used
by a pack to their northeast. Although many of the
sightings were temporary (e.g., one or two Coyotes
observed for 1-2 sec while crossing a road at night),
160 (42.7%) observations occurred at rendezvous
sites averaging 44.5 + 31.4 (SD) min (maximum bout
= 142 min) in duration. These rendezvous site obser-
vations produced all of the observations of play
detailed below.
2007
Coyote pups often played with each other during
observation bouts (n = 80) at the rendezvous sites,
which consisted of chasing, wrestling, and self-play
(Way et al. 2006). It was very apparent when they were
playing because role reversals often occurred where
small pups pinned larger, more dominant ones (Ren-
nicke 2007). Also, chasing often involved different
combinations of chaser(s) and chasee(s) where close
observations indicated open (sometimes panting), re-
laxed mouths. In all observations, colleagues, some of
my students, or the general public (1 usually observed
from a main road and people frequently stopped to
observe with me) agreed with my assessment of play
rather than another behavior (e.g., agonistic). On 15
occasions, I observed adult Coyotes play with one
another. These were frequently games such as group-
chases or wrestling matches where pups also became
involved. The only negative interactions that I observed
between adults was when one probable yearling pinned
another (7 = 9 observations). I did not observe any ag-
gressive acts directed toward Sill or his mate “Mange-
back” but on four occasions Mange-back aggressively
pinned another (probable female) Coyote into a passive
submission posture. Based on subsequent observations,
these Coyotes were likely her yearling daughters.
Sill was a very diligent father and often returned to
the rendezvous sites throughout the year. Many of my
observations consisted of him and 1-2 others (usually
adults) returning from patrolling their territory at night
to joining other Coyotes waiting at these sites (i.e.,
full-sized pups). During the summers of 2005 to 2007,
when I positively identified Mange-back as his mate,
I more often saw Sill (80% of sightings) at the rendez-
vous sites (July-October) with the pups than Mange-
back (~60% of sightings involved her). All Coyotes
acted submissive to Sill except for Mange-back, who
rarely interacted with him, likely because of their
familiarity with one another. Yearlings (n = 20 sight-
ings) and pups (n = 75) often greeted Sill in an active
submission posture which involved repeated licking
of his mouth with exaggerated tail wagging by the
intended recipient (Way et al. 2006). Sill frequently
regurgitated (n = 10-12 observations) to pups when
solicited, especially when returning to the rendezvous
sites from surrounding areas. In fact, the only nega-
tive (agonistic) interactions I observed was when Sill
growled at persistent pups (n = 10-12 observations)
who repeatedly tried to lick his mouth to try stimulat-
ing him to regurgitate. All of the factors detailed
herein indicated that Sill was the breeding male of a
large, very successful, and amiable eastern Coyote
pack.
The evening of 6 July 2007 produced the most mem-
orable observations of amiable and play behavior from
this pack. For 33 minutes I observed three adults (out
of four in the pack at the time) and five pups from the
pack (one of which was later hit and killed by a car on
20 July), including Sill (8 years, 3 months old at the
time), Mange-back (likely >5 years old), and a yearling
Way: SOCIAL AND PLAY BEHAVIOR IN A COYOTE PACK
399
male named “White-band” (for a white band running
laterally on his tail just posterior to the base) that was
the same size as Sill. They were at the northern-most
area of the two rendezvous sites (north of the site from
Way 2003). At 20:15, I arrived at the rendezvous site
and immediately saw eight Coyotes about 250 m NW
of my location on Bog Road in Marstons Mills. They
were at the junction between the northwest part of the
east bog and northeast part of the middle bog (three
bogs comprised this area west of Bog Road) in an open
sand flat (about 25 x 20 m) dubbed “Olivia's” (O) flats
for a homeowner's walking trail that leads southeast
to this location ~75 m from her house.
When | arrived, I immediately set up my tripod and
spotting scope as the pups were particulary active
and running back and forth playing, both play chas-
ing and play wrestling (see Way et al. 2006). At
20:20, Sill walked west about 50-75 m and bedded in
sit] alert position (Way et al. 2006) on a small (1.5 m
high) mound of sand, next to a small shed/pumphouse
just north of the northcentral part of the central bog.
He sat alert and watched the other seven Coyotes from
a distance until 20:25, when Mange-back led the five
pups to the sand mound where Sill was sitting. To get
the pups there, she turned around and faced the pups
and took a few steps toward them until they followed
her. I was too far away to hear if she called them over
or if they just followed her, but she did that two or
three times before completing the short trip. White-
band followed the group west to the sand mound.
After trotting to that location (~15-20 sec.) the pups
immediately re-engaged in intense bouts of play chas-
ing and play wrestling. Sill greeted a couple of the
pups by licking them but mostly stood up when the
group arrived and then proceeded to watch them. At
ca. 20:30, Mange-back, which was watching the pups
play for ca. 2-3 min, got into the games by pinning a
couple of pups. She had a noticeably relaxed play face
(Bekoff 1974a, 1974b; Lehner 1978) and jumped back
and forth as a couple of pups chased after her. Sill and
White-band watched the actions right on the mound
and despite pups bumping into Sill, he maintained his
vigilance by surveying the area for danger.
At 20:32 Sill became active. He gathered a couple of
pups, then trotted east back to the O flats. Mange-back
and White-band followed him. Immediately upon ap-
proaching the O flats, he did a play bow (Way et al.
2006) to initiate play from the two pups. The pups
immediately responded by chasing him and he, like
Mange-back on the sand mound, stayed contained
within a localized area (the O flats) instead of run-
ning from the pups as he sometimes did to get away
from them. In other words, it was obvious during this
particular observation that he was purposefully play-
ing with the pups.
For the next 10 min, Sill engaged in vigorous bouts
of play chasing with groups of two to three pups at a
time, after the other three pups returned to the area as
well. The action occurred quickly, but I believe that
400
he played with all five pups by the end of the session as
pups were constantly changing directions and con-
specifics with which they interacted. I could easily see
Sill’s tail wagging and his mouth open and relaxed,
similar to the pups when they played. By 20:40 it was
nearly dark and becoming difficult for me to see, but
at this point all of the Coyotes gradually diminished
their activity except for White-band which did not
engage in play on that particular night. He often baby-
sat the pups during summer 2007 and may have felt
the need to watch them instead (although I did observe
him = 4 times playing with the pups that summer).
At 20:42 Sill trotted at a steady pace southwest on a
canal separating the east and central bog, leaving the
pups at the O flats area. When he got to the south edge
of the canal (~200 m) he approached the north edge of
a private property that has a ca. 40 kg Labrador/
Golden retriever Domestic Dog mix (“Kota”) with
which he commonly interacted (by growling and bark-
ing at each other, usually from opposite sides of the
bogs) south of the southeastern part of the central bog.
When Sill approached the edge of Kota’s yard (Kota
was not outside) he sniffed then raised-leg urinated on
a bush followed by a ground-scratch, common behav-
ior I observed from him when marking his territory.
He then proceeded to trot west at the south part of the
bog complex, displaying a noticeably different and
more serious disposition than when he was playing
with the pups. He kept his tail at a 45° angle and was
very alert and tense on his route west. At 20:44,
Mange-back and White-band left the pups and trotted
through the central bog in the general (southwest)
direction of Sill, but taking a more northerly counter-
clockwise route, likely to avoid Kota. All three adults
gradually made their way to the south part of the west
bog west of Bog Road and eventually became too
difficult to see at 20:48. Sill’s signal (at 20:58) indi-
cated that he continued south, crossed a secondary road
(River Road), and went to the south of the two cran-
berry bog rendezvous sites (detailed in Way 2003).
Meanwhile, the pups remained around the O flats,
eventually heading northeast into the wooded patch
where they commonly rested and hid. I departed the
area at 20:50 at dark.
Discussion
This Coyote social group behaved in a remarkably
friendly manner toward one another and similar to what
has been described for Wolves, other than that they
preyed on small animals (rodents, rabbits), trash and
other human food, and domestic cats (J. Way, unpub-
lished data) rather than large ungulates (see Mech et
al. 1998: Mech and Boitani 2003). In fact, it was obvi-
ous to me that the Coyotes, including adults, were hav-
ing fun when they played, as demonstrated by such
characteristics as a play face, tail-wagging, and/or
engaging in affiliative activities which have been pur-
ported as nothing but play (Bekoff 1972a, 1974b;
THE CANADIAN FIELD-NATURALIST
Vol. 121
Packard 2003; Way et al. 2006). Such canid play char-
acteristics are obvious and recognizable to dog owners
when their pet dogs are playing.
Bekoff (1999) claimed that social play may pro-
vide more promising evidence of animal minds than
research in many other areas, for it may yield clues
about the ability of animals to understand one anoth-
er’s intentions. While watching Sill, it was difficult
for the pups (and for me as an observer) to misinter-
pret Sill’s intentions from play bowing to initiate play,
to a more serious (tail at 45° angle) posture when he
departed for his nightly travels (when none of the
pups attempted to follow him). It is highly likely that
the success of this social unit was predicated on the
stability of the pack members, mainly Sill and Mange-
back. By being mature older adults and extremely
familiar with the area (see Way 2007*), these two Coy-
otes were never observed getting challenged by their
younger, less experienced offspring.
This particular area (i.e., the open cranberry bog
complexes) has produced some memorable Coyote
sightings, including observations of Coyote pups for
11 years in a row (1997-2007: see Way 2003, 2007*).
However, I do not believe this group to be the excep-
tion to the norm. Other packs in my study area, al-
though less visible on the average, have similar social
units and were occasionally observed in comparable
amiable scenarios where adults were seen playing with
one another (e.g., see Way 2001* and Way 2007*).
Many of these observations of friendliness/play includ-
ed older adults (usually radio-collared) which, like Sill,
probably provided stability and experience to their
pack. The eastern Coyote’s family-oriented nature can-
not be understated. In one instance, I have even
observed two adults in a Coyote pack raising pups
when an older, breeding female was killed when the
pups were young (Way 2004).
Due to its success, the Coyote is typically maligned
by the public and is often treated as vermin or pests by
state wildlife agencies and/or animal control/pest agen-
cies (Way 2007*). However, Coyotes are highly social,
family-oriented, caring animals (Camenzind 1978;
Bekoff and Wells 1980; Gese et al. 1996; Crabtree
and Sheldon 1999;) that probably even enjoy life and
each other (Way 2007*; this study). Once thought to
be too subjective, there is a growing body of literature
that supports this statement (e.g., see Bekoff 2002,
2007), and data from this paper regarding this one pack
studied over eight years (2000 to 2007), provides
irrefutable corroboration of Coyote sociality, includ-
ing amiable/affiliative behaviors such as play.
Acknowledgments
I thank my family (especially my wife Tara), Marie
Thomas, Eric Strauss and Boston College, and Barn-
stable High School for helping me with various aspects
of my research. Two anonymous reviewers provided
helpful comments.
2007
Documents Cited (marked * in text)
Cape Cod Commission. 1998. Cape trends: demographic
and economic characteristics and trends. Cape Cod Com-
mission, Barnstable, Massachusetts.
Way, J. G. 2001. The eastern coyote: documenting the habits
of one of Cape Cod’s newest residents. Conservation Per-
spectives, the online journal of the New England Chapter
of the Society for Conservation Biology. http://www.nescb.
org/epublications/spring2001/coyotes.html.
Way, J. G. 2007. Suburban howls: Tracking the eastern coy-
ote in urban Massachusetts. Dog Ear Publishing, Indi-
anapolis, Indiana. http://www.easterncoyoteresearch.com.
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Received 28 August 2007
Accepted 25 July 2008
Biologie et situation de la Lamproie du Nord, /chthyomyzon fossor, au
Québec |
CHRISTIAN FortINn!, MARTIN OUELLET’, ISABELLE CARTIER!, DANIEL BANVILLE?, et CLAUDE B. RENAUD“)
'FORAMEC, 70, rue Saint-Paul, Québec, Québec G1K 3V9 Canada; e-mail : christian.fortin@foramec.qe.ca
*Amphibia-Nature, 1902, boulevard de Grande-Gréve, Gaspé, Québec G4X 6L6 Canada; e-mail: mouellet@ amphibia-nature.org:
*Ministére des Ressources naturelles et de la Faune, Direction de I’ expertise sur la faune et ses habitats, 880, chemin Sainte-Foy,
Québec, Québec G1S 4X4 Canada; e-mail: daniel.banville@mrnf.gouv.qc.ca |
+Musée canadien de la nature, Service de recherche, C.P. 3443, succursale D, Ottawa, Ontario K1P 6P4 Canada; e-mail: cre-/
naud @mus-nature.ca
|
'
Fortin, Christian, Martin Ouellet, Isabelle Cartier, Daniel Banville, and Claude B. Renaud. 2007. Biologie et situation de lz
Lamproie du Nord, /chthyomyzon fossor, au Québec. Canadian Field-Naturalist 121(4) : 402-411.
La Lamproie du Nord est une espéce propre a 1’ Amérique du Nord et elle ne se rencontre, au Québec, que dans I’extréme suc |
de la province. Cette lamproie a été observée dans le fleuve Saint-Laurent ainsi que dans 11 de ses tributaires. Elle aurait dis--
paru de la riviere Yamaska ot elle avait été recensée entre 1946 et 1959. Poisson non parasite, sa taille au stade adulte nes
dépasse habituellement pas 160 mm. Les larves, appelées ammocétes, et les adultes ne se retrouvent qu’en eau douce. Les!
cours d’eau utilisés sont généralement des ruisseaux et des riviéres a fond graveleux ou sablonneux. La Lamproie du Nore)
requiert deux habitats distincts, soit un pour les adultes reproducteurs et un autre, le long du méme cours d’eau, pour le
développement des ammoceétes. La détérioration de l’habitat et la pollution des eaux représentent les principales menaces 2
la survie des lamproies. La faible fécondité et la mobilité réduite de la Lamproie du Nord font en sorte que cette espéce est
peu adaptable aux modifications de son environnement.
Mots-clés : Lamproie du Nord, /chthyomyzon fossor, Petromyzontidae, rapport de situation, Québec.
The Northern Brook Lamprey is endemic to North America. In Quebec, it is restricted to the southernmost part of the province. |
This lamprey is known from the St. Lawrence River as well as in 11 of its tributaries. It appears to have been extirpated from
the Yamaska River where it was collected between 1946 and 1959. It is a nonparasitic fish and its adult size does not usually
exceed 160 mm. Larvae, called ammocoetes, and adults are found only in fresh water. Watercourses where it occurs are gen-
erally streams and rivers with gravel or sandy bottoms. The Northern Brook Lamprey requires two types of habitats during
its life cycle, one for spawning adults and one, along the same watercourse, for the developing ammocoete. Habitat degrada-
tion and water pollution are the major threats to the survival of lampreys. The low fecundity and low dispersal abilities of the
Northern Brook Lamprey make this species poorly adapted to withstand changes in its environment.
Key Words: Northern Brook Lamprey, Ichthyomyzon fossor, Petromyzontidae, status report, Québec.
Le genre Ichthyomyzon comprend six especes qui
se regroupent en trois paires, chacune étant composée
a été identifiée au Québec comme espéce susceptible
d’étre désignée « menacée ou vulnérable ». Au Cana-
d’une espéce parasite (espéce souche) et d’une espéce
non parasite (espeéce satellite; Vladykov et Kott
1979a). La Lamproie du Nord (Ichthyomyzon fossor),
une espece non parasite, serait issue de la Lamproie
argentée (Ichthyomyzon unicuspis), une espéce para-
site (Vladykov et Kott 1979a). Toutefois, certaines
analyses génétiques récentes ont remis en question le
fait que la Lamproie du Nord soit vraiment distincte
de la Lamproie argentée (COSEPAC 2007*). La Lam-
proie du Nord est un petit poisson d’eau douce. Au
stade larvaire, les larves, que l’on désigne sous le terme
d’ammocétes, vivent enfouies dans les sédiments et se
nourrissent par filtration. Au stade adulte, une fois la
métamorphose terminée, l’espéce ne conserve pas de
tractus intestinal fonctionnel et ne peut ainsi se nourrir.
Au cours de sa vie, elle se reproduit seulement une fois
et meurt peu de temps apres la fraie.
L’aire de répartition de la Lamproie du Nord est
limitée a l’Amérique du Nord. Au Canada, on la
retrouve au Manitoba, en Ontario et au Québec. Elle
da, la Lamproie du Nord n’est pas encore officielle-
ment désignée, mais elle possede le statut d’espéce
« préoccupante » attribué par le Comité sur la situation
des espéces en péril au Canada (COSEPAC). Un rap-)
port de situation, commandé par cet organisme, a été:
réalisé en 1991 (Lanteigne 1991*) et une mise a jour!
a été effectuée en 2007 (COSEPAC 2007*). Le présenti
article constitue une mise a jour du rapport de situa-:
tion officiel soumis au ministere des Ressources’
naturelles et de la Faune du Québec (Fortin et al.l
2005’).
Description
La taxinomie des lamproies holarctiques au stadel
adulte se fonde principalement sur quatre caracteres,
soit la disposition des dents, le nombre de myomeres
du tronc, les proportions du corps et la répartition géo-
graphique de l’espéce (Vladykov et Kott 1979b*). La
longueur totale de la Lamproie du Nord adulte ne
dépasse habituellement pas 160 mm (94 a 146 mm:
402
403
2007 FORTIN ET AL. : BIOLOGIE DE LA LAMPROIE DU NORD AU QUEBE¢
A Site d'observation d'/chthyomyzon fossor
QUEBEC
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&
ORE
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e
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FiGuRE |. Répartition québécoise de la Lamproie du Nord, /chthyomyzon fossor.
Hubbs et Trautman 1937; 128 @ 150 mm: Vladykov
1952; 86 a 166 mm: Morman 1979*; 98 @ 158 mm:
Lanteigne 1981; 130 a 173 mm: Trautman 1981). Son
corps est de forme cylindrique et sa téte est courte et
dépourvue de machoire. Elle possede une bouche
ronde en forme de ventouse dont Il’ ouverture est tou-
jours plus courte que la longueur de la région bran-
chiale. Elle est pourvue de sept paires de branchies.
Les yeux sont petits et situés pres du haut de la téte.
Le nombre de myoméres du tronc varie généralement
entre 50 et 53 (Lanteigne 1988).
Les dents de l’entonnoir buccal sont peu dévelop-
pées. La lamina supraorale consiste en une dent
pourvue de deux pointes alors que la lamina linguale
transversale posséde entre six et onze petites pointes
sur laréte denticulée (Scott et Crossman 1974). Les
dents latérales internes consistent quant a elles en
quatre dents unicuspides situées sur chaque coté du
disque. Des dents latérales externes et de petites
pointes situées dans les champs antérieur et postérieur
completent la dentition. La peau est lisse et ne possede
pas d’écailles. L’adulte est bicolore : coloration gris
foncé a brune sur le dos et les flancs et grise a blanc
argenté sur les parties inférieures (Vladykov 1949*).
La surface ventrale a une teinte orangée, particuliére-
ment chez les femelles sexuellement matures (Leach
1940). Le repli anal est plus visible chez la femelle
sexuellement mature que chez le male et est fusionné
a la nageoire caudale (Scott et Crossman 1974). La
nageoire dorsale est continue, ce qui la différencie de
la Lamproie de |’ Est (Lampetra appendix).
Répartition
La Lamproie du Nord est une espéce propre a
l’ Amérique du Nord. Au Canada, elle se retrouve dans
le bassin hydrographique de la baie d’Hudson, au
Manitoba, et dans le bassin hydrographique des Grands
Lacs et du fleuve Saint-Laurent, en Ontario et au
Québec (Lanteigne 1992). Aux Etats-Unis, elle se ren-
contre dans le bassin du Mississippi depuis le Wis-
consin jusqu a l’Ohio et vers le sud jusqu’au Kentucky
(Lanteigne 1992). Il existe aussi une population dis-
jointe au Missouri.
La connaissance de la répartition de la Lamproie
du Nord au Québec est trés restreinte. Elle se limite a
des mentions rapportées dans le cadre de travaux de
recherches et dinventaires multispécifiques. Cette
espéce n’a jamais fait l'objet d’°un programme d’in-
ventaire systématique et ciblé au Québec, a l’excep-
tion des travaux de Vladykov (1952) dans les années
1940 et 1950. Le manque de données sur sa réparti-
tion réside, entre autres, dans la nature discréte de ses
meceurs tant au stade larvaire qu’adulte, a une faible
longévité des individus une fois métamorphosés, a
404
leur petite taille, a la difficulté d’identifier les lamproies
a l’espece et dans le manque d’inventaires spécifiques.
Les populations actuellement connues sont localisées
dans le Québec méridional (Figure 1). Elles se retrou-
vent uniquement en eau douce dans le fleuve Saint-
Laurent et dans le cours de certaines rivieres tribu-
taires du Saint-Laurent (Tableau 1). Des inventaires
ciblés pourraient permettre de confirmer la présence
de l’espece ailleurs dans le fleuve Saint-Laurent et
dans certains autres de ses tributaires.
La Lamproie du Nord a déja été observée en grand
nombre dans la riviere Yamaska (pres de Saint-Césaire)
entre 1946 et 1950 (Vladykov 1952). Malgré un
effort intensif au moyen de la péche électrique, aucun
spécimen d’ammocete n’a été observé lors de travaux
réalisés en 1990 (Renaud et al. 1995a) et 1992 (C. B.
Renaud, non publié) a cette localité. Des quatre sta-
tions ou Vladykov (1952) a trouvé des spécimens, le
site de Saint-Césaire était qualifié d’endroit par excel-
lence pour la Lamproie du Nord. L’espéce pourrait
s’étre maintenue dans la riviere Saint-Francois, pres
de Pierreville, ot: elle fut observée en 1947, 1951 et
1990. Vladykov (1952) qualifie ce site d’ habitat idéal
pour les ammocetes, en raison de l’abondance des
iles et du substrat de fond favorable. La Lamproie du
Nord a de nouveau été observée dans cette riviere en
2003, mais cette fois dans la localité de Bromptonville,
a environ 130 km en amont de Pierreville (Desroches
et al. 2008).
Biologie générale
Cycle vital
Le cycle vital des lamproies comprend deux stades
: le stade larvaire (ammocéte) et le stade adulte. Tot
aprés |’éclosion, la jeune ammocete émerge du nid et
migre passivement vers |’aval. Apres avoir trouvé un
habitat favorable, elle creuse un trou en forme de U
dans un substrat plutot mou ou elle demeure géné-
ralement cachée. Les ammocétes sont en effet des
animaux fouisseurs relativement sédentaires (Potter
1980). Ce stade larvaire persiste de trois a six ans
chez la Lamproie du Nord (Hardisty et Potter 197 1a).
Une fois qu’elle a atteint une certaine taille (moyenne
de 131 mm: Leach 1940; moyenne de 114 mm: Purvis
1970), la larve se métamorphose entre les mois d’ aout
et novembre. Le début de la métamorphose est syn-
chrone entre les individus d’une méme population;
un accroissement rapide et substantiel de la tempéra-
ture serait un des facteurs impliqués dans I’ initiation
de ce phénomene (Potter 1980). D’une durée de 2 a 3
mois, la métamorphose est un lent processus au cours
duquel individu cesse de se nourrir (Leach 1940).
On observe méme une réduction de la longueur totale
des individus au cours de cette phase. La bouche se
transforme en un entonnoir buccal pourvu de dents.
Cette période est suivie d’un stade adulte immature
qui se déroule généralement entre novembre et févri-
er et au cours duquel l’individu est semi-sédentaire
(Leach 1940). Une période plus active menant A la
THE CANADIAN FIELD-NATURALIST
Vol. 121
maturité sexuelle a lieu de février 4 mai. L’adulte ne
se nourrit jamais. La fraie survient généralement en
mai et est suivie quelques jours plus tard par la mort
des individus reproducteurs.
Alimentation
Le régime alimentaire des ammocétes se compose
principalement de détritus organiques, auxquelles s’a-
joutent des algues unicellulaires (principalement des
diatomées) et des bactéries (Sutton et Bowen 1994).
Les ammocétes utilisent efficacement les détritus
organiques pendant les mois les plus chauds lorsque
la température des cours d’eau et la qualité des ali-
ments sont plus favorables a |’alimentation, la diges-
tion et la croissance (Sutton et Bowen 1994). Elles se
nourrissent tres lentement, ce qui leur permettrait
d’utiliser au maximum une ressource de faible qualité.
Leur taux d’alimentation est d’ailleurs consistant avec
le faible métabolisme observé chez ce groupe de pois-
sons (Hill et Potter 1970). Le taux d’alimentation
dépend de la température et, a un degré moindre, de la
densité des aliments (Moore et Mallatt 1980). La vari-
ation dans l’abondance des algues et des bactéries
dans l’alimentation des ammocétes correspond a leur
disponibilité dans les cours d’eau durant le cycle
annuel (Sutton et Bowen 1994). Les bactéries et les
algues ne peuvent cependant supporter a elles seules
la croissance et le métabolisme des larves; les détritus
organiques sont nécessaires a ces fonctions (Moore
et Mallatt 1980; Sutton et Bowen 1994). La Lamproie
du Nord adulte, tout comme les autres espéces de lam-
proies non parasites, ne conserve pas de tractus intes-
tinal fonctionnel une fois la métamorphose complétée.
Par conséquent, elle ne peut pas manger et n’atteint
jamais la taille de l’espéce parasite a laquelle elle est
apparentée, soit la Lamproie argentée (Vladykov et
Kott 1979b*).
Reproduction
La Lamproie du Nord fraie uniquement en eau
douce et atteint généralement |’4ge de reproduction a
six ans. Les lamproies ne pondent qu'une seule fois
dans leur vie et tous les adultes meurent quelques
jours apres la fraie (Leach 1940). La température de
leau est le facteur critique qui détermine le début de
la période de reproduction chez les lamproies (Hardis-
ty et Potter 1971b; Manion et Hanson 1980). Au
Québec, la reproduction a lieu en mai, avec un pic de
fraie 4 des températures de l’eau entre 13 et 16°C
(Vladykov 1949*). Comtois et al. (2004) ont capturé
une femelle en fraie un 23 juin dans la riviére Gatineau
a une température de 19°C. Au Michigan, le pic de
reproduction s’étend de la fin mai a la mi-juin, a une
température de eau moyenne de 18°C (Morman
1979*). Toujours au Michigan, Reighard et Cummins
(1916) rapportent une température optimale de l'eau
entre 20 et 22°C ; la fraie était rarement observée a
des températures inférieures a 18°C. Elle aurait lieu
vers la fin mai dans la riviere Tippecanoe, en Indiana
(Leach 1940).
2007
FORTIN ET AL. : BIOLOGIE DE LA LAMPROIE DU NORD AU QUEBEC
405
TABLEAU |. Liste des mentions d’ observations de la Lamproie du Nord, /chthyomyzon fossor, au Québec
Cours d'eau Localité
Année d' observation
Source
Fleuve Saint-Laurent
Lac Saint-Louis
Rapides de Lachine
Pointe-Claire
Riviere Chateauguay Athelstan
Riviere des Prairies Lachenaie
Riviere Gatineau Gatineau
Hinchinbrooke
North Hatley
Mitchell Station
Franklin
Saint-Roch-de-Richelieu
Pierreville
Pierreville
Bromptonville
Huntingdon
Saint-Césaire
Riviere Hinchinbrooke
Riviere Massawippi
Riviere Nicolet Sud-Ouest
Riviere aux Outardes Est
Riviere Richelieu
Riviere Saint-Frangois
Riviere Trout
Riviere Yamaska
“ Centre de données sur le patrimoine naturel du Québec.
Viadykov 1952
1950
194] CDPNQ
1990, 1992 Renaud et al. 1995b, 1998
1998 CDPNQ
1999 Comtois et al. 2004
1976 MRNF®
1995 Desroches et al. 2008
195] Vladykov 1952
2002 J. Dubé, communication personnelle
1990 Renaud et al. 1995b
1947, 1951 Vladykov 1952
1990 Renaud et al. 1995a
2003 Desroches et al. 2008
1976 CDPNOQ
1946 — 1950 Vladykoy 1952
1959 CDPNQ
> Ministére des Ressources naturelles et de la Faune du Québec. Direction régionale de la Montérégie. Fichier informatisé
des relevés fauniques en milieu aquatique et riparien.
En plus de la température de l'eau, deux autres fac-
teurs sont essentiels au succes de ponte, soit un sub-
strat adéquat et un courant uniforme et unidirection-
nel s’écoulant au-dessus du nid (Manion et Hanson
1980; Lanteigne 1992). La Lamproie du Nord fraie
dans des ruisseaux et des petites rivieres sur fond de
gros gravier, de galets (25 a 152 mm de diameétre) et
de sable (Reighard et Cummins 1916; Scott et Cross-
man 1974). Bien que la fraie des lamproies soit géné-
ralement associée a des habitats peu profonds sans
couvert de protection, il est maintenant admis que les
especes du genre /chthyomyzon se reproduisent aussi
sous un couvert de grosses roches ou de débris ligneux
(Cochran et Gripentrog 1992). Ce comportement per-
mettrait aux lamproies de se reproduire dans des
milieux ot l’eau est plus profonde et ot le courant
serait autrement trop fort pour frayer dans des milieux
ouverts. I] réduirait aussi la vulnérabilité des repro-
ducteurs a certains types de prédateurs (Cochran et
Gripentrog 1992). Hardisty et Potter (1971b) rappor-
tent que les cours d’eau typiques des lamproies de
ruisseaux ont une vitesse du courant située entre 0,3
et 0,5 m/s dans les aires de fraie.
Les adultes ne semblent pas migrer en masse vers
les aires de fraie (Leach 1940). Une fois arrivés sur les
lieux, les reproducteurs se regroupent habituellement
dans une aire restreinte (Scott et Crossman 1974).
Les males amorcent alors la nidification (Manion et
Hanson 1980; Moyle et Cech 2004). Ils établissent
leurs nids en déplacant de petits cailloux a l'aide de
leur bouche afin de créer une légére dépression. Ils
déplacent ensuite le sable 4 l'aide de mouvements
vigoureux du corps (Hardisty et Potter 1971b). Le nid
aun diamétre d’environ 7 4 10 cm (Scott et Crossman
1974). La profondeur se situe généralement entre 20
et 46 cm, de la surface de l’eau jusqu’au fond du nid
(Hardisty et Potter 1971b; Scott et Crossman 1974).
Leach (1940) rapporte qu'une femelle pond environ
1 195 ceufs, mais cette donnée n’est basée que sur un
échantillon composé de deux femelles. Pour sa part,
Vladykov (1951) mentionne une moyenne de 1524
ceufs (1115 a 1979), a partir de neuf spécimens
récoltés au Québec. Le diameétre moyen d’un ceuf se
situe entre 1,0 et 1,2 mm (Vladykov 1951; Scott et
Crossman 1974). Apres la fertilisation, les ceufs sont
recouverts du substrat environnant le nid (Hardisty et
Potter 1971b). Les oeufs éclosent 2 4 4 semaines plus
tard (Leach 1940).
Déplacements
La répartition des lamproies le long d’un cours d’eau
est le résultat de deux types de comportements liés
aux déplacements des adultes et des larves. Le premier
correspond a la migration vers l’‘amont des adultes
reproducteurs aux aires de fraie. La migration des
lamproies non parasites se ferait sur des distances
beaucoup plus courtes que chez les espéces parasites
(Hardisty et Potter 1971b). Le deuxiéme type de
déplacements concerne la migration des ammocétes
vers l’aval, 1a ot les conditions d’envasement et de
courant sont plus favorables a l’alimentation et a
l’enfouissement dans le substrat. Cette migration est
en partie passive (Potter 1980). Le déplacement des
ammoceétes est fonction de leurs mouvements, des
conditions hydrographiques et hydrologiques locales,
de la température de l’eau, de la saison, de la stabilité
et de la nature du fond du cours d'eau, de la qualité
de l'eau et de leur densité en relation avec la qualité
de habitat (Hardisty et Potter 1971a: Morman et al.
1980: Potter 1980). Certains auteurs suggerent que
cette migration est minimale, tout particuliérement
dans les cours d’eau ot les gradients sont faibles, ou
l’écoulement est stable et ot les habitats sont propices
406
(Morman et al. 1980). Elle pourrait cependant étre plus
importante dans les cours d’eau ou les gradients sont
plus importants, les ammocetes plus agées étant de plus
en plus prédominantes vers |’aval (Hardisty et Potter
1971a; Potter 1980). Bien que les ammocetes nagent
relativement bien, il est improbable qu’elles puissent
soutenir cette grande vitesse au cours d’une longue
période (Potter 1980). Elles peuvent cependant nager
a contre-courant, si ce dernier est faible (Hardisty et
Potter 1971a). Les déplacements volontaires des am-
mocetes s’effectueraient principalement la nuit (Pot-
ter 1980), ce qui réduirait impact des prédateurs
diurnes. La migration vers l’aval des individus méta-
morphosés se réalise aussi généralement la nuit et est
influencée par le débit et la température du cours d’eau
(Potter 1980).
Parasites et maladies
A notre connaissance, aucune étude n’a décrit la na-
ture et la fréquence des parasites et des maladies spé-
cifiques a la Lamproie du Nord. L’espeéce est proba-
blement susceptible a un bon nombre de maladies non
spécifiques que l’on retrouve chez les autres poissons
d’eau douce du Québec (Uhland et al. 2000). Chez la
Lamproie marine (Petromyzon marinus), Wilson et
Ronald (1967) ont observé que les adultes capturés en
lac avaient un taux d’infection parasitaire (19 %) beau-
coup plus faible que les spécimens qui retournaient
en riviere pour frayer (53 %).
Longévité
Chez les lamproies, la durée du stade adulte est
beaucoup plus courte que celle du stade larvaire. Dans
le cas de la Lamproie du Nord, les adultes ne vivent
qu’environ 6 mois, tandis que les ammocetes ont
besoin de 3 a 6 ans pour arriver a leur métamorphose,
selon le lieu et la nourriture (Okkelberg 1922; Leach
1940; Purvis 1970). L’age maximum d’un adulte serait
donc d’au plus 7 a 8 ans. La durée précise du stade
larvaire est difficile a déterminer avec assurance car
estimation est basée sur des classes de fréquence de
longueur totale des individus (Leach 1940; Scott et
Crossman 1974; Potter 1980).
Habitats
La Lamproie du Nord habite généralement les
ruisseaux et les riviéres a fond graveleux ou sablon-
neux (Jyrkkanen et Wright 1979; Morman 1979*).
Au Québec, on la retrouve aussi dans le fleuve Saint-
Laurent. Elle semble éviter les eaux stagnantes et les
lacs (Hubbs et Trautman 1937). Dans le bassin hydro-
graphique du lac Supérieur, Schuldt et Goold (1980)
ont observé la Lamproie du Nord dans des cours d’eau
de taille intermédiaire ot le débit moyen estival vari-
ait de 0,3 a 28,3 m*/s; l’espéce était commune dans
plusieurs cours d’eau aux eaux turbides et habitait les
régions de la téte des eaux. On ne retrouve la Lamproie
du Nord qu’en eau douce; elle ne tolére aucunement
la salinité (Vladykov 1949*). La riviére Yamaska, prés
THE CANADIAN FIELD-NATURALIST
Vol. 121
de Saint-Césaire, représentait un habitat typique de la
Lamproie du Nord selon Vladykov (1952).
En fonction de l’age des individus, la Lamproie du
Nord requiert deux habitats distincts le long d’un cours
d’eau donné, soit un pour les adultes reproducteurs et
un autre pour le développement des ammocétes. Les
adultes reproducteurs ont besoin de petits cours d’ eau
aux eaux Claires et a gradient topographique élevé,
avec présence de rapides et un fond de gravier et de
sable (Trautman 1981). Quant aux ammocétes, elles
préferent la portion des cours d’eau a4 courant plus lent,
la ou le substrat est propice a l’enfouissement (sub-
strat mou; Potter et al. 1986). Ce substrat renferme sou-
vent un contenu élevé en matériel organique (Hardisty
et Potter 1971a; Potter et al. 1986; Beamish et Lowartz
1996). La composition des sédiments, dont la taille
des particules ainsi que hétérogénéité, constitue un
facteur important dans la sélection de l’ habitat (Hardis-
ty et Potter 1971a; Anderson et White 1988; Beamish
et Lowartz 1996). Les ammocétes éviteraient les fonds
composés principalement de tres petites ou de grosses
particules, le fond étant trop compact dans le premier
cas et les particules trop lourdes pour étre délogées par
les larves dans le second (Beamish et Lowartz 1996).
La taille idéale correspondrait a du sable mi-fin.
Au Michigan, Morman (1979*) a observé que la
Lamproie du Nord était plus tolérante a l’eau chaude
que les autres espéces étudiées. Les plus grandes den-
sités d’ammoceétes se retrouvaient habituellement dans
les sections les plus chaudes des cours d’eau. Ce type
de milieux serait généralement non propice a la pré-
sence de l’Omble de fontaine (Salvelinus fontinalis;
Vladykov 1949*). Les ammocétes sont photopho-
biques, ce qui expliquerait pourquoi |’ombre est un
facteur qui influence significativement leur répartition
(Potter et al. 1986). La quantité de chlorophylle a a la
surface du substrat serait aussi une variable environ-
nementale importante expliquant la densité des larves
de lamproie (Potter et al. 1986). Cette observation re-
flete bien importance des diatomées et autres micro-
algues dans le régime alimentaire des ammocétes qui
évitent généralement les eaux stagnantes ou trop riches
en matiere organique, 1a ou la teneur en oxygene est
faible (Potter 1980).
Dynamique des populations
Les connaissances sur la dynamique des populations
de la Lamproie du Nord sont limitées. La fécondité
de cette espéce fut caractérisée au Québec par Vla-
dykov (1951) et est considérée faible par rapport aux
autres espéces de lamproies étudiées. La survie des
ceufs dans les nids n’a pas été déterminée chez la
Lamproie du Nord, mais chez la Lamproie marine,
celle-ci est élevée, pouvant atteindre 90 % (Manion et
Hanson 1980). A partir d’un échantillon de 627 larves
de Lamproie du Nord, Purvis (1970) a observé un
rapport des sexes de | : 1 dans un tributaire du lac
Supérieur, au Michigan. Chez les adultes reproduc-
2007
teurs, il semble que le rapport des sexes favorise les
males (75 %, Purvis 1970; 71 %, Jyrkkanen et Wright
1979). Ce phénomene est observé chez plusieurs autres
espéces de lamproies non parasites (Potter 1980).
Chez la Lamproie du Nord, comme chez d’autres
especes de lamproies, une classe d’4ge peut contri-
buer a plusieurs cohortes de reproducteurs. En effet,
seule une fraction d’une classe d’a4ge se métamorphose
une année donnée, les autres individus de la méme
classe se métamorphosant au cours des années suiv-
antes (Purvis 1970). Ce recrutement de différentes
classes d’age a la population reproductrice pourrait
expliquer les fluctuations de populations observées
chez la Lamproie de Planer (Lampetra planeri), une
espéce européenne (Hardisty 1961). La température de
eau et la disponibilité de nourriture seraient des fac-
teurs déterminants pour la croissance des ammocetes
(Hardisty et Potter 1971a). La croissance variait beau-
coup entre les classes d’age étudiées par Purvis (1970).
L’auteur suggeére que cette différence est principale-
ment causée par une forte diminution de la densité de
la population consécutive a des traitements chimiques
avec des lampricides. I] en résulterait une diminution
de la compétition pour la nourriture et l’espace et une
augmentation de la croissance dans la population
résiduelle.
Pressions sur l’espeéce, facteurs limitants et
adaptabilité
La détérioration de habitat constitue probablement
la principale pression sur l’espece (Vladykov 1973;
Renaud 1997). Différents types de pollution, seule ou
en combinaison avec d’autres facteurs, restreignent la
répartition des lamproies (Morman et al. 1980; Renaud
1997). La pollution de leau lige au développement
agricole semble étre la principale cause de la dispari-
tion apparente de la Lamproie du Nord de la riviere
Yamaska, au niveau de Saint-Césaire (Renaud et al.
1995a). Les programmes de contrdle chimique de la
Lamproie marine introduite dans les bassins hydro-
graphiques des lacs Huron, Michigan et Supérieur
(Smith et Tibbles 1980) ont affecté les populations
locales d’autres especes de lamproies, dont la Lam-
proie du Nord (Schuldt et Goold 1980). Les larves de
cette espece ainsi que de la Lamproie argentée ont dis-
paru de 41 des 81 cours d’eau traités du bassin hydro-
graphique du lac Supérieur qu’elles fréquentaient jadis,
tant au Canada qu’aux Etats-Unis (Schuldt et Goold
1980). Le lampricide utilisé, le 3-trifluorométhyl-4-
nitrophénol (TEM), est destiné a tuer les larves avant
leur métamorphose. Le genre /chthyomyzon serait de
susceptibilité intermédiaire au TEM (Davis 1970).
La Lamproie du Nord était utilisée autrefois comme
appat pour la péche sportive (Vladykov 1973). Vla-
dykov (1952) mentionne des récoltes totales annuelles
d’environ 300 000 ammocétes pour l’°ensemble du
Québec dans les années 1940 et 1950, toutes espéces
confondues. L’impact qu’a eu cette pratique sur les
populations québécoises est cependant inconnu. Cette
FORTIN ET AL. : BIOLOGIE DE LA LAMPROIE DU NORD AU QUEBEC
407
utilisation pour la péche sportive est maintenant inter
dite au Québec. L’accumulation de sédiments trans
portés par l'eau jusqu’aux aires de fraie, les barrages
et autres obstacles bloquant les remontées d’adultes
reproducteurs et l’abaissement du niveau de |'eau,
lesquels constituent une menace pour la survie des
larves dans leurs abris, représentent d'autres pressions
sur l’espéce (Scott et Crossman 1974; Morman et al.
1980; Lanteigne 1992; Renaud 1997).
La mortalité peut parfois étre élevée au tout début
du cycle vital (Potter 1980). Diverses espéces de pois-
sons ont en effet été observées en train de se nourrir
d’ceufs de lamproies ou de trés jeunes larves (Hardis-
ty et Potter 197la; Vladykov 1973; Potter 1980;
Lanteigne 1992). Les jeunes larves sont aussi suscep-
tibles a la prédation lorsqu’elles quittent le nid et
migrent vers |’aval (Hardisty et Potter 1971a; Potter
1980). La prédation sur les ammocétes plus agées ne
semble cependant pas étre un facteur de mortalité
important car elles demeurent enfouies (Potter 1980).
Trautman (1981) considére les ammocétes et les
adultes comme trés vulnérables 4 la prédation par
d'autres poissons. Par ailleurs, des cas de prédation
sur les adultes reproducteurs ont été rapportés pour
une panoplie de prédateurs tels que belette, chien do-
mestique, couleuvre d’eau, goéland, héron, rat musqué,
raton laveur et renard (Morman et al. 1980). Les lam-
proies au nid sont davantage susceptibles a la préda-
tion, car elles sont plus exposées dans des eaux rela-
tivement peu profondes.
La compétition interspécifique, les parasites et les
maladies ne semblent pas influencer de fagon impor-
tante la répartition et l'abondance de la Lamproie
marine (Morman et al. 1980). L’importance de ces
derniers facteurs pour la Lamproie du Nord reste in-
connue. La faible fécondité de la Lamproie du Nord
Suggeére qu ‘il pourrait étre plus difficile pour elle de se
rétablir a la suite d’un déclin des populations (Schuldt
et Goold 1980; Lanteigne 1992). Aussi, la mobilité
restreinte de l’espéce limite ses chances de pouvoir
recoloniser les cours d’eau ov elle aurait disparu. Ces
éléments font de la Lamproie du Nord une espece peu
adaptable aux modifications de son environnement.
Importance particuliére de l’espéce
Méme si les lamproies constituent un mets de choix
dans certaines régions hors Québec (Moyle et Cech
2004), ce type de poissons est actuellement sans grande
valeur économique. N’étant pas un poisson parasite,
la Lamproie du Nord ne présente pas le danger de
blesser ou de tuer d’autres espéces de poissons, au con-
traire de la Lamproie marine par exemple. La Lam-
proie du Nord présente cependant un intérét certain
d°ordre scientifique et écologique. Cette espéce fait
en effet partie d’un groupe de poissons parmi les plus
primitifs (Forey et Janvier 1993) et offre ainsi l’occa-
sion détudier l’évolution des poissons et les raisons
pour lesquelles les lamproies ont continué a se main-
tenir dans un environnement en perpétuel change-
408
ment (Lanteigne 1992; Moyle et Cech 2004). Les
lamproies sont également utiles comme animaux de
laboratoire, particulierement en neurobiologie (Moyle
et Cech 2004). Les ammoceétes filtrent différents types
de micro-organismes et constituent une source de
nourriture pour plusieurs especes de poissons, d’ou
leur importance dans la chaine alimentaire (Vladykov
1973). La présence de ce poisson dans le Québec
méridional contribue aussi a la biodiversité des ruis-
seaux et des riviéres. Comme les ammocetes sont des
poissons relativement sédentaires passant une bonne
partie de leur vie enfouis dans les sédiments, elles
représentent ainsi des bio-indicateurs potentiels des
niveaux de contaminants persistants dans les milieux
d’eau douce. Une étude réalisée au Québec a démon-
tré que les ammocetes, et notamment celles de la Lam-
proie du Nord, sont aussi appropriées que les mol-
lusques bivalves adultes comme barometres des niveaux
de contaminants organochlorés (Renaud et al. 1995b).
Elles sont considérées comme un bon bio-indicateur
de la contamination au mercure en plus de pouvoir
indiquer les différences dans les niveaux de contam1-
nation en métaux entre les rivieres (Renaud et al.
1998).
Bilan de la situation
Taille des populations et tendances démographiques
Aucune estimation de la taille totale des popula-
tions de la Lamproie du Nord n’est disponible pour
le Québec ou ailleurs dans son aire de répartition
(COSEPAC 2007*). On ne connait donc pas sa situa-
tion exacte ni la tendance démographique des popu-
lations identifiées a ce jour. Seules des données
ponctuelles de taille de populations sont disponibles,
lesquelles sont présentées ci-aprés.
A Saint-Césaire, dans la rivire Yamaska, 61 adultes
et 842 ammocetes ont été capturés entre 1946 et 1950,
dont 721 ammocetes entre le 27 juillet et le 30 aott
1949, au cours d’une étude menée par Vladykov (1952).
En dépit d’un effort de péche intensif, aucun spécimen
n’a été capturé au méme endroit en 1990 et 1992
(Renaud et al. 1995a; C. B. Renaud, non publié). L état
actuel de l’espece dans |’ensemble de la riviére reste
a préciser.
Deux individus métamorphosés et plusieurs ammo-
cetes ont été recensés dans la riviére Saint-Fran¢ois
pres de Pierreville entre 1947 et 1951 (Vladykov
1952). Une étude réalisée en 1990 a permis la récolte
de deux ammoceétes dans cette localité, pres de Vile
Cartier (Renaud et al. 1995a). L’espéce était donc tou-
jours présente a l’époque, bien qu’aucune donnée ne
soit disponible quant a l’abondance de la population.
La situation actuelle a cette localité demeure incon-
nue. Toutefois, l’espéce fut de nouveau observée dans
cette riviére le 26 aout 2003 a Bromptonville, soit a
130 km en amont de la localité de Pierreville (Des-
roches et al. 2008). Il s’agissait d’une ammocéte vi-
vante trouvée dans un bassin. De méme, environ douze
THE CANADIAN FIELD-NATURALIST
Vol. 121
ammocetes mortes et deux vivantes ont été observées
le 20 septembre 1995 dans la riviére Massawippi, un
tributaire de la riviere Saint-Francois, a North Hatley
(Desroches et al. 2008). Quelques secteurs du bassin
versant de la riviére Saint-Frangois semblent donc
propices a la Lamproie du Nord.
Une seule femelle adulte a été capturée dans la riv-
iere Gatineau en 1999 (Comtois et al. 2004). Un seul
individu a aussi été capturé dans la riviere aux Out-
ardes Est en 2002 (Jean Dubé, communication per-
sonnelle). La mention de la localité des Rapides de
Lachine concerne six ammocétes capturées le 26 juil-
let 1950 alors que celle du bassin de la riviere Nicolet
(riviere Nicolet Sud-Ouest) se rapporte 4 une ammo-
céte notée le 22 juillet 1951 (Vladykov 1952). Respec-
tivement trois et six individus de sexe et de stade
indéterminé ont été rapportés sur les rivieres Trout et
Hinchinbrooke en 1976 par le Centre de données sur
le patrimoine naturel du Québec (CDPNQ) et le min-
istere des Ressources naturelles et de la Faune du
Québec, ainsi qu’un adulte en 1998 dans la riviére des
Prairies par le CDPNQ. Finalement, la mention de 1990
de la riviére Richelieu concerne cing ammocétes
(Renaud et al. 1995b).
Menaces a la survie de l’espéce au Québec
La détérioration de l’habitat et la pollution des eaux
semblent représenter les principales menaces a la
survie des lamproies (Renaud 1997; Moyle et Cech
2004), incluant la Lamproie du Nord. La pollution de
la riviere Yamaska, associée a une agriculture intensive
et a des rejets industriels et urbains, apparait comme
une cause possible de l’absence de mention de l’espece
dans la localité de Saint-Césaire en 1990 et 1992
(Renaud et al. 1995a; C. B. Renaud, non publié). Luti-
lisation intensive de l’herbicide atrazine appliqué dans
la culture du mais serait également en cause. Lessivé
dans la riviére suite aux fortes pluies, ce produit chi-
mique réduirait de fagon importante le phytoplancton
dont se nourrissent les ammocétes (Renaud et al.
1995a). On retrouve également de nombreux autres
pesticides dans la riviere Yamaska et plusieurs de ses
tributaires, dont le métolachlore, la cyanazine et le
dicamba (Gouvernement du Québec 1998*). I] n’est
pas rare de découvrir plus de six pesticides différents
au méme endroit. La prédominance de la culture du
mais qui nécessite régulicrement |’application de ces
produits expliquerait en bonne partie cette situation.
Malgré tous les efforts qui ont été déployés au cours
des derniéres années, la riviere Yamaska présente tou-
jours la pire qualité d’eau au Québec. Le portrait n’est
pas plus réjouissant dans le secteur particulier de Saint-
Césaire (Gouvernement du Québec 1998*).
D’autres riviéres ot l’espéce fut observée telles
que les riviéres Richelieu et Nicolet, sont aussi d’une
qualité relativement mauvaise (Gouvernement du
Québec 1993*, 1995*). En fait, depuis 1992, le minis-
tere du Développement durable, de 1’ Environnement
et des Parcs mene un programme de suivi annuel des
2007
pesticides dans quatre cours d’eau des régions agri-
coles ot la culture du mais est importante (Gouverne-
ment du Québec, 2006*). Ces sont les rivieres Chibou-
et (bassin de la riviere Yamaska), des Hurons (bassin
de la riviere Richelieu), Saint-Régis (affluent direct
du fleuve Saint-Laurent) et Saint-Zéphirin (bassin de
la riviere Nicolet). Des pesticides ont réguli¢rement
été détectés durant |’été dans ces rivieres. Les sub-
stances trouvées le plus souvent sont l’atrazine, le
métolachlore, le bentazone, le dicamba, le 2,4-D et le
diméthénamide. Ces substances représentent autant
de menaces pour la Lamproie du Nord en affectant la
qualité de l'eau.
Une autre menace importante a la survie de la Lam-
proie du Nord au Québec est la disparition des bandes
riveraines qui provoque |’érosion des sols et augmente
la sédimentation dans les lits des cours d’eau, altérant
ainsi les aires de fraie. Ce phénomeéne constitue une
menace importante pour les lamproies (Moyle et Cech
2004). Les bandes riveraines boisées, par leur role de
filtres et de stabilisateurs des berges, protegent aussi
les cours d’eau contre les apports en fertilisants et en
pesticides (Société de la faune et des parcs du Québec
2003*). Elles procurent aussi de l’ombre, élément im-
portant de habitat des ammocetes (Potter et al. 1986).
Les superficies requises pour épandre les surplus de
lisier de pore et les monocultures comme celle du mais
sont, entre autres, responsables de la disparition des
bandes riveraines (Société de la faune et des parcs du
Québec 2003*). Les régions de la Montérégie, du
Centre-du-Québec et de Chaudiére-Appalaches sont
particuli¢rement touchées a cet égard.
La croissance de l’industrie porcine a provoqué une
surcharge de fumier et de ses substances polluantes.
Lexcés de fertilisant, et tout particuli¢rement le phos-
phore, touche directement la qualité de l’habitat de la
Lamproie du Nord via l’eutrophisation des cours d’eau
(Société de la faune et des parcs du Québec 2003*).
Cette menace touche potentiellement les cours d’eau de
la Montérégie, du Centre-du-Québec et de Chaudiére-
Appalaches, lesquelles ont été particuligrement tou-
chées par la production porcine au cours des derniéres
décennies, comme par exemple les rivieres Nicolet,
Richelieu, Saint-Frangois et Yamaska.
D’autres menaces potentielles pour habitat de la
Lamproie du Nord concernent le redressement des
petits cours d’eau, lequel pourrait endommager |’ habi-
tat utilisé par les adultes reproducteurs, ainsi que la
présence d’ obstacles (ex. barrages) a la libre circula-
tion de l’espéce entre ses habitats de reproduction et
de développement des ammocétes.
Statut actuel et protection légale
La Lamproie du Nord a été identifiée au Québec
comme espéce susceptible d’étre désignée « menacée
ou vulnérable » en vertu de la Loi sur les espéces me-
nacées ou vulnérables. Au Canada, la Lamproie du
Nord figure 4 l’annexe 3 de la Loi sur les espéces en
péril a titre d’espéce « préoccupante », désignation
FORTIN ET AL. : BIOLOGIE DE LA LAMPROIE DU NORD AU QUEBEC
409
donnée par le COSEPAC en 1991 et une mise a jour
a été effectuée en 2007 (COSEPAC 2007*). En tant
qu’espéce de poissons, la Lamproie du Nord est proté-
gée au Québec en vertu de la Loi sur la conservation
et la mise en valeur de la faune. L habitat de I’ espece
n’est protégé au Québec que sur les terres publiques
par l’application du Réglement sur les habitats fau-
niques, de méme qu’a l’échelle fédérale, tant sur les
terres privées que sur les terres publiques, en vertu de
la Loi sur les péches et la Politique de gestion de
U’habitat du poisson. Au Québec, la Loi sur la qualité
de l'environnement et la Politique de protection des
rives, du littoral et des plaines inondables s’ appliquent
également a I’ habitat de l’espeéce autant en terres pub-
liques que privées.
Evaluation
I] existe un urgent besoin de réaliser des travaux
d’inventaire spécifiques et d’effectuer un suivi des
populations de Lamproie du Nord au Québec, en
donnant priorité aux sites actuellement connus. Des
vérifications s’imposent pour savoir si l’espéce a
survécu dans ces cours d’eau et, le cas échéant, si ces
populations et leurs habitats sont en bonne santé. Une
estimation de la taille des populations serait aussi
souhaitable. Cette initiative devrait étre considérée
comme prioritaire dans le contexte de la conservation
de l’espece au Québec. Actuellement, on estime que
55 % des espéces de lamproies de |hémisphére nord
sont en difficulté (Renaud 1997).
Nos connaissances actuelles sur la situation de la
Lamproie du Nord au Québec vis-a-vis sa répartition
et l’état de ses populations ne nous permettent pas de
nous prononcer quant a la catégorie de conservation
a attribuer a cette espece. En effet, contrairement a sa
situation dans la région des Grands Lacs ot leffort
de péche est connu a cause du programme de controle
de la Lamproie marine, Petromyzon marinus, permet-
tant ainsi de déterminer les tendances des populations
de toutes les espéces de lamproies de cette région, les
mentions de la Lamproie du Nord au Québec ne sont
pas le résultat d°un suivi de longue durée mais plutot
la conséquence de travaux d’inventaire ichtyologique
d’ordre généraux (e.g. Comtois et al. 2004). De plus,
ce n’est que tout récemment (Desroches et al. 2008)
que la présence de la Lamproie du Nord est connue
bien en amont de la riviere Saint-Frangois, a environ
130-165 km de la localité originale de Pierreville
(Vladykov 1952), une indication de notre manque
dinventaire a l’échelle de la province. COSEPAC
(2007*) attribua a la Lamproie du Nord de la région
des Grands Lacs et du haut Saint-Laurent le statut
d’espéce « préoccupante » en raison des menaces dues
a l’épandage de larvicides sur environ 50% de son aire
de répartition (i.e. la région des Grands Lacs). Nous
souhaitons donc que notre article serve de point de
référence pour des études de suivi sur la Lamproie du
Nord au Québec et langons un appel pour celles-ci.
410
Remerciements
Nous remercions J. Dubé (ministére des Ressources
naturelles et de la Faune) pour sa contribution de don-
nées inédites ainsi qu’ Alain Chouinard et Marie-France
La Rochelle pour la réalisation de la carte.
Documents cités (identifiés par un * dans le texte)
COSEPAC. 2007. Evaluation et rapport de situation du
COSEPAC sur la Lamproie du Nord (Ichthyomyzon fossor)
(populations des Grands Lacs — du haut Saint-Laurent et
population de la Saskatchewan — Nelson) au Canada —
Mise a jour. Comité sur la situation des espéces en péril
au Canada, Ottawa, Ontario, Canada. vi +34 pages.
Fortin, C., I. Cartier, et M. Ouellet. 2005. Rapport sur la
situation de la Lamproie du Nord (Ichthyomyzon fossor)
au Québec. Ministére des Ressources naturelles et de la
Faune, Direction du développement de la faune, Québec,
Québec, Canada. 23 pages.
Gouvernement du Québec. 1993. Qualité des eaux de la
riviere Richelieu, 1979 a 1992. Ministére de |’ Environ-
nement du Québec, Québec, Québec, Canada. 12 pages.
Gouvernement du Québec. 1995. Qualité des eaux de la
riviere Nicolet, 1979 a 1994. Ministére de |’ Environne-
ment et de la Faune du Québec, Québec, Québec, Canada.
8 pages.
Gouvernement du Québec. 1998. Etat de l’écosystéme aqua-
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Received 8 March 2006
Accepted 20 August 2008
Seasonal Variation in Sex Ratios Provides Developmental Advantages |
in White-tailed Deer, Odocoileus virginianus
SARAH T. SAALFELD!, STEPHEN S. DITCHKOFF’, JOHN J. OZOGA?, and MICHAEL S. MITCHELL*
‘Arthur Temple College of Forestry and Agriculture, Stephen F. Austin State University, Nacogdoches, Texas 75962 USA \|
[Corresponding author e-mail: saalfeldst@titan.sfasu.edu]
*School of Forestry and Wildlife Sciences, Auburn University, Alabama 36849 USA
3N. 6383 Knox Street, Munising, Michigan 49862 USA
4Montana Cooperative Fish and Wildlife Research Unit, University of Montana, Missoula, Montana 59812 USA
Saalfeld, Sarah T., Stephen S. Ditchkoff, John J. Ozoga, and Michael S. Mitchell. 2008. Seasonal variation in sex ratios provides |
developmental advantages in White-tailed Deer, Odocoileus virginianus. Canadian Field Naturalist 121(4):412-419.
Since Trivers and Willard first proposed their hypothesis concerning the adaptive advantages of producing a particular off- |
spring sex in relation to maternal condition in 1973, it has been at the forefront of scientific research concerning sex ratios |
with most subsequent studies focusing on maternal condition as a key contributor to variations in sex ratios. Another factor |
that could greatly influence sex ratios, although has been only infrequently examined in mammalian species, is birth date. We
investigated how birth date influenced offspring sex ratios in White-tailed Deer (Odocoileus virginianus). Since date of birth
can greatly influence an individual’s fitness and reproductive success, we suggest that birth date may be an alternative strate-
gy in determining offspring sex ratios. Since it has been shown that the lifetime reproductive fitness of a mother can be «
increased by producing a particular sex during a particular time, we hypothesized that more male offspring should be born
earlier in the season due to their increased reproductive potential from being born at this time. Offspring born earlier will
have a head start in development and therefore have greater potential for increased body size and dominance later in life,
traits that greatly influence male reproductive success. In this study, we found that maternal condition did not affect offspring +
sex ratio in a captive population of White-tailed Deer in Michigan; however, birth date did. We found that more males tended
to be born during the second and fourth birthing periods, while more females were born during the first, third and fifth peri-
ods. In addition, we found that males born earlier in the season had greater mass the following spring than those born later, a
trend that was not as dramatic in females. These results lend moderate support to our hypothesis that in White-tailed Deer
offspring sex will tend to vary according to timing of birth and relative reproductive advantages gained by a particular sex
being born at that time.
Key Words: White-tailed Deer, Odocoileus virginianus, birth date, early bird hypothesis, local resource competition, season- »
ality, secondary sex ratio, temporal variation, Trivers-Willard hypothesis.
The potential for offspring sex ratios to differ from
parity has been well documented in a number of mam-
malian species, including White-tailed Deer (Odo-
coileus virginianus) (Verme and Ozoga 1981; Kojola
and Eloranta 1989; Kucera 1991; Landete-Castillejos
et al. 2001; Landete-Castillejos et al. 2004; Sheldon
and West 2004). This strategy could enhance the life-
time fitness of a female by producing the sex with the
greatest potential reproductive value. Some factors
shown to influence this variation include: maternal
age, condition, social status, and habitat quality. Tri-
vers and Willard (1973) proposed that females of sex-
ually dimorphic, polygynous species in good condition
would benefit more from producing sons, because the
mother’s condition would more strongly influence the
future reproductive success of sons than daughters.
Conversely, females in poor condition would benefit
most by producing daughters, since female offspring
require less maternal investment (i.e., milk produc-
tion) and will likely reproduce regardless of maternal
condition. Poor quality male offspring likely will not
develop the physical characteristics necessary to com-
pete successfully for females later in life. Assump-
tions for the Trivers and Willard hypothesis include:
condition of offspring is correlated with condition of
the mother after the parental investment period, con-
dition of the young following the parental investment
period will carry over to adulthood, and conditional
advantages will differentially benefit male offspring in
reproductive success (Trivers and Willard 1973). Since
most ungulate species are polygynous, sexually dimor-
phic species which commonly exhibit these three
assumptions (Hewison and Gaillard 1999), numerous '
studies have been conducted to test the Trivers- Willard
hypothesis. In several of these studies, support for the \
Trivers-Willard hypothesis has been documented, !
where older or more dominant individuals gave birth!
to more males, and younger or subordinate females *
gave birth to more females (Landete-Castillejos et al. |
2001; Landete-Castillejos et al. 2004; Sheldon andi
West 2004).
Despite the many findings that support Trivers and
Willard (Landete-Castillejos et al. 2001; Landete-
Castillejos et al. 2004; Sheldon and West 2004), numer-
ous studies have reported data that seem to contradict
their hypothesis. In a review of published papers on |
412
2007
the genus Odocoileus, Verme (1983) found that more
males were produced by younger females, females in
poor condition, and females in areas of poor habitat
quality. Similarly, Mendl et al. (1995) discovered that
more female offspring were produced by dominant
females in domestic pigs (Sus scrofa). These contra-
dicting trends have been explained by alternative
hypotheses for biased sex ratios, including: the local
resource competition hypothesis (Clark 1978), the
advantaged daughter hypothesis (Hiraiwa-Hasegawa
1993), and the advantaged matriline hypothesis (Leimar
1996). According to the advantaged daughter and the
advantaged matriline hypotheses, mothers are better
able to influence the reproductive success of daughters
rather than sons (e.g., by the transfer of rank and/or
access to high quality resources). Therefore, these
hypotheses propose that high quality females are more
likely to produce females than males. Conversely, the
local resource competition hypothesis states that local
resources rather than maternal condition drive biases
in sex ratios. In matriarchal species, Clark (1978)
proposed that females in good condition will produce
more female offspring, whereas females in poor con-
dition will invest more in male offspring. This is be-
cause female offspring normally establish home ranges
that overlap their mother’s home range, while males
tend to disperse, creating less competition for resources
in the future.
While they differ, all of these hypotheses focus on
maternal condition as the driving factor in offspring
sex ratios of mammals. Another factor that should
greatly influence reproductive potential of offspring
and offspring sex ratios is birth date. In many sexually
dimorphic species, it is believed that time at birth dif-
ferentially affects the ability of a particular sex to re-
produce during its first and subsequent years (Suttie
1983; Clutton-Brock et al. 1984; Kruuk et al. 1999:
Festa-Bianchet et al. 2000). Therefore, having biased
sex ratios of offspring according to birth date could in-
crease a female’s lifetime fitness. A number of avian
species have been shown to have varying sex ratios
seasonally, including the Northern Goshawk (Accip-
iter gentiles: Byholm et al. 2002), European Kestrel
(Falco tinnunculus: Pen et al. 1999), Common Sand-
piper (Actitis hypoleucos: Andersson et al. 2003),
American Kestrel (Smallwood and Smallwood 1998;
F. sparverius: Griggio et al. 2002), Spotted Starling
(Sturnus unicolor: Cordero et al. 2001), and Little
Grassbirds (Megalurus gramineus: McIntosh et al.
2003). In their work with a nonmigratory population of
American Kestrels, Smallwood and Smallwood (1998)
found more male-biased sex ratios earlier in the sea-
son compared to later and proposed that males that
fledged earlier could find nest sites and defend terri-
tories better than males that fledged later. Griggio et
al. (2002) found the same results in a migratory popu-
lation of American Kestrels, with more males born ear-
her in the season. Similarly, it was determined that the
SAALFELD, DITCHKOFF, OZOGA, MITCHELL: SEASONAL SEX RATIOS
413
sex ratio variation could be due to acquisition of nest
sites that were limited for this population. In contrast,
competition for mates seemed the most likely expla-
nation for male-biased sex ratios early in season in the
Common Sandpiper, because males born earlier have
an advantage in obtaining mates (Andersson et al.
2003). Although the explanations for these trends dif-
fer, they suggest that varying offspring sex ratios by
birth date has an adaptive advantage to a mother by
enhancing her own future fitness. Smallwood and
Smallwood (1998) coined this trend the “Early Bird
Hypothesis” and predicted that the sex which gains
the greatest advantage by hatching earlier is selected
for in early conception dates in avian species.
The Early Bird Hypothesis (Smallwood and Small-
wood 1998), however, has not been evaluated in mam-
mals. While it has been demonstrated that older indi-
viduals tend to give birth earlier (Johns et al. 1977:
Festa-Bianchet 1988; Christley et al. 2002), older
individuals tend to produce more male offspring (Lan-
dete-Castillejos et al. 2004), and offspring born earli-
er are more likely to survive due to increased food
resources (Festa-Bianchet 1988; Singh et al. 1990;
Festa-Bianchet et al. 1997; Coté and Festa-Bianchet
2001), the link to differential sex ratios based on birth
date has not been well documented in mammalian
species. We hypothesize that birth date is a driving
factor in offspring sex ratios, with females producing
more male offspring earlier in the breeding season
and increasing production of females as the season
progresses. Because offspring born earlier should
have a longer period of time to grow before their first
winter, more males should be produced at this time
period because of the relative importance of size on
future reproductive success in males and females.
Materials and Methods
We analyzed data from the Cusino enclosure, a
252-ha enclosure located near Shingleton, 46°21'N,
86°28'W, in Michigan’s Upper Peninsula from 1973
through 1984 (excluding 1974). The enclosure consist-
ed mainly of northern hardwoods, mixed hardwood-
conifer stands, and pine (Pinus sp.) plantation. Due to
initial overbrowsing and subsequent condition decline,
deer were supplementally fed ad libitum year-round
with a pelleted ration beginning in 1972 and were fed
throughout the study period. Following this, deer
were documented to be in good physical condition,
even during winter months (Ozoga and Verme 1982).
Initially (1972-1976), no deer were intentionally
removed from the enclosure; however, to ensure a
stable population, the number of individuals within
the enclosure was maintained at approximately 40
individuals with the removal of most fawns and older
individuals each year starting in 1977 (Ozoga and
Verme 1982). Although some potential bias could
result from the removal of such individuals, in a nat-
ural population, most mortality would occur among
414
the removed age classes (i.e., young and old individ-
uals). Therefore, we assume the conditions within the
enclosure are representative of a natural population,
and in fact, the ages of mothers observed in this
study (i.e., | to 13 years of age) falls within the natu-
ral range of breeding females (e.g., ages of pregnant
deer darted in Alabama ranged from | to 13 year of
age as determined by annuli in the cementum; S. T.
Saalfeld, unpublished data).
A complete population census was obtained annu-
ally in March by live-trapping deer and placing them
in a holding pen. At this time, all deer were marked
with numbered collars and weighed. In addition, adult
does were X-rayed to determine stage of pregnancy
and number of fetuses (Ozoga and Verme 1982). Age
of fetuses was determined by comparing fetal radi-
ograms with known-age fetal specimens (105 speci-
mens ranging from 76 to 137 days old) determined by
observed breeding dates (Ozoga and Verme 1985).
Following this, birth dates were ascertained by using
a 199-day gestation. Accuracy of this procedure was
assessed by comparing predicted dates with known
birth dates for 90 litters, and was found to be accu-
rate within 3 days in 50% of litters, within 5 days in
66% of litters, and within 9 days in 93% of litters
(Ozoga and Verme 1985). This variability fell within
the natural variation expected in gestation times of
well-nourished females (i.e., 196-213 days; Ozoga
and Verme 1985). Pregnancy in fawns was also mon-
itored at this time. Since fetuses within fawns would
be too small to be detected on radiograms due to late
conception dates, the presence of pregnancy in fawns
was monitored through progesterone assay in periph-
eral plasma (Plotka et al. 1977).
Following the above procedures, deer were released
back into a holding pen where they could be observed
to confirm mother-fawn lineage from the previous year
in some instances. Although most fawns were cap-
tured and tagged whenever possible within the enclo-
sure to identify their mother, some were not captured
until the March census. Therefore, observations at this
time were used to either confirm or determine mother-
fawn lineage as well as to document the dominance
hierarchy among the deer. The dominance number
assigned to each individual corresponded to the per-
centage of the herd that the individual dominated while
inside the holding pen. Dominance was determined by
observing interactions at feeding stations and record-
ing the number of individuals dominated by a partic-
ular individual. All deer were released from the hold-
ing pen in late March or early April, and thereafter,
observations were used to further elucidate behavioral
patterns, social habits, birthing periods, rearing suc-
cess, and sex of offspring. Fawns were captured and
tagged whenever the opportunity arose and therefore
most were already identified prior to the following
March census. Those that were not tagged by the
March census were identified and tagged for visual
THE CANADIAN FIELD-NATURALIST
Vol. 121
identification at this time (Ozoga and Verme 1982).
All research was performed in a humane manner, fol-
lowed ASM guidelines, and was approved by an
institutional animal care and use committee.
In order to determine any possible role of birth date
on offspring sex ratio we analyzed data similar to
Smallwood and Smallwood (1998). We subdivided
fetuses into 10-day intervals for the range of birth
dates determined from fetal radiograms in this study
(13 May—25 July). Since not all groups had adequate
sample sizes (Figure 1), we pooled data of groups with
samples sizes less than 10 litters. This resulted in the
pooling of dates after 22 June. We then tested whether
sex ratios differed among these groups using repeated
measures analysis of variance repeated among years
with a compound symmetric covariance structure and
birthing group nested within year as our error term
(PROC MIXED: SAS Institute 1999). In addition, to
determine differences in physical condition by birth
date between sexes, we tested if mass of fawns meas-
ured the following spring (March) differed among
birthing groups (e.g., birth date) within each sex using
the same repeated measures analysis of variance. To
test the validity of maternal-condition-based hypothe-
ses within this population, we investigated if any dif-
ference existed among maternal condition (1.e., body
mass and dominance) or age for females that produced
male or female fawns using repeated measures analy-
sis of variance repeated among years with a compound
symmetric covariance structure. Although we were un-
able to account for repeatedly sampling females, we
tested whether sex ratios among the different birthing
periods differed from equality using a Pearson chi-
square.
Results
A total of 428 fawns from 251 litters was exam-
ined during the study from fetal radiograms: mean
birth date was 4 June (n = 230; SE = 0.63 days), and
mean number of fawns per doe was 1.85 (n = 230;
SE = 0.04). The offspring sex ratio (50.4%) for the
entire study did not differ from equality (x = 0.02:
P = 0.880). We detected a difference (F _ = 2.86;
P = 0.031; Figure 2) in the proportion of tales born
among time periods, with a greater tendency of males
being born in the second and fourth periods. We detect-
ed a greater proportion of males being born between
12 June and 21 June (70.4%) than between 13 May
and 22 May (38.9%; t,, = —2.30; P = 0.025), 2 June
and 11 June (46.5%; t,, = —2.78; P = 0.007), and 22
June and 25 July (38.9%; t,.= 2.11; P = 0.040). We
also found a tendency for more males being born
between 23 May and | June (58.0%) than between 2
June and 11 June, however only approaching signifi-
cance (t,,= 1.78; P = 0.081). We found no differences
among other time periods (P > 0.050). Additionally,
we found that the second birthing period had a sex
ratio different than equality (y = 3.93; P = 0.047).
2007
100
80
60
Frequency
40
20
2 June —
11 June
13 May —
22 May
23 May —
| June
SAALFELD, DITCHKOFF, OZOGA, MITCHELL: SEASONAL SEX RATIOS
12 June —
21 June
415
22 June —
1 July
2 July -
11 July
12 July —
21 July
22 July -
25 July
FiGurE 1. Frequency of White-tailed Deer litters in relationship to birth date from 1973 to 1984 (excluding 1974) at the Cusi-
no deer research facility, Michigan, USA.
Body mass of fawns when approximately 9 months
of age decreased as birth date became later for males
(F,,, = 5.11; P = 0.003), but not for females (F,, ,. =
1.94; P = 0.130; Figure 3). We also did not detect any
differences between body mass (f;, = 1.60; P =
0.211), dominance (t,, = 0.00; P = 0.973), or age
(ts7 = 0.62; P = 0.434) among mothers who produced
either males or females.
Discussion
Our results support our hypothesis that offspring
sex ratios differ based on birth date in White-tailed
Deer. However, we found only moderate support for
our hypothesis that more male offspring would be pro-
duced earlier in the birthing season as compared to
females. Although we were unable to detect a signifi-
cant difference in the sex ratios of the earliest birthing
periods, we did find that more males tended to be born
in the second birthing period resulting in a sex ratio dif-
ferent from parity. Although we found only a partial
trend for this hypothesis, this trend has been demon-
strated in other studies of White-tailed Deer, where it
was found that more male offspring were likely to be
conceived earlier in the birthing season (S. S. Ditch-
koff, unpublished data; W. V. Underwood, unpublished
data). In ungulates in a natural environment, individu-
als that are born earlier have more time to grow and
access to more food than late-born conspecifics (Festa-
Bianchet 1988; Singh et al. 1990; Festa-Bianchet et
al. 1997; Coté and Festa-Bianchet 2001). While an ear-
lier birth could be beneficial to both sexes, fitness gain
would potentially be greater for males than females in
a sexually dimorphic, polygynous species. Increases
in body size and quality have been shown to strongly
influence a male’s initial reproductive success and life-
time fitness (Suttie 1983; Clutton-Brock et al. 1984:
Kruuk et al. 1999; Festa-Bianchet et al. 2000). In addi-
tion, early births have also been shown to increase
antler quality (Ozoga and Verme 1982; Schmidt et al.
2001; Gray et al. 2002), a trait which strongly affects
416
dominance and ultimately reproductive success in
ungulates (Clutton-Brock and Albon 1980; Bowyer
1986). Also, postweaning male fawns have a greater
dietary protein requirement than females (Ullrey et al.
1967). Therefore, being born earlier would be most
beneficial for males, since earlier born individuals have
better food availability than late born individuals.
Female fitness, however, is not as strongly influenced
by this initial disadvantage in body size. Jorgenson et
al. (1993) found that condition in female Bighorn
Sheep (Ovis canadensis) did not have a major effect
on age at first reproduction. However, they did detect
a minimum mass for young ewes to begin reproduc-
tion. Variation in mass above this minimum, howev-
er, did not effect age at first reproduction. Therefore,
producing a particular sex at a certain time has an
adaptive advantage for a mother which ultimately in-
creases her lifetime fitness.
In White-tailed Deer, most births normally occur
within a short time period (e.g., several weeks). For
example, 61% of litters in our population were born
during a 15 day period (28 May — 11 June). We expect
that in species that have a much longer birthing peri-
od or multiple litters within a year, seasonal sex ratio
biases would be more prominent. This is true in Vir-
ginia Opossums (Didelphis virginiana: Wright et al.
1995), Meadow Voles (Microtus pennsylvanicus: Dob-
son and Myers 1989), and Gray-tailed Voles (Microtus
canicaudus: Bond et al. 2003), and this pattern has
been termed the first-cohort advantage hypothesis
(Wright et al. 1995). In Virginia Opossums, females
had more male-biased first litters than second litters,
possibly due to the increased reproductive fitness of
males born earlier (Wright et al. 1995). Additionally,
in Meadow Voles, more male-biased litters were born
earlier in the season than later, corresponding to a sim-
ilar pattern in litter size, which also declined as the
season progressed (Dobson and Myers 1989). On the
other hand, Bond et al. (2003), when examining Gray-
tailed Voles, found female-biased litters rather than
male-biased litters early in the season. They hypothe-
sized that this trend was seen since females almost
always reproduce their first year, while most late-born
males delay first breeding until the following spring.
Additionally, they speculated that winter survival could
also influence Gray-tailed Vole sex ratios, where males
born later had greater survival through winter than fe-
males born at the same time. However, this does not
discount the importance of seasonal sex-ratio variation
in a species like White-tailed Deer where births are
highly synchronized. Our data demonstrate the relative
importance of birth dates on female and male condi-
tion the following spring in White-tailed Deer. Males
born in the earliest birthing period had 18.1% greater
mass the following spring at 9 months of age than
males born in the last birthing period. Early-born fe-
males also tended to have greater mass the following
spring than late-born individuals, but the difference
THE CANADIAN FIELD-NATURALIST
Vol. 121
between the groups was less (9.7%). Since our study
was conducted on a supplementally fed population,
we suspect these differences to be less than in a natu-
ral population, where food would be limited during
the winter months. Although these data do not meas-
ure future fitness, they provide strong support for the
hypothesis that birth date differentially influences fu-
ture fitness in males and females, as has been demon-
strated in a number of avian species (Smallwood and
Smallwood 1998; Pen et al. 1999; Cordero et al. 2001;
Byholm et al. 2002; Griggio et al. 2002; Andersson
et al. 2003; McIntosh et al. 2003). Despite this appar-
ent trend, we did not detect a larger proportion of
males born during the earliest time period. Although
we are unsure of the specific reason for this, we suspect
that early in the season, resource availability may be
unpredictable. While giving birth early has advan-
tages, it could also have disadvantages if food is un-
available or unpredictable at this time. Therefore, it
may not be advantageous for females to invest in the
more costly sex (1.e., males) if they may not be able to
provide adequate nutrition for them. Low sample size
during this time period (7 = 12) could also have pre-
vented us from detecting this trend.
Another interesting trend seen in our data was that
more males were additionally born later in the season.
This trend was also seen in American Kestrels, where
more male-biased broods were born later in the season,
although low sample size precluded statistical analy-
sis (Smallwood and Smallwood 1998). Giving birth
late in the season increases the likelihood of produc-
ing smaller, disadvantaged offspring of either sex. In
Elk (Cervus elaphus nelsoni: Smith and Anderson
1998), Red Deer (C. elaphus: Clutton-Brock et al.
1987), and Moose (Alces alces: Keech et al. 2000) off-
spring of either sex that were late-born had decreased
survival and reproduction. We propose that the best
evolutionary strategy for a mother that conceives late
may be to produce the dispersing sex. According to the
local resource competition hypothesis (Clark 1978),
a mother improves her own fitness, as well as that of
previous and future offspring that reside within her
home range, by producing the dispersing sex, which,
by leaving, reduce competition for resources. Our data
support this hypothesis, since both males and females
born late in the birthing season had low mass the fol-
lowing spring compared to offspring born early. Despite
this trend, we were unable to detect a greater propor-
tion of males produced in the last time period, possi-
bly due to the low sample size (9 litters).
Differing sex ratios according to birth date is an
overlooked but perhaps very important tactic for in-
creasing lifetime fitness of White-tailed Deer and other
polygynous mammals. Although this is not the first doc-
umented evidence of seasonal variation in sex ratios in
ungulates (Clutton-Brock et al. 1982; Kohlmann 1999)
or large sexually dimorphic mammals (Coulson and
Hickling 1961; Stirling 1971), we believe our data
2007 SAALFELD, DITCHKOFF, OZOGA, MITCHELL: SEASONAL SEX RATIOS 417
Percent males
(83) (82)
13 May — 23 May — 2 June — 12 June — 22 June —
22 May 1 June 11 June 21 June 25 July
FIGURE 2. Litter sex ratio of White-tailed Deer offspring in relationship to birth date from 1973 to 1984 (excluding 1974) at
the Cusino deer research facility, Michigan, USA. Means with the same letters are not different (P > 0.05). Numbers
correspond to sample sizes of litters in each time period.
@ Females
© Males
Percent males
13 May— 23May— 2June— 12June— 22 June-—
22 May 1 June 11 June 21 June 25 July
FiGuRE 3. Mass of 9-month old male and female White-tailed Deer fawns born in relationship to birth date from 1973 to
1984 (excluding 1974) at the Cusino deer research facility, Michigan, USA. Means with the same letters within an
age class are not different (P > 0.05).
418
provides a better explanation for this phenomenon
than has previously been proposed. Although there are
numerous factors that influence sex ratios and timing
of birth, it appears that timing of birth was the main
cause of variation in sex ratios in the population we
studied. Although our data were collected from a cap-
tive herd of White-tailed Deer, we assume that they
are representative of a natural population, and since
food was provided year round and a stable population
was ensured, we expect some of the trends we have
noted to be more dramatic in a natural population. This
seasonal variation would be most beneficial in species
where reproductive success is correlated with initial
body mass and therefore birth date of a particular sex.
This initial advantage for one sex may be based on a
variety of life history traits that differentially affect one
sex more than the other such as in sexually dimorphic,
polygynous species that exhibit sex-biased competi-
tion for mates or territories.
Acknowledgments
We thank all of the individuals that assisted with
data collection at the Cusino Enclosure during the
time the research was conducted. This research was
supported by the Center for Forest Sustainability and
the School of Forestry and Wildlife Sciences at Auburn
University, the Alabama Cooperative Fish and Wildlife
Research Unit, and the Michigan Department of Nat-
ural Resources.
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Notes
Observation of Foliage-roosting in the Little Brown Bat, Myotis lucifugus
CHRISTINA M. Davy! and ERIN E. FRASER?
' Department of Ecology and Evolutionary Biology, University of Toronto, Canada
? Department of Biology, University of Western Ontario, Canada
Davy, Christina M., and Erin E. Fraser, 2007. Observations of folige-roosting in the Little Brown Bat, Mytotis lucifugus.
Canadian Field-Naturalist 121(4): 420.
First report of foliage-roosting behaviour in a Little Brown Bat (Myotis lucifus). The observation is discussed in relation to
similar behaviour in other bat species.
Key Words: Little Brown Bat, Myotis lucifugus, foliage roosting, Ontario.
We report the occurrence of foliage-roosting behav-
iour by a Little Brown Bat, Myotis lucifugus, which
is generally considered a cavity-roosting species (e.g.,
Barbour and Davis 1969; Fenton and Barclay 1980;
Barclay 1980). At 11:00 hrs on 18 July, 2004, in Algon-
quin Provincial Park (Ontario, Canada) we observed
a single M. lucifugus hanging by one foot from a rel-
atively exposed maple leaf (Acer saccharum) at a
height of about 5 m. The bat was torpid, and would
have been well camouflaged had it not been roosting
directly opposite a small deck. When captured and
handled, it woke from its torpid state within 45 s. and
behaved normally. The bat had a number of ectopara-
sites (bat bugs) on it. However, its ectoparasite load
was not higher than the loads observed on several other
bats captured in the area, so we have no reason to as-
sume the bat was in poor health.
To our knowledge this is the first time that day-
roosting in foliage has been reported for M. lucifugus,
or any North American Myotis, although it is known
from other species in the genus (e.g., the Taiwanese
endemic M. formosus; Y. Ho, personal communication).
Other foliage-roosting species in Ontario, such as the
larger and more brightly coloured Lasiurus species are
notoriously difficult to spot in mature forests (Fenton
1998; B. Hickey personal communication), so such be-
haviour could be easily missed. The Eastern Pipistrelle
(Pipistrellus subflavus) was previously believed to be
an obligate cavity rooster until it was recently found
frequently roosting in foliage (Veilleux et al. 2003).
The individual that we observed was less than
100 m from an old building that contained a maternity
colony of M. lucifugus (>50 individuals), While we
cannot discount the possibility that it was a sick or
excluded member of the colony, we observed no indi-
cations of acute illness. We suggest that our observa-
tion represents further evidence that many bat species
are capable of substantial behavioural flexibility.
Acknowledgements
Our observation occurred during a survey of the
bats of Algonquin Provincial Park. We acknowledge
the generous support of M. Brock Fenton, the Ontario
Ministry of Natural Resources, and the Friends of
Algonquin Park, which made the survey possible.
Literature Cited
Barclay, R. M. R. 1980. Night roosting behaviour of the Lit-
tle Brown Bat Myotis lucifugus. Journal of Mammalogy,
63: 464-474.
Barbour, R. W., and W. H. Davis. 1969. Bats of America.
The University Press of Kentucky, Lexington, Kentucky.
Fenton, M. B. 1998. The bat: wings in the night sky. Firefly
books, Buffalo, New York, USA.
Fenton, M. B., and R. M. R. Barclay. 1980. Myotis luci-
fugus. Mammalian Species. 142: 1-8.
Veilleux, J. P., J. O. Whitaker Jr., and S. L. Veilleux. 2003.
Tree-roosting ecology of reproductive female Eastern Pip-
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Received 18 April 2007
Accepted 6 August 2008
420
2007
NOTES
42]
Cowbane, Oxypolis occidentalis, A New Native Vascular Plant Species
for the Queen Charlotte Islands, British Columbia
MICHAEL CHENEY! and KENDRICK L. MARR?
'Box 556, Masset, British Columbia VOT 1MO Canada; deceased November 2007
Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8W 9W2 Canada
Cheney, Michael, and Kendrick L. Marr. 2007. Cowbane, Oxypolis occidentalis, a new native plant species for the Queen Char
lotte Islands, British Columbia, Canadian Field-Naturalist 121(4); 421-422.
We report the recent discovery of Oxypolis occidentalis, a species that is new to both British Columbia and Canada, disjunct
on the Queen Charlotte Islands.
Key Words: Native vascular plants, new records, Oxypolis occidentalis, Cowbane, Apiceae, Queen Charlotte Islands, British
Columbia, Canada.
The Queen Charlotte Islands, a group of islands
lying off the north coast of British Columbia, continue
to yield new and interesting floristic finds. In the sum-
mer of 2001, Michael Cheney found a small colony of
an unknown Apiaceae growing in and around a series
of seepage pools in an old-growth forest margin at
approximately 300 m elevation on south-central Gra-
ham Island in the Queen Charlotte Islands. A specimen
(Cheney s.n.; V190944) was sent to the Royal British
Columbia Museum (V) for identification, as it did not
match any Apiaceae from British Columbia. Here it
was identified as Oxypolis occidentalis J. Coulter &
Rose, using the Flora of California (Constance 1993).
A specimen sent to the Rocky Mountain Herbarium
(RM) (NIS# 2003-133 [DAO]) was also identified as
O. occidentalis by Ronald Hartman (RM). Specimens
were also deposited in the Ministry of Forests herbar-
ium at Smithers (S). Verification of the identification
of O. occidentalis was based upon the original descrip-
tion (Coulter and Rose 1900) as well as comparison
with herbarium specimens (OSU141785, OSU177111,
UC177437) from Oregon and California. The Queen
Charlotte Islands specimen matches these specimens in
the following characters: seed morphology, little varia-
tion in ray length, and the size, shape and serration of
the leaflets.
Oxypolis occidentalis is new to Canada and belongs
to a solely North American genus of six species, found
in eastern and western North America and the Carib-
bean (Constance 1993). In Canada, one other species
in this genus has been collected, O. rigidior (L.) C. &
R., a species of bogs and wet meadows. This species
was collected in southern Ontario in bogs, wet woods
and swamps, only from 1886 to 1901, and possibly is
now extinct (Scoggan 1979).
The closest populations of O. occidentalis to the
Queen Charlotte Islands populations are approximate-
ly 1400 km distant. The northernmost is from eastern
Lane County (Oregon Vascular Plant Database Record:
ORE103195) in westcentral Oregon as well as Jack-
son and Lake counties of southern Oregon (Peck 1961).
The distribution of collections in California is centred
in transmontane California, particularly the high Sierra
Nevada and the foothills regions of Fresno, Mariposa,
Tulare, and Tuolumne counties from 1200-2600 m in
bogs, wet meadows, streamsides and coniferous forests
(Constance 1993).
In the summers of 2002 and 2003, 10 other widely-
dispersed sites were discovered, revealing that O. occi-
dentalis ranges extensively on the south-eastern flank of
the Skidegate Plateau in suitable sites from 250-370 m
elevation. It is quite possible that it is also found through-
out the Skidegate Plateau and in other suitable habitats
in the Queen Charlotte archipelago. Only one lowland
site has been discovered, at approximately 10 m ele-
vation, 3 km north of Port Clements. Specimens of O.
occidentalis at this site are confined to a shady area ap-
proximately 100 m2, though this small area is densely
populated with O. occidentalis.
At the higher elevation sites on the Queen Charlotte
Islands, O. occidentalis is very frequently associated
with Veratrum viride, Senecio triangularis and Carex
stylosa. The lowland population is associated with
Angelica genuflexa, Carex cusickii, Lysichiton ameri-
canus, Menyanthes trifoliata, and Oenanthe sarmen-
tosa. The known distribution of the populations on the
Queen Charlotte Islands, along with its narrow habi-
tat preferences and its presence in forests that have
not been logged or otherwise disturbed by humans,
suggest that it is not a recent introduction. Whether
O. occidentalis is also found in some of the many
suitable habitats on Moresby Island remains, for the
moment, an intriguing question and one that may shed
some light on its status as a native species of the Queen
Charlotte Islands.
The existence of a disjunct Queen Charlotte Island
population of O. occidentalis raises biogeographical
questions. Queen Charlotte Islands specimens of O.
occidentalis are being examined using molecular tech-
niques at the University of Illinois as part of a system-
atic and phylogenetic treatment of the genus. The
results of this investigation should contribute further to
an understanding of the biogeographical relationships
between the Queen Charlotte Islands populations and
those further south.
422
Literature Cited
Constance, L. 1993. Apiaceae. Page 159 in The Jepson Man-
ual: Higher Plants of California. Edited by J. C. Hickman.
University of California Press, Berkeley.
Coulter, J. M., and J. N. Rose. 1900. North American
Umbelliferae. Contributions from the United States Nation-
al Herbarium 7:192-196.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Peck, M. E. 1961. A manual of the Higher Plants of Oregon,
second edition. Metropolitan Printing Company, Portland.
Scoggan, H. J. 1978-1979. The Flora of Canada. Volumes
1-4. National Museums of Canada, Ottawa.
Received 17 September 2005
Accepted 28 June 2008
Apparent Predation of an American Water Shrew, Sorex palustris, by
an American Marten, Martes americana
Topp Powe. !?, THOMAS S. JUNG !3, and KAREN J. CLYDE !
' Fish and Wildlife Branch, Yukon Department of Environment, P.O. Box 2703, Whitehorse, Yukon Y1A 2C6 Canada
? Current address: Fish and Wildlife Division, Alberta Department of Sustainable Resource Development, P.O. Box 9915,
Fort McMurray, Alberta T9H 2K5 Canada
> Corresponding author. e-mail: thomas.jung @ gov.yk.ca
Todd Powell, Thomas S. Jung, and Karen J. Clyde. 2007. Apparent predation of an American Water Shrew, Sorex palustris,
by an American Marten, Martes americana. Canadian Field-Naturalist 121(4): 422-423.
Little is known of the natural history of American Water Shrews, Sorex palustris, including their predators. We document an
apparent case of predation of an American Water Shrew by an American Marten, Martes americana, in the boreal forest of
southeastern Yukon.
Key Words: American Marten, Martes americana, American Water Shrew, Sorex palustris.
Little is known of the natural history of the Ameri-
can Water Shrew (Sorex palustris). For example, inci-
dents of predation are rarely noted, despite the fact
that the relatively large size of its skull should be easy
to identify in the stomachs, scats, or pellets of preda-
tors. The American Water Shrew is semi-aquatic and
the vast majority of records are from within several
metres of a waterbody (Benenski and Stinson 1987,
Nagorsen 1996). As such, most non-volant predators
of American Water Shrews are aquatic or semi-aquatic,
including large frogs, snakes, and fish (Benenski and
Stinson 1987; Nagorsen 1996). The American Mink
(Neovison vison) is the only reported mammalian
predator of the American Water Shrew (Benenski and
Stinson 1987), and it also is semi-aquatic. We are not
aware of any observations of terrestrial mammals prey-
ing upon American Water Shrews. Here, we provide
an observation of apparent predation of an American
Water Shrew by a terrestrial predator, the American
Marten (Martes americana).
Our observation was made during a live-trapping
study of American Marten near Watson Lake, Yukon
(60.2°N, 128.7°W). On 13 July 2006, we captured an
American Marten and found a dead shrew at the
entrance of the trap. The shrew was identified as an
American Water Shrew using the key provided by
Nagorsen (1996). The trap site was in upland boreal
forest, about 350 m from the nearest waterbody.
We surmise that the Marten was carrying the Water
Shrew and dropped it prior to entering the trap. Alter-
natively, the Water Shrew may have been feeding on
the bait in the trap, where the Marten found it and killed
it. The latter hypothesis is not likely, however, because
the Water Shrew was found outside the trap; for this
hypothesis to be more plausible then the Water Shrew
should have been found inside the trap. Therefore, we
believe that the Marten killed the Water Shrew before
it had entered the trap. It is not known, however, if the
Marten would have consumed the Water Shrew.
Several studies of food habits of American Marten
have reported that shrews are among the least pre-
ferred prey item of American Marten (e.g., Cowan
and Mackay 1950; Quick 1955; Douglass et al. 1983;
Slough et al. 1989; Thompson and Colgan 1990;
Nagorsen et al. 1991; Simon et al. 1999). All shrews
found in the stomach contents or scats of Marten have
been terrestrial species (e.g. S. cinereus, S. hoyi, S.
monticolus). Our observation adds the American
Marten as a likely predator of the semi-aquatic Amer-
ican Water Shrew.
We do not know where the Marten killed the Water
Shrew, but we find it interesting that the Marten may
have killed the shrew while it was away from a water-
body. On 8 July 2006, we found a dead American
Water Shrew about 400 m from a small pond near
Whitehorse, Yukon (60.5°N, 135.1°W) that was killed
by a tethered sled dog. In another instance, on 20
September 2005, we trapped an American Water Shrew
in a pitfall trap in a clearcut near Watson Lake, Yukon,
that was 1.1 km from water. Other observations of
American Water Shrews away from water have been
noted (e.g., Kinsella 1967; Wrigley et al. 1979). Taken
together, these observations suggest that American
Water Shrews may make overland movements, and
2007
that they may be susceptible to mortality during these
movements, including predation.
Acknowledgments
We thank S. Barker, J. Walchuk, and H. Slama for
field assistance. Funding was provided by the Yukon
Department of Environment.
Literature Cited
Benenski, J. T., and D. W. Stinson. 1987. Sorex palustris.
Mammalian Species 296: 1-6.
Cowan, I. McT., and G. H. Mackay. 1950. Food habits of
the Marten (Martes americana) in the Rocky Mountain
region of Canada. Canadian Field-Naturalist 64: 100-
104.
Douglass, R. J., L. G. Fisher, and M. Mair. 1983. Habitat
selection and food habits of Marten, Martes americana, in
the Northwest Territories. Canadian Field-Naturalist 97:
71-74.
Kinsella, J. M. 1967. Unusual habitat of the water shrew in
western Montana. Journal of Mammalogy 48: 475-477.
Nagorsen, D. W. 1996. The Mammals of British Columbia.
Volume 2: Oppossums, Shrews and Moles of British
Columbia. Royal British Columbia Museum, Victoria,
British Columbia, Canada.
NOTES
423
Nagorsen, D. W., R. W. Campbell, and G. R. Giannico.
1991. Winter food habits of Marten, Martes americana, on
the Queen Charlotte Islands. Canadian Field-Naturalist
Quick, H. F. 1955. Food habits of Marten (Martes ameri
cana) in northern British Columbia. Canadian Field
Naturalist 69: 144-147.
Simon, N. P. P., F. E. Schwab, M. L. LeCoure, and F. R.
Phillips. 1999. Fall and winter diet of martens, Martes
americana, in central Labrador related to smal! mammal
densities. Canadian Field-Naturalist | 13: 678-680
Slough, B. G., W. R. Archibald, S. S. Beare, and R. H.
Jessup. 1989. Food habits of Marten, Martes americana,
in the south-central Yukon. Canadian Field-Naturalist 103:
18-22.
Thompson, I. D., and P. W. Colgan. 1990. Prey choice by
marten during a decline in prey abundance. Oecologia 83:
443-45].
Wrigley, R. E., J. E., Dubois, and H. W. R. Copeland.
1979. Habitat, abundance and distribution of six species
of shrews in Manitoba. Journal of Mammalogy 60: 505-
520.
Received 21 November 2006
Accepted 19 August 2008
Giant Pacific Octopus, Enteroctopus dofleini, Attacks on Divers
ROLAND C. ANDERSON!, RONALD SHIMEK2, JAMES A. COSGROVE?, and STEVE BERTHINIER*
'The Seattle Aquarium, 1483 Alaskan Way, Seattle, Washington 98101 USA; e-mail: Roland.Anderson @seattle.gov
?P.O. Box 4, Wilsall, Montana USA
3 Royal British Columbia Museum, Natural History Section, 675 Belleville Street, Victoria, British Columbia V8Y 1G2 Canada:
e-mail: jacosgrove @telus.net
*316 15th Avenue, Seattle, Washington 98122 USA; e-mail: chogless @earthlink.net
Anderson, Roland C., Ronald Shimek, James A. Cosgrove, and Steve Berthinier. 2007. Giant Pacific Octopus. Enteroctopus
dofleini, attacks on divers. Canadian Field-Naturalist 121(4): 423-425.
Documentation is given of four instances of attacks on scuba divers by the giant Pacific octopus.
Key Words: Giant Pacific Octopus, Enteroctopus dofleini, Mollusca, Cephalopoda, attacks, Washington, British Columbia.
In the cold waters around the North Pacific live some
truly giant animals. Among them, individuals of one
octopus species, Enteroctopus dofleini (Wiilker, 1910)
have been documented to weigh over 68 kilograms
(150 pounds) (Cosgrove 1987), making it the world’s
largest octopus (Norman 2002). Although octopuses
(the now preferred plural) have been portrayed as mon-
sters in fiction, folklore, and movies such as /t Came
from Beneath the Sea, octopuses are, as a whole, not
monsters, but interesting and intelligent creatures
(Mather and Anderson 1993, 1998*). Although preda-
tory, they are generally shy and retiring animals that
would rather retreat to a den than have any contact
with humans (Hartwick 1983; Hanlon and Messenger
1996). We have emphasized “as a whole” and “gener-
ally” because octopuses, particularly the larger, more
experienced ones, are individuals (Anderson and Wood
2001). As should be expected with individuals, they
may show decided differences in how they react to any
given situation. Sometimes divers will encounter a large
octopus that is bold and aggressive rather than shy.
The motivations for aggression in normally docile ani-
mals are unknown. It is possible that these animals
may want to see if a diver has food or frankly, they may
well be considering that the diver is food. Whatever the
cause, we here relate four instances of octopus attacks
on divers.
Fishy Business
A diving crew from the Seattle Aquarium was col-
lecting animals near Slant Rock on the outer coast of
Washington State. With rocky cliffs plunging dramat-
ically into the cold Pacific, this is one of the more pic-
turesque areas in a beautiful state. Upwelling at the
entrance to the Strait of Juan de Fuca keeps water clear
and as cold as 4.4°C (40°F) in the summer! Six meter
424
(twenty foot) swells are common and the vigorous
wave action on the rocks keeps the water well-oxy-
genated making it prime octopus habitat. While com-
mon, mature male octopuses in this area are general-
ly small, weighing 14 kg (30 pounds) or less; in
nearby Puget Sound mature male octopuses are com-
monly 27 kg (60 pounds) or heavier.
The team was collecting adult rockfish with a bar-
rier net. Two divers direct the fish into the net, where
they are captured. A third diver bags the fish caught in
the net. On this occasion the bagger noticed that a
small octopus, having an arm span of about 2.4
meters (8 feet) and an estimated weight of about 6.8
kg (15 pounds), had crawled onto the net and had
“flared” across it. When the two other divers returned,
the octopus reached out to each and held onto them,
one at a time. It actually pulled off one diver’s dry
suit glove, causing him great aggravation as the cold
water leaked up his sleeve. The divers gently pushed
the octopus away knowing, from past experience,
that this should be enough to scare it off. They then
picked up their gear and moved to a different nearby
area. While floating in the water 1.5 or 1.8 meters (5
or 6 feet) off the bottom, they were surprised to see
the octopus crawling across the bottom toward them.
It attacked them again, doing a flaring webover
pounce on each of them. When one diver had wres-
tled it off, it moved to another diver and repeated the
behavior. The divers could not get rid of the animal.
This so broke up their concentration they decided to
surface to the support boat overhead, laughing about
the horrible octopus attack!
Now, a 6.8 kg (15 pound) octopus is by no means
a monster. The attack these veteran divers experi-
enced was no worse than a puppy worrying a
shoelace. But as a puppy might chew through an
electrical cord, so might a small octopus do damage
in an encounter with a diver. A giant Pacific octopus
has a formidable beak that could tear a diver’s suit or
flesh (Snow 1970; Anderson 1999). The octopus can
also inject a venomous saliva into a bite (all octopus-
es are venomous) (Hanlon and Messenger 1996).
However, if a big octopus were to attack a novice
diver, the outcome could be different. Instead of sur-
facing laughing, the diver might not surface at all. The
following anecdotes of octopus attacks are more seri-
ous and the outcome could have easily been tragedy
but for the experience and professionalism of the
divers plus some good luck.
Kick Me, Will You?
The second incident happened at Small Pox Bay
on the San Juan Island in Northern Puget Sound. Two
divers were swimming out into the bay from the shore.
One diver was about 3 meters (10 feet) ahead of and
above his partner. He looked back at his buddy and saw
him kick a big rock. Immediately the “rock” changed
color to white then dark red! It was a large octopus, at
THE CANADIAN FIELD-NATURALIST
Vol. 121
least 3 meters (10 feet) across its arms. The animal ex-
tended one arm over the unaware diver’s fins and ankle,
immobilizing him. His forward velocity ceased and,
severely startled, he attempted to rise up in the water.
He was securely held by the octopus clinging to him
and onto the rocks beneath it. The diver grabbed his
dive knife and slapped at the octopus’ arm. The mol-
lusk responded by wrapping another arm around the
diver’s legs. He again attempted to ascend and did rise
a bit, but the octopus was now moving up toward him
by shortening the arms holding the diver and length-
ening the arms anchored to the rock beneath it. The
diver was working hard enough that a steady stream of
air came out his regulator; no discrete breaths were
noticeable. The diver approached a nearby cliff and
picked up a boulder. The diver attempted to drop it on
the large mollusk but just as he did so, the octopus let
go, jetted to the bottom and swam off rapidly. During
the approximately two minutes of the encounter, the
victim had exerted enough effort in trying to break free
of the octopus that he had just 20% of his breathing
gas (600 psi) left in his tank. His buddy’s tank had
67% (2000 psi) remaining. This attack appeared to be
the result of simple aggravation at being kicked.
I Don’t Like Your Looks
At the Keystone Jetty on Whidbey Island, Wash-
ington, a diver was signaled by his buddies to come see
a large octopus sitting in front of its den. As he swam
over, the octopus turned and faced this diver for a
moment then it rapidly swam toward him. The diver
used his camera to fend it off. This worked for a
while. The animal then pulled on his camera, tugging
on the strobes, strobe arms and whole assembly before
moving to the diver himself. At first the diver thought:
“This is cool, it’s going to check me out,” then he felt
it pulling his mask off. Suddenly it was decidedly
“uncool.” He grabbed what he could on the bottom
and pushed the animal off, holding tight to his mask
and biting tightly to hold on to his regulator. The
octopus finally let go and retreated back to its den
after the diver’s buddies shone an ultra-bright canis-
ter light at it.
‘‘Come Into My Parlor,’ Said the Spider to
the Fly...
The next attack occurred in Saanich Inlet on Van-
couver Island, British Columbia. The diver was a scien-
tist working for a crew filming octopuses. The scientist
was very experienced at handling and being around
octopuses. He had enticed an octopus to leave its den
and the film crew went off with it. As the diver, now
alone, gathered his equipment, a second octopus came
out of an unseen den. After looking at the diver for a
moment, the octopus crawled quickly to the diver’s
chest and, while remaining firmly anchored to the rocks
below with several arms, it seized his upper arms,
chest and head. The octopus got three suckers onto the
2007
diver’s bare forehead and wouldn’t let go. The pressure
from the suckers created bruises that lasted for about
a week. During the roughly three-minute struggle the
diver fought to keep his regulator in his mouth by bit-
ing down hard on the mouthpiece and using both hands
to cover it. The diver eventually outlasted the octopus
but in the struggle lost his mask, had his hood pulled
partially off and had his dry suit flooded. As might be
expected, he also used up a significant amount of
breathing gas. The octopus finally gave up on this test
of strength and let go. The diver felt that, had he lost
his regulator or had he been low on air, he would have
undoubtedly died. The octopus was just too strong
for him to do anything but hang onto his regulator
and brace himself from being pulled into the den.
Some Thoughts
Most large octopuses are “pussycats.” Usually, the
larger they are, the more placid and relaxed they are.
But as these experiences indicate, there are exceptions.
At certain stages in their lives, octopuses are lean,
mean, eating machines. They can put on 2% of their
body weight per day (Hartwick et al. 1981; Cosgrove
1987) so they are constantly looking for food. “Check-
ing out a diver” with chemotactic suckers may simply
be seeing if he or she is edible. As far as we know,
the octopuses will let go when they find they cannot
subdue a diver, or the diver is not edible, but the diver
needs to remain calm and keep a tight grip on his or
her regulator. So far, we can document no case of a
Giant Pacific Octopus attack that has resulted in a
diver fatality, but there have been diver deaths where
no cause is found to explain the tragedy. Perhaps octo-
puses were involved in these deaths but even it they
were not it is wise for all divers to be respectful of
the potential danger of these powerful animals.
NOTES
425
Acknowledgments
The authors thank the reviewer for constructive com-
ments. We also thank family members and friends for
their thoughts on this paper.
Documents Cited (marked * in text)
Mather, J. A., and R. C. Anderson. 1998. What behavior can
we expect of octopuses? Available online at: http://www.
thecephalopodpage.org/behavior.php.
Literature Cited
Anderson, R. 1999. An octopus bite and its treatment. The
Festivus 31(4): 45-46.
Anderson, R., and J. B. Wood. 2001. Enrichment for Giant
Pacific Octopuses: happy as a clam? Journal of Applied
Animal Welfare Science 4: 157-168.
Cosgrove, J. A. 1987. Aspects of the natural history of Octo-
pus dofleini, the Giant Pacific Octopus. Unpublished Mas-
ter’s thesis, University of Victoria, Victoria, British Colum-
bia, Canada.
Hanlon, R. T., and J. B. Messenger. 1996. Cephalopod
behaviour. Cambridge University Press. 232 pages.
Hartwick, E. B. 1983. Octopus dofleini. Pages 277-291 in
Cephalopod Life Cycles, Volume I. Edited by P. R. Boyle.
Academic Press, New York, New York.
Hartwick, E. B., L. Tulloch, and S. Macdonald. 1981.
Feeding and growth of Octopus dofleini (Wiilker). The
Veliger 24(2): 129-138.
Mather, J. A., and R. C. Anderson. 1993. Personalities of
octopus. Journal of Comparative Psychology 107: 336-340.
Norman, M. 2002. Cephalopods: a World Guide. Conch-
Books, Germany. 320 pages.
Snow, C. D. 1970. Two accounts of the northern octopus
Octopus dofleini, biting scuba divers. Research Report Fish
Commission Oregon (2): pages 103-104.
Received 2 February 2007
Accepted 16 July 2008
426
THE CANADIAN FIELD-NATURALIST
Vol. 121
Details of Eastern Coyote, Canis latrans, Predation on Great
Black-backed Gull, Larus marinus, Eggs on Boot Island National
Wildlife Area, Nova Scotia
CoLiIn M. MacKInnon!, ANDREW C. KENNEDY!, and DONALD W. CoLpPitTs?
‘Environment Canada, Canadian Wildlife Service, Atlantic Region, P. O. Box 6227, Sackville, New Brunswick E4L 1G6 Canada
2500 Route 16, Point de Bute, New Brunswick E4L 2N1 Canada
MacKinnon, Colin M., Andrew C. Kennedy, and Donald W. Colpitts. 2007. Details of eastern Coyote Canis latrans predation
on Great Black-backed Gull Larus marinus eggs on Boot Island National Wildlife Area, Nova Scotia. Canadian Field-
Naturalist 121(4): 426-428.
We detail field observations of Eastern Coyote eating Great Black-backed Gull eggs for the first time in the literature. Pho-
tographic evidence of the remaining egg shells allowed us to identify the Coyote as the predator.
Key Words: Eastern Coyote, Canis latrans, Great Black-backed Gull, Larus marinus, seabird, egg predation, Boot Island
National Wildlife Area, Nova Scotia, Canada.
Parker (1995) discussed in detail the recent colo-
nization of eastern Canada by the Eastern Coyote
Canis latrans. The coyote arrived in New Brunswick
from Maine around 1970 and this expansion contin-
ued eastward; it was found in mainland Nova Scotia
by 1980 and on Prince Edward Island by 1983. Stud-
ies of coyotes in eastern Canada have reported that
White-tailed Deer (Odocoileus viginianus) and Snow-
shoe Hare (Lepus americanus) are important prey
species (Moore and Miller 1986; Parker 1986; Morton
1988; Patterson and Messier 2001). However, through-
out their range Coyotes are opportunistic and have a
diversified diet (Berg and Chesness 1978; Parker 1995).
In recent years, tracks and scats of coyotes have been
found on a number of off-shore islands in eastern
Canada including areas important to nesting seabirds
(MacKinnon et. al. 2006*). However, actual detailed
descriptive information on coyote depredation on sea-
birds (adults, eggs, or chicks) is generally lacking in
the literature.
Colonial nesting birds typically prefer remote
islands to escape terrestrial predators and mammalian
disturbance. One such colonial seabird colony is located
on Boot Island, Kings County, Nova Scotia (45°08'N,
64°16'W). Boot Island is a 144 ha protected area (Boot
Island National Wildlife Area) consisting of 135 ha of
saltmarsh and two upland islands adjacent to the marsh.
The smaller island (0.5 ha), referred to as “Cyril’s
Island”, is surrounded by saltmarsh and is located
along the northwest border of the NWA. The main
island, Boot Island proper (8.5 ha), is bordered by salt-
marsh to the southwest and 5 m cliffs to the northeast.
Boot Island and Cyril’s Island are predominantly
long-abandoned farm land (6.5 ha) dominated by two
species of wild mustard — Wild Radish, Raphanus
raphansitrum, and Hairy-pod Hedge Mustard, Sisym-
brium officinale. The highest ground is dominated by
a small area (2.0 ha) of dead and dying White Spruce,
Picea glauca, with an understory of predominantly
Red Elderberry, Sambucus racemosa, (Newell et. al.
2006). The western side of Boot Island is separated
from the nearest mainland by a narrow creek (340 m
across water at low tide).
In 2006, the island supported the following colonial-
nesting birds: Great Black-backed Gull, Larus marinus;
Double-crested Cormorant, Phalacrocorax auritus:
Great Blue Heron, Ardea herodias; and Herring Gull,
Larus argentatus, with 943, 151, 52 and 22 breeding
pairs, respectively (MacKinnon et. al. 2006*). The
heron and cormorant nests were in trees and restricted
to the forested area while the Great Black-backed Gull
nests were scattered throughout the old field habitat.
Herring Gull nests were relegated to a few small pock-
ets within the much larger Great Black-backed Gull
colony.
Evidence of Coyote activity (tracks and scats) had
been recorded on earlier visits to Boot Island (person-
al observations). On 10 May 2004, the authors direct-
ly observed a Coyote in the act of egg depredation.
We arrived on the island near high tide, around 06:50
AST, and began a nest survey of the gull colony. By
12:50, the survey was completed and C. M. M. and
D. W. C. were at the extreme northwestern edge of the
main island. At that time we observed a Coyote walk-
ing towards us in an easterly direction, across the salt-
marsh. Its coat was wet (the day was sunny and clear,
14°C) suggesting the animal had just swum across the
narrow creek between Boot Island and the mainland.
When the coyote was first observed, D. W. C. was
sitting on the shore behind a large piece of driftwood,
and C. M. M. was standing, in full view, in the salt-
marsh just east of Cyril’s Island, about 100 m from
D. W. C. Observations of Coyote behaviour were made
at ~ 50 m using 10 x 40 Leitz Trinovid binoculars.
The Coyote proceeded east, passing within 25 m of
the senior author, before changing direction towards
the gull colony on the main island. Upon reaching the
high water mark on the main island (site of the Great
Black-backed Gull colony), the Coyote slowed and
purposefully hunted the upper edge of the seaweed
2007
Figure |. Dimple marks made by Coyote’s teeth (probably
the premolars) when carrying a Great Black-backed
Gull egg, Boot Island National Wildlife Area, Nova
Scotia, 2004. (A. Kennedy photo)
“wrack” line apparently searching for gull nests. It
immediately encountered a Great Black-backed Gull
nest, picked up an egg with its mouth and carried it a
few meters carefully cradled in its jaws, before slicing
it open and eating the contents. Upon seizing the egg,
the Coyote left several small tooth impressions (inden-
tations), next to each other, on one side of the egg
(Figure 1). The Coyote opened the egg by position-
ing it in its jaws allowing one side of its canine teeth
to make a “slice” across the thickest portion (Figure
2). As a result of this lateral cut through the egg, the
shell fragments along the edge of the slice appeared
oriented in the direction of the cut as opposed to fac-
ing uniformly inward as would be expected from a
puncture. The Coyote then dropped the damaged egg
on the ground, spilling the contents (mostly yolk) into
the grass where it was quickly lapped up. The Coyote
consumed a number of eggs (> 5) from different nests
in the same manner. It then continued eastward, away
from our point of observation, presumably predating
more nests and paying little attention to the observers
or the hundreds of very vocal Great Black-backed
Gulls. The Coyote was of average size (~15 kg) and
appeared to be healthy and in good physical shape.
The process of how the Coyote opened the gull eggs
prior to consuming the contents, as well as the tell-
tale marks left on the egg remnants, was not expect-
ed. Had we encountered similar broken gull eggs,
without knowing that they had been depredated by a
Coyote, we probably would have attributed this pre-
dation to American Crow, Corvus brachyrhynchos, or
Common Raven, Corvus corax. are frequent-
ly observed foraging in seabird colonies and depredat-
ed eggs with puncture marks are routinely attributed
to these birds by researchers. Causes of predation are
particularly important if one is trying to determine
survival rates of eggs of various seabird species.
The gull colony on Boot Island National Wildlife
Area has been declining in recent years with a near lin-
NOTES 427
Figure 2. Lateral ‘slice’ made by the canine teeth of a Coy-
ote to open a Great Black-backed Gull egg, Boot
Island National Wildlife Area, Nova Scotia, 2004.
(A. Kennedy photo)
ear decrease in Herring Gull nests from 727 in 1986
to only 22 in 2006 (MacKinnon et al. 2006*). Great
Black-backed Gulls have also experienced a less drastic,
although similar, decline. It is as yet unclear to what
extent Coyote predation may have on the decline in
gull numbers being observed on Boot Island. This note
provides recognizable characteristics on depredated
gull eggs which may help identify egg loss from Coy-
otes versus other predators.
Acknowledgments
We thank P. Chamberland, A. J. Erskine, P. W.
Hicklin and G. R. Parker for helpful reviews of the
manuscript.
Documents Cited (marked * in text)
MacKinnon, C. M., A. C. Kennedy, A. Campbell, and K.
Wellband. 2006. Boot Island National Wildlife Area,
wildlife survey and inspection visit, 8 May 2006. Internal
Report, Environment Canada, Canadian Wildlife Service
— Atlantic Region, Sackville, New Brunswick, 10 pages.
Literature Cited
Berg, W. E., and R. A. Chesness. 1978. Ecology of coyotes
in northern Minnesota. Pages 229-247 in Coyotes, Biol-
ogy, Behavior and Management. Edited by M. Bekoff.
Academic Press, New York, San Francisco, and London,
384 pages.
Moore, G. C., and J. S. Miller. 1986. Food habits and the
average weights of a fall-winter sample of eastern coyotes.
Canis latrans. Canadian Field-Naturalist 100: 105-106.
428
Morton, L. D. 1988. Winter Ecology of the Eastern Coyote,
Unpublished M.Sc thesis, University of New Brunswick,
Fredericton, New Brunswick.
Newell, R. E., C. M. MacKinnon, and A. C. Kennedy.
2006. Botanical Survey of Boot Island National Wildlife
Area, Nova Scotia, 2004. Technical Report Series (450).
Canadian Wildlife Service, Atlantic Region, 32 pages.
Parker, G. R. 1986. The seasonal diet of coyotes, Canis
latrans, in northern New Brunswick. Canadian Field-Nat-
uralist 100: 74-77.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Parker, G. R. 1995. Eastern Coyote — The story of its suc-
cess. Nimbus Publishing Limited, Halifax, Nova Scotia,
Canada, 254 pages.
Patterson, B. R. and F. Messier. 2001. Social organization
and space use of coyotes in eastern Canada relative to
prey distribution and abundance. Journal of Mammalogy
82(2): 463-477.
Received 23 April 2007
Accepted 14 August 2008
Red-breasted Nuthatch, Sitta canadensis, Dies Stuck in Resin at
Entrance to a Nest-box
CLIVE F. STRAUSS
975 Ridgeway Street, Victoria, British Columbia V8X 3C2 Canada; e-mail: cstrauss @islandnet.com
Strauss, Clive F. 2007. Red-breasted Nuthatch, Sitta canadensis, dies stuck in resin at entrance to a nest-box. Canadian Field-
Naturalist 121(4): 428-429.
I document a female Red-breasted Nuthatch (Sitta canadensis) found dead and stuck to resin in the opening of a nest-box in
an urban setting. This is the second reported occurrence of such mortality and the first involving a nest-box. This species
applies resin to its nest openings to deter predators and reduce competition or parasitism from other species. Red-breasted
Nuthatches infrequently utilize nest-boxes. At necropsy this female nuthatch was found to be carrying a fully-formed egg with
shell and this increase in abdominal girth may have contributed to her demise.
Key Words: Red-breasted Nuthatch, Sitta canadensis, mortality, resin, nest-box, British Columbia.
On 12 April 2007 I found a dead female Red-breast-
ed Nuthatch (Sitta canadensis) stuck in the opening of
a nest-box about 5 m above the ground under the west-
facing eaves of a city residence in Victoria, British
Columbia (48°20.0'N, 123°22.0'W). The entrance of
the nest-box was covered by a hard transparent acrylic
plate, measuring 6.7 x 6.0 cm with an oval entrance
hole of diameter 3.4 x 3.0 cm, designed to exclude
House Sparrows (Passer domesticus). Earlier a Chest-
nut-backed Chickadee (Poecile rufescens) was seen
building a nest in this nest-box, and it was often inter-
rupted by a Red-breasted Nuthatch pecking at the open-
ing and calling persistently.
Extraction of the dead nuthatch resulted in numerous
chest and abdominal feathers remaining stuck in the
entrance hole of the nest-box (Figure 1). It was ob-
served that the acrylic plate was irregularly covered in
sticky resin that had an odor suggestive of a coniferous
tree. The interior of the box revealed a newly built
chickadee nest made out of moss but devoid of eggs.
Subsequent to the removal of the dead bird, a pair of
Red-breasted Nuthatches was seen “excavating” this
nest-box by removing moss. In previous years, Violet-
green Swallows (Tachycineta thalassina) successfully
raised broods in this nest-box. The acrylic plate was
added only after House Sparrows killed several swal-
low nestlings.
Several observers in British Columbia (Campbell
et al. 1997) and elsewhere (Bent 1948; Ghalambor and
Martin 1999) have reported Red-breasted Nuthatches
daubing resin around the entrance of their nest holes,
it is thought, to protect the nest against competition from
other cavity-nesting birds or mice, as well as against
predation by other species or insect parasitism. In most
cases this seems to pose little risk to the nuthatch
itself but seems to deter competitors (Ghalambor and
Martin 1999). Daubing nest entrances and cavities
with mud or insects is known for other Sitta species
(Pasquet 1998).
A literature search of the nesting behaviour of the
Red-breasted Nuthatch revealed only one previous
report of death at the entrance of nest hole: Kilham
(1972) reported a dead female Red-breasted Nuthatch |
found stuck to the resin in the entrance of a tree cavity.
Red-breasted Nuthatches infrequently utilize nest- |
boxes. Dunn et al. (1975) described two nest-boxes
utilized out of 100 designed for Black-capped Chick-
adees (Poecile atricapillus) in a pine plantation on the ©
north shore of Lake Erie. Those nest-box entrances ||
were also heavily daubed with pine (Pinus sp.) resin. .|
Nest-box sites were utilized in only two out of 143 }
nests in British Columbia reported by Campbell et al.
(1997). The Cornell Laboratory of Ornithology rec- —
ommends a circular nest-box entrance hole with a
diameter of 3.2 cm for Red-breasted Nuthatches
(Cornell Lab of Ornithology, The Birdhouse Net-
work. Habitat Requirements Available: www.birds.
cornell.edu/birdhouse/bios/nestrequire/habitat.
[Accessed 1 May 2007]). The hole in the cover plate
in this instance measured 3.4 x 3.0 cm.
The dead Red-breasted Nuthatch was found to con- |
tain a complete egg with shell as well as three develop-
2007
NOTES $29
FiGuRE |. Photograph of the acrylic nest-hole cover (removed from the nest-box) with an oval entrance hole of diameter
3.4 x 3.0 cm, showing the Red-breasted Nuthatch feathers stuck to sticky resin on the lower half.
ing ova in the ovary. It is possible that the increased
abdominal girth of this gravid female nuthatch con-
tributed to her getting trapped and subsequent demise
Acknowledgments
My thanks to A. E. Burger of the Department of Bio-
logy, University of Victoria, for performing the necrop-
sy of the dead bird and for his helpful assistance in
the preparation of the manuscript.
Literature Cited
Bent. A. C. 1948. Life Histories of North American Nuthatch-
es, Wrens, Thrashers and their Allies. Order Passeri-
formes. U.S. National Museum Bulletin (195), Washing-
ton, D.C.
Campbell, R. W., N. K. Dawe., I. McTaggart-Cowan., J.
M. Cooper., G. W. Kaiser., M. C. E. McNall, and G. E.
J. Smith. 1997. The birds of British Columbia. Volume 3.
University of British Columbia Press, Vancouver, British
Columbia.
Dunn, E. H., H. F. Howkins, and R. V. Cartar. 1975. Red-
breasted Nuthatches breeding in nest boxes in pine plan-
tations on the north shore of Lake Erie. Canadian Field-
Naturalist 89: 467-468.
Ghalambor, C. K., and T. E. Martin. 1999. Red-breasted
Nuthatch (Sitta canadensis). In The Birds of North Amer-
ica, No. 459 (Edited by A. Poole and F. Gill). The Birds of
North America, Inc., Philadelphia, Pennsylvania.
Kilham, L. 1972. Death of a Red-breasted Nuthatch from
pitch around nest hole. Auk 89: 451-452.
Pasquet, E. 1998. Phylogeny of the Nuthatches of the Sitta
canadensis group and its evolutionary and biogeographic
implications. Ibis 140: 150-156.
Received 11 May 2007
Accepted 11 August 2008
430
THE CANADIAN FIELD-NATURALIST
Vol. 121
Attempt to Cross-Foster Gray Wolf, Canis lupus, Pups into Another
Wolf Pack
RONALD N. SCHULTZ!, ADRIAN P. WYDEVEN2, LINDA S. WINN!, and SHERI A. BULLER!
!Wisconsin Department of Natural Resources, 8770 Highway J, Woodruff, Wisconsin 54568 USA
2Wisconsin Department of Natural Resources, 875 South 4" Avenue, Park Falls, Wisconsin 54552 USA
Schultz, Ronald N., Adrian P. Wydeven, Linda S. Winn, and Sheri A. Buller. 2007. Attempt to cross-foster Gray Wolf, Canis
lupus, pups into another wolf pack. Canadian Field-Naturalist 121(4): 430-432.
We attempted to cross-foster four 18-19 week-old Gray Wolf (Canis [upus) pups into another Wolf pack 182 km from their
natal pack territory. The pup introduction was the result of depredation control on a farm in northwestern Wisconsin. Three
pups died within 14 days of release. A fourth pup survived along the edge of the new pack territory over winter, dispersed in
the spring, joined or formed a new pack and was captured on a depredation complaint four years later.
Key Words: Gray Wolf, Canis lupus, introduce, depredation, natal pack, Wisconsin.
Depredation control programs on Gray Wolves,
(Canis lupus) may create concern about how to hu-
manely deal with wolf pups captured at depredation
sites. Such problems can be especially acute when
lethal controls are not allowed.
It has been demonstrated that Black Bear (Ursus
americanus) cubs have been successfully introduced
to other female bears (Carney 1985). Cross-fostering
of week-old captive Coyote (Canis latrans) pups was
highly successful (Kitchen and Knowlton 2006). Cross-
fostering has been successful in captive Red Wolves
(Canis rufus) with all fostered wolves surviving to
weaning (Waddell et al. 2002*), and has been success-
ful for captive Gray Wolves (Canis lupus) (Goodman
1990). Two captive-born Red Wolf pups were fos-
tered into a wild wolf den containing two pups of
identical age in 2002 (U. S. Fish and Wildlife Service
2004 Fostered red wolf recaptured and released. Red
Wolf News 5: 2, US Fish and Wildlife Service, Manteo,
North Carolina, Unpublished report). Two 6-month-old
pups that were given standard soft releases in separate
translocation incidents remained cohesive with adult
pack members after release and survived (Bradley et
al. 2005). In Wisconsin a wolf pup was successfully
reintroduced back into its natal pack after 53 days in
captivity (Schultz et al. 1999).
However, not all cross-fostering pup translocation
attempts have been successful. An attempt was made
to cross-foster six 3 to 4-week-old Mexican wild wolf
pups (Canis lupus baileyi) to a surrogate adult pair at
the Sevilleta National Wildlife Refuge Mexican Wolf
Management Facility that had two pups, but they were
rejected and killed by the adult male (Mexican Blue
Range Reintroduction Project Monthly Update 1 — 31
May 2006. Unpublished report). Two hard release
translocation events, including three 5-month-old pups
in northwestern Montana resulted in abandonment by
adult members after release, and the pups died soon
thereafter (Bradley et al. 2005).
To our knowledge there have never been any attempts
made to cross-foster 18 to 19-week-old weaned free rang-
ing wild Gray Wolf pups into another wild wolf pack.
We describe an experimental cross-fostering of wild
post-weaning Gray Wolf pups at an age before they
began accompanying adults on hunts (Mech 1970) into
another pack territory in Wisconsin during a time when
translocations were used to deal with problem wolves.
Methods
Four 13 to 14-week-old Gray Wolf pups, two females
(F390, F393) and two males (M391, M392) were cap-
tured on a farm in northwest Wisconsin where cattle
were being killed by wolves. Wolf M392 was captured
on 25 July, M391 on 27 July, F390 on 30 July, and
F393 on 31 July 2001. A lactating adult female, F367
was previously caught on this farm on 26 May 2001,
and released with a shock collar (Schultz et al. 2005)
5.3 km from the farm because she was likely nursing a
litter of pups. Three other wolves (one yearling male,
one adult male, one adult female) were also captured
at the farm in August and were translocated 260 km
from the farm. In previous years, two different wolf
packs bordered the farm: the River Side Pack to the
east and Chases Brook pack to the north and west. We
were unable to determine which pack the pups came
from and were concerned the pups might be abandoned
if translocated with the adults, so they were transport-
ed to a holding pen 174.8 km from their capture site.
The holding pen consisted of a 3 m2 enclosure
with an artificial den (46 x 81 x 61 cm) placed 0.3 km
from the nearest human dwelling. The four pups were
fed a mixture of 60% venison, 40% dry dog food
(Tuffy’s, Division of H. J. Heinz, Perham, Minnesota
56573, USA). All four pups were given a 0.5 ml injec-
tion of Gentocin (Schering-Plough Animal Health
Corp. Kenilworth, New Jersey 07033) daily for 21 days
to treat trap-related injuries.
On 16 August 2001, a White-tailed Deer (Odo-
coileus virginianus) carcass was placed by a rendez-
vous site commonly used by the Bootjack Lake Pack
(BJLP). On 20 July 2001 pups and adult wolves
responded on a howling survey from the rendezvous
2007
Distance (km)
NOTES
431
© #390
Oo #391
A #392
O #393
Ore 4 6 8 IOMZ 1416.18 20.22 24 26 28 30:32
Days after release from holding pen
FiGurRE |. Daily distance pups F390, M391, M392 and F393 moved from holding pen, (28 August to 29 September 2001).
site and on 21 July 2001 an industrial forest employee
saw one gray and three black wolf pups 200 m south-
east of the rendezvous site.
On 19 August 2001 a new holding pen of the same
size was erected 30 m from the deer carcass drop site,
next to the rendezvous site. On 20 August 2001 all four
pups were weighed (10.9 — 11.8 kg) and fitted with
radio tracking collars (Telonics Inc., Mesa, Arizona
85204-6699 USA). The radio collars were lined with
Rockford 3319 foam rubber (Rockford Supply Inc.
1500 Washington Avenue, North Minneapolis, Min-
nesota 55411) to compensate for neck growth. The four
pups were then transported to the new holding pen, and
another deer carcass was dropped off. Three adult
wolves from the BJLP spontaneously howled while
the pups were being placed in the new pen.
On 28 August 2001, all four pups were given a |-ml
external treatment of Revolution (Pfizer Animal Health
Exton, Pennsylvania 19341, USA, Division of Pfizer
Inc., New York 10017) and 0.5-ml subcutaneous injec-
tion of Ivermectin (MSD-AGVET, Division of Merck
& Co., Rahway, New Jersey 107065, USA) to kill a
heavy infestation of lice (Trichodectes canis).
Wolves F393 and M392 were released from the
holding pen on 28 August 2001, and wolves F390 and
M391 were released on 3 September 2001. Venison
and dry dog food were placed in the holding pen every
day for the next five days. Deer carcasses were dropped
off every 2-3 days for the next 17 days. All pups were
located one or more times daily for the first 32 days
after release and one or more times weekly thereafter.
Necropsies on all pups that died were performed
by wildlife disease specialists and pathologists at the
United States Geological Survey (USGS) National
Wildlife Health Center in Madison, Wisconsin.
Results
The pups repeatedly returned to feed on the deer
carcasses and remained within 4.0 km of the release
site for the first 14 days (Figure 1). Adult wolf tracks
were observed at the deer carcasses on 5 and 8 Sep-
tember 2001, and on 9 September 2001 an adult wolf
was observed by the deer carcass drop site, confirm-
ing members of the BJLP were visiting the site.
On 3 September 2001, wolf M392 was found dead
0.4 km from the holding pen. The body was partially
submerged in water on the edge of a small marsh. His
carcass looked to be in good condition, did not have
any external wounds, and had gained 1.5 kg from when
he was collared. Necropsy results revealed congested
lungs, and death probably due to bacterial pneumonia
(USGS, National Wildlife Health Center, Case 17366).
On 12 September 2001, wolf F390 was found dead
1.1 km from the holding pen at the edge of a northern
White Cedar (Thuja occidentalis) swamp. F390 had
deep puncture wounds to the head and throat, and a
fractured skull and ribs. F390 had gained 2.4 kg from
when she was collared (USGS, National Wildlife Cen-
ter, Case 17394). Adult male wolf M336 of the BJLP
was located at the same location as F390 on 10 Sep-
tember 2001, so it was assumed he or other pack mem-
bers may have killed the pup.
On 17 September 2001, wolf M391 was found dead
1.9 km from the holding pen on an old Beaver (Cas-
tor canadensis) dam. M391 had superficial puncture
wounds to the throat and lower jaw. The superficial
wounds may have occurred on 10 September 2001
when M391 was located near F390 and M336. Necrop-
sy results revealed congested lungs and death prob-
ably due to bacterial pneumonia (USGS, National
432
Health Wildlife Center, Case 17395). M391 had gained
2.8 kg from when he was collared. The average weight
gain of all three pups from release date until their
deaths was 0.21 kg per day. This was double the stan-
dard weight gain of 17 wild-caught female wolf pups
(Van Ballenberghe and Mech 1975).
Wolf pups F390, M391 and M392 stayed within
1.9 km of the release pen until their death. Wolf pup
F393 moved 3.7 km from the release site two days
after being released but returned 3 days later. She re-
mained within 2.7 km of the release site until 15 Sep-
tember 2001, when she moved 4.1 km (figure 1) and
stayed within a 3.5 km2 area until 4 December 2001.
She then moved 23.2 km southwest of the holding pen.
Pilots observed F393 with two other wolves on 8 Jan-
uary 2002, but she did not appear to establish a new
territory during the winter of 2002.
In spring 2002, wolf F393 dispersed westward and
by summer had settled into the Long Lake Pack area
106 km southwest of the Bootjack Lake Pack. On 1
August 2005, wolf F393 was captured and euthanized
on a depredation complaint. Five placental scars were
observed on F393 necropsy, confirming she apparently
raised pups in 2004 and 2005 (USGS, National Wild-
life Health Center, Case 19680).
Discussion
The cross-fostering attempt was not successful, but
one pup (25%) of the four did survive and eventually
joined up with other wolves. This pup remained alive,
but abandoned the Bootjack Lake Pack’s territory.
None of the pups joined the new pack, and three died
within 15 days of release. Two of the pups (50%) had
injuries which were assumed to have been inflicted by
Wolves from the BJLP. The cross-fostering attempt
might have been more successful if the pups had been
introduced to a pack adjacent to their natal pack’s
territory where there may have been some interaction
between packs (Hamilton 1964). Perhaps if the pups
had been placed in the BJLP’s territory earlier and
kept in the holding pen longer, the BJLP wolves would
have become more bonded to them (Bradley et al.
2005). We also assume that pup mortality may be high
on this type of introduction, as in bears, and should
only be attempted under extreme circumstances.
Acknowledgments
Persons assisting in the capture and monitoring of
the four wolf pups included Michael Weinfurter, Ed-
THE CANADIAN FIELD-NATURALIST
Vol. 121
ward Zwisik and Mathew Nilson. Wolf monitoring
was funded by the Chequamegon Nation Forest, U.S.
Fish and Wildlife Service, Section 6 Grant, Federal
Aid in Wildlife Restoration Project W-145-R and
Wisconsin Endangered Species Fund. Depredating
wolves were captured by personnel of United States
Department of Agricuture-APHIS-WildlifeServices
under the direction of Robert Willging. A special
thanks to Nancy Thomas, Valerie Shearn-Bochsler, and
Grace McLaughlin at the USGS, National Wildlife
Health Center Madison, Wisconsin for necropsy results.
We also thank Ellen Heilhecker for literature research.
Document Cited (marked * in text)
Waddell, W., S. Behrns, C. Lucash, and S. Mclellan. 2002.
Intraspecific fostering in the red wolf (Canis rufus). Poster
at Carnivores 2002, Monterey, California.
Literature Cited
Bradley, E. H., D. H. Pletscher, E. E. Bangs, K. E.
Kunkel, D. W. Smith, C. M. Mack, T. J. Meier, J. A.
Fontaine, C. C. Niemeyer, and M. D. Jimenea. 2005.
Evaluating wolf translocation as a non-lethal method to re-
duce livestock conflicts in the northwestern United States.
Conservation Biology 19: 1498-1508.
Carney, D.W. 1985. Population dynamics and denning ecol-
ogy of Black Bears in Shenandoah National Park, Virginia.
Masters thesis, Virginia Polytechnic Institute and State
University, Blacksburg, Virginia, USA.
Goodman, P. A. 1990. Jn: A history of wolf park 1972-1999.
Ethology Series, volume 5. Institute of ethology, North
American Wildlife Park Foundation, Battle Ground, Indiana.
Hamilton, W. D. 1964. The genetical evolution of social
behavior. Journal of Theoretical Biology 7: 1-52.
Kitchen, A. M., and F. F. Knowlton. 2006. Cross-fostering in
coyotes: evaluation of a potential conservation and research
tool for canids. Biological Conservation 129: 221-225.
Mech, D. L. 1970. The wolf: the ecology and behavior of an
endangered species. University of Minnesota Press, Min-
neapolis, USA. Pages 136, 140-143.
Schultz, R. N., K. W. Jonas, L. H. Skuldt, and A. P.
Wydeven. 2005. Experimental use of dog-training shock
collars to deter depredation by gray wolves. Wildlife Soci-
ety Bulletin 33: 142-148 USA.
Schultz, R. N., A. P. Wydeven, and J. M. Stewart. 1999.
Acceptance of a gray wolf, Canis lupus, pup by its natal
pack after 53 days in captivity. The Canadian Field-Natu-
ralist 113: 509-511.
Van Ballenberghe, V., and L. D. Mech. 1975. Weights,
growth and survival of timber wolf pups in Minnesota.
Journal of Mammalogy 56: 44-63.
Received 15 November 2006
Accepted 7 August 2008
2007 NOTES 433
Abandoned Seabird Eggs as a Calcium Source for Terrestrial
Gastropods
ALEXANDER L. BOND!:? and ANTONY W. DIAMOND!
' Atlantic Cooperative Wildlife Ecology Research Network (ACWERN) and Department of Biology, University of New
Brunswick, P.O. Box 4400, Fredericton, New Brunswick E3B 5A3
* Current address: Department of Biology, Memorial University of Newfoundland, St. John’s, Newfoundland and Labrador,
A1IB 3X9 Canada; Corresponding author; e-mail: abond@mun.ca
Bond, Alexander L., and Antony W. Diamond 2007. Abandoned seabird eggs as a calcium source for terrestrial gastropods
Canadian Field-Naturalist 121(4): 433-435.
Birds’ use of terrestrial gastropods to supplement calcium for egg formation has been well documented, but the reverse,
gastropods using avian eggshells as a calcium source, has not been recorded previously. We collected Arctic Tern, Sterna
paradisaea, eggs, abandoned in July 2006, that were found to have many terrestrial gastropods on the surface, and it
appeared as though the outer eggshell layers were removed. Using scanning electron microscopy, we compared “grazed” and
“ungrazed” sections of the same shell with eggshells collected in 2005 that were not attended by snails. It appears as though
the snails removed the outer eggshell layers, presumably to supplement their own shell growth.
Keywords: Arctic Tern, Sterna paradisaea, White-lip Gardensnail, Cepaea hortensis, Glossy Pillar, Cionella lumbria, calcium,
eggshell, New Brunswick
Although is it true that birds may seek to augment
calcium intake during egg production (Nisbet 1997),
particularly in acidified environments (Beintema et al.
1997; Brenninkmeijer et al. 1997; Mand et al. 2000),
the reverse relationship of snails exploiting an avian
calcium source has not been recorded. This is likely
because access by snails to avian calcium sources (i.e.,
eggs) is either limited during periods of nest attendance
and incubation by adult birds, or because other avian
calcium sources such as bones from dead chicks or
eggshells ejected from the nest decompose or other-
wise disappear from view.
Machias Seal Island (MSI; 44°30'N, 67°06'W) is a
9.5 ha treeless island in the Bay of Fundy, Canada, and
is home to several nesting seabird species, including
Arctic Tern, Sterna paradisaea, and Common Tern, S.
hirundo (Diamond and Devlin 2003; Bond et al. 2007).
In the summer of 2006, the tern colony was aban-
doned on 25 June, before any eggs hatched. Although
the proximate cause likely was intense predation on
tern nests by Herring Gulls, Larus argentatus, and
Great Black-backed Gulls, L. marinus (AWD unpub-
lished data), the ultimate cause is likely a decrease in
available food supply, mainly 0-group Atlantic Her-
ring, Clupea harengus (DFO 2005), which preceded
several years of near reproductive failure for both tern
species (Bond et al. 2007).
On 22 July 2006, eleven tern eggs that had not been
taken by gulls were collected and it was noticed that
many small land snails had attached themselves to the
exterior of the tern eggs, causing surface aberrations
(Figure 1). Through visual comparison with eggs not
attended by snails, we wanted to determine if snails
were “grazing” on the tern eggs. Snails have been ob-
served on abandoned eggs at tern colonies in southern
New England (I. C. T. Nisbet, personal communica-
tion, 2006), but their effects on the eggs’ exterior was
not examined.
Our objective was to use electron microscopy to
compare visually the surface of eggshells attended by
snails with sections of the same eggshell that were un-
attended and with shells of eggs that had no snails
present at all.
Methods
Tern eggs were collected on 22 July 2006 from
Machias Seal Island, Bay of Fundy, Canada. Eggs were
taken from nests located in the central vegetated por-
tion of the island, where Canada Bluejoint, Calama-
grostis canadensis, dominates and is kept short by
regular mowing. Areas immediately surrounding the
tern nests were not mowed to avoid damage to the
nests’ contents. Once the eggs were collected, snails
were removed from the eggs. Eggs were wrapped
individually in cellophane, frozen at -18°C and trans-
ported to the University of New Brunswick, Frederic-
ton, for analysis. Snails were similarly frozen in ster-
ile polythene bags, and transported.
Eggshell samples were prepared and examined
using a JEOL JSM 6400 scanning electron microscope
with an operating voltage of 7 kV at the Microscopy
and Microanalysis Facility at the University of New
Brunswick. Small sections of shell were mounted on
aluminium stubs using carbon tape and carbon paste
to properly ground the specimens, and sputter-coated
with approximately 50 nm of gold. Photographs were
taken at either 90° (from directly above) or at 50° to
the electron beam.
434
10mm
FiGurE |. Abandoned Arctic Tern egg. Light circular marks
on the shell’s surface were the locations of snails.
Results and Discussion
Gastropods were identified at the New Brunswick
Museum (Saint John, New Brunswick) as Cionella
lumbria (Glossy Pillar; Stylommatophora: Cionelli-
dae), a common terrestrial gastropod in the northeast-
erm United States (Martin 2000), and immature or
hatchling Cepaea hortensis (White-lip Gardensnail;
Stylommatophora: Helicidae), which is common on
several islands in the Bay of Fundy (D. McAlpine,
New Brunswick Museum, St. John, personal commu-
nication, 2006).
Individual eggs had between | and 14 gastropods
present (mean + SD, 4.18 + 3.79), with Cepacaea
hortensis being more common (40 of 46 total gas-
tropods found on ten eggs), and the remainder being
Cionella lumbria (6 of 46, present on three eggs). Lin-
ear regression found no relationship between snail
abundance and egg size (breadth or length).
Areas on the tern eggshell that were “grazed” by the
snails had the outer surface layer removed, exposing
the radially oriented crystals immediately beneath the
tangentially-oriented crystals that form the shells’ sur-
face (Becking 1975; Figure 2), whereas ungrazed areas
(Figure 3 top) appeared similar to shells collected in
2005 (Figure 3 bottom). An examination of the inter-
face of the grazed and non-grazed areas with the
photograph taken at 50° to the electron beam shows a
“landscape” view, with the grazed areas lower than
non-grazed areas (Figure 4), indicating the removal of
the outer calcified layer of the eggshell by the snails.
This outer crystalline layer varies in thickness from 1
to 10 mm in other bird species (Tyler 1964; Becking
1975), but it has not been examined previously in any
of the Laridae.
It is unlikely that this method of calcium acquisi-
tion by snails would occur had the terns remained and
incubated their nests through to hatching. Arctic Terns
incubate their clutch of 1-3 eggs for approximately
21-23 days (Hatch 2002). Paquet (2001) found that
THE CANADIAN FIELD-NATURALIST
Vol. 121
FIGURE 2. Grazed (top) and ungrazed areas (extreme bot-
tom) of an Arctic Tern egg. Electron micrograph
taken at 90° to electron beam.
incubating terns spent 41% of their daily time budget
incubating the egg, and that for more than 80% of the
time, at least one adult in the pair was tending the
eggs. This implies that any colonization attempt by
gastropods would be unsuccessful because the egg
would be unattended and accessible by the gastropods
very little of the time. If such colonization did occur,
the contact with the adult during incubation would
likely remove the gastropod from the eggshell.
It is likely that the large number of abandoned eggs
drew our attention to what could be a common phe-
nomenon at the microscopic level. Seabirds can alter
soil chemistry drastically (Mizutani et al. 1991), and
it is likely that eggshells ejected from nests and bones
from dead adults and chicks provide a reliable source
of calcium enrichment, which could be accessed by
the snails.
Rainfall on Machias Seal Island was well above aver-
age in 2006, with 469 mm between 15 May and 18 Au-
gust, compared with the mean (+ S.E.) of 225.0 mm
(+ 29.2) over the previous 12 years (Bond et al. 2007),
and this likely contributed to high productivity among
the snails (Barrientos 2000; Tattersfield et al. 2001).
The gastropods’ use of avian eggs as a calcium
source is a reversal of the more typical relationship be-
tween birds and molluscs, as birds in acidified envi-
ronments are often documented consuming molluscs
to supplement production of calcium for their eggs
(Beintema et al. 1997; Brenninkmeijer et al. 1997; Nis-
bet 1997; Mand et al. 2000). It is our belief, though,
that this behaviour in the snails is likely to be regular
or frequent in occurrence, and that the snails are seek-
ing new and abundant natural sources of calcium to
promote shell growth, including abandoned eggs.
Acknowledgments
G. Bouchard and T. Clarke provided assistance in
the field and A. Patterson provided transportation to
and from the island. B. Bagley, G. Clark, P. Cranford,
2007
FicureE 3. An ungrazed portion of the same shell as Figure 2
(above) and an Arctic Tern eggshell collected in 2005
with no snails observed (below). Electron micrographs
taken at 90° to electron beam.
R. Daley, R. Eldridge and R. Ross provided friend-
ship and company on the island. D. F. McAlpine
identified the snails, S. Belfry assisted with the SEM
photography and S. Martin assisted in locating older
references. K. C. Fraser, J. L. Lavers, A. J. Erskine and
one anonymous reviewer provided helpful comments
on previous drafts. The Atlantic Cooperative Wildlife
Ecology Research Network funded this project. This
is ACWERN publication number UNB-73.
Literature Cited
Barrientos, Z. 2000. Population dynamics and spatial dis-
tribution of the terrestrial snail Ovachlamys fulgens (Sty-
lommatophora: Helicarionidae) in a tropical environment.
Revista de Biologia Tropical 48: 71-87.
Becking, J. H. 1975. The ultrastructure of the avian eggshell.
Ibis 117: 143-151.
Beintema, A. J., T. Baarspul, and J. P. De Krijger. 1997.
Calcium deficiency in Black Terns Chlidonias niger nest-
ing on acid bogs. Ibis 139: 396-397.
Bond, A. L., M.-P. F. McNutt, T. C. Clarke, and A. W.
Diamond. 2007. Machias Seal Island 1995-2006 Progress
Report. Atlantic Cooperative Wildlife Ecology Research
Network, Fredericton, New Brunswick.
NOTES
BAN
»
A
ee
FIGURE 4. Arctic Tern eggshell showing grazed (bottom) and
ungrazed (top) interface. Electron micrograph taken
at 50° to electron beam to provide a “landscape” view.
Brenninkmeijer, A., M. Klaassen, and E. W. M. Stienen.
1997. Sandwich Terns Sterna sandvicensis feeding on shell
fractions. Ibis 139: 397-400.
DFO (Department of Fisheries and Oceans Canada).
2005. Stock Assessment Report on 4VWX_ Herring.
Fisheries and Oceans Canada, Science Advisory Secre-
tariat, Science Advisory Report 2005/033 (Revised).
Diamond, A. W., and C. M. Devlin. 2003. Seabirds as indi-
cators of changes in marine ecosystems: ecological mon-
itoring on Machias Seal Island. Environmental Monitoring
and Assessment 88: 153-175.
Hatch, J. J. 2002. Arctic Tern (Sterna paradisaea). In The
Birds of North America, (707). Edited by A. Poole and F.
Gill. The Birds of North America, Inc., Philadelphia, Pen-
nsylvania.
Mand, R., V. Tilgar, and A. Leivits. 2000. Reproductive
response of Great Tits, Parus major, in a naturally base-
poor forest habitat to calcium supplementation. Canadian
Journal of Zoology 78: 689-695
Martin, S. M. 2000. Terrestrial snails and slugs (Mollusca:
Gastropoda) of Maine. Northeastern Naturalist 7: 33-88.
Mizutani, H., Y. Kabaya, P. J. Moors, and T. W. Speir.
1991. Nitrogen isotope ratios identify deserted seabird
colonies. Auk 108: 960-964.
Nisbet, I. C. T. 1997. Female Common Terns Sterna hirundo
eating mollusc shells: evidence for calcium deficits during
egg laying. Ibis 139: 400-401.
Paquet, J. M. 2001. Time-budget flexibility of breeding
Arctic Terns (Sterna paradisaea): an investigation of the
behavioural buffer. M.Sc. thesis, University of New
Brunswick, Fredericton, New Brunswick.
Tattersfield, P., C. M. Warui, M. B. Seddon, and J. W.
Kiringe. 2001. Land-snail fauna of afromontane forests
of Mount Kenya, Kenya: ecology, diversity and distribu-
tion patterns. Journal of Biogeography 28: 843-861.
Tyler, C. 1964. A study of the egg shells of the Anatidae.
Proceedings of the Zoological Society of London 142:
547-583.
Received 9 March 2007
Accepted 26 August 2008
436 THE CANADIAN FIELD-NATURALIST Vol. 121
Fish-line Entanglement of Nesting Mourning Dove, Zenaida macroura
GLENN H. PARKER and CHRIs G. BLOMME
Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada
Parker, Glenn H., and Chris G. Blomme. 2007. Fish-line entanglement of nesting Mourning Dove, Zenaida macroura. Canadian
Field-Naturalist 121(4): 436-437.
We give a detailed account of monofilament entanglement in nesting material of a gravid female Mourning Dove (Zenaida
macroura) which resulted in death. Such incidents substantiate the hazards of anthropogenic materials and the need to dis-
pose of them with due diligence.
Key Words: Zenaida macroura, Mourning Dove, monofilament, fish line, avian anthropogenic hazards.
Mourning Doves (Zenaida macroura) nest annually
in the town of Val Therese (46°39'N, 81°01'W), Sud-
bury District, Ontario. On 12 April 2005, an unusual
flash of movement attracted my eye to the neighbour’s
15-meter-tall Blue Spruce (Picea pungens). Upon more
careful inspection, I noticed a Mourning Dove hanging
unnaturally among the interior boughs of the spruce. An
additional bird was present, presumed to be the other of
a mated pair. Entering under the boughs, I noticed that
the lifeless dove was hanging from a thread of mono-
filament fish line to which it had succumbed following
an obvious struggle. One meter above the dangling
bird, situated on a branch 1.75 meters from the trunk
and 2.75 meters off the ground, was a fully-formed nest.
The hanging bird was a female Mourning Dove
weighing 124 grams. Autopsy revealed that the bird
was in a healthy state; the crop was full of wild seeds
and normal amounts of adipose tissue were present
throughout the body. A fully-developed egg with a
near-calcified shell was present in the oviduct, indicat-
ing that egg laying would probably have occurred with-
in a day. The nest was well-formed but empty, imply-
ing that nest construction had been completed and egg
laying was about to commence.
The fish line was of ten-pound test and consisted of
two separate lengths each entwined in the nesting mate-
rial. The smaller piece measured 25 centimeters and the
longer piece 2.5 meters. The longer piece was woven
into the nest at mid-level, indicating that it had been
incorporated during nest building rather than added
later. The bird had become entangled with a portion of
the longer line. The fish line ran between the sixth and
seventh primary on the underside of the right wing,
then looped over the dorsal surface of the humerus to
the neck where it was looped completely around the
neck. Subsequently, the line stretched to the left wing
which was wrapped twice very close to the body, and
then extended from the dorsum of the bird as a free end.
The bird was dangling from the line anchored within
the nest, with a single upwardly-extended wing and a
severely down-turned head and neck directed into the
pit of the extended wing. Strangulation as a result of
the initial entanglement and panic response is inferred
as the cause of death.
Various anthropogenic hazards have been reported
to contribute to mortality in Mourning Doves, includ-
ing decapitation from power lines, utility lines and TV
towers while in flight (Stoddard and Norris 1967), win-
dow strikes (Terres 1980), and vehicle, train and aircraft
strikes (Terres 1980). Monofilament fish line entangle-
ment has been well documented in waterfowl, marine
birds. (Manville 2005; King et al. 1979) and loons
Gavia spp. (Sidor et al 2003) as well as noted in doves
on previous occasions (Sadler 1993; Mirarchi and Bas-
kett 1994). The tendency of certain species to in-
corporate foreign anthropogenic materials during nest
building, including fine wires as noted in dove nests
reported by Peck and James (1983), appears to predis-
pose such species to higher risk levels. Specific details
of incident reports appear to be lacking in much of the
literature. This note serves to further document the use
of discarded fish line (a non-natural nesting material)
and its potential contribution to mortality in a common
tree-nesting species.
Literature Cited
King, W. B., R. G. B. Brown, and G. A. Sanger. 1979. Mor-
tality to marine birds through commercial fishing. Pages
95-199 in Conservation of marine birds in northern North
America. Edited by J. C. Bartonek and D. N. Nettleship.
Wildlife Research Report (11). Fish and Wildlife Service,
U.S. Department of the Interior.
Manville, A. M., II. 2005. Seabird and waterbird bycatch in
fishing gear: next steps in dealing with a problem. Pages
1071-1082 in USDA Forest Service General Technical
Report PSW-GTR-191.
Mirarchi, R. E., and T. S. Baskett. 1994. Mourning Dove
(Zenaida macroura). In The Birds of North America (117).
Edited by A. Poole and F. Gill. The Academy of Natural
Sciences, Philadelphia; The American Ornithologists’
Union, Washington, D.C.
Peck, G. K., and R. D. James. 1983. Breeding birds of On-
tario, Nidiology and distribution. Volume 1: Nonpasserines.
Publications in Life Sciences. Royal Ontario Museum,
Toronto. 321 pages.
Sadler, K. C. 1993. Other natural mortality. Pages 225-230 in
Ecology and management of the Mourning Dove. Edited
by T. S. Baskett, M. W. Sayre, R. E. Tomlinson, and R.E.
Mirarchi. Stackpole Books, Harrisburg, Pennsylvania.
Sidor, I. F., M. A. Pokras, A. R. Major, R. H. Poppenga,
K. M. Taylor, and R. M. Miconi. 2003. Mortality of the
~
2007 NOTES 43
Terres, J. K. 1980. The Audubon Society En
North American Birds. Knopf Inc., New York
common loons in New England, 1987 to 2000, Journal of
Wildlife Diseases 39: 306-315.
Stoddard, H. L., and R. A. Norris. 1967. Bird casualties at
a Leon County, Florida TV tower: An eleven year study.
Tall Timbers Research Station. Bulletin (8). Tallahassee,
Florida.
yclopedia of
Received 14 May 2007
Accepted 28 July 2008
Bull Trout, Salvelinus confluentus, and North American Porcupine,
Erethizon dorsatum, Interaction in the Mackenzie Mountains,
Northwest Territories
PETER A. Cott!? and NEIL J. MOCHNACZ?
' Fisheries and Oceans Canada, 101, 5204-50" Avenue, Yellowknife, Northwest Territories X1A 1E2 Canada
? Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Canada
* corresponding author; e-mail: Pete.Cott@dfo-mpo.ge.ca
Cott, Peter A, and Neil J. Mochnacz. 2007. Bull Trout, Salvelinus confluentus, and North American Porcupine, Erethizon dorsat-
um, interaction in the Mackenzie Mountains, Northwest Territories. Canadian Field-Naturalist 121(4): 438-439.
Evidence of an interaction between a Bull Trout (Salvelinus confluentus) and a North American Porcupine (Erethizon dorsatum),
was observed during a fisheries survey in the Mackenzie Mountains, Northwest Territories. A male Bull Trout with porcupine
quills imbedded in its mouth was collected. It is speculated that this occurred when the trout was aggressively defending its
territory from a perceived threat — a swimming porcupine. This is the first documented account of an interaction between a
fish and a porcupine.
Key Words: Bull Trout, Salvelinus confluentus, North American Porcupine, Erethizon dorsatum, prey, interaction, diet, North-
west Territories, Mackenzie Mountains, Mackenzie Valley.
We report an interaction between a Bull Trout
(Salvelinus confluentus) and a North American Porcu-
pine (Erethizon dorsatum), two animals that, due to
different life history requirements, would rarely be in
contact.
The Bull Trout is a native salmonid (genus Salveli-
nus) occurring in streams, rivers and lakes across west-
ern North America. The species’ range extends from
northern California to the central Northwest Territories
[NWT] (Haas and McPhail 1991; Reist et al. 2002;
Mochnacz et al. 2004; Stewart et al. 2007a). Bull Trout
are a top level predator consuming a variety of prey
items ranging from insects to small mammals (Stew-
art et al. 2007b). North American Porcupines are large
rodents, weighing up to 14 kg and are widely distrib-
uted from the Arctic Ocean to northern Mexico (Roze
1989). Porcupines occupy a variety of habitat types and
are considered generalist herbivores (Roze 1989).
Their dorsal surface is covered with modified guard
hairs in the form of barbed quills, which serve as pro-
tection against predators. Predators include the Wolf
(Canis lupus), Coyote (Canis latrans), Wolverine
(Gulo gulo), Fisher (Martes pennanti), Cougar (Puma
concolor), Canada Lynx (Lynx canadensis), Bobcat
(Lynx rufus), and Great Horned Owl (Bubo virgini-
anus) (Roze 1989). When a porcupine is attacked, the
quills are driven into the assailant and released from
the porcupine’s skin, allowing the porcupine opportu-
nity to escape (Roze 2002).
On | September 2007, during a field survey to delin-
eate the distribution and assess habitat use of Bull
FicurE |. A Bull Trout (Salvelinus confluentus) collected
from an unnamed stream in the Mackenzie Moun-
tains, Northwest Territories, impaled by the quills of
a North American Porcupine (Erethizon dorsatum).
Trout in the Mackenzie Mountains, NWT, two male
Bull Trout were angled from a pool in an unnamed
tributary of the Redstone River (63°39'001"N,
125° 23'414"W). The larger fish (fork length 583 mm,
2022 g) had four Porcupine quills embedded in its
mouth. The quills were in the thin skin covering the
upper mandible (Figure 1), with no quills in the
esophagus or stomach, which suggested that the Bull
Trout had bitten a porcupine, rather than scavenged
off a carcass. Only fish remains were found in the
trout’s stomach. The size and colouration of the quills
were consistent with those from the side of a porcu-
438
pines’ tail, suggesting a defensive interaction (U. Roze,
Queens College, Flushing, New York, personal com-
munication 2007). For a Bull Trout to encounter a
porcupine, the latter would either have to be wading or
swimming across a stream, or less likely, floating dead
down a stream. Porcupines are known to swim or
otherwise traverse watercourses to access different
parts of their range (Murie 1926; Roze 1989). When a
porcupine swims, the majority of the body is below
water, with the head and some of the back and tail
visible above the surface, much like a Beaver (Castor
canadensis) (U. Roze, personal communication 2007).
Bull Trout often live in environments that have sparse
food resources and are, therefore, opportunistic preda-
tors (Stewart et al. 2007b). However, given a porcu-
pine’s size it is unlikely that it was a potential food
source. A more plausible explanation is that the Bull
Trout was defending its territory against the porcu-
pine which the fish interpreted as a threat. Bull Trout
are very aggressive, particularly during spawning, and
have attacked SCUBA divers during in situ spawning
surveys (J. Stewart, Fisheries and Oceans Canada,
Winnipeg, personal communication, 2007). The time
frame of the survey and temperature observed (6.0°C)
are both within the preferred ranges for spawning
Bull Trout in the NWT (Mochnacz et al. 2004) and
further south (Baxter and McPhail 1996).
This is the first record of a Bull Trout, or perhaps
any fish, attacking a North American Porcupine. Al-
though an interaction between these two species is cer-
tainly a rare event, the fact that it occurred is evidence
of the aggressive behavior of Bull Trout, and the
unusual and unforeseen linkages that can occur with-
in a given ecosystem.
Acknowledgments
We thank Jim Reist and Stephanie Backhouse, Fish-
eries and Oceans Canada, Winnipeg, Manitoba; Mur-
ray Somers, Fisheries and Oceans Canada, Yellow-
knife, Northwest Territories; Jesse Pierce and Guy
Thibault, Sahtu Helicopters, Norman Wells, NWT;
Richard Popko, Department of Environment and Nat-
ural Resources, Norman Wells, NWT. Uldis Roze,
THE CANADIAN FIELD-NATURALIST
Vol. 121
Queens College, Flushing, New York, and John Baba-
luk, Fisheries and Oceans, Winnipeg, Canada, who
reviewed earlier drafts; and two anonymous reviewers
for helpful, constructive critiques of this manuscript.
Literature Cited
Baxter, J. S., and J. D. McPhail. 1996. Bull Trout spawning
and rearing habitat requirements: summary of the litera-
ture. University of British Columbia, Department of Zool-
ogy, Fisheries Technical Circular (98). 27 pages.
Haas, G. R., and J. D. McPhail. 1991. Systematics and distri-
bution of Dolly Varden (Salvelinus malma) and Bull Trout
(Salvelinus confluentus) in North America. Canadian Jour-
nal of Fisheries and Aquatic Sciences 48: 2191-2211.
Mochnacz, N. J., J. D. Reist, P. Cott, G. Low, and R.
Wastle. 2004. Biological and habitat data for Bull Trout
(Salvelinus confluentus) and associated species from stream
surveys conducted in the southern and central Mackenzie
River Valley, Northwest Territories, 2000 to 2001. Canadian
Data Report of Fisheries and Aquatic Sciences (1131): iv
+ 38 pages.
Murie, O. J. 1926. The porcupine in northern Alaska. Journal
of Mammalogy 7: 109-113.
Reist, J. D., G. Low, J. D. Johnson, and D. McDowell.
2002. Range extension of the Bull Trout, Salvelinus con-
fluentus, to the central Northwest Territories, with notes on
char identification and distribution in the western Canadian
Arctic. Arctic 55: 70-76.
Roze, U. 1989. The North American Porcupine. Smithsonian
Institution Press, Washington, DC. 261 pages.
Roze, U. 2002. A facilitated release mechanism for quills of
the North American porcupine (Erethizon dorsatum). Jour-
nal of Mammalogy 83: 381-385.
Stewart, D. B., N. J. Mochnacz, C. D. Sawatzky, T. J.
Carmichael, and J. D. Reist. 2007a. Fish life history and
habitat use in the Northwest Territories: Bull Trout (Sal-
velinus confluentus). Canadian Manuscript Report of Fish-
eries and Aquatic Sciences (2801): vi + 46 pages.
Stewart, D. B., N. J. Mochnacz, C. D. Sawatzky, T. J.
Carmichael, and J. D. Reist. 2007b. Fish diets and food
webs in the Northwest Territories: Bull Trout (Salvelinus
confluentus). Canadian Manuscript Report of Fisheries
and Aquatic Sciences (2800): iv + 18 pages.
Received 12 December 2007
Accepted 29 July 2008
Book Reviews
Book Review Editor’s Note: We are continuing to use the current currency codes. Thus Canadian dollars are
CAD, United States dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars a
AUD and so on. You will find these are the codes now used by financial institutions and internet currency con-
verters. I will include an updated note for the next few issues as a reminder.
ZOOLOGY
The Ecology & Behavior of Amphibians
By Kentwood D. Wells. 2007. University of Chicago Press,
1427 East 60th Street, Chicago, Iinois 60637. 1148 pages.
75.00 USD.
This massive volume synthesizes knowledge gained
through researcher Wells’ extensive field experience and
total immersion in the literature focussed on habitats
and habits of a generous sampling from the approxi-
mately 6000 species of amphibians now recognized in
the world. Its extended gestation time began with con-
ception and partial drafts in the early 1980s. The period
since has been one of ever-increasing research and pub-
lication by a growing number of researchers resulting
in exponential additions to the number of species recog-
nized and, especially, the volume of information on
them. But simultaneously, it has been a time of realiza-
tion of the fragility of the world’s amphibian fauna
and the decline, sometimes the extinction, of many
species in many parts of the world ascribed to a vari-
ety of causes. Wells has been long-recognized as one
of the most productive and original researchers on am-
phibians of this period.
His book has 16 chapters, the first on phylogeny,
classification, and morphological evolution, followed
by ones on water relations, temperature relations, respi-
tation, metabolism and energetics, movements and ori-
entation, anuran vocal communication, mating systems
and sexual selection in anurans, communication and
social behavior of urodeles and caecilians, the natural
history of amphibian reproduction, parental care, ecol-
ogy and behavior of amphibian larvae, complex life
cycles and ecology of amphibian metamorphosis, am-
phibians and their predators, the ecology of amphib-
jan communities, and, finally, the conservation of am-
phibians. The reference section well reflects the vast-
ness of the information in the volume itself — the
two-column format occupies 238 pages, the length of
many books. Included are many, but by no means a
comprehensive selection of relevant papers by Cana-
dian herpetologists. Finally, a 64-page index concludes
the text. It is unfortunate that the initial chapter sum-
marizing classification was completed before Frost and
his colleagues (2006 American Museum of Natural His-
tory Bulletin 297) published their extensive reassess-
ment of amphibia relationships which proposed many
changes in nomenclature that could not be evaluated
for possible inclusion here. This is not a serious flaw
as the book’s overall emphasis is not on systematics.
Wells has deftly managed both a detailed and a read-
able reference and primer for study of amphibians in
nature. Its price and mass (approximately 11x8.5x2.2
inches or 286x220x60 mm) will doubtless keep it from
many a naturalist’s library shelves, except for the few
primarily focussed on amphibians who have ample dis-
posable income (perhaps not a common combination),
but it is a must for every university biology and federal
and provincial government environment department
and should be read by all biology students and con-
servation-charged public servants. Graduate students
focussed on amphibians or vanishing habitats in par-
ticular, will find this a treasure of the research to date,
and a gold mine of background information to inspire
future research projects.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P
6P4 Canada
The Amphibians and Reptiles of New York State: Identification, Natural History,
and Conservation
By James P, Gibbs, Alvin R. Breisch, Peter K. Ducey, Glenn
Johnson, John L. Behler, and Richard C. Bothner. 2007.
_ Oxford University Press, 198 Madison Avenue, New York,
New York 10016. 422 pages. 34.50 USD.
Lack of an authoritative field guide to the herpetol-
ogy of the state of New York has been a glaring vacu-
um in texts on the northeastern United States through-
|
|
:
out the last century. This, despite its relatively large
size for a northeastern state (49 108 square miles or
127 189 km?), and being the home of many excep-
tionally prominent herpetologists over the last centu-
ry. The state had been among the earliest to be given
a comprehensive scientific treatment when James
DeKay (for whom the snake Storeria dekayi is named)
440
included amphibians and reptiles in his Zoology of New
York in 1842. Almost 100 years later (in 1941) it
received the most through state monograph on a sin-
gle group, the Salamanders of New York by Sherman
Bishop, Rochester, New York. Bishop followed this
with a handbook to all North American salamanders.
Others included, in the first half of the 1900s, Ray-
mond Ditmars, New York, who wrote immensely pop-
ular and influential books on reptiles of North Ameri-
ca and of the world and, Albert and Anna Wright,
Ithaca, who used local studies to produce the first com-
prehensive data on the life histories of eastern North
American frogs and toads then went on to produce
North American handbooks to frogs and toads in 1933,
revised in 1942 and 1949, and to snakes in 1957.
The six authors who finally have filled the need for
a field guide are also New York state residents, scattered
among universities, conservation departments and
organizations: Gibbs at the State University of New
York College of Environmental Science, Syracuse;
Breisch at the New York State Department of Envi-
ronmental Conservation, Albany, Ducey at the State
University of New York at Cortland, Johnson at the
State University of New York at Potsdam, and Bothn-
er at St. Bonaventure University, Olean. The late John
L. Behler (1933-2006) was with the Wildlife Conser-
vation Society, Bronx. The distributional data are from
the Amphibians and Reptile Atlas Project of the New
York Department of Environmental Conservation
which compiled 59 000 reports submitted by 1800 vol-
unteers. The atlas also included 28 O00 pre-1990
records from museums, field notes, graduate theses, and
published literature.
The text covers 69 native species (18 salamanders, 14
frogs and toads, 17 turtles, 3 lizards and 17 snakes), and
includes 65 distribution maps (ranges shaded, but with-
out spot localities) and 72 colour plates (each with one
to four superior, sharp photographs of pattern and life
stage variants contributed by over 30 photographers).
Each species account is headed with English and Sci-
entific names followed by sections on Quick Identifi-
cation, Description, Habitat, Natural History, Status and
Distribution, and Other Intriguing Facts. Statements are
referenced to a 39-page Literature Cited section. Many
leading Canadian herpetologists are cited, among them
classical studies by J. P. Bogart on ambystomid sala-
manders and R. J. Brooks on Snapping and Wood tur-
Extreme Birds
By D. Couzens. 2008. Firefly Books Ltd., 66 Leek Crescent,
Richmond Hill ON L4B 1H1. 281 pages. 45 CAD Cloth.
When Extreme Birds arrived in the mail, it caught
my attention with its dramatic, somewhat comical,
cover photograph of a Shoebill. I started to nose
through, looking first at the remarkable photographs,
and wondering who might review this book. Then the
chapter titles made me take a second look and I was
THE CANADIAN FIELD-NATURALIST
Vol. 121
tles (both at University of Guelph), M. Berrill on pes-
ticide effects and declines (Trent University), J. Gilhen
on Nova Scotia species (Nova Scotia Museum), D. M.
Green on hybridization in toads (McGill University),
P. J. Weatherhead, K. A. Prior, and G. Blouin-Demers
on Rat Snakes (Carleton University, Environment Cana-
da. and Ottawa University, respectively).
But the text is not just species accounts. A brief intro-
duction summarizes the history of state herpetology
and stresses the conservation theme found through-
out the book. It is followed by a discussion of the state
herpetofauna with a table giving, for each species, its
New York Natural Heritage Program State Rank,
NatureServe Global Rank, and (where applicable) its
New York State and Federal Listings, exotic species
sometimes reported (alligators in the sewers of New
York City and beyond) and similarities and differences
between amphibians and reptiles. The is a chapter on
New York’s environment as habitat for amphibians and
reptiles with maps of ecological zones, of major lakes
and rivers, of counties and major cities, and “land
cover’. The ecological zones are described with rep-
resentative habitat photos and a table of all species and
habitat types they occur in. Final chapters are Threats
outlining the effects of historic overexploitation (par-
ticularly commercial harvest) and habitat loss over
time, acid rain, contaminants, disease and deformities.
Following chapters discuss legal protection, habitat con-
servation guidelines, conservation case studies, find-
ing and studying amphibians and reptiles, folklore,
and epilogue. Appendices cover the “Herp Atlas” Sur-
vey Card, and Resources (organizations and identifica-
tion and general reference). A Glossary covers terms
from amplexus to wetland, followed by Literature Cited
and a 20-page index.
Eastern Canadian naturalists will find this book a
must for their bookshelves as an excellent reference not
just to the species included (species that occur in east-
ern Canada are also in New York) but also for conser-
vation approaches and philosophy. It lacks only a sec-
tion on care of captives, and that, I suspect, is not an
oversight but deliberate, making a strong statement by
its omission.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario
K1P 6P4 Canada
hooked! I expected widest wingspan, smallest bird,
longest bill and so on, but heaviest testes, biggest belly
and smelliest bird took me by surprise. In the author’s
view the malodorous claimant is the Crested Auklet.
These cute beasts are said to emit a potent, tangerine-
like stink that carries for miles. I am afraid I missed that
pleasure last year when we sailed by their colonies;
there is an advantage or disadvantage to having a poor
sense of smell.
2007
This book not only covers the record individual char-
acteristics, like largest and smallest, but includes three
other categories. Extreme Ability covers such perform-
ance heights as fastest swimmer, best flock coordina-
tion and best mimic. Extreme behaviour describes the
strangest incubation, the loveliest display and the best
drummer. Finally Extreme families documents the lim-
its of courtship, promiscuity and breeding.
Many of the author’s choices will not surprise the
ardent birder. The widest wingspan belongs to the Wan-
dering Albatross. The biggest eyes [5 cm] are in the
head of an Ostrich and the smallest bird is a Bee Hum-
mingbird [5.7 cm]. So the litthke hummer could almost
squeeze into an ostrich eye. Although the choices are
obvious, the author tucks in many fascinating details.
The ostrich eye is as big as an eye can be, or diffraction
would cause problems. It would take 3000 Bee Hum-
ming bird eggs to equal the weight of one ostrich.
I did not realize there was a poisonous bird, the
Hooded Pitohui of New Guinea. Apparently this bird
absorbs batrachotoxin alkaloids from the Choresine
beetles it eats. This toxin is also found in some poison-
dart frogs and believed to come from their diet too.
Under the title “Classiest Colours” Couzens gives an
excellent discussion of the feather colour of a turaco.
This genus is the only one to make green pigment
[other “green” birds use refracted light to give the
impression of green] and the author covers the process
in simple and elegant language.
For the whitest bird he chooses an Ivory Gull, not
what came to my mind. I would say several egrets are
good contenders, but my choice would be the Snow
Petrel. Not only are these little birds bright white, but
they have a beautiful sheen that gives them a special
glow. Perhaps they did not “win” this category as the
Great Egret is cited for the best grooming aids [its
powder down and pectinated, comb toe] while the Snow
Petrel gets the “Most relaxed attitude to breeding.”
Cranes: A Natural History of a Bird in Crisis
By Janice M. Hughes. 2008. Firefly Books Ltd. 66 Leek
Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 256
pages. 45 CAD Cloth.
Firefly Books produces well-illustrated nature books
featuring larger format, glossy paper, and generally
readable text. Cranes are exciting birds, and they have
captured mankind’s imagination for millennia, so they
make good subjects for a book of this character. It is
written by Janice Hughes, an Associate Professor at
Lakehead University.
With a title like Cranes one might expect the subtitle
to refer to a “Family in Crisis” but in fact the “Bird”
referred to is the Whooping Crane, and despite the em-
phasis on the dust jacket on cranes as a group, two-
thirds of the text is devoted to the history of this one
species. Chapters one and six cover the family as a
whole, the first outlining their natural history, and the
BooK REVIEWS
44)
However the Barn Swallow gets two citations as “Sex
lest tail” and Canniest false alarm.”
The author often chooses one species, but the same
remarks can be made of others in the genus. The longest
legs are not confined to the Lesser Flamingo but are
characteristic of all flamingos. Similarly at least the
Variable Pitohui and the Brown Pitohui [there are six
in the genus] are poisonous as well as the Hooded Pito-
hui. Indeed Blue-capped Ifritas carry the same toxin.
The photos, one for each species, come from pro-
fessional stock companies. Therefore they are superb.
It is futile to try and pick the best ones, but those that
caught my attention were Lesser Spotted Eagle [so
majestic], Clarke’s Nutcracker [so homely], Western
Capercaillie [so haughty] and the Barn Owl [such a
beautiful bird]. The photo of the Red-crowned Crane
is SO artistic it could be a Japanese woodcut by Hokusai
or Hiroshige.
I usually read bird books to learn something. While
this is a pleasurable act, the primary focus is not amuse-
ment, but information. I read this book because it is
fun. Even when I knew the facts, such as the Emperor
Penguin’s unbelievable fast, 1 enjoyed this author’s
version of the tale. It was even better when I was learn-
ing something new, like the Great Crested Grebe’s pas-
sion for eating its own feathers. So if you want to know
the biggest, the fastest, the smallest, best nest builders,
weirdest matings and who dives the deepest or flies
the highest, then this is the book for you. Even non-
birder trivia fans will get great enjoyment.
Oh yes, and the heaviest testes [8% of its body mass
and heavier than its brain] record is held by the Alpine
Accentor. For good reason too, but you really should
read the book to find out why! And then there is the
bird that f*rts to catch its food — but that is another
story.
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada
second profiles of the 15 species. Perhaps appropri-
ately, the natural history section includes extensive coy-
erage of the species’ relationship to man, starting with
a section on mythology, folklore and cultural impor-
tance, and concluding with demographics and threats,
and crane conservation. In chapter 6 each species is
assigned two pages in a smaller type-face, including
a range map and a rather small [~3x5 inch] picture.
Chapters 2 through 5 chronicle, first the decline of
the Whooping Crane, followed by accounts of its recoy-
ery, discussing in turn the bird’s population and migra-
tion, and concluding with a summing-up. Because the
threats to cranes are mentioned in a number of sections,
there tends to be some repetition of material. Two
appendices provide the World Conservation Union rat-
ings of crane status, and wild Whooping Crane peak
winter counts from 1938-1971.
442
I was rather disappointed by the book, mainly
because the Whooping Crane sections are so domi-
nant in it that I had a feeling the rest of the species
were almost an after-thought. The family is a rather
homogeneous one, so treating the natural history of
the birds as a group is reasonable, but the two-page
summaries are just that — summaries — and the illus-
trations in them are inadequate given the scope and
size of the book. In fact, the pictures in general are
not up to the standard one expects from a book of
this type. Many of the ones in my copy are a little
off, lacking sharpness, and some are no more than
cropped versions of pictures that appear earlier.
The text is rather uneven. Some parts are rather plod-
ding. The author lapses periodically into jargon and
then, for those who might not understand, pops in a
THE CANADIAN FIELD-NATURALIST
Vol. 121
brief explanation. Why use the term in the first place?
At its best, however, it is well-written, even eloquent,
especially in sections discussing the plight of cranes.
The Whooping Crane history is given in considerable
detail, even to the extent of outlining the events lead-
ing to the formation of the Audubon movement.
Those seeking a comprehensive treatment of the
world’s cranes will find this book wanting. If you are
interested in the fascinating account of the collapse
and gradual recovery of the Whooping Crane in North
America, coupled with an overview of cranes gener-
ally, then this could be a good book to acquire.
CLIVE E. GOODWIN
1 Queen Street, Suite 405, Cobourg, Ontario K9A 1M8
Canada
Marine Mammals of the Pacific Northwest: A Concise and Comprehensive Waterproof Guide
By P. A. Folkens. 2001. Harbour Publishing, P.O. Box 219,
Madeira Park, British Columbia VON 2HO. 8 pages. 9.95
CAD -— (synthetic film).
The increasing development of the whale-watching
industry has resulted in the demand for, and creation
of, books and identification guides for sea mammals,
seabirds and the marine ecosystem. In this regard, and
looking back on a history of whale-watching for over
90 years, the coast of British Columbia has received
most of the publication activity in Canada.
Despite its catchy title, this guide by Pieter Arend
Folkens is more a leaflet of a guide. It consists of three
text pages and five pages of drawings and photos full
of information for the whale watcher in the field.
Although the leaflet is printed on “waterproof, UV re-
sistant synthetic film made from a 100% recyclable,
environmentally inert material containing no forest
products (similar to milk jugs)”, the user might actu-
ally have difficulties using it on an offshore whale
watching trip during periods of stronger wind or
higher waves; the light leaflet could easily fly away
and the small print is hard to read when on a rolling
boat. However, the compressed text gives a nice sum-
mary and overview on 31 sea mammal species in the
area; it even mentions Steller’s Sea Cow, which was
hunted to extinction by 1768. In addition, major
whale watching locations in British Columbia and
Alaska are named, but none are reported really for
Oregon and Washington (as the title would imply).
Most of the eight pages of the field guide are devoted
to drawings and to fine pictures from the author and
several others. The reader might find the distinction
between Mysticetes (Baleen Whales), Odontocetes
(Toothed Whales) and Small Cetaceans a little unclear
from the arrangements of the drawings. Very helpful
and informative is the page about “Common visible
behaviours and terms” allowing to link sea mammal
sightings to a classified set of behaviour types. Help-
ful also is that images of fluke displays are presented
for species that are known to show such a behaviour.
Even the body sizes of newborn sea mammals are
given. All measurements are made in SI units, and the
conversion factor for feet is provided.
Of interest to the general audience might be the
section “Marine Mammal Watching Guidelines”,
also presented on the web http://www.fakr.noaa.gov/
protectedresources/mmviewingguide.html (Note that
the old URL www.nmfs.gov/prot_res.html and given
in the guide was updated). In addition, contact address-
es and a web address (revised to http://www.fakr.
noaa.gov/protectedresources/strandings.htm) are given
for sightings of stranded sea mammals (Department
of Fisheries and Oceans Canada, 800 4654336; or the
Whale Reporting & Stranding Line 800 665 5939).
As found in many other field guides, the text sug-
gests some field marks and details for species identi-
fication and separation that normal whale watchers
might not be able to apply, or which are not really real-
istic. For instance, Sei Whales are supposed to be dif-
ferentiated from Fin Whales by a fin angle of over 45
degrees; male Beaked Whales (genus Mesoplodon)
ideally can be identified by the location of teeth and jaw
line (which is for most of the time covered by the
ocean); phocids (true seals) differ from otariids (Sea
Lions and Fur Seals) by their hair and small nails on
their foreflippers. Overall, features like these might be
very hard to recognize for the untrained as well as for
the trained observer, particularly when observations are
done for moving animals, from a shaky boat far away
and with binoculars. Rather than focussing on classi-
cal small-scale features, outlining the use of propor-
tions and shapes could be more useful for telling
species apart. Counterproductive for a field guide might
be the point that Beaked Whales, the species group
that lacks most knowledge on distribution and where
whale watchers could indeed contribute greatly to
science, are described as the “most difficult whales to
identify correctly”; no further help or details are given
for the interested whale watcher. For my taste, the
“Habitat and Symbol Keys” that are supposed “to
2007
narrow possibilities in a particular area” and link sea
mammals with “habitat” are not really helpful since
many whales migrate across habitats anyway and
since the regular observer has no real way to tell
“temperate” habitats apart from “cool temperate”
ones. The meaning of the orange W habitat class pre-
sented for the False Killer Whale will likely remain a
mystery to the reader because its meaning is nowhere
explained in the guide. For pinnipeds, their “calls” and
the mentioning of rookeries could have been helpful.
Reptiles and Amphibians of Canada
By Chris Fisher, Amanda Joynt, and Ronald J. Brooks. 2007.
Lone Pine Publishing, 206, 10426 — 81 Avenue, Edmon-
ton, Alberta T6E 1X5. 208 pages. 18.50 CAD.
It has been 24 years since the first comprehensive
one-volume guide to all Canadian amphibians and
reptiles known at the time appeared (Cook 1984) and
that is now badly out-of-date and, fortunately, out-of-
print. There have been new guides to several provinces
since and a highly popularized superficial attempt for
the entire country by Bumstead (2003).
For this new effort, only the third author, the leg-
endary Ron Brooks of the University of Guelph will be
very familiar to most Canadian herpetologists. Brooks
has long been a CITES committee member and crusad-
er and has made an extensive ecological contribution
with a legion of graduate students conducting studies
at Algonquin Park and selected central and southern
Ontario sites. These have produced new insights into
the lives of turtles and aquatic frogs in eastern Canada
and one especially endangered snake (the Blue Racer).
The other two authors have BScs from the University
of Alberta. Chris Fisher is a writer of wildlife articles
and field guides and lecturer on wildlife. Amanda Joynt,
an ecologist from the Okanagan Valley, was a techni-
cian with Parks Canada and Canadian Wildlife Service,
followed by writing full time for Lone Pine publish-
ing, ecological surveys including rare plants in South
Dakota, and direction (2004-2006) of Children in Wild-
erness Malawi, southern Africa. She now is a biologist
for Fisheries and Oceans Canada in Inuvik, North-
west Territories.
The book is traditionally organized and opens with
the mandatory acknowledgements, including one to
Ron Brooks, odd in that he is also a coauthor. There is
a very useful “Species at a glance” which gives a
miniature reproduction of the text illustration, species
by species, together with size, and account page num-
ber. The 12-page Introduction briefly highlights the
antiquity of the groups and their characteristics. A map
plots the location of selected national and provincial
parks in Canada. This is followed by a summation of
major habitats from the temperate west coast to the
Maritimes, a discussion titled “the good, the bad and
the misconceptions”, and another on the general harm-
lessness of most species as well as the negative aspects
of keeping native species (not the least of which is that
BOOK REVIEWS
443
Overall, it appears that this “guide” is an excerpt of
a better and larger guide book from the same author
It is useable in the field, but does not replace the real
and classical guide books
FALK HUETTMANN
Centre for Wildlife Ecology, Biology Department, Simon
Fraser University, 8888 University Drive, Burnaby, British
Columbia VS5A 186 Canada
in most provinces, is illegal to keep many species
except under permit from resource departments). A few
words on conservation are followed by the headings
used in the species accounts. The latter are the bulk
of the book (150 pages) and cover both native and
introduced species: 11 freshwater or terrestrial turtles
and 4 marine ones, 7 lizards, 26 snakes, 21 salaman-
ders, and 25 frogs (including toads and treefrogs, etc.).
Three of the turtles and are introduced or likely so, and
two of these probably no longer occur, one lizard is in-
troduced and one apparently extirpated, and one snake
(Timber Rattlesnake) is extirpated.
Each species account is dominated by an enlarged
colour drawing of an adult, and these vary from very
good to embarrassingly bad. Some of the snakes, turtles
and frogs are among the excellent, some salamanders
and virtually all the lizards are poor, the latter particu-
larly washed-out, among other faults. The Common
Garter Snake represented seems to be a particularly odd
colour pattern (a Pacific region variant?) that will be un-
recognized in most of the country. The Plains Garter
Snake is very pale, typical of the southern and eastern
portions of the range mostly beyond Canada, while the
“Eastern Ribbon Snake” on the next page matches the
majority of Canadian specimens of plains species. In
Canada, Ribbon Snakes rarely have an orange dorsal
stripe (I know of only one report) like the one pictured:
the lateral stripe is usually prominently bordered below
by chocolate, and overall it is more slender. Among
other snakes particularly poorly done are the Red-bel-
lied, Brown, and Green snakes. The species accounts
themselves lead with English and scientific names fol-
lowed by a casual informal introduction, then a para-
graph ID (identification), and one or two lines on
Length, Distribution (with a tiny map of Canadian
range), Habitat, Activity Patterns, Reproduction, Food,
Similar Species, French Name, and a Did You Know
sidebar, the latter focussed on some additional fact re-
garded as particularly remarkable. Each native and ex-
isting species is given two facing pages. The marine tur-
tles (designated “vagrant species”) fare even less well,
with only a half page each, and the introduced or extir-
pated ones only somewhat better at one page each. The
format of necessity means that only the briefest info-
rmation can be given for each. Sometimes these have
little Canadian relevance. An example is the statement
444
that the Small-mouthed Salamander “often” shares
breeding ponds with Spotted Salamanders. Presumably
this applies to United States populations as in Canada
the Small-mouthed Salamander has been recorded only
on Pelee Island where Spotted Salamanders apparently
do not occur. But omitted from the Small-mouthed Sal-
amander account is mention of its extensive hybrid-
ization with Blue-spotted Salamanders on Pelee Island
although elsewhere the hybridization of Blue-spotted
with Jefferson salamanders is recognized.
On the positive side, this book does have all the
species we know in Canada to date with distributions,
many interesting highlights, and very colourful pictures.
These will be found useful by some casually interest-
THE CANADIAN FIELD-NATURALIST
Vol. 121
ed uncritical naturalists. However, it is no substitute
for the many carefully researched and more compre-
hensive, within their more limited areas covered, pro-
vincial guides available for amphibians and reptiles.
Literature Cited
Bumstead, Pat E. 2003. Canadian skin and scales: A complete
encyclopedia of Canadian amphibians and reptiles. Simply Wild
Publications Inc., Calgary. Alberta. 161 pages.
Cook, F. R. 1984. Introduction to Canadian amphibians and reptiles.
National Museum of Natural Sciences, National Museums of Cana-
da. 200 pages.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario
K1P 6P4 Canada
Guides and Reference to the (1) Snakes, (2) Crocodilians, Turtles, and Lizards (3)
Amphibians, of Eastern and Central North America (North of Mexico)
By R. C. Bartlett and Patricia P. Bartlett. 2005 (1), 2006(2&3).
University Press of Florida, 15 NW 15th Street, Gainesville,
Florida 32611. 342 (1), 316 (2), 283 (3) pages. 29.95 USD
each.
The Bartlett team has produced a number of earlier
books: A Field Guide to Florida Reptiles, Reptiles
and Amphibians of the Amazon: An Ecotourists Guide
(2003), and Florida Snakes: A Guide to Their Identifi-
cation and Habits (2003). R. D. Bartlett, veteran her-
petoculturist/herpetologist, founder of the Reptilian
Breeding and Research Institute, boasts over 40 years
writing and photographing, including more than 500
articles in magazines such as The Tropical Fish Hobby-
ist, Reptiles, and Reptile and Amphibian. Patricia Bart-
lett, former director of the Fort Myers Historical Muse-
um, is an editor and writer who, in addition to
coauthoring 12 books with R.D.B., also wrote A Dic-
tionary of Sharks.
In these present volumes, the Bartletts have broad-
ened their horizons and compiled a herpetofauna ref-
erence in three volumes which cover the entire east-
ern and central United States and adjacent Canada
from the Atlantic to the tier of mid-western states
from North Dakota to Texas. However, their southern
bias persists. Species which range far north of the
U.S. border have only the southern portion of their
Canadian range shown on the generally crude distri-
bution maps which only casually outline Canada,
with the northern potions of the Canadian-based
range are often cut off. No Canadian-based authors
are included in the references, despite the many that
have studied species included, some even from U.S.
populations as well as Canadian material.
Each book has a brief Species List, a Preface and
Introduction to the group(s) included. The bulk of the
texts are accounts arranged by family and within these
by related groups of species, and then individual species
accounts. The later includes English and Scientific
name followed by sections on Toxicity/Disposition
(Snakes), Abundance/Range, Habitat, Size, Identifying
features, Voice (in crocodilians and frogs), and Similar
Species, Comments, and Additional Subspecies. There
is a rather eclectic selection here sometimes of sub-
species far to the west of the limits set for the book.
There is a distribution map (see comment above), and
one or more colour photographs for each species (re-
grettably without localities or date), but usually of spec-
tacular quality, and it here that the book shines, and wins
a place on many shelves.
But why, aside from this, would any Canadian nat-
uralist want these books on their shelf? They do include
all the species in eastern Canada, as all of these range
north into this country from the United States. These
accounts are of reference value for depictions of appear-
ance (many species pictured are different from the
Canadian variants) and natural history (contrast from
more southern habitats) of these species in areas adja-
cent to Canada and beyond.
Attractive as it is, it is primarily an add-on for Cana-
dian naturalists unless they are travelling south, and
for practical purposes there are now many Canadian
and adjacent state guides to reptiles and amphibians
combined in single volumes that will be more useful
references for identifying our herpetofauna.
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4
Canada
2007
BOTANY
Ginkgo in China
By Cao Fuliang. 2007. China Forestry Publishing House,
Number 7, Liuhai Hutong, Denei Street, Xicheng Dis-
trict, Beijing, China. 300 pages. 98 CNY.
Ginkgo is a genus of highly unique plants with one
extant species, Ginkgo biloba, which is a long-lived,
stress-tolerant, disease-, pest-, and fire-resistant dioe-
cious tree, referred to as a living fossil by Charles Dar-
win in 1859. The only surviving member of its family,
Ginkgo leaf was first mentioned in Lan Mao’s book
Pharmaceutical Natural History of Southern Yunnan
(1436) in the Ming dynasty of China. Engelbert
Kaempfer (1651-1716), a German botanist, firstly gave
a scientific description of this species in his book
Amoenitatum exoticarum (1712), and formly gave it
its name according to a phonetic pronunciation of a
Chinese or Japanese word.
It was reported that fossils recognizably related to
modern Ginkgo biloba date back to the Permian, some
270 million years ago. The genus diversified and
spread throughout Laurasia during the middle Jurassic
and Cretaceous, but became much rarer thereafter. At
the end of the Pliocene, Ginkgo fossils disappeared
from the fossil record everywhere apart from a small
area of central China where the modern species sur-
vived. Chinese people, especially the monks in history
formally, have long cultivated Ginkgo, thereby con-
tributing to its survival as a species. All the present
Ginkgo trees originated from China and were intro-
duced into Europe in the 1700s, and then spread to
other continents.
Ginkgo not only has scientific value in terms of re-
search on its evolution and genetic tenacity, but also its
medicinal and ornamental applications. Ginkgo has
been used in medicinal botanicals in China for more
than 2000 years. Nowadays, more and more medicinal
functions of its leaves and seeds are being extensively
revealed by scientists all over the world, with a focus
on the standardized Ginkgo biloba extract (GBE) being
prepared from the dried green leaves. At present, Gink-
go has become one of the most common trees along the
sidewalks in many cities of the world due to its urban
tolerant traits and the beautiful shape and colour.
China is the hometown of modern Ginkgo trees. The
small-sized natural Ginkgo populations were exten-
sively distributed over different geographical areas of
China. The cultivation area of Ginkgo covers wider re-
BOOK REVIEWS
445
gions. The germplasm resource of Ginkgo is abundant
Ancient Chinese carried out preliminarily studies or
observations on the trees. However, afterwards, there
were not too many new studies in China due to the un-
stable or abnormal social and political situation. Since
the 1980s, Chinese scientists have carried out a series
of comprehensive studies on the trees. The book Gink-
go in China is a systematic summation of these studies.
As well, the book collected and analysed other abun-
dant information on the Ginkgo at home and abroad.
The book covers almost every aspect of the Ginkgo,
such as its origin, evolution and taxonomy, history of
cultivation and utilization, natural populations and
cultivation areas, biological and ecological character-
istics, cultivation techniques, exploitation and use of
Ginkgo resources, genetics and breeding, and so on.
Chapter one, is an outline of the Ginkgo’s name, and
its economic, ecological, social, cultural, scientific
values, developmental status, and prospects; Chapter
two covers the origin, evolution and taxonomy of the
Ginkgo; Chapter three, the history of cultivation and
utilization of the Ginkgo at home and abroad; Chap-
ter four, the natural populations and cultivation regions
of the Ginkgo; Chapter five: the biological characteris-
tics of ginkgo; Chapter six, the cultivation techniques
of ginkgo; Chapter seven, the comprehensive exploita-
tion and utilization of Ginkgo resources; Chapter eight,
the germplasm and breeding of the Ginkgo; Chapter
nine, reputed varieties, strains or individuals of Gink-
go in China; Chapter 10, key areas of Ginkgo produc-
tion; and Chapter 11, the development of the culture
of the Ginkgo.
The book is well written with few errors, and can be
easily understood by non-scientists. As a comprehen-
sive book, it would be suitable for anyone who is en-
gaged in various areas of research or application in
relation to ginkgo. Hopefully, it would become a valu-
able reference for them.
Li Dezut' and Qin AIL
'Labratory of Urbanization and Ecological Restoration of
Shanghai; National Field Observation and Research Sta-
tion in Tiantong Forest Ecosystem of Zhejiang: Depart-
ment of Environmental Science, East China Normal Uni-
versity, 3663, Zhongshan Road (N), Shanghai 200062
China.
“Jilin Forestry Staff School, Jilin, China 130000
Comprehensive Monograph of Contemporary Medicinal Plants (Volumes I-IV)
By Zhao Zhongzhen, Xiao Peigen. World Book Publishing
House, Number 137, Chaonei Street, Beijing, China.
2008. 4 volumes: total 2180 pages. 368 CNY Cloth.
In a broader sense, the medicinal plants referred to
are not only the so-called traditional medicinal plants
used for prevention and treatment of human diseases,
but also the plant resources used as nutrients, hobby
materials, spices, coloring additives, pesticides and
veterinary drugs and so on. The whole or parts of the
medicinal plants can be used directly in treatment or
as raw material in the pharmaceutical industry.
China is one of the countries with abundant resources
of medicinal plants, and with the long history of use
and cultivation. The discovery of medicinal plants was
446
the result of accumulation of experience and knowledge
in the long-term production and life in ancient times.
In China, literary records on medicinal plants might be
traced to the Spring and Autumn Era (770 BC~221BC)
or much earlier. Till the Ming Dynasty, the world wide
famous comprehensive medicinal book Compendium
of Materia Medica (Li Shizhen 1578), contained more
than 1200 species of the medicinal plants.
Generally speaking, Chinese traditional medicine
as a cultural treasure of China, together with Western
medicine, plays an important role in human health
care, and is the commonwealth of mankind. Howev-
er, not all records in the traditional medicinal litera-
tures were correct, and some of them were proved to be
wrong or inaccurate with long-term clinical practices
or scientific tests. Furthermore, some new pharmaco-
logical effects of these medicinal plants have been
revealed, and some new medicinal plants have been
gradually added to our knowledge. Therefore, the sci-
ence of medicinal plants as an ancient and developing
science needs systematic testing of the records in the
traditional medicinal literature, and also needs a con-
tinual infusion of new knowledge or scientific find-
ings. Actually modern medicine, or Western medi-
cine, as a whole, has persistently absorbed a great deal
of valuable experience and knowledge accumulated
in the course of long-term clinical application and sci-
entific research. However, the science of medicinal
plants itself has been in the shade, and not summed up
the progress of time.
The Comprehensive Monograph of Contemporary
Medicinal Plants is divided into three parts and four
volumes. The Chinese and English versions were pub-
lished successively. The book collected information on
more than 800 frequently used medicinal plant species
from both China and abroad. Each entry of medicinal
plant species includes Chinese, Latin and English
names, main geographical distribution region, main
morphological characteristics, medicinally effective
THE CANADIAN FIELD-NATURALIST
Vol. 121
parts, chemical composition, pharmacological effects
and the progress in clinical applications. Abundant dia-
grams are used to supplement the explanations. The
book adopted the definition of the medicinal plant
species in broader sense, to include not only the tradi-
tional medicinal plant species being used for a long
time both in China and abroad, but also the new prod-
ucts made from the medicinal plants, natural health
plant products, natural plant cosmetics, natural plant
pigment, and so on.
The book reflects the results of the authors’ in-depth
investigations on these medicinal plant species for many
years. On the basis of systematic compilation and
analysis of the illustrations of the efficacies of these
medicinal plants in the voluminous traditional medici-
nal literatures, the book introduced the latest medical
progress in plant biochemistry, pharmacological, toxi-
cological and clinical research. For each medicinal plant
species mentioned, the authors also give pertinent com-
ments and suggestions on sustainable exploitation
and utilization, as well as safety in clinical applica-
tions. All these might contribute greatly to the clarifi-
cation of the mechanism of the role of the traditional
medicine plants, to the modernization of the science of
traditional medicinal plants, and to guidance for future
clinical applications.
The book is well written with few errors. Abun-
dant illustrations help readers understand the expla-
nations. The book is suitable for professionals who
engage in plant science, pharmaceutical research, pro-
duction, development, testing as well as sales staff or
other persons who are interested in these fields.
Li Dezut! and Qin AILP
'Labratory of Urbanization and Ecological Restoration of
Shanghai; National Field Observation and Research Station
in Tiantong Forest Ecosystem of Zhejiang; Department of
Environmental Science, East China Normal University,
3663, Zhongshan Road (N). Shanghai, 200062 China
“Jilin Forestry Staff School, Jilin, China 130000
Book Review Editor’s note: Continuing with our decision to investigate suitable websites and, if appropriate,
include their reviews. I have written the following review. If others know of similar suitable sites and are moved
to submit a review please contact me at r.john@rogers.com.
MushroomExpert.Com website
By Michael Kuo. 2008. http://www.mushroomexpert.com/
index.html.
I recently wanted to identify a mushroom in my yard
and, with help, thought it might be a Turkey Tail. On the
web I found MushroomExpert.Com, a site developed
by an Illinois English teacher and amateur mycologist,
Michael Kuo. He is also the author of two convention-
al books; Morels (2005), and 100 Edible Mushrooms
(2007) [University of Michigan Press]. By following
Dr. Kuo’s keys and his “Totally True Turkey Tail Test”
I confirmed I had a totally true Turkey Tail.
I was very impressed by the individual pages on each
species — the field guide section. This section current-
ly covers 700+ mushroom species using identification
keys. Each species has a web page with identification
information, background material, habitat and some
excellent photographs. Some of the photos are by the
author, but there are many other contributors. The text
is clear and easy to follow and shows a certain sense
of humour. I smiled at phrases like “...let me say this as
plainly as I can: You are stupid if you eat this mush-
room.” It certainly gets to the point! There are web links
to other mushroom sites, so you can see additional pho-
tos and read other versions of the species’ characteris-
tics [these were not as attractively displayed nor as
well illustrated as Kuo’s site].
2007
There are three ways to search. You can step your
way through the key system by clicking “Identification
keys.” For, say, Boletus luridus this is a four-step
process and leads to Kuo’s page [four photos]. For
Boletus flammans you get moved to RogersMushroom
with only one photo.
The second way to get to a species is to select a sci-
entific name from a drop-down box. If you choose
Boletus bicolor you get to Kuo’s page. Oddly, if you go
through the keys and decide this is your species there
is no link to any further page.
The third method is to go to a search box and type in
turkey tail and this will take you out to Google with a
link back to MushroomExpert! This will also happen if
you enter Boletus flammans, and the link will be the
page with the link to RogersMushrooms. This all seems
a little odd, but it does work. I think this is a reflection
on the incomplete status of the site. The author, for ex-
ample, states “We have not yet completed a key to North
American polypores, but we have started the ball rolling
with a key to the pale-fleshed, stemmed polypores”.
I explored the site further and found it covers Col-
lecting for Study, Making Spore Prints, Descriptions
& Journals, Identifying Mushrooms, Determining Odor
and Taste, Pronouncing Latin Names, Testing Chemi-
BooK REVIEWS
447
cal Reactions, Preserving Specimens, Using a Micro-
scope, Mushroom Taxonomy, Introduction to Mush-
room Photography, Digital Photography Tips, Scanning
Mushrooms, Eating Wild Mushrooms, Mushroom Tox-
ins, Meixner Test for Amatoxins.
I read a few of these sections and found them in-
formative and remarkably well done. Where required
they have excellent illustrations or photographs,
including step-by-step sequences. If you intend not
to heed the author’s warning in paragraph two above,
then read the section on toxins. Under Amatoxin he
states , in part, “But the remission is a cruel hoax; in
the meantime, the victim’s liver and kidneys are
being destroyed.” A very unpleasant thought.
Kuo is currently writing, with Andrew Methven, /00
Cool Mushrooms, [University of Michigan Press] and
Mushrooms of the Midwest (University of Illinois
Press. |
I have often been frustratingly confused by mush-
room field guides. This is the first time I have actually
enjoyed the process of resolving identification issues.
This site is now in my list of “favourites” — a rare
event as I only have 20 or so favourites.
Roy JOHN
2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada
Between Earth and Sky: Our Intimate Connections to Trees
By Nalini M. Nadkarni. 2008. University of California Press,
2120 Berkeley Way, Berkeley, California 94704-1012.
322 pages. 24.95 USD Cloth.
Nalini Nadkarni, forest ecologist and author of
Between Earth and Sky, has written a natural and social
history of trees, a tribute to the strong and influential
global inhabitants she dearly loves. As she writes in
the introduction to Between Earth and Sky, “1 love trees:
how they look, how they behave, how they smell and
sound, and how I feel when I am around them.” The
first dedication of her book is to the “maple trees out-
side the front door, where this book began.”
Nine chapters exploring the relationship between
humans and trees cover topics as diverse as “what is a
tree”; tree goods and services; trees as shelter and pro-
tection; trees and health and healing; trees in play and
the human imagination; trees and the expression of
time; trees as signs and symbols; trees in spirituality and
education; and trees and mindfulness. Woven through-
out the chapters is scientific fact, personal reflection
and narrative, and a variety of visual material.
It is a compelling combination. I found the generous
sprinkling of tree quotes, poems and other passages
particularly intriguing. Passages from Shakespeare's
Othello, Homer’s The Odyssey, and a Woodie Guthrie
song. Poems by Wendell Berry, William Blake, John
Clare, E. E. Cummings, Jane Hirschfield, Pablo Neru-
da, Li Po, Rumi, Christina Georgina Rossetti, Percy
Bysshe Shelley, and Rabindranath Tagore. Quotes by
Hermann Hesse, Federico Garcia Lorca, and Antoine
de Saint-Exupéry. One of my favourites is a line from
The Power of Myth by Joseph Campbell and Bill Moy-
ers: “God is the experience of looking at a tree and
saying, “Ah!”
Creative writings aside, the book contains no short-
age of scientific and other facts. I discovered “arboreal
soil” situated on large branches in certain forests, spaces
that epiphytes, insects, earthworms, and spiders call
home. I learned about “witness trees,” which bear per-
manent marks carved by settlers of forested lands to
marks territorial boundaries. I was interested to read
that the sacred architecture of Greek temples and
Gothic cathedrals is thought to be inspired by sacred
groves, and that Hindu and Buddhist temples are in fact
associated with living trees.
Anyone who loves trees, who admires their stead-
fastness, who finds in their presence a certain peace and
calm, will love this book. As Nadkarni writes in the
introductory chapter, “When I place my own strong
brown hand on the trunk of a tree, I feel connected to
something that deserves my curiosity, care, and pro-
tection.” Reading Between Earth and Sky makes you
want to rush outside and follow her example.
RENATE SANDER-REGIER
3, 11th Line, Bristol, Quebec JOX 1G0 Canada
448
MISCELLANEOUS
THE CANADIAN FIELD-NATURALIST
Vol. 121
Contributions to the History of Herpetology Volume 2
Edited by Kraig Adler. Society for the Study of Amphibians
and Reptiles, Post Office Box 58517 Salt Lake City,
Utah 84158-0517 USA. 396 pages. 65.00 USD Cloth.
This is volume 21 in the Society for the Study of
Amphibians and Reptiles series Contributions to
Herpetology begun in 1980. The first history contri-
bution, number 5 in the series, was published in 1989 to
commemorate the First World Congress of Herpetol-
ogy held in Canterbury, England, that year. It contained
biographies of 152 “Herpetologists of the Past” com-
piled by Alder using two criteria for inclusion: (1) sig-
nificant contribution to herpetology and (2) deceased.
Its scope was world-wide and included references for
each biography, signatures, and 149 portraits. The book
concludes with index to 2500 authors in taxonomic her-
petology compiled by John S. Applegarth, and academic
lineages of 1300 herpetologists by Robert Altig. It was
202 pages and priced at $20. It is now out of print but it
has been put on the web for public access at www.ssar-
herps.org (click on “Publications”). The new volume
extends the coverage of the first and has biographies of
285 additional Past Herpetologists (with signatures and
269 portraits) by Adler. Following sections are an index
to 3604 taxonomic authors (again by Applegarth),
and academic lineages of 3810 herpetologists (again
authored by Altig). The price, in keeping with the
times, has more than tripled. Like the first, this is also a
commemorative volume, issued for the Society’s 50th
Anniversary Meeting, St. Louis, 2007.
The new biographies are comprised of both long-
deceased herpetologists omitted in the earlier edition
and recent ones who have died since the first was writ-
ten. The accounts are grouped by 26 variable time
periods from 1400-1500 to 1970-1990 with placement
within each period based on when the person first be-
came prominent in herpetology. Fortunately, there is
an index (pages 259-273) covering both volumes that
includes page numbers not only for the 437 subjects
of individual biographies but also every mention of
these and other herpetologists. The bibliographies in
volume, 2 begin with Leoniceno (1428-1524) and con-
clude with Slowinski (1962-2001). The most recent
additions are John L. Behler (1943-2006) and Mar-
garet M. Stewart (1927-2006). Canadians, represent-
ed by E. B. S. Logier (1893-1979) in the initial vol-
ume, have one more entry in the second, Stanley W.
Gorham (1917-1984). Stan’s career was spent at the
National Museum of Canada (now Canadian Muse-
um of Nature) and the New Brunswick Museum and
he authored, among other contributions, a checklist
of amphibians of the world (1974). Its value is recog-
nized here: “Even when a larger, more comprehen-
sive checklist appeared in 1985, authored by a team
of 59 specialists from 20 countries, Gorham’s single-
authored, simpler list still had the advantage of includ-
ing synonyms and subspecies and remained a useful
reference for many years”. [See also 1991 tribute in
Canadian Field-Naturalist 105: 592-596]. Adler con-
tinues to augment his files and there will be many
more Canadians represented in a future volume 3 and
beyond when the those so active from 1960 to the
present are finally eligible for inclusion.
Interest in all corners of the history of biology has
come of age and editor Adler is well suited to the task
of documenting the herpetological research compo-
nent. He is Professor of Biology and former Vice Pro-
vost for Life Sciences at Cornell University, Ithaca,
New York. He has been President of the SSAR and was
founding Secretary-General of the World Congress of
Herpetology. He has published extensively on behav-
iour and systematics of amphibians and reptiles in sci-
entific journals and books. He has been, and remains,
a driving force in the hyperactive Society for the Study
of Amphibians and Reptiles since its foetal days as
the Ohio Herpetological Society and the publication of
its first journal (1959-1966). He has promoted and edit-
ed both the Facsimile Reprints in Herpetology series
(41 ranging from pamphlets to books) and the Contri-
butions series as well as being a supporter of varied
other publications of the Society: The Catalogue of
Amphibians and Reptiles (840 accounts to date), Her-
petological Conservation (2 volumes dealing with
Canadian amphibians (1) 1997 and reptiles (2) 2007),
The Herpetological Review (1967-, at volume 37 in
2006), Journal of Herpetology (1968-, at volume 40 in
2006) and Herpetological Circulars (1973-, 36 num-
bers to 2007).
FRANCIS R. COOK
Emeritus Curator and Researcher, Canadian Museum of
Nature, PO. Box 3443, Station D, Ottawa, Ontario
K1P 6P4 Canada
Science and Conservation of Vernal Pools in Northeastern North America
Edited by Aram J.K. Calhoun and Phillip G. deMaynadier.
2008. CRC Press Boca Raton, 6000 Broken Sound Park-
way, NW (Suite 300), Boca Raton, Florida 33487. 363
pages. 97.80 USD Cloth.
Some time ago the public began to notice that urban
and private development had started to encroach upon
lands which had traditionally been identified as waste-
land, swamps and bogs. Habitat was being lost with the
destruction of breeding grounds of birds, fish and many
mammals to create shoreline, developed areas in subdi-
visions and agricultural lands where the previous condi-
tion of the land had been determined to be “too wet”.
2007
As we have spent research time, money and bureau-
cracy on the identification and preservation of wetlands,
biologists have begun to see the further encroachment
on lands which are seasonally flooded but do not have
the distinction of being wetlands.
Pools of water and the retention of water in a pro-
tected setting often occurs in the springtime in many
places in forested areas as well as shallow ditches both
altered by humans or left by glaciation. With the pres-
ence of water, different habitats are created which
provide breeding grounds for many invertebrates as
well as amphibians, reptiles and birds, become present
since the spring is breeding season and the above need
habitat which will allow them to go through parts of
their life cycles in standing water. Many plants are also
associated with pools of water or the moist depressions
which are left after the water evaporates. Many of us
recognize these plants as common weeds but weeds
provide seeds for food, shelter for nesting, and habi-
tat for a variety of fauna to move and live beneath.
Conservation is one stated purpose of the book. Some
states have already identified the shallow pools of water
present in the spring, and other provinces and states
could easily identify similar regions using the criteria
outlined here. People who wish to live in country set-
tings often have to alter the natural setting which they
have selected in order to build upon it. Developers build
according to formulae which sell houses, not rustic set-
tings where the best interests of nature are preserved.
Builders choose level versus low-lying ground which
leads to bulldozing depressions into lawns and wood-
lots into open areas. Environmental laws now define the
YOUNG NATURALISTS
Amazing Animal Journeys
By Liam O’Donnell. 2008. Tourmaline Editions Inc./DK Pub-
lishing Canada. 48 pages. 4.99.CAD Paper.
Snow Dogs! Racers of the North
By Ian Whitelaw. 2008. Tourmaline Editions Inc./DK Publish-
ing Canada, 662 King Street West, Suite 304, Toronto,
Ontario, M5V 1M7. 48 pages. 4.99 CAD Paper.
Amazing Animal Journeys and Snow Dogs! Racers of
the North, both by Canadian writers, are books about
animals and travel. Amazing Animal Journeys, rated at
DK Publishing’s Level 3-Reading Alone, is the story of
four long-distance wildlife migrations. Snow Dogs!
Racers of the North, rated at Level 4-Proficient Readers,
is about dogs and people travelling together in the north.
Of the two books, Snow Dogs! Racers of the North is
the most coherent, mainly because it focuses on a single
story line: dog sledding. The book is divided into 10
chapters covering topics that include the history of dog
sledding, human-dog teamwork, descriptions of high-
profile sled-dog races, and profiles of sled-dog racers.
The information provided is comprehensive and inter-
esting. The sentence structure is dynamic, flowing, and
BOOK REVIEWS
449
wetlands but the woodland depressions which form ver-
nal pools have neither been identified nor are builders
inclined to further limit the buildable space by look-
ing for areas to preserve.
All researchers and professionals in the field who
work in areas which are populated by vernal pools can
benefit from this rich source. The available literature
of flora, fauna, conservation and identification of vernal
pools is shown for naturalists and conservationists alike.
It has also built upon an earlier work by Elizabeth
Colburn, Vernal Pools: Natural History and Conserva-
tion (Colburn 2004) and borrows heavily from her fur-
ther research.
This book is a valuable resource in the identification
and ecology of seasonal pools which support many
amphibians, insects and invertebrates. Reptiles, birds
and mammals are also occasional visitors and some-
times breeders. Human building pressures, pesticides,
and conservation opportunities are all parts of the study
forming the most comprehensive work I have seen on
this topic to date. I was only able to read this book slow-
ly because it contains so much research but found my-
self going back to sections for reference around related
topics. It is well researched, carefully prepared and a
valuable resource for conservationists.
Literature Cited.
Colburn, E.A. 2004. Vernal Pools: Natural History and Conservation.
The MacDonald and Woodward Publishing Co., Blackburg, Vir-
ginia USA.
Jim O° NEILL
28718 Five Mile Road, Livonia, Michigan 48154 USA
appropriate to the reading level. And the illustrations,
mainly photographs, as well as maps of dog-sled race
routes, are plentiful and colourful.
All things considered, Snow Dogs! Racers of the
North is an interesting, well-written book. My only
complaint is with the title, which suggests a book about
sled dogs. Yet the content of the book concentrates on
sled-dog racing, races, and mushers. While dogs are
depicted in most of the photographs, and mentioned on
every page, the spotlight is not shone on them very
often, and then only briefly.
Two of the book’s 48 pages deal with breeds of sled
dogs. One paragraph is dedicated to the roles played
by dogs positioned at different points along the gang-
line. One photograph and three sentences portray Balto,
the famous lead dog who guided the sled in the last
leg of the heroic journey that inspired Alaska’s re-
nowned Iditarod Sled Dog Race. Oh yes, the name of
another lead dog involved in that journey. Togo, was
also mentioned.
The book’s title led me to hope for at least one chap-
ter on the various breeds of sled dog developed in differ-
450
ent parts of the world. Another chapter portraying
famous sled dogs, past and present, would have been
fascinating, and would have helped round out the book
and make it truer to its title. And I would have liked
to see more information about the qualities sought for
dogs playing different roles on a sled-dog team. The
dogs, who work so hard, and who give so much to the
sport — without whom the sport would not, in fact, be
possible — deserve as much.
The focus of Amazing Animal Journeys is definitely
on animals. And the stories it tells — about Barren-
ground Caribou, Monarch Butterfly, Gray Whale and
Snow Goose migrations — are indeed amazing. While
the book offers lots of terrific information and colour-
ful illustrations, mainly photographs, it tries, in my
opinion, to do too much.
NEW TITLES
Prepared by Roy JOHN
+ Available for review * Assigned
THE CANADIAN FIELD-NATURALIST
Vol. 121
Covering four complex wildlife migrations in 48
pages is itself an amazing feat. To do so successfully
would have required either telling fewer migration sto-
ries more fully, or telling the same number of stories
more simply. The author, unfortunately, tried to take the
middle road, and the result is stories that jump around
a little too much for smooth reading, and that don’t
provide enough links between details to make the infor-
mation flow nicely. That said, the sentence structure is
appropriate to the reading level, and the stories are cer-
tainly interesting.
Both books, however, left me wanting to know more.
And that, perhaps, is not a bad thing.
RENATE SANDER-REGIER
3, 11th Line, Bristol, Quebec JOX 1GO Canada
Currency Codes — CAD Canadian Dollars, USD U.S. Dollars, EUR Euros, AUD Australian Dollars, GBP Great Britain Pound.
ZOOLOGY
The Question of Animal Culture. Edited by Kevin N. Laland
and Bennett G. Galef. 2009. Harvard University Press, 79 Gar-
den Street, Cambridge, Massachusetts 02138 USA. 298 pages.
49.95 USD Cloth
+ Animal Life. 2008. Tourmaline Editions Inc., DK Publish-
ing Canada, 662 King Street West, Suite 304, Toronto,
Ontario MSV 1M7 Canada. 512 pages. 55.00 CAD.
Oxford Dictionary of Biology [Sixth Edition]. By Elizabeth
Martin and Robert S. Hine. 2008. Oxford University Press,
198 Madison Avenue, New York, New York 10016 USA. 704
pages. 19.95 USD Paper.
* Extreme Birds. By D. Couzens. 2008. Firefly Books Ltd. 66
Leek Crescent, Richmond Hill Ontario L4B 1H1 Canada. 281
pages. 45 CAD Cloth.
Feathered Dinosaurs: The Origin of Birds. By John Long.
2008. Oxford University Press, 198 Madison Avenue, New
York, New York 10016 USA. 208 pages. 43.95 USD Cloth.
Witness to Extinction: How We Failed to Save the Yangtze
Dolphin. By Samuel Turvey. 2008. Oxford University Press,
198 Madison Avenue, New York, New York 10016 USA. 256
pages. 32.95 USD Cloth.
Egg & Nest. By Rosamond Purcell, Linnea S. Hall, and René
Corado, 2008. Harvard University Press, 79 Garden Street,
Cambridge, Massachusetts 02138 USA. 232 pages. 39.95
USD Cloth.
Fish of the Rockies. By Bill Gould, David Propst, Michael G.
Sullivan. 2009. Lone Pine Publishing 206, 10426-81 Avenue,
Edmonton, Alberta TOE 1X5 Canada. 208 pages. 18.95 CAD
Paper.
* Frog. By Thomas Lorent. 2008. Nature / Tourmaline Edi-
tions Inc., DK Publishing Canada, 662 King Street West,
Suite 304, Toronto, Ontario, M5V 1M7 Canada. 280 pages.
33.00 CAD Cloth.
Mountain Gorillas: Biology, Conservation, and Coexis-
tence. By Gene Eckhart and Annette Lanjouw. 2008. The
Johns Hopkins University Press, 2715 North Charles Street,
Baltimore, Maryland 21218-4363 USA. 224 pages. 24.95
USD Cloth.
Organization of Insect Societies: From Genome to Socio-
complexity. Edited by Jiirgen Gadau and Jennifer Fewell.
2009. Harvard University Press, 79 Garden Street, Cambridge,
Massachusetts 02138 USA. 518 pages. 79.95 USD Cloth.
Men: Evolutionary and Life History. By Richard G. Bri-
biescas. 2008. Harvard University Press, 79 Garden Street,
Cambridge, Massachusetts 02138 USA. 320 pages. 17.95
USD Paper.
Prairie Dogs: Communication and Community in an Ani-
mal Society. By C. N. Slobodchikoff, Bianca S. Perla, and
Jennifer L. Verdolin. 2009. Harvard University Press, 79 Gar-
den Street, Cambridge, Massachusetts 02138 USA. 280 pages.
39.95 USD Cloth.
The Beachcomber’s Guide to Seashore Life in the Pacific
Northwest. By J. Duane Sept. 2008. Harbour Publishing, PO
Box 219, Madeira Park, British Columbia VON 2HO Cana-
da. 240 pages. 26.95 USD Paper.
Squirrels of North America. By Tamara Eder. 2009. Lone
Pine Publishing 206, 10426-81 Avenue, Edmonton, Alberta
T6E 1X5 Canada. 160 pages. 16.95 CAD Paper.
* Return to Warden’s Grove: Science, Desire and the Life
of Sparrows. By Christopher Norment. 2008. Iowa Univer-
sity Press, 2121 State Avenue, Ames, lowa 50014-8300 USA.
234 pages. 26.00 USD Cloth.
2007
BOTANY
Alaska Trees and Shrubs. Second edition, By Leslie Viereck
and Elbert L. Little. 2008. University of Alaska Press, P.O.
Box 756240, Fairbanks, Alaska 99775-6240 USA. 370 pages.
24.95 USD,
The Algal Bowl - Overfertilization of the World’s Fresh-
waters and Estuaries. By David W. Schindler and John R.
Vallentyne. 2008. University of Alberta Press, Ring House 2,
Edmonton, Alberta T6G 2E] Canada. 348 pages. 34.95 CAD
Paper
* Glistening Carnivores — the sticky-leaved insect-eating
plants. By Stewart McPherson. 2008. Redfern Natural History
Productions Limited, 61 Lake Drive, Hamworthy, Poole,
Dorset BH15 4LR, England, UK. 385 pages. 29.99 GBP
Cloth.
Edible and Medicinal Plants of Canada. By Amanda Karst,
Linda J. Kershaw, Patrick Owen and Fiona Hammersley
Chambers. 2009. Lone Pine Publishing 206, 10426-81 Ave-
nue, Edmonton, Alberta T6E 1X5 Canada. 448 pages. 29.95
CAD Paper.
Major Evolutionary Transitions in Flowering Plant Repro-
duction. By Spencer C. H. Barrett. 2008. University of Chica-
go Press, 1427 E. 60th Street, Chicago, Illinois 60637 USA.
216 pages. 30.00 USD Paper.
MISCELLANEOUS
Outside in the Interior: An Adventure Guide for Central
Alaska. By Kyle Joly. 2007. University of Alaska Press, P.O.
Box 756240, Fairbanks, Alaska 99775-6240 USA. 160 pages.
19.95 USD Paper.
Living with Wildness — An Alaskan Odyssey. By Bill Sher-
wonit. 2008. University of Alaska Press, P.O. Box 756240,
Fairbanks, Alaska 99775-6240 USA. 232 pages. 21.95 USD.
Crosscurrents North — Alaskans on the Environment. Edit-
ed by Marybeth Holleman and Anne Coray. 2008. Universi-
ty of Alaska Press, P.O. Box 756240, Fairbanks, Alaska 99775-
6240 USA. 320 pages. 26.95 USD.
Audubon: Early Drawings. By John James Audubon. 2008.
Harvard University Press, 79 Garden Street, Cambridge,
Massachusetts 02138 USA. 288 pages. 125 USD cloth.
Sustaining Life: How Human Health Depends on Biodi-
versity. By Eric Chivian and Aaron Bernstein. 2008. Oxford
University Press, 198 Madison Avenue, New York, New York
10016 USA. 528 pages. 39.50 USD Cloth.
Wild Costa Rica: The Wildlife and Landscapes of Costa
Rica. By Adrian Hepworth. 2008. The MIT Press, 55 Hay-
ward Street, Cambridge, Massachusetts 02142 USA. 176
pages. 29.95 USD Cloth.
CO, Rising: The World’s Greatest Environmental Chal-
lenge. By Tyler Volk. 2008. The MIT Press, 55 Hayward
Street, Cambridge, Massachusetts 02142 USA. 264 pages.
22.95 USD or 14.95 GBP Cloth.
The Shadows of Consumption: Consequences for the
Global Environment. By Peter Dauvergne. 2008. The MIT
Press, 55 Hayward Street, Cambridge, Massachusetts 02142
USA. 328 pages. 24.95 USD or 16.95 GBP Cloth.
BOOK REVIEWS 45]
Charles Darwin: The Concise Story of an Extraordinary
Man. By Tim M. Berra. 2008. The Johns Hopkins University
Press, 2715 North Charles Street, Baltimore, Maryland 21218
4363 USA, 136 pages. 9.95 USD Cloth
DDT, Silent Spring, and the Rise of Environmentalism.
By Thomas Dunlap. 2008. The University of Washington
Press, P.O. Box 50096, Seattle, Washington 98145-5096 USA
160 pages. 16.95 USD Paper.
Oxford Dictionary of Environment and Conservation. By
Chris Park. 2008. Oxford University Press, 198 Madison
Avenue, New York, New York 10016 USA. 528 pages. 24.95
USD Paper.
Plains Apache Ethnobotany. By Julia A. Jordan. 2008. Uni-
versity of Oklahoma Press, 1005 Asp Avenue, Norman, Okla-
homa 73019 — 0445 USA. 240 pages. 34.95 USD.
Evolution: The First Four Billion Years. Edited by Michae!
Ruse and Joseph Travis. 2009. Harvard University Press, 79
Garden Street, Cambridge, Massachusetts 02138 USA. 992
pages. 39.95 USD Cloth.
The Better to Eat You With: Fear in the Animal World. By
Joel Berger. 2008. University of Chicago Press, 1427 E. 60th
Street, Chicago, Illinois 60637 USA. 360 pages. 29.00 USD
Cloth.
Global Catastrophes and Trends. By Vaclay Smil. 2008.
The MIT Press, 55 Hayward Street, Cambridge, Massachusetts
02142 USA. 320 pages. 29.95 USD or 19.95 GBP Cloth.
Global Environmental History. By |. G. Simmons. 2008.
University of Chicago Press, 1427 E. 60th Street, Chicago,
Illinois 60637 USA. 288 pages. 49.00 USD Cloth.
The Discovery of Global Warming. Revised and expanded
edition. By Spencer R. Weart. 2008. Harvard University Press,
79 Garden Street, Cambridge, Massachusetts 02138 USA. 240
pages. 16.95 USD Paper.
* Lost Worlds of the Guiana Highlands. By Stewart
McPherson. 2008. Redfern Natural History Productions Lim-
ited, 61 Lake Drive, Hamworthy, Poole, Dorset BHI5 4LR
England, UK 385 pages. 29.99 GBP
The Dominant Animal: Human Evolution and the Envi-
ronment. By Paul R. Ehrlich and Anne H. Ehrlich. 2008. UBC
Press, University of British Columbia, 6344 Memorial Road,
Vancouver, British Columbia V6T 1Z2 Canada. 428 pages.
35.00 CAD Cloth.
The North American Journals of Prince Maximilian of
Wied. Volume 1: May 1832-April 1833. Edited by Stephen
S. Witte and Marsha V. Gallagher. 2008. University of Okla-
homa Press, 1005 Asp Avenue, Norman, Oklahoma 73019 —
0445 USA. 544 pages. 85.00 USD.
The Mermaid’s Tale: Four Billion Years of Cooperation
in the Making of Living Things. By Kenneth M. Weiss and
Anne V. Buchanan. 2009. Harvard University Press. 79 Gar-
den Street, Cambridge, Massachusetts 02138 USA. 268 pages.
35.00 USD Cloth.
The Accidental Mind: How Brain Evolution Has Given
Us Love, Memory, Dreams, and God. By David J. Linden.
2008. Harvard University Press, 79 Garden Street, Cam-
bridge, Massachusetts 02138 USA. 288 pages. 17.95 USD
Paper.
452
A Mountain Year. By Chris Czajkowski. 2008. Harbour
Publishing, P.O. Box 219, Madeira Park, British Columbia
VON 2HO Canada. 192 pages. 36.95 USD Cloth.
Science Magazine’s State of the Planet 2008-2009 with a
Special Section on Energy and Sustainability. By Donald
Kennedy. 2008. Island Press, 1718 Connecticut Avenue, N.W.,
Suite 300, Washington, D.C. 20009-1148 USA. 200 pages.
19.95 USD Paper.
Canadian Rockies Access Guide — Revised and Updated.
By John Dodd, Gail Helgason and Jennifer Groundwater.
2009. Lone Pine Publishing 206, 10426-81 Avenue, Edmon-
ton, Alberta T6E 1X5 Canada. 400 pages. 26.95 CAD Paper.
Chasing Science at Sea: Racing Hurricanes, Stalking
Sharks, and Living Undersea with Ocean Experts. By Ellen
Prager. University of Chicago Press, 1427 E. 60th Street,
Chicago, Illinois 60637 USA. 178 pages. 22.50 USD Cloth.
The Sea. Volume 15, Tsunamis. By Eddie N. Bernard and
Allan R. Robinson [Editors]. 2009. Harvard University Press,
79 Garden Street, Cambridge, Massachusetts 02138 USA.
462 pages. 125 USD Cloth.
Tapping the Riches of Science: Universities and the
Promise of Economic Growth. By Roger L. Geiger and
Creso M. Sa. 2009. Harvard University Press, 79 Garden
Street, Cambridge, Massachusetts 02138 USA. 262 pages.
39.95 USD Cloth.
The Oxford Book of Modern Science Writing. By Richard
Dawkins. 2008. Oxford University Press, 198 Madison
Avenue, New York, New York 10016 USA. 448 pages.
34.95 USD Cloth.
Endocrinology of Social Relationships. Edited by Peter B.
Ellison and Peter B. Gray. 2009. Harvard University Press,
79 Garden Street, Cambridge, Massachusetts 02138 USA.
528 pages. 49.95 USD Cloth.
THE CANADIAN FIELD-NATURALIST
Vol. 121
Eyewitness Companion Weather. 2008. Tourmaline Editions
Inc., DK Publishing Canada, 662 King Street West, Suite 304,
Toronto, Ontario MSV 1M7 Canada. 288 pages. 22.00 CAD.
The Weather of the Pacific Northwest. By Cliff Mass. 2008.
The University of Washington Press, P.O. Box 50096, Seattle,
Washington 98145-5096 USA. 336 pages. 29.95 USD Paper.
The Vanishing Present: Wisconsin’s Changing Lands,
Waters, and Wildlife. Edited by Donald Waller and Thomas
Rooney. 2008. University of Chicago Press 1427 E. 60th
Street, Chicago, Illinois 60637 USA. 536 pages. 40.00 USD
Cloth.
YOUNG NATURALISTS
Animals: A Visual Encyclopedia. 2008. Tourmaline Editions
Inc., DK Publishing Canada, 662 King Street West, Suite 304,
Toronto, Ontario MSV 1M7 Canada. 304 pages. 32.99 CAD.
Animal Families. 2008. Tourmaline Editions Inc., DK Pub-
lishing Canada, 662 King Street West, Suite 304, Toronto,
Ontario MSV 1M7 Canada. 64 pages. 13.99 CAD.
* DK Readers Level 3: Amazing Animal Journeys. By
Liam O’ Donnell. 2008. Tourmaline Editions Inc., DK Pub-
lishing Canada, 662 King Street West, Suite 304, Toronto,
Ontario MSV 1M7 Canada. 48 pages. 4.99 CAD Cloth.
* Earth Matters. 2008. Tourmaline Editions Inc., DK Pub-
lishing Canada, 662 King Street West, Suite 304, Toronto,
Ontario M5V 1M7 Canada. 256 pages. 27.99 CAD Cloth.
Sharks and Other Creatures of the Deep. 2008. Tourma-
line Editions Inc., DK Publishing Canada, 662 King Street
West, Suite 304, Toronto, Ontario MSV 1M7 Canada. 144
pages. 21.99 CAD.
* DK Readers Level 4: Snow Dogs. By Ian Whitelaw. 2008.
Tourmaline Editions Inc., DK Publishing Canada, 662 King
Street West, Suite 304, Toronto, Ontario M5V 1M7 Canada.
48 pages. 4.99 CAD Cloth.
News and Comment
Marine Turtle Newsletter (118)
October 2007. 32 pages: EDITORIALS: Conservation Con-
flicts, Conflicts of Interest, and Conflict Resolution: What
Hopes for Marine Turtle Conservation? (L. M. Campbell) —
ARTICLES: From Hendrickson (1958) to Munroe & Limpus
(1979) and Beyond: An evaluation of the Turtle Barnacle
Tubicinella cheloniae (A. Ross and M. G. Frick) — Nest Re-
location as a Conservation Strategy: Looking from a Different
Perspective (O. Turkkozan and C. Yilmaz) — Linking Micro-
nesia and Southeast Asia: Palau Sea Turtle Satellite Tracking
and Flipper Tag Returns (S. Klain, J. Eberdong, A. Kitalong, Y.
Yalap, E. Matthews, A. Eledui, M. Morris, W. Andrew, D. Albis
and P. Kemesong) — Morphometrics of the Green Turtle at
the Atol das Rocas Marine Biological Reserve, Brazil (A.
Grossman, P. Mendonca, M. Rodriques da Costa, and C.
Bellini) — NOTES: Epibionts of Olive Ridley Turtles Nesting
at Playa Ceuta, Sinaloa, Mexico (L. Angulo-Lozano, P. E.
Nava-Duran and M. G. Frick) Self-Grooming by Logger-
head Turtles in Georgia, USA (M. G. Frick and G. McFall)
— IUCN-MTSG QUARTERLY REPORT ANNOUNCEMENTS
— NEWS & LEGAL BRIEFS — RECENT PUBLICATIONS.
The Marine Turtle Newsletter is edited by Lisa M.
Campbell, Nicholas School of Environment and Earth Sci-
ences, Duke University, 135 Duke Marine Lab Road, Beaufort,
North Carolina 28516 USA; and Matthew H. Godfrey, North
Carolina Wildlife Resources Commission, 1507 Ann Street,
Beaufort, North Carolina 28516 USA. Subscriptions and dona-
tions towards the production of the MTN can be made online
at or postal mail to Michael
Coyne (Managing Editor) Marine Turtle Newsletter, | South-
ampton Place, Durham, North Carolina 27705 USA; e-mail:
mcoyne @seaturtle.org.
The Vermont Reptile and Amphibian Atlas Project Publishes the First Vermont Amphibian
Posters
The Vermont Reptile and Amphibian Atlas Project has pro-
duced two informational posters on the amphibians of Ver-
mont. Amphibians Part | contains the frogs of Vermont and
Amphibians Part 2 contains the salamanders of Vermont.
Photographs of all Vermont amphibians are included on the
posters along with the latest available (2005) distribution
maps, identification information, and the basic natural history
of each species. These posters reflect the most reliable and cur-
rent information available on Vermont amphibians. Each color
poster is approximately 23” by 16” and is printed on durable,
glossy, heavyweight paper with non-fading inks. Posters cost
$9.43 each ($10 with VT sales tax) with a $3 shipping charge.
In 2006, two Vermont reptile posters were printed and all four
posters can be seen at the Atlas website http://community.
middlebury.edu/~herpatlas/index.html. All proceeds from the
sale of the posters will help fund the Vermont Reptile and Am-
phibian Atlas project.
The Vermont Reptile and Amphibian Atlas Project is an
effort begun in 1994 by the Vermont Reptile and Amphibian
Scientific Advisory Group to the Endangered Species com-
mittee. The atlas project initially began as an effort to gather
data for use by this volunteer advisory group in order to make
informed recommendations regarding the appropriate status
and conservation of reptiles and amphibians in Vermont. Since
then, the goals have widened to incorporate public education,
encourage citizen involvement, and promote conservation. The
Atlas Project is coordinated by Jim Andrews, a research her-
petologist at Middlebury College. The ultimate goal of the
Atlas is to gather and disseminate data on reptiles and amphib-
ians in Vermont in a way that involves and informs Vermont
individuals and organization so that they will become more
informed and effective stewards of wildlife habitat. Since the
Atlas began in 1994, over 3,000 individuals, fifteen private
organizations, and many government employees and agencies
have provided over 60 000 records of local reptiles and am-
phibians. With the help of these volunteers and a few part-
time staff members, the atlas continues to collect information
and broaden the knowledge base regarding the natural history,
distribution, and effective conservation of Vermont's reptiles
and amphibians.
Any resident or visitor can contribute valuable distribution
information simply by reporting what they saw, when they saw
it, and where it was. Reports can be forwarded by mail, phone,
or better yet, through the use of online reporting system at
the website http://community.middlebury.edu/~herpatlas/index.
html. Photographs are particularly helpful but not necessary.
With a good description, sketch, or photograph, they can figure
out what species you have observed and enter it into their data-
base. The data help them generate accurate maps of the in-state
distribution of Vermont’s native herptofauna.
Maps at their website show clearly the towns and species
most in need of additional survey work. Some of the towns
most in need of additional reports include Lunenburg, Lyndon,
Andover, Baltimore, Burke, Danville, Granby, Irasburg, Rut-
land, Cabot, Derby, Elmore, Warner’s Grant, Avery’s Gore,
and Pittsfield. Cities most in need of additional surveys in-
clude St. Albans, Barre, Newport, and Vergennes. Species most
in need of additional reports include all reptiles but particularly
snakes other than Gartersnake and all turtles. However. gaps
exist in the distribution maps for even some common species
and in some towns that have been surveyed more thoroughly.
Additional information on the posters, how to contribute to
the survey effort, and resources on identification and natural
history of these species groups can be found at http://commu
nity.middlebury.edu/~herpatlas.
For more information please contact James Andrews, Vermont
Reptile and Amphibian Atlas, Sunderland 114, Middlebury
College, Middlebury, Vermont 05753 jandrews @middlebury.
edu.
453
The Ottawa Field-Naturalists’ Club Awards for 2006
IRWIN M. Bropo, CHRISTINE HANRAHAN, DIANE LEPAGE, and ELEANOR ZURBRIGG
On April 28, 2007, members and friends of the
Ottawa Field-Naturalists’ Club gathered at the Club’s
Annual Soirée at St. Basil’s Church in Ottawa, and once
again, awards were given to members, and one non-
member, who distinguished themselves by accomplish-
ments in the field of natural history and conservation, or
Annie Bélaire — Member of the Year
The OFNC’s Member of the Year award recognizes
the club member judged to have contributed the most
to the club during the previous year. Annie Bélaire is
this year’s Member of the Year for her dedicated, cre-
ative and innovative involvement with the Macoun Club,
as well as her work as a volunteer at the Fletcher Wild-
life Garden.
Annie started working with the Macoun Field Club
four years ago, in the fall of 2003. Annie attends every
meeting and field trip as a knowledgeable leader,
always ready to answer children’s questions and enrich
their experience with her special knowledge. For exam-
ple, when the group found the fresh remains of a big
coyote-killed deer in Stony Swamp, she eagerly seized
the skull and pointed out to all who were gathered
around, “You can tell this was still a young deer,
because the teeth are still sharp and the dentine and
enamel are of equal thickness!” Understanding that part
of Stony Swamp is the Club’s Study Area, she under-
took a resurvey of Eastern Hemlock trees and found
that their population has declined severely over the past
‘by extraordinary activity within the Club. Careful obser-
vation of plants and animals, political savvy, selfless
service, and teaching skills are all in evidence among
this year’s winners. The following citations for those
who received an award were read to the members and
guests assembled for the event.
15 years, owing to a special fondness of porcupines and
deer for their foliage. She then shared her findings with
the children at a Macoun Club meeting, and wrote up
her report for the Club’s Little Bear magazine.
In the face of declining membership in the Macoun
Field Club in recent years, it became clear that one of
the major problems is “getting the word out,” 1.e., pub-
licity. To help the Macoun Club reach families with
young children, Annie designed a vibrantly coloured
new brochure and a poster, which have become signifi-
cant parts of its publicity efforts.
Aside from her energetic efforts with the Macoun
Club, Annie participates as a willing volunteer at the
Fletcher Wildlife Garden as part of the Friday morning
crew working in the Backyard Garden and Butterfly
Garden, and also serving on the FWG committee. In
addition, Annie has done translations for our brochures
and other bilingual Club documents, and she has put
a lot of time into organizing and data-basing the library.
Annie Beélaire is clearly well qualified to be our
Member of the Year for 2006.
Christina Lewis — George McGee Service Award
The OFNC George McGee Service Award is given in
recognition of a member who has contributed signifi-
cantly to the smooth running of the Club over several
years.
Since joining the OFNC in 1995, Chris Lewis has
been an exceptionally busy and productive member of
the club. A long-time member of the Birds Committee,
Chris has served as recording secretary since 2001, a
position she continues to fill. Many club members will
recognize Chris as one of the stalwarts of the Seedathon
team who not only organizes, but participates in this
important event to raise money for the purchase of seeds
for all the OFNC bird feeders in the region.
Chris is the recorder and reporter for the OFNC’s
Bird Status line, a position she has held since 2001. She
is responsible for making sure the data are logged, re-
corded, and ultimately submitted to ONTBIRDS, the
provincial listserv for bird observations. This is no small
task and is vitally important if we are to have reliable
data on bird occurrences in this region.
In 2003, Chris joined the OFNC’s Bird Records Sub-
committee, which is responsible for reviewing all rare
bird reports from the region. A very keen and knowl-
edgeable birder, Chris has also written numerous rare
bird reports since 1995. In addition, she contributes sig-
nificantly to the Bird Records Subcommittee database.
Chris 1s also responsible for many of the regular quar-
terly bird summaries in Trail & Landscape. She has also
authored or co-authored many other articles for T&L,
most recently an excellent report on the first confirmed
nesting of the Double-crested Cormorant in Ottawa.
Chris authored the significant species summaries for
the Ottawa area for the Birder’s Journal (2001-2004),
and for the American Birding Association’s North
American Birds (2001 to the present).
With Bob Bracken, she has co-authored several arti-
cles promoting Ottawa area bird-watching for “OFO
News”, the newsletter of the Ontario Field Ornithol-
ogists.
454
|
2007
When the Ontario Field Ornithologists held their
annual convention in Ottawa in September 2006, Chris
very capably assisted not only with promotion of field
trips for this event, but helped to lead several very suc-
cessful trips.
While many people may connect Chris with birds,
there are many more who know her as an expert on
odonates — dragonflies and damselflies, and on Lepi-
doptera. Over the last decade she has led or co-led
numerous walks for the OFNC on birds, butterflies,
dragonflies (the Morris Island dragon walks are very
popular), fish and general natural history.
CLUB AWARDS 45
WN
Her interest in, and knowledge of, odonates has led
her to publish some important articles on dragonflies
and damselflies of the Ottawa area, and submit an aston-
ishing 2,000+ records of odonates for the Ottawa area,
together with Bob Bracken, to the Ontario Odonata
Summaries. She and Bob received the 2001 Anne Hanes
Natural History award for this work.
These are remarkable contributions, both in terms of
service to the club and to the furthering of our knowl-
edge of natural history in this area. For all of these rea-
sons, the OFNC is pleased to give the 2006 Service
Award to Christina Lewis.
Madeline Kallio — Conservation Award (Non-Member)
The OFNC Conservation Award — Non-Member is
given in recognition of an outstanding contribution by
a non-member in the cause of natural history conser-
vation in the Ottawa Valley, with particular emphasis
on activities within the Ottawa District.
For nine years, Madeline Kallio has been the driving
force behind the National Capital Region Wildlife
Festival. Through its Wildlife Awareness event, library
programs, public forums, guided nature walks and
more, the festival has, for many years, introduced the
general public to natural history and the concept of
conservation.
While the festival appears to run effortlessly, in fact,
a tremendous amount of organization and year-round
planning is required. Madeline Kallio chairs the festi-
val committee which meets approximately 11 times a
year, produces the minutes, and attends to the thousand
and one details that arise. She is the liaison with the
Billings Bridge Shopping Centre (where the wildlife
awareness event takes place), drawing up detailed floor
plans, and ensuring that everything needed to make this
particular event run smoothly is taken care of. She is
also the link with all the exhibitors, contacting them
each year, determining their requirements, making sure
they have the right location for their exhibit, etc. She
staffs the Wildlife Festival booth for the entire three days
and is always available should other exhibitors need
assistance. Coordinating an event of this magnitude and
ensuring that it runs smoothly requires skill, good organ-
izational abilities and huge time commitments.
Madeline also writes all the grant proposals and con-
stantly seeks new ways to fund this event. In addition,
she writes, produces and prints the festival’s events
brochure. A few years ago, she came up with the bril-
liant idea of developing a map of greenspaces in east-
ern Ontario where people can explore nature. Called
“On the Trail of Mother Nature” this large, full-colour
map, prepared, researched, and primarily designed by
Madeline, has been a huge success with several updates
and thousands of copies distributed.
Madeline is full of ideas for making the festival ever
more interesting. One example is the library program
series, which is so popular that presenters from previ-
ous years contact Madeline to make sure they are in-
cluded in the current line-up, while libraries begin ask-
ing for information well in advance of the festival itself.
She encouraged the idea of a public forum (now in its
fifth year), has been instrumental in organizing the fes-
tival’s Environmental Awards ceremony, and probably
has a hundred other ideas just waiting to burst forth!
Endlessly resourceful, energetic, and tireless, Madeline
is always willing to step in and pick up where needed.
On top of all this, she is always a calming, cheerful
presence, seemingly unflappable, and without a doubt
is the glue that holds the festival together!
Long before she was the festival’s Coordinator,
Madeline was a member of the festival committee, join-
ing in 1990. When a coordinator was needed, Madeline
stepped in and has not looked back.
You might think that the festival was enough to
keep Madeline occupied full-time. This would be true
for most people, but Madeline is involved in a num-
ber of other activities, including volunteering for the
Kanata Canada Day program. She is also an accom-
plished writer who has written and published many
articles and brochures. Chances are, you have seen her
writing. If you have picked up brochures on Lanark
County Maple Byway Tours, or browsed the “Coffee
Companion’ newsletter in a Kanata coffee house, read
magazines such as Fifty-Five Plus or Due West, you’ve
seen her work.
Back in 1985, Madeline set up Ottawa Valley Field
Trips, to introduce young and old to the joys of the nat-
ural world. She developed hands-on environmental
programs for schools, organized workshops on natural
history themes for the school boards, and worked
extensively on nature interpretation programs with
the National Capital Commission, Scouts Canada,
Girl Guides, the Home School Association and other
learning and public groups. The company also devel-
oped “backroads tours”, which explored interesting
villages and natural areas on the backroads of Eastern
Ontario, as well as longer trips in Ontario, Quebec and
the ULS.
The OFNC is proud to recognize Madeline Kallio for
her outstanding efforts on behalf of nature and conser-
vation in the Ottawa region, and in particular for her
work with the National Capital Region Wildlife Festival.
456
Frank Pope — Conservation Award (Member)
The Conservation Award for Members is presented
in recognition of an outstanding contribution by a mem-
ber of the OFNC in the cause of natural history con-
servation in the Ottawa Valley, with particular empha-
Sis ON activities within the Ottawa District. Frank Pope
is chosen as the recipient for the 2006 award in recog-
nition his skillful leadership of the Alfred Bog Commit-
tee for over 20 years, which in 2006 achieved its goal
of protection for the Alfred Bog for generations to come.
Alfred Bog, lying 75 kilometres east of Ottawa, is one
of the great conservation success stories of the past two
decades. It took long years of hard work by local groups
and all levels of government to finally save this vulnera-
ble wetland from peat extraction, logging and drainage.
Frank Pope’s involvement in the Alfred Bog con-
servation effort began in earnest in the early 1980s. A
large part of the bog had been re-zoned to Agriculture,
and an appeal to the Ontario Municipal Board seeking
to keep the zoning as Conservation had failed. Not will-
ing to accept defeat, Frank Pope, with Charles Sauriol
of the OFNC, created the Alfred Bog Committee in
1985 to bring together interested folks and strengthen
the bog protection effort. (At the time, Frank could not
have known that it would take over 20 years to achieve
success!) Frank chaired the Alfred Bog Committee in its
ongoing save-the-bog campaign for the ensuing 20+
years, a long, drawn-out struggle that, against the odds,
achieved its goal.
In this exhausting battle, Frank’s commitment never
wavered in the face of setbacks. His efforts spanned a
range of activities — making credible presentations
at Ontario Municipal Board hearings, tribunals, advo-
cating legislative change at the provincial level (for
example, to enable municipalities to control peat extrac-
tion activities), commenting on official plans and by-
THE CANADIAN FIELD-NATURALIST
Vol. 121
laws, working with major NGOs (NCC) in land acqui-
sition efforts, fund-raising, and negotiating with diverse
groups to name but a few.
Under Frank’s leadership, the Alfred Bog Committee »
watched for acquisition opportunities within the bog,
and initiated such activities as the preparation of a man-
agement plan, a major report on the plants, animals and
hydrology of the bog, contacts with other land hold-
ers in the bog, the preparation of educational brochures —
and articles and field trips into the bog. Funding was
obtained and a 1000 ft. boardwalk was constructed in |
the bog, because the impressive natural features of the
bog made it a popular destination for naturalists and ©
outdoor lovers, whereas walking in the bog is destruc-
tive to bog flora and hazardous to walkers.
In the years leading up to 2004, the combined con-
servation work of non-governmental groups and the sev- |
eral levels of government intensified. The Nature Con- |
servancy of Canada purchased a total of 2800 hectares
of the bog, and Ontario Parks now holds title to this»
property, to be managed as a provincial nature reserve.
Peat harvesting within the bog was banned when the:
Prescott-Russell County Council passed the Alfred Bog »
Protection By-Law. In addition, peat-mining operations |
that border the bog and still pose a threat to its integrity |
are now required to operate according to rules designed
to minimize this threat. All appeals of the county offi-
cial plan pertaining to the Bog are withdrawn and the
years-long mediation of the Ontario Municipal Board:
terminated. The Bog now has an official boundary and
wetland designation. Quite a success story!
With this conservation award, the OFNC is recogniz-
ing Frank Pope’s unflagging commitment, leadership
and final success in the effort to gain environmental
protection for Alfred Bog. Congratulations, Frank!
Linda Jeays — Anne Hanes Natural History Award
This award is given in recognition of a member who,
through independent study or investigation, has made
a worthwhile contribution to our knowledge, under-
standing and appreciation of the natural history of the
Ottawa Valley.
Linda Jeays, an enthusiastic birder and longtime
member of OFNC, began to study butterflies as a Mil-
lennium project. Her first identification was a Black
Swallowtail flushed while watching waterfowl at Rich-
mond lagoons. She jotted down a description, later
referred to a field guide, and suddenly had a new nat-
ural history interest.
Linda is a former English teacher with an inclination
towards note-taking. In her first summer studying Lepi-
doptera she began to make detailed documentation on
every butterfly she encountered except where numbers
made this impractical. She also keeps data on topics
such as flight season, habitat, nectaring plants, road-kill
and garden species.
At the end of the 2001 season, Linda learned that her
sightings of Eastern Tailed Blue and Common Buckeye:
were very rare in the Ottawa area. Encouraged by Ross:
Layberry to write up her observations for Trail & Land-'
scape and to check the locations again the following |
year, her detailed notes soon became valuable. In 2002,’
Linda again saw Buckeye briefly, but her real work}
began when she netted Eastern Tailed Blue within 10(
feet of a previous year’s site. In the very few records!
available, no one else had reported the butterfly in a:
repeat location.
What if Eastern Tailed Blue had overwintered? Linda’
began spending many hours each week netting, meas-
uring and individually marking the Blues. Each year
she recorded her findings in T&L, accumulating proof
of overwintering. She has now established the presence
of this species at one site near Richmond for six con-
secutive years. The culmination of Linda’s work came
in the summer of 2006 when she spent over 250 hours
|
2007
in the field documenting the population explosion and
widespread dispersal of Eastern Tailed Blue. She her-
self tallied 30 locations and 160 individuals. Another
25 sites and 170 Blues were reported by independent
contributors to the Annual Eastern Tailed Blue Round-
up that Linda conducts.
Linda Jeays has written feature articles, essays,
humour, book reviews and autobiographical material for
newspapers such as the Ottawa Citizen and the National
Post. She has many magazine credits and has worked
on assignment for the Canada Safety Council. Linda
wrote her first short item for T&L in 1975. In 1977
she called then-editor Anne Hanes about submitting a
piece of free verse. Anne said that unfortunately T&L
Isabelle Nichol —- Mary Stuart Education Award
The Mary Stuart Education Award was established
to recognize members, non-members or organizations,
for their outstanding achievements in the field of natu-
ral history education in the Ottawa Region. The OFNC
is pleased to give the award for 2006 to Isabelle Nicol.
Isabelle has a great affinity for teaching natural his-
tory. She is both knowledgeable and enthusiastic and
conveys this interest to the many lucky people who
have been in her classroom or participated in her many
field trips. Isabelle was a former employee of the NCC
but, before she left that job, she was actively working
on this, her second career that she has pursued so suc-
cessfully for over 15 years.
Isabelle is a freelance teacher of natural history who
is invited back year after year in several local schools
because of her natural ability to teach and because of
CLUB AWARDS
457
did not use poetry, but she would still like to read the
poem. “Metamorphosis” was printed in January 1978,
the first of Linda’s 35 poems in the club magazine
Anne Hanes would surely have been pleased that
her early encouragement of the creative expression of
Linda’s nature observations has developed into metic-
ulous factual documentation of Eastern Tailed Blue
and a worthwhile contribution to the knowledge and
understanding of this butterfly in the Ottawa Valley.
For her outstanding contribution to our further know!-
edge of the Eastern Tailed Blue in the Ottawa Region,
we are very pleased to present Linda Jeays with the
Anne Hanes Natural History Award for 2006.
her careful planning of various natural history modules
that complement the subjects covered in the grade
school curriculum. These include modules on habitats,
plants, hibernation in winter, migration, spring ephemer-
als and active winter wildlife. She works closely with
the teachers to tailor her teaching to fit the interests and
abilities of the students at hand.
Recently Isabelle has branched out and now also
teaches natural history to various senior citizen groups
and garden clubs. Over these many years Isabelle has
also been a much sought after field trip leader for our
Club, where participants have come to appreciate her
quiet style and originality. We have a first rate natural
history teacher in our midst and it is time that she be
appropriately honoured.
Index to Volume 121
Compiled by Leslie Cody
Abies balsamea, 327,329,362
lasiocarpa, 156
Absinth, 184
Acalypha virginica, 184
Accipiter cooperii, 216
gentilis, 329,413
Accipiter gentilis, Exploits a Beagle Hound, Canis familiaris,
as a “Beater” to Catch a Snowshoe Hare, Lepus
americanus, Northern Goshawk, 329
Acer rubrum, 385
saccharinum, 385
saccharum, 256,304,420
spicatum, 359
Achillea millefolium, 77
millefolium ssp. millefolium, 184
Acipenser brevirostrum, 176
Actitis hypoleucos, 413
Advice to Contributors, 478
Aechmophorus clarkii, 50
occidentalis, 50
Agelaius phoeniceus, 245
Agrimonia gryposepala, 184
Agrimony, Hooked, 184
Agrostis gigantea, 184
Albert, A.J., 92
Alberta, The Influence of Air Pollution on Corticolous
Lichens near the Strathcona Industrial Area, 17
Alberta, Twenty-Four-Hour Activity Budgets of Mule Deer,
Odocoileus hemionus, in the Aspen Parkland of
Eastcentral, 299
Alca torda, 275,289
Alces alces, 156,359,391,416
alces americana, 2605
americanus, 121
Alces alces, Winter Browse Use in Central Labrador,
Moose, 359
Alder, 82,329
Mountain, 81
Alectoria spp., 380
Alfalfa, 184
Alisma gramineum, 165
plantago-aquatica, 165
Allan, M., 212
Alliance of Natural History Museums of Canada Annual
Meeting and Awards, 349
Alnus sp., 82,329
crispa, 81,297,362
Alopex lagopus, 73,308,322,330
Ambrosia artemisiifolia, 184
Amelanchier sp., 86
arborea, 89
arborea var. arborea, 203
canadensis, 89
interior, 90
intermedia 89
laevis, 89,203
spicata, 90
stolonifera, 203
xgrandiflora, 89
Amelanchier intermedia, Escaped from Cultivation in
Eastern Ontario, Serviceberry, 89
American-aster, White Heath, 204
Ammodramus bairdii, 244
henslowii, 244
leconteii, 244
savannarum, 244
Ammodytes spp., 285
hexapterus, 325
Amnicola limosa, 92
Anaphalis margaritacea, 184
Anas acuta, 245
americana, 245,325,376
clypeata, 245
crecca, 326
discors, 245
platyrhynchos, 130,245
strepera, 245
Anastrophyllum assimile, 26
Anderson, R.C., R. Shimek, J.A. Cosgrove, and S. Berthinier.
Giant Pacific Octopus, Enteroctopus dofleini, Attacks
on Divers, 423
Andropogon gerardii, 203
Anemone canadensis, 184
cylindrica, 203
Anemone, Canada, 184
Angelica genuflexa, 421
Anodonta implicate, 93,
Antennaria howellii ssp. petaloidea, 203
neglecta, 184
parlinii, 184
parlinii ssp. fallax, 203
Anthelia julacea, 26
Juratzkana, 26
Anthus spragueii, 244
Apalone spinifera, 178
Apocynum androsaemifolium, 203
Aquila chryseatos, 156
Aquilegia canadensis, 203
Arabis divaricarpa, 203
hirsuta var. pycnocarpa, 203
holboellii var. retrofracta, 203
Arctium minus ssp. minus, 184
Arctostaphylos uva-ursi, 203
Ardea herodias, 426
Aries, C.E., 325
Artemia, 130
Artemisia spp., 134
absinthium, 184
campestris ssp. borealis var. scouleriana, 203
Arthonia patellulata, 18
458
2007
Asclepias exaltata, 203
syriaca, 184,203
tuberosa ssp. tuberosa, 203
Ash, Black, 385
Mountain, 359
White, 304
Asio flammeus, 245
Aspen, 256,297,304
Quaking, 150
Trembling, 18,191,261,327
Aster ssp., 182
cordifolius, 184
curtus, 40
ericoides, 184
lanceolatus, 184
lateriflorus, 184
novae-angliae, 185
ontarionis, 186
simplex, 184
umbellatus, 184
urophyllus, 184
Aster, Arrow-leaved, 184
Calico, 184
Flat-topped White, 184
Heart-leaved, 184
Heath, 184
New England, 185,204
Panicled, 184
Sky-blue, 204
Western Willow, 166
White-top, 40
Aster, Sericocarpus rigidus, a Threatened Species from the
Garry Oak Ecosystem in British Columbia, Micro-
Propagation of White-top, 40
Atlas of amphibians and reptiles of Quebec, 2007 inventory,
234
Avens, White, 184
Yellow, 184
Avocet, American, 245
Aythya affinis, 245
americana, 245
collaris, 245
valisineria, 245
Badger, American, 150
European, 265
Ballard, W.B., 71
Banville, D., 402
Barbarea vulgaris, 184
Barbilophozia floerkei, 26
hatcheri, 26
kunzeana, 26
lycopodioides, 26
voerkei, 27
Barley, Wild, 166
Bartramia longicauda, 245
Basil, Wild, 184
Bat, 385
Big Brown, 120
Eastern Red, 124,386
Hoary, 124,208,386
Little Brown, 57,208,386,420
Long-legged, 124
Northern Long-eared, 208,386
Red, 208
INDEX TO VOLUME 12]
459
Silver-haired, 121
Western Long-eared, 124
Bat, Myotis lucifugus, in Manitoba and Northwestern Ontario,
Recent Distribution Records of the Litthe Brown, 57
Bat, Myotis lucifugus, Observation of Foliage-roosting in the
Little Brown, 420
Bat, Myotis septentrionalis (Chiroptera: Vespertilionidae), on
Prince Edward Island: First Records of Occurrence
and Over-Wintering, Northern Long-eared, 208
Bay, California, 305
Bazzania pearsonii, 26
tricrenata, 26
Bear, 385
American Black, 120,387
Black, 156,266,330,362,430
Grizzly, 120,156
Polar, 120
Bearberry, Red, 203
Beardtongue, Hairy, 203
Beaver, 271,330,385,438
American, 120,391
Giant, 330
Beaver, Castoroides ohioensis, Remains in Canada and an
Overlooked Report from Ontario, Giant, 330
Bedstraw, Licorice, 203
Northern, 203
Small, 184
Beech, American, 304,385
Beetle, 182
Bennett, B.A., 295
Berthinier, S., 423
Betula alleghaniensis, 385
glandulosa, 81,297
papyrifera, 81,304,327,359
Bindweed, Black, 184
Field, 184
Low False, 203
Birch, Dwarf, 81
White, 81,304,327,359
Yellow, 385
Bison bison, 121,130,192
bison athabascae, 124
Bison, 192
American, 121,130
Wood, 124
Bittern, American, 245
Blackbird, Brewer’s, 243
Red-winged, 242
Yellow-headed, 245
Blarina brevicauda, 206,386
Blatt, S.E., 182
Blepharostoma trichophyllum, 27
trichophyllum ssp. trichophyllum , 26
Blomme, C.G., 436
Bluegrass, Alpine, 134
Bluejoint, Canada, 184,433
Bluestem, Big, 203
Little False, 204
Blueweed, 184
Bobcat, 437
Bobolink, 243
Bombycilla cedrorum, 244
Bonasa umbellus, 329
Bond, A.L. and A.W. Diamond. Abandoned Seabird Eggs as
a Calcium Source for Terrestrial Gastropods, 433
460
Boneset, 184
Boreal Dip Net/L’epuosette boreal, The, 234,349
Bos taurus, 72
Botaurus lentiginosus, 245
Bouteloua curtipendula, 71
gracilis, 71
Branta canadensis, 245,313
canadensis maxima, 316
canadensis moffitti, 316
Branta canadensis, Management Implications of Molt Migra-
tion by the Atlantic Flyway Resident Population of
Canada Geese, 313
Brazil, J., 81
Brice, J.S., 46
British Columbia, Cowbane, Oxypolis occidentalis, A New
Native Vascular Plant Species for the Queen Charlotte
Islands, 421
British Columbia, Micro-Propagation of White-top Aster,
Sericocarpus rigidus, a Threatened Species from the
Garry Oak Ecosystem in, 40
British Columbia, 1997 2005, Translocation and Recovery
Efforts for the Telkwa Caribou, Rangifer tarandus
caribou, Herd in Westcentral, 155
British Columbia’s Gulf Island Waters, A River Otter’s, Lontra
canadensis, Capture of a Double-crested Cormorant,
Phalacrocorax auritus, in, 325
British Columbia, The Hepatic Flora and Floristic Affinity
of Hepatics Around Takakia Lake, Queen Charlotte
Islands, 24
British Columbia, The Rumsfeld Paradigm: Knowns and
Unknowns in Characterizing Habitats Used by the
Endangered Sharp-tailed Snake, Contia tenuis, in
Southwestern, 142
Brome, Smooth, 184
Bromus inermis ssp. pumpellianus, 183
Brown, J.A., D.F. McAlpine, and R. Curley. Northern
Long-eared Bat, Myotis septentrionalis (Chiroptera:
Vespertilionidae), on Prince Edward Island: First
Records of Occurrence and Over-Wintering, 208
Bruce, L., 212
Bryoria spp., 18,303,380
Bubo virginianus, 437
Bubulcus ibis, 50,330
Bucephala albeola, 370,375
Bucephala albeola) After Twilight in Winter: An Anti-Preda-
tion Tactic?, Offshore Flight of Buffleheads, 375
Bucephala albeola: Autumn Arrivals in Shoal Harbour Sanc-
tuary, Vancouver Island, in Relation to Freeze-up,
The Punctual Bufflehead, 370
Buchlée dactyoides, 72
Buffalo-berry, Russet, 204
Buffalograss, 72
Bufflehead, 370,375
Bufflehead, Bucephala albeola: Autumn Arrivals in Shoal
Harbour Sanctuary, Vancouver Island, in Relation to
Freeze-up, The Punctual, 370
Buffleheads (Bucephala albeola) After Twilight in Winter:
An Anti-Predation Tactic?, Offshore Flight of, 375
Buller, S.A., 430
Bulrush, Great, 166
Hardstem, 54
Small-fruited, 166
Bunchgrass, 134
Bunting, Lark, 242
Snow, 310
THE CANADIAN FIELD-NATURALIST
Vol. 121
Bur-Reed, 166
Burdock, Common, 184
Burrograss, 71
Buteo lagopus, 330
lineatus, 216
platypterus, 216
regalis, 245
swainsoni, 245
Buttercup, Field, 184
Kidney-leaf, 184
Labrador, 204
Thread-leaved, 166
Caenestheriella, 129
belfragei, 131
gynecia, 128
setosa, 130
Caenestheriella gynecia (Crustacea: Conchostraca), in New
York and New Jersey, State Records and Habitat of
Clam Shrimp, 128
Calamagrostis canadensis, 183,433
Calamospiza melanocorys, 244
Calcarius ornatus, 244
Calidris alpina, 376
Calypogeia azurea, 26
integristipula, 26
neesiana, 26
Calystegia spithamaea ssp. spithamaea, 203
Campion, Bladder, 184
Canada and an Overlooked Report from Ontario, Giant
Beaver, Castoroides ohioensis, Remains in, 330
Canada, Arboreal Late Summer Courtship Behaviour of Mari-
time Garter Snake, Thamnophis sirtalis pallidulus, in
Dartmouth, Nova Scotia, 210
Canada, Before and After the Application of the Herbicide
Magnacide, An Inventory of the Aquatic and Sub-
aquatic Plants in SASKWater Canals in Central
Saskatchewan, 164
Canada Geese, Branta canadensis, Management Implications
of Molt Migration by the Atlantic Flyway Resident
Population of, 313
Canada, Human-assisted Movements of Raccoons, Procyon
lotor, and Opossums, Didelphis virginiana, between
the United States and, 212
Canadian Association of Herpetologists/Association Cana-
dienne des Herpetologists Bulletin 14(2) Spring 2007,
234
Canadian Field-Naturalist, Advice for Contributors to The,
118,238,358
Canis familiaris, 268,329
latrans, 71,120,150,156,268,328,364,392,397,426,
430,437
lupus, 120,156,245,256,364,386,397,430,437
lupus baileyi, 430
rufus, 430
Canis familiaris, as a “Beater” to Catch a Snowshoe Hare,
Lepus americanus, Northern Goshawk, Accipiter gen-
tilis, Exploits a Beagle Hound, 329
Canis latrans, in Urbanized Eastern Massachusetts, Move-
ments of Transient, 364
Canis latrans, Pack, Social and Play Behavior in a Wild
Eastern Coyote, 397
Canis latrans, predation on Great Black-backed Gull, Larus
marinus, eggs on Boot Island National Wildlife Area,
Nova Scotia, Details of Eastern Coyote, 426
2007
Canis lupus, Behavior in Areas of Frequent Human Activity,
Wolf, 256
Canis lupus, Pups into Another Wolf Pack, Attempt to Cross-
Foster Gray Wolf, 430
Canvasback, 245
Capelin, 285
Capsella bursa-pastoris, 78,184
Cardamine pensylvanica, 78
umbellata, 24
Carduelis tristis, 245
Carex spp., 134,187,359
aurea, 184
backii, 203
communis, 184
cusickii, 421
debilis, 184
foenea, 184
gracillima, 184,203
granularis, 184
laxiflora, 184
molesta, 203
muehlenbergii var. muehlenbergii, 203
normalis, 184
pensylvanica, 203
richardsonii, 203
rostrata, 166
scoparia, 184
siccata, 203
stylosa, 421
tonsa var. rugosperma, 203
vulpinoidea, 184
Caribou, 82,121,327,379,385
Telkwa, 155
Woodland, 96,155,360,389
Caribou, Rangifer tarandus caribou, Herd in Westcentral Bri-
tish Columbia, 1997-2005, Translocation and Re-
covery Efforts for the Telkwa, 155
Caribou, Rangifer tarandus, Track Decadal Changes in Arctic
Tundra Vegetation, Diets of Overwintering, 379
Carrot, Wild, 184
Carthamus tinctorius, 44
Cartier, I., 402
Castor canadensis, 120,271,330,391,438
Castoroides, 330
ohioensis, 330
Castoroides ohioensis, Remains in Canada and an Overlooked
Report from Ontario, Giant Beaver, 330
Cat, Domestic, 268
House, 208
Wild, 385
Catbird, Gray, 244
Catling, P.M. and B.A. Bennett. Discovery of a Possibly
Relict Outbreeding Morphotype of Sparrow’s-egg
Lady’s-slipper Orchid, Cypripedium passerinum, in
Southwestern Yukon, 295
Catling, P.M. and B. King. Natural Recolonization of Culti-
vated Land by Native Prairie Plants and its Enhance-
ment by Removal of Scots Pine, Pinus sylvestris, 201
Catling, P.M. and G. Mitrow. Serviceberry, Amelanchier inter-
media, Escaped from Cultivation in Eastern Ontario,
89
Cattail, 53
Common, 166
Narrowleaf, 53
Ceanothus americanus, 203
INDEX TO VOLUME 121
46)
Cedar, Eastern White, 304,385
White, 431
Celery, Wild, 164
Cepaea hortensis, 434
Cephalozia ambigua, 26
bicuspidata, 26
leucantha, 26
lunulifolia, 26
macounti, 26
Cepphus grylle, 275,289
Cerastium fontanum ssp. triviale, 184
vulgatum, 78
Ceratophyllum demersum, 93,165
Cervus canadensis, 192
elaphus, 62,121,156,214,334,416
elaphus nelsoni, 133,416
Cervus elaphus nelsoni, Populations, Colonization of Non-
Traditional Range in Dispersing Elk, 133
Cervus Elaphus, Using Tooth Wear/Eruption Patterns and
Counts of Annuli in Tooth Cementum, Correlation
Between Age Estimates for Elk, 214
Cervus elaphus, with Telezol and Xylazine and Reversal with
Tolazine or Yohimbine, Immobilization of Elk, 62
Cetraria spp., 380
Chabot, D. and R. Miller. A Rare Case of Completely Ambi-
coloured Atlantic Halibut, Hippoglossus hippoglossus,
from the Lower St. Lawrence Estuary, Quebec, 35
Chara, 166
Charadrius vociferus, 130
Chelydra serpentina, 130,178
Chelydra serpentina, Hatchlings in New York and New
Hampshire, Post-Emergence Movements and Over-
wintering of Snapping Turtle, 178
Cheney, M. and K.L. Marr. Cowbane, Oxypolis occidentalis,
A New Native Vascular Plant Species for the Queen
Charlotte Islands, British Columbia, 421
Chenopodium simplex, 184
Cherry, Black, 203
Choke, 203
Clammy Ground, 184,203
Pin, 359
Susquehanna Sand, 203
Chhin, S. and G.G. Wang. Growth of White Spruce, Picea
glauca, Seedlings in Relation to Microenvironmental
Conditions in a Forest-Prairie Ecotone of South-
western Manitoba, 191
Chickadee, Black-capped, 428
Chestnut-backed, 42
Chickweed, 185
Common Mouse-ear, 184
Chicory, 184
Chipmunk, 385
Eastern, 271,389
Least, 120
Chisholm, B., 214
Chlidonias niger, 50
Chondestes grammacus, 244
Chordeiles minor, 245
Chrysanthemum leucanthemum, 184
Chrysemys picta, 178
Chrysops sp.. 257
Chubbs, T.E., 359
Chubbs, T.E. and ER. Phillips. A Tribute to Neal Philip Perry
Simon 1973-2006, 96
462
Chubbs, T.E. and F.R. Phillips. Winter Occurrences of Ivory
Gulls, Pagophila eburnea, in Inland Labrador, 327
Chubbs, T.E. and J. Brazil. The Occurrence of Muskoxen,
Ovibos moschatus in Labrador, 81
Cichorium intybus, 184
Cinquefoil, Downy, 184
Erect, 184
Old Field, 184,203
Rough, 184
Silvery, 184
Tall, 203
Cionella lumbria, 434
Circus cyaneus, 245,330
Cirsium arvense, 184,302
Cistothorus palustris, 244
platensis, 244
Cladina spp., 380
Clark in 1804-1806, Were Native People Keystone Predators?
A Continuous-Time Analysis of Wildlife Observations
Made by Lewis and, 1
Clemmys guttata, 178
Clethrionomys gapperi, 271,390
Clinopodium vulgare, 184
Clover, Alsike, 185
Red, 185
Smaller Hop, 185
White Sweet, 166,184
Yellow, 185
Yellow Sweet, 184
Clupea harengus, 433
pallasii, 325
Clyde, K.J., 422
Coad, B.W., Review by, 102
Coccyzus erythropthalmus, 245
Cod, 285
Cody Receives Honorary Doctorate of Science from
McMaster University, June 5, 2007, Bill, 233
Cole, M.J., 379
Colpitts, D.W., 426
Columbine, Red, 203
Comandra umbellata ssp. umbellata, 203
Comptonia peregrina, 203
Condylura cristata, 206
Condylura cristata) in Northeastern Vermont, A High Eleva-
tion Record of the Star-nosed Mole, 206
Contia tenuis, 124
Contia tenuis, in Southwestern British Columbia, The Rums-
feld Paradigm: Knowns and Unknowns in Charac-
terizing Habitats Used by the Endangered Sharp-
tailed Snake, 142
Convolvulus arvensis, 184
Conyza canadensis, 184
Cook, F.R., Reviews by, 100,338,339,342,343,347,439,443,
444,448
Cool, N., 214
Coot, American, 50,245
Copperleaf, 184
Cormorant, 274
Double-crested, 275,288,325,426
Great, 275,288
Pelagic, 325
Cormorant, Phalacrocorax auritus, in British Columbia’s
Gulf Island Waters, A River Otter’s, Lontra cana-
densis, Capture of a Double-crested, 325
THE CANADIAN FIELD-NATURALIST
Vol. 121
Cornus racemosa, 203
stolonifera, 359
Corvus brachyrhynchos, 150,427
corax, 150,427
Cosgrove, J.A., 423
Cota-Sanchez, J.H. and K. Remarchuk. An Inventory of the
Aquatic and Subaquatic Plants in SASK Water Canals
in Central Saskatchewan, Canada, Before and After
the Application of the Herbicide Magnacide, 164
Cott, P.A. and N.J. Mochnacz. Bull Trout, Salvelinus conflu-
entus, and North American Porcupine, Erethizon
dorsatum, Interaction in the Mackenzie Mountains,
Northwest Territories, 437
Cotter, R., 287
Cotter, R. and J.-F. Rail. Third Census of Seabird Populations
of the Gaspé Peninsula, Québec, 2002, 274
Cottongrass, 134
Couchgrass, 184
Cougar, 120,437
Cow, 72
Cowbane, 421
Cowbane, Oxypolis occidentalis, A New Native Vascular
Plant Species for the Queen Charlotte Islands, British
Columbia, 421
Cowbird, Brown-headed, 239
Cowbird, Molothrus ater, Parasitism and Abundance in the
Northern Great Plains, Brown-headed, 239
Coyote, 71,120,150, 156,268,328,364,392,397,430,437
Eastern, 426
Coyote, Canis latrans, Pack, Social and Play Behavior in a
Wild Eastern, 397
Coyote, Canis latrans, predation on Great Black-backed
Gull, Larus marinus, eggs on Boot Island National
Wildlife Area, Nova Scotia, Details of Eastern, 426
Coyotes, Canis latrans, in Urbanized Eastern Massachusetts,
Movements of Transient, 364
Cranberry, 257
Crane, Whooping, 257
Crane’s Bill, Bicknell’s, 184
Crataegus, 86 VA
Crins, W.J., Reviews by, 336,344 /
Crossman, L.M., 92
Crow, American, 150,427
Crowberry, Black, 81
Crowder, A., R. Harmsen, and S.E. Blatt. Notes on Succession
in Old Fields in Southeastern Ontario: the Herbs, 182
Crowell, M., 265
Cuckoo, Black-billed, 245
Curley, R., 208
Cyathura carinata, 168
polita, 168
Cyathura polita in the Saint John River Estuary, New Bruns-
wick: a Species at the Northern Extent of its Range,
Life History of the Marine Isopod, 168
Cyperus houghtonii, 203
lupulinus ssp. lupulinus, 203
Cypripedium passerinum, 295
Cypripedium passerinum, in Southwestern Yukon, Discovery
of a Possibly Relict Outbreeding Morphotype of
Sparrow’s-egg Lady’s-slipper Orchid, 295
Cystophora cristata, 392
Cyzicus, 129
Dactylis glomerata, 184
2007
Dadswell, M.J., 168
Daisy, Ox-eye, 184
Dalén, L., A. Gétherstr6m, T. Meijer, and B. Shapiro.
Recovery of DNA from Footprints in the Snow, 321
Dandelion, 182
Danthonia spicata, 184,203
Daucus carota, 184
Davies, C., 212
Davy, C.M. and E.E. Fraser. Observation of Foliage-roosting
in the Little Brown Bat, Myotis lucifugus, 420
Deer, 151,385
Mule, 121,156,271,299
Red, 416
White-tailed, 121,139,257,265,302,333,391,412,426,
430
Deer, Odocoileus hemionus, in the Aspen Parkland of East-
central Alberta, Twenty-Four-Hour Activity Budgets
of Mule, 299
Deer, Odocoileus virginianus, Fawn in Southcentral Wiscon-
sin, A Late Born White-tailed, 333
Deer, Odocoileus virginianus, Seasonal Variation in Sex Ratios
Provides Developmental Advantages in White-tailed,
412
Deermouse, North American, 120
Northwestern, 120
Dendragapus canadensis, 329
Dendroica petechia, 244
Deveau, A.E., 85
Devil, King, 184
Dezhi, L. and Q. Aili, Reviews by, 445
Dezhi, L., Q. Aili and Q. Hong, Reviews by, 108,111
Dezhi, L., Review by, 225
Diamond, A.W., 433
Dianthus armeria, 184
Dichanthelium columbianum, 203
depauperatum, 203
linearifolium, 203
perlongum, 203
sabulorum var. thinium, 203
Dickcissel, 245
Dicrostonyx groenlandicus, 120
nunatakensis, 120
Didelphis virginiana, 212,416
Didelphis virginiana, between the United States and Canada,
Human-assisted Movements of Raccoons, Procyon
lotor, and Opossums, 212
Diplophyllum albicans, 26
imbricatum, 26
plicatum, 26
taxifolium, 26
Dipoides, 331
Ditchkoff, S.S., 412
Dock, Curly, 184
Dodecatheon jeffreyi, 24
Dog, Domestic, 268,
Dogbane, Spreading, 203
Dogwood, Gray, 203
Red-osier, 359
Dolichonyx oryzivorus, 245
Donovan, D., 212
Dove, Mourning, 245,436
Dove, Zenaida macroura, Fish-line Entanglement of Nesting
Mourning, 436
Draud, M., 178
Dropseed, 184
INDEX TO VOLUME 12]
463
Dubois, J.E. and K.M. Monson. Recent Distribution Records
of the Little Brown Bat, Myotis lucifueus, in Manitoba
and Northwestern Ontario, 57
Duck, Harlequin, 82
Ring-necked, 245
Ruddy, 50
Duck Nests, Do Repugnant Scents Increase Survival of
Ground Nests? A Test with Artificial and Natural,
150
Duckweed, 166
Dumetella carolinensis, 244
Dunlin, 376
Dunn, J., 313
Duskysnail, Squat, 92
Duskysnail, Lyogyrus granum (Mollusca), Widespread in
the Hampton Marsh, New Brunswick, A Freshwater
Hydrobiid, cf. the, 92
Eagle, Bald, 376
Golden, 156
Echinacea pallida, 44
Echinochloa crusgalli, 183
Echinochloa, 184
Echium vulgare, 184
Editor’s Report for Volume 102 (2006), 235
Eel-grass, American, 164
Egret, Cattle, 50,330
Snowy, 50
Egretta caerulea, 54
thula, 50
Ehret, D.L., 40
Eider, Common, 275,288,308
Pacific Common, 308
Eider, Somateria mollissima, Nests on a Central Beaufort Sea
Barrier Island: A Case Where No One Wins, Depre-
dation of Common, 308
Elderberry, Red, 426
Eleocharis acicularis, 166
palustris, 166
Elk, 62,121,133,156,192,214,334,416
Elk, Cervus elaphus nelsoni, Populations, Colonization of
Non-Traditional Range in Dispersing, 133
Elk, Cervus Elaphus, Using Tooth Wear/Eruption Patterns
and Counts of Annuli in Tooth Cementum, Corre-
lation Between Age Estimates for, 214
Elk, Cervus elaphus, with Telezol and Xylazine and Reversal
with Tolazine or Yohimbine, Immobilization of, 62
Elliptio complanata, 93
Elm, American, 385
Elodea canadensis, 164
Elsinger, M., E. Burrell, N. deBruyn, K. Tanasichuk, and
K. Timoney. The Influence of Air Pollution on
Corticolous Lichens near the Strathcona Industrial
Area, Alberta, 17
Elymus repens, 183
trachycaulon ssp. trachycaulon, 183
trachycaulus ssp. subsecundus, 203
trachycaulus ssp. trachycaulus, 184
trachycaulus ssp. unilaterale, 203
Empetrum nigrum, 81
Empidonax minimus, 244
traillii, 244
Emydoidea blandingii, 178
Engelstoft, C., 142
Enhydra lutra, 376
464
Enteroctopus dofleini, 423
Enteroctopus dofleini, Attacks on Divers, Giant Pacific
Octopus, 423
Epigaea repens, 79
Epilobium spp., 359
ciliatum, 166
delicatum, 24
hirsutum, 184
Eptesicus fuscus, 120
Equisetum sp., 359
arvense, 184
hyemale, 184
palustre, 166
Eremontus myriocarpus, 26
Eremophila alpestris, 244
Erethizon dorsatum, 120,266,391,437
Erethizon dorsatum, Interaction in the Mackenzie Moun-
tains, Northwest Territories, Bull Trout, Salvelinus
confluentus, and North American Porcupine, 437
Erigeron annuus, 184
peregrinus, 24
philadelphicus ssp. philadelphicus, 184
strigosus, 184
Eriophorum spp., 134
angustifolium, 24
Ermine, 120,385
Erysimum cheiranthoides, 78
Eumeces, 217
fasciatus, 216
okadae, 216
Eumeces fasciatus, Predator-Prey Interaction Between an
American Robin, Turdus migratorius, and a Five-
lined Skink, 216
Euonymus alata, 85
americana, 85
atropurpurea, 85
europaea, 85
fortunei, 85
nana, 85
obovata, 85
occidentalis, 85
Euonymus europaea L. (Celastraceae): A Newly Naturalized
Shrub in Nova Scotia, The Spindle Tree, 85
Eupatorium maculatum, 184
perfoliatum, 184
rugosum, 184
Euphagus cyanocephalus, 245
Euphorbia helioscopa, 78
Euthamia graminifolia, 184
Everlasting, Pearly, 184
Evernia mesomorpha, 18
Fagus grandifolia, 304,385
Falco columbarius, 330
peregrinus, 82,330,376
sparverius, 216,413
tinnunculus, 413
Falcon, Peregrine, 82,330,376
Felis domesticus, 208,268
Fern, Northern Bracken, 204
Royal, 184
Sensitive, 184
Feverfew, 44
Field-Naturalists’ Club Awards for 2006, The Ottawa, 455
THE CANADIAN FIELD-NATURALIST
Vols121 a
Field-Naturalists’ Club 9 January 2007, Minutes of the 128th
Annual Business Meeting of The Ottawa, 350
Finley, J.K. Offshore Flight of Buffleheads (Bucephala
albeola) After Twilight in Winter: An Anti-Predation
Tactic?, 375
Finley, J.K. The Punctual Bufflehead, Bucephala albeola:
Autumn Arrivals in Shoal Harbour Sanctuary, Van-