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The CANADIAN 
rIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 115, Number 3 July-September 2001 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


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Date of this issue: July-September 2001 (April 2002). 


Cover: Prairie Falcons, Falco mexicanus, Edmonton, Alberta, February 1999. Photo by Robert Gehlert. See article by 
Dick Dekker and Jim Lange pages . 


a. —— 


The Canadian Field-Naturalist 


Volume 115, Number 3 July-September 2001 


Hunting Methods and Success Rates of Gyrfalcons, Falco rusticolus, 
and Prairie Falcons, Falco mexicanus, Preying on Feral Pigeons 
(Rock Doves), Columba livia, in Edmonton, Alberta” ° 


| SMITHSOsy 
Dick DEKKER! and Jim LANGE? iy 


13819-112A Street, Edmonton, Alberta T6J 1K4 Canada 
213216-128 Avenue, Edmonton, Alberta TSL 3H2 Canada 


JUN 1 4 2099 


Dekker, Dick, and Jim Lange. 2001. Hunting methods and success rates of Gyrfalcons, Falco rts 
Falcons, Falco mexicanus, preying on feral pigeons, (Rock Doves), Columba livia) in Edmonton, 
Canadian Field-Naturalist 115(3): 395-401. 


Three adult female Gyrfalcons (Falco rusticolus) and one adult female Prairie Falcon (Falco mexicanus), wintering in 
Edmonton, Alberta, killed 15 and 27 feral pigeons (Rock Doves, Columba livia) respectively in 141 and 104 hunts. The 
Prairie Falcon’s hunting success rate (26.0%) was significantly greater than that of the Gyrfalcons (10.6%). Six kills (40%) 
by Gyrfalcons and 18 kills (67%) by the Prairie Falcon were the result of surprise attacks on sitting pigeons. Open attacks 
on flying pigeons resulted in nine captures each. Gyrfalcons typically attacked airborne flocks of pigeons with upward 
swoops from below, whereas the Prairie Falcon stooped down at them from above. Both species often resumed hunting 
immediately or soon after consuming a pigeon. On one occasion, the Prairie Falcon killed four pigeons of which three were 
abandoned in 20 minutes. Besides capturing live prey, a Gyrfalcon stole a pigeon from the Prairie Falcon, and another 
Gyrfalcon scavenged four dead pigeons. 


Key Words: Gyrfalcon, Falco rusticolus, Prairie Falcon, Falco mexicanus, hunting methods, feral pigeon, Rock Dove, 


Columba livia, Edmonton, Alberta. 


Gyrfalcons (Falco rusticolus) take a wide range of 
avian and mammalian prey including Rock Doves 
(Columba livia) (Palmer 1988; Clum and Cade 1994), 
but descriptions of their hunting methods are general- 
ly based on few field observations and do not include 
hunting success rates (White and Weeden 1966; 
Bengtson 1971; Dobler 1989; White and Nelson 
1991; Garber et al. 1993). The diet of Prairie Falcons 
(Falco mexicanus) includes pigeons but consists 
mainly of small mammals and passerines (Anderson 
and Squires 1997; Steenhof 1998). The literature con- 
tains few data on hunting success rates of Prairie 
Falcons capturing avian prey. Dekker (1982) saw 36 
hunts including one kill. Beauvais et al. (1992) record- 
ed 37 hunts with five captures. To the best of our 
knowledge there is only one published account on the 
foraging habits of Gyrfalcons wintering in an urban 
environment (Jenning 1972), and none on Prairie 
Falcons. This paper presents a relatively large sample 
of hunting flights and success rates of Gyrfalcons and 
a Prairie Falcon preying on pigeons in Edmonton, 
Alberta, over two winters, 1998-2000. 

Like all big cities, Edmonton harbours a large, 
year-round population of feral Rock Doves. They 
commonly attract falcons. Urban Merlins (Falco 


columbarius), which prey mainly on small birds 
(James and Smith 1987), occasionally kill pigeons 
in Edmonton during winter (Lange 1985, and 
unpublished notes). Peregrine Falcons (Falco pere- 
grinus), which nest on high buildings in 
Edmonton, hunt pigeons but are absent from mid 
October to late March (Gordon Court, Alberta 
Environment). Gyrfalcons breed in arctic regions 
and some migrate south in winter (Godfrey 1986; 
Clum and Cade 1994). The earliest arrival date for 
central Alberta is 25 September, and sight records 
for the city of Edmonton range between November 
and March (Dekker 1983; Lange 1995). Prairie 
Falcons are breeding residents in southern Alberta 
up to the latitude of Red Deer (Godfrey 1986), and 
there is one (1978) nesting record from west of 
Edmonton (Gordon Court, Alberta Environment). 
Prairie Falcons are transients in north-central 
Alberta from spring to fall, but they summer in the 
Rocky Mountains up to the latitude of Jasper 
National Park, where they have been sighted in all 
months of the year (Dekker 1982, 1984, and 
unpublished field notes). This paper includes a first 
record of the Prairie Falcon wintering in the city of 
Edmonton. 


595 


396 


Study Area, Methods, and 
Description of Falcons 

The study area is centred on the Alberta Grain 
Loading Terminal (the Granary) that is situated 
along the railway corridor that transects the city of 
Edmonton (Figure 1). The 43 m elevator building 
includes an annex loading-shed. Its flat roof is lit- 
tered with spilled grain and canola seeds on which 
feral pigeons feed. Varying numbers, up to an esti- 
mated 600, roost on the tanks and chutes above the 
annex and on the window sills of the main building. 

At the approach of a falcon, most pigeons fly up 
and draw together in dense flocks that either course 
back and forth over the area or depart. Alerted by 
such anti-predator behaviour, the second author (JL), 
who works in the rail yards and lives near the 
Granary, was each year the first to sight wintering 
falcons. Beginning in mid-December 1998, we 
recorded all foraging activity (hunts). We defined a 
hunt as a completed attempt at capturing prey of 
which the outcome was known (Dekker 1980). A 
hunt could include one or more swoops or stoops at a 
pigeon or a flock of pigeons. If the falcon abandoned 
or interrupted its efforts by a spell of perching or by 
flying off, its next attack was considered another 
hunt. Thus, a hunting sequence could include more 
than one hunt. 

The hunts reported here combine those seen by 
both authors, alone or together. JL’s work often 
placed him in a position to observe falcon activity 


-— — —_—____ ern 
——S Se ee we ee es es es Ss es oe es Se 
SS 8 OS SD ES ES GS SE GS a ee Ge ee es es ees om ee 


Yellowhead Trail 


FIGURE 1. Location sketch of the Alberta Grain Terminal 
near the intersection of Yellowhead Trail and St. 
Albert Trail in northwest Edmonton, Alberta. The 
habitual perches of the white Gyrfalcon, the two 
grey Gyrfalcons, and the Prairie Falcon are indi- 
cated, respectively, by the symbols W, G, and P. 


THE CANADIAN FIELD-NATURALIST 


Vol ids 


- : ae é Wr 
FiGuRE 2. This recognizable, mostly white adult female 
Gyrfalcon, Falco rusticolus, roosted on a grain silo 
in Edmonton, Alberta, over six consecutive winters, 


1995-2000. Photograph by Gordon Court. 


during the week. In addition, on some days he 
watched from a parked vehicle just south of the 
Granary. DD watched from the same parking lot on 
37 days in 1998-1999 and on 7 days in 1999-2000, 
usually from 11:00 to 16:00. Particularly during 
weekends, one or more associate observers (listed in 
the acknowledgments) spent varying amounts of 
time watching for falcons at the Granary. The pigeon 
kills they reported to us contributed to our under- 
standing of the hunting methods used by these fal- 


cons. Weather conditions during observations varied a 
and daytime highs ranged from 5° to —30° C. The . 


ground was covered with 10 to 50 cm of snow. 


We saw at least three different adult female ; 
Gyrfalcons in the study area. One was mostly ~ 


white, the others grey variants (Cade et al. 1998). — 


The “white gyr” was lightly spotted on flanks and 
leg feathers. Dorsally, it was grey-brown with fine 
barring that gave it a “ladderback” appearance. The 
crown of its head was light brown unlike other 
white Gyrfalcons which have white heads (Brown 
and Amadon 1968). The white gyr was first seen on 
15 January 1995 when it was perched on a staircase 
railing near the top of the Granary (Lange 1995). It 
roosted in the same spot over the next six winters 
although it could be absent for several nights in a 
row. Use of specific roosting sites, with occasional 
absences of one or several days, was also typical of 
Gyrfalcons wintering in Washington State (Dobler 
1989). The white gyr usually arrived on its roost at 
dusk and was gone at first light. During the day, it 
was rarely seen until the winter of 1999-2000. 
Then, it often used an antenna on the roof of the 
Granary as a hunting perch (Figure 1). The last date 
on which the white gyr was seen each year varied 
from mid-March to 23 March. This is well past the 
time most adult Gyrfalcons are believed to return to 
arctic breeding sites (Platt 1976). At Beaverhills 
Lake, east of Edmonton, the latest Gyrfalcon sight- 


E 


2001 


ing, possibly of an immature, was 14 April (Dekker 
1983). 

Initially, we assumed the white gyr to be a male 
since it looked smaller, more slender and faster on 
the wing than a heavily streaked, grey-brown 
Gyrfalcon (gyr #2) which frequented the study area 
from 6 December 1998 to mid March 1999. Our 
assumption was proven wrong on | March 2000 
when Erhard Pletz captured and banded the white 
gyr. Its measurements (weight 1780 g, wing chord 
405 mm) placed it well within the range of female 
Gyrfalcons (Palmer 1988; Clum and Cade 1994). 

Gyr #2 was observed during the winter of 
1998-1999, but not in 1999-2000. Then, a different 
grey-brown Gyrfalcon (gyr #3) was present from 2 
December 1999 to mid March 2000. Ventrally, it 
was less heavily streaked and its feet were a paler 
yellow than gyr #2. Gyr #3 was identified as an adult 
by Gordon Court (Alberta Environment). On its left 
leg, it carried a dull grey (as opposed to a new and 
shiny) metal band. On 28 November 1998, a similar- 
looking grey Gyrfalcon, then a first-year immature, 
was banded by Erhard Pletz on Edmonton’s out- 
skirts. Its weight and wing chord were respectively 
1580 g and 400 mm. 

We saw no interactions between the white gyr and 
gyr #2, but often between the white gyr and gyr #3. 
The white gyr was slightly larger than #3 and chased 
it off aggressively. In addition to the individual birds 
described above, we saw two or three other 
Gyrfalcons in the city (Court 1999), but they were 
not in view long or often enough to allow for accu- 
rate details as to colour, age group, or sex. 

The first Edmonton record for the Prairie Falcon 
was 24 December 1995 (Lange, unpublished notes). 
Its yellow feet, large size and the dark axillar mark- 
ings running down much of the wing identified it as 
an adult female (Wheeler and Clark 1995). In this 
paper, we refer to it as “the” Prairie Falcon although 
we have no proof that all sightings involved the same 
individual. During the winters of 1996-1998, JL and 
associate observers EJ and JP saw the Prairie Falcon 


DEKKER AND LANGE: HUNTING AND SUCCESS OF GYRFALCONS AND PRAIRIE FALCONS 


397 


attack and capture at least eight pigeons at the 
Granary, but no detailed tally of hunts and kills was 
kept until the start of this study. During the winter of 
1998-1999 the falcon was present from 5 December 
to 26 February. In 1999-2000, it was first sighted on 2 
December but not after the end of December. 


Results and Discussion 
Gyrfalcon Hunting Methods 

Gyrfalcons commonly hunt very low over the 
ground, and they seize avian prey just after it flush- 
es. The target may be first spotted from an elevated 
perch (White and Weeden 1966; Bengtson 1971; 
Palmer 1988; Clum and Cade 1994). A similar strat- 
egy of surprise was also employed by the Gyrfalcons 
in this study. From a perch on the roof of the 
Granary, the falcons launched attacks on pigeons 
seeking food or grit along the railway or on snow- 
free parking lots in the neighbourhood. The hunt 
could be partially obscured from our view behind 
buildings so that its outcome could not be deter- 
mined unless the falcon came back into view, either 
with or without prey. From their perch on the 
Granary, falcons launched similar surprise attacks on 
pigeons sitting on the flat roofs of adjacent, much 
lower buildings. The above methods resulted in two 
captures (Table 1). Three similar kills were reported 
to us by associate observers EJ, BG and JG. In addi- 
tion, on a day when the number of pigeons at the 
Granary was small, BG saw gyr #2 drop down from 
its high perch and seize a pigeon just as it flushed 
from the roof of the annex. 

Surprise was also the initial strategy if the falcon 
approached the Granary from afar. Flying between 
adjacent buildings or along the railway, its sudden 
arrival caused the pigeons to flush in a panic from 
the annex or the front of the Granary. Typically, the 
falcon shot upward through the dense flock. In four 
instances, it seized a pigeon at once. Associate 
observers DL and JF reported two similar captures. 
In a variant of these surprise tactics, gyr #3 started 
its attack from a perch on the side of the Granary. 


TABLE 1. Captures of feral pigeons by Gyrfalcons and Prairie Falcons wintering in Edmonton, Alberta. 


Low surprise attack on pigeon(s) sitting on the ground 
Surprise attack on pigeon(s) sitting on flat-roofed buildings 


| 
Vertical swoop from high perch on building at pigeons sitting 25 m lower down - 
Sudden attack on flock flushed from building, first pass successful 4 
Repeated attacks on free-flying flocks, prey seized from flock 8 

if 


Pursuit of lone pigeon leaving flocks under attack 
Total 
Total number of hunts (successful and unsuccessful) 


Success rate 


*G-test, G=6.82. P< 0.01 (Sokal and Rohlf 1969) 


Prairie 
Gyrfalcon Falcon 
1 : 
10 
8 
8 
l 
15 27 
141 104 
10.6% 26.0%* 


398 


Flying around the corner of the building, it seized 
one pigeon from a flock that flushed from the annex 
below (JF). 

If the falcon’s first pass failed, it either (1) 
perched on the roof of the Granary; (2) left the area 
to hunt elsewhere; (3) went out of sight behind the 
Granary complex and came back suddenly around 
the corner to again attack flushing pigeons; or (4) 
made open attacks on the flying flock. In the latter 
case, it did so in a deceptively casual manner, flying 
back and forth near the careening pigeons. After 
gaining a strategic position in relation to its target, 
the falcon suddenly descended in a burst of speed 
with beating wings and, carried by its momentum, 
swooped upwards into and through the flock. 
Twisting aside or “standing on its tail,” practically 
stalling at the apex of its swoop, the falcon attempted 
to seize the nearest pigeon. If it failed to make a cap- 
ture, it turned back down and might repeat its 
upward swoops up to six times. Alternately descend- 
ing and ascending, the falcon typically attacked (as 
BG termed it) “like a pendulum.” We saw eight cap- 
tures. Associates reported at least another eight, all 
by the grey falcons. 

The white gyr rarely attacked flocks in the above 
described pendulum-swoops, and instead employed a 
very different hunting method: direct pursuits of lone 
pigeons or small groups passing by the Granary, 
often well over 100 m distant. Starting from its high 
perch on the antenna, the white gyr quickly over- 
hauled the pigeons, which dodged the attack at the 
very last moment. The falcon seldom made a second 
try at the same target. Despite its impressive speed, 
the white gyr was usually unsuccessful. On 17 
February 2000, over a three-hour period, it alternated 
at least 12 attacks on single pigeons and flocks with 
spells of perching, yet it failed to make a kill. On 20 
February, over two hours, it made about 20 futile 
hunts. We saw one kill by the white gyr. Here too, 
surprise may have played a role. Breaking off its 
attack on a flock, the gyr flew about 50 m in direct 
pursuit of a single pigeon that had left the flock. 
Failing to dodge, the pigeon was seized from behind. 
Gyr #2 never tried to capture single pigeons that 
flew by while it was perched on the Granary. Gyr #3 
rarely did so and always without success. 


Prairie Falcon Hunting Methods 

Prairie Falcons take a wide variety of prey, includ- 
ing pigeons (Palmer 1988). They often soar to great 
altitudes and descend in long glides to surprise small 
mammals and ground-dwelling birds, but during 
winter they generally hunt low over open terrain 
(Dekker 1982; Anderson and Squires 1997; Steenhof 
1998). The Prairie Falcon observed in this study 
attacked prey sitting on large city buildings, and sur- 
prise was the initial strategy in 67% of its kills. Its 
- most common method was a sudden approach to the 
Granary. We rarely spotted the falcon before the 


THE CANADIAN FIELD-NATURALIST 


Vol us 


pigeons flushed in alarm. Swooping through the 
flock, the falcon could be immediately successful 
(Table 1). If it failed, it often took up a perch on the 
Granary. Unlike the Gyrfalcons which sat on the 
roof corners or the antenna, the Prairie Falcon 
always perched on a metal bracket supporting the 
vertical pipes that ran down the front of the Granary 
(Figure 1). The falcon’s perch was below the roof 
line but still about 25 m above the annex. After the 
pigeons had settled back onto the flat roof and 
machinery below, the falcon suddenly dropped off 
its perch and stooped down perpendicularly. As the 
pigeons flared away from the building, the falcon 
passed through them and might at once emerge car- 
rying its prey. In a similar attack, when few pigeons 
were present, the stooping falcon passed in between 
the machinery above the annex, levelled off just over 
the flat roof and seized a sitting pigeon from behind, 
carrying it along without pause. 

If its surprise tactics failed, the falcon either (1) 
returned to its bracket perch and tried again 5—25 
minutes later; (2) left the area to hunt elsewhere; or 
(3) pressed its attack on flocks that had already been 
flushed. This latter hunting method did not involve 
surprise as the primary strategy. The falcon, either 
flapping its wings or sailing and soaring, typically 
stayed close to or above the flock or flocks of 
pigeons that were careening back and forth. 
Selecting an individual target on the outside of the 
flock, the falcon then stooped down with rigid 
wings, either fully extended or tucked in close to the 
tail. If the pigeon dodged the attack, the falcon 
pulled up and directed subsequent stoops at other tar- 
gets. Some hunts included five or six stoops. A hunt- 
ing sequence could last five or ten minutes with the 
falcon sailing away some distance and returning to 
attack again, perhaps a different flock. Meanwhile, 
some pigeons might settle back onto the Granary, 
while others had remained there, crouching on win- 
dow sills or machinery. The falcon flushed these 
pigeons by stooping close to the building or passing 
in between the pipes and tanks above the annex. If it 
attacked the flushed pigeons, it seldom chased them 
farther than 10—30 m. 

On 4 January 1999, the Prairie Falcon made a 
series of four kills in about twenty minutes (DD, GC, 
JF). The first three prey were taken from flocks at 
the falcon’s first pass and carried to a nearby snow- 
covered parking lot. They were abandoned a few 
minutes later, probably because of the approach of 
vehicles. Each time, the falcon flew directly back to 
the Granary and seized its next prey from the flush- 
ing flock. After its third kill, the falcon perched 
briefly on a pole by the parking lot. It then returned 
to the Granary and caught a fourth pigeon, carrying 
it out of sight. By searching the parking lot and fol- 
lowing a trail of blood drops, GC and DD located the 
falcon’s three previous kills lying in 30 cm of soft 
snow. 


2001 


Food Habits, Scavenging, and Kleptoparasitism 

On several occasions, a Gyrfalcon attacked the 
pigeons at the Granary immediately after it had 
killed and consumed one. The Prairie Falcon, after 
we had observed it capture and eat a prey, twice flew 
to its usual perch with a slightly bulging crop and 
began attacking the pigeons about 25 min later. On 
both occasions, it killed and fed upon a second 
pigeon within two hours from the first. As Rock 
Doves weigh 250-400 g, the falcons in this study 
apparently killed well in excess of daily food 
requirements which are reported to be 250 g for 
Gyrfalcons and 170 g for Prairie Falcons (Craighead 
and Craighead 1956; Palmer 1988). 

All kills we saw, including those reported by asso- 
ciates, occurred between 10:30 and 16:30. Although 
we saw no hunting or feeding activity before 10:00, 
that possibility remains to be investigated more 
closely. In this regard it may be significant that the 
white gyr, on the day it was trapped, came down to 
the lure pigeon at first light. The previous night it 
had arrived at its roost with a bulging crop as proof 
that it had eaten well (Erhard Pletz, personal com- 
munication). 

Both species of falcon carried their freshly-caught 
pigeons to open terrain such as the parking lot adja- 
cent to the Granary. We did not see them pluck prey 
on poles or buildings. The Gyrfalcons did not avoid 
snow of 20-40 cm deep, but the Prairie Falcon usu- 
ally selected bare ground along the railway. The fal- 
cons sometimes carried their prey to the Municipal 
Airport, about one km away, where the runways 
were mostly clear of snow. 

There appeared to be little overt antagonism 
between the Gyrfalcons and the Prairie Falcon. Gyr 
#2 occasionally chased the Prairie Falcon. At other 
times, both perched on the Granary at the same time. 
Opportunistic klepto-parasitism might be common 
but we saw only one such incident. On 13 February 
1999, the Prairie Falcon caught a pigeon at the 
Granary. Gyr #2 was perched on the roof and took 
off in pursuit. After the loss of its prey, the Prairie 
Falcon at once returned to the Granary and caught 
another pigeon out of the flushing flock. We did not 
see any klepto-parasitism among the Gyrfalcons. 

Gyrfalcons are known to feed on carrion in their 
northern breeding grounds (Dementiew 1960; 
Palmer 1988; Cade et al. 1998). On 19 February 
2000, after eight unsuccessful attacks on flocks of 
pigeons, gyr #3 landed on the flat roof of the annex, 
walked to the remains of a dead pigeon, and flew off 
with it. On 21 February, it retrieved two dead 
pigeons in about forty minutes. As proof that it had 
eaten from the first carcass, the gyr showed a 
bulging crop. On 22 February, it again retrieved a 
dead pigeon from the annex. Clutching the carcass in 
its feet, the gyr soared over the area and drifted 
away. 


DEKKER AND LANGE: HUNTING AND SUCCESS OF GYRFALCONS AND PRAIRIE FALCONS 


399 


On 24 February, in preparation for an unsuccess- 
ful attempt to trap gyr #3, GC removed eight pigeon 
carcasses from the annex. Apparently, their death 
had been accidental, caused by interference with roof 
fans. However, it is possible that some of the 
pigeons had died as a result of wounds caused by fal- 
con attack. Once, as gyr #2 swooped upwards 
through a flock, a pigeon dropped out and fell like a 
stone in between the machinery above the annex. 
The falcon made no effort to retrieve it. Other 
pigeons might have succumbed to injuries sustained 
during attacks. Some pigeons flapped their wings 
while being carried off by the falcon. Both the 
Gyrfalcons and the Prairie Falcon killed some of 
their prey by biting it during flight. One struggling 
pigeon freed itself from the gyr’s grasp after it had 
been carried about 100 m. The gyr did not pursue it 
but returned to the Granary. 

In one instance we saw the Prairie Falcon strike a 
flying pigeon a mortal blow that caused it to stop 
beating its wings and make a tumble. The falcon 
instantly doubled back and caught the pigeon before 
it had fallen more than 3 m (DD and BG). In all 
other captures the Prairie Falcon directly seized the 
prey in its feet. Contrary to reports that Gyrfalcons 
commonly strike prey in the air, so that it drops to 
the ground (Dementiew 1960; Clum and Cade 1994), 
the Gyrfalcons in this study seized their prey direct- 
ly. In close but unsuccessful attacks, the pigeons 
might lose a puff of feathers, or the falcons trailed 
feathers from their claws even though all pigeons 
flew on. Such damage was likely the result of misdi- 
rected attempts at seizing prey. Loose feathering in 
pigeons may be an anti-predator adaptation. Seizing 
prey, as opposed to striking them down, is also the 
most common capture method of the Peregrine 
Falcon (Dekker 1999). 

The avian prey of Gyrfalcons consists mainly of 
grouse and waterfowl (Brown and Amadon 1968; 
Palmer 1988; Clum and Cade 1994), which are less 
manoeuvrable in flight than pigeons. Jenning (1972) 
noted that a Gyrfalcon wintering in Stockholm, 
Sweden, ignored pigeons. Bent (1937-1961: 2) cited 
three eyewitness accounts to the effect that Gyr- 
falcons often tried but “never succeeded in capturing 
a pigeon.... which were more than a match for them.” 
This contention is supported by some of our observa- 
tions, such as the futility of the white gyr’s attacks 
on single flying pigeons. However, the overall suc- 
cess of the Edmonton Gyrfalcons was 10.9%, which 
is close to the mean (12.7%) success rate of 13 stud- 
ies of wintering and migrating Peregrine Falcons 
(Roalkvam 1985). The fair rate of hunting success 
demonstrated by the gyrs in this study was related to 
the effective use of surprise and to their persistence 
in attacks on flying flocks. The “pendulum-style” 
swoops reported in this paper have not been 
described before. 


400 


In its swoops on flocking pigeons, the Prairie 
Falcon seemed capable of capturing its prey at will. 
Flocking by pigeons and other birds is an anti-preda- 
tor strategy. However, in our study it was apparent 
that the pigeons’ habit of flocking together, instead 
of fleeing, rendered them particularly vulnerable, 
both to the Gyrfalcons and the Prairie Falcon. Hunt- 
ing success rates reported for the Peregrine Falcon, 
for which there is a great amount of data from 
around the world, tend to show a wide range (Cade 
1982; Roalkvam 1985) unless sample sizes are large 
and criteria consistent (Dekker 1999). Comparisons 
of hunting success rates, even within one species, let 
alone between different species of falcon, mean little 
unless they hunt the same kind of prey, at the same 
locality, and during the same season. In this study, 
the success rate of the Prairie Falcon in capturing 
pigeons was significantly greater (Table 1) than that 
of the Gyrfalcons. This fits the theory that a falcon 
pursuing prey in the air should have a body mass 
close to that of the prey in order to match as closely 
as possible the speed and manoeuvrability of the 
prey (Andersson and Norberg 1981; Cade 1982). 


Acknowledgments 

Numerous birdwatchers and biologists took 
advantage of the opportunity to watch Gyrfalcons 
and Prairie Falcons at the Granary. The following 
contributed information about the pigeon kills they 
observed: John Acorn (JA), Gordon Court (GC), 
John Folinsbee (JF), Bob Gehlert (BG), John Groves 
(JG), Edgar Jones (EJ), Dennis Lavallee (DL), Jack 
Park (JP), Lisa Takats (LT), and Fred Wiley (FW). 
Tom J. Cade and Gordon Court reviewed the manu- 
script and made helpful suggestions. Gordon Court 
did the statistical test. Erhard Pletz contributed 
weights and measurements of trapped Gyrfalcons. 
Wayne Nelson provided relevant literature. We 
thank Francis Cook, Anthony Erskine, Stuart 
Houston, and one other, anonymous, reviewer for 
their comments. 


Addenda 

During the very mild winter of 2000—2001, the 
white Gyrfalcon failed to return to the Edmonton 
Granary, and sightings of grey gyrs were rare in the 
city. A Prairie Falcon, seen at the Granary on 13 
March by DD, proved to be a different individual, 
based on plumage characteristics, than the bird seen 
in 1998-2000. Yet, it used exactly the same perch on 
the Granary and attacked the pigeons in similar style. 
It appeared to be less accomplished, however, as all 
of its 18 hunts, made over a three-hour period, were 
unsuccessful. 

During the winter of 2001-2002, a Prairie Falcon 
was again frequently seen attacking pigeons at the 
granary, but Gyrfalcon sightings were uncommon 
(JL). 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


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predatory birds, with a size scaling of flight perfor- 
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105-130. 

Anderson, S. H., and J. R. Squires. 1997. The Prairie 
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Bengtson, S. A. 1971. Hunting methods and choice of 
prey of Gyrfalcons, Falco rusticolus, at Myvatn in 
Iceland. Ibis 113: 468-476. . 

Bent, A. C. 1937. Life histories of North American birds 
of prey. Part 2. Dover Publications, New York. Reprint 
1961. 482 pages. 

Brown, L., and D. Amadon. 1968. Eagles, hawks and fal- 
cons of the world. Hamiyn House, Feltham, Great 
Britain. 945 pages. 

Cade, T. J. 1982. The falcons of the world. Cornell 
University Press, Ithaca, New York. 192 pages. 

Cade, T. J., P. Koskimies, and O. K. Nielsen. 1998. 
Gyrfalcon (Falco rusticolus). Birds of the Western 
Palaearctic 2(1). Oxford University Press. 

Clum, N. J., and T. J. Cade. 1994. Gyrfalcon (Falco rus- 
ticolus). In The birds of North America, Number 114. 
Edited by A. Poole and F. Gill. The Academy of Natural 
Sciences, Washington, D.C. 

Court, G. 1999. Edmonton’s year of the gyr. Edmonton 
Naturalist 27(1): 29-30. 

Craighead, J. J.. and F. C. Craighead. 1956. Hawks, 
owls and wildlife. Dover Publications. New York. 443 
pages. 

Dekker, D. 1980. Hunting success rates, foraging habits, 
and prey selection of Peregrine Falcons migrating 
through central Alberta. Canadian Field-Naturalist 94: 
371-382. 

Dekker, D. 1982. Occurrence and foraging habits of 
Prairie Falcons, Falco mexicanus, at Beaverhills Lake, 
Alberta. Canadian Field-Naturalist 96: 477-478. 

Dekker, D. 1983. Gyrfalcon sightings at Beaverhills Lake 
and Edmonton, 1964-1983. Alberta Naturalist 13: 103. 

Dekker, D. 1984. Prairie Falcon sightings in the Rocky 
Mountains of Alberta. Alberta Naturalist 14: 48-49. 

Dekker, D. 1999. Bolt from the blue — Wild peregrines 
on the hunt. Hancock House Publishers, Surrey, British 
Columbia. 192 pages. 

Dementiew, G. P. 1960. Der Gerfalke. Die Neue Brehm- 
Biicherei. Heft 264. Ziemsen Verlag, Wittenberg, 
Germany. 

Dobler, F.C. 1989. Wintering Gyrfalcon (Falco rustico- 
lus) habitat utilization in Washington. Pages 61—70 in 
Raptors in the modern world. Edited by B. Meyburg and 
R. D. Chancellor. World Working Group on Birds of 
Prey. Berlin, London, Paris. 

Garber, C.S., B. D. Mutch, and S. Platt. 1993. Obser- 
vations on wintering Gyrfalcons (Falco rusticolus) hunt- 
ing Sage Grouse (Centrocercus urophasianus) in 
Wyoming and Montana, U.S.A. Journal of Raptor Re- 
search 27: 171-172. 

Godfrey, W. E. 1986. The birds of Canada. Revised 
Edition. National Museum of Natural Sciences, Ottawa. 
595 pages. 


2001 


James, P. C., and A. R. Smith. 1987. Food habits of 
urban-nesting Merlins (Falco columbarius) in Edmonton 
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Jenning, W. 1972. laktagelser rodrande en overwintrande 
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Lange, J. 1985. Merlin kills pigeon. Alberta Naturalist 
15(4): 134. 

Lange, J. 1995. Observing gyrfalcons from the living 
room sofa. Edmonton Naturalist 23(2): 12-13. 

Palmer, R.S. 1988. Handbook of North American birds, 
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Platt, J.B. 1976. Gyrfalcon nest site selection and winter 
activity in the western Canadian arctic. Canadian Field- 
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Roalkvam, R. 1985. How effective are hunting pere- 
grines? Journal of Raptor Research 19: 27-29. 

Sokol, R.R., and F. J. Rohlf. 1969. Biometry. Freeman 
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Steenhof, K. 1998. Prairie Falcon (Falco mexicanus). The 
Birds of North America, Number 346. Edited by A. 
Poole and F. Gill. The Birds of North America, Inc. 
Philadelphia, Pennsylvania. . 

Wheeler, B. K., and W.S. Clark. 1995. North American 
Raptors. Academic Press Limited. London and San 
Diego. 198 pages. 

White, C. M., and R. B. Weeden. 1966. Hunting methods 
of Gyrfalcons and behaviour of their prey. Condor 68: 
517-519. 

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and strategies in a tundra-breeding Peregrine and 
Gyrfalcon observed from a helicopter. Journal of Raptor 
Research 25(3): 49-62. 


Received 7 July 2000 
Accepted 11 May 2001 
Addenda | March 2002 


Biodiverety of Adult Damselflies (Zygoptera) at Eastern Ontario 
Gravel Pit Ponds 


P. M. CATLING and V. R. BROWNELL 
2326 Scrivens Drive, R.R. 3, Metcalfe, Ontario KOA 2PO Canada 


Catling, P. M., and V.R. Brownell. 2001. Biodiversity of adult damselflies (Zygoptera) at eastern Ontario gravel pit 
ponds. Canadian Field-Naturalist 115(3): 402-405. 


Adults of twenty-five species of damselflies were recorded at 41 gravel pit ponds in eastern Ontario. Twenty-four of the 
species recorded are believed to breed in the ponds where they were captured. Species present at 16 or more of the sites 
included Enallagma boreale, E. civile, E. cyathigerum, E. ebrium, E. hageni, Ischnura verticalis, Lestes forcipatus, L. 
unguiculatus, and Nehalennia irene. Two provincially and regionally rare species, Lestes eurinus and Enallagma 
aspersum, were abundant in some of the pit ponds. With 70% of the eastern Ontario zygopteran fauna, these naturally colo- 
nized sites may serve as important habitats for the conservation of biodiversity. The relatively high overall diversity com- 
pared to the much lower within-pond diversity is probably related to variation between ponds in flora, fauna, depth and 
other characteristics. Species specific associations with characteristics of the water body other than chemistry, may make 


damselflies a valuable group of bioindicators. 


Key Words: Biodiversity, bioindicator, ecology, damselfly, Zygoptera, Odonata, Argia, Enallagma, Ischnura, Lestes, 
Nehalennia, rare species, Ontario, gravel pit, pond, wetland, habitat, conservation, protection. 


Wetlands in parts of southern Ontario have been 
reduced to 5% of their previous extent, and the 
decline continues (e.g., Snell 1987, 1989). In some 
areas ponds created through aggregate extraction may 
become significant habitats for wetland flora and 
fauna. For example, 18 species of damselflies were 
recorded over a three-year period from a single 20- 
year-old gravel pit in southwestern Quebec (Pilon et 
al. 1988). It appears that these gravel pit ponds may be 
valuable in conserving aquatic insect diversity. 
Unfortunately pit ponds are often filled to avoid liabil- 
ity, or to meet the requirements of Ontario’s 
Aggregate Act, or as part of general restoration pro- 
cesses. An underlying lack of recognition of the value 
of pit ponds in biodiversity conservation is evident. 
The present study further explores the value of aban- 
doned pit ponds that have been naturally colonized, as 
habitats for damselflies. In addition it is intended to 
contribute to an ecological framework that will enable 
damselflies to be used more effectively as bioindica- 
tors, as well as to provide a basis for ongoing studies 
of biodiversity of man-made ponds that will involve 
large sample sizes permitting objective analysis of 
some of the trends suggested here. 


Methods 

The number of species and the number of individu- 
als of each species of adult damselfly was recorded at 
each of 41 eastern Ontario gravel pit ponds (Table 1) 
during two visitation periods; 15—28 August 1996 
and 14-22 June 1997. The ponds ranged from one to 
30 acres in size, were 2—80 years old and had a pH 
range of 7-8. One-half hour to one hour was spent at 
each site during each visitation period. There had 
been no purposeful rehabilitation of any of the ponds. 


Size, depth, shoreline characteristics and presence of 
frogs and fish (some of which were introduced) were 
recorded at each pond to enable tentative correlations 
with damselfly species occurrences. 

As many adult specimens were captured as possi- 
ble at each site. These were identified and released at 
the end of the sample period. Sore voucher speci- 
mens were retained and placed in the national collec- 
tion at Agriculture Canada (CNC). Specimens were 
identified using Walker (1953) and Westfall and 
May (1996) and tabulated for each site. “Sranks” 
denoting relative abundance in Ontario were taken 
from Oldham et al. (2000): S1 = critically imperiled 
(5 or fewer occurrences), S2 = imperiled (6—20 
occurrences), S3 = vulnerable (21-100 occurrences), 
S4 = apparently secure (more than 100 occurrences) 
and S5 = secure. Authorities for all scientific names 
used in the text are found in Table 2. 


Results and Discussion 
Species and overall biodiversity 

Twenty-five species of damselflies were recorded 
at 41 pit ponds (Table 2). In almost all cases the 
species recorded are believed to have developed in 
the ponds where they were captured. Some species, 
such as Enallagma aspersum were recorded infre- 
quently but were very abundant at some of the sites 
where they were found (Table 2). Other species were 
not found in large numbers but were still associated 
with the pond where captured due to their teneral 
condition or the fact that they were observed in cop- 
ula (e.g., Chromagrion conditum). The low number 
of E. vesperum observed was probably a result of 
observation during mid-day instead of during the 
evening when this species is most active. Amphia- 


402 


2001 CATLING AND BROWNELL: DAMSELFLIES AT GRAVEL Pit PONDS 403 
TABLE |. Names, locations, and numbers of damselfly species recorded for 41 eastern Ontario gravel pit ponds. 
Location Number 
number Name County Township Latitude Longitude of species 
Central dry Ottawa-Carleton Gloucester 45°16'34" 75°34'44" | 
2 shallow pond Ottawa-Carleton Gloucester 45°16'57" 75°34'53" 3 
3 w shallow pond Ottawa-Carleton Gloucester 45°16'50" (pec eal 5 
4 Twin Ponds Ottawa-Carleton Gloucester 45°16'47" 75°34'39" 7 
5 Clear Pond Ottawa-Carleton Gloucester 45°16'31" Taree. 5 
6 Upper Clear Ottawa-Carleton Gloucester 45°16'34" 75°35'30" 5 
7 Church Dry Ottawa-Carleton Gloucester 45°17 3" 1S 30.0 | 
8 Johnstons Corner Ottawa-Carleton Gloucester ASAT ae 75°35'48" 8 
9 Middle Pit Ottawa-Carleton Gloucester 45°16'44" Ta ID Aas 7 
10 North Dry Ottawa-Carleton Gloucester 45°16'44”" 73 3'43" 2 
1] South Dry Ottawa-Carleton Gloucester 45° 16/25" jel wai 5 
12 South Greely Ottawa-Carleton Osgoode 45°16'5” fa ie Ne 6 
13 Kemptville Leeds, Grenville South Gower 45°0'35" ef tars be 14 
14 Old Pond Ottawa-Carleton Gloucester 45°17'20" jake pw low 8 
15 Big Dry Ottawa-Carleton Gloucester 45°16'47" Tse 30 De 7 
16 K-Hwy 43 Leeds, Grenville South Gower AS°2' 32h 75°34'40" 8 
17 K-Shrubby Pond Leeds, Grenville South Gower 45°14” 75°34'34" 8 
18 Herbert-Lake Ottawa-Carleton Osgoode 45°14'22” 75°34'47" 5 
19 Herbert-Pit Ottawa-Carleton Osgoode 45°14'12” 75°34'42" q 
20 Herbert-Dry Ottawa-Carleton Osgoode 45°14’9" 75°34'10" 10 
21 Long Pond Osgood Ottawa-Carleton Osgoode 45°9'16" (ascewlily ii 
22 Loughlin Dry Stormont, Dundas Mountain 45°3'17" T3233) 59" 5 
23 Moore Middle Leeds, Grenville South Gower 45°1'8" 75°33'40" 7 
24 Moore North Leeds, Grenville South Gower AS Ae (Bees 9 
25 Moore Small Leeds, Grenville South Gower Ae Die dae Sage 3 
26 Drew Dry West Ottawa-Carleton Osgoode 45°9'7" 75°34'7" 10 
27 Drew Dry East Ottawa-Carleton Osgoode 45°9'14" 75°34'48" 8 
28 Cory Drew Pond Ottawa-Carleton Osgoode 45°9'29" TS°34, 22) 8 
29 Sunset Lake N Ottawa-Carleton Osgoode 45°15'59" TS 34°57" 3 
30 Greely Bro Pit Ottawa-Carleton Osgoode 45°13'53” 73°33) 29" 5 
31 Big Shallow Leeds, Grenville South Gower 45°1'40” ao 4 55" 8 
32 West Shallow Leeds, Grenville South Gower AS WVAG" 75°34'44" 9 
33 East Shallow Leeds, Grenville South Gower 45°1'46” WSS4 SS 5 
34 Sunset Lake S Ottawa-Carleton Osgoode 45°15'56" 75°34'43" 5 
35 Bruces Pit Ottawa-Carleton Nepean 45°19'31" 75°48'14" 12 
36 Burnside Lake Ottawa-Carleton Nepean 45°13'9" 75°46'14" 5 
37 Costello Pit Ottawa-Carleton Nepean 45°14'18” 75°45'11" 5 
38 Stanstead Pit Leeds, Grenville Nepean 44°47'40" Soule g 
39 Moore South Leeds, Grenville South Gower 4520/52, TS se Cae 7 
40 North Dump Pond Leeds, Grenville South Gower 45°1'40" 75°34'48" 11 
41 Quinn Pond Ottawa-Carleton Gloucester 45°17'55" (eeu S10y 6 


grion saucium, although present in only one pit 
pond, was well established in marshy springs along 
the shoreline. Enallagma exsulans was found at only 
one pond but it was a large pond with much ground 
water flow and shoreline wave action, perhaps simu- 
lating the streams where E. exsulans characteristical- 
ly occurs. Enallagma vernale is the only species that 
may not have developed in the pond where it was 
captured. It is characteristically associated with 
ponds having a deep, organic bottom unlike its close 
relative E. cyathigerum which is characteristic of pit 
ponds with sandy bottoms. 

The occurrence of twenty-four (not including E. 
vernale) of the 34 species occurring in the local area 
(Catling and Brownell 1997*; Ménard 1996) [i.e., 


70% of the eastern Ontario damselfly fauna] indi- 
cates that pit ponds serve as an important habitat for 
maintaining damselfly biodiversity. The number of 
species found at eastern Ontario pit ponds represents 
more than 50% of the damselfly species present in 
all of Ontario (47 species of which eight are con- 
fined to the Carolinian Zone north of Lake Erie). 
The relatively diverse damselfly fauna in pit 
ponds may be explained in terms of ecological vari- 
ability at these sites. Although similar in water 
chemistry, in having few floating and emergent plant 


*See Documents Cited section. 


404 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


TABLE 2. Species recorded in 41 eastern Ontario gravel pit ponds indicating number of ponds recorded in and 
total number of individuals recorded, and “S” rank. 


Number Number of 
Scientific Name Common Name of Ponds Individuals SRank 
Ischnura verticalis (Say) Eastern Forktail Sy 1163 S5 
Lestes unguiculatus Hagen Lyre-tipped Spreadwing 31 424 S5 
Nehalennia irene (Hagen) Sedge Sprite De 367 S5 
Enallagma civile (Hagen) Familiar Bluet 21 287 S4 
Enallagma ebrium (Hagen) Marsh Bluet 18 904 So 
Lestes forcipatus Rambur Sweetflag Spreadwing 16 394 S485 
Enallagma boreaile Sélys Boreal Biuet a 384 S5 
Enallagma cyathigerum (Charpentier) Northern Bluet 17 455 S4 
Enallagma hageni (Walsh) Hagen’s Bluet 16 356 S5 
Lestes congener Hagen Spotted Spreadwing 12 154 S5 
Lestes dryas Kirby Emerald Spreadwing 11 Bg S5 
Enallagma carunculatum Morse Tule Bluet 11 36 S5 
Lestes disjunctus Sélys disjunctus Common Spreadwing 6 54 SS 
Enallagma aspersum (Hagen) Bog Bluet 6 873 S1S2 
Argia fumipennis violacea (Hagen) Violet Dancer 5 PES) S5 
Coenagrion resolutum Sélys Taiga Bluet 5 43 S5 
Lestes eurinus Say Amber-winged Spreadwing 4 45 S1S2 
Enallagma antennatum (Say) Rainbow Bluet 3 19 S5 
Lestes rectangularis Say Slender Spreadwing 2 10 S5 
Amphiagrion saucium (Burmeister) Eastern Red Damsel 1 oe S5 
Chromagrion conditum (Sélys) Aurora Damsel 1 + S5 
Enallagma vernale (Gloyd) Spring Northern Bluet 1 1 33? 
Enallagma exsulans (Hagen) Stream Bluet 1 2 SS 
Enallagma signatum (Hagen) Orange Bluet 1 2 S4 
Enallagma vesperum Calvert Vesper Bluet i 3 S4 


species, and in lacking quaking mats of vegetation 
and deep organic sediments, they differ in age, size, 
shoreline and vegetation structure, depth, perma- 
nence, and presence or absence of predators and/or 
prey including waterfowl, fish, frogs and inverte- 
brates. Different species of damselflies may be 
adapted to different combinations and/or characteris- 
tics of these features. For example, Argia fumipennis 
violacea occurred only in pit ponds with rocky 
shores; Lestes unguiculatus and, to a lesser extent, 
Lestes dryas and L. forcipatus occurred around tem- 
porary ponds which dried out in late summer; and 
Enallagma aspersum occurred in permanent ponds 
that were mostly less than 0.5 m deep when sampled. 
This concept of different pit ponds offering a variety 
of different niches is supported by the relatively 
large number of species recorded overall in compari- 
son with the relatively small number at any one site 
(Table 2). 


Diversity within a pond 

Considering that many damselfly species can be 
common in some ponds, but rare in others, it may 
only be through extensive field study that all of the 
rare species at a particular site will be recorded. With 
an average of only 2 hours spent observing at each 
site in the present study, it is possible that rare species 
were overlooked. Consequently the individual site 
diversity is probably under-represented. It is notable 


in this context that with more extensive field study of 
a single gravel pit from spring to fall over a three-year 
period, Pilon et al. (1988) found 18 species. The num- 
ber of species recorded from a site in the present study 
ranged from | to 14 (Table 1), with an average of 6.41 
and a standard deviation of 2.80. Ponds that were very 
recently created and entirely temporary had the fewest 
species. Other factors influencing diversity at a pond 
appeared to include abundance of frogs and fish as 
well as pond size. 


Characteristic fauna 

Ontario damselflies can be split into two major eco- 
logical groups; those of streams and rivers (including 
the genera Calopteryx, Hetaerina, and most species of 
Argia as well as Enallagma exsulans), and those of 
ponds. The latter group can be subdivided into at least 
two subgroups; (1) widespread and common species 
of many kinds of ponds, (2) species of older ponds 
with deep organic layers and much floating and emer- 
gent vegetation. The latter subgroup includes species 
such as Enallagma geminatum, E. vernale, Lestes 
inaequalis and Lestes vigilax, all of which were con- 
spicuously absent from the present survey, although 
within range (Catling and Brownell 1997*). The 
species of pit ponds may be assigned to the former 
subgroup, and species present at most of the sites may 
be considered as a characteristic fauna. Enallagma 
boreale, E. civile, E. cyathigerum, E. ebrium, E. hag- 


2001 


eni, Ischnura verticalis, Lestes forcipatus, L. unguicu- 
latus, and Nehalennia irene were present at 16 or 
more of the sites. Some of the rarer species may also 
be characteristic of certain kinds of pit ponds; e.g., 
Enallagma aspersum may be characteristic of uni- 
formly shallow pit ponds. 


Rare species 

Until recently Enallagma aspersum was consid- 
ered to be very restricted in Ontario and confined to 
bog ponds, but it has recently been found in a num- 
ber of man-made habitats and is now known to be 
widespread (Catling and Pratt 1997). Although 
widespread and less restricted by habitat than previ- 
ously thought, it is still a rare species that is current- 
ly known from less than 25 locations. Its abundance 
in some pit ponds, where hundreds can be seen with- 
in a few minutes, make these ponds an important 
habitat and also provides an opportunity to study the 
population dynamics of this interesting species. 

Another uncommon species previously associated 
with bogs, Lestes eurinus, is also well established in 
some pit ponds. Although it is more common than 
previously suspected, it is currently known from less 
than 20 localities in southern Ontario and is certainly 
one of the rarest species of Lestes in the province. 
The pit ponds provide an important opportunity to 
study its ecology. 

In eastern Ontario Argia fumipennis violacea is a 
very restricted species. In areas of several townships it 
occurs only on the rocky shores of gravel pit ponds. 
Thus the pit ponds contribute to the protection of rare 
species on both a local and provincial scale. 

The presence of rare species as well as high over- 
all biodiversity of damselflies suggest that naturally 
colonized ponds in abandoned gravel pits are poten- 
tially important sites for biodiversity conservation 
and research. The results provide further support for 
the general idea (Fox and Cham 1994; Chovanec and 
Raab 1997) that “artificial wetlands” created accord- 
ing to specific ecological principles can serve as 
important sites for biodiversity conservation espe- 
cially with respect to Odonata. Since these ponds 
have a high collective overall diversity of damsel- 
flies, but a low within-pond diversity, they appear to 
be more variable habitats than their similarities in 
water chemistry and history would suggest. While 
certain groups of organisms, such as plants, are use- 
ful bioindicators of water chemistry, damselflies 
may prove to be valuable indicators of other water 
body characteristics such as permanence, depth, 
structure, predators and prey. 


CATLING AND BROWNELL: DAMSELFLIES AT GRAVEL PIT PONDS 


Acknowledgments 

B. Ménard and R. Hutchinson assisted in the field. 
G. Drew, G. W. Drummond, R. Fortin, E. Moore, 
and F. Cramp all facilitated access to their proper- 
ties. 


Documents Cited 

Catling, P. M., and V. R. Brownell. 1997. Damselflies 
(Zygoptera) in Ontario from 1900 to 1952. An atlas of 
E. M. Walker’s distributional data for monitoring, biodi- 
versity and biogeography studies. 2326 Scrivens Drive, 
Metcalfe, Ontario. 


Literature Cited 

Catling, P. M. 1997. Evidence for the recent northward 
spread of Enallagma civile (Zygoptera: Coeagrionidae) 
in Ontario. Proceedings Entomological Society Ontario 
127: 1-3. 

Catling, P. M., and P. D. Pratt. 1997. An expanding 
“race” of the Azure Bluet, Enallagma aspersum in 
Ontario? Argia 9(3): 16-17. 

Chovanec, A., and R. Raab. 1997. Dragonflies (Insecta, 
Odonata) and the ecological status of newly created wet- 
lands — examples for long-term bioindication. Limno- 
logica 27: 381-392. 

Fox, A. D., and S. A. Cham. 1994. Status, habitat use and 
conservation of the scarce blue-tailed damselfly [sch- 
nura pumilio (Charpentier) (Odonata: Coenagrionidae) 
in Britain and Ireland. Biological Conservation 68: 
115-122. 

Ménard, B. 1996. Liste annotée des Odonates de la vallée 
de 1’ Outaouais. Fabreries 21: 29-64. 

Oldham, M. J., D. A. Sutherland, and M. L. Holder. 
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Pages 1-7 in Ontario Odonata, Volume |. Edited by 
P. M. Catling, C. D. Jones and P. Pratt. Toronto Ento- 
mologists’ Association, Toronto, Ontario. 153 pages. 

Pilon, S., J.-G. Pilon, and L. Pilon. 1988. Faune odona- 
tologique d’une graviére de la plaine de Montréal, 
Québec, Canada. Notulae Odonatologicae 3: 26-28. 

Snell, E. A. 1987. Wetland distribution and conversion in 
southern Ontario. Inland Waters and Lands Directorate, 
Environment Canada, Ottawa. 

Snell, E. A. 1989. Recent wetland loss trends in southern 
Ontario. Pages 183-197 in Wetlands — inertia or 
momentum? Edited by M. J. Bardecki and N. Patterson. 
Federation of Ontario Naturalists, Toronto. 426 pages. 

Walker, E. M. 1953. The Odonata of Canada and Alaska. 
Volume 1. University of Toronto Press, Ontario. 292 
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Westfall, M. J., and M. L. May. 1996. Damselflies of 
North America. Scientific Publishers, Gainesville, Flor- 
ida. 649 pages. 


Received 11 March 1998 
Accepted 3 January 2002 


String and Net-Patterned Salt Marshes: 
Rare Landscape Elements of Boreal Canada 


KEVIN P. TIMONEY 


Treeline Ecological Research, 21551 Township Road 520, Sherwood Park, Alberta, T8E 1E3 Canada; e-mail: 
ktimoney @compusmart.ab.ca. 


Timoney, Kevin P. 2001. String and net-patterned salt marshes: Rare landscape elements of boreal Canada. Canadian 
Field-Naturalist 115(3): 406-412. 


A rare form of wetland, string and net-patterned salt marshes, exist in and near Wood Buffalo National Park in northeastern 
Alberta and the adjacent Northwest Territories. In two wetlands (Benchmark and Lobstick creeks) examined in August 
2001, a benthic blue-green algae/diatom community covered flark bottoms. Calamagrostis stricta (Narrow Reed Grass) 
and Triglochin maritima (Seaside Arrow-grass) at Benchmark Creek and Atriplex subspicata (Saltbush), Puccinellia nut- 
talliana (Nuttall’s Salt-Meadow Grass), and Triglochin palustris (Slender Arrow-grass) at Lobstick Creek dominated the 
strings. Hordeum jubatum (Foxtail Barley), Puccinellia nuttalliana, Spergularia marina (Salt-marsh Sand Spurry), and 
Triglochin palustris were common to both wetlands. Other noteworthy plant occurrences included Aster ericoides (Tufted 
White Prairie Aster), A. pauciflorus (Few-flowered Aster), Carex mackenziei (Mackenzie’s Sedge), Glaux maritima (Sea 
Milkwort), Monolepis nuttalliana (Spear-leaved Goosefoot), Plantago eriopoda (Saline Plantain), P. maritima (Sea-side 
Plantain), Scirpus paludosus (Prairie Bulrush), and Suaeda calceoliformis (Western Sea-blite). The wetlands observed were 
associated with valleys. In order to form, they may require a gentle elevational gradient for lateral flow of saline surface 
water. The patterns may be related also to soil processes involving excessive soil salts. Soil augering revealed an insuffi- 
cient amount of peat for the sites to be classified as fens; soils appear to classify as Rego Humic Gleysols. These rare wet- 
lands require study of their mode of formation, distribution, and their biota. I suggest a new type designation: interior pat- 


terned saline marsh. 


Key Words: meadow, saline, vegetation, wetland, Wood Buffalo National Park, Alberta, Northwest Territories. 


String or net-patterned fens are fairly common 
features of boreal landscapes in Canada (National 
Wetlands Working Group 1988; Vitt et al. 1996). 
Patterned fens are composed of open, wet flarks and 
drier strings. Strings form perpendicular to surface 
water flow, and are usually sinuous on gently slop- 
ing terrain and net-like on more level terrain where 
surface water flow is multidirectional (Vitt et al. 
1996). In such wetlands the strings or nets are the 
result of peat accumulation. 

Inland saline wetlands in Canada are characteristic 
of the Continental Prairie and Intermountain Prairie 
Wetland regions, wherever hydrogeological condi- 
tions favor their existence (National Wetlands 
Working Group 1988).Typically inland saline wet- 
lands occupy periodically-flooded flats or terminal 
basins where alkali salts concentrate by evaporation. 
The alkali salts (sodium, magnesium, and calcium 
sulphates and chlorides) may be derived from glacial 
deposits overlying Cretaceous bedrock; from alluvial 
and lacustrine soils formed under impeded drainage; 
from soils subjected to seepage from underlying 
shales; and from dissolution of soluble Paleozoic 
evaporite rocks (National Wetlands Working Group 
1988; Rawson and Moore 1944; Last 1995). In 
Wood Buffalo National Park, saline areas may be 
associated with sodium chloride springs that occur 
along the dissolution edge of the Cold Lake 
Formation of marine evaporitic halite (Derry 2001). 


Wood Buffalo National Park supports a diverse 
array of wetlands, ranging from bogs, fens, shrub 
carrs, meadows, and marshes (Raup 1935; PADPG 
1973; Townsend 1973; Cordes 1975; Cordes and 
Pearce 1979; Dirschl et al. 1974; Timoney 1996), 
gypsum-karst wetlands of the Whooping Crane 
(Grus americana) nesting area (Novakowski 1966; 
Kuyt 1981; Timoney et al. 1997; Timoney 1999), 
and saline marshes/meadows or salt plains (Raup 
1935; Airphoto Analysis Associates 1979). The 
saline communities in the park, such as those at the 
Salt Plains west of Fort Smith, are typically feature- 
less or are sometimes dotted by mineral-cored 
upland “islands.” Rarely, saline wetlands in the park 
form string or net patterns reminiscent of patterned 
fens. In this note I describe some of the biological 
and physical features of patterned boreal saline wet- 
lands. 


Observations 

During fixed-wing overflights of the park from 
1993 to 1996 I noted string and net-patterned saline 
wetlands on several occasions (Figure 1). One of these 
wetlands is accessible by foot from the Salt River hik- 
ing trail east of the Pine Lake road, at 59° 48’ 40” N, 
111° 57’ 45” W, in far northern Alberta. I visited this 
site in August 1994 and again in August 2001. 
Calamagrostis stricta, Agropyron trachycaulum 
(Slender Wheat Grass), Juncus (Rush), Puccinellia 


406 


2001 


TIMONEY: STRING AND NET-PATTERNED SALT MARSHES 


407 


Devonian, mainly 
non-karstic rocks 


Northern 
Alberta 


Devonian, mainly 
karstic rocks 


Cretaceous Oilsands 


FIGURE 1. Occurrences of patterned saline marshes described in the paper: L = Lobstick 
Creek, B= Benchmark Creek, and C = Clearwater Springs; all appear to occur in 
association with Devonian karstic rocks. SH = Precambrian Shield; IP = Interior 


Plains. Base map after Ozoray (1976). 


nuttalliana, and Hordeum jubatum meadows, Sali- 
cornia europaea (Samphire) flats, and semi-barren 
areas form a mosaic with patterned wetlands in the 
valley of Benchmark Creek (vascular nomenclature 
follows Moss 1983; common names follow Alberta 
Environmental Protection 1993; voucher specimens 
for difficult or significant species are deposited at the 
herbarium of the Northern Forestry Centre, Natural 
Resources Canada, Edmonton, Alberta). 

The plant taxa of the strings and flarks on 10 
August 2001 (during a wet summer) are listed in 
Table 1. At the time of sampling, the dominants on 
the strings were Calamagrostis stricta and Triglo- 
chin maritima and in flarks was a blue-green algae / 
diatom community with Triglochin maritima and 
Scirpus paludosus (Figure 2). A single moss was 
found: Amblystegium varium (Hedw.) Lindb., grow- 
ing in the shade below the graminoids. Seven years 
previous (7 August 1994, during a dry summer), the 
same spot was visited. Strings were then dominated 
by Scirpus paludosus (Prairie Bulrush), along with 
Calamagrostis stricta and Hordeum jubatum. Pucci- 
nellia nuttalliana, Aster ericoides, Atriplex subspica- 
ta, and Carex mackenziei were also present. Flarks 


(dry at the time of sampling) were barren of vegeta- 
tion, and were covered by salt encrusted mud and 
dead roots and rhizomes. No bryophytes were found. 
Nets in some areas are predominantly mineral, 
devoid of vegetation or peat. It is possible that at 
least some of the patterns might arise from a pat- 
terned ground process associated with excessive soil 
salts. 

Another fine example of a boreal patterned saline 
wetland is located in the Northwest Territories, east 
of the Wood Buffalo park boundary and the 
Whooping Crane nesting area, in the salt flats near 
Kobsticks Creek: (Fieure 3:60" O01’ 55° N;. 112° 33" 
30” W). Atriplex subspicata, Puccinellia nuttalliana, 
Triglochin palustris, and Plantago maritima domi- 
nated the strings, while blue-green algae, diatoms, 
and Triglochin palustris dominated the flarks (Table 
1). There were no bryophytes found. 

The benthic blue-green algae/diatom community 
covering flark bottoms, and Hordeum jubatum, 
Puccinellia nuttalliana, Spergularia marina, and 
Triglochin palustris were common to both wetlands. 

Lobstick appeared to be rich in bird life, with 
flocks of Canada Geese (Branta canadensis) (feed- 


408 


THE CANADIAN FIELD-NATURALIST 


Vokuits 


FIGURE 2. Ground level view of flarks and strings at Benchmark Creek valley, 10 August 2001. Photo cour- 
tesy Mark Bradley, Wood Buffalo National Park. 


ing on Triglochin palustris), Sandhill Cranes (Grus 
canadensis), Least Sandpipers (Calidris minutilla), 
Western Sandpipers (Calidris Maori), American 
White Pelicans (Pelecanus erythrorhynchos), and a 
Bald Eagle (Haliaeetus leucocephalus) observed 
during the hour-long sample. There were abundant 
aquatic invertebrates in the flarks. This saline wet- 
land complex appears distinct ecologically from the 
gypsum-dominated Whooping Crane nesting area. In 
the latter area, the dominant vegetation types are 
shrub bog-marsh, mixed marsh, diatom ponds, and 
shrubby organic terrain; salt marshes are not present 
(Timoney 1999). 


Discussion 

While the study area lies geographically within 
the Continental High Boreal wetland subregion 
(National Wetlands Working Group 1988), the biota, 
saline hydrogeology, and minimal peat accumulation 
in these patterned salt marshes indicate the type to be 
allied with wetlands of the Aspen Parkland Conti- 
nental Prairie subregion. The virtual lack of bryo- 
phytes and characteristic fen species, the predomi- 
nance of marsh halophytes, and the prevalence of 
plants and birds characteristic of the Great Plains 
(Table 1) indicate these communities are similar to 
Prairie type intermittent saline lakes, not boreal fens. 
It is also noteworthy that the Benchmark Creek salt 


marshes lie adjacent to true Prairie grassland disjunct 
occurrences on the south-facing uplands along the 
creek where Stipa, Agropyron, and dryland Carices 
dominate on Dark Grey and Black Solods (Schwarz 
1994; Schwarz and Wein 1997). Soil observations in 
the two wetlands bear on the question of whether 
these communities are salt marshes or fen peatlands 
(Table 2). The minimal peat accumulation and the 
predominance of a heavily-gleyed clayey Cg ally the 
soils with Rego Humic Gleysols (although the sur- 
face horizon is Om, not Ah) (classification after 
Canada Soil Survey Committee 1978). Within the 
Canadian Wetland Classification System (Zoltai and 
Vitt 1995), the patterned saline wetlands would clas- 
sify as a mosaic of tidal marshes and shallow open 
water. I suggest a new type: interior patterned saline 
marsh. 

Looman (1981) classified saline vegetation in the 
Canadian prairie provinces. Prairie Bulrush (domi- 
nant at Benchmark in August 1994) was represented 
in six of eight associations and was the dominant 
species in one association. Kantrud (1996) found that 
two groups of plants are frequent associates of 
Prairie Bulrush. The first group includes tall, often 
long-lived perennial graminoids such as Phragmites 
australis (Reed), Typha latifolia (Common Cattail), 
and S. acutus (Great Bulrush) that may shade Prairie 
Bulrush. The second group of associates includes 


2001 


TIMONEY: STRING AND NET-PATTERNED SALT MARSHES 409 


FiGurRE 3. Aerial view of string and net-patterns and flarks at Lobstick Creek, 10 August 2001. Photo courtesy 


Mark Bradley, Wood Buffalo National Park. 


TABLE |. Plant species composition and percent cover values from Benchmark and Lobstick creeks in 20 by 25 m? releves 
sampled 10 August 2001. Bold face taxa are common to both sites. Distribution notes are based on Scoggan (1978-1979); 
Porsild and Cody (1980); Moss (1983); and Kershaw et al. (2001). 


Taxa 


Aster pauciflorus 

Alberta 

Atriplex subspicata 

Blue-green algae - 
diatom community 

Calamagrostis stricta 

Carex mackenziei 


Hordeum jubatum 
Amblystegium varium 
Monolepis nuttalliana 
Plantago maritima 
Puccinellia nuttalliana 
Scirpus paludosus 
Spergularia marina 
Triglochin maritima 
Triglochin palustris 
Total Vascular Cover 
Total Non-vascular Cover 


Benchmark Creek 
String Flark 
trace 0 

0 0 

0 75 
60 0 
01 0) 
02 0 
01 0 
trace 0 
0) 0 
trace 0 
0 10 
trace 0 
10 25 

trace 0 
~74 B35 

1 TS 


Lobstick Creek 
String Flark 

0 0 
40 0 

0 100 

0 

0 0 
Ol 0 

0 0 

0 0 
15 0 
20 0 

0 0 
trace 0 

0 0 
20 233 
96 She 

0 100 


Distribution Notes 


rare and disjunct in northern 
at n. limit 
need documentation 


circumpolar, coastal species; 
first inland record for Canada? 


near northern limit 

uncommon and local 
uncommon in northern Alberta 
at northern limit 

uncommon and local 


At Benchmark Creek, present outside releve: Agropyron trachycaulum, Aster ericoides (near northern limit), Glaux 
maritima (uncommon in northern Alberta), Plantago eriopoda (uncommon in northern Alberta), Potentilla anserina, and 


Suaeda calceoliformis (near northern limit). 


410 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


TABLE 2. Soil horizon observations at Benchmark and Lobstick Creeks based on 1 m deep soil auger holes. 


Location Soil horizon 
Benchmark Creek String 

0-30 cm Om 
30-100 cm Cg 
Benchmark Creek Flark 

0-43 cm Om 
43-100 cm Cg 
Lobstick Creek String 

0-10 cm Om1 
10-23 cm Om2 
23-100 cm Cg 
Lobstick Creek Flark 

0-8 cm Om 
8-25 cm Om/Cg 
25-100 cm Cg 


Triglochin maritima, Eleocharis palustris (Creeping 
Spike-Rush), Atriplex patula (sensu lato), S. pungens 
(Three-Square Rush), Hordeum jubatum, and several 
species of Juncus. Increases or decreases in the com- 
petitive advantage of these species are related to 
changes in salinity, elevation, disturbance and inun- 
dation regime, substrate texture, and many other fac- 
tors (Kantrud 1996). Prairie Bulrush marshes tend to 
contain surface water for shorter periods than do 
semipermanent fresh or slightly brackish wetlands. 
The patterned saline wetlands in Wood Buffalo and 
vicinity belong to Kantrud’s second group. 

Wallis (1990) noted a Scirpus paludosus “saline 
emergent marsh” type for the Grassland and Park- 
land region of eastern Alberta. No mention was 
made of net or string patterns in any of the saline 
wetlands he studied. Fairbarns (1990) studied saline 
meadows near High Level, Alberta and made no 
mention of net or string patterns. He identified a 
Scirpus paludosus-Eleocharis palustris type of mod- 
erately saline “marsh meadow” and noted that the 
Species was rare in northern Alberta. 

I could find no published accounts of boreal pat- 
terned saline wetlands, but P. Lee (Global Forest 
Watch, Edmonton, Alberta, personal communication 
1999) called my attention to two such wetlands 
along the Clearwater River in northeastern Alberta. 
Clearwater River file notes in the Alberta Natural 
Heritage Information Centre (Alberta Environmental 
Protection, Edmonton) indicated a “wet meadow 
with crescent-shaped ponds” at 56° 40'30” N, 110° 
55’W and “spring and crescent-shaped ponds” at 56° 
44'30"N, 110°30'W. No vegetation data could be 
found. A 1:15 000 airphoto of the area showed string 
patterns in the wetlands. A chemical analysis at one 
of the Clearwater River saline wetlands (dated 28 
March 1984, location not given) reported a pH of 
7.9, conductivity of 14.77 mS/cm, and total dis- 
solved solids 8500, chloride 4600, sodium 2940, sul- 
phate 600, bicarbonate 184, calcium 180, and 


Notes 


Om graminoid peat; Cg strongly gleyed clay 


Om a complex of diatomaceous earth and graminoid 
peat; Cg strongly gleyed clay 


Om1 vascular plant peat; Om? diatomaceous earth; 
Cg strongly gleyed clay includes an apparently frozen 
sand layer at ~86 cm 


Om diatomaceous earth; Om/Cg = mix of Om 
with gleyed C; Cg strongly gleyed clay includes 
apparently frozen sand layer at ~71 cm 


magnesium 71 mg/l. While such figures classify the 
wetland as saline, the wetland does not fit within the 
boreal types described by National Wetlands 
Working Group (1988), and is most closely-related 
to their (unpatterned) saline wet meadow and emer- 
gent marshes of the Canadian Prairies. 

The chemical limnology of some lakes in the area 
of Benchmark and Lobstick has been described by 
Moser et al. (1998). Their most similar site, e.g., was 
lake “WB25”, but conductance, sodium, and chlo- 
ride values were lower by 1-2 orders of magnitude 
at “WB25”. The marshes of Benchmark and 
Lobstick are likely more saline than the gypsum- 
dominated sinkhole lakes they described, and also 
drawdown regularly resulting in widely fluctuating 
solute concentrations. Chemical profiles of several 
ponds and springs described by McNaughton (1991) 
west of Lobstick may be more similar (e.g., 
“Klewil” where he observed a conductivity of 
11.3 mS/cm, somewhat lower values for sodium and 
chloride and higher values for sulphate and calcium). 
Chemical data and zooplankton communities were 
described for one lake near Benchmark Creek (lake 
SP-CL, 59° 48’N, 112° 01’W) and one lake near 
Lobstick (lake GL-D, 60° 00'N, 112° 37’W) by 
Derry (2001). Near Benchmark, she reported a con- 
ductivity of 17.6—20.4 mS/cm, Na 3520-4077 mg/l, 
and Cl 5265-6116 mg/l — very similar to the chem1- 
cal data from the Clearwater River saline wetlands. 

A significant feature of these wetlands is the high 
proportion of rare or uncommon plant species (Table 
1). Most surprising is the occurrence of Carex 
mackenziei, a circumpolar coastal species. Its occur- 
rence at Benchmark may be the first inland record 
for Canada. Scirpus paludosus, Atriplex subspicata, 
and Aster pauciflorus, are prairie wetland plants that 
are rare in northern Alberta. 

Little is known of the benthic algal and diatom 
community. While some characteristic diatoms have 
been noted from the nearby Whooping Crane nesting 


2001 


area (Timoney et al. 1997), the water regime and its 
chemistry may result in a different flora and fauna at 
Benchmark and Lobstick. Near the headwaters of 
Benchmark Creek, M. Rosen (personal communica- 
tion, Fort Smith, Northwest Territories, October 
2001) has found the dried mud crust at Grosbeak 
Lake (59° 47’ 30”N, 111° 59’W) to be dominated by 
blue-green algae, whose matrix contained abundant 
copepods, nematodes, diatoms, and foraging brine 
flies. 

The three interior patterned saline marshes noted 
here were associated with creek or river valleys. In 
addition to saline surface waters, these wetlands may 
require a slope gradient conducive to gradual surface 
water flow. String or net formation may be encour- 
aged by peat accumulation or entrainment of lateral- 
ly-moving mineral or organic matter. Some of the 
patterns might arise from a patterned ground process 
associated with excessive soil salts. 

Interior patterned saline marshes are groundwater 
and runoff fed wetlands which undergo large season- 
al and annual changes in water level and concentra- 
tion of solutes. These wetlands, which appear to be 
rare, may harbor a variety of biota that require better 
documentation. They merit further study, which I 
hope will delight all naturalists who visit them. 


Acknowledgments 

Peter Lee and Lorna Allen made available infor- 
mation on the Clearwater River wetlands. Thanks to 
Dale Vitt, Bill Last, John Smol, Derek Johnson, and 
anonymous reviewers for comments. Thanks to 
Anne Robinson who assisted in August 1994 field- 
work; to Catherine England who identified the moss, 
and to Mark Bradley who assisted in the August 
2001 fieldwork and identified the birds at Lobstick. 
Thanks to Wood Buffalo National Park fire manage- 
ment who provided the helicopter access to Lobstick. 
This paper is dedicated to my late friend Steve Zoltai 
who would have liked these wetlands. 


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Received 3 March 2000 
Accepted 6 November 2001 


Predation on Nesting Woodpeckers in British Columbia 


Eric L. WALTERS! and EDWARD H. MILLER? 


Department of Biology, University of Victoria, P.O. Box 3020, Victoria, British Columbia V8W 3N5 Canada 

‘Current address: Department of Biologicai Science, Florida State University, Tallahassee, Florida 32306-1100 USA 

2Current address: Biology Department, Memorial University of Newfoundland, St. John's, Newfoundland AI1B 3X9 
Canada 


Walters, Eric L., and Edward H. Miller. 2001. Predation on nesting woodpeckers in British Columbia. Canadian Field- 
Naturalist 115(3): 413-419. 


Predation on eggs, nestlings, and breeding adults of Red-naped Sapsuckers, Sphyrapicus nuchalis, Northern Flickers, 
Colaptes auratus, Hairy Woodpeckers, Picoides villosus, and Williamson's Sapsuckers, S. thyroideus, was documented in 
the Hat Creek valley, south-central British Columbia from 1989-1994. Predation by Black Bears (Ursus americanus), Deer 
Mice (Peromyscus maniculatus), and House Wrens (Troglodytes aedon) was observed; and predation by Long-tailed 
Weasels (Mustela frenata) was inferred. , 


Key Words: Red-naped Sapsucker, Sphyrapicus nuchalis, Williamson's Sapsucker, S. thyroideus, Hairy Woodpecker, © 


Picoides villosus, Northern Flicker, Colaptes auratus, Black Bear, Ursus americanus, Deer Mouse, Peromyscus 
maniculatus, House Wren, Troglodytes aedon, nesting, predation, British Columbia. 


Hole nesting in birds has evolved independently in 
many taxonomic groups. The generally accepted 
dogma is that hole nesting offers an advantage over 
open nesting (e.g., von Haartman 1957). Many 
authors (e.g., Lack 1954; Nice 1957) have provided 
evidence to suggest that nesting success is higher in 
hole-nesting species even though there is more con- 
straint on choice of nesting location. Because preda- 
tion can be a major cause of nest failure among 
species that nest in holes (Nilsson 1984), it has obvi- 
ous implications for the evolution of life history 
traits (Martin 1995). Predation, therefore, must be a 
strong evolutionary force with respect to breeding 
biology (Alerstam and Hoégstedt 1981; Nilsson 
1984). However, data concerning predation on hole- 
nesting species are difficult to obtain, requiring 
detailed life-history studies over successive breeding 
seasons (Greene 1986); and usually involve climbing 
trees, many of which are in various stages of decay. 
Recent advances in technology have allowed some 
researchers to utilize cameras to circumvent some of 
the problems associated with observing nests (e.g., 
Martin 1988; Picman and Schriml 1994; Thompson 
et al. 1999). The use of cameras for tree holes that 
are relatively high or for observing nocturnal preda- 
tors, however, is still quite limited. Besides difficul- 
ties associated with nest monitoring, many studies of 
the reproductive success of hole-nesting species have 
relied upon nest box studies (e.g., van Balen and 
Potting 1990; Verhulst et al. 1995) rather than using 
natural cavities. 

Our objectives are-to document predation events, 
to identity the predators, and to describe the evi- 
dence that allows inferences about the species of 
predator that prey upon woodpecker nests. In this 
paper we define nest predation as any event that 


results in the destruction of eggs or the death of a 
chick or adult during the nesting stage without 
regard to whether the “predator” actually ingested 
the egg, chick, or adult (cf. Sealy 1994). Under this 
definition, a predator may be motivated either by 
hunger or by a desire to obtain a nest or roost site. 


Study Area and Methods 

The study site was within the Hat Creek valley near 
Upper Hat Creek (25 km SW of Cache Creek), south- 
central British Columbia (50°46’N 121°38’W), at an 
elevation of approximately 1200 m. The slopes of the 
narrow valley are forested with second-growth 
Interior Douglas-fir (Pseudotsuga menziesii), Interior 
Spruce (Picea engelmannii X glauca), and pines 
(Pinus contorta and P. ponderosa), with some 
Trembling Aspen (Populus tremuloides). On the val- 
ley bottom the same tree species occur, but aspen and 
willow (Salix spp.) are more abundant. Further details 
about the study site are given by Walters (1996). 

Woodpecker nests were studied from late April to 
late July 1989-1994. Nest monitoring varied among 
years. In 1989 and 1991, nests were monitored up to 
three times; in 1990 and 1992, they were visited 
approximately 20 times; and in 1993 and 1994, nests 
were visited daily. Emphasis was on finding nests of 
the Red-naped Sapsucker (Sphyrapicus nuchalis), 
the most abundant breeding species in the area. 
Other common breeding woodpeckers were the 
Northern Flicker (Colaptes auratus), Hairy Wood- 
pecker (Picoides villosus), and Pileated Woodpecker 
(Dryocopus pileatus). Downy Woodpeckers (Pico- 
ides pubescens) and Williamson’s Sapsuckers (Sphy- 
rapicus thyroideus) bred regularly in the area but 
were uncommon. 

Nests were found by various means: nesting signs 


413 


414 


(e.g., recent excavations, wood chips on ground), 
audible cues (e.g., drumming, vocalizations, nestling 
calls), observing adults feeding young, or by direct 
observations of birds (Jackson 1977). In 1992-1994 
we checked nest contents with a mirror and flash- 
light (nest contents were not checked prior to 1992). 
Predation was assumed when all eggs or nestlings 
were missing from the nest (except when fledging 
was expected), or if eggshells, feathers, or other 
signs (e.g., sticks in the hole entrance) were in the 
nest cavity or on the ground below the nest 
(Johnsson 1994). The identity of the predator was 
determined by either observation of the predation 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


event or indirectly by examining the result of the 
predation event (e.g., use of sticks by wrens, bear 
claw marks). All of our observations of predation 
events were opportunistic in nature and occurred 
while we were checking the status of each nest. 


Results 

The number of woodpecker nests monitored var- 
ied among years (Table 1). We found evidence of 23 
cases of nest predation out of a total of 239 nests 
during our study: probable musielid, 12; Black Bear 
(Ursus americanus), 4; House Wren (Troglodytes 
aedon), 3; Deer Mouse (Peromyscus maniculatus), 


TABLE 1. Numbers of woodpecker nests with eggs or young and the outcome associated with each. 


Species Year Total Cause of Failure Outcome 
Predation Unknown Successful Unknown 
Weasel Bear Mouse Wren 

RNSA 1989 39 - - = - - _ 39 
1990 36 ~ — - — 3 DD 11 
1991 19 _ = - - - — 19 
1992 24 1 — - _ 1 21 1 
1993 25 p — 1 — 2 20 — 
1994 30 3 - - 3 5) 19 — 
NOFL 1989 1 -: - - - ~ 1 
1990 9 - 1 ~ _~ 1 4 
1991 2 _ _ _ — ~ = 2, 
1992. 11 1 Z 1 _ 2 _ 5 
1993 9 1 - - _ - ij 1 
1994 11 2, — 1 _ j 7 — 
HAWO 1989 3 - - — - - 3 
1990 3 — _ _ ~ _ 1 2, 
1991 0 - — _ = ~ - - 
1992 3 1 — = = — 2 - 
1993 2 1 _ — = = 1 — 
1994 5 - _ _ - — 3 - 
WISA 1989 1 ~ _ = — ~ 1 — 
1990 0 _ — - ~ - = - 
1991 1 _ = = = = = 1 
1992 i = - = ~ = - 
1993 0 ~ ~ — _ _- _ - 
1994 0 — = = = = _ - 
DOWO 1989 0 — _ = — - — — 
1990 1 — = = = _ 1 - 
1991 0 — - — = = = - 
1992 1 _ = = = = = 1 
1993 0 - _ = = _ ~ — 
1994 1 — = = = = 1 - 
PIWO 1989 0 — _ = =: _ _ = 
1990 0) _ = = = =_ — _ 
1991 0 _ = = = = = ~ 
1992 0 - _ = = - ~ - 
1993 1 _ = ~ = 1 _ - 
1994 2 _ = = = 1 1 - 
Total 239 1% 4 3 3 9 110 90 


RNSA = Red-naped Sapsucker; NOFL = Northern Flicker; HAWO = Hairy Woodpecker, WISA = Williamson’s 
Sapsucker, DOWO = Downy Woodpecker, and PIWO = Pileated Woodpecker. 


2001 


3; and Cooper’s Hawk (Accipiter cooperii), 1. We 
also found 17 nests where the young died of 
unknown causes. 

Twelve occurrences of nest predation (six Red- 
naped Sapsucker, four Northern Flicker, two Hairy 
Woodpecker) were presumed to be by a mustelid, 
probably the Long-tailed Weasel (Mustela frenata) 
given that it was the only mustelid observed in the 
study area. Killed were both adult woodpeckers and 
large nestlings, all within nesting cavities. In the first 
case of predation on sapsuckers, some hairs (light 
brown in colour and >3 cm) were found at the cavity 
entrance and the eggs were gone. In the second, shell 
fragments were observed within a sapsucker cavity 
followed by a dead adult male in the cavity the next 
day. The third event occurred when a large sapsuck- 
er nestling was found partly eaten at the base of a 
nest tree. In the fourth, flesh plus crushed shells were 
present in the sapsucker nest. The fifth nest was 
observed late in the day as both parents were feeding 
the young. Early the following day the male was 
gone and the chicks were found dead in the cavity. 
Finally, of three sapsucker nestlings within a few 
days of fledging, one was found dead in the nest and 
the others alive at the base of the nest tree. The dead 


FiGuRE 1. Portion of a Trembling Aspen trunk used for 
nesting by Red-naped Sapsuckers over many sea- 
sons. Note the extensive scarring on the trunk, 
caused by Black Bears climbing up the tree. 


WALTERS AND MILLER: PREDATION ON NESTING WOODPECKERS 


415 


chick was removed, and the live chicks were 
returned to their nest. The next day one chick was 
dead inside the cavity and the other chick was alive 
at the base of the nest tree. We placed this chick on 
the trunk of the tree; it fledged successfully. Three 
adult Northern Flickers were depredated while incu- 
bating or brooding. Hairy Woodpecker adult males 
were preyed upon at night when brooding large 
young. Both nests were in the same tree in succes- 
sive years. In each case at least one of the young was 
removed from the cavity. Nine of the twelve nests 
where suspected mustelid predation occurred were 
from two areas (<5 ha) within our 80 ha study site. 
In all nests where mustelid predation is suspected, no 
tooth marks were evident around the cavity entrance. 

Evidence of predation or attempted predation by 
Black Bears was of three types: fresh scarring of trees 
by claws; scarring around nest holes by teeth; and 
mortality of chicks or adults. Many old nest trees 
(10/25 in 1993 and 11/30 in 1994 for Red-naped 
Sapsuckers) had numerous scars caused by the claws 
of bears during climbing (Figure 1). We noted four 
instances where apparently successful predation by 
bears had occurred and eight more attempts. In one 
case, a low (approximately 1.5m above ground) 
Northern Flicker nest in a rotten stub of a large 
Interior Spruce had been exposed when the stub was 
ripped open. Bloody pinfeathers (remiges) of the 
nestlings were around the base of the stub. The tall 
grass around the stub was beaten down, suggesting 
the presence of a large mammal, and fresh bear 
feces lay a few meters away. In another case, a 
Williamson’s Sapsucker nested in a Trembling Aspen 
1.8 m above the ground. When the nest was checked, 
bite marks (consistent with a bear) were evident 
around the entrance (Figure 2) and the remains of the 
incubating male were in the intact nest cavity. We 
also found six Red-naped Sapsucker nests, one 
Northern Flicker nest, and one Hairy Woodpecker 


a: : 
aes 


FIGURE 2. Nest hole of Williamson’s Sapsucker, showing 
marks from lower canines of Black Bear; the adult 
male died in the nest cavity from injuries received 
from the bear. 


416 


nest in which a bear had climbed to the cavity and 
clawed at the entrance but was not successful in gain- 
ing access to the nest. The nests had been checked 
less than 24 hours earlier. 

We observed one occurrence of egg predation by a 
Deer Mouse and suspected it in two other nests. A 
female Northern Flicker flew and called agitatedly as 
we checked her nest. A Deer Mouse was visible in 
the nest, amidst the eggs, and two of the six eggs 
were smashed. Six hours later the entire clutch had 
been destroyed and only broken, flattened eggshells 
remained. Single nests of a Northern Flicker and a 
Red-naped Sapsucker were found with broken and 
flattened full or partial clutches in the cavity. 

House Wrens depredated three Red-naped Sap- 
sucker nests. At one, eggshell fragments were inside 
and outside the nest cavity. The next day, House 
Wrens were observed entering and exiting the cavity 
and there were twigs in the cavity (no twigs were 
present the day before). Two other predation events 
occurred such that freshly dead Red-naped 
Sapsucker chicks were found in their nests, sticks 
over them, and House Wrens were nearby. In all 
cases, House Wrens later nested within the sapsucker 
cavities. 

One instance of predation of an adult Red-naped 
Sapsucker by a Cooper’s Hawk was observed. The 
radiotagged sapsucker, five days after successfully 
fledging four young, was found dead in a Cooper’s 
Hawk nest. 


Discussion 

In spite of any extra protection afforded cavity 
nesters, woodpeckers in our study suffered substan- 
tial losses in the breeding season. We cannot, how- 
ever, estimate the proportion of nests that were 
depredated because nest-monitoring effort differed 
among years. 

Evidence suggested that mustelids may be the 
most common predators of woodpecker nests in our 
study area. In England, almost all (96%) of the pre- 
dation on tit (Parus spp.) nests in nest boxes was by 
mustelids (Dunn 1977). Sleeman (1993) even specu- 
lates that many hole-nesting fauna found in Britain 
are not found in Ireland because of predation pres- 
sure by M. erminea. Of known predation events in 
our study, we attribute 55% to mustelids; but one 
would expect relative abundances of predators to 
vary among geographic areas. For example, in 
Sweden, woodpeckers were the chief predator (48%) 
of tits nesting in nest boxes (Nilsson 1984). 

Interestingly, presumed predation by mustelids 
occurred in certain parts of our study area from year 
to year. Individual mustelids learn where nests are 
(Johnson 1947) and revisit them from one year to 
another (Sonerud 1985a,b; 1989). This may explain 
what we attribute to mustelid predation in our study, 
and why (in some species) nestling predation in new 


THE CANADIAN FIELD-NATURALIST 


Volos 


cavities may be less than in old ones (Nilsson et al. 
1991). Because mustelids in our study area tend to 
be nocturnal (Burt and Grossenheider 1980; but see 
Johnson 1947; Pettingill 1976; Kilham 1977b; and 
Daily 1993) and our nest monitoring was diurnal, we 
are not able to state conclusively that mustelids were 
responsible for the predation events we attributed to 
them. However, we found hairs at the entrance to the 
cavity in one case similar to what Kilham (1977b) 
reported after he had observed a weasel depredating 
a Yellow-bellied Sapsucker (Sphyrapicus varius) 
nest. In contrast, Crockett (1975, page 93) observed 
“the total destruction” of a Williamson’s Sapsucker 
nest by M. frenata. Similarly, Erskine and McLaren 
(1972) report several Northern Flicker nests that 
were destroyed by assumed M. erminea. 

Successful predation by Black Bears on nesting 
adult Red-naped Sapsuckers and Northern Flickers 
has been reported by Franzreb and Higgins (1975) 
and DeWeese and Pillmore (1972), respectively. 
Similar to some of the nests in our study, the latter 
authors noted that bears gained entrance to nest cavi- 
ties in living aspen. How Black Bears capture adult 
woodpeckers and probably advanced nestlings is 
largely unknown. Dixon (1927) reported a Black 
Bear trying to gain access to a Black-backed 
Woodpecker (Picoides arcticus) nest by gnawing at 
the entrance hole. Our Williamson’s Sapsucker 
observation suggested that the bear gnawed at the 
nest entrance and caught the inhabitant as it exited. 
Adults and advanced nestlings are prone to scramble 
out of the nest when disturbed (e.g., by a human 
climbing the nest tree). Northern Flickers are partic- 
ularly susceptible to predation by bears at our field 
site, as Northern Flickers nest close to the ground in 
rotten snags. Because it has been assumed that hole 
nesting offers a refuge against predation (Lack 
1968), Redondo and de Reyna (1988) claimed that 
the young of hole-nesting species produce calls with 
wider frequency ranges and less attenuable signals 
than those of open-nesting species (cf. Popp and 
Ficken 1991). The incessant calling of young in 
some species may be a cue to which Black Bears 
(and other predators) are attuned and thus a paradox 
seems apparent. Counter to the views of Redondo 
and de Reyna (i.e., ecological release of nestling 
vocalization), perhaps the vocal cues emitted by the 
young of hole-nesting species are constrained such 
that the signal will carry outside of the nest (i.e., so 
the parents can hear the young). It does not appear 
that. Black Bears randomly climb trees. We com- 
pared the frequency with which available trees (1.e., 
> 12 cm diameter at breast height) in a 1 ha area sur- 
rounding the nest tree (N = 17) exhibited bear claw 
marks compared with nest trees. Frequency of bear 
claw marks differed significantly (Fisher’s Exact 
Test, p<0.001) between nest trees (11 / 30 Red- 
naped Sapsucker nests in 1994) and available trees 


2001 


(190 / 4155), suggesting that bears are selectively 
climbing nest trees. 

Deer Mice have been reported to be significant 
predators on ground-nesting birds (Maxson and 
Oring 1978; Reitsma et al. 1990). However, we are 
only aware of one study that reported predation by 
Peromyscus spp. on a hole-nesting species: Guillory 
(1987) observed predation by P. leucopus and P. 
gossypinus on Prothonotary Warbler (Protonotaria 
citrea) nests. Our findings of Deer Mice predation 
appear to be the first for a woodpecker nest. We esti- 
mate that at least 14% of our predation events were 
due to Deer Mice. 

House Wrens often peck at and perforate eggs, in 
conspecific and heterospecific nests, and then 
remove them (White and Kennedy 1997). One adap- 
tive interpretation (among several) placed on this 
behavior is that it is an interference mechanism 
(Belles-Isles and Picman 1986). In our study, House 
Wrens benefited through such behavior by disrupting 
the nesting cycle of Red-naped Sapsuckers, who 
abandoned their nesting attempt, or spent more time 
away in preparation for another breeding attempt. In 
the latter case, sapsucker re-use of the nest cavity 
was discouraged because the wrens put nesting 
material in the cavity. Kennedy and White (1992) 
have noted the discouraging effect of sticks on other 
species. We suspect that the placement of nesting 
material (e.g., sticks) on sapsucker nestlings within 
our study may have caused their death. 

Other species may have been responsible for the 
unknown cases of predation. For example, both Red 
Squirrels (Tamiasciurus hudsonicus) and Northern 
Flying Squirrels (Glaucomys sabrinus) are present in 
the study area. The former is known to depredate 
Yellow-bellied Sapsucker nests (Lawrence 1967; 
Erskine and McLaren 1972) but we have no evidence 
(e.g., none of the depredated nests was chewed around 
the entrance hole) to suggest that squirrels depredated 
any nests. In fact, we had several nest trees where 
both squirrels (T. hudsonicus and G. sabrinus) and 
sapsuckers coexisted without any apparent negative 
effect on the sapsucker nests. Although Raccoons, 
Procyon lotor, are known to prey upon Yellow-bellied 
Sapsucker nests (Kilham 1971, 1977a), they do not 
occur in our study area. 

We have outlined the nature of predation events 
on four woodpecker species, all of which excavate 
cavities in which to nest. Given the extent to which 
these cavity nesters are susceptible to predation may 
lead one to question the adaptiveness of hole nesting 
as an anti-predation strategy. As some have suggest- 
ed (e.g., Alerstam and Hogstedt 1981), perhaps hole 
nesting is the ancestral trait and open nesting is 
derived. Thus, open-nesting species that are secretive 
in their foraging might overcome the risk of preda- 
tion relative to the cost of finding or constructing a 
suitable hole in which to nest. Under this scenario, 


WALTERS AND MILLER: PREDATION ON NESTING WOODPECKERS 


417 


hole-nesting species are not seeking refuge from 
predatory events but, in fact, have less chance of 
being depredated than if they were to become open 
nesters. Lack (1954) and Nice (1957) both estimated 
that the proportion of eggs in completed clutches that 
give rise to flying young was approximately 45-46% 
in Open-nesting species compared with 66-67% in 
hole-nesting species. One would expect predation to 
be lower in hole-nesters that excavate within rela- 
tively hard trees as opposed to species that use softer 
wood. Supporting this contention is the work of 
Christman and Dhondt (1997) who found that nest 
predation in Black-capped Chickadees (Poecile atri- 
capilla), a species that excavates within soft and 
often rotten wood, is as high as 62%. In our study, 
Northern Flickers tended to nest in softer trees, and 
we recorded a predation event in 21% of our nests. 
On the other hand, Red-naped Sapsuckers tended to 
nest in live aspen (i.e., relatively hard wood; 
Schepps et al. 1999), and we recorded a predation 
event in only 6% of those nests. Neither figure 
should be interpreted as overall predation frequency 
because the number of nest observations differed 
from year to year. Besides the integrity of the sub- 
strate, Northern Flickers may have been exposed to 
higher predation rates than sapsuckers because the 
cavity entrance of Northern Flicker nests is larger. 
Several researchers (e.g., Sandstr6m 1991; Sonerud 
1985b) found that cavities with larger entrance holes 
were more prone to predation. Ironically, Martin and 
Li (1992) did not observe any predation on Northern 
Flicker or Red-naped Sapsucker nests during three 
breeding seasons in Arizona. We hypothesize that 
this finding may be due to the fact that their site dif- 
fered from ours with respect to potential predators 
(e.g., the Arizona field site had less bears, personal 
observation (ELW)) and cavity-nesting species tend- 
ed to nest higher in trees in Arizona (i.e., reducing 
potential for depredation, personal observation 
(ELW)). In fact, Li and Martin (1991) reported that 
nest success was lower for species with lower nest 
height in their study area. 


Acknowledgments 

We are indebted to Ken and Gina Reynolds, 
Brand 88 Ranch, for allowing us to work and stay on 
their property, and for their strong support of our 
work there. The ranch managers, Tim and Lois 
Malpass, helped us on many occasions, and extended 
numerous courtesies. John Cooper was instrumental 
in the initial development of the study and he kindly 
provided data on several bear predation events. 
Financial and logistic support was provided by: 
Natural Sciences and Engineering Research Council 
of Canada (operating grants to EHM); King-Platt 
Memorial Scholarship and Fellowship (awarded to 
ELW); British Columbia Ministry of Environment, 
Parks and Lands (Wildlife Branch); British 


A18 


Columbia Ministry of Forests (Silviculture and 
Research branches); Canadian Wildlife Service; 
Copley Bros. Construction; and British Columbia 
Ministry of Tourism, Recreation and Culture (Royal 
British Columbia Museum). For their assistance, 
encouragement, and advice, we thank Joe Antos, 
Alan Burger, Trudy Chatwin, Don Clark, Andrew 
Derocher, Jakob Dulisse, Michael Dunn, Mike 
Fenger, Elizabeth Hunter, Frances Jones, Sarah 
Jones, Kathy Martin, Peter Miller, Rissa Miller, Ross 
Miller, Cathy Mutter, Bev Mutter, Roy Mutter, Brian 
Nyberg, Karen Walters, Larry Walters, and Ted 
White. Frances James, Julie Jo Walters, Spencer 
Sealy, and Tony Erskine graciously offered con- 
structive criticisms of earlier versions of this 
manuscript for which we are grateful. 


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2001 


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WALTERS AND MILLER: PREDATION ON NESTING WOODPECKERS 


419 


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Received 20 July 1999 
Accepted 8 August 2001 


Timing of Pregnancy, Lactation, and Female Foraging ACUIVAtY 4 in 
Three Species of Bats in Southern Illinois 


GEORGE A. FELDHAMER, TIMOTHY C. CARTER, and STEVEN K. CARROLL 


Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901 USA. 


Feldhamer, George A., Timothy Carter, and Steven K. Carroll. 2001. Timing of pregnancy, lactation, and female foraging 
activity in three species of bats in southern Illinois. Canadian Field-Naturalist 115(3): 420-424. 


Pregnant Northern Long-eared Bats (Myotis septentrionalis), Red Bats (Lasiurus borealis), and Eastern Pipistrelles 
(Pipistrellus subflavus) were taken beginning in mid-May in southern Illinois. For Northern Long-eared and Red Bats, 
pregnant individuals were caught during a two-week period until early June. Pregnancy and lactation in Northern Long- 
eared and Red Bats were fairly synchronized; both species apparently limited their foraging activity during a one-week 
period possibly near and immediately after parturition. Pregnancy in Eastern Pipistrelles was much less synchronized. 
Capture of pregnant females extended over at least an eight-week period from mid-May to mid-July. Volant juvenile 
Northern Long-eared Bats were taken three weeks earlier than any other species; either parturition was earlier or young 
developed more rapidly. Pregnant Northern Long-eared Bats foraged primarily in the early evening; early and late activity 
periods were evident for lactating individuals. Foraging trends for pregnant and lactating Red Bats and Eastern Pipistrelles 
were less apparent. Significant differences in body mass of Northern Long-eared Bats and Eastern Pipistrelles were evident 
based on reproductive status. No differences in body mass associated with reproductive condition were found in female 
Red Bats. 


Key Words: Eastern Pipistrelle, Lasiurus borealis, mist netting, Myotis septentrionalis, Northern Long-eared Bat, 


Pipistrellus subflavus, Red Bat, reproduction. 


Bats in southern Illinois are seasonally monestrus 
and give birth during the late spring and early sum- 
mer when insect populations are most abundant. 
Despite increased research, there is little quantitative 
information on the timing of pregnancy and lactation 
for many species of bats. Hoffmeister (1989), for 
example, notes time of parturition for many species 
simply as “late June or early July.” We collected 
information on temporal patterns of pregnancy and 
lactation for species of bats mist netted throughout 
southern Illinois during a study on the maternity 
roosting ecology of Indiana Bats (Myotis sodalis). 
Specifically, we discuss the first and last dates of 
capture for pregnant and lactating adult Northern 
Long-eared Bats (Myotis septentrionalis), Red Bats 
(Lasiurus borealis), and Eastern Pipistrelles 
(Pipistrellus subflavus), species-specific differences 
in body mass of pregnant, lactating, and nonpregnant 
females, timing of nightly foraging activity, and the 
first date volant juveniles were netted. 


Methods and Materials 

The study was conducted throughout the 109 263- 
ha Shawnee National Forest in southern Illinois. We 
selected a variety of sites to capture bats based on a 
range of habitat variables obtained from the 
Southern Illinois University GIS database and the 
Illinois State Geological Survey. 

Mist netting was conducted on a total of 41 sites 
from 18 May through 18 August 1999, and from 12 
May through 20 July 2000. We used nets and associ- 


ated equipment as described by Gardner et al. 
(1989). Nets were placed along streams or other 
areas of anticipated high bat activity, as well as with- 
in closed canopy interior forest sites. A minimum of 
two nets 5.6 m or higher, was used for two consecu- 
tive nights when they were set over a water source. 
When nets were not set above water, four sets 5.6 m 
or higher were operated for two consecutive nights. 
Nets were stacked up to three high (9 m) with vari- 
ous widths to enclose the flight corridor (Gardner et 
al. 1989). Netting began at dusk and usually contin- 
ued until 0200 h the next morning, with nets checked 
every 20 minutes. Criteria for site selection and the 
netting protocol are described in Carroll (2001) and 
Carroll et al. (2002). 

Pregnant females were identified by palpation of 
their abdomen; lactation was evident by condition of 
the nipples (Foster and Kurta 1999; Kunz 1973; 
Racey 1988). Juveniles and adults were distin- 
guished by examination of phalangeal epiphyses, 
overall body size, condition of teats in females, and 
size of testes in males (Anthony 1988). Bats were 
weighed to the nearest 0.5 g. A 3 mm circular punch 
in each wing was used to collect tissue for genetic 
analyses and also served as a semi-permanent mark 
to determine recaptures. Forage activity periods were 
based on the time of captures, with each six-hr mist 
netting session grouped into three two-hr periods. 


Results 
Mist nets were operated a total of 339 net nights 


420 


2001 


FELDHAMER, CARTER, AND CARROLL: BATS IN SOUTHERN ILLINOIS 


42] 


TABLE |. Species of bats, number of individuals mist netted, and the earliest date volant 
juveniles were caught, at 41 sites throughout Shawnee National Forest during spring and 


summer 1999 and 2000. 


Species 


Northern Long-eared Bat (Myotis septentrionalis) 


Red Bat (Lasiurus borealis) 

Eastern Pipistrelle (Pipistrellus subflavus) 
Big Brown Bat (Eptesicus fuscus) 

Evening Bat (Nycticeius humeralis) 

Indiana Bat (Myotis sodalis) 

Little Brown Bat (Myotis lucifugus) 
Southeastern Bat (Myotis austroripareous) 
Hoary Bat (Lasiurus cinereus) 

Silver-haired Bat (Lasionycteris noctivagans) 


during the two years of sampling. A total of 10 
species and 417 individual bats was captured (Table 
1). Only two individuals were recaptured. Because 
equal numbers of bats were captured each year, we 
combined data from both years, with analyses limit- 
ed to Northern Long-eared Bats, Red Bats, and 
Eastern Pipistrelles based on sample sizes. 


Timing of Pregnancy, Lactation, and Volant 
Juveniles 

Northern Long-eared Bats — Pregnant females were 
taken from 16 May through 2 June. Lactating females 
were captured from 5 June through 20 July. During 
the period when pregnant females were no longer 
taken, but before lactating bats appeared (3-8 June 
1999 and 31 May-5 June 2000), females appeared to 
limit their foraging activity. We netted 13 male M. 
septentrionalis during these two periods, but no 
females (x? = 13.00, P < 0.001). Of 41 adult females 
netted during May and June, 40 (97.6%) were either 
pregnant or lactating. The only adult considered non- 
pregnant was taken on 12 May. From 2 July until the 
end of fieldwork, only 4 of 26 females (15.4%) cap- 


Number of Earliest Date 
Individuals Volant Young 
174 25 June 
Hie. 13 July 
B 15 July 
31 15 July 
20 15 July 
14 7 July 
13 17 July 
10 17 July 
4 2 
2 bis 


tured were still lactating. The mean body mass of 20 
pregnant females was 7.99 g. This was significantly 
heavier than the body mass of 32 lactating individuals 
(x= 7.05 g; t= 2.86, P = 0.01), as well as 28 nonpreg- 
nant adult females captured after the reproductive sea- 
son (X= 6.73 g; t= 5.04, P < 0.0001). The first volant 
juveniles were netted on 25 June; 20 days after the 
earliest lactating females were taken. 

Red Bats — Pregnant females were taken from 15- 
30 May. Lactating females were netted from 5 June 
through 7 July. As with M. septentrionalis, female 
Red Bats also apparently limited activity during the 
week between captures of the last pregnant individu- 
als and the first females that were lactating. Only 
eight Red Bats were taken during this period; seven 
were males (7? = 4.50, P < 0.05). The mean body 
mass of 13 pregnant Red Bats (16.4 g) was not sig- 
nificantly heavier than that of nine lactating individ- 
uals (kX = 14.1 g; t= 2.03, P = 0.076). Volant young 
were first taken on 13 July, three weeks after 
Northern Long-eared Bats, and 38 days after the ear- 
liest lactating females were taken. 


TABLE 2. Number of female bats mist netted during each of three 2-hour time periods, based 
on reproductive condition, on Shawnee National Forest, southern Illinois, during spring and 


summer 1999 and 2000. 


Species/ 
Reproductive 
Condition 20:00-21:59 
Northern Long-eared Bat 
Pregnant 18 
Lactating 12 
Nonpregnant 12 
Red Bat _ 
Pregnant 8 
Lactating 5 
Nonpregnant 1 
Eastern Pipistrelle 
Pregnant 13 
Lactating ri 


Nonpregnant 5 


Time Periods Captured 


22:00-23:59 24:00-02:00 

2 0 
8 13 
10 3 
5 1 
3 I 
0 

4 7 
3 3 
4 3 


422 


Eastern Pipistrelles — Pregnant females were taken 
throughout an eight-week period from 19 May 
through 17 July. Lactating females were taken from 
25 June through 17 July. Lactation in Eastern 
Pipistrelles in southern Illinois continues well into 
August given that pregnant individuals occurred as 
late as 17 July. However, we did not take any later 
than this during the first field season, even though 
netting continued until mid-August. The mean body 
mass of 21 pregnant females was 7.43 g, not different 
than the body mass of 10 lactating individuals (6.7 g; 
t= 1.44, P = 0.182). Considered together, Pipistrelles 
that were either pregnant or lactating had a greater 
mean body mass than six nonpregnant females taken 
following the reproductive season (X= 6.67 g; t = 
4.03, P = 0.01). Volant young were first netted on 15 
July. As in Northern Long-eared Bats, this was 20 
days after the earliest lactating females were taken. 
Volant young P. subflavus emerged about the same 
time as Red Bats and most other bat species (Table 1) 
we took in southern Illinois. 


Female Foraging Activity Periods 

Based on reproductive condition, trends were evi- 
dent in the timing of foraging activity for some 
species (Table 2). Pregnant Northern Long-eared 
Bats were taken significantly more often early in the 
evening (y? = 29.14, P < 0.0001), with no individu- 
als mist netted after midnight. Within the time frame 
we mist netted each evening, a bimodal activity pat- 
tern was evident for lactating Northern Long-eared 
Bats, with reduced activity from 2200 — 2400 h, 
although the differences were not statistically signifi- 
cant (7 = 5.63; 0.05 < P < 0.10). Trends were less 
apparent for Red Bats and Eastern Pipistrelles, 
although a bimodal activity pattern was also suggest- 
ed for pregnant Pipistrelles (vy? = 5.24; 0.05 < P< 
0.10). 


Discussion 

Cause and effect in the timing of reproductive 
events in mammals often is difficult to identify 
because of the interrelationships of numerous life 
history characteristics. This is especially true of bats 
because they do not fit the typical reproductive pat- 
tern for similar-sized terrestrial mammals. Bats have 
small litter size, relatively long gestation, and large 
neonatal body mass (Kunz and Hood 2000). 
Lactation is particularly demanding energetically. As 
noted by Racey (1982: 63) “. . . adequate food sup- 
ply during lactation and weaning is the most impor- 
tant selection pressure in the timing of mammalian 
reproductive cycles.” Vespertilionids in Ilinois, 
because they are insectivorous, are ultimately con- 
strained reproductively by the seasonality of insect 
abundance. Thus, pregnancy and lactation must be 
timed to coincide with the abundance of energy 
available in the spring and early summer (Racey and 
Entwistle 2000). 


THE CANADIAN FIELD-NATURALIST 


Volos 


Some early information on the timing of reproduc- 
tion in Northern Long-eared Bats is anecdotal 
(Brandon 1961; Easterla 1968). Most investigators 
report later breeding dates in M. septentrionalis than 
we found. For example, Kunz (1971) found pregnant 
females in central lowa from 20 May until 23 June, 3 
weeks later than we did. He also reported the first 
volant young occurred on 23 July — a month later 
than we took them. Whitaker and Hamilton (1998: 
101) stated for this species that “... the single young 
is often born in July, or later than in most other east- 
ern U. S. bats.” Lactating females in South Dakota 
(Turner 1974) were taken in mid-August, a month 
later than we found them. There appears to be much 
geographic variation in the timing of pregnancy and 
lactation in Northern Long-eared Bats, with more 
northern populations active later than in southern 
Illinois. 

Based on previous studies, less variation may 
occur in the timing of reproductive events in Red 
Bats (Shump and Shump 1982) than in Northern 
Long-eared Bats. Hoffmeister (1989) reported that 
parturition occurs in late May or early June in 
Illinois, which coincides with when we found preg- 
nant females. Similar timing is reported for the adja- 
cent states of Kentucky (Barbour and Davis 1974) 
and Missouri (Schwartz and Schwartz 1981). 

Parturition and lactation in P. subflavus in southern 
Illinois was much less synchronized than in Northern 
Long-eared or Red Bats. Reduced body size and roost 
temperatures are two strong selection pressures for 
synchrony of parturition (Tuttle and Stevenson 1982). 
Because they are smaller than Red Bats and more 
affected by ambient temperature, Eastern Pipistrelles 
could be expected to come out of hibernation later, 
with associated affects on ovulation, pregnancy, and 
lactation (Racey 1982). Later initiation of reproduc- 
tive activity and high synchrony was not the case in 
southern Illinois, however, although it may occur in 
populations at higher latitudes (Lane 1946; Davis 
1963; Fujita and Kunz 1984). We first encountered 
pregnant Eastern Pipistrelles about as early as 
Northern Long-eared and Red Bats, but also took 
them much later in the summer, as was the case with 
lactating females. Young Eastern Pipistrelles begin to 
forage by the time they are one month of age (Fujita 
and Kunz 1984). We took juveniles in mid-June, sug- 
gesting parturition occurred in mid-May. Given the 
lengthy period during which pregnant and lactating 
Pipistrelles were taken, it is not surprising that there 
was no apparent week-long cessation of foraging 
activity as noted in female Northern Long-eared and 
Red Bats. 

The activity patterns we noted for pregnant and 
lactating females of each species are typical of ves- 
pertilionids in temperate regions, and insectivorous 
bats generally (Erkert 1982). For example, Kunz 
(1974) reported that pregnant Cave Myotis (Myotis 
velifer) in south-central Kansas emerged to forage 


2001 


before lactating females. Feeding activity was bi- 
modal with a secondary period prior to sunrise. A 
later secondary foraging period probably occurred 
with our populations too; however, we ended mist 
netting at 0200 h prior to the second peak. Bimodal 
foraging of lactating bats, suggested most strongly in 
our study for Northern Long-eared Bats, also might 
be expected because most bat species return to their 
roost during the night to suckle young (Racey 1982) 
and are not available to mist net. 

Significant differences in body mass of Northern 
Long-eared Bats were evident based on reproductive 
status. Considered together, pregnant and lactating 
Eastern Pipistrelles also were heavier than nonrepro- 
dutive females. Conversely, no differences in body 


mass associated with reproductive condition were. 


found in female Red Bats, the largest of the three 
species. We suspect our results are influenced by a 
number of interacting selective factors, including the 
relationship between adult body mass and neonatal 
size and litter size, as well as wing loading and asso- 
ciated aerodynamic constraints on flight and forag- 
ing ability (Hayssen and Kunz 1996; Kunz and Hood 
2000). 

Size is also a factor determining when young 
bats first became volant, as is roost temperature 
(Tuttie and Stevenson 1982). The 38-day time peri- 
od to volancy we estimated in juvenile Red Bats 
was almost twice as long as the 20-day period for 
juvenile Northern Long-eared Bats and Eastern 
Pipistrelles. Like most species of bats, M. septentri- 
onalis have singleton litters. Eastern Pipistrelles 
usually have twins, whereas Red Bats have litters 
of 3-4 pups, one of the largest litter sizes in bats 
(Hayssen et al. 1993). Neonates of Northern Long- 
eared Bats and Pipistrelles probably develop faster 
because with smaller litters, they are born at a rela- 
tively advanced size. Smaller species also produce 
relatively larger young (Tuttle and Stevenson 1982; 
Hayssen and Kunz 1996). Even if only one or two 
of the Red Bat pups in a litter survive to volancy, it 
still takes them longer because they are relatively 
small at birth. 

Because this project was not designed specifically 
to study reproduction, pregnant females may have 
been flying earlier in the spring than when we began 
mist netting. Nonetheless, even had we started earli- 
er, we probably would not have been able to detect 
females in the earliest stages of pregnancy by palpa- 
tion. Our field seasons certainly encompassed the 
last dates of pregnancy and the duration of the lacta- 
tion period for each spécies. 


Acknowledgments 

This study was part of a project funded by the 
U.S. Forest Service, Shawnee National Forest, 
Challenge Cost-Share Agreement No. 98-0908-24. 
We thank Steve Widowski and Mike Spanel, 


FELDHAMER, CARTER, AND CARROLL: BATS IN SOUTHERN ILLINOIS 


423 


Shawnee NF, for their loan of equipment and help in 
all phases of this study, and to numerous individuals 
who participated in mist netting. 


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Received 25 November 2000 
Accepted 19 December 2001 


Breeding Bird Declines in the Boreal Forest Fringe of 
Western Canada: Insights from Long-term BBS Routes 


EnID E. CUMMING!, KEITH A. HOBSON?2, and STEVEN L. VAN WILGENBURG2 


11542 Empress Avenue, Saskatoon, Saskatchewan S7K 3G3 Canada 
2Environment Canada, Prairie and Northern Division, 115 Perimeter Road, Saskatoon, Saskatchewan S7N 0X4 Canada 


Cumming, Enid E., Keith A. Hobson, and Steven L. Van Wilgenburg. 2001. Breeding bird declines in the boreal forest 
fringe of western Canada: Insights from long-term BBS routes. Canadian Field-Naturalist 115(3): 425-435. 


We examined North American Breeding Bird Survey (BBS) data for five routes affected by loss and fragmentation of local 
forest habitat and drainage or degradation of wetlands. We focused on the Brightsand, Saskatchewan (1979-1996) route 
since it once recorded the highest species richness for routes in Canada and has now dropped dramatically. We also 
examined other routes, Athabasca, Alberta (1972-1996), Two Hills, Alberta (1972—1996), Clouston, Saskatchewan (1973-— 
1996), Tyndall, Manitoba (1971-1996), and one route located (as a control) in contiguous forest at Bird River, Manitoba 
(1976-1996), to determine how changes in land use influenced relative abundance and diversity of bird species. Over the 
18-year history of the Brightsand route, species diversity declined from a maximum of 105 species in 1987 to 67 species in 
1995. In addition, 43 species recorded regularly prior to 1990, have disappeared, and 13 other species showed significant 
negative population trends. This pattern was also seen in Athabasca and Tyndall, as population trends for eight species 
declined, as did four species on the Two Hills and Clouston routes between 1971 and 1996. Analysis of land cover data 
along the Brightsand BBS route indicated a 48 to 55% loss of forest cover from 1963 to 1994. Significant declines were not 
restricted to a particular guild of birds, as forest, grassland and wetland species all declined. In contrast, none of the species 
on the Bird River control route showed significant population declines. We conclude that habitat loss, due primarily to 
forest clearing and wetland drainage and modification for agriculture in the forest fringe region of the Canadian prairies, 
has contributed to the decline of species and disappearance of others on several BBS routes. 


Key Words: Breeding Bird Survey, species declines, forest loss, habitat fragmentation, Manitoba, Saskatchewan, Alberta. 


The forested regions of the Aspen Parkland and 
the Boreal Forest Transition zone (Acton et al. 1998) 
host breeding bird communities among the most 
diverse in North America (Robbins et al. 1986; Price 
et al. 1995). The amount of natural habitat available 
to birds and other wildlife in this area is decreasing, 
however, as forest land is being converted for agri- 
culture, the remaining forests are fragmented, and 
wetlands are drained. Estimates using Landsat TM 
imagery of the southern boreal mixedwoods of 
Saskatchewan showed the rate of deforestation in the 
last 20 years as approximately 1.3% per year; which 
exceeds the world average (Fitzsimmons et al. 
1997*; Hobson and Van Wilgenburg, unpublished 
data). The effect of such habitat loss on bird commu- 
nities is difficult to assess because few long-term 
datasets exist (Kirk et al. 1997). The North American 
Breeding Bird Survey (BBS), however, is a source of 
long-term data that may provide insight into popula- 
tion trends at various temporal and geographic scales 
(Sauer et al. 1997*). 

The BBS, is a roadside survey with more than 
4100 permanent routes many of which are surveyed 
annually in early summer. Each route is 39.4 km 
(24.5 miles) long, with 3-minute point counts con- 
ducted at 0.8 km (0.5 mile) intervals for a total of 50 
point-count stops. All birds seen or heard within a 
0.4 km (0.25 mile) radius of each stop are recorded. 
These surveys begin 30 minutes before sunrise and 


normally require 4-5 hours for completion (Sauer et 
al. 1997*). We examined data from five long-term 
BBS routes in Alberta, Saskatchewan and Manitoba 
in order to evaluate the effects of local habitat 
changes, particularly forest loss, on breeding bird 
communities. Four of these areas had undergone 
substantial forest clearing and fragmentation. We 
contrast these data with a single long-term BBS 
route located in continuous forest. 

The BBS was originally designed to monitor pop- 
ulation change at broad levels. Several factors con- 
tribute to “noise” in BBS trend analyses including 
varying degrees of expertise and ageing among 
observers, as well as weather and random events on 
breeding, wintering, and stopover sites that may 
affect birds in one BBS route and not others. We rec- 
ognize these drawbacks but maintain that trend anal- 
yses along single, long-term, BBS routes, when com- 
bined with interpretations of overall habitat change 
along those routes, represents a valuable course of 
investigation. While conclusions from such analyses 
must be considered preliminary, this is the nature of 
scientific investigation, particularly as it relates to 
pressing conservation issues. 

At one time the BBS route at Brightsand, 
Saskatchewan, had the highest species diversity 
recorded on any Canadian BBS route, and one of the 
highest in North America (Johns 1990*). This route, 
one of few in the forest fringe area of Saskatchewan, 


425 


426 


was surveyed consistently for 18 years (1979-1996) 
by the same observer (Muriel Carlson). The number 
of species detected per year on the Brightsand route 
declined sharply between the 1980s and 1990s. 
Starting in 1990, species richness began to decrease, 
and by 1995, only 67 bird species were recorded on 
this route compared to a high of 105 species in 1987. 
Of those species still recorded on this route, almost 
all are declining and many of these declines are sig- 
nificant. 

In order to determine if species declines were 
related to local changes in habitat, or were a more 
widespread phenomenon, we examined other BBS 
routes surveyed consistently for long periods (..e., 
>20 years) both in forest fringe and in continuous 
forest habitat. Four additional routes with a similar 
suite of species were added to the analysis; 
Athabasca, Alberta, surveyed for 26 years by John 
Kinnaird; Two Hills, Alberta, surveyed for 14 years 
by Graham Greenle and the last 12 years by David 
Ealey; Clouston, Saskatchewan surveyed for 23 
years by Maurice Mareschal; and Tyndall, Manitoba, 
surveyed for 26 years (22 by Rudolf Koes). We 
chose the BBS Bird River, Manitoba, as a control 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


route. Bird River is the longest running BBS route 
(surveyed for 21 years by Peter Taylor) located in 
continuous forest habitat in the Prairie Provinces. 


Study Area : 

Brightsand, Saskatchewan (53° 30’ N, 108° 40’ 
W), Clouston, Saskatchewan (53° 06’ N, 105° 51’ 
W), Athabasca, Alberta (54° 50’ N, 113° 06’ W), 
Two Hills, Alberta (53° 44’ N, 111° 32’ W ), and 
Tyndall, Manitoba routes (50 04’ N, 96 36’ W), are 
all located in the Boreal Plain Ecozone (Acton et al. 
1998). This ecozone extends across Prairie Canada 
from southeastern Manitoba to northwestern Alberta 
and is bounded by the Precambrian Shield to the 
north and Aspen Parkland to the south (Figure 1). 
The Boreal Plain Ecozone is a gently rolling plain 
that was originally covered by boreal mixedwood 
forest. This forest is composed, of both deciduous 
[Trembling Aspen (Populus tremuloides), Balsam 
Poplar (P. balsamifera), and White Birch (Betula 
papyrifera)| and coniferous [Jack Pine (Pinus 
banksiana), White Spruce (Picea glauca), Black 
Spruce (P. mariana) and Balsam Fir (Abies bal- 
samea)| (Rowe 1972; Kabzems et al. 1986). The 


Boreal Plain 
Athabasca 


\ 


. Alberta 


Saskatchewan 


Two Hills 
Brightsand 
Clouston 
Tyndall 
Bird River 


PX OP HO 


Manitoba ,: 


300 
__——— SS _ 


Figure 1. Location of BBS routes in the study. 


600 Kilometers 


2001 


southern portion of the Boreal Plains is categorised 
as the Boreal Transition Zone (Kabzems et al. 1986; 
Acton et al. 1998). This area is an ecotone between 
the continuous forest of the north and the Aspen 
Parkland of the south. Like most ecotones, this 
blending of two different habitats promotes a high 
species diversity in the area (Smith 1992) and nearly 
300 species of birds breed here (Robbins et al. 1986; 
Acton et al. 1998). However, much of the Boreal 
Plains, especially the transition zone, has been 
cleared for agriculture (Kabzems et al. 1986; Acton 
et al. 1998; Hobson and Van Wilgenburg, unpub- 
lished data). In some areas, more than 80% of the 
forest has been cleared (Acton et al. 1998). Although 
most of this clearing took place with settlement, 60 
or more years ago, in other areas, agriculture contin- 
ues to expand into the forest, especially in more 
northern areas. 

Our continuous forest site, Bird River, Manitoba 
(50° 25’ N, 95° 42’ W) is located in the Boreal 
Shield Ecozone. This ecozone has similar forest 
cover to the Boreal Plains, but with more conifer and 
fewer deciduous trees. Bird species diversity is lower 
than in the Boreal Plains, but many of the same 
species occur (Smith 1993). 


Methods 

We examined North American Breeding Bird 
Survey (BBS) data for four routes assumed to be 
affected by loss and fragmentation of local forest 
habitat: Brightsand, Saskatchewan (1979-1996), 
Clouston, Saskatchewan (1973-1996, except 1976), 
Athabasca, Alberta (1972-1996), and Two Hills, 
Alberta (1971-1996). Tyndall, Manitoba 
(1971-1996), occurred in a region where local forest 
cover had increased (R. Koes, personal communica- 
tion) and Bird River, Manitoba (1976-1996), occur- 
ring in continuous forest that had not been impacted 
by agriculture or forestry were also chosen to deter- 
mine how changes in land use influenced relative 
abundance and diversity of bird species. Data for 
trends of individual bird species and raw data on 
Species numbers were obtained from the BBS 
WebPage (Sauer et al. 1997*). This source also sum- 
marised those species on each route that had declined 
significantly. We also analysed trends of bird species 
by guilds. The categories used were; forest, shrub- 
land, grassland, wetland and farm/urban (cosmopoli- 
tan) birds. Species were assigned to the different cat- 
egories using the criteria of the Canadian Breeding 
Bird Survey (Downes and Collins 1996). Trends in 
populations by species guilds were analysed by Brian 
Collins of the Canadian Wildlife Service in Hull, 
Quebec, using the program BBSANALYS (Collins 
1998*). Route trends are estimated using the methods 
described in Link and Sauer (1994). 

In addition to analyses of trends, we examined 
habitat change along the Brightsand BBS route. We 


CUMMING, HOBSON, AND VAN WILGENBURG: BREEDING BIRD DECLINES 


427 


quantified habitat for the route using two methods. 
First, the amount of habitat in 1963 was obtained by 
digitizing the 73F/10 National Topographic Series 
(NTS) 1:50 000 scale map using Atlas GIS V. 2.0 
(ESRI, Redlands California). That coverage was 
based on photo interpretation from 1963. We 
obtained Landsat TM satellite imagery from 1994 for 
the same area, and reclassified 26 cover types to 
three; forest, water and agriculture/cleared land, to 
approximate the classifications used in the NTS map- 
sheet. A georeferenced digital map of Forest 
Management Agreement areas (FMA) was used to 
compare loss of forest cover due to agriculture and 
forestry. Amount of each of the three habitat types 
was then assessed using the GIS system Idrisi for 
Windows v. 2.01 (Clark Labs, Clark University, 
Worcester, Massachusetts). Secondly, in order to 
more directly link habitat change with bird trends as 
determined by the BBS, we analysed a buffer area of 
400 metres on each side of the BBS route. This dis- 
tance was chosen because BBS stops are 800 metres 
apart; therefore, 400 metres on each side of a BBS 
stop represents the approximate distance over which 
data for a stop are recorded. 


Results 

The total numbers of species detected each year 
on the six BBS routes are shown in Figure 2. Species 
richness declined at Brightsand (slope =-1.65, 
r?=0.51, F=17.55, P<0.001), Clouston (slope = -0.47, 
r=0.27, P—8., P<0:05),.and Two Hills (slope= 
-0.92, r2=0.55, F=29.23, P<0.001) and increased at 
Tyndall, (slope= +0.42, r2=0.44, F=18.54, P<0.001). 
No trends were detected on the Athabasca or Bird 
River (control) routes. At Brightsand, the average 
number of species (+SD) detected per year was 95.7 
+ 2.7 before 1990 but this declined to 74.7 + 2.2 
after 1990 (Mann-Whitney U; U=4.0, Z=-3.13, 
P=0.0017). Almost every species on this route had a 
negative population trend, and thirteen species 
decreased significantly (Table 1). No species was 
significantly increasing on the Brightsand route. 
Athabasca and Tyndall each had eight species with 
significantly negative population trends (Table 1). 
Only Common Raven significantly increased on the 
Athabasca route (trend = +19.9, P=0.04 n=24). Four 
species showed a significantly increasing population 
trend on the Tyndall route (Savanna Sparrow, trend 
= +3.1, P=0.03, n=24; Chipping Sparrow, trend = 
+6.2, P=0.003, n=24; Red-eyed Vireo, trend = +9.6, 
P=0.002, n=24; and Yellow Warbler, trend = + 4.5, 
P=0.03, n=24). Clouston and Two Hills each had 
four species with significantly negative population 
trends (Table 1). Neither Clouston or Two Hills had 
any species with significantly increasing trends, 
although two species approach significance on the 
Clouston route (American Robin, trend = +5.3, 
P=0.06, n=25; Brown-headed Cowbird, trend = +9.8, 


428 THE CANADIAN FIELD-NATURALIST Vol. 115 
TABLE |. Species showing significant negative population trends for all routes examined. 
Route Species Average Trend P 
#/year 
Brightsand, Saskatchewan Blue-winged Teal 6.6 -11.2 aa 
n=18 years (1979-1996) Common Snipe Suk -10.5 i 
Sora Dd -12.4 sae 
Veery 1.6 -21.4 ae 
Swainson’s Thrush 4.6 -9.9 aay 
Red-eyed Vireo 63.7 -2.1 A 
Ovenbird 7 -9.9 ae 
Connecticut Warbler 13.4 -5.0 
Clay-coloured Sparrow 51.8 -5.8 ae 
Chipping Sparrow 3031 -11.2 a 
Song Sparrow 38.0 -5.2 ae 
Athabasca, Alberta Pied-billed Grebe 1.4 -6.4 her 
n=25 years (1972-1996) Blue-winged Teal 1.9 -5.6 oa 
Ruddy Duck 0.2 -8.3 $5 
Sora Ds -4.4 * 
Common Nighthawk 0.3 -9.8 e 
Red-eyed Vireo 31.0 -1.0 a 
Warbling Vireo 0.2 -14.2 Be 
Tennessee Warbler 2 -7.1 ah 
Two Hills, Alberta Red-necked Grebe 1.8 -7.2 huey 
n=26 years (1971-1996) Ruddy Duck lek -9.4 5 
Northern Flicker 2.6 -10.7 aces 
Least Flycatcher 22 -0.7 a 
Clouston, Saskatchewan American Coot 3) ai -2.0 nee 
n=23 years (1973-1996) Northern Flicker 3.9 -2.5 Aone 
Pied-billed Grebe 3.4 -4,2 = 
American Bittern 5.9 -4.8 bree 
Tyndall, Manitoba American Bittern 0.6 = IA ics 
n=26 years (1971-1996) Blue-winged Teal 0.4 -15.3 a 
Red-headed Woodpecker 0.4 -12.2 ee 
Least Flycatcher eS -2.4 Pax 
Gray Catbird 9.8 -2.0 = 
Veery 0.5 -8.7 co 
Northern Oriole 10.4 -3.4 pe 


BeOS. F< Ol: sse< 001 


P=0.07, n=25). On the Bird River route, no species 
had a significant trend, positive or negative. 

To assess whether any particular group of birds 
was affected more than others, route trend analysis 
was also done by grouping species into guilds using 
the program BBSANALYS (Collins 1998*). Species 
showing significant declines were not restricted to 
any particular habitat. On the Brightsand BBS route, 
all guilds showed negative population trends (Table 
2). Although forest and wetland birds showed the 
largest annual declines (-5.0 % and -8.7 %, respec- 
tively), grassland and cosmopolitan birds (magpies, 
crows etc.) also declined. Athabasca showed similar 
results to Brightsand, except that cosmopolitan birds 
increased (Table 2). While Two Hills and Clouston 
were similar to Brightsand and Athabasca in declines 
of grassland and wetland birds, both routes showed 
no declines in forest birds and shrubland birds were 


relatively stable (Table 2). Guild trends were differ- 
ent on Tyndall than they were on the other four for- 
est fringe routes. Wetland species on Tyndall 
declined as they had elsewhere, however, forest and 
cosmopolitan (disturbance-tolerant) species 
increased, whereas shrubland and grassland birds 
remained stable. 

Forest birds declined on both the Brightsand and 
Athabasca BBS routes with 10 year annual trends of 
-40.2% and -18.1%, respectively (Table 2). Even the 
abundant Red-eyed Vireo (species names given in 
Appendix A) declined on both these routes (Table 
1). Overall, forest birds increased slightly on the 
Tyndall route but Veery declined. In contrast, on the 
Bird River route (continuous forest), forest birds 
increased 12.4% over a 10-year average (Table 2). 
Species such as Swainson’s Thrush, Veery, Oven- 
bird, and Red-eyed Vireo that were declining on the 


2001 CUMMING, HOBSON, AND VAN WILGENBURG: BREEDING BIRD DECLINES 429 


110 
100 
90 
80 
70 
60 
50 


* Brightsand 


1970 1975 1980 1985 1990 1995 2000 


110 
100 


an as BS 
— i — a — er —) 


Athabasca 


40 
1970 1975 1980 1985 1990 1995 2000 


Number of Species Detected 


110 
100 
90 
80 
70 
60 
50 


fa Clouston 


1970 1975 1980 1985 1990 1995 2000 


110 
100 
90 
80 
70 
60 
50 


Bird River 
40 


1970 1975 1980 1985 1990 1995 2000 


110 
100 
90 
80 
70 
60 
50 


ao | Lyndall 


1970 1975 1980 1985 1990 1995 2000 


110 
100 
90 
80 
70 
60 
50 


Two Hills 


4 
1970 1975 1980 1985 1990 1995 2000 


Year 


FIGURE 2. Number of species observed each year on BBS routes. 


fragmented BBS routes, were either stable or slightly 
increasing on the Bird River route. 
Wetland-associated birds showed the largest 
decline in all five fragmented BBS routes. Many 
wetland species such as Pied-billed Grebe, American 


Bittern, Blue-winged Teal, Common Snipe and Sora 
all declined significantly on one or more routes. 
Blue-winged Teal showed significant declines on 
three of the BBS routes in the forest transition habi- 
tat (Table 1). All five observers (Carlson, Kinnaird, 


430 


THE CANADIAN FIELD-NATURALIST 


Vols 


TABLE 2. Population trends by species guild, in fragmented vs continuous forest habitat for all BBS routes examined. The 
10% change refers to an extrapolated ten-year trend based on the regression line for the existing data (B. Collins, personal 


communication). 
Route Trend Forest 
Fragmented 
Brightsand, Annual % Change -5.0 
Saskatchewan 10 year % Change -40.2 
Athabasca, Annual % Change -1.9 
Alberta 10 year % Change -18.1 
Two Hills, Annual % Change 0.1 
Alberta 10 year % Change lel 
Clouston, Annual % Change 0.8 
Saskatchewan 10 year % Change 8.6 
Tyndall, Annual % Change 0.7 
Manitoba 10 year % Change 6.8 
Continuous 
Bird River, Annual % Change 12 
Manitoba 10 year % Change 12.4 


Mareschal, Ealey and Koes), informed us that many 
of the marshes and small ponds along their BBS 
routes had been drained and broken for crops. Only 
at Bird River, the contiguous forest site, did all 
guilds remain stable or increase. This was the only 
BBS route examined where wetland birds increased. 
By contrasting landscape changes along the 
Brightsand BBS route with changes occurring within 
the larger Landsat mapsheet (inside and outside of 
the FMA) we were able to examine to what extent 
our BBS analysis was typical of a much larger area. 
A significant amount of habitat change occurred 
along the Brightsand BBS route between 1963 and 
1994 (Table 3). Of forest cover within the 400 metre 
buffer in 1963, only 48.2% remained in 1994. 
Similarly, only 54.7% of forest outside of FMAs 
remained in 1994 when the entire mapsheet was 
examined (Figure 3).Within the BBS buffered area, 
87.2% of the water remained in 1994, whereas 
96.1% remained of all waterbodies outside of the 
FMA over the entire mapsheet. For waterbodies 
inside the FMA however, 84.9% remained in 1994. 
Amount of land under cultivation increased by 


Habitat Guild 
Shrubland Grassland Wetland Cosmopolitan 

-4,2 -2.9 -8.7 -1.2 
-34.6 -25.6 -59.8 -11.5 
-4.1 -0.4 -6.8 0.8 
-34.0 -3.9 -50.5 7.9 
Or -2.9 -8.5 -2.1 
1.20 -25.3 -58.8 -19.0 
-0.10 -2.7 -1.9 -0.4 
-1.01 -23.8 -17.6 -3.5 
-0.2 0.1 -3.5 Se 
-2.2 2 -29.9 77.6 

5) n/a Sul -0.2 
16.4 n/a 36.2 -1.9 


85.6% over the same time period within the 400 
metre buffer, and by 117.6% for the entire mapsheet. 


Discussion 

The number of species recorded along the 
Brightsand BBS route has decreased significantly 
since the 1980s and many of the species remaining 
have declined significantly in relative abundance. 
The most likely reasons for these declines are a com- 
bination of forest and wetland habitat loss and frag- 
mentation. Our GIS analyses showed that along the 
Brightsand BBS route in the past 31 years, 51.8% of 
the forest and 12.8% of the wetlands have been lost. 
In addition, many of the pastures in this area have 
been broken for crops (Muriel Carlson, personal 
communication). This may explain declines in abun- 
dance of forest, grassland and wetland species. 

Similar patterns of species declines were also 
detected on the Athabasca, Two Hills, and Clouston 
BBS routes. Coinciding with the opening of the 
Alpac Mill in 1993, many farm woodlots were 
cleared and much of the forest habitat disappeared 
from this area (John Kinnaird, personal communica- 


TABLE 3. Habitat change for the Brightsand BBS route and mapsheet, determined from a digitized NTS map (1963) and 
Landsat TM satellite imagery (1994). FMA refers to Forest Management Area. 


Within 400 meters of BBS route _ Entire Mapsheet 

1963 1994 Change Change/year 1963 1994 Change Change/year 

(Ha) (Ha) % (%) (Ha) (Ha) To (%) 
Water 79.8 69.6 -12.8 -2.81 IWiG29%3 11180.6 -3.9 -3.10 
Forest 1905-6 ae SOD -51.8 -1.56 43845.0  23973.3 -45.3 -1.76 
Agriculture/Cleared Land 1163.7 2160.3 85.6 5.99 728035 (676009) 7 allio 7.02 
Water inside FMA = - - ~ 1349.6 1145.9 -15.1 -2.74 
Forest Cover in FMA = — ~ - TATOO eb 97 3.8 ilail 3.26 


2001 


ie 


~ el ase * - x. 

Jue 4 a od Res #. 3 
03 6 

EE 


9 kilometers 


CUMMING, HOBSON, AND VAN WILGENBURG: BREEDING BIRD DECLINES 


43] 


Figure Legend: 


Water 
M) Forest Outside FMA 


[J Agriculture 
MFMA 


FiGuRE 3. The Brightsand BBS route and the buffered area analysed from digital coverage for 1963 (NTS) and 1994 


(Landsat TM). FMA refers to Forest Management Area. 


tion). On the Two Hills route, dramatic changes in 
species richness and abundance occurred since 1985. 
This involved a period of road straightening and 
widening corresponding with the removal of trees 
and shrubs along roadways. A major tent caterpillar 
infestation took place in the late 1980s which was 
followed by extensive clearing of aspen forest. A 
period of dry years in the late 1980s resulted in many 
wetlands being cleared for agriculture. During the 
past five years, farmers have taken advantage of pri- 
vate timber sales to the Alpac mill and this has 
resulted in loss of woodlots, especially mixedwoods. 
Finally, a recent increase in livestock production, 
including elk ranching, has resulted in intensive land 
use that has impacted bird habitat (David Ealey, per- 
sonal communication). On the Tyndall BBS route, 
most of the forest clearing took place 60 or more 
years ago. Some of the farms in the area have since 
been abandoned and patches of young aspen are 
starting to mature (Rudolf Koes, personal communi- 
cation). This may explain why overall species diver- 
sity increased on this route. 

On the Brightsand route, several species occurring 


in the 1980s disappeared in the 1990s. Many of these 
declines were not statistically significant due to low 
statistical power since most of these species occurred 
in low numbers. From a biological perspective, how- 
ever, it is worth noting that species which formerly 
bred in this area have since disappeared. In 1982, the 
Brightsand BBS route had the second highest num- 
ber (n=25) of Connecticut Warblers recorded for any 
BBS route in North America (Price et al. 1995), by 
1995, only five were recorded. Ovenbird declined 
from 20 in 1979 to 2 by 1996, and Veery declined 
from 10 in 1979 to zero by 1992. 

Roadside habitat loss along the Brightsand BBS 
survey route is representative of landscape change at 
a much larger scale. For example, in the area south 
of Prince Albert National Park, an area also in the 
forest transition zone, estimates using Landsat satel- 
lite images have shown that the rate of deforestation 
in the last 20 years has been approximately 1.3% per 
year (Hobson and Van Wilgenburg, unpublished 
data; Environment Canada 1991*; Fitzsimmons et al. 
1997*; Alberta Environmental Protection 1998*). 
The rate of deforestation found in this and the above 


432 


mentioned studies far exceed the world average of 
0.3% per year and come close to the highest rates in 
the world (FAO 1999*). Remaining forest patches, 
are becoming more fragmented and isolated. Birds 
requiring forest interiors for breeding do not occur in 
small woodlots, and other species that breed in these 
woodlots may find them to be “ecological sinks” 
(Pulliam 1988; Robbins et al. 1989; Donovan et al. 
1995; Hobson and Bayne 2000). Birds nesting in iso- 
lated forest fragments can suffer from higher rates of 
nest depredation and nest parasitism reducing their 
reproductive success (Wilcove 1985; Wilcove and 
Robinson 1990; Robinson et al. 1995). 

Some species declines may be exacerbated by 
habitat degradation in the Neotropics (Askins et al. 
1990; Rappole and McDonald 1994; Sherry and 
Holmes 1996), however, many of the species which 
have declined or disappeared on the BBS routes are 
not restricted to this group. (e.g., American Bittern, 
Common Snipe, and Northern Flicker). Neotropical 
migrants such as Veery, Swainson’s Thrush and 
Ovenbird, declined in the forest transition areas, but 
were stable on the continuous forest Bird River BBS 
route. This strongly suggests an influence of breed- 
ing habitat change at the BBS route level. Kirk et al. 
(1997) in their comparison of surveys conducted at 
forested sites in boreal Manitoba and Saskatchewan 
between the 1970s and 1990s also present evidence 
for changes in breeding habitat as causing population 
declines of Neotropical migrants. 

Canadian BBS records from 1966 to 1994 show 
that in the Boreal Plains ecoregion, 15 species had a 
significant negative population trend; Pied-billed 
Grebe, Northern Pintail, Ruddy Duck, Northern 
Harrier, Killdeer, Lesser Yellowlegs, Franklin’s 
Gull, Black-billed Cuckoo, Short-eared Owl, Horned 
Lark, Veery, Chipping Sparrow, Clay-coloured Spar- 
row, Song Sparrow and Bobolink (Downes and 
Collins 1996). Many of these were the same species 
that also declined or have disappeared from 
Brightsand and the other BBS routes we examined in 
the forest fringe portion of this ecoregion. Similarly, 
American Bittern, Common Snipe, Sora, Black Tern, 
and Veery, species showing declines across North 
America (1966-1993, Price et al. 1995), also 
declined or disappeared at Brightsand, but not on our 
Bird River control route. 

We recognize that our control route was inade- 
quate for all species recorded along those routes sub- 
ject to habitat loss (i.e. our “experimental” routes). 
Ideally, the use of control routes that closely 
matched habitat characteristics at the beginning of 
the sampling period for all experimental routes 
would have been more useful but such a paired 
design was impossible in our region. Our control 
route was located in a different ecoregion than the 
experimental routes but the forest and wetland bird 
communities were similar. Thus, at the level of 


THE CANADIAN FIELD-NATURALIST 


Vols 


species by species comparisons, we feel that the con- 
trol area was valid. | 

Habitat alteration and fragmentation in the south- 
ern boreal forest transition zone is a serious conser- 
vation problem that has largely been overlooked by 
the scientific and conservation community. 
Expansion of agriculture, clearing of forest, and 
draining of wetlands is undoubtedly contributing to 
the decline of many avian species. For example, in 
the Aspen Parkland and southern boreal mixedwoods 
of Saskatchewan, many forest interior species did 
not occur in small, isolated woodlots (Johns 1993; 
Hobson and Bayne 2000). We can expect declines in 
these species as forest is removed and fragmented. 
Similarly, intensification of agriculture also results 
in wetland loss and alteration as well as the loss of 
meadows and parkland with clear impacts on birds 
associated with these habitats. We have shown that 
such habitat alterations correspond well with BBS 
data from this area. We encourage similar examina- 
tion of other BBS datasets, particularly at the region- 
al level where habitat alterations can be quantified. 


Acknowledgments 

We thank the numerous volunteers across North 
America who participate every year in the Breeding 
Bird Survey. Our special thanks to Muriel Carlson 
for the Brightsand BBS route, and to Peter Taylor for 
Bird River, Rudolf Koes for Tyndall, Maurice 
Mareschal for Clouston, John Kinnaird for Atha- 
basca, and Graham Greenle and David Ealey for 
Two Hills routes. We also thank Connie Downes and 
Brian Collins (C.W.S.) for analysing the species 
guild data and Gregg Babish (C.W.S.) for assistance 
with digitizing. Alan Smith and Brian Johns 
(C.W.S.), the current and former BBS co-ordinators 
for the province of Saskatchewan, brought to our 
attention the dramatic declines of the Brightsand 
route. Funding provided by a Canadian Wildlife 
Service operating grant to KAH. The critical com- 
ments of W. Easton, A. J. Erskine and one anony- 
mous reviewer improved an earlier draft of the 
manuscript. 


Documents Cited (marked * in text) 

Alberta Environmental Protection. 1998. The Final 
Frontier: protecting landscape and biological diversity 
within Alberta’s boreal forest natural region. Protected 
area report number 13. Natural Resources Service, 
Recreation and Protected Areas Division. Natural 
Heritage Planning and Evaluation Branch, Edmonton, 
Alberta. 

Collins, B. T. 1998. BBSANALYS: a program to analyse 
BBS data. National Wildlife Research Centre. Hull, 
Quebec. 

Environment Canada. 1991. The state of Canada’s envi- 
ronment-1991. Ottawa, Ontario. 

FAO. 1999. The state of the World’s forests. Food and 
Agriculture Organisation of the United Nations. Rome, 
Italy. 


2001 


Fitzsimmons, M., L. Patino, P. MacTavish, and P. 
Farrington. 1997. Estimating changes in forest area in 
central Saskatchewan, Canada. Poster, presented at the 
3rd National Ecological Monitoring and Assessment 
Network Conference, Saskatoon, Saskatchewan, 
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Johns, B. W. 1990. Saskatchewan breeding bird survey. 
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Sauer, J. R., J. E. Hines, G. Gough, I. Thomas, and B. G. 
Peterjohn. 1997. The North American breeding 
bird survey: results and analysis. Version 96.4. 
Patuxent Wildlife Research Centre, Laurel, Maryland. 
http://www.mbr.nbs.gov/bbs/bbs. html 


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Donovan, T. M., F. R. Thompson 3rd, J. Faaborg, and J. 
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Downes, C. M., and B. T. Collins. 1996. The Canadian 
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Hobson, K. A., and E. Bayne. 2000. Effects of forest 
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Johns, B. W. 1993. Influence of grove size on bird 
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Kabzems, A., A. L. Kosowan, and W. C. Harris. 1986. 
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Kirk, D. A., A. W. Diamond, A. R. Smith, G. E. Holland, 
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CUMMING, HOBSON, AND VAN WILGENBURG: BREEDING BIRD DECLINES 


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est birds in Saskatchewan and Manitoba. Wilson Bulletin 
109: 1-27. 

Link, W. A., and J. R. Sauer. 1994. Estimating equations 
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atlas of North American birds. Academic Press, London. 

Pulliam, J. R. 1988. Sources, sinks, and population regu- 
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Rappole, J. H., and M. V. McDonald. 1994. Cause and 
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111: 652-660. 

Robbins, C. S., D. Bystrak, and P. H. Geissler. 1986. 
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Sherry, T. W., and R. T. Holmes. 1996. Winter habitat 
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Academic Publishing, The Hague, The Netherlands. 


Received 10 November 1999 
Accepted 29 November 2001 


APPENDIX A. All birds recorded on the Brightsand BBS route (141 species). Species in bold are those which disappeared 


in the 1990s. 


Non-Passerines 


Common Name 


Common Loon 
Pied-billed Grebe 
Western Grebe 

Eared Grebe 

Horned Grebe 
Red-necked Grebe 
American White Pelican. 
Double-crested Cormorant 
American Bittern 
Great-blue Heron 
Black-crowned Night-Heron 
Sandhill Crane 

Canada Goose 


Scientific Name 


Gavia immer 

Podilymbus podiceps 
Achmorphorus occidentalis 
Podiceps nigricollis 
Podiceps auritus 

Podiceps grisengena 
Pelecanus erythrorhynchos 
Phalocrocorax auritus 
Botaurus lentiginosus 
Ardea herodias 

Nycticorax nycticorax 
Grus canadensis 

Branta canadensis 


Passerines 
Common Name 


Olive-sided Flycatcher 
Western Wood-Pewee 


Scientific Name 


Contropus borealis 
Contropus sordidulus 


Alder Flycatcher Empidonax alnorum 
Least Flycatcher Empidonax minimus 
Eastern Phoebe Sayornis phoebe 
Say’s Phoebe Sayornis saya 
Great-crested Flycatcher Myarchus crinitus 
Eastern Kingbird Tyrannus tyrannus 
Horned Lark Eremophila alpestris 
Purple Martin Progne subis 


Tree Swallow 
Bank Swallow 
Barn Swallow 


Tachycineta bicolor 
Ripara ripara 
Hirundo rustica 


Continued 


434 


APPENDIX A. (Continued). 


Non-Passerines 
Common Name 


Green-winged Teal 
Mallard 

Northern Pintail 
Blue-winged Teal 
Nothern Shoveler 
Gadwall 

American Widgeon 
Canvasback 
Redhead 
Ring-necked Duck 
Lesser Scaup 
White-winged Scoter 
Common Goldeneye 
Bufflehead 
Red-breasted Merganser 
Ruddy Duck 

Yellow Rail 

Sora 

American Coot 
American Avocet 
Killdeer 

Willet 

Spotted Sandpiper 
Common Snipe 
Franklin’s Gull 
Bonaparte’s Gull 
Ring-billed Gull 
Herring Gull 
California Gull 
Common Tern 
Black Tern 
Northern Harrier 
Red-tailed Hawk 
American Kestrel 
Merlin 

Ruffed Grouse 
Sharp-tailed Grouse 
Gray Partridge 
Mourning Dove 
Black-billed Cuckcoo 
Great-horned Owl 
Common Nighthawk 
Ruby-throated Hummingbird 
Belted Kingfisher 
Yellow-bellied Sapsucker 


Downy Woodpecker 
Hairy Woodpecker 


Pileated Woodpecker 
Northern Flicker 


THE CANADIAN FIELD-NATURALIST 


Scientific Name 


Anas crecca 

Anas platyrhynchos 
Anas acuta 

Anas discors 

Anas clypeata 

Anas strepera 

Anas americana 

Aythya valisineria 
Aythya ferina 

Athya collaris 

Aythya affinis 
Melanitta perspicillata 
Bucephala clangula 
Bucephala albeola 
Mergus serrator 

Oxyura jamaicensis 
Coturnicops noveboracensis 
Porzana carolina 
Fulica americana 
Recurvirostra americana 
Charadrius vociferus 
Catoptrophorus semipalmatus 
Actitis macularia 
Gallinago gallinago 
Larus pipixcan 

Larus phildelphia 

Larus delawarensis 
Larus argentatus 

Larus californicus 
Sterna hirundo 
Chlidonias niger 

Circus cyaneus 

Buteo jamaicensis 

Falco sparverius 

Falco columarius 
Bonasa umbellus 
Tympanuchus pallidicinctus 
Perdix perdix 

Zenaida macroura 
Coccyzus erythropthaimus 
Bubo virginianus 
Chordeiles minor 
Archilochus colubris 
Ceryle alcyon 
Sphyrapicus varius 


Picoides pubescens 
Picoides vilosus 


Dryocopus pileatus 
Colaptes auratus 


Passerines 
Common Name 


Gray Jay 

Blue Jay 

Black-billed Magpie 
American Crow 
Common Raven 
Black-capped Chickadee 
Boreal Chickadee 
Red-breasted Nuthatch 
House Wren 

Marsh Wren 
Ruby-crowned Kinglet 
Mountain Bluebird 
Veery 

Swainson’s Thrush 
Hermit Thrush 
American Robin 

Gray Catbird 

Brown Thrasher 
Sprague’s Pipit 

Cedar Waxwing 
European Starling 
Solitary Vireo 

Warbling Vireo 
Philadelphia Vireo 
Red-eyed Vireo 
Tennessee Warbler 
Orange-crowned Warbier 
Yellow Warbler 
Chestnut-sided Warbler 
Magnolia Warbler 
Yellow-rumped Warbler 
Palm Warbler 
Black-and-White Warbler 
American Redstart 
Ovenbird 

Northern Waterthrush 
Connecticut Warbler 
Mourning Warbler 
Common Yellowthroat 
Rose-breasted Grosbeak 
Spotted Towhee 
Chipping Sparrow 
Clay-coloured Sparrow 
Vesper Sparrow 
Savanna Sparrow 


LeConte’s Sparrow 
Sharp-tailed Sparrow 


Song Sparrow 

Lincoln’s Sparrow 
Swamp Sparrow 
White-throated Sparrow 
Dark-eyed Junco 
Bobolink 

Red-winged Blackbird 
Western Medowlark 
Yellow-headed Blackbird 


Vol yiis 


Scientific Name 


Perisoreus canadensis 
Cyanocitta cristata 
Pica pica 
Corvus brachyrhynchos 
Corvus corax 
Parus atricapillus 
Parus hudsonicus 
Sitta canadensis 
Troglodytes aedon 
Cistothorus palustris 
Regulus satrapa 
Sialia currucoides 
Catharus fuscescens 
Catharus ustulatus 
Catharus guttatus 
Turdus migratorius 
Dumetella carolinesis 
Toxostoma rufum 
Anthus spragueii 
Bombycilla cedrorum 
Sturna vulgaris 
Vireo solitarius 
Vireo gilvus 
Vireo philadelphicus 
Vireo olivaceus 
Vermivora peregrina 
Vermivora celata 
Dendroica petechia 
Dendroica penslyvanica 
Dendroica magnolia 
Dendroica cornata 
Dendroica palmatum 
Minotilta varia 
Setophaga ruticilla 
Seiurus aurocapilus 
Seiurus noveboracensis 
Oporornis agilis 
Oporornis philadelphia 
Geothlysis trichas 
Pheucticus ludovicianus 
Pipilo erthrophthalmus 
Spizella passerina 
Spizella pallida 
Pooecetes gramineus 
Passerculus 
sandwichensis 
Ammodramus leconteii 
Ammodramus 
caudacutus 
Melospiza melodia 
Melospiza lincolnii 
Melospiza georgiana 
Zonotrichia albicollis 
Junco hyenalis 
Dolichonyx oryzivorus 
Agelaius phoeniceus 
Sturnella neglecta 
Xanthocephalus 
xanthocephalus 


Continued 


2001 CUMMING, HOBSON, AND VAN WILGENBURG: BREEDING BIRD DECLINES 435 


APPENDIX A. (Concluded). 


Non-Passerines 
Common Name 


Scientific Name 


Passerines 
Common Name 


Rusty Blackbird 
Brewer’s Blackbird 


Common Grackle 
Brown-headed Cowbird 
Northern Oriole 

Purple Finch 

Pine Siskin 

American Goldfinch 
Evening Grosbeak 


House Sparrow 


Scientific Name 


Euphagus carolinus 
Euphagus 
cyanocephalus 
Quiscalus quiscula 
Molothrus ater 
Icterus galbula 
Carpodacus purpureus 
Carduelis pinus 
Carduelis tristis 
Coccothrausters 
vespertinus 
Passer domesticus 


The Spring and Fall Migrations of Scoters, Melanitta spp., at 
Confederation Bridge in the Northumberland Strait between 
New Brunswick and Prince Edward Island | 


PETER HICKLIN! and KATHERINE BUNKER-POPMA2 


‘Canadian Wildlife Service, Atlantic Region, P.O. Box 6227, Sackville, New Brunswick E4L 1G6 Canada 
240 Weldon Street, Sackville, New Brunswick E4L 4N4 Canada 


Hicklin, Peter, and Katherine Bunker-Popma. 2001. The spring and fall migrations of scoters, Melanitta spp., at 
Confederation Bridge in the Northumberland Strait between New Brunswick and Prince Edward Island. Canadian 
Field-Naturalist 115(3): 436-445. 


With assistance from volunteer observers on both sides of the Confederation Bridge, we counted and identified the three 
species of scoters migrating through the Northumberland Strait between New Brunswick and Prince Edward Island in 
spring and fall in order to assess whether the presence of Confederation Bridge affected the migration of scoters through 
the Strait. The numbers of scoters using Northumberland Strait during migration were three times greater in spring than 
fall. In both spring and fall, Surf Scoters were the most abundant species followed by Black and White-winged scoters. 
Only 13% of the scoters flew across Confederation Bridge in spring, and 22% in the fall. It is assumed that the remainder 
of the birds flew either (1) around Prince Edward Island in order to reach the Gulf of St. Lawrence in spring and the Strait 
of Canso in the fall or (2) high above Confederation Bridge and were not seen by the observers. 


Key Words: Black Scoter, Surf Scoter, White-winged Scoter, seaducks, migration, Northumberland Strait, Confederation 


Bridge, Cape Jourimain, New Brunswick, Borden, Prince Edward Island. 


Confederation Bridge, spanning 13 kilometers 
across Northumberland Strait between New 
Brunswick and Prince Edward Island (Figure 1), was 
begun in 1995 and completed in spring 1997. In 1990, 
the Canadian Wildlife Service (CWS) conducted sur- 
veys of seaducks and other seabirds in Northumber- 
land Strait from Cape Jourimain National Wildlife 
Area (MacKinnon et al. 1991). The present study 
(conducted from the same observation site) was initi- 
ated in 1997, immediately following completion of 
construction, to determine if the presence of the 
bridge affected migration of scoters through the 
Northumberland Strait during the birds’ spring and 
fall migrations. 


Materials and Methods 
Observation Sites 

In 1990, seaduck observations were conducted 
from Money Point (46°09'N, 63°49’W) on 
Jourimain Island on the New Brunswick side of 
Northumberland Strait (see MacKinnon et al. 1991). 
In spring 1997, a single observer conducted surveys 
from Money Point. During the fall, two observers 
conducted the observations, one at each end of the 
bridge: Money Point at Cape Jourimain, New 
Brunswick, and Borden Point in Borden, Prince 
Edward Island (see Figure 2). 


A. Cape Jourimain, New Brunswick 

Principally a farming area, Cape Jourimain may 
have been settled as early as 1720. In 1933, Jouri- 
main Island and Trenholme island were appropriated 
by the province of New Brunswick. By that time, the 


Jourimain area consisted of six farmsteads: two on 
the mainland and the two on each island, owned by 
members of the Allen and Trenholme families since 
1810 (Harries 1996). The road now leading to 
Confederation Bridge (including causeways between 
the two islands) was built in 1965 prior to its designa- 
tion as a National Wildlife Area by the CWS in 1979. 

Except for observations conducted during early 
morning, the sun was usually behind the observers in 
1997 thus allowing a clear view of the birds. All 
species seen from this observation site were recorded. 
A total of 49.8 person-hours of observations were 
conducted at Cape Jourimain in spring from 14 April 
to | May, inclusive, and similar surveys were under- 
taken from the same site over 156 person-hours dur- 
ing the fall migration between 18 September and 18 
November, inclusive. 

Volunteers used 7 X 35 mm and 10 X 42 mm 
binoculars and a Bausch & Lomb 15-60% telescope. 
The start and end times of observations were noted so 
that the rates at which species crossed the bridge 
(number of birds/hr) could be computed. 


B. Borden, Prince Edward Island 

The town of Borden was the former Prince Edward 
Island docking site of the Northumberland ferries 
(Figure 2). Observations from Prince Edward Island 
were carried out only during fall (nine days in October 
and nine days in November) for a total of 41.9 person- 
hours of observations. No surveys were undertaken 
from Prince Edward Island in 1990 (MacKinnon et al. 
1991). An observer was placed at Borden Point in 
1997 after a substantial number of seaducks first 


436 


2001 


HICKLIN AND BUNKER-POPMA: MIGRATIONS OF SCOTERS 


437 


FiGuRE 1. The Confederation Bridge across the Northumberland Strait between New Brunswick and Prince Edward Island. 


sighted from New Brunswick were seen to fly parallel 
to the bridge towards Prince Edward Island until they 
were lost from view and before they could be accu- 
rately identified. Thus, an observer on the Prince 
Edward Island side was necessary to count and identi- 
fy all the birds that reached visible range from the 
island. This observer was located under the bridge at 
Borden, looking towards New Brunswick to docu- 
ment only the numbers and species of seaducks, pri- 
marily scoters, reaching and crossing the bridge. 


Results 
1. Spring 

Forty-one species of birds were seen from Cape 
Jourimain at Confederation Bridge in spring (Table 
1). As this project was primarily concerned with scot- 
ers, only the observations of Black Scoters Melanitta 
nigra, Surf Scoters M. perspicillata and White- 
winged Scoters M. fusca are presented here. The 
numbers of scoters seen in the spring, and their rates 
of movement are shown in Table 2. 

The northward migration of scoters was already in 
progress when the observations began on 14 April, 
when 209 scoters were recorded (Table 2). The 
largest flocks of scoters seen in spring consisted of 
1200 Surf Scoters and 320 Black Scoters resting on 
the water, south of the bridge, on 17 April. The first 
White-winged Scoter (1 bird) was seen on 26 April 
and scoter numbers peaked the following day (150 
birds) with the last 16 White-wings observed on 1 
May, the final day of observations (Table 2). 

In the spring, a relatively low proportion of the 
scoters which reached the bridge actually crossed it. 
Many landed in nearby waters or continued flying 


along the bridge towards Prince Edward Island. On a 
daily basis, birds which were seen flying over 
Confederation Bridge ranged between 3.6%—100% of 
the totals which had migrated into the area. Overall, 
out of a total of 3986 scoters observed at the bridge in 
spring, 1997, 12.8% were seen to cross over the 
bridge (Table 3). 


2. Fall 

From the New Brunswick side, 47 species of birds 
were seen during fall observations between 18 
September and 18 November 1997 (see Table 1). The 
numbers of scoters seen from Cape Jourimain are 
shown in Table 4. The proportions of those birds 
which were seen to cross Confederation Bridge are 
presented in Table 5 and the numbers of scoters 
observed at Borden are shown in Table 6. 

Observations began in New Brunswick on 18 
September when 20 Surf Scoters and 7 White- 
winged Scoters were first observed flying towards 
the bridge (Table 4). They sharply veered away 
from the bridge and appeared to be disturbed by the 
presence of the structure. Clearly, the southward 
migration of scoters was already well underway by 
mid-September. In 1990, small numbers of Black 
and Surf scoters were seen on 7 September, the first 
of fall observations in that year (MacKinnon et al. 
1991). In 1997, between 6 and 18 October, the 
largest flocks seen throughout the 28-day observa- 
tion period at Cape Jourimain totalled 689 scoters 
(identified and unidentified). These included 220 
Surf Scoters (31.9%), 74 Black Scoters (10.7%), 71 
White-winged Scoters (10.3%) and 324 (47.1%) 
unidentified scoters (see Table 4). 


438 


New 
Brunswick 


Baie Verte 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


a4) 


Prince 
Edward 
Island | 


FIGURE 2. Location of the Confederation Bridge between New Brunswick and Prince Edward 


Island. 


The observations from the New Brunswick side 
showed that not all the scoters flew across the 
bridge once they reached it: on 14 of the 28 days 
when scoters were seen from New Brunswick (50% 
of the observations), birds were seen to fly over the 
bridge (see Table 5). Of the total numbers seen 
reaching the bridge on those days, between 4.5% 
and 66.1% of the birds recorded actually crossed 
over Confederation Bridge (Table 5). Overall, only 
22% of all scoters seen from New Brunswick dur- 
ing autumn, flew over Confederation Bridge. The 
remainder either turned away from the bridge or 
flew high, parallel to it towards Prince Edward 
Island, until they were lost from view by the New 
Brunswick observer. On the Prince Edward Island 


side, once scoters were seen, they were observed to 
fly over the bridge. However, we were unable to 
determine whether the two observers actually 
watched the same birds or if some of those birds 
flying high from New Brunswick were ever seen on 
the Prince Edward Island side because of the great 
heights they reached. 

From the Prince Edward Island side, observations 
were not begun until late in the migration period 
when maximum numbers were already being record- 
ed in New Brunswick. From 17 October to 18 
November, a total of 814 scoters were seen from 
Cape Jourimain (see Table 5). At Borden, from 17 
October to 26 November, 475 birds (58.4% of the 
New Brunswick total) were sighted (Table 6). 


2001 


Surf Scoter 
13 
11 
9 
7 
5 
3 
1 
260 270 280 290 300 310 320 
5 
2 Black Scoter 
= 13 
= 11 
® 9 
2) 7 
Ewch 
Oo 3 
o ; 
5 260 270 280 290 300 310 320 


White-winged Scoter 


= oO awn © 


260 270 280 290 300 310 320 
Day of Year (1 = 1 Jan) 


FiGurE 3. The numbers of scoters seen at Confederation 
Bridge from Cape Jourimain, New Brunswick, 17 
September (day 260) to 20 November (day 322) 
1997. 


In New Brunswick, all three species were first 
encountered on 18—20 September and Surf and Black 
scoters were still present on the last day of observa- 
tion (18 November); the last White-winged Scoters 
(2) were seen on 6 November (Table 4). Of the total 
numbers of scoters seen over the 28 days of observa- 
tions at Cape Jourimain, the percentages of the three 
species seen were 26.8% Surfs, 20.1% Blacks, 11.1% 
White-winged while 41.9% remained unidentified. 

The rates of movement (number of scoters seen per 
hour) on the New Brunswick side of the Strait ranged 
from a peak of 37.1/hr on 11 October to 1.3/hr on 13 
November (Table 4). From Borden, the rates of 
movement ranged from 44.6/hr on 20 October to 
1.0/hr on 3 and 4 November (Table 6). 

Based on the daily total numbers of birds seen at 
Cape Jourimain, the migration of scoters through the 
Northumberland Strait peaked in early to mid- 
October, between 6 and 20 October (Table 4). Peaks 


HICKLIN AND BUNKER-POPMA: MIGRATIONS OF SCOTERS 


439 


of White-winged Scoters remain unclear, possibly 
17-25 October, and Black Scoters peaked later, 
around 26 October to 3 November (Table 4). Hence, 
based on those birds identified by the observers, Surf 
Scoters reached Confederation Bridge in early 
October, White-winged Scoters in mid-October and 
Black Scoters in late October-early November 
(Figure 3). On the Prince Edward Island side, the 
largest numbers of scoters were seen on 22-28 
October, slightly later than when peak numbers were 
recorded in New Brunswick, although most could not 
be identified to species (see Table 6). 

Of all three species of scoters which were identi- 
fied from Cape Jourimain in October and November, 
Surf Scoters were clearly the predominant species 
using the Northumberland Strait during the fall migra- 
tion. Of these (836), 46.2% were Surf, 34.7% Black, 
and 19.1% White-winged scoters (see Table 4). 

Of the 1441 scoters observed in the fall from Cape 
Jourimain, only 316 (21.9%), flew across the bridge 
(Table 5). This proportion varied daily between 4.5% 
and 66.1% (Table 5). 


Discussion 

The numbers of scoters seen using Northumber- 
land Strait during northward migration were nearly 
three times greater than those in fall (3986 vs. 1441, 
respectively; see Tables 2 and 4). That the bridge 
proved to be a “barrier” to scoters migrating through 
the Northumberland Strait is suggested by the low 
proportions seen to fly over the bridge (12.8% in 
spring, 21.9% in fall). However, as the total numbers 
seen migrating through the area in daytime were rela- 
tively low (< 5000 birds), the bridge cannot be con- 
sidered a major impediment to the migration of the 
eastern breeding North American populations of Surf 
and Black scoters which number over 62 000 pairs 
(see Bordage and Savard 1995; Savard et al. 1998). 
We speculate that some birds which do not fly over 
Confederation Bridge detour around Prince Edward 
Island to reach the Strait of Canso where other sead- 
ucks (Common Eiders Somateria mollissima) have 
been seen to cross high over the Canso causeway 
(A. J. Erskine, personal communication; see Figure 
1). Birds which leave the New Brunswick coast and 
fly high towards Prince Edward Island along the 
bridge may similarly cross high over land once they 
reach the island. 

The 475 scoters seen on the Prince Edward Island 
side between 17 October and 26 November (Table 6) 
represent only 37% of the numbers counted on the 
New Brunswick side (1320) on the same dates. 
Furthermore, the numbers seen from Prince Edward 
Island do not correspond closely with the numbers 
seen on the New Brunswick side on the same days 
(see Tables 4 and 6). Therefore, these observations 
suggest that the remaining 54.5% either (1) crossed 
high overland at Borden, Prince Edward Island, and 


440 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


TABLE |. The species of birds seen in spring (°) and fall (©) off Cape Jourimain, New Brunswick, in spring (14 April and 
1 May, inclusive) and fall (18 September and 18 November, inclusive) 1997. 


GAVIFORMES 

Gaviidae 
Common Loon Gavia immer © 
Red-throated Loon, Gavia stellata *O 


PODICIPEDIFORMES 

Podicipedidae 
Horned Grebe Podiceps auritus © 
Red-necked Grebe Podiceps grisegena © 


PROCELLARIIFORMES 
Procellariidae 
Leach’s Storm Petrel Oceanodroma leucorhoa © 


PELECANIFORMES 
Sulidae 
Northern Gannet Sula bassanus *© 
Phalacrocoracidae 
Double-crested Cormorant Phalacrocorax auritus * © 
Great Cormorant Phalacrocorax carbo « 


CICONITIFORMES 
Ardeidae 
Great Blue Heron Ardea herodias *© 


ANSERIFORMES 

Anatidae 
Canada Goose Branta canadensis *© 
American Black Duck Anas rubripes *© 
Northern Pintail Anus acuta © 
Scaup Aythya spp. ° 
Common Eider Somateria mollissima ° 
Harlequin Duck Histrionicus histrionicus 
Oldsquaw Clangula hyamalis * © 
Black Scoter Melanitta nigra ° 
Surf Scoter Melanitta perspicillata *© 
White-winged Scoter Melanitta fusca * © 
Common Goldeneye Bucephala clangula ° 
Common Merganser Mergus merganser °© 
Red-breasted Merganser Mergus serrator *© 


FALCONIFORMES 
Accipiditricidae 
Osprey Pandion haliaetus ° © 
Bald Eagle Haliaeetus leucocephalus * © 
Northern Harrier Circus cyaneus *© 
Sharp-shinned Hawk Accipiter striatus * © 
Falconidae 
American Kestrel Falco sparverius 
Merlin Falco columbarius ° 


GALLIFORMES 
Phasianidae 
Grouse spp. ° 


were not seen by the observer there, or (2) turned 
away from the bridge and flew around the north shore 
of Prince Edward Island to get to the Strait of Canso 
and cross to the Atlantic shore. Both explanations are 


CHARADRITFORMES 

Scolopacidae 
Spotted Sandpiper Actitis macularia 
Sanderling Calidris alba © 

Laridae é 
Bonaparte’s Gull Larus philadelphia © 
Iceland Gull Larus glaucoides «© 
Greater Black-backed Gull Larus marinus ¢© 
Herring Gull Larus argentatus *© 
Caspian Tern Sterna caspia © 
Common Tern Sterna hirundo © 


STRIGIFORMES 

Strigidae 
Great Horned Owl Bubo virgianus ° 
Short-eared Owl Asia flammeus ° 


CORACIIFORMES 
Alcedinidae 
Belted Kingfisher Ceryle alcyon ° 


PICIFORMES 
Picidae 
Woodpecker spp. 0 
Northern Flicker Colaptes auratus ° 


PASSERIFORMES 
Hirundinidae 

Tree Swallow Tachicinetta bicolor 
Corvidae 

Blue Jay Cyanocitta cristata 

American Crow Corvus brachyrynchus * © 

Common Raven Corvus corax *© 
Paridae 

Black-capped Chickadee Parus atricappillus * 
Turdidae 

American Robin Turdus migratorius ¢© 
Mimidae 

Gray Catbird Dumetella carolinensis ° 
Sturnidae 

European Starling Sturnus vulgaris *© 
Parulidae 

Yellow-rumped warbler Dendroica coronata *© 
Icteridae 

Common Grackle Quiscalus quiscula « 
Emberizidae 

Savannah Sparrow Passerculus sandwichensis © 

Song Sparrow Melospiza melodia °© 

Dark-eyed Junco Junco hyemalis © 

Snow Bunting Plectrophenax novalis © 

Crossbill spp. © 

Common Redpoll Carduelis flammea © 


likely although we are unable to determine the relative 

proportions which might follow either potential route. 
There is also the distinct possibility that some scot- 

ers may cross the Botsford Peninsula and into Baie 


2001 


HICKLIN AND BUNKER-POPMA: MIGRATIONS OF SCOTERS 44] 


TABLE 2. The numbers of scoters seen in spring 1997 from Money Point at the base of Confederation 


Bridge in the Cape Jourimain National Wildlife Area. 


Date Time Period (Hrs) Total 
14 April 1015-1045 0 
y 1145-1215 200 
2 1230-1315 9 
Mean no./hr. (2.0 hours) - 
15 April 1100-1130 50 
e 1145-1230 0 
y 1330-1500 250 
Mean no./hr. (2.75 hrs) - 
16 April 1400-1500 20 
Mean no./hr. (1 hr) - 
17 April 1300-1700 1520 
Mean no./hr. (4 hrs) - 
18 April 153-1900 95 
Mean no./hr. (3.5 hrs) - 
20 April 1400-1800 56 
Mean no./hr. (4.0 hrs) - 
22 April 1300-2030 180 
Mean no./hr. (7.5 hrs) - 
23 April 1600-2015 109 
Mean no./hr. (4.25 hrs) _ 
26 April 0915-1315 536 
Mean no./hr. (4 hrs) - 
26 April 1500-1930 378 
Mean no./hr (4.5 hrs) ~ 
27 April 1500-1930 400 
Mean no./hr. (4.5 hrs) ~ 
1 May 1200-1400 0 
; 1700-1900 183 
Mean no./hr. (4 hrs) 183 
Totals & 
Mean No./hr. 46 hours 3986 


Verte, New Brunswick, a large bay just south of Cape 
Tourmentine and the bridge (see Figure 2). For exam- 
ple, one of us (K. B-P.) visited Baie Verte on 8 
November 1997, and noted White-winged Scoters “in 
the hundreds all strung out along and in the bay”. On 
that same day, very few scoters were observed at 
Cape Jourimain. This observation suggests that some 
scoters may cross to Baie Verte over the Botsford 
Peninsula, between Shemogue Harbour and Baie 
Verte, and were therefore not recorded by the observ- 
er at the base of Confederation Bridge. 

In 1990, over 22 hours of field observations, 
MacKinnon et al. (1991) recorded a total of 1405 
scoters migrating through Northumberland Strait in 
spring and fall. As in the present study, they were 
unable to identify 41% of these birds to species. Of 
the 824 that they did identify, 493 (59.8%) were Surf, 
191 (23.2%) Black, and 140 (16.9%) White-winged 


Numbers of Scoters 


Surf Black WW Unid. No./hr 

- - - - 0.0 

_ _ - 200 400.0 

- ~ - 9 12.0 

- _ -~ - 137.3 

_ - — 50 100.0 

_ — _ _ 0.0 

L235 125 _ - 89.2 

ee = = _ 174.1 

= 20 - - 20.0 

- - - - 20.0 

1200 320 — _ 380.0 

- - — - 380.0 

40 55 _ _ Zbl 

- - - - PAI | 

- 56 - — 14.0 

at = = = 14.0 
90 90 — — 24 

ns = ae = 24.2 

88 - _ oH PG sas 

_ - - - 25.6 
58 352. 1 125 134 
14.5 88 0.25 31.25 134 

50 95 6 D7). 84.4 

11.1 21.1 1.3 50.4 84.4 

150 100 150 _ 88.8 

33.3 222 33.3 _ 88.8 

- - - ~ 0.0 

_ 3 16 164 91.5 

- 3 16 164 45.8 

1801 1216 173 796 86.7 


scoters. The larger proportions of Surf Scoters seen 
by MacKinnon et al. (1991) were due to the relatively 
larger numbers of this species seen there in spring. 

The total numbers of scoters seen in the fall by 
MacKinnon et al. (1991) in 1990 and by the 
observers in the present study, were almost identical 
(1405 vs 1441 birds, respectively) even though 
MacKinnon et al. (1991) spent only 14% of the hours 
spent by the main observer (K. B-P.) in the present 
study (22 hours over both spring and fall seasons vs 
156 hours- fall season only). Hence, the great differ- 
ence in methodology most likely accounts for the dif- 
ferences between their results. 

For example, the migration rate for Surf Scoters on 
23 October 1990, was 237 birds/hr (MacKinnon et al. 
1991), or six times the rate we recorded (1.3-37.1 
birds/hr). The rates recorded for Black Scoters were 
18/hour on 12 October (MacKinnon et al. (1991: page 


442 


TABLE 3. The numbers of scoters flying to Confederation 
Bridge in spring 1997 from New Brunswick and the num- 


bers, and percent of those numbers, flying over the bridge. 


Total No. No. Over Per Cent 

Date Scoters Bridge Over Bridge 
14 April 209 9 4.3 
15 April 300 0 0 

16 April 20 0 0 

17 April 1520 0 0 

18 April 95 47 49.5 
20 April 56 2 3.6 
22 April 180 180 100.0 
23 April 109 Ph 193 
26 April (i) 536 92 2 
26 April (ii) ~ 378 138 36.5 
27 April 400 0 0 

1 May 183 23 12.6 
Total 3986 Si? 12.8 


i) 0915-1315 hrs 
ii) 1500-1930 hrs 


THE CANADIAN FIELD-NATURALIST 


Volts 


26, section 4.10.2), and for White-winged Scoters 
39/hour on 17 October 1990 (ibid). These are closer to 
the rates computed in 1997 for all three species com- 
bined in either New Brunswick or Prince Edward 
Island. | 

Using only the data on rates of southward migra- 
tion for 1990 and 1997, the post-Confederation Bridge 
results suggest that Surf Scoters were significantly 
and negatively affected by presence of the bridge but 
Black Scoters and White-winged Scoters were not. 
However, here too, the great differences in observa- 
tion hours between the two projects may be a factor. 

Nonetheless, both studies show that the North- 
umberland Strait was most important to Surf Scoters 
during fall migration and least to White-winged 
Scoters. Clearly, the scoters are affected by the pres- 
ence of the Confederation Bridge and some of the 
birds treat it as a complete barrier. White-winged 
Scoters may be the species which has most easily 
adapted to this change by choosing to cross overland 
either on the New Brunswick or the Prince Edward 
Island, as it was the one that appeared to fly across 
the bridge with the least hesitation. 


TABLE 4. The numbers (and mean number/hr) of scoters seen in the fall from Money Point at Cape Jourimain, New 


Brunswick, in 1997. 


Surf Black White-winged — Unidentified 

Date Time Period (hrs) Total No./hr Total No/hr Total No./hr Total No./hr Total No./hr 
18 Sept. 1400-1700 & 

1600-1700 (4) 27 6.8 20 4 — — 7 1.8 - — 
20 Sept. 0900-1000 & 

1030-1200 (2.5) 9 3.6 - - 9 3.6 - — - — 
21 Sept. 1645-1930 (2.75) 39 14.2 5) 1.8 10 3:6 _ - 24 8.7 
25 Sept. 1400-1530 (1.5) 4 Dah 1 0.7 ~ ~ - - 3 2.0 
29 Sept. 1100-1300 (2.0) 42 21.0 6 3.0 ~ - 1) 55) 23 NZS 
DOck 1100-1700 (5) 37) As P| 4.2 - - 16 BP = - 
3 Oct. 1300-1615 (3.25) 26 8.0 10 3.1 9 2.8 ~ - i DD, 
6 Oct. 1500-1700 (4) 70 IS) 26 6.5 - - - = 44 22.0 
9 Oct. 1200-1600 (4) 107 26.8 48 12.0 5 1163) — - 54.) 1355 
11 Oct. 0830-1300 (4.5) 167 Sail aD) 4.9 13 2.9 19 4.2 LIS ese 
13 Oct. 1445-1854 (4) 64 16.0 25 GS - - = — 39 OFS 
14Oct. 1430-1830 (4) 35 8.8 35 8.8 — — — = - - 
17 Oct. 0915-1315 (4) |P>) 3.3 34 8.5 3 0.8 39 9.75 AQ 1223 
18 Oct. 0835-1300 (4.5) 121 26.9 30 6.7 53 11.8 13 2.9 Des 5.6 
20 Oct. 0930-1230 (3) 54 18.0 — - D Om 23 Tal 29 Oey 
21 Oct. 1500-1830 (3.5) ») 1.4 3 0.9 1 0.3 — - 1 0.3 
23 Oct. 0900-1300 (4) 24 6.0 24 6.0 - — - - - - 
25 Oct. 0900-1300 (4) 64 16.0 6 ES) — — 20} 5.0 38 9.5 
26 Oct. 1400-1730 (5.5) 90 Dei 5 1.4 40 114 1 0.3 =: AG A AORG 
29 Oct. 0900-1300 (4) 89 DES 14 4.3 36 9.0 5 1.3 Sil 7.8 
30 Oct. 0900-1345 (4.75) 48 10.1 4 1S) Al 8.6 - — — = 
1 Nov. 0840-1240 (4) 45 11.3 6 iPS 25 6.3 4 1.0 10 2D 
3 Nov. 0920-1320 (4) 60 15.0 10 BES 30 ILS) — — 20 5.0 
6 Nov. 0815-1215 (4) Di 6.8 18 4.5 1 0.3 D 0.5 6 1.5 
8 Nov. 0815-1215 (4) 12 3.0 1 0.3 ~ — - ~ il 2.8 
11 Nov. 1230-1530 (3) 28 9.3 1 0.3 - — — - 27 9.0 
13 Nov. 0730-1130 (4) 5) 1:3 4 1.0 - — - - 1 0.3 
18 Nov. 0935-1335 (4) 119 4.3 1 0.3 12 3.0 - - 4 1.0 
Total 103.75 hours 1441 13.9 386 3.8 290 2.6 160 ES 605 5.8 


2001 HICKLIN AND BUNKER-POPMA: MIGRATIONS OF SCOTERS 443 


TABLE 5. The numbers of scoters flying to Confederation Bridge in fall 1997 from New Brunswick and the numbers and 
per cent flying over the bridge. 


Total No. No. Over Per Cent Total No. No. Over Per Cent 

Date Scoters Bridge Over Bridge Date Scoters Bridge Over Bridge 
18 September 27 0 0 21 October 5 0 0 

20 September 9 0 0 23 October 24 0 0 

21 September oe 0 0 25 October 64 8 ee, 
25 September 4 0 0 26 October 90 12 [334 
29 September 42 21 50.0 29 October * 89 4 4.5 

2 October 37 0 0 30 October 48 7 14.6 

3 October 26 0 0 1 November 45 4 8.8 

6 October 70 26 ail 3 November 60 0 0 

9 October 107 17 15.9 6 November 27 2 7.4 
11 October 167 49 29.3 8 November 12 0 0 

13 October 64 13 20.3 11 November 28 0 0 

14 October 35 0 0 13 November 5 0 0 

17 October 125 70 56.0 18 November Ie, 0 0 

- Se a is Total (mean) 1441 (51.5) 316(11.3) (21.9) 


TABLE 6. The numbers (and numbers per hour) of scoters seen in the fall from Borden Point, Prince Edward Island, in 1997. 


Numbers of Scoters 


Date Time Period (Hrs) Total Surf Black WW Unid. No./hr 

17 October 1045-1100 15 - ~ - 15 60 
1115-1103 0 — - ~ - 0 
1145-1200 9 - _ - 9 36 
1215-1230 10 - ~ — 10 40 
1245-1300 3 - - - 5 12 

Average (1.25 hrs) 29.6 
20 October 1000-1015 0 - - = - 4 
1030-1045 10 9 ~ - 1 40 
1115-1130 0 - - — - 0 
: 1145-1200 4 _ - 4 ~ 16 
4 1215-1230 5 - _ = 20 
1245-1300 - 47 20 _ Det 188 

Average (1.5 hrs) 44.6 
21 October 0915-0930 34 _ _ 34 136 
0930-0945 4 _ - _ 4 16 
1000-1015 0 - - - - 0 
1015-1030 0 _ = - 0 
1045-1100 0 - _ - 0 
1100-1115 0 - - - 0 
1115-1130 0 - ~ - 0 
1130-1145 0 — — - 0 
1220-1230 l _ _ | 4 
1230-1245 ] _ - l 4 
1245-1300 1 - - 1 4 

Average (2.45 hrs) 14.9 
22 October 0900-1000 20 - - 20 20 
1000-1100 4 _ _ 4 4 
1100-1200 3 - — 3 32 35 
1200-1300 e — _ _ 2 2 

Average 0900-1300 (4 hrs) 61 - - 0.75 14.5 1535 
23 October 0900-1000 0 — — _ 0 
1000-1100 4 — 1 3 16 
1100-1200 8 _ = 4 4 32 
1200-1300 3 - - - 3 i 
Average 0900-1300 (4 hrs) 15 - - 5 10 15 


continued 


444 


TABLE 6. Concluded 


THE CANADIAN FIELD-NATURALIST 


Date Time Period (Hrs) Total Surf 
28 October 0915-1015 24 = 
1015-1115 15 _ 
1115-1215 D4} = 
1215-1300 21 - 
Average 0915-1300 (3.75 hrs) 87 - 
29 October 0900-1000 8 = 
1000-1100 0 ~ 
1100-1200 0 - 
1200-1300 14 ~ 
Average 0900-1300 (4 hrs) 22 - 
30 October 1100-1200 2 — 
Average 1100-1200 (1 hr) 2 - 
31 October 0930-1030 3 - 
1030-1130 4 - 
1130-1230 0 -- 
1230-1300 0 - 
Average 0930-1300 (4 hrs) 1.75 - 
3 November 0915-1015 4 — 
1015-1115 0 — 
1115-1215 0 ~ 
1215-1300 0 - 
Average 0915-1300 (3.75 hrs) 1 ~ 
4 November 0915-1015 3 - 
1015-1115 1 — 
1115-1215 1 _ 
1215-1300 0 - 
Average 0915-1300 (3.75 hrs) 1 - 
6 November 0930-1030 0 - 
1030-1130 0 _ 
1130-1215 0 - 
Average 0930-1215 (2.75 hrs) 0 _ 
7 November 1015-1115 > — 
1115-1215 10 — 
1215-1245 60 — 
Average 1015-1245 (2.75 hrs) 26 - 
12 November 0900-1000 42 _ 
Average 0900-1000 (1 hr) 42 - 
19 November 0830-0900 5 - 
Average 0830-0930 (0.5 hr) 5 ~ 
20 November 0815-0845 1 1 
Average 085-0915 (0.75 hr) 1 1 
21 November 0815-0845 5 - 
Average 0815-0915 (0.75 hr) 5 - 
26 November 0830-0845 0 - 
Average 0830-0930 (1 hr) 0 - 
OVERALL TOTALS 41.95 hours 475 30 


Acknowledgments 

Students from Holland College in Prince Edward 
Island conducted many surveys from the Prince 
Edward Island side of Confederation Bridge and we 
are particularly grateful to Ben Hotelling for training 
and supervisory assistance. Ruth Miller, Stan Bunker, 
Nelson Poirier, Vivian Beale, Ron Arsenault, Harold 
Popma, John Reardon, Melanie Kotef, Shannon 
Atkinson and Jane Roma all assisted at Cape 
Jourimain. We sincerely thank Richard D. Elliot and 


Volts 
Numbers of Scoters 
Black WW Unid. No./hr 
1 = D3 24 
~ — 1S 15 
- — 27 20 
_ = DA 21 
1 — 86 PSPS 
li 3 4 8 
— — -- 0 
— — — 0 
i! — 13 14 
Z 3 17 5.5 
— = 2 2, 
— - 2 2 
= DF it 3 
A = ~ 4 
- - — 0 
-- - — 0 
1 0.5 0.25 _ 1.75 
1 - 3 4 
- — - 0 
- - 0 
— — _ 0 
0.25 - 0.75 1 
_ — 3 3 
1 = - 1 
1 = = 1 
— - - 0 
0.5 - 0.75 1 
a es € OF 
- ~ _ 0 
= = _ 0 
- - _ 0 
- —- — 5 
= = 10 10 
= = 60 100 
_ 2 20 38.3 
~ 3 39 42 
- 3 39 42 
— 3 2 10 
- 3 Z 10 
= = = 2 
_~ - _ 2 
2 = 5 10 
= - 5 10 
= = _ 0 
= _ - 0 
10 55 380 11.3/hr 


John W. Chardine who reviewed earlier drafts of the 
manuscript and A. J. Erskine and Allan Keith who 
provided valuable comments which served to 
improve the contents and presentation of this paper. 
Julie Paquet drafted the map. 


Documents Cited 

Harries, H. 1996. Plant Cover and History of Cape 
Jourimain, New Brunswick — Part I. Oldfields and 
Forests. Unpublished Report to the Canadian Wildlife 
Service — Atlantic Region. 61 pages. 


2001 HICKLIN AND BUNKER-POPMA: MIGRATIONS OF SCOTERS 445 


Webster, J.C. 1930. The Forts of Chignecto. J.C. MacKinnon, C. M., R. W. Daury, and R. J. Hicks. 1991. 
Webster, Shediac, New Brunswick [includes a 1685 map Seabird and Seaduck Movement through the North- 
by de Meulles]. umberland Strait, 1990. Technical Report Number 130, 

Canadian Wildlife Service — Atlantic Region. 86 pages. 
P 7 Savard, J.-P. L., D. Bordage, and A. Reed. 1998. Surf 

Literature Cited Scoter (Melanitta perspicillata). In The Birds of North 

Bordage D. and J. L. Savard. 1995. Black Scoter (Mela- America, Number 363. Edited by A. Poole and F. Gill. 
nitta nigra). In The Birds of North America, Number The Birds of North America, Inc., Philadelphia, Phila- 
177. Edited by A. Poole and F. Gill. The Academy of delphia and The American Ornithologists’ Union, 
Natural Sciences, Philadelphia, and The American Orni- Washington, D.C. 
thologists’ Union, Washington, D.C. 

Clark, A. H. 1959. Three Centuries and The Island. Uni- —_ Received 18 February 2000 
versity of Toronto Press, Toronto, Ontario, Canada. Accepted 7 June 2001 


New Plant Records for Prince Edward Island 


KATE MACQuarRRIE! and HEIDI SCHAEFER? 


1Executive Director, Island Nature Trust, P.O. Box 265, Charlottetown, Prince Edward Island C1A 7K4 Canada 
2153 Spring Street, Summerside, Prince Edward Island C1N 3G2 Canada 


MacQuarrie, Kate, and Heidi Schaefer. 2001. New plant records for Prince Edward Island. Canadian Field-Naturalist 


115(3): 446-450. 


The first records of Apios americana (Groundnut), Polystichum braunii (Braun’s Holly Fern), Hieracium piloselloides 
(King Devil) and Impatiens glandulifera (Glandular Touch-me-not) from Prince Edward Island are reported. Large num- 
bers of A. americana were found in two populations on the Lennox Island Mi’Kmaq reserve, Malpeque Bay, Prince 
County while P. braunii was restricted to a few individuals along the bank of the Mill River, Bloomfield, Prince County. H. 
piloselloides was first identified in retired farmland at Greenwich, Kings County, and /. glandulifera was found along the 


Barbara Weit River at New Annan, Prince County. 


Key Words: Apios americana, Groundnut, Polystichum braunii, Braun’s Holly Fern, Hieracium piloselloides, King Devil, 
Impatiens glandulifera, Glandular Touch-me-not, Prince Edward Island, new records. 


As Canada’s smallest and most densely populated 
province, Prince Edward Island has few unexplored 
or undisturbed habitats. Since the arrival of European 
settlers in the early 1700s, land clearing for agricul- 
ture, shipbuilding, and habitation has consumed 
much of the Island’s original Acadian forest. It is 
estimated that by 1900, over 70% of the land had 
been cleared (P.E.I. Department of Agriculture and 
Forestry 1997) and much of the remnant forest had 
been previously burned or harvested in varying 
degrees. Today, roughly 50% of the province is 
wooded, but less than one fifth of this resembles the 
original Acadian upland forest of Fagus grandifolia 
(American Beech), Betula alleghaniensis (Yellow 
Birch), Acer saccharum (Sugar Maple), Acer rubrum 
(Red Maple), Tsuga canadensis (Eastern Hemlock), 
Picea rubens (Red Spruce), and Pinus strobus (White 
Pine) (Arsenault 1997*). 

Despite this history of land use, there is much to 
be discovered about the flora of Prince Edward 
Island. While The Plants of Prince Edward Island 
(Catling, Erskine and MacLaren 1985) and The Rare 
Vascular Plants of Prince Edward Island (Day and 
Catling 1991) remain the primary publications on the 
topic, many new locations for some of the rarest 
species have been found in the past two years 
(MacQuarrie, Schaefer and Schoenrank 1999; 
MacQuarrie et al. 2000; MacQuarrie, Schaefer, 
Schoenrank and Lewis 2000*). Creation and revision 
of a vascular plant tracking list for P.E.I. (Blaney, 
MacQuarrie and Curley 2000) and the 2000/01 
update of the provincial forest inventory are adding 
to our knowledge of this province’s flora. 

Recently, two apparently indigenous species previ- 
usly unknown from Prince Edward Island have been 
discovered: Apios americana (Groundnut), found on 
Lennox Island on 27 July 1999 and Polystichum brau- 


nii (Braun’s Holly Fern), found in Bloomfield on 12 
August 1999. Two additional exotic species have also 
been identified: Hieracium piloselloides (King Devil) 
found at Greenwich in June 1999 and Impatiens glan- 
dulifera (Glandular Touch-me-not) found in New 
Annan in October 1999. 


Apios americana (Groundnut) 

Located in the western end of Malpeque Bay, 
Prince County, Lennox Island is home to the Lennox 
Island First Nation of Mi’Kmaq peoples (Figure 1). 
Encompassing approximately 540 hectares, Lennox 
Island is among the largest of the province’s many 
offshore islands. Nearly two-thirds of the island is 
forested, primarily in disturbed hardwood character- 
ized by Acer rubrum (Red Maple), Betula papyrifera 
(White Birch), Populus tremuloides (Trembling 
Aspen) and P. grandidentata (Large-toothed Aspen). 
There are pockets of Picea mariana (Black Spruce) 
scattered throughout, as well as sandy beaches on the 
north shore and tidal salt marshes on the south. Non- 


FiGuRE 1. Locations of four new provincial floral records 
for Prince Edward Island. 


446 


2001 


A Locations of Apios americana 


FiGuRE 2. Locations of Apios americana on Lennox Island, 
Prince County, Prince Edward Island. 


forested land is a combination of inhabited areas and 
lands being used for commercial blueberry produc- 
tion. The topography is generally flat, with the high- 
est elevation only eight metres. Soils range from 
sand to sand phase till, with a handful of calcareous 
mudstone brecchia lenses (Prest 1973). 

Apios americana was first found in the northwest- 
ern end of Lennox Island, in a region known locally 
as “the Cove” (46°37'30"N, 63°52’30”W) (Figure 2). 
The first specimen was identified in an area where 
the disturbed hardwood forest blended into Picea 
glauca (White Spruce), P. mariana, and Abies bal- 
samea (Balsam Fir) which in turn bordered the 
coastal salt marsh. The plant was growing vine-like 
among the conifers, climbing to heights in excess of 
two metres. Although not in flower, the plant was 
easily identified by its trailing growth and pinnate, 
entire leaves with five lanceolate leaflets. Leaflets 
varied from 2 to 4 centimetres in width and 4 to 6.4 
cm in length on the specimens collected. The root 
was found to have the characteristic tubers from 
which the plant gets its common name, Groundnut. 

A second trip to Lennox Island on 9 August 1999 
found Apios americana fully in flower (Figure 3). 
During this second visit, individual plants were pho- 
tographed and collected, the extent of the population 
was roughly mapped and a second population was 
found near the southwestern tip of the island 
(43°36'30'N, 63°52'30"W) (Figure 2). 

The first population was found to extend into the 
upper salt marsh, growing in areas of dense and vig- 
orous Toxicodendron rydbergii (Poison Ivy). At the 
transition zone between marsh and woodland, Apios 
was growing up all available vegetation including 
Myrica pensylvanica (Bayberry), Nemopanthus 
mucronata (False Holly), Prunus pensylvanica (Pin 
Cherry), Viburnum nudum (Wild Raisin) and Rosa 
spp. (wild roses). The population extended along this 
coastal area for more than 750 metres, varying in 
width from a few to nearly 100 metres. Other plants 
of this area included the provincially rare Elymus 
trachycaulus (Slender Wheat Grass), Distichlis spi- 
cata (Seashore Salt Grass), and Teucrium canadense 
(American Germander). Voucher material for each 


MACQUARRIE AND SCHAEFER: NEW PLANT RECORDS FOR PRINCE EDWARD ISLAND 


447 


of these species was collected and remains on file 
with Island Nature Trust. 

The second population was growing in a roadside 
ditch in the southwestern region of the island, 
extending up the “Welcome to Lennox Island” sign. 
It covered a much smaller area than the aforemen- 
tioned population, most likely because regular road- 
side maintenance has prevented its spread. 

During the potato famine of 1845, Apios was con- 
sidered as a possible blight resistant substitute for 
Solanum tuberosum (Potato) (Reynolds et al 1990). It 
was at this time that Island geologist Abraham Gesner 
visited the islands of Richmond (now Ma!peque) Bay 
in search of native plants that could be cultivated as 
substitutes for the blight-stricken potato. In a letter to 
The Islander on 12 August 1846, Gesner wrote of a 
plant shown to him by the Mi’ Kmagq people: 

“{Saa-gaa-ban] was found on several of the Islands in 

Richmond Bay, but is most plentiful at the bases of the 

sand mounds of Fish Island. Its favourite site seems to be 

along the skirts of the sandhills that form the lagoons 
along the coast, where it is nourished by decomposed sea 
weed and shells. It occurs in the midst of matted grass 
and wild tares, and frequently occupies patches of sever- 
al square rods. The leaf of the saa- gaa-ban resembles the 
leaf of the cultivated potato. The stock is like a small 


FIGURE 3. Apios americana on Lennox Island, Prince 
Edward Island, 9 August 1999. Photo by J. Waddell, 
Island Nature Trust. 


448 


vine; the roots are situated two inches below the surface 

of the soil, and the bulbs of oval figures, are strung 

together like beads, being attached to each other by a 

strong ligament. They are of a blackish brown color, and 

also resemble potatoes in their general character, being 
dry, farinaceous and nutritive”. 

Gesner’s saa-gaa-ban has characteristics of Apios 
and occupies habitat similar to that in which it was 
found on Lennox Island, also in Malpeque Bay. 
Hinds (2000) reports Segabun as a Mi’ Kmaq name 
for Apios and this plant is well-known as a food of 
eastern North America’s early aboriginal peoples 
who both gathered it from the wild and transplanted 
it near campsites (Reynolds et al. 1990). It is possi- 
ble that this accounts for the species’ presence on 
Lennox Island. 

Apios is common in southwestern Nova Scotia 
and scattered elsewhere in that province (Zinck 
1998). In New Brunswick, it is scattered in southern 
areas, reaching as far north as Kouchibouguac 
National Park (Hinds 2000). Its range extends into 
southern Quebec and Ontario (Scoggan 1979) and 
the eastern half of the United States to the Gulf of 
Mexico (Reynolds et al. 1990). 

Voucher material and photographs of the newly- 
discovered Prince Edward Island population are on 
file with Island Nature Trust. A voucher specimen 
has been sent to the Department of Agriculture 
Herbarium in Ottawa. 


Polystichum braunii (Braun’s Holly Fern) 
Polystichum braunii is an easily recognized fern 
(Figure 4). Its stipe is conspicuously chaffy, covered 
in light brown to golden scales. The blade is leathery 
at both ends, twice pinnate and chaffy beneath and 
on the rachis. Each pinnule has spiny edges and a 
sharp lobe at its base, characteristic of the holly 
ferns. Sori are arranged in two ranks along the 
midrib and spores are present from June to 
September (Zinck 1998). In Canada, P. braunii is 
found from Newfoundland west to southern Ontario, 


ie aoe =. a = 
is " a 3 ‘ fe ‘ec = . 
en alae 8 be > we. sa OS & f 


FIGURE 4. Polystichum braunii at Mill River, Prince 
Edward Island, 20 September 1999. Photo by J. 
Waddell, Island Nature Trust. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


FiGureE 5. Location of Polystichum braunii along the Mill 
River, Bloomfield, Prince County, Prince Edward 
Island. 


while in the United States it ranges from Maine, 
south to Connecticut and west to Wisconsin. It has 
also been recorded in St. Pierre and Miquelon. 
Western populations are known in British Columbia, 
Yukon, Alaska and Idaho (Kartesz 1999). P. braunii 
is not considered common anywhere in its North 
American range. Kartesz (1999) cites Polystichum 
braunii as rare in Idaho, Minnesota, Wisconsin, 
Pennsylvania, Massachusetts and Ontario, and it is 
listed as endangered in Minnesota (Minnesota 
Department of Natural Resources 1999. www.dntr. 
state.mn.us/fish_and_wildlife/endangered_species/en 
vasend.ht) and threatened in Wisconsin (Wisconsin 
Department of Natural Resources 1999. www.dnr. 
state.wi.us/org/land/er/factsheets/plants/brafern. 
htm). 

In neighbouring Nova Scotia, Roland (1941) 
recorded P. braunii only in the richest hardwood 
areas which include lands along the Bay of Fundy, in 
the Cobequid Hills and in the hardwood areas of 
Inverness and Victoria counties of Cape Breton. 
Nichols (1918) found P. braunii characteristic of the 
deciduous climax forest, sandy flood plains and espe- 
cially in the ravines of northern Cape Breton. Zinck 
(1998) added seepy hillsides as a habitat type for this 
species. P. braunii is considered uncommon to rare in 
New Brunswick, where it is associated with calcare- 
ous soils (Hinds 2000). With the exception of soil 
type, the habitat in which the Prince Edward Island 
specimens were found is consistent with records from 
the mainland. 

Approximately six to eight specimens of P. brau- 
nii were found in a steeply sloped area of mature for- 
est along the Mill River in Bloomfield, Prince 
County (46°44'30"N, 64°11'30”"W) (Figure 5). The 
topography of the Mill River area is primarily gently 
rolling hills that rarely exceed 45 metres in height, 
however the river itself flows through a steep ravine 
en route to Cascumpec Bay on the Island’s north 
shore. Land use in the area is divided between forest 
(40%) and agriculture (60%), with forest dominating 
the upper watershed area and lands along the river 
itself. The soils are classed as clay and clay-phase till 


2001 


and the area in which the fern was found is the 
largest of a very few pockets of ablation moraine in 
Prince County (Prest 1973). 

Information based on Samuel Holland’s 1767 sur- 
vey of Prince Edward Island suggests that land in the 
Mill River area was classed as good farmland. By 
1833, this was among the most extensively cleared 
areas in the province (Clark 1959). Clark’s map of 
forest cover prior to land clearances delineates a 
hardwood forest association more typical of Central 
P.E.I. in this western area of the province. This 
pocket of hardwood forest was an. oddity among 
what Clark called the White Cedar association which 
dominated most of the western end of the Island. 
Maps of the Mill River area estimating forest cover 
circa 1900 (P.E.I. Department of Agriculture and 
Forestry 1997) show that while much of the land was 
cleared, pockets of forest remained in the general 
area where P. braunii was found. 

The ravine in which the fern was found includes 
trees typical of the Acadian Forest: Betula 
alleghaniensis, Fagus grandifolia, Acer saccharum 
and Tsuga canadensis. The size of these trees, the 
extreme slope, and the estimates of forest cover in this 
area a century ago, suggest that this land has never 
been cleared. This is consistent with the knowledge 
that P. braunii is sensitive to forest harvest practices 
(Wisconsin Department of Natural Resources 1999. 
www.dnr.state.wi.us/org/land/er/factsheets/plants/ 
brafern.htm). It is probable that P. braunii was once 
more widespread on P.E.I, and that the specimens 
found in 1999 remained simply because they were 
restricted to an area too difficult to harvest. 

In addition to the new Island record, Mill River is 
also the location for other provincially rare vascular 
plants. Clematis virginiana (Virgin’s Bower), Lapor- 
tea canadensis (Wood Nettle) and Solidago flexicaulis 
(Zigzag Goldenrod) were all found in the immediate 
area, adding to its interest and significance. Day and 
Catling (1991) report that Erskine’s specimen of L. 
canadensis had been lost; Zinck (1998) reports this 
species as absent from Prince Edward Island. Voucher 
material for L. canadensis, C. virginiana and P. brau- 
nii is on file with Island Nature Trust, and a voucher 
specimen for the latter species has been forwarded to 
the Department of Agriculture Herbarium in Ottawa. 


Impatiens glandulifera 
(Glandular Touch-me-not) and 
Hieracium piloselloides (King Devil) 

In addition to the two indigenous species, we have 
identified two exotic species that are absent from the 
Island’s floral literature. Hieracium piloselloides 
(King Devil) was found by the authors in retired farm- 
land at Greenwich, Prince Edward Island National 
Park, Kings County, in June 1998. Impatiens glan- 
dulifera (Touch-me-not) was found and collected in 
New Annan, Prince County, by Peter Stewart on 6 


MACQUARRIE AND SCHAEFER: NEW PLANT RECORDS FOR PRINCE EDWARD ISLAND 


449 


October 1999, and subsequently identified by Kate 
MacQuarrie. Voucher material is on file with Island 
Nature Trust. 


Discussion 

There are few current threats to either Apios amer- 
icana or Polystichum braunii. A. americana is not in 
a region of Lennox Island that is likely to be under 
pressure for commercial blueberry production, hous- 
ing, or other development. Preliminary contact with 
the Band Council suggests that there is an interest in 
conservation of this species on the reserve, and we 
look forward to working with band members in an 
effort to identify the plant’s provenance on Lennox 
Island. 

The reasonably inaccessible location of P. braunii 
suggests that it will not be disturbed by logging or 
development, although recreational use of the area 
may be a concern. Local anglers have worn a path 
through the Laportea canadensis, not far from the 
location of P. braunii. As community interest in 
watershed enhancement increases across P.E.I., it is 
possible that stream clearing activities designed to 
improve fish habitat could result in elimination of 
the very few fern individuals found. We have 
advised the provincial watershed enhancement coor- 
dinator of the presence of the fern and rare vascular 
plants along this system. 


Acknowledgments 

The authors thank former Lennox Island Chief 
Charlie Sark and Brian McHattie for the opportunity 
to investigate the flora of Lennox Island; Jackie 
Waddell of Island Nature Trust for photographing 
the finds; the late Hal Hinds of the University of 
New Brunswick for verification of P. braunii and for 
all his support of local botanical investigations; Ben 
Hoteling for forwarding Peter Stewart’s specimen of 
I. glandulifera; and David Erskine for inspiration 
and encouragement. 


Documents Cited 

Arsenault, Mark. 1997. Protected woodlands of Prince 
Edward Island and the forest communities they repre- 
sent. Unpublished report prepared for Island Nature 
Trust, P.O. Box 265, Charlottetown, Prince Edward 
Island C1A 7K4. 12 pages. 


MacQuarrie, K. E., H. L. Schaefer, K. A. Schoenrank, 
and W.K. Lewis. 2000. Results of floral inventories 
from 51 Acadian-type forest stands on Prince Edward 
Island. Unpublished data collected for Island Nature 
Trust, P.O. Box 265, Charlottetown, Prince Edward 
Island C1A 7K4. 


Literature Cited 

Blaney, S., K. MacQuarrie and R. Curley. 2000. Prince 
Edward Island Vascular Plant Tracking List. Atlantic 
Canada Conservation Data Centre, Mount Allison 
University, Sackville, Nova Scotia. 


450 


Catling, P., D. Erskine and B. MacLaren. 1985. The 
plants of Prince Edward Island with new records, 
nomenclatural changes and corrections and deletions. 
Agriculture Canada Research Branch Publication Num- 
ber 1798. 272 pages. 

Clark, A. 1959. Three Centuries and the Island. Uni- 
versity of Toronto Press, Toronto, Ontario. 287 pages. 
Day, R., and P. Catling. 1991. The rare vascular plants of 
Prince Edward Island. Syllogeus of the Canadian Muse- 

um of Nature (67). 65 pages. 

Gesner, A. 1846. Letter to the editor of The Islander, 12 
August 1846. P.E.I. Government Archives Microfilm, 
The Islander 1846-1849, Volume 3, page 3. 

Hinds, H. 2000. The flora of New Brunswick, second edi- 
tion. University of New Brunswick, Fredericton. 695 
pages. 

Kartesz, J. and C. Meacham. 1999. Synthesis of the 
North American Flora. North Carolina Botanical Gar- 
den. Chapel Hill, North Carolina. 

MacQuarrie, K., H. Schaefer, and K. Schoenrank. 2001. 
A floral inventory of the Central and Schooner Pond 
Areas, Greenwich, Prince Edward Island National Park. 
Parks Canada technical report in ecosystem science. 
Number 031. 40 pages plus appendix. 

MacQuarrie, K., H. Schaefer and K. Schoenrank. 1999. 
A floral inventory of the Western Area, Greenwich, 
Prince Edward Island National Park. Parks Canada tech- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


nical report in ecosystem science. Number 021. 42 pages 
plus appendices. . 

Natural Resources Division, P.E.I. Department of Agri- 
culture and Forestry. 1997. Map of Prince Edward 
Island forest area circa 1900. 

Nichols, G. 1918. The vegetation of Northern Cape Breton 
Island Nova Scotia. Transactions of the Connecticut 
Academy of Arts and Sciences 22: 249-467. 

Prest, V. 1973. Map of surficial deposits of Prince Edward 
Island. Geological Survey of Canada Map Number 
1366A. 

Reynolds, B., W. Blackmon, E. Wickremesinhe, M. 
Wells and R. Constantin. 1990. Domestication of 
Apios americana. Pages 436—442 in Advances in new 
crops. Edited by J. Janick and J. E. Simon. Timber Press, 
Portland, Oregon. pages 436-442. 

Roland, A. 1941. The Ferns of Nova Scotia. Nova Scotia 
Agricultural College, Truro, Nova Scotia. 

Scoggan, H. 1979. The flora of Canada. National Muse- 
um of Canada Publications in Botany Numbers 7(1) to 
7(4). 1711 pages. 

Zinck, Marian. 1998. Roland’s Flora of Nova Scotia. 
Nimbus Publishing and the Nova Scotia Museum: 
Halifax, Nova Scotia. 2 volumes. 1297 pages. 


Received 7 April 2000 
Accepted 10 December 2001 


Status of the Deltoid Balsamroot, Balsamorhiza deltoidea 
(Asteraceae) in Canada? 


GEORGE W. DouGLAS! and MICHAEL RYAN2 


‘Conservation Data Centre, Ministry of Environment, Lands and Parks, Resource Inventory Branch, P.O. Box 9344, 
Station Provincial Government, Victoria, British Columbia V8T 9M1 Canada 
*Present address: 801 Frayne Road, RR # 1, Mill Bay, British Columbia VOR 2P0 Canada. 


Douglas, George W., and Michael Ryan. 2001. Status of the Deltoid Balsamroot. Balsamorhiza deltoidea (Asteraceae) in 
Canada. Canadian Field-Naturalist.115(3): 451-454. 


In Canada, Deltoid Balsamroot, Balsamorhiza deltoidea has been collected only on southeastern Vancouver Island. 
Populations have been confirmed at six sites in recent years while the species has probably been extirpated at eight other 
previous collection sites. The confirmed populations collectively represent the northern range limit of B. deltoidea. 
Population sizes range from five to 1600 plants. Threats to the existing populations include habitat destruction and the 
increasing invasion of introduced species. In addition, sites in ecological reserves or municipal and regional parks are 
sometimes at risk due to management activites. Considering these threats to the habitat of B. deltoidea, we recommend a 


status of threatened. 


Key Words: Deltoid Balsamroot, Balsamorhiza deltoidea, British Columbia, threatened, distribution, population size. 


The Deltoid Balsamroot, Balsamorhiza deltoidea 
Nuttall is a member of a genus of about 12 species 
found in western North America (Cronquist 1955). 
Only two of these species, the latter and Arrowleaf 
Balsamroot (Balsamorhiza sagittata), occur in Brit- 
ish Columbia and Canada (Scoggan 1979; Douglas 
et al. 1998b). Taxonomy and nomenclature follows 
Douglas et al. (1994, 1998b, 1998c, 1999a, 1999b, 
2000, 2001). 

Balsamorhiza deltoidea is a leafy, ascending plant, 
ranging from 20-100 cm tall with long—stalked, trian- 
gular, stiff-hairy and glandular leaves with the blades 
10-50 cm long and 10-20 cm wide (Figure 1, 
Douglas et al. 1998b). The few to several flower 
heads are borne on long stalks. The involucral bracts 
are lanceolate to oblong-lanceolate and the 13 to 21 
tay flowers are bright yellow. The numerous disk 
flowers are also yellow. The fruits (achenes) are 7-8 
mm long, smooth and lack a pappus. 


Distribution 

Balsamorhiza deltoidea occurs on the west coast of 
North America from southwestern British Columbia 
along the western slopes of the Cascade Mountains in 
Washington and Oregon to the western slopes of the 
Sierra Nevada in California. In Canada, it is restricted 


sees A 
*This paper is based primarily on a COSEWIC status 


report by the authors. It has been revised to include more 
recent information. The species was designated threatened 
by COSEWIC in April 1996. COSEWIC Reports are avail- 
able from the COSEWIC Secretariat, c/o Canadian Wildlife 
Service, Environment Canada, Ottawa, Ontario K1A OH3 
Canada. 


to southeastern Vancouver Island (Figure 2, Douglas 
et al. 1998a). 


Habitat 

Balsamorhiza deltoidea populations are restricted 
mainly to very dry, exposed or partially shaded sites 
where soils are shallow. Trees such as Garry Oak 
(Quercus garryana) are frequent. Other associated 
species include: Scotch Broom (Cytisus scoparius), 
Snowberry (Symphoricarpos albus), Broad-leaved 
Stonecrop (Sedum spathifolium), Nooding Onion 
(Allium cernuum), Menzies’ Larkspur (Delphinium 
menziesit), Sweet Vernalgrass (Anthoxanthum odor- 
atum), and several species of Brome (Bromus). Two 
of the populations are on quite different sites from 
the latter. The largest population, at Campbell River, 
dominates an open meadow on marine sediments 
adjacent to the ocean (Figure 3) while another popu- 
lation, on Mt. Tzuhalem, occurs in a relatively moist 
ravine with Symphoricarpos albus and Viola prae- 
morsa Ssp. praemorsa. 


General Biology 

Balsamorhiza deltoidea emerges in the spring 
from the perennial taproot and flowers by early sum- 
mer. By mid-summer, when drought conditions are 
prevalent, seed set has occurred and the leaves with- 
er and turn brown. Although seeds are usually rela- 
tively easy to germinate, the cultivation of young 
plants in the Victoria area appears to be difficult 
since they seem to be very sensitive to soil moisture 
conditions and predators during the growing season. 
Compared to the large number of flowers comprising 
the floral head, viable seed production is low. No 
information is available regarding the population 


45] 


452 


THE CANADIAN FIELD-NATURALIST 


TABLE |. Location of Balsamorhiza deltoidea sites in British Columbia. 


Collection Last 
Site Observation 
Tolmie Farm (Victoria) 1891 
Lost Lake (Victoria) 1916 
Lake Hill (Victoria) 1926 
Royal Oak (Victoria) 1935 
Campbell River, south of WS 2) 
Witty’s Lagoon (Victoria) 1965 
Fort Rodd Hill (Victoria) 1966 
Portage Inlet (Victoria) 1976 
Francis-King Park, south of (Victoria) 1993 
Mill Hill (Victoria) 1998 
Tyee Spit (Campbell River) 1998 
Mount Tzuhalem (Duncan) 1999 
Thetis Lake (Victoria) 1999 
Beacon Hill (Victoria) 1999 


dynamics of this species including the extent to 
which seed remains viable in the soil, the frequency 
with which recruitment occurs from established 
seedlings, and the average life-span of mature plants. 


Population Size and Trends 

Of the 14 Canadian sites from which Balsamor- 
hiza deltoidea has been collected, only six sites have 
been confirmed recently (Table 1). The status of the 
species at the additional sites remains uncertain but it 
is believed that all of the populations and some of 
the sites have been extirpated. Little is known 
regarding the specific size of past populations 
although it is likely some of these were larger at one 
time. The largest population (1600 plants) is located 
on an Indian Reserve near Campbell River and has 
remained stable between 1992 and 1999. All other 
populations number less than 250 plants in total. 
Three of the sites (Beacon Hill, Mt. Tzuhalem and 
Tyee Spit), visited on a yearly basis between 1992 
and 1999, have remained stable, or, in the case of 
Mt. Tzuhalem, have increased slightly in numbers 
since 1992. 


Limiting Factors 

The most direct and immediate threat to Balsa- 
morhiza deltoidea is habitat destruction. This is of 
particular concern on the rock outcrops often associ- 
ated with Quercus garryana stands that are limited 
to the southeastern side of Vancouver Island. This 
type of vegetation is believed to have been much 
more common before European settlement. Its 
destruction has continued to the present resulting in 
the elimination of many sites occurring outside parks 
or ecological reserves. At this time, pressures to 
develop the remaining unprotected Quercus gar- 
ryana stands for the expansion of the urban infras- 
tructure of Victoria, and other population centres on 
Vancouver Island, are intense. The population at the 


Volts 

Population 
Observer - (number/area) 
Newcombe Extirpated 
Newcombe Extirpated 
Walker Extirpated 
Goddard Extirpated 
Beamish Extirpated 
Carl Extirpated 
Ashlee Extirpated 
Brayshaw Extirpated 
Ryan 70/20 m2 
Douglas 50/100 m2 
Douglas 1600+/1260 m2 
Douglas 55/40 m2 
Douglas 72/60 m2 
Douglas Sy lem 


Campbell River Indian Reserve may currently be 
under even greater threat since new development 
plans are under consideration for the site. 

The suppression of fire within the past century 
may have also contributed to the demise of 
Balsamorhiza deltoidea populations. Many of the 
sites in which this species has been collected may 
have been maintained in the past as a result of peri- 
odic fires, both natural and intentionally set, aborigi- 
nal peoples probably set fire to these stands to main- 
tain them as an important habitat for wildlife and for 
the continued harvesting of Camas (Camassia spp.), 
a member of the Liliaceae (Roemer 1972; Turner 
and Bell 1971). Since that time, these sites have 
experienced little disturbance, resulting in the inva- 
sion of many other species, especially introductions. 

The introduction of exotic species has resulted in 
substantial changes, not only to the grass-dominated 
meadows associated with Quercus garryana, but 
also to the dry, rocky sites north and west of 
Victoria where Balsamorhiza deltoidea has been 
collected in the past. One of the most devastating 
species, over the past 100 years, is Cytisus scopar- 
ius which has become a dominant shrub on dry, 
exposed sites throughout much of eastern 
Vancouver Island and the Gulf Islands. Much of the 
vegetation is also now dominated by introduced 
grasses. These species include Early hairgrass (Aira 
praecox), Anthoxanthum odoratum, Hedgehog 
Dogtail (Cynosurus echinatus) and Orchard Grass 
(Dactylis glomerata). 


Special Significance of the Species 
Balsamorhiza deltoidea is a member of a relative- 
ly large group of species with a Western Cordilleran 
range that have their northern limits in western 
Canada. The significance of these peripheral popula- 
tions, especially with respect to their genetic charac- 
teristics, has yet to be studied adequately. This and a 


2001 


Figure 1. iHustration of Balsamorhiza deltoidea. (Line 
drawing by Elizabeth J. Stephen in Douglas et al. 
(1998a, 1998b). 


number of other species with similar ranges may 
prove to be a fruitful subjects for genetic research. 


Protection 

Balsamorhiza deltoidea has been globally ranked 
by The Nature Conservancy of the United States as 
“G5,” or “common to very common with an exis- 
tence that has been demonstrated to be secure and 
essentially ineradicable under present conditions.” 

The British Columbia Conservation Data Centre 
has ranked this species as S2 and placed it on the 
Ministry of Environment, Lands and Parks Red list 
(Douglas et al 1998a). The S2 rank indicates that the 
plant is “imperiled because of rarity (typically 6-20 
extant occurrences or few remaining individuals) or 
because of some factor(s) making it vulnerable to 
extirpation or extinction”. Balsamorhiza deltoidea 
has a national rank in Canada of N2. 

There is no specific legislation for the protection of 
rare and endangered vascular plants in British 
Columbia. Some populations of Balsamorhiza del- 
toidea are protected to a certain extent by their loca- 
tion on public property. Of all the B. deltoidea sites 
known in British Columbia, those located on Mount 
Tzuhalem receive the greatest degree of protection 


DOUGLAS AND RYAN: STATUS OF DELTOID BALSAMROOT 


453 


Us 
VANCOUVER 
ISLAND 


124. 423°w 
/ 


/ 
/ 
/ 
/ 
j 


VICTORIA 


FIGURE 2. Distribution of Balsamorhiza deltoidea in 
British Columbia (0 — extirpated sites, * — recently 
confirmed sites). 


because of their location within an ecological reserve. 
The Mount Tzuhalem Ecological Reserve encompass- 
es 18 ha of Quercus garryana woodland, spring-flow- 
ering meadows, and rock outcrops which have been 
preserved to represent an example of Q. garryana 
woodlands and associated spring-flowering herbs. 
Unfortunately, Cytisus scoparius has become a domi- 
nant species at this site and threatens many herba- 
ceous species, including Balsamorhiza deltoidea 

A number of extant Balsamorhiza deltoidea sites 
are in small regional parks in the Greater Victoria 
area. These include populations at Beacon Hill, Mull 
Hill and Thetis Lake. These parks receive little 
active management, at least with respect to their rare 


454 


FiGureE 3. A dense population of Balsamorhiza deltoidea 
on the Campbell River Indian Reserve. (Photo by 
Sylvia M. Douglas). 


plants. Park enhancement projects, road and trail 
developments and heavy recreational use by humans 
often result in the destruction of the native vegeta- 
tion and rare plant species. The largest population 
(1600 plants), located on an Indian Reserve at 
Campbell River, is also the most seriously threatened 
population. The loss of Balsamorhiza deltoidea at 
this site seriously jeopardizes the future of this 
species in Canada considering that the number of 
individuals comprising the remaining populations 
number less than 250 plants in total. 


Evaluation of Status 

Balsamorhiza deltoidea is considered, by the 
British Columbia Conservation Data Centre (Douglas 
et al. 1998a) to be endangered in Canada and is 
known only from six extant colonies restricted to 
southeastern Vancouver Island. Except for the large 
population (1600 plants) at Campbell River, the 
remaining populations range from five to 72 individu- 
als and may be in danger of extirpitation. The progno- 
sis for this species is not good considering the threats 
posed by potential developments and by aggressive 
competitive species such as Cytisus scoparius which 
dominate many suitable habitats and directly threaten 
some colonies. Therefore, even if all colonies were 
protected from human interference, many colonies 
may eventually disappear as a result of aggressive 
introduced species. Likewise, much of the Quercus 
garryana vegetation in which Balsamorhiza deltoidea 
is usually found has been extensively altered or 
destroyed, thus limiting the potential of this species to 
become established at new sites. 


Acknowledgments 

We thank Carmen Cadrin for information 
acquired at one of the sites and Betty Brooks, Marta 
Donovan, Sylvia Douglas, Tracy Fleming, Marie 
Fontaine, Sharon Hartwell and Jenifer Penny for aid- 


THE CANADIAN FIELD-NATURALIST 


Volishis 


ing the senior author at three other sites. Gail F. 
Harcombe prepared the map. Funds for this project 
were provided jointly by COSEWIC and the British 
Columbia Conservation Data Centre. 


Literature Cited 

Cronquist, A. 1955. Vascular plants of the Pacific North- 
west. Part 5: Compositae. University of Washington 
Press, Seattle. 343 pages. 

Douglas, G.W., D. Meidinger, and J. Pojar. 1999a. Illus- 
trated flora of British Columbia. Volume 3. Dicotyledons 
(Diapensiaceae through Onagraceae). Ministry of 
Environment, Lands and Parks, British Columbia Ministry 
of Forests, Victoria, British Columbia. 436 pages. 

Douglas, G.W., D. Meidinger, and J. Pojar. 1999b. II- 
lustrated flora of British Columbia. Volume 4. Dicotyle- 
dons (Orobanchaceae through Rubiaceae). Ministry of 
Environment, Lands and Parks, British Columbia Min- 
istry of Forests, Victoria, British Columbia. 427 pages. 

Douglas, G.W., D. Meidinger, and J. Pojar. 2000. 
Illustrated flora of British Columbia. Volume 5. Dicotyle- 
dons (Salicaceae through Zygophyllaceae). Ministry of 
Environment, Lands and Parks, British Columbia Min- 
istry of Forests, Victoria, British Columbia. 396 pages. 

Douglas, G.W., D. Meidinger, and J. Pojar. 2001. 
Illustrated flora of British Columbia. Volume 6. Mono- 
cotyledons (Acoraceae through Najadaceae). Ministry of 
Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 361 
pages. 

Douglas, G.W., G.B. Straley, and D. Meidinger. 1994. 
The vascular plants of British Columbia. Part 4 — Mono- 
cotyledons. Special Report Series 4. British Columbia 
Ministry of Forests, Victoria, British Columbia. 257 
pages. 

Douglas, G. W., G. B. Straley, and D. V. Meidinger. 
1998a. Rare native vascular plants of British Columbia. 
British Columbia Conservation Data Centre, Ministry of 
Environment, Lands and Parks, Victoria, British Colum- 
bia. 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). Ministry of Environment, Lands 
and Parks, British Columbia Ministry of Forests, Vic- 
toria, 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 Cucurbitaceae). 
Ministry of Environment, Lands and Parks, British 
Columbia Ministry of Forests, Victoria, British Columbia. 
436 pages. 

Roemer, H.L. 1972. Forest vegetation and environments 
of the Saanich Peninsula, Vancouver Island. Ph.D. the- 
sis. University of Victoria, Victoria. 405 pages. 

Scoggan, H. J. 1979. The flora of Canada. Part 4 — 
Dicotyledoneae (Loasaceae to Compositae). National 
Museum of Natural Sciences Publications in Botany 
Number 7. 

Turner, N. and M. A.M. Bell. 1971. The ethnobotany of 
the coast Salish Indians of Vancouver Island. Economic 
Botany 25: 63-104. 


Received 14 March 2000 
Accepted 12 July 2001 


Status of Scouler’s Corydalis, Corydalis scouleri (Fumariaceae) 
in Canaday 


GEORGE W. DOUGLAS and JUDY A. JAMISON! 


Conservation Data Centre, British Columbia Ministry of Environment, Lands and Parks, Victoria, British Columbia V8V 
1X4 Canada 
'!Present address: Copy Editor, Nature Biotechnology, NewYork, New York 10010. 


Douglas, George W., and Judy A. Jamison. 2001. Status of Scouler’s Corydalis, Corydalis scouleri (Fumariaceae) in 
Canada. Canadian Field-Naturalist 115(3): 455-459. 


In Canada, Corydalis scouleri is restricted to the Klanawa River/Nitinat River watersheds on southwestern Vancouver 
Island. There are 20 known populations of Corydalis scouleri in British Columbia, 19 of which were confirmed in the 
1997/1998 field season. One site in the Klanawa River valley, reported in 1992 was not visited. Historical growth or 
decline in size of populations is unknown. Existing populations represent the northern range limits of C. scouleri. British 
Columbia populations of this species occur in an area that is subject to human disturbance in the form of extensive logging 


operations, as well as natural disturbance in the form of erosional damage from flooding. 


Key Words: Scouler’s Corydalis, Corydalis scouleri, British Columbia, threatened, distribution, population sizes. 


Scouler’s Corydalis, Corydalis scouleri Hook., is 
a member of a cosmopolitan genus of about 100 
species, mostly in the Northern Hemisphere 
(Hitchcock et al. 1969). Ownbey (1947), in his 
monograph of the genus Corydalis, notes that annual 
species predominate in North America. Scoggan 
(1979) reports five species indigenous to Canada, of 
which three are annuals or biennials. Four of these 
five species occur in British Columbia: Corydalis 
aurea and C. sempervirens are annuals or biennials, 
while C. pauciflora and C. scouleri are perennials 
(Douglas et al. 1999a). 

Corydalis scouleri is a tall (60-120 cm), rhi- 
zomatous perennial with hollow, somewhat 
branched stems (Figure 1). Three blue-green, glau- 
cous, tri- to quadripinnate leaves project near or 
above the middle of the stem, the lowest often 
20-30 cm long (Figure 1). There is a delicate termi- 
nal raceme of 15-20 rosy-pink zygomorphic flow- 
ers These flowers are 2—3 cm long and have two 
lateral outer petals, one spurred or hooded, and two 
inner, dorsi-ventrally placed petals opposite the 
quickly deciduous sepals. The bilobed stigma and 
six stamens, fused in two groups alternating with 
the petals, are sheltered by a second hood formed 
by the inner petals. The bicarpellate capsule sepa- 
rates elastically when jarred, scattering the seeds a 
considerable distance. For each plant, the large dis- 
sected leaves form a delicate blue-green canopy, 


{This is a Status Report of a species restricted in Canada to 
British Columbia. This report has been submitted to the 
National Committee on the Status of Endangered Wildlife 
in Canada (COSEWIC), but a status designation has not yet 
been made. 


which intermingles in dense stands with other 
canopies to form a raised carpet of lush foliage 
about 1 m above the forest floor. The genus name is 
Greek for “crested lark”, possibly referring to the 
shape of the flower. 


Distribution 

Corydalis scouleri is limited in distribution to the 
region west of the Cascades (mostly coastal) from 
northwestern Oregon (near Tillamook) northward 
through Washington’s Olympic Peninsula to south- 
western Vancouver Island (Ownbey 1947; Hitchcock 
et al. 1969; Douglas et al. 1999a). In Canada, known 
sites are limited to the Klanawa River/Nitinat River 
watersheds (Figure 2; Douglas et al. 1999a). There 
has also been a recent report from the upper headwa- 
ters of the Cowichan River, about 0.5 km east of the 
Nitinat River watershed, but this has yet to be con- 
firmed. Another report from the Saanich Peninsula 
(Szczawinsky and Harrison 1973) has been refuted 
by Pavlick (1989), who reidentified the specimen to 
another species. 


Habitat 

Lush stands of C. scouleri are invariably found in 
wet, cool habitats associated with watercourses — 
from moderately large rivers to small tributary 
streams. Sites range in elevation from sea level to 
close to 1000 m; in British Columbia the highest ele- 
vation is about 200 m, the lowest about 5 m. Fine 
floodplain silts as well as coarser floodplain materi- 
als provide ideal habitat for C. scouleri. Typical 
overstory trees include old-growth Bigleaf Maple 
(Acer macrophyllum) and Sitka Spruce (Picea 
sitchensis), as well as Red Alder (Alnus rubra), 
Western Hemlock (Tsuga heterophylla) and Western 


455 


456 


Redcedar (Thuja plicata). Major associates in the 
understory include Western Swordfern (Polystichum 
munitum), Devil’s Club (Oplopanax horridus), Red 
Elderberry (Sambucus racemosa), Palmate Coltsfoot 
(Petasites frigidus var. palmatus), Stink Currant 
(Ribes bracteosum) and Salmonberry (Rubus 
spectabilis). The climate is cool and mesothermal, 
and these plant communities are commonly found in 
early seral forests in a nitrogen-rich moder and mull 
humus ranging from merely moist to very wet. In 
regions where Corydalis scouleri is locally abun- 
dant, such as lower elevations of Washington State’s 
Mt. Rainier National Park (Brockman 1947) and the 
Nitinat River watershed, high annual precipitation is 
also characteristic. 


General Biology 

Corydalis scouleri is a perennial herb producing 
annual stems apically from thick rhizomes. Single 
clones have been identified by the authors with 
numerous annual stems spreading for at least tens of 
square meters. Only a single leaf is produced annu- 
ally until the plants reach flowering age, which 
apparently only occurs after four or more years. 
Flowering in C. scouleri takes place in late spring to 
early summer, usually May and June in the Nitinai 
River watershed. Both anthers and stigma are 
enclosed by the determinately shaped and arranged 
petals, and flowers of all species studied by Ownbey 
(1947) show evidence of germination of the pollen 
that is clustered around the stigma. Therefore, self- 
fertilization in C. scouleri is a distinct possibility. 
Although Liden (1986) reports that most of the 
species in the tribe Corydaleae (to which Corydalis 
belongs) are strongly self-sterile; no studies are 
known to have investigated the status of C. scouleri 
in this regard. The species reproduces very well 
asexually by underground rhizomes and is capable 
of sexual reproduction by seed. However, it is 
known that sometimes only the terminal flower of 
the raceme develops (Hitchcock et al. 1969), and 
this would severely compromise total seed produc- 
tion. In transplanting rhizomes to suitable habitats, 
material from different clones should be chosen; 
planting sexually produced seed would ensure a 
multiclone population that, consequently, could 
itself reproduce sexually. 

Corydalis scouleri is a perennial, rhizomatous 
herb that, in addition to reproducing asexually to 
form large clones, has the potential for considerable 
sexual reproduction: each flower in the raceme of 
15-20 flowers has two multiseeded carpels. 
Although C. scouleri’s would seem to be fairly com- 
mon, field surveys of potential sites in seemingly 
ideal habitats have yielded no occurrences outside 
the immediate Nitinat River/Klanawa River region 
on Vancouver Island. That a number of pollinators 
visit C. scouleri indicates the likelihood of cross-fer- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


tilization; however, since few flowers develop, this 
may well not be an effective means of reproduction. 
Pollination experiments indicated that Corydalis 
ambigua, a spring ephemeral species, is self-incom- 
patible; moreover, seed production was increased by 
a few bee visits of long duration (over 60 seconds), 
whereas it seemed unaffected by many short visits 
(less than 60 seconds). Furthermore, the observation 
that plants with larger inflorescences were visited 
more often argues for natural selection for a larger 
inflorescence (Onara and Higashi 1994). Ideal habi- 
tat for C. scouleri appears to be perennially moist 
streamside or riverside floodplain soils that are often 
very fine and silty, although sometimes coarser. 
Periodic flooding likely facilitates seed or rhizome 
dispersal. 


Population Size and Trends 

There are 20 known populations of Corydalis 
scouleri in the Nitinat River/Klanawa River water- 
sheds, 19 of which were confirmed in the 1997/1998 
field season (Table 1). Population numbers range 
from a single plant to several thousand stems stretch- 
ing over a 0.5 ha area. Several extremely large popu- 
lations were documented, always in perennially moist 
riverbank silt. However, no new sites were found far- 
ther north along nearby watercourses that offer similar 
habitats. Pavlick (1989) searched along the Gordon 
and San Juan drainages without locating additional 
populations of Corydalis scouleri. Historical growth 
or decline in size of populations is unknown. 


Limiting Factors 

Extensive logging operations in the Nitinat and 
Klanawa River Valleys pose the greatest threat to 
extant populations of C. scouleri. A formidable net- 
work of logging roads keeps the region open to 
heavy equipment and tourist traffic alike. Roads 
require bridges across watercourses, which in turn 
threatens C. scouleri populations along the water- 
course—not only when the bridge is constructed but 
also when tourists gain access to the riverbank to 
hunt, fish, or hike. Catastrophic flooding could also 
pose a threat to C. scouleri populations, although as 
mentoned earlier, the plants could also achieve prop- 
agation by more moderate flood dispersal. There is a 
dearth of research evidence suggesting low genetic 
diversity as a threat to the survival of-extant C. 
scouleri populations; however, as suggested in this 
report, extensive rhizomatous growth accompanied 
by ineffectual sexual reproduction could be a limit- 
ing factor in the species’ long-term survival. Lack of 
genetic heterogeneity in a species can lead to such 
threats as reduced resistance to disease. 


Special Significance of the Species 
Gardeners in both North America and Europe 
value highly several species of Corydalis for both 


2001 


Fiure 1. Illustration of Corydalis scouleri (Line drawing 
by Jane Lee Ling in Douglas et al. [1998a, 1999a)). 


flowers and foliage. Ownbey (1947), in his mono- 
graph on the genus noted that European gardeners 
were using Corydalis scouleri, C. aurea, and C. sem- 
pervirens. Cultivation of a rare species for its aes- 
thetic value is, of course, to be encouraged. In addi- 
tion, the alkaloidal properties of a large number of 
members of both the fumitory and the closely related 
poppy family are of great interest to plant tax- 
onomists, plant chemists, and agronomists. Each 
species has been found to contain a unique set of 
alkaloids, some of which are common to other 
species, but not in the same combinations. Agrono- 
mists consider these properties significant in that 
they probably render the plants toxic to livestock; a 
bitter taste likely makes them unpalatable in any 
event (Ownbey 1947). 


Protection 

There is no specific legislation for the protection 
of rare native vascular plant species in British 
Columbia. The British Columbia Conservation Data 
Centre has ranked this species as S2 and placed it on 
the Ministry of Environment, Lands and Parks Red 
list (Douglas et al. 1998a). The S2 rank is that of The 
Nature Conservancy, United States and indicates the 
species is “imperiled because of rarity (typically 
6—20 extant occurrences or few remaining individu- 


DOUGLAS AND JAMISON: STATUS OF SCOULER’S CORYDALIS 


457 


BRITISH 


COLUMBIA 


48°55'N 


124°51'N 126°30'N 
FIGURE 2. Distribution of Corydalis scouleri in British 
Columbia (¢ - recently confirmed sites). 


als) or because of some factor(s) making it vulnera- 
ble to extirpation or extinction.” 

It has been globally ranked by The Nature Con- 
servancy of the United States as G4. The G4 rank 
indicates that it is “frequent to common (greater than 
100 occurrences); apparently secure but may have a 
restricted distribution; or there may be perceived 
future threats.” 


Evaluation of Status 

Corydalis scouleri occurs rarely in British Colum- 
bia in wet, cool, usually shady habitats. As yet we 
cannot confirm its distribution beyond the Nitinat 
River/Klanawa River watersheds, where several 
large populations have been documented (Table 1). 
These lands are within the South Island forest dis- 
trict, and are owned and administered by timber 
companies such as Timber West. Responsible log- 
ging practices that leave an anti-erosion margin 
along watercourses should spare most known British 
Columbia populations of C. scouleri, which are 
apparently limited to streambank and riverbench 
sites. Propagation of plants (preferably seedlings 
rather than rhizome cuttings to preserve genetic 
diversity) from populations that do come under 
threat would be desirable. In addition, ecological and 
genetic studies of C. scouleri populations would add 
considerably to our knowledge regarding the reasons 
for the rarity of this species, thus increasing the 
potential for conserving viable populations. 


458 


THE CANADIAN FIELD-NATURALIST 


TABLE |. Locations and population sizes of Corydalis scouleri sites in British Columbia. 


Collection Site 


Nitinat watershed 
Old Camp 3 road (old Nitinat campsite) 
Nitinat Lake Road, 1 km west of Vernon Creek 


Nitinat River, ca. 5 km northeast of northeast end of Nitinat Lake 


Caycuse River, mouth of 

Nitinat Lake Road, north of motel 

Nitinat Lake/Bamfield—Carmanah road junction, south of 
Nitinat Lake Road, northwest of motel 

Nitinat River Road, north of visitor centre cut-off 


Nitinat Lake, northwest side, southwest of hatchery turnoff, site A 
Nitinat Lake, northwest side, southwest of hatchery turnoff, site B 
Nitinat Lake, northwest side, southwest of hatchery turnoff, site C 


Nitinat Main Road, 2 km west of Cowichan Lake 

Lake Cowichan/Nitinat—Bamfield road junction, south of 
Jasper Creek , unnamed creek SE of 

Jasper Creek bridge 

Jasper Creek, 1.9 km NW of 


Klanawa watershed 

W of N Klanawa at confluence 
Klanawa River, Moon Creek 
Klanawa River, NE of Corry Creek 
Klanawa River, NE of West Fork of 


Corydalis scouleri is considered by the authors 
and the British Columbia Conservation Data Centre 
to be threatened in Canada. Its restricted range and 
the threat of logging combine to make this species 
highly vulnerable in the near future. 


Acknowledgments 

The authors would like to extend their gratitude to 
Marie Fontaine for her able assistance in the field. In 
addition, thanks are due to the curators of the follow- 
ing herbaria for their cooperation during the research 
phase of the COSEWIC study: Royal British 
Columbia Museum, Victoria; University of 
Washington, Seattle; Washington State University, 
Pullman; and Oregon State University, Corvallis. 
Funding for this project was provided by the British 
Columbia Conservation Data Centre. 


Literature Cited 

Abrams, L. 1944. Flora of the Pacific States, Volume (Poly- 
gonaceae to Krameriaceae). Stanford University Press, 
Stanford. 635 pages. 

Brockman, C.F. 1947. Flora of Mount Rainier National 
Park. United States Government Printing Office, Wash- 
ington. 170 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. 
Illustrated flora of British Columbia. Volume 3. Dicot- 
yledons (Diapensiaceae through Onagraceae). Ministry 
of Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 436 
pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. 
Illustrated flora of British Columbia. Volume 4. Dicot- 


Vol. 115 

Last Collector/ Population 
Observation Observer (Number of stems/area) 

1997 Jamison 16/60 m2 

1997 Jamison 1100/4000 m2 

1997 Jamison 3000/2.7 ha 

1997 Jamison 500—700/4000 m2 

1997 Jamison 100/100 m2 

1997 Jamison 570/1950 m2 

1997 Jamison 1300/3250 m2 

1997 Jamison 100/60 m2 

1997 Jamison 400/2020 m2 

1997 Jamison 1000+/1000 m2 

1997 Jamison 30/50 m2 

1998 Douglas 8000/5000 m2 

1998 Douglas 2000/7000 m2 

1998 Douglas 100/5000 m2 

1998 Douglas 100000+/6 ha 

1998 Douglas 50/150 m2 

1992 Roemer unknown 

1998 Douglas 1/1 m2 

1998 Douglas 2/5 m2 

1998 Douglas 14/10 m2 


yledons (Orobanchaceae through Rubiaceae). Ministry 
of Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 427 
pages. 

Douglas, G. W., G. B. Straley, and D. Meidinger. 1994. 
The vascular plants of British Columbia. Part 4 — 
Monocotyledons. Special Report Series 4. British 
Columbia Ministry of Forests, Victoria, British Colum- 
bia. 257 pages. 

Douglas, G. W., G. B. Straley, and D. V. Meidinger. 
1998a. Rare-native vascular plants of British Columbia. 
British Columbia Conservation Data Centre, Ministry of 
Environment, Lands and Parks, Victoria, 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). Ministry of Environment, Lands and Parks, 
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. Vol- 
ume 2. Dicotyledons (Balsaminaceae through Cucurbita- 
ceae). Ministry of Environment, Lands and Parks, 
British Columbia Ministry of Forests, Victoria, British 
Columbia. 436 pages. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and J. W. 
Thompson. 1969. Vascular Plants of the Pacific North- 
west—Part 2: Salicaceae to Saxifragaceae. University of 
Washington Press, Seattle. 914 pages. 

Liden, M. 1986. Synopsis of Fumarioideae (Papavera- 
ceae) with a monograph of the tribe Fumarieae. Opera 
Botanica 88: 1-133. 

Ohara, M., and S. Higashi. 1994. Effects of inflores- 
cence size on visits from pollinators and seed set of 


2001 DOUGLAS AND JAMISON: STATUS OF SCOULER’S CORYDALIS 459 


Corydalis ambigua (Papaveraceae). Oecologia (Berlin) Dicotyledoneae (Loasaceaee to Compositae). National 

98: 25-30. Museum of Natural Sciences Publications in Botany 
Ownbey, G. B. 1947. Monograph of the North American (7): 1711 pages. 

species of Corydalis. Annals of the Missouri Botanical Szezawinski A., and A. S. Harrison. 1973. Flora of the 


Garden 34: 187-259. Saanich Peninsula. Occasional Papers of the British 
Paviick, L. E. 1989. Scouler’s Corydalis — one of Columbia Provincial Museum Number 16. Victoria. 

Canada’s rare and beautiful plants. The Victoria Natur- 

alist 45: 17. Received 29 May 2000 


Scoggan, H. J. 1979. The flora of Canada. Part 4. Accepted 12 July 2001 


Status of the Purple Sanicle, Sanicula bipinnatifida (Apiaceae), 
in Canadat | 


JENIFER L. PENNY and GEORGE W. DOUGLAS 


Conservation Data Centre, British Columbia Ministry of Environment, Lands, and Parks, Resources Inventory Branch, Wildlife 
Inventory Section, P.O. Box 9344 Station Provincial Government, Victoria, British Columbia V8W 9M1 Canada 


Penny, Jenifer L., and George W. Douglas. 2001. Status of Purple Sanicle, Sanicula bipinnatifida (Apiaceae), in Canada. 
Canadian Field-Naturalist 115(3): 460-465. 


In Canada, there are 21 extant (>1949) populations of S. bipinnatifida on southeastern Vancouver Island and the Gulf 
Islands, 14 of which have been verified in recent years, three of which have been extirpated, and four of which the present 
status is unknown. There are five historic sites, two of which have been extirpated. These populations represent the north- 
ern range limit for the species, which extends south to Baja California in northwestern Mexico. Population sizes range from 
a single plant to over 1100 plants. The habitat of S. bipinnatifida on southeastern Vancouver Island (Garry Oak stands and 
dry Douglas-fir forests) are rapidly being converted to residential and commercial developments. Although several of the 
populations receive some protection in parks and ecological reserves, they are still threatened by competition from intro- 
duced species. In addition, sites in ecological reserves, regional parks, and municipal parks are at risk due to some manage- 
ment activities and developments. Considering these threats to the habitat of S. bipinnatifida, we recommend a status of 


threatened. 


Key Words: Purple Sanicle, Sanicula bipinnatifida, threatened, distribution, population size, British Columbia. 


Purple Sanicle (Sanicula bipinnatifida Douglas ex 
Hooker) is one of approximately 40 species of 
Sanicula known world-wide (Bell 1954). [Taxonomy 
and nomenclature follows Douglas et al. (1994, 
WOE Sb L999 8es 199947 1999bs 2000; 2001} The 
genus is semi-cosmopolitan, well-represented in both 
the old and the new worlds and mainly found in the 
north temperate zone. There are five sections in the 
genus with S. bipinnatifida classified in the “Sani- 
coria” (Shan and Constance 1951). The species in 
this section exhibit a great variety of developmental 
trends in their vegetative and reproductive charac- 
ters. Eight species occur in Canada with five in 
British Columbia (Scoggan 1979; Douglas et al. 
1998b). 

Sanicula bipinnatifida is an erect, stout, branch- 
ing, perennial herb from a vertical, elongated tap- 
root (Figure 1; Hitchcock et al. 1961). It stands 
10-60 cm in height with moderately to widely 
spreading branches that originate from the base. The 
leaves, which are extremely variable in form, are 
basal or lower stem, numerous, forming a flat 
rosette, somewhat thick and leathery in texture, and 
petiolate, the petioles flattened and usually about as 
long as the blade. The blades are 4-13 cm long and 
3-12 cm wide. The cauline leaves are similar to the 
basal, but reduced upwards with some subsessile. 
The range of leaf form variation noted in Bell (1954) 


+This report was submitted to the Committee on the Status 
of Endangered Wildlife in Canada (COSEWIC), and the 
species designated Threatened, April 2001. 


was also observed in the field in British Columbia. 
Leaves are toothed and once or twice pinnately- 
divided, sometimes with an obtuse terminal leaflet, 
or simple. Leaf axes are toothed and winged. The 
inflorescence is comprised of several to many com- 
pact, 3- to 5-radiate umbels. The umbels are 20- 
flowered with 10-12 staminate flowers and 8-10 
perfect flowers. Individual flowers are wine-colored 
(purple). The involucral bracts are usually two, 
lanceolate, 0.2—2.5 cm long, foliaceous, and trisect 
or pinnatifid. The involucels are inconspicuous, 
composed of six to eight lanceolate, 2.5 mm long 
bractlets, which are slightly fused at the base with 
wax-papery margins. The fruits are dry schizocarps 
(fruits which split into separate carpels at maturity), 
egg-shaped to sub-globose, 3-6 mm long, and cov- 
ered with stout, hooked prickles. When this plant is 
vegetative or fruiting, it may look similar to some 
Pacific Sanicle (Sanicula crassicaulis) plants that 
have deeply incised to pinnate leaves and reddish 
flowers. 


Distribution 

In Canada, Sanicula bipinnatifida occurs on 
southeastern Vancouver Island and the adjacent Gulf 
Islands (Figure 2; Douglas et al. 1998a). Globally, S. 
bipinnatifida occurs on the west coast of North 
America from southern British Columbia to northern 
Baja California, Mexico. 


Habitat 
Throughout its distribution, S. bipinnatifida 
occurs in several different habitats. In Canada, this 


460 


2001 


Ficure 1. Illustration of Sanicula bipinnatifida (Line draw- 
ing by Karen Uldall-Ekman). 


species occurs in the rainshadow of the Coast and 
Olympic Mountains, in a unique Mediterranean-type 
climate where dry Douglas-fir (Pseudotsuga men- 
ziesii) forests and Garry Oak (Quercus garryana) 
stands predominate. Sanicula bipinnatifida is rela- 
tively shade-intolerant, occurring on very dry to 
moderately dry, nitrogen-rich, moder or mull humus 
form soils (Klinka et al. 1989). Early spring condi- 
tions are moist, facilitating germination and growth 
before the onset of summer drought. Specific habi- 
tats include grass-forb meadow openings in 
Pseudotsuga menziesii-Arbutus (Arbutus menziesii) 
forests, Quercus garryana-Arbutus menziesii stands, 
eroding, sandy banks on seashore cliffs, and shrub- 
by, grassy knolls. On southeastern Vancouver Island, 
Sanicula bipinnatifida is mainly a component of the 
spring flora (April-May), but may persist into late 
June. Many of the associated species in all sites are 
introduced. 

In one instance, S. bipinnatifida was found on a 
shrubby-grassy knoll dominated by Idaho Fescue 
(Festuca idahoensis), Common Hawthorn (Crata- 
egus monogyna), Scotch Broom (Cytisus scoparius), 
and Kentucky Bluegrass (Poa pratensis), the latter 
three of which are introduced. 


PENNY AND DOUGLAS: STATUS OF THE PURPLE SANICLE 


46] 


f ALBERNI ASSLAND < 
VANCOUVER \) 
ISLAND 


/ 
ee | 
/ 
/ 
/ 


oe / 


FIGURE 2. Distribution of Sanicula bipinnatifida in British 
Columbia (© — extirpated sites, ® — recently con- 
firmed sites, @ — present status unknown). 


In the grass-forb meadows, dominants in the month 
of May include brome (Bromus spp.), Sweet Vernal 
Grass (Anthoxanthum odoratum), Long-stoloned 
Sedge (Carex inops), and Many-flowered Woodrush 
(Luzula multiflora). 

In the Quercus garryana-Arbutus menziesii stands, 
grassy slopes are dominated by Hedgehog Dogtail 
(Cynosurus echinatus), California Oatgrass 
(Danthonia californica), Orchard Grass (Dactylis 
glomerata). Shrubby trailside habitats for Sanicula 
bipinnatifida are dominated by Cystisus scoparius and 
Great Camas (Camas leitchtlinii). 

The eroding sea cliff site at Dallas road is dominat- 
ed by Beach Pea (Lathyrus japonicus var. maritimus), 
Rip-gut Brome (Bromus rigidus), Soft Brome (B. 


462 


hordeaceus), and Kentucky Bluegrass, whereas the 
Macaulay Point site is dominated by Sanicula bipin- 
natifida, S. crassicaulis, Narrow-leaved Plantain 
(Plantago lanceolata) and Perennial Ryegrass 
(Lolium perenne). 

A final notable habitat is in the Mount Tzuhalem 
Ecological Reserve where feral livestock have never 
grazed and consequently there is a rich diversity of 
native forbs and grasses. In this site, Sanicula bipin- 
natifida grows on south-facing ledges in a Quercus 
garryana community with Deltoid Balsamroot (Bal- 
samorhiza deltoidea), another rare taxon. Spring wild- 
flowers that occur at this site include Sanicula crassi- 
caulis, Sierra Sanicle (S. graveolens), Henderson’s 
Shootingstar (Dodecatheon hendersonii), Spring Gold 
(Lomatium utriculatum), Blue-eyed Mary (Collinsia 
parviflora), Sea Blush (Plectritis congesta var. con- 
gesta) and several others. 


Biology 

There is little information available on the biology 
of Sanicula bipinnatifida in Canada. This species is 
briefly mentioned by Shan and Constance (1951) 
and Bell (1954) in a work on the entire genus of 
Sanicula in the old and the new worlds and a study 
of variation and polyploidy in the S. crassicaulis 
complex respectively. 


Phenology and Growth 

Sanicula bipinnatifida is a short-lived perennial. 
In Canada, it germinates in mesic, but not wet habi- 
tats of the early spring season on southeastern 
Vancouver Island. Flowering occurs by the begin- 
ning of May and may continue to the end of June. 
Experiments conducted by Bell (1954) revealed that 
germination time for seed planted in the winter or 
early spring was 41 days. According to Bell (1954), 
when seed of S. crassicaulis and the closely-allied 
species of it (including S. bipinnatifida) germinate, 
the old seed coats are shed and the small green 
cotyledons grow rapidly and usually reach their 
maximum size within a week or 10 days. At this 
time the true leaf has emerged from the sheath 
formed by the petioles of the cotyledons. The first 
year seedling results from the production of seedling 
leaves, each slightly larger than the last resulting in a 
loose rosette of small foliage leaves. Growth stops 
with the beginning of the dry season. Most species in 
the complex do not flower in the first year, but 
instead in the second or third year. 

Flowers of members of the Apiaceae are fairly 
unspecialized with little variation from species to 
species. Pollinators are likely generalist insects. 
Most species are visited by a large range of pollina- 
tors exhibiting little plant-pollinator specificity. The 
tiny and inconspicuous flowers of Sanicula bipinnat- 
ifida and others makes them less attractive to polli- 
nators. General floral features of the Apiaceae 
include: a prominent stylopodium, exposed nectar, 


THE CANADIAN FIELD-NATURALIST 


Vols 


promiscuous (non-specific) pollination, perfect flow- 
ers, protandry (male flowers mature before female), 
regular corollas, sexual reproduction, and semi-com- 
pact umbels (Heywood 1971). Within Sanicula 
species there are some exceptions to these general 
features and some floral specialization. Some of the 
species including 

S. bipinnatifida, are protogynous (female flowers 
mature before male) rather than protandrous. 
Furthermore, Sanicula species have functional 
unisexual flowers taking on the role of pistils or sta- 
mens. The stylopodium is a structure that is charac- 
teristic of the Apiaceae; it is a swollen, often color- 
ful, nectar-secreting style-base which appears to be 
significant in reproduction. In Sanicula species, the 
stylopodium is somewhat specialized; it is strongly 
flattened and reduced, thus protecting the nectar. In 
addition, Sanicula species are visited by a relatively 
high percentage of Hemipterans (True Bugs) which 
may represent a line of specialization (Heywood 
NST). 


Population Size and Trends 

There are 21 extant (>1949) populations of S. 
bipinnatifida on southeastern Vancouver Island and 
the Gulf Islands, 14 of which have been verified in 
recent years, three of which have been extirpated, and 
four of which with the present status unknown. There 
are five historic sites, two of which have been extir- 
pated. Population sizes range from a single plant to 
over eleven hundred plants. The largest population 
observed occurs at Macaulay Point with approxi- 
mately 1100 individuals occupying approximately 
130 m2 (Table 1). Some sites have as few as one to 
six plants. The historic sites listed are presently areas 
of heavy urbanization where all native vegetation has 
been destroyed including S. bipinnatifida (Table 1). 
Only limited information is known for population 
trends of S. bipinnatifida. Populations observed in 
Metchosin and on Mount Tzuhalem have not 
changed appreciably over the past five and nineteen 
years respectively Adolf Ceska, Conservation Data 
Centre- Conservation Biology Section, Resources 
Inventory Branch, Ministry of Environmental Lands 
& Parks, Victoria. (Personal Communication 1999) 
The junior author has also observed that the popula- 
tion at Cattle Point has remained stable over the last 
eight years. 


Limiting Factors 

Sanicula bipinnatifida is mainly limited by human 
disturbance. There are no apparent biological limi- 
tations. Pollination mechanisms are fairly non- 
specific, and pollinators appear not to be limiting. 
Fruit dispersal is likely effective due to the numerous 
prickles found on the schizocarps. Ecologically, pop- 
ulations of this species are somewhat restricted with- 
in Canada. Populations only occur on southeastern 


2001 


PENNY AND DOUGLAS: STATUS OF THE PURPLE SANICLE 


TABLE |. Locations and Population Sizes for Sanicula bipinnatifida in British Columbia. 


Population 


Cedar Hill, Saanich (historic) 

Cloverdale District, Saanich (historic) 

Sidney (historic) 

Blenkinsop road, Saanich (historic) 

Ten Mile Point, Saanich (historic) 

Mount Douglas vicinity, Saanich 

Flora Islet, E of Hornby Island 

Golf Hill, Esquimalt 

Little D’Arcy Island (S of Sidney Island) . 

Alpha Islet, Oak Bay Islands 

Uplands Park, N of Beach Drive, Victoria 

East Point, Saturna Island 

Dionisio Point Park, Galiano Island 

Brown Ridge, Saturna Island 

Mount Tzuhalem Ecological Reserve 

Uplands Park, Cattle Point, Victoria 

Metchosin, off Happy Valley road, on Neild road, 
2 subpopulations 

Glencoe Cove Park, Gordon Head 

Mill Hill, near summit (3 subpopulations — 
Calypso trail, Summit trail, and SE slope) 

Rithet’s Bog, strata complex south of bog 

Esquimalt, Macaulay Point, E side 

Albert Head, Metchosin 

Mount Tzuhalem (Cowichan Bay Indian Reserve 1) 

Holland Point, Victoria 

Francis King Park, SE of 

Thetis Lake Regional Park, Seymour Hill 


Vancouver Island and in the Gulf Islands in a unique 
Mediterranean-type climate which is characterized 
by warm, very dry summers and mild, wet winters. 

More important than biological or ecological con- 
siderations is the threat of urbanization. Human 
development has reduced this species historic range 
and the trend continues in the present day. The most 
threatened occurrence of S. bipinnatifida occurs on 
private land, on a tract of land southeast of Francis 
King Park (Table 1). At least one population of rare 
plants has already been destroyed on this property. 
In the absence of provincial or good federal endan- 
gered species legislation, they will not be required to 
protect the Sanicula bipinnatifida population. 

It is likely that S. bipinnatifida has been extirpated 
from five of the sites (Table 1). Sanicula bipinnatifi- 
da was observed northwest of East Point on Saturna 
Island in 1984 and has not been seen again (Harvey 
Janszen, personal communication 1999, Saturna 
Island, British Columbia). In addition, the senior 
author and the original collector of the specimen 
from Golf Hill confirmed that S. bipinnatifida has 
disappeared from the site. 

Populations in the municipal and regional parks 
are likewise not secure. Benches or trails are some- 
times constructed over rare plant populations. In the 
regional parks, there are procedures in place for 


463 


Last 
observation Collector Number of Plants/area 
1897 Macoun Extirpated 
1919 Newcombe Extirpated 
1927 Goddard Unknown 
1939 Unknown Unknown 
1942 Eastham Unknown 
1953 Melburn Unknown 
1976 Pojar Unknown 
1976 Ceska Extirpated 
1977 Ceska Unknown 
1981 Ceska Unknown 
1983 van Dieren Extirpated 
1984 Janszen Extirpated 
1993 Roemer 1/1 m2 
1999 Janszen 140/200 m2 
1999 Penny & Douglas 94/36 m2 
1999 Penny & Douglas 215 plants/54 m2 
1999 Penny 446/60 m2 & 
184/100 m2 =630 total 
1999 Penny 6/3 m2 
1999 Penny & Fleming 34/24 m2, 54/72 m2, & 
39/141 m2= 127 total 
1999 Penny & Hartwell 24/4 m2 
1999 Penny 1138/170 m2 
1999 Penny & Donovan 1014/11250 m2 
1999 Penny & Douglas 75/16 m2 
1999 Penny 63/56 m2 
1999 Penny 13/18 m2 
2000 Ussery & Fleming 152/20 m2 


determining if rare plants occur in areas slated for 
alteration via environmental impact assessments. 
However, this is not true in the municipal parks. One 
of the municipal park sites contains a very small 
population of S. bipinnatifida (six plants) and the 
other ones have moderate-sized populations. 
Populations in ecological reserves could be adverse- 
ly affected by the activities of groups attempting to 
manage for introduced species. For instance, during 
Cystisus scoparius removal at Mount Tzuhalem in 
the past, the exotic plants have been piled up and 
burned on top of a rare plant population. 

In addition, all sites are heavily infested with 
introduced species, some highly competitive. Many 
of the dominant grasses in southeastern Vancouver 
Island communities are exotic. There are also a vari- 
ety of introduced forbs present as well. 


Special Significance of the Taxon 

Sanicula bipinnatifida on Vancouver Island and 
the Gulf Islands may represent genetically distinct 
individuals because they are the northernmost popu- 
lations of the species (peripheral populations). 
Peripheral populations may be regarded as important 
pools of genetic information, but since they are not 
globally endangered, there is much less concern for 
them in terms of conservation. Some empirical evi- 


464 


dence exists indicating that isolated peripheral popu- 
lations are genetically and morphologically distinct 
from populations in the center of the distribution 
(Lesica and Allendorf 1995). These populations may 
serve as a reservoir of genetic material capable of 
expressing unique adaptations brought on by their 
peripheral environments that could be of use for 
future speciation events. For this reason, many 
authors consider these populations important for the 
long-term survival and evolution of the species 
(Mayr 1982; Lesica and Allendorf 1995). 

Another consideration of the importance of S. bip- 
innatifida is as a culinary or medicinal herb. All 
members of the Apiaceae are aromatic plants and 
include many culinary herbs. Many well-known 
essential oil constituents were first isolated from 
members of this family (Heywood 1971). Although 
not important internationally as food or medicine, 
the Miwok Indians of the Yosemite area in 
California used S. bipinnatifida as a cure-all by the 
decoction of root (Barrett and Gifford 1933). 


Protection 

Sanicula bipinnatifida has been globally ranked 
by The Nature Conservancy of the United States as 
“G5,” or “common to very common with an exis- 
tence that has been demonstrated to be secure and 
essentially ineradicable under present conditions.” 

The British Columbia Conservation Data Centre 
considers S. bipinnatifida a Ministry of Environment 
“Red-listed,” or a threatened/endangered taxon 
(Douglas et al. 1998a). This taxon is ranked as an 
“$2,” in British Columbia, or “critically imperilled 
because of rarity (typically 6-20 extant occurrences or 
few remaining individuals) or because of some fac- 
tor(s) making it vulnerable to extirpation or extinc- 
tion.” Sanicula bipinnatifida has a national rank in 
Canada of N2. 

There is no legislation that protects S. bipinnatifida 
in British Columbia and the up-coming federal endan- 
gered species legislation will only provide protection 
for plants on federal lands that are listed by the federal 
government. Currently, S. bipinnatifida is not listed 
and occurs on federal land in only two sites. 


Evaluation of Status 

Sanicula bipinnatifida has a restricted range in 
Canada, occurring only on southeastern Vancouver 
Island and the Gulf Islands. The 14 recently con- 
firmed populations fall within relatively rare ecosys- 
tems for Canada, the dry coastal Douglas-fir forests 
and the Garry Oak stands. Most of the populations 
face serious long-term threats or consist of only a 
single or a few individuals. The extensive develop- 
ment that has occurred over the last century within 
the range of S. bipinnatifida leaves little undisturbed 
habitat left to which it could further spread. In the 
absence of adequate rare species legislation and 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


active stewardship, the five populations that occur on 
private land have an uncertain future. Even in the 
parks and ecological reserves, rare plant populations 
are potentially imperilled. Activities such as con- 
structing trails, roads, or boat ramps are always pos- 
sible threats. For these reasons, the British Columbia 
Conservation Data Centre and the authors recom- 
mend that Sanicula bipinnatifida be designated as a 
threatened taxon in Canada. 


Acknowledgments 

We thank Adolf Ceska and Hans Roemer for con- 
sultation regarding biological, ecological and conser- 
vation information. In addition, we are grateful to 
Nan Spohn of Cowichan Bay for leading us to a pre- 
viously unconfirmed locality on Mt. Tzuhalem and 
Clare Allen for allowing us to access her property to 
survey a population. We also thank the Department 
of National Defence for allowing us to survey the 
population at Albert Head. 


Literature Cited 

Barrett, S. A. and W. A. Gifford. 1933. Indian life of the 
Yosemite region: Miwok. Yosemite Association. 
Milwaukee, Wisconsin. 376 pages. 

Bell, C.R. 1954. The Sanicula crassicaulis complex 
(Umbelliferae). A study of variation and polypoidy. 
University of California Press. Berkeley and Los Angeles. 
228 pages. 

Douglas, G.W., D. Meidinger, and J. Pojar. 1999a. 
Illustrated flora of British Columbia. Volume 3. Dico- 
tyledons (Diapensiaceae through Onagraceae). Ministry 
of Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 436 
pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. 
Illustrated flora of British Columbia. Volume 4. Dicot- 
yledons (Orobanchaceae through Rubiaceae). Ministry 
of Environment, Lands and Parks, 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. Dicotyle- 
dons (Salicaceae through Zygophyllaceae). Ministry of 
Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 388 
pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 2001. 
Illustrated flora of British Columbia. Volume 6. Mono- 
cotyledons (Acoraceae through Najadaceae). Ministry of 
Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 361 
pages. 

Douglas, G. W., G. B. Straley, and D. Meidinger. 1994. 
The vascular plants of British Columbia. Part 4 — Mono- 
cotyledons. Special Report Series 4. British Columbia 
Ministry of Forests, Victoria, British Columbia. 257 
pages. 

Douglas, G. W., G. B. Straley, and D. V. Meidinger. 
1998a. Rare native vascular plants of British Columbia. 
British Columbia Conservation Data Centre, Ministry of 
Environment, Lands and Parks, Victoria, British Colum- 
bia. 423 pages. 


2001 


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). Ministry of Environment, Lands and Parks, 
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 
Cucurbitaceae). Ministry of Environment, Lands and 
Parks, British Columbia Ministry of Forests, Victoria, 
British Columbia. 436 pages. 

Heywood, V.H. 1971. The biology and chemistry of the 
Umbelliferae. Academic Press, New York. 438 pages. 
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. 
Thompson. 1961. Vascular plants of the pacific north- 
west part 3: Saxifragaceae to Ericaceae. University of 

Washington Press. Seattle, Washington. 614 pages. 


PENNY AND DOUGLAS: STATUS OF THE PURPLE SANICLE 


465 


Klinka, K. V. J. Krajina, A. Ceska, and A.M. Scagel. 
1989. Indicator plants of coastal British Columbia. Uni- 
versity of British Columbia Press, Vancouver, British 
Columbia. 288 pages. 

Lesica, P., and F. W. Allendorf. 1995. When are periph- 
eral populations valuable for conservation? Conservation 
Biology 9: 753-760. 

Mayr, E. 1982. Adaptation and selection. Biologie 
Zentralblatt 101: 161-174. 

Scoggan, H. J. 1979. The Flora of Canada. Part 4 —Dico- 
tyledoneae (Loasaceae to Compositae). National 
Museums of Canada. Ottawa, Ontario. 594 pages. 

Shan, R. H., and L. Constance. 1951. The genus Sanicula 
(Umbelliferae) in the old world and the new. University of 
California Press. Berkeley and Los Angeles. 


Received 12 June 2000 
Accepted 12 July 2001 


Status of Snake-root Sanicle, Sanicula arctopoides (Apiaceae) 
in Canadat | | 


MARTA T. DONOVAN and GEORGE W. DOUGLAS 


Conservation Data Centre, British Columbia Ministry of Environment, Lands, and Parks, Resource Inventory Branch, 
Wildlife Inventory Section, P.O. Box 9344 Station Provincial Government, Victoria, British Columbia V8W 9M1 
Canada 


Donovan, Marta T., and George W. Douglas. 2001. Status of Snake-root Sanicle, Sanicula arctopoides (Apiaceae) in 
Canada. Canadian-Field Naturalist 115(3): 466-471. 


In Canada, Sanicula arctopoides is restricted to dry rocky outcrop or coastal grassy bluff habitats unique to the Victoria 
area and adjacent small islands. The species ranges along the coast from central California to its northern limit in British 
Columbia. Of the twelve sites at which S. arctopoides has been collected, the populations at six sites have been were con- 
firmed in 1999-2001, whereas the populations at the remaining sites are likely extirpated. Historic records suggest that the 
taxon has declined in abundance since the early part of the century. Population numbers vary from only fifty plants to in 
excess of six thousand. Although several populations of S. arctopoides are partially protected from direct habitat destruc- 
tion at sites in which access to the general public is restricted, introduced herbaceous species threaten the continued exis- 
tence of most populations, especially those that have few plants. The potential for dispersal of this taxon to other sites is 
limited and opportunities for colonization are constrained by the scarcity of favorable habitats. Accordingly, a status of 


“endangered” is recommended. 


Key Words: Snake-root Sanicle, Sanicula arctopoides, endangered, peripheral populations, British Columbia. 


Snake-root Sanicle (Sanicula arctopoides Hooker 
& Arnott) [Taxonomy and nomenclature follows 
Douglas et al 1994, 1998a, 1998b, 1999a, 1999b], is 
a member of a world-wide genus of about 40 species 
(Bell 1954). It is one of five species of the genus 
Sanicula occurring in British Columbia (Douglas et 
al. 1998b) and eight in Canada (Scoggan 1979). 

Sanicula arctopoides is a herbaceous taprooted 
biennial with stems widely branching at the base into 
prostrate or ascending branches 5-30 cm long 
(Figure 1; Douglas et al. 1998b). The basal leaves 
form a rosette, are somewhat succulent and often 
yellowish-green. The leaves are 3-cleft and irregular- 
ly toothed and leaf blades are 2.5-6 cm long and 
2.5—9 cm wide. The inflorescence consists of several 
to many compact umbels, which are borne by a sin- 
gle apical shoot, and develop in the axils of the 
uppermost rosette leaves. The corollas are bright yel- 
low with a conspicuous involucel that surpasses the 
flower heads. The flowers of S. arctopoides produce 
seeds in an egg-shaped schizocarp that is 2-5 mm 
long and covered with stout, hooked prickles. 

In the field, Sanicula arctopoides may be confused 
with Purple Sanicle (S. bipinnatifida) or Pacific 
Sanicle (S. crassicaulis) ) both of which may grow 


+This is a Status Report of a species restricted in Canada to 
British Columbia. This report was submitted to the 
Committee on the Status of Endangered Wildlife in Canada 
(COSEWIC) and in April 2001 the species was designated 
“Endangered”. 


nearby. Both of these plants are distinguished from S. 
arctopoides by an erect growth habit and inconspicu- 
ous involucels. Sanicula bipinnatifida has a distinctly 
toothed leaf axis, leaves that are pinnately divided and 
purple flowers. Sanicula crassicaulis has yellow flow- 
ers and leaves that are palmately or pinnipalmately 
divided and tends to grow in more sheltered grassy 
niches at sites with slightly deeper soils that retain 
moisture for longer periods of time. 


Distribution 

Sanicula arctopoides ranges from southeastern 
Vancouver Island in British Columbia to Santa 
Barbara County in central California. In Canada, the 
species occurs in the Victoria area and nearby 
islands (Figure 2; Douglas et al. 1998a). The nearest 
known extant occurrence south of Victoria is in 
Pacific County, southern Washington State (Wash- 
ington Natural Heritage Program 1999*). 


Habitat 

The restricted range of S. arctopoides in Canada 
appears to be is a result of the distinctive climate 
found along the coast of southeastern Vancouver 
Island and the southern Gulf Islands. Limited to low 
elevations, this unique area is sheltered by the rain 
shadow of the Vancouver Island and Olympic 
Mountains and is warmed by air from the Pacific 
Ocean. The rain shadow effect is responsible for a 


*See Documents Cited section. 


466 


FiGureE 1. Illustration of of Sanicula arctopoides. (Line 
drawing by Karen Uldal-Eckman) 


x 


Mediterranean climate that differs markedly from the 
rest of the province, contributing to the area’s warm, 
dry summers and mild, wet winters. The dominant 
forest vegetation in this region is Douglas-fir 
(Pseudotsuga menziesii), a fire-climax species that 
occurs in a wide range of sites from rocky outcrops 
to moist valley bottoms. In areas characterised by 
low rainfall, shallow soils and rock outcrops, Garry 
Oaks (Quercus garryana) form open stands of trees 
mixed with grass-dominated meadows. In addition to 
S. arctopoides, the rock outcrop and vernal seep 
habitats found within the rain shadow zone favour 
the growth of many other rare plants which are also 
at the northern limit of their distribution. 

Sanicula arctopoides occurs on low, dry, grassy 
coastal bluffs along the shoreline in the Victoria 
area. All sites are in vernal seeps on gravelly or 
rocky outcrops near the ocean where the plants are 
exposed to salt spray, sun and wind. Growing in 
shallow soils over bedrock, the plants have a low 
cushion form that provides protection from wind and 
desiccation and concentrates solar energy at soil 
level, warming the roots. 

Trial Island Ecological Reserve supports the 
largest and most vigorous extant population of S. 
arctopoides. The plants grow on open, west facing, 
grassy banks close to the shore and on rocky moss 
ledges with spring seepage. The population is part of 
a rich meadow community with Nodding Onion 
(Allium cernuum), Spring Gold (Lomatium utricula- 
tum) and Yarrow (Achillea millefolium). This site is 
particularly important in terms of long-term conser- 
vation because the habitat is not as fragmented as at 
other sites and it supports the growth of other rare 
plant species. 

At the Alpha Island Ecological Reserve, Sanicula 
arctopoides is associated with Barestem Desert-pars- 
ley (Lomatium nudicaule) and Beach Pea (Lathyrus 
japonicus) with Sweet Vernal Grass (Anthoxanthum 
odoratum), Red Fescue (Festuca rubra) and Hairy 


DONOVAN AND DOUGLAS: STATUS OF SNAKE-ROOT SANICLE 


467 
51°N 


50°N 


a; 
VANCOUVER G 
ISLAND 


423°W 


| 


- 
- 


128W 127°W126°W SO 125°W 124 / 
ai Yeo 
/ / 


Ze 7 


FIGURE 2. Distribution of Sanicula arctopoides in British 
Columbia (© — extirpated sites, @ — recently con- 
firmed sites, ™ — present status unknown). 


Cat’s-ear (Hypochaeris radicata). Although the 
overall site quality is fair to good, introduced species 
appeared to be more abundant here than at the Trial 
Islands Ecological Reserve. 

The population of S. arctopoides at Saxe Point 
Park is located on a gently sloping south facing 
grassy bluff in shallow soil over bedrock with Thrift 
(Armeria maritima), Sanicula crassicaulis, and 
Common Camas (Camassia quamash). Seaside 
Plantain (Plantago maritima) and Cytisus scoparius 
were also observed at this site. 

Two small populations of S. arctopoides were 
observed at Bentinck Island on a grassy bluff above a 
rocky shore. The first population, located on the 
southwest side of the island was dominated by Early 


468 


Hairgrass (Aira praecox), Idaho Fescue (Festuca ida- 
hoensis), Soft Brome (Bromus hordeaceus) and 
Barren Fescue (Vulpia bromioides). Common Velvet- 
grass (Holcus lanatus), Hypochaeris radicata and 
Garden Sorrel (Rumex acetosella) were also present. 
The second, larger population at the southern tip of 
the unnamed point that is located west of George 
Point, was associated with Aira praecox, Festuca 
rubra, Festuca rubra, Small-flowered Birds-foot 
Trefoil (Lotus micranthus), Armeria maritima, Sand 
Clover (Trifolium willdenowii), Cladina and other 
lichen species. 

The small population of S. arctoipoides at Harling 
Point is located along a footpath within a few metres 
of the ocean bluff. The habitat at this location is the 
most degraded of all the sites. Dominant species 
include the introduced grasses: Rip-gut Brome 
(Bromus rigidus), Perenniel Ryegrass (Lolium 
perenne) and Dactylis glomerata. Native species pre- 
sent included Entire-leaved Gumweed (Grindelia inte- 
grifolia) and Hooker’s Onion (Allium acuminatum). 

At Mary Tod Islet, two small populations of 
approximately one hundred mature plants and anoth- 
er hundred seedlings were found in a dry grassy 
meadow dominated by Aira praecox, Bromus rigi- 
dus, Hedgehog Dogtail (Cynosurus echinatus) and 
Holcus lanatus. Lomatium nudicaule and Two- 
coloured Lupine (Lupinus bicolor) were also present. 


Biology 

Sanicula arctopoides is a biennial, that flowers 
during a single season, in March and April of its sec- 
ond year. Umbels are produced in a hierarchical pat- 
tern and may be divided into ranks or orders based 
on their spatial location on the flowering stalk and 
the timing of flowering. Umbel orders mature in 
9-18 day intervals and each order develops in the 
axils of leafy bracts subtending the previously 
formed order. Secondary umbels are axillary to the 
single apical primary umbel, tertiary umbels are axil- 
lary to the secondary umbels and, if present, quater- 
nary umbels are axillary to the tertiary ones. Each 
umbel bears a mixture of bisexual and staminate 
flowers. The single apical primary umbel is com- 
posed almost entirely of staminate flowers. 

No information is available on the biology and 
ecology of S. arctopoides in British Columbia. There- 
fore it is not possible to compare British Columbia 
populations with those found in the main range of the 
species in Oregon and California. 

There has been some research conducted on popu- 
lations of S. arctopoides in California where the plant 
is more common. Lowenberg (1994) published the 
results of a study into the effects of floral herbivory 
on maternal reproduction in natural populations of S. 
arctopoides. The removal of inflorescences, both nat- 
urally by grazing Black-tailed Deer (Odocoileus 
hemionus Rafinesque ssp. columbianus Richardson) 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


and by artificial clipping early in the flowering sea- 
son, resulted in no loss of maternal reproduction as 
measured by seed number and seed mass. 

Lowenberg (1994) found that when primary and 
secondary umbels bearing up to one-third of a 
plant’s flowers were removed at the stage when 
plants are normally grazed, full compensation for 
seed number occurred. There are, however, thresh- 
olds both in timing and in severity of removal 
beyond which plants were unable to compensate 
fully. It is possible that an herbivore-induced change 
in allocation patterns is the most likely mechanism 
for the compensatory response observed, allowing S. 
arctopoides to shift seed production to either earlier 
or later maturing umbels. 

Gender expression in hermaphroditic plant 
species is often affected by a variety of environmen- 
tal factors, which may include herbivory. Although 
the effects of herbivory on gender display have not 
been well documented, several studies have demon- 
strated that herbivory may reverse or reduce the ten- 
dency of unmanipulated plants to produce a greater 
proportion of staminate (male) flowers as the flower- 
ing season progresses. 

Working with andromonoecious dicots, Hendrix 
and Trapp (1981), Hendrix (1984) and May and 
Spears (1988) reported that the destruction of ovaries 
in bisexual flowers led to increased bisexual flower 
production later in the reproductive season. 
Similarly, the results of a study by Diggle (1993) 
indicated that lack of pollination in the andromonoe- 
cious annual, Solanum hirtum, produced more bisex- 
ual and almost no staminate flowers throughout the 
course of the flowering season. In each of these stud- 
ies, the flowers of plants undamaged by herbivory 
and those that were heavily pollinated typically 
become increasingly male as the flowering season 
progresses. 

Unlike the andromonoecious species cited above, 
in Sanicula arctopoides, the proportion of staminate 
flowers on later umbels declines as the season pro- 
gresses. Lowenberg (1997) assessed the role of both 
herbivory and lack of pollination in determining the 
proportion of staminate and hermaphroditic flowers 
in S. arctopoides. Although neither factor appeared 
to affect gender expression to any significant degree, 
the probability of producing staminate flowers on 
later umbels was positively related to plant size, or, 
more specifically to rosette area. It was suggested 
that staminate flowers on late umbels may be of 
greater benefit to large early-blooming plants than to 
small late blooming plants because more mating 
opportunities exist when these flowers release their 
pollen. 


Population Number, Sizes and Trends 
There are twelve reported locations of S. arc- 
topoides in the Victoria area (Table 1). Of the twelve 


2001 


reported locations, only six sites could be verified 
during this study. Populations range from fifty to 
over six thousand individuals. The largest population 
observed to date occurs on Trial Island (Table 1). 

Sanicula arctopoides has been recorded in the 
Victoria area since 1897. Records at the British 
Columbia Conservation Data Centre report that it 
occurred historically in Victoria at Clover Point, at 
Cadboro Bay, at Foul Bay, on the edge of cliffs at 
Beacon Hill Park, and on the waterfront at the foot 
of Menzies Street. Although S. arctopoides may 
once have occurred at each of these-sites, all have 
been extensively disturbed and no populations have 
been verified in recent years. They have probably 
been extirpated. Two other historic records are 
unmappable because the locality is too vague (i.e., 
Victoria). . 

Although there have been no long-term studies of 
the population dynamics of S. arctopoides in the 
Victoria area, the population numbers at Trial Island 
and Harling Point have been relatively stable since 
1992 and 1977, respectively. 


Limiting Factors 

The primary and most immediate threat to 
Sanicula arctopoides in British Columbia is the loss 
and degradation of waterfront habitat on both public 
and private property in the Greater Victoria region. 
All extant populations occur in small, isolated frag- 
ments of rocky outcrop and grassy meadow habitats. 
The species is also threatened by the cultivation of 
non-native plants and competition from aggressive 
introduced species. Lawn grasses and ornamental 
horticultural plants have often been planted in parks 
where suitable habitat for S. arctopoides may once 
have existed. Human manipulation of moisture 
regimes through landscaping practices may also have 
contributed to the disappearance of S. arctopoides in 
suitable habitats. The rapid pace of land transforma- 
tion in the Victoria area may have has also resulted 
in barriers to seed dispersal. 

The most secure populations of S. arctopoides 


DONOVAN AND DOUGLAS: STATUS OF SNAKE-ROOT SANICLE 


469 


occur in protected Ecological Reserves and on 
Department of National Defense (DND) property 
where access by the general public is restricted. 

Saxe Point Park is a popular and frequently visit- 
ed Municipal Park located along the waterfront in 
Esquimalt. Although the populations of S. arcto- 
poides in the park are in an area of relatively heavy 
pedestrian traffic, they appear not to be threatened 
by park visitors at the present time. Although the site 
is subject to some trampling, this activity may, in 
fact, serve to reduce competition from introduced 
grass species. However, the population at this site 
could be at risk if park development, such as trail or 
bench construction, is scheduled in the future. 

The outlook for the small population at Harling 
Point appears to be the most precarious. Although 
the property is presently a privately owned cemetery, 
there is no assurance of protection for this oceanfront 
site from development in the future. Once a popula- 
tion becomes small because of habitat fragmentation, 
it becomes more vulnerable to demographic and 
environmental variation and loss of genetic variabili- 
ty. In some cases, small populations are at risk of 
inbreeding depression, genetic drift and loss of fit- 
ness (Primack 1998). 

Populations of aggressive weedy exotics such as 
Cytisus scoparius, Dactylis glomerata and Bromus 
sterilis have become established at all sites and 
threaten the long-term ecological integrity of each 
population. The naturally open structure of rocky 
outcrops and grassy bluffs has likely contributed to 
these habitats being somewhat predisposed to weed 
invasion, especially where the soil is disturbed. 
These invasive species have altered the composition 
and structure of large areas of the Garry Oak 
(Quercus garryana) landscape on southeastern Van- 
couver Island and in some areas have completely 
replaced the native vegetation. 


Special Significance of the Taxon 
The rock outcrop and vernal seep habitats on 
southeastern Vancouver Island are especially signifi- 


TABLE 1. Locations and population sizes for S. arctopoides in the Victoria area, British Columbia. 


Collection site 


Chain Island (Victoria) 1897 
Cadboro Bay (Victoria) 1913 
Clover Point (Victoria) 1913 
Menzies Street (Victoria) 1917 
Beacon Hill Park (Victoria) 1938 
Foul Bay (Victoria) 1942 
Alpha Islet Ecological Reserve 1999 
Bentinck Island, Rocky Point (Victoria) 1999 
Harling Point, Chinese Cemetery (Victoria) 1999 
Saxe Point Park (Victoria) 1999 
Trial Island Ecological Reserve (Victoria) 1999 
Mary Tod Islet 2001 


Last observation 


Collector Number of plants/area) 
J. R. Anderson unknown 

W. Taylor extirpated 

J. Macoun extirpated 

C. F. Newcombe extirpated 

J. W. Eastham extirpated 

G. A. Hardy extirpated 

M. Donovan & G. Douglas 52/52 m2 

M. Donovan & J. Penny 71/21 m? 

M. Donovan & G. Douglas 81/250 m2 

M. Donovan 1145/215 m2 
M. Donovan & J. Penny 6015/3076 m2 
M. Donovan & G. Douglas 100/150 m2 


470 


cant because they support the growth of a number of 
plants, in addition to S. arctopoides, that are rare in 
British Columbia. Efforts to protect rare species 
associated with this unique ecosystem may, in turn, 
help to protect many other native species associated 
with this particular habitat. 

As peripheral populations at the northern extent 
of their geographic range, these taxa present an 
interesting dilemma. On the one hand, taxa that are 
not globally endangered can be viewed as being of 
lesser conservation significance than those which 
are globally endangered. However, isolated periph- 
eral populations are often genetically and morpho- 
logically divergent from central populations and 
may have an evolutionary and ecological signifi- 
cance out of proportion to the percentage of the 
species they represent (Mayr 1982; Lesica and 
Allendorf 1995). The protection of genetically dis- 
tinct peripheral populations may be important for 
the long-term survival of the species as a whole 
(Lesica and Allendorf 1995). 


Protection 

The British Columbia Conservation Data Centre 
ranks S. arctopoides as an S1 or Red-listed species in 
British Columbia (Douglas et al. 1998a). 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 extreme rarity (typi- 
cally five or fewer occurrences or very few remain- 
ing individuals) or because of some factor(s) making 
it especially vulnerable to extirpation or extinction”. 
Since the species is restricted to British Columbia it 
has a national rank in Canada of N1. 

Although there is no specific legislation for the 
protection of rare and endangered vascular plants in 
British Columbia, the populations at Trial Island and 
Alpha Islet are located within Ecological Reserves 
that provide the plants with the greatest degree of 
legal protection currently available in British 
Columbia. The populations at Bentinck Island are 
located on land controlled by the Department of 
National Defense (DND) where public access is pro- 
hibited and, although portions of the island are 
utilised for training purposes, there are no immediate 
plans to extend these exercises to the area in which 
S. arctopoides occurs. An Environmental Risk Man- 
agement Division monitors land use on DND proper- 
ties but no endangered species legislation exists at 
the national level in Canada. The population of S. 
arctopoides at Mary Tod Islet is on property that is 
owned by the Municipality of Oak Bay. No plans for 
development at this site are known at this time and 
are considered unlikely in the future. 

The Nature Conservancy of the United States has 
designated a global rank of “GS” for the species, a 
ranking which indicates that, on a global scale, it is 
considered to be “common to very common; demon- 


THE CANADIAN FIELD-NATURALIST 


Volum) 


strably secure and essentially ineradicable under pre- 
sent conditions”. In the southern portion of its range 
along the central and north coast of California, S. 
arctopoides is common. In Oregon, the plant is not 
common but is also not considered rare by the 
Oregon Natural Heritage Program (Scott Sundberg, 
personal communication 1999, Department of 
Botany and Plant Pathology, Oregon State Univer- 
sity, Corvallis, Oregon). In the state of Washington, 
where one site remains extant, S. arctopoides is 
ranked S1S2, with a state status indicating that the 
plant is “Sensitive” (Washington State Natural 
Heritage Program 1999*),. 


Evaluation of Status 

Sanicula arctopoides is considered an endangered 
taxon in Canada by the authors and the British 
Columbia Conservation Data Centre. Suitable habi- 
tats for Sanicula arctopoides are rare in Canada and 
are restricted to the Victoria area and adjacent local 
islands. The potential for dispersal and opportunities 
for colonisation are extremely limited. 

Only six extant populations are known in British 
Columbia. The existence of populations at risk 
depends on the protection of the marginal habitats in 
which they occur. In the absence of federal or 
provincial rare species legislation or active steward- 
ship, populations of rare plants on private lands are 
much less secure than those in protected areas. 
Although three of the extant populations are protect- 
ed in Ecological Reserves, an increasing abundance 
of aggressive introduced species at these sites may 
threaten the survival of S. arctopoides in the future. 
The populations at Trial Island and Alpha Islet are 
also at risk from potential oil spills since they are 
alongside one of the most active oil shipping lanes in 
North America. 

Populations of S. arctopoides in British Columbia 
are at the northern extent of their range and may rep- 
resent populations that are genetically distinct and 
important for the long-term survival and evolution of 
the species. The exclusion of S. arctopoides and 
other peripheral species at the geographic margins of 
their range from legal protection could result in a 
significant and irreversible loss of Canada’s genetic 
resources. 


Acknowledgments 

Special thanks to the following people who assist- 
ed in this project: Jenifer Penny for her assistance 
with field work; Chris Kissinger and Doug Biffard at 
British Columbia Parks for access permits and trans- 
portation to Ecological Reserves (ER) #94 and #132. 
Transportation to ER #94 and #132 was also gener- 
ously provided by Volunteer Warden, Marilyn Lam- 
bert. We thank the Department of National Defense 
for permission to survey Bentinck Island and for 
transportation to the site. 


2001 


Documents Cited [marked* in text] 

Washington State Natural Heritage Program. 1999. 
Washington State Department of Natural Resources, 
Olympia, Washington. (Unpublished report). 


Literature Cited 

Bell, R.C. 1954. The Sanicula crassicaulis complex 
(Umbelliferae). A study of variation and polypoidy. Uni- 
versity of California Press. Berkeley and Los Angeles. 

Diggle, P. K. 1993. Developmental plasticity, genetic 
variation, and the evolution of andromonoecy in Solan- 
um hirtum (Solanaceae). American Journal of Botany 
80: 967-973. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. 
Illustrated flora of British Columbia. Volume 3. Dicot- 
yledons (Diapensiaceae through Onagraceae). Ministry 
of Environment, Lands and Parks, British Columbia 
Ministry of Forests, Victoria, British Columbia. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. 
Illustrated flora of British Columbia. Volume 4. Dicotyle- 
dons (Orobanchaceae through Rubiaceae). Ministry of 
Environment, Lands and Parks, British Columbia Min- 
istry of Forests, Victoria, British Columbia. 

Douglas, G. W., G. B. Straley, and D. Meidinger. 1994. 
The vascular plants of British Columbia. Part 4 — Mon- 
ocotyledons. Special Report Series 4. British Columbia 
Ministry of Forests, Victoria, British Columbia. 

Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. 
Illustrated flora of British Columbia. Volume 1. Gymno- 
sperms and Dicotyledons. (Aceraceae through Aster- 
aceae). Ministry of Environment, Lands and Parks, 
Wildlife Branch and Resources Inventory Branch and 
Ministry of Forests Research Branch. 

Douglas, G. W., G. B. Straley, and D.V. Meidinger. 
1998b. Rare native vascular plants of British Columbia. 


DONOVAN AND DOUGLAS: STATUS OF SNAKE-ROOT SANICLE 


47] 


BC Conservation Data Centre, Ministry of Environment, 
Lands and Parks, Victoria. 

Hendrix, S. D., and E. J. Trapp. 1981. Plant-herbivore 
interactions: insect-induced changes in host plant sex 
expression and fecundity. Oecologia (Berlin) 49: 
119-122. 

Hendrix, Stephen D. 1984. Reactions of Heracleum 
lanatum to floral herbivory by Depressaria pastinacella. 
Ecology 65: 191-197. 

Lesica, P., and F. W. Allendorf. 1995. When are periph- 
eral populations valuable for conservation? Conservation 
Biology 9: 753-760. 

Lowenberg, G. J. 1994. Effects of floral herbivory on 
maternal reproduction in Sanicula arctopoides (Api- 
aceae). Ecology 75: 359-369. 

Lowenberg, G. J. 1997. Effects of floral herbivory, limit- 
ed pollination, and intrinsic plant characteristics on phe- 
notypic gender in Sanicula arctopoides. Oecologia 109: 
279-285. 

May, P. G., and E. E. Spears, Jr. 1988 Andromonoecy 
and variation in phenotypic gender of Passiflora incar- 
nata (Passifloraceae). American Journal of Botany 75: 
1830-1841 

Mayr, E. 1982. Adaptation and selection. Biologisches 
Zentralblatt 101: 161-174 

Primack, R. B. 1998. Essentials of Conservation Biology. 
24 Edition. Sinauer Associates Inc., Sunderland, 
Massachusetts. 

Scoggan, H. J. 1979. The flora of Canada. Part 4 — 
Dicotyledoneae (Loasaceae to Compositae. National 
Museums of Natural Sciences Publication in Botany 
Number 7. 


Received 12 June 2000 
Accepted 12 July 2001 


Spatial Scales of Trapping in Small-mammal Research 


JEFF BOWMAN!?, CRISTINE V. CORKUM?, and GRAHAM J. ForBes! 


'New Brunswick Cooperative Fish and Wildlife Research Unit, P.O. Box 44555, University of New Brunswick, 
Fredericton, New Brunswick E3B 6C2 Canada 

"Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada 

Present address: Wildlife Research and Development Section, Ontario Ministry of Natural Resources, 300 Water Street, 
3rd Floor North, Peterborough, Ontario K9J 8M5 Canada, e-mail: jbowman@canada.com 


Bowman, Jeff, Cristine V. Corkum, and Graham J. Forbes. 2001. Spatial scales of trapping in small-mammal research. 
Canadian Field-Naturalist 115(3): 472-475. 


We surveyed 127 published small-mammal trapping studies (of either Peromyscus maniculatus or Clethrionomys gapperi) 
to assess the range of sizes used in the design of trap arrays. The distribution of trap spacings was bimodal, with peaks at 
10 and 15 m. The mean extent of trapping arrays was 1.8 ha for grids and 358 m for transects. Only seven manuscripts 
mentioned prebaiting. The results suggest that many small-mammal studies are designed in a similar way, at small spatial 
scales. Larger-scale patterns and processes may occur undetected without larger-scale sampling designs. We present and 
discuss some recent examples from the literature. 


Key Words: Clethrionomys, dispersal, landscape context, Peromyscus, prebaiting, sampling, scale, small mammal, spatial 
autocorrelation, synchrony, trapping, winter mortality. 


Ecological research has often been carried out 
over a narrow range of relatively small spatial scales 
(Kareiva and Anderson 1989; Brown and Rough- 
garden 1990). This broad observation also may apply 
to small-mammal research. One consequence of 
working at small spatial scales is that larger-scale 
patterns and processes can be overlooked or misin- 
terpreted (Wiens et al. 1993). For example, Wegner 
and Merriam (1990) showed that White-footed Mice, 
Peromyscus leucopus, use agricultural fields adja- 
cent to forest fragments; a phenomenon that could 
have been overlooked without considering landscape 
context. Empirical small-mammal field studies com- 
monly employ grid or transect arrays of traps, the 
design of which often establishes the spatial scale of 
the study. We surveyed the literature to assess the 
range of trapping array sizes, and thus spatial scales, 
used in small-mammal research. 


Methods 

We surveyed 127 studies published in five jour- 
nals (American Midland Naturalist, The Canadian 
Field-Naturalist, Canadian Journal of Zoology, 
Ecology, and Journal of Mammalogy) between 1960 
and 1998. We selected only field-based studies of 
two common species: the Red-backed Vole, 
Clethrionomys gapperi, and the Deer Mouse, Pero- 
myscus maniculatus. From each published manu- 
script, we tabulated information on trap spacing, spa- 
tial extent of grid or transect, total trap nights of the 
study, length of trapping period, length of prebait 
deriod, and the number of grid or transect replicates. 
Replication was a difficult issue to evaluate as 
authors often did not clearly discuss assumptions 
about the spatial independence of sampling sites. 


Grids or transects were assumed to be replicated if 
multiple sites were sampled within a single study, 
whether or not these sites were truly independent in 
the statistical sense. This included the few studies 
where large-scale questions were addressed by the 
spatially-explicit juxtaposition of small replicates 
(e.g., Morris 1996). Areal units were converted to ha 
and length was converted to m. Some authors pub- 
lished multiple manuscripts from one field study, 
and in these cases we only included one sample. 
When multiple designs (e.g., transect and grid) were 
used in one study they were considered as separate 
samples. We carried out an exploratory analysis of 
the tabulated data. 


Results and Discussion 

The majority of studies in our survey (N = 80; 
Table 1) employed trapping grids rather than tran- 
sects. The mean extent of the grids was 1.8 ha while 
the mean extent of transects was 358 m (Figure 1; 
Table 1). More than 50% of transects were < 300 m, 
and more than 50% of grids were 1 ha or smaller 
(Figure 1). 

Although the mean trap spacing was 14 m, the dis- 
tribution of spacings was bimodal, with peaks at 10 
and 15 m (Figure 2; Table 1). The convention of using 
10- or 15-m trap spacing seems to be based in part on 
papers by Burt (1940), Calhoun (1948), Kikkawa 
(1964) and Smith et al. (1975). Calhoun (1948) pre- 
sented a standardized protocol for the “North 
American Census of Small Mammals”, which used 
trap stations spaced 20, 50, or 100 ft apart. Note that 
50 ft is approximately 15 m. Kikkawa (1964) suggest- 
ed a 10-m spacing in a deciduous woodland, while 
Smith et al. (1975) indicated that 15 m is a good 


472 


2001 BOWMAN, CORKUM, AND FORBES: SPATIAL SCALES OF TRAPPING 473 
TABLE 1. Descriptive statistics of small-mammal trapping designs published between 1960 and 1998 in five journals*. 
Variable N Mean Median SE Min Max 
Trap spacing (m) 114 14 15 0.6 2 45 
Extent (grid; ha) 80 1.8 1.0 0.3 < 0.1 18 
Extent (transect; m) 30 358 294 35.5 16 1309 
Number of replicates 116 24 6 4.6 l 429 
Length of trapping period (# nights) 68 4 3 0.3 2 14 
Length of prebait period (# nights) 121 0.1 0.0 <0.1 0 5 
Total trap nights 112 11238 5346 1403 154 90000 


*Field studies of Peromyscus maniculatus and/or Clethrionomys gapperi published in American Midland Naturalist, 
Canadian Field-Naturalist, Canadian Journal of Zoology, Ecology, or Journal of Mammalogy. 


compromise distance for studying a range of species. 
However, we agree with Tew et al. (1994) that there 
should be no a priori standard distance between traps. 


The selection of a trap spacing should be based on the 
question of interest and on the site-specific biology of 
the study species. For example, a spacing might be 


selected so that each individual has a trap within its 
home range. This is balanced against the extent of the 
trapping design and the number of traps logistically 


no te a8 feasible. In practice, the extent of the trapping unit 
= 10 x (grid or transect) and the spacing of traps are chosen 
= Transects 0.3 3 as a compromise between wanting a large area covy- 
7) ie ered in traps and wanting adequate coverage of that 
— 8 N= 30 mi P g 
fe) Sh area. For example, Tew et al. (1994) demonstrated 
o 6 0.2 is that a 10-ha grid with a 24-m spacing was an efficient 
r= ens: way to sample a low density, widely dispersed Wood 
= = Mouse (Apodemus sylvaticus) population. 
= 0.1 © We were surprised at how rarely prebaiting has 
9 : been practised by small-mammal researchers. Only 7 
of 127 studies indicated that traps were prebaited. 
Other authors either did not mention prebaiting or 
0 500 4000 4500 specifically indicated that it did not take place. 
Extent (m) 
50 
0.4 
be 60 @ 40 
© 07 DU 8 
= Grids oS = 30 
5 40 N = 80 O52 5 
at = r= 0.2 
® 30 045 §& 20 
o Zz 
5 > 5 ef ae 
ms Ga) oy ln 1G ne 
sia 0.1 
0 0.0 
0 5 10 15 20 0 10 20 30 40 50 


Extent (ha) 


Figure 1. Spatial extent of trapping arrays used in studies of 
Clethrionomys gapperi and/or Peromyscus manicu- 
latus published in five journals (American Midland 
Naturalist, Canadian Field-Naturalist, Canadian 
Journal of Zoology, Ecology, and Journal of Mam- 
malogy) between 1960 and 1998. 


Trap spacing (m) 


Figure 2. Trap spacing used in studies of Clethrionomys 
gapperi and/or Peromyscus maniculatus published 
in five journals (American Midland Naturalist, Can- 
adian Field-Naturalist, Canadian Journal of Zool- 
ogy, Ecology, and Journal of Mammalogy) between 
1960 and 1998. 


474 


Chitty and Kempson (1949) suggested that prebait- 
ing could be an important method for avoiding the 
“new object reaction” by rodents. Though there is 
limited evidence that more animals can be captured 
during a given trapping period by employing this 
technique (Chitty and Kempson 1949) it seems that 
prebaiting is not widely used, or at least not widely 
reported, in small-mammal studies. 

Our survey revealed that between 1960 and 1998 
most empirical studies of C. gapperi or P. manicula- 
tus used similar, small-scale designs. Trapping grids 
were mostly < 2 ha in extent (or transects < 500 m), 
and traps were spaced 10- or 15-m apart. Replicate 
trapping grids often were used (mean number of 
replicates = 24.0; median = 6; Table 1). These repli- 
cates usually were considered to be spatially inde- 
pendent and were used to generate variance esti- 
mates. It is apparent that few of the studies in our 
review have been designed to address questions 
about large-scale spatial processes. There is, howev- 
er, a need for such questions since we cannot assume 
that population processes are restricted to small 
areas. 

This last point has been empirically demonstrated 
in recent years, by the few studies that have been 
designed to look at large-scale spatial processes in 
small-mammal populations. For example, Morris 
(1992) and Knight and Morris (1996) have used the 
spatially-explicit juxtaposition of small, replicate 
trapping grids to measure density-dependent habitat 
selection. This approach is different from many oth- 
ers in that the replicate grids are not considered as 
independent samples. Rather, their juxtaposition in 
space is used to measure spatial processes. Bowman 
et al. (2001a,b) also used explicitly juxtaposed rephi- 
cates. They have demonstrated dynamic temporal 
and spatial structure in Clethrionomys and Peromy- 
scus populations over a spatial scale that corresponds 
to dispersal distance and they have hypothesized that 
winter extinctions and spring recolonizations play an 
important role in the spatial dynamics of these popu- 
lations. Finally, a number of authors have used very 
large, spatially-explicit grid or transect designs to 
measure regional synchrony in small-mammal popu- 
lation dynamics (e.g., Steen et al. 1996; Bjornstad et 
al. 1999; Mackinnon et al. 2001). 

A common feature of these recent studies is that 
they use sampling designs that measure space over 
relatively large areas as a surrogate for spatial popu- 
lation processes. This is either done by using a sys- 
tematic design that controls the distance between 
replicates (e.g., Morris 1992) or by varying the spa- 
tial extent of environmental samples taken around 
replicates (e.g., calculating landscape metrics over 
buffers of varying radii; Bowman et al. 2001c). A 
major challenge currently facing small-mammal 
ecologists is to link these relatively large-scale pro- 
cesses with the kind of well-studied, local popula- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


tion dynamics that have long been sampled using 
the traditional methods described in our literature 
review. 


Acknowledgments 

The authors acknowledge the Sustainable Forest 
Management Network (SFMN) for financial support. 
JB and GF received additional funding from Fraser 
Papers Inc., and the Sir James Dunn Wildlife 
Research Centre. JB received scholarship funding 
from NSERC and the University of New Brunswick, 
and CC received support from the University of 
Alberta. John Bissonette, Tony Diamond, Tim Dil- 
worth, Doug Morris, Roger Powell, Marc-André 
Villard, and two anonymous reviewers commented 
on the manuscript and Jan Murie gave some wel- 
come assistance with the literature search. 


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Received 19 March 2000 
Accepted 21 August 2001 


Diets of Nesting Boreal Owls, Aegolius funereus, 
in Western Interior Alaska 


JACKSON S. WHITMAN!2 


1Alaska Department of Fish and Game, Wildlife Conservation Division, P.O. Box 230, McGrath, Alaska 99627 USA 
2Current address: Alaska Department of Fish and Game, Wildlife Conservation Division, 304 Lake Street, Room 103, 
Sitka, Alaska 99835 USA 


Whitman, Jackson S. 2001. Diets of nesting Boreal Owls, Aegolius funereus, in western interior Alaska. Canadian Field- 
Naturalist 115(3): 476-479. 


Boreal Owls, Aegolius funereus, a circumboreal species, are relatively common throughout interior Alaska where suitable 
habitat exists. Although research has been conducted on the conspecific Tengmalm’s Owl in Eurasia, little is known con- 
cerning their requirements in North America, especially in Alaska. Along with other aspects of Boreal Owl ecology, I inves- 
tigated prey selection and predation rates using nest boxes in western interior Alaska during springs of 1995-1997 based on 
a total of 778 prey items found in nest boxes. As with studies elsewhere, microtines were the most important dietary compo- 
nent. I hypothesize that plant phenology would influence prey selection during the nesting season because of variations 
noted in proportions of Northern Red-backed Voles, Clethrionomys rutilus, a forest-dwelling species, and Meadow Voles, 
Microtus pennsylvanicus, generally preferring more open areas. From the differences noted in proportions of these two 
species in the diet, I suspect that Meadow Voles are selected for when grasses and sedges are short. When vegetation growth 


obscures this prey item, Boreal Owls apparently switch to a higher proportion of Northern Red-backed Voles. 


Key Words: Aegolius funereus, Alaska, Boreal Owl, Clethrionomys, diet, Microtus, nest boxes, polygyny. 


Boreal Owls (Aegolius funereus L.) are listed as 
common throughout interior Alaska (Armstrong 
1980). Few natural nest sites have been documented, 
although Boreal Owls and the Eurasian conspecific, 
Tengmalm’s Owl, readily use artificial nesting struc- 
tures (Franz et al. 1984; Sonerud 1989; Korpimaki 
1992). Because of their status as predators at or near 
the top of the food chain, owls are often viewed as 
bio-indicators of the health of an ecosystem. In part 
because the U.S. Forest Service has designated the 
Boreal Owl as a “sensitive species” requiring special 
management, basic ecological data are important to 
collect, analyze, and understand, especially in eco- 
systems that have not been subjected to man’s vari- 
ous modifications. These baseline data are useful for 
measuring and understanding changes due to habitat 
alteration or modification. 

As part of a larger investigation of Boreal Owl 
ecology in the Upper Kuskokwim River Basin, 
Alaska, direct observations of prey items in nest 
boxes during incubation and brood-rearing periods 
were used to quantify changes in the nesting-season 
diet. Other aspects of this investigation will be pre- 
sented elsewhere (Whitman, in preparation). 


Study Area 

This study was conducted in western interior 
Alaska near the village of McGrath (62°55'N, 
155°30'W). Elevations ranged from 90 m to 150 m 
above MSL. Bottomlands associated with the 
Kuskokwim River floodplain were dominated by 
White Spruce (Picea glauca), Balsam Poplar (Popu- 
lus balsamifera), or Paper Birch (Betula papyrifera), 


with understories consisting largely of woody shrubs 
(Salix alaxensis, Vaccinium spp. and Alnus crispa). 
Poorly drained sites were often dominated by 
Eastern Larch (Larix laricina). Upland sites away 
from the river were generally Black Spruce (P. mari- 
ana) with small copses of Quaking Aspen (Populus 
tremuloides) and Paper Birch. Generally, understory 
vegetation in the upland sites was mat and cushion 
lichens (largely Cladina spp.) and Red Cranberry 
(Vaccinium vitis-idaea). 

Climate in the area was largely continental, with 
cold winters (mean January temperature —23°C, 
extremes to —60°C) and moderate summers (mean 
July temperature +15°C, extremes of +30°C). Mean 
annual precipitation was 41 cm, most falling as 
snow. Snow cover at McGrath generally lasts from 
early October through April, and sometimes accu- 
mulates to depths over 125 cm. 

Human influence in the area has been light. There 
were less than 500 residents of McGrath, and no 
roads access the village. Travel was usually facilitat- 
ed by small boats or aircraft in the summer. In win- 
ter, snowmachines were the primary mode of trans- 
portation. Only very light logging has occurred, with 
most products being firewood and saw lumber for 
local use. 


Methods 

In the study area segment where Boreal Ow! diets 
were investigated, 36 artificial nesting boxes, con- 
structed of rough-cut 2.5 cm spruce lumber, were 
placed in suitable trees near access corridors (along 
navigable rivers or roadways). In 1995, only one 


476 


2001 


active site was used for the diet investigation. During 
both 1996 and 1997, all boxes were visited two or 
three times early in the incubation period to deter- 
mine use. In 1996, 14 active nest sites were visited 
an average of 8.6 times during incubation and brood- 
rearing periods, for a mean of 3.9 days between vis- 
its. In 1997, 14 active nest boxes were visited at a 
mean rate of 8.2 days between examinations. Access 
to the boxes was aided by use of an aluminum exten- 
sion ladder and cordless electric screw-gun for 
removing and replacing the tops of boxes. Following 
the nesting season, boxes were cleaned and repaired, 
and a 5-7 cm layer of clean wood chips was placed 
in each box for nesting duff. 
All prey items were examined to determine species, 


and feet of rodents and birds were removed with fin- - 


gernail clippers and discarded to assure against future 
duplication. Mammal identifications were usually 
based on gross characteristics, although dental exami- 
nations were required on four young specimens. Birds 
were identified based on feather characters. With the 
exception of bird and mammal feet, no prey items 
were removed from the nest boxes. Due to time con- 
straints, no attempt was made to examine prey 


WHITMAN: DIETS OF NESTING BOREAL OWLS 


477 


remains based on analyses of the detritus “brick” in 
the floor of nest boxes following fledging. 

Changes in proportions of prey items in. Boreal 
Owl diets during the nesting season were based on 
frequency of occurrence, and were analyzed and pre- 
sented in 5-day increments. Samples were collected 
between 11 May—15 June in 1995, and 20 April—5 
June during 1996. In 1997, diets were examined only 
during 20—24 April and 16 May-—5 June. 


Results 

Due to differences in number of active boxes as 
well as number of nest site visits, we documented 37 
individual prey items during 1995, 530 items during 
1996, and 201 during 1997. Most prey found in nest 
boxes was mammalian, comprising 95%, 98% and 
96% of the identified prey in 1995, 1996, and 1997, 
respectively (Table 1). In his summary of Boreal Ow] 
diets, Hayward (1994) indicated a heavy dependence 
on microtines throughout their North American range 
where studies have been undertaken. Insects, amphib- 
ians, and reptiles were non-existent in the Alaska prey 
remains, and birds were represented only rarely. 

During 1995, the nesting season diet depicted in 


TABLE 1. Diet of nesting Boreal Owls (Aegolius funereus) in western Interior Alaska during 1995-1997 based on prey 


items recorded from nest boxes. 


1995 1996 1997 
no. % no. % no. % 
MAMMALIA 
Microtidae 
Clethrionomys rutilus 4 10:3 72 271 a0 ~ 101 50.2 
Microtus pennsylvanicus 30 Slt yee 40.9 72, 35.8 
Microtus xanthognathus 10 128 wi 8.5 
Lemmus sibiricus w 1.3 
Subtotal 34 91.9 509 94.3 190 955 
Zapodidae 
Zapus hudsonius 1 eg 2 0.4 
Subtotal 1 1.7 Zz 0.4 
Soricidae 
Sorex spp. 19 33) 2 1.0 
Subtotal 19 35 2 1.0 
Leporidae 
Lepus americanus 1 0.5 
Subtotal 1 0.5 
TOTAL MAMMALIA 35 94.6 530 98.1 193 96.0 
AVES 
Spruce Grouse Falcipennis canadensis 1 0.2 
Gray Jay Perisoreus canadensis 1 0.2 ] 0.5 
Boreal Chickadee Poecile hudsonica i 0.2 1 0.5 
Swainson’s Thrush Catharus ustulatus 1 0.2 
Varied Thrush Ixoreus naevius | 0.2 
American Robin Turdus migratorius 1 0.2 
Yellow-rumped Warbler Dendroica coronata 1 Bol, 1 0.2 
White-crowned Warbler Zonotrichia leucophrys ] 0.2 
Common Redpoll Carduelis flammea 1 2 6 3.0 
unknown passerine not identified 2 0.4 
TOTAL AVES 2 5.4 10 1 8 4.0 
GRAND TOTAL 37 100.0 540 100.0 201i 100.0 


478 


Table 1 was from a single nest box. This box was 
visited daily from hatching of the first owlet on 11 
May until fledging on 15 June. No fresh prey items 
were found in this box after 28 May, although obvi- 
ously, the attendant male continued to deliver food 
to the box. As the dietary needs of the brooding 
female and her five young were high, prey items 
were probably consumed soon after the prey was 
delivered by the male. Although the primary reason 
for daily nest visits was to gather growth data on 
the hatchlings (Whitman, in preparation), prey 
occurrence was recorded. During both 1996 and 
1997, diet data were collected from 14 active nest 
boxes. As with 1995, fresh prey found in boxes 
during 1996 and 1997 declined during later brood- 
ing periods. 

When frequency of occurrence of prey items was 
analyzed by 5-day periods during 1996, incidence of 
Northern Red-backed Voles (Clethrionomys rutilus) 
was relatively high during early incubation (late 
April), then declined. During the last half of May 
and early June, however, their incidence was again 
quite high (Figure 1). During the late-April through 
mid-May period when incidence of Northern Red- 
backed Voles in the diet declined, their occurrence 
was replaced by a higher incidence of Meadow 
Voles (Microtus pennsylvanicus). Other prey items 


100% : EZ = 
80% | 
60% 
40% = 
20% 
o, _ . 


20-24 April 25-30 April 
n=135 n=32 


NUMERICAL PERCENT OF PREY 


RIMIE (PERIOD 250 joys 0 acs 6 lie pln 
lM %Microtus pennsylvanicus O%Other prey 


fm % Clethrionomys rutilus 


THE CANADIAN FIELD-NATURALIST 


1-5May 6-10May 11-15May 16-20May 21-25 May 26-31 May 
n=86 n=87 n=39 n=137 n=52 


Voly iS 


made up a relatively small proportion of the diet 
throughout the nesting period. 


Discussion 

In North America, data from three investigations 
indicates that Boreal Owls rely heavily on microtines. 
In Canada, Bondrup-Nielsen (1978) found 76% of 
prey items were Clethrionomys or Microtus. In 
Colorado, these two genera made up 79% of the diet 
(Palmer 1986), and in Idaho, Hayward et al. (1993) 
indicated microtines made up 55% of the prey. Not 
unlike our observations, birds made up less than 10% 
of the diet in all previous North American studies. 

In western Finland, Korpimaki (1986, 1988) and 
Korpimaki and Norrdahl (1989) found that Microtus 
and Clethrionomys collectively made up 77% of the 
diet of Tengmalm’s Owls. It appears, however, in 
both North America and Europe, diets become more 
varied as latitude decreases, probably simply reflect- 
ing a greater diversity of available prey (Kloubec 
and Vacik 1990). Seasonal and annual variations 
occur as well in Boreal Owl diets (Hayward 1994). 

I suspect that Boreal Owls prefer to hunt in small 
openings in the forest (either natural or man-made) 
where grasses and sedges are prevalent. In years of 
late snow cover, April and early May hunting is prob- 
ably restricted to sloughed river banks, roadways, and 


1-5 June 


n=140 n=33 


Ficure 1. Percent frequency of occurrence of major prey items in the diet of nesting Boreal Owls (Aegolius funereus) dur- 
ing 5-day intervals during 1995-1997 from western interior Alaska. 


2001 


small bare patches around the bases of spruce trees. In 
these situations, Red-backed Voles are probably the 
predominant prey available. As open meadows 
become snow-free, these areas appear to be hunted 
more intensively, as evidenced by the increasing inci- 
dence of Meadow Voles as prey items. Then, in late 
May, as forest environs become devoid of snow (and 
visibility of Meadow Voles declines) hunting efforts 
shift back to more heavily forested habitats where 
understory vegetation is sparse, and Red-backed 
Voles become the primary target. Amount of snow 
and plant phenology certainly influences hearing and 
visibility and thus, hunting success. 

Most, owl diets are determined from analysis of 
nesting cavity “bricks” (Ted Swem, personal com- 
munication), from observations of foraging owls, or 
from regurgitated pellets. From these analyses, 
investigators are able to document a wider variety of 
prey over a longer period of the year, but it is diffi- 
cult to document seasonal shifts in prey selection. 
Therefore, the data herein are helpful in understand- 
ing the seasonal importance of micro-habitats for 
foraging. Small clearings in otherwise monoculture 
forests (resulting from small-scale clear-cut logging, 
remote homestead clearing, small wildfires, and 
rights-of-way, among other things) may be benefi- 
cial to Boreal Owls if they result in habitats that 
become snow-free earlier in the year. 

On one occasion during 1997, the rear half of a 
juvenile Snowshoe Hare (Lepus americanus) was 
found in a nestbox. Another active nest site, about 
400 m from the first, held the front half of a hare. 
Comparison of the halves and the rareness of hares 
in the diet confirmed it was but one animal. I suspect 
one male was responsible for maintaining two 
females with broods at these particular sites, strongly 
Suggesting polygyny. Although most brooding 
females were captured at the nest boxes and banded, 
no attempts were made to mark adult males, so con- 
firmation was not possible. In Europe, polygyny has 
been documented in Tengmalm’s Owl (Solheim 
1983; Korpimaki 1991), especially during years of 
high food abundance. 


Acknowledgments 

Several people contended with insatiable mos- 
quitoes and rickety ladders to assist in the collection 
of field data. I am particularly indebted to Natalia and 
Valentin Whitman for their help. Tashi and Norv 
Dallin were always enthusiastic about owls, and 
assisted on numerous occasions. Patrick Snow col- 
Jected diet information during periods when the 
author was otherwise unavailable. The McGrath High 
School biology class built many of the original nest 
boxes and assisted in their placement. Ted Swem and 
Jeffrey Hughes encouraged continuation of the pro- 
ject after the first year of preliminary work. Beverly 
Minn, John Wright, A. J. Erskine, Jeff Hughes, and 


WHITMAN: DIETS OF NESTING BOREAL OWLS 


479 


an anonymous reviewer provided many helpful com- 
ments and constructive criticism. The Alaska Depart- 
ment of Fish and Game periodically provided vehicle 
fuel and support for the project. 


Literature Cited 

Armstrong, R. H. 1980. Guide to the birds of Alaska. 
Alaska Northwest Publishing Company, Edmonds, 
Washington, USA. 

Bondrup-Nielsen, S. 1978. Vocalizations, nesting, and 
habitat preferences of the boreal owl (Aegolius funereus) 
in North America. M.S. thesis, University of Toronto, 
Ontario. 151 pages. 

Bondrup-Nielsen, S. 1984. Vocalizations of the Boreal 
Owl, Aegolius funereus richardsoni, in North America. 
Canadian Field-Naturalist 98: 191-197. 

Franz, A., T. Mebs, and E. Seibt. 1984. Several aspects 
of the population biology of Tengmalm’s Owl (Aegolius 
funereus) in southern Westphalia and adjacent areas on 
the basis of ringing results. Die Vogelwarte 32: 
260-269. 

Hayward, G. D., P. H. Hayward, and E.O. Garton. 1993. 
Ecology of Boreal Owls in the Northern Rocky 
Mountains, USA. Wildlife Monographs 124: 1-59. 

Hayward, G. D. 1994. Review of technical knowledge: 
boreal owls. Pages 92-127 in Flammulated, Boreal, and 
Great Gray Owls in the United States: A Technical 
Conservation Assessment. Edited by G. D. Hayward and 
J. Verner. General Technical Report RM-253. Fort 
Collins, Colorado. U.S. Department of Agriculture, Forest 
Service, Rocky Mountain Forest and Range Experiment 
Station. 214 pages 3 maps. 

Kloubec, B., and R. Vacik. 1990. Outline of food ecology 
of Tengmalm’s owl (Aegolius funereus L.) in Czech- 
oslovakia. Tichodroma 3: 103-125. 

Korpimaki, E. 1986. Seasonal changes in the food of the 
Tengmalm’s Owl Aegolius funereus in western Finland. 
Annales Zoologici Fennici 23: 339-344. 

Korpimaki, E. 1988. Diet of breeding Tengmalm’s Owls 
Aegolius funereus: long-term changes and year-to-year 
variation under cyclic conditions. Ornis Fennica 65: 
21-30. 

Korpimaki, E., 1991. Poor reproductive success of poly- 
gynously mated female Tengmalm’s Owls: are better 
options available? Animal Behaviour 41: 37-47. 

Korpimaki, E. 1992. Fluctuating food abundance deter- 
mines the lifetime reproductive success of male 
Tengmalm’s Owls. Journal of Animal Ecology 61: 
103-111. 

Korpimaki, E. and K. Norrdahl. 1989. Predation of 
Tengmalm’s Owls: numerical responses, functional 
responses and dampening impact on population fluctua- 
tions of microtines. Oikos 54: 154-164. 

Palmer, D. A. 1986. Habitat selection, movements and 
activity of Boreal and Saw-whet Owls. M.S. thesis. 
Colorado State University, Fort Collins, Colorado, USA. 

Solheim, R. 1983. Bygyny and biandry in the Teng- 
malm’s Owl. Ornis Scandinavica 14:51—57. 

Sonerud, G. A. 1989. Reduced predation by pine martens 
on nests of Tengmalm’s Owl in relocated boxes. Animal 
Behaviour 37: 332-333. 


Received 24 May 2000 
Accepted 23 July 2001 


Influence of Predation on Piping Plover, Charadrius melodus, 
and Least Tern, Sterna antillarum, Productivity along the Missouri 
River in South Dakota 


CASEY D. KRUSE!, KENNETH F. HIGGINS2, and BRUCE A. VANDER LEE? 


1U.S. Army Corps of Engineers, P.O. Box 710, Yankton, South Dakota 57078 USA 

2U.S. Geological Survey, South Dakota Cooperative Fish and Wildlife Research Unit, Box 2140B, Brookings, South 
Dakota 57007 USA 

3Department of Wildlife and Fisheries Sciences, South Dakota State University, Box 2140B, Brookings, South Dakota 
57007 USA 


Kruse, Casey D., Kenneth F. Higgins, and Bruce A. Vander Lee. 2002. Influence of predation on Piping Plover, 
Charadrius melodus, and Least Tern, Sterna antillarum, productivity along the Missouri River in South Dakota. 
Canadian Field-Naturalist 115(3): 480-486. 


Predation along the Missouri River in South Dakota was examined from May—August during 1991 and 1992 to determine 
its influence on Piping Plover (Charadrius melodus) and Least Tern (Sterna antillarum) productivity. Egested raptor pellet 
collections, track and trail surveys, time-lapse photography, and visual observations were used to identify predators at 
active colony sites. Predation was the leading cause of nest and chick loss. American Crow (Corvus corvus), Raccoon 
(Procyon lotor), and Mink (Mustela vison) caused 98.0% of known nest losses. American Kestrels (Falco sparverius) and 
Great Horned Owls (Bubo virginianus) accounted for 93.0% of the documented chick mortalities. Chick escape shelters 
and wire mesh predator exclosures were evaluated as a means of increasing nest success and chick survival. Piping Plover 
apparent nest success increased significantly (P< 0.001) from 34.4% to 61.6% with the use of predator exclosure cages. 
Chick shelters were not used by either species and appeared to provide no benefit to chick survival. High predation rates on 
the Missouri River may be the result of severe habitat deterioration and increased predator effectiveness. Management 
activities for Piping Plovers and Least Terns should be based on thorough knowledge of predator community composition 
and dynamics. 


Key Words: Piping Plover, Charadrius melodus, Least Tern, Sterna antillarum, productivity, predation, Missouri River, 


South Dakota. 


Predation is one of several factors that limits the 
productivity of Great Plains Piping Plovers (Char- 
adrius melodus) and interior Least Terns (Sterna 
antillarum) within the Missouri River Basin (Dirks 
1990; Higgins and Brashier 1993; Higgins et al. 
1999; Mayer and Ryan 1991; USFWS 1991%, 
1992*). Changes in land-use practices within the 
basin have redistributed predator communities and 
inflated local predator populations (Sargeant et al. 
1993). High rates of predation on Piping Plover and 
Least Tern eggs and chicks within this region are 
also due to habitat loss as a result of vegetation 
encroachment (Kruse 1992*), inundation, flow sta- 
bilization (Schwalbach 1988), and sandbar degrada- 
tion (Dirks 1990). Habitat loss concentrates nesting 
efforts into areas that otherwise would be avoided 
due to predation (Dolan 1973; Godfrey and 
Godfrey 1973; Patterson et al. 1991). Concentrating 
birds on limited habitats may actually attract preda- 
tors through changes in social interactions (Burger 
1984; Hunt et al. 1986; Rodgers 1987). Piping 
Plovers are territorial nesters (Cairns 1982; Haig 
and Oring 1988) that depend on cryptic coloration 
and isolation as defenses against predation 
(Tinbergen et al. 1966). Piping Plover territorial 
behavior in limited habitat may increase their sus- 


ceptibility to predation through changes in activity 
patterns caused by intraspecific intolerance, or 
through changes in nest site selection (Burger 
1987). 

Several avian [Ring-billed Gulls (Larus dela- 
warensis) and California Gulls (L. californicus), 
McCracken et al. 1981; Prindiville-Gaines and Ryan 
1988; Northern Harriers (Circus cyaneus), Whyte 
1985; Great Horned Owls (Bubo virginianus), Dirks 
1990] and mammalian [Striped Skunks (Mephitis 
mephitis) and Red Fox (Vulpes vulpes), Haig and 
Oring 1988; Mink (Mustela vison) and Raccoon 
(Procyon lotor), Dirks 1990] predators may prey on 
Piping Plover and Least Tern eggs and chicks. 
However, no research in the Northern Great Plains 
has identified specific predator species that cause 
impacts to Piping Plovers and Least Terns, deter- 
mined timing of the impacts, and evaluated nonde- 
structive methods to reduce predator effectiveness. 
Our objectives were: (1) to identify predator species 
having an impact on Piping Plover and Least Tern 
nest success and chick survival along the Missouri 
River in South Dakota, and (2) to develop and evalu- 
ate techniques to minimize the impacts of predation 
on Piping Plover and Least Tern nest success and 
chick recruitment. 


480 


2001 


Study Area 

Our study area was the Fort Randall and Gavins 
Point reaches of the Missouri River in southeastern 
South Dakota. The Fort Randall reach (80.5 km) lies 
between Fort Randall Dam, at Pickstown, South 
Dakota (43°03'21”"N, 98°32'58”W), and Lewis and 
Clark Lake at Springfield, South Dakota (42°51'09’N, 
97°53'25"W). The Gavins Point reach (96.6 km) lies 
between the Gavins Point Dam at Yankton, South 
Dakota (42°51'01”N, 97°28'52”W), and Ponca State 
Park, Dixon County, Nebraska (42°36'03’N, 
96°42'36"W ). Lewis and Clark Lake, a 22-km artifi- 
cial impoundment, connects these two reaches. About 
98% of the Piping Plovers and 80% of the Least Terns 
that breed in South Dakota are found on these two 
river reaches (Schwalbach 1988). Since 1986, annual 
populations within the study area have averaged 90 
pairs of Piping Plovers and 130 pairs of Least Terns 
(Kruse 1993). Habitat in the study area is character- 
ized by barren to sparsely vegetated inter-channel 
sandbars. Adjacent landscape consists of a forest cor- 
ridor (0-850 m) dominated by mature cottonwood 
trees (Populus deltoides). The remaining river flood- 
plain is used for agricultural purposes. 


Methods 
Productivity Surveys 

Piping Plover and Least Tern productivity surveys 
were conducted from 15 May to 15 August in 1991 
and 1992. Methods used for collection of productivity 
data were those of Schwalbach (1988) and Dirks 
(1990). Piping Plover and Least Tern nests were 
found by observing adult birds or by systematic 
searches of nesting areas. Once found, nests were vis- 
ited every 5—7 days to determine fate. Nest fate was 
determined on the basis of nest condition, presence of 
pipping fragments, egg shells, adult behavior, and 
presence of chicks (Dirks 1990). Because nests were 
easily detected and visited frequently, apparent nest 
success (hatched nests / nest attempts) was used to 
quantify nest success. Chicks were monitored at 5—7 
day intervals until they were lost or became capable of 
sustained flight (fledged). Because they have high 
probability of survival, Piping Plover chicks greater 
than 20 days old and Least Tern chicks greater than 
15 days old were counted as fledged (USFWS 1988%*). 
Fledging success was calculated as fledged chicks / 
breeding pair of adults. The number of adult breeding 
pairs was determined by the number of nests and an 
annual adult census conducted during late June. 


Predator Surveys 
Raptor Pellet Surveys 

Regurgitated pellets were used to indicate predation 
by large raptors (Errington 1930; Craighead and 
Craighead 1956) on Piping Plovers and Least Terns. 
Pellets were collected on Piping Plover and Least 
Tern nesting areas during productivity surveys and 
from raptor nests adjacent to the river along the 


KRUSE, HIGGINS, AND VANDER LEE: PREDATION ON PLOVER AND TERN 


481 


Gavins Point Reach. Efforts focused on large raptors 
common to the study area, including Great Horned 
Owls and Red-tailed Hawks (Buteo jamaicensis) 
(Dirks 1990). An aerial survey was conducted on 9 
March 1991 to locate potential raptor nests within the 
floodplain forest corridor. This date corresponded to 
the initiation of nesting activity by Great Horned 
Owls in the region (Stewart 1975) and occurred 
before spring leaf-out of the forest canopy (Gaines 
and Kohn 1982), maximizing nest visibility from the 
plane. All potential nest sites were visited during 
26-30 April 1991 to determine occupation and 
remove accumulated pellets prior to arrival of Piping 
Plovers and Least Terns. Pellets and unconsumed prey 
parts were subsequently collected within a 10 m diam- 
eter circle beneath each active nest every 10 to 14 
days until the raptor nest site was vacated. Number of 
young hatching and surviving to fledge (capable of 
sustained flight) was recorded for each nest. 

Pellets were collected, preserved and dissected 
according to Korschgen (1980). Skulls, feet, and 
feathers were used to identify Piping Plovers and 
Least Terns in pellets. If remains of more than one 
Piping Plover or Least Tern were identified in a pellet, 
large bones (e.g., skulls and femurs) were grouped to 
quantify individuals. Each Piping Plover or Least 
Tern chick identified from evidence in a pellet was 
recorded as a take, which we defined as a mortality 
directly caused by a predator. 


Occurrence Surveys 

Evidence of predator occurrence on Piping Plover 
and Least Tern nesting areas was documented during 
systematic searches conducted concurrently with 
productivity surveys. Predators observed, track trails 
(Murie 1974; Rimmer and Deblinger 1990), feces 
and feathers were used as evidence of predator 
occurrence. Evidence related to nest, egg, and chick 
remains that could be attributed to specific predators 
(Reardon 1951) were recorded as takes. Both com- 
plete and partial clutch losses were recorded as sin- 
gle takes. Evidence of occurrence was collected or 
removed and track trails were mapped to prevent 
double-counting during subsequent visits. Feces 
were used to document predator occurrence, but 
were generally not examined for Piping Plover or 
Least Tern remains unless a specific take incident 
was being investigated. 


Visual Observation 

Visual observation and time-lapse photography 
were used to document predator activity and its effect 
on Piping Plover and Least Tern behavior and sur- 
vival during the peak hatching period (6—13 June). 
Visual observations of three randomly selected nest- 
ing areas were conducted in 1991, with binoculars and 
spotting scopes from secluded shoreline blinds. Areas 
were observed during crepuscular and diurnal periods 
for seven consecutive days. Any takes or interactions 
between Piping Plovers or Least Terns and predators 


482 


were recorded. An interaction was defined as any 
predator activity causing a behavioral change in 
Piping Plover or Least Tern incubation, brooding or 
foraging activity. Minolta XL-401 super-G time-lapse 
cameras, set at 15-second exposure intervals, were 
used to record predator occurrence on five additional 
randomly selected nesting areas. Films were replaced 
every 24h for seven consecutive days. Cameras were 
placed on sandbars with the field of view encompass- 
ing the entire nesting area. 


Predator Deterrence 
Chick Shelters 

Chick shelters were placed on sandbars with 
active nests to provide escape cover for pre-fledged 
chicks during 1990. Shelters were constructed of 
wooden snow fence materials (Nan-Jenks 1982), and 
secured by burying the bottom edge in the sand to 
the first wire support (10 cm) and staking it through 
the center. Chick shelters were placed on exposed 
substrates near chick foraging and loafing areas. 
Chicks, tracks or feces found inside a shelter during 
productivity surveys indicated shelter use. 


Predator Exclosures 

Piping Plover nests were randomly assigned to 
control (uncaged) or treatment (caged) groups. 
Exclosures were installed on treatment nests imme- 
diately after locating each nest. Predator exclosures, 
designed for portability and ease of assembly, con- 
sisted of 90 X 120cm top and sides and 90 x 90 cm 
ends constructed from 5 X 10cm galvanized weld- 
wire mesh fencing (Kruse 1993). Exclosures were 
placed around each treatment nest by pressing them 
into the sand (2.5 cm) and securing them with wire 
J-hook stakes (35 cm). Exclosures were observed 
until an adult entered to attend the nest. If an adult 
did not enter within 15 minutes, the exclosure was 
removed. A Wilcoxon signed-ranks test (Wilkinson 
1990) was used to test for differences in apparent 
nesting success between control and treatment nests. 


Results 
Productivity Surveys 

Two hundred fifty-six Piping Plover and 398 
Least Tern nests were monitored during 1991 and 
1992. Eighteen Piping Plover nests and 44 Least 
Tern nests with unknown fates were excluded from 
analysis. Apparent nest success was 46.2% for 
Piping Plovers and 49.4% for Least Terns (Table 1). 
Predation was the leading cause of nest loss, 
accounting for nearly half of all destroyed nests. The 
remaining nest losses were attributed to inundation, 
weather events, adult abandonment, human distur- 
bance, and unknown causes. Of 368 Piping Plover 
and 330 Least Tern chicks that hatched, only 57 
plovers and 81 tern chicks survived to fledge (Table 
1). Fledge ratios (chicks fledged per pair of breeding 
adults) were 0.33 for Piping Plovers and 0.32 for 
Least Terns. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


TABLE |. Productivity summary for Piping Plovers and 
Least Terns nesting on the Gavins Point and Fort Randall 
river reaches of the Missouri River, South Dakota, 1991 
and 1992. 


Piping Plover Least Tern 
Total Nests 238 354 
Total Destroyed 128 (53.8%) 179 (50.6%) 
Predation 61 (25.6%) 80 (22.6%) 
Chicks Hatched 368 330 
Chicks Fledged a7 (las) 81 (24.5%) 
Raptor Pellet Surveys 


Nineteen potential raptor nest locations were 
mapped during aerial surveys of the study area. 
During subsequent field visits we confirmed that five 
pairs of Great Horned Owls and six pairs of Red- 
tailed Hawks were occupying nests on the Gavins 
Point reach. Of these, three Great Horned Owl and 
five Red-tailed Hawk nests successfully fledged 
young with a mean productivity rate of 1.2 fledg- 
lings per nesting pair for both species. Juvenile Great 
Horned Owls fledged as early as 16 May and all owl 
fledglings vacated nests by 29 May. Mean fledge 
date was 23 May, approximately two weeks before 
peak hatching (9 June) for Piping Plovers and Least 
Terns. Red-tailed Hawks had a mean fledge date of 
13 July with young fledging from 8 to 19 July, well 
after the peak hatching period for Piping Plovers and 
Least Terns. 

Forty-one Great Horned Owl and Red-tailed Hawk 
pellets were collected from the ground beneath nest 
sites. One (8%) of 13 Great Horned Owl pellets con- 
tained remains of an adult Piping Plover with no evi- 
dence of any Piping Plover or Least Tern remains in 
28 Red-tailed Hawk pellets. Eleven Great Horned 
Owl pellets were found on sandbars, 9 (82%) of 
which contained the remains of seven juvenile Piping 
Plovers and 12 juvenile Least Terns. 


Predator Surveys 

Ten avian and five mammalian predator species 
were detected on Piping Plover and Least Tern nest- 
ing areas (Table 2). Evidence of 190 occurrences of 
11 predator species were noted during weekly produc- 
tivity surveys, and 13 intrusions of nesting areas by 
three predator species were photographed during 
402 h of time lapse film exposure (Table 2). During 
267 h of visual observations, nine species of avian 
predators were observed 264 times. These avian 
predators interacted with Piping Plovers and Least 
Terns 47 times (Table 2). Responses to predator intru- 
sions included vocalized warning calls, aggressive 
“mob bombing” by terns, and “broken wing” distrac- 
tion displays and “charge” threats by plovers. 
Warning vocalizations by adults elicited escape 
behavior, with chicks either lying still or running to 
vegetative cover. Great Blue Herons (Ardea hero- 


2001 


KRUSE, HIGGINS, AND VANDER LEE: PREDATION ON PLOVER AND TERN 


483 


TABLE 2. Evidence of predator occurrence, interactions with Piping Plovers and Least Terns, and takes of Piping Plover 
and Least Tern nests and chicks on nesting colonies on the Gavins Point and Fort Randall river reaches of the Missouri 


River, South Dakota, 1991 and 1992. 


Observation Technique 


Occurrence Surveys 


Species Occurrence Takes Occurrence 
Great Blue Heron + 0 154 
Great Horned Owl 47 15 12 
Ring-billed Gull ~ 0 0 Be 
Red-tailed Hawk 30 0 8 
American Crow 10 8 9 
American Kestrel 7 ] 5 
Caspian Tern 0 0 13 
Bald Eagle 0 0 3 
Golden Eagle> 0 0 l 
Northern Harrier ] 0 0 
Raccoon 4] 15 0 
Mink 30 22 0 
Domestic Dog 13 i 0 
Coyote + 0 0 
Striped Skunk 3 0 0 
Total 190 62 264 


Interactions; Aquila chrysaetos; ‘Canis latrans 


dias), Ring-billed Gulls, and Caspian Terns (Sterna 
caspia) frequented mud flats surrounding the fringes 
of the nesting areas, but rarely elicited any behavioral 
changes by Piping Plovers and Least Terns other than 
vocalizations. In contrast, Great Horned Owls, 
American Kestrels (Falco sparverius), American 
Crows (Corvus brachyrhynchos), and Bald Eagles 
(Haliaeetus leucocephalus) were aggressively 
approached by adult Piping Plovers and Least Terns 
when they landed on or near a nesting colony. 

During predator surveys, we recorded 66 takes by 
five species. American Crows (9), Raccoons (15), 
and Mink (19) caused 43 of 44 (97.7%) documented 
nest losses. A Domestic Dog (Canis familiaris) 
accounted for one nest take. Mink (3), American 
Kestrels (4), and Great Horned Owls (15) caused 22 
known chick mortalities. An American Kestrel took 
two chicks from one colony within two hours. This 
site was completely devoid of Piping Plover and 
Least Tern chicks seven days later and it was deter- 
mined that the chicks were too young to have 
fledged and left the site. In other incidents, a pair of 
Ring-billed Gulls were observed pursuing 20 day-old 
Piping Plover chicks, and a brood of 10 to 14 day- 
old Piping Plovers were observed feeding passively 
within 2 m of a Great Blue Heron. 


Predator Deterrents 

Nine chick shelters were placed on seven sandbars 
during the 1990 nesting season. Only one Piping 
Plover chick was seen using a shelter. No other evi- 
dence of chick shelter use by Piping Plovers or Least 


Visual Observations 


Time-lapse Pellet 
Photo Analysis Total 

Int? Take Occurrence Takes Occurrence Takes 
ry 0 p 160 0 
8 0 9 20 68 35 
5) 0 0 59 0) 
l 0 0 0 38 0 
8 | 0 19 9 
5 3 2 14 4 
0 0 0 13 0 
3 0 0 3 0 
0 0 0 | 0 
0 0 0 | 0 
0 0 0 4] 15 
0 0 0 30 22 
0 0 0 13 | 
0 0 0 4 0 
0) 0 0 3 0 
47 4 13 20 467 86 


Terns was found. Chicks threatened on shoreline 
feeding areas or other exposed substrates were 
observed traveling past shelters to use vegetated 
areas or driftwood as escape cover. 

Apparent nesting success of caged nests (n=86, 
62%) was higher (P< 0.001) than for control nests 
(n=122, 34%). Of 33 caged nests that were unsuc- 
cessful, 6 (18%) were inundated, 16 (48%) were lost 
to predation, 2 (6%) were abandoned for unknown 
reasons, and 9 (27%) were lost due to human distur- 
bance, sandbar erosion, weather, or unknown causes. 
Of the 16 caged nests that were lost to predators, 11 
(69%) were destroyed by Mink, one (6%) by a 
Domestic Dog, one (6%) by a Raccoon, two (13%) 
by unknown species, and one (6%) by a Great 
Horned Owl that struck the side of cage and caused 
the adult to abandon the nest. Mink predation of 
caged nests occurred primarily after July. Nests initi- 
ated prior to 10 June and protected by cages were 
more successful (67%) than nests initiated and pro- 
tected by cages after 10 June (46%). 

Piping Plovers readily adapted to predator exclo- 
sures, with few behavioral changes noted during 
post-installation observations. Nesting Piping Plovers 
with incomplete clutches or nests in early stages of 
incubation often circled a cage before entering. Adult 
Piping Plovers with eggs in advanced stages of incu- 
bation showed greater nest affinity and did not hesi- 
tate to enter a cage. Time between final installation of 
an exclosure and return of the incubating adult 
(n= 15) ranged from 7 to 195 seconds, and averaged 


484 


60 seconds. Piping Plovers left and returned to caged 
nests by walking or running through openings in the 
wire mesh; none attempted to exit or enter a cage 
while flying. 


Discussion 

We found that predation suppressed Piping Plover 
and Least Tern productivity along the Gavins Point 
and Fort Randall reaches. Piping Plover (0.33 
chicks/pair) and Least Tern (0.32 chicks/pair) 
recruitment during our study was below that deter- 
mined necessary to recover the populations (Piping 
Plovers 1.44 chicks/pair, Least Terns 0.70 chicks/ 
pair) (USFWS 1990*). We attribute high rates of 
predation within our study area to habitat changes 
within the Missouri River Basin (Schwalbach 1988; 
Kruse 1992*). Although dynamic sandbar complexes 
are critical to Piping Plover and Least Tern produc- 
tivity, Missouri River habitat in 1991 and 1992 con- 
sisted of small, isolated semi-permanent islands. 
Because of their longevity, these islands were largely 
vegetated, limiting available nesting habitat to nar- 
row bands of bare substrate between the water and 
vegetation. In addition, because of the lack of alter- 
native nesting areas, most nesting areas had also 
been used for several years prior to our study (Dirks 
1990). Under these conditions, predator efficiency 
was extremely high. When Mink or Raccoons 
accessed a site, losses were rarely limited to a single 
nest. Predator efficiency under natural sandbar cre- 
ation/degradation cycles would likely be lower than 
those we observed due to the presence of large 
dynamic sandbar complexes that changed in size, 
location, and composition annually and the presence 
of alternative/unused nesting substrates. 

Mink, Raccoons, and American Crows were 
responsible for most of the nest predations we 
observed. We noted marked changes in the use of 
nesting areas by these predators throughout the 
breeding season. Single sets of Mink tracks were 
often found early in the season (May and June), but 
were not seen in early and mid-July. In late July, 
multiple sets of Mink tracks, which we presumed to 
be family groups, were found on nesting areas. 
Raccoon tracks, which were common during spring 
surveys, were rarely found after mid-June. We sug- 
gest that these periodic shifts in mammalian predator 
occurrences are attributable to predator reproductive 
chronology (young beginning to forage with adults) 
or the availability of alternative prey. Beginning in 
mid-July, we observed small groups (<4) of 
American Crows on nesting areas feeding on com- 
mon carp (Cyprinus carpio) carcasses. We believe 
American Crows were initially attracted to the sand- 
bars to forage on carp, and we believe any predation 
of Piping Plover and Least Tern eggs by them during 
this period was incidental or secondary in purpose. 

Predator exclosures enhanced Piping Plover nest- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


ing success on the Gavins Point and Fort Randall 
reaches in 1991 and 1992. In particular, we stress the 
importance of protecting nests in May and early 
June, when Mink and Raccoons were most frequent- 
ly observed on colony sites and overall nest preda- 
tion rates were highest. While only 19% (16/86) of 
the caged nests were destroyed by predators, 69% 
(11) of them were destroyed by Mink. Thus, we urge 
caution when considering exclosures for areas where 
Mink are the dominant nest predator, particularly 
late in the season when Mink are foraging as family 
groups, and young Mink may be able to enter exclo- 
sure cages. Reducing mesh size of the exclosure to 
5 X 5cm as suggested by Rimmer and Deblinger 
(1990), may have the potential to deter Mink better. 
In contrast to reports by Nol and Brooks (1982) and 
Reynolds (1985), we found no evidence that preda- 
tors were attracted to nests by exclosures. Although 
this may become a concern with continued applica- 
tion of this method. Although our exclosures were 
comparatively small, predators were not able to 
reach eggs inside of cages. Raccoons reached inside 
of exclosures on two occasions, but failed to take 
any eggs there. Other predators deterred by cages 
included Domestic Dogs, Striped Skunks, and Great 
Horned Owls. 

Great Horned Owls and American Kestrels were 
the major predators of Piping Plover and Least Tern 
chicks in 1991 and 1992, and we likely underestimat- 
ed their effect as evidenced by the number of our 
unexplained chick disappearances. Except for talon 
strike marks in the sand, Great Horned Owls left little 
evidence of a take. Pellet analysis provided the most 
conclusive evidence of their influence on Piping 
Plover and Least Tern chicks. Likewise, our only evi- 
dence of American Kestrel takes was by direct obser- 
vation of their predatory events. Chick loss to these 
avian predators tended to be site-specific and related 
to the presence of nesting predators along the proxi- 
mal shoreline. We often saw family groups of fledged 
Great Horned Owls loafing in the trees along the 
shore during the day and moving to perches (mostly 
snags) on colony sites in the evening. 

Chick shelters are a non-destructive deterrent 
technique that has been successfully used on barren 
Atlantic Coast beaches (Nan-Jenks 1982). In this 
study, Piping Plover and Least Tern chicks did not 
use shelters, and we have no evidence they reduced 
chick loss to predators. With brood rearing habitat in 
this region of the Missouri River Basin currently 
containing an abundance of naturally occurring 
escape cover, we do not recommend providing pre- 
fledged chicks with escape structures as a manage- 
ment technique. However, this type of artificial 
structure should be re-evaluated if natural escape 
cover is reduced or ephemeral barren sandbars 
become the dominant habitat type available. 

Although various techniques have been developed 


2001 


to survey predators and to determine predator 
impacts on wildlife populations (Schemnitz 1980), 
determining nest and chick losses on our study area 
was difficult. We believe a combination of monitor- 
ing techniques is necessary to provide a complete 
picture of predator activity and effects. We found 
that each predator survey technique had its advan- 
tages and disadvantages during our study. Although 
track trail surveys provided evidence of predator 
occurrence, unstable substrate conditions in our 
study area (e.g., unconsolidated sand and washover 
mud) enabled identification of predators from tracks 
for only short periods of time. Time-lapse photogra- 
phy of entire nesting areas provided limited evidence 
of predation, targeting specific nests may have pro- 
vided more information. Although time consuming, 
field observations of colony sites provided the most 
quantitative and qualitative data on predation and 
predator interactions with Piping Plovers and Least 
Terns. Pellet collection provided the best evidence of 
chick takes by raptors. 


Management Implications 

Our results demonstrate the importance of under- 
standing the structure and dynamics of the predator 
community in making management decisions for 
Piping Plovers and Least Terns on the Missouri 
River. We believe the high predation rates we 
observed were a direct result of sandbar habitat 
deterioration and the resulting increase in predator 
foraging efficiency. Thus, we propose habitat 
improvement as a major management goal to reduce 
predation and increase productivity of Piping Plovers 
and Least Terns. Habitat improvements to reduce 
predator efficiency should include the creation of 
large, dynamic sandbar complexes that change in 
size, location, and vegetation composition, as well as 
the presence of alternate/unused nesting substrates. 
We also recommend the application of predator 
deterrents on sites with marginal habitats that receive 
intense predator pressure. In particular, predator 
exclosure cages have the potential to increase nest 
success significantly. Because predator pressure 
appears to be site specific, application of other 
predator management techniques (e.g., predator 
removal, transplant, manipulation of alternative prey 
resources) to some areas may be justified in specific 
management situations. 


Acknowledgments 

Thanks to J. Jenks for assistance with statistical 
analysis. R. Renken and L. Flake provided earlier 
reviews of this paper. For field assistance we thank 
R. Heirmieir, J. Van’t Hull, J. Higgins, C. Kruse, G. 
Morris, M. Dorhout, and K. Pedersen. Funding for 
this project was provided by the U.S. Army Corps of 
_ Engineers (Contract Number DACW4591P1327), 
the U.S. Fish and Wildlife Service (Cooperative 
Agreement Number 14-16-0009-1549, Research 


KRUSE, HIGGINS, AND VANDER LEE: PREDATION ON PLOVER AND TERN 


485 


Work Order Number 1|7), and the South Dakota 
Department of Game, Fish and Parks through the 
South Dakota Cooperative Fish and Wildlife 
Research Unit. Cooperating agencies included the 
U.S. Fish and Wildlife Service, the U.S. Geological 
Survey, the National Biological Service, South 
Dakota Department of Game, Fish and Parks, South 
Dakota State University, and the Wildlife Manage- 
ment Institute. None of the funding or cooperating 
agencies or institutions endorse any trade names of 
equipment or supplies presented in this paper. 


Documents Cited [marked * where cited] 

Kruse, C.D. 1992. Evaluation of small plot habitat 
enhancement projects and their effects on Piping Plover 
and Least Tern productivity. Unpublished Project Report 
1992. U.S. Army COE, Omaha District Omaha, 
Nebraska. 

U.S. Fish and Wildlife Service. 1988. Great Lakes and 
Northern Great Plains Piping Plover recovery plan. U.S. 
Fish and Wildlife Service, Twin Cities, Minnesota. 

U.S. Fish and Wildlife Service. 1990. Biological opinion 
on Missouri River operations. Unpublished Report. 
1990. U.S. Fish and Wildlife Service, Denver, Colorado. 

U.S. Fish and Wildlife Service. 1991. Least Tern and 
Piping Plover annual surveys on the Missouri River. 
Unpublished Annual Report. 1991. U.S. Fish and 
Wildlife Service, Ecological Services Office, Pierre, 
South Dakota. 

U.S. Fish and Wildlife Service. 1992. Least Tern and 
Piping Plover annual surveys on the Missouri River. 
Unpublished Annual Report. 1992. U.S. Fish and 
Wildlife Service, Ecological Services Office, Pierre, 
South Dakota. 


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Received 4 April 2000 
Accepted 3 January 2002 


Population Status of Shorebirds Nesting at Churchill, Manitoba 


JOSEPH R. JEHL, JR.!, and WINLI LIN? 


'Hubbs Sea World Research Institute, 2595 Ingraham Street, San Diego, California, 92106, USA 
2717 Alder Street, Pacific Grove, California, 93950 USA 


Jehl, Joseph R., Jr., and Winli Lin. 2001. Population status of shorebirds nesting at Churchill, Manitoba. Canadian Field- 
' Naturalist 115(3): 487-494. 


We present a quantitative assessment of shorebird populations breeding in the vicinity of Churchill, Manitoba, in 1997 and 
compare it with qualitative data amassed since 1930. Our study was based on extensive ground surveys, supplemented by 
data from long-term studies of several individual species. Over the past seven decades the status of most local shorebirds 
has changed importantly, and species that were once abundant (Semipalmated Sandpiper, Red-necked Phalarope) have 
almost vanished. Currently, American Golden-Plover, Whimbrel, Semipalmated Plover, and Dunlin predominate. Future 
surveys at approximately 10-year intervals are warranted to maintain this exceptional long term record of birdlife in the 


subarctic. 


Key Words: Shorebirds, populations, subarctic, Churchill, Manitoba. 


Areas bordering the Hudson Bay coast provide 
important nesting habitats for a large variety of shore- 
birds and staging sites for other species migrating to 
or from more northerly destinations. The Churchill, 
Manitoba, area (58° N, 94° W) has played an espe- 
cially prominent role in developing information about 
shorebird biology because of its easy access and the 
abundance and diversity of the 15 breeding species, 
many of which have been studied in some detail 
locally: American Golden-Plover (Byrkjedal and 
Thompson 1998; scientific names given in species 
accounts), Semipalmated Plover (Nol and Blanken 
1999), Whimbrel (Skeel 1976, 1983; Lin 1997), Stilt 
Sandpiper (Jehl 1970; Klima and Jehl 1998), Least 
Sandpiper (Jehl 1970), Short-billed Dowitcher (Jehl 
et al. 2001), Hudsonian Godwit (Hagar 1966; Elphick 
and Klima in press), Semipalmated Sandpiper 
(Gratto-Trevor 1992), Red-necked Phalarope 
(Reynolds 1987), and Dunlin (Jehl, unpublished). 

Churchill’s importance became apparent in 1934 
with the publication of Taverner and Sutton’s classic 
The Birds of Churchill, Manitoba. Over the ensuing 
seven decades, however, the status of many species 
changed so greatly that the standard literature 
(Taverner and Sutton 1934; Jehl and Smith 1970) is 
out of date. Our goal in this paper is to quantify the 
numbers of shorebirds breeding in the immediate 
Churchill area to provide a baseline for further stud- 
ies. Such information is salient because Churchill 
has a long ornithological history (Jehl and Smith 
1970), is well studied, and is easily accessible, and 
thus is an important area in which to conduct long 
term monitoring studies. Shorebirds are important 
because they are a dominant component of high lati- 
‘ude avifaunas and tend to nest in or near coastal 
wetlands, the very areas that will be most affected by 
changes in moisture regimes or sea level that are pre- 
dicted to accompany global warming. 


There has been no previous effort to produce a 
comprehensive assessment of local shorebird popu- 
lations, although there are incomplete data for sever- 
al species (e.g., Skeel 1983; Gratto-Trevor 1993/94). 
To put our findings in perspective we compared 
them with estimates of relative abundance provided 
by, or interpreted from, earlier reports. 


Study Area and Methods 

In June 1997 we surveyed approximately 7000 ha 
in the immediate Churchill area for shorebird nests 
and territories. The study area extended from the 
coast of Hudson Bay from the southern edge of the 
Churchill townsite east to Gordon Point, and inland 
to varying distances south to Landing Lake and the 
large fen several kilometers north of Twin Lakes 
[Figure 1A; for detailed maps and place names see 
Chartier (1994)]. It contains a variety of habitats 
used by shorebirds including hummock bog, dry and 
wet sedge meadows, dry and wet tundra, and gravel 
coastal beaches (and their anthropogenic equivalents 
— the shoulders of roads and borrow pits). 

We divided the study area into five sections: 

Section 1: Extends from the southern limits of the 
townsite to the airport, north to the coast and south 
to Landing Lake. The habitat between town and the 
start of Goose Creek Road is primarily well-drained 
meadow (much modified by humans since 1930), 
whereas that along the Landing Lake road is wet 
sedge meadow and hummock-bog. 

Section 2: From the dump, north to the coast, east 
to the Auroral Observatory (“Golfballs’”), and south 
of the Launch Road on the Restricted Access Road 
to the end of the airport runway. Includes two long- 
term study plots, one established in 1964 (36 ha), the 
other in 1992 (40 ha). The habitat is primarily sedge- 
meadow, with dry areas predominating. 

Section 3: From the Auroral Observatory east to 


487 


488 THE CANADIAN FIELD-NATURALIST 


Bird Cove Road, north to the coast and south to 
Stygge Lake. Includes two plots, one established in 
1965 (72 ha), and one in 1993 (40 ha). Habitat 
along the Launch Road is primarily heath-tundra, 
rocky ridge and dry plains, with localized patches 
of wet sedge meadow between the road and Stygge 
Lake. 

Section 4: From Bird Cove Road east to 
Churchill Northern Studies Centre (CNSC) and 
Gordon Point, and southward through the Spruce 
Ridge and Camp Nanuk areas. The habitat is main- 
ly dry tundra with scattered sedge meadow. From 
about 3 km east of CNSC, we concentrated obser- 
vations between the coast and 2 km inland, princi- 
pally along the main tracks leading to Gordon Point 
but also including ali of the two major peninsulas 
that project into Hudson Bay. Other observations 
were made along the major trails that lead south 
onto Christmas Lake Esker. 

Section 5: Starts approximately 4 km south of 
CNSC, extends 1-4 km east and west of the Twin 
Lakes Road, and continues southward through to the 
large Twin Lakes fen, which includes a 64-ha plot 
established in 1992. Habitat east of the road is pri- 
marily wet sedge meadow, to the west hummock- 
bog. Areas of continuous stands of boreal forest 
were excluded. 

Surveys were made by WL and JRJ (occasionally 
assisted by other biologists) walking approximately 
100 m apart on parallel transects through all suitable 
habitat and recording the presence of territorial birds 
and nests. Transect data were obtained between 7 
and 22 June, by which time local breeders had estab- 
lished territories or were on eggs. For most species 
the peak of hatching occurred 28 June—5 July (earli- 
est 26 June). Because of the large distances involved, 


Vol. 115 


it was not possible to replicate transects. This was 
not a problem because in previous seasons (Jehl 
1964-1967, 1977, 1991-1996; Lin 1994-1996), we 
had already amassed detailed information on the dis- 
tribution and status of shorebirds. This allowed us to 
concentrate transects work in good habitat and mini- 
mize efforts in dry upland areas that support few or 
no shorebirds. Highly productive areas, however, 
were visited repeatedly—some almost daily—from 
early June into mid-July, either because they con- 
tained long term census plots or were under special 
study by us or other researchers. 


Results 

Of the 15 shorebird species nesting near 
Churchill, 13 occur in the survey area (Table 1). 
Two others, Spotted Sandpiper (Actitis macularia) 
and Solitary Sandpiper (Tringa solitaria), are rare 
and nest locally around lakes or in spruce woods 
farther inland. 


American Golden-Plover (Pluvialis dominica). 
Golden-Plovers nest from the south end of the town- 
site (scarce) and eastward along the coast, and inland 
to the Twin Lakes fen and about | km north of 
Landing Lake. Most use well drained coastal tundra 
or dry areas the sedge marshes, although in recent 
years increasing numbers have been found in wetter 
situations. The species is conspicuous and territories 
are easily detected. We found 94 territories, which 
constitute about 90% of the population to the west 
and south of CNSC. While our studies to the east 
were less thorough, shorebirds are so scarce there 
that we cannot have overlooked any significant num- 
ber of plovers. We estimated the total population at 
110-120 pairs. 


TABLE 1. Territorial pairs of shorebirds (and nests) found in Churchill area (see Figure 1) 1997. 


Section! 

Estimated 
Species 1 2 e) + 5) Total Population? 
American Golden-Plover 23°) 9 (1) 10 (1) 40 (30) 10 (6) 94 (43) 110-120 
Semipalmated Plover 2Pi(22) Bi) CD) 46 (46) 0 (0) 12s) 80 
Killdeer 0 (0) 1 (O) 0 (0) 2, le) 0 (0) a9 4-5 
Lesser Yellowlegs 9 (0) 0 (0) 2 (0) 3 (0) 3 (2) Weg 40 
Whimbrel 29 (9) 8 (2) 15 (1) 15 (14) 28 (14) - 95 (40) 105-110 
Hudsonian Godwit 18 (0) TAGy) 6 (1) 0 (0) 3G) 36 (5) 50-55 
Semipalmated Sandpiper 0 (0) 0 (0) 2 (2) 0 (0) 6 (1) 8 (3) 10 
Least Sandpiper 8 (6) (3) 10 (4) 3 (1) 8 (3) 36 (17) 80-100 
Dunlin 3 (2) 26 (10) 17 (9) - Abe Ae) 19 (18) 76 (50) 80-100 
Stilt Sandpiper 8 (5) 9 (1) 1) 2 (0) 6 (4) 36 (12) 50-60 
Short-billed Dowitcher 122) 3 (0) Sn) 1 (0) 3: (3) 24 (6) 50-60 
Common Snipe 13 (0) 3 (0) 2 (0) 0 (0) 2 (0) 20 (0) 40 
Red-necked Phalarope 2 (0) 8 (0) 2 (1) 6 (1) 1 (1) VAMR)) 30-40 
Total 149 (51) 84 (20) 83 (23) 129(104) 91(55) 398.4253) 719-820 


'For locations of Sections see Figure 1. 


2See text for details. 


2001 JEHL AND LIN: SHOREBIRDS NESTING AT CHURCHILL, MANITOBA 489 


FicurE 1. A The Churchill, Manitoba, area showing the five areas censused for nesting shorebirds in 1997. Dark areas 
are lakes. Areas studied (cross-hatched) by earlier investigators: B Taverner and Sutton in 1930-1933, and 
Grinnell and Palmer in 1940; C Allen in 1944; D Breckenridge et al. in 1954; E Jehl in 1964-1967. 


490 


Semipalmated Plover (Charadrius semipal- 
matus). This conspicuous species nests almost exclu- 
sively in very dry habitats, typically coastal beaches 
and adjacent lichen tundra, and occasionally the 
gravel shoulders of roads. Studies from 1988 through 
1996 emphasizing coastal sites indicated a popula- 
tion of 45-55 pairs (E. Nol, personal communica- 
tion). Our broader surveys detected 72 territories and 
71 nests, the majority on the beaches in the eastern 
part of Section 1 (mainly the gravel extraction plant 
below old Fort Churchill) and in Section 4 (Halfway 
Point). Very few birds (< than 3% of the population) 
breed inland (E. Nol, personal communication). We 
estimated about 80 pairs in the survey area. A few 
others nest in the townsite and along the flats of the 
Churchill River. 


Killdeer (Charadrius vociferus). We found three 
territories, including one nest. Other studies 
(1991-1998) indicated that the local population 
averages 4—5 pairs (up to 6-8 pairs in some years; 
E. Nol, personal communication). Killdeers nested 
usually in disturbed situations (e.g., roadsides, 
dumps), occasionally on undisturbed tundra. An 
additional 1—2 pairs nested in the townsite (not sur- 
veyed). 


Lesser Yellowlegs (Tringa flavipes). Yellowlegs 
are fairly common in brushy or forested areas, but 
only rarely breed in the open habitats we studied. 
Most territories were near the forest edge around 
Landing Lake and Twin Lakes. A few pairs were 
scattered through the wooded areas (not surveyed) 
along the Twin Lakes Road between Sections 4 and 
5. Yellowlegs that forage in the Twin Lakes fen do 
not breed there but in nearby wooded areas. 

With obvious flight displays and noisy and per- 
sistent alarm calls, yellowlegs are hard to overlook. 
Our count of 14 pairs in Sections 1-4 is probably 
80% accurate; data for Section 5 are less good. We 
estimated 30—40 pairs in the area surveyed. We 
cannot make a confident estimate of the total popu- 
lation because we did not survey forest edge situa- 
tions farther inland, as at Twin Lakes or along 
Goose Creek. 


Whimbrel (Numenius phaeopus). Whimbrel are 
widespread, especially in wet sedge meadows or in 
hummock-bogs near the treeline; few breed on the 
coastal tundra or east of CNSC. Territories are easy 
to detect because of the birds’ noisy flight displays 
and propensities for chasing aerial predators 
(ravens, gulls, jaegers) that approach their nest. We 
determined 95 territories and found 40 nests. Given 
the Whimbrel’s conspicuous behavior and Lin’s 
detailed studies in 1994-1996 (Lin 1997), we are 
confident of having found at least 90% of the terri- 
tories and estimate the local population at about 
105-110 pairs. 


Hudsonian Godwit (Limosa haemastica). Hud- 


THE CANADIAN FIELD-NATURALIST 


sonian Godwits breed most commonly in sedge | 


meadows near the treeline. The general location of 
territories is easy to determine by watching flight 
displays early in the season. By mid-season the birds 


Semipalmated Sandpiper (Calidris pusilla). In 
1993-1996, the entire population west of CNSC con- 
sisted of 5—9 pairs, all in Section 3 (Jehl unpub- 
lished). When the nesting area flooded in 1997, only 
two pairs bred. At Gordon Point (Section 4) we 
found about five pairs, all restricted to the outermost 
beach. Between CNSC and Gordon Point the species 
is virtually absent (one pair). 


Least Sandpiper (Calidris minutilla). Least 
Sandpipers are hard to census because they do not 
display as conspicuously as some other species and 
sit tightly on nests. Based on our experience in find- 
ing nests in intensively studied areas in good habitat, 
we estimate that combined study plot/transect proce- 
dures in 1997 located 40-50% of the population (36 
territories, 17 nests), which would total about 
80-100 pairs. 


Dunlin (Calidris alpina). Intensive studies (JRJ 
unpublished) through the 1990s delineated the local- 
ized breeding distribution of the Dunlin. Nests can 
be found fairly easily, and territories are evident 
from the flight displays of males, which often 
involve other species. From 199i—1996 Jehl estimat- 
ed the local population at about 80 pairs; we estimat- 
ed 76. Very few can remain undetected. 


Stilt Sandpiper (Calidris himantopus). Stilt Sand- 
pipers have been studied locally since the 1930s 
(Jehl 1970, summarized in Klima and Jehl 1998). 
Flight displays involving several individuals or other 
species may be continued rather late in the season, 
making it easy to locate territories. In 1997, as in 
other recent years, Stilt distribution was patchy. Our 
finding of 36 territories (12 nests) agreed with other 
estimates in the 1990s (Jehl unpublished), which 
indicated a population of 50-60 pairs. We did not 
find this species in Section 4 east of CNSC. 


Short-billed Dowitcher (Limnodromus griseus). 
Dowitchers nest in wet sedge meadows, occasional- 
ly north to the Launch Road. Males do-not sing as 
much or as long as other species, making territories 
harder to detect, and after nesting starts dowitchers 
become even less conspicuous; nests are found only 
by accident. We found 24 territories and six nests. 
Although occurring in all areas, dowitchers are 
commonest in Section 1, where we probably detect- 
ed only about half of the population. Counts in 
Sections 2-5, however, are probably accurate to 
within 25%. We estimated the population at 50-60 
pairs. 


Vol. 115 


become less obvious. We located 36 territories (but — 
only five nests, which are extremely difficult to find) | 
and estimate our success at about 70%. 


JEHL AND LIN: SHOREBIRDS NESTING AT CHURCHILL, MANITOBA 49] 


2001 


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492 


Common Snipe (Gallinago gallinago). The distri- 
bution of snipe was quite local. We found them 
mainly in the deeper marshes near willow thickets 
along the treeline; a few pairs occurred north to the 
Hudson Bay shore (Bird Cove). Because snipe have 
crepuscular displays, they cannot be censused reli- 
ably by our techniques, and we found no nests. We 
identified 20 territories, of which 13 were in Section 
1, and estimated 40 pairs in the survey area. 


Red-necked Phalarope (Phalaropus lobatus). 
Phalaropes are scarce and local, usually nesting in 
pond habitats in association with Arctic Terns 
(Sterna paradisaea). In Sections 1—3 and 5 we found 
only 13 pairs. That figure may be slightly conserva- 
tive, because a few pairs sometimes breed in areas of 
the large Akudlik Marsh (Section 1) that we could 
not study in detail because of its designation as a 
sanctuary. In Section 4, 2—3 pairs nested near Bird 
Cove and perhaps 6—10 pairs at Gordon Point, which 
we could only survey briefly. We doubt that there 
are more than 30—40 pairs in the survey area. 
Otherwise we know only of several pairs that some- 
times nest on the townsite ponds and 1-2 pairs at 
West Twin Lake. 


Discussion 

Approximately 800 pairs of shorebirds (slightly 
greater than 0.1 pair ha!) nested in the study area in 
1997. That total, and the estimate by species, is con- 
sistent with other data obtained by JRJ throughout 
the 1990s. Historical data are scarce and allow only 
qualitative comparisons, but several kinds of evi- 
dence including (1) writings of early naturalists (e.g., 
Taverner and Sutton 1934; Grinnell and Palmer 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


1941; Jehl and Smith 1970; and others below), (2) 
large specimen holdings in many museums (Jehl per- 
sonal observations), and (3) data from plots censused 
in the 1960s and 1990s (Jehl unpublished) leave no 
doubt that shorebirds are far scarcer than they were 
in the 1930s and 1960s. The dearth is so obvious that 
visiting birders often inquire “where are the shore- 
birds?” — a question that would have been ludicrous 
only three decades ago. For instance, the Semi- 
palmated Sandpiper, once the most abundant species 
(Taverner and Sutton 1934; Allen 1945), is virtually 
extirpated. The Least Sandpiper, locally fairly com- 
mon in the 1960s, occurs in greatly reduced num- 
bers. And observations that one or more Red-necked 
Phalaropes could be found on every lakelet or pool 
(Taverner and Sutton 1934: 49), or that Stilt Sand- 
pipers nest “quite commonly” (Farley 1936), “with 
“scores” displaying at the same time (Sutton 1961), 
are so contrary to current status that future ornitholo- 
gists may find these impressions hard to reconcile. 
On the other hand, Hudsonian Godwits, thought to 
be on the verge of extinction as late as the 1940s, 
have increased, and the Golden-Plover has gone 
from being one of the least common species to per- 
haps the commonest. 

Some of these differences can be appreciated in 
the writings of ornithologists who have commented 
on general status (Table 2), or have either ranked rel- 
ative abundance or provided enough information that 
we have tried to do so (Table 3). No two studies are 
fully comparable because no two involved the same 
localities or habitats (Figure 1 B-E). The rankings of 
Breckenridge et al. (1954) are not very informative 
because that party was based near the treeline at 


TABLE 3. Ranking of relative abundance of shorebirds at Churchill, Manitoba. 


Taverner 

and Sutton Allen Breckenridge Jehl and Smith. This 
Species (1934)! (1945) et al. (1954) (1970)4 paper 
American Golden-Plover 11 — 11 1 
Semipalmated Plover 2 3 2 + 5 
Killdeer 12 11 13 12 
Lesser Yellowlegs 6 3 2 95 
Whimbrel 4 1 3 2 
Hudsonian Godwit 13 6 10 8 
Semipalmated Sandpiper 1 1 - 9 13 
Least Sandpiper 5) 2 ul 1 3 
Dunlin 5 ~ 8 3 
Stilt Sandpiper + 8 10 2) 6 
Short-billed Dowitcher 10 8 6 6 
Common Snipe 7 4 7 10 
Red-necked Phalarope 3 3) 11 ae 


‘Based on our interpretation of Taverner and Sutton 1934. 


2Allen ranked Spotted Sandpipers (not treated herein) as 13, Hudsonian Godwit 14. 
3Breckenridge et al. inexplicably ranked Sanderling as 9. That species does not breed within hundreds of miles 


of Churchill. 


‘Based on Jehl and Smith 1970, and unpublished field notes. 
SRelative abundance underestimated because survey areas did not include major habitat. 


2001 


~ 


Landing Lake, where many species are absent. Jehl’s 
observations from the 1960s (summarized in Jehl 
and Smith 1970) provide the best basis for compari- 
son, as they encompassed nearly all the areas 
searched by earlier workers, as well as most of the 
1997 survey area; the exceptional areas were Section 
5 and the tundra east of CNSC (Section 4), to which 
access was then severely limited owing to military 
operations. 

In general, it appears that the relative abundance 
of most species was fairly stable from the 1930s 
through the 1960s, but that important changes took 
place over the following three decades. The causes 
are probably complex and vary by species. Many 
breeding areas that were productive in the 1960s 
now hold few shorebirds, having been heavily 
modified by overgrazing geese (Abraham and 
Jeffries 1997; Jehl unpublished). In other cases 
habitat changes are surely involved, particularly 
near areas of human habitation. Yet the declines in 
some species are not simply local phenomena. 
Semipalmated Sandpipers and Red-necked 
Phalaropes have declined at La Pérouse Bay, 
20-25 km east of CNSC (Gratto-Trevor 1993/94). 
Other population effects likely involve conditions 
on wintering grounds or migration routes. 
Censuses made every decade or so may resolve the 
relative importance of local vs. regional factors 
affecting shorebird numbers. Such data are impor- 
tant because of indications that shorebirds are 
undergoing severe declines (Morrison et al. 1994; 
Morrison 2001). They will also help maintain and 
extend what may be the best long-term record of 
birdlife anywhere in the subarctic (Houston et al. 
2002; Jehl unpublished). 


Acknowledgments 

This study received major support through a 
grant to W. Lin from the Churchill Northern 
Studies Centre. It is an outgrowth of long-term 
studies that have been supported by Hubbs- 
SeaWorld Research Institute (through J. and T. 
Morris, and the Joan Irvine Smith and Athalie R. 
Clarke Foundation), the National Geographic 
Society, and the Canadian Wildlife Service. The 
success and thoroughness of this survey was 
enhanced by the contributions of many field col- 
leagues, especially J. Williams, K. Krijgsveld, R. 
Ricklefs, E. Nol, and Y. Zharikov, and the gra- 
Ciausness of LL. Turgeon, J. Wilson and D. 
Bellerive, who facilitated access to restricted areas. 
Our efforts were greatly enhanced by researchers 
from Trent University (2-3 persons), who were 
engaged in a long-term study (1988-present) of 
Semipalmated Plover populations, and from the 
University of Missouri and the University of 
Groningen (6-8 persons), who were gathering 
detailed information on nesting distribution of all 


JEHL AND LIN: SHOREBIRDS NESTING AT CHURCHILL, MANITOBA 


493 


species in Sections 4 and 5. E. Nol, C. Gratto- 
Trevor, G. Morrison, and A. J. Erskine provided 
helpful comments on a draft manuscript. 


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Nol, E., and M.S. Blanken. 1999. Semipalmated Plover 
(Charadrius semipalmatus). In The Birds of North 
America number 444. Edited by A. Poole and F. Gill. 
The Birds of North America, Inc., Philadelphia, 
Pennsylvania. 

Reynolds, J. D. 1987. Mating system and nesting biology 
of the Red-necked Phalarope Phalaropus lobatus: what 
constrains polyandry. Ibis 129: 225-242. 

Skeel, M. A. 1976. Nesting strategies and other aspects of 
the breeding biology of the Whimbrel (Numenius phaeo- 


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Vol. 115 


pus) at Churchill, Manitoba. M. Sc. thesis, University of 
Toronto. 

Skeel, M. A. 1983. Nesting success, density, philopatry 
and nest-site selection of the Whimbrel (Numenius 
phaeopus) in different habitats. Canadian Journal of 
Zoology 61: 218-225. 

Sutton, G.M. 1961. Stilt Sandpiper. Pages 340-346. In 
The birds of the British Isles, Volume 9. Edited by D. A. 
Bannerman). Oliver & Boyd, London. 

Taverner, P. A., and G. M. Sutton. 1934. The birds of 
Churchill, Manitoba. Annals of the Carnegie Museum 
23: 1-83. 


Received 14 December 2000 
Accepted 24 August 2001 


Grizzly Bear, Ursus arctos, Usurps Bison Calf, Bison bison, Captured 
by Wolves, Canis lupus, in Yellowstone National Park, Wyoming 


DANIEL R. MACNULTY!, NATHAN VARLEY2, and DOUGLAS W. SMITH? 


'Department of Ecology, Evolution and Behavior, University of Minnesota, St. Paul, Minnesota 55108 USA 
2Symbiosis Consulting, P.O. Box 490 Gardiner, Montana 59030 USA 
3Yellowstone Center for Resources, P.O. Box 168, Yellowstone National Park, Wyoming 82190 USA 


MacNulty, Daniel R., Nathan Varley, and Douglas W. Smith. 2001. Grizzly Bear, Ursus arctos, usurps Bison calf, Bison 
bison, captured by Wolves, Canis lupus, in Yellowstone National Park, Wyoming. Canadian Field-Naturalist 
115(3): 495-498. 


We describe an adult Grizzly Bear (Ursus arctos) usurping a Bison (Bison bison) calf from a pack of five Wolves (Canis 
lupus) attempting to kill the Bison in Yellowstone National Park during early spring. Five Wolves grabbed the hind end 
and neck of the calf while it was trailing behind two adult male Bison. In 3 minutes a Grizzly Bear arrived and displaced 
the two Wolves attacking the hind end. For 1 minute the Grizzly Bear attacked the rear of the Bison while three Wolves 
attacked the front end. The Grizzly Bear subsequently pulled the struggling calf from the Wolves and made the kill. The 
Wolves were unable to displace the Grizzly Bear from the carcass. Our observation demonstrates the capacity for Grizzly 
Bears to exploit the predatory abilities of Gray Wolves restored to Yellowstone National Park. Kleptoparasitism by Grizzly 
Bears on Wolf-captured ungulates may be a selective pressure promoting group living in Wolves, and could provide an 


important new food resource to threatened Grizzly Bears in the Greater Yellowstone Ecosystem. 


Key Words: Grizzly Bear, Ursus arctos, Wolf, Canis lupus, Bison, Bison bison, predation, kleptoparasitism. 


Wolves (Canis lupus) and Grizzly Bears (Ursus 
arctos) commonly interact in the defense of offspring 
or in competition for carcasses (Murie 1944; Ballard 
1982; Hornbeck and Horejsi 1986; Hayes and Mossop 
1987; Hayes and Baer 1992; Servheen and Knight 
1993; Kehoe 1995; Mech et al. 1998). Lent (1964) 
reported tolerance between a Grizzly Bear and a Wolf 
feeding simultaneously on a Caribou (Rangifer taran- 
dus) carcass. During competition for carcasses, Murie 
(1944:204) observed that in general, “the Wolves are 
the losers and the meat-hungry bears are the gainers”. 
The ability of Grizzly Bears to contest carcasses suc- 
cessfully could be a factor favoring the evolution of 
group living in Wolves, and may be an important con- 
sideration in the conservation of Grizzly Bears in the 
Greater Yellowstone Ecosystem (GYE). Here we 
describe the first observation of a Grizzly Bear usurp- 
ing a Bison captured by Wolves. 

The Wolf-Grizzly Bear interaction was observed 
near Pelican Valley, Yellowstone National Park, 
Wyoming (44° 38’ N, 110° 13’ W) on 24 March 
2000. The interaction occurred in an open creek 
drainage bordered by forests composed primarily of 
Lodgepole Pine (Pinus contorta). The area lies at 
2432 m and is subjected to long, cold winters, with 
snow thickness from 33-160 cm. To survive in 
Pelican Valley during winter, Bison rely upon an 
archipelago of snow-free and nearly snow-free areas 
created by geothermal activity (Meagher 1976). 
Thermal features (hotsprings, fumaroles, etc.) and 
datches of thermally influenced warm ground are 
scattered throughout the area. South-facing and 
wind-blown slopes provide additional habitat for 


bison. Winter forage includes various species of 
sedge (Carex spp.) and grass (Meagher 1973). 

From a prominent hilltop, we observed the Wolf- 
Grizzly interaction using a 56X Nikon spotting 
scope from a distance of 6.0 km. Observations were 
timed with a digital stopwatch and recorded on a 
portable voice recorder. The pack of five adult 
Wolves first encountered the herd of 59 Bison on 23 
March at 09:08, and a Grizzly Bear first appeared at 
12:31 while running toward the Wolves as they 
made a failed attempt to kill a Bison. In each of two 
subsequent attacks observed that day, the Grizzly 
Bear ran immediately behind or alongside the pursu- 
ing Wolves. Two additional adult-size Grizzly Bears 
appeared traveling together toward the Wolves at 
18:38. Subsequent play behavior suggested that the 
bears were a two-year-old sibling pair. 

The next day, 24 March, 12:13-18:19, 1-5 (x = 3) 
Wolves made a series of 14 attacks, alternately 
attacking and resting, on a Bison group containing 
nine adult/juvenile bulls and one 11-month-old calf 
of unknown sex. The group of 10 Bison had 
remained in the area while the remainder of the herd 
fled when the pack pursued a single unidentified 
Wolf approaching the Bison at 11:19. The unidenti- 
fied Wolf was not wearing a radio-collar and was 
presumed to be either an unrelated trespasser or an 
unwelcome relative. The pack mobbed the unidenti- 
fied Wolf, but allowed it to escape. 

Wolf attacks on Bison lasted 2—11 minutes (x = 6 
minutes), and involved Wolves lunging at Bison 
while they stood grouped on a thermal feature 
(n= 12) or running single file through deep snow 


495 


496 


between thermal features (n=2). Wolves targeted 
the calf during all attacks. In four attacks 1-2 wolves 
grabbed the calf and inflicted visible damage to the 
rump and flanks but failed to make the kill. In all but 
the final attack 1—5 bulls (x = 2) defended the calf by 
charging and kicking at the Wolves. A bull success- 
fully kicked a Wolf in only two attacks. 

During Wolf attacks Grizzly Bears watched at a 
distance greater than 15m (n= 2), approached and 
watched from less than 15 m (n = 2), approached and 
walked the perimeter of the attack area (n= 3), and 
approached and grabbed prey (n = 1). When walking 
the perimeter of the attack area Grizzly Bears often 
displaced resting Wolves not participating in the 
attack. Grizzly Bears were not visible in the remain- 
ing six attacks due to vegetation and topography; 
however, their ongoing presence was inferred when 
visible between attacks. For example, at 15:07, dur- 
ing a pause between the fifth and sixth attacks, one 
grizzly reappeared from behind a screen of trees and 
rushed the wounded calf, which stood in the center 
of nine resting bulls. The grizzly halted and walked 
back behind the trees when the nine bulls stood up 
and gathered around the calf. 

The final attack was preceded by a 3.5 hour stand- 
off between Wolves and Bison on a thermal feature 
at the foot of a small snow covered ridge. During 
this period Wolves made periodic attempts to grab 
the calf but were repelled each time by the bulls. The 
standoff ended when seven bulls walked away from 
the thermal feature and ascended the ridge, leaving 
two bulls to defend the calf. After a failed attempt to 
capture the calf, the Wolves left the thermal feature 
at 18:05 and trotted toward the seven bulls. 
Meanwhile, one Grizzly Bear appeared and began 
walking toward the two bulls and calf. 

For clarity the events involving the final attack are 
listed below chronologically. 

18:14 The two bulls, followed closely by the calf, 
walked away from the thermal feature and 
began ascending the ridge single file toward 
the seven bulls and five resting Wolves. The 
grizzly continued to approach the three 
Bison. 

Sighting the oncoming Bison the pack arose 
and approached the two bulls and calf from 
the west while the grizzly approached from 
the east. The calf moved with difficulty 
through the snow and trailed 10-15 m 
behind the two bulls. 

All five Wolves walked past the two bulls 
and four Wolves grabbed the caif; three 
grabbed the hind end and one grabbed the 
flank. Seconds later a fifth Wolf grabbed the 
neck. The two bulls continued walking and 
joined the seven other bulls. 

The grizzly continued to approach from the 
east, nearing the calf and Wolves. 


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18:20 


THE CANADIAN FIELD-NATURALIST 


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18:23 


18:24 


18:26 


18:41 


18:53 


18:54 


18:58 


19:00 


Vol. 115 


Three Wolves released the calf and charged 
the grizzly while the remaining two Wolves 
continued to grab the hind end of the calf. 
As the Wolves started to lunge at the grizzly 
it rose to its hind legs, quickly fell back to 
all four feet, and fled with the three Wolves 
in pursuit. Within seconds, the grizzly 
turned, ran past the pursuing Wolves, and 
back to the calf. 

The grizzly forced the two Wolves from the 
calf’s hind end and began to attack the calf 
with its forepaws. The two Wolves, joined 
by a third, then grabbed the front end of the 
calf. 

Three Wolves bit the front end of the calf 
while the grizzly swatted at the hind end. 
The grizzly pulled the calf from the Wolves, 
dragged it to the ground, and began to feed. 
The Wolves did not contest the carcass any 
longer. Four Wolves disappeared over the 
ridge while one Wolf stood by and watched 
the grizzly feed. 

Two Wolves returned from behind the ridge 
and rested along side the other Wolf approx- 
imately 5 m from the carcass and the feeding 
bear. Meanwhile, the nine bulls began to 
walk single file back along the ridge toward 
the thermal area, the feeding grizzly, and the 
resting Wolves. 

As the bulls approached to within 5 m, the 
grizzly abandoned the carcass and disap- 
peared over the ridge. Nearly simultaneous- 
ly, two Wolves reappeared, and all five 
Wolves repossessed the carcass and started 
feeding. The first bull to run past the carcass 
briefly displaced the Wolves. Once the bull 
was past the Wolves returned and continued 
to feed. Five more bulls ran past the feeding 
Wolves singly or in pairs. Two Wolves 
briefly chased a pair of Bison as they ran 
past and then returned to the carcass. 

The pair of sibling grizzlies appeared at the 
thermal area and began to approach the car- 
cass and the feeding Wolves. 

The two grizzlies veered and began walking 
away from the carcass. 

The last bull, which was also the smallest, 
ran past the feeding Wolves. Again, two 
Wolves briefly pursued the bull and returned 
to the carcass. 

The single grizzly returned to the carcass 
from behind the ridge, displaced all five 
Wolves, and resumed feeding. Two Wolves 
disappeared over the ridge immediately, 
while three Wolves remained standing 
around the carcass, watching the grizzly 
feed. 


2001 


19:06 The single grizzly continued to feed on the 
carcass without interruption until observa- 
tion ended due to darkness. 


Initial observation at 07:00 the following day 25 
March, found one grizzly feeding on the carcass, 
while three Wolves were sleeping 25 m away. The 
grizzly fed until 11:51, and then walked over the 
ridge and out of view. During this period the Wolves 
approached the grizzly to within 5 m but did not 
actively contest the carcass. Once the grizzly left the 
carcass three Wolves returned and fed until at least 
12:29, at which time the carcass was pulled over the 
ridge and out of view. 

The Wolves’ failure to seriously challenge the 
grizzly once the Bison was usurped may have been 
influenced by the tearing of the carcass into two or 
more parts during the struggle for possession. This 
explanation is supported by our inspection of the kill 
site, which revealed the remains of a front and rear 
leg over the ridge where the Wolves and the grizzly 
disappeared. It is possible that the grizzly fed on 
these parts when it was displaced from the carcass 
by the nine passing bulls. Likewise, when the grizzly 
was feeding on the carcass and only one to three 
Wolves were visible, the remaining Wolves were 
probably feeding on these same parts. Based on 
examination of two mandibles found at the kill site 
the age of the Bison was confirmed to be 11 months. 

The sequence of events that led to the Grizzly Bear 
usurping the Bison calf from the Wolves began with 
the unusual association between the calf and the 
bulls. While bulls are known to defend calves from 
Wolves (Carbyn and Trottier 1988; Carbyn et al. 
1993), the occurrence of a calf within a Bison group 
composed exclusively of bulls has not been previous- 
ly reported for a wild Bison population. Bison calves 
are usually found in large mixed herds containing 
females of all ages, yearlings, and most males two to 
three years old (Meagher 1986). Explanations include 
(1) random association, (2) calf was orphaned, and 
(3) if a male, the calf deliberately joined the bull 
group. The cow-calf bond breaks earlier for males 
than for females (Lott and Minta 1983; Green et al. 
1989), and calves not associating with a cow are sub- 
ject to more aggression than calves with cows 
(Coppedge et al. 1997). Frequent feeding displace- 
ment during early spring food scarcity (Rutberg 
1986) may have led the 11-month-old calf to join the 
smaller bull group. 

Kleptoparasitism by Grizzly Bears and other scav- 
engers on Wolf-captured ungulates could be a factor 
favoring the evolution of group living in Gray 
Wolves. In African Wild Dogs (Lycaon pictus) klep- 
toparasitism by Lions (Panthera leo) and Spotted 
Hyenas (Crocuta crocuta) is considered an important 
factor in the evolution of group living (Fanshawe 
and FitzGibbon 1993; Creel and Creel 1996; 
Carbone et al. 1997). Wolves might respond to klep- 


MACNULTY, VARLEY, AND SMITH: BEAR Usurps BISON CALF 


497 


toparasitism from Grizzly Bears by increasing group 
size and thereby increase access time at carcasses. 
Future studies should attempt to assess whether larg- 
er packs of Wolves are more successful at defending 
carcasses from Grizzly Bears and whether increases 
in access time fully compensate for reductions in per 
capita intake due to larger pack size (Schmidt and 
Mech 1997). Interestingly, the Wolves reported in 
this observation rejected an opportunity to increase 
the size of their group when they ran off an unidenti- 
fied Wolf, suggesting that factors unrelated to prey 
capture or carcass defense may be more important to 
group formation. 

While an earlier assessment predicted that Wolves 
restored to Yellowstone National Park would not 
affect the Yellowstone Grizzly Bear population 
(Servheen and Knight 1993), the capacity for Grizzly 
Bears to usurp Wolf-killed ungulates suggests the 
potential for a positive effect. Wolf-killed ungulates 
may be an important new high-quality food resource 
for Grizzly Bears in the GYE, where the future of 
many traditional Grizzly Bear foods is uncertain 
(Mattson and Reid 1991). An increase in the avail- 
ability of meat to Grizzly Bears could positively 
influence several population parameters including 
adult female body mass, litter size, and population 
density (Hilderbrand et al. 1999). 

Whether Yellowstone Grizzly Bears realize a 
population level benefit from Wolf-killed ungulates 
may largely depend on seasonal and demographic 
variation in access to Wolf-killed ungulates. 
Grizzly Bear population densities have been found 
to correlate positively with fall meat availability, in 
part because meat consumed in fall is more impor- 
tant for successful cub production and hibernation 
than meat consumed in spring (Hilderbrand et al. 
1999). Fall is also a period when Wolf persistence 
at kills may be greater due to lower kill rates com- 
pared to spring (Smith unpublished), thus potential- 
ly limiting grizzly access to Wolf-killed ungulates. 
The ability of Grizzly Bears to usurp and maintain 
possession of a Wolf-killed ungulate may also be 
affected by Grizzly Bear age, sex, and reproductive 
status. Specifically, females with offspring or 
smaller bears of either sex may be less capable than 
adult males in usurping a carcass from Wolves, and 
subsequently keeping the carcass from other griz- 
zlies. Adult male grizzlies are more aggressive than 
other age and sex classes and tend to dominate 
localized high-quality food resources (Craighead et 
al. 1995). As a result, adult male grizzly dominance 
at Wolf-killed ungulates may restrict access to 
Wolf-killed ungulates among other members of the 
GYE grizzly population (i.e., reproductive 
females). 

If Wolf-killed ungulates are unavailable to griz- 
zlies in the fall and/or consumed mainly by adult 
male grizzlies, a population level benefit to GYE 


498 


Grizzly Bears due to Wolf-killed ungulates seems 
unlikely. However, further research is necessary to 
understand the seasonal and demographic variation 
in Grizzly Bear access to Wolf-killed ungulates. 


Acknowledgments 

We appreciate the support and cooperation of 
Yellowstone rangers John Lounsbury and Lloyd 
Kortge, and the administrators and staff at the 
Yellowstone Center for Resources. 


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Received 15 August 2000 
Accepted 16 July 2001 


Notes 


An Observation of a Mallard, Anas platyrhynchos, 
Feeding on a Wood Frog, Rana sylvatica 


BRIAN R. EATON! and ZACHARY C. EATON 


221 RH Michener Park, Edmonton, Alberta T6H 4M5 Canada 


'Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada T6G 2E9. Author to whom corre- 


spondence should be addressed. 


Eaton, Brian R., and Zachary C. Eaton. 2001. An observation of a Mallard, Anas platyrhynchos, feeding on a Wood Frog, 
Rana sylvatica. Canadian Field-Naturalist 115(3): 499-500. 


Mallards (Anas platyrhynchos) will feed opportunistically on vertebrates, and have been reported feeding on amphibians in 
Europe; similar reports in North America are rare. Here we report on an observation of a Mallard feeding on an adult Wood 


Frog (Rana sylvatica) in April 1999, in Edmonton, Alberta. 


Key Words: Mallard, Anas platyrhynchos, Wood Frog, Rana sylvatica. 


On 24 April 1999, we were searching for Wood 
Frogs (Rana sylvatica) and Boreal Chorus Frogs 
(Pseudacris triseriata maculata), and their eggs, in a 
series of pools in Whitemud Park in Edmonton, 
Alberta (53° 33’ N, 113° 28’ W). These stagnant 
pools seemed to be the remnants of a braided stream, 
with a width of approximately 1 metre. Both Wood 
and Chorus Frogs were calling in the area, and male 
Wood Frogs were visible on the surface of the water; 
we found no egg masses. 

While searching, we disturbed a female Mallard 
Duck (Anas platyrhynchos) from under vegetation 
overhanging a pool; she swam slowly away from us. 
Several minutes later, the same Mallard was seen 
biting at what proved to be a Wood Frog. She 
flipped the frog out of the water and on to the bank, 
and, after some struggle grasped the frog in her bill, 
tipped her head up and back, and swallowed the frog 
head first. We were able to train binoculars on the 
duck as she swallowed, and based on the size and 
shape of the hind legs were able to identify the prey 
as an adult Wood Frog. The entire sequence of 
events, from attack to complete consumption of the 
frog, lasted approximately 30 seconds. 

After eating the Wood Frog, the Mallard settled 
on the bank where she had captured the frog. She 
remained there several minutes and reacted once 
more to movement in the water, probably the activity 
of another frog. When reacting she jerked her head 
up and forward from its resting position and stood. 
She did not, however, pursue more frogs. 

Predation on amphibians by members of the 
Anseriformes is not often reported in the literature. 
In a review of amphibians and reptiles as prey of 


birds in southwestern Europe, Martin and Lopez 
(1990) only listed one general reference, and one 
specific reference, for predation on amphibians by 
ducks, geese, and related taxa. In North America, 
McAtee (1918) found remains of frogs in one of 622 
Black Duck (Anas rubripes) stomachs. Mabbott 
(1920) found frog remains in one of 790 Pintail 
(Anas acuta), and two of 413 Wood Duck (Aix spon- 
sa) stomachs. Mallory and Lariviére (1998) collected 
a Wood Duck on 15 September 1997 with three 
Mink Frogs (Rana septentrionalis) in her esophagus; 
SVL of all three frogs was approximately 45 mm. 
Trochell and Watermolen (1995) observed two adult 
Canada Geese (Branta canadensis) feeding on newly 
metamorphosed American Toads (Bufo americanus) 
emerging from a small pond in Wisconsin in June 
1994. These few reports suggest that, although 
amphibians may be taken by anseriforms, they do 
not usually form a large part of the diet. 

To our knowledge, there are no published reports 
of Mallards eating Wood Frogs, although the use of 
these frogs as prey by Mallards is not surprising. 
McTee (1918) found frog remains in 19 of 1725 
Mallard stomachs. Bent (1923) stated that Mallards 
take the “occasional slug, snail, frog, or lizard’, and 
Bannerman (1958) mentioned that “Frogs, tadpoles, 
and spawn are greedily devoured” by Mallards; nei- 
ther author supplied specific information. Cramp 
(1977) also listed amphibians among prey taken by 
Mallards. Sugden and Driver (1980) observed a 
Mallard with a brood of full-grown young capture 
and swallow a salamander (Ambystoma sp.) on 2 
August 1978; the authors did not specify whether the 
encounter occurred on land or in the water, or the 


499 


500 


size of the salamander. Based on distribution maps, 
the salamander must have been a subspecies of Tiger 
Salamander (Ambystoma tigrinum); adults of that 
species range in size from 75 to 162 mm SVL 
(Stebbins 1985). 

More recently, Mjelstad and Setersdal (1989) 
observed Mallards eating the Common Frog (Rana 
temporaria) in western Norway. Mijelstad and 
Sztersdal (1989) were not able to measure the frogs 
being taken by Mallards, but Common Frogs in 
Switzerland range in size from 59 to 94 mm in 
snout-to-vent length (SVL) (Ryser 1996). We were 
unable to measure the Wood Frog we observed being 
consumed by the Mallard, but adult Wood Frogs in 
Alberta range in size from 30 to 60 mm SVL 
(Russell and Bauer 2000). Captive Mallards will eat 
Boreal Toad (Bufo boreas) tadpoles (Jones et al. 
1999), even though toad tadpoles are generally con- 
sidered noxious to most potential predators. These 
observations suggest that Mallards may prey upon a 
variety of amphibians, some of which are quite large. 

Amphibians may represent an important protein 
source for both male and female Mallards during mat- 
ing and egg-laying (Mjelstad and Setersdal 1989). 
Eldridge and Krapu (1988) showed experimentally 
that female Mallards on a high protein diet produced 
larger clutches and eggs than Mallards on a protein- 
poor diet. Some species of amphibians contain a high- 
er concentration of protein in their bodies than either 
birds or small mammals, and represent higher-quality 
prey than these other two groups (Burton and Likens 
1975). In some areas, this high-quality food becomes 
available to Mallards during the egg-formation period, 
as was the case in Norway. As a result, frogs were 
taken regularly by both male and female Mallards 
during diving, gleaning, and kleptoparasitic activities 
(Mjelstad and Szetersdal 1989). 

Observations of waterfowl preying on amphibians 
are relatively rare, but indicate that a wide range of 
amphibians, both in terms of size and species, will 
be taken when available. Even if the observed level 
of predation by waterfowl on amphibians is an accu- 
rate reflection of the actual rate at which they are 
consumed by ducks and geese in the wild, amphib- 
ians are high quality prey that may be important pro- 
tein and energy sources for individual waterfowl. 


Acknowledgments 

Thanks are extended to A. J. Erskine, Cynthia 
Paszkowski, and an anonymous reviewer, who pro- 
vided comments on an earlier version of this paper. 
These observations were made during a field trip 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


with the Michener Park Young Naturalist Club; 
thanks to Mayrse Maurice for leading the trip. 


Literature Cited 

Bannerman, D. A. 1958. The birds of the British Isles. 
Volume 7. Oliver and Boyd, London. | 

Bent, A. C. 1923. Life histories of North American wild 
fowl. United States National Museum, Smithsonian 
Institution, Bulletin Number 126. 

Burton, T. M., and G. E. Likens. 1975. Energy flow and 
nutrient cycling in salamander populations in the Hub- 
bard Brook Experimental Forest, New Hampshire. 
Ecology 56: 1068-1080. 

Cramp, S. Editor. 1977. Handbook of the birds of 
Europe, the Middle East, and North Africa. The birds of 
the western palearctic. Volume 1: Ostrich to ducks. 
Oxford University Press, Oxford. 

Eldridge, J. L., and G. L. Krapu. 1988. The influence of 
diet quality on clutch and laying pattern in Mallards. 
Auk 105: 102-110. 

Jones, M. S., J. P. Goettl, and L. J. Livo. 1999. Bufo 
boreas (Boreal toad): predation. Herpetological Review 
0791: 

Mabbott, D. C. 1920. Food habits of seven species of 
American shoal-water ducks. United States Department 
of Agriculture Bulletin Number 862. 

Mallory, M. L., and R. Lariviére. 1998. Wood Duck, Aix 
sponsa, eats Mink Frogs, Rana septentrionalis. Canadian 
Field-Naturalist 112: 714-715. 

Martin, J., and P. Loépez. 1990. Amphibians and reptiles 
as prey of birds in southwestern Europe. Smithsonian 
Herpetological Information Service Number 82. 43 pages. 

McAtee, W. L. 1918. Food habits of the Mallard Ducks 
of the United States. United States Department of 
Agriculture, Bulletin Number 720. 

Mjelstad, H., and M. Setersdal. 1989. Mallards Anas 
platyrhynchos utilizing frogs Rana temporaria as a pro- 
tein resource during breeding. Fauna norvegica Series C. 
12: 47-48. 

Russell, A. P., and A. M. Bauer. 2000. The amphibians 
and reptiles of Alberta. Revised (second) edition. 
University of Calgary Press, Calgary. 

Ryser, J. 1996. Comparative life histories of a low- and a 
high-elevation population of the common frog Rana 
temporaria. Amphibia-Reptilia 17: 183-195. 

Stebbins, R. C. 1985. A field guide to western amphib- 
ians and reptiles (2nd edition). Houghton Mifflin 
Company, Boston. 336 pages. 

Sugden, L. G., and E. A. Driver. 1980. Natural foods of 
mallards in Saskatchewan parklands during late summer 
and fall. Journal of Wildlife Management 44: 705-709. 

Trochell, P., and D. J. Watermolen. 1995. Report of 
Canada geese feeding on American toads. Passenger 
Pigeon 57: 199. 


Received 8 May 2000 
Accepted 26 October 200i 


2001 NOTES 501 


First Record of an Anomalously White Killer Whale, Orcinus orca, 
Near St. Lawrence Island, Northern Bering Sea, Alaska 


SUZANN G. SPECKMAN!:2 and GAY SHEFFIELD! 3 


[Institute of Marine Science, University of Alaska Fairbanks, Fairbanks, Alaska 99775 USA 

2Present address: School of Aquatic and Fishery Sciences, University of Washington, 1122 NE Boat St., Seattle, 
Washington 98105 USA; e-mail Speckman @u.washington.edu 

3Present address: Alaska Department of Fish and Game, 1300 College Road, Fairbanks, Alaska 99701 USA 


Speckman, Suzann G., and Gay Sheffield. 2001. First record of an anomalously white Killer Whale, Orcinus orca, near 
St. Lawrence Island, Northern Bering Sea, Alaska. Canadian Field-Naturalist 115(3): 501-502. 


An anomalously white Killer Whale (Orcinus orca) was sighted swimming with a group of normally-pigmented Killer 
Whales near St. Lawrence Island in the northern Bering Sea. The white whale had a tall, straight dorsal fin, indicating a 
mature male. Most sightings of white or albinistic Killer Whales have been of individuals that were smaller than their com- 
panions. This sighting of an adult white male is unusual, given that albinistic individuals may not survive as long or attain 
as large a size as their normally-pigmented conspecifics due to the wide range of physical and physiological abnormalities 


commonly associated with partial or complete albinism. 


Key Words: Killer Whale, white cetacean, Orcinus orca, albinism, Bering Sea. 


On 27 August 1993, we sighted an all-white Killer 
Whale (Orcinus orca) swimming with approximate- 
ly 11 other Killer Whales 5.5 km southwest of St. 
Lawrence Island, in the northern Bering Sea, Alaska 
at 63°47’N, 171°45’W. The water depth was <50 m 
and sea state was calm. The whales were first spotted 
about 500 m from our vessel, swimming slightly 
towards us and parallel to our course, and we could 
see them clearly through 10 x 40 binoculars. The 
white whale was large with a tall, straight dorsal fin, 
indicating a mature male. It was a creamy yellowish 
color over all of its exposed body—the head, back, 
and dorsal fin. There did not appear to be any dark 
pigmentation. The placing of the typical pigmenta- 
tion pattern of a Killer Whale was visible, with the 
normally white saddle behind the dorsal fin showing 
as a lighter, whiter color. We were unable to see the 
whale’s eyes so cannot confirm it was a true albino 
(pigmentless pink eyes) or leucistic (normal dark 
eyes). This represents the first report of a white or 
possibly albinistic Killer Whale for the Bering Sea. 

The group we encountered included a second 
mature male with a tall dorsal fin. This second male 
was normally-pigmented and its fin was slightly 
taller than that of the white whale. The remaining 
whales were normally-pigmented adult females or 
subadults. The closest approach of the group was 
about 400 m, and all of the whales, including the 
white one, were behaving normally, swimming and 
surfacing for air. 

Anomalously white cetaceans are reported infre- 
quently. Hain and Leatherwood (1982) and Fert] et 
al. (1999) reviewed the literature and compiled 
unpublished observations, and found that such indi- 
viduals have been sighted in 20 cetacean species. 
Although details are scarce, most sightings of white 
Killer Whales have been of individuals that were 


reported as “small” or smaller than their companions 
(Carl 1959; Pilleri and Pilleri 1987; Scheffer and 
Slipp 1948). There are two reports of white female 
Killer Whales that appeared to have calves, both of 
which were also white (although one calf had a black 
border on the dorsal fin: Carl 1959). These breeding 
females must have been at least 15 years old 
(Olesiuk et al. 1990). The tall dorsal fin of the white 
male we sighted indicates that he was physically 
mature and therefore at least 21 years of age 
(Olesiuk et al. 1990). 

Our sighting of such a long-lived albinistic Killer 
Whale is unique, because on average, albinistic indi- 
viduals may not survive as long or attain as large a 
size as their normally-pigmented conspecifics. Hain 
and Leatherwood (1982, from Searle 1968) noted a 
wide range of pathological conditions that are com- 
monly associated with partial or complete albinism. 
These included lowered fertility, anemia, defects of 
the eyes and ears, defects of the central nervous sys- 
tem, and increased susceptibility to infection. Other 
conditions also occur that can mimic albinism. A 
captive “white” Killer Whale with pale, ghostly out- 
lines of the usually white markings was diagnosed 
with Chediak-Higashi syndrome (Ridgway 1979), 
which presumably caused or contributed to its death. 
Most of the conditions that accompany lack of pig- 
ment would increase mortality rates, diminishing the 
average body size and shortening the average lifes- 
pan. 


Acknowledgments 

This sighting occurred during a research cruise 
aboard the Russian R/V Okean as part of the Program 
of Long Term Ecological Investigations of the Bering 
Sea and Other Pacific Ocean Ecosystems (BERPAC). 
Logistical support was provided by the U.S. Fish and 


502 


Wildlife Service. We thank C. Peter McRoy and 
Jackie Grebmeier for additional support and encour- 
agement, and Robert Suydam for comments on the 
manuscript. We also appreciate the critical reviews by 
Robin W. Baird and an anonymous reviewer. 


Literature Cited 

Carl, C. 1959. Albinistic killer whales in British 
Columbia. Report of the Provincial Museum, British 
Columbia 1959: 29-36. 

Fertl, D., L. T. Pusser, and J. J. Long. 1999. First record 
of an albino bottlenose dolphin (Tursiops truncatus) in 
the Gulf of Mexico, with a review of anomalously white 
cetaceans. Marine Mammal Science 15: 227-234. 

Hain, J. H. W., and S. Leatherwood. 1982. Two sight- 
ings of white pilot whales, Globicephala melaena, and 
summarized records of anomalously white cetaceans. 
Journal of Mammalogy 63: 338-343. 

Olesiuk, P. F., M. A. Bigg, and G. M. Ellis. 1990. Life 
history and population dynamics of resident killer 
whales (Orcinus orca) in the coastal waters of British 


THE CANADIAN FIELD-NATURALIST 


Vole nis 


Columbia and Washington state. Pages 209-243 in 
Individual recognition of cetaceans: use of photo identi- 
fication and other techniques to estimate population 
parameters. Edited by P: S. Hammond, S. A. Mitzroch, 
and G. P. Donovan. Report of the International Whaling 
Commission, Special Issue 12. 

Pilleri, G., and O. Pilleri. 1987. Records of cetaceans in 
the Mediterranean Sea and North Atlantic Ocean in the 
period 1982-1986. Investigations on Cetacea 20: 
267-280. 

Ridgway, S. H. 1979. Reported causes of death of captive 
killer whales (Orcinus orca). Journal of Wildlife 
Diseases 15: 99-104. 

Scheffer, V. B., and J. Slipp. 1948. The whales and dol- 
phins of Washington State with a key to cetaceans of the 
West Coast of North America. American Midland 
Naturalist 39: 257-337. 

Searle, A. B. 1968. Comparative genetics of coat color in 
mammals. Logos Press Ltd., London. 


Received 4 April 2000 
Accepted 17 December 2001 


Evidence for Double Brooding by a Mallard, Anas platyrhynchos, 


in Eastern South Dakota 


JOSHUA D. STAFFORD!:3, LESTER D. FLAKE!, and PAUL W. MAMMENGA2 


1Department of Wildlife and Fisheries Sciences, NPB 138 Box 2140B, South Dakota State University, Brookings, South 


Dakota 57007 USA 


2South Dakota Department of Game, Fish and Parks, 5850 East Highway 12, Aberdeen, South Dakota 57401 USA 


3Current Address: Department of Wildlife and Fisheries, Box 9690, Mississippi State University, Mississippi State, 
Mississippi 39762 USA 


Stafford, Joshua D., Lester D. Flake, and Paul W. Mammenga. 2001. Evidence for double brooding by a Mallard, Anas 
platyrhynchos, in eastern South Dakota. Canadian Field-Naturalist 115(3): 502—504. 


Documentation of double brooding by ducks is uncommon in the northern hemisphere. We report double brooding by a 
Mallard (Anas platyrhynchos) in eastern South Dakota during 1999. A radio-marked female hatched her clutch, reared her 
brood to 15-25 d post-hatch, and renested. The female successfully hatched and reared her second brood to 50 days post- 
hatch. These data plus a band recovery in October of 2001 from the first brood provide circumstantial evidence of double 


brooding by this female. 


Key Words: Mallard, Anas platyrhynchos, brood rearing, double brooding, second nesting. 


Renesting by ducks after disturbance during lay- 
ing, incubation, or after clutch loss is not uncommon 
(Sowls 1955; Swanson et al. 1986). Some duck 
species in North America will renest after brood 
loss, especially if brood mortality occurs soon after 
hatch (Bjarvall 1969; Doty 1975). Few cases in 
North America, with the exception of Wood Ducks 
(Aix sponsa)(Fredrickson and Hansen 1983; 
Moorman and Baldassarre 1988; Fielder 1992), doc- 
ument the production of a second brood when at 
least one duckling from the first brood is known to 
have survived (double brooding). In the southern 
hemisphere, evidence for double brooding exists for 
several Australian duck species including the Pink- 


eared Duck (Malacorhynchus membranaceus), 
Chestnut Teal (Anas castanea), Grey Teal (Anas gib- 
berifrons) and Australian Wood Duck (Chenonetta 
jubata) (Braithwaite 1976a, 1976b). A White- 
cheeked Pintail (Anas bahamensis) in the Bahamas 
re-nested and hatched a second clutch after success- 
fully fledging her first brood (Sorenson et al. 1992). 
Eleven female Mallards (Anas platyrhynchos) in 
an unusually dense population near the Delta Water- 
fowl Research Station in Manitoba abandoned 
broods to renest, but fates of individual broods were 
not presented; this population consisted largely of 
artificially maintained and hatchery-reared Mallards, 
many of which overwintered on site (Titman and 


2001 


Lowther 1975). Based on age class data and repeated 
observations, Bjarvall (1969) documented double 
brooding by two Mallards in Sweden, where duck- 
lings from early and late broods of both females 
were known to have fledged. The rearing of two 
broods to fledge by a Mallard in North America dur- 
ing a single breeding season, in a natural setting, has 
not, to our knowledge, been documented. Here we 
report evidence of double brooding by a Mallard in 
eastern South Dakota. 

In 1999, we conducted research on Mallard duck- 
ling survival at the Redetzke Game Production Area 
(Redetzke) in southeastern Day County, South 
Dakota (45°15’45’’N, Longitude 97°25’00’’W). 
Redetzke is a 160-ha semi-permanent wetland with 
many overwater nesting structures (Stewart and 
Kantrud 1971). About 70-80% of this wetland con- 
sisted of dense interspersed emergent vegetation, pri- 
marily cattails (Typha spp.), whereas the center 
remained free of emergent vegetation. 

Female Mallards were captured from overwater 
nesting structures during incubation and fitted with 
4.5-g radio transmitter using a modified Mauser 
attachment (Pietz et al. 1995). Ducklings from some 
of our marked females were also fitted with 1.5 g 
transmitters of the same type while still at the nest. 
After nest exodus, females with broods were moni- 
tored until ducklings fledged (approximately 55 days 
post-hatch, Bellrose 1976), contact was lost, or the 
adult died. Marked ducklings were also monitored 
until death or loss of the transmitter. Counts of duck- 
lings within broods were scheduled every seven days 
but were recorded opportunistically. Female behav- 
ior was monitored daily and used to determine the 
presence or absence of a brood when ducklings were 
not seen (Bergmann 1992; Rotella and Ratti 1992). 

On 13 May, a radio-marked female hatched all of 
13 eggs and left her nest on 14 May; two of these 
ducklings were radio-marked and the remaining 11 
were fitted with plasticine-filled leg bands (Blums et 
al. 1994). On 15 May we determined that the two 
radio-marked ducklings were not with the female. 
Upon investigation, on 18 May, we found both 
radio-marked ducklings and one banded duckling 
dead on a Muskrat (Ondatra zibethicus) lodge near 
the nesting structure. The presence of an egg tooth 
and progress of decomposition of the ducklings indi- 
cated that they died shortly after leaving the nest. 
Consistent use of dense emergent vegetation by the 
female compromised our ability to observe the brood 
regularly. Based on her movements and behavior we 
assumed the female was with her brood from the 
time of hatch until 15 days post-hatch when we first 
observed her accompanying a brood of eight. During 
the period from 29 May until 7 June we did not 
observe the female again, but she remained highly 
localized in the same marsh where she had last been 
observed with a brood; other radio-marked females 


NOTES 


503 


that lost their broods greatly increased their home 
range and moved erratically between wetlands. Such 
erratic movements among wetlands were clearly dif- 
ferent than for females with broods (Rotella and 
Ratti 1992). 

On 7 June, 25 days post-hatch, this female shifted 
her activities from the brood rearing area to the vicini- 
ty of the nesting structure where she had hatched her 
first brood. The female laid a second clutch of nine 
eggs in the original nesting structure and initiated 
incubation. On 12 July, 60 days after hatching the first 
brood, 8 of 9 eggs hatched from the second clutch. 
We observed ducklings with the female several times 
during the second rearing period; most notably, six 
ducklings were present on 16 August, 36 days post- 
hatch. Additionally, female behavior and partial brood 
observations suggested the presence of the second 
brood until 50 days post-hatch. 

Survival of at least one duckling from the first 
brood was confirmed in October of 2001 when we 
received a band recovery report for a female mallard 
shot near Rugby, North Dakota. We believe that sur- 
vival to flight of ducklings, from the 8 alive at 15 
days post-hatch was probable. Mallard duckling 
mortality in the company of the female is low after 
10-18 days post-hatch (Ball et al. 1975; Talent et al. 
1983; Orthmeyer and Ball 1990; Mauser et al. 1994). 
Additionally, there were no catastrophic events (e.g., 
hail storms) generally associated with sudden and 
severe ducling losses. Non-catastrophic weather 
events (e.g., prolonged periods of rain or cold) may 
also adversely affect duckling survival (Korschgen et 
al. 1996; Krapu et al. 2000). We do not discount the 
possibility that periodic rainfall and temperature 
fluctuations may have contributed to duckling mor- 
tality in our study. 

During our two-year study, radio-marked duck- 
lings from a different brood fledged after the brood 
female was killed at approximately 10 days post- 
hatch. In a California study (Mauser et al. 1994), 
three radio-marked Mallard ducklings from two 
females that appeared to lose their entire brood 
joined and were fledged by other brood females. 
Frequent amalgamations of Mallard broods may 
occur in crowded populations (Titman and Lowther 
1975). Fifty-two successful nests were produced 
from overwater nesting structures at Redetzke in 
1999. Other Mallard broods used this wetland during 
brood rearing and the ducklings in the first brood 
may have joined another brood. 

The female’s daily behavior suggested nearly con- 
tinuous brood care from hatching until the renesting 
attempt. Because there was no noticeable time-lag 
between leaving the first brood and initiating the sec- 
ond nest, copulation must have taken place while the 
female was rearing the first brood. We cannot 
account for the timing of this event. Throughout our 
study we observed several females with broods con- 


504 


currently accompanied by drakes and occasionally 
forced away from their broods. Although the exact 
behavioral circumstances are unknown, the evidence 
documents an unusual case of double brooding by 
this Mallard. 

Oring and Sayler (1992) identify double brooding 
as one strategy in female ducks to adjust (increase) 
reproductive output. For example, double brooding 
by Wood Ducks in Missouri increases production by 
an estimated 2% annually (Fredrickson and Hansen 
1983). Reports of double brooding by temperate 
nesting waterfowl, though, are uncommon and poor- 
ly documented. Therefore, we do not suspect that 
double brooding is common in Mallards, and the low 
frequency of occurrence precludes a significant 
influence on annual Mallard production. 


Acknowledgments 

We thank D. Alexander, M. Grovijahn, and C. 
Langer for field assistance. A. J. Erskine, L. 
Fredrickson, P. Pietz, and J. Rotella reviewed this 
manuscript and provided helpful comments. Many 
thanks to the Waubay National Wildlife Refuge for 
providing field housing. This study was funded by the 
Federal Aid to Wildlife Restoration Fund adminis- 
tered through the South Dakota Department of Game, 
Fish and Parks; the Delta Waterfowl and Wetlands 
Research Station; and Ducks Unlimited, Inc. 


Literature Cited 

Ball, I. J.. D.S. Gilmer, L. M. Cowardin, and J. H. 
Riechmann. 1975. Survival of wood duck and Mallard 
broods in north-central Minnesota. Journal of Wildlife 
Management 39: 776-780. 

Bellrose, F.C. 1976. Ducks, geese & swans of North 
America. Second edition. Stackpole Books, Harrisburg, 
Pennsylvania. 

Bergmann, P. J. 1992. Movements, survival, and habitat 
use of Mallard broods hatched from predator reduced 
nesting habitats in eastern South Dakota. M.S. thesis, 
South Dakota State University, Brookings. 

Bjarvall, A. 1969. Unusual cases of re-nesting Mallards. 
Wilson Bulletin 81: 94-96. 

Blums, P., A. Mednis, and J. D. Nichols. 1994. Retention 
of web tags and plasticine-filled leg bands applied to 
day-old ducklings. Journal of Wildlife Management 58: 
76-81. 

Braithwaite, L.W. 1976a. Breeding seasons of waterfowl 
in Australia. Proceedings of the International Ornitho- 
logical Congress 16: 235-247. 

Braithwaite, L.W. 1976b. Environment and timing of 
reproduction and flightlessness in two species of 
Australian ducks. Proceedings of the International 
Ornithological Congress 16: 489-501. 

Doty, H. A. 1975. Renesting and second broods of wild 
Mallards. Wilson Bulletin 87: 115. 

Fielder, P.C. 1992. Double brooding by a wood duck in 
Washington. Northwest Naturalist 73: 26-27. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Fredrickson, L.H., and J. L. Hansen. 1983. Second 
broods in wood ducks. Journal of Wildlife Management 
47: 320-326. . 

Korschgen, C.E., K.P.. Kenow, W.L. Green, D. H. 
Johnson, M.D. Samuel, and L. Sileo. 1996. Survival 
of radiomarked Canvasback ducklings in northwestern 
Minnesota. Journal of Wildlife Management 60: 
120-132. 

Krapu, G.L., P. J. Pietz, D. A. Brandt, and R. R. Cox. 
2000. Factors limiting Mallard broods survival in prairie 
pothole landscapes. Journal of Wildlife Management 64: 
553-561. 

Mauser, D.M., R.L. Jarvis, and D.S. Gilmer. 1994. 
Survival of radio-marked Mallard ducklings in north- 
eastern California. Journal of Wildlife Management 58: 
82-87. 

Moorman, T. E., and G. A. Baldassarre. 1988. Incidence 
of second broods by wood ducks in Alabama and 
Georgia. Journal of Wildlife Management 52: 426-431. 

Oring, L. W., and R. D. Sayler. 1992. The mating sys- 
tems of waterfowl. Pages 190-231 in Ecology and man- 
agement of breeding waterfowl. Edited by B. D. J. Batt, 
A.D. Afton, M.G. Anderson, C.D. Ankney, D.H. 
Johnson, J. A. Kadlec, and G.L. Krapu. University of 
Minnesota Press, Minneapolis, Minnesota, USA. 

Orthmeyer, D. L., and I. J. Ball. 1990. Survival of Mallard 
broods on Benton Lake National Wildlife Refuge in 
northcentral Montana. Journal of Wildlife Management 
54: 62-66. 

Pietz, P. J.. D. A. Brandt, G. L. Krapu, and D. A. Buhl. 
1995. Modified transmitter attachment method for adult 
ducks. Journal of Field Ornithology 66: 408-417. 

Rotella, J. J., and J. T. Ratti. 1992. Mallard brood sur- 
vival and wetland habitat conditions in southwestern 
Manitoba. Journal of Wildlife Management 56: 499— 
507. 

Sorenson, L. G., B. L. Woodworth, L. M. Ruttan, and F. 
McKinney. 1992. Serial monogamy and double brood- 
ing in the White-cheeked (Bahama) Pintail Anas baha- 
mensis. Wildfowl 43: 156-159. 

Sowls, L. K. 1955. Prairie ducks. A study of their behavior, 
ecology, and management. Stackpole Co. Harrisburg, 
Pennsylvania, and Wildlife Management Institute, Wash- 
ington, D.C. 

Stewart, R.E., and H. A. Kantrud. 1971. Classification 
of natural ponds and lakes in the glaciated pothole 
region. U.S. Fish and Wildlife Service Resource Publi- 
cation 92. 49 pages. 

Swanson, G.A., T. L. Shaffer, J. F. Wolf, and F. B. Lee. 
1986. Renesting characteristics of captive Mallards on 
experimental ponds. Journal of Wildlife Management 
50: 32-38. 

Talent, L.G., R. L. Jarvis, and G.L. Krapu. 1983. Sur- 
vival of Mallard broods in south-central North Dakota. 
Condor 85: 74-78. _ ; 

Titman, R.D., and J. K. Lowther. 1975. The breeding 
behavior of a crowded population of Mailards. Canadian 
Journal of Zoology 53: 1270-1283. 


Received 30 May 2000 
Accepted 20 July 2001 


2001 


NOTES 


505 


Opportunistic Foraging at American Elk, Cervus elaphus, 
Droppings by Clark’s Nutcracker, Nucifraga columbiana 


PAUL HENDRICKS and LISA N. HENDRICKS 


Montana Natural Heritage Program, 909 Locust Street, Missoula, Montana 59802 USA 


Hendricks, Paul, and Lisa N. Hendricks. 2001. Opportunistic foraging at American Elk, Cervus elaphus, droppings by 
Clark’s Nutcracker, Nucifraga columbiana. Canadian Field-Naturalist 115(3): 505-506. 


In the Uinta Mountains, Utah, a Clark’s Nutcracker (Nucifraga columbiana) was observed in August 1998 picking through 
a large pile of fresh soft droppings of American Elk (Cervus elaphus). During a five-minute period the nutcracker 
swallowed items recovered from the droppings 33 times, but items were too small to identify from 3-6 m distant using 
binoculars. Dissected droppings revealed only a few small grass and sedge seeds among undigested plant fragments. No 
insects were observed on or under the droppings, and the nutcracker never consumed a large quantity of the droppings, as it 
might have if it were seeking rare minerals or vitamins. The nutcracker pried apart some individual droppings, indicating it 
was searching for items contained within them, and may have been after the smail, undigested seeds. 


Key Words: Clark’s Nutcracker, Nucifraga columbiana, foraging behavior, American Elk, Cervus elaphus, Utah. 


Ingestion of fecal material is a common attribute 
of nest sanitation activity of passerine birds raising 
nestlings (Skutch 1976), including Clark’s 
Nutcracker, Nucifraga columbiana (Mewaldt 1956). 
Fecal sacs of nestling birds may be ingested for their 
water, energy, and nutrients by the parents (Morton 
1979). Birds also sometimes feed on the feces of 
another species to extract undigested food (e.g., 
Takenaka 1992), but feeding on another species’ 
feces by Clark’s Nutcracker is apparently unreported 
(Bent 1946; Tomback 1998). The European 
Nutcracker (N. caryocatactes) is reported to feed on 
dung (Cramp 1994), but it is not clear if the birds 
ingested dung or food items in or on the dung. 

On 18 August 1998 we observed a Clark’s 
Nutcracker as it foraged at fresh droppings of 
American Elk (Cervus elaphus). Our observation 
was made in Summit County, Utah near treeline 
(40°47°N, 110°36’W) at 3292 m elevation in the 
Uinta Mountains. Forest canopy at the site was 
Subalpine Fir (Abies lasiocarpa), Lodgepole Pine 
(Pinus contorta), and Engelmann Spruce (Picea 
engelmannii). Between 1015-1020 MDT we 
observed the nutcracker at close range (3-6 m) with 
binoculars as it probed through the droppings and 
swallowed 33 times unidentified items that were too 
small to see. The bird jabbed at the pile of droppings 
with a closed bill, flipped the loosened droppings 
aside, and sometimes pried individual droppings 
apart by piercing them and then slightly opening its 
bill, as it would open a conifer cone to extract a seed. 
The nutcracker’s bill was covered with fecal materi- 
al, and when finally flushed from the droppings it 
went te a nearby rock and wiped off its bill. 

The elk droppings were fresh, and the soft type 
(Murie 1954) typically produced from a diet of green 
succulent vegetation. We collected a random grab- 
sample of 20 elk droppings (mean length < width = 
25 X 15 mm), 10 of which (total dry weight = 8.3 g) 


we later dissected and examined with a microscope. 
The pellets were composed largely of coarse and fine 
undigested plant fragments. No insect eggs, larvae or 
large seeds were present, but we recovered eight 
small seeds (mean length X width= 1.5 * 1.0 mm) 
of sedge (Carex) and grass (Poa). We probably 
missed a few small seeds during our dissections, but 
the droppings were composed mostly of non-seed 
material. 

Nutcrackers feed on a diverse variety of items, 
including seeds, carrion, insects, and small verte- 
brates (Bent 1946; Goodwin 1976; Mulder et al. 
1978; Cramp 1994; Tomback 1998), and are consid- 
ered opportunistic foragers. There are at least three 
explanations for the apparent behavior of the 
nutcracker we observed, two of which we exclude. 

First, the bird could have been searching for 
insects associated with the droppings. Nutcrackers 
have been reported hunting insects hidden under dry 
cow dung by flipping over individual droppings 
(Goodwin 1976). No insects were observed on, 
under, or in the elk droppings, nor did we observe 
the nutcracker actively hunting and consuming 
insects even though we were close enough to detect 
this activity. This eliminates the possibility that the 
bird was hunting for insects on or under the drop- 
pings. Second, the nutcracker could have been seek- 
ing nutrients or vitamins in the elk droppings. 
Because we never saw the bird consume a whole 
dropping or large dropping fragment, even though 
the bird could easily have done so, this is an unlikely 
explanation for why the nutcracker was feeding at 
the elk droppings, and remained at them for five 
minutes. Furthermore, the nutcracker’s behavior of 
splitting open some of the individual droppings indi- 
cated it was searching for items contained in the 
droppings and was not after the droppings them- 
selves. Third, the nutcracker could have been search- 
ing for undigested plant material and seeds in the elk 


506 


droppings. We favor this third explanation as the 
most plausible explanation for the unusual foraging 
behavior we observed, or at least what initially 
attracted the nutcracker to the droppings. 

The few seeds we found in the droppings were 
quite small, however, seemingly a very small food 
reward for the nutcracker. Nevertheless, small seeds 
have been reported previously in stomachs of Clark’s 
Nutcracker (Giuntoli and Mewaldt 1978). It has been 
suggested that some of these seeds may have been 
ingested with fruits (Giuntoli and Mewaldt 1978), but 
our observation indicates that nutcrackers might also 
actively seek small seeds from novel sources. Perhaps 
foraging at animal dung is more likely to occur when 
the fall harvest of ripening conifer seeds (Tomback 
1978) coincides with years of low cone production, 
inducing nutcrackers to seek food from other sources 
to supplement their diet of conifer seeds. 
Nevertheless, absence of additional observations of 
foraging behavior at ungulate dung indicates it is 
probably a rare activity at any time. 


Acknowledgments 

D. Tomback and A. J. Erskine made several useful 
suggestions on earlier drafts that improved the presen- 
tation. We thank P. F. Stickney for identifying the 
seeds, and extend a special thanks to J. Neeling for 
planning our trip into the High Uintas Wilderness. 


Literature Cited 
Bent, A. C. 1946. Life histories of North American jays, 


THE CANADIAN FIELD-NATURALIST 


Vol) 115 


crows, and titmice. U. S. National Museum Bulletin 
Number 191. 

Cramp, S. 1994. Handbook of the birds of Europe, the 
Middle East and North Africa. Oxford University Press, 
Oxford, United Kingdom. 

Giuntoli, M., and L. R. Mewaldt. 1978. Stomach con- 
tents of Clark’s Nutcracker collected in western 
Montana. Auk 95: 595-598. 

Goodwin, D. 1976. Crows of the world. Cornell 
University Press, Ithaca, New York. 

Mewaldt, L. R. 1956. Nesting behavior of the Clark 
Nutcracker. Condor 58: 3-23. 

Morton, M. L. 1979. Fecal sac ingestion in the Mountain 
White-crowned Sparrow. Condor 81: 72-77. 

Mulder, B. S., B. B. Schultz, and P. W. Sherman. 1978. 
Predation on vertebrates by Clark’s Nutcracker. Condor 
80: 449-451. 

Murie, O. J. 1954. A field guide to animal tracks. 
Houghton Mifflin Company, Boston, Massachusetts. 

Skutch, A. F. 1976. Parent birds and their young. 
University of Texas Press, Austin, Texas. 

Takenaka, M. 1992. A flock of Rosy Finch Leucosticte 
actoa feeding on Gray Starling Sturnus cineraceus feces 
in downtown Sapporo. Japanese Journal of Ornithology 
40: 77-79. 

Tomback, D. F. 1978. Foraging strategies of Clark’s 
Nutcracker. Living Bird 16: 123-161. 

Tomback, D. F. 1998. Clark’s Nutcracker (Nucifraga 
columbiana). In The Birds of North America. Number 
331. Edited by A. Poole and F. Gill. The Birds of North 
America, Inc., Philadelphia, Pennsylvania. 


Received 18 July 2000 
Accepted 26 October 2001 


Gillnet Survival and Healing by a Porbeagle, Lamna nasus 


GEORGE W. BENZ!, ANDY KINGMAN?, and JOANNA D. BORUCINSKA?3 


'Tennessee Aquarium and Southeast Aquatic Research Institute, P.O. Box 11048, Chattanooga, Tennessee, 37401-2048 


USA 


71805 Mar West Street, Apartment C, Tiburon, California 94920 USA 


3Department of Biology, University of Hartford, 200 Bloomfield Avenue, West Hartford, Connecticut, 06117-1559 USA 


Benz, George W., Andy Kingman, and Joanna D. Borucinska. 2001. Gillnet survival and healing by a Porbeagle, Lamna 
nasus. Canadian Field-Naturalist 115(3): 506—509. 


A 182 cm fork-length female Porbeagle, Lamna nasus, was captured on the Scotian Shelf south of Halifax (42°53’N, 
62°53’W) with a square of monofilament gillnet mesh almost totally embedded in its snout. Besides this shark’s strange 
“whiskered” appearance, gross inspection revealed it to be healthy. To our knowledge this report is the first documenting 


the healing of a shark after a net contact injury. 


Key Words: Porbeagle, Lamna nasus, Scotian Shelf, net contact injury, healing. 


Porbeagles, Lamna nasus, are sharks with large 
eyes, short snouts, long gill slits, and chubby appear- 
ances that are relatively common in cooler waters 
throughout the North Atlantic. While they are often 
found about the offshore fishing banks, they are also 
occasionally seen inshore (Bigelow and Schroeder 
1953; Compagno 1984). Like all lamnids, the 


Porbeagle is a swift and voracious predator, and the 
design of its circulatory system retains metabolic 
heat such that its body temperature is often higher 
than that of the surrounding water (see Carey and 
Teal 1969; Block and Carey 1985; Carey et al. 1985; 
Wolf et al. 1988). This “warm-bloodedness” could 
give the Porbeagle a metabolic advantage over its 


2001 


NOTES 


507 


le soe 


FIGURE 1. Right lateral (A), dorsal (B), left lateral (C), and ventral (D) views of a Porbeagle, Lamna nasus, with a gillnet 
mesh square embedded around its snout. Arrows indicate locations of exposed monofilament knots; s = thin linear 
scar across pineal window caused by the embedded mesh strand. Pineal window (p) appears as a lightly colored 
oval region at the dorsal midline of the head. Note that the jaws of this fish were removed prior to taking these pho- 


tographs. 


colder and more sluggish prey — which include 
mackerels, herrings, cods, and squids (Bigelow and 
Schroeder 1953; Compagno 1984). 

Porbeagles have been fished throughout much of 
their range (Bigelow and Schroeder 1953; 
Lineaweaver and Backus 1970; Compagno 1984) and 
in the northwestern Atlantic they have been the target 
of a longline fishery since 1961 (Hurley 1997). This 
fishery is mainly seasonal, March through November, 
with fishing beginning in early spring on the edge of 
the Scotian Shelf, followed by movement onto the 
Shelf itself and then into the Gulf of Saint Lawrence 
and onto the Grand Banks by July (Hurley 1997). 
Some fishermen working in late fall have followed the 
fish back into deeper water (Hurley 1997). Castro et 
al. (1999) noted that Porbeagle stocks have been 
depleted within a few years by intensive fisheries 


wherever they have existed, but that the species is not 
currently rare. 

In the spring of 1994, AK was aboard the Fishing 
Vessel Aquatic Pioneer collecting biological sam- 
ples during routine longline operations targeting 
Porbeagles. On 4 April an unusual Porbeagle was 
landed while fishing along the Scotian Shelf (42°53’ 
North latitude, 62°53’ West longitude). This female, 
182 cm in fork length, had a mesh square of mono- 
filament gillnet almost totally embedded in her snout 
with only the eight corner strands protruding, like so 
many stout whiskers (Figure 1). The snout of this 
shark was removed and fixed in 10 percent formalin 
for later parasitological studies by GWB. The 
unweighed shark was not noted in AK’s detailed 
deck log as being otherwise abnormal (i.e., thin 
body, discolored or small liver, sickly looking, etc.). 


508 


In the laboratory the “whiskered” snout raised 
enough interest to initiate a closer inspection, includ- 
ing a histologic examination by JDB. 

The net mesh square was 8.25 cm on each side 
(16.5 cm stretch mesh) and was composed of mono- 
filament nylon 0.6 mm in diameter. The stretched 
mesh square encircled the snout, coursing from just 
anterior to the opening of the right orbit (Figure 1A), 
across the pineal window (Figure 1B), to just anteri- 
or to the left orbit and then just posterior to the left 
external naris (Figure 1C), and finally across the 
ventral snout anterior to the upper jaw and just pos- 
terior to the right external naris (Figure 1D). Only 
four small portions of the net square remained 
exposed, each associated with one of the four corner 
knots (Figure 1). Where the monofilament was 
embedded, linear scars were visible on the overlying 
skin surface (Figure 1). Microscopic examination of 
these scars revealed them to be approximately 
2-10 mm wide, consisting of up to 33 rows of pla- 
coid scales whose crown ridges were not properly 
aligned with those of other scales about them. It is 
likely that they were formed as the lesions caused by 
the net healed and the placoid scales were haphaz- 
ardly regenerated. The monofilament was embedded 
up to 8 mm in places and it resided in the epidermis 
or more often in the dermis, except along the dorsal 
midline where it was embedded in the clear tissue of 
the pineal window (see Figure 1B). The oddly 
“whiskered” appearance of the fish was created by 
the two short lengths of monofilament associated 
with each of the four corner knots that at one time 
composed portions of an intact gillnet’s adjacent 
mesh squares. Some of the tips of these free lines 
appeared as if they had been stretched and broken 
while others looked as if they had been cleanly sev- 
ered. The location of the mesh square about the 
snout seemingly would not have permitted this fish 
to bite these lines. Besides the aforementioned com- 
plications, histologic examination of the tunnels 
within which the mesh square resided revealed a 
chronic inflammatory and healing response. The 
inflammation was associated with fragments of gran- 
ular gray-translucent material, evenly spaced at 
5—10 um intervals, that appeared to be tiny pieces of 
net material. At first we thought that the regular 
spacing of these fragments might be explained by 
them having been carved by the evenly spaced ridges 
on the crowns of the shark’s placoid scales. How- 
ever, subsequent measurements of the distance 
between these ridges corroborated the results of Reif 
(1985) concerning the Porbeagle by revealing the 
ridges on most head scales to be spaced about 
70-80 um apart. The inflammation about the frag- 
ments consisted of macrophages and lymphocytes 
within necrotic collagen or in perivascular aggre- 
gates. The healing response was marked by exces- 
sive collagen deposits, i.e., scar tissue, and numerous 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


elongated fibroblasts in close association with the 
net fragments. All of the above lesions were local- 
ized within the dense collagen layer of superficial 
dermis, i.e., in the upper stratum compactum. 

As mentioned by Jones (1993), although skin 
lesions have been observed worldwide on a large 
assemblage of fishes, only scattered reports exist of 
fish with healing wounds that were caused by net 
contact. With few exceptions (e.g., see Hampton et 
al. 1991; Jones 1993), the majority of these reports 
concerned net contact injuries to various flatfishes 
(e.g., see Shelton and Wilson 1973; Bucke et al. 
1983; Dethlefsen 1990) or salmonids (e.g., see 
Taylor 1985; Hansen 1988). The healing of teleosts 
after net contact injuries often consists of scrapes 
about the head, operculum, and along the trunk as 
well as lesions on and about fins (e.g., see 
Sindermann et al. 1978; Jones 1993). 

To our knowledge, this is the first report of a heal- 
ing net contact injury in a shark, and this case is 
furthermore unusual because it documents a net 
escapement event resulting in a shark becoming per- 
manently encircled by an almost totally embedded 
mesh square. Many reports exist of fishes having 
been permanently encircled by a wide variety of 
man-made items (e.g., Gudger 1928; Schwartz 1963; 
Honma 1964). As noted by Overstreet and Lyles 
(1974), part of the interest in these reports stems 
from the fact that encircled fishes can sometimes 
avoid natural predators, capture prey, and otherwise 
survive even though encirclement can cause severe 
deformities. Regarding sharks, Overstreet (1978) and 
Bird (1978) reported on the encirclement of several 
species of carcharhinids (Carcharhinidae) by box 
straps. Details provided by Bird (1978) indicated that 
this type of encirclement can cause significant 
lesions that may eventually result in death, and 
although signs of healing were observed, none of the 
encirclement lesions was fully resolved. 

Regarding the present instance it is possible that 
the Porbeagle tore itself free from a gillnet only to 
have the remaining mesh square eventually embed 
itself in its snout as the fish grew larger. It is also 
possible that a fisherman may have partially untan- 
gled and released this toothy fish from a net target- 
ing other species. The literature contains reports of 
Porbeagles being taken as by-catch in gillnet opera- 
tions targeting ground fish (Bigelow and Schroeder 
1953) and also of these sharks making a mess of nets 
not meant for them (Lineaweaver and Backus 1970). 
With adult female Porbeagles commonly reaching 
152 to 219 cm in total length and possibly up to 
370 cm in total length (Compagno 1984) it is likely 
that the 182 cm fork length individual reported on 
here was mature, but still possessed some remaining 
scope for growth. How or if the embedded mesh 
square impacted the growth of this fish is unknown. 
However, its survival and the healing of its net 


2001 


wound is significant given that net injuries have been 
shown to lead to an increased susceptibility to viral 
and bacterial diseases in some fish (e.g., see 
LeTendre et al. 1972; Dethlefsen 1990). Certainly 
the Porbeagle reported on here was unlucky, being 
netted and freed from a net only to later be caught by 
a longline, and this case may represent a conse- 
quence of high fishing pressure in the northwestern 
Atlantic. 


Acknowledgments 

We thank captain Albert Lawrence, first mate 
Perry Morris, and the “boys” of the Aquatic Pioneer 
for their help in the field and for allowing the junior 
author to participate in their fishing operations; H. L. 
“Wes” Pratt, Jr. (National Marine Fisheries Service) 
for logistic support; S. A. Bullard (Gulf Coast 
Research Laboratory) for assistance with literature; 
and Todd Stailey and Jeff Worley (both Tennessee 
Aquarium) for help with the figure. 


Literature Cited 

Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the 
Gulf of Maine. Fishery Bulletin 74, Fishery Bulletin of 
the Fish and Wildlife Service, Volume 53, U.S. 
Government Printing Office, Washington, D.C. 577 
pages. 

Bird, P.M. 1978. Tissue regeneration in three car- 
charhinid sharks encircled by embedded straps. Copeia 
1978: 345-349. 

Block, B. A., and F. G. Carey. 1985. Warm brain and eye 
temperatures in sharks. Journal of Comparative Physiol- 
ogy B 156: 229-236. 

Bucke, D., S. W. Feist, M. G. Norton, and M.S. Rolfe. 
1983. A histopathological report of some epidermal 
anomalies of Dover sole, Solea solea L., and other flat- 
fish species on coastal waters off South-east England. 
Journal of Fisheries Biology 23: 565-578. 

Carey, F. G., J. G. Casey, H. L. Pratt, D. Urquhart, and 
J. E. McCosker. 1985. Temperature, heat production 
and heat exchange in lamnid sharks. Memoirs of the 
Southern California Academy of Sciences 9: 92-108. 

Carey, F. G., and J. M. Teal. 1969. Mako and porbeagle: 
warm bodied sharks. Comparative Biochemistry and 
Physiology 28: 199-204. 

Castro, J. I., C. M. Woodley, and R. L. Brudek. 1999. A 
preliminary evaluation of the status of shark species. 
Fisheries Technical Paper number 380, Food and Agri- 
culture Organization of the United Nations, Rome. 72 
pages. 

Compagno, L. J. V. 1984. Sharks of the World: Hexan- 
chiformes to Lamniformes. Fishery Synopsis number 
125, Volume 4, Part 1. Food and Agriculture Organ- 
ization of the United Nations, Rome. 249 pages. 

Dethlefsen, V. 1990. Ten years fish disease studied of the 
Institut fiir Kusten- und Binnenfischerei. Archiv fiir 
Fischereiwissenschaft 40: 119-132. 

Gudger, E. W. 1928. A mackerel (Scomber scombrus) 


NOTES 


509 


with a rubber band rove through its body. American 
Museum Novitates 310: 1-6. 

Hampton, J., T. Murray, and K. Bailey. 1991. South 
Pacific albacore observer programme on troll vessels, 
1989-1990. South Pacific Commission Tuna and Bill- 
fish Assessment Programme Technical Report number 
25. 25 pages. 

Hansen, L. P. 1988. Status of exploitation of Atlantic 
salmon in Norway. Pages 143-161 in Atlantic salmon: 
planning for the future. Edited by D. Mills, and D. Pig- 
gins. Proceedings of the 3rd International Atlantic 
Salmon Symposium. Biarritz, France 1986. 

Honma, Y. 1964. Notes on the specimens of halfbeaks, 
Hemirhamphus sajori (T. & S.), with heads or trunks 
bound with rubber bands. Collecting and Breeding 
26(2): 1-3. [in Japanese] 

Hurley, P. 1997. Review of the porbeagle shark fishery in 
Atlantic Canada. Pages 12-13 in The Shark Tagger 1996 
Summary. Edited by N. E. Kohler, L. J. Natanson, H. W. 
Pratt, Jr., P. A. Turner, and R. Briggs. Cooperative Shark 
Tagging Program, National Marine Fisheries Service, 
Narragansett, Rhode Island. 

Jones, J. B. 1993. Net damage injuries to New Zealand 
hoki, Macruronus novaezelandiae. New Zealand Journal 
of Marine and Freshwater Research 27: 23-30. 

LeTendre, G. C., C. P. Schneider, and N. F. Ehlinger. 
1972. Net damage and subsequent mortality from furun- 
culosis in smallmouth bass. New York Fish and Game 
Journal 19: 73-82. 

Lineaweaver, T.H., III, and R. H. Backus. 1970. The 
Natural History of Sharks. J. B. Lippincott Company, 
Philadelphia, Pennsylvania. 256 pages. 

Overstreet, R. M. 1978. Marine Maladies? Worms, 
Germs, and Other Symbionts from the Northern Gulf of 
Mexico. MASGP-78-021, Mississippi-Alabama Sea 
Grant Consortium, Ocean Springs, Mississippi. 140 
pages. 

Overstreet, R. M., and C. H. Lyles. 1974. A rubber band 
around an Atlantic croaker. Gulf Research Reports 4: 
476-478. 

Reif, W-E. 1985. Squamation and ecology of sharks. 
Courier Forschungsinstitut Senckenberg 78: 1-255. 

Schwartz, F. J. 1963. Bluefish from Chesapeake Bay 
deformed by plastic band. Chesapeake Science 4: 196. 

Shelton, R. G., and K. W. Wilson. 1973. Epidermal 
lesions in Irish Sea flatfish. Nature 241: 140-141. 

Sindermann, C. J., J. J. Ziskowski, and V. T. Anderson, 
Jr. 1978. A guide for the recognition of some disease 
conditions and abnormalities in marine fishes. National 
Marine Fisheries Service Technical Series Report num- 
ber 14: 1-60. 

Taylor, S. G. 1985. Scarred Pacific salmon Oncorhynchus 
spp., at freshwater recovery sites in southeastern Alaska. 
Marine Fisheries Review 47: 39-42. 

Wolf, N. G., P. R. Swift, and F. G. Carey. 1988. Swim- 
ming muscle helps warm the brain of lamnid sharks. 
Journal of Comparative Physiology B 157: 709-715. 


Received 6 July 2000 
Accepted 28 December 2001 


510 THE CANADIAN FIELD-NATURALIST Volts 


Hyperthermia Induced Mortality of Gravid Snapping Turtles, 
Chelydra serpentina, and Eggs in a Wood Chip Pile 


SHANE R. DE SOLLA!, DOUGLAS CAMPBELL2, and CHRISTINE A. BISHOP?4 


1158 Herkimer Street, Hamilton, Ontario L8P 2H4 Canada 

2Ontario Veterinary College, University of Guelph, Guelph, Ontario N1G 2W1 Canada 

3Canadian Wildlife Service, Environment Canada, Canada Centre for Inland Waters, P.O. Box 5050, Burlington, Ontario 
L7R 4A6 Canada 

4 Present address: Canadian Wildlife Service, Environment Canada, 5421 Robertson Road, Delta, British Columbia 
V4K 3N2 Canada 


de Solla, Shane R., Douglas Campbell, and Christine A. Bishop. 2001. Hyperthermia induced mortality of gravid 
Snapping Turtles, Chelydra serpentina, and eggs in a wood chip pile. Canadian Field-Naturalist 115(3): 510-512. 


Two gravid female Snapping Turtles (Chelydra serpentina) were observed attempting to nest in a composting wood chip 
pile, on the shoreline of the west end of Coote’s Paradise, Hamilton, Ontario, on 7-10 June 1999. Instead of laying eggs, 
both females buried themselves within the wood chip pile, and were found dead the following day. An autopsy revealed 
that both females were still gravid, and that the likely cause of death was hyperthermia and dehydration. Four clutches of 
eggs were also found in the wood chip pile after the cessation of nesting; one clutch was incubated in laboratory conditions 
but none of the eggs hatched. Subsequent dissection of all of the clutches revealed no development. Subsurface tempera- 
tures from the wood chip pile on 10 July 1999 ranged from 28 °C to 60 °C. On 8 August 2000, one clutch was found in the 
same wood chip pile, in which some eggs successfully hatched when incubated artificially. Artificial compost heaps near 


nesting sites may pose a risk for nesting turtles, through both adult mortality and reduced hatching success. 


Key Words: Snapping Turtle, Chelydra serpentina, Hamilton Harbour, composting, mortality, oviposition, wood chip. 


Snapping Turtles (Chelydra serpentina) nest in a 
variety of artificial substrates, such as dams, railway 
and road embankments (Loncke and Obbard 1977), 
and vegetable gardens (de Solla, unpublished data). 
In many parts of their range, the majority of females 
may nest in artificial substrates because natural areas 
may not be available, and they may travel consider- 
able distances to nest on embankments or dams 
(Obbard and Brooks 1980). There are no published 
reports of females dying of causes other than preda- 
tion and road mortality during oviposition, regard- 
less of the substrate. We report here the death of two 
gravid Snapping Turtles during a nesting attempt, 
and the failure of four clutches of eggs, in a com- 
posting wood chip pile in Hamilton, Ontario. 


Study Site 

The turtles were found in a single 6m X 18m 
pile of wood chips located approximately 30m from 
the northwestern shoreline of Coote’s Paradise, 
Hamilton, Ontario (43°17’ N, 79°53’ W), a 90 ha 
eutrophic wetland at the western end of Hamilton 
Harbour on Lake Ontario. There is a large popula- 
tion (>400; Galbraith et al. 1988) of Snapping 
Turtles in this wetland, despite exposure to 
organochlorine pesticides and other chemicals 
(Bishop et al. 1998; de Solla et al. 1998). 

Along this shoreline, an average of approximately 
40 nests has been observed annually from 1986 to 
1999 (Bishop and de Solla, unpublished data). Most 
females nested either in a large community vegetable 
garden, or in the wood chip pile, although a few 
females nested in other areas such as grass fields, 


embankments, or gravel roads. Snapping Turtles 
have been observed attempting to nest in the wood 
chip pile every year from 1986-2000, some of whom 
successfully deposited eggs. Unlike the nests at the 
vegetable garden, no occurrences of nest depredation 
have been observed at the wood chip pile, nor any 
successfully hatched nests. No deaths of nesting 
females were detected by the authors, or otherwise 
reported, from 1986-2000, with the exception of ani- 
mals run over by motor vehicles or deliberately 
killed by gardeners. 

It is unknown how long the wood chip pile has 
been present. The last known time an appreciable 
amount of new material had been added was in 1994. 
Although the surface and the core of the pile were 
poorly decomposed, in 1999 there was a layer that 
extended from approximately 2 cm depth down as 
deep as 30 cm depth that was well composted and 
soil-like. Fungi were present throughout and large 
numbers of invertebrates occurred where conditions 
were favourable. In 2000, the wood chip pile was 
much reduced, and was only about half of its size in 
Oe). 


Site Observations 

The area was searched for nests for about three to 
five hours at a time, both in the morning and 
evening, from 31 May to 2 June 1999, and 5 June to 
11 June 1999. The wood chip pile was rarely unob- 
served for longer than 30 minutes during these times, 
and often would be under almost continuous obser- 
vation. On 7 June 1999, at 1950 hrs a large female 
Snapping Turtle was sighted at the wood chip pile, 


2001 


and by 2010 hrs was buried within it. At 2050 hrs, 
she had moved and was buried in a different location 
in the pile, with her nose and the top of her head 
exposed. No change in position was seen when 
observed on 8 June at 0800 and again at 2240 hrs. 
The ambient air temperature that day reached 31.8 
°C. At 0740 hrs on 9 June, the turtle was dug out of 
the pile and was found to be dead, with the carcass 
appearing bloated. Eggs were palpable within the 
abdomen. The body was placed in a freezer at -20°C 
at approximately 1145 hrs on 9 June. 

On 9 June, a second female was found completely 
buried in the wood chip pile at 0800 hrs, approxi- 
mately 3m from the location of the first female. 
Ambient air temperature that day reached 28.5°C. 
This turtle was partially dug out later that evening, 


but she was still alive. The first author reburied her 


in the position in which she was found, with an 
approximately 10-15 cm thick layer of wood chips 
covering the carapace. At 2150 hrs, she was still 
buried in the same position. At 1040 hrs, 10 June she 
was excavated, found to be dead, and was placed in a 
freezer at -20°C at approximately 1200 hrs. 


Post-Mortem Results 

The post-mortem examination of carcasses of both 
dead turtles were performed at the Canadian 
Cooperative Wildlife Health Centre (CCWHC) labo- 
ratory in Guelph, Ontario, to determine cause of 
death. Both turtles were kept frozen until the necrop- 
sy. The carcasses were thawed at room temperature 
and a necropsy examination was done on 24 June. 
Both turtles were in fair body condition, with moder- 
ate stores of fat. Turtle A (found dead 9 June 1999) 
weighed 6.2 kg, with a carapace length of 31.0 cm. 
The carcass was autolysed. The stomach was empty 
except for a small quantity of brown fluid. The lungs 
were congested and the urinary bladder was empty. 
The uterus was intact and contained 45 calcified 
eggs. 

Turtle B (found dead 10 June 1999) weighed 7.25 
kg and had a carapace length of 31.5 cm. The carcass 
was extremely autolysed with all viscera in a state of 
advanced decomposition. The uterus had ruptured 
post-mortem and 54 eggs lay free in the abdominal 
cavity. 

Sections of heart, lung, liver, spleen, kidney, 
adrenal, brain, skeletal muscle, thyroid, intestine and 
stomach were taken from Turtle A and fixed in 10% 
neutral buffered formalin, sectioned for histology 
and stain with hematoxylin and eosin using routine 
methods (Luna, 1968). Sections of skeletal muscle, 
lung and kidney from Turtle B were collected and 
processed in similar fashion. 

Significant histological abnormalities detected in 
the two turtles included acute multifocal myocardial 
degeneration in Turtle A and acute degeneration of 
skeletal muscle myofibres and acute degeneration 
and mineralization of renal tubular epithelium in 


NOTES 


511 


Turtle B. These changes are consistent with dehydra- 
tion and hyperthermia. In the absence of other signif- 
icant gross or microscopic lesions, or evidence of 
infectious disease, it was concluded that death was 
due to hyperthermia. 


Temperature and the fate of clutches 

Temperature readings were taken with a mercury 
stick thermometer on 10 July in the wood chip pile 
in areas that had the greatest concentrations of turtle 
tracks and holes excavated by turtles. Readings were 
taken between 1300 and 1400 hrs on a sunny day, 
and the air temperature was 27.5°C. Temperature 
readings were taken at the top of the woodchip pile, 
1-2 m above the bottom of the pile (locations 1-5, 
Table 1), and near the base of the pile, approximate- 
ly 0.5 m above the bottom of the pile (locations 6-9, 
Table 1). Unless otherwise noted, each temperature 
reading was taken at about 10 cm deep in the wood 
chips, at approximately the depth of the centre of 
gravity of the turtles that were found dead, and was 
also the approximate depth of the top of the nests. 
Temperatures at the top of the pile were appeared to 
be higher than at the bottom, and the wood chips 
were drier to the touch than at the base. 

Although no more dead turtles were found, one 
clutch of eggs (N=34) was found and excavated in 
mid-July. The eggs appeared grossly abnormal with 
blotchy areas of discoloration. The temperature was 
37°C where this clutch was located, and 46°C 
approximately 20cm from the clutch at the same 
depth. The clutch was incubated in a water and ver- 
miculite mixture (1.1:1 mass ratio) at 25°C until late 
fall; no eggs hatched. There was no evidence of 
development of the embryos in these eggs, upon dis- 
section. 

On 15 October 1999 the wood chip pile was sifted 
with rakes and three more clutches were found with 
all the eggs intact. No viable eggs were found, and 
some eggs had fungal growth on their shells. 
Internally, only yolk was present with no develop- 
ment of embryos. 


TABLE |. Temperature readings from different locations on 
the woodchip pile in early July 1999. The depths of the 
readings were approximately 10 cm from the surface except 
where otherwise noted. 


Top of Temp. (°C) Base of Temp. (°C) 
Woodchip pile Woodchip pile 
[le 42.0 6 35.0 
2 45.5 7 28.0 
Se 43.0 8 39.0 
4 S255 9 38.0 
5 at 10cm 43.0 
5 at 20 cm 56.0 
5 at 30 cm 62.5 


*Tocations where clutches were found. 


a2 


None of the four clutches had been disturbed and 
eggs were desiccated in only one clutch. All four 
clutches were found near the top of the wood chip 
pile, where both temperature and moisture were 
greatest. 

On 8 August 2000 the wood chip pile was sifted 
with rakes, and one clutch was found. The wood 
chip pile was greatly reduced in size compared to 
1999, and there appeared to be less well composted 
material. Although no temperature measurements 
were taken, the temperature of the compost appeared 
to be lower than in 1999. One egg from the clutch 
was already dead and broken, but the rest appeared 
to be potentially alive. The clutch was used in an 
unrelated project, and was incubated in a water and 
vermiculite mixture (1.1:1 mass ratio) at 25°C at the 
University of Guelph, and a large (but unknown) 
proportion of the eggs successfully hatched (S. 
Ashpole, personal communication). 


Discussion 

Both turtles died in the wood chip pile, without 
having deposited any eggs. Normally, Snapping 
Turtles do not bury completely themselves during 
Oviposition, although some turtles that were 
observed to lay eggs in the wood chip pile had their 
posterior end lightly covered with wood chips. Thus, 
the act of burial by the two gravid females was 
unlikely directly linked to oviposition. There was no 
apparent effort by the turtles to abandon the nesting 
attempt and leave the wood chip pile, based upon the 
available observations of their behaviour. This fail- 
ure to leave the nest site may have been due to the 
clinical effects of hyperthermia, to some behavioral 
cause, or to some other undetected factor. 

The temperatures recorded in the wood chip pile 
in 1999 exceeded the critical thermal maximum for 
both adults and eggs of this species (Cloudsley- 
Thompson 1971; Yntema 1976). In this setting, with 
no method of heat dissipation available, and an addi- 
tional increment of heat stress due to the muscular 
exertion of digging, the animals were extremely vul- 
nerable to heat stroke (Simon 1993). Consequences 
of hyperthermia include cardiovascular failure, 
metabolic abnormalities and neurological impair- 
ment, including stupor (Simon 1993). These effects 
may be sufficient to account for the turtles’ failure to 
leave the nest site. The removal of some compost 
from the wood chip pile in 2000 may have reduced 
the temperature to a sub-lethal level. 

High temperatures in the wood chip pile likely 
make it a reproductive sink for turtles because of 
clutch failure. Mortality of adult females on the nest 
is likely a rare event. However, the mortality of adult 
females would have a much larger effect on the pop- 
ulation dynamics of Snapping Turtles than decreased 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


hatching success (Cunnington and Brooks, 1995). 
Female Blanding’s Turtles (Emydoidea blandingii) 
and Painted Turtles (Chrysemys picta) have been 
seen in the study area, but neither was seen to nest in 
the pile. Improved success of nesting attempts and a 
reduced risk of death of adult animals could be 
achieved by blocking access to the pile during the 
nesting period. 


Acknowledgments 

We thank Tana McDaniel and Sara Ashpole for 
help looking for nests in the woodchip pile, Sara 
Ashpole for incubating the clutches, Ontario Hydro 
for permission to enter their property, and the 
National Climatic Data Center for the air tempera- 
ture data. 


Literature Cited 

Bishop C. A., P. Ng, K. E. Pettit, S. W. Kennedy, J. J. 
Stegeman, R. J. Norstrom, and R. J. Brooks. 1998. 
Environmental contamination and developmental abnor- 
malities in eggs and hatchlings of the common snapping 
turtle (Chelydra serpentina serpentina) from the Great 
Lakes-St Lawrence River basin (1989-91). Environ- 
mental Pollution 101: 143-156. 

Cloudsley-Thompson, J. L. 1971. The temperature and 
water relation in reptiles. Merron Publications Co. Ltd., 
Watford Herts, England. 

Cunnington, D. C. and R. J. Brooks. 1996. Bet-hedging 
theory and eigenelasticity: a comparison of the life histo- 
ries of loggerhead sea turtles (Caretta caretta) and snap- 
ping turtles (Chelydra serpentina). Canadian Journal of 
Zoology 74: 291-296. 

de Solla,. S. R., C. A. Bishop, G. Van Der Kraak, and R. 
J. Brooks. 1998. Impact of organochlorine contamina- 
tion on levels of sex hormones and external morphology 
of common snapping turtles (Chelydra serpentina ser- 
pentina) in Ontario, Canada. Environmental Health 
Perspectives 106: 253—260. 

Galbraith, D. A., C. A. Bishop, R. J. Brooks, W. L. 
Simser, and K. P. Lampman. 1988. Factors affecting 
the density of populations of common snapping turtles 
(Chelydra serpentina serpentina). Canadian Journal of 
Zoology 66: 1233-1240. 

Loncke, D. J. and M. E. Obbard. 1977. Tag success, 
dimensions, clutch size and nesting site fidelity for the 
snapping turtle, Chelydra serpentina, (Reptilia, 
Testudines, Chelydridae) in Algonquin Park, Ontario, 
Canada. Journal of Herpetology 11: 243-244. 

Obbard, M. E. and R. J. Brooks. 1980. Nesting migra- 
tions of the snapping turtle (Chelydra serpentina). 
Herpetologica 36: 158-162. 

Simon, H. B. 1993. Hyperthermia. New England Journal 
of Medicine 329: 483-487. 

Yntema C. L. 1976. Effects of incubation temperatures 
on sexual differentiation in the turtle, Chelydra serpenti- 
na. Journal of Morphology 150: 453-462. 


Received 31 July 2000 
Accepted 16 November 2001 


2001 


NOTES 


515 


A Significant New Record of the Pygmy Shrew, Sorex hoyi, on the 


Montana-Alberta Border 


PAUL HENDRICKS 


Montana Natural Heritage Program, 909 Locust Street, Missoula, Montana 59802, USA; email-phendricks @state.mt.us 


Hendricks, P. 2001. A significant new record of the Pygmy Shrew, Sorex hoyi, on the Montana-Alberta border. Canadian 


Field-Naturalist 115(3): 513-514. 


Remains of an adult Pygmy Shrew, Sorex hoyi, skull were recovered from a deteriorated raptor pellet on 20 July 2000 at 
Wild Horse Lake, Hill County, Montana about 2.4 km S of the Montana-Alberta border and 17.7 km WSW of the 
Montana-Saskatchewan border. Wild Horse Lake is within a large hiatus in the known distribution of this Boreal- 
Cordilleran shrew species, indicating there is a high probability that a relict population occurs in the immediate vicinity of 
Wild Horse Lake or in one of the isolated forested uplands (Sweet Grass Hills or Bears Paw Mountains in Montana; 


Cypress Hills in Alberta-Saskatchewan) in this prairie region. 


Key Words: Pygmy Shrew, Sorex hoyi, Montana, distribution. 


The Pygmy Shrew (Sorex hoyi) is widespread 
across boreal and subarctic North America, with 
populations also extending south in the United States 
along the Rocky Mountains in the west and Appala- 
chian Mountains in the east (Long 1974; Diersing 
1980; Hall 1981). Although found in a variety of 
microhabitats (including bogs, marshes and dry 
grassy clearings), the species generally seems to be 
associated with mesic forested habitats (Long 1972; 
van Zyll de Jong 1983) and riparian corridors con- 
nected to these. In Montana, the Pygmy Shrew so far 
has been found in the western Rocky Mountain 
region near, or west of, the Continental Divide 
(Foresman 1999) usually in forested habitats, but 
also in sagebrush-steppe at one collection site. 

On 20 July 2000 I found the remains of a Pygmy 
Shrew skull (intact lower jaw plus upper jaw and skull 
except the cranium) in a deteriorated raptor pellet at 
Wild Horse Lake, Hill County, Montana (48°59’N, 
110°10’W; 853 m elevation), about 2.4km S of the 
Montana-Alberta border and 17.7 km WSW of the 
Montana-Saskatchewan border, in T37N, R12 E, Sect. 
8NENE. The shrew was an adult, based on the degree 
of tooth wear (K. Foresman, personal communica- 
tion); M?-M? breadth (breadth across the second 
molars) and molariform toothrow length (P?-M?) were 
3.81mm and 3.62 mm, respectively. Also in the pellet 
were the remains of one Meadow Vole (Microtus 
pennsylvanicus). 

The area where the skull was retrieved is adjacent to 
an extensive lakebed depression largely of treeless 
prairie habitat of a variety of native grassland species 
(e.g. Bouteloua gracilis, Carex filifolia, Koeleria 
macrantha, Poa secunda, Stipa comata), scattered sil- 
ver sage (Artemisia cana), introduced species such as 
crested wheatgrass (Agropyron cristatum), and crop- 
land of wheat (Triticum aestivum). The lake basin has 
experienced several years of drought, supporting no 
standing water by early summer, but during some 
years in the early 1990’s the lakebed held water and 


was surrounded by a lush growth of prairie vegetation 
(L. Lund personal communication). 

The Wild Horse Lake site falls within a large 
northward hiatus in the known distribution of the 
Pygmy Shrew, between the known limits in the 
Rocky Mountains of western Montana and Alberta 
and the woodlands of eastern North Dakota and 
South Dakota (Long 1974; Hall 1981). Nearest 
records (van Zyll de Jong 1983; Smith 1993; 
Foresman 1999) are Helena, Montana (about 315 km 
SW), Polebridge, Montana (about 315 km W), near 
Calgary, Alberta (about 340 km NW), and Swanson, 
Saskatchewan (about 418 km NE). 

Because the Pygmy Shrew skull was found in a 
raptor pellet, the exact location where the shrew was 
caught is unknown. The most obvious possibility is 
that the shrew was captured in the immediate vicinity 
of Wild Horse Lake in mesic grassy habitat in associ- 
ation with Meadow Voles. However, a migrating rap- 
tor could have ejected the pellet containing the shrew 
skull a considerable distance from where the shrew 
was captured, depending on the mean daily distance 
moved and length of the meal to pellet interval. 

Little published data is available on daily move- 
ments of raptors during migration or their meal to 
pellet intervals. Schmutz et al. (1996) reported that 
Swainson’s Hawks (Buteo swainsoni) traveled 
straight-line distances of about 190 km/day during 
southward migration, and Beske (1982) observed 
juvenile Northern Harriers (Circus cyaneus) that 
moved 14-106 km/day. Both species nest in the 
vicinity of Wild Horse Lake and the harrier is also a 
known shrew predator (Bent 1937). However, mean 
meal to pellet intervals for these two raptor species 
are not available. For Short-eared Owls (Asio flam- 
meus) and Great Horned Owls (Bubo virginianus), 
species also present at Wild Horse Lake and known 
to feed on shrews and voles, ingested bones of prey 
take about 10-16 h to be ejected in a pellet (Grimm 
and Whitehouse 1963, Duke et al. 1976). 


514 


Assuming a hypothetical distance of 190 km/day 
moved during migration, a 10-16 h meal to pellet 
interval, and a spring/autumn daily diurnal period of 
16 h, the Pygmy Shrew could have been carried 
119-190 km from where it was caught, if it was con- 
sumed immediately in the morning before migration 
commenced for the day. The above calculated trans- 
port distances still leave the Pygmy Shrew found at 
Wild Horse Lake far removed from the nearest 
known locations 315—418 km distant; the distance 
transported during migration could be much less if 
the shrew was captured by an owl. 

Pygmy Shrews are often found in mesic forested 
habitats (Long 1972; van Zyll de Jong 1983), there- 
fore I suggest that the specimen collected at Wild 
Horse Lake more likely came from one of the near- 
by forested uplands, such as the Sweet Grass Hills 
(about 72 km W), Bears Paw Mountains (about 82 
km S), or Cypress Hills (about 65 km N). Relict 
populations of Pygmy Shrews with little or no gene 
flow to the Rocky Mountain and boreal populations 
might persist in one or more of these isolated 
ranges, as appears to be the case for Dwarf Shrew 
(Sorex nanus) in the Sweet Grass Hills and Bears 
Paw Mountains (Thompson 1977). My suggestion 
is supported by limited data on skull measurements. 
M?-M? breadth in the Wild Horse Lake specimen 
was 3.81 mm which is well below the mean value 
of 3.93 mm for specimens from the Rocky 
Mountains of Alberta, British Columbia, Montana, 
and NE Washington (Diersing 1980). M?-M? 
breadth was also below the range of values for indi- 
viduals with equivalent molariform toothrow length 
in that sample. The Wild Horse Lake specimen 
could come from a population with a skull mor- 
phology significantly narrower than the main 
Rocky Mountain population. However, cranial size 
of shrews is known to diminish in old individuals 
(Pruitt 1954), and this alone could explain the 
exceptionally small M?-M? measurement of the 
Wild Horse Lake specimen. Clearly, additional sys- 
tematic sampling for Pygmy Shrews is desirable in 
the forested uplands of the Sweet Grass Hills, Bears 
Paw Mountains, and Cypress Hills, as well as at 
Wild Horse Lake. 


Acknowledgments 

The work during which the shrew specimen was 
found was funded by a contract to the Montana 
Natural Heritage Program from the Natural 
Resources Conservation Service, U.S.D.A; I thank 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Dave Heilig (NRCS) for promoting the survey. I also 
thank Kerry Foresman (University of Montana) for 
verifying my identification of the shrew, determining 
the age of the animal, and providing the skull mea- 
surements. The specimen (MTHP 4280) is now 
deposited in the Philip L. Wright Zoological 
Museum, the University of Montana. I benefited 
from the comments of two anonymous reviewers on 
an earlier version of this paper. 


Literature Cited 

Bent, A. C. 1937. Life histories of North American birds of 
prey. Part 1. United States National Museum Bulletin 
167. 

Beske, A. E. 1982. Local and migratory movements of 
radio-tagged juvenile harriers. Raptor Research 16: 
39-53. 

Diersing, V. E. 1980. Systematics and evolution of the 
Pygmy Shrews (subgenus Microsorex) of North Amer- 
ica. Journal of Mammalogy 61: 76-101. 

Duke, G. E., O. A. Evanson, and A. Jegers. 1976. Meal 
to pellet intervals in 14 species of captive raptors. Com- 
parative Biochemistry and Physiology 53A: 1-6. 

Foresman, K. R. 1999. Distribution of the Pygmy Shrew, 
Sorex hoyi, in Montana and Idaho. Canadian Field- 
Naturalist 113: 681-683. 

Grimm, R. J., and W. M. Whitehouse. 1963. Pellet forma- 
tion in a Great-horned Owl: a roentgenographic study. 
Auk 80: 301-306. 

Hall, E. R. 1981. The mammals of North America. Second 
edition, Volume IJ. John Wiley & Sons, New York, New 
York. 600 pages. 

Long, C. A. 1972. Notes on habitat preference and repro- 
duction in Pigmy Shrews, Microsorex. Canadian Field- 
Naturalist 86: 155-160. 

Long, C. A. 1974. Microsorex hoyi and Microsorex thomp- 
soni. Mammalian Species Number 33: 1-4. 

Pruitt, W. O., Jr. 1954. Aging in the Masked Shrew, Sorex 
cinereus cinereus Kerr. Journal of Mammalogy 35: 
35-39. 

Schmutz, J. K., C. S. Houston, and G. L. Holroyd. 1996. 
Southward migration of Swainson’s Hawks: over 
10,000 km in 54 days. Blue Jay 54: 70-76. 

Smith, H. C. 1993. Alberta mammals, an atlas and guide. 
Provincial Museum of Alberta, Edmonton, Alberta. 239 
pages. 

Thompson, L. S. 1977. Dwarf Shrew (Sorex nanus) in 
north-central Montana. Journal of Mammalogy 58: 248— 
250. 

van Zyll de Jong, C. G. 1983. Handbook of Canadian mam- 
mals. 1. Marsupials and Insectivores. National Museum of 
Natural Sciences. Ottawa, Ontario. 210 pages. 


Received 8 August 2000 
Accepted 24 May 2001 


2001 NOTES 515 


Observation of a Golden Eagle, Aguila chrysaetos, Attack on a 
Harlequin Duck, Histrionicus histrionicus, in Northern Labrador 


JOEL P. HEATH!25, GEOFF GOODYEAR?, and JOE BRAZIL* 


'Biopsychology Programme, Departments of Biology and Psychology, Memorial University of Newfoundland, St. John’s, 
Newfoundland A1B 3X9 Canada 

Present address: Centre for Wildlife Ecology/Behavioural Ecology Research Group, Department Biological Sciences, 
Simon Fraser University, 8888 University Drive, Burnaby, British Columbia V5A 1S6 Canada 

3Universal Helicopters, P.O. Box 529, Station C, Happy Valley — Goose Bay, Labrador, Newfoundland AOP 1C0 Canada 

‘Inland Fish and Wildlife Division, Department of Tourism, Culture and Recreation, P.O. Box 2006, Fortis Tower, Corner 
Brook, Newfoundland A2H 6J8 Canada 

°Corresponding author: joel@cs.mun.ca — 


Heath, Joel P., Geoff Goodyear, and Joe Brazil. 2001. Observation of a Golden Eagle, Aquila chrysaetos, attack on a 
Harlequin Duck, Histrionicus histrionicus, in northern Labrador. Canadian Field-Naturalist 115(3): 515-516. 


During an aerial survey on the Kingurutik River, northern Labrador, a Golden Eagle (Aquila chrysaetos) was observed 
attacking a female Harlequin Duck (Histrionicus histrionicus). Although the attack ended unsuccessfully, we believe this 
was only a result of disturbance by the presence of the helicopter. We overview previous reports of predation on Harlequin 
Ducks, and emphasize the need for further research investigating the importance of the influence of predators on popula- 


tions of Harlequin Ducks. 


Key Words: Harlequin Duck, Histrionicus histrionicus, Golden Eagle, Aquila chrysaetos, predation, Labrador 


On 21 June 1999, we conducted a survey for 
Harlequin Ducks on the Kingurutik River system (57 
0.5’ N, 63 4.0’ W) in northern Labrador using a Bell 
206L helicopter. This river system is known to hold 
moderate densities of both Harlequin Ducks and var- 
ious raptor species (Heath 2001). During this survey, 
we observed a Golden Eagle, Aquila chrysaetos, 
attacking a female Harlequin Duck, Histrionicus 
histrionicus, in mid-stream. It appeared that the 
Golden Eagle had captured the female; however, she 
managed to escape as the Golden Eagle, presumably 
startled by the helicopter, flew away. We believe that 
this would have been a successful predation event 
had the birds not been disturbed. We were unable to 
determine if the Harlequin Duck had been injured in 
the encounter; however, this is the first documented 
interaction between a Golden Eagle and Harlequin 
Duck. 

Current literature on birds of prey as predators of 
Harlequin Ducks is limited to incidental reports, but 
indicates that predation can occur on both adults and 
broods. In Forillon Park, Quebec, Brodeur et al. 
(1998) located two adult female Harlequin Ducks 
(carrying satellite radios) killed and consumed by a 
Red-tailed Hawk (Buteo jamaicensis) and Great 
Horned Owl (Bubo virginianus). In Hebron Fiord, 
northern Labrador, a white-phase Gyrfalcon (Falco 
rusticolus) was observed circling and stooping on 
two adult female Harlequin Ducks (Rodway et al. 
2000); however, the females avoided capture by div- 
ing (J. Gosse, personal communication). Bald Eagles 
(Haliaeetus leucocephalus) have been reported as a 
threat on both breeding and winter grounds (Dzinbal 
1982; McEneaney 1997; Robertson and Goudie 
1999). Smith (2000*) reports that 10 of 13 predation 


events on ducklings were most likely by raptors (4 by 
Red-tailed Hawk, | by Northern Goshawk Accipiter 
gentilis, 1 by unknown Owl spp., and 4 by unknown 
raptor spp.), and that fledging by Harlequin Ducks 
coincided with high numbers of raptors and their 
fledged young. Raven (Corvus corax), Arctic Skua 
(Stercorarius parasiticus) and several mammalian 
predators including Mink (Mustela vison), Marten 
(Martes americana) and Arctic Fox (Alopex lagopus) 
have also been identified as potential predators dur- 
ing the breeding season (Bengtson 1966, 1972, 
Kuchell 1977, Bruner 1997, Smith 2000*) and 
numerous Mink attacks have been observed on 
Harlequin Duck broods in southern Labrador (K. 
Squires and R. I. Goudie, personal communication). 

Despite these incidental reports, the influence of 
predation on breeding populations of Harlequin 
Ducks has yet to be addressed in the literature (but 
see Heath 2001 and forthcoming publications). 
Previous studies have emphasized that Harlequin 
Ducks may be primarily limited by prey availability 
on the breeding grounds (Bengtson and Ulfstrand 
1971; Gardarsson and Einarsson 1994; Rodway 
1998); however, the growing list of direct predation 
encounters suggests that the role of predation should 
receive further attention. 


Acknowledgments 

Funding for this project was provided by the 
Newfoundland and Labrador Provincial Wildlife 
Division via. JB, a Northern Scientific Training 


*See Documents Cited Section 


516 


Program grant to JPH and W. Montevecchi, and a 
World Wildlife Fund of Canada Endangered Species 
Recovery Fund grant to W. Montevecchi. Thanks to 
John Gosse (Parks Canada), Kelly Squires (Univer- 
sity of British Columbia) and R. Ian Goudie (Mem- 
orial University of Newfoundland) for providing 
information on their observations of predation 
encounters in Labrador. 


Documents Cited (marked * after date in text) 

Smith, C. M. 2000. Population dynamics and breeding 
ecology of Harlequin Ducks in Banff National Park, 
Alberta, 1995-1999. Unpublished Technical Report, 
Parks Canada, Banff National Park, Banff, Alberta, 
Canada. 


Literature Cited 

Bengtson, S.-A. 1966. Field studies on the Harlequin 
Duck in Iceland. Wildfowl 17: 79-94. 

Bengtson, S.-A. 1972. Breeding ecology of the Harlequin 
Duck Histrionicus histrionicus (L.) in Iceland. Ornis 
Scandinavica 3: 1-19. 

Bengtson, S.-A. and S. Ulfstrand. 1971. Food resources 
and breeding frequency of the Harlequin Duck Histrioni- 
cus histrionicus in Iceland. Oikos 22: 235-239. 

Brodeur, S., A. Bourget, P. Laporte, S. Marchant, G. 
Fitzgérald, M. Robert, and J.-P. L. Savard. 1998. 
Etude des déplacements du Canard Arlequin (Histrionicus 
histrionicus) en Gaspésie, Québec. Canadian Wildlife 
Service Technical Report Series Number 331, Québec 
Region, Ste-Foy, Québec, Canada. 

Bruner, H. J. 1997. Habitat use and productivity in the 
central Cascade Range of Oregon. M.Sc. thesis, Oregon 
State University, Corvallis, USA. 


THE CANADIAN FIELD-NATURALIST 


Vol. 1S 


Dzinbal, K. A. 1982. Ecology of Harlequin Ducks in 
Prince William Sound, Alaska, during summer. M.Sc. 
thesis, Oregon State University, Corvallis, USA. 

Gardarsson, A. and A. Einarsson. 1994. Responses of 
breeding duck populations to changes in food supply. 
Hydrobiologia 279/280: 15-27. 

Heath, J.P. 2001. Factors influencing breeding distribu- 
tions of Harlequin Ducks Histrionicus histrionicus in 
northern Labrador: a multi-scale approach. M. Sc. thesis, 
Memorial University of Newfoundland, St. John’s, 
Newfoundland, Canada. 

Kuchell, C. R. 1977. Some aspects of the behaviour and 
ecology of Harlequin Ducks breeding in Glacier National 
Park, Montana. M.Sc. thesis, University of Montana, 
Missoula, USA. 

McEneaney, T. 1997. Harlequins: noble ducks of turbu- 
lent waters. Yellowstone Science 5: 2-7. 

Robertson, G. J. and R. I. Goudie. 1999. Harlequin 
Duck (Histrionicus histrionicus) in The Birds of North 
America, number 466. Edited by A. Poole and F. Gill. 
The Birds of North America, Inc., Philadelphia, Pennsyl- 
vania, USA. 

Rodway, M.S. 1998. Activity patterns, diet, and feeding 
efficiency of Harlequin Ducks breeding in northern 
Labrador. Canadian Journal of Zoology 76: 902-909. 

Rodway, M.S., J. W. Gosse Jr., I. Fong, W. A. Monte- 
vecchi, S. G. Gilliland, and B. C. Turner. 2000. Abun- 
dance, habitat use, activity patterns and foraging behavi- 
our of Harlequin Ducks breeding in Hebron Fiord, 
Labrador in 1996. Canadian Wildlife Service Technical 
Report Series Number 350, Atlantic Region, Sackville, 
New Brunswick, Canada. 


Received 7 May 2001 
Accepted 17 December 2001 


A Remembrance of John Clifton Ward, 1921—1999 


DONALD R. FLOOK! and JosEPH E. BRYANT? 


1971 Isabella Point Road, Salt Spring Island, British Columbia V8K 1T7, Canada (Retired from the Canadian Wildlife 


Service) 


2447 Thessaly Circle, Ottawa, Ontario K1H 5W7, Canada — (Retired from the Canadian Wildlife Service) 


Flook, Donald R., and Joseph E. Bryant. 2001. A remembrance of John Clifton Ward, 1921-1999. Canadian Field- 


Naturalist 115(3): 517-519. 


John Clifton Ward (Clift) was born 10 January 
1921 in Sexmith, Alberta, and died 21 November 
1999 in Victoria, British Columbia. He was prede- 
ceased by his sisters Hazel McCann and Lona Waite, 
and his step-son Cleve. Mourning his passing are his 
wife, B.J., now living in Courtenay, British Columbia, 
his sons Scott in Canmore, Alberta, and Dean in 
Chesterville, Ontario, and step-daughter Cori-Lee in 
Victoria and their families including nine grand- 
children. 

Clift assumed responsibility early. He was only 7 
years old when his father died and 13 when his 
mother died. He and his sisters, both older, helped 
each other while he lived with them first in Grand 
Prairie and then in Hythe, Alberta, where he com- 
pleted high school. He joined the RCAF in 1940 at 
19 years of age and served overseas. 

On his discharge in 1945 he enrolled at the Uni- 
versity of Alberta and received his B.Sc. in zoology 
in 1949. He had an excellent undergraduate record 
and went on to complete an M. Sc. in zoology in 
1951, also at the University of Alberta. He special- 
ized in Fisheries Management and Conservation 
under Dr. R. B. Miller. His thesis was on the biology 
of Arctic Grayling in the southern Athabasca 
drainage. (Ward 1951*) 

Clift joined the Canadian Wildlife Service in 1951 
with the title of “Limnologist,” to work as an advisor 
to the National Parks Service, later named Parks 
Canada. He continued in that role until he retired in 
1976. During his career he was headquartered sequen- 
tially in Ottawa, Banff, Edmonton, Jasper, Calgary, 
Ottawa, and finally in Edmonton from where he 
retired. 

Clift was a kind and modest man. He loved his 
work and counted his co-workers among his best 
friends. He worked closely with park wardens and the 
relationship was mutually helpful. Clift was pleased 
to pass along his knowledge of fish and limnology 
and gratefully acknowledged information provided 
by wardens from their field observations. Wardens 
were always ready to help him with field projects and 
he was a favourite lecturer at the warden’s training 
schools where he frequently instructed. 

The official role of the Canadian Wildlife Service 


VA ~ aN, 


ie en 
sai iets 


Cliff Ward in retirement, 1982, Victoria, British Columbia. 
Photographer: Master Portriature of Victoria, 
British Columbia. 


in the National Parks was advisory to Parks Canada 
but during the period when Clift was assigned to the 
mountain parks he was called upon, because of his 
expertise, to oversee fish hatchery operations and to 
perform hands-on supervision of stocking and other 
fish management activities. That program contribut- 
ed greatly to angling opportunities in the parks. 
When trout hatcheries in Waterton Lakes and 
Banff National Parks were closed the production of 
trout for stocking in the mountain parks was concen- 
trated in Jasper. Clift devoted considerable attention 
to improving trout culturing methods and water sup- 
ply at the Jasper hatchery (Ward 1962d; Ward and 
Cuerrier 1967a; Ward 1969a*). Under Clift’s guid- 
ance the hatchery became a popular destination 


ah 


518 


among park visitors and he encouraged its use to 
inform the pubic about the aquatic environment 
(Ward and Cuerrier 1967a; Ward 1972a*). In retire- 
ment Clift met several people who spoke warmly of 
visits they had made to the Jasper trout hatchery. On 
one occasion when he was working on the British 
Columbia Ferry between Vancouver and Victoria a 
group of tourists from Arkansas recognized him 
from a visit to the Jasper hatchery. The hatchery had 
been the highlight of their trip to Jasper and for the 
rest of the voyage they and Clift discussed the hatch- 
ery in great detail. 

Clift and Jean-Paul Cuerrier of the Canadian Wild- 
life Service pioneered a technique of transporting live 
trout anaesthetized and sealed in plastic bags filled 
with pure oxygen and a small amount of water and 
cooled by ice in separate containers (Ward 1958, 
1962c; Ward and Cuerrier 1967c). The method was 
used successfully in National Park projects to trans- 
port live trout by plane, train, truck and packhorse. 
Survival was high and in contrast to tank transport no 
heavy equipment and much less water were required. 
The method has been used widely in trout manage- 
ment. 

Clift and his colleagues experimented with other 
innovative techniques. They oxygenated ice-covered 
lakes by pumping air or water in order to prevent 
winter kill of fish (Ward and Kooyman 1967b). They 
also tested the effects of toxicants applied to lakes to 
kill non-game fish in order to reduce predation on or 
competition with trout) Ward 1962a; Ward and 
Cuerrier 1967d). Clift also participated in a project 
in which television was used to observe conditions 
on the bottom of a deep lake. 

Clift evaluated experimental plantings in the 
mountain parks of some salmonid species not native 
to those parks (Ward 1962b, 1962c, 1974c*; Ward 
and Cuerrier 1967b; Ward and Kooyman 1967a). 
Species with which he experimented were: Atlantic 
salmon, Quebec red trout, golden trout, and splake, 
the latter a fertile hybrid of Lake Trout and Eastern 
Brook Trout developed in 1946 by J. E. Stenton, 
Park Warden in Banff. Among those plantings, only 
the splake established naturally reproducing popula- 
tions and they did so in two lakes. Because splake 
exhibited good survival, growth, longevity and sport- 
ing characteristics, they were also maintained for a 
number of years by repeated plantings in some lakes 
where they did not reproduce. 

The emergence of ecological awareness in the 
Canadian public that began in the late 1960s led to 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


changes in the philosophy and policies of Parks 
Canada. By the early 1970s the priority for manage- 
ment of park waters had shifted from enhancement 
of angling to preservation of aquatic ecosystems. 
Clift responded by undertaking comprehensive basic 
limnological inventories or Pukaskwa National 
Park, Ontario (Ward 1972b*, 1974b*, 1975c*) and 
La Mauricie National Park, Quebec (Ward 1975a*), 
writing his major report on the fishes of the moun- 
tain parks (Ward 1974c*), and performing other 
limnological investigations all with the same high 
degree of dedication and professionalism as he had 
exhibited in his earlier projects to enhance angling. 

By combining his 25 years of service with the fed- 
eral government and his five years of service over- 
seas in World War II, Clift was able to retire on full 
pension at the age of 55. “Retirement” did not mean 
an arm chair in front of the fire. All his life Clift had 
practised a broad variety of “extra curricular” activi- 
ties, including such things as hooking magnificent 
huge rugs. Retirement simply allowed more time to 
devote to his broad span of interests. First came a 
couple of courses in geology at Cariboo College in 
Kamloops, B.C. to increase the excitement of the 
extensive rock hounding and gold panning that he 
and his wife undertook all over B.C. He and his wife 
also served for four years on B.C. Ferries where Clift 
obtained Life Boat Certification for International 
Waters and, with an Industrial First Aid Certificate, 
became a First Aid Officer. To keep things active, 
Clift and his wife built a beautiful modern home on 
Salt Spring Island and a few years later another on 
Vancouver Island near Duncan. During some of their 
years at Duncan they operated the “JoyClift Coffee 
Stop” where Clift became locally famous as “the 
best muffin maker on the Island.” Unfortunately, 
Alzheimer’s disease gradually took over and Clift 
and his wife moved to Victoria to be closer to appro- 
priate medical services. Eventually this bright, active 
and all-round great guy succumbed to the ravages of 
the disease — a very sad ending to a productive and 
happy life. 


Acknowledgments 

We thank the librarians at the Environment 
Canada libraries, especially Jean-Francois Bélanger 
in Ottawa, Susan Blackman in Edmonton and 
Deborah Lister in Calgary for their assistance in 
locating the unpublished reports and we greatly 
appreciate the assistance of Clift’s widow, B. J. 
Ward, in the preparation of this remembrance. 


2001 


FLOOK AND BRYANT: JOHN CLIFTON WARD, 1921-1999 


319 


Publications and Selected Unpublished Documents by J. Clifton Ward 


Much of Clift’s work was not published but was 
presented in manuscript reports that are are deposit- 
ed in the libraries of Environment Canada in Ottawa 
and Edmonton and Parks Canada in Calgary. We 
hope that the following listing of his published 
papers and unpublished reports will make them bet- 
ter known and increase their use as sources of valu- 
able baseline information. 


Publications 

Ward, J.C. 1958. Alive, alive all. Intercom. Winter 1958. 
[Newsmagazine of the Department of Indian Affairs and 
Northern Development, Ottawa] Pages 7-8. 

Ward, J.C. 1962a. The use of fish toxicants in manage- 
ment of trout waters. Page 15 in Canadian Wildlife Ser- 
vice Research Progress Report — 1961. Queen’s Printer, 
Ottawa, Catalogue Number R65-5/1961. 

Ward, J.C. 1962b. Experimental introduction of Atlantic 
salmon with Vibert boxes. Page 16 in Canadian Wildlife 
Service Research Progress Report — 1961. Queen’s 
Printer, Ottawa, Catalogue Number R65-5/1961. 

Ward, J.C. 1962c. Splake plantings in the mountain 
parks. Page 16 in Canadian Wildlife Service Research 
Progress Report — 1961. Queen’s Printer, Ottawa, Cata- 
logue Number R65-5/1961. 

Ward, J.C. 1962d. Fish culture activities. Page 16 in 
Canadian Wildlife Service Research Progress Report — 
1961. Queen’s Printer, Ottawa, Catalogue Number R65- 
5/1961. 

Ward, J.C. 1962e. Transfer of trout in closed-circuit 
plastic bags. Page 17 in Canadian Wildlife Service 
Research Progress Report — 1961. Queen’s Printer, 
Ottawa, Catalogue Number R65-5/1961. 

Ward, J.C. 1962f. Limnological surveys of mountain park 
waters. Page 17 in Canadian Wildlife Service Research 
Progress Report — 1961. Queen’s Printer, Ottawa, Cata- 
logue Number R65-5/1961. 

Ward, J. C., and J.-P. Cuerrier, 1967a. Fish culture activ- 
ities. Page 29 in Canadian Wildlife Service ’66. Queen’s 
Printer, Ottawa, Catalogue Number R66-3766. 

Ward, J.C., and J.-P. Cuerrier. 1967b. Splake trout 
plantings in the mountain parks. Page 29 in Canadian 
Wildlife Service “66. Queen’s Printer, Ottawa, Catalogue 
Number R66-3766. 

Ward, J.C., and J.-P. Cuerrier. 1967c. Transfer of trout 
in closed-circuit plastic bags. Page 30 in Canadian Wild- 
life Service “66. Queen’s Printer, Ottawa, Catalogue 
Number R66-3766. 

Ward, J.C., and J.-P. Cuerrier. 1967d. Use of fish toxi- 
cants in management of trout waters. Page 31 in Can- 
adian Wildlife Service ‘66. Queen’s Printer, Ottawa, Cat- 
alogue Number R66-3766. 

Ward, J.C., and A.H. Kooyman. 1967a. Experimentai 
introductions of exotic species to national parks waters. 
Page 31 in Canadian Wildlife Service ‘66. Queen’s 
Printer, Ottawa, Catalogue Number R66-3766. 

Goldberg, E., J.-P. Cuerrier, and J.C. Ward. 1969. Lac- 
tate dehydrogenase ontogeny, paternal gene activation, and 
tetramer assembly in embryos of brook trout, lake trout, 
and their hybrids. Biochemical Genetics 2: 335-350. 

McAllister, D. E., and J.C. Ward. 1972. The deepwater 


sculpin, Myoxocephalus guadricornis thompsoni, new to 
Alberta, Canada. Journal of the Fisheries Research Board 
of Canada 29: 344-345. 


Unpublished Documents (marked* where cited) 

Ward, J.C. 1951. The biology of the Arctic Grayling in 
the southern Athabasca drainage. M. Sc. thesis, 
University of Alberta, Edmonton, Alberta. 71 pages. 


Canadian Wildlife Service Manuscript Reports 

[Copies of these reports are deposited variously in 

the libraries of Environment Canada in Ottawa (EC- 

O), Edmonton (EC-E) and Parks Canada in Calgary 

(PC-C).] 

Ward, J.C. 1960. A preliminary survey of the waters suit- 
able for angling in Banff National Park. 167 pages. (EC- 
O, EC-E). 

Ward, J.C. 1967. Annual progress report, mountain 
national parks, fiscal year 1966-67. 22 pages (EC-E). 

Ward, J.C. 1969a. Summary of the investigations carried 
out on the disease problem at Jasper National Park Malign 
River trout hatchery, 1969. 10 pages, (EC-O, EC-E micro- 
fiche). 

Ward, J.C. 1969b. Fishery investigations: mountain 
national parks; annual progress report, 1968-69 fiscal 
year. 28 pages, (EC-E, PC-C). 

Ward, J.C. 1970. Fishing waters in Mount Revelstoke 
National Park. 9 pages (EC-E, PC-C). 

Ward, J.C. 1972a. Two decades of fisheries management 
in the mountain national parks. 26 pages. (EC-E, PC-C, 
National Parks Documentation Centre, Ottawa.) 

Ward, J.C. 1972b. Aquatic resource survey Pukaskwa 
National Park, Ontario, 1972. 215 pages, (EC-O, EC-E). 
Ward, J.C. 1973a. Fish population and water depths of 
the marshes in Point Pelee National Park. 10 pages, (EC- 

O, EC-E). 

Ward, J.C. 1973b. Dundee marsh studies. 20 pages. (EC- 
O microfiche, EC-E Microfiche). 

Ward, J.C. 1974a. Salmon streams in Forillon National 
Park (EC-O microfiche, EC-E microfiche). Also listed as 
Riviéres 4 saumon dans le Parc National Forillon, 15 
pages. 

Ward, J.C. 1974b. Aquatic resources survey: Pukaskwa 
National Park, 1973 and 1974. 125 pages plus maps. 
(EC-O, EC-E). 

Ward, J.C. 1974c. The fishes and their distribution in the 
mountain national parks of Canada. 43 pages plus maps 
and appendices. (EC-O, EC-E, PC-C). 

Ward, J.C. 1975a. Aquatic resources surveys, 1973 and 
1974, La Mauricie National Park. 354 pages plus maps. 
(EC-O, EC-E). 

Ward, J.C. 1975b. Aquatic resources survey of Fairy and 
Goblin Lakes in Georgian Bay Islands National Park. 17 
pages plus maps. (EC-O, EC-E). 

Ward, J.C. 1975c. Aquatic resources of coastal streams in 
Pukaskwa National park. 45 pages plus maps. (EC-O, 
EC-E). 

Ward, J.C. 1975d. Progress report Waskesiu Lake wall- 
eye study, 1975. 41 pages. (EC-E). 

Ward, J.C. 1976. Waskesiu Lake walleye study: progress 
report 1976. 34 pages and maps. (EC-E). 


Accepted 17 February 2001 


Book Reviews 


ZOOLOGY 


Cuckoos, Cowbirds and Other Cheats 


By N. B. Davies. 2000. T and A. D. Poyser, London, UK. 
ix + 310 pp., illus. 


As a lad I frequently heard the bizarre tale of the 
(Common) Cuckoo. This tale was a relatively new 
discovery that lent itself to wiles of the photogra- 
pher. With the arrival of television, the struggles of 
the juvenile Cuckoo were just the fodder for the 
newly created BBC’s nature shows. Images of the 
tiny cuckoo baby’s struggles with its host’s young 
were just what this new medium wanted. So I 
thought I knew this narrative well. I was wrong. This 
piece of biological history involves more cunning 
and deceit than most of us could ever imagine. 

Nicholas Davies describes in great detail the well- 
studied life of the Common Cuckoo. This Eurasian 
bird has developed some remarkable tactics to live its 
life as a parasite. How it has achieved these strategies, 
why it has adopted this way of life and the response of 
the host victim are the major questions researched by 
the author. He analyses experiments he and others 
performed to test hypotheses proposed to explain each 
segment of behaviour. He is very careful not to jump 
at the most obvious explanation, but to lay out all pos- 
sibilities. He then examines these possibilities in light 
of logic and the observed facts. The result is an insight 
into a world of intrigue, betrayal, fakery, deception, 
and success on the part of the cuckoo. The hosts retal- 
iate with rejection, abandonment and, finally, accep- 
tance, when acceptance is the more productive tactic. 

The author then repeats his study for Great Spotted, 
Diederik, and Bronze cuckoos in the Old World. For 
the New World he looks at the cowbird clan, especial- 
ly the abundant Brown-headed Cowbird. In the case 
of the latter, he outlines how our human activities 
have benefited the cowbird at the expense of other 
species, and that their current abundance and the prob- 
lems they create are primarily our fault. 

For each species the author examines in detail the 
strategy that each bird uses to ensure success for its 
parasitic lifestyle. In fact, it involves just as much 
work, more stealth and speed and often considerably 
more bravery to be a parasite than to raise a brood of 
one’s own. Davies looks at the reasons why some 
species adopt this approach and reviews the advan- 
tages and disadvantages. Not suprisingly there are 
added benefits both to the individual and to the 
species for parasitism. There is also a point where it is 
more productive for the host to accept parasitism. 


Naturally hosts that have been parasitised for ages 
have developed defense strategies. 

So far Davies has dealt with the “classical” para- 
sites, birds that lay eggs in other species’ nests. There 
are however many more cases of parasitism amongst 
birds than I ever thought possible. I did remember 
reading of Wood Ducks and goldeneyes behaving 
parasitically, but never dreamed that swallows and 
Starlings, Moorhens and Coots are included in the 
avian freeloaders. Finding parasitic eggs that are laid 
by the same species has proven very difficult. 
Researchers have had to be innovative just to become 
aware of the possibility. It still seems hard to believe 
that a bird like a Cliff Swallow has been videotaped 
zipping next door to throw out her neighbour’s egg 
and replace it with one of her own. 

The author has added a final chapter on brood 
parasitism in other animals. The examples range 
from the bizarre Lake Tanganyika catfish, an obli- 
gate parasite on cichlid mouthbreeders, to the 
demonical slave-owning ants. 

The book is illustrated with line drawings by 
David Quinn and with both black-and-white and 
coloured photographs. Quinn’s drawings are first 
rate, showing accurate attitude, body shape, and fine 
detail while being gently artistic. Sadly the only two 
coloured illustrations by Quinn are on the dust cover. 
A few of Quinn’s colour plates inside the text would 
add much charm to this book. The photographs are 
good quality and very illustrative. 

The author’s writing style is clear and informative 
and this is an easy book to read. My sole criticism is 
that he repeats some of his key findings rather too 
often. I was also surprised that he used Hispaniola 
instead of the Dominican Republic and Haiti until I 
learned that that is still the name of the island itself. 

This is a book that should interest all naturalists. 
In many ways it reads as much like a detective story 
as a biology text. The parasites are a devious bunch 
and the author and his fellow researchers have had to 
be really creative with their methods of discovering 
and chronicling their strategies. I’m sure Agatha 
Christie would have been fascinated. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario K1J 6K5 Canada 


520 


2001 


BOOK REVIEWS 


The Nature of Hummingbirds: Rainbows on Wings 


By Harry Thurston. 1999. Greystone Books, Vancouver/ 
Toronto. 112 pp., illus. $34.95. 


Everyone who knows them loves hummingbirds. 
Tiny, beautiful, and feisty, they seem too fragile to be 
real. This author has captured their appeal both in 
words and stunning photographs, and hummingbird 
lovers can learn a great deal from the detailed profes- 
sional text. They are the second-largest family 
(Apodiformes) in the Western Hemisphere and are 
closely related to swifts. They breed from Alaska to 
Terra del Fuego. Of the 320 species (which are found 
exclusively in the Americas) only the Ruby-throated 
spends the summer in eastern Canada, although occa- 
sionally a vagrant of another species will stray east. 
The Rufous is common in British Coumbia, and a 
growing number of Anna’s are found in Vancouver, 
Victoria, and environs, some even over-wintering at 
well maintained feeders. The Bee hummingbird is the 
smallest bird in the world at 6.25 cm but you will 
have to go to Cuba to see that one. 

The text strikes a good balance between science 
and description of every aspect of their lives, and 
much of it is hard to believe: 1260 heartbeats and 
250 breaths per minute; a thimble-sized nest, bound 
with spider web; raisin-sized eggs; a migration of 
2500 km from the Canadian west coast to Mexico; a 
migration stage flight of 1000 km on 2g of fat; a 
wingbeat of 80 times per second, sometimes rising 


Albatrosses 


By W.L.N. Tickell. 2000. Yale University Press, New 
Haven and London. 448 pp., illus. + 52 plates. U.S.$60. 


It was a pleasant surprise to find there is a wealth 
of knowledge on albatrosses. I have often been dis- 
mayed to read of our lack of knowledge for some 
even relatively common birds. On reflection, I real- 
ized albatrosses nest on open, vegetation-free head- 
lands and islands in large colonies rather than scat- 
tered in deep woods or dense wetlands. This means 
counting nests and observing behaviour is simply a 
question of devoted work. Getting to the colony, 
however, can be quite a task. The weather too can be 
irksome; two researchers we met recently had 
endured two weeks of rain (on Albatross Island, 
South Georgia). Nevertheless, there are good data 
sets for most of the world’s albatross. Now this 
information is accumulated into a single volume. 

So if you want to know the details of the 
land—based portion of an albatross’s life this is the 
book for you. The author avoids the issue of species 
definition and treats each population separately. This 
way it is not an issue whether the birds are split or 
lumped in the future. For example, the Black-browed 


to 200. To help accomplish these feats, they have 
two extra pairs of ribs and an enlarged sternum. 
Since they have very little insulating down, they can 
reduce their body temperature from 97.5° F to 53.5° 
F to conserve heat overnight. Their metabolism is so 
efficient that we would have to eat half our weight in 
sugar to metabolize food at the same rate, but there 
would not be enough flowers to feed us in any case - 
nor would we be able to fly. Hummingbirds prefer to 
drink nectar from flowers, which is high sucrose and 
low fructose. Because the nectar is at the base of a 
tubular flower — not always red, but purple and blue 
as well — the flowers on which they feed have 
evolved for bird pollination and most of them 
exclude bees or butterflies as pollinators, but the 
hummingbirds are adaptive and eat insects if neces- 
sary. Unlike many other species, Hummingbird pop- 
ulations are healthy, partly because their fledging 
rate is higher than most birds — between 20 and 
50%, and perhaps partly because they are impossible 
for predators to catch when flying. 

This is a beautiful book with a good deal of seri- 
ous scientific information about a bird which thrills 
many people. If you have a passion for humming- 
birds, this is for you. 


JANE E. ATKINSON 


255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 


Albatross and the Campbell Island Black-browed 
Albatross (from Campbell Island, naturally) are list- 
ed separately. 

The book begins with information on the alba- 
trosses’ habitat. There are some fine descriptions of 
the southern, equatorial Pacific and north Pacific 
oceans themselves, in terms of current and winds 
that affect albatross. The portrayal of the relevant 
islands includes a location and description and pro- 
vides a history of the voyages of discovery (mostly 
from a European perspective). These fine birds were 
exploited for the feathers, quills, meat, and oil as 
soon as these islands became known. They were 
even used as target practice. Friends of humans such 
as rats, cats, and pigs, ravaged their islands. Mother 
Nature added storms, tempests, ice, and volcanic 
eruptions. Now albatrosses face the perils imposed 
by longlining for tuna. Despite all these desecrations 
we still have these aristocratic birds gracing the 
oceans skies. 

The author organizes the birds into three groups; 
southern, tropical, and northern albatrosses. He pre- 
sents an organized compilation of all the information 


522 


he could gather on each species. This includes anec- 
dotal evidence from voyagers, particularly from the 
early years, as well as more structured scientific 
studies. The most recent studies use electrical, radio, 
and satellite detectors to give some information on 
the birds’ activities away from the nest. 

While the author devotes considerable attention to 
the bird’s breeding history I found the information 
on geographic distribution of more immediate use. 
The island location is given for each colony and the 
colony positions are pinpointed on each island. An 
appendix gives the size of the population over the 
years for which data are available. This enabled me 
to compare my own estimates with more rigorously 
conducted counts. Typically my assessments were 
very low. This was truer where the terrain was rocky 
and birds could be hidden from the viewer. These 
chapters also contained information on albatross 
habits that challenged long held beliefs. Wandering 
Albatross that were tracked by satellite did not go 
around and around the world but made a series of 
erratic zigzags to and from feeding areas. Albatross 
do not always nest on the windy side of the island. 
Another interesting aspect of albatross behaviour is 
their partition of the sea’s food resources. While they 
all feed on fish, krill, and squid, they do so in a way 
that reduces unwarranted competition. 

The writer includes a fascinating chapter on alba- 
tross flight. They are such impressive, graceful birds 
on the wing that you are forced to wonder how they 
do it. The author has summarized what is known (or 


THE CANADIAN FIELD-NATURALIST 


Vols 


speculated) so far. In reality a little magic still 
remains in the albatross’s flying skill. The following 
chapter on behaviour absorbed me less. Not that 
albatross behaviour is dull; far from it. I admit I have 
given descriptions of some incidences of behaviour 
in similar terms to the author. Reading about 
behaviour, even with neat line drawings is, however, 
a bit dry. 

A little more unusual in a book of this type is a 
chapter on the poetry written about the albatross. I 
was surprised to learn that Samuel Taylor Coleridge 
who wrote the classic “Rime of the Ancient 
Mariner” never saw an albatross. Such is the 
romance of these birds. 

This book will be primarily of interest to scientists 
and the more serious visitors to albatross islands. It 
is written in a scholarly style, but is easily read by 
the non-scientist. The text is well supported by 
numerous charts, graphs, maps, tables, and drawings. 
There are several black-and-white photos, including 
some taken in the late 1800s, scattered throughout 
the text and a special section of modern, high-quality 
colour photos at the back. The 21-page bibliography 
must contain over 1000 references. This book is a 
good reference source for both naturalists and histo- 
rians. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario K1J 6K5 Canada 


The Nature of Frogs: Amphibians With Attitude 


By Harry Parsons. 2000. Douglas & McIntyre Publishing 
Group, Vancouver. x + 102 pp., illus.,Cloth $34.95; 
paper $24.95. 


One of the perennial questions in conservation is 
how to inoculate the unenlightened with a passion for 
biophilia. Gesticulating one’s arms wildly while effus- 
ing about the endearing qualities of a particular species 
rarely is an effective strategy. For the most part people 
need an emotional connection to something before 
they can embrace it more intellectually. Books lavishly 
endowed with colour photographs are one way to try 
and capture the hearts and minds of people and The 
Nature of Frogs succeeds admirably well at this task. 

Almost every other page of the book is devoted to 
a full-page, colour photograph. Not surprisingly, 
many of the photos are of colourful tropical species, 
but in addition many North American species are 
featured. Photos are also used to illustrate various 
activities (e.g. calling, amplexus, feeding) as well as 
life stages (e.g. egg mass and tadpoles). The images 
are crisp, the colours are rich and the quality of the 
photos ranges from good to exceptional. 


The accompanying text provides a broad intro- 
duction to frogs, but of necessity it skims over most 
topics. The text is divided into five chapters. 
“Consider the Frog” introduces amphibians, frogs 
and the major groups. “Kermit and the Devil” deals 
with the role of frogs in human societies. 
Reproduction and the anuran life cycle are exam- 
ined in “A Frog He Did A-Wooing Go.” “Food or 
Foe” deals with frogs as predators and prey. The 
last chapter, “The Future of the Order Anura” dis- 
cusses amphibian decline. 

The text is filled with many little gems. For exam- 
ple, according to Parsons the Egyptian hieroglyph 
for 100 000 is a tadpole. Readers are also introduced 
to the Paradoxical Frog (Pseudis paradoxa) whose 
tadpoles are larger than the adult. Inevitably, with so 
many examples mentioned in such a short text, the 
reader is left wanting to know more about particular 
topics. According to Parsons there are tropical bats 
which specialize on frogs, but no other detail is pro- 
vided. On rare occasions, the text becomes little 
more than a list: one short paragraph covers cultural 


2001 


roles of frogs in Nepal, Korea, Vietnam, Australia, 
and New Zealand. For the most part, Parsons has 
skillfully skimmed the highlights of anuran biology, 
weaving his own experiences into the text, adding a 
personal touch to the many topics. Equally impor- 
tant, his passion for frogs and his concern for their 
survival is evident throughout the text. It’s hard to 


BOOK REVIEWS 


523 


imagine anyone reading this book and not coming 
away smitten with frogs. 


DAVID SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, 
Oxford Mills, Ontario KOG 1S0 Canada 


Gatherings of Angels: Migrating Birds and their Ecology 


Edited by Kenneth P. Able. 1999. Cornell University Press, 
Ithaca, New York. 193 pp., illus. U.S.$29.95. 


Perhaps once every five years I read a book that 
presents scientific facts in such a rivetting way that 
one wonders why other writers fall so far short. Able 
offers such a winning formula. His book provides 
insight into the complexities of bird migration, one 
of the most “extraordinary of natural spectacles.” 

Ken Able, a biology professor at the State Univer- 
sity of New York, Albany, has devoted his life to the 
study of bird migration. He has contributed two 
introductory chapters and one concluding chapter, 
together with a short essay to introduce each of the 
other chapters: two on trans-gulf migration and six 
about migrations of eight bird species, the Rufous 
Hummingbird, Blackpoll Warbler, Broad-winged 
Hawk, Sandhill Crane, White-rumped and Baird’s 
Sandpipers, Dunlin and Western Sandpiper. 

Readers cannot help but be impressed by the 

“immensity, elegance and inherent risk that attend 
the great migrations” described by Able and his eight 
collaborators. For some species, challenges to a 
species’ survival may be greater during migration 
than on either the breeding or wintering grounds. 
To understand the complex, multi-factorial, interact- 
ing pressures that weigh on the survival of each indi- 
vidual migrating bird it is “critical to keep a clear 
view of what we know, ... the degree of certainty with 
which we know it, and the inferences that can confi- 
dently be made.” Nor should we, Able suggests, “rely 
too heavily on strictly economic arguments,” for this 
dooms us to “the bean counter’s dilemma — knowing 
the cost of everything but the value of nothing.” 

Radar studies have helped to confirm that Black- 
poll Warblers, after building up body fat in 
Massachusetts, wait for a cold front and then fly over 
ocean for about 85 hours, passing over Bermuda, and 
not landing until they reach Northern South America; 
the hazardous but largely predator-free trans-oceanic 
route is 1500 miles shorter than the land route via 
Florida. Flocks of Broad-winged Hawks time their 
migrations to take advantage of topography and sunny 


days to soar, with low energy consumption, 80% of 
the way to Colombia. Near the Platte River in 
Nebraska about 500 000 Sandhill Cranes during each 
spring migration benefit from the 6 to 8% of corn ker- 
nels left in fields after harvest; they gain sufficient fat 
there to aid their reproduction when they reach their 
breeding grounds farther north. After putting on fat at 
Cheyenne Bottoms, Kansas, and a non-stop flight of 
2000 miles from there to the arctic breeding ground, 
White-rumped Sandpipers lay four eggs equal to 90% 
of the female’s body weight. Two species arrive in the 
Cooper River Delta of Alaska with their energy 
reserves depleted: the long beak of the Dunlin gives it 
access to tiny clams deep in the mud, with which it 
can satisfy its energy needs in a few hours; the smaller 
Western Sandpiper, in contrast, must feed almost con- 
tinuously to satisfy itself. In terms of body lengths 
(almost 49 million), the Rufous Hummingbird makes 
what is relatively the longest migration of any bird, in 
spring up the Pacific coast from central Mexico as far 
as Alaska, and in fall down the alpine meadows of the 
Coast Range or the Rocky Mountains; nectar from 
flowers may increase their body mass from 2.7 g to 
5.7 g, but to get it they must hover, the most energy- 
intensive form of flight. 

Each chapter is illustrated with appropriate photos 
and each migration is illustrated by a superb map. It 
is a treat to read a book that is coherent, understand- 
able, and almost free from mathematical formulae, 
with each chapter written by a single author. 
Complicated scientific hypotheses are distilled into 
understandable English, reminiscent of the prose of 
men like P. A. Taverner and A. C. Bent in the first 
half of the 20 century. What a striking contrast to 
the typical heavy, overwhelmingly scientific (“‘tech- 
nospeak”) papers in some ornithology journals 
today! I recommend this book to everyone with an 
interest in birds. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


524 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Kingbird Highway: The Story of a Natural Obsession that Got a Little Out of Hand 


Kenn Kaufman. 2000. Mariner Book, Houghton Mifflin, 
Boston MA. Paperback. 318 pp., illus. U.S.$13.00. 


The gestation period of a book is said to be longer 
than that of an elephant (up to 22 months). This book 
is another example; he wrote the first draft in 1975, 
resurrected it in 1990, obtained help from skilful edi- 
tors, and finally published the hard-cover edition in 
1997. Three years later it was re-issued as an inex- 
pensive paperback, the subject of this review. 

Keen birders recognize Kaufman as one of the top 
authorities on bird identification. The first six chap- 
ters of Kingbird Highway tell how he got his start. 

In 1973, after completing high school in Wichita, 
Kansas, Kaufman determined that he would set a 
new North American record for the number of bird 
species seen in a year. Guy Emerson of the National 
Audubon Society had been the first; he identified 
497 species in 1939, followed by Bob Smart with 
510 species in 1952, Roger Tory Peterson (accompa- 
nied much of the way by James Fisher) with 572 in 
1953, Stuart Keith with 598 in 1956, and Ted Parker 
with 626 in 1971. Kaufman determined to spend 
1973 surpassing the Parker record; this quest occu- 
pies the last 20 chapters. To run up a large total for 
the United States and Canada, Kaufman had, like his 
predecessors, to crisscross North America from 
California to the Florida Keys, to Gambell Island off 
the western coast of Alaska. Unlike his predecessors, 
he made his trips by hitch-hiking. 

The previous year, 1972, had seen Richard 
Stallcup edge out Richard Webster, for the largest 
California-only list, 428 species to 427. Kaufman 
was aiming higher, for a United States and Canada 
list of 635 or 640. By 26 January he had ticked off 
his 200" species for the year, somewhat ahead of his 
expectations. Yet he was somewhat non-plussed two 
days later in Portsmouth, New Hampshire, to meet 
Floyd Murdoch, on an identical quest. Murdoch was 
researching the history of the National Wildlife 
Refuges, and in visiting each was certain to gain a 
large list. Now Kaufman had a competitor, not mere- 
ly a number to surpass. 

Some destinations were for a specific purpose, such 
as San Juan island off the Washington coast, to see the 
Skylark. From there he went back to the Florida Keys. 
Who should be with him on a boat trip to the Dry 
Tortugas but Floyd Murdoch, whose year list by then 
was about 50 species behind Kaufman’s 440. 

Kaufman shares with us the travails of hitch- 
hiking, including days without food and being wet 
for several days in succession. He travelled light, but 
met some of the keenest birders on the continent, 
including Ted Parker and the author-artist, Roger 
Tory Peterson, guest speaker at the first-ever con- 
vention of the newly formed American Birding 
Association in Kenmare, North Dakota in June. 
Kaufman’s year-to-date tally was 575. He then 


hitched a ride straight through from near Whitehorse, 
Yukon, to Fairbanks, Alaska. He passed the Parker 
record of 626 in late July, had a total of 630 when he 
hitched his way back to New Jersey in August, and 
640 when he reached El Paso, Texas. Persuaded to 
take a holiday from his quest, Kaufman spent late 
August in Mexico with Ted Parker and two friends 
in search of the Eared Trogon. 

Now he had four months to go. The species not 
yet on his list were widely scattered. A phone mes- 
sage concerning a Spotted Redshank at Brigantine 
Refuge, New Jersey, sent Kaufman hitch-hiking 
2500 miles in the rain in five days. When he got 
there, he saw the bird in question, already viewed by 
a thousand or more birders, all of whom had ticked 
off Spotted Redshank. But a shock awaited him. 
Harold Axtell, an acknowledged expert, had been to 
see the bird and two days earlier had written a full 
page description of it pointing out that it was merely 
a Greater Yellowlegs stained by oil. 

This long and futile trip to see a single species 
that was not as advertised, was a turning point in 
Kaufman’s life. He realised that in future he must 
learn the skills that Axtell had developed over a life- 
time of study and that “long-distance hitching ... was 
hypnotically mindless.” Another trip across the con- 
tinent to Seattle gained him a Sharp-tailed 
Sandpiper, then he took work picking apples in a 
Washington orchard to earn money. With him on a 
Westport boat trip on 7 October was his friendly 
competitor, Floyd Murdoch, who was now several 
species ahead. Kaufman hitched to Oklahoma to add 
the Lesser Prairie-Chicken, but he was slowing 
down. He took another trip into Mexico, outside the 
area of his list. In December he took part in the 
Christmas Bird Counts at Phoenix, and in the Yuma 
count added a Black Rail and the first-ever Rufous- 
backed Robin recorded in California. Then at 
Freeport, Texas, on the final CBC of the year, he fell 
off the breakwater rocks and nearly drowned, losing 
the precious telescope loaned him by Edgar Kincaid. 

During 1973 Kaufman had thumbed his way 
69 000 miles, paying only for a few short boat trips 
and the flight to Kimball Island. On 31 December he 
did not yet know that Floyd Murdoch had edged him 
by 669 species to his 666. Participating in a field 
sport that “had few fans, no professionals and no ref- 
erees,” Kaufman had come to the realization that 
“list-chasing was not the best way to learn birds,” 
though it had been “an incentive for getting to a lot 
of places.” And unequivocally he had set a record 
that would likely never be broken, “the most birds 
for a buck.” His total living and travelling expenses 
for the year had come to less than $1000. 

This book is a travelogue that tells of interesting 
birds in fascinating places, watched by idiosyncratic 
people, some of them “big names,” (Stuart Keith, Jim 


2001 


Tucker, Guy McCaskie, Harold Axtell, and Victor 
Emanuel, to name but a few). Though listing of new 
species eventually becomes humdrum, this book 
would make a good present for any fanatic birder of 
your acquaintance, whether or not that person is 
familiar with the term IDIOT (Incredible Distances In 


Reproductive Biology of Bats 


Edited by Elizabeth G. Crichton, and Philip H. Krutzsch. 
2000. Academic Press, San Diego. xi + 510 pp., illus. 
US. $99.95. 


This edited volume brings together 16 
researchers summarizing 11 specific topics on the 
reproductive biology of bats. Although there are 
general reviews and works on various aspects, there 
is no single comprehensive source that compiles the 
full breadth of biological knowledge of bat repro- 
duction. In terms of extant species and global distri- 
butions, bats are the second most successful group 
of mammals, next to rodents. The reproductive 
potential of bats is considered constrained, howev- 
er, with typically one young produced only once or 
twice per year. This deficiency is compensated by a 
diverse array of reproductive strategies that have 
benefited them in radiating to most environments 
throughout the world. But to understand and study 
the unique and fascinating mechanisms employed 
by bats and their associated life history traits is dif- 
ficult but challenging because of their small size, 
nocturnal behaviour, and ability to fly. 

The first three contributions deal with endocrine 
control of reproduction in bats. These cover the 
interactions of the hypothalamic-pituitary complex 
(written by E.L.P. Anthony), circulating gonadal 
hormones (L. Martin and R.T.F. Bernard), and 
peripheral endocrines (G. G. Kwiecinski and D. A. 
Damassa). At present, the current state of knowledge 
of bat endocrinology is considered in its infancy. As 
knowledge on the captive breeding of bat colonies 
improves, increased breadth in experimental design 
will shift research from the accumulation of baseline 
data to an emphasis on the role and mechanisms of 
reproductive hormones. 

Two chapters are devoted to the morphology and 
physiology specific to the male (P.H. Krutzsch) 
and female (J. J. Rasweiler, [V and N. K. Badwaik) 
reproductive tract of bats. More so than other 
groups of mammals, bats have a higher frequency 
of anatomical asymmetry such as dominance of one 
ovary. The adaptive significance of this may be to 
avoid more than one young per litter. There is a 
limit to the amount of extra weight that a pregnant 
female can carry while maintaining flight and con- 


BOOK REVIEWS 525 


Ornithological Travel), introduced to us with 
Kaufman’s quiet humour. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 


tinuing to forage, a size constraint worthy of in- 
depth study in relation to reproductive form and 
function. Pregnancy (N.K. Badwaik and J. J. 
Rasweiler, IV) is also thoroughly discussed includ- 
ing preimplantation development, implantation of 
the blastocyst, and development of placental organs 
and foetal membranes. 

Male activity patterns in bats are usually closely 
synchronized to female reproductive cyclicity. 
Reproduction seems to be timed so that lactation, 
the most energetically expensive part of reproduc- 
tion, occurs during the peak of food availability. 
This can be facilitated by sperm storage, a subject 
covered by E.G. Crichton, and a phenomenon 
unique in bats because of the extended time inter- 
vals involved. Considering the potential benefits 
of sperm preservation in humans and domesticated 
animals, research into the mechanisms and physi- 
ology in bats is an area of study deserving atten- 
tion. 

There is an overview of life history traits and 
reproductive strategies of bats (P. A. Racey and 
A.C. Entwistle). Topics include seasonality and 
reproductive patterns, reproductive delays and their 
adaptive significance, and the limits of reproduc- 
tion. A complementary paper discusses the effects 
of environmental regulation on the reproductive 
ecology of bats (P.D. Heideman). Separate chap- 
ters also deal with mating systems in bats (G. F. 
McCracken and G. S. Wilkinson), and parental 
care and postnatal growth (T.H. Kunz and W.R. 
Hood). 

The content ranged nicely from general reviews 
to more specific in-depth studies; however, I 
thought the order of chapters could have been bet- 
ter organized. Reproductive strategies and environ- 
mental regulation, two chapters near the end, would 
have been good introductory discussions for the 
book. This would have given a firm basis to the 
more specialized sections dealing with endocrine 
control, which were presented first, but the chapters 
are nonetheless independent works of study. One 
apparent editorial oversight is the absence of refer- 
ences for literature cited in the preface. Overall, the 
information is attractively presented with each 


526 


chapter having a table of contents, appropriate 
headings and subheadings, discussion or summary, 
and extensive up-to-date references. However, 
based on content and price, the book is definitely 
geared to the professional bat researcher as opposed 
to the amateur bat naturalist. It will be the source to 
consult for years to come on reproductive biology 
of bats. Not only is it a summary of the current 


BOTANY 


Phycology 


By R. E. Lee. 1999. 3rd Edition. Cambridge University 
Press, Cambridge. 614 pp. Cloth U.S.$202.60, paper 
U.S.$72.85. 


Algae 


By L. E. Graham and L. W. Wilcox. 2000. Prentice Hall, 
Upper Saddle River, New Jersey. 640 pp. U.S.$100.95. 


It is a rare event when one can welcome the return 
of an old friend dressed up in new clothes (Lee) 
along with a brand new kid on the block (Graham 
and Wilcox). Lee’s text has arguably been the best 
algal textbook for the last 20 years. The new edition 
updates a strong second edition (from 1989) and 
improves the general presentation. Graham and 
Wilcox (henceforth G&W) have used a new 
approach, and produced a stunning book that will 
replace Lee for many people as the textbook of 
choice. Both of these books offer excellent systemat- 
ic surveys, great introductions to comparative ultra- 
structure and the basics of algal physiology, ecology, 
and cell biology. G&W offer a more comprehensive 
ecological perspective and have emphasized molecu- 
lar analyses in putting algae into a phylogenetic con- 
text. 

The 61-page introductory chapter in Lee describ- 
ing the basic characteristics of algae may be the sin- 
gle best summary available for algae in that it encap- 
sulates algal diversity in the contexts of morphology, 
ultrastructure, biochemistry, and ecology. Thus, if 
you are teaching plant diversity or aquatic ecology, 
here is the essential information on these organisms. 
Lee, at 614 pages, is not as comprehensive as G&W 
with 699 pages (with 50% greater print area per 
page). This might make Lee more attractive in algal 
survey courses where one doesn’t want students con- 
fronted with quite so much information. One of the 
features of Lee that I have liked from the beginning 
is the photographs of prominent phycologists. This 
helps convince students that science is done by peo- 
ple, and that the material in textbooks is not divine 
revelation! 

G&W is simply the best algal textbook to date. 
The strongest feature is that in addition to the 15 
chapter systematic survey of algal groups that is 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


state of knowledge, but areas in need of research 
are also identified. . 


3 BurRTON K. LIM 


Centre for Biodiversity and Conservation Biology, Royal 
Ontario Museum, 100 Queen’s Park, Toronto, Ontario 
MS5S 2C6 


standard in algal texts, there are eight chapters that 
are conceptually based and integrate across all algal 
classes. This includes chapters on biogeochemistry, 
technological applications, biotic associations, 
molecular methods and phylogeny reconstruction, 
endosymbiosis, as well as chapters on phytoplank- 
ton, macroalgal, and periphyton ecology. The chap- 
ter on phytoplankton ecology contributed by J. M. 
Graham is one of the highlights of the book. In addi- 
tion to the word-based descriptions of ecological and 
biophysical processes, Graham provides quantitative 
accounts of these phenomena, as well as sufficient 
background to make it possible to teach this material 
to undergraduates. This material may not be for 
everyone, but it helps justify why we make biology 
students take calculus! One general complaint about 
this book it is that there are five chapters dedicated 
to the green algae; i.e., one for each of the classes. 
Thus the pagination dedicated to brown, red and 
green algae is 32:54:146. Since the background of 
the authors is with freshwater algae, this bias does 
not surprise me. The chapter on biotic associations is 
weak on marine algal symbioses, and fungi are men- 
tioned only in the context of lichens and microalgae. 

The classification in G&W is much more current. 
Thus the seven orders that Lee recognizes give a 
very different perspective on phaeophytes than the 
14 of G&W. Similarly, the larger number of red 
algal orders (10 versus 18) gives a more current 
aspect to G&W. The additional orders of G&W are 
not a case of splitting, but a recognition that modern 
morphological and molecular systematics have 
helped define these monophyletic groups. This dif- 
ference in conceptual underpinnings is epitomized 
by the fact there are only two phylogenetic trees in 
Lee, compared to the numerous cladograms in 
G&W. These two books are extensively referenced, 
although G& W has many more bibliographic entries. 
In addition, G&W appears to be more current with 
over 40% of the references post 1990. This contrasts 
with Lee where only 20% of the references are post 
1990. 

Both books are well illustrated with lots of func- 
tional to high quality line drawings, and good to 
excellent black and white photographs. The line draw- 


2001 


ings in G&W are much crisper and are drawn in a 
consistent style. In terms of production values the 
designers at Prentice Hall have put a lot more thought 
into G&W than Cambridge did for Lee. Even with 
similar content, students and teachers will find G&W 
esthetically more attractive. Both books could have 
been improved with even a few plates of colour pho- 
tographs, or simply using a different colour ink to 
highlight some of the figures and tables. In both books 
the text is clearly written and is free of typos. Price 
provides an important distinction: the hard cover ver- 
sion of Lee is out of sight, whereas the hard cover of 


EVIRONMENT 


BOOK REVIEWS 


527 


G&W is consistent with many university texts. On the 
one hand, the paper bound version of Lee is a relative 
bargain, and might decide the choice of text by itself. 
On the other hand, if you were to have one algal text- 
book or reference book on your shelf, G&W is the 
one I would recommend. 


DAVID GARBARY 


Department of Biology, St. Francis Xavier University, 
Antigonish, Nova Scotia, B2G 2W5 Canada; e-mail dgar- 
bary @stfx.ca 


Something New Under the Sun: An Environmental History of the Twentieth-Century World. 


J. R. McNeill. 2000. W. W. Norton, New York. N.Y. Cloth. 
421 pp., illus. 


Ecclesiastes is out of date; there is something new 
under the sun. The 20" century has been “a gigantic 
uncontrolled experiment. ... the world economy grew 
14-fold. Humankind has begun to play dice with the 
planet without knowing the rules of the game.” We 
have accepted the century’s short-lived abundance of 
cheap energy and cheap fresh water which have led 
to rapid and unsustainable growth of human popula- 
tion and of the economy. 

When historians discuss the environment, they 
often do so with little understanding of it. Environ- 
mentalists in turn rarely have much knowledge of 
history. McNeill approaches his topic from the view- 
point of a historian, allowing his abounding and 
well-documented treasury of facts and figures in 
most cases to speak for themselves, without extreme 
alarmism. The thoroughness of two solid years of 
research into the environmental history of our planet 
is evident from his 42-page bibliography. He claims 
to take an anthropocentric view, yet he refers to 
Homo sapiens as a “rogue animal.” 

McNeill emphasizes the complexities of the inter- 
relationships between soil, atmosphere, water, forests, 
and humans, and the many unforeseen and unintended 
effects of human interventions. He explains the 
sources, amounts, circumstances, and locations of 
industrial pollutants and of ecologic catastrophes 
around the world. Coal smoke, sulphur dioxide (the 
main constituent of acid rain), carbon dioxide (the 
main greenhouse gas), methane, copper, freon, and 
other chlorofluorocarbons, you name it, all are docu- 
mented here. Of 1.8 billion city dwellers, world-wide, 
one billion breathe unhealthy levels of something 
harmful. 

Follies continue apace. In the United States, Ogal- 
lala Aquifer water continues to be pumped above 
ground by 150 000 pumps at ten times the rate of 
recharge, half of its water already extracted by 1993. 


In Saudi Arabia, it takes a thousand tons of water to 
grow a ton of wheat. In Libya, water costs four times 
the value of the crops produced. Engineers have 
designed expensive monuments, some of them exam- 
ples of short-sightedness and stupidity. Massive dams 
have destroyed forests and fisheries, and raised the 
rates of malaria and schistosomiasis. The Colorado 
River now delivers only useless brine to its mouth in 
Mexico. Although Norman Borlaug won the Nobel 
Peace) Prize.in. 1970. for fatherime, the Green 
Revolution, his plant breeding has led inevitably to 
monoculture crops, reduced biodiversity, use of incre- 
mental amounts of fertilizer and pesticides, and the 
chemicalization and mechanization of agriculture. 
These in turn have led to salinization of soils, rural 
instability, poverty, recurrent famines, and massive 
movements of people into ever-larger cities that are 
unable to cope. The most valuable fisheries — 
pilchard, herring, sardine, cod, and anchoveta — have 
experienced massive collapse. During the century, fin 
whale populations dropped from 750000 to 20 000 
and blue whales from 200 000 to 500. 

One-time resources are being consumed at an 
unconscionable rate. In 1700 there were five cities 
with over half a million population, each with a large 
“ecological footprint” of food, water, and energy 
input and of waste disposal output; now there are 
over 1800 cities of this size. Now automobiles use 
up 5 to 10 percent of the surface area in Europe, the 
United States, and Japan. In Germany, for every ton 
of automobile produced, 29 tons of waste result. 

While economists ignored nature almost complete- 
ly, ecologists in general had no political or economic, 
and hence no ecologic, impact. Rachel Carson in 1963 
was the first to get her message across; the first Earth 
Day in 1970 marked the beginning of attention to 
ecology. Nevertheless, an international sampling in 
1997 revealed that citizens of only five countries, 
New Zealand, Canada, Switzerland, Austria, and the 
Netherlands, were willing to give environmental pro- 


528 


tection priority over economic growth. The effects of 
genetic engineering and of computers on the environ- 
ment remain to be experienced. The effects of global 
warming, should this process continue, cannot be pre- 
dicted with any certainty. A further loss of biodiversi- 
ty seems inevitable. 

There have been a very few successes: elimina- 
tion of lead from gasoline; the cleanup of air and 
water in Pittsburgh and London; the development of 
model public transportation and waste recycling in 
the new city of Curitiba in Brazil. 

This book has added to my knowledge and cor- 
rected some of my prejudices. In his epilogue, 
McNeill tells us, “It is impossible to know whether 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


mankind has entered a genuine ecological crisis. ... 
By the time we do know, it will be far too late to do 
much about it.” He hopes for new and cleaner energy 
regimes and for formal education of girls in poor 
countries, since “female education is the strongest 
determinant of fertility.” History and ecology “need 
to integrate with one another.” Yes. Read this book 
and you will better understand the history of the 
world’s environment. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


You Are the Earth, From Dinosaur Breath to Pizza from Dirt 


By David Suzuki and Kathy Vanderlinden. 1999. 
Greystone Books, Douglas & McIntyre, Vancouver. 128 
pp., illus. $24.95. 


I like books for young people. They’re quick to 
read. They present clear ideas. They convey just the 
right amount of information to grasp basic concepts. 
And they make you want to learn more. 

You Are the Earth is no exception. It’s a dynamic 
book, featuring lots of great photographs, illustra- 
tions, annotated diagrams, cartoons, activities, glos- 
sary and quiz, complementing clear text that makes 
every possible connection — some I would never 
have thought of — between Homo sapiens and the 
natural world. 

The chapter on air, for example, talks about argon 
gas. It tells us that the 30 zillion or so argon atoms 
we exhale with every breath travel through our 
neighbourhood within minutes, and that within a 
year they spread around the planet, with about 15 of 
them returning to our own noses. We also learn that 
argon atoms never die or change, meaning, as the 
book points out, that “thousands of years ago, an 
Egyptian slave building the pyramids breathed some 
of the same argon atoms that later Joan of Arc, 
Napoleon, and his horse breathed. And some of 
those were argon atoms exhaled by dinosaurs that 
lived 70 million years ago.” It’s a mind-boggling and 
playful notion. 

Yet the book isn’t all fun and games. It also points 
out the negative impacts humans have on the envi- 


ronment. The chapter on energy tells us that plants 
and animals in the wild use and pass on energy in a 
continuous cycle, but that humans use energy in a 
linear manner that leads to waste. The chapter on soil 
addresses topsoil loss, factory farming, and the use 
of chemical pesticides and fertilizers. The chapter 
titled “Depending on Our Relatives” illustrates bio- 
diversity (including the bacteria we host on our own 
bodies) and emphasizes how human activities are 
killing off species up to 10 000 times faster than ever 
before in the history of the planet. 

But the book isn’t all doom and gloom either. The 
last chapter provides encouragement. It tells stories 
of young people who are making a positive contribu- 
tion to the environment. It gives a list of things kids 
can do to help the natural world. 

And it provides hope: “Our survival depends on 
remembering who we are. We are the Earth — part 
of the air, water, soil, and energy of the world; 
beings with love in our hearts, life in our souls, and a 
kingdom of kin at our doorstep. It is up to us to pro- 
tect those things so that they will be around for many 
generations to come.” 

[f, as a result of reading the book, readers end up 
believing these things and taking action, it will have 
fulfilled its purpose. 


R. SANDER-REGIER 


RR5 Shawville, Quebec JOX 2Y0 Canada 


2001 


BOOK REVIEWS 


529 


The Alvars of Ontario: Significant Alvar Natural Areas in the Ontario Great Lakes Region 


By V.R. Brownell and J.L. Riley. 2000. Federation of 
Ontario Naturalists, Don Mills, Ontario. 269 pp., illus. 


Alvars are areas with low tree cover on shallow 
soils over limestone bedrock. These highly diverse 
habitats have recently received much attention from 
the conservation and scientific communities. The 
purpose of this volume is to assess the conservation 
value of 92 of the natural areas containing alvars in 
southern Ontario. The first section provides a general 
review of the methods the authors used to evaluate 
sites and details of many aspects of alvar natural his- 
tory. The second half of the book is devoted to the 
site-by-site evaluation summaries. While there is 
much overlap with the final report of the 


International Alvar Conservation Initiative (Reschke. 


et al. 1999), this report differs in its specificity to 
Ontario, its coverage of a greater range of natural 
history information, and the inclusion of more 
Ontario sites. Being an FON publication, it should 
also prove easier for the public to get hold of this 
document. 

Generally, the descriptive natural history is excel- 
lent and detailed throughout, providing sufficient 
context for their evaluation of sites. A section on the 
classification of alvar habitats quickly became overly 
complicated, however, with the presentation of three 
separate classification systems. For example, shore- 
line alvars were discussed as such in the text but 
classified out of the alvar group into “limestone 
shorelines” in the displayed table. It became obvious 
that a definitive classification of alvar habitats is not 
yet available. 

A detailed discussion of the evaluation criteria 
(representation, diversity, site condition, special fea- 
tures, and ecological function) follows with more 
reference to natural history details and provides the 
framework for the site summaries. Having analyzed 
data from a variety of sources, the authors conclude 
that 9 of 13 physiographic regions containing alvars 
do not have adequate protection for significant alvar 
sites. The analysis thus provides a solid basis for pri- 
oritizing conservation actions on Ontario alvars. 

Parts of the conclusion section were puzzling, 


especially the strong prescriptive statements about 
fire as a management tool for alvars. The authors 
claim that “[t]he burning of alvar habitats regardless 
of presence or absence of past fire, is an appropriate 
consideration because the vast majority of alvars 
have burned” page 66. This is out of place in a report 
that is not primarily about management techniques, 
especially since the topic of fire receives only one 
page of text before the conclusions. This idea seems 
to come from two sources: a single descriptive study 
on one site (Catling and Brownell 1998) which 
shows high species richness several decades after a 
burn, and the evidence of past fires on a number of 
alvars. Fire evidence in alvar habitats does not imply 
that alvar community structure is maintained by fire. 
There is probably evidence that most of Ontario has 
burned at one time or another, does this mean that all 
present-day habitats are maintained by fire or should 
be managed using fire? This reader wanted either a 
much more tentative statement about fire as a recom- 
mended management tool or much more detailed 
evidence for both the relationship between fire and 
the maintenance of alvar communities and the 
threats to which burning would putatively respond. 
Much research needs to be done on the threats to 
alvars and the potential management actions neces- 
sary to counter them before burning should be uni- 
versally considered for Ontario alvars. Despite prob- 
lems in the classification and management sections, 
the site summaries and natural history details in this 
book will provide a useful reference for future 
research, conservation, and management initiatives 
on alvars in Ontario. 


References 

Catling, P. M. and V. R. Brownell. 1998. Importance of fire in 
alvar ecosystems—evidence from the Burnt Lands, eastern 
Ontario. Canadian Field-Naturalist 112: 661-667 

Reschke, C. et al. 1999. Conserving Great Lakes Alvars. The 
Nature Conservancy, Great Lakes Program, Chicago. 230 pages. 


JEREMY T. LUNDHOLM 


Department of Botany, University of Guelph, Guelph, 
Ontario NIG 2W1 Canada 


530 THE CANADIAN FIELD-NATURALIST Vol. 115 


MISCELLANEOUS 


The Essential Aldo Leopold: Quotations and Commentaries 


Edited by Curt Meine and Richard L. Knight. 1999. 
University of Wisconsin Press, Madison. 362 pp., illus. 
U.S.$27.95. 


Just as Henry David Thoreau is identified with 
Walden, so too is Aldo Leopold with A Sand Country 
Almanac. But Leopold’s conservation classic was 
published a year after his death, the work of a scientist 
and scholar at the pinnacle of his career. He had spent 
the better part of his life observing, learning, and 
thinking about many of the ideas that he presented so 
eloquently, so forcefully, in the essays in the 
Almanac. And although millions of readers around the 
world are familiar with Leopold’s Almanac and the 
concept of a land ethic, his earlier writing on a wide 
range of related subjects is largely unknown. 

The Essential Aldo Leopold seeks to bring togeth- 
er in a single volume some of the acclaimed conser- 
vationist’s best work over a thirty-year period — 
from his days in the U.S. Forest Service to his tenure 
at the University of Wisconsin. Most are brief 
extracts from his many articles, essays, and reviews; 
there are only a few unpublished sources. The quota- 
tions have been organized into twenty-one chapters, 


NEw TITLES 


Zoology 


*Amphibians and reptiles of Pennsylvania and the 
northeast. 2001. By A.C. Hulse, J.C. McCoy, and E. 
Censky. Cornell University Press, Ithaca. 432 pp., illus. 
WES2 539195: 


The animal kingdom: a guide to vertebrate classifica- 
tion and diversity. 2000. By K. Whyman. Raintree Steck- 
Vaughn, Austin, Texas. 48 pp., illus. U.S. $25.69. 


Animal minds: beyond cognition to consciousness. 
2001. By D.R. Griffin. University of Chicago Press, 
Chicago. Xvi + 356 pp. U.S. $27.50. 


Antelopes, deer, and relatives: fossil records, behavioral 
ecology, and conservation. 2000. Edited by E. S. Vrba and 
G. B. Shaller, Yale, New Haven. 340 pp., illus. U.S. $65. 


The astonishing elephant. 2000. By S. Alexander. 
Random House, New York. 300 pp., illus. U.S. $25.95. 


+Avian research at the Savannah River site: a model for 
integrating basic research and long-term management. 
2000. Edited by J. B. Dunning, Jr. and J.C. Kilgo. Studies 
in Avian Biology No. 21. Cooper Ornithological Society, 
Camarillo, California. 170 pp., illus. U.S. $20. 


Biology of the plant bugs (Hemiptera: Miridae): pests, 
predators, opportunists. 2001. By A.G. Wheeler, Jr. 
Cornell University Press, Ithaca. 456 pp., illus. U.S. $90. 


grouped around three major themes: conservation 
science and practice; conservation policy; and con- 
servation and culture. Each chapter is introduced by 
a specialist, who explains the nature and significance 
of Leopold’s work to that particular field or topic — 
be it game management or advocacy. 

The Essential Aido Leopold confirms Leopold’s 
position as America’s pioneering wildlife ecologist, 
and a gifted writer. In reading the quotes and com- 
mentaries, one is immediately struck by Leopold’s 
ability to express “big” ideas or concepts with clarity 
and precision. In fact, those looking for that perfect 
quote for a lecture will find many here. But anyone 
trying to understand and appreciate how Leopold’s 
thinking evolved will have to look elsewhere. If any- 
thing, the collection might encourage readers to go 
back to the Almanac and the simple joys of Wis- 
consin’s sand counties. 


BILL WAISER 


Department of History, University of Saskatchewan, 
9 Campus Drive, Saskatoon, Saskatchewan S7N 5A5 


*Birds, mammals, ard reptiles of the Galapagos Islands. 
2000. By A. Swash and R. Still. Yale University Press, 
New Haven. 168 pp., illus. U.S. $24.95. 


*Birds of British Columbia, Volume 4: wood warblers 
through old world sparrows. 2001. By R. W. Cambell. 
N.K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. 
Kaiser, A.C. Stewart, and M.C.E. McNall. UBC Press, 
Vancouver. 744 pp., illus. $125. 


*The birds of Ecuador: Volume 1, status, distribution, 
and taxonomy; and Volume 2, field guide. 2001. By 
R.S. Ridgely and P.J. Greenfield. Cornell University 
Press, Ithaca. 768 pp., illus. and 816 pp., illus. 2 volume 
set U.S. $110. 


Birds of the Seychelles. 2001. By A. Skerrett and I 
Bullock. Princeton University Press, Princeton. 320 pp., 
illus. U.S. $39.50. 


+Butterflies of British Columbia. 2001. By C.S. Guppy 
and J. H. Sheppard. UBC Press, Vancouver. 414 pp., illus. 
$95. 


Cetacean societies: field studies of dolphins and whales. 
2000. Edited by J. Mann, et al. University Chicago Press, 
Chicago. xiv + 433 pp., illus. Cloth U.S. $80; paper U.S. 
$35. 


Crows and jays. 2001. By S. Madge and H. Burn, 
Princeton University Press, Princeton. 216 pp., illus. U.S. 
$24.95. 


2001 


*The destruction of the bison. 2000. By A.C. Isenberg. 
xii + 206 pp., illus. U.S. $24.95. 


*Ecology and management of large mammals of North 
America. 2000. Edited by S. Demarais and P. Krausman. 
Prentice Hall, New York. 


+The ecotraveller’s wildlife guide: Alaska. 2001. By D. 
Paulson and L. Belestsky. Academic Press, San Diego. xiv 
+ 436 pp., illus. U.S. $29.95. 


*Enjoying moths. 2001. By R. Leverton. Poyser Natural 
History, London, England. xi + 276 pp., illus. 


+Evolution, ecology, conservation, and management of 
Hawaiian birds: a vanishing avifauna. 2001. Edited by 
J.M. Scott, S. Conant, and C. vanRiper, III. Cooper 
Ornithological Society, Camarillo, California. 428 pp., 
illus. Cloth U.S. $48.50; paper U.S. $29. 


+Estimates of shorebird populations in North America. 
Piteeby hk. 1G. Morrison, R.E. Gill, Jr., B.A. 
Harrington, S. Skagen, G. W. Page, C.L. Gratto-Trevor, 
and S. M. Haig. Canadian Wildlife Service, Ottawa. 64 pp. 


A field guide to the amphibians and reptiles of the Maya 
world: the lowlands of Mexico, northern Guatemala, 
and Belize. 2000. By J.C. Lee. Cornell, Ithaca. xi+402 
pp., illus. Cloth U.S. $ 59.95; paper U.S. $35. 


Fish watching with Eugenie Clark. 2000. By M.E. 
Ross. Lerner, Minneapolis. 48 pp., illus. U.S. $19.93. 


*Flying foxes: fruit and blossom bats of Australia. 2000. 
By L. Hall and G. Richards. Krieger, Melbourne, Florida. 
viii + 135 pp., illus. Cloth U.S. 29.50; paper U.S. $21.50. 


*Four wings and a prayer: caught in the mystery of the 
monarch butterfly. 2001. By S. Halpern. Knopf Canada, 
Mississauga. 212 pp. $29.95. 


*Handbook of birds of the world, volume 6: mousebirds 
to hornbills. 2001. Edited by J. del Hoyo, A. Elliot, and J. 
Sargatal. Lynx Edicions, Barcelona. 589 pp., illus. U.S. 
$185. 


tIn search of the golden frog. 2000. By M. Crump. Uni- 
versity of Chicago Press, Chicago. xiv + 299 pp., illus. 
$27. 


yAn introduction to the invertebrates. 2001. By J. 
Moore. Cambridge University Press, New York. xv + 355 
pp., illus. Cloth U.S. $64.95; paper U.S. $22.95. 


Kangaroos in outback Australia: Comparative ecology 
and behavior of three coexisting species. 2000. By D.R. 
McCullough and Y. McCullough. Columbia, New York. 
Xvii + 308 pp., illus. Cloth U.S. $50; paper U.S. $30. 


Katydids and bush-crickets: reproductive behavior and 
evolution of the Tettigoniidae. 2001. By D. T. Gwynne. 
Cornell University Press, Ithaca. 400 pp., illus. U.S. $39.95. 


Keepers of the wolves: the early years of wolf recovery 
in Wisconsin. 2001. By R. P. Thiel. University Wisconsin 
Press, Madison. 256 pp., illus. Cloth U.S. $50; paper U.S. 
$50; paper U.S. $19.95. 


BOOK REVIEWS 


53] 


+Life underground: the biology of subterranean rodents. 
2000. Edited by E. A. Lacey, J. L. Patton, and G.N. 
Cameron. University Chicago Press, Chicago. xii + 450 pp., 
illus. Cloth U.S. $65; paper U.S. $24. 


Medieval birds in the Sherborne missal. 2001. By J. 
Backhouse. University of Toronto Press, Toronto. c64 pp., 
illus. $19.95. 


+A natural history of sex: the ecology and evolution of 
mating behaviour. 2001. By A. Forsyth. Firefly, 
Willowdale, Ontario. 192 pp., illus. $14.95. 


+The New York City Audubon Society guide to finding 
birds in the metropolitan area. 2001. By M. T. Fowle 
and P. Kerlinger. Cornell University Press, Ithaca. xix + 
230 pp., illus. U.S. $ 17.95 


The philosophy and practice of wildlife management. 
2001. By F. F. Gilbert and D. G. Dodds. 3rd edition. 
Krieger Publishing, Melbourne, Florida. 370 pp., U.S. 
$34.50. 


*Pigeons and doves: a guide to the pigeons and doves of 
the world. 2001. By D. Gibbs and E. Barnes. Yale 
University Press, New Haven, 615 pp., illus. U.S. $60. 


*Prairie birds: fragile splendor in the Great Plains. 
2001. By P. A. Johnsgard. University Press of Kansas, 
Lawrence. xvii + 331 pp., illus. U.S. $29.95. 


Salmon country: a history of the Pacific salmon. 2000. 
By. R.H. Bush. Firefly, Willowdale, Ontario. 176 pp., 
illus. U.S. $ 17.95. 


+Sea stars of British Columbia, southeast Alaska, and 
Puget Sound. 2000. By P. Lambert. Revised edition. 
UBC Press, Vancouver. 186 pp., illus. $25.95. 


*The Sibley guide to birds. 2000. By D.A. Sibley. 
National Audubon Society. Knopf (Random House), New 
York. 544 pp., illus. 


The snakes of Trinidad and Tobago. 2001. By H.C.A 
Boos. Texas A & M University Press, College Station. 
328 pp., illus. U.S. $47.95. 


*Snipes of the western palearctic. 2000. By R. Rouxel. 
Eveil Nature, Saint Yrieux sur Charente, France. 304 pp., 
illus. 


*Sylvia warblers. 2001. By H. Shirihai, G. Gargallo, and 
A.J. Helbig. Princeton University Press, Princeton. 576 
pp., illus. U.S. $75. 


Thrushes. 2001. By P. Clement. Princeton University 
Press, Princeton. 424 pp., illus. U.S. $ 49.50. 


+Towards conservation of the diversity of Canada Geese 
(Branta canadensis). 2000. Edited by K. M. Dickson. 
Occasional Paper No. 103. Canadian Wildlife Servide, 
Ottawa. 165 pp., illus. 


Wildlife study design. 2001. By M. L. Morrison. W. M. 
Block, M. D. Strickland, and W. L. Kendall. Springer- 
Verlag, New York. 255 pp., illus. U.S. $69.95. 


532 


Botany 


+The Cambridge illustrated glossary of botanical terms. 
2001. By M. Hickey and C. King. Cambridge University 
Press, New York. xii + 208 pp., illus. Cloth U.S. $ 85; 
paper U.S. $ 29.95. 


*Flora of Florida, Volume 1: pteridophytes and gym- 
nosperms. 2000. By R. P. Wunderlin and B. F. Hansen. 
University Press of Florida, Gainesville. xii + 365 pp., 
illus. U.S. $49.95. 


*Flora of New Brunswick. 2000. By H.R. Hinds. 2nd edi- 
tion. Department of Biology, University of New 
Brunswick, Fredericton. 695 pp., illus. $50. 


The interactive identification of native and naturalized 
new world Salix using intkey. 2001. By G. Argus. 
Available from author, 310 Haskins Road, Merrickville R 
3, Ontario KOG 1N0. 104 pp., illus. + diskette. $20. 


tLichens of Antarctica and South Georgia: a guide to 
their identification and ecology. 2001. By D.O. Ovstedal 
and R. I. L. Smith. Cambridge University Press. New 
York. xii + 411 pp., illus. U.S. $ 100. 


*North woods wildflowers. 2001. By D. Ladd. Globe 
Pequot Press (Canadian distributor or General Publishing, 
North York). 280 pp., illus. $39.95. 


Painting flowers in watercolour: a naturalistic 
approach. 2001. By C. G. Guest. Timber Press, Portland, 
Oregon. 160 pp., illus. U.S. $ 29.95. 


+Pondweeds, bur-reeds, and their relatives of British 
Columbia. 2000. By T.C. Brayshaw. 2nd edition. Royal 
British Columbia Museum, Victoria. 250 pp., illus. 
$24.95. 


Weeding through the wilderness: common and scien- 
tific names of weeds in Canada. 2000. By Agriculture 
Canada. Canadian Government Publishing, Ottawa. 132 
pp., illus. $22. 


Environment 


*AAAS atlas of population and environment. 2000. By 
P. Harrison and F. Pearce. University of California Press, 
Berkley. Xi + 204 pp., illus. Cloth U.S. $65; paper U.S. 
52009) 


*The alvars of Ontario: significant alvar natural areas in 
Ontario Great Lakes Region. 2000. By V.R. Brownell 
and J. L. Riley. Federation Ontario Naturalists, Toronto. x + 
269 pp., illus. 


+Cognitive ecology of pollination: animal behavior and 
floral evolution. 2001. Edited by L. Chittka and J. D. 
Thomson. Cambridge University Press, New York. xiii + 
344 pp., illus. U.S. $95. 


*Ecosystem dynamics of the boreal forest. 2001. Edited 
by C.J. Krebs, S. Boutin, and R. Boonstra. Oxford 
University Press, New York, 512 pp., illus. U.S. $152. 


Encyclopedia of environmental science. 2000. Edited by 


THE CANADIAN FIELD-NATURALIST 


Vols 


J. Morgillo and L. Zierdt-Warshaw. Oryx Press, Phoenix. 
v + 450 pp., illus. U.S. $112.25. 


*Environmentalism unbound: exploring new pathways 
for change. 2001. By R. Gottlieb. MIT Press, Cambridge, 
Massachusetts. xvii + 396 pp., U.S. $29.95. — 


*Finding order in nature: the naturalist tradition from 
Linnaeus to E.O. Wilson. 2000. By P. L. Farber. Johns 
Hopkins University Press, Baltimore. x + 136 pp., illus. 
Cloth U.S. $39.95; paper U.S. $15.95. 


+Genetics, demography, and viability of fragmented 

populations. 2000. Edited by A.G. Young and G.M 
Clarke. Cambridge University Press, New York. xviii + 
438 pp., illus. Cloth U.S. $110, paper U.S. $39.95. 


Iniand flood hazards: human, riparian, and aquatic 
communities. 2000. By E.E. Whohl. Cambridge, New 
York. xiv + 498 pp., illus. U.S. $110. 


+Journeys through paradise: pioneering naturalists in 
the southeast. 2001. By G. Fishman. University of 
Florida Press, Gainesville (Canadian distributor Scholarly 
Book Services, Toronto). 297 pp. $41.25. 


The Laguna Madre of Texas and Tamaulipas. 2001. 
Edited by J. W. Tunnell, Jr. and F. W. Judd. Texas A & M 
University Press, College Station. 384 pp., illus. U.S. $60. 


*+Learning to manage global environmental risks: Vol- 
ume 1, a comparative history of social responses to cli- 
mate change, ozone depletion, and acid rain; Volume 2, 
a functional analysis of social responses to climate 
change, ozone depletion, and acid rain. 2001. By the 
Social Learning Group. MIT Press, Cambridge, 
Massachusetts. 376 pp., illus. Cloth U.S. $75; paper U.S. $ 
30 and 226 pp., illus. Cloth U.S. $60; paper U.S. $ 24. 


Naturalist’s Big Bend: an introduction to the trees and 
shrubs, wildflowers, cacti, mammals, birds, reptiles and 
amphibians, fish, and insects. 2001. By R. H. Waver and 
C. M. Fleming. Revised edition. Texas A & M University 
Press, College Station. 232 pp., illus. Cloth U.S. $29.95; 
paper U.S. $15.95. 


*Natural selections: national parks in Atlantic Canada, 
1935-1970. 2001. By A. MacEachern. McGill-Queen’s 
University Press, Montreal. 328 pp., illus. $49.95. 


*Northern wild: best contemporary Canadian nature 
writing. 2001. Edited by D.R. Boyd. Greystone (Douglas 
and McIntyre, Vancouver). ix + 278 pp., $22.95. 


Picturing tropical nature. 2001. By N. L. Stepan. Cornell 
University Press, Ithaca. 256 pp., illus. U.S. $35. 


+A primer of ecology. 2001. By N.J. Gotelli. 3"¢ edition. 
Sinauer, Sunderland, Massachusetts. xxi + 265 pp., illus. 
U.S. $ 31.95. 


+Snow ecology: an interdisciplinary examination of 
snow-covered ecosystems. 2001. Edited by H.G. Jones, 
J.W. Pomeroy, D. A. Walker, and R. W. Hoham. Cam- 
bridge University Press, New York. xx + 378 pp., illus. 
U.S. $80. i 


2001 


+Thieves, deceivers, and killers: tales of chemistry in 
nature. 2001. By W. Agosta. Princeton University Press, 
Princeton. 241 pp., illus. U.S. $26.95. 


Miscellaneous 


Cabins: a guide to building your own nature retreat. 
2001. By D. and J. Stilles. Firefly Books, Willowdale, 
Ontario. 232 pp., illus. Cloth $29.95; paper $19.95. 


+The complete guide to walking in Canada. 2001. By E. 
Katz. Firefly Books, Willowdale, Ontario. 360 pp., illus. 
$16.95. 


+Cradle of life: The discovery of earth’s earliest fossils. 
2001. By J. W. Schopf. Princeton University Press, 
Princeton. xv + 367 pp., illus. U.S. $17.95. 


+Memorabilia of activities of William W. Judd while with — 


the Canadian Meteorological Service during World 
War II, 1942 to 1945. 2001. By W. W. Judd. Phelps 
Publishing, London, Ontario. 98 pp., illus. $10. 


+Memorabilia of Robert Elliott (1858-1902) poet and 
naturalist of Plover Mills, Middlesex County, Ontario. 
2001. By W.W. Judd, 50 Hunt Club Drive, London, 
Ontario N6H 3Y3. 75 pp., illus. $10. 


*+Minutes of meetings, 1956 to 1960, of the Mcilwraith 
Ornithological Club London, Ontario, Canada. 2001. 
By W.W. Judd. Phelps Publishing (order from author, 
Greenpeace, 50 Hunt Club Drive, London, Ontario N6H 
3Y3) 81 pp. $10. 


*Permafrost: a guide to frozen ground in transition. 
2001. By N Davis. University of Alaska Press, Fairbanks. 
xvi + 351 pp., illus. U.S. $35.95. 


*Sir William Jardine: a life in natural history. 2000. By 
C.E. Jackson and P. Davis. Leicester University Press, 
London. 256pp., illus. £55. 


*Their father’s work: casting nets with World’s fisher- 
men. 2000. By W. McCloskey. McGraw-Hill, Scar- 
borough. 307. pp., $20.95. 


Books for Young Naturalists 


About insects: a guide for children. 2000. By C. Sill. 
Peachtree, Atlanta. 48 pp., illus. U.S. $14.95. 


The age of dinosaurs. 2000. By S. Parker. Grolier, 
Danbury, Connecticut. 58 pp., illus. Each of 12 Volume 
set. U.S. $249 for set. 


Animal lives: the frog and Animal lives: the rabbit. 
2000. By S. Tagholm. Kingfisher, New York. Each 32 pp., 
illus. U.S. $9.95. 


Animals among us: living with suburban wildlife. 2000. 
By F. Hodgkins. x + 117 pp., illus. U.S. $ 19.50. 


Animals in cold places. 2000. By M. Butterfield. Raintree 
Steck-Vaughn, Austin, Texas. 32 pp., illus. U.S. $22.83. 


BOOK REVIEWS 


533 


Beetles; Flies. 2000. By E. Pascoe. Blackbirch, Wood- 
bridge, Connecticut. Each 48 pp., illus. U.S. $ 18.95. 


Destination Australia. 2000. By J. Gupper. National 
Geographic Society, Washington. 32 pp., illus. U.S. $ 
16.95. 


Eagles. 2000. By D. Hodge. Kids Can Press, Niagara 
Falls, New York. 32 pp., illus. Cloth U.S. $10.95; paper 
US. $ 5.95. 


HEco-fun: great projects, experiments, and games for 
a greener earth. 2001. By D. Suzuki and K. Vander- 
linden. Greystone Books, Douglas and McIntyre, 
Vancouver. 128 pp., illus. $14.95. 


Exploring the universe: science activities for kids. 2000. 
By A.D. Fredericks. Fulcrum, Golden, Colorado. 128 pp., 
illus. U.S. $16.95. 


Feet that suck and feed. 2000. By D. Swanson. 
Greystone, Vancouver. 30 pp., illus. 


A fish caught in time: the search for the coelacanth. 
2000. By S. Weinberg. Harper Collins Children’s, New 
York. xx + 220 pp., illus. U.S. $24. 


Great grizzly wilderness: a story of the Pacific rain 
forest. 2000. By A. Fraggalosch. Soundprints, Norwalk, 
Connecticut. 35 pp., illus. U.S. $15.95. 


My monarch journal: parent-teacher edition. 2000. By 
C. Muther. Dawn, Nevada City, California. 52 pp., illus. 
US239:95: 


Partners and parents; Plants and planteaters; Poisoners 
and pretenders; Predators and pray. 2000. By M. 
Chinery. Secrets of the Rainforest Series. Crabtree, New 
York. Each 32 pp., illus. Cloth U.S. $14.97; paper U.S. 
$7.16. 


Realm of the panther: a story of south Florida forests. 
2000. By E. Costello. Soundprints, Norwalk, Connecticut. 
35 pp., illus. U.S. $15.95. 


The Rocky Mountains. 2001. By L. Bogard. Ecosystems 
of the World Series. Benchmark Books, Tarrytown, New 
York. 64 pp., U.S. $18.95. 


Science experiments. 2000. By A. Hansen. Lowell 
House, Los Angeles. 112 pp., illus. U.S. $8.95. 


Sockeye’s journey home: the story of the Pacific salmon. 
2000. By B.G. Winkelman. Soundprints, Norwalk 
Connecticut. 32 pp., illus. U.S. $15.95. 


Taking root. 2000. by A. Fowler. Children’s Press, 
Danbury, Connecticut. 32 pp., illus. Cloth U.S. $19; paper 
US. $4.95. 


The winking, blinking sea: all about bioluminescence. 
2000. By M Batten. Millbrook Press, Brookfield, Con- 
necticut. 30 pp., illus. U.S. $20.99. 


+Available for review 
* Assigned for review 


Minutes of the 122nd Annual Business Meeting of 
the Ottawa Field-Naturalists’ Club, 9 January 2001 


Place and time: 
Chairperson: 
Attendance: 


Eleanor Zurbrigg, President 


Canadian Museum of Nature, Ottawa, Ontario, 7:30 pm 


Thirty-one 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 7:56 pm. 


1. Minutes of the Previous Meeting 

In the heading of the minutes, the date should read 
“January 11, 2000”. 

In section 5, “Committee Reports’, the motion to 
approve the report of the Education & Publicity 
Committee, second sentence, the final words in 
parentheses should read “slide shows”. 

Moved by Dave Moore and seconded by Colin 
Gaskell, that the minutes be accepted as amended. 


(Motion Carried) 


2. Business Arising from the Minutes 

In response to a recommendation by Claudia 
Burns at the previous meeting, that the Club make an 
effort to be more accessible to new members, 
Eleanor Zurbrigg described several initiatives that 
had been taken during the year. 


3. Communications Relating to the Annual 
Business Meeting 
There were no communications relating to the An- 
nual Business Meeting. 


4. Treasurer’s Report 

Frank Pope reviewed the financial report for the 
year ending September 30, 2000, noting a favourable 
report from the Auditor and the fact that the Club’s 
net assets had increased by $8154. In response to a 
question from Fenja Brodo, he explained that “other 
revenue” in the note for the Fletcher Wildlife Garden 
included funds raised from activities such as the 
Taverner Cup Competition. 

Moved by Frank Pope and seconded by Colin 
Gaskell that the Financial Report be accepted. 


(Motion Carried) 


5. Committee Reports 

Eleanor Zurbrigg introduced each of the 
Committee reports and a representative of the appro- 
priate Committee and asked for questions and 
comments. 

Dave Moore commented that future reports of the 
Macoun Club should include more information about 


the activities of the Club. Fenja Brodo asked whether 
participation in the Macoun Club had increased. There 
was no representative to give a definitive answer, but 
Eleanor Zurbrigg said she thought that it had. 

Ron Bedford gave a number of numerical changes 
to the report of the Publications Committee. 

Moved by Ken Allison, seconded by Dave Moore, 
that the reports as amended be accepted. 


(Motion Carried) 


6. Nomination of the Auditor 
Moved by Frank Pope, seconded by Bill Cody, 
that Janet Gehr continue as Auditor for another year. 


(Motion Carried) 


7. Report of the Nominating Committee 

The Committee recommended the following list of 
candidates for the 2001 Council (new members are 
indicated by an asterisk) 


President Eleanor Zurbrigg 
Vice-President Roy John 
Recording Secretary Ken Allison 
Treasurer Frank Pope 
Other Members: 

Ron Bedford Terry Higgins 
Roseanne Bishop* David Hobden 
Fenja Brodo Bev McBride 
Irwin Brodo* Garry McNulty* 
John Cameron* Dave Moore 
Bill Cody Rita Morbia* 
Francis Cook Bob Roach 


Barbara Gaertner Stan Rosenbaum 
Tony Halliday Dave Smythe 

Six members of the 2000 Council decided not to 
stand for re-election: Sarah Coulber, Ellaine 
Dickson, John Martens, Philip Martin, Jim Sutton, 
and Dorothy Whyte. Colin Gaskell thanked these 
members for their contribution to the Club. 
Moved by Colin Gaskell, seconded by Frank Pope, 
that the list of nominations for the 2001 Council be 
accepted. 


(Motion Carried) 


534 


2001 


8. New Business 

Eleanor Zurbrigg noted that the year 2001 is the 
“International Year of Volunteers” and welcomed 
suggestions for increasing the volunteer involvement 
of members in the administration of the Club. 


9. Presentation by the Birds Committee 

Bev McBride gave a presentation about the struc- 
ture and activities of the Birds Committee, noting 
that it is the largest of the present committees. For 
the past twenty years the Birds Committee has con- 


MINUTES OF 122ND ANNUAL BUSINESS MEETING 


a5 


ducted the Christmas Bird Count, an important activ- 
ity which was initiated in the Ottawa area around the 
turn of the century. 


10. Adjournment 
Moved by David Hobden, seconded by Stan 
Rosenbaum that the meeting be adjourned at 9:10 pm. 


(Motion Carried) 


DAVE SMYTHE 
Acting Recording Secretary 


The Ottawa Field-Naturalists’ Club Committee Reports for 2000 


Awards Committee 
The following awards were presented at the Annual Soiree, 
held on 14 April 2000: 


MEMBER OF THE YEAR AWARD: Claudia Burns for her 
work at the Fletcher Wildlife Garden and in particular with 
the backyard garden project. 


GEORGE MCGEE SERVICE AWARD: Betty Campbell for her 
extensive Club activities including work at the Fletcher 
Wildlife Garden, Club displays at various venues, work 
with the Education and Publicity Committee, and organiza- 
tion of the Club's huge collection of slides. 


CONSERVATION AWARD FOR Member: Stan Rosenbaum for 
his leadership as chair of the OFNC Conservation 
Committee, revitalized the committee to the point where it 
is Once again one of the largest and most active of the 
OFNC committees. 


CONSERVATION AWARD FOR NON-MEMBER: Friends of 
Petrie Island, a community-based group, for its remark- 
able job in the past two years of focusing public and politi- 
cal attention on this special area in the Ottawa River. 


HONORARY MEMBER: Ted Mosquin for his many years of 
significant contributions to both the OFNC and to Canadian 
natural history and its conservation. 


The awards nomination form was re-designed for inclu- 
sion each year with the fourth issue of Trail & Landscape, 
rather than with membership renewal notices, in an effort to 
provide the membership with a more visible solicitation for 
input into the awards process. 

S. DARBYSHIRE 


Birds Committee 

The Birds Committee participated jointly with the Club 
des Ornithologues de l'Outaouais to run a successful 
Christmas Bird Count at the end of 1999. Plans are in place 
for the 2000 count. We also had a well-attended fall bird 
count in October 2000. We participated in the Peregrine 
Falcon Watch at the downtown Ottawa nest site. One 
youngster survived the fledgling season, and an adult bird 
appears to be wintering downtown. The Bird Records Sub- 
committee has reviewed records of rare birds for the area 
and will publish the findings in an upcoming Trail and 
Landscape. We added a new ad hoc sub-committee this 
year, as plans for the 2nd Ontario Breeding Bird Atlas are 
underway. The chair of the sub-committee is the regional 


coordinator for the atlas. Our Seed-a-thon raised around 
$800.00 for the Club's bird feeders. Our bird study group 
held two well-attended training sessions during the year. 
We also revised the rare bird alert telephone tree and con- 
tinued to operate the Ottawa Bird Status Line (a recorded 
telephone message service indicating current bird sight- 
ings). 

B. McBRIDE 


Computer Management Committee 
The Committee provided support services for the 
Membership database, the accounting system, the computer 
systems for Trail & Landscape and The Canadian Field- 
Naturalist, and maintained the OFNC web site. In particu- 
lar, it upgraded the accounting system to make it Y2K com- 
patible. The Committee operated without a chairperson for 
most of the year and was unable to undertake any new ini- 
tiatives. 
D. SMYTHE 


Conservation Committee 

Alfred Bog: The Prescott-Russell Official Plan (protect- 
ing against draining of adjacent land for peat extraction) 
took effect December 31, but is being appealed to OMB. 
OFNC has status as a participant at hearings. Meeting held 
between Frank Pope and Minister John Baird regarding 
need for peat legislation. Letters drafted for FON and 
OFNC signatures. Bird Studies Canada: Contribution of 
$1,000 from the de Kiriline-Laurence Fund to Norfolk 
Field Naturalists new headquarters. Green Map: the 
Committee recommended OFNC support, but a later invita- 
tion that OFNC become its main sponsor was declined by 
Council. The Committee co-operated with Green Map to 
mount a display at the OFNC June 4th Open House. Petrie 
Island: comments made on City of Cumberland proposed 
interpretive trail system. Petrie Island is now better known 
to the public and has many defenders. New reserves: letters 
sent to OMNR supporting proposals for Burnt Lands 
Nature Reserve, and White Lake Conservation Reserve. 
Canadian Environmental Assessment Act (CEAA) 5-year 
review: Written submission made. Consultants’ Manage- 
ment Proposals for Shirley's Bay: comments made on first 
draft and on full report. Comments were printed in the July- 
Sept. issue of Trail & Landscape. Leitrim Wetlands: 
increased concern as a result of Regional Council allocation 
of $6M funds for infrastructure in the area of the proposed 
subdivision. Letters sent to Region and to the Department 


536 


of Fisheries and Oceans. Goulbourn Township: letters 
exchanged with OMNR regarding applications for subdivi- 
sions — Upper Poole Creek and Fernbank wetland. The 
former application was denied, but subject to OMB appeal. 
Fernbank application is presently denied owing to wetland 
encroachment, but OMNR advised that wetland draining by 
private landowners is not prohibited in all cases. A follow- 
up response was drafted. Species At Risk Act (SARA): 
Letters of support sent, but Bill C-33 died with election 
call. Ottawa Transition Team: Three OFNC members 
attended a meeting with transition staff on Environmental 
Advisory Committees. Our comments were made verbally 
and by letter. 

S. ROSENBAUM 


Education and Publicity Committee 

Education on natural history was promoted through pro- 
vision of volunteer leaders for nature outings and talks, 
including four outings in Mer Bleue with Beavers, Cubs 
and Girl Guides, and four outings in Gatineau Park and 
along the Ottawa River with other organizations. 

OFNC displays were presented at the Annual OFNC 
Soirée, National Wildlife Week at Carlingwood Shopping 
Centre (18 volunteers), Environment Week at Place du 
Portage, OFNC Volunteer Day at the Fletcher Wildlife 
Garden, and for the FON at the Carleton Teachers’ 
Federation PD day. A special display was prepared for the 
Macoun Field Club and presented at the Greenbank Middle 
School Environmental Conference. The Committee also 
provided judges for the Ottawa Regional Science Fair. 
Three OFNC prizes were awarded. Monthly lectures were 
advertised through notices to public libraries. 

Sales of club merchandise totaled approximately 
$500.00. At the sale table at the Monthly Lecture series, the 
Committee started featuring publications which are on the 
topic of that evening's presentation, with good results. 


B. CAMPBELL 


Excursions and Lectures Committee 

The Excursions and Lectures Committee arranged 42 
outings. There were 16 outings to study birds, six for 
plants, four for insects, and six to observe amphibians, fish, 
fungi, geology and astronomy, while another 10 outings 
were general in nature. The bird outings included two pop- 
ular annual excursions to Presqu'ile and Derby Hill. One 
further birding event was organized in conjunction with the 
Fletcher Wildlife Garden. Our program also included nine 
regular monthly meetings at the Canadian Museum of 
Nature and the Annual Soiréée. 


P. MARTIN 


Executive Committee 

The Executive Committee, with other interested mem- 
bers of Council, met once in April 2000, to reach a decision 
on the Green Map Project proposal. This proposal invited 
the OFNC to accept sponsorship and responsibility for the 
project and its website. It was decided that, while the Club 
supported "in principle" the project, it was not in a position 
to become directly involved in management of the project. 
The Club could be involved as a provider of available natu- 
ral history data for the project, and would anticipate being a 
user of Green Map products in the future. 


E. ZURBRIGG 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Finance Committee 

The Committee met four times. In December 1999, the 
Committee examined the preliminary (pre-audit) Financial 
Statement for 1998/1999. The draft statement was present- 
ed by the Treasurer to the Council's December meeting and 
approved at the Club's Annual Business Meeting. The 
Statement showed small surpluses on both the OFNC and 
CEN accounts. 

At two meetings held in the Spring, the Committee pre- 
pared a recommendation to Council on the disposition of 
the Manning bequest. The recommendation was adopted by 
Council at its June 2000 meeting. 

The Committee continued its examination of ways to 
improve the transparency and usefulness of the accounting 
system. The Treasurer prepared a numbered break-down of 
budget revenue and expenditure heads to facilitate accurate 
recording of expenditures by Committee Chairs and the 
Treasurer's Assistant. 

In August, the Committee prepared a draft budget for 
2000/2001. This was presented to Council in September 
2000 and, after adjustments, accepted at its October meet- 
ing. 

The Committee recommends no change in the member- 
ship fee structure. 

A. HALLIDAY 


Fletcher Wildlife Garden Committee 

The level of activity at the garden remained steady in 
2000 compared to the previous year, with a total of about 
2,500 volunteer hours contributed. The Friday morning 
Backyard Garden crew again gave unstintingly of their time 
every week to improve the showpiece for the Fletcher gar- 
den. Regular clean ups were held using OFNC volunteers. 
The spring plant sale and exchange raised funds for garden 
operations. 

In addition to several special events, including a work- 
shop on invasive species and a course on wildflower gar- 
dening by Philip Fry, the Interpretive Centre was open to 
the public every Sunday over the summer. A new project 
was started in partnership with a professor at Carleton 
University to explore methods to control the very invasive 
Black Swallowwort. Wildlife gardening publications were 
produced and disseminated to the public supported mainly 
by a grant from the Canada Trust Friends of the Environ- 
ment Program. 

The Taverner Cup birding competition was hosted at the 
FWG on May 27. About $900 was raised for the garden, 
with six competitive and 12 recreational teams participat- 
ing. 

The FWG continued to be a central location for OFNC 
activities, with a Club membership drive hosted at the gar- 
den in June. Club Council and committees, including the 
Birds Committee, meet regularly in the Interpretive Centre. 


P. HALL 


Macoun Field Club Committee 

The Committee met five times to plan the weekly pro- 
grams for Club members. In addition to the regular sched- 
ule of speakers and natural history workshops, committee 
members led 21 field trips and four camping trips. 


B. GAERTNER 


Membership Committee 
The distribution.-of memberships for 2000 is shown in 


2001 MINUTES OF 122ND ANNUAL BUSINESS MEETING 537 


the table (below), with the comparable numbers for 1999 in 
brackets. These statistics do not include four complimenta- 
ry memberships awarded to winners of the 2000 Science 
Fair competition, nor the 22 affiliate organizations which 
receive copies of the Club's publications. 

The Club awarded an Honorary Membership to Ted 
Mosquin for his contributions to the Club and to Canadian 
natural history (see T&L volume 34, number 3). 

Two prominent and long-time members of the Club 
passed away this year. Verna Ross McGiffin had been a 
member of the Club since 1944, and was given Honorary 
Membership in 1983. Dr. Clarence Frankton had been a 
member of the Club since 1946, and was given Honorary 
Membership in 1979. 


D. SMYTHE 


Publications Committee 

The Committee met twice. Three issues of The Canadian 
Field-Naturalist were published: Volume 114, #1,2,3. 
These three issues contained: 554 pages; 45 articles; 2] 
notes; four COSEWIC articles; 40 book reviews; 198 new 
titles; one commemorative tribute; and 22 pages of News 
and Comment. There were no changes in the panel of 
Associate Editors. However, Warren Ballard indicated that 
he will step down with the completion of volume 114, and 
a replacement has been approached. Three articles pub- 
lished in Volume 113 qualified for support from the 
Manning Memorial Fund. 

Four issues of Volume 34 of Trail & Landscape contain- 
ing 164 pages were published. 


Distribution of memberships in The Ottawa Field-Naturalists’ Club 


CANADIAN 

Type Local Other 
Family 350 (360) 23 (23) 
Individual 340 (344) 122,425) 
Honorary 15 (14) 8 (8) 
Life 22(17) 21 (21) 
Sustaining 8 (8) 1 (0) 
Total 735 (743) 175 (178) 


R. BEDFORD 
FOREIGN 
USA Other Total 
2{2) 1 (2) 376 (387) 
25 (23) 4 (3) 491 (495) 
0 (0) 0 (0) 23 (22) 
5 (5) 1) 49 (44) 
0 (0) 0 (0) 9 (9) 
32 (30) 6 (6) 948 (957) 


538 


Auditor’s Report 


THE CANADIAN FIELD-NATURALIST 


To The Members of THE OTTAWA FIELD-NATURALISTS’ CLUB 
The Ottawa Field-Naturalists’ Club . 


I have audited the balance sheet of THE OTTAWA 
FIELD NATURALISTS' CLUB as at September 30, 
2000, the statement of changes in net assets, and the 
statements of operations. These financial statements 
are the responsibility of the organization's manage- 
ment. My responsibility 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 generally 
accepted auditing standards. Those standards require 
that I plan and perform an audit to obtain reasonable 
assurance whether the financial statements are free of 
material misstatement. An audit includes examining 
evidence supporting the amounts and disclosures in 
the financial statements. An audit also includes assess- 
ing the accounting principles used and significant esti- 
mates 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 rev- 
enue from memberships, 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 concerning 
the completeness of the revenues referred to in the 
preceding paragraph, these financial statements pre- 
sent fairly, in all material respects, the financial posi- 
tion of the OFNC as at September 30, 2000, and the 
results of its operations and changes in net assets for 
the year then ended in accordance with generally 
accepted accounting principles. 


JANET M. GEuHR, C.A. 


Chartered Accountant 


North Gower, Ontario 
December 28, 2000 


Balance Sheet 
September 30, 2000 


ASSETS 


Cee e ccc essere ne seesseuscesescsssssseseseese 
ee cccccrccccccce 


eceesccccccesssces 


CAPITAL ASSETS (Note 3) 


eeccccseccsens 


Land — Alfred'Bog 2.5...) oes: 


LIABILITIES AND FUND BALANCES 


CURRENT 
Accounts payable and 
accrued: lta biUtiteSic10).eeeee ees 
Deferred Teveniew. eee 


Life membersiupsacee liane os 


NET ASSETS 
Whiresthicte dies. eicchs heat a ue 
Clubireseives,..... eS eee 
Manning interest - OFNC 

—CFN 


pecccsseee 


eco ccecoveeee 


2000 


wees) IS) 
106,939 
185,765 
19,689 


1,000 


322,012 


3,348 
$ 326,260 


$ 2,000 


133532, 
153532 
11,249 


166,721 
100,000 
1,203 
4,813 
1,308 
870 
22,941 
1,039 
584 


299,479 
$ 326,260 


Vols 


199 


$ 28,378 
206,435 
79,568 
9,287 


1,000 


324,668 


3,348 
$ 328,016 


$ 14,000 


13,800 
27,800 
8,891 


162,676 
100,000 


DONESDS) 
$ 328,016 


2001 MINUTES OF 122ND ANNUAL BUSINESS MEETING 539 


The Ottawa Field-Naturalists’ Club The Ottawa Field-Naturalists’ Club 
Statement of Operations Statement of Operations — 
For the Year Ended September 30, 2000 The Canadian Field-Naturalist 
2000 1999 For the Year Ended September 30, 2000 
2000 1999 
REVENUE 
TASPIOCESIIDS oo... c.c.-e.cscsecssceseecess $15,355 $ 14,646 REVENUE 
ivaiane Landscape...............0..... 224 226 Members Tipit ic. sccsasckceoneesayee $10,253 $ 9,776 
MURR 12s hoc se Acewaee ace issece 1,433 1725 SUL G) FOL CY eb Met Ee a 26,509 30,734 
RT TG eee canis iececascustecicacasaene 915 1132 FRE ORGS? a5 s cesccisisioteaventenbend ene cco 7,033 12,823 
Fletcher Wildlife Garden .............. ~ 1,245 Publication charges.............::000 39,888 34,498 
ON ee gcc vecueeadavacsescues 680 1,573 Interest and exchange .................. 8,519 11,198 
GrSlte ates. oct 0h 4,590 3,839 
peel 01 A? ESE Tea RR Ae 706 3,595 
OPERATING EXPENSES 97,498 106,463 
PRUMMITARIOM TOES <.ccccccoscces-ceesssescees. 1,309 1,094 
(22) 1 444 (66) EXPENSES 
2 2S L752 1,656 Jeg FES| nce sete Oe eine ee 58,461 58,707 
MPEEMCRPASSISEOUE 200. <deccccececceecos..eeeee 1,000 1,000 FRC PUGUELES secs s ose eer cence noes ayes 6,135 5,423 
Me MOTI erence cnarenca--.sencs+eassees00s 1,206 1,202 (IRC TILATIOM ec nee scsceerccte tives 6,070 12,048 
OS Od TES a 655 871 PAGANO ely cope NG sane us Sateaesetaa zak 3,258 4,541 
UCT aa 1,000 — Office Assistanit i s.ce-.dsccsecnteseean 5,000 5,000 
Ce 1,494 1710 lon OraGl aon sive noe eee eee 6,000 6,000 
WU ET DOS ie 2,037 925 CSE ee aS ge Nts 6 6,045 6,348 
Others eee ee et 2,020 3,609 


92,989 101,676 


CLUB ACTIVITY EXPENSES 
aw eee 215 165 EXCESS REVENUE OVER 
ite 875 400 EXPENSES _ $4,509 _ $4,787 _ 
Education and Publicity............... 387 217 
Excursions and lectures ............... (98) (160) 
iMacoun Field Club...................... (TS lis 783 
SOUP EE 429 304 
Tirastvand Pandscape...............:..... 8,256 7,788 
Fletcher Wildlife Garden............. Dies ve _ 
DUES 0 ea (66) 170 
13,281 9,667 
__ 24,158 __ 18,059 _ 
EXCESS REVENUE 


OVEBRCEXPENSES.................+.+- $ (5,550) $ 2,488 


540 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


The Ottawa Field-Naturalists’ Club Statement of Changes In Net Assets For the Year Ended September 30, 3000 


2000 

Net Beginning Excess Excess Donations | Ending 
Assets Balance Revenue CFN Revenue OFN Contributions Expenses Balance 
Unrestricted $ 162,676 $ 4,509 $ (5,550) $ 5,086 $ Sh $ 166,721 
Club Reserve 100,000 - - - - 100,000 
Manning — OFNC - - - 1,203 - 1,203 
Manning — CFN - - - 4,813 - 4,813 
Seedathon 1,193 - - 633 518 1,308 
Anne Hanes 
Memorial 870 . - - - 870 
de Kirilin- 
Lawrence D5 2 - - 1,404 3,674 Payot 
Macoun Baillie 
Birdathon 826 - . 213 - 1,039 
Alfred Bog 549 - - 35 - 584 

$ 291,325 $ 4,509 $ (5,550) $ 13,387 $ 4,192 $ 299,479 


The Ottawa Field-Naturalists’ Club 
Summary of Significant Accounting Policies 


September 30, 2000 
1. Nature of Business 

The organization is non-profit and incorporated under 
the laws of Ontario (1884). The organization promotes the 
appreciation, 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 information on these fields as widely as possible. It 
also supports and cooperates with other organizations 
engaging in preserving, maintaining or restoring environ- 
ments of high quality for living things. 


2. Financial Instruments 

The organization’s financial instruments consist of cash, 
accounts receivable, marketable securities, and accounts 
payable. Unless otherwise noted, it is the management’s 
opinion that the organization is not exposed to significant 
interest, currency, or credit risks arising from these finan- 
cial instruments. The fair value of these instruments 
approximate their carrying values. 


3. Capital Assets 

Capital assets acquired after 1989 are expenses. Capital 
assets acquired prior to 1990 were 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 
information. 


5. Foreign Currency 

Transactions during the year in U.S. dollars have been 
converted in the accounts to Canadian dollars at the 
exchange rate effective at the dateof the transaction. All 
monetary assets in U.S. dollars at year end have been con- 
verted to Canadian doilars at the rate effective on Sept. 30, 
2000. Gains or losses. resulting therefrom are included in 
revenue or expenses. 


oy wet 


News and Comment 


Froglog: Newsletter of the Declining Amphibian Populations Task Force (45, 46) 


Number 45, June 2001, contains: Contemporary conserva- 
tion considerations for Nigerian amphibians (G. C. Akani 
and L. Luiselli) — The DAPTF Captive Breeding Working 
Group (Andrew Gray) — Limb deformities in the Marine 
Toad in Peru and Bermuda (Donald W. Linzey) — Amphib- 
ian Status in Serbia and Montenegro (FR Yugoslavia) (Milos 
Kalezic and Georg Dzukic) — Froglog Shorts — Publi- 
cations of Interest. 

Number 46, August 2001, contains: Successful treatment 
of Chytridiomycosis (Donald K. Nichols and Elaine W. 
Lamirande) — The frog princess and other projects (Mark 
Pestov and Vladimir Anufriev) — Parasitic copepods 
responsible for limb abnormalities? (Leong Tzi Ming) — 
Reduced genetic diversity in Swiss populations of the Italian 
Agile Frog, Rana latastei (Trent Garner and Peter Pearman) 
—- Habitat fragmentation and amphibian species richness in 


Marine Turtle Newsletter (93) 


The July 2001 issue, 52 pages, contains: ARTICLES: 
Preliminary evaluation of helicopter survey as a method of 
assessing sea turtle nesting distribution in the Ten 
Thousand Islands of Florida (Ahjond S. Garmestani, H. 
Franklin Percival, and Kenneth M. Portier) — The National 
Marine Park of Zakynthos: A refuge for he Loggerhead 
Turtle in the Mediterranean (Dimitrios Dimopouls) — 
Notes: Record of pelagic East Pacific Green Turtles asso- 
ciated with Macrocystis mats near Baja California Sur, 
Mexico (Wallace J. Nichols, Louise Brooks, Melania 
Lopez, and Jeffrey A. Seminoff — Organisational profile: 
Ghana Wildlife Society (Richard Adjei, Gerard Boakye, 
and Samuel Adu) — Organisational profile: Save Our Sea 
Turtles (SOS) Tobago: A research, education and action 
programme (REAP) (Nicole Leotaud) — Third Leather- 
back Turtle stranding in Belgium (Jan Haelters, Francis 


Canadian Species at Risk May 2001 


Issued by the Committee on the Status of Endangered 
Wildlife in Canada (COSEWIC), the list is 31 pages con- 
taining [1] About COSEWIC, (mandate, membership and 
definitions); — [2] Summary Tables (COSEWIC species at 


’ risk, not at risk, and data deficient; results of May 2001 


» COSEWIC meeting), [3] COSEWIC Lists (Explanation of 


aye 


* symbols, Geographical occurrence and abbreviations; List 
» 1 Species designated in the five “risk” categories, List 2. 


Recovery: June 2001 


This issue contains: Recovery Highlights — Profile: Exper- 
ience and respect central to team’s success (David Wylynko) — 
Special Report: ‘Canadian’ Burrowing Owls found in Mexico: 
Culmination of a ten-year search (Geoffrey L. Holroyd and 
Helen Trefry — News Bites — Field Notes — Announcements 
(New Publications, Site seeing, Upcoming events, Awards) — 
Featured Species: First Nation’s involvement is Key (William 
Andrew: Oregon Spotted Frog Rana pretiosa in Frazer River 
Lowlands of southwestern British Columbia). 


riparian areas of the Parana River, Argentina (Paola M. 
Peltzer and Rafael. C. Lajmanovich) — Froglog Shorts — 
Publications of Interest. 

Froglog is the bi-monthly newsletter of the Declining 
Amphibian Populations Task Force of The World Conser- 
vation Union (IUCN)/Species Survival Commission (SSC) 
and is supported by The Open University, The World 
Congress of Herpetology, The Smithsonian Institution, and 
Harvard University. The newsletter is Edited by John W. 
Wilkins on, Department of Biological Sciences, The Open 
University, Walton Hall, Milton Keynes, MK7 6AA, 
United Kingdom; e-mail: daptf@ open.ac.uk. Funding for 
Froglog is underwritten by the Detroit Zoological 


- Institute, P. O. Box 39, Michigan 48068-0039, USA. 


Froglog can be accessed at http:// www2.open.ac.uk/biolo- 
gy/froglog/ 


Kerckhof and Thierry Jauniaux) — BooK REVIEW — 
OBITUARIES: Dr. Elvira Carrillo Cardenas (1941-2001) Fred 
H. Berry — ANNOUNCEMENTS — MEETING REPORTS — 
News & LEGAL BRIEFS — RECENT PUBLICATIONS. 

The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, School of Biological Sciences, University of Wales, 
Swansea, SA2 8PP Wales, United Kingdom; e-mail 
MTN @swan.ac.uk; Fax +44 1792 295447. Subscriptions to 
the MTN and donations towards the production of MTN 
and its Spanish edition NTM [Noticiero de Tortugas 
Marinas] should be sent to Marine Turtle Newsletter c/o 
Chelonian Research Foundation, 168 Goodrich Street, 
Lunenburg, Massachusetts 01462 USA; e-mail 
RhodinCRF@aol.com; fax + 1 978 582 6279. MTN web- 
site is: <http://www.seaturtle.org/mtn/> 


Species examined and designated in the NOT AT RISK cate- 
gory; List 3 Species examined and designated in the DATA 
DEFICIENT category) — [4] Record of Status Re-examina- 
tions — [5] List of name changes. 

It is available from COSEWIC Secretariat, Chief, Coleen 
Hyslop, c/o Canadian Wildlife Service, Environment 
Canada, Ottawa, Ontario K1A 0H3. See Web site: 
http://www.cosewic.gc.ca 


Recovery is a free newsletter providing information and 
views on the recovery of species at risk published by the 
Canadian Wildlife Service, and edited and designed by 
West Hawk Associates, Inc. It is available in either english 
or french [as Sauvegarde] from Canadian Wildlife Service, 
Environment Canada, Ottawa, Ontario, KIA 0H3 and is 
accessible at www.cws-scf.ec.gc.ca/es/recovery/archive. 
html 


541 


542 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Amphipacifica: Journal of Aquatic Systematic Biology 3(1) 16 May 2001 


ConTENTS: Norma E. Jarrett, 1931-2001: A tribute — 
The amphipod superfamily Leucothoidea on the Pacific 
coast of North America: Family Amphilochidae: systemat- 
ics and distributional ecology (P. M. Hoover and E. L. 
Bousfield) — The genus Anisogammarus (Gammaroidea: 
Anisogammaridae) on the Pacific coast of North America 
(E. L. Bousfield) — An updated commentary on phyletic 
classification of the amphipod Crustacea and its application 
to the North American fauna (E. L. Bousfield). 


Alberta Wildlife Status Reports: (32 to 36) 


The Fisheries and Wildlife Management Division of the 
Alberta Natural Resource Status and Assessment Branch, 
Alberta Environmental Protection, has released new 
Wildlife Status Reports. The Series Editor is Isabelle M. 
G. Michaud, the Senior Editor is David R. C. Prescott, 
and the illustrations are by Brian Huffman. For a listing 
earlier numbers in the series, see The Canadian Field- 
Naturalist 112(1): 169 for 1-11; 113(2): 311 for 12-17; 
113(4): 686 for 18-21; 114(1): 151 for 22-25; 115(2): 
390 for 26-31. 

Reports issued March—May 2001 are: 

32. Status of the Bay-breasted Warbler (Dendroica cas- 
tanea) in Alberta, by Michael Norton. 21 pages. 

33. Status of the Cape May Warbler (Dendroica tigrina) in 
Alberta, by Michael Norton. 20 pages. 


Amphipacifica is published by Amphipacifica Research 
Publications. Dr. E. L. Bousfield, Managing Editor, Ottawa; 
Dr. D. G. Cook, Technical Editor, Greely, Ontario. Sub- 
scriptions (4 numbers per volume) are renewable 
at $50 (Can) or $40 (US) including surface postage. Author 
charges are $25 per printed page, subject to change. For fur- 
ther information please contact Dr. E. L. Bousfield, Managi 
ng Editor, 1710-1275 Richmond Road, Ottawa, Ontario, 
Canada K2B 8E3; e-mail: elbousf@magma.ca 


34. Status of the Whooping Crane (Grus americana) in 
Alberta, by Jennifer L. White. 21 pages. 
35. Status of Soapweed (Yucca glauca) in Alberta, by 
Donna Hurlburt. 18 pages. 
36. Status of the Harlequin Duck (Histrionicus histrioni- 
cus) in Alberta, by Beth MacCallum. 38 pages. 
For copies contact the Information Centre - Publications, 


Alberta Environmental Protection, Natural Resources . 


Service, Main Floor, Great West Life Building, 9920-108 t 
Street, Edmonton, Alberta TSK 2M4, Canada (telephone: 


(780) 422-2079), OR Information Service, Alberta 
Environmental Protection, #100, 3115-12 Street NE, 
Calgary, Alberta T2E 7J2, Canada (telephone: (403) 


297-3362); or visit web site at: http//www.gov.ab.ca/ . — 


env/fw/status/reports/index.html 


Point Pelee Natural History News 1(2) Summer 2001 


This newsletter for Point Pelee, Ontario, is edited by 
Alan Wormington (e-mail: wormington@juno.com). 
Editorial Assistants are M. Lea Martell and Matthew J. 
Smith. The wed site is www.wincom.net/~fopp/ 
Natural_History_News.htm 
ARTICLES: The Colonial Waterbirds of Middle Island, 
Western Lake Erie (D. V. “Chip” Weseloh — Noteworthy 
bird records: March to May 2001 (Alan Wormington) — 
Early Migration of Bonaparte’s Gull at Point Pelee (Alan 


Wormington — IN THE FIELD — UPCOMING EVENTS AND 

OUTINGS. 
Subscription rates are Canada: CAN $15 (one year) or 

$30 (two years); International: US $15 (one year) or $30 


(two years). Send payment (and e-mail address, optional) to | : 


The Friends of Point Pelee, 1118 Point Pelee Drive, Leam- 
ington, Ontario N8H 3V4. Issues will be mailed in March, 


June, September, and December, and back issues will be a 


available for $15 per Volume/ $5 per issue (postage paid). 


$2nd Annual Meeting of the American Society of Mammalogists 


The 82nd Annual Meeting of the American Society of 
Mammalogists will be held 15-19 June 2002 at McNeese 
State University, Lake Charles, Louisiana. In addition to 
contributed oral and poster presentations covering all aspects 
of mammalian biology, this year’s program will feature two 
symposia. “Wildlife capture, handling and release: large and 
small” will be conducted by Mark Johnson DVM; 
“Macroecology of mammals: patterns, processes and possi- 
bilities” will be convened by Dawn M. Kaufman and 
Michael R. Willig. Special addresses will be offered by Drs. 
Timothy E. Lawlor (Joseph Grinnell awardee), and Theodore 


Fleming (C. Hart Merriam awardee). Also included are the 
usual ASM socials, ideal for professional interaction. 
Non-members who are interested in attending the meet- 
ings and/or presenting papers should request materials from 
the Chairman of the Local Program Committee, Dr. Gale 
Haigh (337-475-5667). For information regarding confer- 
ence arrangements, contact Dr. Haigh or Dr. Greg Hartman 
(337-475-5672). Additional information and electronic reg- 
istration is available at http://www.mcneese.edu/asm2002; 
for more information about the ASM, see their website at 
http://www.mammalsociety.org. 


TABLE OF CONTENTS (concluded) 


otes 


1 observation of a Mallard, Anas platyrhynchos, feeding on a Wood Frog, Rana sylvatica 
BRIAN R. EATON and ZACHARY C. EATON 


‘st record of an anomalously White Killer Whale, Orcinus orca, near St. Lawrence Island, 
Northern Bering Sea, Alaska SUZANN G. SPECKMAN and GAY SHEFFIELD 


idence for double brooding by a Mallard, Anas platyrhynchos, in eastern South Dakota 
JOSUA D. STAFFORD, LEASTER D. FLAKE, and PAUL W. MAMMENGA 


yportunistic foraging at American Elk, Cervus elaphus, droppings by Clark’s Nutcracker, 
Nucifraga columbiana PAUL HENDRICKS and LISA HENDRICKS 


IInet survival and healing by a Porbeagle, Lamna nascus 
GEORGE W. BENZ, ANDY KINGMAN, and JOANNA D. BORUCINSKA 


yperthermia induced mortality of gravid Snapping Turtles, Chelydra serpentina, and eggs 
in a wood chip pile SHANE R. DE SOLLA, DOUGLAS CAMPBELL, and CHRISTINE A. BISHOP 


significant new record of the Pygmy Shrew, Sorex hoyi, on the Montana-Alberta border PAUL HENDRICKS 


servation of a Golden Eagle, Aquila chrysaetos, attack on a Harlequin Duck, 
Histrionicus histrionicus, in Northern Labrador JOEL P. HEATH, GEOFF GOODYEAR, and JOE BRAZIL 


ibutes 


emembrance of John Clifton Ward, 1921-1999 DONALD R. FLOOK and JOSEPH E. BRYANT 


0k Reviews 


ology: Cuckoos, Cowbirds and Other Cheats — The Nature of Hummingbirds: Rainbows on Wings 
— Albatrosses — The Nature of Frogs: Amphibians with Attitude — Gatherings of Angels: 
Migrating Birds and their Ecology — Kingbird Highway: The Story of a Natural Obsession that Got 
a Little Out of Hand — Reproductive Biology of Bats 


ttany: Phycology — Algae 


‘vironment: Something New Under the Sun: An Environmental History of the Twentieth-Century 
World — You are the Earth, From Dinosaur Breath to Pizza from Dirt — The Alvars of Ontario: 
Significant Alvar Natural Areas in the Ontario Great Lakes Region 


scellaneous: The Essential Aldo Leopold: Quotations and Commentaries 


Ww Titles 


nutes of the 122nd Annual Business Meeting of The Ottawa Field-Naturalists’ Club 
Tuesday 9 January 2001 


vs and Comment 

roglog: Newsletter of the Declining Amphibian Populations Task Force (45, 46) — Marine Turtle 
lewsletter (93) — Canadian Species at Risk May 2001 — Recovery: June 2001 — Amphipacifica: 
ournal of Aquatic Systematic Biology 3(1) 16 May 2001 — Alberta Wildlife Status Reports: (32 to 
36) — Point Pelee Natural History News 1(2) Summer 2001 — 82nd Annual Meeting of the 
American Society of Mammalogists 


ailing date of the previous issue 115(2): 14 December 2002 


499 


501 


502 


505 


507 


510 
513 


315 


517 


520 
526 


Spa) 
530 
5) 


534 


541 


THE CANADIAN FIELD-NATURALIST Volume 115, Number 3 


Articles | 


Hunting methods and success rates of Gyrfalcons, Falco rusticolus, and Prairie Falcons, 
F. mexicanus, preying on feral pigeons (Rock Doves), Columba livia, in Edmonton, Alberta 
DICK DEKKER and JIM LANGE 


Biodiversity of adult damselflies (Zygoptera) at eastern Ontario gravel pit ponds 
PAUL M. CATLING and V. R. BROWNELL 


String and net-patterned salt marshes: rare landscape elements of boreal Canada 
KEVIN P. TIMONEY 


Predation on woodpeckers in British Columbia Eric L. WALTERS and EDWARD H. MILLER 


Timing of pregnancy, lactation, and female foraging activity in three species 
of bats in southern Illinois 
GEORGE A. FELDHAMER, TIMOTHY C. CARTER, and STEVEN K. CARROLL 


Breeding bird declines in the boreal forest fringe of western Canada: 
Insights from long-term BBS routes 
ENID E. CUMMING, KEITH A. HOBSON, STEVEN L. VAN WILGENBURG 


The spring and fall migrations of scotters, Melanitta spp., at Confederation Bridge 
in the Northumberland Strait between New Brunswick and Prince Edward Island 
PETER HICKLIN and KATHERINE BUNKER-POPMA 


New plant records for Prince Edward Island KATE MACQUARRIE and HEIDI SCHAEFER 


Status of the Deltoid Balsamroot, Balsamorhiza deltoidea (Asteraceae) in Canada 
GEORGE W. DOUGLAS and MICHAEL RYAN 


Status of Scouler’s Corydalis, Corydalis scouleri (Fumariaceae), in Canada 
GEORGE W. DOUGLAS and JUDITH A. JAMISON 


Status of Purple Sanicle, Sanicula bipinnatifida (Apiaceae), in Canada 
JENIFER L. PENNY and GEORGE W. DOUGLAS 


Status of Snake-root Sanicle, Sanicula arctopoides (Apiaceae), in Canada 
MARTA T. DONOVAN and GEORGE W. DOUGLAS 


Spatial scales of trapping in small-mammal research 
JEFF BOWMAN, CRISTINE V. CORKUM, and GRAHAM J. FORBES 


Diets of nesting Boreal Owls, Aegolius funereus, in western interior Alaska 
JACKSON S. WHITMAN 


Influence of predation on Piping Plover, Charadrius melodus, and Least Tern, 
Sterna antillarum, productivity along the Missouri River in South Dakota 
CASEY D. KRUSE, KENNETH F. HIGGINS, and BRUCE A. VANDER LEE 


Population status of shorebirds nesting at Churchill, Manitoba 
JOSEPH R. JEHL, Jr., and WINLI LIN 


Grizzly Bear, Ursus arctos, usurps Bison, Bison bison, captured by Wolves, Canis lupus, 
in Yellowstone National Park, Wyoming 
DANIEL R. MACNULTY, NATHAN VARLEY, and DOUGLAS W. SMITH 


2001 


B95 
402 


406 
413 


420 


425 


476 


480 


487 


495 


(continued on inside back cover) 


ISSN 0008-3550 


/ 


Cé 


ui 


OH 


iS 


H §=©6©9°'The CANADIAN 
FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 115, Number 4 October—December 2001 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. 
Governor General of Canada 
His Excellency John Ralston Saul, C.C. 


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 infor- 
mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- 
ing or restoring environments of high quality for living things. 


Honorary Members Robert W. New 
Edward L. Bousfield Bruce Di Labio George F. Ledingham E. Franklin Pope 
Donald M. Britton R. Yorke Edwards John A. Livingston William O. Pruitt, Jr. 
Irwin M. Brodo Anthony J. Erskine Stewart D. MacDonald Joyce and Alan Reddoch 
William J. Cody John M. Gillett Hue N. MacKenzie Mary E. Stuart 
Francis R. Cook W. Earl Godfrey Theodore Mosquin Sheila Thomson 
Ellaine Dickson C. Stuart Houston Eugene G. Munroe 


2001 Council 


President: Eleanor Zurbrigg Ronald E. Bedford Francis R. Cook Gary McNulty 
sane . f Rosanne Bishop Barbara Gaertner David W. Moore 
Bee eer eau Fenja Brodo Anthony Halliday Rita Morbia 
Recording Secretary: Ken Allison Irwin Brode Terry Higgins Robert Roach 
Treasurer: Frank Pope John Cameron David Hobden Stanley Rosenbaum 
William J. Cody Beverly McBride David Smythe 
Dorothy Whyte 


To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, Box 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 http//www.achilles.net/oinc/index.htm 


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 toward our mailing cost through the Publication 
Assistance Program (PAP), Heritage number 09477. 


Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211; e-mail: fcook @achilles.net 
Copy Editor: Wanda J. Cook 
Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 159- 1374 
Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1JO; e-mail: edith@netcom.ca 
Associate Editors: 

Robert R. Anderson Robert R. Campbell Brian W. Coad W. Earl Godfrey 

Charles D. Bird Paul M. Catling Anthony J. Erskine William O. Pruitt, Jr. 


Chairman, Publications Committee: Ronald E. Bedford 


Ail manuscripts intended for publication should be addressed to the Editor and sent by postal mail (no courier, no 
post requiring signature on delivery). Exception: book reviews should go directly to Book Review Editor. 


Subscriptions and Membership 

Subscription rates for individuals are $28 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 $28 (individual) $30 (family) $50 
(sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and mem- 
bers (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 $28 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 2001 (July 2002). 


Cover: Great Horned Owl (Bubo virginianus) holding a recently captured Richardson’s Ground Squirrel (Spermophilus 
richardsonni) at 1315 h Mountain Standard Time on 17 July 1999 near Picture Butte, Alberta, Canada. Photograph 
courtsey of Gail R. Michener. See pages 543-548. 


The Canadian Field-Naturalist 


Volume 115, Number 4 October—December 2001 


Ground Squirrels, Spermophilus richardsonii 


GAIL R. MICHENER 


Department of Biological Sciences, University of Lethbridge, Lethbridge, 
michener@uleth.ca 


Michener, Gail R. 2001. Great Horned Owl, Bubo virginianus, predation on Richardson’s Ground Squirrels, Spermo- 
philus richardsonii. Canadian Field-Naturalist 115 (4): 543-548. 


Although diurnal squirrels often form a minor component of the prey found in regurgitated pellets of Great Horned Owls, 
direct observation of the circumstances that result in a normally diurnal species appearing in the diet of a normally noctur- 
nal predator is rare. During a long-term study of the behavioural ecology of Richardson’s Ground Squirrels (Spermophilus 
richardsonii) in southern Alberta, I observed Great Horned Owls capture 14 adult ground squirrels in late February-early 
March and six juveniles in mid-summer. Temporal overlap between predator and prey occurred in two ways, through a 
shift toward twilight activity by estrous female and male ground squirrels during their mating season in spring and through 
a shift to daylight hunting by owls in summer. Twilight attacks were launched from the tops of objects such as grain augers 
and fence posts, whereas daylight attacks were launched from within the canopy of a tree. 


Key Words: Bubo virginianus, diurnal hunting, Great Horned Owl, owl pellets, predator-prey relations, Richardson’s 


Ground Squirrel, Spermophilus richardsonii. 


Common mammalian prey items in the diet of 
Great Horned Owls (Bubo virginianus) range in size 
from mice (Peromyscus) and voles (Microtus) to rab- 
bits (Sy/vilagus) and hares (Lepus) (Houston et al. 
1998). Although almost all sciurids fall within this 
size range, squirrels generally account for a minor 
fraction (usually <2% of prey items) of the diet of 
Great Horned Owls, even in areas where such squir- 
rels are abundant (Weir and Hanson 1989) and form a 
major component of the diets of sympatric buteos 
(Fitch 1947; Craighead and Craighead 1956; 
MclInvaille and Keith 1974; Gilmer et al. 1983). With 
the exception of the nocturnal flying squirrels 
(Glaucomys), other North American sciurids, whether 
tree-dwelling (Sciurus, Tamiasciurus) or ground- 
dwelling (Tamias, Spermophilus, Cynomys, 
Marmota), are diurnal. Great Horned Owls are gener- 
ally crepuscular and nocturnal hunters (Marti 1974; 
Rudolph 1978), so the low representation of diurnal 
sciurids in the diet of Great Horned Owls is attributed 
to the minimal temporal overlap in times of day at 
which squirrels and owls are active (Jaksic and Marti 
1984). Furthermore, most ground-dwelling squirrels 
in North America are hibernators, so are available to 
above-ground predators for only a portion of the year. 

Descriptions of Great Horned Ow] diets are rarely 
based on direct observation of hunting, but depend 


primarily on analyses of regurgitated pellets and sec- 
ondarily on prey remains at nests or in stomach con- 
tents. Diurnal hunting by Great Horned Owls has 
been inferred when ground-dwelling squirrels 
appear in pellets (Bent 1938, page 337; Fitch 1947, 
page 144), but the circumstances associated with 
such captures are unknown. Here I report observa- 
tions of Great Horned Owls successfully hunting 
Richardson’s Ground Squirrels (Spermophilus 
richardsonii) under two conditions, during evening 
twilight in the ground squirrels’ mating season in 
late February-early March and in full daylight during 
summer in June and July. 


Methods 

Information on Great Horned Owl predation was 
obtained incidentally during a study of the behay- 
ioural ecology of Richardson’s Ground Squirrels 
(Michener 1992, 1998, 2000) at a site located 5 km E 
and 1 km S of Picture Butte in southern Alberta, 
Canada (49°52'N, 112°43'W). The focus of the study 
was a 1.4-ha site inhabited by Richardson’s Ground 
Squirrels, but squirrels also encroached into adjacent 
cultivated fields and a farm yard. Except for a solitary 
Narrow-Leaf Cottonwood (Populus angustifolia), 
hereafter referred to as the Lone Tree, the study site 
was open grassy habitat with no cover. In the adjacent 


543 


544 


farm yard, equipment such as metal grain augers pro- 
vided potential perches for owls. Great Horned Owls 
commonly nested and roosted in a shelter belt (pre- 
dominantly cottonwoods Populus spp.) 200 m from 
the Lone Tree. 

All Richardson’s Ground Squirrels, both within 
the main site and in adjacent areas, were permanent- 
ly marked with a metal tag in each ear. All adults 
and some juveniles were individually dye-marked 
for daily visual censussing throughout the active sea- 
son from 1987 to 2000 and most adults were individ- 
ually dye-marked in 2001. The number, age, and sex 
of squirrels active in the above-ground population 
varied seasonally (Michener 1998). Generally, adult 
(21 year old) male Richardson’s Ground Squirrels 
emerged from hibernation in mid- to late February 
and adult females emerged in late February to mid- 
March. Females mated, usually in the late afternoon, 
2-5 days after emergence from hibernation 
(Michener and McLean 1996), then gave birth 
underground 23 days later (Michener 1989). Litters 
(typically of 6-8 juveniles) first emerged above 
ground when 29-30 days old, in late April and early 
May. Adult males entered hibernation in early to 
mid-June, adult females in mid-June to early July, 
juvenile females in early to mid-August, and juvenile 
males in early to mid-October (Michener 1998). 
Squirrels were live-trapped at intervals throughout 
the active season to record body mass (Michener 
1998). 

Information on daily termination of diurnal activi- 
ty of Richardson’s Ground Squirrels was obtained by 
observing dye-marked animals retire into their sleep- 
ing burrows. Observations were made either from a 
farm house adjacent to the study site or wooden 
booths within the study site. 

Because the focus of the study was on behaviour 
of ground squirrels, collection of data on the pres- 
ence and activity of Great Horned Owls was usually 
limited to periods when owl activity overlapped with 
that of ground squirrels. However, in summer 2000, 
I scanned for the presence of Great Horned Owls at 
twilight. Additionally, when owls were known to be 
frequenting the site, searches for pellets were made 
beneath perches. Pellets were inspected, and fur and 
skeletal remains of Richardson’s Ground Squirrels 
were identified with respect to reference specimens 
from the study site. Remains of other species were 
noted but not classified to species. 

All times are reported as Mountain Standard Time 
(MST). 


Results 

Retirement times were known for Richardson’s 
Ground Squirrels on 27 evenings between 17 March 
and 13 June 1996 and 59 evenings between 7 March 
and 3 July 1997. On average, retirement into the 
sleeping burrow occurred 70 + 37 (SD) min before 


THE CANADIAN FIELD-NATURALIST 


Vol-115 


sunset (n = 86 evenings on which retirement times 
were averaged for 5—12 adult females per evening). 
Ground squirrels tended to retire latest relative to 
sunset from mid-May to mid-June, but they still 
entered their sleeping burrows 35 + 21 min before 
sunset (n = 22 evenings). In June, squirrels usually 
retired between 1940 and 2020 h MST. Conse- 
quently, a Great Horned Owl that arrived between 
2045 and 2147 h (mean = 2111 h) and perched on 
grain augers on 16 evenings in June 2000 did not 
overlap with the active period of ground squirrels. In 
contrast, a Great Horned Owl that perched during the 
daytime in the Lone Tree on 20 of 43 observation 
days between 13 June and i2 August 1999 and on 4 
of 17 observation days between 26 June and 16 July 
2000 overlapped with the daily activity period of 
ground squirrels. The times of day at which the owl 
was first noted depended on concurrent studies of 
ground squirrels, but ranged from 0545 h to 1940 h 
MST. On at least seven days, the owl remained in 
the Lone Tree for minimum periods of 4-13 daylight 
hours. Only one Great Horned Owl, in adult 
plumage, was observed at any time in the Lone Tree; 
it was assumed to be the same individual throughout 
June to August 1999 and, given the similarity in 
behaviour, may have been the same individual in 
June and July 2000. 

In the Lone Tree, the owl perched within the 
canopy, 4.25—5.75 m above the ground and 1.25-— 
2.75 m below the highest point of the tree. In this 
location, the owl was partially or completely obscured 
by leaves and branches from most viewing angles 
and often blended against the bark on the tree trunk. 
Richardson’s Ground Squirrels usually assume an 
upright alert posture or give alarm calls when a 
predator is nearby, but such behaviours were rarely 
given when the owl was in the Lone Tree, suggesting 
that ground squirrels either failed to detect the owl or 
habituated to its prolonged presence. On four days 
in 2000 when the owl’s position in the Lone Tree 
enabled me to observe it with a telescope from 
115 m away, it frequently (13/21 observations) either 
changed its orientation on the branch, moved its 
head, or had its eyes open, suggesting that the owl 
was visually scanning the site. 

On three occasions, at 0700 h MST on 24 June 
1999, 1842 h on 10 July 1999, and 0712 h on 5 July 
2000, I witnessed a Great Horned Owl capture a 
Richardson’s Ground Squirrel by swooping down 
from the Lone Tree to distances of 10—15_m. 
Additionally, I saw the owl in the Lone Tree holding 
a recently captured ground squirrel in its talons (see 
cover photograph) on three other days, at 1300 h on 
17 July 1999, 1115 h on 27 June 2000, and 1510 h on 
16 July 2000, and I saw it launch unsuccessful attack 
flights of 10-30 m from the Lone Tree on five other 
days. Following the witnessed captures, the owl 
remained on the ground grasping the prey in its talons 


2001 


for several minutes before flying. After the first of 
these captures, the owl initially landed on nearby 
farm machinery, then in an adjacent crop field, before 
flying out of sight with the intact ground squirrel. 
After the other two witnessed captures, the owl took 
the prey back to the Lone Tree. Of the five ground 
squirrels carried to the Lone Tree, two were still 
intact when the owl flew with them in its talons to the 
nearby shelter belt on my approach after 15 and 60 
min. The anterior end of the other three squirrels was 
partially eaten when I checked after 70, 105, and 270 
min, confirming Errington’s (1932) observation that 
captive Great Horned Owls given live ground squir- 
rels regularly consumed the head first. The owl even- 
tually flew off with the remaining carcass in its talons 
after the elapse of a further 230, 55, and 120 min, 
respectively, suggesting that consumption of an entire 
ground squirrel usually spans several hours or is 
delayed until nightfall. However, 105 min after the 
departure of the owl with prey on 24 June 1999, an 
owl (presumed to be the same individual, but without 
prey) perched in the Lone Tree and, 11 min later, 
attempted to capture another ground squirrel. 

Adult male Richardson’s Ground Squirrels had 
already entered hibernation by early June, and the 
mean + SD date of entry into hibernation for adult 
females was 18 June + 11 days (n = 64) in 1999 and 
23 June + 11 days (n = 73) in 2000. The mean + SD 
date of birth for ground squirrels was 5 April + 6 
days in 1999 (n = 80 litters) and 3 April + 7 days in 
2000 (n = 131 litters). Consequently, when Great 
Horned Owls began frequenting the study site in late 
June, few adults were active and juvenile 
Richardson’s Ground Squirrels were 210 weeks old 
and weighed >300 g (mean + SD mass: 360 + 54 g, 
n= 38, 3 July 1999; 348 + 60 g, n = 46, 6 July 2000). 
Based on pelage colour, all six of the ground squirrels 
I saw captured by Great Horned Owls in late June 
to mid-July were juveniles, of which one (a female) 
was individually identifiable from its dye mark. 

I collected 10 Great Horned Owl pellets beneath 
the Lone Tree (eight pellets) and a grain auger 40 m 
from the tree (two pellets). Two pellets contained 
only ground squirrel remains, five pellets contained 
both ground squirrel and mouse remains, and three 
pellets contained only mouse remains. Five of the 
seven pellets with ground squirrel remains also con- 
tained either a single ear tag (four pellets) or a pair 
of ear tags (one pellet), positively identifying the 
Richardson’s Ground Squirrels as three juvenile 
males and two juvenile females. The seven pellets all 
contained cranial elements and one or both dentaries 
of a Richardson’s Ground Squirrel, and six pellets 
also contained 1-3 cervical vertebrae; no pellets 
included remnants of the appendicular skeleton of 
Richardson’s Ground Squirrels. Two pellets collect- 
ed on the same day contained mutually exclusive 
sets of similar-sized bones, so were assumed to rep- 


MICHENER: GREAT HORNED OWL PREDATION ON RICHARDSON’ S GROUND SQUIRRELS 


545 


resent two castings from the same ground squirrel. 
Another pellet contained the ear tag and remains of 
one of the observed kills. Adjusting for these pellets 
and allowing for the possibility that remains in some 
other pellets derived from ground squirrels seen cap- 
tured before the pellet was collected, the minimum 
number of Richardson’s Ground Squirrels represent- 
ed by observed captures and in pellets in the sum- 
mers of 1999 and 2000 was nine juveniles and the 
maximum was | 1 juveniles. 

In addition to diurnal hunting of Richardson’s 
Ground Squirrels in June and July of 1999 and 2000, 
a Great Horned Owl was observed capturing ground 
squirrels after sunset during the squirrels’ mating 
season in 1998 and 2001. The only time of year at 
which Richardson’s Ground Squirrels remain active 
until sunset and later is during the mating season, 
when animals engage in courtship and copulation 
late in the day (Michener and McLean 1996). Males 
and estrous females retired, on average, at 1811 h 
MST between 22 February and 4 March 1998 
(SD =15 min, range: 1740-1840 h, n= 37 retire- 
ments on 10 evenings), whereas sunset occurred at 
1800-1817 h during this period. During the next 
bout of mating activity, following a snowstorm, 
males and estrous females retired, on average, at 
1838 h between 14 and 21 March (SD = 14 min, 
range: 1810-1859 h, n = 31 retirements observed on 
five evenings), whereas sunset occurred at 
1833-1844 h. A Great Horned Owl arrived, on aver- 
age, at 1836 h on six evenings from 24 February to 
16 March 1998 and perched on fence posts, grain 
augers, or wooden booths. Four captures, assumed to 
be Richardson’s Ground Squirrels from their size, 
were observed 22-34 min after sunset, at 1822 h 
MST on 24 February, 1845 h o.n 1 March, 1849 h on 
3 March, and 1903 h on 15 March. Based on a com- 
bination of the known location of the capture and 
which individual was missing from that area the next 
morning, Great Horned Owls were assumed to have 
captured one adult female and three adult male 
Richardson’s Ground Squirrels in 1998. 

In conjunction with observations of Richardson’s 
Ground Squirrels during the mating season in 2001, 
a particular effort was made to monitor the 
behaviour of Great Horned Owls every evening 
from 3 to 19 March, the period during which 97% of 
females mated, and to videorecord predation events 
when light levels permitted. Female Richardson’s 
Ground Squirrels rarely copulated before 1745 h, 
and those females that copulated with a second male 
usually did so after 1830 h. Although most copula- 
tions occurred below ground, courtship behaviours 
occurred above ground and squirrels usually re- 
surfaced after termination of underground copula- 
tions before finally retiring for the night. 

A Great Horned Owl perched on the study site on 
14 of the 17 evenings from 3 to 19 March 2001. Only 


546 


a single owl was seen on any evening, and it arrived 
between 1838 and 1905 h MST (mean arrival time = 
1851 h, SD = 9 min, n = 13 evenings with recorded 
times), on average 21 min after sunset (SD = 6 min, 
range: 6-29 min, n= 13). Except for one evening 
with no attempts to capture prey on site, the owl was 
successful on each of the other 13 evenings, with 
captures of 10 Richardson’s Ground Squirrels, one 
vole or mouse, and two prey that could not be identi- 
fied due to low light levels. Of evenings with con- 
firmed capture of a squirrel, the owl was successful 
on either its first (n = 5 evenings) or second attempt 
(n = 5). Average elapsed time between arrival of the 
Great Horned Owl and capture of a Richardson’s 
Ground Squirrel was 11 min (SD = 4 min, range: 
2-16 min, n = 9 evenings with recorded times). 
Captures occurred, on average, at 1859 h (SD = 9 
min, range: 1848-1918 h, n = 9), 30 + 6 min after 
sunset. Of 10 successful attacks, eight were were 
launched from metal grain augers (heights = 3.32— 
4.13 m), one from a wooden booth (height = 2.14 
m), and one from a wooden fence post (height = 
1.15 m); although the Lone Tree was the tallest 
object on the site, it was never used to launch twilight 
attacks on squirrels. The average distance between 
the perch from which successful attacks were 
launched and the capture point was 41 m (SD = 28 m, 
range: 14-94 m, n = 10). Except for one capture on 
which the owl scooped up the ground squirrel in its 
talons and flew off without landing, the Great Horned 
Owl remained on the ground at the capture site, 
grasping the Richardson’s Ground Squirrel in its 
talons while occasionally pecking and tugging with 
the beak. After 0.7—2.7 min on the ground, the owl 
then flew 125-238 m (n=6 evenings when light lev- 
els permitted visual tracking), landed briefly on a 
wooden booth or on irrigation equipment in an adja- 
cent field before finally flying out of sight still car- 
rying the squirrel in its talons. The 10 witnessed 
captures included two squirrels of unknown identity 
that resided in peripheral locations, one male, and 
seven estrous female Richardson’s Ground Squirrels. 
Thus, of 13 female squirrels known to have disap- 
peared within one week of their emergence from 
hibernation, owl predation accounted for at least 
54% of those losses. Mean + SD mass of the eight 
individually identified Richardson’s Ground 
Squirrels, based on weights obtained 1—3 days before 
capture by the owl, was 269 + 63 g (range: 
195-395 g). Although a Great Horned Owl contin- 
ued to frequent the site in the evenings in late March, 
after the squirrels’ mating season had terminated, 
Richardson’s Ground Squirrels were no longer avail- 
able as prey because they all retired before sunset. 


Discussion 
Successful hunting by owls requires overlap of the 
active periods of both predator and prey (Reynolds 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


and Gorman 1999). Consequently, the most com- 
monly reported mammalian prey of Great Horned 
Owls in North America are nocturnal species such as 
Lepus, Sylvilagus, Neotoma, Thomomys, Sigmodon, 
Dipodomys, Microtus, and Peromyscus (Fitch 1947; 
Korschgen and Stuart 1972; Marti 1974; McInvaille 
and Keith 1974). Diurnal hunting has been inferred 
from daylight activity by Great Horned Owls and 
from remains of diurnal species in pellets or at nests 
(Sherman 1912; Dixon 1914; Bent 1938; Fitch 1947; 
Vaughan 1954; Maser and Brodie 1966). Although 
ground squirrels (Spermophilus) are frequently 
reported to form a minor component of the diet of 
Great Horned Owls (Bird 1929; Hamerstrom and 
Mattson 1939; Errington et al. 1940; Alcorn 1942; 
Fitch 1947; Orians and Kuhlman 1956; Seidensticker 
1968; Maser et al. 1970; McInvaille and Keith 1974; 
Gilmer et al. 1983; Jaksic and Marti 1984; Knight 
and Jackman 1984; Zimmerman et al. 1996; Murphy 
1997; Rohner et al. 2001), direct information on the 
circumstances that result in a normally diurnal 
species appearing in the diet of a normally nocturnal 
predator is rare. My observations establish that tem- 
poral overlap between Richardson’s Ground Squirrels 
and Great Horned Owls occurred in two ways, 
through a shift toward twilight activity by squirrels 
during their mating season in spring, the only time of 
year at which this otherwise strictly diurnal sciurid is 
active after sunset, and through a shift to daylight 
hunting by owls in summer. 

Seidensticker (1968) reported that Richardson’s 
Ground Squirrels accounted for 42 of 774 prey items 
(5.4%) in three collections of Great Horned Ow! pel- 
lets gathered in southern Montana, whereas 
MclInvaille and Keith (1974) found that Richardson’s 
Ground Squirrels were rarely fed to Great Horned 
Owl chicks in central Alberta despite their common- 
ness in the diets of sympatric Red-tailed Hawks 
(Buteo jamaicensis). If, as my observations suggest, 
ground squirrels tend to be captured only at certain 
periods within their active season, representation in 
the diet is likely to vary seasonally even in areas 
where some owls hunt ground squirrels. Vulner- 
ability of ground squirrels to predation by Great 
Horned Owls also may vary with species and locale. 
Richardson’s Ground Squirrels and Arctic Ground 
Squirrels (S. parryii) mate in the late afternoon 
(Michener and McLean 1996; Lacey et al. 1997), 
and thus may be at risk from predation at twilight in 
spring, whereas those species of ground-dwelling 
squirrels that mate in the morning or early afternoon 
(Murie 1995; Hoogland 1998) may be less at risk. At 
the northern limit of the geographic range of Great 
Horned Owls, where owls prey heavily on Snowshoe 
Hares (Lepus americanus) and summer daylight is 
lengthy, Arctic Ground Squirrels form a consistent 
component of owl diets (usually about 5% of prey 
items) and can account for 12-22% of prey biomass 


2001 


in years with low population density of hares (Rohner 
et al. 2001). 

Great Horned Owls are perch-and-pounce hunters 
(Houston et al. 1998) that usually perch on tall, often 
isolated, vantage points when foraging (Marti 1974; 
Rudolph 1978). Owls that hunted on my site after 
sunset perched on exposed objects, both in spring and 
summer. Even when perched in conspicuous loca- 
tions, owls are likely not conspicuous to ground 
squirrels under twilight conditions because ground 
squirrels have a pigmented eye lens and cone- 
dominant retina (Yolton et al. 1974; Jacobs 1978). 
Indeed, by remaining active after sunset during the 
mating season, estrous female and male Richardson’s 
Ground Squirrels were particularly vulnerable to pre- 
dation, to the extent that an owl hunting during the 
squirrels’ mating season in 2001 was able to capture a 
squirrel almost every evening within a few minutes of 
its arrival. During summer daylight, when visual acu- 
ity is high for ground squirrels, Great Horned Owls 
perched inconspicuously within the canopy of a tree. 
Sovern et al. (1994) suggested that diurnal hunting of 
chipmunks (Tamias) by Spotted Owls (Strix occiden- 
talis) was an opportunistic response to prey detected 
while owls were roosting. Hunting of Richardson’s 
Ground Squirrels on my study site in summer may 
initially have been an incidental outcome of the 
choice of a daytime roost, but repeated use of the tree 
and visual scanning suggest a deliberate hunting strat- 
egy by at least one Great Horned Owl. The range of 
capture times in summer (0700-1842 h MST) indi- 
cates that responsiveness to potential prey occurred 
throughout the daylight hours. 

Great Horned Owls have a long-standing reputa- 
tion (e.g., Errington 1932; Houston et al. 1998) as 
versatile predators. Bosakowski et al. (1989) noted 
that the diet of Great Horned Owls nesting in decidu- 
ous forests in north-eastern USA included more 
birds and fewer lagomorphs than is typical in more 
open habitats, and they reported that two species of 
diurnal arboreal squirrels (Sciurus carolinensis and 
Tamiasciurus hudsonicus) formed 10% of prey 
items. Bosakowski et al. (1989) did not discuss diur- 
nal hunting, but Packard (1954) noted Great Horned 
Owls striking the leaf nests of Fox Squirrels (S. 
niger) during daylight, sometimes flushing squirrels 
from the nest. My observations of twilight hunting of 
Richardson’s Ground Squirrels in the squirrels’ mat- 
ing season in spring and daylight hunting throughout 
the day in summer confirm the categorization of 
Great Horned Owls as flexible hunters by describing 
two more circumstances in which diurnal squirrels 
appear in the diet. 


Acknowledgments 

I thank J. O. Murie for helpful comments on the 
manuscript, D. R. Michener and M. D. Kujath for 
videotaping predation events in 2001, and T. D. 


MICHENER: GREAT HORNED OWL PREDATION ON RICHARDSON’S GROUND SQUIRRELS 


547 


Charge for assistance with observations in 1998. 
This research project was supported by the Natural 
Sciences and Engineering Research Council of 
Canada. 


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August 2000 
Accepted 5 March 2001 
Revised with 2001 data, 17 April 2001 


Inability to Predict Geographic Origin of Yellow-headed Blackbirds, 
Xanthocephalus xanthocephalus, During Migration 


DANIEL J. TWEDT!, GEORGE M. LINZ?, and WILLIAM J. BLEIER? 


'USGS Patuxent Wildlife Research Center, 2524 South Frontage Road, Vicksburg, Mississippi 39180 USA 

°U.S. Department of Agriculture, National Wildlife Research Center, Great Plains Field Station, 2110 Miriam Circle, Suite 
B, Bismark, North Dakota 58501 USA 

’Department of Zoology, North Dakota State University, Fargo, North Dakota 58105 USA 


Twedt, Daniel J., George M. Linz, and William J. Bleier. 2001. Inability to predict geographic origin of Yellow-headed 
Blackbirds, Xanthocephalus xanthocephalus, during migration. Canadian Field-Naturalist 115(4): 549-554. 


Yellow-headed Blackbirds (Xanthocephalus xanthocephalus) collected at different breeding locations in Alberta, 
Saskatchewan, Manitoba, and North Dakota exhibit clearly discernable morphometric differences with larger bodied birds 
found at more northern and western locations. We reduced eight skeletal measurements and body length from adult female 
and male Yellow-headed Blackbirds to their first two principal components. Principal component scores progressively 
increased at more northwestern locations. Principal component scores were also derived from measurements of birds col- 
lected in central North Dakota throughout summer and fall. We hypothesized an increase in principal component scores of 
Yellow-headed Blackbirds from summer through fall within central North Dakota as larger bodied migrants arrived and 
displaced local breeding birds. However, we were unable to detect such an increase in principal component scores from 
mid-June though mid-September over two years of study. Discriminant models that were developed to distinguish birds 
breeding in Canada from those breeding in the USA were thus poor predictors of the migratory status of Yellow-headed 
Blackbirds. Consequently, we were unable to exploit the morphometric differences inherent among Yellow-headed 
Blackbirds breeding at different geographic locations to quantify the timing or the magnitude of their migration through 
central North Dakota. 


Key Words: Yellow-headed Blackbird, Xanthocephalus xanthocephalus, principal component analysis, Alberta, 


Saskatchewan, Manitoba, North Dakota, morphology, migration. 


Identification of the patterns, timing, and duration 
of migration for passerine birds has been hindered by 
an inability to identify the origin or destination of 
birds encountered during migration. Historically, 
encounters with banded birds have been used to 
ascertain migration routes and the timing of migra- 
tion (e.g., Royall et al. 1971; Dolbeer 1982). How- 
ever, encounters of banded birds are rare. Encounters 
at two locations within the same year, that may elu- 
cidate the timing of migration, are even rarer. 
Indeed, only 246 of 40 855 Yellow-headed Black- 
birds banded between 1937 and 1968 were subse- 
quently encountered, and only 36 of these were 
direct recoveries at separate locations during the 
same year (Royall et al. 1971). Encounters of banded 
birds, however, can be markedly increased if bands 
are accompanied by auxiliary markers that can be 
detected without the recapture of individuals (Bray 
et al. 1977). 

Mass-marking of birds at en-route migration 
roosts using aerially applied micro-tags or dyes 
(Linz et al. 1991) has been successfully employed to 
mark large numbers of migratory birds (Otis et al. 
1986) that were subsequently encountered at breed- 
ing or wintering localities (Knittle et al. 1987). Even 
with this vastly increased efficiency in marking 
birds, recovery rates of marked birds remained rela- 
tively low. For example, Knittle et al. (1987) found 


that only 770 of 8880 Red-winged Blackbirds 
(Agelaius phoeniceus) collected at breeding locali- 
ties across the northern Great Plains had been 
marked during mass-marking of blackbirds at en- 
route roosts during spring. 

An alternative to physically marking individual 
birds is to use their inherent morphometric or genetic 
variation to discern their geographic origin or desti- 
nation. If geographically distinct subpopulations can 
be identified among breeding sites, these data could 
be used to identify the geographic origin of individu- 
als encountered elsewhere. Indeed, morphometric 
data have been used to suggest breeding location of 
species encountered on wintering grounds (Ramos 
and Warner 1980) and during migration (James et al. 
1984; Atwood 1989; Linz et al. 1993). Inability to 
assign individuals of widely distributed, panmictic 
species to subspecies or races — either through mor- 
phometric or genetic techniques — has hampered the 
development of these methods in the study of migra- 
tion. However, both morphometric (Zink and 
Remsen 1986; Aldrich and James 1991) and genetic 
(Zink et al. 1987) techniques have been successfully 
employed in assessing geographic variation within 
otherwise monotypic species. 

Yellow-headed Blackbirds (Xanthocephalus xan- 
thocephalus) breeding on the northern Great Plains 
of North America exhibit discernable clinal variation 


549 


550 


in morphology: larger bodied birds with relatively 
shorter limbs breed at more northern and western 
locations (Twedt et al. 1994). We attempted to 
exploit these inherent morphological differences to 
infer the geographic origin of individuals that were 
encountered during migration. Our objective was to 
determine if and when Yellow-headed Blackbirds 
that breed within central North Dakota are displaced 
by migrants. 


Methods 

From 28 May to 16 June 1987 and from 14 May to 
17 June 1988, we collected adult male and female 
Yellow-headed Blackbirds within breeding locales 
in Alberta, Saskatchewan, Manitoba, and North 
Dakota as part of a study on geographic variation 
(Twedt et al. 1993, 1994). Because after-second-year 
males can easily be distinguished from second-year 
males under normal field conditions and because few 
second-year males hold breeding territories, we only 
collected after-second-year males during May and 
June. Additionally, we collected adult male and 
female Yellow-headed Blackbirds within Benson, 
Ramsey, and Wells Counties in central North Dakota 
(48° O01’ N, 99° 40’ W) between 21 June and 18 
September during 1987 and 1988 for studies on diet 
(Twedt et al. 1991) and molt (Twedt 1990). Because 
after-second-year males cannot be separated from 
second-year males by plumage characteristics after 
their pre-basic molt is completed in mid-summer, we 
collected both adult age classes after | July. 

From collected Yellow-headed Blackbirds, we 
recorded 13 morphometric measurements (Twedt et 
al. 1994). Collection site locations, methods of 
skeletal preparation, and measurement procedures 
were described by Twedt (1990). Because pre-basic 
molt and pre-migratory fat deposition during late 
summer result in temporal changes in mass, wing 
chord, total length, and tail length, we dropped all 
morphometric variables that were temporally unsta- 
ble (Twedt and Linz 2002). However, we derived 
one additional measurement — body length. Body 
length was the difference between total length and 
tail length. Both total length and tail length were 
temporally unstable due to loss and re-growth of the 
tail during molt. However, because their difference 
negated this temporal instability, body length was 
temporally stable. 

Using principal components analysis in SAS (SAS 
Institute 1989) we reduced body length and eight 
temporally stable skeletal measurements (skull 
width, and lengths of skull, keel, ulna, humerus, 
femur, tarsus, and tibiotarsus) to their principal com- 
ponents. Sex-specific principal components were 
obtained separately for birds collected at breeding 
locations and for birds collected in central North 
- Dakota during summer and fall. Before deriving 
principal components, we replaced missing data that 


SS 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


resulted from broken or deformed bones (4% of total 
data) using regressions against the most highly cor- 
related variables (Chan et al. 1976): To assess the 
relationship between these morphometric mensura- 
tions and date of collection, we plotted principal 
component scores against date of collection. We 
reduced the variation among collection dates by con- 
structing 5-day running-averages. These 5-day run- 
ning averages were subjected to regression analysis 
to relate collection date to principal component 
scores. Finally, we discriminated between hypothe- 
sized subpopulations in Canada and the USA using 
discriminant analysis. 


Results 

We collected 176 female and 1481 male Yellow- 
headed Blackbirds from breeding locations across 
the northern Great Plains (Twedt et al. 1994). An 
additional 624 females and 865 males were collected 
after 21 June from central North Dakota. The first 
two principal components derived from morphomet- 
ric measurements were consistently, biologically 
interpretable among sexes and seasons. The first 
principal component (PC 1) represented generalized 
size whereas PC 2 represented the birds’ shape by 
contrasting axial dimensions with appendicular 
dimensions (Table 1). Generally, PC 3 contrasted 
skull and body measurements but varied slightly in 
interpretation among sexes and seasons. Because of 
similarity in interpretation between seasons, we pre- 
sent sex-specific principal component scores for 
combined data from all collected birds. We restricted 
further analyses to PC 1 and PC 2 within each sex. 
These first two principal components accounted for 
68% and 61% of the variability in females and 
males, respectively (Table 1). 

Multivariate analysis of variance of principal com- 
ponent scores detected significant differences among 
breeding locations (F, ,,,=7.04, P<0.01 female; 
F oes = 46.2, P<0.01 male). Principal component 
scores from central North Dakota were negative but 
became increasingly positive at more northern and 
western locations as larger bodied birds with rela- 
tively shorter appendages were encountered (Table 
2). Thus we hypothesized that migrants encountered 
in central North Dakota, that had originated in more 
northwestern breeding locations, would have greater 
PC scores than did Yellow-headed Blackbirds that 
bred in North Dakota. — : 

To examine the hypothesis of increasing PC 
scores over time, we plotted 5-day running averages 
of PC scores against dates of collection (Figure 1). 
Although average PC scores derived from female 
measurements slightly increased over time, the slope 
of neither PC 1 nor PC 2 differed significantly from 
zero (t> 1.19, P>0.21). Similarly, PC scores for 
males exhibited no significant trend over time 
(¢>:1.58,P S012): 


2001 


TWEDT, LINZ AND BLEIER: GEOGRAPHIC ORIGIN OF YELLOW-HEADED BLACKBIRDS 


551 


TABLE |. Principal component (PC) loadings on nine temporally stable variables for Yeliow-headed Blackbirds. 


PC | 

Variable Female Male 
Body length 0.105 0.157 
Skull length 0.236 0.261 
Skull width 0.214 0.164 
Keel length 0.357 0.244 
Femur length 0.381 0.424 
Tarsus length 0.415 0.419 
Tibiotarsus length 0.413 0.436 
Humerus length 0.421 0.370 
Ulna length 0.309 0.371 
Eigenvalue 5.03 4.40 
% variance explained 33529 48.9 
Cumulative variance 55.9 48.9 


Because a previous analysis (Twedt et al. 1994) 
detected two discernable subpopulations within the 
overall morphometric cline exhibited by Yellow- 
headed Blackbirds, we attempted to predict mem- 
bership in these subpopulations using discriminant 
analysis. We first used stepwise discriminant analy- 
sis to reduce the number of morphometric variables 
to four for females (body length, skull length, hu- 
merus, and femur) and five for males (body length, 
skull length, skull width, keel, and tibiotarsus). Using 
birds collected at breeding locations and discriminat- 
ing between birds breed ing in Canada and those 
breeding in the USA, the four variable discriminant 
model for females had a 0.70 (CI,,., = 0.65—0.75) 
probability of correctly classifying birds (i.e., 70% of 
birds were correctly classified to their collection 
location). Similarly, the five variable model we used 
for males had a 0.68 (CI,,., = 0.66—0.70) probability 
of correct classification. Kappa statistics (Titus et al. 
1984) for both female and male models were signifi- 
cantly greater than zero (k > 0.35, SE <0.11, 
z>3.37, P<0.01) which indicated that both models 
predicted significantly better than chance. 

When we applied these discriminant models to 
Yellow-headed Blackbirds collected in central North 
Dakota during summer and fall, we classified 232 of 
601 females and 190 of 936 males as migrants (i.e., 


PC loadings 
PC 2 PC 3 

Female Male Female Male 

0.441 0.617 0.831 -0.577 

0.480 0.392 -0.475 0.009 

0.670 0.542 -0.202 0.767 
-0.153 0.184 -0.038 -0).273 
-0.074 -0.155 -0.032 0.009 
-0.175 -0.160 0.028 0.007 
-0.186 -0.158 0.023 -0.012 
-0.180 -0.178 0.029 0.019 

0.015 -0.177 0.196 0.055 

1.09 1.06 0.95 0.86 
[2h On 10.5 9.6 
68.0 60.6 78.5 70.2 


birds with geographic origins to the northwest of the 
collection location). The proportion of migrants 
within 5-day collection periods ranged from 0.09 to 
0.59 for females and from 0.04 to 0.39 for males 
(Figure 2). However, regression slopes of the pro- 
portion of migrants against time did not differ from 
zero (t= 1.376, P=0.17) for females and had a sig- 
nificant negative slope (t=-4.857, P<0.01) for 
males. 


Discussion 

We were unable to detect our hypothesized in- 
crease in principal component scores over time with- 
in central North Dakota. Indeed, average PC scores 
tended to decrease slightly from June through 
September. Discriminant models constructed to distin- 
guish birds breeding in Canada from those breeding in 
the USA similarly were unable to detect trends in the 
migratory status of Yellow-headed Blackbirds in cen- 
tral North Dakota. Failure of these models was likely, 
in part, due to their relatively poor ability to discrimi- 
nate between local breeding birds and birds breeding 
at more northwestern locations — only 70% of birds 
could be correctly classified. In fact, the inferred 
proportion of migrants that resulted from application 
of our discriminant models remained constant or 
declined during summer and fall. Neither of these 


TABLE 2. Mean principal component scores for Yellow-headed Blackbirds collected at breeding locations during 1987 and 


1988. 


Location 


Alberta and western Saskatchewan 
Central Saskatchewan 

Southern Saskatchewan and Manitoba 
Central North Dakota 


alncludes all females collected in Saskatchewan and Manitoba. 


PET i Gee 
Female Male Female Male 
0.546 0.751 0.365 0.205 
0.1438 0.573 -0.078@ 0.065 
-0.015 0.206 
-0.822 -0.909 -0.341 -0.339 


52 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


female 


S 
A. 
0 
2 
205 
O 
A4 
al 
ak 


Thule! en Senal 


1 Aug 


15 Aug 15 Sep 


1 Sep 


FIGURE |. Five-day running average of principal component (PC) scores derived from 8 
skeletal mensurations and body length of Yellow-headed Blackbirds collected in cen- 
tral North Dakota during 1987 and 1988. For any 5-day period, mean sample sizes 
were 63 (range 7-172) for females and 55 (range 7-161) for males. 


scenarios conformed with our hypothesis of an 
increase in the proportion of migrants over time. 
Several possible explanations could account for 
our inability to detect late-summer migrants passing 
through central North Dakota. Most likely, our prin- 
cipal component and discriminant function models 
had insufficient power to discriminate along the 
morphometric cline exhibited by Yellow-headed 
Blackbirds. Our ability to detect migrants may have 
- also been hindered by a failure of migrants to dis- 
place locally breeding birds but instead congregating 


in heterogeneous populations within central North 
Dakota throughout late summer. 

Alternatively, despite data from returns of banded 
birds that indicate a southeasterly migration route is 
used by Yellow-headed Blackbirds, it is possible 
that most migrants passing through our central 
North Dakota study area may have originated in 
more proximate locations in southern Manitoba and 
eastern Saskatchewan. Populations within these rel- 
atively nearby breeding locations share greater mor- 
phometric similarity with birds breeding in central 


2001 


0.6 


0.5 


Proportion Migrants 
oO 


So 
N 


0.1 
Females 


Males 


TWEDT, LINZ AND BLEIER: GEOGRAPHIC ORIGIN OF YELLOW-HEADED BLACKBIRDS 


553 


September 


FIGURE 2. Proportion of male and female Yellow-headed Blackbirds classified as migrants in 
central North Dakota during 1987 and 1988. Plotted proportions are running averages 
based on 5-collection-day samples. Mean sample sizes were 54 (range 15-134) for 
females and 78 (range 29-166) for males. 


North Dakota (Twedt et al. 1994), thereby increas- 
ing the difficulty of distinguishing differences 
between these populations using either principal 
components analysis or discriminant function analy- 
SiS. 

In summary, although distinct morphometric dif- 
ferences exist between Yellow-headed Blackbirds 
breeding in central North Dakota and those breeding 
at more northwestern locations in Canada, we were 
unable to exploit these differences to elucidate 
either the duration or magnitude of migration 
through central North Dakota. Unless refinement of 
these morphometric techniques can be achieved, it 
appears that elucidating migratory movements of 
this species will continue to rely on physically 
marking individual birds or the development of 
improved genetic markers. 


Acknowledgments 

We thank B. Osborne for his dedicated assistance 
in the field and laboratory. Additional laboratory 
assistance was provided by J. Lindlauf, and B. 
Mautz. We are deeply indebted to the numerous 
landowners who granted access to their property. 
Additionally we thank personnel from state, provin- 
cial, and federal wildlife agencies for their coopera- 
tion. Funding was provided by the USDA National 
Wildlife Research Center and North Dakota State 
University. 


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Bray, O. E., W. C. Royall, Jr., and J. L. Guarino. 1977. 
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Knittle, C. E., G. M. Linz, B. E. Johns, J. L. Cummings, 
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Linz, G. M., C. E. Knittle, J. L. Cummings, J. E. Davis, 
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THE CANADIAN FIELD-NATURALIST 


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Twedt, D. J. 1990. Diet, molt, and geographic variation in 
Yellow-headed Blackbirds, Xanthocephalus xantho- 
cephalus. Ph.D. dissertation, North Dakota State Univer- 
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Twedt, D. J.. W. J. Bleier, and G.M. Linz. 1991. 
Geographic and temporal variation in the diet of Yellow- 
headed Blackbirds. Condor 93: 975-986. 

Twedt, D. J., W. J. Bleier, and G. M. Linz. 1993. 
Genetic variation in male Yellow-headed Blackbirds 
from the northern Great Plains. Canadian Journal of 
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Twedt, D. J., W. J. Bleier, and G. M. Linz. 1994. Geo- 
graphic variation in Yellow-headed Blackbirds (Xantho- 
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Received 9 February 2000 
Accepted 1 September 2001 


Review of the Status of the Northern Abalone, 
Haliotis kamtschatkana, in Canadat 


G. S. JAMIESON 
Department of Fisheries and Oceans, Pacific Biological Station, Nanaimo, British Columbia V9R 5K6 Canada 


Jamieson, G. S. 2001. Review of the status of the Northern Abalone, Haliotis kamtschatkana, in Canada. Canadian Field- 
Naturalist 115(4): 555-563. 


The Northern Abalone (Haliotis kamtschatkana) is the only abalone species in Canada and is widely distributed along the 
entire outer coast of British Columbia. In the late 1970s and 1980, there was a commercial fishery, but following a peak in 
landings in 1978-1979, landings rapidly declined. Its distribution in commercially exploitable concentrations is patchy, 
making stock assessments difficult to conduct. However, in the late 1980s, consistently low estimated abundance levels 
and increasing evidence for a substantial illegal harvest of abalone raised conservation concerns, resulting in aboriginal, 
recreational and commercial fisheries closures in 1990. Studies in other abalone species suggest that abalone larvae dis- 
perse over relatively short distances. If true for Northern Abalone as well, then this species may be particularly vulnerable 
to localised extirpations. Through the 1990s, there was no evidence of significant Northern Abalone recruitment along 
Canada’s Pacific coast, and because of a high price, abalone poaching is still occurring. Expansion in the range of Sea 
Otters (Enhydra lutris), a major abalone predator, is also increasing natural mortality. Given these circumstances, a 


COSEWIC status of “threatened” was recommended. 


Key Words: Northern Abalone, Haliotis kamtschatkana, British Columbia, distribution, stock status. 


Jamieson (1989) reviewed the status of Northern, 
or Pinto, Abalone, Haliotis kamtschatkana Jonas, 
1845, along the coast of British Columbia, Canada, 
but while the paper was accepted and reviewed by 
COSEWIC in April, 1988, no status could be 
assigned as COSEWIC at that time had no mandate 
to assess invertebrates. COSEWIC’s mandate 
changed in 1994, however, and COSEWIC is now 
able to assign status to molluscs. Here, I review the 
current status of this species, and note changes in 
abundance and fisheries management which have 
occurred since my first report (Jamieson 1989). That 
report included sections on Northern Abalone gener- 
al biology, limiting factors and special significance 
that are still relevant, and so are not repeated here. 

The Northern Abalone (Figure 1) is the world’s 
northernmost abalone species. Although eight 
abalone species occur in the Northeast Pacific off 
California (Cox 1962), there is only one species in 
the marine waters of Washington State, British 
Columbia, and Alaska. McLean (1966) provided the 
following description for it: “Shell relatively small, 
thin, elongate-oval, low. Open holes 3 to 6, on tubu- 
lar projections. Broad channel present on the body 
whorl between the suture and row of holes. 
Sculpture of irregular bumps superimposed over spi- 
ral sculpture of broad ribs with weak spiral ribs in 
interspaces. Colour mottled reddish or greenish with 
areas of white or blue. Shell margin narrow. Muscle 
scar lacking, interior pearly white with faint irides- 
cence of pink and green.” British Columbia speci- 


+Threatened status assigned by COSEWIC in April 1999. 


mens differ from Californian specimens in not hay- 
ing a broad channel in the body whorl and in having 
a less complex distribution of lumps in the spiral 
sculpture and a muscle scar on the anterior of some 
shells (McLean 1966). 


Distribution 

As noted by Sloan and Breen (1988) and Jamieson 
(1989), the Northern Abalone is found from Sitka 
Island, Alaska (57° N; Paul and Paul 1981) to Turtle 
Bay, Baja California (27.5° N; McLean 1966). In 
central California, the typical form merges into the 
subspecies Haliotis kamtschatkana assimilis Dall 
(Threaded Abalone), which occupies the southern 
part of the range (McLean 1966). The type locality 
for Haliotis kamtschatkana, “near Unalaska, Kam- 
chatka Sea’, is evidently in error since there are no 
records of Haliotis occurring at any point along the 
Aleutian Islands (McLean 1966). At its northern 
range limit, the Northern Abalone occurs from the 
lower intertidal zone to a depth of at least 100 m, 
whereas near its southern range limit, it is strictly 
subtidal, with most individuals occurring at depths 
between 10 to 20 m (McLean 1966). In British 
Columbia, most of the adult population is found at < 
10 m depth (Mottet 1978). Northern Abalone prefer 
a firm substrate, usually rock, and are generally 
found in areas of moderate water exchange, such as 
occurs on exposed or semi-exposed coasts. They are 
patchily distributed within this habitat. 


Habitat 
There is no indication that there has been any per- 
manent physical deterioration in Northern Abalone 


IID 


556 


FIGURE |. Northern Abalone shells. A. dorsal view. B. ven- 
tral view. 


habitat in British Columbia over time. Fluctuation in 
biotic factors such as food availability and predators 
may vary substantially on a local scale, and catastro- 
phes such as oil spills and human-induced, increased 
sedimentation can have short-term deleterious conse- 
quences. In British Columbia, greatest deterioration 
in marine water quality or disruption of substrate by 
either industry or urbanisation has occurred in shel- 
tered waters (e.g., harbours), either around estuaries 
or in the Strait of Georgia. Northern Abalone are not 
greatly affected by this as they prefer cool, exposed 
waters. However, the eradication of Sea Otters 
(Enhydra lutris) in the late 1800s through hunting 
had ecosystem implications that likely affected 
abalone. Herbivorous prey items of Sea Otters, 
notably sea urchins (Stronglyocentrotus spp.) and 
abalone, likely subsequently increased dramatically 
in abundance. This increase resulted in the establish- 
ment of “sea urchin barrens’, which are large areas 
populated with urchins and virtually devoid of 
marine macroalgae. Sea urchins and Northern 
Abalone may compete for food, resulting in food- 
limited environments where abalone growth can be 
stunted (stunted abalone are locally called “surf” 
abalone) and larval dispersal from local populations 
can be reduced. When “surf” abalone are transplant- 
ed to kelp-abundant habitats, they commence grow- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


ing again (Emmett et al. 1988), suggesting that food 
limitation can result in growth rate reduction. 


Historical First Nation Utilisation 

Diversity was a key to sustainable resource use by 
coastal First Nations peoples. Along with other 
species, abalone were used for food and the shells 
were used for decoration and as currency (Turner 
1997). Virtually all areas of the land, water and shore- 
line fell under the specific “ownership” of one indi- 
vidual, usually a hereditary chief, who, while the 
recipient of the benefits of using the area, also had the 
responsibility of maintaining and sustaining the 
resources and sharing them equitably among other 
community members. All productive areas were 
closely watched and managed to ensure resource sus- 
tainability because the availability of species such as 
abalone directly affected the survival of these hunter- 
gatherers. There is no documented evidence that 
Canadian indigenous peoples dove for abalone, so the 
abalone that were harvested came only from the inter- 
tidal or shallow subtidal zones (Campbell 2000). The 
lack of diving limited the ability of native peoples to 
exploit abalone concentrations extensively, as most 
abalone concentrations occur at a depth of 0 to 10 m 
in high wave-energy areas (Sloan and Breen 1988). 
Historically, most waters deeper than 1 m would have 
functionally represented a natural abalone refugium 
from human harvesting (Jamieson 2000). 

Historically, the major predator of adult abalone 
was the Sea Otter, not humans, as abalone are a pre- 
ferred prey of these otters (Johnson 1982; Estes and 
Van Blaricom 1985). However, because native peo- 
ple hunted Sea Otters as well as abalone and Sea 
Otters likely avoided the immediate areas around 
native villages (Russ Jones, Haida Fisheries 
Program, Skidegate, British Columbia, personal 
communication). Coastal human habitations were 
historically relatively abundant along the coast, as 
indicated by the many coastal middens that can be 
seen today, and their presence may have created 
many refugia from Sea Otters where abalone could 
have occurred in at least modest abundance subtidal- 
ly. Both Sea Otters and indigenous human popula- 
tions were devastated almost simultaneously in the 
early 1800s by the arrival of Europeans, the former 
by hunting and the latter by diseases such as small- 
pox. 


Population Size and Trends 

Sloan and Breen (1988) and Jamieson (1989) 
both described factors influencing the abundance of 
abalone in British Columbia up until the mid-1980s. 
Campbell (2000) has summarised Northern Abalone 
stock status in British Columbia more recently. 
Briefly, the suggested relatively high abundance of 
abalone in the early 1970s (Figure 2) may have been 
influenced by both the absence at that time of sig- 


2001 


nificant abalone fisheries and the eradication of Sea 
Otters. In areas where Sea Otters have been re- 
introduced in British Columbia, almost all surviv- 
ing abalone are found only in rock crevices, whereas 
outside the range of Sea Otters, abalone are abundant 
on open rock faces outside crevices (Hines and 
Pearse 1982; Watson 1993). It is hypothesised that 
after Sea Otters were eradicated and First Nations 
peoples decreased by disease, abalone populations 
may have increased substantially in abundance by 
moving into more open habitats. The potentially 
numerous, albeit likely small, and geographicaily 
isolated abalone populations that might have existed 
along the coast would have been released from 
major predation pressures and may have gradually 
expanded in both size and number to ultimately sup- 
port the recent commercial fisheries. 


500 


JAMIESON: STATUS OF THE NORTHERN ABALONE IN CANADA S57 


The modern industrial (i.e., export) abalone fish- 
ery began in British Columbia around 1975. Surveys 
of abundance have been conducted since 1978 
(Table 1). Reported annual landings peaked in 1977 
to 1978 at around 400 t (Figure 2), before being 
reduced by a quota to 226.8 t in 1979 and 113.4 t in 
1980. Quotas continued to be reduced until they lev- 
elled out at 47.2t from 1985 to 1989. At the time, any 
smaller quota was considered to be impractical, as 
existing limited-entry fishers would then be unable to 
make a desirable living. The only other option was 
fishery closure. Although efforts were being made to 
manage the fishery to a sustainable level of produc- 
tion, there were little data to indicate what this level 
might actually be. The abundance of legal-size 
abalone may have declined by as much as 60 to 90% 
by 1978 (Sloan and Breen 1988) and by the end of 


@ Reported Landing (t) 


400 


@ Quota (t) 


300 


Landings (t 


200 


100 


Year 


FIGURE 2. Reported Northern Abalone landings and quotas between 1972—1998 in British Columbia. 


558 THE CANADIAN FIELD-NATURALIST Vol. 115 

TABLE |. Surveys of abalone abundance by region in northern British Columbia. 
Queen Charlotte 

Year Central Coast Islands Reference 
1978 x Breen and Adkins 1979 
1979 X Breen and Adkins 1981 
1983 x Boutillier et al. 1984 
1984 X Boutillier et al. 1985 
1985 X Farlinger and Bates 1986 
1987 x Carolsfeld et al. 1988 
1989 X Farlinger et al. 1991 
1990 x Thomas et al. 1992 
1993 x Thomas and Campbell 1996 
1994 K Winther et al. 1995 
1995—96 Xx Cripps and Campbell 1998 
1997 X Campbell et al. 1998 


1980 was only ~450t (Breen 1986). Annual produc- 
tion was initially assumed to be substantial because 
of the large initial biomass, but data from the mid- 
1980s indicated that annual recruitment was occur- 
ring at less than predicted levels (Breen 1980). 

Concern about the apparent continuing decline in 
abalone abundance was regularly expressed in scien- 
tific advice to resource managers. The concern was 
never that abalone as a species would go extinct, but 
rather that local, accessible populations might either 
be extirpated or become so small that abalone would 
be difficult to find. At the least, abundance might 
become so low that abalone fisheries of any type 
could not be rationalised. This concern was reflected 
by reductions in annual quotas in the early 1980s, 
but logistic difficulties in sampling this widely and 
contagiously distributed species made the collection 
of comprehensive fisheries-independent abundance 
data difficult. In addition, biologists studying the 
species were not sure if the observed low recruitment 
at that time (Tables 2, 3) was part of a natural cycle 
in abundance determined by natural causes, or if it 
was indicative of over-exploitation. Because limited 
time-series data made scientific analysis ambiguous, 
and because of strong lobbying by industry for con- 
tinuation of at least a reduced fishery, resource man- 
agers were hesitant to close the fishery. Instead, they 
gradually reduced annual quotas through to 1985 to 
the minimum level that would support existing fish- 
ers (Adkins 2000). 


Abalone are broadcast spawners, and Northern 
Abalone recruitment characteristics were initially 
assumed to be similar to that of other local molluscs, 
notably bivalves. However, abalone recruitment 
appears to be different, and it is now recognised that 
their biology must also be considered in attempting to 
understand why Northern Abalone recruitment was 
not as high as was initially expected. World-wide, 
many abalone species have declined significantly in 
abundance coincident with the establishment of inten- 
sive export fisheries. It is now recognised that as a 
group abalone seem relatively vulnerable to over- 
exploitation (Tegner et al. 1996), and that particular 
attention must be given to monitoring and controlling 
their harvest (Tegner and Butler 1985; Prince et al. 
1987, 1988; McShane et al. 1988; McShane 1992, 
1995a,b; Campbell 2000). Reasons for this are not 
well understood for most abalone species, but Prince 
et al. (1988) suggested why this might be so in 
Haliotis rubra in Australia. Somewhat unique among 
exploited marine invertebrates, abalone have a partic- 
ularly short larval period, typically being less than 10 
days. In the early 1980s, it had been generally 
assumed that free-swimming meroplanktonic larvae 
dispersed relatively widely (Fedorenko and Sprout 
1982). However, after considering a simple experi- 
ment, Prince et al. (1987) suggested that Haliotis 
rubra recruitment from a specific population was lim- 
ited to the immediate vicinity of conspecifics. They 
hypothesised that dispersal of larvae for this species 


TABLE 2. Abalone densities (number m~) from 25 comparable sites in the Central Coast resurveyed in 1983, 
1985, 1989 and 1993 (from Thomas and Campbell 1996). 


Year 
Cohort 1983 1985 1989 1993 1997 
Total abalone AS a) 0.56 0.53 0.53 
Legal size (100+ mm) O22 0.34 0.11 0.09 0.09 
Prerecruits (94-101 mm) 0.18 0.25 0.08 0.06 0.06 
Recruits (102-107 mm ) 0.10 0.14 0.03 0.03 0.03 


2001 JAMIESON: STATUS OF THE NORTHERN ABALONE IN CANADA 


TABLE 3. Abalone densities (number m-*) from 28 comparable sites in the Queen Charlotte Islands resurveyed 


in 1984, 1987, 1990 and 1994 (from Winther et al. 1995). 


Cohort 1979 1984 
Total abalone 3.54 0.69 
Legal size (100+ mm) 0.30 0.09 
Prerecruits (92-99 mm) 0.27 0.08 
Recruits (100-106 mm ) 0.14 0.04 


Year 

1987 1990 1994 
0.79 0.44 0.33 
0.13 0.07 0.06 
0.07 0.04 0.03 
0.05 0.04 0.02 


559 


was generally limited to < 50 m. Prince et al. (1988) 
then conducted a more extensive study to further test 
this hypothesis and they concluded that larval disper- 
sal for this species could be as small as 10 to 100m 
and that limited dispersal was indeed the most likely 
explanation for their observations. Thus, widespread 
dispersal of meroplanktonic larvae for species where 
this has not been shown to occur should not always be 
assumed, particularly if nothing is known about larval 
behaviour under field conditions. Prince et al. (1988) 
noted that concentration of fishing within specific 
areas may have serious negative effects on abalone 
populations and could lead to recruitment overfishing. 

We currently have no data about the dispersal 
characteristics of Northern Abalone larvae (Campbell 
2000), but it seems appropriate to assume that this 
species also has limited larval dispersal. Abalone 
fishing in Canada involves divers finding concentra- 
tions of abalone, and then harvesting all they can 
find above the minimum legal size limit. As previ- 
ously cryptic animals become accessible over time, a 
population may gradually be depleted of most of its 
spawning adults, particularly if the minimum legal 
size limit is not the most appropriate. Poaching, 
where size limits would be less likely considered, 
would only make the situation worse. 

Specific details of the decline in abundance may 
also be important. While 50% of Northern Abalone 
were estimated to be mature at a 55-mm shell length, 
the importance of large mature female abalone in 
contributing to total population fecundity has proba- 
bly been underestimated. For example, at an eastern 
Moresby Island study site in 1990, 20% of mature 
female abalone were above the legal size of 100 mm 
shell length, yet these large abalone were estimated 
to produce 50% of the total potential eggs released 
by that population (Campbell et al. 1992). The fish- 
ery targeted larger animals (Sloan and Breen 1988), 
which may have had a greater impact on population 
fecundity than realised in the early 1980s. There is no 
described stock-recruitment relationship for the species 
(Campbell 2000), but a precautionary approach 
(Richards and Maguire 1998) argues that management 
should maintain gamete production at a level that 
accommodates uncertainty. 

Another major problem was illegal harvesting, or 
poaching (Farlinger and Thomas 1989; Farlinger 


1990; Campbell 2000; Jubinville 2000). Coincident 
with a reduction in the legal quota for abalone, dive 
fisheries for both Geoduck (Panopea abrupta) and 
Red Sea Urchins (Stronglyocentrotus franciscanus) 
were expanding. Expansion of sea urchin fishing 
was particularly important because urchins and 
abalone occur in the same habitat and depth range. 
Many commercial and recreational divers were thus 
encountering abalone on a regular basis. Reduced 
quotas for abalone increased their price per kilogram, 
making it increasingly attractive for poachers to har- 
vest abalone illegally. 

Subsequent to the summary documents of Sloan 
and Breen (1988) and Jamieson (1989), resurveys of 
harvested areas continued in northern British 
Columbia (Table 1), with either the south-east Queen 
Charlotte Islands or the Central Coast sampled in 
any year. In 1987, it looked like abalone recruitment 
in the Queen Charlotte Islands might be finally 
increasing (Table 3), but no similar pattern was 
observed in the 1989 survey of the Central Coast 
(Table 2). Conservation of the species was now con- 
sidered to be a significant issue, but before the fish- 
ery was closed, it was decided to conduct one more 
survey in the Queen Charlotte Islands to determine if 
the recruitment pulse observed in 1987 was being 
sustained. Results of the 1990 survey (Table 3) indi- 
cated it was not, and so the Department of Fisheries 
and Oceans (DFO) took the then-unprecedented step 
of closing all abalone fishing, initially for five years 
(Farlinger 1990). However, subsequent surveys 
(Tables 2, 3) demonstrated no stock rebuilding 
(Campbell 2000), and so aboriginal, recreational and 
commercial fisheries remain closed indefinitely 
(Adkins 2000). 

Abalone abundance in southern British Columbia 
was also reduced to a low level by overfishing in 
the early years of the fishery (Adkins 1996), with 
again little sign of population recovery to date. There 
remains a population of large, relatively old abalone 
in the immediate vicinity of William’s Head 
Penitentiary near Victoria, British Columbia, as pen- 
itentiary guards discourage nearshore access to fish- 
ers and poachers in an effort to minimise opportunity 
for inmate escape (Wallace 1997). 

Establishing recreational and commercial fishery 
closures does not mean that all abalone fishing 


560 


stopped. Illegal fishing is still considered to be sub- 
stantial in British Columbia (Adkins 2000; Campbell 
2000; Jubinville 2000). Undocumented and poten- 
tially unreliable sources suggested that the illegal 
fishery in 1990 at the time of the fishery closure may 
have been up to five times the size of the then-legal 
fishery (47.2 t). There is still an indication that the 
current illegal harvest is up to several times the pre- 
closure legal harvest, and enforcement of a closure 
by the DFO is proving difficult. DFO fishery officers 
try to enforce the law (Jubinville 2000), but abalone 
volumes landed at any one time are often small and 
hidden with other much larger volumes of other 
legally-landed species. There have been a few signif- 
icant arrests and convictions involving thousands of 
abalone (Jubinville 2000). Other fishery issues, 
notably management of herring and salmon fishing 
during the spring, summer or fall when most abalone 
are likely poached, have been of higher priority for 
fishery officers and there has been relatively little 
allocation of resources to address effectively the 
illegal abalone harvest. Many Asian restaurants in 
Vancouver in particular have openly listed abalone 
on their menus, but the paper trail to identify their 
sources is time-consuming and difficult to follow 
because the species can be harvested elsewhere in 
the north-east Pacific. The problem is a little like the 
illegal drug trade - value and potential gain are now 
so high that with the relatively limited law enforce- 
ment resources available, all illegal fishing cannot 
realistically be stopped or curtailed (Jamieson 2000). 
Existing marine protected areas (Jamieson and 
Lessard 2000) in British Columbia appear to offer 
little additional protection for Northern Abalone, as 
enforcement of fishing regulations at these locations 
is generally no better than the existing enforcement 
of fishery restrictions anywhere on the entire coast. 

Washington State has never had commercial fish- 
ing for Northern Abalone, and Alaska’s commercial 
fishery was closed in 1995 (Woodby et al. 2000). 
The recreational fishery for abalone in Washington 
was closed in 1994, also because of conservation 
concern (B. Sizemore, Washington Department of 
Fish and Game, Olympia, Washington, personal 
communication). An Alaskan sport fishery for 
abalone (minimum legal size = 89 mm (3.5 in)) cur- 
rently exists, but abalone cannot be fished with 
SCUBA gear, which means that harvests are only 
through snorkelling or handpicking in the intertidal 
one. The magnitude of abalone poaching in Alaska 
and Washington State is undocumented. 

In recent years, methodology used to survey 
abalone stocks in British Columbia has been evaluat- 
ed (Campbell 1996) and criteria proposed that if met, 
could justify reopening Northern Abalone fisheries 
in British Columbia (Campbell 1997). A common 
method has been used in all surveys to date to pro- 
vide a series of comparable data, but concern exists 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


that this method is not sufficiently precise to detect 
small changes in population density (Farlinger and 
Campbell 1992). Campbell (1996) described the 
advantages of modifying the present survey tech- 
nique, but these changes have been only minimally 
implemented, in part because major a methodology 
change would mean starting a new data series and 
result in a short-term inability to compare data. 
Criteria for reopening the fishery can not be fully 
defined at the present because of a lack of data on 
the frequency and patch size of adult Northern 
Abalone concentrations that would be required to 
maintain sufficient recruitment for a healthy popula- 
tion. As mentioned above, studies on other abalone 
species indicate that larval dispersal is not as exten- 
sive as with other mollusc broadcast spawners. This 
makes the spatial distribution of abalone patches 
important. Also, dilution of gamete concentration 
through reduced adult spawner densities can reduce 
fertilisation success (Clavier 1992; McShane 1995a, 
1995b; Shepherd and Partington 1995), making the 
number of spawning abalone and their localised spa- 
tial distribution in any concentration important. 

The range of Sea Otters in British Columbia is 
expanding and their abundatice is increasing (aver- 
age rate of 18.6% per year on the west coast of 
Vancouver Island; Watson et al. 1997; Watson 2000), 
following their reestablishment at a few isolated loca- 
tions in the period 1969-1972. The population has 
increased from 89 animals introduced between 1969 
to 1972 to over 2500 in 1998 in two disjunct popula- 
tions (Watson 2000): about 2000 along northwest 
Vancouver Island and about 500 in the central por- 
tion of the British Columbian coast in Queen 
Charlotte Sound. Ultimately, unless extensively har- 
vested in certain localities, Sea Otters can be expect- 
ed to regain all of their original range, which includes 
most, if not all, habitats presently occupied by 
Northern Abalone. Abalone can coexist with Sea 
Otters, but at a relatively low density as cryptic indi- 
viduals (Watson and Smith 1996; Watson 2000). 
Abalone fisheries and unharvested Sea Otters are 
unlikely to coexist (Watson 2000) because humans 
with SCUBA equipment and Sea Otters are both very 
efficient predators and competitors for abalone. 


Protection 

Haliotis kamtschatkana is still afforded no special 
treatment in law, although the provisions of the 
Fisheries Act provide some measure of control to 
legal aboriginal, recreational and commercial 
exploitation. Because of significant reductions in 
regional abundance, legal fisheries for abalone have 
been closed by Fisheries regulation in Canada since 
1990. However, because demand for abalone world- 
wide and particularly in Canada is high, the closure 
simply resulted in a dramatic increase in price per 
pound (estimated to now be about $100 per kg 


2001 


(Jubinville 2000)), with the result that poaching of 
abalone continues to be a serious problem. Campbell 
(1997) suggested that obtaining a COSEWIC classi- 
fication of either “Endangered” (species facing 
imminent extirpation or extinction) or “Threatened” 
(a species likely to become endangered if limiting 
factors are not reversed) would help to emphasise the 
serious poaching problem with this species and help 
to justify the resources required for conservation of 
this species. 


Management Options 

Given the current high value of abalone, other 
higher fishing resource management priorities, and 
the logistic difficulties of restricting abalone poach- 
ing, it seems unlikely that abalone poaching can be 
significantly reduced below the current level unless 
it is made illegal to possess abalone in Canada, 
including imports from other countries. Even this 
action may not stop the illegal export of poached 
abalone, but it would curtail local demand for the 
product. Establishing marine protected areas to fur- 
ther protect abalone is likely to only work if people 
in local communities assist in trying to rebuild local 
abalone stocks by reporting, and hopefully prevent- 
ing, local fishing infractions (Jamieson 2000). With 
fishing now illegal but poaching still occurring, 
reliance on enforcement officers alone to prevent 
abalone poaching can not guarantee the long-term 
protection of specific populations. 

Abalone can be cultured, and a number of groups, 
particularly First Nations, have proposed the estab- 
lishment of abalone hatcheries to enhance local 
abalone populations and thereby allow the reestab- 
lishment of local abalone fisheries. While probably 
technically feasible, the economics may not be 
viable, and it may be more profitable to hold cultured 
abalone in closed pens until they reach a marketable 
size. Even in the absence of Sea Otters, the mortality 
of cultured Northern Abalone released as juveniles 
for grow-out in the wild and later harvest may be 
high, from natural predators when small and from 
poachers when large. It may also be difficult to fully 
separate wild from “cultured-and-released” individu- 
als at the marketplace, meaning that to protect wholly 
“wild” stocks, released animals would not likely be 
harvestable from the wild until they reached the mini- 
mum legal size for wild abalone (100 mm in 1990). 
Continuous holding of cultured abalone in tanks 
might mean that smaller abalone could be marketed, 
but poachers might then increasingly target smaller 
wild abalone if practical techniques to separate wild 
from cultured abalone are not available. Any abalone 
culture is thus likely to open the door to further, or at 
least continued, abalone poaching. 

From a sustainable population perspective, deter- 
mining an historical population size baseline for 
Northern Abalone may be a major COSEWIC issue. 


JAMIESON: STATUS OF THE NORTHERN ABALONE IN CANADA 


56] 


Humans have impacted abalone abundance twice — 
initially by eliminating a major natural predator, 
which is assumed to have increased adult abundance, 
and then by overharvesting, which decreased adult 
abundance and perhaps returned it to more historic 
levels. Over the extreme long term (i.e., centuries), 
Northern Abalone abundance over much of its range 
may have been relatively low (at least compared to 
the 1950s and 1960s) if predation of abalone by Sea 
Otters was high. From a COSEWIC status perspec- 
tive, humans seem to have had a mixed impact on 
Northern Abalone compared to that of other severely 
depleted species. However, we have no scientific 
data on Northern Abalone abundance in the long ago 
past. All we know 1s that while past predation of 
abalone may have been high, recent removals by 
fishers have also been high, and with both these 
combined sources of mortality likely to continue, 
abalone seem unlikely to occur in abundance any- 
where on the coast in the near future. The ranges of 
Sea Otters and Northern Abalone may never have 
totally overlapped on a microhabitat scale, but the 
ranges of fishers and Northern Abalone almost cer- 
tainly do. Sea Otters also prey on many other 
species, and do not target abalone in the same man- 
ner than fishers do. 

Given the criterion of relative change in abun- 
dance used to determine status, the speculated histor- 
ical abundance of abalone may be an issue that needs 
debate. The issue of abalone poaching can be consid- 
ered of immediate ecosystem importance because 
even in the long run poaching removes potential 
food items from Sea Otters as they expand their 
range, thereby affecting the possible rate of Sea 
Otter range expansion. Alternatively, poaching can 
be considered as simply replacing one efficient 
predator (Sea Otters) with another (humans), which 
makes little difference to an abalone that is killed. 


Evaluation 

From a COSEWIC perspective, I suggest that only 
recent decadal changes in abundance be considered 
in assigning status, as who knows what the past real- 
ly was and what the future will bring. For a variety 
of reasons, Sea Otters may never, or at least not in 
the foreseeable future, reoccupy their historical 
range, and so Northern Abalone abundance should 
be considered on the basis of its current potential 
level and how it is being impacted by humans. The 
effects of illegal fishing are an immediate serious 
problem, and all factors influencing abalone abun- 
dance need consideration. 

Given the above, I suggested that Northern 
Abalone be listed as “Threatened”. Northern Abalone 
abundance over most of its range in British Columbia 
has been greatly reduced by fishing, and is probably 
less than 5% of the level that occurred immediately 
prior to the commencement of the industrial fishery. 


562 


There is no evidence that populations are increasing 
in abundance, and population size may still be 
decreasing. Over the past decade, information about 
the Northern Abalone fishery closure has been made 
available to the public, but there may still be oppor- 
tunities to better educate people on the need to pro- 
tect abalone. 


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Received 14 February 2000 
Accepted 11 February 2002 


Rare and Endangered Fishes and Marine Mammals of Canada: 


COSEWIC Fish and Marine Mammal Subcommittee 
Status Reports XIV 


R.R. CAMPBELL 


131 Dahlia Avenue, Ottawa, Ontario K1H 6G1 Canada 


Campbell, R.R. Editor. 2001. Rare and endangered fish and marine mammals of Canada: COSEWIC Fish and Marine 
Mammal Subcommittee Status Reports: XIV. Canadian Field-Naturalist 115(4): 564-572. 


Ten status reports representing the 1999 fish and marine mammal status assignments have been prepared for publication as 
well as two additional reports, one each from 1998 and 2000. Committee (COSEWIC) and Subcommittee (Fish and Marine 
Mammals) activities, including the implications of proposed legislation for the protection of wildlife species at risk in 
Canada, are briefly discussed. 


Dix rapports de statut relatevement aux poissons et aux mammiféres marins auxquelles ont été attribués un statut en 1998 
ont été préparés pour publication; il y a aussi deux rapports additionnels de 1998 et 2000 respectivement. Les activitités du 
Comité (CSEMDC) et du sous-comité (des poissons et des mammifeéres marins), sont briévement discutées, méme que les 


implications de la loi proposée concernant la protection des espéces sauvages en péril au Canada. 


Key Words: Rare and Endangered species, fish, marine mammals, COSEWIC. 


As indicated in previous submissions (Campbell 
1984 through 2001), the intent of the Subcommittee 
on Fish and Marine Mammals has been to publish 
(as funding permits) the status reports (on those 
species of fish and marine mammals) which the 
Committee on the Status of Endangered Wildlife in 
Canada (COSEWIC) has reviewed, approved and 
used as a basis of assigning status to species in jeop- 
ardy in Canada. The group of reports presented here- 
in represent the seven fishes and the three marine 
mammals considered by COSEWIC and assigned 
status in 1999 (see Tablel; COSEWIC 1999). An 
additional report on the Redfin Pickerel (Esox ameri- 
canus americanus) is included. This report is from 
the 1998 status assignments, but not included in the 
publication of that set (see Campbell 2001). This 
series also contains one update from the group con- 
sidered in 2000 at the request of the authors. 
Summaries of these (and any and all other status 
reports) are available from the COSEWIC Secre- 
tariat (Canadian Wildlife Service, Environment 
Canada, Ottawa, Ontario K1A 0H3). 


Progress 

COSEWIC has undertaken to make available to 
all Canadians supporting information on each 
species classified (see Cook and Muir 1984). The 
Fish and Marine Mammal Subcommittee has been 
able to use this journal as one step in achieving the 
goal. A series of reports have appeared in various 
volumes and numbers from 1984 through 1998 [see 
Canadian Field-Naturalist 98(1): 63-133; 99(3): 
404-450; 101(2): 165-309; 102(1): 81-176 and 
102(2): 270-398; 103(2): 147-220; 104(1): 1-145; 


105(2): 151-250; 106(1): 1-72; 110(3): 462-532; 
111(2): 249-307; 112(1): 94-157; 115(1): 115-167]. 

As of April 1999, COSEWIC has reviewed the 
status of 109 fish species and 61 marine mammals 
(see Tablel; COSEWIC 1999). Of the 170 species 
(or discrete populations) investigated 11 are indeter- 
minate (seven fish, four marine mammals), 65 (30 
fish, 35 marine mammals) have been found not to 
require status designation and another 50 (42 fish, 
eight marine mammals) have been designated as 
vulnerable leaving 33 species (22 fish, 11marine 
mammals) of immediate concern (threatened and 
endangered), and 11 species (eight fish, three marine 
mammals) extinct or extirpated. 

As of April 1999 there are 27 status reports on 
fish species (includes 15 updates), and 6 on marine 
mammal species (three updates) under review or in 
preparation (Table 2). Several of these were present- 
ed to the Committee for status assignment in 2000 
and 2001. 

As well, some 73 additional species of fish (plus 
23 to be updated) and one marine mammal (plus 
four to be updated) are on the list for possible 
future consideration (Table 3). A few may be found 
to not require status designation, but the process 
serves to bring together the information necessary 
to make the appropriate determination and satisfy 
the need to fill those knowledge gaps. Although 
some of these may be of no immediate concern, the 
Subcommittee will, as opportunity allows, attempt 
to document the status of these species to determine 
their status in Canada. 

In addition to soliciting further status reports on 
species of concern, the Subcommittee continues to 


564 


2001 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS XIV 


Table 1. Fish and Marine Mammal Species with Assigned COSEWIC Status to April 1999. 


Species 


FISH 

Lake Sturgeon 

Bloater 

Blueback Herring 
Cutlips Minnow 
Eastern Silvery Minnow 
Striped Shiner 

Redfin Shiner 
Hornyhead Chub 

River Chub 

Ghost Shiner 

Blackchin Shiner 

Weed Shiner 

Bluntnose Minnow 
Leopard Dace 

Central Stoneroller 
Mountain Sucker 
Golden Redhorse 
Redfin Pickerel 

Chain Pickerel 

Least Darter 

Tesselated Darter 

River Darter 

Green Sunfish 

Longear Sunfish 
Spoonhead Sculpin 
Brook Silverside 
Y-Prickleback 

Darktail Lamprey 
Bering Cisco 

Mira Whitefish 
Chiselmouth 

Flathead Catfish 
Spinynose Sculpin 

Pixy Poacher 

Lake Lamprey 

Chestnut Lamprey 
Northern Brook Lamprey 
Shortnose Sturgeon 
Green Sturgeon 

White Sturgeon 

Spotted Gar 

Spring Cisco 

Squanga Whitefish 

Kiyi 

Pacific Sardine 

Atlantic Cod 

Redside Dace 

Western Silvery Minnow 
Silver Chub 

Pugnose Shiner 

Bridle Shiner 

Bigmouth Shiner 

Silver Shiner 

Roseyface Shiner (Manitoba) 
Pugnose Minnow 
Speckled Dace 
Umatilla Dace 

Banded Killifish (Newfoundland) 
Blackstripe Topminnow 


Scientific Name 


Acipenser fulvescens 
Coregonus hoyi 

Alosa aestivalis 
Exoglossum maxillingua 
Hybognathus nuchalis regius 
Luxilis chyrsocephalus 
Lythrurus umbratilis 
Nocomis biguttatus 
Nocomis micropogon 
Notropis buchanani 
Notropis heterodon 
Notropis texanus 
Pimephales notatus 
Rhinichthys falcatus 
Campostoma anomalum 
Catostomus platyrhynchus 
Moxostoma erythrurum 
Esox americanus americanus 
Esox niger 

Etheostoma microperca 
Etheostoma olmstedi 
Percina shumardi 
Lepomis cyanellus 
Lepomis megalotis 
Cottus ricei 

Labidesthes sicculus 
Allolumpenus hypochromus 
Lethenteron alaskense 
Coregonus laurettae 
Coregonus sp. 
Acrocheilus alutaceus 
Pylodictis olivaris 
Asemichthys taylori 
Ocella impi 

Lampetra macrostoma 
Ichthyomyzon castaneus 
Ichthyomyzon fossor 
Acipenser brevirostrum 
Acipenser medirostris 
Acipenser transmontanus 
Lepisosteus oculatus 
Coregonus sp. 
Coregonus sp. 
Coregonus kiyi 
Sardinops sagax 

Gadus morhua 
Clinostomus elongatus 
Hybognathus argyritis 
Macrhybopsis storeriana 
Notropis anogenus 
Notropis bifrenatus 
Notropis dorsalis 
Notropis photogenis 
Notropis rubellus 
Opsopoeodus emiliae 
Rhinichthys osculus 
Rhinichthys umatilla 
Fundulus diaphanus 
Fundulus notatus 


Status 


Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Indeterminate 
Indeterminate 
Indeterminate 
Indeterminate 
Indeterminate 
Indeterminate 
Indeterminate 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 


Date Assigned 


April 1986 
April 1988 
April 1980 
April 1994 
April 1997 
April 1993 
April 1988 
April 1988 
April 1988 
April 1993 
April 1994 
April 1999 
April 1998 
April 1990 
April 1998 
April 199] 
April 1989 
April 1998 
April 1997 
April 1989 
April 1993 
April 1989 
April 1987 
April 1987 
April 1989 
April 1989 
April 1991 
April 1990 
April 1990 
April 1999 
April 1997 
April 1993 
April 1997 
April 1991 
April 1998 
April 1991 
April 1991 
April 1980 
April 1987 
April 1990 
April 1994 
April 1992 
April 1988 
April 1987 
April 1987 
April 1998 
April 1987 
April 1997 
April 1985 
April 1985 
April 1999 
April 1985 
April 1987 
April 1994 
April 1985 
April 1980f 
April 1988 
April 1989 
April 1985 


continued 


566 THE CANADIAN FIELD-NATURALIST 


Table 1. continued 


Species 


Lake Chubsucker 

Bigmouth Buffalo 

Black Buffalo 

Spotted Sucker 

River Redhorse 

Greenside Darter 

Brindled Madtom 

Northern Madtom 

Redbreast Sunfish 

Warmouth 

Orangespotted Sunfish 

Cultus Pygmy Sculpin 

Fourhorn Sculpin (Arctic Islands) 
Giant Stickleback 

Unarmoured Stickleback 
Blackline Prickleback 

Bering Wolffish 

Morrison Creek Lamprey 

Lake Simcoe Whitefish 

Blackfin Cisco 

Shortnose Cisco 

Shortjaw Cisco 

Lake Utopia Dwarf Smelt 

Black Redhorse 

Copper Redhorse 

Eastern Sand Darter 

Channel Darter 

Enos Lake Sticklebacks (Species Pair) 
Paxton Lake Sticklebacks (Species Pair) 
Vananda Creek Sticklebacks (Species Pair) 
Margined Madtom 

Shorthead Sculpin 

Deepwater Sculpin (Great Lakes) 
Atlantic Whitefish 

Aurora Trout 

Nooksack Dace 

Salish Sucker 

Paddlefish 

Gravel Chub 

Longjaw Cisco 

Deepwater Cisco 

Banff Longnose Dace 

Blue Walleye 

Hadley Lake Sticklebacks (Species Pair) 


MARINE MAMMALS 
Sea Otter 
Sea Mink 
Northern Fur Seal 
Hooded Seal 
Bearded Seal 
Grey Seal 
Northern Elephant Seal 
Harbour Seal (Lac des Loups Marins) 
Harbour Seal (Atlantic & Arctic) 
Harbour Seal (Pacific) 
Steller Sea Lion 
California Sea Lion 
Atlantic Walrus 
— Eastern Arctic 
— Northwest Atlantic 


Scientific Name 


Erimyzon sucetta 

Ictiobus cyprinellus 
Ictiobus niger 

Minytrema melanops 
Moxostoma carinatum 
Etheostoma blennioides 
Notorus miurus 

Noturus stigmosus 
Lepomis auritus 

Lepomis gulosus 

Lepomis humilis 

Cottus sp. 

Myoxocephalus quadricornis 
Gasterosteus sp. 
Gasterosteus sp. 
Acantholumpenus mackayi 
Anahichas orientalis 
Lampetra richardsoni marifuga 
Coregonus clupeaformis ssp. 
Coregonus nigripinnis 
Coregonus reighardi 
Coregonus zenithicus 
Osmerus sp. 

Moxostoma dusquesnei 
Moxostoma hubbsi 
Ammocrypta pellucida 
Percina copelandi 
Gasterosteus spp. 
Gasterosteus spp. 
Gasterosteus spp. 

Noturus insignis 

Cottus confusus 
Myoxocephalus thompsoni 
Coregonus huntsmani 


Salvelinus fontinalis timagamiensis 


Rhinichthys cataractae ssp. 
Catostomus sp. 

Polyodon spathula 

Erimystax x-punctata 
Coregonus alpenae 
Coregonus johannae 
Rhinichthys cataractae smithi 
Stizostedion vitreum glaucum 
Gasterosteus sp. 


Enhydra lutris 

Mustela macrodon 
Callorhinus ursinus 
Cystophora cristata 
Erignathus barbatus 
Halichoerus grypus 
Mirounga angustirostris 
Phoca vitulina mellonae 
Phoca vitulina richardsi 
Phoca vitulina concolor 
Eumetopias jubatus 
Zalophus californianus 
Odobenus rosmarus rosmarus 


Status. 


Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Vulnerable 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Threatened 
Endangered 
Endangered 
Endangered 
Endangered 
Extirpated 
Extirpated 
Extinct 
Extinct 
Extinct 
Extinct 
Extinct 


Threatened 
Extinct 

Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Not At Risk 
Vulnerable 
Not At Risk 


Indeterminate - 


Not At Risk 
Not At Risk 


Not At Risk 


Extirpated 


Vols 


Date Assigned 


April 1994 
April 1989 
April 1989 
April 1994 
April 1987 
April 1990 
April 1985 
April 1998 
April 1989 
April 1994 
April 1989 
April 1997 
April 1989 
April 1980 
April 1983 
April 1989 
April 1989 
April 1999 
April 1987 
April 1988 
April 1987 
April 1987 
April 1998 
April 1988 
April 1987 
April 1994 
April 1993 
April 1988 
April 1999 
April 1999 
April 1989 


November 1983 


April 1987 
April 1983 
April 1987 
April 1996 
April 1986 
April 1987 
April 1987 
April 1988 
April 1988 
April 1987 
April 1985 
April 1999 


April 1996 
April 1985 
April 1996 
April 1986 
April 1994 
April 1999 
April 1986 
April 1996 
April 1999 
April 1999 
April 1987 
April 1987 


April 1987 
April 1987 


continued 


2001 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS XIV 


Table 1. Concluded 


Species Scientific Name Status Date Assigned 
Baird’s Beaked Whale Berardius bairdi Not At Risk April 1992 
Beluga Delphinapterus leucas 

— Beaufort Sea Not At Risk April 1986 

— Western and Southern Hudson Bay Not At Risk April 1993 

— High Arctic Vulnerable April 1992 

— Eastern Hudson Bay Threatened April 1988 

— St. Lawrence River Endangered April 1997 

— S.E. Baffin Island Endangered April 1990 

— Ungava Bay Endangered April 1988 
Striped Dolphin Stenella coeruleoalba Not At Risk April 1993 
Ringed Seal Phoca hispida Not At Risk April 1989 
Common Dolphin Delphinus delphis Not At Risk April 1991] 
Grey Whale Eschrichtius robustus 

— Northeast Pacific Not At Risk April 1987 

— Northwest Atlantic Extirpated April 1987 
Risso’s Dolphin Grampus griseus Not At Risk April 1990 
Short-finned Pilot Whale Globicephala macrohynchus Not At Risk April 1993 
Long-finned Pilot Whale Globicephela malaena Not At Risk April 1994 
Northern Bottlenose Whale Hyperoodon ampullatus 

— Northwest Atlantic Not At Risk April 1993 

— Gully Population Vulnerable April 1996 
Pygmy Sperm Whale Kogia breviceps Not At Risk April 1994 
Dwarf Sperm Whale Kogia simus Indeterminate April 1997 
Atlantic White-sided Dolphin Lagenorhynchus acutus Not At Risk April 1991 
White-beaked Dolphin Lagenorhynchus albirostris Not At Risk April 1998 
Pacific White-sided Dolphin Lagenorhynchus obliquidens Not At Risk April 1990 
Northern Right Whale Dolphin Lissodelphis borealis Not At Risk April 1990 
Hubbs’ Beaked Whale Mesoplodon carlhubbsi Not At Risk April 1989 
Blainville’s Beaked Whale Mesoplodon densirostris Not At Risk April 1989 
True’s Beaked Whale Mesoplodon mirus Not At Risk April 1989 
Stejneger’s Beaked Whale Mesoplodon stejnegeri Not At Risk April 1989 
Narwhal Monodon monoceros Not At Risk April 1987 
Dall’s Porpoise Phocoenoides dalli Not At Risk April 1989 
False Killer Whale Pseudorca crassidens Not At Risk April 1990 
Killer Whale Orcinus orca 

— North Atlantic and Arctic Indeterminate April 1999 

— North Pacific Transients Vulnerable April 1999 

— North Pacific Residents Threatened April 1999 
Sperm Whale Physeter macrocephalus Not At Risk April 1996 
Bottlenose Dolphin Tursiops truncatus Not At Risk April 1993 
Cuvier’s Beaked Whale Ziphius cavirostris Not At Risk April 1990 
Blue Whale Balaenoptera musculus Vulnerable April 1983 
Fin Whale Balaenoptera physalus Vulnerable April 1987 
Sowerby’s Beaked Whale Mesoplodon bidens Vulnerable April 1989 
Harbour Porpoise Phocoena phonoeca 

— Northwest Pacific Indeterminate April 1991 

— Northwest Atlantic Threatened April 1991 
Humpback Whale Megaptera novaeangliae 

— Northwest Atlantic Vulnerable April 1985 

— Northeast Pacific Threatened April 1985 
Bowhead Whale Balaena mysticetus Endangered April 1986 
Right Whale Eubalaena glacialis Endangered April 1990 


obtain updates on the status of selected species as Future Direction 

new information becomes available, or during the The introduction to Series XI referred to proposed 
10-year review process initiated in 1993 (Table 2) federal legislation in regards to a Species At Risk 
for those species which had not already received fur- Act (Campbell 1997), but the Bill was not passed 
ther examination following the initial assignment of _ prior to the dissolution of Parliament in June 1997 
status. and thus became non-existent (Campbell 1998). 


568 THE CANADIAN FIELD-NATURALIST Vol. 115 


However, the federal, provincial and territorial Min- 
isters responsible for wildlife in Canada, recognizing 
that species are not respective of arbitrary jurisdic- 
tional boundaries, and that cooperation is crucial to 
the conservation of species at risk, reached a 
National Accord for the Protection of Species at Risk 
in June of 1996. The Ministers agreed to form a 
national council [Canadian Endangered Species 
Conservation Council (CESCC)] to coordinate their 
activities (in relation to the conservation of species at 
risk in Canada); to recognize COSEWIC as a source 
of independent advice on the status of species at risk; 
and to establish complementary legislation and pro- 


grams to provide for the conservation and protection 
of species at risk throughout Canada. Such legisla- 
tion and programs will, among other things: provide 
legal designation and protection for threatened and 
endangered species and their habitat; provide for the 
development and implementation of recovery plans; 
ensure multi-jurisdictional cooperation for the con- 
servation of species that cross borders; ensure that 
species at risk are a part of the environmental assess- 
ment processes; monitor, assess and report on the 
status of wild species; and provide effective enforce- 
ment. 

The Ministers went on to develop “A National 


TABLE 2. Fish and Marine Mammal Species for which Status Reports are in preparation, or under review — to April 1999. 


Species 


FISH 

Updated Reports 
Chestnut Lamprey 
Lake Sturgeon 

Aurora Trout 

Shortjaw Cisco 

Silver Shiner 

Silver Chub 

Pugnose Shiner 
Bigmouth Shiner 
Pugnose Minnow 
Redside Dace 
Speckled Dace 
Blackstripe Topminnow 
Brindled Madtom 
Enos Lake Stickleback 
Pacific Sardine 


Scientific Name 


Ichthyomyzon castaneus 
Acipenser fulvescens 
Salvelinus fontinalis timagamiensis 
Coregonus zenithicus 
Notropis photogenis 
Macrhybopsis storeriana 
Notropis anogenus 
Notropis dorsalis 
Opsopoeodus emiliae 
Clinostomus elongatus 
Rhinichthys osculus 
Fundulus notatus 
Noturus miurus 
Gasterosteus sp. 
Sardinops sagax 


New Reports (Species Not Previously Considered) 


Atlantic Sturgeon 
Arctic Char 
Round Whitefish 
Lake Herring 


Lake Whitefish 


Grass Pickerel 

Pearl Dace 

Jasper Longnose Sucker 
Greater Redhorse 
Mottled Sculpin 
Shorthead Sculpin 
Bluefin Tuna 


MARINE MAMMALS 
Updated Reports 
Bowhead Whale 
Blue Whale 
Humpback Whale 
— Northwest Atlantic 
— Northeast Pacific 


Acipenser oxyrhynchus 
Salvelinus alpinus 
Prosopium cylindraceum 
Coregonus artedi 


Coregonus clupeaformis 


Esox americanus vermiculatus 
Margariscus margarita 
Castostomus castostomus lacustris 
Moxostoma valenciennesi 

Cottus bairdi 

Cottus confusus 

Thunnus thynnus 


Balaena mysticetus 
Balaenoptera musculus 
Megaptera novaeangliae 


New Reports (Species Not Previously Considered) 


Minke Whale 
Sei Whale 
Harbour Seal 


Balaenoptera acutorostrata 
Balaenoptera borealis 
Phoca vitulina 


Proposed Status 


Current Status 

Vulnerable 199] 
Not At Risk 1986 
Endangered 1987 
Threatened 1987 


Vulnerable 1987 
Vulnerable 1985 
Vulnerable 1985 
Vulnerable 1985 
Vulnerable 1985 
Vulnerable 1987 
Vulnerable 1980 
Vulnerable 1985 
Vulnerable 1985 
Threatened 1988 
Vulnerable 1987 
Proposed Status 
Vulnerable 

9 
Vulnerable 
Great Lakes and BC population — 
may be Threatened 


Populations of lakes Erie and 
Ontario may be Threatened 
Vulnerable 

Vulnerable — BC, NT, NS 
Vulnerable 

Vulnerable 

Vulnerable — BC, Alberta 


Threatened 
? 


Current Status 
Endangered 1986 


Vulnerable 1983 
Vulnerable 1985 
Threatened 1985 
Proposed Status 


? 
? 
Lake Ontario — Extirpated 


2001 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS XIV 


TABLE 3. Fish and Marine Mammal Species of Possible Interest to COSEWIC — April 1999 (Not listed by Priority) 


Species 


SPECIES UPDATES 
Fish 

Salish Sucker 

Lake Simcoe Whitefish 
Blackfin Cisco 

Black Redhorse 
Margined Madtom 
Great Lakes Deepwater Sculpin 
Shorthead Sculpin 
Shortnose Sturgeon 
Green Sturgeon 
Squanga Whitefish 
Kiyi 

Umatilla Dace 
Bigmouth Buffalo 
Black Buffalo 

River Redhorse 
Redbreast Sunfish 
Orangespotted Sunfish 
Banded Killifish 
Fourhorn Sculpin (Arctic Islands) 
Charlotte Stickleback 
Giant Stickleback 
Blackline Prickleback 
Bering Wolffish 


Marine Mammals 
Beluga — Ungava Bay 

— Eastern Hudson Bay 
Sowerby's Beaked Whale 
Fin Whale 


SPECIES YET TO BE CONSIDERED 


Fish 

Nass River Lamprey Species | 
Nass River Lamprey Species 2 
Copper River Lamprey 
American Eel 
American Shad 
Atlantic Herring 
Pacific Herring 

Creek Chubsucker 
Brassy Minnow 

Pygmy Smelt 

Emerald Shiner 
Spottail Shiner 

Creek Chubsucker 
Rainbow Darter 
Stonecat 

Capelin 

Broad Whitefish 

Least Cisco 

Giant Pygmy Whitefish 
Pygmy Whitefish 
Ninespine Stickleback - 
Pacific Salmonids 
Atlantic Salmon 
Euchalon 

Spiny Dogfish 
Barndoor Skate 

Thorny Skate 


Scientific Name 


Catostomus sp. 
Coregonus clupeaformis 
Coregonus nigripinnis 
Moxostoma dusquesnei 
Noturus insignis 
Myoxocephalus thompsoni 
Coitus confusus 
Acipenser brevirostrum 
Acipenser medirostris 
Coregonus sp. 
Coregonus kiyi 
Rhinichthys umatilla 
Ictiobus cyprinellus 
Ictiobus niger 
Moxostoma carinatum 
Lepomis auritus 
Lepomis humilis 
Fundulus diaphanus 


Myoxocephalus quadricornis 


Gasterosteus sp. 
Gasterosteus sp. 
Acantholumpenus mackayi 
Anarichus orientalis 


Delphinapterus leucas 


Mesoplodon bidens 
Balaenoptera physalus 


Lampetra sp. 
Lampetra sp. 
Lampetra sp. 

Anquilla rostrata 
Alosa sapidissima 
Clupea harengus 
Clupea pallasi 
Erimyzon oblongus 
Hybognathus hankinsoni 
Osmerus spectrum 
Notropis atherinoides 
Notropis hudsonius 
Erimyzon oblongus 
Etheostoma caeruleum 
Noturus flavus 
Mallotus villosus 
Coregonus nasus 
Coregonus sardinella 
Prosopium sp. 
Prosopium coulteri 
Pungitius pungitius 
Oncorhynchus spp. (5) 
Salmo salar 
Thaleichthys pacificus 
Squalus acanthias 
Raja laevis 

Raja radiata 


569 

Possible Status 

Current Status 

Endangered 1986 
Threatened 1987 
Threatened 1988 
Threatened 1988 
Threatened 1989 
Threatened 1987 
Threatened 1984 
Vulnerable 1980 
Vulnerable 1987 
Vulnerable 1987 
Vulnerable 1988 
Vulnerable 1988 
Vulnerable 1989 
Vulnerable 1989 
Vulnerable 1987 
Vulnerable 1989 
Vulnerable 1989 
Vulnerable 1989 
Vulnerable 1989 
Vulnerable 1983 
Vulnerable 1980 
Vulnerable 1989 
Vulnerable 1989 
Endangered 1988 
Threatened 1988 
Vulnerable 1989 
Vulnerable 1987 


Possible Status 
BC 

EBC? 

BC ? 

Vulnerable 

? 

? 

A 

Extirpated — NB 
Vulnerable 
Vulnerable — QC 
Vulnerable — BC 
Vulnerable — BC 
Threatened — ON 
Vulnerable — QC 
Vulnerable — SK, QC 
5 

Vulnerable — BC 
Vulnerable 
Threatened — BC 
Threatened — AB 
Vulnerable — BC 
: 

9 

9 

9 

Threatened 

= 


continued 


570 


TABLE 3. continued 
Species 


Smooth Skate 


THE CANADIAN FIELD-NATURALIST 


Scientific Name 


Malacoraja senta 


Monkfish Lophius americanus 
Tomcod Microgadus tomcod 
Haddock Melanogrammus aeglefinus 
Silver Hake Merluccius bilinearis 
Pollock Pollachius virens 

Red Hake Urophycis chuss 


Rock Grenadier 
Pacific Ocean Perch 


Coryphaenoides rupestris 
Sebates alutus 


Aurora Rockfish Sebastes aurora 
Redbanded Rockfish Sebastes babcocki 
Silvergray Rockfish Sebastes brevispinis 
Copper Rockfish Sebastes caurinus 
Splitnose Rockfish Sebastes diploproa 
Widow Rockfish Sebastes entomelas 
Chilipepper Sebastes goodei 
Shortbelly Rockfish Sebastes jordani 
Quillback Rockfish Sebastes maliger 
Black Rockfish Sebastes melanops 
Vermillion Rockfish Sebastes miniatus 
Blue Rockfish Sebastes mystinus 
China Rockfish Sebastes nebulosus 
Golden Rockfish Sebates norvegicus 
Bocaccio Sebates paucispinis 
Canary Rockfish Sebates pinniger 
Yelloweye Rockfish Sebates ruberrimus 
Sripetail Rockfish Sebates saxicola 
Shortspine Thornyhead Sebastolobus alascanus 
Longspine Thornyhead Sebastolobus altivelis 
Lingcod Ophiodon elongatus 
Lumpfish Cyclopterus lumpus 
Wolffish Anarhichas lupus 
Atlantic Mackerel Scomber scombrus 
Swordfish Xiphias gladius 
Summer Flounder Paralichthys dentatus 
Witch Flounder 


American Plaice 
Atlantic Halibut 
Pacific Halibut 
Yellowtail Flounder 
Winter Flounder 


Greenland Halibut (Turbot) 


Marine Mammals 
Harp Seal 


Glyptocephalus cynoglossus 
Hippoglossoides platessoides 
Hippoglossus hippoglossus 
Hippoglossus steolepis 
Limanda ferrugineus 
Pseudopleuronectes americanus 
Reinhardtius hippoglossoides 


Phoca groenlandica 


Vol. 115 


Possible Status 

? 

2 

Vulnerable — Quebec 


ulnerable 


Not At Risk 


Framework For The Conservation Of Species At 
Risk” to provide a coordinated national approach for 
the conservation of species at risk in Canada. 
CESCC is to be responsible for the National 
Framework , its implementation and the resolution of 
issues for the protection of species at risk in Canada. 
CESCC recognized COSEWIC as an independent 
committee of scientific experts mandated to assess 
the status of species which may be at risk nationally 
and provided terms of reference for COSEWIC. The 
organization will also provide the lead for the 
“Recovery of Species at Risk Nationally” (RENEW) 
organization which develops and implements recov- 


ery plans for those species assessed by COSEWIC as 
threatened and endangered. The National Framework 
also includes provisions for the development of 
authority over wild species in each of the federal, 
provincial, and territorial jurisdictions which do not 
currently have such legislation. Endangered species 
legislation is under consideration by the federal par- 
liament as well as by those provinces and territories 
which do not have such legislation. 

Under the Accord and the National Framework, 
COSEWIC now reports to CESCC and is supported 
by a secretariat funded by Environment Canada. The 
secretariat is provided with a budget to facilitate 


2001 


meetings, fund production and publication of status 
reports and communications. The basic structure and 
operation of COSEWIC remains much as it was (see 
Cook and Muir 1984) with some exceptions. The 
major changes are that members are now appointed 
by the (Federal) Minister of the Environment, as rec- 
ommended by CESCC and the creation of a new 
subcommittee structure which will include the addi- 
tion of an Aboriginal Traditional Knowledge 
Specialist Group. 

There will now be eight subcommittees which are 
referred to as “Species Specialist Groups” (SSGs). 
An invertebrates subcommittee had been recognized 
in 1995 and that is why marine invertebrates no 
longer (since 1996) appeared as part of the fish and 
marine mammals reports. To-date the invertebrates 
SSG has only been mandated to examine lepidoptera 
and molluscs. The fish and marine mammal subcom- 
mittee is to be split into three SSGs, freshwater fish- 
es, marine fishes and marine mammals. The four 
remaining specialists groups are the existing groups 
representing birds, “terrestrial” (1.e, non-marine) 
mammals, plants, and amphibians and reptiles. 

COSEWIC is also in the process of adopting 
quantitative scientific assessment criteria based on 
the IUCN Red List Criteria (UCN 1994). This will 
permit a process based on science and traditional or 
local knowledge to assess species at risk according 
to the probable risk of extinction. Future status 
assignments will be made using these criteria. 


Concluding Remarks 

The 12 reports included in the following series are 
based on reports on the status of the respective 
species in Canada. Status was assigned by consensus 
of the COSEWIC Committee based on the originals. 
Here they are published under the name(s) of the 
original author(s). The present reports have under- 
gone minor editing to provide some degree of con- 
sistency in format and presentation. 

As indicated above, COSEWIC is in a state of 
transition and has undergone some change in its 
structure and operation to pave the way for its taking 
on a legal basis under federal endangered species 
legislation. COSEWIC will maintain its indepen- 
dence and continue to provide assessments of status 
of species considered to be at risk in Canada based 
on the best scientific, traditional and local knowl- 
edge available. It is still mandated to make that 
information available to the public; however, at this 
time the possible continuation of this series in some 
form is uncertain. The publication of all reports, 
from all SSGs, in some format is under considera- 
tion. However, copies of all original reports are 
available from the COSEWIC Secretariat (c/o 
Canadian Wildlife Service, Environment Canada, 
Ottawa, Ontario K1A 0H3) or the COSEWIC web 
site at http://www.cosewic.gc.ca. 


CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS XIV 


57] 


The author has remained as co-chair of the fresh- 
water fishes group and, if circumstances permit, 
would like to continue presenting the reports from 
that group and to provide updated information on 
COSEWIC and its operations and the progress of 
endangered species legisiation in this forum. 


Acknowledgments 

The members of COSEWIC and the Fish and 
Marine Mammal Subcommittee would like to extend 
their thanks to the various authors who have so gen- 
erously contributed their time and talents in support 
of COSEWIC. The Committee also wishes to 
acknowledge the members of the Subcommittee for 
their unstinting efforts in reviewing the reports and 
their helpful comments. 

COSEWIC is grateful to the Canadian Wildlife 
Federation, World Wildlife Fund Canada, the Can- 
adian Wildlife Service, the Canadian Museum of 
Nature and the Royal Ontario Museum for assistance 
provided in cash and kind. A special thanks are due 
the Canadian Field-Naturalist for assistance in publi- 
cation and editing and to all members of COSEWIC 
for their dedication and interest in the future of 
Canada’s flora and fauna. We gratefully acknowl- 
edge the financial support provided by Environment 
Canada which permitted the contracting of several 
new reports and publication of this series. 


Literature Cited 

Campbell, R.R. 1984. Rare and endangered fish of Can- 
ada: The Committee on the Status of Endangered 
Wildlife in Canada (COSEWIC) Fish and Marine 
Mammal Subcommittee. Canadian Field-Naturalist 
98(1): 71-74. 

Campbell, R. R. 1985. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammals Subcommittee Status Reports: II. 
Canadian Field-Naturalist 99(3): 404408. 

Campbell, R. R. 1987. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: III. 
Canadian Field-Naturalist 101(2): 165-170. 

Campbell, R.R. 1988. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: IV. 
Canadian Field-Naturalist 102(1): 81-86. 

Campbell, R.R. 1989. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: V. 
Canadian Field-Naturalist 103(2): 147-157. 

Campbell, R.R. 1990. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: VI. Can- 
adian Field-Naturalist 104(1): 1-6. 

Campbell, R.R. 1991. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: VII. 
Canadian Field-Naturalist 105(2): 151-156. 

Campbell, R.R. 1992. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: VIII. 
Canadian Field-Naturalist 106(1): 1-6. 


572 


Campbell, R.R. 1993. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: IX. 
Canadian Field-Naturalist 107(4): 395-401. 

Campbell, R.R. 1996. Rare and endangered fish and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: X. 
Canadian Field-Naturalist 110(3): 454-461. 

Campbell, R. R. Editor. 1997. Rare and endangered fish 
and marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: XI. 
Canadian Field-Naturalist 111(2): 249-257. 

Campbell, R.R. Editor. 1998. Rare and endangered fish 
and marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: XII. 
Canadian Field-Naturalist 112(1): 94-97. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Campbell, R.R. Editor. 2001. Rare and endangered fish 
and marine mammals in Canada. COSEWIC Fish and 
Marine Mammals Subcommittee Status Reports: XIII. 
Canadian Field-Naturalist 115(1): 115-117. 

Cook, F.R., and D. Muir. 1984. The Committee on the 
Status of Endangered Wildlife in Canada (COSEWIC): 
History and progress. Canadian Field-Naturalist 98(1): 
63-70. 

COSEWIC. 1999. Canadian species at risk: April 1999. 
COSEWIC Secretariat, C/o Canadian Wildlife Service, 
Environment Canada, Ottawa Ontario K1A 0H3. 

IUCN. 1994. IUCN Red List Categories. Prepared by the 
IUCN Species Survivial Commission, IUCN, Gland, 
Switzerland. 


Accepted 19 February 2002 


Status of the Morrison Creek Western Brook Lamprey, 
Lampetra richardsoni, in Canada* 


R. J. BEAMISH, J. H. YOUSON!, and L. A. CHAPMAN 


Department of Fisheries and Oceans, Pacific Biological Station, 3190 Hammond Bay Road, Nanaimo, British Columbia 
VOR 5K6 Canada 
'Department of Zoology, Scarborough Campus, University of Toronto, West Hill, Ontario MIC 1A4 Canada 


Beamish, R. J., J. H. Youson, and L. A. Chapman. 2001. Status of the Morrison Creek Western Brook Lamprey, Lampetra 
richardsoni, in Canada. Canadian Field-Naturalist 115{4): 573-578. 


The Morrison Creek Western Brook Lamprey, is a distinct and rare population of the widely distributed Western Brook 
Lamprey, Lampetra richardsoni, and it is endemic to a small creek on Vancouver Island, Canada. This special population 
produces both non-parasitic and potentially parasitic adult life history types, representing a unique type of polymorphism in 
jampreys. The Morrison Creek population demonstrates that a linkage between parasitic and non-parasitic lampreys exists 
and provides an opportunity to study lamprey speciation and evolution. However, this rare population is at risk because its 
habitat is currently facing substantial development pressures. The Morrison Creek Lampetra richardsoni population 
should be protected because of its important implications for our understanding of lamprey taxonomy and evolution, and 
because the potentially parasitic variety needs further study as it represents a life history that is difficult to classify with our 
existing knowledge. 


La lamproie du ruisseau Morrison, Lampetra richardsoni, est endémique dans un petit cours d’eau de l’ile de Vancouver, 
Canada; elle représente une forme distincte et rare de la lamproic de l’ouest, Lampetra richardsoni, qui, elle, jouit d’une 
grande distribution. Le ruisseau abrite les deux formes de L. richardsoni et celles-ci forment une population unique de lam- 
proies qui produit des types biologiques adultes a la fois parasitiques et non parasitiques. Cette population polymorphe est 
un phénoméne unique en son genre chez la lamproie dont l’ascendance remonte a au moms 350 millions d’années. Ce qui 
se passe dans le ruisseau Morrison revét donc une grande pertinence pour ce qui est de la compréhension de |’ évolution des 
premiers vertébrés. Toutefois, cette population rarissime est en danger car son habitat subit actuellement d’ importantes 
pressions d’aménagement. I] est impératif que nous préservions le complexe des deux formes de la lamproie du ruisseau 
Morrison puisqu’il présente des implications importantes pour la compréhension de la taxonomie et de |’évolution de la 
lamproie et que la variété est un cas qu’il est difficile de classer dans |’ état actuel des connaissances. 


Key Words: Morrison Creek Western Brook Lamprey, lamproie du ruisseau Morrison, Lampetra richardsoni, polymorphic 
lampreys, parasitic lampreys, British Columbia, rare and endangered species, Petromyzontiformes. 


Lampreys (Petromyzonidae) are a successful 
group of vertebrates that have survived for close to 
350 million years and have had a conservative evolu- 
tion (Forey and Janvier 1994). The reason for their 
evolutionary success is not known but may be 
attributed to their ability to change among the three 
adult life history types; anadromous and parasitic, 
nonanadromous and parasitic, and, nonanadromous 
and non-parasitic (Beamish 1987a). Direct evidence 
for this possibility comes from a rare population of 
Lampetra richardsoni that is presently known only 
from Vancouver Island, Canada. This population 
produces a potentially parasitic and a non-parasitic 
adult life history type each year, both of which are 
nonanadromous. The parasitic form is an unde- 
scribed variety of Lampetra richardsoni which we 
are tentatively calling the Morrison Creek variety. It 
should be noted that the variety has not been given 


*Reviewed and approved by COSEWIC April 1999, status 
assigned — Threatened 


official taxonomic status, despite its distinctive life 
history and morphology relative to Lampetra 
richardsoni. The variety is parasitic in the laboratory 
and the taxonomy of lampreys uses adult life history 
type as a species specific character (Zanandrea 1959; 
Vladykov and Kott 1979; Potter 1980). Therefore, in 
a sense, this population produces two species accord- 
ing to conventional lamprey taxonomy. However, 
although the Morrison Creek variety could be con- 
sidered a new species on the basis of morphology 
and life history type, the genetic similarity which 
exists between the two forms indicates that the vari- 
ety is probably not a species but rather a unique 
morph of a single population (Beamish and Withler 
1986). 

It is thought that the Morrison Creek variety rep- 
resents an intermediate step in the evolution of 
Lampetra richardsoni from an anadromous parasitic 
ancestor (Beamish 1985; Beamish and Withler 1986; 
Youson and Beamish 1991). The parasitic morph 
offers a rare opportunity to study the product of an 
evolutionary transition and to improve our under- 
standing of lamprey taxonomy and evolution. For 


a3 


574 


this reason, it is imperative that the Morrison Creek 
Lampetra richardsoni population is preserved and 
protected. We term this population a “variety” of 
Lampetra richardsoni, as variety marifuga which 
relates to the inability of this lamprey to survive in 
the ocean. 


Distribution 

The known distribution is extremely restricted, for 
this polymorphic population of Lampetra richardsoni 
has only been found in the Morrison Creek watex- 
shed, located on Vancouver Island, British Columbia, 
Canada (Figures | and 2). Morrison Creek is a small 
freshwater stream which originates from a series of 
springs (Beamish 1985). Stream channels in the high- 
er gradient headwaters are typically 1-2 m wide and 
about 3—4 m in width at the lower reaches (Lough 
1995a*,b*; Beamish 1985). Morrison Creek runs for 
approximately 35 kilometers until its confluence with 
the Puntledge River system, a few kilometers 
upstream of the ocean (Beamish 1985). Small num- 
bers of the anadromous Lampetra tridentata have 
also been found in Morrison Creek (Beamish and 
Withler 1986). 


Protection 

There are currently no protection provisions for the 
Morrison Creek Western Brook Lamprey. However, 
due to the problems of declining wild Coho stocks in 
the Strait of Georgia, there has been an increased 
effort to protect and enhance fish habitat. A sensitive 
habitat atlas was created for the Comox-Strathcona 
region. It highlights the areas which contain sensitive 
aquatic and terrestrial habitat so that they are consid- 
ered in any proposed land developments (Comox- 
Strathcona Sensitive Habitat Atlas 1995). Fish habitat 
in these areas are subject to fisheries protection regu- 
lations and guidelines; however, sometimes encroach- 
ment into sensitive habitats can not be avoided, and 
development plans must resort to mitigation and com- 
pensation strategies to replace the losses. 


Population Numbers, Sizes and Trends 

There are no reliable population estimates of 
Lampetra richardsoni in Morrison Creek. The num- 
ber of spawning individuals (considered to be typical 
Lampetra richardsoni) caught in traps exceeded the 
number of the variety that have been captured 
(Beamish and Withler 1986). Furthermore, a sample 
of 18 metamorphosed lampreys collected in 
Morrison Creek produced two silver forms (the 
Morrison Creek variety) and the rest matured, 
spawned and died in the spring as would typical 
Lampetra richardsoni (Beamish 1985). Data suggest 
that the Morrison Creek variety was relatively stable 
during the initial studies which ran from 1978 to 
1984 (Beamish 1985). However, it is possible that 
their numbers have declined in recent years 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


FIGURE 1. Location of Morrison Creek in British Columbia. 


(Beamish unpublished data). The adults of the 
Morrison Creek variety range in size from 10 to 
15 cm, and it is thought that their length increases 
after metamorphosis are small (Beamish 1985). The 
sex ratio of the variety appears to be about 80% male 
and the males contain gonads in an advanced stage 
of maturity at a time when females are immature or 
just beginning to mature (Beamish 1985). A difficul- 
ty has been the inability to catch adults after the 
summer (or rather to distinguish them from recently 
metamorphosed lampreys), thus the biology and fate 


Puntledge 
River : 
Courtenay .” 


Morrison / 
Creek . 
wf Strait of 
“1 Georgia 


FIGURE 2. Location of Morrison Creek on Vancouver Island, 
the scale represents 4.2 km. 


2001 


of the variety in Morrison Creek is unknown after 
the summer period (Beamish 1985). 


Habitat 

Habitat studies conducted on the upper water- 
shed (Lough et al. 1995a*,b*; Knight and Blood 
1997) indicate that the Morrison Creek area is char- 
acteristic of interlinking wetlands, with meadows, 
thick brush, beaver dams and open beaver ponds. 
The stream bed is dominated by compressed till 
with limited patches of small graveis and an abun- 
dance of stream debris which provide habitat diver- 
sity. : 
The creek is known for its Coho Salmon, Oncor- 
hynchus kisutch, habitat and is a major contributor of 
Coho to the Puntledge River system (personal com- 
munication: Brian Allen, Department of Fisheries 
and Oceans, 148 Port Agusta, Comox, British 
Columbia). The specific habitat features required to 
support a polymorphic population of lamprey are not 
known. 

The area surrounding Morrison Creek has, in the 
past, been disturbed by logging and mining activities 
which contribute to its complex hydrology (Knight 
and Blood 1997). Evidence of development pres- 


BEAMISH, YOUSON, CHAPMAN: STATUS OF MORRISON CREEK WESTERN BROOK LAMPREY 


ey 


sures are also seen in the mainstem of the creek 
where streambank degradation and the removal of 
in-stream debris has resulted in a loss of the pool/ 
riffle complex (personal communication: Brian 
Allen, Department of Fisheries and Oceans, 148 Port 
Agusta, Comox, British Columbia). 


General Biology 

The biology of the Morrison Creek variety is not 
fully understood. Aside from an extended post- 
metamorphic period and the ability to be parasitic, 
its biology is very similar to that of typical Lampetra 
richardsoni (Beamish 1985). Typical Lampetra 
richardsoni from Morrison Creek remain in fresh 
water throughout their entire life cycle and begin to 
reproduce in May and June (Youson and Beamish 
1991). The lamprey spawn only once and their eggs 
are deposited in river bed gravel. After hatching, the 
young quickly burrow into the soft bottom sediments 
where they spend an unknown time (possibly three 
to seven years) as filter feeding ammocoetes before 
metamorphosing into juveniles. 

The population of Lampetra richardsoni in 
Morrison Creek begins metamorphosis in July or 
August and two adult forms appear in the spring of 


FiGURE 3. (A) Male of the Morrison Creek variety showing counter shading which develops in the spring fol- 
lowing metamorphosis. The variety can live for one year longer than typical Lampetra richardsoni and 
is capable of feeding during this time under laboratory conditions. Total length is 13cm. (B) Mature 
male of the Morrison Creek variety brought into the laboratory in June 1981 and regard to maturity in 
1982. Total length is 18.5 cm. (C) Mature female and (D) mature male, typical Lampetra richardsoni 
from Morrison Creek, which spawn and die in the spring after metamorphosis. Total lengths are 12.5 cm 


and 11 cm respectively. 


576 


the following year: a spawning variety (Figure 3: C 
and D) with advanced signs of sexual maturation, 
typical of Lampetra richardsoni, and a parasitic vari- 
ety (Figure 3: A and B), which is not completely 
mature and could delay another year before spawning 
(Beamish 1987a; Youson and Beamish 1991). No 
distinction has been made, however, between the two 
morphs of the population when they are ammocoetes. 

Electrophoretic and morphological evidence 
(Beamish and Withler 1986; Beamish 1987b) have 
shown that the two forms of Lampetra richardsoni in 
Morrison Creek belong to a single population, thus 
forming a species complex. For instance, the non- 
parasitic morph is more genetically similar to the 
parasitic morph than it is to other Lampetra richard- 
soni populations from different streams. It is uncer- 
tain, however, whether the two morphs breed as 
independent lines. It is possible that the Morrison 
Creek variety does not survive to reproduce as its 
internal development at metamorphosis is not com- 
plete in all individuals (Youson and Beamish 1991). 
However, some individuals of the variety spawn suc- 
cessfully in the laboratory and produce viable eggs 
which hatch into larvae (Beamish 1985). Given that 
spawning was successful in the laboratory, it is also 
possible that the variety is reproducing in Morrison 
Creek. Identical allelic frequencies indicate that 
genetic exchange occurs between the two forms of 
Lampetra richardsoni in Morrison Creek (Beamish 
and Withler 1986). 

The Morrison Creek variety is able to live for one 
year longer than typical Lampetra richardsoni as 
demonstrated in the laboratory (Beamish 1987a). 
Youson and Beamish (1991) found that the delayed 
maturation in males of the Morrison Creek variety 
allowed the retention of a functional digestive sys- 
tem which would provide an opportunity for feeding. 
The variety fed (on both live and dead fishes) and 
grew under laboratory conditions from July until 
mid-November and matured over the following 
spring (Beamish 1985). It is uncertain, however, 
whether the Morrison Creek variety feeds in the 
stream. Feeding may occur for only a short period 
and on food items other than live fishes (Beamish 
1985). However, it is also possible that the variety 
does not feed in Morrison Creek and simply perishes 
before spawning. 

The Morrison Creek variety, when not in spawn- 
ing condition, is easily distinguished from typical 
Lampetra richardsoni by its silver color and counter 
shading, which develops in early spring, and by its 
prominent teeth (Beamish 1985). It was noted that 
the feeding variety maintained in the laboratory lost 
the silver color and became uniformly dark by the 
end of September (Beamish 1985). 

The variety appears to be very similar to the 
closely related Lampetra ayresi but it differs in the 
number of cusps on some teeth, some aspects of its 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


morphology, in colour, and its inability to osmoregu- 
late in salt water (Beamish 1985). It is possible that 
the Morrison Creek variety may be a natural hybrid 
of Lampetra richardsoni and Lampetra ayresi; how- 
ever, Lampetra ayresi have not been found in the 
Morrison Creek or surrounding watersheds (Beamish 
1985; Beamish and Withler 1986). Furthermore, no 
ammocoetes in Morrison Creek with pigmentation 
patterns typical of Lampetra ayresi crosses have 
been found (Beamish and Withler 1986). 

Youson and Beamish (1991) compared the inter- 
nal morphology of the two morphs and suggested 
that the intermediate parasitic lifestyle is due to a 
slower sexual maturation and, possibly, a slower and 
incomplete metamorphosis. The incomplete meta- 
morphosis was particularly evident in females, 
which were far less developed than males, and this 
may account for the abnormal sex ratio (Youson and 
Beamish 1991). In addition, males of the parasitic 
variety were found to contain gonads in an advanced 
stage of maturity with no external signs of matura- 
tion and were capable of feeding (Beamish 1985; 
Beamish and Withler 1986; Beamish 1987a; Youson 
and Beamish 1991). This is the first documentation 
of a lamprey that is actively feeding in an advanced 
stage of maturity (Beamish and Withler 1986; 
Youson and Beamish 1991). The unusual biology of 
the Morrison Creek variety appears to represent a 
key phase of lamprey evolution. The inability to 
osmoregulate in salt water, the feeding habit in the 
laboratory, the precocious development in males, 
and the delayed organ development indicate that the 
variety is intermediate between a parasitic and non- 
parasitic form (Beamish 1985; Beamish and Withler 
1986; Beamish 1987a; Youson and Beamish 1991). 


Special Significance 

The existence of a lamprey population in Canada 
that produces two distinct life history types has only 
been described from Morrison Creek. This rare pop- 
ulation represents an important transition in the evo- 
lution of adult life history types in lampreys and may 
be the key to understanding why lampreys have been 
successful for over 350 million years. 

Lampetra richardsoni is a close relative of the 
anadromous parasitic Lampetra ayresi, and they are 
considered to be a species pair (Zanandrea 1959; 
Vladykov and Kott 1979; Potter 1980). The evolu- 
tionary relationship between members of a species 
pair remains uncertain, although there is a widely 
accepted view that the non-parasitic form evolved 
from. an anadromous parasitic form (Vladykov and 
Kott 1979; Potter 1980). The Morrison Creek vari- 
ety is intermediate between the non-parasitic 
Lampetra richardsoni and the parasitic Lampetra 
ayresi in its biology and morphology (Beamish and 
Withler 1986). According to Zanandrea (1959, 
1961), a freshwater feeding phase may be an inter- 


2001 


mediate step in the evolution of freshwater non- 
parasitic lampreys from anadromous parasitic 
forms. The existence of the Morrison Creek variety 
provides evidence of this intermediate freshwater 
feeding phase. 

Histological evidence indicates that the freshwater 
parasitic lifestyle of Lampetra richardsoni in Mor- 
rison Creek is the result of a delayed sexual matura- 
tion and possibly a delayed and incomplete meta- 
morphosis (Youson and Beamish 1991). Based on 
this evidence from the Morrison Creek population, it 
has been suggested that the appearance of the two 
morphs within a single population may not necessar- 
ily represent an intermediate evolutionary transition 
but rather the sensitivity of lamprey metamorphosis 
to environmental factors (Youson 2002). It is 
hypothesized that the two forms may be the result of 
a recent heterochrony during metamorphosis which 
has effected the post metamorphic rate of sexual 
maturation (Youson 2002). This sensitivity to envi- 
ronmental factors and the change in developmental 
timing may explain both the success of lampreys and 
the presence of two adult life history types among 
lampreys as a group. 

In any case, the presence of a parasitic variety of 
Lampetra richardsoni is of special significance to 
the scientific community as it demonstrates the very 
close relationship between the two life history types 


and has important implications to lamprey systemat- | 


ics (Beamish 1985; Beamish and Withler 1986). It 
appears to show that during metamorphosis, it is 
possible to direct the life history type into two direc- 
tions. The occurrence of polymorphic populations 
such as the Lampetra richardsoni population found 
in Morrison Creek also confounds the use of life his- 
tory type as a basis for defining species (Beamish 
and Withler 1986). 


Limiting Factors 

The survival of the Morrison Creek Lampetra 
richardsoni complex depends on the protection of the 
entire lamprey population and its habitat. There has 
been concern that rapid development in the area has 
overwhelmed the ability to protect sensitive habitat 
(Comox-Strathcona Sensitive Habitat Atlas 1995). 
Residential development has encroached on the 
mainstem of the creek and has resulted in an alter- 
ation of the riparian vegetation which poses a definite 
threat to fish habitat. A recent concern is the short- 
term and long-term effects of highway construction 
on fish habitat in Morrison creek. The British 
Columbia Ministry of Transportation and Highways 
has proposed to cross two sections of Morrison Creek 
sometime during 1999 (Lough et al. 1995a*,b*). The 
major fisheries concerns at these crossings are the 
potential loss of a considerable amount of fish habitat 
and the possible loss of fish passage at one of the 
crossings (Ship Environmental Consultants 1993). 


BEAMISH, YOUSON, CHAPMAN: STATUS OF MORRISON CREEK WESTERN BROOK LAMPREY 


577 


Although mitigation options have proposed to 
replace habitat loss with an equal area of new habi- 
tat, it should not be assumed that all habitat conser- 
vation needs will be met in this way (Lough et al. 
1995a*,.b*; Knight and Blood 1997). It is important 
that existing fish populations are not compromised 
by mitigation and enhancement projects and that 
consideration is given to the effects of these activi- 
ties on all species within the system. 

Given the potential impacts of development pres- 
sures, the survival of Lampetra richardsoni in 
Morrison Creek is highly at risk. A significant loss 
of fish numbers in a presumably small population, 
combined with a considerable loss of habitat may 
prevent the survival of this rare species complex. 


Evaluation 

Due to its restricted distribution and threatened 
habitat, the polymorphic population of Lampetra 
richardsoni in Morrison Creek is both threatened and 
endangered. There is an urgent need to protect this 
most unusual and poorly understood species complex. 


Acknowledgments 

This report was funded through a contract with 
COSEWIC on behalf of the Canadian Wildlife 
Service, Government of Canada. We would like to 
thank Robert Campbell, whose pleasantly persistent 
manner was responsible for initiating this work; Ray 
Scarsbrook, for his technical assistance in producing 
the necessary maps; Bonita Wallace for her secretar- 
ial assistance, Edward Matko and Michael Folkes for 
their assistance with computer graphics. 


Documents Cited 

Lough, M. J., & Associates, and Thurber Engineering 
Ltd. 1995a. Morrison Creek Fish Habitat Study Maple 
Lake To Lake Trail Road). Report to Vancouver Island 
Highway Project, Ministry of Transportation and High- 
ways, Courtenay, British Columbia. 

Lough, M. J., & Associates. 1995b. Morrison Creek Fish 
Habitat Study — Addendum — Prepared for Vancouver 
Island Highway Project, Ministry of Transportation and 
Highways, Victoria, British Columbia. 

Comox-Strathcona Sensitive Habitat Atlas. 1995. A Co- 
operative project of the Province of British Columbia, 
and the Regional District Comox-Strathcona. 

Ship Environmental Consultants. 1993. Vancouver In- 
land Island Highway Study. Fisheries Impact Assessment 
Vol. IV, Section 442. Vancouver Island Highway Project 
British Columbia Ministry of Transportation and High- 
ways, Victoria, British Columbia. (sub-projects and 
schedules). 1998. Ministry of transportation and High- 
ways, Victoria. 


Literature Cited 

Beamish, R. J. 1985. Freshwater parasitic lamprey on 
Vancouver Island and a theory of the evolution of the 
freshwater parasitic and non-parasitic life history types. 
Pages 123-140. in Evolutionary biology of primitive 


578 


fishes. Edited by R. E. Foreman, A. Gorbman, J. M. 
Dodd, and R. Olson. Plenum Publishing Corporation, 
New York, NY. 463 pages. 

Beamish, R. J. 1987a. Evidence that parasitic and non- 
parasitic life history types are produced by one popula- 
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THE CANADIAN FIELD-NATURALIST 


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Potter, I. C. 1980. the Petromyzontiformes with particular 
reference to paired species. Canadian Journal of Fish- 
eries and Aquatic Sciences 37: 1641-1657. 

Viladykov, V. D., and E. Kott. 1979. Satellite species 
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Youson, J. H., and R. J. Beamish. 1991. Comparison of 
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ogy 69: 628-637. 

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Pearson, and G. B. Muller, MIT Press, Cambridge, 
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Evolution 15: 523-534. 


Accepted 19 February 2002 


Status of the Stickleback Species Pair, Gasterosteus spp., 
in Hadley Lake, Lasqueti Island, British Columbia* 


TODD HATFIELD 


Solander Ecological Research, 1324 Franklin Terrace, Victoria, British Columbia V8S 1C7 Canada 


Hatfield, Todd. 2001. Status of the stickleback species pair, Gasterosteus spp., in Hadley Lake, Lasqueti Island, British 
Columbia. Canadian Field-Naturalist 1 15(4): 579-583. 


Two species of threespine stickleback (Gasterosteus spp.) formerly inhabited Hadley Lake on Lasqueti Island, British 
Columbia, and are described briefly in this report. One species (referred to as “limnetic”) was a pelagic zooplankton feeder, 
and the other (referred to as “benthic”) exploited littoral zone benthic invertebrates. The species were unique British 
Columbia endemics restricted to this single lake. The species were first discovered in the late 1980s, but have recently 
become extinct following the unauthorized introduction of catfish (Ameirus nebulosus) to the lake. Independently evolved 
yet morphologically similar species pairs exist elsewhere on nearby islands. Measures should be taken, such as transplant- 
ing to inaccessible fishless lakes, to protect the remaining species pairs. 


Key Words: Gasterosteidae, sticklebacks, Gasterosteus, épinoches, Hadley Lake, Lasqueti Island, British Columbia, 


endangered species. 


The Threespine Stickleback (Gasterosteus aculea- 
ius complex) is a small (usually 35-55 mm) fish com- 
monly found in marine and freshwater throughout the 
northern hemisphere. Marine sticklebacks are essen- 
tially identical throughout their range, whereas fresh- 
water sticklebacks are ecologically, behaviorally and 
morphologically extremely variable. In general, G. 
aculeatus has a laterally compressed body and deli- 
cate pectoral and caudal fins. It is generally well- 
armored, getting both its latin and common names 
from different aspects of its armor. Sticklebacks have 
retractable pelvic and dorsal spines, and the body is 
protected by calcified lateral plates. Freshwater popu- 
lations are highly variable in extent of armor but gen- 
erally have many fewer lateral plates than the marine 
form. Body color also varies considerably from sil- 
very to mottled green and brown. Sexually mature 
males develop bright red throats, though in some 
freshwater populations males turn completely black 
instead (McPhail 1969; Reimchen 1989). 

At the end of the Pleistocene marine sticklebacks 
colonized freshwater repeatedly (Bell 1976). That is, 
each extant stream or lake population of sticklebacks 
is assumed to have arisen by an independent inva- 
sion from the sea (as opposed to a single coloniza- 
tion of one or a few bodies of freshwater followed by 
range expansion into previously uncolonized streams 
or lakes). Thus, the marine form is presumed to be 
the most recent ancestor to freshwater forms (Bell 
and Foster 1994). This scenario suggests that the 
tremendous variability we currently see in British 


*Reviewed and approved by COSEWIC April 1999, status 
assigned to benthic and limnetic forms — Extinct. 


Columbia freshwater sticklebacks has arisen in 
approximately the last 10 000 to 13 000 years. 

The British Columbia coastline is dotted with 
hundreds of small, low elevation lakes. Many of 
these lakes have been surveyed for sticklebacks (e.g., 
Lavin and McPhail 1985; Reimchen et al. 1985; 
Schluter and McPhail 1992; McPhail 1993) and most 
contain a single, albeit variable, form of stickleback 
(see e.g., Bell 1976; Lavin and McPhail 1985; 
Riemchen et al. 1985; Schluter and McPhail 1992). 
However, several lakes on islands in the Strait of 
Georgia are especially noteworthy because they each 
contain two distinct species (McPhail 1984, 1992, 
1993, 1994; Schluter and McPhail 1992). The lakes 
are found on Texada Island, Lasqueti Island and 
Vancouver Island. 

We refer to the two species in each lake as a 
“species pair’. The pattern of morphological and 
ecological divergence is remarkably similar in each 
of the lakes (Schluter and McPhail 1992). In each 
case, one of the species (referred to as “limnetic’’) 
primarily exploits plankton, and has morphological 
traits such as a fusiform body, narrow mouth and 
many, long gill rakers (Figure 1). These traits are 
considered adaptations to a zooplankton-consuming 
lifestyle (Kliewer 1970; Magnuson and Heitz 1971; 
Sanderson et al. 1991; Schluter 1995). The other 
species (referred to as “benthic”) mainly exploits 
benthic invertebrates in the littoral zone, and has a 
robust body form (Figure 1), wide gape and few, 
short gill rakers, traits considered to be advantageous 
in benthic feeding (Schluter 1995). 

Although limnetics and benthics have yet to be 
assigned formal scientific names there is no argu- 
ment among researchers studying these fish that they 
warrant taxonomic status as distinct species. In each 
of the lakes limnetics and benthics co-exist and are 


579 


580 


THE CANADIAN FIELD-NATURALIST 


Vol. 


lille 


FIGURE 1. Limnetic (A) and benthic (B) forms of stickleback, Gasterosteus spp. (drawn by L. Nagel). 


reproductively isolated by behavioral and genetic 
differences (McPhail 1984, 1992; Ridgway and 
McPhail 1984; Nagel 1994; Hatfield 1995, 1997; 
Hatfield and Schluter 1996). They therefore meet a 
conservative definition of species (Mayr 1942, 
1963). One reason for the delayed naming of the 
species is that we have been awaiting good evidence 
of whether the species pairs are independently 
evolved or represent a single speciation with subse- 
quent dispersal. That is, are we dealing with two 
species, or more? Geography and geological history 
are consistent with the hypothesis of independent 
evolution (McPhail 1993; 1994), but the best evi- 
dence will be a DNA-based phylogeny. Studies to 
construct such a phylogeny are underway and initial 
evidence suggests that the species pairs are indeed 
independently derived (McPhail 1993; Taylor et al. 
1997; Taylor, unpublished data). Thus, the similar 
morphology and ecology of the pairs is due to paral- 
lel evolution, and we must name more than two 
species. Species pairs have some opportunity for dis- 
persal through streams to adjacent lakes in the same 
watershed, though little opportunity for dispersal to 
other watersheds. It is therefore assumed that, since 
Hadley Lake is the only lake in its watershed, its 
sticklebacks represent distinct species. 


Taxonomy 

Taxonomic classification of British Columbia 
sticklebacks is highly complex, and presents one of 
the greatest challenges to systemicists of British 
Columbia fish fauna. Freshwater populations of stick- 
lebacks are invariably distinct from the marine form, 
yet they show both parallel evolution and remarkable 
phenotypic variation among sites (e.g., Hagen and 
Gilbertson 1972; Lavin and McPhail 1985; Reimchen 


et al. 1985; Schluter and McPhail 1992; Bell and 
Foster 1994; McPhail 1994). Classical approaches to 
systematics rely on the measurement of morphologi- 
cal traits and assume that traits are never (or at least 
very rarely) evolved in parallel. Because parallel evo- 
lution is rampant in freshwater sticklebacks their tax- 
onomy has been in chaos for some time (see e.g., 
Hagen and McPhail 1970). It is precisely this mix of 
parallel and independent phenotypic evolution that 
confounds taxonomists while at the same time 
intrigues ecologists and evolutionary biologists. It is 
now generally agreed that freshwater populations of 
sticklebacks have been derived from marine stickle- 
backs multiple times. 

Modern molecular genetic techniques offer the 
most promising approach for resolving taxonomic 
issues in British Columbia sticklebacks. When con- 
structing phylogenies, molecular data are used in 
much the same way as morphological data. The 
major difference is that the basic assumption of no 
parallel evolution is much more likely to be valid at 
a molecular level. 

Studies using molecular approaches to stickleback 
taxonomy are currently underway. Regional patterns 
have been reported in Withler and McPhail (1985) and 
Orti et al. (1994). Finer scale issues (the most impor- 
tant issues for conservation biologists) are being 
researched primarily at labs at the University of British 
Columbia. It will be some time before data are com- 
plete and the issues resolved but early results have 
been reported in Taylor et al. (1997). These and 
unpublished data suggest that stickleback species pairs 
are genetically distinct units. However, more data are 
needed in order to understand the relationships among 
the different pairs and to determine which of the pairs 
are independently derived from the marine form, and 


2001 


which represent replicate populations of the same 
species. Unfortunately, the Hadley Lake species pair 
became extinct before scientists could sample DNA 
from these fish. It is assumed that the considerable 
geographic isolation of the Hadley Lake sticklebacks 
indicates that they evolved independently from other 
species pairs. They should thus be named as two 
species distinct from other species pairs. 

In reality the decision to award species status to 
an organism is subjective (McPhail and Carveth 
1992). All of the limnetics and benthics studied to 
date meet conservative definitions of biological 
species (see Mayr 1942, 1963). For example, they 
maintain their morphological differences over sever- 
al generations in a common environment (McPhail 
1992; Hatfield 1995, 1997), they do not interbreed in 
the lab if given a choice among mates (Ridgway and 
McPhail 1984; Nagel 1994), and they have remained 
distinct in the wild despite some very large distur- 
bances to their habitat (Larson 1976; McPhail 1994). 
Molecular data corroborate the view that benthics 
and limnetics are genetically different. Most biolo- 
gists would not dispute that these are true species, 
albeit extremely young species. 


Distribution 

Hadley Lake, known locally as “Pete’s Lake’, is 
near False Bay, Lasqueti Island, British Columbia 
(Figure 2). To our knowledge the two species from 
Hadley Lake were restricted to this site (49° 30’ N, 
124° 20’ W). Other lakes on the island have been 
surveyed and do not contain species pairs. 


} Vancouver 


Island de. 


FiGuRE 2. Map of the Strait of Georgia, British Columbia. 
Indicated are the major lakes in each of the four 
watersheds with species pairs. 


HATFIELD: STATUS OF STICKLEBACK SPECIES PAIR IN HADLEY LAKE 


58] 


Protection 

There is currently no specific protection for any of 
the extant stickleback species pairs. Much of the 
potential for legislated protection of sticklebacks 
appears to depend on whether they are designated 
“endangered species” and whether proposed legisla- 
tion gets enacted. The Fisheries Act, federal legisla- 
tion that extends protection to fish and fish habitat, 
was originally written for the protection of extractive 
fisheries (i.e., commercial, recreational, aboriginal) 
and it is doubtful that it can be successfully applied 
to the extant stickleback species pairs. There is very 
little political support for protection of non-salmonid 
fish or non-game species in British Columbia. 


Population Sizes And Trends 

Sticklebacks have not been observed in Hadley 
Lake since the early 1990s and both species are now 
presumed extinct. Two separate expeditions have 
failed to observe a single stickleback despite consider- 
able effort. Hadley Lake limnetics and benthics have 
been designated extinct by the British Columbia 
Conservation Data Centre. Specimens of Hadley Lake 
limnetics and benthics are held in the Fish Museum of 
the University of British Columbia. 

Sometime in the early 1990s catfish (Ameirus neb- 
ulosus) were introduced to Hadley Lake. Nocturnal 
nest predation by the catfish is the likely mechanism 
for the stickleback extinctions. Male sticklebacks will 
vigorously defend their nests during the day, but are 
poorly equipped to defend their nests at night. Catfish 
are not indigenous to this area; thus, sticklebacks 
have not co-evolved with this predator. Stickleback 
extinctions have occurred in other lakes following the 
introduction of catfish (McPhail 1989). The common 
pattern is that catfish numbers build until stickleback 
recruitment fails, leaving only an adult population 
that subsequently goes extinct within one or two 
years. Extinction can be alarmingly rapid: in the case 
of Hadley Lake it occurred before any conservation 
effort could be mounted. 

Prior to catfish introductions the sticklebacks were 
under no apparent threat. Hadley Lake is small 
(~ 10 ha), but it likely supported large populations of 
both limnetic and benthic sticklebacks. However, no 
quantitative estimates of population size were 
attempted. McPhail (1989) estimated population sizes 
on the order of 100 000 for each of the species in Enos 
Lake, which is roughly twice the size of Hadley Lake. 
If Hadley Lake had half this number it would have 
been considered locally abundant by any standard. 


Habitat 

Hadley Lake is a small coastal lake typical of this 
region. It is ~ 50 m above sea level and is connected 
to the sea by ~ 1.2 km of stream. There is no perma- 
nent inlet stream for the lake and the outlet creek has 
been dammed and culverted. The hydrology of the 
outlet stream has been substantially modified by 


582 


road building and housing development. It is possi- 
ble that the stream and lake had Cutthroat Trout 
(Oncorhynchus clarki) prior to these alterations. 

The lake is lightly stained, and in the summer the 
littoral region is covered in dense beds of Chara, 
Potamogeton and Utricularia. There is a distinct 
pelagic zone in the lake, and in the summer plankton 
productivity is high. Catfish are presently the only 
fish species in the lake. 


General Biology 

The Hadley Lake stickleback species are the least 
studied of the known species pairs. They were not 
subjected to intense study in the way that species 
pairs from Enos and Paxton lakes have been. 
Although scientists believe that Hadley Lake limnet- 
ics and benthics were unique British Columbia 
endemics they were likely similar in ecology and 
behavior to the other species pairs. It is also assumed 
that they followed the general life history pattern of 
most Gasterosteus populations (for reviews of gener- 
al stickleback biology see Wooton 1976; Bell and 
Foster 1994). The following brief description is 
based on the few observations of Hadley Lake stick- 
lebacks in addition to observations of the other 
species pairs (see McPhail 1994 for review). 

During the spring and summer months Hadley 
Lake limnetics were found in the open water where 
they foraged for plankton, and benthics were found in 
the littoral zone where they foraged for benthic inver- 
tebrates. During fall and winter months both species 
dispersed to deeper water. At maturity benthics were 
bigger on average than limnetics, and it is assumed 
that they lived longer than limnetics, perhaps forgoing 
reproduction in their first year in favor of growth. 
Limnetics reproduced at the end of their first year, and 
usually died before their second reproductive season. 

Male sticklebacks are the sole providers of 
parental care. In the spring, both limnetics and ben- 
thics acquired territories in the littoral region where 
they built nests and mated (sometimes with many 
females). Following fertilization eggs took approxi- 
mately 7—10 days to hatch, depending on the temper- 
ature. During this time males actively aerated the 
eggs by forward thrusts of their pectoral fins; 
embryos die if inadequately aerated (van den Assem 
1967; Sargent and Gebler 1980). Males of both 
species vigorously defended their nests and territo- 
ries from invaders (most often other sticklebacks) 
and continued to defend their young for about a 
week after they hatched. Sexual dimorphism was 
absent or minimal among benthics, but limnetic 
males tended to be bigger on average than limnetic 
females. Large male size enabled greater nest protec- 
tion and territory defence (Rowland 1989). 


Limiting Factors 
_ Prior to the introduction of catfish to Hadley Lake 
there were no piscivorous fish, though herons (Ardea 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


herodias), kingfishers (Megaceryle alcyon) and 
loons (Gavia immer) are regularly seen in the area. 
Their effect on populations was likely minimal with 
the primary limiting factor being the capacity of the 
lake to produce plankton and benthos. 


Special Significance of the Species 

For more than 100 years one of the great ques- 
tions in biology has been how diversity originates 
and is maintained. Speciation, the division of one 
species into two or more species, is the ultimate pro- 
cess giving rise to diversity. One major reason for 
our remarkably slow progress in understanding spe- 
ciation has been that it usually occurs over vast time 
frames. It is therefore difficult to identify popula- 
tions undergoing speciation until the process is com- 
plete or nearly complete. Furthermore, forces that 
maintain two species as distinct today are not neces- 
sarily the same forces that drove speciation several 
millennia ago. 

For the student of speciation there are two excep- 
tionally fascinating aspects of the biology of stickle- 
back species pairs. The first is that the species are so 
young that it is reasonable to expect that the forces 
that drove speciation are the same forces that main- 
tain their present reproductive isolation. These pro- 
cesses can therefore be studied in situ (e.g., Hatfield 
1995; Schluter 1995). The second aspect is that simi- 
lar speciation events have produced limnetic-benthic 
species pairs elsewhere on the coast, thus providing 
unheard of natural replication in studies of specia- 
tion. The loss of opportunity for further study of the 
Hadley Lake species pair is widely regarded as a sci- 
entific tragedy. 

At a more local scale the ramifications of the loss 
of Hadley Lake sticklebacks depend on one’s point 
of view. The loss of local biodiversity has been 
mourned from a purely aesthetic point of view. 
However, more tangible effects may be that there are 
more mosquitoes and midges in the region, and 
fewer piscivorous birds to watch. 


Evaluation 

Hadley Lake sticklebacks were among the most 
rare and endangered species in the world. The 
British Columbia Conservation Data Centre has des- 
ignated them extinct. They should now be designated 
extinct by COSEWIC. For the reasons noted above 
these designations are justified. 


Aknowledgments 

Preparation of the report was funded by the 
British Columbia Conservation Data Centre. The 
project was coordinated by Syd Cannings, British 
Columbia Conservation Data Centre, Victoria, 
British Columbia. 


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Accepted 19 February 2002 


Status of the Stickleback Species Pair, Gasterosteus spp., in the 
Vananda Creek watershed of Texada Island, British Columbia* 


TODD HATFIELD 


Solander Ecological Research, 1324 Franklin Terrace, Victoria, British Columbia V8S 1C7 Canada 


Hatfield, Todd. 2001. Status of the stickleback species pair, Gasterosteus spp., in the Vananda Creek watershed of Texada 
Island, British Columbia. Canadian Field-Naturalist 115(4): 584-590. 


A pair of threespine stickleback species (Gasterosteus spp.) inhabit Balkwill, Emily and Priest Lakes (Vananda Creek 
watershed) on Texada Island, British Columbia. One species (referred to as “limnetic’’) is a pelagic zooplankton feeder, and 
the other (referred to as “benthic’”’) is adapted for feeding on littoral zone benthic invertebrates. Current data suggest that 
limnetics and benthics are unique biological species and are restricted to this single watershed made up of three lakes. 


Key Words: Gasterosteidae, sticklebacks, Gasterosteus, épinoches, Balkwill, Emily, Priest lakes, Vananda Creek, endan- 


gered species. 


Threespine sticklebacks (Gasterosteus aculeatus) 
form a large group of species made up of thousands 
of phenotypically diverse populations. Typically 
sticklebacks are small (usually 35 to 55 mm) fish 
that are abundant in coastal marine and freshwater 
throughout the northern hemisphere. Marine stickle- 
backs are phenotypically similar throughout their 
range, whereas freshwater sticklebacks are ecologi- 
cally, behaviorally and morphologically extremely 
variable. In general, G. aculeatus has a laterally 
compressed body and delicate pectoral and caudal 
fins. Individuals in most populations are well- 
armored—sticklebacks get both their latin and com- 
mon names from different aspects of their armor. 

Sticklebacks have retractable pelvic and dorsal 
spines, and their bodies are covered with calcified 
lateral plates. Freshwater populations are highly 
variable in extent of armor but generally have many 
fewer lateral plates than the marine form. Body color 
also varies considerably, from silvery to mottled 
green and brown. Sexually mature males in most 
populations develop bright red throats, although in a 
few freshwater populations males turn completely 
black instead (McPhail 1969; Reimchen 1989). 

Many extant stream and lake populations of stick- 
lebacks arose at the end of the Pleistocene (the most 
recent glaciation or “Ice Age”) via independent inva- 
sions from the sea (as opposed to a single coloniza- 
tion of one or a few bodies of freshwater followed by 
range expansion into previously uncolonized streams 
or lakes). Thus, the marine form is generally pre- 
sumed to be the most recent ancestor to most fresh- 
water forms (Bell and Foster 1994). This scenario 
suggests that the tremendous variability we currently 


*Reviewed and approved by COSEWIC April 1999, status 
assigned — Threatened. 


see in British Columbia freshwater sticklebacks has 
arisen within approximately the last 13 000 years. 

The British Columbia coastline is dotted with hun- 
dreds of small, low elevation lakes. Many of these 
lakes have been surveyed for sticklebacks (e.g., Lavin 
and McPhail 1985; Reimchen et al. 1985; Schluter 
and McPhail 1992; McPhail 1993) and most contain a 
single, albeit variable, form of stickleback (see e.g., 
Bell 1976; Lavin and McPhail 1985; Reimchen et al. 
1985; Schluter and McPhail 1992). However, several 
lakes on islands in the Strait of Georgia (see Figure 1) 
are especially noteworthy because they each contain 
two distinct species (McPhail 1984, 1992, 1993, 1994; 
Schluter and McPhail 1992). Balkwill, Emily, Priest 
and Paxton lakes are located on Texada Island (see 
Figure 2), Hadley Lake is located on Lasqueti Island, 
and Enos Lake is located on Vancouver Island. This 
report documents the current status of the species pair 
from Balkwill, Emily, and Priest lakes. 


Description 

We refer to the two species in each lake as a “spe- 
cies pair” (Figure 2). The pattern of morphological 
and ecological divergence is remarkably similar in 
each of the lakes (Schluter and McPhail 1992). In 
each case, one of the species (referred to as “limnet- 
ic’) primarily exploits plankton, and has morphologi- 
cal traits such as a fusiform body, narrow mouth and 
many, long gill rakers. These traits are considered 
adaptations to a zooplankton-consuming lifestyle 
(Magnuson and Heitz 1971; Kliewer 1970; Sanderson 
et al. 1991; Schluter and McPhail 1992, 1993). The 
other species (referred to as “benthic”) mainly 
exploits benthic invertebrates in the littoral zone, and 
has a robust body form, wide gape and few, short gill 
rakers, traits considered to be advantageous in benthic 
feeding (Schluter and McPhail 1992, 1993). 

Although limnetics and benthics have yet to be 
assigned formal scientific names there is no argu- 


584 


Vancouver 


Island “&. 


125 124 123 


FiGuRE 1. Map of the Strait of Georgia, British Columbia. 
Indicated are the major lakes in each of the four 
watersheds with species pairs. 


ment among researchers studying these fish that they 
warrant taxonomic status as distinct species. In each 
of the lakes limnetics and benthics co-exist and are 
reproductively isolated by behavior and genetic dif- 
ferences (McPhail 1984, 1992; Ridgway and 
McPhail 1984; Nagel 1994; Hatfield 1995, 1997; 
Hatfield and Schluter 1996). They therefore meet a 
conservative definition of species (Mayr 1942,1963). 


HATFIELD: STATUS OF STICKLEBACK SPECIES PAIR IN VANANDA CREEK 


585 


A primary reason for the delayed naming of the 
species is that we have been awaiting good evi- 
dence of whether the species pairs are independent- 
ly evolved or represent a single speciation event 
with subsequent dispersal. Researchers are present- 
ly trying to determine if we are dealing with two 
species, twelve, or an intermediate number. 
Geography and geological history are consistent 
with the hypothesis of independent evolution 
(McPhail 1993; 1994), but the best evidence will be 
a DNA-based phylogeny. 

Preliminary evidence suggests that the species 
pairs in several lakes are indeed independently 
derived (McPhail 1993; Taylor et al. 1997; Taylor 
unpublished data), while others likely represent 
replicate populations. We must therefore name more 
than two species. Species pairs have some opportuni- 
ty for downstream and upstream dispersal to adja- 
cent lakes in the same watershed suggesting the pos- 
sibility of genetic exchange (see Figure 3). Current 
data suggest that the limnetics and benthics in 
Balkwill, Emily and Priest lakes (hereafter the 
Vananda Creek watershed) should be considered 
replicate populations of the same species pair. There 
is a pressing need to protect this species pair due to 
its extreme rarity. 


Taxonomy 

Taxonomic classification of British Columbia 
sticklebacks is highly complex, and presents one of 
the greatest challenges to systemicists of B.C. fish 
fauna. Freshwater populations of sticklebacks are 
invariably distinct from the marine form, yet they 
show both parallel evolution and remarkable pheno- 


FIGURE 2. Limnetic (A) and benthic (B) forms of stickleback, Gasterosteus spp. (drawn by L. Nagel). 


586 


FIGURE 3. Northern portion of Texada Island showing the 
four lakes with Stickleback species pairs. 


typic variation among sites (e.g., Lavin and McPhail 
1985; Schluter and McPhail 1992; McPhail 1994; 
Hagen and Gilbertson 1972; Reimchen et al. 1985; 
Bell and Foster 1994). Classical approaches to sys- 
tematics rely on the measurement of morphological 
traits and assume that traits are never (or at least very 
rarely) evolved in parallel. Because parallel evolu- 
tion is rampant in freshwater sticklebacks their tax- 
onomy has been in chaos for some time (see e.g., 
Hagen and McPhail 1970). It is precisely this mix of 
parallel and independent phenotypic evolution that 
confounds taxonomists while at the same time 
intrigues ecologists and evolutionary biologists. It is 
now generally agreed that freshwater populations of 
sticklebacks have been derived from marine stickle- 
backs multiple times. 

Modern molecular genetic techniques offer the 
most promising approach for resolving taxonomic 
issues in B.C. sticklebacks. When constructing phy- 
logenies, molecular data are used in much the same 
way as morphological data. The major difference is 
that the basic assumption of no parallel evolution is 
much more likely to be valid for molecular traits 
than for morphological traits. 

Studies using molecular approaches to stickleback 
taxonomy are currently underway. Regional patterns 
have been reported in Withler and McPhail (1985) 
and Orti et al. (1994). Finer scale issues (the most 
important issues for conservation biologists) are 
being researched primarily at labs at the University 
of British Columbia. It will be some time before data 
are complete and the issues resolved but early results 
have been reported in Taylor et al. (1997). These and 
other unpublished data suggest that several of the 
stickleback species pairs are genetically distinct 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


units. More data are needed in order to understand 
the relationships among the different pairs and to 
fully understand which of the pairs are independent- 
ly derived from the marine form, and which repre- 
sent replicate populations of the same species. 

In reality the decision to award species status to 
an organism is subjective (McPhail and Carveth 
1992). All of the limnetics and benthics studied to 
date meet the definition of biological species (see 
Mayr 1942, 1963). For example, they maintain their 
morphological differences over several generations 
in a common environment (McPhail 1992; Hatfield 
1995, 1997), they do not interbreed in the lab if 
given a choice among mates (Ridgway and McPhail 
1984; Nagel 1994), and they have remained distinct 
in the wild despite some very large disturbances to 
their habitat (Larson 1976; McPhail 1994). Most 
biologists would not dispute that these are true 
species, albeit extremely young species. 

Molecular data corroborate the view that benthics 
and limnetics are genetically different. However, the 
preliminary molecular genetic data are consistent 
with geographic data and suggest that Balkwill, 
Emily and Priest Lakes are inhabited by replicate 
populations of the same species pair. Together the 
three lakes represent the complete distribution of this 
species pair. 


Distribution 

Stickleback species pairs are currently confined to 
five low elevation coastal lakes on islands in the 
Strait of Georgia, southwestern British Columbia 
(Figures | and 3). A species pair formerly inhabited 
a sixth lake (Hadley Lake) on Lasqueti Island, but 
went extinct in the mid-1990s. Preliminary molecu- 
lar genetic data suggest that limnetics and benthics in 
Balkwill Lake (49° 44’ 90” N, 124° 35'), Emily Lake 
(49° 44’ 80” N, 124° 32' 30”), and Priest Lake 
(49° 44’ 80” N, 124° 33’ 70"), of the Vananda Creek 
watershed, are genetically similar to one another. 
However, they are genetically distinct from species 
pairs found in other lakes. A conservative interpreta- 
tion of these data is that these fish should be consid- 
ered replicate populations of the same species pair. 
There are some differences among the three popula- 
tions, but the differences are relatively small. 
Stickleback species pairs identical to those in the 
Vananda Creek watershed may exist elsewhere, but 
biologists have surveyed hundreds of lakes along the 
British Columbia, Washington, and Alaska coasts for 
these fish, and found them nowhere else. These 
sticklebacks thus appear to be unique British 
Columbia endemics. 


Protection 

There is currently no specific protection for stick- 
lebacks in the Vananda Creek watershed. Much of 
the potential for legislated protection appears to rest 


2001 


on whether they are designated “endangered species” 
and whether proposed legislation gets enacted. The 
Fisheries Act, federal legislation that extends protec- 
tion to fish and fish habitat, was originally written 
for the protection of extractive fisheries (i.e., com- 
mercial, recreational, aboriginal) and it is doubtful 
that it can be successfully applied to sticklebacks in 
the Vananda Creek watershed. 

The Forest Practices Code of British Columbia 
Act provides guidelines for riparian management 
around streams, lakes and fisheries sensitive zones 
on lands managed for forest harvest. However, the 
Code does not apply to private lands, and most lands 
around Balkwill, Emily and Priest lakes are privately 
owned. 

The government of British Columbia has enacted 
the Fish Protection Act, and amended the Wildlife 
Act, and Water Act in an effort to protect fish and 
fish habitat. This legislation may substantially 
enhance the protection of the Vananda Creek water- 
shed sticklebacks if they are designated “endangered 
species”. Once designated the Convention on 
International Trade in Endangered Fauna and Flora 
would also apply. 


Population Sizes and Trends 

Balkwill, Emily and Priest lakes each support 
large populations of both limnetic and benthic stick- 
lebacks that have not been quantified. McPhail 
(1989) estimated population sizes on the order of 
100,000 for each of the stickleback species in Enos 
Lake, which is larger than Emily and Balkwill lakes 
although smaller than Priest Lake. It is possible that 
Priest Lake supports more sticklebacks than Enos 
Lake. 

Stickleback species pairs have been intensively 
studied by zoologists at University of British 
Columbia for the last two decades. However, the 
species pairs within the Vananda Creek watershed 
have received the least attention. Occasional trap- 
ping and study throughout this period suggests that 
populations have remained abundant and easy to trap 
in large numbers in baited Gee traps. This species 
pair should be considered locally abundant and 
under no immediate threat from population decline. 


Habitat 

Balkwill, Emily and Priest lakes are small 
(< 25 ha) coastal lakes typical of this region. Balk- 
will and Priest lakes are approximately 80 m above 
sea level, while Emily Lake is about 40 m in eleva- 
tion. Although some maps show Balkwill and Priest 
lakes joined by a small stream, they are in fact con- 
nected by about 100 m of marsh that is sufficiently 
submerged during high water as to be passable by 
fish. Emily Lake is about 1 km downstream of Priest 
Lake, and connected to the sea by a further 2 km of 
stream. The hydrology of the entire system has been 
substantially modified by small dams, water extrac- 


HATFIELD: STATUS OF STICKLEBACK SPECIES PAIR IN VANANDA CREEK 


587 


tion and land development (housing, road-building, 
mining). 

Riparian areas in the watershed consist of mixed 
second growth forest. The lakes are lightly stained, 
and usually turbid. In summer the littoral region is 
covered in dense beds of Chara, Potamogeton and 
Utricularia. Emergent littoral vegetation is dominat- 
ed by Juncus and Nuphar. There is a distinct pelagic 
zone in each lake, and in the summer plankton pro- 
ductivity is high. All three lakes have a good supply 
of large woody debris in the littoral zone. 


General Biology 
Life History — Limnetics 

Sticklebacks in the Vananda Creek watershed are 
the least-well studied of all the species pairs. 
However, based on the studies that have been under- 
taken (e.g., Schluter and McPhail 1992, 1993; Nagel 
1994; McPhail unpublished data) their life history is 
assumed to be very similar to the other species pairs. 
Much of the following description is therefore based 
on studies carried out on the species pairs found in 
Paxton and Enos lakes. 

The stickleback species pairs in Balkwill, Emily 
and Priest lakes follow the general life history pat- 
tern of most Gasterosteus populations (for reviews 
of general stickleback biology see Wooton 1976; 
Bell and Foster 1994), but some aspects of limnetic 
and benthic life histories are divergent from each 
other. Limnetics can be considered “live fast and die 
young” species, whereas benthics devote consider- 
ably more energy to growth and longevity. 

Limnetics are generally sexually mature after one 
year and rarely live beyond two years. There is con- 
siderable sexual dimorphism: reproducing males 
tend to be bigger on average than gravid females. 
Large male size enables greater nest protection and 
territory defense (Rowland 1989). Fecundity rela- 
tionships for Paxton Lake limnetics are shown in 
Hatfield (1995), with typical fecundity about 30-40 
eggs per clutch. In the lab, females produce several 
clutches per season, usually in close succession if 
food availability is high (personal observation). 
Female life history is likely similar in the wild. 

As with other sticklebacks, limnetic males are the 
sole providers of postzygotic parental care. In the 
spring, they acquire territories in the littoral region 
where they build nests and mate (sometimes with 
many females). Limnetics prefer unvegetated, open 
nesting locations (McPhail 1994; Hatfield and 
Schluter 1996). They often nest in less than 1 m of 
water on submerged logs, in shallow bays with grav- 
el or rocky substrates, and on firm muddy substrate. 
Because preferred spawning habitat is not uniformly 
distributed in the littoral zone, nesting males are 
clumped in their distribution. Despite the fact that 
limnetics and benthics breed at the same time of year 
they rarely interbreed (McPhail 1994). 

Following fertilization, eggs take approximately 


588 


7-10 days to hatch, depending on temperature. 
During this time males actively aerate the eggs by 
thrusts of their pectoral fins; embryos die if inade- 
quately aerated (van den Assem 1967; Sargent and 
Gebler 1980). Male sticklebacks vigorously defend 
their nests and territories from invaders (most often 
other sticklebacks) and continue to defend their 
young for about a week after they hatch. The young 
then disperse into the littoral vegetation where they 
feed under cover. By late summer limnetics have 
become large and swift enough to escape predators, 
and their spines have grown big enough to act as a 
deterrent. At this time they school up and forage for 
plankton in the open water. 


Life History — Benthics 

As with the above section on limnetics, much of 
the following description of benthic life history is 
based on studies carried out on the species pairs 
found in Paxton and Enos lakes. Benthic stickle- 
backs in the Vananda Creek watershed have a gener- 
al life history pattern consistent with most Gaster- 
osteus populations. However, benthics are divergent 
from limnetics. On average, benthics live longer and 
reproduce less often than limnetics. Determining the 
age of sticklebacks is difficult, but benthics do not 
seem to become sexually mature after one year, and 
appear to live well beyond two years, perhaps as 
long as seven years (McPhail, personal communica- 
tion). There is little or no sexual dimorphism in ben- 
thics, if present it tends to be in the opposite direc- 
tion from the limnetics: reproductive males tend to 
be smaller on average than gravid females. 
Fecundity relationships for Paxton Lake benthics are 
shown in Hatfield (1995); but are likely representa- 
tive of benthics from other lakes. In the lab, females 
produce only one or two clutches per season, regard- 
less of food availability (personal observation). 
Females probably have a similar life history in the 
wild. 

Benthics prefer densely vegetated nesting loca- 
tions, usually among beds of Chara (McPhail 1994; 
Hatfield and Schluter 1996). Their nests are highly 
concealed and difficult to find in the field. They tend 
to nest in water of greater depth than limnetics, 
though usually less than 2m depth. As with limnet- 
ics, preferred spawning habitat for benthics is not 
uniformly distributed so nesting benthic males are 
clumped in their distribution. 

Benthics are similar to limnetics in all aspects of 
parental care and development. About a week after 
they hatch, the young disperse into the littoral vege- 
tation where they feed. Juvenile benthics continue to 
feed in the shallow littoral zone under cover of, or 
within close proximity to vegetation cover. 


Diet 
Diets of limnetics and benthics have been well 


studied for later life history stages during spring 
summer and early fall (Schluter and McPhail 1992; 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Schluter 1995). Very little is known about diets dur- 
ing the initial life stages of the two species, or what 
all life stages eat during late fall and winter months. 

As adults, limnetic and benthic sticklebacks eat 
quite different foods. Limnetics feed primarily in the 
surface waters away from the lake margins. There 
they hunt in loose schools for copepods, cladocera 
and insect larvae. Males will often forage for ben- 
thos when nesting in the littoral zone. As young 
juveniles, limnetics feed at the lake edges among 
reeds and submerged plants where they can seek 
cover if approached by a potential predator. 

Benthics on the other hand forage along the shal- 
low margins of the lake for larger prey such as 
snails, clams, dragonfly nymphs, amphipods, and 
chironomids. These invertebrates are found among a 
variety of substrates including plants, rocks or mud. 
Benthics likely eat similar food types throughout 
their life, but gradually shift to larger prey as they 
get bigger themselves. 


Species Movement 

There are three movement trends of note within 
other lakes with species pairs that are assumed to 
reflect movements within Balkwill, Emily and Priest 
lakes. In early to mid fall most individuals are found 
in deeper waters. where they remain during the win- 
ter months. During the spring spawning months 
adults of both species are abundant in the shallow lit- 
toral zone; however limnetic females tend to forage 
in the surface pelagic zone and move into the littoral 
zone to seek nesting males. During the summer 
months limnetics are found in the pelagic zone and 
benthics are found in the shallow littoral zone. These 
movements are minor, and are unlikely to have sig- 
nificant bearing on management options for the 
species, at least at present. 


Adaptability 

Historically the Vananda Creek watershed has 
been impacted by significant human disturbance, 
including damming the outlet stream draining Priest 
Lake, water extraction for mining and residential 
use, and land clearing associated with mining, forest 
harvest and human settlement. Limnetics and ben- 
thics within each of the lakes have been tolerant of 
these disturbances. Water extraction has likely been 
the most significant disturbance — lake level fluctu- 
ations can affect water quality, spawning habitat, 
predation, food resources, and cover. In reality, we 
know little about the response of the sticklebacks to 
these and other disturbances except that they were 
resilient enough to persist. 

Introduction of non-native species likely presents 
the most significant immediate threat to the species 
pair in Balkwill, Emily and Priest lakes. The only 
disturbance known to have led to the extinction of a 
species pair is the unauthorized introduction of cat- 
fish (Ameirus nebulosus) to Hadley Lake on Las- 
queti Island. In that case extinction was swift. 


2001 


Limiting Factors 

Current limits to stickleback abundance in 
Balkwill, Emily and Priest Lakes are poorly under- 
stood. For example, it is not known whether abun- 
dance is limited by rearing habitat, food production, 
cover, predation, spawning habitat or other factors. 
Currently limnetics and benthics are locally very 
abundant in each lake, and are not apparently in 
decline. 

The lakes are inhabited by numerous invertebrates 
that feed on young sticklebacks, and are regularly 
visited by piscivorous birds (e.g., herons (Ardea 
herodias), kingfishers (Megaceryle alcyon) and loons 
(Gavia immer)). Cutthroat Trout (Oncorhynchus 
clarki) also live in the lakes and are pretatory. 
However, their presence is not a threat to the stickle- 
backs. 

Based on anecdotal evidence it seems unlikely 
that the abundance of sticklebacks in these lakes is 
limited by spawning habitat. Both species have pre- 
ferred spawning habitat types within the littoral zone 
(McPhail 1994; Hatfield and Schluter 1996), but dur- 
ing the spawning season nesting males are very 
abundant, and nesting success and recruitment 
appear to be high (personal observation). 

The primary limiting factor at present is most 
likely food supply, the capability of the lake to pro- 
duce plankton and benthos. Balkwill, Emily and 
Priest lakes are very productive and therefore can 
and do support large numbers of sticklebacks. 


Special Significance of the Species 

The special significance of the Balkwill, Emily 
and Priest lakes species pair is primarily of aesthetic 
and scientific value. It is very unlikely that the 
species will be of commercial use and value. 

Stickleback species pairs are widely regarded as a 
scientific treasure, in large part because they are 
among the youngest species on earth. Scientists 
believe they have evolved since the end of the last 
glaciation, approximately 13 000 years ago. The pro- 
duction of a new species usually takes in the order of 
millions of years. The speed with which these dis- 
tinct fish species evolved has intrigued and excited 
scientists from around the world. They are a remark- 
able research subject that will help us understand the 
biological and physical processes that have given us 
the tremendous diversity of organisms we see around 
us. Newspapers, magazines and scientific journals 
have published the story of the discovery of these 
species, and have followed the results of ongoing 
scientific studies. 


Evaluation 

Limnetic and benthic sticklebacks in Balkwill, 
Emily and Priest Lakes are among the most rare and 
endangered species in the world, and should be des- 
ignated endangered species (i.e., Red Listed) by 
COSEWIC. Unlike many species that are rare in 


HATFIELD: STATUS OF STICKLEBACK SPECIES PAIR IN VANANDA CREEK 


589 


Canada but found elsewhere, stickleback species 
pairs exist nowhere else. Their entire worldwide dis- 
tribution is highly restricted, with no potential for 
natural dispersal to suitable habitat elsewhere. All of 
the species pairs occur only in a fast-growing region 
of southwestern British Columbia. Although the 
lakes are in relatively undeveloped areas of the 
province, they have already been affected by human 
activities such as logging, land clearing, water 
removal, road building, and septic tank inputs. The 
sudden extinction of the Hadley Lake species pair 
emphasizes the extreme susceptibility of these fish to 
introductions of exotic species. 

The British Columbia Conservation Data Centre 
has placed stickleback species pairs, including the 
species pair in Balkwill, Emily and Priest Lakes, in 
the highest risk category, Category 1: “critically 
imperiled because of extreme rarity”. The species 
have been placed on the provincial Red List as 
threatened or endangered. For the reasons noted 
above these designations are justified. 


Acknowledgments 

Preparation of the report was funded by the 
British Columbia Habitat Conservation Trust Fund. 
The project was coordinated by Juanita Ptolemy, 
Fisheries Branch, Ministry of Environment, Lands 
and Parks, Victoria, British Columbia. 


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Fluid dynamics in suspension-feeding blackfish. Science 
251: 1346-1348. 

Sargent, R. C., and J.B. Gebler. 1980. Effects of nest 
site concealment on hatching success, reproductive suc- 
cess, and paternal behavior of the threespine stickleback, 
Gasterosteus aculeatus. Behavioral Ecology and Socio- 
biology 7: 137-142. 

Schluter, D. 1995. Adaptive radiation in sticklebacks: 
trade-offs in feeding performance and growth. Ecology 
76: 82-90. 

Schluter, D., and J. D. McPhail. 1992. Ecological char- 
acter displacement and speciation in sticklebacks. 
American Naturalist 140: 85-108. 

Schluter, D., and J. D. McPhail. 1993. Character dis- 
placement and replicate adaptive radiation. Trends in 
Ecology and Evolution 8: 197-200. 

Taylor, E. B., J.D. McPhail, and D. Schluter. 1997. 
History of ecological selection in sticklebacks: uniting 
experimental and phylogenetic approaches. Pages 
511-534 in Molecular evolution and adaptive radiation. 
Edited by T.J. Givnish and K. J. Sytsma. Cambridge 
University Press, Cambridge, UK. 

van den Assem, J. 1967. Territory in the three-spined 
stickleback Gasterosteus aculeatus (L.). Behaviour 
Supplement 16: 1-164. . 

Withler, R. E., and J. D. McPhail. 1985. Genetic vari- 
ability in freshwater and anadromous sticklebacks 
(Gasterosteus aculeatus) of southern British Columbia. 
Canadian Journal of Zoology 63: 528-533. 

Wooton, R. J. 1976. The biology of the sticklebacks. 
Academic Press, London, UK. 


Accepted 19 February 2002 


Status of the Stickleback Species Pair, Gasterosteus spp., 
in Paxton Lake, Texada Island, British Columbia* 


TopD HATFIELD! and JUANITA PTOLEMY2 


'Solander Ecological Research, 1324 Franklin Terrace, Victoria, British Columbia V8S 1C7 Canada 
2B.C. Ministry of Fisheries, Victoria, British Columbia V8W 9M1 Canada 


Hatfield, Todd, and Juanita Ptolemy. 2001. Status of the stickleback species pair, Gasterosteus spp., in Paxton Lake, 
Texada Island, British Columbia. Canadian Field-Naturalist 115(4): 591-596. 


A pair of threespine stickleback species (Gasterosteus spp.) coexists in Paxton Lake on Texada Island, British Columbia. 
One species (referred to as “limnetic”) forages on pelagic zooplankton, and the other (referred to as “benthic”) feeds on lit- 
toral zone benthic invertebrates. The morphology of each species reflects divergent resource use strategies. For example, 
benthics are larger, have wider mouths and fewer, shorter gill rakers than limnetics. Benthics also have considerably less 
armour than limnetics. Current data suggest Paxton Lake limnetics and benthics are unique biological species. Although 
both species are numerous, their entire distribution is restricted to this single lake. The immediate threat to these stickle- 
back species is the unauthorized introduction of a non-native predator such as catfish (Ameirus nebulosus) or Pumpkinseed 


(Lepomis gibbosus). 


Key Words: Gasterosteidae, sticklebacks, Gasterosteus, épinoches, Paxton, endangered species 


Threespine sticklebacks (Gasterosteus aculeatus 
complex) form a taxonomically difficult group com- 
posed of thousands of phenotypically diverse popu- 
lations. They are abundant, small fish (usually 
35—55 mm) found in coastal marine and fresh water 
throughout the Northern Hemisphere. Marine stick- 
lebacks are phenotypically similar throughout their 
range, whereas freshwater sticklebacks are ecologi- 
cally, behaviourally, and morphologically variable 
(e.g., Lavin and McPhail 1985; Schluter and 
McPhail 1992; McPhail 1994; Hagen and Gilbertson 
1972; Reimchen et al. 1985; Bell and Foster 1994). 
In general, G. aculeatus has a laterally compressed 
body, well armoured by calcified lateral plates and 
retractable pelvic and dorsal spines, with delicate 
pectoral and caudal fins. Freshwater populations are 
highly variable in extent of armour but generally 
have fewer lateral plates than the marine form. Body 
colour also varies considerably, from silvery to mot- 
tled green and brown. Usually sexually mature males 
develop bright red throats, although in a few fresh- 
water populations males turn completely black 
(McPhail 1969; Reimchen 1989). 

The marine form is the most recent ancestor to 
most freshwater forms (Bell and Foster 1994). 
Sticklebacks did not colonize fresh water in one 
single event followed by a range expansion; they 
invaded fresh water repeatedly following deglacia- 
tion. This suggests that the tremendous variability 
currently seen in British Columbia’s freshwater 


*Reviewed and approved by COSEWIC April 1999, status 
assigned — threatened. 


Gasterosteus has arisen in the last 10 000 to 13 000 
years. 

Many of the hundreds of small, low elevation, 
coastal lakes found in British Columbia have been 
surveyed for sticklebacks (e.g., Lavin and McPhail 
1985; Reimchen et al. 1985; Schluter and McPhail 
1992; McPhail 1993). Most contain a single, variable, 
form of stickleback (see e.g., Bell 1976; Lavin and 
McPhail 1985; Riemchen et al. 1985; Schluter and 
McPhail 1992). Six lakes found on islands in the 
Strait of Georgia (see Figure 1) are especially note- 
worthy. Two distinct species evolved in each of these 
lakes (McPhail 1984, 1992, 1993, 1994; Schluter and 
McPhail 1992). The pattern of morphological and 
ecological divergence shown by the sticklebacks is 
remarkably similar in each of the lakes (Schluter and 
McPhail 1992). In each case, one of the species 
(referred to as “limnetic”’) displays morphological 
traits (such as a fusiform body, narrow mouth and 
many, long gill rakers) best suited for foraging on 
zooplankton (Magnuson and Heitz 1971; Kliewer 
1970; Sanderson et al. 1991; Schluter and McPhail 
1992, 1993). The other species (referred to as “benth- 
ic’) exploits benthic invertebrates in the littoral zone. 
It has a robust body form, wide gape and a few, short 
gill rakers, traits considered advantageous for benthic 
feeding (Schluter and McPhail 1992, 1993). 

These “species pairs” have been discovered in 
Paxton, Priest, Balkwill and Emily lakes located on 
Texada Island, Hadley Lake on Lasqueti Island, and 
Enos Lake on Vancouver Island (Figures 2 and 3). 
The pair that occurred in Hadley Lake is now con- 
sidered extinct. In each of the remaining occurrences 
limnetics and benthics co-exist and are reproductive- 
ly isolated by behaviour and genetic differences 


591 


é Vancouver 


stand Sepa 
A aes 


125 124 123 


FIGURE |. Map of the Strait of Georgia, British Columbia. 
Indicated are the major lakes in each of the four 
watersheds with species pairs. 


(McPhail 1984, 1992; Ridgway and McPhail 1984; 
Nagel 1994; Hatfield 1995, 1997; Hatfield and 
Schluter 1996) meeting a conservative definition of 
species (Mayr 1942, 1963). Species description and 
naming have been delayed until completion of a 
DNA-based phylogeny. Preliminary genetic evi- 
dence suggests that the species pairs in several lakes 
are independently derived (McPhail 1993; Taylor et 
al. 1997, Taylor unpublished data). 


BRITISH COLUMBIA 
MAINLAND. 2 


TEXADA \ eee 
ISLAND ,-~ X 


‘FiGuRE 2. Northern portion of Texada Island showing the 
four lakes with Stickleback species pairs. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


This report documents the current status of the 
species pair found in Paxton Lake. Reports on the 
current status of stickleback species found in Hadley 
Lake, and in Balkwill, Emily and Priest lakes have 
also been prepared. A status report on the Enos Lake 
species pair was completed earlier (McPhail 1989) 
and should be updated. 


Distribution 

Paxton Lake (49° 42’ 30” N, 124° 31’ 30” W) 
shares a watershed with two smaller lakes, Myrtle 
and Case lakes, as well as Rumbottle Creek (the out- 
let of Paxton). Only Paxton Lake contains a species 
pair. Based on surveys of hundreds of lakes along 
the coast of Alaska, British Columbia, and Wash- 
ington, both Paxton Lake species appear to be 
unique British Columbia endemics. 


Protection 

Currently, there is no specific protection for 
Paxton Lake sticklebacks. The provisions of the fed- 
eral Fisheries Act, originally intended to protect 
extractive fisheries, do not apply to the protection of 
fish that do not contribute to a fishery. The govern- 
ment of British Columbia has enacted the Fish 
Protection Act containing consequential amendments 
to the Wildlife Act designed to provide some protec- 
tion for species designated under the Act. The 
amendments extend provisions for endangered and 
threatened wildlife to fish, and to aquatic inverte- 
brates and plants that are a factor in fish habitat. For 
example, causing lasting harm to, or intentionally 
causing harm to endangered or threatened species 
would become an offence. Once designated the 
Convention on International Trade in Endangered 
Species of Wild Fauna and Flora could also be 
applied, if it became necessary. 


Population Size(s) and Trends 

Paxton Lake supports unquantified but large pop- 
ulations of both limnetic and benthic sticklebacks. It 
is roughly the size of Enos Lake where McPhail 
(1989) estimated population sizes on the order of 
100,000 per species. Current abundance in Paxton 
may be higher than historic levels as a dam on the 
outlet has prevented reproduction of piscivorous 
native Cutthroat Trout (Oncorhynchus clarki). Over 
the last decade both sticklebacks have remained 
abundant and easy to trap in large numbers, indicat- 
ing the populations are currently stable. Severe 
declines may have occurred historically. 


Habitat 
General Description 

Paxton Lake is a small (17 ha) coastal lake with a 
maximum depth of about 15 m. The lake has distinct 
littoral and pelagic zones. It lies about 90 m above 
sea level and is connected to the sea by roughly 


2001 


HATFIELD AND PTOLEMY: STATUS OF STICKLEBACK PAIR IN PAXTON LAKE 


593 


-_ 
a 
ss 
a 


gas 


FiGuRE 3. Limnetic (A) and benthic (B) forms of stickleback, Gasterosteus spp., (drawn by L. Nagel). 


4.5 km of stream. There is no permanent inlet stream 
to the lake. Rumbottle Creek was dammed in 1956 to 
provide a source of water for a nearby mine. Prior to 
dam construction Paxton Lake was actually two sep- 
arate lakes, joined by about 100 m of stream. Raising 
the water level by 1.5 m combined the two lakes into 
one with a narrow waist and two fairly equal basins 
(McPhail 1992). 

Rumbottle Creek drops about 80 m, in a series of 
small falls, over the last 2km of its length before 
entering Malaspina Strait. The falls effectively iso- 
late the upper portions of the creek and the lake from 
the sea. Much of its length has been affected by 
development (housing, road-building, culverts, etc.) 
and logging. 

Paxton Lake lies in limestone overlain by post- 
glacial marine sediments. Substrate at the south end 
is predominantly marl (Larson 1976). Riparian areas 
consist of mixed second growth forest. Lake water is 
lightly stained. Summer conditions described by 
Larson (1976) indicate surface water temperatures 
can reach 23°C, a thermocline develops between 2 
to 5m. The littoral region becomes covered in dense 
beds of Chara, Potamogeton and Utricularia. 
Emergent littoral vegetation is dominated by Juncus 
and Nuphar. Plankton productivity is high. 


Habitat Use 

Limnetics feed primarily in the pelagic zone. In 
the spring males move inshore and select unvegetat- 
ed, open nest sites (McPhail 1994; Hatfield and 
Schluter 1996). They often nest in less than 1 m of 
water on submerged logs, in shallow bays with grav- 
el or rocky substrates, or on firm muddy substrate. 
Patchy distribution of preferred spawning habitats 
may lead to clustering of nest sites. 


Benthics feed over open mud bottom or sub- 
merged aquatic vegetation. Males build their nests in 
dense vegetation, usually in beds of Chara (McPhail 
1994; Hatfield and Schluter 1996). Their nests are 
highly concealed and difficult to find. They tend to 
nest in water of greater depth than limnetics, though 
usually less than 2 m. Spawning males congregate in 
areas of preferred habitat. 

About a week after hatching, the young of both 
species disperse into the littoral vegetation where 
they feed under cover. Juvenile benthics continue to 
feed in the shallow littoral zone or within close prox- 
imity to vegetation. By late summer the limnetics 
school up and move into the open water since they 
have the speed and spines to escape predators. 

By late fall most individuals of both species have 
moved to deeper water where they spend the colder 
months. 


General Biology 
Life History 

Both Paxton Lake stickleback species follow the 
general life history pattern of most Gasterosteus 
populations (for reviews of general stickleback biol- 
ogy see Wooton 1976; Bell and Foster 1994). 
Although limnetics and benthics breed at the same 
time of year (April-June), they rarely interbreed 
(McPhail 1992). As with other sticklebacks, males 
are the sole providers of postzygotic parental care. In 
the spring, males acquire territories in the littoral 
region where they build nests and mate (sometimes 
with many females). Following fertilization, eggs 
take approximately 7-10 days to hatch, depending 
on water temperature. During this time males active- 
ly aerate the eggs by thrusts of their pectoral fins; 
embryos die if inadequately aerated (van den Assem 


594 


1967; Sargent and Gebler 1980). Male sticklebacks 
vigorously defend their nests and territories from 
invaders (often other sticklebacks) and continue to 
defend their young for about a week after they hatch. 
In other aspects limnetic and benthic life histories 
are divergent from each other. Limnetics are rela- 
tively short lived, whereas benthics devote consider- 
ably more energy to growth and longevity. Limnetics 
are generally sexually mature after one year and 
rarely live beyond two years. There is considerable 
sexual dimorphism; reproducing males tend to be 
bigger on average than gravid females. Fecundity 
relationships are shown in Hatfield (1995), but typi- 
cal fecundity is about 30-40 eggs per clutch for a 
limnetic female. In the lab, females produce several 
clutches per season, usually in close succession if 
food availability is high (Hatfield, personal observa- 
tion). 

Benthics generally live longer and reproduce less 
frequently than limnetics. Determining the age of 
sticklebacks is difficult, but Paxton Lake benthics do 
not seem to mature until their second year, and may 
live as long as seven years (McPhail, personal com- 
munication). There is little or no sexual dimorphism 
in benthics, if present it tends to be in the opposite 
direction from the limnetics (i.e.,reproductive males 
may be slightly smaller than gravid females). 
Fecundity relationships for benthics are shown in 
Hatfield (1995); females often carry more than 150 
eggs. In the lab, females produce only one or two 
clutches per season, regardless of food availability 
(Hatfield, personal observation). 


Diet 

Diets of both species have been well studied dur- 
ing the spring, summer, and early fall for later life 
history stages (Schluter and McPhail 1992; Schluter 
1995). Little is known about diet during early life 
stages, or the late fall and winter months. 

Adult limnetic and benthic sticklebacks consume 
quite different foods. Limnetics feed primarily in the 
surface waters away from the lake margins. They 
hunt in loose schools for copepods, Daphnia and 
insect larvae. Males forage for benthos while nesting 
and often consume eggs from the nests of other 
males. 

Benthics forage along the shallow margins of the 
lake for larger prey such as snails, clams, dragonfly 
nymphs, amphipods, and chironomids. These inver- 
tebrates are found among a variety of substrates 
including plants, rocks or mud. Benthics likely eat 
similar items throughout their life, selecting larger 
sized prey as they grow. 


Limiting Factors 

Limits to Paxton Lake stickleback abundance are 
- poorly understood. Piscivorous cutthroat trout are no 
longer a consideration. Other predators include 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


numerous invertebrates that feed on young stickle- 
backs, and piscivorous birds [e.g., herons (Ardea 
herodias), kingfishers (Megaceryle alcyon) and 
loons (Gavia immer)]. Their presence does not 
appear to be a significant factor. Each species has 
different spawning habitat preferences (McPhail 
1994; Hatfield and Schluter 1996). However, recruit- 
ment does not seem to be restricting population num- 
bers as nesting males are very abundant, and nesting 
success appears high (Hatfield, personal observa- 
tion). Food supply may be the current limiting factor. 


Threats 

Historically Paxton Lake has been subjected to 
tremendous human disturbance. Although Paxton 
Lake limnetics and benthics have survived, each of 
the disturbances undoubtedly had an influence on 
their numbers and relative abundance. Water extrac- 
tion for a nearby mine and the introduction of 
salmonids likely had the most profound effects, and 
may have led to considerable hybridization between 
the two species (McPhail, personal communication). 
Fluctuations in water level would affect water quali- 
ty, spawning habitat, predation, food resources, and 
cover. Water removal ceased following closure of 
the mine but could recur. The introduction of large 
numbers of Coho Salmon (Oncorhynchus kisutch) 
may have had a significant effect on food resources 
and predation. Provincial fisheries managers are now 
aware of the presence of the sticklebacks and will 
not authorize further fish introductions. 

However, the unauthorized release of non-native 
fish species in the watershed is a significant immedi- 
ate threat to both species. The species of most con- 
cern are catfish (Ameirus nebulosus) or Pumpkin- 
seeds (Lepomis gibbosus). Both exotics appeared on 
Vancouver Island and the mainland in the early 
1900s, and continue to spread through unauthorized 
public transplants. Catfish are nocturnal nest preda- 
tors; pumpkinseeds act as predators and competitors 
at different life history stages. The only disturbance 
known to have led to the extinction of a stickleback 
species pair was the release of catfish (Ameirus neb- 
ulosus) to Hadley Lake on Lasqueti Island, immedi- 
ately west of Texada Island. In that case extinction 
was swift. Populations of both exotic species have 
become established in close proximity to Texada 
Island. 


Special Significance of the Species 

Paxton Lake sticklebacks are significant for aes- 
thetic and scientific reasons. Stickleback species 
pairs are widely regarded as scientific treasures. 
Scientists believe they have evolved since the end of 
the last glaciation, approximately 13 000 years ago, 
making them some the youngest species on the plan- 
et. The speed with which these species pairs evolved 
has intrigued and excited scientists around the world. 
They are remarkable subjects for the study of specia- 


2001 


tion and evolution. The public has also followed sto- 
ries about the species pairs in the popular press and 
on television. 


Evaluation 

The sudden extinction of the Hadley Lake species 
pair emphasizes the extreme susceptibility of these 
fish to introductions of exotic species. Unlike many 
species that are rare in Canada, the Paxton Lake 
stickleback species occur nowhere else. There is no 
potential for natural dispersal to suitable habitat else- 
where. They occur in a region of southwestern 
British Columbia experiencing rapid urbanization. 
Although the lake lies in a relatively undeveloped 
area, the drainage has been affected by various 
human activities including mining, logging, land 
clearing, water removal, road building, septic efflu- 
ent, and introduction of non-native species. 

The British Columbia Conservation Data Centre 
has ranked both Paxton Lake sticklebacks as G1/S1 
(critically imperiled because of extreme rarity), 
using the system developed by The Nature 
Conservancy. These species have also been placed 
on the provincial Red List, the highest risk category. 


Acknowledgments 

Funding for the status report was provided by the 
British Columbia Habitat Conservation Trust Fund, 
World Wildlife Fund (Canada), and the Department 
of Fisheries and Oceans. The compilation and sum- 
mary provided by J. Houston was most helpful. 


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THE CANADIAN FIELD-NATURALIST 


Vol. 115 


History of ecological selection in sticklebacks: uniting 
experimental and phylogenetic approaches. Pages 
511-534 in Molecular evolution and adaptive radiation. 
Edited by T. J. Givnish and K. J. Sytsma. Cambridge 
University Press, Cambridge, UK. 

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Academic Press, London, UK. 


Received 19 February 2002 


Rapport sur la Situation du Brochet d’ Amérique, 
Esoex americanus americanus, au Canada* 


STEPHANIE LACHANCE 


Société de la faune et des parcs du Québec, Direction de !a recherche sur la faune, 675 boulevard René-Lévesque Est, Boite 
92, Québec, Québec GIR 5V7 Canada 


Lachance, Stephanie. 2001. Rapport sur la situation du brochet d’amérique, Esox americanus americanus, au Canada. 
Canadian Field-Naturalist 115(4): 597-607. 


In Canada, the Redfin Pickerel (Esox americanus americanus), is restricted to Quebec. The first Canadian record of Redfin 
Pickerel dates back to 1944. Its distribution is limited to the St. Lawrence River, between the Verchéres Islands and the 
mouth of the Godefroy River; Lake St. Paul; and the drainages of the Richelieu River, including Lake Champlain, the St. 
Francois River and the Yamaska River. The abundance of the Redfin Pickerel is difficult to assess because it is able to 
escape from several types of gear commonly used in fish surveys. Further, it prefers shallow grass beds, in which intensive 
inventories are seldom conducted. The only well-established populations seem to be those of the Upper Richelieu River 
and the Berthier Islands, where its abundance is stable. The primary threats to the survival of the Redfin Pickerel are 
believed to be dyke construction on farmland to prevent spring flooding and the drainage of grass beds. This subspecies has 
not been given any special protective status and is not the subject of specific regulations. There are, however, a few pro- 
tected areas within its range, and the Quebec Act on the Conservation and Development of Wildlife protects fish habitat 
located on public land. The Redfin Pickerel has no commercial value. 


Au Canada, la sous-espéce de la famille des Esocidés connue sous le nom de brochet d’ Amérique (Esox americanus ameri- 
canus) est exclusive au Québec. La premiére mention du brochet d’ Amérique au Canada date de 1944. Son aire de réparti- 
tion est limitée au fleuve Saint-Laurent entre les iles de Verchéres et l’embouchure de la riviere Godefroy, au lac Saint- 
Paul, ainsi qu’aux réseaux hydrographiques des rivieres Richelieu, incluant le lac Champlain, Saint-Frangois, et Yamaska. 
L’abondance du brochet d’ Amérique est difficile 4a évaluer puisqu’il échappe a plusieurs engins de péche couramment util- 
isés lors des inventaires ichtyologiques. De plus, il préfére les herbiers de faibles profondeurs qui ne font pas toujours 
V objet d’inventaires trés intensifs. Les seules populations bien établies semblent étre celles du Haut-Richelieu et des iles de 
Berthier, ot: l’abondance serait stable. Les principales menaces 4 la survie du brochet d’ Amérique seraient |’endiguement 
des terres agricoles pour prévenir |’inondation printaniére et l’assechement des herbiers. Cette sous-espéce ne jouit pas de 
protection particuliére et ne fait pas l’objet de réglementation spécifique. Il existe quelques territoires protégés sur son aire 
de répartition, et la Loi sur la conservation et la mise en valeur de la faune du Québec protége I’habitat du poisson sur les 
territoires publics ot elle est présente. Le brochet d’ Amérique n’a pas de valeur économique. 


Mots clés: brochet, brochet d’ Amerique, Redfin Pickerel, Esox americanus americanus, Québec, Esocidae espéces en péril. 


Le présent rapport vise a établir le statut du brochet 
d’Amérique (Esox americanus americanus Gmelin 
1788) au Canada. Cette sous-espéce a été placée sur la 
liste des espéces susceptibles d’étre désignées men- 
acées ou vulnérables par le ministére de |’ Environ- 
nement et de la Faune du Québec (Beaulieu 1992). 
Les raisons qui ont justifié cette classification sont les 
suivantes : elle est a la limite de son aire de réparti- 
tion, elle est rare, et 11 existait un doute sur son déclin 
possible. Elle fait aussi partie de la liste des espéces 
prioritaires dans le cadre du Plan Saint-Laurent Vision 
2000. 

Les principaux documents sur lesquels se base le 
présent rapport sur la situation du brochet 
d’Amérique sont le mémoire de maitrise de M™« 
Michéle Lapointe (1980), les inventaires ichty- 
ologiques effectués, depuis le début des années 


*Reviewed and approved by COSEWIC April 1998., status 


assigned — Not At Risk. 


1960, par le Service de l’aménagement et de 
V’exploitation de la faune du ministére de |’ Environ- 
nement et de la Faune des régions de Montréal, de 
l’Estrie et de Trois-Riviéres, ainsi que sur les infor- 
mations contenues dans l’ouvrage de Scott et 
Crossman (1974). 


Classification et nomenclature 

Le brochet d’ Amérique appartient a la famille des 
Esocidés qui ne comprend qu’un seul genre, Esox. 
Au Canada il n’existe que quatre especes de bro- 
chets, le grand brochet, Esox lucius, le maskinongé, 
E. masquinongy, le brochet maillé, E. niger, et enfin 
E. americanus, qui présente deux sous-espéces, soit 
le brochet d’ Amérique, E. americanus americanus, 
et le brochet vermiculé, E. americanus vermiculatus 
(Scott et Crossman 1974). Historiquement, le bro- 
chet d’Amérique a été considéré tour a tour comme 
une espéce distincte du brochet vermiculé, et comme 
la forme typique dont le brochet vermiculé est une 
sous-espéce. Aujourd’hui, les deux formes sont con- 


597 


598 


sidérées comme des sous-espéces (Scott et Crossman 
1974). Les populations du sud des Etats-unis seraient 
intermédiaires entre les deux sous-espéces, c’est-a- 
dire qu’il y a hybridation (Reist et Crossman 1987; 
Scott et Crossman 1974; Crossman 1966). Il peut 
aussi y avoir hybridation avec le brochet mailié et le 
grand brochet, mais dans ce dernier cas les hybrides 
sont stériles (Scott et Crossman 1974). 

Le nom de brochet des marais est parfois utilisé 
pour désigner le brochet d’ Amérique (Mongeau et al. 
1974). Le nom anglais de ce poisson, Red-finned 
Pickerel, fait référence 4 sa taille, pickerel signifiant 
petit brochet « little pike » et a la couleur rouge- 
orangée de ses nageoires inférieures (Scott et Cross- 
man 1974). 

Le brochet d’Amérique est arrivé récemment au 
Canada. La premiére mention date de 1944 (Cuerrier 
1947). Il semble qu’il aurait pénétré en eaux canadi- 
ennes par la voie de communication artificielle con- 
struite en 1819 entre la riviére Hudson et le régime 
du lac Champlain-riviére Richelieu (Scott et 
Crossman 1974). 


Description 

Les informations suivantes sont tirées de Scott et 
Crossman (1974). Le brochet d’ Amérique présente un 
corps de forme allongée, plutdt cylindrique, et aplati 
dorsalement devant la nageoire dorsale. Les adultes 
dépassent rarement la longueur de 300 mm (Figure 1). 
La téte est modérément longue, plate, nue sur le 
dessus et large. Les joues et les opercules sont entiére- 
ment recouverts d’écailles. Le museau est court, large 
et convexe entre |’oeil et le bout. La bouche est 
grande et horizontale, et la machoire inférieure est 
légérement plus longue que le museau. Les nageoires 
sont arrondies. Il n’y a qu’une seule nageoire dorsale, 
a rayons mous, placée loin derriére. La nageoire cau- 
dale est fourchue. La nageoire anale est placée juste 
sous la nageoire dorsale. Les nageoires pelviennes 
sont abdominales, au centre du corps. Enfin, les 
nageoires pectorales sont placées sous le volet opercu- 
laire. Les nageoires inférieures; pectorales, pelviennes 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


et anale présentent une couleur allant du rouge a 
l’ orange. 

Les brochetons d’une longueur de 50 mm présen- 
tent une coloration plut6t uniforme d’un brun foncé. 
A partir de 100 mm de longueur, les prolongements 
verticaux d’une bande latérale pale commencent a 
séparer les flancs en barres verticales sombres. Le 
patron de coloration des adultes fait son apparition 
alors que le brochet aiteint une tailie entre 100 mm et 
150 mm. Alors la face dorsale du corps et de la téte, 
ainsi que la partie supérieure des flancs vont du brun 
pale au vert olive. Il existe une faible bande pale sur 
le milieu du dos, de la nuque a l’origine de la nage- 
oire dorsale. Les flancs sont marqués de 20 a 36 bar- 
res verticales ondulées, allant du vert olive au brun 
foncé. Les espaces clairs entre les barres sont plus 
étroits que celles-ci. La face ventrale du brochet varie 
de l’ambre pale au blanc laiteux. La téte est marquée 
de deux barres noires, l’une sous-orbitaire, |’ autre 
postorbitaire. La surface inférieure du mandibule est 
pigmentée et ce, de fagon plus importante chez les 
femelles. La pupille de I’ oe11 varie du jaune au vert. 

Le brochet d’ Amérique se distingue du grand bro- 
chet et du maskinongé, par un moins grand nombre 
de pores sous-mandibulaires (ordinairement 7 ou 8). 
Il présente moins de rayons branchiostéges que le 
brochet maillé, 11 a 13 contre 14 a 17. Enfin, on le 
sépare de la sous-espéce du brochet vermiculé, par 
son museau court a profil supérieur convexe, un plus 
grand nombre d’écailles cardioides entre les 
nageoires pelviennes (plus de 5) et la coloration 
rouge-orangée de ses nageoires inférieures. 


Répartition 
Répartition générale 

La Figure 2 présente la distribution nord-américaine 
des deux sous-espéces de Esox americanus. On 
remarque que les deux sous-espéces sont largement 
répandues aux Etats-Unis. On les rencontre dans les 
eaux douces de la plaine cOtiére atlantique jusqu’au 
lac Okeechobee en Floride, dans les tributaires du 
golfe du Mexique jusqu’a la riviére Brazos au Texas 


' Ficure 1. Brochet d’ Amérique, Esox americans americanus, Male 190 mm; Québec, riviére Godefroy (A. Odum d’aprés 


Scott et Crossman 1975 avec permission). 


2001 


LACHANCE: RAPPORT SUR LA SITUATION DU BROCHET D’ AMERIQUE 


599 


% 

~ 

Ww 
2 
c 
% 
© 
© 
o 


® Sous-espece 


americanus 


Ea wv Sous-espéce 


vermiculatus 


Eee Hybride des 


deux sous-espéces 


FiGuRE 2. Aire de répartition mondiale des deux sous-espéces dEsox americanus modifiée daprés Crossman 


1966 (tirée de Lapointe 1980). 


a l’ouest et dans le Mississippi et ses tributaires. Les 
sous-espéces sont séparées par la chaine de mon- 
tagnes des Appalaches, ie brochet d’ Amérique occu- 
pant l’est et le brochet vermiculé occupant |’ ouest, 
sauf au sud ot elles s’hybrident. On remarque, de 
plus, que la limite septentrionale de la répartition 
géographique des deux sous-espéces est le systéme 
des Grands Lacs inférieurs et le fleuve Saint-Laurent 
et quelques-uns de ses tributai]res. 
Répartition au Canada 

Le brochet d’Amérique au Canada est limité au 
Québec. Sa répartition (Figure 3) comprend le fleuve 


Saint-Laurent depuis les iles de Contrecoeur jusqu’a 
l’embouchure de la riviére Godefroy, y compris le 
lac Saint-Pierre et plus particuli¢rement la région 
autour des iles de Berthier; le bassin de la riviére 
Richelieu, y compris le lac Champlain, la Baie 
Missisquoi et la riviére du Sud; ainsi que les rivieres 
Yamaska, Saint-Francois, Maskinongé et Godefroy, 
y compris le lac Saint-Paul (Mongeau et al. 1974; 
Scott et Crossman 1974; Massé et Mongeau 1974; 
Mongeau 1979 a et b; Mongeau et al. 1981; Dubé 
1986; Dubé et al. 1988; P.N. Mellado, étudiant a la 
maitrise au département de biologie de |’ Université 


600 


BROCHET D'AMERIQUE 
( brochet des marais) 


ESOX AMERICANUS AMERICANUS 
REDFIN PICKEREL 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


FIGURE 3. Aire de répartition du brochet dAmérique, Exos americanus americanus, au Canada d’ apres les sites de capture 


(modifiée de Mongeau et al. 1974). 


du Québec a Montréal, Montréal, QC; communica- 
tion personnelle.). Il s’agit de la répartition minimale 
inventoriée, puisque le brochet d’ Amérique échappe 
souvent aux engins de péche couramment utilisés 
lors des inventaires ichtyologiques. De plus, |’ habi- 
tude qu’il a de se tenir dans les herbiers denses com- 
plique sa capture. 

Enfin, il n’a pas fait l’ objet d’étude de répartition 
spécifique. Il est 4 noter que la mention de |’ espéce 
dans les iles de Contrecoeur est récente (1994; 
Mellado 1996) et que c’est la premiére fois que les 
brochets ont été inventoriés si loin en amont de 
V’embouchure de la riviere Richelieu. Il y aurait donc 
une certaine extension de l’aire de répartition vers 
l’ouest. Ceci engendre la possibilité de recoupement 
avec l’aire de répartition de |’autre sous-espéce, le 
brochet vermiculé, dont l’extrémité est de l’aire de 
répartition se trouve autour de I’ile Perrot. 


Biologie et écologie 
Biologie générale 
La biologie du brochet d’ Amérique au Canada est 


peu connue. Les renseignements suivants ont été tirés 
de l’oeuvre de Scott et Crossman (1974), de l’étude 
de Crossman (1962) en Caroline du Nord, ainsi que 
de celles de Dubé et al. (1988) et de Lapointe (1980) 
dans le Haut-Richelieu, Québec. Cette derniére étude 
faisait le point sur les connaissances acquises a cette 
époque et la revue informatisée de la littérature de 
1978 a 1995, effectuée dans le cadre du présent rap- 
port de statut, n’a ajouté que quelques informations 
supplémentaires. Afin de compléter le portrait 
général brossé ci-dessous, des données sur la crois- 
sance en longueur et en poids, sur l’alimentation, et 
sur le rapport gonado-somatique des brochets 
d’Amérique du Haut-Richelieu sont présentées en 
Figures 4—6. 

Le brochet d’ Amérique fraie généralement au 
printemps, mais une fraie automnale est possible 
comme chez les autres Esocidés. Il n’y a pas eu 
d’ observation de la fraie du brochet d’ Amérique au 
Canada, mais on a retrouvé des oeufs aux mémes 
endroits et au méme moment que ceux pondus par le 
grand brochet. La période et les aires de fraie 


2001 


(6) saiod 


€ 
E 
Ww 
< 
kK 
Oo 
F 
4 
~ 
Ww 
2 
oO 
Zz 
(e} 
- 


Valeurs moyennes pondérées par leur effectif 
(A= longueur observée; = poids observé; © = valeur calculée) 


FiGuRE 4. Courbes de croissance en longeur (-) et en poids 
(- -) des brochets d’ Amérique males et femelles cal- 
culées d’aprés le modéle de von Bertalanffy pour le 
Haut-Richelieu (tiré de Lapointe 1980). 


seraient donc probablement les mémes que celles du 
grand brochet. 

Typiquement, au printemps, la fraie débute en 
période d’inondation des rives, alors que la tempéra- 
ture de l’eau atteint 10°C. La durée de la fraie serait 
d’environ deux semaines (Scott et Crossman 1974). 
La fraie aurait lieu pres des rives inondées couvertes 
de végétation en eau peu profonde. Des données 
récoltées par Dubé et al. (1988) indiquent a cet effet 
que la répartition des ceufs était équivalente entre les 
zones a plus de 60 cm de profondeur et celles a 
moins de 60 cm (Lapointe 1980). La prairie humide 
a phalaris, et le marais a scirpe et a typha, seraient 
caractéristiques de habitat de fraie. Les oeufs sont 
pondus au hasard et abandonnés sur le fond et sur la 
végétation ou ils adherent. 

Le nombre total moyen d’oeufs est de 3716, dont 
186 a 542 sont mis, c’est-a-dire préts a étre pondus. 
Le nombre total d’oeufs dans un brochet d’ Amérique 
est environ un tiers du nombre total contenu dans un 
brochet vermiculé de méme taille. La fécondité du 
brochet d’ Amérique serait donc moindre que celle 
du brochet vermiculé. En tout temps, avant la fraie, il 
y a, en plus des oeufs miirs, des oeufs a deux autres 
stades de recrutement. Selon des découvertes 
récentes, il semble que ces autres oeufs, qui ne sont 
pas expulsés lors de la fraie printaniére, pourraient 
mdrir au cours de |’été et servir 4 une fraie autom- 


LACHANCE: RAPPORT SUR LA SITUATION DU BROCHET D’ AMERIQUE 


601 


nale, sinon il y aurait atrésie (E. J. Crossman, 
Curator Emeritus, Royal Ontario Museum, Toronto, 
Ontario; communication personnelle). De fait, des 
larves d’Esocidés portant encore des vestiges de la 
resicule utelline, ont été capturées aux mois d’octo- 
bre et de novembre !996 au marais de lile St- 
Eugene. Ces larves ont été identifiées comme etant 
des brochets de Amérique de taille allant de 14 a 
32 mm ce qui suggére fortement une reproduction a 
la fin de l’été ou au début de l’automne (Letendre et 
Dumas 1999). Les oeufs mirs sont transparents, de 
couleur jaune doré, et présentent un diamétre moyen 
de 1,9 mm. Les oeufs éclosent aprés une période de 
10 a 14 jours, et les alevins demeurent inactifs pen- 
dant le période de résorption de la réserve vitelline, 
soit de 10 a 12 jours. Par aprés ils commencent a se 
nourrir activement. 

A la sortie de lV oeuf, les alevins mesurent de 5,8 a 
6,1 mm. La croissance est trés rapide le premier été, 
et peut atteindre de 20 a 30 mm par mois. Dans le 
Haut-Richelieu, les jeunes de l’année atteignent une 
taille moyenne d’un peu plus de 100 mm 4a la fin de 
septembre. Par la suite, la croissance differe selon le 
sexe. Les femelles croissent plus vite et plus 
longtemps que les males. Les longueurs rétrocal- 
culées aux différents annuli pour les femelles d’age 
1, 2, 3, 4, et 5 sont respectivement de 106—120 mm, 
de 163-179 mm, de 202—220 mm, de 244-247 mm 
et de 269 mm. Pour les males ces longueurs sont de 
99-110 mm, de 141—156 mm, de 182—193 mm, de 


(6) salod 


E 
E 
Ww 
z 
i al 
ie) 
re 
a 
= 
Ww 
= 
ro} 
= 
6 
pa) 


Valeurs moyennes pondérées par leur effectif 
(A= longueur observée; (J= poids observé; * = valeur calculée) 


FIGURE 5. Croissance des jeunes brochets d’ Amérique de 
l'année capturés dans le Haut-Richelieu en 1976. 


602 


ANNEES 
REGROUPEES 
N= 846 


g 
a 
=) 
& 
3) 
a 
ze 
2 
2 
re) 
Qa 
Ww 
a 
uw 
a 
ao 
= 
ro} 
4 


i} 
GROUPE D'AGE 


FicureE 6. Age des spécimens capturés 4 chaque année 
d’échantillonnage dans le haut-Richelieu (tiré de 
Lapointe 1980). 


204—228 et de 244 mm respectivement (Lapointe 
1980). 

Le taux de croissance du brochet d’Amérique du 
Haut-Richelieu est tres semblable a celui noté pour 
la sous-espéce en Caroline du Nord, soit a 10° de lat- 
itude plus au sud (Lapointe 1980). Des mesures som- 
maires relevées sur les brochets d’ Amérique de la 
baie Lavalliére confirment ce rythme de croissance 
(Lepage et Gélinas 1996). Aucune mention de |’ age 
ou de la taille 4 maturité sexuelle n’a été relevée 
dans les informations disponibles pour le brochet 
d’ Amérique. On sait par contre, que le brochet ver- 
miculé devient mature lorsqu’il atteint 157 mm chez 
les femelles et 141 mm chez les males, soit vers 
lage de deux ans. Il est plausible que ce soit le cas 
pour le brochet d’ Amérique, étant donné un rythme 
de croissance similaire (Lapointe 1980). Les femel- 
les semblent vivre plus longtemps, et l’4ge maximal 
atteint par les brochets d’ Amérique serait de 7 ans. 
Dans le Haut-Richelieu, les individus capturés les 
plus agés atteignaient 5 ans. 

Les brochets d’ Amérique sont des poissons omni- 
vores se nourrissant principalement de crustacés, de 
mollusques, d’insectes, de poissons et d’amphibiens 
(Lapointe 1980). Bien qu’on puisse retrouver des 
proies de toutes catégories chez des individus de 
toutes les tailles, il existe cependant des différences 
quant a la proportion des différents groupes de proies 
utilisés selon la taille du brochet. Ainsi les jeunes 
individus de moins de 100 mm consomment une 
grande proportion de crustacés (cladocéres, ostra- 


THE CANADIAN FIELD-NATURALIST 


Vor E15 


ANNEES 
REGROUPEES 
N= 846 


| | 
i) 


Bei Wipes 
Oe ae CR ei eet 


Mt 


gi 
a 
=) 
& 
3) 
7) 
2 
oO 
7) 
2 
fo) 
a 
uu 
a 
Ww 
or 
oO 
= 
(e) 
2 


GROUPE D’AGE 


FIGURE 7. Végétation des terres humides au lac Saint- 
Pierre (MLCP 1986). Rapport Sur La Situation Du 
Brochet D’amérique (Esox americanus ameri- 
canus), Au Canada* 


codes, amphipodes et isopodes) et dans une moindre 
mesure de larves d’insectes aquatiques tels les 
éphéméropteres, les dipteéres, les hémipteéres et les 
odonates (Lapointe 1980). Chez les brochets de 100 a 
150 mm, les crustacés demeurent une part importante 
du régime, mais les insectes au printemps, et les pois- 
sons en été et en automne deviennent omniprésents 
dans le régime alimentaire. A partir de la taille de 
150 mm, le poisson devient |’élément le plus impor- 
tant du régime des brochets d’ Amérique bien que les 
autres éléments soient aussi présents. Les espéeces de 
poissons consommeées par les brochets d’ Amérique 
sont nombreuses et se regroupent dans les familles 
suivantes : Centrarchidés, Cyprinidés, Esocidés, 
Percidés, Ictaluridés, Cyprinodontidés et Umbridés. II 
peut y avoir du cannibalisme. 

Le brochet d’ Amérique étant une espéce plutot 
discrete et peu étudiée, il n’existe pas de données sur 
son comportement. Il semble par contre, selon les 
patrons de captures, que ce soit un poisson plutdt 
sédentaire, se déplacant trés peu. On sait que les bro- 
chetons demeurent a proximité des aires d’alevinage 
jusqu’a la fin de l’été (Lapointe 1980). Comme on 
l’a vue, il est un prédateur de plusieurs especes de 
poissons, mais il peut aussi servir de proie. Son ali- 
mentation omnivore le met en situation de compéti- 
tion potentielle avec un grand nombre d’espeéeces 
ichtyologiques. Les jeunes entrent en concurrence 
avec les espéces qui consomment des invertébrés 
(Scott et Crossman 1974). Par contre, il est probable- 


2001 


ment la proie d’autres espéces telles que les crapets, 
la perchaude, la barbotte brune, la barbue de riviere 
et les autres brochets qui partagent son habitat. 

De ce résumé de la biologie du brochet 
d’ Amérique, on retient que ce poisson présente une 
faible fécondité, a un cycle de vie court, entretient 
des relations de compétition potentielles avec 
plusieurs espéces, et est tour a tour une proie et un 
prédateur. Ces facteurs peuvent contribuer a limiter 
son abondance. 


Habitat 

L’habitat typique du brochet d’ Amérique, tant des 
adultes que des juvéniles, et ce, tout au long de leur 
cycle vital, se trouve dans les cours d’eau ordinaire- 
ment acides, 4 eaux mortes et a végétation dense. On 
peut aussi le retrouver dans des étangs, des eaux de 
retenue riche en végétation et les baies tranquilles 
des grands lacs et rivieres. 

Lapointe (1980) décrit habitat dans lequel on 
retrouve le brochet d’ Amérique dans le Haut- 
Richelieu. I] s’agit de ruisseaux, de canaux artificiels 
ou de petites baies de faible profondeur, alimentés 
par les eaux de ruissellement des terres agricoles. 
Tous sont caractérisés par des eaux peu profondes, 
plutot riches, et quelque peu turbides, et les courants 
y sont faibles ou nuls. La végétation aquatique y est 
trés abondante et composée de plantes aquatiques 
submergées, telles Myriophyllum exalbescens, 
Elodea canadensis, et Potamogeton spp., ainsi que 
de plantes aquatiques 4a feuillage flottant, telles 
Nuphar variegatum ou rubrodiscum et Nymphea 
tuberosa. La température de |’eau peut atteindre 
26°C, et le pH serait de 8,4. Cette description de 
habitat du brochet d’Amérique dans le Haut- 
Richelieu correspond assez bien a celle faite pour la 
sous-espece en Caroline du Nord, a l’exception du 
fait que les cours d’eau en question aux Etats-Unis 
sont alimentés par des marécages dont les eaux a 
forte teneur en matiére humique présentent du pH 
acide, soit de 4,2 a 4,9 (Crossman 1962). 

Dans le fleuve Saint-Laurent, l’archipel du lac 
Saint-Pierre est la zone la plus utilisée par le brochet 
d’Amérique selon Massé et Mongeau (1974). Elle 
est caractérisée par la présence d’herbiers aquatiques 
et de marais de faible et de grande profondeur 
(Figure 7). Les canaux entre les différentes files de 
Varchipel sont les lieux ot on le capture le plus 
fréquemment. Dans le fleuve Saint-Laurent, le pH de 
l’eau est aussi généralement alcalin. L’acidité de 
l’eau ne semble donc pas étre une caractéristique 
essentielle de habitat du brochet d’ Amérique. 

L’ étude de Dubé et al. (1988) a Vile Sainte-Marie 
dans le Richelieu décrit le milieu de fraie du brochet 
d’ Amérique. II s’agit de prairie humide a phalaris et 
de marais a scirpe et a typha, a des profondeurs vari- 

ant de 32 a 95 cm. 
Le brochet vermiculé et le brochet d’ Amérique, 
tolérent et méme préférent des températures élevées. 


LACHANCE: RAPPORT SUR LA SITUATION DU BROCHET D’ AMERIQUE 


603 


Ils peuvent aussi tolérer de trés faibles concentra- 
tions en oxygéne (0,3 ppm) caractéristiques des eaux 
tres chaudes (Scott et Crossman 1974). Par contre, 
les milieux ot l’oxygénation est faible 4 cause d’une 
forte pollution organique ne sont pas habités par les 
brochets (E. J. Crossman, communication person- 
nelle). Il semblerait que les jeunes de |’année et les 
adultes partagent tous le méme habitat puisqu’ils ont 
été capturés aux mémes endroits lors des inventaires 
récents dans la baie Lavalliére (Lepage et Gélinas 
1996). 


Dynamique des populations 

Il existe tres peu de données concernant |’ abon- 
dance passée ou présente du brochet d’ Amérique. 
Cependant, il semble que la fertilité de l’espéce soit 
faible (Scott et Crossman 1974). La fécondité du 
brochet d’ Amerique serait moindre que celle du bro- 
chet vermiculé : le nombre total d’ceufs dans un bro- 
chet d’ Amérique est d’environ le tiers du nombre 
total contenu dans un brochet vermiculé de méme 
taille (Scott et Crossman 1974). Par ailleurs, le suc- 
cés de la reproduction est probablement fonction de 
la durée et du niveau des inondations, tout comme 
c’est le cas pour le grand brochet (Machniak 1975; 
Massé et al. 1988). 


Facteurs limitatifs 

Compte tenu que le brochet d’ Amérique présente 
une faible fertilité (Scott et Crossman 1974), est peu 
mobile (Lapointe 1980), exige un habitat carac- 
térisée par des herbiers trés dense qui sont des 
milieux souvent perturbés et que le Québec soit a la 
limite septentrionale de sa répartition géographique, 
il semble peu probable que |’espéce puisse prendre 
beaucoup d’expansion dans I’ avenir. 

Par contre, la survie des populations est possible 
par le maintien de I’habitat. L’asséchement des her- 
biers et le contréle des inondations printaniéres sur 
les terres agricoles peuvent s’avérer fatals pour la 
survie des adultes, des oeufs et des alevins. 

Outre les modifications physiques de l’habitat et 
la pollution organique, il est difficile de préciser 
quels autres facteurs sont limitatifs pour le brochet 
d’Amérique, les éléments réglant la dynamique des 
populations de cette sous-espéce n’ayant pas été 
étudiés. Il est par contre permis de croire que la pré- 
dation et la compétition pourraient jouer un rdle dans 
l’expansion limitée du brochet d’ Amérique. 


Adaptabilité 

Il existe peu d’information concernant I’ adaptabil- 
ité de l’espéce, par exemple, aux changements dans 
les conditions du milieu. Bien que ce ne soit pas doc- 
umenté, il semble peu probable que le brochet 
d’ Amérique soit une sous-espéce trés flexible au 
niveau de ses exigences d’habitat, puisqu’il n’a pas 
pris d’expansion importante au Québec depuis 50 
ans. Un exemple inverse, serait celui de |’éperlan 
arc-en-ciel (Osmerus mordax) qui a envahi le lac 


604 


Ontario de fagon impressionnante sur une période de 
15 ans (Christie 1972). Les températures froides des 
eaux du Québec, ainsi que les grandes étendues 
d’eau claire, sans herbiers importants, ne sont pas 
propices a l’expansion de cette sous-espeéce. De plus, 
la grande détérioration des herbiers aquatiques 
autour des iles prés de Montréal depuis la deuxieme 
moitié de ce siécle n’aura sans doute pas aidé a 
l’extension de l’aire de répartition du brochet 
d’ Amérique vers |’ ouest. 

Aujourd’hui, avec les divers programmes visant la 
restauration et la dépollution des milieux humides 
(par example le Plan Saint-Laurent Vison 2000 et le 
Plan d’intervention de Montréal), la sous-espece 
pourra vraisemblablement étendre son aire dans cette 
direction. Dans ce cas, le brochet d’ Amérique se 
retrouvera sur le méme territoire que le brochet ver- 
miculé au Québec. L’extrémité est de l’aire de répar- 
tition se situe a l’fle Perrot, et il pourra y avoir hybri- 
dation comme dans le sud des Etats-Unis. 


Importance particuliére 

Le brochet d’Amérique, bien qu’il puisse présen- 
ter une certaine abondance localement, n’est pas un 
poisson d’importance pour la péche commerciale, 
sportive ou de subsistance au Québec. C’est une 
sous-espéce qui n’est pas trés connue aupres du 
grand public, qui la confond souvent avec des jeunes 
grands brochets et la remet a l’eau ou la jette. Aux 
Etats-Unis, elle fait l’objet d’une pécherie sportive 
(Scott et Crossman 1974). 

L’impact de l’avenement du brochet d’ Amérique, 
au niveau de la chaine trophique, est inconnu. II est 
probable qu’il joue un role similaire a celui du grand 
brochet, en utilisant des proies de moins grande 
taille. 

A en juger par la faible quantité d’ informations 
disponibles a son sujet, il ne semble pas attirer 
Vintérét des milieux scientifiques québécois, canadi- 
en et américain. Vu sa faible abondance et sa réparti- 
tion restreinte, il ne présente pas d’intérét comme 
bio-indicateur, les espéces utilisées a cet effet devant 
présenter une large répartition et une abondance 
élevée. 

Son arrivée récente au Canada, et le faible intérét 
du public pour cette sous-espece méconnue, ne font 
pas du brochet d’ Amérique un poisson d’importance 
culturelle ou sociale. 

Le brochet d’ Amérique est donc peu connu et de 
peu d’intérét pour le grand public, contrairement au 
grand brochet et au maskinongé qui sont extréme- 
ment populaires comme espéce d’intérét sportif. A 
cet €égard, son importance est semblable a celle de 
autre sous-espeéce, soit le brochet vermiculé. 
L’importance du brochet d’ Amérique réside en sa 
rareté, son adaptation aux milieux humides et aux 
petits cours d’eau, en son caractére énigmatique, en 
sa beauté et en son apport au patrimoine faunique du 
Québec. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Bilan de la situation 
Etat des populations 

Il n’existe que trés peu de données concernant 
V’abondance passée ou présente du brochet 
d’Amérique au Québec. Son aire de répartition a été 
établie a partir de mentions de captures, et trés sou- 
vent il s’agissait de la capture d’un seul individu. Les 
populations les plus abondantes se retrouveraient 
dans l’archipel du lac Saint-Pierre. 

Dans le Haut-Richelieu, la population semble sta- 
ble depuis une quarantaine d’années (R. Fortin, 
J. Dubé et P. Gosselin, communication personelle). 
Scott et Crossman (1974) mentionnent que I’ abon- 
dance de l’espéce dans le fleuve Saint-Laurent a la 
hauteur du lac Saint-Pierre aurait diminué suite a 
lV’aménagement de la voie maritime du Saint- 
Laurent. Un inventaire aupres des pécheurs commer- 
ciaux utilisant des verveux au lac Saint-Pierre en 
1983 a révélé que le brochet d’ Amérique formait 
moins de 0,1% de la biomasse annuelle capturée, soit 
environ 53 kg (Roy 1985, 1986). Les captures, a 
cette Epoque, étaient concentrées dans l’archipel du 
lac Saint-Pierre et dans la baie de Maskinongé. Les 
captures, qui sont accidentelles puisque les espéces 
visées par ces pécheurs sont surtout la perchaude et 
la barbotte, étaient plus fréquentes en avril et mai 
(Roy 1985, 1986). Des entrevues téléphoniques avec 
trois pécheurs actifs ayant au minimum 20 ans 
d’expérience dans le secteur, ont confirmé que les 
captures sont rares (moins de cing par année) et se 
font plutot au printemps (R. Michaud, J. Michaud, 
J.-C. Adant, communication personelle). Ces person- 
nes semblent d’accord pour dire que l abondance est 
légérement plus élevée depuis cing ou six ans. 

A ce jour, il est impossible de statuer de facon 
rigoureuse sur |’état des populations du brochet 
d’ Amérique. II semblerait que les seuls endroits ou il 
soit modérément abondant soient l’archipel du lac 
Saint-Pierre et le Haut-Richelieu. A ces endroits, les 
populations seraient stables. Par contre, des inven- 
taires récents ont permis d’établir qu’il existe dans la 
baie Lavalli¢re et dans le marais Saint-Eugene, des 
populations indépendantes qui s’y reproduisent. 
D’aprés les indications, l’abondance du brochet 
d’ Amérique dans la baie Lavalliére serait environ 5 a 
6 fois inférieure a celle du grand brochet (Lepage et 
Gélinas 1996). Ailleurs, les captures ne sont 
qu’occasionnelles et peu nombreuses. Bien qu’il soit 
probable que le nombre de brochets d’ Amérique soit 
faible, il existe aussi un biais dans |’évaiuation de son 
abondance, causé par les méthodes et les lieux 
d’échantillonnage qui ne visent pas cette sous-espece. 


Menaces a la survie de l’espéce 

Avec le peu de connaissances disponibles sur le 
brochet d’ Amérique, il est difficile d’établir les men- 
aces a sa survie. Par contre, les caractéristiques de son 
habitat préférentiel étant assez bien cernées, il est pos- 
sible d’affirmer que |’asséchement des herbiers et 


2001 


l’endiguement pour empécher I’ inondation printaniére 
des terres agricoles sont des menaces 4a Sa survie, ainsi 
qu’a celle de nombreuses autres espéces. II est a noter 
que de tels ouvrages ont déja été effectués dans le 
Haut-Richelieu 1a ot le brochet d’ Amérique est abon- 
dant (Dubé 1986). En ce moment, aucun projet poten- 
tiellement dangereux pour le brochet n’est a |’ étude 
que ce soit dans le Haut-Richelieu ou dans |’archipel 
du lac Saint-Pierre. 

Certains aménagements fauniques visant la 
sauvagine entrainent la formation de petits plans 
d’eau isolés 1a ot il y avait avant communication 
entre les marais et l’eau libre. Ces. aménagements 
devraient tenir compte des exigences du brochet 
d’ Amérique lorsqu’il est présent, c’est-a-dire perme- 
ttre la libre circulation des poissons, favoriser la 
présence des herbiers aquatiques et permettre |’inon- 
dation de la prairie humide au printemps, pour des 
profondeurs entre 30 et 90 cm. 

La pollution est une menace pour toutes les 
espéces vivantes et donc pour le brochet d’ Améri- 
que. A ce niveau, il semble que la pollution diffuse 
originant des pratiques agricoles intensives lui soit 
particulicrement néfaste. 

Sur le plan des facteurs naturels comme la mal- 
adie, la prédation ou la compétition, peu d’informa- 
tions sont disponibles. Par contre, il semble que le 
brochet d’ Amérique ait trouvé une niche dans la 
chaine trophique qu’il est en mesure de conserver 
dans les milieux ou il est plus abondant et ou les 
populations se maintiennent depuis plusieurs années. 


Protection légale et mesures de conservation 

Le brochet d’ Amérique n’est pas mentionné spé- 
cifiquement dans les reglements québécois de péche 
sportive, mais est groupé avec les autres Esocidés 
sous l’appellation de brochets. On le retrouve dans 
les zones de péche sportive 5, 6, 7 et 8 du Québec. 
La saison de péche aux brochets est fermée au print- 
emps afin de protéger les poissons en période de 
reproduction. Partout, la limite de prise quotidienne 
est de six, toutes espéces confondues (MEF 1995). 

Il est défendu de pécher le brochet d’ Amérique 
commercialement ou de l’utiliser comme poisson- 
appat. Par contre, les individus capturés a la péche 
sportive peuvent étre vendus. Cependant, les proba- 
bilités de capture sont extrémement faibles. 

Pour ce qui est de la protection de son habitat, il y 
a un site protégé sur son aire de répartition, soit la 
réserve écologique Marcel-Raymond dans le Haut- 
Richelieu (Hone 1988). De plus, le MEF est présen- 
tement en cours de procédures pour donner a un 
secteur du lac Saint-Paul le statut de réserve 
écologique (réserve écologique projetée Léon- 
Provancher) (G. St-Onge, communication person- 
elle). 

La Loi sur la conservation et la mise en valeur de 
la faune protége |’habitat du poisson sur les terres 
publiques du Québec. Aux fins législatives, |’ habitat 


LACHANCE: RAPPORT SUR LA SITUATION DU BROCHET D’ AMERIQUE 


605 


du poisson inclut les cours d’eau, les marais et les 
marécages ou |’on retrouve du poisson jusqu’a la cote 
de récurrence d’inondation de 2 ans, ou, si celle-ci 
n’est pas disponible, jusqu’a la ligne naturelle des 
hautes eaux. Entre autres, le terme poisson comprend 
en plus des poissons eux-mémes, leurs produits sexu- 
els et leurs oeufs de méme que les crustacés et les 
mollusques. Certaines interventions sont interdites 
dans cet habitat alors que d’autres peuvent étre réa- 
lisées conformément au Réglement sur les habitats 
fauniques. 

La Loi sur les péches du gouvernement fédéral 
protege aussi l’habitat du poisson qu’elle définit 
comme les frayéres, les aires d’alevinage, les aires 
d’alimentation et les voies de migration dont dépend, 
directement ou indirectement, la survie des poissons 
(article 34). L’esprit de cette loi est d’interdire 
exploitation d’ouvrages ou d’entreprises qui entrai- 
nent la détérioration, la destruction ou la perturbation 
de l’habitat du poisson (article 35[1]), a moins que 
ces activités ne soient autorisées par le ministre des 
Péches et des Océans ou faites conformément aux 
reglements (article 35[2]). 

Enfin, tout aménagement visant a améliorer ou a 
agrandir les herbiers et 4 permettre |’inondation des 
basses terres lors des crues printanieéres, sur |’ aire de 
répartition de cette sous-espéce, devrait aider a main- 
tenir sinon augmenter les populations de brochets 
d’ Amerique. 


Statuts actuels, légaux ou autres 

A ce stade, outre le fait que le brochet d’ Améri- 
que ait été placé sur la liste des espéces susceptibles 
d’étre désignées menacées ou vulnérables par le 
ministére de |’Environnement et de la Faune du 
Québec (Beaulieu 1992), et sur la liste des espéces 
prioritaires du Plan d’action Saint-Laurent, aucun 
statut légal n’a été accordé a cette sous-espéce, que 
ce soit au niveau international, national ou provin- 
cial. Le présent document vise cet objectif au niveau 
provincial et national. 


Conclusion/evaluation 
Le présent rapport a mis en lumieére plusieurs faits 
importants concernant le brochet d’ Amérique. Lors- 
que l’on considére cette sous-espéce, il est essentiel 
de tenir compte des éléments suivants : 
il existe trés peu d’informations au sujet du brochet 
d’Amérique au Québec outre certains travaux réalisés 
dans le Haut-Richelieu, les informations contenues dans 
Scott et Crossman (1974) et les résultats des inventaires 
ichtyologiques effectués au cours des 50 derniéres 
années dans le cadre de diverses études; 


V’aire de répartition du brochet d’Amérique au Canada 
n’inclut que le Québec; 

la sous-espéce est a l’extrémité nord de son aire de 
répartition mondiale; 


l’aire de répartition au Québec est restreinte au secteur 
du fleuve Saint-Laurent entre les iles de Contrecoeur et 
l’embouchure de la riviére Godefroy, ainsi qu’aux 


606 


réseaux hydrographiques des riviéres Richelieu (incluant 
le lac Champlain), Godefroy (incluant le lac saint-Paul), 
Yamaska et Saint-Franc¢ois; 


Vabondance de la sous-espéce semble faible en général, 
trois populations bien établies étant connues, soit celles 
du Haut-Richelieu, de |’archipel du lac Saint-Pierre et de 
la baie Lavalliére; 


V’abondance de la sous-espéce est possiblement sous- 
estimée puisque tres peu d’études lui ont été consacrées 
et que les inventaires ichtyologiques classiques ne font 
pas appel a des méthodes propices a sa capture: 


Vhabitat du brochet d’ Amérique, tant au niveau des 
juvéniles que des adultes, est caractérisé par des eaux 
propres, des herbiers denses de faible profondeur et des 
températures estivales élevées; 


les aires de fraie seraient identiques a celles du grand 
brochet, soit la zone riparienne inondée, ce qui rend la 
sous-espéce sensible aux travaux de remblaiement, 
d’endiguement ou d’asséchement; 


outre son apport au patrimoine faunique québécois et sa 
valeur intrinséque en tant qu’ élément de la biodiversité, le 
brochet d’Amérique n’a pas de valeur économique, cul- 
turelle ou sociale; il est trés peu connu du grand public; 


cette sous-espéce ne jouit d’aucune protection légale 
particuliére. 


Ainsi le brochet d’ Amérique est un poisson plutét 
rare, a propos duquel peu de choses sont connues et 
qui est trés sensible aux modifications qui peuvent 
étre apportées a son habitat. La conservation de cette 
sous-espéce passe par la protection de son habitat, 
par l’aquisition de connaissances supplémentaires a 
son sujet et par la sensibilisation du grand public a 
son existence, ainsi qu’a sa valeur intrinséque, 
écologique et esthétique. 

Au Canada, la sous-espéce le brochet d’ Amérique 
ne présente pas une grande aire de répartition, est a 
la limite septentrionale de son aire de répartition, et 
est généralement peu abondante. Il s’agit en fait d’un 
poisson rare. Les seules populations d’importance se 
trouvent dans |’archipel du lac Saint-Pierre et dans le 
Haut-Richelieu, et ou il semble que l’abondance soit 
stable. Le brochet d’ Amérique n’a pas d’ importance 
économique, mais sa beauté et sa rareté en font un 
membre important du patrimoine faunique canadien. 
Bien qu’on ait peu de connaissances sur le brochet 
d’ Amérique, il est plutdt évident que les menaces a 
son habitat que sont les endiguements et les asséche- 
ments d’herbiers aquatiques, peuvent étre détermi- 
nants pour la survie de la sous-espéce. 

La protection de son habitat, plus particulierement 
de ses aires de fraie et d’alevinage, sera déterminante 
pour sa survie future. La mise en oeuvre d’un projet 
d’ acquisition de connaissances sur cette sous-espéce, 
et la surveillance de l’expansion de son aire de répar- 
tition seraient souhaitables. 


Remerciements 
L’auteure tient 4 remercier toutes les personnes 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


qui ont consacré un tant soit peu de temps a la 
recherche d’informations concernant le brochet 
d’ Amérique. Soulignons entre autres la participation 
de E. J. Crossman, de Réjean Fortin, de Mme 
Suzanne Lepage, des gestionnaires des différentes 
régions du ministere de |’Environnement et de la 
Faune, soient Yves Mailhot, Jacques Bergeron, 
Martin Léveillé, Jean Dubé, Gérard Massé, Pierre 
Dumont et Bernard Bergeron, de Michéle Lapointe, 
des pécheurs commerciaux Pierre Gosselin, Roger 
Michaud, Jude Michaud et Jean-Claude Ardant et de 
M. Pedro Nilo Mellado. L’auteure remercie égale- 
ment M. Pierre East pour ses commentaires critiques 
du rapport et son aide a I’ édition finale. 


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l’exploitation de la faune. 135 pages. 

Roy, C. 1986. Importance relative de la biomasse péchée, 
commercialisée et rejetée pour les espéces de poissons 
capturés au verveux au lac Saint-Pierre en 1983. Mini- 
stére du Loisir, de la Chasse et de la Péche, Direction 
régionale de Trois-Rivieres, Service de l’aménagement 
et de l’exploitation de la faune. 44 pages. 

Scott, W.B., et E. J. Crossman. 1974. Poissons d’eau 
douce du Canada. Office des recherches sur les pécheries 
du Canada, Ottawa, Bulletin 184. 1026 pages. 


Accepté 19 février 2002 


Status of the Weed Shiner, Notropis texanus, in Canada* 


J. HOUSTON 
743 Fireside Drive, Constance Bay, Ontario KOA 3MO Canada 


Houston, J. 2001. Status of the Weed Shiner, Notropis texanus, in Canada. Canadian Field-Naturalist 115(4): 608-613. 


The Weed Shiner (Notropis texanus) is known in Canada only from Manitoba, where it was first collected in 1982. It has, no 
doubt, occurred in the area for some time, but was misidentified as the Blackchin (Notropis heterodon) or Blacknose Shiner 
(Notropis heterolepis). It does not seem likely that its presence is a result of introductions, but by natural dispersal through 
the headwaters of the Mississippi to the Winnipeg-Rainy River headwaters. Canadian populations appear to be stable. 


Le méné diamant (Notropis texanus) est connu au Canada seulement du Manitoba d’ou il était premiérement recueilli en 
1982. Sans doubte il se produisait dans la région depuis quleque temps, mais mal identifié avec le menton noir (Notropis 
heterodon) et/ou le museau noir (Notropis heterolepis). 11 n’ est pas probalble que sa présence est au cause de une introduc- 
tion d’homme, mais par une dispersion naturelle au travers les eaux a |’amont de la riviére Mississippi de l’amont de la riv- 
iére Winnipeg-Rainy. Au Canada I’espéce semble stable. 


Key Words: Cyprinidae, cyprinids, minnows, shiners, Weed shiner, méné diamant, Notropis texanus, rare fishes, endan- 


gered species, Manitoba. 


The Weed Shiner, Notropis texanus (Girard, 
1856), is a small cyprinid which has a nearly discon- 
tinuous North American distribution. There is a 
northern group in the upper Mississippi watershed in 
northern Illinois, lowa, Wisconsin, Missouri and 
Minnesota, with rare records in Red River headwa- 
ters in northwest Minnesota. This group is not com- 
mon. A lower Mississippi group starts at about the 
confluence of the Mississippi and Ohio Rivers and 
goes south from there, spreading along the Gulf 
Coast from the Florida Panhandle to coastal Texas 
(Swift in Lee at al. 1980 et seq.) Becker (1983) and 
Eddy and Underhill (1974) report that the range of 
Weed Shiners is decreasing in Wisconsin and 
Montana, respectively. Although it ranges north to 
Minnesota, and Illinois (Swift 1980) it is unknown in 
the Lake Superior drainage and until 1982 had not 
been recorded in Canadian waters. Stewart (1988) 
provides a record of the first Canadian collections in 
Manitoba. 


Description 

The Weed Shiner (Figure 1) is a small cyprinid 
rarely exceeding 6.5 cm in length. It is cylindrical in 
form and slightly compressed with an intense black 
lateral stripe which extends from the snout to the 
caudal peduncle where it ends in a black spot. Dark 
pigmentation may continue into the caudal fin rays. 
Dark pigment spots may be found on scales below 
the lateral line. Overall the Weed Shiner is straw 
coloured and is lighter on the sides and the belly. 
There is an obvious diffuse light stripe just above the 


*Reviewed and approved by COSEWIC April 1999, status 
assigned — Not At Risk. 


dark lateral band. The dorsal fin originates anterior 
to the posterior insertion of the pelvic fins and the 
anal has seven rays. The mouth reaches the anterior 
margin of the eye and the upper jaw does not pro- 
trude beyond the tip of the snout, the lower jaw 
being included in the upper. There are two rows of 
pharyngeal teeth (2, 4—4,2 modal count), the pres- 
ence of two on the inner row being diagnostic for 
Notropis texanus (K.W. Stewart, Department of 
Zoology, University of Manitoba, Winnipeg, Mani- 
toba; personal communication). The lateral line has 
34 to 37 scales (Eddy and Underhill 1974; Smith 
1979; Becker 1983). 

Weed Shiners are very similar to the Blackchin 
Shiner (Notropis heterodon) and the Blacknose Shiner 
(Notropis heterolepis) which have been collected in 
the Winnipeg River as far downstream as Great Falls 
(Stewart 1988). It also superficially resembles the 
Spottail Shiner (Notropis hudsonius) and the 
Bluntnose Minnow (Pimephales notatus). All four 
species can be distinguished from the Weed shiner 
and each other by use of morphometric and meristic 
features described above and elsewhere (e.g. Scott and 
Crossman 1973). However, the character values are 
variable and distinguishing the species from other 
local “blackline” shiners is extremely difficult. It also 
resembles the River Shiner (Notropis blennius), but it 
lacks the lateral prominent black stripe and the River 
Shiner is not known from the Winnipeg River water- 
shed (Stewart 1988). 


Distribution 

This is a lowland species which ranges from the 
Suwannee River of Florida and Georgia west to the 
Neuces River in Texas, north along the Mississippi 
Valley to the Red River of the North in Minnesota 
and in drainage of lakes Michigan and Huron (Figure 


608 


2001 


HOUSTON: STATUS OF THE WEED SHINER 


609 


10 mm 


FiGurE |. Left side of a 29 mm specimen of the Weed Shiner, Notropis texanus, collected from the Winnipeg 
River on 20 September 1986 [reproduced from Stewart (1988) by permission]. 


2). It is not found in the upper Ohio River Basin or 
in the Lake Superior drainage (Swift 1980). The 
species has not been found in northwestern Ontario 
despite attempts to find it there (E. Holm, Royal 
Ontario Museum, Toronto, Ontario; personal com- 
munication). 

In Canada the species is known only from 
Manitoba (Figure 3), where it was first collected in 
1982 from the Ochre River (51E17’N 99E48’W), 
although the first recorded collection was upstream 
of the Great Falls Dam in the Winnipeg River 
(S0E28’N, 96E00’W) in 1986 (Stewart 1988, Table 
1). Specimens had previously been taken from Lake 
Dauphin (51E12’N, 99E34’W) between 1982 and 
1986, but misidentified as Blackchin Shiners 
(Stewart, personal communication). Further collec- 
tions have been made in the Icelandic River, south of 
Riverton, Manitoba, in Dauphin Lake, and the 
Winnipeg River above and below the Pine Falls Dam 
(Stewart, personal communication, Table 1) and 
north along the east side of Lake Winnipeg to Poplar 
River. 


Protection 

In the U.S. the species has declined in Wisconsin 
where it has been given Protected Status, and is con- 
sidered of Special Concern in Iowa (Johnson (1987). 
A survey of state agencies in 1995 indicated that the 
Weed Shiner was endangered in Illinois, Iowa and 
Michigan and of special concern in Wisconsin. 

In Canada, the fish are not subject to any protect- 
ed status. General protection could be afforded if 
required under Manitoba provincial wildlife and 
endangered species legislation. The Provincial 
Conservations Status Rank is S4 (secure). 


Population Sizes and Trends 
The species is apparently common in the south, but 


rare north of Arkansas (Swift 1980). It has declined in 
Illinois (although never abundant there), apparently 
due to siltation and general deterioration of water 
quality (Smith 1979). It also declined in Wisconsin 
(Becker 1983), Minnesota (Eddy and Underhill 1974), 
Illinois, lowa and Michigan. 

There is no evidence on which to base informa- 
tion on populations sizes and trends from Manitoba. 
The 1986 collections (Table 1) yielded 53 under- 
yearlings, and in 1987, 292 specimens ranging in 
length from 18.5 to 39.1 mm Standard Length (S.L.) 
were taken from two locations (50°E28’N, 
96°E00’W and 50°E26’N, 96°E00’W) on the south 
shore of the forebay of the dam during the same col- 
lections (212 in 1986 and 80 in 1987). Both sexes 
were present in these collections and fish over 
30mm S.L. showed evidence of sexual maturity 
(Stewart 1988). No information on specimens taken 
in other collections is available. 

The Weed Shiner is apparently widely distributed 
in tributaries of the east side of Lake Winnipeg, 
extending northward to almost the north end of the 
lake. In addition, it has been found in the Icelandic 
River, a tributary of the west side of the south basin 
of Lake Winnipeg, and west to Lake Dauphin. In 
suitable habitat, it seems to be abundant. Manitoba, 
may in fact, be a refuge for the northern stocks of 
this species, given that its distribution in the North- 
Central United States seems to be shrinking mainly 
due to habitat loss. By contrast, its distribution in 
Manitoba suggests that it has been there for a long 
time and the known populations seem to be stable. 
The more remote of those populations also seem to 
be protected, at least for the present, from human 
impact. The spread of commercial wild rice culture 
could become a threat to Weed Shiner habitat in the 
future, but has not affected any of the known loca- 
tions thus far. 


610 


een 


THE CANADIAN FIBLD-NATURALIST 


Vol. 115 


A 


ee i noes nine 


FiGURE 2. North American distribution of the Weed Shiner, Notropis texanus (Swift 1980; Stewart 1988). 


The site of the nearest Manitoba collections is 
435 km linear distance north-northeast of the Otter 
Tail River (tributary to the Red River of the North) 
in Minnesota, the previously known extent of the 
northern occurrence of the species (Stewart 1988). It 
is not currently known from elsewhere in the Hudson 
Bay drainage (Stewart 1988), although Hubbs and 
Greene (1928) reported Weed shiners from the St 
Croix River, but Eddy and Underhill (1974) have 
found no evidence of the species there. Its presence 
in Manitoba is most likely the result of a post-glacial 
invasion from the south (Crossman 1991) as apposed 


to transport by man since it is not commonly used as 
a bait fish as it does not survive well in a bucket and 
is smaller than minnows usually used as bait 
(Stewart 1988), although ilt may be more commonly 
used for bait in southern states. 

Given its known distribution in Manitoba, the 
species has undoubtably been there for some time 
and has gone unnoticed or been misidentified as the 
Blackchin or Blacknose Shiner. The distribution of 
the species suggests rather recent post-glacial disper- 
sion from the Mississippi River headwaters in 
Minnesota into the headwaters of the Winnipeg- 


2001 


roe 


FiGuRE 3. Manitoba sites of Canadian collections of the 
Weed Shiner, Notropis texanus (Stewart 1988). 


Rainy River system and subsequent downstream dis- 
persal (see Crossman and McAllister 1985; Stewart 
1988). Although it is not now present in the 
Minnesota headwaters of the Mississippi it apparent- 
ly once was (Hubbs and Greene 1928) and should be 
looked for there and in the Rainy River, although 
attempts to find it there have so far proved fruitless 
(Holm, personal communication). 


Habitat 

The Weed Shiner is commonly found in sand bot- 
tomed streams of varying sizes and low gradient and 
in the slower moving regions of higher gradient 
streams in the south (Heins 1977; Smith 1979). In 
the north they are most often found in clear, protect- 
ed weedy streams, lakes and larger rivers (Eddy and 
Underhill 1974). 

In Manitoba the species was found in the Win- 
nipeg River in waters of pH 8.5, 0.8 to 1.2 m deep 
over silty or muddy substrate with thick beds of sub- 
merged aquatic vegetation (Stewart 1988). The water 
was stained slightly brown and varied from clear to 


Houston: STATUS OF THE WEED SHINER 


611 


turbid depending on wind direction. Water tempera- 
tures were 21°EC in August and 16.5°EC in 
September. 

Ross and Baker (1983) found that Weed Shiners 
were a flood exploitive species, being more abundant 
in years following non-destructive floods on low 
gradient streams than in low flood years. This sug- 
gests that the species may utilize floodplains for 
breeding. Kwak (1988) also found the Weed Shiner 
to be a flood exploitive species using the floodplain 
as a nursery area and returning to backwater areas 
when forced off the floodplain by receding water 
levels. 


Biology 

Little information is available on the biology of 
the species. It apparently breeds during March to 
April in the extreme south, during May in Missouri 
and June in Minnesota (Swift 1980). Heins (1977) 
discussed age and growth in Mississippi and found 
that the life span is two to three years, 60% of the 
growth is attained in the first year of life. 


Limiting Factors 

The species appears to have a narrow range of 
habitat requirements and responds quickly to 
changes in habitat and water quality. Smith (1979) 
indicated that Weed Shiners declined in Illinois 
waters at locations where human or other distur- 
bances resulted in increases in turbidity and siltation 
or decreased aquatic vegetation. Similar results have 
been noted in Wisconsin (Becker 1983) and 
Minnesota (Eddy and Underhill 1974). 


Special Significance of the Species 

The Weed Shiner is probably an important forage 
species where abundant, it is not suitable as a bait 
fish as it does not survive well in a bucket and is 
smaller than preferred minnow size (Stewart 1988). 

The recent discovery of the Weed Shiner in 
Canada, its disjunct distribution, and habitat require- 
ments are of interest to science in relation to the zoo- 
geographic history and distribution of species subse- 
quent to the Wisconsinan Period of glaciation. Its 
critical habitat requirements could also make the 
species a useful indicator of changing water quality, 
if the previous occurrence at a specific site were 
known. 


Evaluation 

The Weed Shiner is probably threatened in every 
northern state in which it occurs because of loss of 
the clear, weedy habitat it requires as the species is 
intolerant of turbidity, pollution or habitat degrada- 
tion (Stewart, personal communication). The species 
has obviously been in Manitoba for some time, and 
has been misidentified as the Blackchin and/or 
Blacknose Shiner until recently. It may be that the 


612 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


TABLE |. Manitoba collection records for Notropis texanus, the Weed Shiner [from University of Manitoba, Royal Ontario 


Museum (ROM) and National Museum of Natural Sciences (NMNS) collections. 


Number of 

Date Location Coordinates Specimens 
~/07/82 25 km (by air) northwest of Ochre River (NMNS 83 0186) 51°E17'N 99°E48'W 

14/06/84 Ochre River, 0.2 km from mouth 51°E06'N, 99°E45'W 3 
12/07/84 Ochre River mouth 51°E06’N 99°E45'W 5) 
20/09/86 Winnipeg River just above Great Falls Dam 50°E28'N, 96°E04'W 47 
11/08/87 Winnipeg River 3.6 km above of Great Falls Dam (ROM52608) 50°E28'N, 96°E00’W DAD 
20/09/87 Winnipeg River just above Great Falls Dam (ROM51886) 50°E27’N, 96°E01’W 

23/09/89 Icelandic River at Highway 8 51°E02’N, 97°E43'W 39 
12/08/91 Poplar River, sandy bay 2 km above Lake Winnipeg 53°E00'N, 97°E24'W 1 
13/08/91 Poplar River, below lowermost falls 53°E00'N, 97°E22’W py 
15/08/91 Mouth of Berens River 52°E21’'N, 97°E03’W 6 
15/08/91 Berens River, just east of IR. 52°E21'N, 96°E58’W 98 
15/08/91 Etomani River, just below lowermost rapids 52°E19’N, 96°E54’W 34 
21/09/91 Winnipeg River 3.6 km upstream of Great Falls Dam 50°E28'N, 96°E00’W 3 
21/09/91 Winnipeg River at Abbitibi boat dock, Pine Falls 50°E34’N, 96°E12’W D 
03/06/92 Fisher River at rock bridge on IR. JL °B26N, 97°E17 W. 4 
03/06/92 Icelandic River just below dam upstream of Highway 8 51°E02’N, 96°E58’W 47 
12/08/92 Bloodvein River, 0.5 km above Lake Winnipeg 51°E47'N, 96°E43'W 1 
13/08/92 Bloodvein River, | km downstream of Wolf Rapids 51°E43’N, 96°E43’W 2 
13/08/92 Bloodvein River just upstream of IR. 51°E45’N, 96°E43’W 92 
18/08/92 Pigeon River at mouth 52°E16'N, 97°E02'W Zt 
19/08/92 Pigeon River, in abandoned channel 52°E16'N, 96°ES59’W 9 
29/06/93 Icelandic River between Riverton and Highway 8 51°E00'N, 96°ES1°W 4 
28/07/93 Icelandic River at Highway 8 51°E00’N, 97°E00’W 10 
18/09/93 Winnipeg River 3.6 km upstream of Great Falls Dam 50°E28’N, 96°E00’W 126 
24/09/94 Winnipeg River 3.6 km upstream of Great Falls Dam 50°E28’N, 96°E00’W 70 
~/09/95 Winnipeg River at St Georges, just above Pine Falls Dam 50°E31'N, 96°E12’W 

~/09/96 Winnipeg River at St Georges, just above Pine Falls Dam 50°E31'N, 96°E12°W 

19/09/97 Winnipeg River 3.6 km upstream of Great Falls Dam 50°E28'N, 96°E00'W 80 
~/09/97 Winnipeg River at St Georges, just above Pine Falls Dam 50°E31'N, 96°E127W 


species has a refuge here, but industrial, urban and 
agricultural activity may result in habitat degradation 
as in the northern U.S. 


Acknowledgments 

The author would like to thank R. Campbell, 
Subcommittee Chairman of the COSEWIC Fish 
and Marine Mammals Subcommittee for helpful 
comments and advice in preparing this report. 
Thanks also to K. W. Stewart, Department of 
Biology, University of Manitoba, for his critical 
review and comments and permission to use his fig- 
ure. Financial support was provided by World 
Wildlife Fund (Canada) and the Department of 
Fisheries and Oceans. The late Don McAllister, for- 
merly of the Canadian Museum of Natural 
Sciences, and E. Holm of ROM helped with collec- 
tion records. 


Literature Cited 
‘Becker, G.C. 1983. Fishes of Wisconsin. University of 
Wisconsin Press, Madison, Wisconsin. 


Crossman, E. J., and D. E. McAllister. 1985. Zoogeog- 
raphy of freshwater fishes of the Hudson Bay drainage, 
Ungava Bay and the Arctic Archipelago. Pages 54-101 
in The Zoogeography of North American freshwater 
fishes. Edited by C.H. Hocutt and E.O. Wiley. John 
Wiley and Sons, New York, New York. 

Crossman, E. J. 1991. Introduced freshwater fishes: A 
review of the North American perspective with emphasis 
on Canada. Canadian journal of Fisheries and Aquatic 
Sciences 48(Supplement 1): 46-57. 

Eddy, S., and J.C. Underhill. 1974. Northern fishes. 
University of Minnesota Press, Minneapolis, Minnesota. 

Heins, D.C. 1977. The age and growth of the weed shin- 
er, Notropis texanus (Girard). American Midland Natur- 
alist 98: 491-495. 

Hubbs, C. L., and G. W. Greene. 1928. Further notes on 
the fishes of the Great Lakes and tributary waters. 
Papers of the Michigan Academy of Sciences, Arts and 
Letters (1927)8: 317-392. 

Johnson, J. E. 1987. Protected fishes of the United States 
and Canada. American Fisheries Society, Bethesda, 
Maryland. 

Kwak, T. J. 1988. Lateral movement and use of flood- 
plain habitat by fishes of the Kankakee River, Illinois. 
American Midland Naturalist 120: 241-249. 


2001 HOUSTON: STATUS OF THE WEED SHINER 613 


Ross, S. T., and J. A. Baker. 1983. The response of fishes Swift, C.C. 1980. Notropis texanus (Girard), Weed shin- 


to periodic spring floods in a southeastern stream. er. Page 316 in Atlas of North American freshwater fish- 

American Midland Naturalist 109: 1-14. es. Edited by D.S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. 
Smith, P.W. 1979. The fishes of Illinois. University of Jenkins, D. E. McAllister, and J. R. Stauffer Jr. North 

Illinois Press, Urbana Illinois. Carolina State Museum of Natural History, North Caro- 
Stewart, K. W. 1988. First collections of the Weed Shiner, lina Biological Survey Publication 1980-12. 


Notropis texanus, in Canada. Canadian Field-Naturalist 
102: 657-660. Accepted 19 February 2002 


Status of The Bridle Shiner, Notropis bifrenatus, in Canada*+ 


E. HoLm!, P. DUMOoNT?, J. LECLERC2, G. Roy2, and E. J. CROSSMAN! 


‘Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queens Park, Toronto, Ontario MSS 2C6 
Canada 

2Direction régionale de la Montérégie, Ministére de Environnement et de la Faune, 201 place Charles-Lemoyne, 
Longueuil, Quebec J4K 2T5 Canada 


Holm, E., P. Dumont, J. Leclerc, G. Roy, and E. J. Crossman. 2001. Status of the Bridle Shiner, Notropis bifrenatus, in 
Canada. Canadian Field-Naturalist 115(4): 614-622. 


The Bridle Shiner is a small minnow that is restricted to the Atlantic drainage of eastern North America. In Canada, it is 
found in lowland areas of eastern Ontario and southwestern Québec where it is generally rare. It is usually found associated 
with aquatic vegetation where it breeds and finds food and protection from predators. Although it appears to be stable in a 
few areas, the Bridle Shiner has apparently declined in most of the river systems that have been recently resurveyed. The 
status of this species in many other waters where it has been captured is unknown because there have been no surveys there 
since the 1970s. Its decline is probably as a result of poor water quality, high turbidity, and decrease in suitable aquatic 
vegetation particularly in small rivers of the agricultural zone. It is recommended that the Bridle Shiner be classified as 
Vulnerable in Canada. 


Le méné d@’herbe est un petit cyprinidé dont la répartition est restreinte au bassin de drainage Atlantique de 1’ Amérique du 
Nord. Au Canada, sa présence a été rapportée dans les basses terres de |’est de l’Ontario et du sud-ouest du Québec; il est 
généralement rare. Ce cyprin est associé a la végétation aquatique, au sein de laquelle il se reproduit, s’alimente et s’abrite. 
La présence de cette espéce n’a été reconfirmée que dans une minorité de cours d’eau ou sections de cours d’eau ayant été 
l’ objet d’inventaires répétés au cours des 50 derniéres années. Son déclin est probablement relié a la dégradation de la qual- 
ité de l’eau, a l’accroissement de la turbidité et 4 une diminution de la disponibilité des herbiers aquatiques propices a 
l’espéce, et ce particuliérement dans les petits cours d’eau en zone agricole. Au Canada, le méné d’herbe devrait étre con- 


sidéré comme une espéce vulnérable. 


Key Words: Notropis bifrenatus, Bridle Shiner, méné d’herbe, Ontario, Québec, status, vulnerable. 


The Bridle Shiner, Notropis bifrenatus (Cope) 
(Figure 1), is a small member of the minnow family 
(Cyprinidae) that reaches a maximum size of 50 mm 
standard length. It is one of five species of Notropis 
in Canada with a prominent black lateral band which 
extends from the tail and continues on to the snout. 
There is often a bold caudal spot which is confluent 
with the midlateral stripe. The black band is particu- 
larly obvious in most preserved specimens but may 
be obscured in living specimens by the silvery 
scales. The Bridle Shiner gets both its common and 
scientific names from the appearance of the black 
pigment on the snout and upper lip. The black band 
narrows toward the tip of the snout and, at its end, is 
restricted to the upper lip. The lower lip has little or 
no pigment. The eye is one of the largest in 
Canadian cyprinids, its diameter ranging from 31.2 
to 38.8% of head length (Scott and Crossman 1973). 
The mouth usually extends to below the posterior 
half of the nostril. It is terminal to subterminal with 
the tip of the upper jaw projecting ahead of the tip of 


*Reviewed and approved by COSEWIC , April 1999, sta- 
tus assigned — Vulnerable 

Contribution Number 93 of the Centre for Biodiversity 
and Conservation Biology, Royal Ontario Museum 


the lower jaw and the snout rarely protruding beyond 
the tip of the upper jaw. Principal anal rays are usu- 
ally seven, although Scott and Crossman (1973) 
recorded 32% of specimens with eight. Male 
Notropis bifrenatus develop minute nuptial tubercles 
on the head, nape and pectoral fin (Jenkins and 
Burkhead 1993). 

Life colours of N. bifrenatus, have been described 
by Scott and Crossman (1973) and Jenkins and 
Burkhead (1994). In life, the back is straw coloured 
and there is a green blue iridescence on the sides giv- 
ing rise to the name “bluesides” used by one bait 
fisherman in the Lake St. Francis area. During the 
breeding season there is sexual dimorphism, males 
are bright yellow or golden on the lower sides, and 
the first 5 or 6 pectoral rays are margined with 
brown. The back is darker than in spawning females 
and non-breeding males. When breeding, both sexes 
develop yellow fins (Harrington 1947). 


Taxonomy 

Class: Actinopterygii 

Order: Cypriniformes 

Family: Cyprinidae 

Scientific Name: Notropis bifrenatus (Cope, 1869) 

English Common Name: Bridle Shiner 

French Common Name: méné d’herbe 

Comments: Gilbert (1980) suggested that the closest rela- 


614 


2001 


YY MAASAI XY TY 


Ho_mM, DUMONT, LECLERC, ROY, AND CROSSMAN: STATUS OF BRIDLE SHINER 


615 


TO gg 


Ficure |. Bridle Shiner, Notropis bifrenatus, female, 53 mm TL, ROM 21671. Fall Creek, 
Thompkins County, New York State. Drawn by Anker Odum, from Scott and 


Crossman (1973), by permission. 


tive of Notropis bifrenatus is the Blackchin Shiner, 
Notropis heterodon. Although one study (Mayden 1989) 
referred N. bifrenatus to Hybopsis, this classification is 
subject to change and we follow Robins et al. (1991) 
until the interrelationships between the species of 
Notropis and Hybopsis are better known. Coburn and 
Cavender (1992) retain Notrepis for N. bifrenatus. 


Distribution 

The Bridle Shiner is restricted to eastern North 
America in the Atlantic drainage from western Lake 
Ontario east to Maine and south to South Carolina 
(Gilbert 1980; Jenkins and Burkhead 1994). (See 
inset, Figure 2) 

In Canada, it is restricted to the eastern part of 
what is known as the Mixedwood Plains ecozone 
(Wiken 1986; Crossman and Holm 1997). It is found 
from the Bay of Quinte, Lake Ontario, east and north 
to Lac St-Paul, near Trois-Riviéres, Québec and 
south to Lac Memphrémagog (Figure 2) but not 
south in Vermont (see inset, Figure 2). It is found in 
lowland areas and does not occur far inland from the 
St. Lawrence River or Riviére Richelieu. 

Because the species is difficult to identify, some 
specimens in the collections of the ROM and the 
Canadian Museum of Nature (CMN) were re- exam- 
ined. Two lots in the ROM collection were question- 
able but their identity as Notropis bifrenatus could not 
be ruled out completely and they have been included 
as valid records: (1) The only record from the 
Napanee River has poor locality data but may be west 
of the Bay of Quinte. The single specimen (ROM 
24732) captured by the Department of Planning and 
Development in 1950, lacks a prominent lateral band 
and has 8 anal rays which is uncommon for Notropis 
bifrenatus. Notropis volucellus and Notropis 
stramineus are shiners that are similar to Notropis 
bifrenatus but lack a prominent lateral band. 
However, it is not Notropis volucellus because it has 
anterior lateral line scales which are not elevated, and 
it is probably not Notropis stramineus because it has 


eight anal rays. (2) One hundred and forty-nine of the 
154 specimens identified as Notropis bifrenatus cap- 
tured in Browns Bay in 1959 (ROM 23628) were rei- 
dentified (by EH) as Notropis stramineus (140), 
Pimephales notatus (8), and Cyprinella spiloptera (1). 
The remaining five specimens were cleared and 
stained and it was not possible for us to confirm their 
identification. However, those cleared and stained 
specimens could be Notropis bifrenatus as they were 
larger than the rest and therefore would have been 
easier to identify originally. Four of the five speci- 
mens had seven anal rays and the fifth had eight, like- 
ly counts for Notropis bifrenatus but also for Notropis 
stramineus. A small specimen at the Canadian 
Museum of Nature (CMN) identified as Notropis 
bifrenatus (NMC 74-0106) from Lac a la Péche in 
Parc National de la Mauricie, Québec, was reidenti- 
fied as Semotilus corporalis (by EH). 

The range of the Bridle Shiner in Canada has 
changed little from what was known in the 1940s. 
The species was first documented in Québec by 
Cuerrier et al. (1946) from the Montréal and Lac St- 
Pierre regions. In Ontario, it was first captured in 
1928 in the Bay of Quinte at the east end of Lake 
Ontario (Hubbs and Brown 1929). By 1938, collec- 
tions by G. C. Toner (ROM collection) had extended 
the range northeastward into an unnamed tributary of 
the Rideau Canal near Brewer’s Mill, and eastward 
into the Gananoque River, the St. Lawrence River 
near Gananoque, and in a tributary of Lake St. 
Francis. Radforth (1944) favoured the idea that 
Notropis bifrenatus had dispersed into Ontario from 
the Atlantic refugium either through the Mohawk- 
Hudson outlet or through the Champlain outlet. Its 
common occurrence in the upper Richelieu suggests 
that it used the latter means of post-glacial dispersal. 
Radforth (1944) suggested that it had arrived in 
Ontario only recently and had possibly not reached 
its limit owing to insufficient time for dispersal. 
However, Scott and Crossman (1973) predicted that 
the expansion of the range of the Bridle Shiner in 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Kilometres 


FIGURE 2. Distribution of the Bridle Shiner, Notropis bifrenatus, in Canada. A point may represent more than one capture 
at different sites within the area of the circle. Inset: North American distribution modified from Gilbert (1980). 


Canada would be prevented owing to sewage dispos- 
al problems caused by high population pressures and 
industrialization. Further range expansion is likely 
prevented by other factors such as water tempera- 
ture, intolerance to acidity and lack of suitable habi- 
tat (clear, vegetated creeks and rivers). 


Protection 

The Bridle Shiner is a species of concern in North 
Carolina (Johnson 1987). Jenkins and Burkhead 
(1994) recommended that the species be granted spe- 
cial concern status in Virginia and suggested that the 
status should be ascertained in other states. Global 
and North American federal, state and provincial 
conservation status and ranks were obtained from the 


Eastern Regional Office of the Nature Conservancy, 
Boston, dated 9 June 1997. The ranks assigned to the 
Bridle Shiner indicate that for those states where 
abundance is classified, abundance ranges from being 
very common to very rare throughout its range: 


Global Rank: G5 

National Ranks: United States: N5; Canada: N4 

Regional Ranks: Connecticut: $3; Washington, DC: 
unknown; Delaware: unknown; Massachusetts: unknown; 
Maryland: $3; Maine: $2; North Carolina: Historical!; New 


'Historical = not captured in the recent past, but not consid- 
ered extirpated. 


2001 


Hampshire: S4; New Jersey S4; New York: S5; Ontario: 
S3; Pennsylvania: $2; Québec: $4; Rhode Island: S5; 
Virginia: $3; Vermont: $3S4; | = extremely rare, 2 = very 
rare, 3 = rare to uncommon, 4 = common, 5 = very 
common 

In Canada, there is no specific protection, but the 
Bridle Shiner receives nominal protection from the 
federal Fisheries Act, particularly section 35(1) 
which states that a development proposal must not 
cause a “Harmful Alteration, Disruption, or Destruc- 
tion” of fish habitat (Minns et al. 1995). Habitat may 
also be protected by other federal legislation includ- 
ing the Environmental Assessment Act, Environ- 
mental Protection Act and Water Act. 

In Québec, habitat is generally protected by “Loi 
sur la qualité de l'environnement” (Environmental 
Quality Act). Fish habitat is also protected by the “Loi 
sur la conservation et la mise en valeur de la faune” 
(Act respecting the conservation and development of 
wildlife) which, under articles 128.1 to 128.18, con- 
trols activities that could modify biological, physical 
or chemical components peculiar to fish habitat. The 
“Loi sur les espéces menacées ou vulnérables” (Act 
respecting threatened or vulnerable species) makes 
additional provision for the protection of the habitat of 
threatened or vulnerable species. 

Ontario legislation which may protect habitat 
includes the Environmental Protection Act, 
Environmental Assessment Act, Game and Fish Act, 
Planning Act, and Water Resources Act. Aquatic 
habitats in Ontario tributaries of Lake St. Francis 
below Highway 2 which contain populations of Bridle 
Shiners (e.g., Gunn and Wood creeks) are protected 
against wetland fill-in by an Act administered by the 
Raisin River Conservation Authority (Anne Bendig, 
Ontario Ministry of Natural Resources (OMNR), per- 
sonal communication). See also Habitat. 


Population Size and Trend 

Population sizes for the Bridle Shiner have not 
been estimated in Canada. However, changes in pop- 
ulation size may be inferred from sampling data 
(summarized details of records of occurrence of the 
Bridle Shiner in Ontario and Quebec are on deposit 
with the COSEWIC Secretariat). Although the num- 
ber of specimens is sometimes given, this number is 
dependent on effort and method of capture and is 
only a rough indicator of relative abundance com- 
pared to other species. Population trends are 
obscured by the difficulty of identification of the 
species and many of the specimens are no longer 
available for identification confirmation. Records of 
occurrence of the species (number of records in 
brackets) were obtained from Ministére de 
l’Environnement et de la Faune du Québec (MEF), 
formerly Ministére du Loisir, de la Chasse et de la 
Péche (MLCP) and Ministére de 1’ Environnement du 
Québec (MENVIQ) (425), Royal Ontario Museum 
(ROM) (22), OMNR (12), University of Michigan 


Ho_M, DUMONT, LECLERC, ROY, AND CROSSMAN: STATUS OF BRIDLE SHINER 


617 


Museum of Zoology (UMMZ) (7) and Canadian 
Museum of Nature (NMC) (3). 

Jenkins and Burkhead (1994) documented several 
areas in the United States where the Bridle Shiner 
has declined or been extirpated. They consider that 
the Neuse River population in North Carolina is 
probably extirpated. In Virginia, some populations 
are extirpated or nearly so and the range of the 
species has receded sharply in Massachusetts, New 
Jersey, Pennsylvania and Maryland. Cooper (1983) 
indicated that the Bridle Shiner was once abundant 
in eastern Pennsylvania but is now taken only rarely, 
and only in the Delaware River drainage. Page and 
Burr (1991) indicated that it was fairly common but 
decreasing in some areas of its range (see also 
Nature Conservancy ranks under Protection). 

In Canada, evidence suggests that populations 
have declined in several river systems such as the 
St-Frangois, aux Brochets, Richelieu, Chateauguay, 
St-Laurent (Lac St-Louis and Lac St-Francois). 
There is no evidence of decline in recently sur- 
veyed waters in the St. Lawrence River around the 
Thousand Islands, two Ontario tributaries of Lake 
St: Francis, Lac St-Pierre and Lac St-Pierre 
Archipelago. Lack of adequate recent sampling 
makes it difficult to determine its status in other 
waters of its Canadian range such as lakes in the 
Rideau canal system, other tributaries of Lake St. 
Francis (on the Ontario side), Lac Memphrémagog, 
and Lac St-Paul. 

In Lac St-Frangois, the species was reported at the 
mouth of Riviere a la Guerre in 1941 and 1945 and 
in the northeastern part of the lake in 1968 
(Mongeau 1979a). No Bridle Shiners were captured 
during the fall of 1996 in 40 seine stations evenly 
distributed along the Québec shore of the lake 
(Fournier et al. 1997). The identification of the 1941 
specimens were confirmed by two of us (JL and PD). 

It would appear that Notropis bifrenatus was most 
abundant in the channels of Iles de Sorel and Iles de 
Berthier (Lac St-Pierre Archipelago) where 5387 
specimens were captured in 143 of 294 stations sam- 
pled between 1970 and 1971 (Massé and Mongeau 
1974). In that region it was more common than the 
related Blackchin Shiner and Blacknose Shiner, 
Notropis heterolepis. The Bridle Shiner was also 
abundant in Lac St-Pierre where 727 specimens were 
captured in 65 of 134 stations. Twenty-seven sites on 
12 tributaries of the south shore of Lac St-Pierre 
were sampled in 1982. The species was not recorded 
in those tributaries, although the Blacknose Shiner 
which is easily confused with the Bridle Shiner, was 
recorded from two of the sites (McFarlane and 
DuRocher 1984). Lac St-Pierre has also been sam- 
pled by MEF during fall 1995: 330 specimens were 
captured in 15 of the 36 seine stations distributed on 
the littoral zone of the lake; 84% of these specimens 
were captured at three stations. In the Lac St-Pierre 
Archipelago, 61 specimens were captured in 8 of the 


618 THE CANADIAN FIELD-NATURALIST 


40 stations evenly distributed along the numerous 
channels. This recent sampling indicates that the 
Bridle Shiner is still well-established in Lac St- 
Pierre and Lac St-Pierre Archipelago (Fournier et al. 
1996). 

In the Riviere Yamaska basin this species occur- 
red only in the lower part where it was found in 12 
of 210 seine hauls made between 1963 and 1971 
(Mongeau 1979b). A total of 16 specimens of the 
species were captured on 22 August 1989 in that 
river near its mouth (ROM 57019). However, it was 
not captured at any of four sites within the known 
range of the species in a 1995 electrofishing survey 
of 39 sites in the Riviére Yamaska (La Violette 
1997). 

It appears to have been reduced in the Riviere 
Chateauguay system. It was found mainly in the 
lower part of the basin in 1968. Between 1975 and 
1976 it occurred in 21 of 217 seine hauls in that sys- 
tem. However, in 1993 it was not recorded at any of 
the 21 sites electrofished by the Ministére de 
l’Environnement et de la Faune (La Violette and 
Richard 1996). 

The species was caught during the 1940s in Rivi- 
ére Saint-Francois, but was not reported in this river 
by Mongeau and his colleagues in the 1960s and 
1970s (Mongeau and Legendre 1976) or in any of 
the 26 sites sampled on the Riviere Saint-Franc¢ois 
during 1991 (Richard 1996). Its presence has also 
been reported in 1941, in Riviere aux Brochets, near 
the outlet, but not in the 1970s, during the systematic 
survey of this river by Mongeau (1979c). However, 
during spring 1990, six Bridle Shiners were captured 
in la Baie Missisquoi of Lac Champlain near the 
mouth of this river. 

The species was frequently encountered between 
1965-1970 in the Riviere Richelieu where it was 
found at 98 of 623 stations (Mongeau 1979c). In the 
lower Richelieu, between Chambly and Saint-Marc, 
27 specimens were collected in August 1970 (in 49 
seine hauls). In 1989, six specimens were captured 
by the ROM in the upper basin near the United 
States border (ROM Accession 5518). In 1993, none 
were caught in 129 seine hauls, during a survey in 
August and September (Jean Leclerc, unpublished 
data). In 1995, one specimen was caught in one of 21 
electrofishing stations 100 km from the mouth of the 
Richelieu (Saint-Jacques and Richard 1997). There- 
fore, the Bridle Shiner is still present in the Richelieu 
but recent surveys suggest that its abundance has 
decreased since 1970. 

Notropis bifrenatus was reported in the 1940s, 
1960s and 1970s in Lac Saint-Louis but was absent 
from the 1982-1983 seine catch of Beaulieu (1988) in 
the littoral and vegetated zone of that lake (n = 61 
seine hauls). During fall 1997, La Société de la faune 
et des Parcs du Québec (FAPAQ) seined 46 seine sta- 
tions evenly distributed along the shoreline and 


Vol. 115 


islands of Lac St-Louis. Of a total of 16 424 fishes, 
only one Bridle Shiner was reported from a site in Les 
Iles de la Paix Archipelago, along the southern shore. 
During fall 2001, FAPAQ seined 115 stations evenly 
distributed in the section of the St. Lawrence River 
between Montreal and Contrecoeur, 25 km down- 
stream of Montreal. Of a total of 23 514 fishes, 103 
Bridle Shiners were captured at three of the stations. 

In Ontario, there are few records and the species ha. 
always been rarely encountered. It appears to be stable 
in some areas but insufficient recent sampling makes it 
impossible to ascertain its status in other areas. In 
1994, a survey by ROM sampled 13 sites in the 
Ontario range of the species. The Bridle Shiner had 
been previously recorded from seven of those sites. It 
was captured at three of the 13 sites and constituted 
3.9-23.7% (= 6.6%) of the catch at those three sites. 
Attempts to capture the species in Sutherland and 
Finney creeks, and an unnamed creek near Brewers 
Mills failed but the Bridle Shiner was caught in Wood 
Creek, the St. Lawrence River in the Thousand Islands 
region, and in Jones Creek, a new location. 

Recent fieldwork by ROM, OMNR, and the New 
York Department of Environmental Conservation 
(see Carlson 1995) have documented several cap- 
tures of Notropis bifrenatus from the St. Lawrence 
and some of its tributaries. In 1991-1994, Bridle 
Shiners were recorded at 12 of 59 sites seined in a 
juvenile musky study. They constituted 0.03—70.1% 
(k= 7.2%) of the catch at each site (Anne Bendig, 
OMNR, St. Lawrence River Fisheries Unit, unpub- 
lished data). Some of these capture records may have 
represented Blackchin, Blacknose and/or Pugnose 
Shiners (Notropis anogenus). In 1994, the senior 
author sampled three of the OMNR sites where 
Bridle Shiners had been recorded and found them at 
two of the three sites. It is therefore likely that a 
majority of Bridle Shiner records documented by the 
OMNKR are correct and we have considered all of 
them valid. 

According to surveys in New York in the 
Thousand Islands region of the St. Lawrence River, 
the species has fluctuated in abundance over the last 
60 years. It was common in the 1930s when it was 
found in 64% of collections. But in 1976 it was found 
in only 2% of collections. It was more common in 
1993-1994 when it occurred in 26% of collections 
(Carlson 1995). The reason for its resurgence may 
relate to the increase in the clarity of the water (see 
Habitat) although differences in sampling personnel, 
gear and sites sampled could also be factors. 


Habitat 

Notropis bifrenatus is a warmwater fish that is 
found in quiet areas of streams and occasionally in 
standing water. It is found over a soft bottom of 
sand, silt, and detritus. It prefers colourless or mod- 
erately stained water and avoids turbid areas (Scott 


2001 


and Crossman 1973; Smith 1985; Jenkins and 
Burkhead 1994). It is tolerant to brackish water but 
is not acid tolerant which will likely prevent its 
spread in acid sensitive areas on the Canadian 
Shield. Carlson (1995) noted that in the Thousand 
Islands region of the St. Lawrence River, Notropis 
bifrenatus and Notropis heterolepis are more com- 
mon than Notropis heterodon and Notropis ano- 
genus, other “blackline” shiner species with similar 
habitat requirements occurring there. 

In Ontario, the Bridle Shiner is primarily restrict- 
ed to quiet areas of creeks and the St. Lawrence 
River, but has also been found in small lakes. It is 
usually associated with submerged, floating or emer- 
gent aquatic macrophytes. The substrate does not 
seem to be critical as it occurs over a variety of bot- 
tom types including organic detritus, clay, silt, grav- 
el, rubble and rocks. Although it is reported to be 
intolerant to turbidity it was captured at two sites 
where the water was described as turbid and at two 
other sites with secchi disk readings of 0.5 and 
0.7 metres (ROM collection records). 

In Québec, this minnow was found in relatively 
high frequency in sectors characterized by slow cur- 
rent, dense aquatic vegetation and highly developed 
shoreline perimeter along the numerous islands of 
Fleuve Saint-Laurent, the upper part of Riviére 
Richelieu and Riviére des Milles Iles and Lac Saint- 
Pierre. In the Lac St-Pierre Archipelago, during fall 
1995, 3 or the 8 stations where the Bridle Shiner was 
captured had low transparency (0.5—0.7 m). Fifteen 
of 61 Bridle Shiners collected in the Archipelago 
were captured in these stations. 

Harrington (1947) reported that the Bridle Shiner 
spawned among submerged aquatic plants (primarily 
Myriophyllum and Chara adjacent to other types of 
submerged and floating vegetation) where there is 
15—46 centimetres of free water above the vegeta- 
tion. Some spawning occurred in relatively barren 
areas later in the spawning season. 

Aquatic macrophytes are probably essential for 
nursery areas. Harrington (1947) found that the 
young of the Bridle Shiner were restricted to areas 
where spawning had occurred and were found 
among strands of Myriophyllum. Larvae have 
cement glands that allow adhesion to plants (Jenkins 
and Burkhead 1994). 

In the Ontario portion of the St. Lawrence River, 
there has been an increase in water clarity which 
occurred about the same time that the exotic Zebra 
Mussel (Dreissena polymorpha) was first discovered 
there in 1989. It would be expected that aquatic veg- 
etation would increase with increased clarity of the 
water, as it has in many other areas invaded by zebra 
mussels, but reports on abundance of aquatic vegeta- 
tion have been conflicting. Anglers have reported 
reduced levels of macrophytes in May and June 
1994. On the other hand, field work conducted by 


HoLM, DUMONT, LECLERC, ROY, AND CROSSMAN: STATUS OF BRIDLE SHINER 


619 


the OMNR in August found aquatic vegetation 
unchanged from previous years. Reduced levels of 
aquatic vegetation may have resulted from severe ice 
scouring and reduced phosphorus levels. Phosphorus 
levels have declined over the last 30 years as the 
result of improved sewage treatment, decreased lev- 
els of industrial pollution, and less agricultural run- 
off. Reduction of aquatic vegetation will be offset by 
habitat restoration or habitat compensation programs 
which are designed to comply with the Fisheries 
Act’s requirement for causing “no net loss or net 
gain” of fisheries habitat. (Anne Bendig, personal 
communication). 

Some populations of Bridle Shiners may receive 
protection because they occur in the upper St. 
Lawrence River in the Thousand Islands National 
Park. However, they would suffer from the effects of 
turbidity and water pollution created by any con- 
struction projects and toxic effluent upstream of the 
Park. 


General Biology 
Reproductive Capability 

The breeding of the Bridle Shiner in New Hamp- 
shire and New York was described in detail in several 
papers by Harrington (1947, 1948a, 1950, 1951). No 
individuals older than two years of age were found. 
Males normally spawned only once and most fre- 
quently in their first year. Females spawned in their 
first year if they reached 30 mm SL, but the majority 
spawned in their second year. The number of eggs 
ranged from 1062 for a 34 mm SL female to 2110 for 
a 44mm SL female but many of these eggs, with 
diameters ranging from 0.2 to 0.8 mm, may not have 
matured in time to be spawned. This size difference 
in the eggs was thought to indicate an extended 
spawning season. In New Hampshire, the breeding 
season ranged from the last week of May to mid-July. 
In New York breeding activity began on 2 May and 
lasted until August but the height of activity occurred 
in mid-June. Spawning usually took place at 17—22°C 
but occurred at temperatures as low as 14°C and as 
high as 27°C. Eggs were broadcast on vegetation and 
no parental care was provided. Most or all eggs were 
eaten by parents before they reached the bottom in 
aquaria (Harrington 1951) but it is not known if a sig- 
nificant number of eggs are eaten in nature. The eggs 
are adhesive but it is not known if they attach to 
plants (Jenkins and Burkhead 1994). 


Movement 

No information on movement or migrations of 
the Bridle Shiner is available. It is not likely that 
this frail, slow-swimming fish has a large home 
range. 


Behaviour and Adaptability 

The Bridle Shiner is a sight feeder and feeds dur- 
ing daylight hours on microcrustaceans, aquatic 
insects, detritus and living plant material. Much of 


620 THE CANADIAN FIELD-NATURALIST 


this food is found either on or above submerged 
aquatic plants. It feeds on the bottom when and 
where the vegetation is sparse and lacking (see 
Harrington 1948b). 

It is presumed that the Bridle Shiner is subject to 
heavy depredations by predators such as Northern 
Pike (Esox lucius), Grass and Redfin Pickerels (Esox 
americanus), Muskellunge (Esox masquinongy), 
Smallmouth Bass (Micropterus dolomieu), Yellow 
Perch (Perca flavescens), White Perch (Morone 
americana) and Black Crappie (Pomoxis nigromacu- 
~ latus) (Scott and Crossman 1973) because of its 
small size and weak swimming ability (Harrington 
1948a). 


Limiting Factors 

Increased turbidity adversely affects the ability of 
this species to locate its food and hinders the growth 
of submerged aquatic plants essential for feeding, 
reproduction and cover (Jenkins and Burkhead 1994). 
Wetland fill-in and physical removal of aquatic vege- 
tation would also be expected to reduce populations. 
The composition of the aquatic macrophyte commu- 
nity is probably also important for suitable spawning 
areas and food. It may prefer to spawn over watermil- 
foil (Myriophyllum) and Chara. The presence of a 
clear area above the submerged vegetation (cf macro- 
phytes that extend to the surface) is probably impor- 
tant for spawning activities (Harrington 1947). Thus 
any plant which has a tendency to grow to the surface 
before spawning occurs will have a deleterious effect 
on spawning success. The decline of the closely relat- 
ed Blackchin and Blacknose shiners have been asso- 
ciated with an explosion of the exotic Eurasian 
Watermilfoil, Myriophyllum spicatum in several 
Wisconsin lakes (Lyons 1989). Watermilfoil has been 
recorded in surveys by the OMNR (Anne Bendig, 
personal communication) but it is unknown whether 
this is a native Myriophyllum species or the intro- 
duced M. spicatum. 

Since the 1960s and 1970s, characteristics of most 
of the rivers of the Saint-Laurent lowlands in Québec 
have been modified by urbanization and by an exten- 
sive agricultural development of corn cropping and 
hog-rearing. Pesticides and nutrient loading 
increased in many small rivers of these lowlands. 
With the exception of Fleuve Saint-Laurent, most of 
the sectors where Notropis bifrenatus has been 
reported now suffer from severe sedimentation and 
eutrophication. In Ontario, the streams affected by 
agricultural development are in the watershed of 
Lake St. Francis where feed-lot and dairy cattle are 
raised and mixed pasture and corn crops are grown. 
These streams (Wood, Gunn, Finney creeks) have 
been channelized for drainage of fields and have 
high loadings of pesticides, nutrients, and sediment 
(M. Eckersley, OMNR, Kemptville, personal com- 
munication). 


Special Significance of the Species 

Wherever it occurs in sufficient numbers, the 
Bridle Shiner is presumably an important forage fish 
for a variety of important game fish (see Behaviour 
and Adaptability). Predators probably find the Bridle 
Shiner an easy target and a decline in numbers 
would have a negative impact on these sport and 
commercial fishes. 

Because of its small size and rarity it has limited 
use as a bait species in Ontario. In Québec, this fish 
is not used as live bait for sport fishing. 

The Bridle Shiner, is one of several “blackline” 
shiners-Notropis species which are superficially very 
similar and have a prominent black lateral band 
which extends from the tail and on to the snout (e.g. 
Blacknose, Blackchin, and Pugnose shiners). These 
species are sensitive to environmental change and 
thus all threatened by decreases in water clarity and 
aquatic vegetation and excessive loading of nutrients 
and pesticides. 


Evaluation 

There is evidence of decline in many waterbodies 
where the Bridle Shiner was formerly more abun- 
dant, particularly in the rivers of Québec. Factors 
which appear to be causing the decline of the Bridle 
Shiner may include decreases in water clarity and 
removal and/or change in species composition of the 
aquatic macrophyte community. Water quality dete- 
rioration may also have had an impact in urbanized 
and industrialized areas. Populations in Lac St- 
Pierre and the Lac St-Pierre archipelago appear to 
be relatively stable. The population in the Thousand 
Islands region of both Ontario and New York seems 
to have increased since the 1970s. Increase in Zebra 
Mussel populations has increased water clarity 
which would presumably have a beneficial effect on 
Bridle Shiner populations. Many areas such as Lac 
Memphrémagog, la Baie Missisquoi tributaries, Lac 
St-Paul and several sites in Ontario have not been 
adequately sampled since the 1960s and 1970s and 
it is not possible to determine its current status in 
these areas. 


Acknowledgments 

The authors acknowledge the following people 
for help with this status report: William R. 
Ramshaw for checking and plotting over four hun- 
dred distribution points and preparing figure 1; 
Anne Bendig for providing OMNR data on sam- 
pling between 1991-1994 on the St. Lawrence 
River, and for providing other information; Nathalie 
La Violette for providing recent data on sampling 
by MEF in several large tributaries of Fleuve Saint- 
Laurent and for reviewing this manuscript; Marty 
Rouse for field assistance during the 1994 ROM 
trip; Douglas Carlson for providing information on 
surveys on the New York side of the St. Lawrence 


Vol. 115 
| 


2001 


River; Sylvie Laframboise and Brian Coad for infor- 
mation on and loan of the specimen from Lac a la 
Péche. Financial support for fieldwork and prepara- 
tion of this report has been provided by the Royal 
Ontario Museum, the Ministére de |’ Environnement 
et de la Faune, World Wildlife Fund (Canada), and 
the Canadian Wildlife Service. 


Documents Cited 

Carlson, D. M. 1995. Status of the pugnose and blackchin 
shiners in the Thousand Islands region of the St. 
Lawrence River, 1993-1994. Manuscript report, New 
York State Department of Environmental Conservation, 
Region 6, Watertown. 

Crossman, E. J., and E. Holm. 1997. Freshwater Fishes. 
In: Assessment of species diversity in the Mixedwood 
Plains ecozone. Edited by |. M. Smith. http://www.cciw. 
ca/eman-temp/reports/publications/Mixedwood/fish/intr 
o.htm#toc (October 1997). 

Fournier, D., F. Cotton, Y. Mailhot, D. Bourbeau, J. 
Leclerc, et P. Dumont. 1996. Rapport d’opération du 
réseau de suivi ichtyologigue du fleuve Saint-Laurent: 
Echantillonnage des communautés ichtyologiques des 
habitats lentiques du lac Saint-Pierre et de son archipel 
en 1995. Ministére de |’Environnement et de la Faune, 
Direction de la faune et des habitats, Direction régionale 
de la Mauricie — Bois-Francs, Direction régionale de la 
Montérégie. 

Fournier, D., J. Leclerc, P. Dumont, et B. Bélanger. 
1997. Rapport d’opération du réseau de suivi ichty- 
ologigue du fleuve Saint-Laurent: Echantillonnage des 
communautés ichtyologiques du lac Saint-Francois en 
1996. Ministére. de |!’Environnement et de la Faune, 
Direction de la faune et des habitats. 


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Harrington, R. H. 1951. Notes on spawning in an aquari- 
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Hubbs, C. L., and D. E. Brown. 1929. Materials for a 
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Jenkins, R. E., and N. M. Burkhead. 1994. Freshwater 
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Johnson, J. E. 1987. Protected fishes of the United States 
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La Violette, N., et Y. Richard. 1996. Le bassin de la riv- 
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La Violette, N. 1999. Le bassin de la riviére Yamaska : 
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Lyons, J. 1989. Changes in the abundance of small littoral 
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MacFarlane, A., et L. Durocher. 1984. Inventaire ichthy- 
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Minns, C. K., J. D. Meisner, J. E. Moore, L. A. Greig, 
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THE CANADIAN FIELD-NATURALIST 


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Envirodog EN960254, rapport no EA-3, 70 pages + 10 
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Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, 
E. A. Lachner, R.N. Lea, and W.B. Scott. 1991. 
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Accepted 19 February 2002 


The Status of the Mira River Population of Lake Whitefish, 
Coregonus clupeaformis, in Canada* 


CHERYL D. GOODCHILD 


2064 Esson Line, RR 1, Indian River, Ontario KOL 2B0 


Goodchild, Cheryl D. 2001. Status of the Mira River population of Lake Whitefish, Coregonus clupeaformis, in Canada. 
Canadian Field-Naturalist 115(4): 623-634. 


A unique and possibly genetically distinct population of Lake Whitefish is found in the Mira River and its tributary the 
Salmon River, Cape Breton Island, Nova Scotia. The Mira River population of Lake Whitefish is another interesting exam- 
ple of the plasticity that can be expressed within the Coregonus clupeaformis ‘species complex’. Previously, it was thought 
that the Mira River population became established as a result of introductions of Lake Whitefish stock originating from the 
Great Lakes. Recent genetic studies indicate, however, that it is most closely related to the Acadian geographic race of 
Lake Whitefish (found only in Maine, Gaspé peninsula of Québec, New Brunswick and Nova Scotia), suggesting that the 
Mira River population may in fact be indigenous. Unfortunately, genetic studies conducted to date have not been at a suffi- 
cient level of resolution to confirm whether the Mira River population has diverged sufficiently to be considered a geneti- 
cally discrete or distinct population. The Mira River population of Lake Whitefish is probably indigenous, differs morpho- 
logically from other Lake Whitefish populations in Nova Scotia and is geographically isolated from other populations in 
eastern North America. Therefore further genetic study at a sufficient level of resolution to verify whether the population is 
an evolutionarily significant unit is warranted. 


Key Words: Lake Whitefish. Corégone de la riviére Mira, Coregonus clupeaformis , Mira River, Salmon River, Cape 


Breton Island, Nova Scotia, Canada, endangered species 


The population of Lake Whitefish (Corégone de 
la riviere Mira), Coregonus clupeaformis, (Mitchell 
1818), that occurs in the Mira River and its tributary 
the Salmon River, Cape Breton Island, Nova Scotia 
was identified as a potentially genetically discrete 
stock during studies of morphometric variation in 
whitefish populations from the Canadian Maritime 
provinces (Edge 1987). Lake Whitefish from the 
Mira River area exhibit distinctive characteristics for 
the species, particularly unusually low lateral line 
scale counts and low gill raker counts (Edge 1987; 
Edge et al. 1991). 

The Lake Whitefish has been shown to exhibit a 
wide range of phenotypic and genotypic variation 
across North America (Koelz 1931; Lindsey et al. 
1970; Scott and Crossman 1973; Ihssen et al. 1981), 
which has resulted in considerable taxonomic confu- 
sion. In recognition of the systematic problems 
resulting from the immense amount of ecological 
and morphological plasticity within the species, dis- 
tinctive populations are often referred to as part of 
the Lake Whitefish “species complex”. 

Recognition of the threatened Lake Simcoe 
Whitefish, Coregonus clupeaformis spp. (Evans et 
al. 1988) and the rare Squanga Whitefish, Coregonus 
sp., which occurs sympatrically with other Lake 
Whitefish populations in four lakes (Bodaly et al. 


*Reviewed and approved by COSEWIC April 1999, status 
assigned Indeterminate 


1988), as distinct provide precedents for considera- 
tion of protection for other distinct populations of 
Lake Whitefish within Canada. A distinct population 
can be defined as a group of interbreeding individu- 
als with similar gene frequencies (Li 1976). Such 
unique, genetically distinct populations represent 
evolutionary significant units of biodiversity that 
warrant identification and protection. 

Recent genetic evidence suggests that the popula- 
tion of Lake Whitefish found in the Mira River area 
should be reevaluated, possibly in the context of a 
larger, evolutionarily distinct, Acadian geographic 
race!, of Lake Whitefish (Bernatchez 1995; 
Bernatchez and Dodson 1990; Bernatchez and Dodson 
1991; Bernatchez et al. 1991; Bodaly et al. 1992). 


Description 

The population of Lake Whitefish found in the 
Mira (Figure 1) and Salmon Rivers (subsequently 
referred to as the Mira River population) has unusu- 


'The Acadian geographic race and the Atlantic or 
Mississippi-Misouri (Atlantic) geographic race are geneti- 
cally distinct stocks of the Lake Whitefish, Coregonus clu- 
peaformis. They should not be confused with the endemic 
Atlantic Whitefish, Coregonus huntsmani [formerly C. 
canadensis], a distinct species that has also been referred to 
by the common name Acadian Whitefish in a previous 
COSEWIC report. In this report naming follows the accept- 
ed common and scientific names used by the American 
Fisheries Society in Robins et al. (1991). 


623 


624 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Bbne 


by Meh 

VW 

tia 

Vy ids 
4, 


FIGURE |. Mira River Whitefish (drawing by D. Brammail, courtesy Don McAllister). 


ally low lateral line scale counts (Mira River 64—75, 
mean 70.1; Salmon River mean 68.0) and low gill 
raker counts (Mira River 20-24, mean 22.6; Salmon 
River mean 23.2) (Edge 1987; Edge et al. 1991). For 
Canadian populations of Lake Whitefish, lateral line 
scale counts reportedly range from 70 to 97 and gill 
raker counts range from 19 to 33 with seldom fewer 
than 22 in eastern Canada (Scott and Crossman 
1973). 

Many adults collected from the Mira River are 
characteristically large, robust and have a different 
external appearance from other Lake Whitefish in 
Nova Scotia (John Gilhen; Nova Scotia Museum of 
Natural History, Halifax, Nova Scotia; personal 
communication). 

For a detailed description of the Lake Whitefish in 
Canada, refer to Scott and Crossman (1973). Overall 
coloration of the Lake Whitefish in Canada 1s sil- 
very, grey or greenish with darker grey, greenish, 
light brown or dark brown back, becoming silvery 
on the sides and silvery or white below. The mouth 
is predominantly inferior and overhung by the snout, 
usually with weak teeth on the lingual plate and 
there are also teeth on the dentary in young fish but 
these are absent in adult specimens. Nuptial tuber- 
cles are present in both sexes, although they are 
fewer on females and more developed on males. 
Average length of adults in Canada is 38 cm total 
length but species can attain up to 73 cm in length. 

Considerable confusion over Lake Whitefish and 
Atlantic Whitefish, Coregonus huntsmani, has led to 
previous mis-identifications of the two species in 
Nova Scotia. Mouth shape, lateral line scales and 
presence or absence of teeth in adults are often good 
distinguishing characteristics (Edge 1987; Scott and 
Crossman 1973). Atlantic Whitefish differ from 
Lake Whitefish in generally having more lateral line 
scales (91 to 100), mouth usually terminal with small 


well-developed teeth on premaxillaries, palatine and 
vomer in adults, and teeth on tongue at all sizes 
(Scott and Scott 1988). With respect to mouth shape, 
Atlantic Whitefish tend to have a rounder snout pro- 
file and a more terminal mouth whereas Lake 
Whitefish tend to have an inferior mouth overhung 
by a more pointed snout. However, considerable 
variation in these characters has been reported for 
whitefish of both taxa from Nova Scotia (Edge 1987; 
Edge et al. 1991). 

Edge et al. (1991) suggested that the best diagnos- 
tic character to reliably distinguish Atlantic 
Whitefish from Lake Whitefish in the Maritimes is 
vertebral number, which ranges between 64 and 67 
for Atlantic Whitefish and 58 and 64 for Lake 
Whitefish. Location of capture may also help to 
identify the endemic Atlantic Whitefish, since it has 
an extremely limited and disjunct distribution in 
southern Nova Scotia. The Atlantic Whitefish has 
only been reported from the Tusket River, vicinity of 
the Tusket River in the Annis River (a tributary), 
Yarmouth Harbour (Yarmouth County), Halls 
Harbour (Kings County), off Wedgeport; as well as 
in Milipsigate, Minamkeak and Hebb Lakes, Petite 
Riviere watershed, (Lunenburg County), (ROM and 
CMN collection data; Scott and Crossman 1973; 
Scott and Scott 1988; Edge et al. 1991). A compari- 
son of characters of Atlantic Whitefish, Lake White- 
fish and Mira River Whitefish are presented in 
Table 1. 


Taxonomic Status 

A study of variation in Coregonus sp. from the 
Maritimes suggested that the Mira and Salmon River 
population of Lake Whitefish is a potentially geneti- 
cally distinct stock in eastern North America. 
With unusually low lateral line scale counts and low 
gill raker counts and can be distinguished using 


2001 


GOODCHILD: STATUS OF MIRA RIVER LAKE WHITEFISH 


625 


TABLE 1. A Comparison Of Characters Of Atlantic Whitefish, Lake Whitefish in Canada, and Mira River Population Of 


Lake Whitefish. 

Atlantic 
Characters Whitefish 
Gill Rakers 23-27, usually 25 

or 26 

Branchiostegal 6-9 
Rays 
Dorsal Rays 10-12 
Anai Rays 9-12 
Pelvic Rays 11 or 12 
Pectoral Rays 15 or 16 
Lateral Line 91-100 
Scales 
Pyloric caeca 
Vertebra 63 or 64 


canonical variates analysis of morphometric data 
(Edge 1987). Yet in a subsequent study of meristic 
and morphometric variation between the Lake 
Whitefish and the Atlantic Whitefish from the 
Maritimes, Edge et al. (1991) concluded that; while 
it was possible to readily distinguish the Atlantic 
Whitefish from the morphological variation found 
within the Lake Whitefish species complex, no sta- 
tistically significant morphological basis was found 
for recognizing any Lake Whitefish population from 
the Maritimes region as taxonomically distinct from 
other Maritimes populations. 

However, morphometric analysis is not a defini- 
tive method of identifying genetically different pop- 
ulations in studies of relationships of Lake Whitefish 
(except at the species level) because, like other 
members of the subfamily Coregoninae, whitefish 
exhibit a large degree of morphological plasticity 
(Imhof 1977). Therefore, morphometric analysis of 
Lake Whitefish populations cannot be relied upon to 
illustrate whether there are significant genetic differ- 
ences among populations. Although differences in 
gill rakers (in this case counts, but in others length) 
and other meristic and morphometric features may 
often reflect genetic differences, variation in the lat- 
ter can also occur in a relatively short time period 
through environmental modification (Lindsey 1981). 
Laboratory reared progeny of Lake Whitefish reared 
at different temperatures had fewer gill rakers than 
their parents (Todd 1997). 

Scott and Crossman (1964) looked at variation 
that occurred in Lake Whitefish populations 75 years 
after being introduced to Hogans Pond (47°35’N, 
52°51’ W), Newfoundland, from stock that originated 
from Lake Erie, Ontario. Gill raker counts from 
Hogans Pond Lake Whitefish were higher than those 
reported for Lake Erie, although only a very small 
sample of fish taken from Hogans Pond were ana- 
lyzed. Similarly, Loch (1974) studied induced phe- 
notypic changes in Lake Whitefish by comparing 


Mira River 
Lake Whitefish Population of 
in Canada Lake Whitefish 
19-33 20—24 ( mean 
22.6) 
8—10 (mean 8.9) 
11-13 (mean 13.9) 
10-14 (mean 14.9) 
11, sometimes 12 (mean 12.0) 
14-17 (mean 16.6) 
70-97 64—75 (mean 70.1) 
140-222 
54-64 (mean 60.5) 


second generation stock transplanted to Lyons Lake 
(49°44’N., 95°10’W.), Manitoba, with parental 
stock. Although gill raker number and lateral line 
scale counts were essentially constant, gill raker 
length was significantly different. He also found dif- 
ferences in other meristic and morphological charac- 
ters, as well as in electrophoretic phenotype frequen- 
cies of isozymes of GPDH. 

Populations of Lake Whitefish that feed on plank- 
tonic organisms and terrestrial insects tend to have a 
greater number of longer gill rakers than benthic 
feeding stocks (Lindsey 1981). Large differences in 
gill raker number and length often occur between 
sympatric pairs of Lake Whitefish with different 
ecological niches. Sympatric pairs tend to be repro- 
ductively isolated from each other and exhibit differ- 
ent protein allele frequencies. Yet sympatric pairs 
have not diverged as much genetically as the differ- 
ent geographic races of Lake Whitefish where there 
is no evidence of reproductive isolation or sympatric 
coexistence (Bodaly et al. 1992). In the geographic 
races there are instead many instances of introgres- 
sion among the different races (Foote et al. 1992). 
However, unlike sympatric pairs, the geographic 
races of Lake Whitefish have diverged little in eco- 
logical and morphological traits (Bodaly et al. 1992). 

Molecular genetic studies indicate that there are at 
least four (possibly five) geographic races of Lake 
Whitefish in North America — Nahanni, Berring, 
Mississippi-Missouri (Atlantic) and Acadian 
which have origins in different Wisconsinan glacial 
refuges (Bernatchez and Dodson 1994; Bernatchez et 
al. 1991; Bodaly et al. 1992). Lake Whitefish found 
in the northeastern United States and eastern Canada 
are from two geographic races. They may be part of; 

(a) an Atlantic race proposed by (Bernatchez and 

Dodson 1991)[said to be found only in parts 
of Maine and southern Quebec and originat- 
ing from an Atlantic glacial refugium], or the 
Mississippi-Missouri (Atlantic) race, and 


626 


(b) the Acadian race (Bodaly et al. 1992). 

The Acadian race, to which the Mira River popu- 
lation reportedly belongs, is present in the northeast- 
ern United States in Maine, and in eastern Canada in 
the Gaspé peninsula of Québec, New Brunswick and 
Nova Scotia (Bodaly et al. 1992). The Acadian race 
presumably survived Wisconsinan glaciation in the 
Northeastern Banks refugium (Schmidt 1986), 
although there are other areas that have been sug- 
gested as possible refugia including the area around 
the present-day Magdalen Islands, Gulf of St. 
Lawrence. 

The Mira River population is genetically most 
similar to other populations of Lake Whitefish that 
belong to the Acadian race (Bernatchez and Dodson 
1991), which suggests that it may be native. 
Although some genetic divergence from the parental 
stock could possibly have occurred in the approxi- 
mately 100 years since Lake Whitefish were intro- 
duced to Cape Breton Island, it would not be expect- 
ed to be as great as the genetic differences expressed 
in races with origins in different glacial refugia dur- 
ing the Wisconsin glaciation. If the Mira River popu- 
lation exists only as the result of previous introduc- 
tions of stocks that reportedly originated from the 
Great Lakes region it would be expected to exhibit 
greater genetic similarity to other members of the 
Mississippi-Missouri race. 

Edge (1987) provides a detailed discussion of 
whether Lake Whitefish populations in Nova Scotia 
are entirely the result of previous introductions or are 
native. (Refer to Distribution section for information 
on Lake Whitefish introductions in Nova Scotia). It 
has also been suggested that some Lake Whitefish 
introduced into the Maritimes region around the turn 
of the century may have come from stocks taken 
from areas in New Brunswick or Maine which 
belong to the Acadian race. However, some early 
natural history accounts suggest that at least some of 
the existing populations may be native to this region 
(Edge 1987). 

Another consideration that will shed light on the 
evolutionary history of Lake Whitefish in Nova 
Scotia is whether the species would have been able 
to naturally disperse from the hypthesized refugia 
after the last glacial period. 

The Wisconsin glaciation ended between 12 000 
and 10 000 years ago. Possibly a strip of unglaciated 
land or a series of islands on the outer continental 
shelf connected Nova Scotia to New England during 
the last glacial ice advance. Native freshwater 
species probably dispersed to Nova Scotia from this 
refugium since the retreat of the last ice sheet. 
Movement from one watershed to another was prob- 
ably as a result of river capture or the freshening of 
coastal waters by the melting ice sheet which 
enabled freshwater species to move around the coast 
from one river to another. Species with some degree 
of tolerance to salt water, including Lake Whitefish, 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


may have been able to move from one river system 
to another via estuaries. (Davis and Browne 1998). 
Nova Scotia has a relatively depauperate freshwater 
fish fauna with only 17 species considered to be 
native, fewer in Cape Breton and in northern Cape 
Breton Island there are no purely freshwater species. 
(Davis and Browne 1998). Low diversity of freshwa- 
ter fish species in parts of Nova Scotia suggests that 
there were impediments to dispersal after the last 
glacial period and may be a reflection of the lack of 
freshwater colonization routes. Perhaps in some 
areas, only species with tolerance to salt water were 
able to move from one river system to another via 
estuaries (Davis and Browne 1998). 

Deglaciation of lowland areas in the southeastern 
part of Cape Breton Island has been estimated to 
have occurred at about 13 000 years ago. In the 
northern upland areas of Cape Breton deglaciation 
probably occurred much later, with successive read- 
vances of ice possibly reoccurring until approximate- 
ly 10 000 years ago (Roland 1982). The lack of 
freshwater species in the northern part of Cape 
Breton Island may be accounted for by the later 
retreat of the ice sheet there. 

Considerable morphological and meristic variation 
has been shown among populations of Lake Whitefish 
in the Maritimes region (Edge et al. 1991). 
Apparently, the Mira River population has been 
known from the Mira River area for generations (J. 
Gilhen; personal communication). It is not known 
whether the species occurs there as a result of previ- 
ous unauthorized introductions or whether it may 
have arrived as a result of post-glacial dispersal and it 
has not been shown whether the Mira River popula- 
tion represents a unique genetic form. To resolve 
these phylogenetic relationships, it is necessary to 
combine morphometrics, molecular genetics and geo- 
logical history, bearing in mind that each of these 
methods may result in conflicting phylogenetic 
hypotheses for coregonids (Sajdak and Phillips 1997). 


Distribution 

Lake Whitefish have been reported from Atlantic 
coastal watersheds west throughout most of Canada 
and parts of the northern United States (Figure 2) to 
British Columbia, Yukon Territory and Alaska 
(Scott and Crossman 1973: Lee et al. 1980). In the 
United States, they have been introduced into parts 
of Montana, Washington and reportedly also in the 
Adirondacks region with fish from Labrador, Canada 
(Smith 1985). 

In Canada, populations are found in most large 
lakes and rivers, from Nova Scotia, New Brunswick, 
and Labrador, west throughout Quebec, Ontario 
including the Great Lakes, coastal waters of Hudson 
Bay, throughout Manitoba, Saskatchewan (especially 
in the north), Alberta, British Columbia, and general- 
ly distributed throughout both Territories (Scott and 
Crossman 1973). 


2001 


GOODCHILD: STATUS OF MIRA RIVER LAKE WHITEFISH 


627 


FiGuRE 2. Distribution of Lake Whitefish, Coregonus clupeaformis, in North America [from Scott and Crossman (1973) 
by permission]. 


Lake Whitefish have been introduced widely in 
Canada, including parts of insular Newfoundland, as 
a forage fish and in attempts to establish commercial 
fisheries (Scott and Crossman 1973). In Nova Scotia, 
early attempts to introduce Lake Whitefish (alleged 
to be from the Great Lakes) around 1877 were 
reportedly unsuccessful (Piers 1927). Ultimately, 
over 24 million Great Lakes Lake Whitefish fry were 
planted in 22 lakes in 8 counties in Nova Scotia 
between 1890 and 1901 (Semple 1973, from Annual 
Reports of the Department of Marine and Fisheries). 

In Cape Breton Island, Lake Whitefish were 
apparently only stocked in Lake Ainslie (46°08’N, 
61°11’ W) and Lake O’Law Brook (46°20°N, 
61°01’ W), Inverness County (Edge 1987). Evidently 
the stocking attempts in Cape Breton Island were 
unsuccessful as no Lake Whitefish were found in 
Lake Ainslie and Lake O’Law Brook during field 
survey’s conducted during the early 1980s (Edge 
1987). 

Currently, disjunct populations of Lake Whitefish 
are now found throughout Nova Scotia including the 
Mira River population in Cape Breton Island, 
although whether any are native populations or the 
result of previous stocking attempts is unclear. Until 


the early 1920s no indigenous form of the genus 
Coregonus had been reported from Nova Scotia even 
though Coregonus clupeaformis was known to occur 
in Québec, Labrador and New Brunswick (Piers 
1927). However, ichthyological surveys were virtu- 
ally unknown prior to this time. Subsequently, 
whitefish, presumed to be Lake Whitefish, were 
reported from a number of disjunct locations in Nova 
Scotia. Many of the localities where Lake Whitefish 
have been found are in locations for which there are 
no records of early introductions, although it is pos- 
sible that there were unrecorded or unauthorized 
introductions or that the original stocks dispersed 
and became established elsewhere (Edge 1987). 
Historical stocking records are considered to be 
unreliable and it is possible that stocking was more 
widespread that records indicate. 

Some of the early collections of whitefish in Nova 
Scotia, thought to be Lake Whitefish, were actually 
the endemic Atlantic Whitefish, Coregonus hunts- 
mani. Errors in identification resulted from confu- 
sion that existed prior to the recognition of the 
Atlantic Whitefish as a distinct species (Scott 1967). 

Lake Whitefish populations are now found in 
Guysborough, Lunenburg, Queens, Yarmouth, and 


628 


Cape Breton Counties and have been reported from 
Annapolis, Halifax and Richmond Counties (Semple 
1973; Edge 1987). 

Although not normally associated with brackish 
and saltwater habitats, the Lake Whitefish has been 
recorded along the east coast of James Bay, shores of 
Hudson Bay, and in brackish water adjacent to 
Ungava Bay (Scott and Scott 1988). It also occurs in 
brackish water in Arctic Ocean drainages of the 
Northwest Territories (Scott and Crossman 1973). 
The species also may occur in Atlantic coastal 
drainages but reports of this species in salt water off 
the Atlantic Coast of Canada have generally been in 
reference to the Atlantic Whitefish, with the excep- 
tion of one Lake Whitefish specimen caught in salt 
water off Black’s Harbour, New Brunswick (Scott 
and Scott 1988). 

The Mira River population of Lake Whitefish has 
been found only in the Mira River (46°02’N 
59°58’ W) and the Salmon River (45°55’N 60°18’ W), 
Cape Breton Island, Nova Scotia (Figure 3). The 
Salmon River is a tributary of the Mira River. Col- 
lection sites reported by Edge (1987) were approxi- 
mately 10 km apart. 

The population of Lake Whitefish from the Mira 
and Salmon Rivers is geographically separated from 
other extant populations of this species in the rest of 
Nova Scotia by St. Georges Bay and the Strait of 
Canso. Although Lake Whitefish are tolerant of salt 
water and may sometimes act in an anadromous 
fashion, they rarely stray too far from their rivers of 


Y 


Bay of Fundy 
Nova Scotia 


ei . 


o 


Atlantic Ocean 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


origin and therefore would not likely disperse to 
other areas through marine waters. Despite relatively 
extensive sampling including field surveys conduct- 
ed during 1982, 1983 and 1985, Lake Whitefish have 
not been found at any other localities in Cape Breton 
Island, particularly the Lake Ainslie and Lake 
O’Law Brook localities where they were reportedly 
stocked in the later part of the last century (Edge 
1987). 

The Mira and Salmon Rivers are approximately 
94 km and 62 km (measured “as the crow flies” from 
the mouth of each river) southeast of the Ainslie 
Lake location and are separated from it by Bras 
D’Or Lake, a large inland sea with surface salinity 
between 20.0 to 25.0 ppt but no significant tide 
(Natural History of Nova Scotia). It is unlikely that 
Bras D’Or Lake is a barrier to dispersal of Lake 
Whitefish since populations of the species elsewhere 
are tolerant of brackish and salt water. However, the 
lack of freshwater connections and opposite direc- 
tion of flow between the watersheds to the northwest 
of Bras D’Or Lake (1.e., Lake Ainslie and Lake 
O’Law Brook) and those to the southeast (i.e., Mira 
and Salmon Rivers) would impede dispersal of Lake 
Whitefish. Lake Ainslie empties into the Southwest 
Margaree River (46°20’N, 61°05’) and Lake O’ Law 
Brook empties into the Northeast Margaree River 
(46°20’N 61°05’W) which at their confluence flow 
northward into the Gulf of St. Lawrence at Margaree 
Harbour (46°26’N, 61°06’W). Although the Salmon 
River flows in a northeast direction parallel to Bras 


Breton Is 


FiGurE 3. Location of Capture of Mira River Lake Whitefish Populations in Cape Breton 


Island, Nova Scotia. 


2001 


D’Or Lake for part of its course, at its most northern 
part it turns and flows south away from Bras D’Or 
Lake and into the Mira River (Roland 1982). The 
Mira River empties into the Atlantic Ocean at Mira 
Bay on the southeastern coast of Cape Breton Island. 

It is possible that transfer between these two 
watersheds could have been assisted by human inter- 
vention. Lake Whitefish are a favoured food species 
and therefore are sought extensively on both a com- 
mercial and recreational basis in many parts of 
Canada (Scott and Crossman 1973). There are 
numerous instances where fish species have appar- 
ently been transferred through unauthorized and 
often unreported human actions. In Nova Scotia, 
unauthorized introductions of other freshwater fish 
species such as Chain Pickerel, Esox niger, and 
Smallmouth Bass, Micropterus dolomieu, have had 
profoundly negative effects on trout populations 
already marginalized by environmental degradation 
(R. Bancroft, Nova Scotia Department of Fisheries 
and Aquaculture, Pictou, Nova Scotia; personal com- 
munication). 


Protection 

The Mira River population Whitefish has not been 
formally recognized as a distinct species or evolu- 
tionarily significant population unit, has not been 
designated any status and it receives no specific pro- 
tection at this time. The status of other unique popu- 
lations of whitefish have been evaluated by the 
Committee on the Status of Wildlife in Canada 
(COSEWIC) with the result that the Lake Simcoe 
Whitefish, Coregonus clupeaformis ssp. is listed as 
threatened and the Squanga Whitefish, Coregonus 
sp. is listed as vulnerable by COSEWIC. However, 
COSEWIC status by itself does not currently confer 
special protection. 

Other populations of Coregonus clupeaformis 
receive no special protection in Canada other than 
the general habitat protection afforded by the Federal 
Fisheries Act. 

The Lake Whitefish does not receive protection in 
Nova Scotia. For recreational fishing, there is no 
catch and possession limit and no closed season, 
except that no one is permitted to angle in inland 
(non-tidal) waters unless the season is open for 
salmon, trout or Smallmouth Bass (Nova Scotia 
1998 Angling Summary of Regulations). On the 
other hand, the endangered Atlantic Whitefish, 
Coregonus huntsmani is a prohibited species under 
the Maritime Provinces Fishery Regulations promul- 
gated under the Fisheries Act. Atlantic Whitefish 
habitat is protected in the Petite Riviére watershed 
through designation as a Protected Water Area since 
Minamkeak, Milipsigate and Hebb Lakes provide 
the domestic and industrial water supply to the town 
of Bridgewater (Edge 1984 from Lunenburg County 
District Planning Commission 1980). 


GOODCHILD: STATUS OF MIRA RIVER LAKE WHITEFISH 


629 


In the United States, the Lake Whitefish is pro- 
tected only in Illinois (Johnson 1987) but Lake 
Whitefish are found only in some northern parts of 
the United States, where the species is at the extreme 
southern limit of its range. 


Population Size and Trends 

The size of the Mira River population of Lake 
Whitefish has not been estimated. Over a hundred 
specimens have been collected in the Mira and 
Salmon Rivers but only sporadic sampling has 
occurred over the past 20 years. The relative abun- 
dance of Mira River population was considered to be 
above average for whitefish from the Canadian 
Maritimes based solely on a comparison of the rate 
of catch in gill nets (Edge 1987); however the study 
was not specifically designed to estimate population 
SIZe. 

The Mira River population population may have 
been large enough to support a small local fishery 
since historically they were reportedly caught in nets 
set in the Mira River. Generally in Nova Scotia, 
Lake Whitefish are not considered to be an important 
species in recreational or commercial fishing 
(Semple 1973), probably a result of generally low 
population numbers. The lack of early accounts of 
the presence of Lake Whitefish in Nova Scotia 
waters (Piers 1927) suggests that population num- 
bers may have always been low or alternatively that 
current populations are primarily or entirely the 
result of introductions. Largely overlooked in the 
past, Lake Whitefish are now known from several 
watersheds in Nova Scotia and are more common 
than previously recognized (Edge 1987). Lake 
Whitefish have eluded capture in other areas. For 
example, despite extensive angling for Brook Trout, 
Lake Whitefish had never been recorded from Kerr 
Lake, New Brunswick, until they were taken in gill 
nets (Smith 1952). 


Habitat 

Very little is known about the habitat of the Mira 
River population of Lake Whitefish. The Mira River 
is generally very wide (up to 1.5 km) with little cur- 
rent until approximately 3 km from the mouth. There 
the river suddenly becomes more narrow and flows 
through a channel (with a steep embankment on the 
north shore) into Mira Bay. Collection data from a 
field input sheet indicates that Lake Whitefish from 
the Mira River were caught upstream in a gill net set 
150 m offshore at depths of between 12 to 25 m dur- 
ing July (collection data; Canadian Museum of 
Nature). The Mira River at the collection site is 
approximately a kilometer wide with almost imper- 
ceptible current and the habitat is more similar to a 
large lake. Although considerably narrower, the 
Salmon River meanders through lowland areas (less 
than 100 m above sea level). 


630 


The Lake Whitefish is a cool water species (Scott 
and Crossman 1973). In lakes in the southern part of 
its range, it occupies cooler waters in the hypo- 
limnion during the summer months (Scott and 
Crossman 1973). As reported by Smith (1952) for 
Kerr Lake, New Brunswick, Lake Whitefish were 
taken in summer in gill nets set approximately 7.5 m 
deep. Lake stratification was present and Lake 
Whitefish appeared to avoid oxygen depleted waters 
and shallow warmer waters. 

Lake Whitefish may be tolerant of a relatively 
wide range of habitats in Canada as indicated by col- 
lection records (unpublished habitat data, ROM). 
They have been found in water with currents ranging 
from still to swift and a wide range of water temper- 
atures up to a maximum of 22°C. However, cooler 
temperatures are required for successful incubation 
of Lake Whitefish eggs. As determined under experi- 
mental conditions, normal development occurs over 
a temperature range of 0.5°C to 6.1°C whereas at 
higher temperatures an increase in mortality and 
abnormality of eggs occurs (Scott and Crossman 
1973). At temperatures greater than 10°C only a very 
small percentage (1%) of eggs survive (Scott and 
Crossman 1973). 

Reported substrates at collection sites include 
rock, rubble, gravel sand, silt, clay, mud, muck and 
detritus in areas where aquatic vegetation ranged 
from absent to some submerged macrophytes pre- 
sent. Many of the differences in habitat among col- 
lection sites are probably related to capture during 
different live history stages, food availability or food 
preferences of individual stocks. 


General Biology 
Reproductive Capability 

No information on Lake Whitefish reproduction 
in the Mira and Salmon Rivers is currently available. 
Generally throughout Canada, Lake Whitefish 
spawning occurs in the fall or winter on sand, gravel 
or stones on reefs in lakes or streams and no redds 
are prepared (McAllister and Crossman 1973). In the 
Great Lakes region, spawning usually occurs in 
November and December but in more northern parts 
of Canada, spawning is usually earlier in September 
to October. Differences in time of spawning appear 
to be temperature dependent with first spawning usu- 
ally commencing after water temperatures drop 
below 8°C and peak spawning occurring at even 
lower temperatures (Scott and Crossman 1973). 

In Nova Scotia, Lake Whitefish spawning was 
observed in Scots Lake (44°47’N, 63°11’ W) (for- 
merly Scotch Pond), Petpeswick Lake area [formerly 
Petpeswick River](44°45’N,63°11’W ), Halifax 
County, by Semple (1973). Spawning commenced in 
early December after water temperatures had 
- dropped to 4.5°C and probably continued into late 
December or possibly early January. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Spawning adults generally move to shallow areas 
near shore and spawn in water less than 7.6 m deep 
(Scott and Crossman 1973; Smith 1985). It appears 
that Lake Whitefish resident in streams remain there 
to spawn and similarly lake resident fish probably do 
not generally move into tributary streams to spawn. 
From observations of Lake Whitefish in Kerr Lake, 
New Brunswick, Smith (1952) suggests that spawn- 
ing occurred in the lake since none were captured in 
trap nets set at the mouths of tributary streams dur- 
ing late summer and fall. It is possible that the fish 
moved upstream to spawn after nets were removed, 
since spawning in areas of Nova Scotia with similar 
climate occurs later, sometimes in late fall or early 
winter (Semple 1973). However, specimens of both 
sexes observed in July had well developed gonads 
suggesting they would have spawned earlier in the 
fall. In Nova Scotia, no mature ripe fish were cap- 
tured until just over a week prior to commencement 
of spawning in mid-December (Semple 1973). 

For Lake Whitefish from Scots Lake (Scotch 
Pond), Nova Scotia, average fecundity was deter- 
mined to be 15 698 eggs/kg of fish (Semple 1973). 
Fecundity was not consistently related to length or 
weight for the 22 fish sampled. Fecundity was lower 
than generally reported in other parts of Canada 
except for Hogans Pond, Newfoundland where it 
was determined to be only 11 650 eggs/kg (Chen 
1967). 

Semple (1973) determined maturity and size of 
Lake Whitefish during the spawning period in Scots 
Lake. Average fork length of spawning adults was 
21 cm and maximum length was 26cm. All fish 
greater than 18 cm FL were mature and members of 
both sexes began maturing at lengths greater than 16 
cm FL. According to age estimates, 80% of all fish 
were mature at age 1, 90% at age 2 and all were 
mature at age 3. Age at maturity in Scots Lake is 
less than that reported from other parts of Canada 
but similar to that of Lake Whitefish in Maine. Sex 
composition was approximately 1:1 male to female 
ratio. 

The rate of growth of Lake Whitefish in Canada is 
generally quite rapid and they generally reach a com- 
mercially acceptable size of over 1 kg between 3 and 
10 yrs (Scott and Crossman 1973). In Scots Lake, 
the largest fish (age 6) weighed only 0.2 kg. The 
average weight for all Lake Whitefish captured in 
the study was 117 g. (Semple 1973), suggesting the 
rate of growth was relatively slow in Scots Lake. 


Species Movements 

Movements are related to both spawning and tem- 
perature preferences. Deeper cooler hypolimnetic 
waters in lakes are preferred during summer (Scott 
and Crossman 1973). Lake Whitefish often move 
into shallow areas in spring (Smith 1985). During 
the fall breeding season spawning adults generally 
move inshore to shallow water areas over hard, stony 


2001 


bottom or sand (Scott and Crossman 1973). Young 
whitefish may remain in shallow water areas where 
they feed primarily on plankton until early summer 
when they generally move into deeper water with 
increasing dependence on benthic organisms (Scott 
and Crossman 1973). Lake Whitefish may remain in 
deeper offshore waters during the warmer summer 
months and then migrate inshore when water tem- 
peratures decline). In Lake Erie, Ohio, they were 
observed moving into inshore reefs when water tem- 
peratures dropped to 10°C and swimming near the 
surface on windless days (Trautman 1981). 

One study suggests that Lake Whitefish may, at 
least occasionally, travel considerable distances. An 
individual from the Great Lakes was reportedly 
recaptured 150 miles from where it was tagged 
(Budd 1957). In northern parts of Canada, Lake 
Whitefish may run to sea (McAllister and Crossman 
1973). There is a possibility that Mira River Lake 
Whitefish may travel out to sea in Mira Bay, perhaps 
to feed (J. Gilhen; personal communication), 
although this has not yet been confirmed through 
analysis of stomach contents or by checking scales 
for evidence of anadromy. 


Behaviour/Adaptability 

Edge (1987) studied the diet of Lake Whitefish in 
the Canadian Maritimes and found that there was 
considerable variation between populations, ranging 
from selection of almost exclusively planktonic 
organisms to almost exclusively benthic organisms. 
This is consistent with what has been observed in 
other parts of Canada. Planktonic organisms and 
even terrestrial insects form the major part of the diet 
of some populations of Lake Whitefish (Scott and 
Crossman 1973). Most Lake Whitefish in Nova 
Scotia, including the Mira River population, appar- 
ently fed primarily on benthic organisms. Mira River 
Lake Whitefish stomach contents were found to con- 
tain the following food items in order of abundance: 
Amphipoda, sphaeriid clams, insect larvae, Clado- 
cera, Gastropoda, Hydracarina, and Ostracoda (Edge 
1987). Small sample size may account for the lack of 
other food items, such as terrestrial insects or fish, 
found in stomach contents (Edge 1987) or it may be 
related to timing of collections, food preferences or 
seasonal variability in food selection. 

Fish species found in association with Lake 
Whitefish in Nova Scotia included Atlantic salmon, 
Salmo salar, Rainbow Smelt, Osmerus mordax, White 
Sucker, Catostomus commersoni, Brown Bullhead, 
Ameiurus nebulosus, White Perch, Morone ameri- 
cana, Yellow Perch, Perca flavescens, American Eel, 
Anguilla rostrata, and Alosa sp. (Edge 1987). In the 
Mira River, Lake Whitefish were found in association 
with Atlantic Salmon, Rainbow Smelt, White Perch, 
White Sucker in gill net catches (Edge 1987). The 
Alewife, Alosa pseudoharengus, was also captured in 
gill nets in the Mira River. 


GOODCHILD: STATUS OF MIRA RIVER LAKE WHITEFISH 


63] 


Limiting Factors 
Human Disturbance 

Mira River Lake Whitefish populations have pos- 
sibly been exploited. Historically, nets were set in 
the Mira River specifically to catch this species (D. 
MacLean; personal communication). Currently, 
unauthorized gill netting for Lake Whitefish in the 
Mira River appears to be a problem. Recently, speci- 
mens caught illegally were confiscated and these are 
now held in museum collections (J. Gilhen; personal 
communication). High levels of unemployment in 
the region may be contributing to the exploitation of 
fish stocks. There are several large population cen- 
tres (i.e., Syndney, Glace Bay) within approximately 
20 km of the mouth of the Mira River. 

Other whitefish populations are know to be 
exploited in Nova Scotia. Edge (1984) proposed that 
populations of the endangered Atlantic Whitefish 
were being depleted by severe overfishing and 
poaching. 

Perhaps populations of Lake Whitefish are also 
being depleted as a result of angling, either when 
specifically targeted or through incidental catches 
when targeting other species. Unlimited recreational 
fishing for Lake Whitefish is permitted throughout 
Nova Scotia when the open season for Atlantic 
Salmon, trout (Rainbow Trout, Oncorhynchus my- 
kiss, Brown Trout, Salmo trutta, Brook Trout, Sal- 
velinus fontinalis, Lake Trout, Salvelinus namaycush) 
and Smallmouth Bass, is in effect in particular 
waters. However, there is only conflicting anecdotal 
information available regarding possible captures of 
Lake Whitefish from either the Mira or Salmon 
Rivers in recent years. There is very little information 
available since the Lake Whitefish is not considered 
an important species throughout most of Nova Scotia. 
This is the opposite of what occurs in many other 
parts of Canada where Lake Whitefish is a major 
recreational and commercial species. 

Creel census catch statistics for Nova Scotia do 
not include Lake Whitefish; however, the Mira River 
area of Cape Breton Island appears to be one of the 
more heavily fished areas of Nova Scotia (ASE 
Consultants Inc. 1995). Cape Breton County repre- 
sents 11% of the total angling effort (measured in 
days fished) for 18 counties in Nova Scotia, the 
majority of effort from resident anglers (Nova Scotia 
Department of Fisheries and Aquaculture 1997). 
Greater angling effort only occurs in two other 
Counties, Halifax and Guysborough which account 
for 15.2% and 11.6% respectively. The majority of 
inland fishing effort in Cape Breton County is direct- 
ed primarily towards Brook Trout and Rainbow 
Trout followed by Smelt, Brown Trout, White Perch 
and Yellow Perch (Nova Scotia Department of 
Fisheries and Aquaculture 1997). 


Habitat Loss/Environmental Contamination 
Since there is so little known about the habitat 


632 


preference of the Mira River population, it difficult 
to determine if habitat degradation is occurring or 
affecting populations in the Mira and Salmon 
Rivers. Similarly, there is very little information 
regarding Lake Whitefish populations in Nova 
Scotia. However, the related Atlantic Whitefish was 
probably once a more widespread or wide ranging 
species in coastal waters of northeastern North 
America and populations have probably been 
reduced by the effects of deforestation, dam build- 
ing, over-exploitation and introductions on aquatic 
ecosystems (W. B. Scott, Huntsman Marine 
Science Centre, St. Andrews, New Brunswick; per- 
sonal communication). Assuming some of the cur- 
rent Lake Whitefish populations in Nova Scotia are 
native, they may also have been reduced by similar 
ecological factors. Evans et al. (1988) suggest that 
decline of Lake Simcoe Whitefish may have been 
partly related to environmental factors such as 
eutrophication and contaminants. However, these 
factors are probably not significantly affecting pop- 
ulations in the Mira and Salmon Rivers. 

In many parts of Nova Scotia, particularly in 
southern and southwestern areas, acidification of 
surface waters is a problem that has affected fish- 
eries, especially salmonids. Edge (1984) expressed 
concern over increasing acidification of rivers affect- 
ing survival of Atlantic Whitefish. Although the 
Cape Breton Island highland area is considered to be 
sensitive to the effects of acid rain, surface waters 
throughout Cape Breton are greater than pH 5.4 
(Davis and Browne 1998) and therefore it is unlikely 
that acidification is affecting fish productivity in the 
Mira River area. 


Predation 

There are no known reports of predation by other 
species on Lake Whitefish in Nova Scotia. 
Throughout Canada, a number of fish species report- 
edly prey on Lake Whitefish (Scott and Crossman 
1973) but of these only Lake Trout and Yellow 
Perch, Perca flavescens, are found in Nova Scotia. 
Lake Whitefish also reportedly consume their own 
eggs (Scott and Crossman 1973). American Eel, 
Anguilla rostrata,may also prey on Lake Whitefish. 
American Eels are important predators of young 
salmon, trout and other fish including cyprinids 
(Scott and Crossman 1973), although there are no 
known reports of predation on Lake Whitefish. 


Parasites 

No parasites have been reported from Lake 
Whitefish found in the Mira River. Lake Whitefish 
elsewhere in Canada are host to a large number of 
parasitic organisms (Hoffman 1967; Lawler 1970; 
Scott and Crossman 1973). In Kerr Lake, New 
Brunswick, Lake Whitefish were reportedly being 
‘heavily scared by eels (Smith 1952), possibly a ref- 
erence to Sea Lamprey, Petromyzon marinus. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Special Significance 

The Mira River Whitefish is known only from the 
Mira and Salmon Rivers, Cape Breton Island. 
Whether it represents a genetically discrete stock of 
Lake Whitefish has not been adequately determined 
and it is unclear whether it occurs in the area as a 
result of previous stocking efforts or has arrived 
there as a result of post-glacial dispersal. In any case, 
it represents an interesting example of the plasticity 
of members of the Coregonus clupeaformis “species 
complex”. 

In other parts of Canada, Lake Whitefish are con- 
sidered a valuable recreational and commercial 
freshwater species. However, they are susceptible to 
environmental deterioration and exploitation. 
Generally throughout Nova Scotia, Lake Whitefish 
are not considered to be an important recreational or 
commercial fish, except perhaps in southeastern 
Cape Breton Island, where there appears to be a lim- 
ited fishery for Lake Whitefish from the Mira River. 


Evaluation 

If the Mira River population of Lake Whitefish is 
determined to be an indigenous and discrete stock of 
Lake Whitefish, then it would warrant some form of 
protection because it is geographically and reproduc- 
tively isolated, has an extremely limited distribution, 
and has unique characteristics. Unless further genetic 
studies indicate differently, no emendation of the sci- 
entific name is suggested or required since distinct- 
ness is probably at the population level and not rep- 
resentative of a new subspecies or species. 

There is insufficient scientific information avail- 
able for status designation at this time. 


Acknowledgments 

Financial support was provided by the World 
Wildlife Fund (Canada). I am grateful to Erling 
Holm, Royal Ontario Museum for access to collec- 
tion records and verification of species identifica- 
tions. I would also like to thank the following indi- 
viduals who provided information on Lake Whitefish 
in Nova Scotia: Thomas Edge, Environment Canada; 
Don MacLean, Robert Bancroft, Nova Scotia 
Department of Fisheries and Aquaculture; Mark 
Elderkin, Nova Scotia Department of Natural 
Resources; John Gilhen, Andrew Hebda, and Leslie 
Pezzack, Nova Scotia Museum of Natural History. 


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Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, 
E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. 
Common and scientific names of fish from the United 
States and Canada. American Fisheries Society Special 
Publication 20. 183 pages. 


634 


Roland, A. E. 1982. Geological background and physiog- 
raphy of Nova Scotia. Institue of Science, Halifax, Nova 
Scotia. 311 pages. 

Scott, W. B. 1967. Freshwater fishes of eastern Canada. 
Second edition. University of Toronto Press, Toronto, 
Ontario. 137 pages. 

Scott, W. B. 1987. A new name for the Atlantic whitefish: 
Coregonus huntsmani to replace Coregonus canadensis. 
Canadian Journal of Zoology 65: 1856-1857. 

Scott, W. B., and E. J. Crossman. 1964. Fishes occurring 
in the freshwaters of insular Newfoundland. Department 
of Fisheries, University of Toronto, Royal Ontario 
Museum Contribution Number 58. 124 pages. 

Scott, W. B., and E. J. Crossman. 1973. Freshwater fish- 
es of Canada. Fisheries Research Board of Canada Bul- 
letin 184: 1-966. 

Scott, W. B. and M. G. Scott. 1988. Atlantic Fishes of 
Canada. University of Toronto Press in cooperation with 
the Minister of Fisheries and Oceans and the Canadian 
Government Publishing Centre, Supply and Services 
Canada. 731 pages. 

Semple, J. R. 1973. The lake whitefish, Coregonus clu- 
peaformis (Mitchell) of Scotch Pond, Petpeswick River, 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Nova Scotia. Transactions Canadian Society of Environ- 
mental Biology: 21. 

Smith, C. L. 1985. The incland fishes of New York State. 
New York State Department of Environmental Conser- 
vation. Albany, New York. 522 pages. 

Smith, M. W. 1952. The lake whitefish in Kerr Lake, New 
Brunswick. Journal of the Fisheries Research Board of 
Canada 8: 340-346. 

Todd, T. N. 1997. [Abstract] Environmental modification 
of gill raker number in Coregonine fishes. American 
Society of Ichthyologists and Herpetologists, 77th 
Annual Meeting, Seattle, Washington, June 26 — July 2, 
1997. 

Trautman, M. B. 1981. The fishes of Ohio with illustrat- 
ed keys. Revised edition. Ohio State University Press, 
Columbus, Ohio. 782 pages. 

Williams, J. E., J. E. Johnson, D. A. Hendrickson, S. 
Contreras-Balderas, J. D. Williams, M. Navarro- 
Mendoza, D. E. McAllister, and J. E. Deacon. 1989. 
Fishes of North America endangered, threatened, or of 
special concern: 1989. Fisheries 14(6): 2— 20. 


Accepted 19 February 2002 


Updated Status Report on the Endangered Atlantic Whitefish, 
Coregonus huntsmani* 


THOMAS A. EDGE! and JOHN GILHEN?2 


‘National Water Research Institute, Environment Canada, Burlington, Ontario L7R 4A6 Canada 
2Nova Scotia Museum of Natural History, Halifax, Nova Scotia B3H 3A6 Canada 


Edge, Thomas A., and John Gilhen. 2001. Updated status report on the endangered Atlantic Whitefish, Coregonus hunts- 
mani. Canadian Field-Naturalist 1115(4): 635-651. 


The Atlantic Whitefish, Coregonus huntsmani, is an endemic Canadian species known only from the Tusket and Petite 
Riviére watersheds in southwestern Nova Scotia. The species is anadromous in the Tusket River and its tributary the 
Annis River. It has declined drastically in these rivers as a result of factors related to a hydro-electric dam, acidification, 
overfishing and poaching. The construction of a hydro-electric dam on the Tusket River has prevented Atlantic 
Whitefish from migrating upstream to spawn and exposed them to poachers in fish ladders. More recently the spread of 
Chain Pickerel, Esox niger, has contributed to the decline of Atlantic Whitefish in the Annis River. The decline in the 
Tusket River watershed has continued since the last COSEWIC report in 1983, although the report of an Atlantic 
Whitefish in the Tusket River in 1996 may indicate a remnant population still exists. Atlantic Whitefish have been regu- 
larly caught by anglers in the Petite Riviere watershed since the late 1800s although population trends are uncertain. 
Atlantic Whitefish are landlocked in three lakes in this watershed and there appears to be a small population in the lower 
Petite Riviére. The Atlantic Whitefish populations in the Petite Riviere lakes may be threatened by acidification and the 
spread of non-indigenous fish predators like Smallmouth Bass. Field studies in 1982—1985 provided some knowledge of 
the habitat and biology of Atlantic Whitefish although the ecological requirements of the species are still poorly known. 
The Atlantic Whitefish Conservation and Recovery Team was created in 1999 and is actively investigating requirements 
to conserve the species. The Atlantic Whitefish is threatened with imminent extinction and its status should remain as 
endangered. 


Key Words: Coregoninae, whitefish, Atlantic Whitefish, Corégone atlantique, Coregonus huntsmani, rare and endangered 


fishes, Tusket River watershed, Petite Riviere watershed, Nova Scotia. 


The status of the Atlantic Whitefish, Coregonus 
huntsmani,! was first reviewed following a field sur- 
vey conducted in Nova Scotia in the fall of 1982 
(Edge 1984). This field survey found Atlantic 
Whitefish surviving in the Annis River, Yarmouth 
County, and in three lakes in the Petite Riviére water- 
shed, Lunenburg Co. The species appeared to be 
threatened with extinction and the Committee on the 
Status of Endangered Wildlife in Canada designated 
the Atlantic Whitefish as an endangered species in 
1983 (McAllister et al. 1985; Campbell 1997). 

The status of the Atlantic Whitefish has not been 
updated since its original designation as an endan- 
gered species in 1983. The systematics, distribution, 
ecology and zoogeography of the Atlantic Whitefish 
were studied in subsequent years, including addition- 
al field studies of its habitat in the Petite Riviére 
lakes during 1983-85 (Edge 1987). The results of 
these field studies and other more recent information 
obtained from fisherman, residents and the Atlantic 
Whitefish Conservation & Recovery Team 
(AWCRT) are summarized in this update on the sta- 
tus of the Atlantic Whitefish. 


*Reviewed and approved by COSEWIC November 2000, 
status assigned — Endangered, no change. 


Description 

The Atlantic Whitefish, Coregonus huntsmani, 1s 
a member of the whitefish subfamily (Coregoninae), 
trout family (Salmonidae) (Figure 1). It has silvery 
sides, a white belly, and a back that is dark bluish- 
black or dark green. The Atlantic Whitefish can be 
readily distinguished from most other trout-like fish- 
es by its larger more prominent scales. The species is 
described by Scott and Scott (1988) as having more 
than 90 scales in the lateral line, a terminal mouth 
(lower jaw and upper jaw equal) and small but well 


[The species was first described as the Atlantic Whitefish, 
Coregonus canadensis, by Scott (1967). This species name 
was subsequently found to be preoccupied by D.E. 
McAllister and the replacement name C. huntsmani was 
proposed by Scott (1987). A number of publications have 
referred to the species by the more geographically precise 
common name Acadian rather than Atlantic (e.g., Edge, 
1984; McAllister et al., 1985; McAllister, 1990; Edge et al., 
1991; Bernatchez et al., 1991; Coad, 1995). However, sci- 
entists, government officials, and residents from the Petite 
Riviére and Tusket River communities met in Halifax, 
Nova Scotia in September 1999 to discuss the status of the 
species and concluded the species should be referred to by 
the common name Atlantic Whitefish as outlined in Robins 
et al. (1991). 


635 


636 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


FIGURE 1. The Atlantic Whitefish, Coregonus huntsmani (by permission of Don E. McAllister). Drawn by P. 
Drukker Bramwell, 1984. 


developed teeth on the tongue, and premaxillaries, 
vomer, palatines, and lower jaw bones at all sizes. 
Edge et al. (1991) found Atlantic Whitefish to have 
25-29 gill rakers. 

The Atlantic Whitefish has been confused in the 
past with the Lake Whitefish, Coregonus clupea- 
formis, which is also found in Nova Scotia. However, 
the Atlantic Whitefish and Lake Whitefish have been 
clearly distinguished based upon both morphological 
and genetic characteristics. Scott and Scott (1988) 
indicate that Lake Whitefish lack teeth on premaxil- 
laries, palatines and vomer bones except in juvenile 
fish under 100mm. Edge et al. (1991) found the best 
distinguishing characteristics for Atlantic Whitefish 
were its more terminal mouth and a higher number of 
vertebrae (64-67, x = 65.3) than Lake Whitefish from 
the Canadian Maritime Provinces and the U.S. State 
of Maine. The Lake Whitefish from this area were 
found to have a subterminal mouth and 58-64 (x = 
60.6) vertebrae. The number of lateral line scales was 
also useful in distinguishing 93% of the whitefish 
specimens examined (Figure 2). Atlantic Whitefish 
had a higher number of lateral line scales (88-100, x 
= 93.8) than Lake Whitefish (63-95, x = 76.6). Other 
morphological characteristics for distinguishing 
Atlantic Whitefish and Lake Whitefish, were provid- 
ed by Edge et al. (1991) including comparative pho- 
tographs. Genetic characteristics have also proven 
useful for distinguishing Atlantic Whitefish from 
Lake Whitefish. Bernatchez et al. (1991) found 
Atlantic Whitefish possessed a unique isozyme and a 
mitochondrial DNA genotype different from that of 
Lake Whitefish which indicated that these two 
species are genetically distinct. 


Distribution 
North American and Canadian Range 

The Atlantic Whitefish is an endemic Canadian 
species known only from southwestern Nova Scotia 


(Figure 3). Field studies conducted in 1982, 1983 
and 1985 found the Atlantic Whitefish restricted to 
two disjunct watersheds: the Tusket River water- 
shed, Yarmouth County, and the Petite Riviere 
watershed, Lunenburg County (Edge 1987). The 
populations in the Tusket River, and its tributary 
the Annis River, are anadromous and known to 
venture into seawater. The populations in the Petite 
Riviere watershed occur in three lakes (Mina- 
mkeak, Milipsigate, and Hebb lakes) (Figure 4). 
Atlantic Whitefish have been angled in Fancy Lake 
although it is not known whether there is a resident 
population there. What appears to be a small 
anadromous population exists in the lower Petite 
Riviere. 

Populations of the Atlantic Whitefish have not 
been reported outside the Tusket and Petite Riviére 
watersheds despite extensive commercial and recre- 
ational fisheries in fresh and coastal waters through- 
out Nova Scotia. In addition, extensive province- 
wide fish surveys have failed to identify new 
Atlantic Whitefish populations outside these two 
watersheds. For example, fish populations were sur- 
veyed in 744 lakes throughout Nova Scotia between 
1964 and 1981 by the federal Department of 
Fisheries and Oceans, the Canadian Wildlife Service, 
and the Nova Scotia Department of Lands and 
Forests (Alexander et al. 1986). While 14 lakes were 
found to contain Lake Whitefish, there were no 
reports of Atlantic Whitefish. Similarly, fish surveys 
conducted for the Atlantic Whitefish throughout 
Nova Scotia in 1982 and 1983 found four lakes con- 
taining Lake Whitefish, but Atlantic Whitefish were 
not found outside of the Annis and Petite Riviére 
watersheds (Edge 1987). 

At one time, the Atlantic Whitefish may have 
occurred in the Medway River watershed. Mina- 
mkeak Lake, the uppermost lake containing Atlantic 
Whitefish in the Petite Riviére watershed, previously 


2001 


Lake whitefish 
n= 198 


F 
r 
5 
q 
u 
e 
n 
Cc 
y 


63 65 67 69 7I 73 75 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


77 #79 8! 
-64 -66 -68 -70 -72 -74 -76 -78 


637 


Atlantic whitefish 
n= 32 


83 85 87 89 91 93 958 97 99 


-8O -82 -84 -86 -88 -90 -92 -94 -96 -98 -I00 


Lateral line scales 


FIGURE 2. The frequency of lateral line scale counts on Atlantic Whitefish and Lake Whitefish specimens from the 
Canadian Maritime Provinces and the State of Maine, U.S.A. 


flowed into the Medway River. Dynamite was used 
to create an outflow from Minamkeak Lake into 
Milipsigate Lake around 1905, and the original out- 
flow of Minamkeak Lake into the Medway River 
watershed was blocked. It is not known whether the 
population of Atlantic Whitefish in Minamkeak Lake 
occurred there before this diversion or whether it is 
the result of upstream dispersal from Milipsigate 
Lake. 

Atlantic Whitefish specimens have been caught 
on five occasions outside the Tusket and Petite 
Riviere watersheds. A specimen was caught in sea- 
water on 12 June 1940, in Yarmouth Harbour, 
Yarmouth County, and another on 31 May 1958, ina 
herring weir at Halls Harbour, Kings County. Two 
specimens are also reported to have been caught in 
full seawater at the mouth of the Sissiboo River, 
Digby County on 8 September 1919 (Scott and Scott 
1988). More recently, an Atlantic Whitefish speci- 
men was caught in the smelt fishery in the Lahave 
River estuary, Lunenberg County in February, 1995 
(D.R. Bell, in litt., February 1997) and another one 
was caught there on 24 May 1997 (A. Hebda, per- 
sonal communication, March 1998). The Lahave 
River watershed is adjacent to the Petite Riviére 


watershed. The isolated nature of all these captures 
as well as the migratory habits and salt water toler- 
ance of Atlantic Whitefish would suggest these spec- 
imens were likely strays from the Tusket and Petite 
Riviere watersheds. 

While the Atlantic Whitefish appears to be 
restricted to the Tusket and Petite Riviére water- 
sheds, it should be noted that populations of white- 
fish in the Maritimes Region have gone undetected 
for long periods of time. Smith (1952) found a pop- 
ulation of Lake Whitefish in Kerr Lake, New 
Brunswick, that was poorly known among local 
anglers and residents. Similarly, by setting gillnets 
in the deeper waters of Nova Scotian lakes, Edge 
(1987) found Lake Whitefish populations that were 
unknown to local anglers, residents or fishery offi- 
cers. Deep water habitat preferences, small popula- 
tion sizes, and poor angling potential may enable 
whitefish populations to evade detection. This 
should be borne in mind for future lake surveys for 
Atlantic Whitefish, since Alexander et al. (1986) 
indicated that a limitation of their provincial lake 
surveys was the likely undersampling of deeper 
waters (> 10m) because of the difficulty of setting 
gillnets in these habitats. 


638 


Tusket » 
River - 


Annis 
River 


FIGURE 3. Distribution of the Atlantic Whitefish in Nova 
Scotia. The Tusket, Annis and Petite Riviére water- 
sheds known to support Atlantic Whitefish are shown 
as well as four records (@) outside these watersheds 
where specimens have been captured. 1 = Hall’s 
Harbour, Kings County, 1958; 2 = Sissiboo River 
mouth, Digby County, 1919; 3 = Yarmouth Harbour, 
Yarmouth County, 1940; 4 Lehave River mouth, 
Lunenberg County, 1995/97. 


Habitat 
Definition 

(1) Tusket River Watershed. Very little is 
known about the habitat requirements of the 
Atlantic Whitefish in the Tusket River watershed. 
The Atlantic Whitefish is known to be anadromous 
in the Tusket River and its tributary, the Annis 
River. The Tusket River watershed provides an 
extensive estuarine habitat for the Atlantic 
Whitefish and the species has been commonly 
caught in these brackish waters in the past. Atlantic 
Whitefish are known to venture into full seawater 
and specimens have been caught off Wedgeport, 
Yarmouth County, at a salinity of 31 ppt (Scott and 
Scott 1988). 

Atlantic Whitefish are not known to enter fresh 
water much above the influence of tidal waters in the 
Tusket and Annis rivers. In the past, Atlantic 
Whitefish have been captured in winter through the 
ice in Lake Vaughn just above the Tusket River dam 
(Patrick Patten, personal communication, March 
2000). While the spawning and nursery requirements 
for Atlantic Whitefish are unknown, Smith (1962) 
indicated the Tusket River provided over 138 000 m2 
of spawning areas and over 668 000 m2 of nursery 
areas for Atlantic Salmon. A potential annual return 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


FIGURE 4. The Petite Riviére watershed, Lunenburg County, 
Nova Scotia, showing water chemistry sampling sta- 
tions in 1983-85 (i — St. George Brook; 2 — Sull 
Brook; 3 — Frederick Lake outlet; 4 — Birch Brook; 5 
— Wildcat Creek; 6 — Petite Riviere at Conqueral 
Mills). 


of between 1600 and 3200 adult salmon was consid- 
ered possible from the river although the annual run 
was considered very small by the 1960s. 

The Tusket River is quite acidic compared to 
many other rivers in Nova Scotia (Farmer et al. 
1980; Watt et al. 1983). Farmer et al. (1980) sur- 
veyed the Tusket River at Gavelton in 1979-1980 
and found pH values from 4.55 to 4.85. They consid- 
ered the Tusket River to be one of seven mainland 
Nova Scotian rivers unsuitable for the successful 
reproduction of Atlantic Salmon. They also suggest- 
ed that a small salmon population had probably sur- 
vived in the Tusket River because water quality had 
remained acceptable in the Carleton River tributary. 
Watt et al. (1983) reported the mean pH of the 
Tusket River to be 4.8 in 1980-81 (4.6 in the East 
Branch). 

Water samples collected from the Tusket River 
area in 1983 and 1984 were analyzed by the Depart- 
ment of Fisheries and Oceans laboratory in Halifax 
(Edge 1987). These water samples were obtained 
from the Annis River and Tusket River (at Lake 
Vaughn reservoir) just above the influence of tidal 
waters in September 1983, and January 1984. In 
September 1983 when Atlantic Whitefish would 
begin migrating into fresh water, the pH of the 
Tusket River at Lake Vaughn was 5.23 and the pH 
of the Annis River was 6.05. In January 1984, the 
pH at Lake Vaughn was 4.59 and the pH of the 


2001 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


639 


Oo Minomkeak Loke 
@ Milipsigate Lake 
& Hebb Lake 


o Petite R. (Conqueral) 


7.0 ° 
@ St. George Brook 
a A Still Brook 
6.0 Gc Oa 6 
pH ee ee apa 8 


4.0 


7.0 


a 
& a 
—_— ee, 


© Birch Brook 
@ Frederick L. Outlet 
& Wildcat Creek 


M J 


Jv AS ON D JS F M A M 


F/85 


Month 


FIGURE 5. Seasonal water pH variation throughout the Petite Riviere watershed, Lunenburg County, Nova 
Scotia from 1983 to 1985. See Figure 4 for water sampling locations. 


Annis River was 5.02. pH data from the Tusket 
River were collected around 1980 in the Carleton 
tributary and 1995-96 in the main Tusket branch 
(Wilsons Bridge) (data from ENVIRODAT (Atlantic 
Database Water Chemistry Data). These data indi- 
cate the average annual pH in the Carleton River 
based upon 11 monthly samples was 5.65 in 1980. 


The average annual pH in the main Tusket branch 
based upon 12 monthly samples was 4.68 in 1995, 
and 4.64 based upon seven monthly samples in 1996. 
Low pH values like those in the Tusket River east 
main branch have been shown to be detrimental to 
the survival of Atlantic Salmon (Lacroix and Towns- 
end 1987), and they have probably had an adverse 


640 THE CANADIAN FIELD-NATURALIST Volv 15 
© Minamkeok Lake 
@ Milipsigate Lake 
& Hebb Lake 
4 Q 
Bi fe a 
: 4.0 
k ° © Petite R. (Conqueral) 
a @ St George Brook 
| 30 A Still Brook 
j 
n 
{ 2.0 ° 
Voted 
1.0 ° e@ 
/L ° o 
mg e\, = 
e=6 ° ° 
CaCos 0.0 s—A—s— 4-4 a— 
3.0 
oO Birch Brook 
@ Frederick L. Outlet 
2.0 A Wildcat Creek 
1.0 
e—e—e 
0.0 Neate cece qn 
i A.Ss "O° WN <8 dot ol fea F/85 


Month 


FIGURE 6. Seasonal water alkalinity (mg/l of CaCO,) variation throughout the Petite Riviére watershed, 
Lunenburg County, Nova Scotia from 1983 to 1985. See Figure 4 for water sampling locations. 


impact on the ability of Atlantic Whitefish to repro- 
duce.in the Tusket River. 


(2) Petite Riviére Watershed. The Atlantic 
Whitefish has adapted to a completely freshwater 
_ habitat in lakes within the Petite Riviére watershed. 
The area around the Petite Riviére was first settled 


in the 1600s and the subsequent building of mill 
dams along the river was extensive (Dunfield 1985). 
While these mill dams would have obstructed the 
river to varying degrees, the construction of a 
hydro-electric dam at the foot of Hebb Lake as early 
as 1901 effectively blocked any upstream migration 
of fishes. There is no record of fish ladders or any 


2001 


means to allow fish passage upstream at this point 
since construction of the dam. One of the earliest 
saw mills in the Petite watershed was formerly 
located at the site. If Atlantic Whitefish in the Petite 
Riviere lakes were once anadromous, these popula- 
tions have had no access from the sea since at least 
1901. At present, dams without fish ladders obstruct 
the Petite Riviére at the outlets of Minamkeak, 
Milipsigate and Hebb lakes. A power dam built 
around 1939 at Conqueral Mills below Fancy Lake 
was breached in 1977. 

The Petite Riviére watershed drains an area of 
about 233 square kilometers. Farms and a few 
small towns are found within the watershed, 
although forested areas predominate, particularly 
surrounding the lakes. A characteristic of the 
watershed is the abundance of bogs which impart 
a tea-colour to the water. Bathymetric surveys of 
Minamkeak, Milipsigate, and Hebb lakes indicated 
that these lakes had maximum depths in 1983 of 
13m, 16m, and 14m respectively, although much 
of the area of these lakes is more shallow. The 
lake bottoms are silt in the deeper water areas, 
whereas shoals and shoreline areas are rocky. A 
study of the temperature profile in Hebb Lake in 
1983 indicated the warm water nature of this lake. 
While the lake did stratify to a degree during the 
summer, a typical coldwater hypolimnion was not 
present. The temperature at the bottom of the lake 
in May was 14°C, and it rose to almost 20°C by 
late August. 

The Petite Riviere has been less affected by acidi- 
fication than the Tusket River. Watt et al. (1983) 
indicated the river had a mean pH of 5.6 between 
1980-81. Watt (1986) categorized the Petite Riviére 
as a river where Atlantic Salmon stocks had not yet 
been affected by acidification and where there was 
no sign of an impact of acidification on angling 
returns. 

Water chemistry data were collected from the 
Petite Riviére watershed in 1983-84 and analyzed by 
the Department of Fisheries and Oceans laboratory 
in Halifax (Edge, 1987). The Petite Riviere showed a 
trend in seasonal pH variation similar to other rivers 
in Nova Scotia (Figure 5). pH was high over the 
spring and summer and then fell quickly in the fall to 
a midwinter pH minimum. pH also varied widely 
throughout the watershed such that while it was usu- 
ally above 5.5 in the lower Petite Riviére at 
Conqueral, pH was rarely above 4.5 in Still Brook, a 
darkly coloured brook draining a bog area. The lakes 
within the Petite Riviére watershed usually had a pH 
above 5.0. During February 1985, however, pH val- 
ues were recorded from Hebb and Minamkeak lakes 
as low as 4.5 and 4.8 respectively, indicating pH 
fluctuations which could adversely affect aquatic 
life. pH data from the lower Petite Riviére available 
from ENVIRODAT (Atlantic Database Water 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


641 


Chemistry Data) indicate the average annual pH in 
the Petite Riviere based upon 11 monthly samples 
was 5.62 in 1985. 

Alkalinity values, indicative of the buffering capa- 
city of water, also varied considerably throughout the 
watershed (Figure 6). Alkalinity was usually 
detectable in the Petite Riviere at Conqueral whereas 
Sull Brook lacked measurable alkalinity throughout 
the year. The Petite Riviere lakes had low alkalinity 
values (usually below 1.0 mg/l of CaCO.,), particular- 
ly over the winter, and alkalinity was not measurable 
in Minamkeak and Hebb lakes in February 1985. It 
remains uncertain how the Petite Riviere watershed 
will retain its buffering capacity to counter acidifi- 
cation. 

An indication of habitat preferences of Atlantic 
Whitefish in the Petite Riviere lakes is provided by 
gillnet capture data (Table 1). Based upon catch per 
unit effort, Atlantic Whitefish were caught more than 
twice as often in surface waters compared to midwa- 
ter or bottom habitats. This finding was in contrast to 
other fish species which were found to be more 
abundant near the bottom than in surface waters. 
Atlantic Whitefish caught at the surface comprised 
27% of the catch of all fishes whereas midwater and 
bottom captures comprised 20% and 2%, respective- 
ly. Those Atlantic Whitefish caught near the bottom 
were almost invariably caught in the deeper parts of 
the lakes. Only 7% of the bottom catches of Atlantic 
Whitefish were made in gillnets set on the bottom at 
a depth of less than 8 m. 

Habitat requirements for Atlantic Whitefish in the 
lower reaches of the Petite Riviére are poorly 
known. It is uncertain if this is a resident anadro- 
mous population or whether Atlantic Whitefish have 
continuously passed downstream over the Hebb 
Lake dam to the sea. While Atlantic Whitefish have 
been regularly caught over many years in the brack- 
ish waters of the Petite Riviére estuary, gaspereau 
fishermen from this area have not heard of captures 
in fresh water in the lower Petite Riviére nor in near- 
by seawaters (D.R. Bell, personal communication, 
1997). 


Trends 

The Tusket River hydro-electric dam has signif- 
icantly changed the habitat of the Atlantic 
Whitefish since it was built in 1929. It is uncertain 
what impact the current dam and associated fish 
ladders are having on any remaining Atlantic 
Whitefish. However, there continue to be concerns 
about inadequate waterflows in fish ladders during 
Atlantic Whitefish migration periods. There con- 
tinues to be threat of habitat loss through acidifica- 
tion in both Tusket and Petite Riviére watersheds. 
In addition, the recent spread of Chain Pickerel in 
the Annis River and introduction of Smallmouth 
Bass into the Petite Riviére watershed have posed 
significant new threats to maintaining suitable 


642 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


TABLE 1. Habitat preferences of Atlantic Whitefish and other fish species in the Petite Riviere lakes, Nova Scotia, 


expressed as number of specimens caught in gill nets. 


Fish species 


gill net Coregonus Catostomus Ameiurus Morone Perca 
habitat effort (hrs) huntsmani commersoni nebulosis americana flavescens 
Fall 1982 
Surface 60 12 0 0 a2 12 
Midwater 48 3 1 0 6 Th 
Bottom < 8 m 85 1 68 9 129 7 
Bottom > 8 m 115 8 64 5 270 40 
Spring 1983 
Surface 12 5 0 ] 0 0 
Midwater 1 2 DZ, 0 4 0 
Bottom > 8 m i, 8 2 0 5 0 
Summer 1983 
Bottom < 8 m 36 0 10 i 18 0 
Bottom > 8 m 18 3 il 5 0) 
Total 
Surface 78 iW 0 1 By 12 
Midwater 60 5 3 0 10 7 
Bottom < 8m 121 1 78 10 147 4 
Bottom > 8 m 145 14 76 5) 280 42 


habitat for Atlantic Whitefish. Any potential 
shoreline development or angling activities around 
the Petite Riviere lakes will require careful moni- 
toring. 


Protection/Ownership 

The habitat of the Atlantic Whitefish in the Petite 
Riviére lakes is protected to some degree by provin- 
cial legislation. The town of Bridgewater (population 
7248 in 1991) has received its water supply from 
Hebb Lake since the turn of the century and mainte- 
nance of water quality in the watershed has been a 
concern. The area surrounding Hebb Lake and 
Milipsigate Lake was designated a Protected Water 
Area under provisions of the Water Act in 1964. The 
area around Minamkeak was similarly designated in 
1975. The Minamkeak Lake and Milipsigate-Hebb 
Lakes protected water areas cover approximately one 
half of the total watershed and protect against 
lakeshore development, recreational activities, and 
the dispersal of sewage, biocides or garbage near the 
lakes. In addition, the Petite Riviere Watershed 
Advisory Group has expressed a concern about 
ensuring the protection of Atlantic Whitefish in the 
watershed (Peter Oickle, Bridgewater, in litt., 1985). 
This group was created in 1977 by the Nova Scotia 
Department of Environment to advise the Province of 
Nova Scotia, the Town of Bridgewater, and the 
Municipality of the District of Lunenberg on prob- 
lems and solutions to water quality, levels, and flows 
in the watersheds of Minamkeak Lake, Milipsigate 
Lake, Hebb Lake, Fancy Lake and the Petite Riviere 
system. 


Biology 
General 

Many aspects of the general biology of Atlantic 
Whitefish are still poorly known. There has not been 
a growth study of Atlantic Whitefish, but anadro- 
mous specimens from the Tusket River are known to 
grow larger than the landlocked ones from the Petite 
Riviére watershed. Scott and Scott (1988) indicated 
that Atlantic Whitefish were reported by a fishery 
officer to have been as large as 3.63 kg in the Tusket 
River and that specimens caught in the Tusket River 
salmon trap in 1954 ranged from 0.45 to 2.27 kg 
according to local observers. A fisherman caught an 
Atlantic Whitefish wedged into an eel pot on the 
Annis River in the fall of 1981 that was about 2.5 kg 
and amongst the largest he had ever caught (Limon 
Earl, Lake Pleasant, personal communication, 
October 1982). 

Anglers have reported Atlantic Whitefish speci- 
mens from the Petite Riviére lakes were usually 
17.5cm to 40cm and from 0.57 kg to 0.68 kg 
although rare individuals as large as 45 cm have 
been caught (Piers 1927; Gilhen 1999). Atlantic 
Whitefish in the lower Petite Riviere appear to grow 
larger than the lake specimens. A gaspereau fisher- 
man caught an Atlantic Whitefish in the estuary in 
May, 1996 that was about 50 cm in length and about 
1.82 kg in weight (D.R. Bell, personal communica- 
tion, 1997). The largest specimens studied by Edge 
(1987) from the Tusket and Petite Riviere water- 
sheds were 507 mm and 317 mm in total length, 
respectively. 


2001 


Reproduction 

The exact spawning period and spawning loca- 
tions for Atlantic Whitefish are not known. Atlantic 
Whitefish in the Tusket River probably spawn in the 
late fall or winter. A female specimen caught in the 
Annis River on 12 October 1982, when water tem- 
perature was 12°C, had well-developed ovaries but 
was not yet ready to spawn. A specimen caught at 
the Tusket River dam on 4 November 1967, also had 
well-developed gonads but was not ready to spawn. 
Adult specimens caught in the Tusket River on 24 
May 1940, and 24 June 1966, had poorly developed 
gonads suggesting spawning had occurred. 

Spawning probably takes place in the Petite Riviere 
lakes in the winter months. Atlantic Whitefish in 
Hebb Lake had not spawned as late as 13 November 
1982 when water temperature was 10° C. The gonads 
of these specimens were well developed although the 
fish were not yet ready to spawn. Specimens caught in 
Hebb Lake on 22 May 1983, when water temperature 
was 14° C had much smaller gonads suggesting 
spawning had already occurred. Semple (1973) found 
Lake Whitefish from Scotch Pond, Nova Scotia, start- 
ed to spawn on 10th December when water tempera- 
ture had dropped to 4.5° C. 

Very little information is available on the spawn- 
ing behavior or early life history stages of the 
Atlantic Whitefish. Despite extensive fish surveys in 
the Petite Riviere lakes in the 1980s and many years 
of observations by fishermen in the Tusket and Petite 
Riviére watersheds, young Atlantic Whitefish were 
never noted. Gaspereau fisherman familiar with 
Atlantic Whitefish in the Annis River and the Petite 
Riviere have indicated they have never seen young 
Atlantic Whitefish (L. Earl, personal communica- 
tion, 1983; D.R. Bell, personal communication, 
1997). More recently, members of the Atlantic 
Whitefish Conservation and Recovery Team have 
reported observing juvenile fish in the Petite Riviere 
lakes in the spring of 2000. 


Survival 

There is no information available on survival of 
the Atlantic Whitefish. Young have rarely been 
reported and Atlantic Whitefish population age 
structures have not been studied. Little can be said 
about important factors affecting mortality or recruit- 
ment rates. The lack of ability to find young Atlantic 
Whitefish in field surveys in 1982-1983 could have 
been the result of recruitment failure, or more likely, 
inadequate sampling methods. 


Physiology 

There is no information on the physiology of the 
Atlantic Whitefish. Like some species of Coregonus 
occurring in coastal drainages of the Nearctic and 
Palearctic, it has a capacity to tolerate full seawater. 
It also appears to be able to tolerate relatively warm 
water conditions in the Petite Riviere lakes. Its abili- 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


643 


ty to tolerate low pH at different life history stages is 
unknown. 


Movement 

The Atlantic Whitefish is anadromous in the 
Tusket and Annis rivers. Scott and Scott (1988) indi- 
cated that the whitefish migrate upstream in the 
Tusket River in October and then downstream to the 
estuary in the spring. A former fishery officer in the 
Tusket River area indicated that Atlantic Whitefish 
move upstream to spawn from mid-September to 
early November, and then moved downstream to the 
sea about mid-February to late March, with a few 
stragglers in early April (Gilhen 1977). One fisherman 
on the Annis River called Atlantic Whitefish “frost 
fish” because they usually started to move into fresh 
water around the time of the first frost in the fall. 

Atlantic Whitefish are not known much above the 
Lake Vaughn reservoir in the Tusket River or much 
upstream of the small Pleasant Lake on the Annis 
River, and thus do not appear to migrate much into 
fresh water. A fisherman on the Annis River indicat- 
ed Atlantic Whitefish did not venture as far into 
fresh water as Atlantic Salmon, and that Atlantic 
Whitefish were not known to reach Salmon Lake on 
this river (Limon Earle, Pleasant Lake, October1982, 
personal communication). Scott and Scott (1988) 
indicated that Atlantic Whitefish were often accom- 
panied by Atlantic Salmon on their upstream migra- 
tion in the Tusket River in the fall. 

Atlantic Whitefish are known to move down- 
stream in the Tusket River in the spring and venture 
well out of the estuary into full seawater. Specimens 
have been caught in seawater at Yarmouth Harbour, 
Hall's Harbour, and probably at the mouth of the 
Sissiboo River. Available capture records indicate 
that Atlantic Whitefish are at sea in coastal waters 
during summer months and suggest that they may 
disperse widely in the marine environment from the 
Tusket River. Although the movements of Atlantic 
Whitefish in seawater are not well known, people 
from the Tusket River area believed their move- 
ments were similar to Atlantic Salmon (W. B. Scott, 
in litt., 1965). 

The Atlantic Whitefish populations in the Petite 
Riviere lakes are landlocked and it is not known 
whether they were anadromous at one time or not. 
There are no fish ladders around the dams at the out- 
lets of Minamkeak, Milipsigate and Hebb lakes so 
upstream movements are probably restricted. The 
hydro-electric dam at the foot of Hebb Lake has 
effectively blocked any upstream migration of fish 
since its construction in 1901. While Atlantic 
Whitefish may be swept downstream from Hebb 
Lake, the dam precludes any upstream movements. 
It is unknown if the Atlantic Whitefish in the lower 
reaches of the Petite Riviére have a regular anadro- 
mous migration. 


644 THE CANADIAN FIELD-NATURALIST Vol. 115 


TABLE 2. Feeding preferences of Atlantic Whitefish and Lake Whitefish from Nova Scotia. Values expressed as per cent 
of the number of all food items eaten at each location. 


Atlantic Whitefish Lake Whitefish 

Hebb Milipsigate Minamkeak Tusket Mira Lake Pringle 

Lake Lake Lake River River George Lake 
Food item n=8 n=6 n=4 n=10 ines) n=14 n=7 
Cladocera 45.2 Da eo 0 Ont ONS) 99.8 
Isopoda 0 0 0 0.9 0 2 0 
Amphipoda 0 0 0.1 0.3 32, 1523 0 
Sphaeriid clam 0 0 0) 0 25.4 14.6 0 
Gastropoda 0 0 0 0 1.6 DS 0 
Fish 0 | 0 0.2 0 0 0 
Insect larva 0 2) 0.1 W2LS 19.2 30.9 0.1 
Insect nymph 3 Dy 0.06 24.6 0 1.4 0 
Insect pupa Ihe Dice 0 333 4.2 0.1 
Insect adult 52.4 ile 1.3 0 13.9 0 
Other 0 0 0.04 0.1 9.4 Dai) 0 


The saltwater tolerance of Atlantic Whitefish and 
records of incidental captures outside the Tusket and 
Petite Riviére watersheds indicate there is some 
potential for movements between watersheds in 
Nova Scotia. 


Nutritional and Interspecific Interactions 

Atlantic Whitefish from the Tusket and Petite 
Riviere watersheds are known to feed upon a wide 
variety of food items based upon a stomach content 
analysis of collected specimens (Edge, 1987). 
Atlantic Whitefish captured in Hebb Lake had fed on 
many flying ants (Hymenoptera) while smaller spec- 
imens from Minamkeak had fed mostly on plankton 
(Cladocera). Atlantic Whitefish from Muilipsigate 
Lake had fed most commonly on dragonfly nymphs 
(Odonata), although they had also eaten adult Hemi- 
ptera and beetles (Coleoptera), Cladocera, mayfly 
nymphs (Ephemeroptera), Diptera pupae and Banded 
Killifish (Fundulus diaphanus). Variation in stomach 
contents probably reflects the seasonal variation in 
available food items at the time the Atlantic 
Whitefish specimens had been caught. 

The stomach contents of Atlantic Whitefish from 
the Tusket River differed from that of the Petite 
Riviere lake specimens. Diptera pupae were general- 
ly the most commonly occurring food item for 
Tusket River Atlantic Whitefish, although stonefly 
nymphs (Plecoptera), blackfly larvae (Simuliidae) 
and other diptera larvae were also commonly eaten. 
Some Tusket River specimens had ingested large 
numbers of stonefly nymphs and blackfly larvae. 
Those Atlantic Whitefish caught in brackish or salt 
water had been feeding upon shrimp (Decapoda), 
amphipods, and fish (Ammodytes sp.). Scott and 
Scott (1988) indicated that the stomachs of Atlantic 
Whitefish caught off Wedgeport in seawater were 
‘found to contain the remains of Atlantic Herring 
(Clupea harengus), periwinkles (Littorina littorea), 


amphipods, decapods, and a few blades of eelgrass 
(Zostera marina). Atlantic herring remains were 
thought to have come from feeding on refuse from a 
nearby herring processing plant. Atlantic Whitefish 
captured in the Petite Riviere estuary have been 
reported to have been feeding upon amphipods and 
marine worms (D.R. Bell, personal communication, 
September 1999). 

A comparison of the major food classes consumed 
by Atlantic Whitefish and Lake Whitefish found in 
Nova Scotia lakes is presented in Table 2. While 
there is considerable variation between populations 
within a species, there were notable differences 
between Atlantic and Lake Whitefish in Nova 
Scotia. Whereas Lake Whitefish commonly fed on 
molluscs, Atlantic Whitefish never fed on this food 
item in fresh water. Similarly Lake Whitefish often 
contained many amphipods and isopods whereas 
Atlantic Whitefish very rarely consumed these 
organisms. Atlantic Whitefish were found to feed 
more heavily on insects than Lake Whitefish, partic- 
ularly Coleoptera and winged insects characteristic 
of surface waters. Atlantic Whitefish also fed on 
fish, unlike Lake Whitefish. 

The data on gillnet capture and stomach contents 
indicate that Atlantic Whitefish commonly occur in 
surface waters in the Petite Riviere lakes. They 
rarely consumed typical benthic food items such as 
amphipods, isopods, and molluscs, but instead had 
fed upon plankton, fish and a variety of foods which 
had presumably fallen onto the surface or were per- 
haps taken above the water surface. These data 
appear consistent with behavioral reports about 
Atlantic Whitefish from anglers who have caught 
them with hooks baited with small minnows and 
with a fly usually before the fly touched the water 
(S. E. March, in litt., 1925). 

There is little information about Atlantic White- 


2001 


fish parasites. Atlantic Whitefish in the Petite Riv- 
iére estuary are reported to lack sea lice which are 
found on Atlantic Salmon in the same estuary (D.R 
Bell, personal communication, September 1999). 


Behaviour/Adaptability 

Little is known about the behavior and adaptabili- 
ty of the Atlantic Whitefish. They have been report- 
ed to occur in schools in the Petite Riviere water- 
shed lakes and they have been caught in swift 
currents by anglers using a hook baited with a worm 
or small minnow, or using a small artificial fly or 
tiny natural flies on very small baited hooks (Piers 
1927). The Atlantic Whitefish would usually take 
the bait just below the surface in swift running 
water. They would take a fly from two to six inches 
above the water and occasionally on the surface. 
Anglers have described them as gamey fighters that, 
when hooked, almost always leap from the water 
until exhausted. 

Some anglers have caught Atlantic Whitefish in 
the Petite Riviere lakes early in the spring as the ice 
was leaving, but considered it impossible to catch 
these fish after the ice was gone (D.R. Bell, in litt., 
1997). In the lower Petite Riviere, a gaspereau fish- 
erman has indicated he had never heard of an 
Atlantic Whitefish entering a net during daylight 
hours in the estuary; only after dusk could they be 
caught (D. R. Bell, in litt., 1997). 

Very little is known about the adaptability of 
Atlantic Whitefish and requirements for maintaining 
the species in captivity for re-stocking purposes. 
Some preliminary efforts to maintain Atlantic 
Whitefish at a hatchery were not successful. Four 
Atlantic Whitefish were captured alive from gillnets 
set in Hebb Lake in November 1982. These four spec- 
imens were transported to a Department of Fisheries 
and Oceans fish hatchery on the Mersey River within 
two hours of removal from gillnets. They were trans- 
ported in barrels containing water from Hebb Lake 
that were aerated using a scuba tank. The four 
Atlantic Whitefish were placed in a holding tank (8 
foot diameter) at the hatchery. Two Atlantic Whitefish 
died the following day, while a third survived for sev- 
eral days. The last specimen survived over the winter 
months but died in the spring probably because it 
would not eat hatchery food (T. Goff, personal com- 
munication). More recently in 2000, the Atlantic 
Whitefish Conservation and Recovery Team trans- 
ported whitefish to the Mersey River fish hatchery to 
renew captive breeding efforts with some success. 
More understanding of the biology and reproductive 
requirements of Atlantic Whitefish will be needed to 
develop a successful hatchery program for the species. 


Population Size and Trends 
Atlantic Whitefish have been known to fishermen 
in the Tusket River, Annis River, and Petite Riviére 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


645 


watersheds for many years. The populations of 
Atlantic Whitefish in the lakes and lower reaches of 
the Petite Riviere watershed appear to be small based 
upon available habitat. There is little information on 
population trends. The numbers of Atlantic 
Whitefish in the Tusket River, and its tributary the 
Annis River, have declined drastically in recent 
decades and it is possible the species may have dis- 
appeared from this watershed. There is insufficient 
information available at this time to enable accurate 
estimates of the population sizes and trends of 
Atlantic Whitefish in the Tusket and Petite Riviere 
watersheds. 


(1) Tusket River 

The Atlantic Whitefish was once very abundant in 
the Tusket River watershed (Gilhen 1977; Scott and 
Scott, 1988). Large numbers of these whitefish were 
known to migrate up the river during anadromous 
migrations in the months of October and November, 
and anglers on the river and in Wedgeport and 
Yarmouth harbours considered them abundant. Prior 
to 1940, it was not uncommon for a gaspereau fish- 
erman on the Tusket River to accidentally catch 200 
specimens in a gaspereau net (Gilhen 1977). Scott 
and Scott (1988) indicated that an Atlantic Salmon 
trap located on the Tusket River in 1954 caught the 
following numbers of Atlantic Whitefish moving 
upstream that year: 

18 to 24 October 

2) to 315* October — 47 whitefish; 

1 to 7 November — 15 whitefish. 

The construction of a hydro-electric dam on the 
Tusket River at Tusket falls in 1929 appears to have 
had a significant impact on the abundance of the 
Atlantic Whitefish. The Tusket River dam is near the 
limit of the influence of high tide, and before con- 
struction of the dam, was an area of fast flowing 
rapids just below a lake that supported good fishing 
(Erskine 1971). The Tusket River power facility is 
composed of two dams; the hydro-electric dam 
which generates electricity and, about | km to the 
east, a holding dam which helps to control the water 
levels in the Lake Vaughn head pond reservoir. 
When the hydro-electric dam was originally built, 
there was no means to deter fish from migrating 
through the sluices of the dam and, as a result, many 
Atlantic Whitefish were probably killed by the tur- 
bine blades (Gilhen 1977). 

Studies conducted at the Tusket River hydro- 
electric dam in the autumn of 1960 and also 1961 
indicated that salmon and gaspereau passing through 
the turbines could suffer considerable mortality from 
abrasion, impact or internal haemorrhaging from 
pressure changes. The average mortality rate for 
young salmon descending through the turbines in 
1960 was 16.5% while the rates for descending 
young gaspereau in 1960 and 1961 were found to be 
52.9% and 50.3% respectively (Smith 1960;1961). 


— 24 whitefish; 


646 


Smith (1962) indicated that the Tusket River had a 
fairly good-sized salmon population prior to the dam 
construction but that very few salmon spawners were 
successfully passing the hydro-electric dam. These 
studies conducted at the Tusket River dams in the 
early 1960s did not indicate any knowledge of the 
Atlantic Whitefish or its migrations. 

A series of fish ladders was built over the years to 
permit fish passage around the Tusket River dams, 
although many were probably ineffective for 
Atlantic Whitefish migrations. Smith (1960;1961) 
indicated that the fish ladder at the Tusket River 
hydro-electric dam in the early 1960s was effective 
for gaspereau and that only a small percentage of 
gaspereau likely descended through the turbines. 
However, the fish ladder was considered very inade- 
quate for spawning Atlantic Salmon passage and it 
was suggested as the major cause for reduction in the 
salmon production of the Tusket River (Smith 1962). 
Its entrance was difficult for salmon to find, and 
even when they did find it, they would rarely enter 
because of the relatively small outflow of water. 
While it is not known how effective the fish ladders 
were for Atlantic Whitefish, those that were effective 
exposed the whitefish to poachers. During the early 
1950s, large numbers of Atlantic Whitefish were 
taken by poachers from the upper pools of the fish 
ladder constructed at the hydro-electric dam and, at 
least once, were loaded into a dump truck to be used 
as fertilizer (Scott and Scott 1988). A new fish lad- 
der was placed at the Tusket River holding dam in 
1997. It is believed to be effective, although water 
flows are not adequately maintained during Atlantic 
Whitefish migration periods (Patrick Patten, personal 
communication, March, 2000). 

The Atlantic Whitefish population in the Tusket 
River watershed appears to have declined most 
rapidly during the 1940s and 1950s. Gilhen (1977) 
suggested that the abundance of Atlantic Whitefish 
in the Tusket River started to drop noticeably in the 
1940s due to the installation of more turbines in the 
Tusket River hydro-electric dam, the lack of screens 
to prevent fish entry into dam sluices, and some inef- 
fective fish ladders. Scott and Scott (1988) suggested 
that after extensive fish ladder poaching in the early 
1950s, the Tusket River population never recovered 
to former levels of abundance. 

It is probable that acidification has also contribut- 
ed to the decline of the Atlantic Whitefish population 
in the Tusket River. While the tolerance of Atlantic 
Whitefish to acidification is not known, the impacts 
were likely comparable to impacts on Atlantic 
Salmon in this same river. Farmer et al. (1980) sug- 
gested that Atlantic Salmon reproduction in rivers 
like the Tusket could have been adversely affected 
by acidity by the early 1950s. Watt et al. (1983) indi- 
cate that records of angling success for Atlantic 
Salmon in the Tusket River suggest no trend until 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


the late 1940s and then declines in the 1950s. Watt 
(1986) suggested the decline in adult salmon returns 
during the 1950s in rivers like the Tusket was due to 
a large increase in acidic deposition during the 
1945-1955 decade sufficient to overwhelm natural 
acid neutralization and buffering capacity in these 
rivers. Nova Scotian rivers with lesser declines in 
pH, like the Annis River and Petite Riviére, did not 
show an impact of acidification on angling returns 
(Watt 1986). 

By the 1970s, it was a novelty for one Atlantic 
Whitefish to be captured in a single season by a 
gaspereau fisherman on the Tusket River (Gilhen, 
1977). When Gilhen questioned gaspereau fishermen 
in 1977 about when they had last seen an Atlantic 
Whitefish, the answers ranged from | to 15 years. A 
field survey in the fall of 1982 failed to find any 
Atlantic Whitefish in the Tusket River despite the 
operation of a Department of Fisheries and Oceans 
fish trap on the fish ladder at the Tusket River hydro- 
electric dam from 5th October to 20th November 
(Edge 1984). This fish ladder would have been the 
only means for Atlantic Whitefish to migrate into 
fresh water since the other fish ladder at the Tusket 
River holding dam was not functioning because of 
insufficient water flow. Many gaspereau fisherman 
indicated in the fall of 1982 that the Atlantic 
Whitefish had probably disappeared from the Tusket 
River. However, an Atlantic Whitefish was reported 
to have been caught by a gaspereau fisherman in 
April 1996 just below the Tusket River holding dam 
(Patrick Patten, personal communication, March 
2000). We are not aware of any reports of Atlantic 
Whitefish around the Tusket River dam since that 
time. 


(2) Annis River 

Atlantic Whitefish are reported to have been 
abundant in the Annis River which is a tributary 
flowing into the Tusket River estuary. A gaspereau 
fisherman on the Annis River indicated that, years 
ago, it was common to accidentally catch 50 to 100 
Atlantic Whitefish each year in gaspereau nets 
(Gilhen 1977). By the early 1980s, fishermen on the 
Annis River indicated that catches were declining 
and that they had been getting a combined catch of 
fewer than ten whitefish each year for roughly the 
last ten years. 

A field survey in October 1982 caught two 
Atlantic Whitefish in the lower Annis River (Edge, 
1984). One specimen was caught on 12th October in 
a gillnet set in brackish water at the Annis River 
mouth. The other Atlantic Whitefish specimen was 
caught, while moving upstream on 11th October, in a 
hoopnet located in fresh water just above the head of 
high tide on the Annis River. This hoopnet operated 
for about 210 hours from 4 to 19 October and the 
Atlantic Whitefish specimen comprised about 4% of 
all fishes caught. A small trapnet which operated for 


2001 


216 hours during the same period in brackish water 
at the Annis River mouth caught eels (Anguilla ros- 
trata) and tomcod (Microgadus tomcod) but failed to 
catch any Atlantic Whitefish. 

The Atlantic Whitefish appears to have declined 
significantly in the Annis River over the last 15 
years. A fisherman indicated that a small population 
of Atlantic Whitefish was surviving in the Annis 
River in the early 1980s, and that he had caught sev- 
eral specimens in a gaspereau net at the river mouth 
in the spring of 1983 (Limon Earle, Pleasant Lake, 
1983, personal communication). However, this fish- 
erman indicated in September 1995, that he had not 
caught any Atlantic Whitefish in the Annis River in 
the last four or five years, and believed that the 
Atlantic Whitefish had probably disappeared from 
the river. He suggested that the recent spread of 
Chain Pickerel, Esox niger, into the lower reaches of 
the Annis River probably had a significant impact on 
Atlantic Whitefish. At present, it appears that the 
Atlantic Whitefish population levels have declined in 
the Annis River since the last COSEWIC report, 
although it is possible that a remnant population still 
exists. 


(3) Petite Riviere Watershed 

Little is known about the population size and 
trends of the Atlantic Whitefish in the Petite Riviere 
watershed. A small recreational fishery for Atlantic 
Whitefish has existed around Milipsigate and Hebb 
lakes since at least the 1870s (J. March, in litt., 
1983). Although anglers have not considered the 
Atlantic Whitefish to be plentiful in these lakes, 
these fish were known to concentrate in schools 
below the Milipsigate Lake dam outlet in the spring. 
On these occasions, anglers could fill their creel in a 
short time. One angler and his son caught 24 Atlantic 
Whitefish at this location on 9 May 1925 (S. March, 
in litt., 1925). While historical knowledge of popula- 
tion trends in the Petite Riviere lakes is scant, Mr. J. 
March indicated there were persistent rumours in the 
early 1920s that persons dynamited schools of 
Atlantic Whitefish and gathered them up in nets. 
This type of activity could have had an influence on 
population sizes. 

In the fall of 1982, a field survey found Atlantic 
Whitefish surviving in Hebb, Milipsigate, and 
Minamkeak lakes within the Petite Riviere water- 
shed (Edge 1984). This survey suggested popula- 
tions of Atlantic Whitefish were small based upon 
average catches of 0.75, 1.25, and 2.0 whitefish per 
75m gillnet set for 18 hours in Milipsigate, Mina- 
mkeak, and Hebb lakes respectively. Subsequent 
field studies focused on Hebb Lake in the summer of 
1983, and minnow traps, seines, trapnets and gillnets 
were used to study fish populations (Edge 1987). 
Fishing methods other than gillnets indicated Banded 
Killifish (Fundulus diaphanus), Golden Shiners 
(Notemigonus crysoleucas), Ninespine Sticklebacks 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


647 


(Pungitius pungitius) and young Yellow Perch 
(Perca flavescens) were common in shallow waters. 
While these methods also caught eels (Anguilla ros- 
trata), Brown Bullheads (Ameiurus nebulosus), 
White Perch (Morone americana) and White 
Suckers (Catostomus commersoni), they did not 
catch any Atlantic Whitefish. Atlantic Whitefish 
were caught only in gillnets. 

The results of gillnetting efforts in the Petite 
Riviére lakes in 1982 and 1983 are shown in Table 
3. This table also shows Lake Whitefish gillnet cap- 
ture data from other lakes in Nova Scotia for com- 
parison. The gillnets used in 1982-1983 were 75 m 
in length with 15 m panels of stretched mesh sizes of 
25, 37.5, 50, 75 and 100 mm that were set overnight 
in lakes. The gillnet data indicate the Petite Riviere 
lakes appear to have a predominance of White Perch, 
White Sucker and Yellow Perch, while Atlantic 
Whitefish were much less abundant. The results indi- 
cate that the 37 specimens of Atlantic Whitefish 
caught in 1982 and 1983 accounted for only 4.9% of 
all fish specimens caught in Minamkeak, Milipsigate 
and Hebb lakes. When considering combined fishing 
effort from bottom and surface set gillnets, Atlantic 
Whitefish appeared more abundant in Hebb Lake 
(0.14 specimens caught per hour of gillnet) than 
Minamkeak Lake (0.07 specimens), or Milipsigate 
Lake (0.06 specimens). The data in Table 3 also sug- 
gest Atlantic Whitefish populations in the Petite 
Riviére lakes are likely smaller than Lake Whitefish 
populations in larger bodies of water like the Mira 
River, Cape Breton County and Lake George, Yar- 
mouth County Lake Whitefish were caught at a rate 
of 0.53 and 0.30 whitefish per hour of gillnet set in 
the Mira River and Lake George, respectively. 

Available information would suggest the popula- 
tions of Atlantic Whitefish in Minamkeak, Milipsi- 
gate, and Hebb lakes are small. A small school of 
Atlantic Whitefish still concentrates at the Milipsi- 
gate Lake dam outlet every spring. A visit to this 
location did not find any Atlantic Whitefish between 
9-15 May 1983, when water temperature was 14° C. 
However, a small gillnet set overnight on 17 Feb- 
ruary 1985, caught five whitefish specimens at the 
site. Members of the Atlantic Whitefish Conser- 
vation and Recovery Team observed whitefish 
around the Milipsigate Lake outlet in the spring of 
1999 (about 25-30 fish). In the spring of 2000 there 
were an estimated 200-250 Atlantic Whitefish at this 
dam outlet. An Atlantic Whitefish was also reported 
to have been caught by an angler in Fancy Lake 
around 1997-1998. 

Little is known about the population size or trends 
of Atlantic Whitefish in the lower reaches of the 
Petite Riviére. Captain D. R. Bell, Petite Riviere, 
Lunenburg County has provided helpful insights on 
the history of Atlantic Whitefish that suggest popula- 
tion levels may have declined over the last few 


THE CANADIAN FIELD-NATURALIST Vol. 115 


648 


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2001 


decades. Captain Bell indicated that Atlantic 
Whitefish have occurred in the Petite Riviere estuary 
since at least the 1930s and that they were caught in 
drift nets in the harbour for use as lobster bait. Thirty 
to fifty Atlantic Whitefish could be caught in ten 
nets over one night’s fishing. Prior to the late 1940s, 
it was relatively common to catch them in the 
gaspereau fishery from mid-April to June. Only one 
gaspereau fisherman is now left on the Petite Riviere 
and he catches and releases only about two or three 
Atlantic Whitefish on average over an entire season. 
He caught and released three Atlantic Whitefish in 
the spring of 1999. A Department of Fisheries and 
Oceans trap net placed in the Petite Riviere estuary 
in the fall of 1999 did not capture any Atlantic 
Whitefish. While the numbers of Atlantic Whitefish 
appear to have declined since the 1930s, Captain 
Bell indicated it is still not uncommon at times to see 
Atlantic Whitefish jumping, much like a salmon on a 
rising tide, in the Petite Riviére estuary. It is not cer- 
tain whether this is a resident population or these are 
strays from the Petite Riviére lakes. 


Limiting Factors and Threats 
(1) The Tusket River Watershed 

If Atlantic Whitefish are still surviving in the 
Tusket River watershed, they are likely to be limited 
by a number of factors. Atlantic Whitefish migra- 
tions in the Tusket River may be limited by the 
Tusket River dams and ineffective fish ladders. 
Inadequate fish ladders for fish to pass around the 
Tusket River hydro-electric dam have been a prob- 
lem for species like Atlantic Salmon since at least 
the 1960s (Smith 1962). While the fish ladder at the 
hydro-electric dam on the Tusket River that was con- 
structed in 1979 has been used by salmon and 
gaspereau, it is unknown whether it is effective for 
Atlantic Whitefish. During the migration period of 
the Atlantic Whitefish in October 1982, there was 
little attraction water coming down this fish ladder 
which may have limited Atlantic Whitefish from 
finding the ladder. The only other fish ladder on the 
Tusket River at that time was at the holding dam and 
waterflow was so small that no fish passage was pos- 
sible. While an effective new fish ladder was con- 
structed at the holding dam in 1997, inadequate 
water flows during Atlantic Whitefish migratory 
periods likely do not permit any passage 

Water flows in the Tusket River can fluctuate 
widely depending upon hydro-electric power 
demands. At times the river can become almost dry 
below the holding dam potentially adversely impact- 
ing fish spawning and nursery habitats. It is un- 
known how these fluctuations in water flows, includ- 
ing those down fish ladders have affected Atlantic 
Whitefish movements or habitats. 

Atlantic Whitefish in the Annis River are increas- 
ingly threatened by the spread of at least two intro- 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


649 


duced fish species which may pose competitive or 
predation risks. One specimen of Brown Trout, 
Salmo trutta, was caught migrating upstream in a 
hoopnet on 9 October 1982, three days before the 
capture of an Atlantic Whitefish specimen in this 
same net. The Brown Trout was a female (580 mm 
TL) in spawning condition. Gilhen (1974) indicated 
Brown Trout were introduced into the Annis River 
around the 1920s or 1930s. 

Five Chain Pickerel, Esox niger, were caught in 
October 1982, in Salmon Lake, Yarmouth County, 
the first major lake upstream from the Annis River 
mouth. Chain Pickerel were also caught in this lake 
by a provincial fish survey in 1977. Since the late 
1980s, however, the Chain Pickerel has spread 
downstream to the lower reaches of the Annis River. 
Fishermen have recently been catching Chain 
Pickerel at least as large as 55 cm in the lower reach- 
es of the Annis River where they never occurred pre- 
viously (L. Earl, 1995, personal communication). 
These Chain Pickerel have apparently spread down- 
stream from Salmon Lake, and they now occur 
throughout the lower reaches of the Annis River 
where Atlantic Whitefish enter fresh water. Chain 
Pickerel were introduced into this area many years 
ago and could pose a threat to salmonids (Gilhen 
1974: Alexander et al. 1986; Coffie 1998). 

Another limiting factor for Atlantic Whitefish in 
the Tusket River watershed is acidification. Many 
rivers in southwestern Nova Scotia have become 
increasingly acidified to the point where Atlantic 
Salmon can no longer reproduce and their native 
salmon populations are considered extinct (Watt et 
al. 1983; Watt 1986; Watt 1989). Watt (1986) indi- 
cated that a large area of Nova Scotia has already 
been rendered barren of Atlantic Salmon, and more 
is expected to be lost over the next 20 years. Watt 
(1986) categorized the Tusket River as a river 
where only remnant populations of Atlantic Salmon 
survive in one or two higher pH tributaries and 
where angling catch has declined to about 10% of 
levels prevalent during the 1936-1950 period. 
While the Annis River appears better buffered from 
threats of acidification, the pH in this river can drop 
quite low at times. Watt (1986) categorized the 
Annis River as a river with salmon stocks depleted 
by acidification of some of the smaller tributaries, 
but in the main stem and most tributaries, salmon 
production rates are normal. The tolerance of 
Atlantic Whitefish to acidification is unknown and 
it remains uncertain whether the species could con- 
tinue to successfully reproduce in acidified waters 
like the Tusket River. 


(2) Petite Riviere Watershed 

The Atlantic Whitefish populations in the Petite 
Riviere watershed also face significant limiting fac- 
tors. The Petite Riviére occurs in an area of Nova 
Scotia which is better buffered from the impacts of 


650 


acidification. However, the time of annual water pH 
minimum in the Petite Riviere lakes may coincide 
with the presence of acid-sensitive life history stages 
of the Atlantic Whitefish. In southwestern Nova 
Scotia, annual pH minimums typically occur from 
midwinter to early spring, presumably when Atlantic 
Whitefish eggs and newly hatched larvae would be 
present. The pH of Hebb Lake in February 1985 was 
4.5 while the pH of the Annis and Tusket rivers in 
March 1985 was 5.15 and 4.66 respectively. While 
the failure to find young Atlantic Whitefish during 
surveys of the Petite Riviere Lakes in the 1980s was 
probably the result of inappropriate fishing methods 
and habitat sampling, it could also have indicated 
recruitment failures. 

While Atlantic Whitefish could be protected 
from acidification by liming, such efforts may not 
be feasible over long periods. White et al. (1984) 
conducted an experimental neutralization of an 
acidified lake and indicated that in Nova Scotia, 
where retention times of most lakes are very short, 
neutralization of acid waters will have to be carried 
out at frequent intervals or continuously. Similarly, 
other experiments in Nova Scotia indicate that the 
pH of salmon streams can be adjusted to satisfacto- 
ry levels (pH above 5.0) by liming, but that fresh 
limestone must be added annually and, in some 
cases, twice annually making it very expensive 
(Watt 1986). 

The Atlantic Whitefish may also be affected by 
water level fluctuations in the Petite Riviere lakes 
and introductions of non-indigenous fish predators. 
While Brook Trout (Salvelinus fontinalis) are native 
to Nova Scotia, they have also been stocked into 
many watersheds in Nova Scotia, including the 
Petite Riviére. While the authors are not aware of 
such hatchery introductions in Minamkeak, 
Milipsigate or Hebb Lakes, other lakes such as 
Fancy, Wallace and Andrew have been regularly 
stocked with Brook Trout. Spread of hatchery Brook 
Trout could represent a threat to populations of 
Atlantic Whitefish and their regular introduction to 
Fancy Lake could contribute to the apparently only 
sporadic presence of Atlantic Whitefish in this lake. 
In addition, an unauthorized introduction of 
Smallmouth Bass (Micropterus dolomieu) took place 
in Wallace Lake within the Petite Riviere watershed 
around 1994 (A. Hebda, personal communication, 
1999). These Smallmouth Bass have survived (it is 
reported there is now a regular bass fishing tourna- 
ment), and appear to be spreading within the Petite 
Riviere watershed posing a potentially serious threat 
to Atlantic Whitefish. 

It is likely there could be some continued threat 
from angling in the Petite Riviere watershed. While 
there appears to be little angling on Milipsigate and 
Hebb Lakes, there is more lakeshore development on 
Minamkeak Lake and anglers fish from boats on this 


THE CANADIAN FIELD-NATURALIST 


Vol £15 


lake. For example, there is a white perch fishing sea- 
son in Minamkeak Lake. It will be important to 
ascertain if inadvertent captures of Atlantic 
Whitefish while angling for other species poses a 
serious threat. 


Special Significance of the Species 

The Atlantic Whitefish is a uniquely Canadian 
species found nowhere else in the world. It was the 
first fish species to be designated as endangered by 
COSEWIC. It has also been featured on a Canadian 
postage stamp. 

The Atlantic Whitefish may have some signifi- 
cance as a food and sport fish. Atlantic Whitefish 
from the Petite Riviere lakes have been described as 
an excellent table fish with flesh that is about the 
same colour as that of a herring when cooked (Piers 
1927). Scott and Scott (1988) also described the 
Atlantic Whitefish as a good food fish of fine 
flavour. Anglers have described them as gamey 
fighters that, when hooked, almost always leap from 
the water until exhausted. 

The Atlantic Whitefish is genetically very distinct 
from other whitefish species and appears to be some- 
what unique in ecological aspects such as its toler- 
ance of full seawater and warm waiters. The Atlantic 
Whitefish likely offers a unique source of genetic 
diversity among salmonid species. Further study of 
this species will be important for understanding the 
biology, behavior, ecology, evolution and zoogeog- 
raphy of the commercially important coregonids. 
The extinction of the Atlantic Whitefish would be a 
significant loss of biological diversity. 


Evaluation 
Existing Legal Protection and Other Status 

The Nova Scotia Fishery Regulations under sec- 
tion 34 of the Fisheries Act were amended 17 
February 1970, to prohibit the taking of Atlantic 
Whitefish from all waters of the province by any 
method at any time of the year. A Variance Order 
under these Regulations was issued in 2000, that 
closes the area below Milipsigate Lake to any fishing 
from 1 April to 30 May for ten years. 

The Atlantic Whitefish was classified as an endan- 
gered species by the Committee on the Status of 
Endangered Wildlife in Canada in 1983. It is identi- 
fied on the 2000 International Union for Con- 
servation of Nature and Natural Resources (IUCN) 
Red List of Threatened Species with a designation of 
VU D2 (Hilton-Taylor 2000). This designation 
implies the species is not endangered but is facing a 
high risk of extinction in the wild in the medium term 
future. The designation indicates the population is 
very small and is characterized by an acute restriction 
in its area of occupancy. The 2000 IUCN Red List 
identifies the species by the previously used common 
name of Acadian rather than Atlantic Whitefish. The 


2001 


List incorrectly describes the distribution of the 
Atlantic Whitefish as being the Great Lakes region, 
including within the United States. 


Assessment of Status 

Since the last assessment by COSEWIC in 1983 
and the IUCN in 1996, the numbers of Atlantic 
Whitefish have declined significantly in the Annis 
River. The species faces new threats from the spread 
of introduced Chain Pickerel into the lower reaches 
of the Annis River and from the spread of introduced 
Smallmouth Bass in the Petite Riviére watershed. In 
addition, the threat of acidification in both Tusket 
and Petite Riviere watersheds continues. The Atlan- 
tic Whitefish should be considered as an endangered 
species. 


Acknowledgments 

This COSEWIC report is dedicated to the memo- 
ry of the late Don E. McAllister and his tremendous 
contributions towards the conservation of biological 
diversity. 

Many thanks to Robert Campbell and COSEWIC 
for assistance in preparing this report. Thanks also to 
Don E. McAllister, Ocean Voice International and 
Canadian Museum of Nature; Brian Coad, Canadian 
Museum of Nature; and Bev Scott, Kingston, 
Ontario, for support and encouragement over a num- 
ber of years. Support from John Loch, Department of 
Fisheries and Oceans, Halifax, Nova Scotia; and the 
members of the Atlantic Whitefish Conservation and 
Recovery Team was greatly appreciated. Captain 
D. R. Bell, Petite Riviére, Lunenburg County; Limon 
Earl, Pleasant Lake, Yarmouth County; Trevor Goff, 
Milton, Nova Scotia; Andrew Hebda, Nova Scotia 
Museum of Natural History; and Patrick Patten, 
Tusket River Environmental Protection Association 
R.R.#4, Yarmouth County, provided very valuable 
insights and historical accounts of Atlantic Whitefish 
behavior and population trends. Comments on the 
draft report from Dick Cutting, Halifax, Nova Scotia, 
and Don E. McAllister, Perth, Ontario, and other 
anonymous reviewers were much appreciated. We 
thank Fred Scott, Nova Scotia Museum of Natural 
History, for assistance in preparing the distribution 
map. 


Documents Cited 

Gilhen, J. 1977. A report on the status of the Atlantic 
Whitefish, Coregonus canadensis, in the Tusket River 
watershed, Yarmouth County, including recommenda- 
tions to ensure its future survival. Report submitted to 
the Ichthyology Section, National Museum of Natural 
Sciences, Ottawa, Ontario. October 30, 1977. 18 pages. 

Hilton-Taylor, C. Compiler. 2000. 2000 IUCN Red List 
of Threatened Species. IUCN, Gland Switzerland and 
Cambridge, UK. xviii + 61 pages. [Downloaded on 
November 28, 2001.] 

Smith, K. E. H. 1960. Tusket River Power Dam, Yar- 
mouth Co., N.S. — Mortality Tests — Young Salmon 


EDGE AND GILHEN: UPDATED STATUS OF THE ATLANTIC WHITEFISH 


651] 


and Gaspereau. Annual Report, Fish Culture Branch, 
Halifax. (Unpublished) 

Smith, K. E. H. 1961. Mortality Tests, Yearling Gas- 
pereau at Tusket River Power Dam. Annual Report, Fish 
Culture Branch, Halifax. (Unpublished) 

Smith, K. E. H. 1962. Tusket River Survey, Yarmouth 
Co., N.S. Department of Fisheries, Halifax, N.S. (Un- 
published) 


Literature Cited 

Alexander, D.R., J. J. Kerekes, and B. C. Sabean. 1986. 
Description of selected lake characteristics and occur- 
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ceedings of the Nova Scotia Institute of Science 36: 
63-106. 

Bernatchez, L., T. A. Edge, J. J. Dodson, and S. U. Qadri. 
1991. Mitochondrial DNA and isozyme electrophoretic 
analyses of the endangered Acadian Whitefish, Core- 
gonus huntsmani Scott, 1987. Canadian Journal of Zoo- 
logy 69: 311-316. 

Campbell, R. R. Editor. 1997. Rare and endangered fish- 
es and marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: XI. 
Canadian Field-Naturalist 111: 249-257. 

Coad, B. W. 1995. Encyclopedia of Canadian fishes. 
Canadian Museum of Nature and Canadian Sportfishing 
Productions Ltd., Ottawa. 928 pages. 

Coffie, P. A. 1998. Status of the Chain Pickerel, Esox niger, 
in Canada. Canadian Field-Naturalist 112: 133-140. 

Dunfield, R. W. 1985. The Atlantic Salmon in the history 
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Fisheries and Aquatic Sciences Number 80. 181 pages. 

Edge, T.A. 1984. Preliminary status of the Acadian 
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Edge, T. A. 1987. The systematics, distribution, ecology 
and zoogeography of the endangered Acadian Whitefish, 
Coregonus canadensis Scott, 1967, in Nova Scotia, Can 
ada. M.Sc. thesis, University of Ottawa, Ottawa, Ontario. 
175 pages. 

Edge, T. A., D. E. McAllister, and S. U. Qadri. 1991. 
Meristic and morphometric variation between the endan- 
gered Acadian Whitefish, Coregonus huntsmani, and the 
Lake Whitefish, Coregonus clupeaformis, in the Canadian 
Maritime Provinces and the State of Maine, USA. Can- 
adian Journal of Fisheries and Aquatic Sciences 48: 
2140-2151. 

Erskine, J.S. 1971. The archaeology of some Nova 
Scotian indian campsites. Proceedings of the Nova 
Scotia Institute of Science 27: 1-10. 

Farmer, G. J., T. R. Goff, D. Ashfield, and H. S. Samant. 
1980. Some effects of the acidification of Atlantic 
Salmon rivers in Nova Scotia. Canadian Technical 
Report, Fisheries and Aquatic Sciences 972. 13 pages. 

Gilhen, J. 1974. The fishes of Nova Scotia’s lakes and 
streams. Nova Scotia Museum, Halifax, N.S. 49 pages. 

Gilhen, J. 1999. Fishes of Nova Scotia: Species Recorded 
in the Accession Books of Harry Piers From 1899 to 
1939, Nova Scotia Museum. Curatorial Report Number 
89: 153 pages, 107 plates. 

Keddy, P. A. 1985. Lakeshores in the Tusket River Val- 
ley, Nova Scotia: Distribution and status of some rare 
species, including Coreopsis rosea Nutt. and Sabatia 
kennedyana Fern. Rhodora 87: 309-320. 

Lacroix, G.L., and D.R. Townsend. 1987. Responses of 


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juvenile Atlantic Salmon (Salmo salar) to episodic in- 
creases in acidity of Nova Scotia rivers. Canadian Journal 
of Fisheries and Aquatic Sciences 44: 1475-1484. 

McAllister, D. E. 1990. A list of the fishes of Canada. 
Syllogeus Number 64: 310 pages. 

McAllister, D. E., B. J. Parker, and P.M. McKee. 1985. 
Rare, endangered and extinct fishes in Canada. Syl- 
logeus 54, National Museums of Canada. 192 pages. 

Piers, H. 1927. Coregonus labradoricus, the Sault White- 
fish, an interesting addition to the freshwater fish fauna 
of Nova Scotia. Proceedings of the Nova Scotia Institute 
of Science 16: 92-95. 

Robins, C.R., R. M. Bailey, C.E. Bond, J. R. Brooker, 
EK. A. Lachner, R.N. Lea, and W.B. Scott. 1991. 
Common and scientific names of fishes from the United 
States and Canada. Special Publication 20. American 
Fisheries Society. Bethesda, Maryland. 

Scott, W.B. 1967. Freshwater fishes of Eastern Canada, 
Second Edition, University of Toronto Press, Toronto, 
Ontario. 

Scott, W. B. 1987. A new name for the Atlantic Whitefish: 
Coregonus huntsmani to replace Coregonus canadensis. 
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Canada. Canadian Bulletin of Fisheries and Aquatic Sci- 
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THE CANADIAN FIELD-NATURALIST 


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Semple, J. R. 1973. The Lake Whitefish, Coregonus clu- 
peaformis, (Mitchill) of Scotch Pond, Petpeswick River, 
Nova Scotia. Transactions of the Canadian Society of 
Environmental Biologists. Pages 1—21. 

Smith, MW. 1952. The Lake Whitefish in Kerr Lake, New 
Brunswick. Journal of the Fisheries Research Board of 
Canada 8(5): 340-346. 

Watt, W. D. 1986. The case for liming some Nova Scotia 
salmon rivers. Water Air Soil Pollution 31: 775-789. 
Watt, W.B. 1989. The impact of habitat damage on 
Atlantic Salmon (Salmo salar) catches. Proceedings of 
the National Workshop on Effects of Habitat Alteration 
on Salmonid Stocks. Edited by C. D. Levings, L. B. 
Holtby and M. A. Henderson. Canada Special Publica- 

tion Fisheries and Aquatic Sciences 150: 154-163. 

Watt, W.D., C. D. Scott, and W. J. White. 1983. Evi- 
dence of the acidification of some Nova Scotian rivers 
and its impact on Atlantic Salmon (Salmo salar). 
Canadian Journal of Fisheries and Aquatic Sciences 40: 
462-473. 

White, W. J., W. D. Watt, and C. D. Scott. 1984. An 
experiment on the feasibility of rehabilitating acidified 
Atlantic salmon by the addition of lime. Fisheries 9: 
25-30. 


Accepted 19 February 2002 


The Status of the Grey Seal, Halichoerus grypus, in the Northwest 


Atlantic* 


VERONIQUE LESAGE and MIKE O. HAMMILL 


Department of Fisheries and Oceans, Maurice Lamontagne Institute, P.O. Box 1000, 850 Route de la Mer, Mont-Joli, 


Québec GSH 324 


Lesage, Véronique, and Mike O. Hammill. 2001. The status of the Grey Seal, Halichoerus grypus, in the Northwest 


Atlantic. Canadian Field-Naturalist 115(4): 653-662. 


Northwest Atlantic Grey Seals form a single population. For management purposes a Sable Island component increasing at 
a rate of 12.6% per year, and a non-Sable Island component increasing at a rate of 3.4 + 1.5% per year, have been identi- 
fied. Modelling population size using information on changes in pup production and age specific reproductive rates, there 
were an estimated 191 800 animals in 1997. Grey Seals are found in northwestern Atlantic waters throughout the year. 
Largely coastal in habit, they feed on a variety of pelagic and demersal fish including several commercially important 
species. With the exception of hunting and possibly predation from sharks no significant sources of natural mortality have 
been identified. Grey Seals could have a negative impact on commercial fisheries through the consumption of fish, trans- 
mission of the nematode parasite Pseudoterranova decipiens, and damage to fishing gear. Global climate changes would be 
expected to affect Grey Seals whelping on the pack-ice in the southern Gulf of St Lawrence. Currently there is no commer- 


cial hunt for Grey Seals. 


Les Phoques Gris du nord-ouest Atlantique constituent une seule et unique population. Pour des fins de gestion, une com- 
posante dite ‘de I’Tle de Sable’, s’accroissant a un taux de 12.6% par année, et une composante dite “non de I’Ile de Sable’, 
augmentant a un taux de 3.4 + 1.5% par année ont été identifiées. La modélisation de la taille de la population a partir de 
l'information concernant les changements de la production de nouveau-nés et du taux de reproduction 4 l’age estime a 


191 800 le nombre d’animaux en 1997. Les Phoques gris oc 


cupent les eaux du nord-ouest Atlantique durant toute I’ année. 


Principalement cOtiers de par leurs habitudes, ils se nourrissent d’une variété de poissons pélagiques et démersaux dont 
plusieurs espéces importantes commercialement. Mis a part la chasse et possiblement la prédation par les requins, aucune 
source de mortalité naturelle significative n’a été identifiée. Les phoques gris pourraient avoir un impact négatif sur les 


pécheries commerciales par leur consommation de poisson 


decipiens, et les dommages infligés aux agres de péche. Les 


les Phoques Gris lors de la reproduction sur les glaces flott 


aucune chasse commerciale dirigée vers les Phoques Gris. 


s, la transmission du nématode parasitaire Pseudoterranova 
hangements globaux de climat sont présumés pouvoir affecter 
antes du sud du Golfe Saint-Laurent. II n’existe présentement 


Key Words: Pinnipedia, Phocid, Halichoerus grypus, Grey Seal, Phoque Gris, Northwest Atlantic. 


The Grey Seal (family Phocidae) was first 
described by Fabricius (1791). Its name Halichoerus 
comes from the Greek meaning “sea pig”, and gry- 
pus from the Latin meaning hook-nosed (Bonner 
1981). In Canada, they are sometimes referred to as 
horse-head seals owing to the elongated snout of the 
males (Figure 1). Males tend to be darker in colour 
than females, in some cases almost black. They may 
reach a length of 231 cm (McLaren 1993), and have 
a mean weight of 298 kg (SD = 29.5) during the 
breeding season (Tinker et al. 1995). Females are 
smaller, with a length of 201 cm (McLaren 1993) 
and a weight of 227 kg (SD = 26) (Baker et al. 
1995). The pups are born with a white lanugo, which 
they begin to shed approximately nine days after 
birth, and is completely replaced by a black spotted, 
silver coat by the 25th day (Davies 1949; Bowen and 
Stobo unpublished in Renouf 1990). 


eS 
*Reviewed and approved by COSEWIC, April 1999, status 
assigned — Not At Risk. 


Grey Seals are considered to be a coastal species. 
They may forage far from shore, but do not appear to 
leave the continental shelf regions (Gosselin and 
Hammill, unpublished data). They haul out on 
exposed reefs or on beaches of undisturbed islands. 
These concentrations of animals are typically quite 
noisy (Lesage and Hammill, personal observation), 
and are associated with vocalisations resembling 
growls and roars (Schneider 1974). These noises 
sometimes resemble the calling of a wolf, and this 
may be the source of the general French term ‘loups 
marins’ (which means sea wolves). 


Distribution 

Grey Seals are found throughout the temperate 
North Atlantic. Three distinct populations are recog- 
nised: the Baltic Sea population, a North-Eastern 
Atlantic population that extends from Iceland to 
Norway, but with the main concentration around the 
United Kingdom, and a North-Western Atlantic popu- 
lation (Davies 1957). An examination of mitochondri- 
al DNA variation in samples from Canada, Norway 


653 


654 


and the Baltic Sea supports the hypothesis that seals 
from these areas do represent three distinct popula- 
tions. Eastern and western Atlantic Grey Seals 
diverged first, possibly 1.0—1.2 million years ago, 
while the Baltic Sea animals diverged much later 
(Boskovic et al. 1996). 

In the Northwest Atlantic, Grey Seals have been 
found as far north as Cape Chidley, at the northern 
tip of Labrador, and as far south as Virginia in the 
USA (Katona et al. 1993). In Canada, the largest 
concentrations of animals outside of the breeding 
season are found around Sable Island, and in the 
northern Gulf of St Lawrence around Anticosti 
Island (Clay and Nielsen 1985; Stobo et al. 1990; 
Lavigueur and Hammill 1993) (Figure 2). With the 
increase in the Grey Seal population there has been 
an increase in sightings along the New England, 
United States coast (Katona et al 1993), along the 
Lower North Shore of Quebec (G. Beck, Toronto 
Ontario, personal communication) and in the Strait 
of Belle Isle (G. Stenson, DFO St John’s Newfound- 
land, personal communication). 

The Northwest Atlantic Grey Seals form a single 
population (Boskovic et al. 1996), but for manage- 
ment purposes, they are divided into two major 
groups based on the locations of the whelping patch- 
es. The largest group whelps on Sable Island, a 40 
km long sand spit located approximately 250 km to 
the east of Nova Scotia (Figure 2). The second 
group, known as non-Sable Island Grey Seals, is 
largely made up of animals that whelp on drifting 
pack ice in Northumberland Strait and along the west 
coast of Cape Breton Island in the southern Gulf of 
St Lawrence. Smaller groups of non-Sable Island 
Grey Seals also whelp on small islands or reefs 
located in the southern Gulf of St Lawrence, and 
along the eastern shore of Nova Scotia (Mansfield 
and Beck 1977). In recent years, a new colony num- 
bering 600+ pups has formed on Hay Island off the 
Cape Breton coast (Figure 2). New pupping colonies 
have also formed in Maine and in Massachusetts 
(Hannah 1998). Although Sable and non-Sable 
Island animals show strong philopatry to their 
whelping sites, some exchange of adults between 
breeding colonies has been documented and may be 
in the range of 5% (Zwanenburg and Bowen 1990). 
Considerable overlap occurs between the two groups 
outside of the breeding season (Stobo et al. 1990; 
Lavigueur and Hammill 1993). 


Movements 

Information on movements of Grey Seals is avail- 
able from anecdotal sightings, recovery of tags from 
shot animals or strandings and more recently from 
satellite telemetry. 

Sable Island Grey Seals have a post-breeding 
pelagic phase (February—April) when they disperse 
from the island. This is followed by a spring moult 
(May-June), when animals return to Sable Island. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


After they have completed their moult, there is a sec- 
ond, summer dispersal away from the island (July- 
September). At this time, animals may move towards 
the Nova Scotia, Maine, and Newfoundland coasts, 
and into the Gulf of St Lawrence. They return to 
Sable Island during the fall and early winter 
(October— December) (Stobo et al. 1990). 

This general movement pattern is also true for 
non-Sable Island Grey Seals. Tag returns and satel- 
lite telemetry indicate that after breeding, the majori- 
ty of adult Grey Seals move out of the Gulf of St 
Lawrence onto the Scotian shelf (Hammill et al. 
1993; Lavigueur and Hammill 1993; Goulet et ai. 
1995), where they remain offshore until the spring 
moult. The pups tend to remain with the pack ice as 
it drifts around the west side of the Cape Breton 
coast into the Atlantic. However, in some years, ice 
drift is slow, or there is little ice available which 
breaks up early in the season. When this occurs, 
many animals move ashore onto the beaches of 
Prince Edward Island, Cape Breton Island and main- 
land Nova Scotia, where mortality may be high. 
During May-June, both adults and juveniles move 
ashore to moult. Although some animals moult 
throughout the Gulf of St Lawrence, most appear to 
move into the northern Gulf of St Lawrence, around 
Anticosti Island, and along the Lower North Shore 
(Clay and Nielsen 1985). After the moult, animals 
disperse, with many animals moving into the St 
Lawrence Estuary (Lavigueur and Hammill 1993). 
In late fall, Grey Seais return to the southern Gulf of 
St Lawrence or to Sable Island for the breeding sea- 
son. This movement, which often occurs in 
November-December, usually takes only a few days 
(Goulet et al. 1995; Gosselin and Hammill 1998). 


Protection 
International Protection Measures 

In the United States of America, all marine mam- 
mals are protected from hunting, except for subsis- 
tence hunting by aboriginal people, under the Marine 
Mammal Protection Act of 1972. Grey seals are 
completely protected from hunting in France, 
Denmark, Finland, Sweden and the Netherlands. In 
the United Kingdom and Ireland they are protected 
from hunting, but exemptions exist for killing seals 
to protect fisheries or for scientific purposes. In 
Iceland, Grey Seals are not protected and may still 
be hunted. In Norway, Grey Seals are protected in 
areas south of 62°N. Hunting is permitted from | 
November to 30 April between 62—67°N, and from 1 
December to 30 April north of 67°N. In the Faeroe 
Islands, Grey Seals may only be killed using guns. 
Licences for guns are mainly restricted to aquacul- 
ture operations. 


National Protection Measures 

In Canada marine mammals fall under federal 
jurisdiction. Grey Seals were managed under the 
Seal Protection Regulations of the Fisheries Act 


i et Hie, 


“i 


vy 


LESAGE AND HAMMILL: STATUS OF GREY SEAL IN NORTHEAST ATLANTIC 


655 


era 


“ee 


7 td a i 


FIGURE 1. The Grey Seal (Halichoerus grypus) [photo by Mike Hammill]. 


since 1966. In February 1993, the Seal Protection 
Regulations were superseded by the Marine 
Mammal Regulations of the Fisheries Act. These 
regulations govern scientific research activities, 
marine mammal ecotourism and observation, and 
hunting activities. Licences are currently issued to 
non-natives to hunt Grey Seals in Canadian waters. 
There is also a provision for personal sealing 
licences, which allows the holder to take six seals 


Newfoundland 


New Brunswick 


© a Nova Scotia Y White Is. 
eb >Bowen's Atlantic Ocean 


Ledge 


Sable Is. 
65° W 60° W 


FIGURE 2. Main whelping sites of land-breeding (G) and 
ice-breeding (dotted area) Grey Seals. 


annually. The harvest, either for commercial or per- 
sonal purposes, is restricted by an obligation to make 
full use of the entire carcass of the killed seal. 
Hunting is not permitted during the breeding season, 
i.e. in January and February. Furthermore, it is ille- 
gal to hunt Grey Seals during the summer (May/1 
June to 30 September) to the west of 67°23’ (Pointe- 
des-Monts) in the St Lawrence Estuary and 
Saguenay River, in certain regions adjacent to the 
coast of New Brunswick, the Magdalen Islands, and 
Murray Harbour (Prince Edward Island), and 
throughout the year near the Gaspé coast. The 
Minister of Fisheries and Oceans can modify closure 
dates by the release of ordinances and variances. 


Population Sizes and Trends 

Grey seals were at one time abundant and widely 
distributed along the Canadian east coast and in the 
Gulf of St Lawrence where they were first hunted by 
Amerindians. Hunting by Europeans, particularly 
after the disappearance of the Walrus (Odobenus ros- 
marus) in the Gulf and on Sable Island, resulted in 
the depletion of the Grey Seal population by the mid- 
1800s (Lavigueur and Hammill 1993). By the early 
1900s, Grey Seals were still considered to be widely 
distributed, but there was no particular hunt for them, 
owing to their small numbers (Comeau 1945). During 
the 1950s, the Grey Seal was considered to be 
uncommon or rare in eastern Canada. Nevertheless, 
Grey Seals continued to be hunted. In 1927, the 
Canadian government initiated a bounty program 
directed towards another seal species, the Harbour 


656 


Seal (Phoca vitulina), under which a reward was paid 
to fishermen upon receipt of a seal snout. In 1949, the 
system was changed to require presentation of the 
lower jaw to receive the bounty. Since it is possible 
to identify the species by the lower jaw, it became 
apparent that some Grey Seals had been collected in 
small numbers (Mansfield and Beck 1977). Between 
1967 and 1984, the Canadian Department of 
Fisheries and Oceans conducted an annual cull for 
Grey Seals at breeding colonies in the Gulf of St 
Lawrence, and along the Nova Scotia eastern shore, 
removing 114 to 2375 animals annually (Zwanenburg 
and Bowen 1990). From 1978 until 1990, a bounty 
was paid to licensed fishermen who submitted lower 
jaws from Grey Seals, and information on date and 
location of capture. A total of 4379 individuals were 
taken during this program. Captures were initially 
quite high, but with the exception of a large number 
of jaws received in 1989 (753), they declined steadily 
until 1990, when only 79 jaws were received 
(Lavigueur and Hammill 1993). 

In spite of continued hunting, the Northwest 
Atlantic Grey Seal population has increased. On 
Sable Island, where hunting pressure has been much 
less intense than elsewhere, Grey Seal pup produc- 
tion was determined by complete enumeration 
between 1977 and 1990. Counts on Sable Island 
indicate that pup production has increased at a rate 
of 12.6% per year from 2200 pups in 1977 to 9700 
in 1989 (Stobo and Zwanenburg 1990). Zwanenburg 
and Bowen (1990) modelled the dynamics of the 
Sable Island component of the population to esti- 
mate total population size, and to monitor changes 
in this component of the population since the 1960s. 
They modified estimates of first year survival and 
age-specific pregnancy rates (Mansfield and Beck 
1977) and assumed that during the 1960s, the Sable 
Island component represented a closed stable popu- 
lation. Using a first year survival of 0.787, and an 
annual adult survival rate of 0.96 (Zwanenburg and 
Bowen 1990) to generate the observed rate of 
increase in pup production, the Sable Island Grey 
Seal component of the population would have 
increased from 2200 animals (rounded to the nearest 
hundred) in 1962 to about 42 000 animals in 1987 
(Zwanenburg and Bowen 1990). Expanding the 
period covered by Zwanenburg and Bowen (1990) 
to 1994, and applying a different model, the Sable 
Island component of the population has increased to 
85 300 (78 000 — 95 000 95% confidence interval) 
animals (Mohn and Bowen 1996). Recent surveys to 
determine Sable Island pup production were flown 
in 1997. These surveys estimated pup production to 
be 25 200 (23 700 — 26 700 95% confidence inter- 
val) (Bowen et al. 1999) which would result in a 
total population of 144700 animals (Figure 3) 
’ (Hammill 1999). 

Much less information is available concerning the 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


size of the non-Sable Island component of the popu- 
lation, owing to the difficulties of working on the 
unstable pack-ice in the southern Gulf of St 
Lawrence. Non-Sable Island pup production has 
been determined from mark-recapture experiments 
conducted between 1984 and 1990, where the pups 
were marked on the whelping patch, and later shot 
during scientific collection programs or by recaptur- 
ing the animals live on Sable Island 3-10 months 
later (Stobo and Zwanenburg 1990; Hammill et al. 
1998). Early estimates suggested that pup production 
during the period 1984-1986 could have been as low 
as 5300 + 2600 or as high as 11 700 + 3000 animals 
(Stobo and Zwanenburg 1990). However, because 
several of the assumptions related to the Peterson 
mark-recapture model were violated, 1t was conclud- 
ed that the true pup production probably lay in 
between these estimates (Stobo and Zwanenburg 
1990). A re-analysis of these estimates and some 
new data from 1989 and 1990 indicated that pup pro- 
duction of the non-Sable Island component would 
have increased from an estimated 5900 pups in 1984 
to 9300 in 1990 for an annual rate of increase of 
7.4% (SE=2.2) (Hammill et al. 1998). More recent 
estimates of pup production from aerial surveys esti- 
mated a pup production of 11 800 (6800-16 900) in 
1996 and 7400 (4600-10 400) in 1997 resulting in a 
rate of increase of 3.4% (SE=1.5) (Hammill et al. 
1999). The lower estimates obtained in 1997 are 
attributed to high pup mortality owing to poor ice 
conditions encountered in that year (Hammill et al. 
1999). The dynamics of this component of the popu- 
lation have been modelled using a model similar to 
the one developed by Zwanenburg and Bowen 
(1990). Assuming that differences in population 
growth between the Sable and non-Sable compo- 
nents of the population are due to differences in pup 
survival rates, then the 3.4% rate of. increase is 
achieved using adult survival rates of 0.96 
(Zwanenburg and Bowen 1990), and pup survival 
rates of 0.329. This results in a 1984 population of 
28 400 in 1984 increasing to 47 100 in 1997 (Figure 
3) (Hammill 1999). Combining the estimates for the 
Sable Island and non-Sable Island components 
results in a total Northwest Atlantic Grey Seal popu- 
lation of 191 800 in 1997 (Hammill 1999). 

It is evident that the Sable Island and non-Sable 
Island components of the population must have 
undergone very different trajectories since the 1970s. 
During that decade, roughly 69% of the population 
was of non-Sable Island origin, consisting mostly of 
animals from the Gulf of St Lawrence. However, by 
1993 less than 43% of the total population would 
have been of Gulf origin (Mohn and Bowen 1996). 
Differences between the two groups likely result 
from the effects of the government sponsored cull of 
non-Sable Island animals in the whelping areas, and 
also from higher mortality rates for pups born on the 


2001 


unstable pack-ice in the Gulf of St Lawrence 
(Hammill et al. 1995). 


Habitat 

Grey Seals inhabit temperate cold waters through- 
out their range. Whelping occurs during late 
December—February (Mansfield and Beck 1977) in 
two very different environments: the unstable drift- 
ing pack-ice in the southern Gulf of St Lawrence, 
and on beaches of isolated islands (Figure 2) 
(Mansfield and Beck 1977). After breeding, animals 
move offshore onto the Scotian Shelf or over the 
Laurentian Channel between the Magdalen Islands, 
Newfoundland and Cape Breton Island, presumably 
to feed (Stobo et al. 1990: Hammill et al. 1993; 
Goulet 1996). Dives to deeper than 400 m have been 
recorded, but the majority of dives are to depths of 
70-100 m (Goulet 1996). In May and June, they 
move ashore to moult on beaches of remote, and 
often small islands or on reefs exposed at low tide 
(Mansfield and Beck 1977; Clay and Nielsen 1985). 
They remain in these areas throughout the summer 
and the fall, partitioning their time between periods 
of hauling out, and periods at sea. Movements in the 
Gulf of St Lawrence appear to be largely coastal 
(Goulet 1996), but a different pattern may occur off 
Sable Island. In european waters, animals at this time 
of year tend to move between a restricted number of 
haul out sites (Thompson et al. 1991; Sj@berg et al. 
1995). 


General Biology 
Reproduction 

The mean age for females giving birth for the first 
time is 5.5 y (SD = 0.12). Pregnancy rates for female 
Grey Seals, using the presence or absence of a foe- 
tus, are 0.18, 0.86 and 0.88 for females aged 44+, 5+ 
and 6+ years respectively (Hammill and Gosselin 
1995). Among males, a marked increase in testis 
weight is observed at age 3+ y. Based on testis mass, 
the mean age of sexual maturity is 5.6 y, and by age 
7 virtually all males are sexually mature (Hammill 
and Gosselin 1995). However, animals do not appear 
to be able to hold tenure in the whelping patch until 
the age of 11-12 y (Godsell 1991). 

Pupping occurs during late December to 
February. A female Grey Seal usually gives birth to 
a single pup after a 12-month gestation period, 
which includes a 3-month period of delayed implan- 
tation and a 9 month active gestation period. At 
birth, the pups weigh 15-17kg, and gain 
2.4—3.0 kg/d. They are weaned at a mass of 
51-56 kg after a 15—16 day lactation period (Bowen 
et al. 1992; Iverson et al. 1993; Baker et al. 1995). 
Males can be heavier at birth, grow faster, and 
weaned at a greater mass (Baker et al. 1995), but this 
has not been observed in all studies (Bowen et al. 
1992; Iverson et al. 1993). During lactation, females 
lose approximately 3—9 kg/d (Iverson et al. 1993; 


LESAGE AND HAMMILL: STATUS OF GREY SEAL IN NORTHEAST ATLANTIC 


657 


160 ~———- —---——— 


I 
@ Non-Sable Island Grey Seals: r=:3.4% 
71 OQ Sable Island Grey Seals: =12.6% 


©) e 
e@ 
eco? 


— 
”n 
S 
i=) 
oO 
— 
1) 
N 
” 
& 
BS 
= 
& 
=] 
Qa 
Oo 
oO. 


O° 
cecscsnccesessZooweee” 


fone) 
090000000° 


.— 


aa i T T T me 
1970 1975 1980 1985 1990 1995 


Year 


FIGURE 3. Changes in abundance of NW Atlantic Grey 
Seals (modified after Hammill 1999). 


Lydersen et al. 1994; Baker et al. 1995). Females in 
land-breeding colonies fast during the lactation peri- 
od (Iverson et al. 1993), but some females breeding 
on the pack-ice may feed (Lydersen et al. 1994; 
Baker et al. 1995). 

Grey Seals are polygynous (Cameron 1967). 
Males do not defend fixed territories, but instead 
compete for access to a shifting population of lactat- 
ing females (Boness and James 1979). The degree of 
polygyny is affected by habitat variability (Anderson 
and Harwood 1985; Bédard 1993; Tinker et al. 
1995). Males attending females lose 2—5 kg/d during 
the breeding season (Godsell 1991; Tinker et al. 
1995). Copulation occurs on land or in the water at 
the end of lactation (Bonner 1981). 

Grey Seal mortality rates have not been measured 
directly. Animals as old as 46+ y have been collect- 
ed from the wild (Hammill, unpubl. data). As out- 
lined above, adult survival rates of 0.96 and pup sur- 
vival rates of 0.79 were required to obtain a rate of 
increase in pup production of 12.6% for Sable 
Island. For the non-Sable Island population, the 
observed rate of increase in pup production was 
obtained with adult survival rates of 0.96 and pup 
survival rates of 0.33. These represent mean survival 
rates. 


Food Habits and Feeding 

Over 40 different prey species, including many 
commercially important species, have been identi- 
fied in the diet of Northwest Atlantic Grey Seals 
(Benoit and Bowen 1990a). Like most pinnipeds, 
strong regional and seasonal changes in Grey Seal 
diets have been observed. In the northern Gulf of St 
Lawrence, Capelin (Mallotus villosus), Lumpfish 
(Cyclopterus lumpus), Herring (Clupea harengus), 
and Atlantic Cod (Gadus morhua) are the most 
important prey species, accounting for over 60% of 
the diet by frequency of occurrence (Benoit and 
Bowen 1990b; Murie and Lavigne 1992; Proust 
1996). Seasonal changes in diet are evident, with 


658 


Capelin and Lumpfish being important prey from 
May to July. During August and September, Cod 
and Herring are the dominant prey species (Benoit 
and Bowen 1990b; Proust 1996). In the southern 
Gulf of St Lawrence, Cod, Herring and flatfish 
(Pleuronectiformes) are the most important prey 
(Benoit and Bowen 1990a). In Grey Seals collected 
from the Atlantic side of Nova Scotia and Sable 
Island, Cod, Herring, Silver Hake (Merluccius bilin- 
earis), Sand Lance (Ammodytes americanus) and 
flatfish form the most important prey (Bowen et al. 
1993; Bowen and Harrison 1994). Near Sable 
Island, Sand Lance are an important component of 
the diet throughout the year, but account for a 
greater percentage of the diet by weight during the 
winter than in summer. Cod and Silver Hake are 
consumed primarily during late summer when these 
species move into the shallower water over the off- 
shore banks surrounding Sable Island (Bowen and 
Harrison 1994). Some differences occur between 
nearshore diets from the Nova Scotia Eastern Shore 
and offshore diets of animals from around Sable 
Island (Bowen and Harrison 1994). In Grey Seals 
collected from nearshore areas, Herring and 
Mackerel (Scomber scombrus) replace Sand Lance 
and flatfishes as the most important foods (Bowen 
et al. 1993). 

Grey Seals feed primarily on fish <40cm in 
length, which for most species represent fish too 
small for the commercial fishery (Benoit and Bowen 
1990b; Murie and Lavigne 1992; Bowen et al. 1993; 
Bowen and Harrison 1994; Proust 1996). Some 
notable differences have been observed between 
studies or within studies between years. Bowen et al. 
(1994) observed that Grey Seals consumed larger 
Herring during the fall on the Scotian Shelf (mean 
length = 34.5 cm) than did Grey Seals feeding on 
Herring during summer in the northern Gulf (mean 
length = 24.9 cm) (Benoit and Bowen 1990b). More 
recently, Proust (1996) observed that Grey Seals ate 
smaller Cod in 1988 (mean length = 32.1 cm) than in 
1992 (mean length = 39.6 cm). These differences in 
the length-frequency distributions of prey consumed 
may be related to the relative abundance of particular 
year classes (Proust 1996). 


Behaviour and Adaptability 

While at sea, Grey Seals appear to travel singly. 
Haul-out sites and whelping areas are normally 
located in remote areas where access is limited 
(Mansfield and Beck 1977; Clay and Nielsen 1985; 
Lesage et al. 1995). Grey Seals react to approaching 
boats or low flying aircraft, even when these sources 
of disturbance may still be a considerable distance 
away, by entering the water and dispersing from the 
site (Lesage and Hammill, personal observation), but 
this may vary between regions. For example, on 
Sable Island Grey Seals appear to be tolerant to dis- 
- turbance and do not enter the water until approached 
closely (Lesage and Hammill, personal observation). 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Limiting Factors 

Past exploitation greatly reduced Grey Seal num- 
bers in eastern Canada (Lavigueur and Hammill 
1993). In spite of government cull programs and 
bounty programs, Grey Seals numbers appear to 
have been increasing since the early 1960s 
(Zwanenburg and Bowen 1990). Catch statistics 
(Department of Fisheries and Oceans, Ottawa) indi- 
cate that current levels of hunting remove less than 
500 animals per year from the population. Grey 
Seals can be sensitive to human disturbance. During 
the breeding season, this can result in the abandon- 
ment of the pup by the female and consequent pup 
mortality (Lesage and Hammill, personal observa- 
tion). 

Quality of the whelping habitat can have an 
important impact on the species. On small islands 
that are repeatedly disturbed, Grey Seals may aban- 
don these sites (e.g., Basques Island). For Grey Seals 
whelping on the shifting pack-ice, thin or otherwise 
poor ice will lead to high pup mortality, which will 
result in very limited or no increase in population 
size in some years. Climatic changes resulting in 
changes in ice cover in the Gulf of St Lawrence will 
affect the availability of whelping habitat. A decline 
in ice cover will force animals to whelp onshore in 
the southern Gulf of St Lawrence where they will be 
exposed to disturbance or hunting by humans. This 
could lead to a reduction in whelping activity in the 
Gulf of St Lawrence. 

Grey Seals are subject to predation by sharks. 
Little information is available, but it has been pro- 
posed that one factor contributing to the rapid 
increase in Grey Seal numbers on Sable Island has 
been an apparent decline in shark abundance (Brodie 
and Beck 1983). 

It has been hypothesised that the northern limits 
of Grey Seals are limited by the effects of cold tem- 
peratures on the thermoregulatory abilities of pups 
during their post-weaning fast (Hansen and Lavigne 
1997). Hansen and Lavigne (1997) found that the 
lower critical temperature of pups during the post- 
weaning fast was -7.1°C. The current distribution of 
the -7.5°C January-February mean air temperature 
isotherm would exclude breeding Grey Seals from 
the western portion of Northumberland Strait, and 
much of the Gulf of St Lawrence lying to the west 
and north-west of a line extending from Cape North 
(northwestern Prince Edward Island) to the Bay of 
Islands on the west coast of Newfoundland (Hansen 
and Lavigne 1997). 

High contaminant burdens could have a negative 
impact on Grey Seals. A reduction in reproductive 
rates due to uterine stenosis and uterine occlusions has 
been associated with PCB levels of 73 + 6.6 to 100 + 
13 mg kg"! wet weight in blubber from Baltic Sea 
Grey Seals (Helle et al. 1976; Bergman and Olsson 
1985). However, published information on contami- 


2001 


nants in Northwest Atlantic Grey Seals indicates that 
organochlorine levels are much lower, and some are 
declining (Addison et al. 1984). A decline is occurring 
rapidly in DDT-group insecticides, while PCB 
residues show little evidence of change, or may have 
increased. In a sample from lactating females collect- 
ed on Sable Island, DDT concentrations declined 
markedly from 12.7 + 6.2 mg kg"! in 1976 to 3.7 + 
1.4 mg kg’! in 1985, while PCB concentrations 
changed little (11.7 + 4.4 in 1976 compared to 16.2 + 
6.8 mg kg"! in 1984, and 30.3 + 17.0 mg kg"! in 1985) 
(Addison et al. 1984; Addison and Brodie 1987). In a 
more recent sample from Sable Island, total DDT lev- 
els were 1.4 + 0.4 mg kg"! and 2.5 + 1.5 mg kg"! in 
weaned pups and yearlings, respectively. PCB levels 
were 2.4 + 1.3 mg kg"! and 4.1 + 2.1 mg kg"! in pups 
and yearlings, respectively (Addison and Stobo 1993). 
No information is available for Grey Seals from the 
Gulf of St Lawrence, but in the St Lawrence Estuary 
area, levels of PCB in immature grey seals were 4.212 
+1.580mg kg"! (n=14) and in adults 9.978 
Pee moko” (n=5)). Levels of DDT were 
2.030 1.280 mg kg"! (n=14) in immature and 2.379 
+1.247 mg/kg (N=5) in adult Grey Seals (Bernt et al. 
1999), 

In 1988, a morbillivirus, phocine distemper virus 
(PDV), killed more than 18 000 Harbour Seals, and a 
small number of Grey Seals in northern Europe 
(Heide-Jérgensen et al. 1992). Morbillivirus neutral- 
ising antibodies have been identified in Grey Seals 
from the Northwest Atlantic, with the highest titers 
against phocine distemper virus (Duigan et al. 1995). 
It has been proposed that a PDV-like morbillivirus is 
enzootic in Grey Seals, and that the virus constantly 
circulates within the population. Although some 
mortality would occur, the overall effect would be to 
maintain a certain level of herd immunity, which 
would limit the potential for an epizootic outbreak 
(Duigan et al. 1995). 


Special Significance of the Species 

Grey Seals are found in north-western Atlantic 
waters throughout the year. There are conflicts with 
commercial fishermen owing to consumption of 
commercially important species of fish, damage to 
fishing gear, and through their role as a terminal host 
for the nematode parasite Pseudoterranova decipiens 
(Bowen 1990). 

Atlantic groundfish stocks collapsed during the 
early 1990s, and have shown little sign of recovery. 
Grey Seals may consume significant quantities of 
commercially important species, but the impact of 
Grey Seals on fish stocks cannot be assessed until 
mortality rates due to seal predation can be placed 
within the context of total natural mortality experi- 
enced by the fish (Mohn and Bowen 1994; Hammill 
et al. 1995). 

Damage to fishing gear in the Atlantic region was 
examined by the Royal Commission on Seals and 


LESAGE AND HAMMILL: STATUS OF GREY SEAL IN NORTHEAST ATLANTIC 


659 


Sealing (Malouf 1986). Based on a series of ques- 
tionnaires and/or personal interviews conducted in 
Nova Scotia, it was concluded that damage to fishing 
gear by both Harbour Seals and Grey Seals in 1983 
could have been as much as $1 241 000. Assuming 
that damage to gear in New Brunswick and Prince 
Edward Island was approximately one third of the 
Nova Scotia value for each province, the total value 
would have been $2 068 333. 

Four species of pinnipeds are abundant through- 
out Atlantic Canada, but the Grey Seal is the most 
important as a vector for the nematode parasite 
Pseudoterranova decipiens, known also as codworm 
or sealworm (Mansfield and Beck 1977; Bowen 
1990). Sealworm is considered to be mildly patho- 
genic in humans if consumed in raw or poorly 
cooked fish. However, their major impact on fish- 
eries is considered to be a cosmetic one, with high 
infections rendering fish unappealing to consumers. 
The cost of removing larvae from cod fillets alone 
was estimated to be in excess of $29 million in 
Atlantic Canada in 1982 (Bowen 1990). Surveys 
conducted during the 1980s and in 1990 indicate that 
worm burdens in fish have increased in both the Gulf 
of St Lawrence and on the Scotian Shelf 
(McClelland et al. 1985; Boily and Marcogliese 
1995). These increases may be linked to an increas- 
ing Grey Seal population (Marcogliese et al 1996). 

Owing to its scarcity during the 1800s and in the 
present century, Grey Seals have not been hunted 
commercially. However, with the recent expansion 
of the Grey Seal population, there is some interest in 
hunting these animals. Grey Seals offer limited 
potential for ecotourism in the breeding colonies 
because of their large size and the aggressive nature 
of both adults and pups. 

The Grey Seal shows perhaps the greatest vari- 
ability of all pinnipeds in the selection of whelping 
habitat. Habitat types vary from sandy beaches, 
caves accessible only from the sea, to rocky islands 
and pack-ice. The occurrence of Grey Seals breeding 
on the pack-ice in the southern Gulf of St Lawrence 
is very unusual and occurs in only one other place in 
the world, the Baltic Sea. 


Evaluation 

Grey Seals appear to be susceptible to disturbance 
at whelping and haul-out sites. Consequently, they 
tend to be more abundant in sparsely inhabited areas. 
Presently, there is no commercial hunt for this 
species, although this could change in the near future. 

The Northwest Atlantic Grey Seal population is 
probably more abundant now than at any previous 
time in the present century. Information on trends in 
pup production (an index of population size) indicate 
that the population is continuing to expand. Grey 
Seal populations whelping on the pack-ice in the 
southern Gulf of St Lawrence and on Sable Island 
are both abundant, but a series of winters with poor 


660 


ice conditions will have a negative impact on the 
Gulf component of the Northwest Atlantic Grey Seal 
population, possibly leading to no or negative 
growth. It is recommended that, for the present time, 
the species should not be given any COSEWIC cate- 
gory, but the Gulf component should be monitored 
for any change in abundance. 


Acknowledgments 

We thank Gerry Conway, Department of Fisheries 
and Oceans for his continued monitoring of Grey Seal 
pup production along the Nova Scotia Eastern shore, 
and for reporting the new colony on Hay Island. 


Literature Cited 

Addison, R.F., and P. F. Brodie. 1987. Transfer of 
organochlorine residues from blubber through the circu- 
latory system to milk in the lactating grey seal Hali- 
choerus grypus. Canadian Journal of Fisheries and 
Aquatic Sciences 44: 782-786. 

Addison, R.F., and W. T. Stobo. 1993. Organochlorine 
residue concentrations and burdens in grey seal (Hali- 
choerus grypus) blubber during the first year of life. 
Journal of Zoology, London 230: 443-450. 

Addison, R.F., P. F. Brodie, and M. E. Zinck. 1984. 
DDT has declined more than PCBs in eastern Canadian 
seals during the 1970s. Environmental Science Tech- 
nology 18: 935-937. 

Anderson, S.S., and J. Harwood. 1985. Time budgets 
and topography: how energy reserves and terrain deter- 
mine the breeding behaviour of grey seals. Animal 
Behaviour 33: 1343-1348. 

Baker, S. R., C. Barrette, and M.O. Hammill. 1995. 
Mass transfer during lactation of an ice-breeding pin- 
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Accepted 19 February 2002 


Status of Harbour Seals, Phoca vitulina, in Canada* 


ROBIN W. BAIRD 


Biology Department, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada 
Current address: National Marine Fisheries Service, NOAA, 101 Pivers Island Road, Beaufort, North Carolina 28516 USA 


Baird, Robin W. 2001. Status of Harbour Seals, Phoca vitulina, in Canada. Canadian Field-Naturalist 115(4): 663-675. 


The Harbour Seal (Phoca vitulina) inhabits all three of Canada’s coastlines, as well as a number of fresh-water systems. 
Three subspecies are recognized from Canadian waters, Phoca vitulina richardsi from the Pacific coast, Phoca vitulina 
concolor from the Atlantic and Arctic coasts, and Phoca vitulina mellonae from several freshwater lakes on the Ungava 
Peninsula, Quebec. This report reviews the status and management of Phoca vitulina richardsi and Phoca vitulina concolor 
in Canadian waters, discussing distribution, movements, population discrimination, population size and trends, and threats 
to this species in Canada. The Harbour Seal population in western Canada is large and has been increasing in size. While 
there are a number of actual or potential anthropogenic threats, including: overfishing, immunosuppresion due to accumu- 
lation of toxins, and illegal killing associated with aquaculture operations, the western Canadian population should proba- 
bly be listed as not at risk. Little recent research has been undertaken on Harbour Seals in the Canadian Arctic or for most 
areas off eastern Canada, and insufficient information is available to assess the status of these populations. 


Key Words: Harbour Seal, Phoque Commun, Loups Marin, Kasigiak, Phoca vitulina richardsi, Phoca vitulina concolor, 


Pacific, Atlantic, Arctic, Phocid, status, pinniped. 


The Harbour Seal (Phoca vitulina Linnaeus, 
1758, Figure 1), known in Quebec as Phoque 
Commun or Loups Marin, in parts of northern 
Canada as the Ranger Seal or Kasigiak, and in 
Europe as the Common Seal, is a small coastal 
Phocid, and is one of the most widely distributed 
pinnipeds in the northern hemisphere. On both the 
east and west coasts of Canada Harbour Seals aver- 
age about 80 cm at birth (McLaren 1993). The 
average adult length is about 1.5 m, and they have 
been recorded to reach a maximum length of about 
1.9 m (Boulva 1971; Bigg 1981; McLaren 1993). 
Males are usually slightly larger than females (5% 
off Nova Scotia, 13% off British Columbia; 
McLaren 1993). Harbour Seals have been described 
as “large-headed, short-bodied and short-limbed” 
(Figures 1 and 2), and have an extremely variable 
pelage pattern with spots, rings and blotches which 
range in color from light grey to dark brown or 
black on either a light or dark background (Stutz 
1967; Bigg 1981). Harbour Seals usually lose their 
lanugo (white) coat in utero, though Boulva (1971) 
and Oftedal et al. (1991) noted 16% and 6% 
(respectively) of the pups born at Sable Island have 
a lanugo coat. Individuals with rust-coloured coats, 
particularly on the head and upper body, have also 
been observed (Jefferson et al. 1993). The eyes are 
large and set close together, and the nostrils form a 
distinct “V” shape. Harbour Seals are usually very 


*Reviewed and approved by COSEWIC, April 1999, status 
assigned — Atlantic and Arctic coastal waters Indeter- 
minate, North Pacific coastal waters Not At Risk. 


difficult to approach on shore. They are gregarious 
on land, but usually solitary or in very small groups 
when seen in the water. Most of time Harbour Seals 
spend on shore they just lay on the substrate with 
periodic checks of their surroundings, but often 
also lie with their head and hind flippers elevated in 
a characteristic crescent position (Katona et al. 
1993; Figure 2). 

Considerable taxonomic uncertainty has existed 
regarding Harbour Seals, both at the sub-specific and 
specific levels (e.g., McLaren 1966; Shaughnessy 
and Fay 1977; Bigg 1981; Smith et al. 1994; 
O’Corry-Crowe and Westlake 1997). Until recently, 
Harbour Seals and Largha Seals (Phoca largha) 
were frequently lumped as one species, Phoca vituli- 
na, resulting in some confusion as to, for example, 
geographic range (O’Corry-Crowe and Westlake 
1997). Three subspecies of the Harbour Seal are cur- 
rently recognized from Canadian waters, with Phoca 
vitulina richardsi in the Pacific, Phoca vitulina con- 
color in the Atlantic and Arctic, and Phoca vitulina 
mellonae, the Lacs des Loups Marins Harbour Seal, 
found exclusively in freshwater lakes in Quebec's 
Ungava Peninsula (Smith et al. 1994). The status of 
this latter population has been recently reviewed 
(Smith 1997) and classified by COSEWIC (the 
Committee on the Status of Endangered Wildlife in 
Canada) as “Vulnerable”’. 

The purpose of this report is to review available 
information and assess the status of the other two 
subspecies of Harbour Seal found in Canadian 
waters, on behalf of COSEWIC. While they are one 
of the most well-studied pinnipeds (Cottrell 1995), 
surprisingly little recent information is available 
regarding population size, trends, and sources and 


663 


664 


THE CANADIAN FIELD-NATURALIST 


Nolmals 


FiGurRE 1. Harbour Seal hauled out on rocky substrate off southern Vancouver Island. Photo by the 
author. 


levels of anthropogenic mortality in Canadian 
waters. 


Distribution 

Harbour Seals have a coastal distribution in tem- 
perate, sub-Arctic and some Arctic waters through- 
out the northern hemisphere, with some populations 
also occurring in fresh-water systems. In the Pacific, 
they are found from northern Japan (Hokkaido), 
through the Aleutians and the Gulf of Alaska, and 
south along the coast of North America as far as 
western central Baja California, Mexico. Records 
reported from Alaskan waters of the Beaufort Sea 
(e.g., Mansfield 1967) are of Largha Seals. In the 
Atlantic, Harbour Seals have been documented as far 
south as Florida (Reeves et al. 1992), and occur reg- 
ularly from New York and New England north to 
Greenland, Iceland, the UK, Norway and Svalbard 
(Wiig 1989), and south as far as Brittany, France. 

In Canada, Harbour Seals are found in coastal 
waters in the Pacific and Atlantic oceans, as well as 
in parts of Hudson Strait, Ungava Bay, Hudson Bay 
and around Baffin Island (Figure 3; Mansfield 1967; 
Beck et al. 1970; Bigg 1981). While sightings in 
Canadian waters have been reported as far north as 
Ellesmere Island (79°N), no recent information is 
available on their distribution in the Canadian Arctic, 
and there is some evidence to suggest that their dis- 
tribution in the Arctic may have changed. Mansfield 
(1967) noted that Harbour Seals had been eliminated 
from some areas in the Canadian Arctic by native 
hunting. 

As mentioned above, Harbour Seals can also be 
found in fresh water, including not only the popula- 
tion which resides year-round in the Seal Lakes 
(Lacs des Loups Marin) in northern Quebec, but also 


rivers and lakes on other coastlines. In western and 
northwestern Hudson Bay Harbour Seals have been 
recorded as far as 240 km inland in several rivers and 
lakes (Mansfield 1967; Beck et al. 1970), and on the 
Pacific coast, Harbour Seals also enter small rivers 
and lakes, occasionally as far as 300 km inland 
(Fisher 1952). In eastern Canada, Harbour Seals 
were once found in a number of fresh water systems, 
including Lake Champlain and Lake Ontario (Allen 
1880). It appears their use of fresh water systems in 
eastern Canada has declined; Boulva and McLaren 
(1979) noted that Harbour Seals seldom reach 
Montreal in the St. Lawrence River. Similarly, 
Lesage et al. (1995b) report relatively few in the 
Saguenay River in 1994, while Boulva and McLaren 
(1979) reported a population of approximately 100 
individuals in that river in 1973 (it should be noted 
however that the methods of these two studies differ 
greatly, and numbers in each are not comparable). 


Movements and Population Discrimination 

On land, Harbour Seals exhibit fairly poor mobili- 
ty, with movements typically restricted to tens of 
meters. However, Renouf and James (1975) state 
that pregnant females have been known to travel 
more than a kilometer overland on Sable Island, to 
give birth on the shores of inland lakes. 

Harbour Seals are generally considered to be non- 
migratory (Bigg 1981), being present in most areas 
year-round, and showing considerable site fidelity 
(e.g., Olesiuk et al. 1995). However, extensive 
movements do occur. Based on seasonal decreases in 
counts at haul-outs in the Bay of Fundy, and corre- 
sponding increases in numbers hauled out in Maine, 
Rosenfeld et al. (1988) suggested that there is a sea- 
sonal movement of Harbour Seals between these two 


2001 


BAIRD: STATUS OF HARBOUR SEALS 


665 


FiGurE 2. Harbour Seals hauled out on dock, Victoria Harbour. Note the individual on the left is lying with head and tail 
raised, a typical posture for Harbour Seals. Photo by the author. 


areas. Using satellite tags on individuals in the St. 
Lawrence estuary, Lesage et al. (1995a) documented 
a migration of one individual which moved 520 km 
to overwinter, and returned to the same summering 
area the next year. In terms of dispersal from breed- 
ing areas, Beck (1983) noted that juveniles tagged at 
Sable Island, off Nova Scotia, moved to various 
mainland sites, and included one individual which 
moved to New Jersey, a straight line distance of 
1475 km. 

Despite such long-ranging movements, there is a 
variety of evidence that suggest Harbour Seals on the 
west coast of Canada may be subdivided into two or 
more populations (Stutz 1967; Temte et al. 1991; 
Burg 1996; Lamont et al. 1996). Stutz (1967) noted 
geographic variation in pigmentation patterns for sev- 
eral areas on the coast of British Columbia and south- 
east Alaska and suggested these reflect subdivision of 
the population. Yochem et al. (1990) documented 
geographic variation in pigmentation patterns off 
California which suggest limited movements between 
areas. Differences in pupping seasonality between 
various areas on the west coast (Bigg 1969b) also 
imply more than one population (Temte et al. 1991). 
Burg (1996) recently examined mitochondrial and 
microsatellite data from animals throughout British 
Columbia and in southeast Alaska, and concluded 
that evidence suggests population segregation. Based 
on pigmentation patterns and dentition, suggestions 
of population discreteness have been made for 
Harbour Seals on Sable Island (Boulva and McLaren 
1979), though recent microsatellite data from Sable 
Island (Coltman, Bowen and Wright, unpublished, 
cited in Whitehead et al. 1998) suggest mixing with 
mainland populations. 


Protection 
International 

Harbour Seals are not listed under the Convention 
on International Trade in Endangered Species of 
Wild Fauna and Flora (CITES), thus international 
trade is not monitored or regulated. 


National 

Canada: Two factors are important in the protec- 
tion of a species, the legal system in place which 
prohibits or regulates hunts or kills, and the system 
of monitoring and enforcement of the regulations or 
rules which exist. In Canada, management authority 
for Harbour Seals lies with the federal Department of 
Fisheries and Oceans through the Marine Mammal 
Regulations of 1993 (promulgated under the 
Fisheries Act of Canada, 1867). These regulations 
theoretically control the hunting of this species in 
Canadian waters. All individuals except aboriginals 
are required to obtain a “Fishing Licence” to hunt 
seals, and fees for such a licence are low ($5). 
Issuance of licences is at the discretion of the 
Minister of Fisheries and Oceans, but for Harbour 
Seals, all areas are currently closed for hunting 
(Marine Mammal Regulations; J. Conway, personal 
communication). D. Petrachenko (in litt.) notes that 
Fisheries and Oceans has issued licences to aborigi- 
nals in British Columbia for a small scale harvest. In 
Arctic regions all hunting is likely to be by aborigi- 
nals (who do not require licences), thus no protection 
appears to be in place. In British Columbia, aquacul- 
ture operators are permitted to kill nuisance animals 
around their net pens (D. Petrachenko, in litt.). Some 
nuisance animals were killed at aquaculture opera- 
tions in New Brunswick as part of a pilot licencing 


666 


system, but such killing is currently prohibited (J. 
Conway, personal communication). “Disturbance” is 
also prohibited through these regulations, except 
when hunting under licence. The operation of air- 
craft within 600 m of any live seal on land is also 
prohibited. Smith (1997) notes that freshwater seals 
north of the 55th parallel are listed as protected 
under the James Bay and Northern Quebec agree- 
ment, but that this protection does not have the force 
of law. Some haul-out and breeding sites are protect- 
ed from development by both provincial and federal 
governments through provincial parks and/or ecolog- 
ical reserves as well as federal parks (such designa- 
tions also provide limited protection from land-based 
disturbance). 


In terms of monitoring or enforcement of regula- 
tions, little information is available. Disturbance of 
animals regularly occurs at haulout sites by boats, 
planes, and people on land (Baird, personal observa- 
tions), and no monitoring or enforcement action is 
taken. As well, there have been numerous reports of 
illegal shooting of animals around aquaculture oper- 
ations on the west and east coasts, as well as shoot- 
ing by fishermen, though no prosecutorial action has 
been taken (H. Breen, J. Conway, D. Tobin, personal 
communication). Such a lack of monitoring or 
enforcement of these regulations leaves their effec- 
tiveness in question. 


United States: Killing or disturbance of Harbour 
Seals in the United States is prohibited under the 
Marine Mammal Protection Act. 


Greenland: Since 1960, hunting of adult Harbour 
Seals in Greenland has been prohibited from May 
through September. Some specific regions have 
more restrictive regulations, with hunting of adults 
completely forbidden. Restrictions have been recent- 
ly reviewed by Teilmann and Dietz (1994). 


Population Numbers, Sizes and Trends 
Populations on both the Pacific and Atlantic 
coasts of Canada were substantially reduced due to 
long-term bounty or culling programs and/or com- 
mercial hunts (Bigg 1969a; Boulva and McLaren 
1979; Olesiuk et al. 1990a), which ended in the late 
1960s and early 1970s (1969 in British Columbia, 
1976 in the Atlantic). On the Pacific coast of 
Canada, the Harbour Seal is the most abundant 
marine mammal in the province, and the population 
has been increasing since the end of culling and may 
be near original levels (Olesiuk et al. 1990a). The 
most recent published data from British Columbia 
(from 1988) suggests the population was between 
75 000 and 88 000 animals. The trend in population 
growth at that time suggested a continued increase 
(Olesiuk et al. 1990a), though Smith (1994) noted 
' that there is some indication from recent surveys that 
the increase is beginning to level off. Between 1973 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


and 1988, the British Columbia population was esti- 
mated to increase at about 12.5% per year (Olesiuk 
et al. 1990a). 

Populations which border British Columbia 
appear to be increasing or stable. From 1978 through 
1993, counts of Harbour Seals in the neighboring 
waters to the south, in Washington state, increased at 
an average rate of almost 8% (Huber 1995); surveys 
from neighboring waters to the north, in southeast 
Alaska, have generally shown increases or stable 
populations (Hill et al. 1997). 

Off Canada's east coast, information on popula- 
tion sizes or trends is less complete. Numbers from 
eastern Canadian waters south of Labrador in 1973 
were estimated by Boulva and McLaren (1979) 
using questionnaire surveys, interviews and distribu- 
tion of bounty kills. The total population off eastern 
Canada at that time was estimated to be 12 700 indi- 
viduals, with some local areas showing decreasing 
numbers and other areas having stable numbers. 
Stobo and Fowler (1994) present data from aerial 
surveys in the Bay of Fundy and off southwest Nova 
Scotia from 1985-1987 and from 1991-1992, and 
suggest that Harbour Seal abundance has increased 
between those periods. However, the rate of growth 
and current population size for that area is unknown 
(Stobo and Fowler 1994). In the late 1980s, the 
Sable Island population was the largest in eastern 
Canada. However, numbers of animals on Sable 
Island have decreased drastically in recent years (D. 
Bowen, W. Stobo, personal communication; Ellis 
1998). Trends in the number of pups born on Sable 
Island between 1978 and 1996 have been presented 
by Ellis (1998). The number of pups born annually 
have shown a steady decline since 1989, when about 
600 pups were born, through to 1997, when only 30 
pups were born (Ellis 1998; D. Bowen, personal 
communication), and the number of adults of both 
sexes has also dramatically declined (Ellis 1998). 
Causes of this decline are unknown, but could 
include predation by sharks, competition with Grey 
Seals, and/or movements of individuals away from 
Sable Island (D. Bowen, W. Stobo, personal commu- 
nication; Ellis 1998). Recent (1994) information on 
Harbour Seals in the St. Lawrence Estuary has been 
provided by Lesage et al. (1995b). A total of 389 
Harbour Seals were counted, but it is unclear what 
proportion of the population this count represents. 
While this number is. substantially lower than the 
710 seals estimated to live in the area in 1973 
(Boulva and McLaren 1979), Lesage et al. (1995b) 
note that differences in methods between the two 
studies make any direct comparison impossible. 
Some distributional data, from a mail-out survey to 
fishermen, was recently collected for Prince Edward 
Island (D. Cairns, Department of Fisheries and 
Oceans, PEI, personal communication). No recent 
information on Harbour Seals in Newfoundland, 
Labrador or the outer Gulf of St. Lawrence has been 


2001 


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BAIRD: STATUS OF HARBOUR SEALS 


FiGurRE 3. Distribution of Harbour Seals in Canadian waters. 


collected (J. Lawson, J. Lien, G. Stenson, personal 
communication). 

No substantive information is available regarding 
population size or trends of Harbour Seals in the 
Canadian Arctic (though see Remnant 1997 for a 
recent assessment of numbers in the Lower Churchill 
River). 

On the east coast there are three countries/territo- 
ries which likely exchange Harbour Seals with 


Canadian populations: the United States, Greenland, 
and the French islands of St. Pierre and Miquelon. 
The Harbour Seal population in adjoining U.S. 
waters appears to be increasing, with an 8.7% annual 
rate of increase in Maine coastal waters based on 
counts between 1981 and 1993 (Kenney and Gilbert 
1994; Waring et al. 1997). In Greenland, Teilmann 
and Dietz (1994) report an apparent decline in 
Harbour Seals numbers over the last 100 years, and 


667 


668 


suggest that hunting may have been the primary 
cause for this decline. Harbour Seal numbers at the 
French islands of St. Pierre and Miquelon, off the 
south coast of Newfoundland, appeared to increase 
between 1970 and 1982 (Ling et al. 1974; Davis and 
Renouf 1987), but no recent information is available. 


Habitat 

Like all pinnipeds, Harbour Seals utilize both 
aquatic and terrestrial habitats. In terms of aquatic 
habitats, Harbour Seals are generally a near-shore, 
coastal species, though movements in pelagic waters 
have been documented (Beck 1983). As noted 
above, Harbour Seals also inhabit fresh-water sys- 
tems and may spend considerable time in river estu- 
aries. Use of the aquatic environment, i.e., which 
areas of the water column Harbour Seals tend to use 
most, may differ depending on time of year (breed- 
ing versus non-breeding, or in response to local con- 
centrations of prey), sex (since males during the 
breeding season may remain closer to breeding sites; 
e.g., Coltman et al. 1997; Van Parijs et al. 1997), and 
location, due to bathymetry, or since predation pres- 
sure may differ between east and west coasts of 
Canada, thus affecting diving behaviour (cf. Le 
Boeuf and Crocker 1996). In the Arctic, Mansfield 
(1967) noted that Harbour Seals are largely restricted 
to areas of high current flow where the surface is 
kept clear of ice. 

In terms of terrestrial habitats used, Harbour Seals 
haul out on both sand and rock substrates, usually on 
isolated rocks or islets (without land-based preda- 
tors), on sand bars, and occasionally in small sea 
caves, including some on large islands with terrestri- 
al predators, e.g., Vancouver Island. Harbour Seals 
also make use of man-made structures such as log 
booms and recreational floats for pupping and haul- 
ing out (e.g., Cottrell 1995; Figure 2). In some areas 
(e.g., southeast Alaska) Harbour Seals will also pup 
on ice calved off tidal glaciers (e.g., Matthews 
1995). Areas used for pupping are also used during 
the non-breeding season as haul-out sites. Age- and 
sex-based segregation occurs at some haul-out sites, 
and may vary according to season (e.g., Allen et al. 
1988; Kovacs et al. 1990; Whitman and Payne 
1990). Understanding which factors influence haul- 
ing out behaviour is important both for calibrating 
surveys (e.g., Olesiuk et al. 1990), as well as for 
understanding how disease transfer at haulout sites 
might be influenced by environmental fluctuations 
(Lavigne and Schmitz 1990; Grellier et al. 1996). A 
number of factors seem to influence hauling out 
behaviour, including haul-out substrate, tide height, 
time of year (relative to breeding and moulting peri- 
ods), time of day, temperature, wind speed, precipi- 
tation, cloud cover, the occurrence of storms, distur- 
bance, El Nifio events, and location (Boulva and 
~ McLaren 1979; Pauli and Terhune 1987a, 1987b; 
Yochem et al. 1987; Watts 1992, 1993: Grellier et al. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


1996; Hanan 1996), thus it seems that location- 
specific studies are needed for developing survey 
calibration factors. Use of land-based sites is typical- 
ly restricted to a few tens of meters from shore 
(though see exception above in Distribution and 
Movements). 


General Biology 

Harbour Seals can be quite gregarious on land 
(Figure 4). Olesiuk et al. (1990) note a mean haul- 
out size in the Strait of Georgia of about 22 individu- 
als, but groups of several thousand have been record- 
ed (Bigg 1981). While in groups Harbour Seals 
typically maintain some distance between individu- 
als (Sullivan 1982). 

Harbour Seals give birth to a single pup, and have 
a Clearly defined pupping season that typically lasts 
one to two months in any particular area (Bigg 
1981). Timing of pupping varies geographically 
(Bigg 1969b; Temte et al. 1991), but generally 
occurs between May and July. Unlike most other 
phocids, Harbour Seal pups follow their mothers into 
the water within hours of birth (Lawson and Renouf 
1985). Pups on Sable Island are weaned at 24 days 
of age (Muelbert and Bowen 1993), and females 
come into estrous within two weeks after weaning 
(Bigg 1969a). Females mature at between 3 and 6 
years of age, with most maturing by 5 years (Bigg 
1969a). Mortality rates have recently been summa- 
rized by Heide-Jorgensen and Harkonen (1988). 
Mortality of animals in the first year can be quite 
high, ranging from 0.20 to 0.60. For individuals 
older than one year, annual mortality ranges betwen 
0.05 and 0.20, and males older than five years show 
higher mortality than females (e.g., Pitcher 1990). 
Maximum recorded longevity is 32 years of age 
(Pitcher and Calkins 1979). 

While numerous studies have been undertaken on 
the diet of Harbour Seals in Canadian and adjoining 
waters, characterization of the diet is exacerbated by 
biases in techniques to study diet (Harvey 1989; 
Cottrell et al. 1996), as well as strong seasonal, geo- 
graphical, age and habitat-based variation (Bigg 
1973; Bigg et al. 1990; Olesiuk 1993; Cottrell 1995; 
Bowen and Harrison 1996; Iverson et al. 1997; Tollit 
et al. 1998). In general, Harbour Seals have an 
extremely diverse diet (Bigg 1981), usually taking 
advantage of locally abundant prey. In the Strait of 
Georgia, Pacific Hake (Merluccius productus) and 
Pacific Herring (Clupea pallasi) account for 75% of 
the diet both in terms of energy and biomass, while 
Salmon (Oncorhynchus spp.), Plainfin Midshipman 
(Porichthys notatus), Lingcod (Ophiodon elongatus) 
and others comprise the remaining prey (Olesiuk et 
al. 1990b; Olesiuk 1993). Their tendency to move 
into river mouths following salmon runs (Fisher 
1952; Bigg et al. 1990; Cottrell 1995) has caused 
considerable conflict with fishermen. Off eastern 
Canada, recent work by Bowen and Harrison (1996) 


2001 


BAIRD: STATUS OF HARBOUR SEALS 


669 


FIGURE 4. Group of Harbour Seals on sandy beach at Smith Island, Washington State. Photo by the author. 


in two areas off Nova Scotia and New Brunswick 
suggest that Atlantic Herring (Clupea harengus), 
Atlantic Cod (Gadus morhua), Polluck (Pollachius 
virens), and Short-finned Squid (e.g., Illex illecebro- 
sus) appear to comprise the majority of prey. Bowen 
and Harrison (1996) document geographic and inter- 
annual variability in prey taken in their study, and 
other evidence also suggests strong geographic or 
temporal variation. For example, Payne and Selzer 
(1989) note that American Sandlance (Ammodytes 
americanus) dominated the diet of Harbour Seals off 
Cape Cod, and a small sample of stomach contents 
from Sable Island examined by Walker and Bowen 
(1993) contained only Sandlance. Little has been 
reported on their diet in Arctic waters, though Beck 
et al. (1970) report stomach contents of one individ- 
ual taken in fresh water, which contained Lake Trout 
(Salvelinus namaycush) and Whitefish (Coregonus 
clupeaformis). 


Sources of Mortality and Potentially 
Limiting Factors 

Potentially limiting factors can be from either 
anthropogenic or natural sources. Such factors could 
either directly or indirectly cause the death of ani- 
mals, or result in decreased reproductive rates. 
Anthropogenic factors which may contribute to pop- 
ulation declines or limits include: incidental mortali- 
ty in fisheries, direct killing (illegal or permitted 
culling associated with aquaculture operations and 


fisheries, as well as small-scale harvesting by 
natives), oil spills, accumulation of persistent toxins, 
disturbance at breeding colonies by tourism, coastal 
development, vessel traffic or researchers, displace- 
ment from feeding or breeding areas by acoustic 
harassment (e.g., high amplitude seal “scarers” at 
aquaculture operations), and depleted food sources 
from competition with human fisheries. 

Historically Harbour Seal populations in both 
eastern and western Canada were drastically reduced 
from direct kills in both control or bounty programs 
and/or for harvesting of pelts (Bigg 1969a; Boulva 
and McLaren 1979). Re-initiation of such bounty or 
culling programs have frequently been suggested by 
fishing groups, in response to perceived or actual 
conflicts with fisheries. In British Columbia, direct 
takes occur from a number of sources. In terms of 
harvesting by natives, Fisheries and Oceans Canada 
has issued licences for a coast-wide harvest totaling 
less than 100 individuals (D. Petrachenko, in litt.). 
Aquaculture operations in both eastern Canada and 
British Columbia are licenced to shoot nuisance 
seals. In British Columbia a total of about 500 
Harbour Seals are reported to be killed annually 
under these permits (D. Petrachenko, in litt.). 
However, reports of substantial illegal kills at aqua- 
culture operations suggest that the number killed 
may be much greater (H. Breen, personal communi- 
cation). In 1997 Fisheries and Oceans Canada under- 
took a small cull (approximately 25 seals) in one 


670 


area on the British Columbia coast. Levels of direct 
takes in Arctic and eastern Canadian waters are 
unknown. Remnant (1997) notes that some hunting 
of seals occurrs in the freshwater portion of the 
Churchill River, and that Manitoba Natural Resour- 
ces annually harvests some seals in the same area for 
bait for live-trapping Polar Bears (Ursus maritimus). 
I. McLaren (personal communication) noted that 
pelts of Harbour Seals are prized by Arctic hunters, 
and Mansfield (1967) stated that given the extremely 
localized distribution of Harbour Seals in Arctic 
waters, this species is an easy target for native 
hunters, and their “future [is] somewhat precarious”. 

Harbour Seal bycatch in gillnet fisheries has been 
well-documented in California, Washington and 
Alaska (Barlow et al. 1994; Read 1994), but little 
information is available from Canadian waters, prob- 
ably due to the lack of observer programs on fishing 
vessels. Reports have been made of Harbour Seals 
being taken on longlines in eastern Canada, and such 
seals drown, since the lines are weighted to the bot- 
tom (W. Stobo, personal communication). Harbour 
Seal mortality in the Smelt cage fishery off Prince 
Edward Island have also been reported (P.-Y. 
Daoust, personal communication). Off Norway, 
Bekkby and Bjorge (1998) suggest that mortality in 
fishing gear may control population growth, thus 
some effort in assessing mortality in fishing gear in 
Canadian waters is probably warranted. Hooking of 
Harbour Seals on sports fishing lines also occurs (R. 
Bates, personal communication). While it seems 
unlikely these animals are directly killed as a result, 
retention of hooks and trailing lines in the digestive 
tract of Harbour Seals could cause problems associ- 
ated with feeding or infection. 

In heavily-populated areas, pups are occasionally 
“kidnapped” by well-meaning people who assume 
they have been abandoned, while a mother is out for- 
aging or when she is disturbed into the water by 
approaching humans. Such pups are often raised at 
rehabilitation facilities and released back into the 
wild, but little information is available on survival 
rates. Vessel collisions with Harbour Seals do occur 
in Canadian waters (K. Langelier, personal communi- 
cation), though there is no information to assess how 
frequently such collisions occur or whether they 
always result in death. A. Morton (personal commu- 
nication) has noted an increase in the frequency of 
vessel collisions in one area on the British Columbia 
coast, which has corresponded with the initiation and 
use of high-intensity acoustic harassment devices at 
nearby aquaculture operations (suggesting that 
Harbour Seals might be deafened by such devices). 
Disturbance of Harbour Seals at pupping or haul out 
sites, resulting in sudden movement of all hauled-out 
animals into water, occurs regularly in some areas 
(e.g., Kovacs et al. 1990). Such disturbance may 
potentially result in separation of mothers and pups, 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


as well as injury of individuals, or increased vulnera- 
bility to predation by sharks or killer whales, due to 
increased time in the water. There are numerous 
sources of such disturbance, including close 
approaches by private or commercial vessels engaged 
in fishing, wildlife viewing, or transiting through an 
area, overflights by aircraft (military and private), 
and approaches by dogs or people on land (including 
researchers). In one area, Race Rocks. at the southern 
tip of Vancouver Island, military activity at a site 
used for explosive testing regularly results in distur- 
bance of hauled out seals. No information is available 
to estimate the magnitude of the impacts of distur- 
bance on Harbour Seals in Canada, but its effects 
may not be trivial. For example, Johnson (1977) esti- 
mated that overflights by aircraft may have been 
responsible for the deaths of 10% of approximately 
2000 Harbour Seal pups born on an island in Alaska 
in 1976. A review of Harbour Seal reactions to air- 
craft and other sources of disturbance is presented in 
Richardson et al. (1995). 

The role of contaminants in immunosuppression 
of Harbour Seals and other marine mammals has 
received considerable attention in recent years (e.g., 
Ross et al. 1995, 1996, 1997; see also Schandorff 
1997a, 1997b). As a result of captive studies which 
demonstrated that ambient contaminant levels in 
Baltic Sea herring were immunotoxic when fed to 
Harbour Seals, it is now thought that the 1988 
Morbilivirus-associated mass mortality among north- 
ern European Harbour Seals may have been exacer- 
bated by contaminants (Ross et al, 1996, 1997; de 
Swart et al. 1996). From the British Columbia coast, 
Ross et al. (1998) report on levels of a number of 
contaminants (PCBs, PCDDs and PCDFs) in 
Harbour Seals, and note that 2 of 24 recently weaned 
pups sampled had levels higher than those found to 
produce immunotoxicity in captive Harbour Seals. 
Ross (personal communication) also notes that male 
Harbour Seals above about seven years of age in the 
Strait of Georgia also have levels above the point 
which produces immunotoxity. Given the exponen- 
tial rate of increase of the Harbour Seal population 
on much of the British Columbia coast (Olesiuk et 
al. 1990), it is clear that such toxins do not have a 
substantial impact on reproduction or longevity, 
although their potential role in mass mortalities (and 
the huge impacts such mortalities may have on popu- 
lation size, see below) does warrant concern. 

The wide-spread distribution of Harbour Seals 
along Canada’s coasts make them less likely to be 
seriously impacted on a population level by an oil 
spill than species whose distributions are more limited. 
However, their habit of hauling out in inter-tidal areas 
would bring them into direct contact with spilled oil, 
and spills (and related spill cleanup activities) occur- 
ring during pupping periods could have the potential 
to result in separations of mothers and offspring. Frost 


2001 


et al. (1994) discuss a variety of impacts on Harbour 
Seals from a major oil spill in Prince William Sound, 
Alaska, and conclude that seals became coated with 
oil, oil was incorporated into tissues, seals behaved 
abnormally, and pathological damage occurred. 
Comparisons of counts within Prince William Sound 
before and after the spill showed a significant decline 
in numbers at oiled sites compared to non-oiled sites 
(Frost et al. 1994). 

Indirect effects including competition with human 
fisheries may be important, though little information 
is available to assess such threats (see Baird et al. 
1992; Lavigne 1995; Trites 1997). Thompson et al. 
(1997) note that in times of shortages of preferred 
prey, switches to alternate prey items may result in 
haematological changes leading to anemia. | 

A number of natural sources of mortality have 
been identified, including predation, separation of 
dependent pups from their mothers, injury during 
storms, and diseases. Predation by Killer Whales 
(Orcinus orca) and sharks has long been known as 
a source of mortality (Scheffer and Slipp 1944). 
Mortality due to predation may be quite high in 
some areas. Harbour Seals appear to be the primary 
prey taken by “transient” Killer Whales around 
southern Vancouver Island (Baird and Dill 1996), 
and Watts (1996) noted that an individual seal may 
face a 50-80% chance of being eaten by a Killer 
Whale before reaching reproductive age. Off Sable 
Island, Nova Scotia, sharks are thought to be a 
major source of mortality (Beck 1983; Ellis 1998; 
W. Stobo personal communication). Some preda- 
tion by Northern Sea Lions, Eumetopias jubatus 
(Pitcher and Fay 1982) as well as Coyotes (Steiger 
et al. 1989) has been documented, and aggression 
by Northern Elephant Seals (Mirounga angu- 
stirostris) towards Harbour Seals has been seen at 
one site in California (Mortenson and Follis 1997). 
Interspecific competition with other rapidly 
increasing pinniped populations may also influence 
population growth (Hanan 1996). Injury during 
storms has been reported as a source of injury 
(Wilke 1943), and separations of mothers and pups 
during storms can be quite frequent (Boness et al. 
1992). Pup mortality may also increase during E] 
Nifio events (Koons 1998). Disease outbreaks, 
including phocine distemper virus and influenza 
(Geraci et al. 1982; Hinshaw et al. 1984; Osterhaus 
and Vedder 1988; Duignan 1995), have been 
responsible for large-scale die-offs in some areas 
(e.g., North Sea and New England). The die-off in 
the North Sea in 1988 resulted in a population 
reduction of over 50%. Such die-offs are clearly 
unpredictable, but given their magnitude and appar- 
ently increasing frequency of occurrence, they must 
be taken into account in conservation planning and 
population viability analyses (Young 1994; 
Simmonds and Mayer 1997). 


BAIRD: STATUS OF HARBOUR SEALS 


671 


Special Significance 

In some areas of Canada (e.g., British Columbia 
and the Bay of Fundy) Harbour Seals are important 
components of commercial wildlife viewing excur- 
sions, and thus contribute economically to tourism, 
although the level of economic input from such 
tourism has not been quantified. Since Harbour Seals 
co-inhabit many developed coastal regions in 
Canada, they are often frequently viewed by residents 
and may be given considerable aesthetic value as an 
important component of the natural environment. 
Conversely, due to actual or perceived conflicts with 
fisheries in many parts of country, Harbour Seals 
(and other pinnipeds) are viewed primarily as a pest. 
As a species which appears to show considerable 
site-fidelity, feeds relatively high on the food web, 
and lives for relatively long periods, they may also be 
viewed as a potential indicator species in the marine 
environment (Calambokidis et al. 1991). One popula- 
tion in Canada, at Sable Island, is unique, being the 
only truly offshore breeding population of this 
species in the world (Whitehead et al. 1998). 


Evaluation 

Based on the large population and evidence of 
increasing trends in numbers (Olesiuk et al. 1990a), 
the Pacific coast population of Harbour Seals should 
probably be classified as Not at Risk (NAR) by 
COSEWIC. However, recent evidence of contami- 
nant levels high enough to cause immunosuppression 
(Ross et al. 1998), and reports of high levels of ille- 
gal kills (H. Breen personal communication) both 
warrant further study, as does the potential impact of 
overfishing or natural fluctuations in prey popula- 
tions on Harbour Seal population dynamics. No one 
is doing research to establish the pesent status of 
Harbour Seals in the Canadian Arctic, including 
Hudson Bay and James Bay, because they have little 
economic importance (P. Richard, personal commu- 
nication). Yet, they are more vulnerable to overex- 
ploitation than other species because they live in 
small sedentary pockets of population (P. Richard, 
personal communication). 

For the same reason (ie., lack of economic impor- 
tance), little is known about the status of Harbour 
Seals off Labrador, Newfoundland, the outer Gulf of 
St. Lawrence, or off mainland Nova Scotia. The pop- 
ulation which has received the most research atten- 
tion off eastern Canada, at Sable Island, has drasti- 
cally declined in recent years, although the causes 
are unknown. Insufficient information is available to 
assess the status of eastern Canadian or Arctic popu- 
lations, and these populations should be classified as 
Indeterminate by COSEWIC. 


Acknowledgments 
I thank the Canadian Wildlife Service (Environ- 
ment Canada) for providing funds, and R.R. 


672 


Campbell (Chair, COSEWIC Fish and Marine 
Mammal Subcommittee) for assistance in the prepa- 
ration of this report. A number of individuals provid- 
ed unpublished information or personal communica- 
tions, including R. Bates, D. Bowen, H. Breen, D. 
Carins, J. Conway, P.-Y. Daoust, J. Lawson, J. Lien, 
R. Palm, D. Petrachenko, R. Remnant, P. Richard, 
G. Stenson, and W. Stobo. D. Bowen, P. Cottrell, S. 
Hooker, V. Lesage, Z. Lucas, M. Hammil, P. Ross, 
W. Stobo, and P. Watts provided constructive com- 
ments on the manuscript. 


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Accepted 19 February 2002 


Status of Killer Whales, Orcinus orca, in Canada* 


ROBIN W. BAIRD 


Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada. 
Current address: National Marine Fisheries Service, NOAA, 101 Pivers Island Road, Beaufort, North Carolina 28516 USA 


Baird, Robin W. 2001. Status of Killer Whales, Orcinus orca, in Canada. Canadian Field-Naturalist 115(4): 676-701. 


Killer Whales can be found in all three of Canada’s oceans, as well as occasionally in Hudson Bay and in the Gulf of St. 
Lawrence. Little is known about their occurrence or biology in the Atlantic or Arctic, but Killer Whales appear to be 
uncommon in most parts of these areas. In the Canadian Arctic and western Atlantic small numbers were killed historically 
in commercial whaling operations (or shot incidentally to such operations), and small numbers have been documented 
taken by natives. Predictable concentrations of killer whales are found in British Columbia, and populations in British 
Columbia’s nearshore waters are among the most well-known populations of cetaceans world-wide. Killer whales off the 
Pacific coast can be classified into two distinct “types” or “forms” (termed residents and transients), which differ in diet 
(residents feed on fish, transients feed on marine mammals), morphology, genetics and behaviour. The exact taxonomic 
relationship between these two types is unclear, though some authors have termed them “races”, others consider them sepa- 
rate species. Regardless, from both a scientific and management perspective these populations should be treated as distinct. 
Within British Columbia waters residents appears to be sub-divided into three geographic communities or populations 
(termed the “northern” and “southern” residents, and “offshore” killer whales), based on association patterns, genetics and 
morphology. Relatively little is known of the “offshore” population of Killer Whales. All populations (including transients 
and the three resident populations) are small (in the low hundreds), and have low potential rates of increase. No trend infor- 
mation is available for “offshore” or transient killer whales. The “northern” resident population has been growing steadily 
in size since the 1970s (when live-capture fisheries stopped and shooting declined), while the “southern” resident popula- 
tion has been growing only sporadically, and is currently smaller than the pre-live-capture population estimate from the 
1960s. Given the small population sizes and their low potential rates of growth, Killer Whales are potentially at risk from 
anthropogenic influences in two primary ways: due to immunotoxic affects of persistent toxic chemicals (levels in “south- 
ern” residents are three times higher than levels known to cause immunotoxicity in harbour seals), and due to a reduction 
in prey availability. It is also possible that the large and growing commercial and recreational whale watching industry on 
the west coast may be having an impact, though such impacts are as yet unclear. In terms of natural factors, periodic events 
such as mass strandings or entrapments in narrow inlets or ice have the potential to drastically reduce local populations. 
Since virtually all of these factors should impact Killer Whales throughout Canadian waters, all populations, at the least, 
should be considered vulnerable, that is, as “species of special concern because of characteristics which make them espe- 
cially sensitive to human activities or natural events”. As the “southern” resident population is extremely small (89 individ- 
uals in 1998), has declined by 10% in the last three years due to an increase in mortality rates (primarily of adult females), 
is more subject to anthropogenic influences than other populations, and these influences are not expected to decrease in the 
foreseeable future, it should be listed as threatened by COSEWIC. Further research, particularly on Arctic, Atlantic and 
“offshore” populations, is clearly needed. 


Key Words: Killer Whale, Orcinus orca, Epaulard, Canada, British Columbia, status, cetacean, sympatric populations. 


The Killer Whale or épaulard, Orcinus orca 
(Linnaeus 1758) is found in all three of Canada’s 
oceans (Figure 1). In the Pacific they are the most well- 
known cetacean both to the scientific community and 
to the general public. In fact, off the British Columbia 
coast long-term studies of Killer Whales have led to a 


*Reviewed and approved by COSEWIC, April 1999 — sta- 
tus assigned North Pacific “resident populations” 
Threatened, North Pacific “transient” populations Vul- 
nerable, North Atlantic and Arctic populations Indeter- 
minate. 

Status reviewed again by COSEWIC, November 2001 — sta- 
tus assigned “southern resident” population Endangered, 
“northern resident” population Threatened, NE Pacific off- 
shore population special concern, Northwest Atlantic 
(Eastern Arctic populations Data Deficient). 


greater understanding of these animals than of almost 
any other species of cetacean (Baird 2000). In the 
Canadian Arctic and Atlantic, Killer Whales are seen 
only occasionally and no in-depth scientific studies 
have been undertaken. Yet, because of their relatively 
large size, distinctive appearance, and the publicity this 
species has garnered in books, magazine articles, tele- 
vision, and in aquaria, Killer Whales are known by and 
recognizable by virtually everyone. In this report I 
review what is known of the general biology and ecolo- 
gy of Killer Whales in Canadian waters, including pop- 
ulation discrimination, sizes and trends, behaviour, life 
history and limiting factors. Gaps in the available data 
are identified that may be relevant to their long-term 
status assessment. This review has been undertaken on 
behalf of the Fish and Marine Mammal Subcommittee 
of COSEWIC, the Committee on the Status of 
Endangered Wildlife in Canada. 


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677 


2001 
ZIONS PEL SS 
4, uit IZ FE 
FIGURE 1. Map showing world-wide distribution of Killer Whales. Shaded areas indicate 
records of sightings or strandings; however unshaded areas may be part of the normal 
range with no sightings documented. Map courtesy of Marilyn Dahlheim, National 
Marine Mammal Laboratory, NMFS, Seattle. 
Description 


The distinctive black and white pattern, blunt 
head, and tall dorsal fin in the middle of the back, 
are the primary identifying characteristics of Killer 
Whales (Figure 2). Adult males are substantially 
larger than females, although very few accurate mea- 
surements are available. Reports of maximum 
lengths of 9.75 m for males and 8.53 m for females 
given in the literature (e.g., Perrin and Reilly 1984) 
are actually estimates. The maximum length mea- 
sured for males and females is 9.0 m and 7.7 m, 
respectively (Heyning and Brownell 1990). There is 
some suggestion of differences in size for individu- 
als from different populations (cf. Berzin and 
Vladimirov 1983; Heyning and Brownell 1990), and 
more accurate measurements from different parts of 
their range are needed before average lengths of 
individuals from any one population can be charac- 


FiGuRE 2. An adult female transient Killer Whale porpois- 
ing off Victoria, British Columbia, showing the 
main features for identification; the striking black 
and white colouration, a blunt head and a tall, cen- 
trally-placed dorsal fin. Photo© by the author. 


terized. From the few measurements available for 
adult individuals from British Columbia (e.g., Bigg 
and Wolman 1975), it is clear that the average length 
of adult individuals is much smaller than the maxima 
noted above. Adults are sexual dimorphic in 
appendage size, with adult males having a tall trian- 
gular dorsal fin which may reach up to 1.8 m in 
height, while in juvenile males and adult females it 
reaches 0.9 m or less and is generally more falcate 
(Figure 3). Pectoral fins and tail flukes are also sexu- 
ally dimorphic, being much larger in adult males, 
with the fluke tips also bending downwards. As well, 
pigmentation in the genital area differs between 
males and females (Bigg et al. 1987). 


Population Discrimination 

The question of population segregation or division 
(fragmentation) is critical to any evaluation of status 
(IUCN 1996). If more than one distinct population 
exists, and factors which affect each population dif- 
fer in any way, then each population must be moni- 
tored and managed independently. 


“Residents” and “Transients” 

In the case of Killer Whales in the Canadian 
Arctic and North Atlantic, no information is avail- 
able to assess whether any population differentiation 
has occurred (Mitchell and Reeves 1988; Anony- 
mous 1993). For the Pacific coast of Canada, clear 
evidence is available for differentiation of Killer 
Whales into two distinct “types” or “forms”, termed 
“resident” and “transient” (Table 1; Figure 4; Bigg et 
al. 1976). The names “resident” and “transient” have 
become entrenched in the literature even though it 
has been demonstrated that they are not accurate as 
descriptions of the site fidelity and movement pat- 
terns of the two forms (Guinet 1990; Baird et al. 
1992). As the names are frequently mis-interpreted 
as descriptive categorizations, they are referred to 


678 


SS... 
RRR 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


FIGURE 3. Photograph of two transient Killer Whales off Victoria, British Columbia, showing the clear sexual-size dimor- 
phism, with the male (right) having a tall, straight dorsal fin and the female (left) having a shorter, falcate, dorsal 


fin. Photo© by the author. 


hereafter as resident and transient to try to prevent 
such confusion. 

Several studies have documented a variety of 
behavioural, ecological, morphological and genetic 
differences between transients and residents (Table 
1; Bigg et al. 1987; Baird and Stacey 1988; Morton 
1990; Baird et al. 1992; Baird and Dill 1995, 1996; 
Barrett-Lennard et al. 1996; Hoelzel et al. 1998; 
Matkin et al. 1998; Ford et al. 1998). One of the 
most important differences is diet; residents appear 


to feed almost entirely on fish, while transients 
appear to feed almost entirely on marine mammals 
(for more detailed discussion, see Feeding Habits, 
below). Association patterns, in terms of observa- 
tions of individuals traveling together in a group, and 
vocal dialects, are also used to discriminate residents 
from transients (Black et al. 1997). Interactions 
between residents and transients have only been 
reported on a small number of occasions (Jacobsen 
1990; Morton 1990; Barrett-Lennard 1992; Baird 


TABLE 1. Characteristics which differ between resident and transient-type Killer Whales in the nearshore waters of the 


eastern North Pacific. 


MORPHOLOGY/GENETICS 
Shape of the dorsal fin (Bigg et al. 1987; Bain 1989) 
Saddle patch pigmentation (Baird and Stacey 1988) 


Possibly eye patch pigmentation (D. Ellifrit, personal communication, cited in Baird 1994) 
Mitochondrial and nuclear DNA (Stevens et al. 1989; Hoelzel and Dover 1991; Hoelzel et al. 1998; Matkin et al. 1998) 


BEHA VIOUR/ECOLOGY 


Diet (Bigg et al. 1987; Morton 1990; Baird and Dill 1996; Ford et al. 1998) 
Travel patterns/habitat use (Heimlich-Boran 1988; Morton 1990; Baird and Dill 1995) 


Respiration patterns (Morton 1990) 


Vocalizations (Ford and Hubbard-Morton 1990; Morton 1990) 


Echolocation (Barrett-Lennard et al. 1996) 
Amplitude of exhalations (Baird et al. 1992; Baird 1994) 
Possibly diving patterns (Baird 1994) 


Group size (Bigg et al. 1987; Morton 1990; Baird and Dill 1996) 
Pattern and extent of natal philopatry (Bigg et al. 1987; Baird and Dill 1996; Baird and Whitehead 2000) 
Seasonal occurrence (Guinet 1990; Morton 1990; Baird and Dill 1995) 


Geographic range (Bigg et al. 1987) 


2001 


BAIRD: STATUS OF KILLER WHALES 


679 


FicurE 4. Morphological differences between residents (left) and transients (right) include differences in dorsal fin shape 
(with residents typically having more rounded fins) and saddle patch pigmentation patterns (with transients typical- 
ly having less complex patterns). Overlap in the characteristics exist, so they cannot always be used to distinguish 
type. Photos© by the author. 


and Dill 1995). On eight of those occasions no 
change was recorded in the behaviour of either form 
as they passed within a couple of kilometers of each 
other. Transients have been seen changing direction 
away from residents (avoiding them) on eight occa- 
sions, residents avoiding transients twice, and both 
avoiding each other twice. Since residents vocalize 
more frequently than transients (Morton 1990; 
Barrett-Lennard et al. 1996), transients may detect 
the presence of residents much sooner, and more fre- 
quently than the other way around (Baird and Dill 
1995). Cases of residents showing no reaction when 
near transients may be due simply to them being 
unaware that transients were nearby. One observa- 
tions of aggression between the two forms involved 
a large group of residents attacking a small group of 
transients (Ford and Ellis 1999), and one other 
observation of apparent aggression has also been 
observed (P. Spong, H. Symonds, personal commu- 
nications). 

The exact taxonomic relationship of these two 
forms is unclear. Bigg et al. (1987) termed these two 
types of Killer Whales “races”, and this term has 
been adopted, uncritically, by many investigators. 
“Races” are usually defined in a geographic sense, 
implying geographically isolated populations which 
are typically given subspecific designation (Mayr 
and Ashlock 1991). Baird et al. (1992) outlined how 
these two forms may have evolved, and termed them 
incipient species. Baird (1994) subsequently argued 
that they should be considered separate species, 
although no formal description of each species has 


been presented. Heyning and Dahlheim (1993) have 
argued that insufficient information is available to 
determine the level of isolation between them. 
Hoelzel (personal communication) estimated genetic 
migration between these two forms at one male per 
five generations and one female per 20 generations 
(see Hoelzel et al. 1998). 

Regardless of such disagreements and uncertainty 
in taxonomic relationship between these two forms, 
there is sufficient evidence to suggest that these 
forms or types should be treated as separate popula- 
tions for management. Considering the differences in 
behaviour and ecology which have been documented 
(e.g., Table 1), it is also prudent not to apply 
behavioural or life history characters from one form 
to another, nor indeed from these populations of 
Killer Whales to Killer Whales elsewhere. 

The suggestion that there may be more than one 
species in the genus Orcinus is not new. Mikhalev et 
al. (1981) and Berzin and Vladimirov (1983) 
described two species in the Southern Ocean, O. 
nanus and O. glacialis, respectively, both of which 
seem to refer to the same population of smaller indi- 
viduals (Heyning and Dahlheim 1988). As well as 
differences in body size, other differences in mor- 
phology, behaviour and diet were noted, with one 
species feeding primarily on fish and the other feed- 
ing primarily on marine mammals, similar to the sit- 
uation off Canada’s west coast (Berzin and 
Vladimirov 1983). Neither of these species designa- 
tions have been generally accepted (Perrin 1982; 
Heyning and Dahlheim 1988). 


680 


“Southern”, “Northern” and “Offshore” Residents 

A further level of population differentiation 
appears to exist within the resident form. Based on 
association patterns, pigmentation patterns and genet- 
ics, residents within British Columbia appear to be 
divided into three distinct, largely geographically 
based communities or populations (Bigg et al. 1987; 
Baird and Stacey 1988; Bain 1989; Ford et al. 1994a; 
Hoelzel et al. 1998; Matkin et al. 1998). One popula- 
tion, found generally around southern Vancouver 
Island and in Washington state, has been termed the 
“southern” resident community, one found generally 
off northern Vancouver Island and in southeast 
Alaska has been termed the “northern” resident com- 
munity, and a third putative resident population is 
termed “offshore” Killer Whales, which appear to 
inhabit offshore waters along the entire coast. It 
should be noted that while based on mitochondrial 
DNA “offshore” Killer Whales are closely related to 
northern and southern residents (Hoelzel et al. 1998; 
Matkin et al. 1998), relatively little is known about 
other aspects of their biology, and it is unclear 
whether “offshore” Killer Whales share behavioural 
or ecological characteristics with northern or southern 
residents. British Columbia northern residents have 
been observed associating, and share the same mito- 
chondrial DNA haplotype, with other resident-type 
Killer Whales in southeast Alaska (Dahlheim et al. 
1997; Hoelzel et al. 1998). The resident-type whales 
from Alaska have not been documented in British 
Columbia, but based on both association patterns and 
genetics are likely part of the same population. These 
southeastern Alaska residents have in turn been 
observed interacting with residents in Prince William 
Sound, Alaska (Matkin et al. 1997), suggesting that 
gene-flow between northern residents and these other 
whales may exist. Similarly, both “offshore” Killer 
Whales and transients documented in British 
Columbia have also been seen off Alaska, 
Washington and/or California (Dahlheim et al. 1997; 
Black et al. 1997), suggesting that these individuals 
are part of larger populations. The U.S. National 
Marine Fisheries Service evaluates each of the resi- 
dent populations, and the transient population, inde- 
pendently (Barlow et al. 1997; Hill et al. 1997). 

The northern and southern resident communities 
have been reported to have ranges which do not 
overlap (e.g. Bigg et al. 1990; Felleman et al. 1991), 
but there are data which indicate their ranges overlap 
by over 120 km on both the east and west coasts of 
Vancouver Island (Bigg et al. 1976; M. A. Bigg, per- 
sonal communication 1990; Ford et al. 1994a). 
“Offshore” Killer Whales similarly overlap in range 
with both northern and southern residents, though 
observations of “offshore” Killer Whales in or near 
the core areas of the other two groups are rare (e.g., 
‘Walters et al. 1992; Ford et al. 1994b*). Regardless, 
behavioural interactions have not been observed 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


between individuals from northern, southern and 
“offshore” resident communities, and differences in 
mitochondrial DNA and physical appearance suggest 
the communities are reproductively isolated (Baird 
and Stacey 1988; Stevens et al. 1989; Hoelzel and 
Dover 1991; Walters et al. 1992; Ford et al. 1994a:; 
Hoelzel et al. 1998; Matkin et al. 1998). The north- 
ern and southern resident communities also appears 
to have distinct behavioural characteristics (Osborne 
1986; Hoyt 1990); whether “offshore” Killer Whales 
exhibit such distinctive behavioural characteristics is 
unknown, simply due to the relative paucity of work 
that has been undertaken on that population. Regions 
identified as high use areas (“core areas” ) for north- 
ern and southern residents are separated by about 
400 km (two and a half days of travel at 3.5 knots — 
Bigg 1982). 


Distribution and Movements 

Killer Whales are cosmopolitan, having been 
observed in all oceans of the world (Leatherwood 
and Dahlheim 1978; Dahlheim and Heyning 1998). 
However, concentrations generally occur in colder 
regions and in areas of high productivity (Bigg et al. 
1987; Heyning and Dahiheim 1988; Guinet and 
Jouventin 1990). In polar areas the occurrence of 
Killer Whales is thought to be limited by the pres- 
ence of pack ice in winter months (Reeves and 
Mitchell 1988a), thus some north-south movements 
would have to occur in such areas. A recent sighting 
by Gill and Thiele (1997) of Killer Whales deep in 
Antarctic sea ice in winter indicates that not all indi- 
viduals move away from the poles. Gill and Thiele 
(1997) suggest that the extreme seasonal differences 
in the number of observers in polar regions could be 
party responsible for the perception that Killer 
Whales do migrate. In general, no clear evidence of 
seasonal north-south migrations is available. In the 
southern hemisphere, based on sightings from whal- 
ing vessels, Mikhalev et al. (1981) described season- 
al migrations from low-latitude areas in the winter 
months to higher latitude areas in summer. However, 
no information was presented on potential seasonal 
biases in effort, so it is difficult to judge the validity 
of such conclusions (Perrin 1982). 

Within British Columbia, Killer Whales have been 
documented throughout virtually all salt-water (and 
some-fresh water) regions, including many long 
inlets, narrow channels and deep embayments. Both 
resident and transient Killer Whales have been 
recorded year-round in British Columbia. Presence of 
resident Killer Whales seems to be closely tied with 
peak abundance of various species of salmon, one of 
their primary prey (Heimlich-Boran 1986; Bigg et al. 
1987; Nichol and Shackleton 1996). Several authors 
have suggested that residents are rare in the core 
study areas of Johnstone Strait and Haro Strait during 
winter months. However, in both areas one pod (A5 


2001 


in Johnstone Strait, J1 in Haro Strait) is recorded dur- 
ing most winter months (D. Ellifrit, P. Spong, H. 
Symonds, personal communications). Broad scale 
shifts in distribution are apparent, though they are 
more conclusive for northern residents than southern 
residents, since there are two year-round land-based 
research projects being undertaken in or near the core 
area for northern residents (Morton 1990; P. Spong, 
H. Symonds, personal communications). There are 
several seasonal biases in effort which should be 
taken into account in terms of the seasonal distribu- 
tion of southern residents, and the seasonal distribu- 
tion of northern residents outside of the core area of 
Johnstone Strait. Inclement weather conditions and 
low daylight hours during winter months likely 
decrease the probability of visually detecting Killer 
Whales when they are present, and little winter work 
has ever been undertaken. Some evidence is available 
to suggest that northern residents decrease their fre- 
quency of vocalizing during winter months (Bain per- 
sonal communication); this may confound examina- 
tions of winter occurrence using this method. 
Similarly, southern residents appear to travel further 
from shore during winter months (Baird unpublished; 
D. Ellifrit, personal communication), biasing detec- 
tion based on shore-based observations. As such, 
more thorough examinations of seasonal movements 
(perhaps using satellite telemetry) and winter habitat 
use are warranted. 

Seasonal influxes of Killer Whales into near-shore 
areas where pinnipeds are abundant have been noted 
at Marion Island, the Crozet Archipelago, and Punta 
Norte, Argentina (Condy et al. 1978; Guinet 1992; 
Hoelzel 1991). Baird and Dill (1995) showed that a 
strong seasonal peak in occurrence of transient Killer 
Whales in southern British Columbia coincided with 
the period when harbour seal pups were being 
weaned. However, only some pods appeared to pref- 
erentially use the area during that time, while others 
were seen regularly year-round (Baird and Dill 1995). 
Those pods which used the area year-round also tend- 
ed to travel further from shore, where land-based 
observers or spotters were less likely to detect them. 
Because of this seasonal difference in use of near- 
shore areas, Baird and Dill (1995) and Baird (1995a) 
suggested that many studies which are shore-based 
may be biased when examining seasonal presence. 

Killer Whales have been documented moving 
long distances, with some individual transients and 
“offshore” Killer Whales identified both in central 
California and southeastern Alaska, a 2660 km one- 
way distance (Goley and Straley 1994; Black et al. 
1997). Actual home range sizes are unknown, since 
virtually no photo-identification work has been done 
in offshore areas (though see Black et al. 1997), and 
no animals have been satellite-tagged. Using the 
northern- and southern-most sightings of particular 
individuals, combined with the limited knowledge of 
onshore-offshore movements, the largest docu- 


BAIRD: STATUS OF KILLER WHALES 


68 | 


mented range for a transient in British Columbia is 
140 000 km2, while the largest documented range for 
a resident is approximately 90 000 km2 (Baird 
2000). Both residents and transients have been docu- 
mented to move up to 160 km in one 24 hour period, 
but pods of both types also spend extended periods 
in small areas. 

A comprehensive review of all records available 
for Killer Whales in the eastern Canadian Arctic and 
the western North Atlantic was last undertaken in the 
1980s (see papers in Sigurjonsson and Leatherwood 
1988). Sergeant and Fisher (1957) stated that Killer 
Whales migrated northwards in the spring along the 
coasts of Labrador and Newfoundland, though a more 
comprehensive review by Mitchell and Reeves (1988) 
concluded that biases in effort precluded the determi- 
nation of any obvious pattern of distribution or move- 
ments. Killer Whales are occasionally recorded in vir- 
tually all areas off eastern Canada, including Nova 
Scotia (Katona et al. 1988), in the Gulf of St. 
Lawrence (Wenzel and Sears 1988), off Newfound- 
land and Labrador (Lien et al. 1988), and in Hudson 
Bay and the Canadian Arctic (Reeves and Mitchell 
1988a), with one record from 81°N. Records from 
these compilations end in the early 1980s, thus anoth- 
er review incorporating more recent records is war- 
ranted. Based largely on records collected since the 
earlier review, it appears that there are only a couple 
of areas where Killer Whales appear to be somewhat 
regular in their occurrence. These include the Mingan 
Islands, Quebec, where R. Sears (Mingan Island 
Cetacean Study, personal communication) has 
observed the same small group of whales a number of 
times since 1984 (see Wenzel and Sears 1988), the 
western end of the Strait of Belle Isle (R. Sears, per- 
sonal communication), off Battle Harbour, Labrador 
(S. Todd, College of the Atlantic, personal communi- 
cation), and around Pond Inlet, Cumberland Sound, 
and the Lancaster Sound region, where regular, and 
possible annual visitation has been noted (Reeves and 
Mitchell 1988a). In the western Canadian Arctic, 
some published distribution maps show the presence 
of Killer Whales (Jefferson et al. 1991; Dahlheim and 
Heyning 1998; Figure 1) in the Canadian side of the 
Beaufort Sea. According to two sources (T. Barry, L. 
Harwood, personal communications), native elders 
recall sightings of Killer Whales in the area in the 
1940s or 1950s, however numerous researchers who 
have undertaken surveys there in the last thirty years 
have never seen this species (M. Fraker, L. Harwood, 
D. Lyungblad, S. Moore, W. J. Richardson, personal 
communications). Any Killer Whales which do travel 
into the Canadian Beaufort Sea are likely part of the 
Bering Sea population (Dahlheim 1997). 


Protection 

Two factors are important in the legal protection 
of a species, the system that is in place to prohibit or 
regulate hunts or other threats, and the system for 


682 


monitoring and enforcing regulations. Where infor- 
mation is available, each of these is discussed 
below. 


International 

Two international management measures/agencies 
are relevant to the protection of Killer Whales, 
CITES (the Convention on International Trade in 
Endangered Species of Wild Fauna and Flora in 
1973) and the IWC (International Whaling Com- 
mission). 

All species of cetaceans are listed by CITES 
under one of two appendices. Appendix I includes 
species threatened with extinction (and which may 
be affected by trade), while Appendix II includes 
species which may become threatened with extinc- 
tion unless trade is regulated, as well as species 
which must be subject to regulation in order that 
trade in threatened species of similar appearance 
may be controlled (Klinowska 1991). Killer Whales 
(and all species of cetaceans not listed under 
Appendix I) are listed under Appendix II for the lat- 
ter of the above reasons. As such, international trade 
of Killer Whales or parts thereof by any countries 
which are Parties to CITES requires export permits 
from the country of origin. According to Klinowska 
(1991) the European Community treats all cetaceans 
as if they were listed in CITES Appendix I — thus 
trade requires permits from both exporting and 
importing countries and such trade must not be pri- 
marily for commercial purposes. Some other coun- 
tries (e.g., USA) also have similar domestic rules, 
requiring both export and import permits for 
Appendix II species. As of October 1998 there were 
144 Parties to CITES, leaving approximately 90 
countries world-wide which were not members 
(CITES Secretariat statistics). This latter group 
includes Iceland, which has been actively involved 
in trade (see Limiting Factors below). Listing on 
CITES Appendix II does not provide protection per 
se, though it does mandate recording of international 
trade. In recent years, the only international trade of 
Killer Whales documented through CITES has been 
in small numbers of live animals between aquaria, a 
few scientific samples, and small numbers of teeth 
and carvings. Trade in teeth and carvings have pri- 
marily involved the transfer of these items between 
Greenland (a dependency of Denmark) and Denmark 
(CITES Secretariat statistics). Although all trade in 
Appendix II species from a CITES member should 
be documented, during a recent review of all Killer 
Whales kept in captivity (Hoyt 1992), E. Hoyt (per- 
sonal communication) noted that some trade involv- 
ing CITES countries had not been reported. 

Killer Whales are considered “small cetaceans” 
by the IWC, and there is currently considerable dis- 
agreement within the Commission as to whether 
small cetaceans are covered by the Convention. 
However, in 1980, in response to a large Russian 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


take of Killer Whales in the Antarctic in the 1979/80 
season, the [WC added a new sentence to Schedule 
paragraph 9(d), officially including Killer Whales in 
their moratorium on factory ship whaling (IWC 
1981). Other IWC management measures (e.g., the 
Southern Ocean Sanctuary, moratorium on commer- 
cial whaling, etc) do not apply to Killer Whales. 


National 

Canada: Within Canada, management of Killer 
Whales has varied considerably over time, and both 
the federal government and one provincial govern- 
ment (British Columbia) have been involved in man- 
agement activities. Prior to 1970 no laws were in 
place to control or regulate captures or other interac- 
tions. Hoyt (1992) notes that news reports of deaths 
during captures and the out-of-country destinations 
of captured Killer Whales in the 1960s prompted 
wide-spread public pressure for the implementation 
of protective legislation. Such legislation was first 
introduced in 1970. Prior to 1982, Killer Whales 
were considered “wildlife” by the British Columbia 
provincial government’s Wildlife Branch, and pos- 
session permits could be issued for holding these 
animals in captivity. In 1982 the provincial Wildlife 
Branch re-wrote the “Wildlife Act’, and deleted 
Killer Whales from the list of wildlife, in response to 
a federal move to include all cetaceans under the 
“Cetacean Protection Regulations” (under the 
Fisheries Act of Canada of 1867). These regulations 
prohibited “hunting” without a license. “Hunting” 
was defined as “to chase, shoot at, harpoon, take, 
kill, attempt to take or kill, or to harass cetaceans in 
any manner’. No scheme, however, was in place to 
enforce such regulations, and aboriginal hunting 
could be undertaken without a license. In 1993, the 
federal government consolidated various marine 
mammal regulations, including the Cetacean 
Protection Regulations, under the new “Marine 
Mammal Regulations”. These regulations stated that 
“no person should disturb a marine mammal except 
when under.... the authorities of these regulations”, 
with “marine mammal” defined as all species listed 
under a particular appendix. However, many species 
of cetaceans, including Killer Whales, were not list- 
ed under that appendix, and thus no legal protection 
appears to have been in place. The definition of 
“marine mammal” was revoked in 1994, thus 
extending coverage to all species of marine mam- 
mals. Currently, hunting of Killer Whales can occur 
if a “Fishing License” is obtained (except for Abor- 
iginals who can hunt without a license), but fees for 
such licenses are low ($5). However, no such licens- 
es have been issued, and issuance is at the discretion 
of the federal Minister of Fisheries and Oceans. It is 
unlikely any would be issued in areas such as British 
Columbia, due to widespread public interest in these 
animals. 

In terms of minimizing negative interactions 


2001 


between boats and Killer Whales, “whale watching 
guidelines” have been produced and disseminated 
by the Department of Fisheries and Oceans. There 
are also several ongoing efforts of self-regulation by 
the commercial whale watching industry in British 
Columbia, involving the production of guidelines 
and codes of conduct (Baird et al. 1998b). Among 
commercial operations in certain specific areas, the 
levels of awareness of and adherence to these guide- 
lines is fairly high, though awareness and adherence 
by general members of the public (which make up 
the majority of boats with whales in some areas — 
see Figure 5) is currently unknown. As with the 
Cetacean Protection Regulations, virtually no offi- 
cial monitoring or enforcement activities take place, 
and enforcement itself is complicated by the diffi- 
culty in defining and measuring “harassment” in the 
field (see Limiting Factors, below). N. Bhaloo 
(DFO Conservation and Protection, Enforcement 
Unit, personal communication) notes that no viola- 
tions of the Marine Mammal Regulations involving 
Killer Whales have been documented between 1993 
and 1997, although there is one charge of harass- 
ment, involving a sports fishing operation outside of 
either of the two core areas for residents, pending 


Nm jw 
on — 


Nm 
— 


BOATS IN AREA 
Ss 


(WHALES PRESENT) 
an 


m 
0 
4 
W 
a 
2 
a 
Zz 
Ww 
0 
< 
x 
> 
< 


1990 1991 1992 1993 1994 1995 1996 1997 


YEAR 
CCOMMERCIAL BOATS [NON-COMMERCIAL BOATS 


FiGureE 5. Trend in the average number of boats (both com- 
mercial whale watching operations and total boats) 
with southern resident Killer Whales as they pass 
the Lime Kiln Lighthouse on San Juan Island, 
Washington state, from 1990 through 1997. Data 
were collected from mid-May through mid-August 
each year, seven days per week, from 0900 till 1700 
h. No such increasing trend was apparent when no 
whales were present. Data from R. Otis (see Baird et 
al. 1998b). 


BAIRD: STATUS OF KILLER WHALES 


683 


from 1998 (E. Lochbaum, DFO, personal communi- 
cation). 

The 1997 Oceans Act provides for the establish- 
ment of marine protected areas (MPAs) in federal 
waters. One of the specific justifications listed for 
establishing MPAs is to conserve and protect marine 
mammals and their habitats. However, as with other 
federal legislation regarding marine mammals, estab- 
lishment of marine protected areas and exclusion of 
activities which might jeopardize Killer Whales or 
other marine mammals are up to the discretion of the 
Minister of Fisheries and Oceans, rather than man- 
dated. Regardless, there are general concerns about 
the efficacy of using MPAs to “protect” cetaceans 
(see below, as well as Phillips 1996; Whitehead et al. 
2000), due primarily to the large range of most 
species and the lack of boundaries in the marine 
environment. Whitehead et al. (2000) note that most 
marine protected areas have provided little or no 
change in the level of threats faced by cetaceans in 
an area. 

One example of an MPA specifically created to 
“protect” Killer Whales is the Robson Bight/Michael 
Bigg Ecological Reserve, a provincial designation in 
a core area for northern residents. This designation 
provides some protection to the shoreline habitat and 
limits human access by land. Its main relevance to 
Killer Whales is the protection of the terrestrial por- 
tion of several “rubbing beaches” which are regular- 
ly used by northern residents. However, its validity 
as a “whale sanctuary” has been questioned. Duffus 
and Dearden (1992) state that this designation “holds 
a fairly limited potential to protect a marine area”, 
since it is the federal government that has jurisdic- 
tion over marine shipping and marine fisheries, and 
this is a provincial designation. They also note that 
the “boundary is highly permeable, and buffers of 
outside impacts are almost non-existent” (Duffus and 
Dearden 1992). They warn against the “fallacy of 
tokenism — that is, giving the public the appearance 
of protecting an important whale habitat, when nei- 
ther the importance of the site to the whales, nor the 
veracity of the protection is established — creat[ing] 
a political “success” that may mask an ecological 
failure” (Duffus and Dearden 1992). Other than this 
effort by the British Columbia provincial govern- 
ment, no other province or territory within Canada 
has legislated protection for this species. 

Other Countries: Considering that Killer Whales 
regularly move between Canada and other countries 
(the U.S. on both coasts and almost certainly Green- 
land, see Heide-Jorgensen 1988, 1993; Mitchell and 
Reeves 1988), protection measures in these countries 
are directly relevant to the conservation of Killer 
Whales in Canada. In the United States, all cetaceans 
are protected through the Marine Mammal 
Protection Act of 1972, as well as through the 
Packwood-Magnuson Amendment of the Fisheries 
and Conservation Act and the Pelly Amendment of 


684 


the Fisherman’s Protective Act. These regulations 
allow for observers on fisheries that have a high 
probability of killing marine mammals, and also pro- 
vide for limited monitoring and enforcement activi- 
ties regarding boat/whale interactions. There are no 
management measures in Greenland that provide 
protection for Killer Whales, and kills in that area 
(see Limiting Factors, below) could affect Canadian 
populations. 


Population Sizes and Trends 

No world-wide population estimates are available. 
Regional population estimates, where available, have 
been derived through photo-identification surveys 
and/or line-transect surveys. Line-transect surveys 
have generally been used in areas where more inten- 
sive photo-identification studies are impractical (e.g., 
the Southern Ocean), but are also accompanied by 
large confidence limits (see e.g., Matkin and Saulitis 
1994). Furthermore line-transect surveys do not 
allow for discrimination of individuals from sym- 
patric populations. 

In British Columbia, four separate populations 
must be considered, transients, “northern” and 
“southern” residents, and “offshore” Killer Whales 
(which should probably be considered “offshore” 
residents, see discussion above). The most detailed 
information is available for northern and southern 
residents, as all of the southern resident pods and 
many of the northern resident pods are censused 
each year. As all individuals are recognizable, the 
census provides an actual count of the number of 
individuals in the population. As of 1998, the south- 
ern resident population numbered 89 individuals 
(van Ginneken and Ellifrit 1998). While the popula- 
tion has grown since the cessation of the live-capture 
fishery in 1973, the current population is smaller 
than the population near the start of that fishery 
(Figure 6), and has declined for the last three years 
(1996-1998). Such a decline is not unprecedented 
(Olesiuk et al. 1990; Figure 6); since the cessation of 
the live-capture fishery the population showed a sim- 
ilar decline from 1980 through 1984. As discussed 
under Limiting Factors (below), this earlier decline 
was likely due in part to the removal of animals in 
the live-capture fishery (Olesiuk et al. 1990). The 
most recent decline appears to have resulted from an 
increased death rate. Using data presented by van 
Ginneken and Ellifrit (1998), the average per capita 
death rate between the years 1995 to 1998 (mean of 
0.052) is significantly higher (Mann-Whitney U-test, 
p = 0.0084) than the average for the preceding 19 
years (mean of 0.021 from 1976 through 1994). The 
per capita birth rate for this same period (mean of 
0.034) is similar to the average for the previous 19 
years (mean of 0.038). In the period from 1995 to 
1998, age-specific mortality rates for mature females 
between 35 and 65 years of age are four to five times 


THE CANADIAN FIELD-NATURALIST 


Volkas 


higher than reported by Olesiuk et al. (1990; see 
Table 2). It is unclear however whether this current 
population decline may be due to demographic 
stochasticity, or even perhaps delayed effects of the 
removal of animals in the live-capture fishery, and a 
study modeling the probability of such effects occur- 
ring by chance or due to selective removals is war- 
ranted. 

As of 1993 the number of northern residents 
thought to occupy British Columbia waters (if only 
seasonally) numbered approximately 200 individuals 
(Ford et al. 1994a). More recent surveys have been 
undertaken, but the Department of Fisheries and 
Oceans, Pacific Region, which undertakes these sur- 
veys, has not released current data. This “popula- 
tion” has been growing at a relatively stable rate 
since the 1960s (Oleisuk et al. 1990; Ford et al. 
1994a). Because of the associations and shared 
mtDNA haplotypes with residents in Alaska, the 
effective population size for northern residents 
should probably be considered larger than the abso- 
lute number recorded within British Columbia 
waters, and the overall trend in the larger population 
is unknown (since less complete information is 
available for Alaskan residents). Population esti- 
mates for several regions of Alaska are available and 
are summarized by Matkin and Saulitis (1994). 
Taking into account only those whales documented 
in British Columbia, there was no evidence of densi- 
ty-dependent effects as of the late 1980s (Meyers 
1990), but data collected since then have not yet 
been examined. Brault and Caswell (1993) examined 
pod-specific demography of residents, and conclud- 
ed that most of the variance in individual pod growth 
rates was due to variance in adult reproductive out- 
put, rather than effects of pod size or structure. 

Population size is not known for “offshore” Killer 
Whales. The first “offshore” Killer Whales groups 
were encountered in the mid-1980s, and about 200 
“offshore” Killer Whales had been documented as of 
1993 (Ford et al. 1994a). These whales have been 
identified over a relatively short period of time, thus 
while natural mortality should only have accounted 
for a few deaths, new individuals are being regularly 
documented (Walters et al. 1992; Ford et al. 1994a). 
Of 56 “offshore”-type whales documented off 
California, 23 were direct matches with “offshore” 
Killer Whales recorded off Oregon, Washington, 
British Columbia and southeast Alaska (Black et al. 
1997). No trend information is available for this pop- 
ulation. 

For transient Killer Whales, the total population 
size is unknown but probably numbers in the low 
hundreds. Seventy-nine transients had been photo- 
identified in British Columbia and Washington up to 
1986 (Bigg et al. 1987), and a further 90 or more 
have been documented in the 10 years since (Ford et 
al. 1994a). Considering their wide-ranging move- 


2001 


BAIRD: STATUS OF KILLER WHALES 


685 


FIGURE 6. Population numbers and trends for southern resident Killer Whales. Data in this graph are from van 
Ginneken and Ellifrit (1998) and Olesiuk et al. (1990). Where differences exist between the two sources I 
have followed van Ginneken and Ellifrit (1998). The left arrow marks the beginning of large catches of 
whales from this population in the live capture fishery (small numbers had been taken prior to that point), 
while the right arrow marks the cessation of that fishery. Data from 1975 on are counts of animals using 
photo-identification; prior to that point numbers given are projections from a matrix model (Olesiuk et al. 
1990). While there has been an increase since the end of live-captures, readers should compare the unsteady 
growth of this population since 1973, with the relatively steady continual increase of the northern resident 


population (Olesiuk et al. 1990; Figure 26). 


ments, known associations, and shared genetic hap- 
lotypes, transients from bordering areas should be 
considered part of the same population which uses 
B.C. waters (Black et al. 1997). Of 79 transients 
documented in southeast Alaska, 69 have been 
observed in British Columbia (Dahlheim et al. 1997; 
Ford and Ellis 1999). One hundred and five tran- 
sients have been documented off California, and at 
least 10 of those have been documented in British 
Columbia, Washington, or further north in Alaska 
(Black et al. 1997). New individuals are occasionally 
being recorded in some areas (e.g., van Ginneken et 
al. 1998). Because of the long-resighting interval for 
some transients, it is not possible to determine 


deaths in the same way as for residents, thus some of 
the whales already documented are probably no 
longer living. The use of mark-recapture models for 
estimating transient population size is not appropri- 
ate, as the probability of encountering transients in 
any particular area differs between groups (see Baird 
and Dill 1995, and differences between regional cat- 
alogues, e.g., Black et al. 1997; Palm 1997; van 
Ginneken et al. 1998). Population trend information 
is unavailable. 

No population estimates are available for the 
Canadian Arctic or Atlantic waters, though compila- 
tions of records have been presented by Lien et al. 
(1988), Mitchell and Reeves (1988), Reeves and 


TABLE 2. Comparison of age-specific mortality rates for all mature female southern residents from 1995-1998 (data from 
van Ginneken and Ellifrit 1998) with values presented by Olseiuk et al. (1990). 


Age group Olesiuk et al. 1990 residents 1995-1998 southern residents 
15.5-24.5 0.0000 0.0 
25.5-34.5 0.0036 0.0 
35.5-44.5 0.0109 0.05 
45.5-54.5 0.0250 0.125 
55.5-64.5 0.0328 0.14 
>65 0.069 0.05 


Ee ee ee 


Note: While sample sizes are small, and this is a post-hoc comparison looking only at a four-year period where a decline 
in the population has been noted, mortality rates for mature females between 35 and 65 years of age are four to five times 


higher than the rates calculated by Olesiuk et al. (1990). 


686 


Mitchell (1988a) and Wenzel and Sears (1988). 
Some anecdotal evidence suggests that the number 
of whales which utilize the St. Lawrence has de- 
clined in the last 60 years; Vladykov (1944) reports 
that large numbers of Killer Whales (including 
groups up to 40 individuals) were found in the area 
in spring and fall, feeding on belugas. It is clear that 
numbers which utilize the area today are much 
smaller (Mitchell and Reeves 1988; Wenzel and 
Sears 1988). The largest group reported off eastern 
and Arctic Canada in the last 20 years appears to be 
of 22 individuals (Finley 1990). Mitchell and Reeves 
(1988) note that Killer Whales appear to be “uncom- 
mon in the western North Atlantic relative to other 
medium-sized and large cetaceans, and that they may 
be numerically few” (this view is supported by data 
presented by Lien et al. 1988). As noted above, a 
review of western North Atlantic records subsequent 
to those mentioned above is warranted. Regional 
estimates for some portions of the eastern North 
Atlantic suggest relatively large populations (Anony- 
mous 1993). Off West Greenland, considerable sur- 
vey efforts have been undertaken since 1984, yet few 
Killer Whales have been recorded, suggesting that 
Killer Whales are not abundant in that area 
(Anonymous 1993). 


Habitat 

Killer Whales do not appear to be as limited by 
such habitat considerations as depth, water tempera- 
ture, or salinity, as do some other cetaceans. Killer 
Whales are found in all oceans, in water ranging in 
temperature from below zero degrees Celsius 
(among ice floes) to warm tropical waters. They 
have been recorded in depths from as shallow as a 
few meters, to open ocean depths. Killer Whales will 
also occasionally spend considerable time in brack- 
ish water and will even enter rivers (e.g., Scheffer 
and Slipp 1948), including ascending into the lower 
reaches of the Fraser River in British Columiba. 
Resident Killer Whales around northern Vancouver 
Island have been documented using shallow inter- 
tidal and subtidal pebble beaches as rubbing sites 
(Hoyt 1990). 

Habitat use by residents and transients does differ 
(Heimlich-Boran 1988; Morton 1990; Felleman et al. 
1991; Baird et al. 1992; Ford et al. 1994a; Baird and 
Dill 1995). Both residents and transients frequent a 
wide range of water depths, but residents tend to 
spend more time in deeper water. Residents will 
occasionally move into water less than five meters 
deep, but some transient pods spend considerable 
time in even shallower depths, often foraging in 
inter-tidal areas at high tides. There appears to be 
considerable variability in habitat use among tran- 
sient pods, with some spending significantly greater 
time foraging in very near-shore areas than others 
(Baird and Dill 1995). Patterns of habitat use of resi- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


dent Killer Whales in Washington state and southern 
British Columbia were examined by Heimlich-Boran 
(1988) and Hoelzel (1993). Both studied the same 
population, but over different time periods; and 
results presented from the two studies differ some- 
what. Heimlich-Boran (1988) noted that residents 
fed more in specific areas with high relief 
bathymetry along the major routes for salmon migra- 
tion. Hoelzel (1993) found no correlation between 
behaviour, bottom topography or specific areas. The 
reasons for these discrepancies are unclear; habitat 
use may have shifted somewhat between the two 
study periods, or differences in methodology may be 
responsible. For both residents and transients, given 
the matrifocal nature of groups, movement patterns 
and habitat use are likely strongly influenced by 
learning within the matrilineal unit. 


General Biology 
Life History/Reproduction 

The most detailed information on life history of 
Killer Whales world-wide is available for northern 
and southern residents. Life history characteristics 
determined by Olesiuk et al (1990) and summarized 
below were calculated using data from both northern 
and southern residents (and all references to resi- 
dents in this section apply to both populations except 
where specified). As many of the characteristics may 
vary between populations, application of these val- 
ues to other populations, including “offshore” Killer 
Whales or transients, should be done with caution. 
Some information, such as gestation period, has best 
been established with captive animals. 

Gestation periods has been reported variously to 
be between 12—17 months. Using hormone levels of 
captive animals, gestation period has been measured 
at 517 days (17 months, SD = 20 days; Walker et al. 
1988; Duffield et al. 1995). Duffield et al. (1995) 
note that successful pregnancies with viable calves 
occurred from 15—18 months (468-539 days). Length 
at birth of residents (based on the smallest and 
largest newborn animals documented stranded) 
ranges from 218 to 257 cm (Olesiuk et al. 1990). The 
largest fetus recorded world-wide was 270 cm in 
length (Nishiwaki and Handa 1958). Calving occurs 
year-round, but there appears to be a peak in births 
between fall and spring (Olesiuk et al. 1990). Precise 
age at weaning is not. known, but Killer Whale 
calves begin taking solid food at a very young age. 
Heyning (1988) noted solid food in the stomach of a 
2.6 m long animal from California. No milk was vis- 
ible in the stomach of that animal, but contents were 
not tested for the presence of milk lactose. Using the 
ages at which southern residents begin spending 
more time away from their mother, as well as when 
they are observed taking fish, Haenel (1986) estimat- 
ed weaning to occur at between 1.0-1.5 to 2 years of 
age. 


2001 


Age at sexual maturity for females can be report- 
ed in a variety of ways, including age of first ovula- 
tion, age at first pregnancy, and age at first parturi- 
tion. Olesiuk et al. (1990) defined age at sexual 
maturity for females as the age at which they first 
give birth to a viable offspring, and noted that it 
varies between 12 and 16 years (mean = 14.9 
years). Onset of sexual maturity for males, defined 
as when the dorsal fin shape changes sufficiently 
enough to distinguish males from females, ranged 
between 10—-17.5 years (mean = 15 years) (Olesiuk 
et al. 1990). Dorsal fin growth for males continues 
for at least six years after onset of sexual maturity, 
and Olesiuk et al. (1990) suggest that physical 
maturity is reached at the end of that period. These 
criteria need to be evaluated using hormone levels 
of captive animals. Calving interval, defined as the 
interval between births of viable calves, ranges 
from 2 to 12 years, with a mean of about 5 years 
(Olesiuk et al. 1990). Calving interval increases 
slightly with age, but is extremely variable (Olesiuk 
et al. 1990). Fecundity rate (defined as the propor- 
tion of mature females which gave birth to viable 
calves each year) declines linearly with age 
(Olesiuk et al. 1990). Olesiuk et al. (1990) provide 
evidence of reproductive senescence in older 
females, and note that the mean age of onset of 
post-reproduction is about 40 years, although one 
female has given birth at 51 years of age. 

Mortality rates vary with age. Neonate mortality 
(birth to six months of age) is high. Using survival 
rates of calves first encountered during winter, and 
the presence of stranded animals, Olesiuk et al. 
(1990) estimated neonatal mortality of residents at 
37% and 50%, respectively. Bain (1990) indepen- 
dently estimated neonatal mortality in northern resi- 
dents at 42%, based on the distribution of calving 
intervals. Because of the high mortality rate during 
the first six months of life, longevity has usually 
been reported as the average lifespan of an animal 
which reaches six months of age. Average longevity 
has been estimated for male residents to be 29.2 
years (maximum estimated at 50-60 years), and for 
females to be 50.2 years (maximum estimated at 
80-90 years) (Olesiuk et al. 1990). At birth, average 
life expectancy is about 29 years for females and 17 
years for males (Olesiuk et al. 1990). 


Feeding Habits 

Information on diet composition is relevant to sta- 
tus assessment both in terms of potential limiting 
factors, as well as in population delineation. In gen- 
eral, Killer Whales world-wide are apex predators, 
with a wide range of prey reported, including squid, 
octopus, bony and cartilaginous fish, sea turtles, sea 
birds, sea and river otters, dugongs, pinnipeds, other 
cetaceans, and occasionally terrestrial mammals such 
as deer, moose and pigs (Heyning and Dahlheim 
1988; Guinet 1992; Jefferson et al. 1991). However, 


BAIRD: STATUS OF KILLER WHALES 


687 


individual populations of Killer Whales appear to 
specialize on particular types of prey, rather than 
exhibit opportunistic predation (Felleman et al. 
1991; Jefferson et al. 1991; Baird et al. 1992). In 
some areas, Killer Whales regularly steal fish from 
commercial fisheries (e.g., Leatherwood et al. 1990; 
Sivasubramanium 1965; Yano and Dahlheim 1994, 
1995), or scavenge discards thrown overboard (e.g., 
Couperus 1994). 

Both northern and southern residents appear to 
feed primarily on fish (Ford et al. 1998). More infor- 
mation is available for northern residents (126 pre- 
dation events) than for southern residents (35 preda- 
tion events), and most records of predation are from 
during the summer months (Ford et al. 1998). Based 
on collection and identification of scales from fish 
captured, Ford et al. (1998) note that 96% of the fish 
kills observed were salmonids, and of these, 65% 
were of Chinook (Oncorhynchus tshawytscha), the 
largest species occurring in that area. Stomach con- 
tents from eight residents also indicate this prefer- 
ence for salmon (7 of 8 individuals) and for Chinook 
(at least 4 individuals), though a variety of other 
species, including Pacific Halibut (Hippocampus 
stenolepis), Lingcod (Ophiodon elongatus) and a 
variety of other bottom fish were noted from one or 
two animals, and from occasional observations of 
predation (Ford et al. 1998). Ford et al. (1998) note 
that there are several potential biases that could 
affect diet composition in this type of study. It is 
possible that prey caught at great depths may be con- 
sumed prior to a whale returning to the surface. The 
limited information available on the depth distribu- 
tion of salmon in British Columbia and Washington 
suggest that most species spend the majority of their 
time in the upper levels of the water column (i.e., 
less than 30 m — Felleman 1986; Quinn and terHart 
1987; Quinn et al. 1989; Ruggerone et al. 1990; 
Olson and Quinn 1993), though Chinook are the 
deepest of the species captured. Thus most salmon 
captured likely have a relatively high probability of 
being observed, compared to prey which usually 
inhabit deeper portions of the water column. 
Southern resident Killer Whales regularly dive to 
depths greater than 100 m (Baird et al. 1998a), and 
may take a minute or more to return to the surface. 
While details on handling time of salmon have not 
been reported, it seems likely that similar-sized fish 
caught at such depths may be consumed prior to the 
whale’s surfacing. Information on handling time for 
various species and sizes of fish would be of value 
for assessing this possibility. Also, there may be a 
bias towards detecting captures of large prey. Small 
prey are likely swallowed whole, while large prey 
may be broken up prior to consumption or shared 
between individuals, thus increasing the chance that 
scales may be recovered at the surface. Little infor- 
mation is available on diet of “offshore” Killer 


688 


Whales, though they have not been observed feeding 
on marine mammals (Ford et al. 1994b). 

I believe that diet information from most observa- 
tional studies of transients is probably less biased, 
since marine mammal prey tend to be fairly large 
and often come to the surface to breathe during 
attacks, where they are easily seen (Baird and Dill 
1996; Ford et al. 1998). Even when live prey are not 
observed at the surface, prey species can often be 
inferred, based on location (e.g., a Harbour Sea! 
haulout), and the presence of large quantities of 
blood or blubber in the water. However, methods in 
many studies focused on acoustic recordings have 
required positioning of a research vessel relatively 
far away (e.g., 100s of meters) from whales under 
observation, and it may therefore not be possible to 
compare kill rates from one study to another (Baird 
2000). In one observational study, calculated food 
intake rates were more than sufficient to meet the 
animals’ predicted energetic needs, thus the vast 
majority of prey actually captured were probably 
documented (Baird and Dill 1996). Harbour Seals 
seem to be the preferred prey for transients, being 
both very abundant in many areas of British 
Columbia, as well as relatively easy to capture 
(Baird and Dill 1996; Ford et al. 1998). Virtually all 
other common species of marine mammals whose 
range overlaps with that of transients have also been 
documented as their prey, as have occasional sea 
birds and even terrestrial mammals (Pike and 
MacAskie 1969; Bigg et al. 1987; Jefferson et al. 
1991; Stacey et al. 1990; Baird and Dill 1996; Ford 
et al. 1998). 

Killer Whales in the Canadian Arctic and Atlantic 
have primarily been documented feeding on other 
marine mammals (Whitehead and Glass 1985; 
Mitchell and Reeves 1988), though some evidence of 
Killer Whales scavenging fish from around longlin- 
ing vessels is also available (Sergeant and Fisher 
1957): 


Social Organization 

Globally, Killer Whales have been observed trav- 
eling alone and in groups of up to several hundred 
individuals (Perrin 1982). However, larger groups 
appear to be temporary associations of smaller, more 
stable groups. In all areas where long-term studies 
have been carried out, evidence suggests stable 
multi-year associations between individuals with 
limited dispersal from maternal groups (Lopez and 
Lopez 1985; Bigg et al. 1990; Guinet 1991a; Simila 
and Ugarte 1993; Baird and Dill 1996; Baird and 
Whitehead 2000). Such evidence is most conclusive 
for Killer Whales in British Columbia and 
Washington (both transients and northern and south- 
ern residents), where there are extensive data on 
variability in group size, structure and stability. 
Differences in these characteristics do occur between 
the sympatric residents and transients. Little is 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


known about the social organization of “offshore” 
Killer Whales. 

Bigg et al. (1990) studied the social organization 
of northern and southern residents, and noted the 
average pod size in the two populations combined 
was about 12 individuals (range of 3—59 individu- 
als). No dispersal from resident pods has been docu- 
mented. Resident pods are thought to form by the 
gradual splitting of a single pod into two (Ford 
1990). Ford’s (1984; 1990) research on Killer Whale 
acoustics demonstrated the existence of stable pod- 
specific dialects, and showed that some pods shared 
a number of calls. He suggested that these reflected 
common ancestry. Ford (1990) defined acoustic 
clans, comprised of pods which share one or more 
calls, and identified four acoustic clans from the 
British Columbia coast, three within the northern 
resident community, and one in the southern resident 
community. 

For transient Killer Whales, average pod size 
reported by Baird and Dill (1996) was of about two 
individuals, with a range in pod size from one to four 
individuals. Those pods which consist of only a sin- 
gle individual (ie., individuals who do not associate 
with others for more than 50% of their time) appear 
to be of two types, either lone adult males which 
tend to spend much of their time alone (and only 
occasionally associates with other groups; Baird 
1994), or adult females which always associate with 
other groups, although none in a stable manner 
(Baird and Whitehead 2000). Transient pods are fair- 
ly stable, with some associations between individu- 
als documented lasting 15 years or more (Baird and 
Whitehead 2000). However, dispersal from transient 
pods has been recorded on two occasions (Bigg et al. 
1987; Baird and Dill 1996), and extensive indirect 
evidence of dispersal exists (Baird 2000). Transient 
pods often associate with one another, and no evi- 
dence of transient communities, as noted for resi- 
dents, has been found. Associations between tran- 
sient pods do not appear to be completely random 
however; they depend in part on pod size and the age 
and sex of all pcd members (Baird 2000), and in part 
on the foraging tactics exhibited by the pod, which 
appear to be pod-specific (Baird and Dill 1995). 


Limiting Factors 
Natural Mortality 
Potential sources of natural mortality fall into sev- 
eral categories: predation, parasitism, disease, 
biotoxins, accidental beaching, entrapment, and star- 
vation. No predators of Killer Whales have been 
recorded, but young or sick whales are potentially at 
risk from attacks by large sharks in some areas, and 
attacks by other Killer Whales may also pose a risk 
(see above). The relatively high incidence of scarring 
on animals also suggests that intraspecific aggression 
occurs (see Visser 1998). 


2001 


A variety of endoparasites have been recorded 
from Killer Whales, including trematodes, cestodes, 
and nematodes (see review in Heyning and 
Dahlheim 1988). Transmission of such parasites is 
primarily through ingestion of infected food items, 
but the role and extent of such parasites in causing 
natural mortality is unknown. External parasites 
have not been documented in British Columbia 
Killer Whales, but Killer Whales elsewhere have 
been seen with barnacles on the rostrum and trail- 
ing edge of flukes, and with a species of cyamid 
ectoparasite. The current understanding of the dis- 
eases and disease processes (e.g., Ridgway 1979) 
affecting Killer Whales is relatively advanced, as a 
result of the study of animals in aquaria (J. McBain, 
Sea World San Diego, personal communication). 
Relatively little of this research has been published 
however. Mortality due to biotoxins has not been 
reported for Killer Whales, though a number of 
large-scale mortality events in other cetaceans have 
been linked to this source (e.g., Geraci et al. 1989). 
Large-scale mortality events due to viral infections 
have been recorded in several populations of 
marine mammals in recent years (Osterhaus and 
Veder 1988; Duignan 1995), and while the occur- 
rence of such die-offs is unpredictable, given their 
magnitude and apparently increasing frequency of 
occurrence, they should be taken into account in 
conservation planning and population viability 
analysis (Young 1994; Simmonds and Mayer 
1997). 

Accidental beaching and entrapments of Killer 
Whales are an occasional source of natural mortali- 
ty. Several cases of beaching of live animals have 
been reported in British Columbia and off eastern 
Canada, both with large groups (Carl 1946; Dearden 
1958; Emery 1960) and lone individuals (Hoyt 
1990). There is one recent unpublished record of 
two adult individuals which live stranded and died 
during a storm on 28 January 1998, near 
Terranceville, Fortune Bay, Newfoundland (R. 
Hudson, personal communication). Mass strandings 
have also been reported from Alaskan waters 
(Hanson and Spraker 1996). The cause(s) of large 
group strandings are usually unclear (though 
Dearden 1958 reports animals being forced ashore 
by ice), but it seems more likely to occur for “off- 
shore” whales traveling on a rare occasion in 
inshore waters, than it does for inshore groups. Hoyt 
(1990) noted one transient individual apparently 
accidentally stranded while chasing porpoise in 
shallow water. Ice entrapments have been reported 
in the Canadian Arctic (Reeves and Mitchell 
1988a), off Newfoundland (Lien et al. 1988) and in 
the Antarctic (Taylor 1957). Several cases of ani- 
mals becoming entrapped in tidal lakes or inlets 
with narrow, shallow openings have also been noted 
(Emery 1960; Mitchell and Reeves 1988; Bain 


BAIRD: STATUS OF KILLER WHALES 


689 


1995). In many cases such entrapment has led to 
mortality of all or part of a group. Considering the 
small size of inshore populations of Killer Whales, 
such periodic events could seriously affect popula- 
tions. Temporary “entrapment” in narrow inlets has 
been documented for southern resident Killer 
Whales twice in recent years (Shore 1995, 1998). In 
the most recent case, occurring in southern Puget 
Sound in 1997, the whales’ reluctance to move 
under a bridge across the mouth of the inlet was 
suggested as a possible factor preventing their leav- 
ing the enclosed area (Shore 1998). Two of the 19 
whales which were in the inlet died at some point in 
the six months after they left the inlet, though it is 
unknown whether the “entrapment” was a contribut- 
ing factor to this mortality (Anonymous 1998). 


Anthropogenic Influences 

Potentially negative interactions with humans fall 
under two broad categories. Some impacts may have 
acute (immediate) effects on individuals or a popula- 
tion, such as directed takes (whaling, culling), live- 
capture fisheries, entanglement in fishing gear, colli- 
sions with vessels, or exposure to acute pollutants 
(e.g., oil spills). Immunotoxic affects due to accumu- 
lation of persistent toxic chemicals may also have an 
acute impact by increasing susceptibility to diseases, 
thus causing an increase in mortality. Besides these 
acute impacts, there are a number of less tangible, 
longer-term potentially negative human influences, 
including a reduction in reproductive rates due to 
accumulation of persistent toxic chemicals, reduced 
prey availability due to human activities, and distur- 
bance or displacement by vessel traffic or other 
sources of underwater sounds (Table 3). While each 
of these is treated independently below, it should 
also be taken into account that cumulative impacts of 
all of these factors could be important (or in the case 
of longer-term stressors, synergistic interactions 
between impacts could occur; Whitehead et al. 
2000). 

Killer Whales have been hunted for oil and meat 
(for human or animal consumption, fertilizer or bait) 
in many areas (e.g., Reeves and Mitchell 1988b; 
Oien 1988; Berzin and Vladimirov 1983; Miyazaki 
1983; Anonymous 1992; Kishiro and Kasuya 1993; 
Price 1985; Bloch and Lockyer 1988; Barnes 1991; 
Yu 1995, though the largest fisheries were discontin- 
ued in the early 1980s (Norway and Russia) or early 
1990s (Japan). Small numbers are probably still 
taken elsewhere however. Information on catches is 
currently reported by many countries that are mem- 
bers of the International Whaling Commission 
through their Annual Progress Reports, though not 
all countries submit Annual Progress Reports, and 
some countries may not include Killer Whales in 
their lists of catches, as they are considered “small 
cetaceans” and are thus not covered under the aus- 
pices of the IWC. Small numbers of animals may be 


690 


THE CANADIAN FIELD-NATURALIST 


Vol 5 


TABLE 3. A summary of causes of mortality or disturbance to Killer Whales and their potential role in population limita- 
tion. Those sections which are bold are considered to be the most serious threats or limiting factors for Killer Whale popu- 


lations in Canada. 


Known or thought to cause 


Potential to cause substantial 


population decline 


No 
Yes 


Yes 


Unlikely on west 


Trend in threat 
NA 
Unknown 


Steady? 


Decreasing on 


Threat population decline 
NATURAL 
Predation No 
Die-offs No (but yes with 
marine mammals 
elsewhere) 

Mass stranding or Yes 

entrapment in ice 

or narrow inlets 
ANTHROPOGENIC 
Culling/direct killing Yes 
Incidental mortality No 
Live capture Yes 
Vessel harassment No 
Vessel collision No 
Acoustic deterrents No 

from aquaculture 

operations 
Immunotoxicity No (but yes with 

marine mammals 
elsewhere) 

Oil spills No (but yes with 


other populations) 

Reduction in prey No (but yes with 

base marine mammals 
elsewhere 


taken by non-member countries, and such catches 
would probably be largely unreported. In Canadian 
Arctic waters, Killer Whales are only taken very 
occasionally by native people; Reeves and Mitchell 
(1988a) noted that there was no tradition of hunting 
of Killer Whales in this area. Fourteen Killer Whales 
were killed by native hunters in the Canadian Arctic 
in 1977, but these were individuals that had been 
trapped in a tidal lake (Mitchell 1979). Small num- 
bers were taken commercially off of Newfoundland 
and Nova Scotia in the 1940s through 1960s 
(Sergeant and Fisher 1957; Mitchell and Reeves 
1988), and it is possible that more animals were 
taken but not reported in an early fishery for pilot 
whales (Sergeant 1962; Mitchell and Reeves 1988). 
Catches in the 1960s and 1970s off Norway were 
both age- and sex-biased, and impacts on the current 
population may still be apparent (Vongraven and 
Bisther 1995). 

Culling (intentional shooting) of animals, because 
of their perceived (or documented) threat to fish- 
eries, has also occurred in British Columbia (Carl 


coast, possible in west coast, 
Arctic Canada unknown in 
eastern and Arctic 
No Steady? 
No Decreasing 
No (but possible Increasing 
contributing) 
No Increasing 
Unlikely Steady but 
potential to 
increase 
Yes Steady or 
increasing? 
Yes Increasing? 
Yes Decreasing for 
mammal-eating 
Killer Whales, 


unknown for fish-eating 


1946; Olesiuk et al. 1990), off eastern Canada (Lien 
et al. 1988), and elsewhere (e.g., Heide-Jorgensen 
1988). In British Columbia this included the 
Canadian Air Force using Killer Whales as practice 
targets (Carl 1946), as well as opportunistic shooting 
by fishermen and the federal fisheries department. 
That such culling may have had an impact on popu- 
lations earlier in this century is apparent in the popu- 
lation growth curves shown by Olesiuk et al. (1990; 
see also Figure 6), since both the northern and south- 
ern resident populations were growing (presumably 
recovering) prior to the initiation of the live-capture 
fishery. As such, it is possible that the populations 
today are still recovering. Elsewhere, as in Prince 
William Sound, Alaska, direct killing by fishermen 
in recent years in response to losses of fish may still 
be having significant effects on the local population 
(Dahlheim and Matkin 1994). Lien et al. (1988) note 
that the shooting of Killer Whales which congregat- 
ed around whaling ships to feed on captured baleen 
whales may have significantly decreased the popula- 
tion in that area (though whaling ceased there in 


2001 


1972). Kills in Greenland (Heide-Jorgensen 1988) 
may be from populations which share their range 
with Canada (Heide-Jorgensen 1993), and whether 
the ongoing occurrence of such kills is currently hav- 
ing an impact on populations off eastern and Arctic 
Canada should be investigated. Killer Whales in 
British Columbia are not reported to regularly take 
fish off fishing gear (G. Ellis, personal communica- 
tion), as has been reported elsewhere (e.g., Yano and 
Dahlheim 1995), thus it seems unlikely that illegal 
shooting because of perceived threats to fisheries is 
currently occurring to any great degree. One fatal 
shooting of an adult female northern resident was 
documented in 1983 (which also appeared to result 
in the death of her calf), and at least one other north- 
ern resident has a bullet hole in the dorsal fin (Ford 
et al. 1994). Because of the extremely low potential 
growth rate of Killer Whale populations, even occa- 
sional shooting could limit population growth, and 
some monitoring of such activities is warranted. 
How such monitoring would be undertaken is diffi- 
cult to envision. 

Incidental mortality in fisheries through acciden- 
tal entanglement in fishing gear appears to be rare 
for this species. A few gear entanglements have been 
reported in British Columbia, though not all have 
resulted in death of the entangled animals (Pike and 
MacAskie 1969; Jamieson and Heritage 1988; Ford 
et al. 1994b; Guenther et al. 1995*; Baird et al. in 
press). Entanglements have also been reported from 
other areas where individuals from the British 
Columbia population range (e.g., Alaska — Barlow 
et al. 1994; California — Heyning et al. 1994). Off 
the eastern coast of North America, no reports of 
incidental mortality in Canadian waters appear to 
have been published (Read 1994), though there is 
one record of a Killer Whale entangling in a ground- 
fish gillnet (though it did not die) in the U.S. waters 
of the Gulf of Maine (Waring et al. 1997), and one 
record of a Killer Whale entangling (but released 
alive) in a swordfish longline in international waters 
off of Newfoundland (A. Williams, NMFS, personal 
communication). As with direct killing, some efforts 
to estimate the numbers of animals which are killed 
through incidental mortality are warranted. While 
questionnaire surveys of fishermen are known to be 
extremely biased (Lien et al. 1994), reporting of such 
accidental events might be more likely than report- 
ing of deliberate activities like shooting. 

Live-capture fisheries for public display in ocea- 
naria have been focused in two areas, British 
Columbia/Washington and off Iceland (Asper and 
Cornell 1977; Bigg and Wolman 1975; Hoyt 1992). 
The last permit for captures in British Columbia was 
issued in 1982, although no animals were taken; the 
last animal taken from British Columbia was in 
1977, and it is unlikely any further captures would 
be allowed, due to widespread public opposition 


BAIRD: STATUS OF KILLER WHALES 


691 


(Hoyt 1990). Of 63 residents estimated to be 
removed from British Columbia/WA, 48 were 
thought to originate from the southern resident com- 
munity (Olesiuk et al. 1990), which currently num- 
bers only 89 animals (van Ginneken and Ellifrit 
1998). These captures substantially reduced the 
southern resident population and it took approxi- 
mately 20 years to return to levels similar to prior to 
the fishery (Figure 6). Besides a reduction in num- 
bers, the live-captures also resulted in a skewed age- 
and sex-composition of the population (Olesiuk et al. 
1990), due to selective cropping. Olesiuk et al. 
(1990) discuss a number of factors which may have 
resulted in the slow recovery of the southern resident 
population betwen 1970 and 1985, including the pos- 
sibility that the selective cropping of males (23 of 35 
known-sex individuals were males) may have 
reduced the number of mature males to below a criti- 
cal number for optimal productivity (Olesiuk et al. 
1990). 

In recent years, whale watching focusing on Killer 
Whales has become particularly prominent in 
Washington state and British Columbia, and vessel 
activity of all types (e.g., sports fishing, whale 
watching, ferry and freighter traffic) has been 
increasing (e.g., Osborne 1991). Whale watching in 
particular has raised a variety of concerns among 
researchers and members of the public about the 
potential for disturbance (Kruse 1991; Osborne 
1991; Duffus and Dearden 1992, 1993; Phillips and 
Baird 1993; Duffus and Baird 1995; Burgin and Otis 
1995; Baird et al. 1998b; Williams et al. 1998). 
Numerous behavioural changes have been reported 
in response to close approaches by boats, although 
some of the studies undertaken have serious method- 
ological problems, causing researchers to question 
their validity (Duffus and Dearden 1993). Studies 
have focused both on northern residents in Johnstone 
Strait and southern residents in Haro Strait. A num- 
ber of differences between these sites, the popula- 
tions of whales which use them, the number and 
types of boats found in the two areas, and the 
research methodologies being used in each site, pre- 
clude any simple comparison of results from the two 
areas. Changes in behaviour in response to approach 
by boats have been demonstrated for northern resi- 
dents (Trites et al. 1996; R. Williams, personal com- 
munication). Unfortunately the implications of such 
changes in behaviour on reproduction or mortality 
are unclear. While similar behavioural changes have 
not yet been demonstrated for southern residents 
(Osborne 1991; Burgin and Otis 1995; Baird et al. 
1998b), there does appear to have been a substantial 
decrease in the proportion of time southern residents 
engage in resting behaviour during daylight hours, 
coincident with the large increase in whale watching 
activity (Osborne 1986; K.C. Balcomb, R. W. 
Osborne, personal communications). For transient 


692 


Killer Whales, Barrett-Lennard et al. (1996) suggest- 
ed that vessel noise might impair their ability to 
detect prey. The impact of a single boat would 
appear negligible, as Baird and Dill (1996) found 
that under such circumstances observed food intake 
rates of transients were more than sufficient to 
account for the whale’s energetic needs. However, at 
least in some areas of the province and at some times 
of the year, such impacts could be serious. In the last 
few years (since 1993), it is not uncommon for small 
groups of transient Killer Whales to be accompanied 
by 5-10 boats when travelling off the Victoria area 
during summer months (Baird, personal observa- 
tions), and such large numbers of boats seem more 
likely to impact foraging success in the way suggest- 
ed by Barrett-Lennard et al. (1996). In one area in 
Washington state, the number of vessels found 
around groups of southern resident Killer Whales 
has been increasing (Figure 5), and in 1997 groups 
were accompanied by an average of 25 vessels (only 
one quarter of which are commercial whale watching 
vessels) during daylight hours in the summer months 
(Baird et al. 1998b; Figure 6). The commercial 
whale watching fleet in the area (including the ports 
of Victoria, Sidney and Sooke in British Columbia, 
and Bellingham and Friday Harbor in Washington) 
has been increasing rapidly, and numbered over 80 
boats in 1997 (Baird et al. 1998b). As noted above 
(see Protection, National), levels of awareness of, 
and adherence to, whale watching guidelines are 
largely unknown (except in a few specific localities 
during summer months), and virtually no official 
monitoring or enforcement of whale watching guide- 
lines takes place. 

A more direct impact of boats on whales involves 
injuries or deaths from collisions. Considering the 
large number of vessels interacting with Killer 
Whales during the summer months in British 
Columbia, vessel collisions are extremely rare. One 
well-documented case in British Columbia has been 
reported (Anonymous 1974), with an animal appar- 
ently fatally wounded after a collision with a large 
vessel (a ferry). Ford et al. (1994a) note the animal 
struck may have been part of the northern resident 
population. Several other live animals have been 
seen with scars that might be attributable to vessel 
interactions, although the evidence for this is incon- 
clusive. One vessel collision with a southern resident 
Killer Whale in Haro Strait, Washington, was wit- 
nessed in 1998 (V. Shore, personal communication), 
but the vessel was moving slowly and the animal did 
not appear to be injured as a result of the collision. A 
northern resident was struck by a speed boat in 1995 
and received a wound to the dorsal fin, which 
appeared to heal quickly (P. Spong, H. Symonds, 
personal communications). 

The generation of loud underwater sounds through 
such sources as acoustic deterrent devices (“seal scar- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


ers” or ADDs) at aquaculture operations also has the 
potential to cause disruption of movement patterns or 
even abandonment of an area (Morton and Symonds 
1998*). Morton and Symonds (1998*) noted a drastic 
reduction of use of the Broughton Archipelago, off 
northeastern Vancouver Island, by both resident and 
transient Killer Whales, coincident with the installa- 
tion of several high amplitude ADDs in the area. 
Residents have shown a decline in use of the area 
(measured as number of days per year observed) of 
over 75%, while transient use of the area has de- 
clined by over 50%. The lesser decline by transients 
may be due to their use of specific channels in the 
area away from the ADDs (Morton and Symonds 
1998*). Morton and Symonds (1998*) compared use 
of the Broughton Archipelago with the nearby 
Johnstone Strait, and area where no ADDs were in 
use, and where resident use of the area has been 
more-or-less stable over the same periods. Their 
study provides evidence of Killer Whale avoidance of 
areas where ADDs were in operation (Nichol and 
Sowden 1995 present information on avoidance of 
another species), thus potential effects by their use by 
other aquaculture facilities on the British Columbia 
coast should be studied, and increased use of such 
devices should be strictly regulated. 

More critical conservation problems concern two 
general areas, the effect of pollutants and the reduc- 
tion of the prey base due to anthropogenic activities. 
Two general groups of pollutants warrant discussion: 
(1) persistent toxic chemicals which bioaccumulate; 
and (2) petroleum products. Killer Whales from 
British Columbia and neighboring areas have been 
shown to accumulate high levels of persistent toxic 
contaminants (Calambokidis et al. 1990; Jarman et 
al. 1996; Matkin et al. 1998; Ross et al. 1998). 
Populations of resident Killer Whales in British 
Columbia spend a large proportion of their time in 
near-shore waters in close proximity to various 
sources of pollutants. A recent study using samples 
collected from free-ranging southern residents 
demonstrated that levels of PCBs and PCDD/Fs in 
these animals were three times higher than levels 
known to be immunotoxic for harbour seals (Ross et 
al. 1998). Levels in individual southern residents 
were three to five times higher than in individuals of 
equivalent age/sex classes of northern residents (P. 
Ross, personal communication), which live in an 
area with far fewer sources of pollutants (see 
Evaluation). Transients appear to spend less time in 
highly polluted areas, but feed almost exclusively on 
marine mammals, so may accumulate higher levels 
of many toxins. Matkin et al. (1998) note that levels 
of PCBs and DDTs in transients from Prince 
William Sound were 14 and 22 times higher, respec- 
tively, than for residents from the same area. They 
also note that the group of transients with high levels 
have shown no recruitment since 1984, suggesting 


2001 


that there may be a linkage between the low rate of 
reproduction and the high contaminant levels. 
Although sample sizes are smaller and based on 
stranded animals, levels of mercury appear to be 
higher in the tissues of residents than transients 
(Langelier et al. 1990). One possible candidate for 
these high levels of toxins in residents is consump- 
tion of heavily contaminated prey, but consumption 
of such prey has not become apparent in the studies 
of foraging undertaken to date. The high levels 
found in southern residents could affect reproduc- 
tion, immune function and endocrine function (Ross 
et al. 1996a, 1996b, 1998). Reproductive or en- 
docrine function impacts fall into the long-term cate- 
gory of effect. Immune function affects can result in 
acute (immediate) impacts on individuals or on the 
population. For example, it is thought that the 1988 
morbilivirus-associated mass mortality of harbour 
seais in northern Europe, which resulted in a popula- 
tion reduction of over 50%, was exacerbated by such 
immuotoxic effects of contaminants (Ross et al. 
1996a, 1996b; de Swart et al. 1996). 

Large-scale dumping of oil has the potential for 
detrimental effects on Killer Whale populations. 
Concurrent with the Exxon Valdez spill in Prince 
William Sound, Alaska, was the unprecedented loss 
of 14 Killer Whales from one pod which was seen in 
the area immediately following the spill (Matkin et 
al. 1994). Dahlheim and Matkin (1994) reviewed the 
evidence for a cause-and-effect relationship between 
the spill and the deaths of these whales, and con- 
clude that while there was a strong spatial and tem- 
poral correlation between these events, insufficient 
evidence is available to determine whether the spill 
caused the deaths, or other factors, particularly fish- 
eries interactions, were responsible. Given the large 
amount of tanker traffic on the British Columbia and 
Washington coasts, there is a potential for the loss of 
a large proportion of a population. In Juan de Fuca 
Strait, tanker traffic has been increasing, and tankers 
are aging (existing tankers are not expected to be 
phased out and replaced with double-hulled tankers 
until 2015; F. Felleman, personal communication). 
Thus a population such as the southern residents 
(considering their tendency to congregate in one area 
during the summer months) may be at risk from a 
major spill. 

It is beyond the scope of this review to undertake 
an assessment of trends in the abundance of all the 
potential prey species of Killer Whales, and in fact it 
is likely that such an assessment is impossible (or 
nearly so) given the data that are available. Re- 
gardless, in terms of a reduction in the prey base 
available for Killer Whales, it is clear that at least 
some of the populations of prey species of Killer 
Whales are substantially smaller today than histori- 
cally. In terms of salmon, anthropogenic influences 
on populations have included destruction, degreda- 


BAIRD: STATUS OF KILLER WHALES 


693 


tion and/or prevention of access to breeding habitat 
through urbanization, slides associated with road or 
railroad building, dam building, forestry and agricul- 
ture, as well as a reduction in numbers through fish- 
ing (Groot and Margolis 1991; Nehlsen et al. 1991; 
Slaney et al. 1996). Recent reviews of stocks from 
southeast Alaska through to California document 
large-scale reductions in many stocks, and extinction 
of others (Nehlsen et al. 1991; Baker et al. 1996; 
Slaney et al. 1996; see also e.g., Holtby and 
Finnegan 1997*; Wood et al. 1997*; Bradford 1998; 
Rutherford et al. 1998*). These reviews focus pri- 
marily on evaluating which stocks are at risk of 
extinction. Killer Whales will be affected by a sim- 
ple reduction in numbers, rather than only an extinc- 
tion of stocks. Within British Columbia, the salmon 
populations most drastically impacted are those in 
the southern part of the province, particularly the 
Strait of Georgia (Slaney et al. 1996), coinciding 
with the population of Killer Whales (southern resi- 
dents) which seems most at risk (see Population Size 
and Trends, and Evaluation). Some evidence is also 
available for declines of other potential prey species 
(e.g., Fargo 1997*; Leaman and McFarlane 1997*; 
Stanley and Haist 1997*; Ware 1997*; Yamanaka 
and Kronlund 1997*). Reduced prey availability 
could result in an increase in the amount of time 
whales would need to spend foraging, potentially 
leading to reduced reproductive rates and/or 
increased mortality rates. Insufficient information is 
available to assess whether such impacts are current- 
ly manifest. Given the inherent difficulty of deter- 
mining such impacts even if they exist, and the 
potentially large role they might have on increasing 
mortality or decreasing reproductive rates, a precau- 
tionary approach is warranted (Richards and 
Maguire 1998). 


Special Significance 

Among the cetaceans, Killer Whales exhibit sev- 
eral unusual features related to social organization 
and behaviour. One is the presence of the two sym- 
patric populations (residents and transients) in the 
nearshore waters of the eastern North Pacific, each 
specializing on different prey types, and differing in 
behaviour, acoustics, and morphology (Baird and 
Stacey 1988; Bain 1989; Ford and Hubbard-Morton 
1989; Morton 1990). Such a situation, with foraging 
specializations occurring among sympatric popula- 
tions, is unusual for mammals, as well as for verte- 
brates in general (see Mayr 1996; Otte and Endler 
1989). This system may provide valuable informa- 
tion on the causes and consequences of reproductive 
isolation between populations (Baird 1994). 

One apparent consequence of the differences in 
diet between the two forms are differences in disper- 
sal patterns. For residents no dispersal of either sex 
occurs; individuals travel in long-term stable groups 


694 


comprised of several maternal lineages. This situa- 
tion has not been documented for any other popula- 
tion of cetacean, or any other species of non-human 
mammal. For transients, dispersal of most individu- 
als of both sexes from the maternal group occurs, 
though not all male offspring seem to disperse (Baird 
1994, 1995b, 1998; Baird and Whitehead 2000). 
Such variability in dispersal patterns between sym- 
patric populations of closely related animals pro- 
vides a unique opportunity for examining some of 
the costs and benefits of group living. 

The types of foraging specialization found in pop- 
ulations in the eastern North Pacific may also occur 
elsewhere in the world, though research efforts else- 
where have been generally insufficient to determine 
whether sympatric forms specializing on different 
prey types exist. Individuals of some populations 
feed almost exclusively on other marine mammals. 
Such predation on marine mammals makes the study 
of foraging behaviour easier than perhaps for any 
other species of cetacean. Several interesting find- 
ings have come from these studies, including evi- 
dence that females teach hunting skills to their off- 
spring (Lopez and Lopez 1985; Guinet 1991b; 
Guinet and Bouvier 1995), and also a strong rela- 
tionship between group size and foraging success in 
one population (Baird and Dill 1996). 

Besides these intrinsic characteristics, Killer 
Whales also hold an unusual fascination for humans. 
Such fascination is reflected in the large attendance 
figures at aquaria which hold Killer Whales around 
the world, through the demand for commercial 
excursions to see these animals in the wild, and 
through the large number of popular books, maga- 
zine articles and films which have been devoted 
towards these animals. In the Haro Strait region, a 
trans-boundary area between Washington state and 
British Columbia, a large and growing whale watch- 
ing industry focused on this species exists (Baird et 
al. 1998b; Figure 5). Ticket sales for this area (in 
both the U.S. and Canada combined) were estimated 
to be approximately 5.5 million (U.S.) dollars in 
1997 (R.W. Osborne, personal communication). 


Evaluation 

The taxon Orcinus has been evaluated by the 
IUCN Cetacean Specialist Group, and designated as 
Lower Risk: Conservation Dependent (IUCN 1996). 
This category, effectively one level below the IUCN 
Vulnerable category (which includes species facing a 
high risk of extinction in the wild in the medium- 
term future), includes species which are the focus of 
a continuing conservation program, the cessation of 
which would result in qualifying for one of the high- 
er (e.g., Vulnerable) categories within a period of 
five years IUCN 1996). The COSEWIC classifica- 
tion of “Endangered” is for species “facing imminent 
extirpation or extinction”, “Threatened” is for 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


species that are “likely to become endangered if lim- 
iting factors are not reversed”, and “Vulnerable” is 
for those species “of special concern because of 
characteristics which make [them] particularly sensi- 
tive to human activities or natural events” (Campbell 
1996). The COSEWIC definition of “species” is par- 
ticularly important in evaluation of the status of 
Killer Whales, as it explicitly includes any “sub- 
species, variety or geographically defined popula- 
tion[s]”. Evidence is summarized below regarding 
Killer Whale populations in Canada relevant to such 
classification. 

Off the British Columbia coast, Killer Whales are 
subdivided into a number of populations, and these 
populations are distinct genetically, morphologically, 
and behaviourally (see e.g., Table 1). Based both on 
these biological characteristics and the COSEWIC 
“species” definition, it is clear that these populations 
could warrant independent evaluation and classifica- 
tion, where appropriate (it should also be noted that 
these populations are evaluated and listed indepen- 
dently in the U.S. — Barlow et al. 1997; Hill et al. 
1997). One of the British Columbia populations, the 
“northern” residents, has been growing since the end 
of live-capture fisheries in the early 1970s, but the 
population in British Columbia only numbers just 
over 200 individuals. The “southern” residents have 
not shown a steady increase, and the population size 
has declined by 10% in the last three years (1996— 
1998), to a level below that prior to live-captures 
(Figure 6). This decline is due to an increase in mor- 
tality rate, particularly mortality of adult females. The 
cause or causes of this increase in mortality are 
unclear, but there are several possibilities (Table 3). 
The core area for southern residents (Haro Strait) is 
bounded by the cities of Vancouver, Victoria and 
Seattle, with over 5.5 million people living in the 
area, increasing numbers of commercial and recre- 
ational vessel traffic, and numerous sources of pollu- 
tants. It seems unlikely that either pollution of these 
waters or vessel traffic will decrease in the near 
future. Southern residents have toxic chemical levels 
three times higher than levels known to cause 
immunotoxicity in Harbour Seals, and the most 
immediate anthropogenic risk to these populations is 
likely immunotoxic effects from this accumulation of 
persistent toxic chemicals (see Ross et al. 1996a, 
1996b, 1998). Potential impacts of a reduction in 
prey populations and increasing numbers of commer- 
cial and recreational whale watching boats (Figure 5) 
may also be serious threats, although insufficient 
information is available to evaluate the magnitude of 
these threats. In terms of reduction of salmon popula- 
tions, numbers in the Strait of Georgia, where the 
southern residents spend a large proportion of their 
time, have been reduced to a larger extent than popu- 
lations elsewhere in British Columbia (Slaney et al. 
1996). 


2001 


For transient and “offshore” Killer Whales in 
British Columbia, no population trend information is 
available, though, as with both northern and southern 
residents, population sizes appear to be small. 
Transients feed high on the food web, and are likely 
also at risk from high levels of contamination by per- 
sistent toxins. For Killer Whales in the Canadian 
Arctic and Atlantic, no information on population 
identity or trends is available, though populations 
appear to be very small, and the threats which face 
British Columbia populations likely also impact east- 
ern Canadian and Arctic populations. Because of 
their small population sizes (in the low hundreds), 
Killer Whales are also at risk from natural events 
(e.g., entrapment or mass stranding) which could 
drastically impact a local population. 

From the above, it is clear that all populations of 
Killer Whales in Canadian waters should, at the min- 
imum, be considered vulnerable. The only question 
that remains is whether one of the populations, the 
southern residents, should be considered threatened. 
The population is extremely small (89 individuals in 
1998), has declined by 10% in three years due to an 
increase in mortality rates, and several threats have 
been identified which have the potential to cause this 
population to become endangered. As noted, it is 
unlikely that at least some of these threats (pollu- 
tants, vessel traffic) will be reversed in the foresee- 
able future. While it is unclear whether the recent 
(1996-1998) decline is directly due to these anthro- 
pogenic factors, or whether the population will con- 
tinue to decrease, the rate at which this population 
has declined demonstrates how quickly such a popu- 
lation could become in danger of extirpation. That 
the threats to the population are insidious, difficult to 
quantify, and even harder to rectify, all warrant a 
conservative (precautionary) approach to manage- 
ment (see Richards and Maguire 1998). Without a 
COSEWIC designation, it seems unlikely that any- 
thing will be done regarding mitigation of these 
impacts, and the population could become endan- 
gered well before another evaluation is undertaken. 
Since the population is a trans-boundary stock, 
efforts to coordinate actions with U.S. management 
agencies are also required. 


Acknowledgments 

I thank Bob Campbell for his patience, as well as 
his support and the support of COSEWIC. The World 
Wildlife Fund (Canada) provided financial support in 
the form of a contract for the production of this 
report. [ would like to thank Mercedes Guerrero for 
her help with an earlier incarnation of this 
manuscript. Janet Mann, Richard Connor and Peter 
Tyack provided many helpful suggestions on a relat- 
ed manuscript which provided the basis for this one. I 
also thank John Caldwell of the Wildlife Trade 
Monitoring Unit, World Conservation Monitoring 


BAIRD: STATUS OF KILLER WHALES 


695 


Centre, for providing trade statistics from the CITES 
Secretariat as well as answering numerous questions 
regarding CITES and its coverage, as well as com- 
menting on the relevant section of this manuscript. A 
large number of individuals reviewed all or part of 
various versions of the manuscript, and/or provided 
unpublished data or copies of manuscripts or reports, 
including Masao Amano, Dave Bain, Ken Balcomb, 
Lance Barrett-Lennard, Marilyn Dahlheim, Dave 
Duffus, David Ellifrit, Graeme Ellis, Candi Emmons, 
Fred Felleman, Kerry Finley, John Ford, Ray 
Gambell, Astrid van Ginniken, Brad Hanson, Mads 
Peter Heide-Jorgensen, Rus Hoelzel, Sascha Hooker, 
Erich Hoyt, Michael Kingsley, Jon Lien, Ed 
Lochbaum, Chris Malcolm, Tony Martin, Craig 
Matkin, Lena Measures, Robert Michaud, Patrick 
Miller, R.V. Miller, Rich Osborne, Bob Otis, Rod 
Palm, Jake Rice, Peter Ross, Randy Reeves, Naomi 
Rose, Richard Sears, Val Shore, Paul Spong, Helena 
Symonds, Sean Todd, Hal Whitehead, Rob Williams, 
Pam Willis, and several anonymous reviewers. My 
apologies for any names accidentally omitted. 


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Report Number 175135023, Marine Mammal Com- 
mission, Washington, D.C. 

Matkin, C. O., G. M. Ellis, M. E. Dahlheim, and J. Zeh. 
1994. Status of Killer Whales in Prince William Sound, 
1985-1992. Pages 141-162 in Marine mammals and the 
Exxon Valdez. Edited by T.R. Loughlin. Academic 
Press, San Diego. 

Matkin, C. O., D. R. Matkin, G. M. Ellis, E. Saulitis, and 


BAIRD: STATUS OF KILLER WHALES 


699 


D. McSweeney. 1997. Movements of resident Killer 
Whales in southeastern Alaska and Prince William 
Sound, Alaska. Marine Mammal Science 13: 469-475. 

Matkin, C. O., D. Scheel, G. Ellis, L. Barrett-Lennard, 
H. Jurk, and E. Saulitis. 1998. Comprehensive Killer 
Whale investigation, Exxon Valdez oil spill restoration 
project annual report (Restoration Project 97012). North 
Gulf Oceanic Society, Homer, Alaska. 

Mayr, E. 1996. The modern evolutionary theory. Journal 
of Mammalogy 77: 1-7. 

Mayr, E., and P. D. Ashlock. 1991. Principles of system- 
atic zoology. Second Edition. McGraw-Hill, Inc. New 
York. 

Meyers, D. F. 1990. Model comparison for two popula- 
tions of Killer Whales, Orcinus orca. M.Sc. Thesis, 
Humboldt State University, Arcata, California. 

Mikhalev, Y. A., M. V. Ivashin, V. P. Savusin, and F. E. 
Zelenaya. 1981. The distribution and biology of Killer 
Whales in the southern hemisphere. Reports of the 
International Whaling Commission 31: 551-566. 

Mitchell, E. 1979. Canada progress report on cetacean 
research June 1977—May 1978. Reports of the Interna- 
tional Whaling Commission 29: 111-114. 

Mitchell, E., and R. R. Reeves. 1988. Records of Killer 
Whales in the western North Atlantic, with emphasis on 
eastern Canadian waters. Rit Fiskideildar 11: 161-193. 

Miyazaki. N. 1983. Catch statistics of small cetaceans 
taken in Japanese waters. Reports of the International 
Whaling Commission 33: 621-631. 

Morton, A. B. 1990. A quantitative comparison of the 
behaviour of resident and transient forms of the Killer 
Whale off the central British Columbia coast. Reports of 
the International Whaling Commission Special Issue 12: 
245-248. 

Nehlsen, W., J. E. Williams, and J. A. Lichatowich. 1991. 
Pacific salmon at the crossroads: stocks at risk from Cali- 
fornia, Oregon, Idaho, and Washington. Fisheries 16(2): 
4-21. 

Nichol, L. M., and D. M. Shackleton. 1996. Seasonal 
movements and foraging behaviour of northern resident 
Killer Whales (Orcinus orca) in relation to the inshore 
distribution of salmon (Oncorhynchus spp.) in British 
Columbia. Canadian Journal of Zoology 74: 983-991. 

Nichol, L. M., and M. J. Sowden. 1995. The effect of an 
acoustic deterrent device on the relative abundance and 
distribution of harbour porpoise (Phocoena phocoena) at 
a study site in coastal British Columbia. Department of 
Fisheries and Oceans Contract Report Number FP597-4- 
3004/01/XSA. 

Nishiwaki, M., and C. Handa. 1958. Killer Whales 
caught in the coastal waters off Japan for recent ten 
years. Scientific Reports of the Whales Research Insti- 
tute 13: 85-96. 

Oien, N. 1988. The distribution of Killer Whales (Orcinus 
orca) in the North Atlantic based on Norwegian catches, 
1938-1981, and incidental sightings, 1967-1987. Rit 
Fiskideildar 11: 65-78. 

Olesiuk, P. F., M. A. Bigg, and G. M. Ellis. 1990. Life 
history and population dynamics of resident Killer 
Whales (Orcinus orca) in the coastal waters of British 
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Olson, A. F., and T. P. Quinn. 1993. Vertical and hori- 
zontal movements of adult chinook salmon Oncorhy- 


700 


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Osborne, R. W. 1986. A behavioral budget of Puget Sound 
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Osborne, R. W. 1991. Trends in Killer Whale move- 
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Osterhaus, A.D.M.E., and E.J. Vedder. 1988. 
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Otte, D., and J. A. Endler. Editors. 1989. Speciation and 
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Perrin, W. F., and S.B. Reilly. 1984. Reproductive 
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Phillips, C. 1996. Conservation in practice: agreements, 
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Phillips, N. E., and R. W. Baird. 1993. Are Killer Whales 
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Pike, G. C., and I. B. MacAskie. 1969. Marine mammals 
of British Columbia. Bulletin of the Fisheries Research 
Board of Canada 171: 1-54. 

Price, W.S. 1985. Whaling in the Caribbean: historical 
perspective and update. Reports of the International 
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Quinn, T. P., and B.A. terHart. 1987. Movements of 
adult sockeye salmon (Oncorhynchus nerka) in British 
Columbia coastal waters in relation to temperature and 
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Quinn, T.P., B. A. terHart, and C. Groot. 1989. 
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Read, A. J. 1994. Interactions between cetaceans and gill- 
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of the International Whaling Commission Special Issue 
15: 133-147. 

Reeves, R. R., and E. Mitchell. 1988a. Distribution and 
seasonality of Killer Whales in the eastern Canadian 
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Reeves, R. R., and E. Mitchell. 1988b. Killer Whale 
sightings and takes by American pelagic whalers in the 
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Rice, D. W. 1968. Stomach contents and feeding behavior 
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Hvalfangst-Tidende 57: 35-38. 

Richards, L. J., and J.-J. Maguire. 1998. Recent interna- 
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directions for fisheries management science. Canadian 


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Vol. 115 


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Ridgway, S. H. 1979. Reported causes of death of captive 
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Ross, P. S., M. G. Ikonomou, G. M. Ellis, L. G. Barrett- 
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2001 


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701 


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Accepted 19 February 2002 


Book Reviews 


ZOOLOGY 


Birder's Mexico 


By Roland Wauer. 1999. Texas A&M University Press, 
College Station, Texas. xxvi + 304 pp., illus. Paper 
$18.95. 


The author is a retired National Park Service 
ranger from Texas. During his life he has had ample 
opportunity to visit Mexico and to study the Mexican 
specialties that spill over onto the U.S. side of the 
border. In this book he displays his affinity to 
Mexico’s flora, fauna, and people. As this book is 
about wild places, the issues that normally pepper 
travel accounts, like crime and poverty, are replaced 
by a sense of hospitality and friendliness from 
Mexico’s rural inhabitants. 

The book is divided into sections that describe the 
four major regions of Mexico, running from north to 
south. The author has selected to describe, within each 
region, a handful of specific areas, visited by him in 
the past. Although the book’s title suggests this work 
is on birds, it contains a lot of information on plant 
life. While this is not a detailed plant account, the 
author gives a good sense of the general habitat type. 
Some plants are specifically identified because they 
are key species, and they help in defining the distinct 
environment used by particular species of birds. The 
book has a number of half page, black-and-white 
illustrations. Most of these are habitat photographs or 
scenery shots related to vegetation. Wauer also notes 
any interesting mammal or reptile sightings. 

Wauer describes the trips he has made during 30 
years to some of Mexico’s best birding places, both 
as a professional and an ardent birder. Often he 
searched for a specific rarity or group of rarities. He 
describes the locality and how to get there. The 
author adds practical travel advice on restaurants, 
hotels, ferries, gas stations, and the like. He warns of 
rough roads and any problem associated with a 
remote locality. He does not mention any problem 
with wild beasts, not even ticks and mosquitoes. I 
presume the area is too dry for bugs and the other so- 
called dangerous animals rarely bother anyone. 

When searching for a bird the author’s description 
reveals much about the bird’s habits and habitat. He 
generally adds a short list of significant birds; local 
specialties, rare and unusual finds, etc. Indeed he has 


added a plant list just before the bibliography but the 
book does not contain an exhaustive bird checklist. 

The author has one habit I found tedious. He 
repeats references and similar material in full, every 
time. For example, he uses Jerry Radz Flora de 
Mexico as a reference throughout his travels. This 
book is referenced in full repeatedly. | would have 
preferred the more normal superscript ['] method, or 
at least “(Radz 19)”. Similarly he makes reference to 
other chapters as “Chapter X The slender-billed 
Wren” rather than “(chapter X)”. The author 
includes the names and home localities of all his 
companions for every trip. While it is nice to be 
mentioned in print, for those of us outside this group 
of friends name repetition is tiresome. The trip par- 
ticipants could have been covered in an appendix. I 
would have preferred the author had used capitals for 
each species. I recognize this is not considered cor- 
rect grammar by some, but it makes the species 
name more obvious (that is a “brownish graceful 
mockingbird” becomes a “brownish Graceful Mock- 
ingbird.” In the second case it is evident which is an 
adjective and which is the bird’s name. ) 

This book was lots of fun to read. The author 
clearly enjoyed the exploration of Mexico’s wild 
places and his contact with the locals and this sense 
of endearment came through in his writing . Like any 
good book it was easy to put yourself in the author’s 
shoes and feel a part of his adventure. The descrip- 
tions of the habitat give a good sense of the country- 
side, without being heavy and cumbersome. The 
reader can easily share his delight and sense of dis- 
covery when he finds a target bird. 

While this is a good fireside book, it will also be 
useful to anyone visiting Mexico (and probably Mexi- 
can residents too.) However for some of the species at 
risk his directions are quite spartan. I suspect this is 
deliberate. Any worthy birder should be able to fill in 
the gaps, whereas a less desirable visitor would find 
the route only with great difficulty. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario K1J 6K5 Canada 


702 


2001 


The Destruction of the Bison 


By Andrew C. Isenberg. 2000. Cambridge University 
Press, New York. 206 pp., illus. U.S. $29.95. 


As I began to read this book, I wondered, “What 
could possibly be added to the story of bison ecology 
and extermination not already presented in Frank 
Roe’s masterful 957-page compendium, The North 
American Buffalo: A Critical Study of the Species in 
its Wild State (University of Toronto Press, 1951)?” 
The answer is: Quite a lot. Isenberg deals more 
extensively with the socio-cultural aspects, especial- 
ly recent studies in social, cultural, labour, and gen- 
der history. As he says, “Insights from those fields 
suggest far greater complexities in the relationships 
of Indians and Euroamericans to the bison than his- 
torians formerly supposed.” 

Unfortunately, Isenberg’s summations are almost 
exclusively restricted to affairs south of the 49th par- 
allel. He tells nothing about the more symbiotic rela- 
tions between Indians and fur traders in Canada, and 
makes no mention of the arrival of the Canadian 
Pacific Railway. His sweeping generalization that 
“The fur trade on the western plains emerged from 
the spread of horses” would be less true for western 
Canada, where the appearance of guns and steel 
traps was perhaps more important than the advent of 
the horse, and furs were more readily transported by 
canoe. 

Isenberg believes that ecological and economic 
changes were inseparable on the plains in the 18th 
century. “Horses and smallpox — an Old World ani- 
mal and an Old World disease — destroyed the dom- 
inance of the Missouri River villagers and levered 
the nomads to power in the grasslands. ... The bison 
liberated and empowered the plains nomads. ...[its] 
flesh served as food, its skin as clothing and lodging, 
its sinew as thread, its horns and bones as tools, even 
dried bison manure served as fuel.” The annual har- 
vest was six or seven bison per person per year. But, 
by selling their furs to the Euroamerican market, the 
aboriginals “bound their fate to the Euroamerican 
economic and ecologic context.” Nevertheless, the 
ethic of communal cooperation, necessitated by the 


BOOK REVIEWS 


703 


precariousness of subsistence, together with their 
nomadic habits, caused the Indians to disdain the 
accumulation of property. 

During the 19th century, bison robes supplanted 
beaver pelts as the prime object of trade. In 1825, 
184,000 robes were shipped through New Orleans; 
in 1847, 110,000 hides through St. Louis; in 1872- 
73, about 400,000 hides out of Dodge City. Hunters 
wounded two or three bison for every one they 
killed. This carnage, augmented by anthrax among 
bison, 1821-31, smallpox among Indians in 1837-40 
and cholera in 1849-50, and by drought in the decade 
after 1846, all combined to speed the eradication of a 
bison-based way of native life. The extension of the 
Northern Pacific Railroad into Montana, in 1881, 
administered the coup-de-grace for the northern pop- 
ulations of bison. With the advent of railroads, bison 
were replaced by domestic cattle, viewed by many as 
a “victory for civilization. ... not justice but destiny.” 
The result for natives was poverty, misery and loss 
of a way of life. 

By 1904, bison were so scarce that surviving ani- 
mals were worth $1000 each. Michel Pablo sold his 
Montana bison herd to Elk Island and Buffalo 
National Parks in Alberta for $245 apiece. The dom- 
inant animal species in the largest biome of the con- 
tinent had disappeared. Isenberg concludes that the 
debacle of bison disappearance “exposed the fragili- 
ty of all societies ... that rely on the unsustainable 
exploitation of nature.” 

This thoroughly researched book is highly recom- 
mended. It provides a wide literature review, is com- 
plementary to Roe’s 1951 book, will be a resource 
for geography, history, or native studies courses, and 
iS an appropriate gift for anyone interested in the 
broad range of topics of bison, horses and native 
Americans. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


Prairie Birds: Fragile Splendor in the Great Plains 


Paul A. Johnsgard. 2001. University Press of Kansas. 
Lawrence, Kansas. 331 pp., illus. $29.95 U.S. 


Paul Johnsgard is the most prolific author in 
ornithology today, writing more than a book per 
year. He obviously enjoys the literature searches and 
the writing, a winning formula. He writes well, often 
with a rhapsodical, poetic flavour. Reminiscences of 
his personal experience with each species are espe- 
cially well done. His talents do not end there, for he 


is an artist as well; his sketches of each species are 
exquisite. The maps and tables are helpful, especial- 
ly data on breeding densities in tables 10 and 11. The 
blurb on the dust-cover of this book tells the truth: 
“written from the heart by a master of both birds and 
words.” 

No one is perfect. Johnsgard is reviled by a 
wildlife librarian of my acquaintance; when graduate 
students bring her lists of references containing erro- 


704 


neous years or volume or page numbers, or mis- 
spelled author’s names, she asks at once, “Which 
Johnsgard book did you get these from?” Perhaps his 
speed of preparation militates against careful check- 
ing and revision. In this volume, Borrer, Growe, 
Means, and Weins are misspellings for Borror, 
Grewe, Mearns, and Wiens. The chapter on furtive 
grass sparrows lists Knapton’s three important 
papers on Clay-colored Sparrows under both Grass- 
hopper and Vesper Sparrow, but not under Clay- 
colored Sparrow where they belong. David Duncan 
and Margaret Skeel’s data on Sprague’s Pipit num- 
bers in Saskatchewan are not cited in references or in 
acknowledgements. There are minor errors in several 
tables (e.g., the highest North American densities 
involve six not seven North Dakota species and four 
not five Saskatchewan species). An occasional read- 
er will be confused by the incorrect name of Breed- 
ing Bird Surveys, correctly used elsewhere, also 
being given to Breeding Bird Censuses (a study of a 
small area done several times each year). 

Johnsgard admits to being highly selective in his 
choice of references, but in general he has surveyed 


THE CANADIAN FIELD-NATURALIST 


Voly ids 


the literature well. Nevertheless, I detected occasion- 
al sweeping generalizations (“grassland soils are the 
most fertile on earth”), historical misrepresentations 
(the common or vernacular name of both the 
Franklin’s Gull and Wilson’s Phalarope derived 
from Dr. John Richardson’s 1820s Saskatchewan 
River specimens, even though both later lost their 
priority as “type specimens”), physiological misun- 
derstandings (owls cannot see in total darkness), 
habitat omissions (he fails to tell us that the 
Bobolink prefers wet meadows), and population 
oversights (no mention of the precipitous decline of 
Burrowing Owl populations in western Canada). 

More than for any other bird guild, a high propor- 
tion of grassland bird species are showing population 
declines. This fact alone makes it important for bird- 
ers and ecologists to understand these grassland 
species. Caveats aside, this readable and informative 
book is an excellent place to start. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


Ecology and Management of Large Mammals in North America 


Edited by Stephen Desmarais and Paul R. Krausman. 2000. 
Prentice-Hall, Upper Saddle River, New Jersey. 778 pp., 
illus. U.S. $59.95. 


The Wildlife Management Institute (WMI), based 
in Washington, DC, is a private, nonprofit, scientific, 
and educational organization committed to the con- 
servation, enhancement, and professional manage- 
ment of North America’s wildlife and other natural 
resources. In 1978, the WMI brought together a 
group of North American wildlife biologists to pro- 
duce a classic in the field of wildlife management: 
Big Game of North America — Ecology and Man- 
agement, compiled and edited by John Schmidt and 
Douglas Gilbert. This text, published by Stackpole 
Books (Emmaus, Pennsylvania), was standard issue 
through three print runs to students in Wildlife 
Management courses from the time it first appeared 
until it went out of print a few years ago. I first 
encountered the book as a Wildlife Technician in 
Labrador in the early 1980s, but then went on to read 
it cover-to-cover during a course in Management of 
Large Mammals at the University of Minnesota in 
1988. Schmidt and Gilbert’s Big Game was a com- 
pendium that strove to be the ‘authoritative volume’ 
on the status, management, and future of large mam- 
mals in North America. It certainly succeeded, and 
was deserving of the praise and awards that it has 
received over the past two decades. 

However, the science of wildlife management (or 


wildlife ecology, whichever you prefer) has pro- 
gressed considerably since 1978, and the status of 
many of our large mammals has changed. Even the 
name ‘big game’ has fallen out of use and has been 
replaced with ‘large mammals’ — for example, the 
course I took at Minnesota had been called Big 
Game Management until 1987. The WMI decided 
that too much of the information in Schmidt and 
Gilbert was dated, or even misleading, and did not 
proceed with a fourth printing, much to the conster- 
nation of those university lecturers and professors 
that taught wildlife management and ecology. 

Paul Krausman (Professor, Wildlife and Fisheries 
Ecology, University of Arizona) and Stephen Des- 
marais (Associate Professor, Wildlife Management, 
Mississippi State University), were among those uni- 
versity faculty that were frustrated by the “out of 
print” status of Schmidt and Gilbert’s text. The cur- 
rent WMI Vice-President, Richard McCabe, suggest- 
ed to Desmarais and Krausman that they spearhead a 
project to produce a replacement text for Big Game 
with a commercial publisher. Following Schmidt and 
Gilbert’s lead, Desmarais and Krausman solicited a 
select group of the continent’s experts on individual 
species or management-related topics that were will- 
ing to share their time and expertise by writing sepa- 
rate chapters. They provided this list, along with a 
proposal and book outline, to the WMI and received 
its wholehearted encouragement and endorsement. 


2001 


The result is Ecology and Management of Large 
Mammals and the editors have again succeeded in 
providing “a valuable textbook and reference for stu- 
dents and a record of contemporary practices for 
wildlife historians.” 

Ecology and Management of Large Mammals 
consists of 778 pages and 33 chapters written by 52 
contributors (including the two editors). It can be 
divided into two main sections: Chapters | to 15 
cover a variety of topics related to various aspects of 
large mammal ecology and management and 
Chapters 16 to 32 are detailed accounts of individual 
species native to North America, while Chapter 33 
covers exotic species. In contrast, Schmidt and 
Gilbert (1978) had 494 pages, 27 chapters and 2 
appendices, and was written by 28 authors. 

The topic chapters include: Taxonomy and Con- 
servation of Biodiversity, Human Values, Population 
Parameters and Estimation, Population Modelling, 
Nutritional Ecology, Carrying Capacity, Behaviour, 
Harvest Management, Genetics, Ranching, and 
Management on Tribal Lands. Chapters on Predators 
and Predator Control and on Habitat Management 
that were included in the 1978 book are noticeably 
absent from the new volume. The former is hardly 
surprising, given changes to the North American 
public’s view of predators and predation in natural 
systems, whereas the lack of an overview on man- 
agement of habitats does seem to be a considerable 
oversight on the editors’ part. After all, the founder 
of the discipline we now call wildlife management, 
Professor Aldo Leopold, pointed out that habitat 
management (i.e., control of food and cover) is the 
highest and most intensive of his five levels of 
wildlife management. 

Some of the first 15 chapters are more “cook- 
book” in nature and provide overviews of the meth- 
ods used by wiidlife biologists to obtain estimates of 
various population parameters, such as population 
size, age and sex ratios, mortality, etc. Much of this 
information can be found in other texts aimed at 
undergraduate and graduate students in wildlife and 
ecology. However, it is still valuable that these top- 
ics are included in this volume since so much of the 
efforts of wildlife biologists and managers across the 
continent are devoted to these areas. I found the 
chapters on Human Values, Human Dimensions and 
Conflict Resolution, Game Ranching, Management 
on Tribal Lands, and the history of large mammal 
management to be the most interesting, although the 
treatment of some of the subjects is rather sparse 
and, naturally, has the United States as the main 
focus. For example, Czech’s chapter on aboriginal 
peoples and wildlife management barely touches on 
the co-management regimes now in place across 
much of northern Canada and where he does write 


BOOK REVIEWS 


705 


about Canada, he refers to the Inuit as ‘Inuits.’ 
White’s chapter on ranching and sport hunting pro- 
vides much food-for-thought, with statements such 
as: “Big game farming has a bright future in 
Canada” providing a lively springboard for debate 
among both wildlife professionals and the general 
public! 

The individual species chapters are likely to be 
the most appealing part of the book for most people. 
The account for each species generally includes tax- 
onomy and distribution, life history, diet, behaviour, 
reproduction, population dynamics, habitat require- 
ments, and management. The chapters for some 
species, such as moose and white-tailed deer, are 
more detailed than others on jaguar and muskox, for 
example, simply because so much more work has 
been done on moose and deer. Again, there is an 
understandable lack of detail in the information pro- 
vided, since entire books have already been pub- 
lished on species such as moose, mountain sheep, 
white-tailed deer, and elk. The reference lists will 
point the student or reader who wishes to know more 
in the right direction to obtain information that could 
not be covered in each chapter. Overall, the chapter 
authors are to be commended for finding a balance 
between brevity and scope. 

One feature I particularly liked in the layout of the 
book is that references are provided numerically in 
parentheses, as opposed to the more familiar 
author(s) and year within parentheses. This reference 
format makes reading the text substantially easier 
and a complete list of references is provided at the 
end of each chapter. Figures and tables are clear and 
generally add to the text. An appendix lists all com- 
mon and scientific names of animals mentioned in 
the text and there is a comprehensive index. 

Despite its rather daunting cost, this book will 
take its proper place as a much-referred to text on 
the shelves of wildlife professionals across Canada, 
the United States, and Mexico. It should also find 
its way into the hands of keen naturalists who wish 
to have a comprehensive overview of our large 
mammal species and the whole business we call 
wildlife management on our continent. The editors, 
chapter authors, and the WMI are all to be com- 
mended for providing this volume that should see 
us through at least the next two decades, or until 
such time as it becomes apparent that another revi- 
sion is required. 


ALASDAIR M. VEITCH 


Supervisor, Wildlife Management (Sahtu Region), Depart- 
ment of Resources, Wildlife, and Economic Development, 
Government of the Northwest Territories, P.O. Box 130, 
Norman Wells, Northwest Territories XOE 0VO Canada 


706 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Handbook of the Birds of the World: Volume 6 Mousebirds to Hornbills 


Edited by Josep del Hoyo, Andrew Elliot, and Jordi 
Sargatal. 2001. Lynx Edicions, Barcelona, Spain. 589 
pp., illus. 


The Handbook of the Birds of the World (HBOW) 
series marches solidly on. The new volume, number 
six (HBOW6), covers mousebirds to hornbills. This 
grouping includes the mousebirds, an African family 
of grey or brown cardinal-like birds with long tails. 
Then follows the stunningly coloured trogons and 
kingfishers. The next group is the very similar 
todies. These five species of red-throated green birds 
are found on Caribbean islands. The book continues 
with the motmots and their cousins the bee-eaters 
and rollers. Ground-rollers and Cuckoo-roller of 
Madagascar are unique genera in that unique island. 
All by itself in the genus Upupa is the Hoopoe. The 
six species of African woodhoopoes are the least 
flamboyant birds in this book, somewhat overshad- 
owed by the hornbills who bring up the close of the 
book. 

What can one say about such a book? The editors 
set their high standards five volumes earlier, and vol- 
ume six maintains their pace. The species covered in 
this volume are among the most beautiful on earth. 
From the legendary Resplendent Quetzal to the 
graceful paradise-kingfishers you can expect to see 
some wonderful photographs and artwork. I searched 
thoroughly to find some errors, without success. If I 
did raise a question in my mind it generally revolved 
around the interpretation of a data set. For example, I 
checked many of the range maps and I did find some 
minor differences with my perceptions. Who was to 
say what is right? Birds move about and ranges 
change. 

The text is authoritative, well researched and writ- 
ten in a clear, uniform style. The systematics, mor- 
phology, habitat requirements, the habits including 
feeding and reproduction, and the current conserva- 
tion and status are described for each family. 

If you have not encountered the mousebirds 
before, try reading the introductory section to this 
family. While the six species have a certain physical 
charm they are not as glamorous as their closest rela- 
tives, the trogons. Their image also suffers too from 
their being fairly common, and therefore “ordinary.” 
Yet their habits set them apart from most other birds 
far more than brilliant colours. They are social ani- 


mals. They are the meerkats of the avian world. 
They stay in groups, bathe in groups, and cuddle 
together for warmth and companionship. One will 
stand guard when others feed: a duty they share in 
turn. There is a fascinating photograph of a Red- 
faced Mousebird laid on its back on a branch, being 
preened by a second bird. Happily these intriguing 
little birds are not at risk. Indeed, their numbers are 
expanding as a result of human activity. 

By contrast the flamboyant hornbills are much 
better known than the mousebirds. Their size and 
appearance have made them filmmaker’s favourites. 
These big bizarre birds appear on nature shows more 
often than the more striking bee-eaters and trogons. 
While hornbills are sometimes cooperative like 
mousebirds, there are species in which the female 
walls herself into solitary confinement for the nest- 
ing period. The text gives a thorough account of 
these traits. 

With each edition of HBOW the editors have 
included “Forewords” of increasing complexity. In 
HBOW6 the foreword is a 41-page essay on bioa- 
coustics and a tribute to the late Luis Baptista. My 
own awakening to bioacoustics came when a 
researcher played the “insignificant” song of a 
Henslow’s Sparrow (usually quoted as tslik). Played 
at one-eighth speed it is a complex and beautiful 
song. How do birds hear this? The foreword looks at 
a multiplicity of these types of questions and reveals 
how little we know about bird song. Indeed the mar- 
gin is filled with a sequence of boxes containing 
questions raised by researchers from around the 
world. The summary of what we do know, or at least 
think we know, is most intriguing. This essay is up 
to date, containing references to the work of 
Dabelsteen, McGregor and others using Automatic 
Location Systems. However it does not include the 
nocturnal migrant work using automatic acoustical 
transient detection or the new and exciting 
autonomous airborne monitoring system. The essay 
is written from the avian viewpoint, so it does not 
cover the difficulties we humans have in perceiving 
birdsong. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario K1J 6K5 Canada ; 


2001 


Threatened Birds of the World 


Edited by A. Stattersfield and D. Capper. 2001. Lynx 
Edicions and Birdlife International, Barcelona, Spain. 
xii + 852 pp., illus. $U.S.115. 


“Tt was the best of books, it was the worst of books, 
it was the doctrine of wisdom, it was the appreciation 
of foolishness.” (With full apologies to Charles 
Dickens.) This is certainly one of the best of books. It 
is perhaps the easiest-to-use reference book [ have 
ever read. It is the worst of books because it contains 
accounts of well over a thousand species at risk. There 
are an additional 700 or so near threatened species, 
meaning there are close to 2000 species for which we 
should be concerned. The authors show wisdom in 
clearly stating we are the solution to the problem. 
They give us an appreciation of the foolishness with 
which we have treated our planet. 

This book covers the species at risk from a global 
perspective and received input from a huge list of 
experts and collaborating organisations. As it 
responds to the global position, the Canadian list is 
only seven species long (Eskimo Curlew, Whooping 
Crane, Piping Plover, Mountain Plover, Marbled 
Murrelet, Sprague’s Pipit, and Bicknell’s Thrush). 
Some of the other species on the Committee on the 
Status of Endangered Wildlife in Canada’s 
(COSEWIC) Canadian Species at Risk list of Novem- 
ber 2000 are not included (for example Sage Grouse, 
Barn Owl, and Burrowing Owl). Obviously, while we 
are concerned over our heritage these birds are doing 
better elsewhere in the world. This list does clearly 
show where Canadian responsibilities lay. 

This book also uses a more complex ranking of 
species status. These include “Extinct in the Wild” 
but with survivors in captivity, and “Critically 
Endangered” where extinction could occur in the 
near future. Spix’s Macaw falls into the latter cate- 
gory with only one male in the wild (there are 60 
captive-bred birds). 

The species are shown two to a 30-cm X 21-cm 
(12 X 8 inch) page. There is a masthead with the 
species name in English and Latin and symbols to 
show its current status. There is a small 4 to 5cm 
portrait of each bird, generally a male in breeding 


BOOK REVIEWS 


707 


plumage, and a range map. A brief introductory note 
is followed by an even briefer identification note. 
Basic statistics (population estimate, current range, 
habitat, etc.) are placed in a yellow box for easy ref- 
erence. A more detailed discussion of range and pop- 
ulation, ecology, threats, and conservation follows. 
Finally the authors present the short range targets to 
be achieved if the birds are to receive help. Lower 
risk species are covered by a short discussion only. 

The authors have also summarized the status of 
species-at-risk by country, listing each bird according 
to their category. A world map to show the country 
location accompanies these summaries. I found this 
section very useful in providing a broad perspective. 
It is immediately obvious that the northern countries 
of South America (Brazil, Columbia, Ecuador, etc.) 
are the ones with the long lists of species at risk. The 
principle reason is loss of habitat. The book con- 
cludes with approximately 3000 references. 

The opening 30 pages contain a summary of the 
whole conservation issue and how to use the many 
features of this book. Beside the vignette illustrations 
of each species, there are several full-page illustra- 
tions. 

This book is well thought out, beautifully illustrat- 
ed with crisp text and clear maps. What I like most 
is the ease of use. It is simple to find a bird and 
rapidly get a good understanding of its status and 
the issues involved. It will be of great value to 
researchers and workers in the conservation move- 
ment throughout the world. It will encapsulate an 
important milestone in our knowledge and, I sincere- 
ly hope, can be used in the efforts to preserve our 
global heritage. Birds are not constrained by our 
political boundaries. Therefore understanding their 
status on a global scale is essential and should pro- 
mote cooperation between countries. We owe a 
round of applause to the authors and contributors for 
this important book. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario K1J 6K5 Canada 


Research Techniques in Animal Ecology, Controversies and Consequences 


Edited by Luigi Boitani and Todd K. Fuller. 2000. 
Columbia University Press, New York. 442 pp., illus. 


This is definitely a book for scientists. But it is 
also a book for naturalists interested in animal ecolo- 
gy and involved in wildlife conservation issues. 
Though the language and concepts can, in places, be 
intimidating for non-scientists, the greater part of the 
book is straightforward enough to provide excellent 


background to common issues and research tech- 
niques in animal ecology. 

The editors — Luigi Boitani (University of Rome) 
and Todd K. Fuller (University of Massachusetts) — 
reveal in the preface that the book grew out of persis- 
tent frustrations they felt in their teaching and 
research work: that scientific literature regarding 
ecology is full of false assumptions and methodologi- 


708 


cal errors. This is not surprising, they concede, given 
the enormous complexity of ecological systems. But 
they also point out that ecology is rooted in the scien- 
tific method applied to observation and experimenta- 
tion of natural facts, and that like any other discipline 
in the natural sciences, ecology can only benefit from 
scientific rigour. 

And so they helped organize a workshop in 1996 
which grew into this book: eleven chapters on such 
intriguing and diverse ecology research topics as 
hypothesis testing, effects of marking, delusions in 
habitat evaluation, GIS modelling of species distri- 
bution, estimating home ranges and territories, inves- 
tigating food habits, detecting stability and causes of 
change in population density, monitoring popula- 
tions, modelling predator-prey dynamics, analysing 
population viability, and measuring the dynamics of 
mammalian societies. 

Any attempt on my part to describe this material 
would not do it justice, so let me tell you which 
chapters I found the most enlightening. 

The first, “A Critical Review of the Effects of 
Marking on the Biology of Vertebrates” by Dennis 
L. Murray (University of Idaho) and Mark R. Fuller 
(USGS Forest and Rangeland Ecosystem Science 
Center) made me aware that data from marked indi- 
viduals (distinguished, for example, by tagging, 
mutilation, or radiotransmitters) may be biased if the 
effects of the markers on that animal’s life (mobility, 


The Sibley Guide to Birds 


By David Allen Sibley. The National Audubon Society. 
2000. Alfred A. Knopf, New York. 544 pp. illus. 


Very occasionally a guide appears that so clearly 
sets a new standard both in concept and execution 
that the potential user decides at once to acquire it. 
There is no need to look over reviews, and balance 
its worth against existing volumes: it is clearly supe- 
rior. This is such a book. 

In case there are some birders out there who have 
not had an opportunity to examine the Sibley guide, 
some basic facts will set the stage. It covers 810 
species, plus 350 regional forms, has over 6600 illus- 
trations, and at 16 X 24 X 3cm and over 1200 g is 
larger and heavier than any other North American 
guide, although no more unwieldy than, for example, 
the Panamanian guide, which I have carried around 
tucked into the back of my belt for days on end. 

The most innovative parts of the book are the 
individual species accounts. Here, instead of the tra- 
ditional layout with blocks of text on the page facing 
the plates, the two are combined. Typically up to 
four species are covered on each pair of facing 
' pages, with the images arranged in vertical columns. 
Similar plumages and postures appear in the same 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


feeding, courtship, etc.) are not detected or dealt 
with. If generalizations regarding unmarked individ- 
uals and populations, as well as conservation deci- 
sions, are based on possibly biased data, the conse- 
quences could be significant. ) 

From “Delusions in Habitat Evaluation’, by David 
L. Garshelis (Minnesota Department of Natural 
Resources), I learned that the concept of habitat — its 
availability, the use animals make of it, the prefer- 
ences they have, and the way these things are mea- 
sured, sometimes based on false assumptions — is 
much more complex than meets the eye. For exam- 
ple, the most valuable habitat is not necessarily the 
place where animals spend the most time, and the 
best habitat is not necessarily where the highest den- 
sities of animals are found. Habitat research built on 
problematic assumptions can have considerable 
repercussions if it is the basis for management deci- 
sions and habitat manipulations. 

These are the sorts of things naturalists should 
know. But they don’t come across in the more popu- 
lar, less scientific literature. That’s why, every so 
often, it’s worthwhile to work through a book like 
Research Techniques in Animal Ecology, Controver- 
sies and Consequences. 


R. SANDER-REGIER 


RRS5 Shawville, Quebec JOX 2Y0 Canada 


positions across the page. The top rows are usually 
devoted to the birds in flight, with images showing 
both upper and underwing. Then we find the birds 
seated, with young bird, adult female, and male posi- 
tioned one above the other. This approach also 
allows for great flexibility, and Sibley uses it to the 
full. On owls, for example, the young birds pictured 
are fledglings, and subsequent images may portray 
colour variations, as in Barn and Barred, different 
postures, as in Long-eared, rear and front views, as 
in Boreal and Great Gray, and age variation, as in 
Snowy. Where distinctive races occur, as in Great 
Horned, then more space can be allocated, and a full- 
page treatment portrays four different regional 
forms. The Red-tailed Hawk, with a huge range of 
variation, receives two full pages and over 39 differ- 
ent images, most of them of the bird in flight. 

The actual text is relatively limited. After a head- 
ing giving common and scientific names and mea- 
surements [including weight], there is a brief sum- 
mary statement on the species’ identification. Each 
account ends with a section on voice, with a small 
range map, and there is often an additional paragraph 
at the bottom of the page on some aspect of the 


2001 


species not covered by the preceding material. The 
identification criteria, however, are covered princi- 
pally by terse notes placed at appropriate points 
among the images themselves, usually with a line to 
indicate the feature being mentioned. 

There are other innovations. I liked the family 
summaries, giving an overview of the genera and 
species being covered, and the small images in this 
section usually show the less-striking plumages of 
the family; hence, female ducks and basic-plumaged 
shorebirds. Then each page has a heading naming 
the species group covered, with a couple of lines of 
general information about them. In some cases 
where the text is inadequate to cover some special 
aspect of a species or group there are short added 
sections, on such topics as the identification fo 
scaup, eiders and buteos, fall warblers, aerial dis- 
plays of snipe and woodcock, and so on. I’m very 
type-oriented, and there are times when I felt some 
nuance seemed to call for more text, but generally 
the approach works very well. 

There is a useful introduction, and bird topogra- 
phy is discussed and well-illustrated in an excellent 
seven page section showing shorebirds, gulls and 
duck, as well as the usual passerines, and including 
page on moult cycles. 

The book is not without its faults, although most 
are minor. However, perhaps the most striking prob- 
lem on first view is that some colour tints are inaccu- 
rate, with many of the russet shades too red and the 
grays too bluish. The latter is most serious on the 
gulls, and detracts from an otherwise excellent treat- 
ment. One can only hope that the publisher will cor- 
rect this in the next edition. 

Some difficulties are a product of the approach. 
The book is bulky, but after all, nowadays most field 
guides spend more time on a car seat than in a pocket. 
On the other hand, I have heard criticism that the 
images are too small. Indeed, many are small, but they 
do the job, and their abundance gives the user the lux- 


BOOK REVIEWS 


709 


ury of seeing a range of options from a difficult 
species, and not just one or two. There may be a few 
places where size might have been increased without 
loss of clarity, but on the whole space seems very well 
used. 

I was less satisfied with the approach to “regional 
populations”, as Sibley describes them. These are 
identified mainly by geographic terms, e.g. “West 
Taiga”’, or “Pacific”, on the basis that the scientific 
names can be more confusing than helpful. Perhaps, 
but the birder will encounter these names anyway, 
and Sibley has now introduced yet another set of 
terms [with attendant confusion] to describe these 
plumages. For example, last fall I saw a very pale 
Great Horned Owl which was quite unlike any of 
Sibley’s images, but very closely resembled the sub- 
arcticus illustration in another reference. Do I con- 
clude that this race is not shown in Sibley, or [more 
likely] that the striking difference was just normal 
variation? I’d welcome the scientific names in small 
print somewhere near the appropriate images. 

While quibbling, I'll note that some Canadian 
ranges are inaccurate [no winter Greater Scaup on the 
Great Lakes!], and it’s possible to second-guess the 
handling of extreme rarities [no Smew, for example]. 
There are also other errors here and there, and some 
statements I disagree with, but such things are 
inevitable in a guide of this scope. None of this serious- 
ly detracts from what is a masterful accomplishment. 
Overall this is an outstanding book that can and will be 
used constantly by novice and veteran alike. Even the 
most experienced birder will learn much from it, and 
its abundance of new illustrations will be of enormous 
value in expanding the scope of field identification. I’m 
just sorry I didn’t have it forty years ago! 


CLIVE E. GOODWIN 


1 Queen Street Suite 401, Cobourg, Ontario K9A 1M8 
Canada 


Ecotravellers' Wildlife Guide to Ecuador and its Galapagos Islands 


By D.L. Pearson and L. Beletsky. 1999. Academic Press, 
San Diego. xiii + 485 pp., illus. U.S. $27.95. 


I was very fortunate to have the opportunity to 
take a two-week birding tour to the eastern Andes of 
Ecuador, along with my wife, Valerie Wyatt, in Feb- 
ruary of this year. Despite finding out that a 
“National State of Emergency” had been declared 
the day before we had left, we had a wonderful time 
tooling around the peaceful countryside in our four- 
Wheel drive rental and would have been non- 
the-wiser of the “emergency” (mostly peaceful 
demonstrations) had we not read about it. 

Being a very avid birder, I sometimes forget to 


look at the “whole package” that nature offers. I was 
very pleased to be given a copy of the Ecotravellers’ 
Wildlife Guide to Ecuador and its Galapagos Islands 
before I departed, for the purpose of this review. 
Perusing this book before I left, reading more about 
the plants, animals and geography of Ecuador while 
I was there, and finally, when wishing I was still in 
Ecuador, instead of just pining to return, | was happy 
and eager to read more about the wildlife I had just 
recently seen. I can say, with out a doubt, that this 
book enhanced my experience. 

The book was written with the goal of providing 
the ecotourist one book to take along that would 


710 


allow field identification of the most commonly 
encountered plants and animals. It is a guide of 
breadth, not depth. The authors also wanted to give 
information on life history, behaviour and conserva- 
tion status. Unlike most field guides, they also 
endeavoured to make it a good read; this it is. 

Knowing more about the general landscape has 
many benefits. For example, instead of trying to 
point out the aracari, “in the tree...the TREE...the 
funny looking tree with umbrella like leaves” — we 
could say, “the aracari in the cecropia tree”. A much 
more accurate and pleasurable way to observe and 
one that leads to a deeper understanding of interac- 
tions in the environment. 

The book is split equally between the background 
information, 249 pages worth, and the identification 
plates, 96 colour plates in all. The text covers the 
geography and selected parks of Ecuador as well as 
natural histories of the amphibians, reptiles, birds, 
mammals, insects, and other arthropods with a sepa- 
rate section dealing with the Galapagos wildlife. The 
plates are nicely coloured and detailed with accom- 
panying text to aid in identification. Although defi- 
nitely not as bulky or awkward as carrying bird, 
plant, mammal, insect, reptile... guides, it is still a 
hefty book and too big to fit in any pocket. 

Species descriptions include icons that, at a glance, 
indicate if the species is found in the habitat region(s) 
you plan to visit. As a planning tool, you could 
decide to visit a particular region based on the plants 
or animals you would be likely to see. Of further aid 
to your travel plans, parks and reserves are listed for 
simple cross-reference in each habitat region. 

The book does not, and was not meant to stand 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


alone. As stated by the authors, a companion travel 
guide is essential so you will know how to get to the 
various reserves, what time of the year would be best 
to go and what to expect in terms of accommoda- 
tions and amenities. 

Although we only visited a few of the habitat 
regions, I found the selection of common species to 
be fairly representative, for our trip. However, with 
only 300 of Ecuador’s nearly 1600 species of birds 
(for example) represented, it may have been useful 
to list similar species with the descriptions with a 
quick explanation of the relevant difference(s) in 
appearance. An example would be the Golden- 
headed (pictured) and the very similar Crested 
Quetzal (not pictured); a simple mentioning of the 
white undertail (male) or barred tail (female) of the 
latter in the text would have identified two birds with 
one picture. This could have been done without 
adding extra pages as most of the descriptive pages 
facing the plates are well endowed with white space. 

If you are a serious nature observer, this is not the 
only book you will take. But as a supplement to fill 
in the gaps that all focused field guides leave, this is 
an excellent choice. The book provides a basic 
understanding of the ecology of Ecuador and the 
Galapagos in an enjoyable to read format. With a 
deeper understanding of the rich tapestry that makes 
up Ecuador, this book can help you to enhance what 
is almost sure to be a wonderful visit. 


PAUL GRANT 


Wild Birds Unlimited, 951 Gordon Street, Guelph, Ontario 
N1G 4S1 Canada 


Belize and Northern Guatemala: The Ecotravellers' Wildlife Guide 


By Les Beletsky. 1999. Academic Press, San Diego, 
California. 488 pp., illustrated. U.S. $27.95 


The purpose of this book is to both encourage 
“properly conducted” nature tourism in Belize and 
Northern Guatemala and to “provide ecotravellers 
with sufficient information to identify many common 
animal species and to learn about them and the fami- 
lies of animals to which they belong.” To these ends, 
Beletsky is abundantly successful. The first few chap- 
ters consist of a detailed description of habitats, tropi- 
cal ecology and terminology, major reserves, and the 
benefits of ecotourism, which provides an excellent 
base from which to start planning or preparing for a 
nature-oriented trip. The remaining bulk of the book 
focuses on meeting the second objective, in the form 
of a selective field guide. It includes 104 colour plates 
of the more common or spectacular species of birds, 
mammals, reptiles, amphibians, fish, coral, and jelly- 
fish and line drawings of plants, as well as tantalizing 


colour photographs of the main habitats. The text 
describes natural history, behaviour, status, descrip- 
tion, and interesting notes on folklore and trivia for 
each animal family. The overall result is an easy- 
reading resource that creates excitement for the coun- 
try before leaving on the trip and a better appreciation 
of the diversity of wildlife when there. I am not aware 
of any other book that fills such a niche for this geo- 
graphical region. 

Given the impossibility of including all species of 
wildlife, there is bound to be disagreement as to 
which species should have been included in the plates. 
I believe Beletsky has been as thorough as is feasible. 
Generally, the guide appears to include all the rela- 
tively abundant species as well as some Yucatan 
endemics, however, for more difficult groups of 
species the guide is less complete. For example, only 
one-quarter of the approximately 50 possible species 
of flycatchers are illustrated, whereas all but one of 


2001 


the seldom seen tinamous are shown, presumably 
because they are especially interesting. I had the 
opportunity to bring this guide along on a recent trip 
to Belize and Tikal, a popular Guatemalen day trip for 
visitors to Belize. While I, as a serious birdwatcher, 
was sure to bring along a more comprehensive field 
guide to the birds, carrying a book that exceeds one 
kilogram is not for everyone. We used the Beletsky 
guide (at less than half the size) to identify numerous 
mammals, fish, and a snake. 

The plates and text are well-done and accurate. 
However, I found some of the plates to be mislead- 
ing in terms of relative proportion. Some, such as 
plates 40, 46, and 64, portray very differently sized 
birds, next to one another and in equal dimensions. 
Instead, actual measurements in centimetres and 
inches, are given in the text opposite each plate. 

The book is attractive and appealing to naturalists 


BOOK REVIEWS 


711 


as well as less-experienced travellers. I made the 
mistake of taking this book out in the lunchroom at 
work, and was not able to retrieve it until everyone 
had a chance to flip through the plates and read the 
exotic names aloud. Although the experienced 
neotropical birdwatcher or researcher should consid- 
er this book as supplemental to more comprehensive 
resources, I would recommend this book as an excel- 
lent all-purpose field guide for the naturalist who 
wishes to be informed regarding the wildlife they are 
likely to see, as well as for the traveller who is inter- 
ested in getting off the beaten path and wonders 
about what might be beyond the beach. 


VALERIE WYATT 


Wild Birds Unlimited, 951 Gordon Street, Guelph, Ontario 
N1G 4S1 Canada 


Millions of Monarchs, Bunches of Beetles: How Bugs find Strength in Numbers 


By Gilbert Waldbauer. 2000. Harvard University Press, 
Cambridge, Massachusetts. 264 pp., illus. U.S. $24.95. 


Dr. Waldbauer will be well-known to many bird- 
watchers for one of his other books: “The Birder’s 
Bug Book’, which has helped to make birders aware 
that insects of the earth have functions other than pro- 
viding meals for birds. His current book delves deeper 
into the lives of those Bugs, and what fascinating lives 
they are. The ingenuity and adaptability of insects to 
enhance their lifespan, to ensure their reproduction, to 
avoid predators and outwit those they predate upon is 
wondrous. The text is scientific and readable. It can be 
readily understood by someone who is observant and 
has spent time in the field or garden, and will enhance 
an understanding of the scheme of nature — and you 
certainly wonder if there is not a scheme from the 
descriptions in this text. The interdependency of all 
living things is well illustrated. 

Separate chapters deal with the structures of the 
insect world such as: controlling the climate of their 
group homes — bees controlling hive temperature, 
for instance; defences; group living; and how insects 
find partners. One chapter describes the methods 
used to obtain and subdue food — they sting, mimic, 
trap, ambush, make group attacks, and build webs. If 
their target has defences, they devise strategies either 
to avoid them or to defeat rivals. Breeding is often 
timed to ensure that young insects will have suitable 
food available when they hatch. Four insects have 
chapters of their own — cicadas, tent caterpillars, 
mayflies, and ladybirds. If you have ever contem- 
plated using ladybirds as a biological control in your 
garden, don’t bother — they disperse away from the 
release site almost as soon as they are freed. 


The longest chapter is devoted to the Monarch 
butterfly and reveals results of new research. In 
recent years, scientists have learned a great deal 
from radio transmitters attached to Monarch wings 
— an attachment which is in itself an incredible feat. 
It is believed that on migration they navigate by the 
sun and a “magnetic compass” in the body, similar to 
birds. 

The butterflies which move south to Mexico or 
California in the fall die and it is their progeny which 
return north in spring, laying eggs en route. 

The hatched young continue north. Most of the 
males die en route. There are concerns about a new 
threat: genetically modified corn is dangerous to 
Monarchs since the pollen, which contains Bt, 
blows onto milkweed leaves and kills the caterpil- 
lars. It is now conceded that all efforts to preserve 
the overwintering sites of monarchs in Mexico have 
failed. The area is supposed to be a protected eco- 
logical site. Guards have failed to protect the sites 
from illegal logging, and at the same time have 
kept out film crews, tourists, and scientists from 
witnessing the logging. The Monarch wintering 
area is only 60 square miles, but even that cannot 
be safeguarded. 

The illustrations by Kathleen Brown-Wing are 
excellent. They are drawings in grey/white/black of 
magnified insects on their food or hatching plant. 
This is a fascinating book which will appeal to a 
wide audience. 


JANE E. ATKINSON 


255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 


gA2 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Behaviour and Conservation, Conservation Biology Series 2 


Edited by L. Morris Gosling and William J. Sutherland. 
2000. Cambridge University Press, London, UK. 438 
pp., illus. Cloth U.S. $90; paper U.S. $39.95. 


Number two in a series that links the advances 
made in behavioural ecology over the last 30 years 
with the relatively new discipline of conservation 
biology, this book will appeal to naturalists and sci- 
entists alike. It contains 19 papers by leading 
researchers working in particularly active areas of 
conservation biology. The first paper, written by the 
editors, outlines the advances in behavioural studies 
and their role in conservation. The remaining contri- 
butions are divided into the following four areas: 
conservation impact of people; habitat loss and frag- 
mentation; sexual selection, threats and population 
viability; and conservation applications of behaviour. 

“Life history characteristics and the conservation 
of migratory shorebirds,” co-written by Theunis 
Piersma, of the Netherlands Institute for Sea 
Research and Centre for Ecological and Evolu- 
tionary Studies at the University of Groningen, and 
Allan J. Barker, of the Centre for Biodiversity and 
Conservation Biology at the Royal Ontario Museum 
(the only Canadian contributor to the book), is par- 
ticularly interesting and easy to understand. It looks 
at various life history characteristics of shorebirds — 
from productivity, lifespan and gregariousness, to 
trophic specialization, immunospecialization, orien- 
tation mechanisms, and geographic bottlenecks — 
and what it all means in a world that is changing at 
an increasing rate. The authors finish by identifying 
pressing conservation issues and research needs. 


Turtle Conservation 


Edited by Michael W. Klemens. 2000. Smithsonian 
Institution Press, Washington. xv + 334 pp., illus., 
S57 7D: 


Turtles first evolved over 200 million years ago. 
They out-lived the dinosaurs, but there is increasing 
evidence that their future is bleak. Turtles’ slow-and- 
steady-wins-the-evolutionary-race (late maturing, 
high egg and juvenile mortality, but very low adult 
mortality) approach to life is now working against 
them. Michael Klemens, director of the Metropolitan 
Conservation Alliance for the Bronx Zoo-based 
Wildlife Conservation Society, along with seventeen 
other experts on turtles and/or conservation provide 
a comprehensive analysis of the current threats fac- 
ing turtles. 

Turtle Conservation consists of 10 peer-reviewed 
papers. The first three chapters focus on major 
threats: habitat alteration, human use of turtles, and 
disease. The next four chapters deal with the major 


“Controversy over behaviour and genetics in chee- 
tah conservation,” by Tim Caro, of the Department of 
Wildlife, Fish & Conservation Biology at the 
University of California at Davis, sheds new light on 
the environmental and genetic problems facing the 
world’s small cheetah populations. Caro starts by out- 
lining the genetic problems facing the cheetah, then 
goes on to examine misgivings over the genetic find- 
ings, review behavioural and demographic data in the 
wild, and discuss findings regarding juvenile mortality 
and disease among captive cheetah. It appears that the 
cheetah’s ecological problems are more serious than 
its genetic uniformity. 

“Retaining natural behaviour in captivity for re- 
introduction programmes,” by Michael P. Wallace, 
of the San Diego Zoo, is a good overview, along 
with a number of case histories, of the behavioural 
considerations (life history strategy, sociality, 
imprinting) and management techniques (enriched 
environment, rearing approaches and soft vs. hard 
releases) that must be thoroughly understood for 
captive-bred progeny to be successfully released into 
the wild. 

Scientists will be better able to understand the 
technical details of the more scientifically oriented 
papers, but naturalists who take the time to work 
their way through the book — at time labouriously, I 
must confess —- will find the knowledge they gain 
well worth the effort. 


R. SANDER-REGIER 


RR5 Shawville, Quebec JOX 2Y0 Canada 


ecological groups of turtles: sea turtles, river turtles, 
non-river freshwater turtles, and tortoises and other 
terrestrial turtles. The final three chapters examine 
various conservation actions: the use of genetics and 
demography, manipulating populations, and a sum- 
mary chapter on developing effective conservation 
Strategies. 

The chapter on human use of turtles is particularly 
staggering. Many species are being eaten into extinc- 
tion because of the growing demand in Asia for tur- 
tle meat. Vietnam alone exports over 200 O00 turtles 
per year (page 42)! It is not just Asian species which 
are at risk. The USA commonly exports more than 
30 000 softshell turtles a year, mainly to China and 
Japan (page 36). Those turtles which avoid the soup 
pot may still end up in the pet trade. Even a “devel- 
oped” nation like the US legally imports over 30 000 
turtles a year for the pet trade (page 56). This is par- 
ticularly troubling when most adults sold in the pet 


2001 


trade are wild-caught individuals. The level of illegal 
trade is unknown, but likely far greater than the legal 
amount. 

Is this a perfect book? No. Although the focus of 
the book is global all the authors are American. In 
fairness, some of the authors have international expe- 
rience, particularly Edward Moll who has worked on 
turtle conservation in Asia for over 20 years, and the 
book has many non-US case studies. Another oddity 
is that one-third of the authors all work for the 
Wildlife Conservation Society. A greater diversity of 
voices would have been beneficial. _ 

In terms of content, all the papers are strong, but 
there are occasional weaknesses. The chapter on dis- 
eases suggests that chemical contaminants have been 
found in “at least one species” of turtle (page 86). 
This vague statement is just one example of less than 
exhaustive review of the existing literature on a par- 
ticular subject. The chapter on demography and 
genetics rightly stresses the importance of carefully 
examining the age structure of a population before 
determining whether or not the population is viable. 
However, there is no mention whatsoever of the 
growing evidence that plastral growth lines can dra- 
matically underestimate age. There is also virtually 
no mention anywhere in the book of the importance 
of hibernacula as critical habitat. Many species over- 
winter communally, possibly because optimal over- 
wintering habitat is limited. Granted this may be a 


BOOK REVIEWS 


713 


temperate zone bias of my own, but it is relevant for 
many species around the world. 

Despite these minor complaints this is a definitive 
guide to turtle conservation. Klemens’ final chapter, 
“From information to action: developing more effec- 
tive strategies to conserve turtles” is clear, insightful, 
and thorough. He makes a number of key recom- 
mendations, at the heart of which is controlling 
exploitation of turtles. A global moratorium on the 
turtle trade should be considered before species are 
lost. Ultimately, though, successful conservation of 
turtles (or other species) must be focused at the 
ecosystem or landscape level, not just the population 
level. Klemens provides the example of the Great 
Swamp in New York’s Harlem Valley as an example 
of effective turtle conservation. He has involved 
local politicians in biological survey work. For these 
people, environmental rhetoric is replaced with the 
personal experience of seeing turtles moving far 
from wetlands into upland forests. This has resulted 
in growing interest in protecting the wetlands with 
large, meaningful buffer zones. If Klemens’ book 
can encourage such actions elsewhere around the 
world it will definitely have served its purpose. 


DAVID SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, 
Oxford Mills, Ontario KOG 1S0 Canada 


The Pheasants of the World: Biology and Natural History 


By Paul A Johnsgard. 2000. Second edition, Smithsonian 
Institution Press, Washington D.C. xvii + 398 pp., illus. 
U.S. $50. 


Paul Johnsgard is a prolific author of bird books. 
He has made a great contribution to the birding 
world by writing books on groups of birds such as 
Cranes, Grouse, Hummingbirds and so on. Indeed, 
about half of his vast output has been of this type of 
generic book. Pheasants follows in this tradition, 
covering 49 species of pheasants, jungle fowl, 
peafowl (Phasianini) and other large, colourful 
species most often thought of as pheasants. This 
book does not cover the smaller members of the 
pheasant family (Phasianidae) such as quail, grouse, 
ptarmigan, francolin, and snowcock. The English 
names used by the author may not conform to those 
used in other well-known guides, but there is a list of 
alternative names for each species. 

The format is similar to many of his other books. 
He has collected all the information he could on each 
species, and massaged it into a logical format. In this 
case, this is a second edition and incorporates infor- 
mation from a database that has increased forty per- 


cent since the original publication. Distribution maps 
and drawings have also changed substantially. The 
author has dropped the detailed plumage description 
of the first edition, feeling that this information is 
fully covered elsewhere. 

The book is divided into two parts. The first twen- 
ty percent contains a generic description of pheasant 
family biology. This includes chapters on reproduc- 
tion, distribution and behaviour, plus one chapter on 
aviculture — or more correctly the role of raised 
pheasants in conservation. (However, details of rear- 
ing pheasants are not given as these are more fully 
covered in books dedicated to aviculture.) 

The second section gives the individual species 
accounts. In all, 49 species are covered by a detailed 
description of range, subspecies and variation, 
behaviour, ecology, and biology. The author also 
summarizes the current status and conservation out- 
look. For, despite mankind’s fascination with these 
exotic birds, almost seventy percent are in some dan- 
ger and there is still much that is not known. About 
forty-five of the species accounts have a range map 
showing distribution, while the rest are covered by a 
text description of their range. Not suprisingly the 


714 


most extensive account is that of the Ring-necked 
Pheasant. The information on the other species is 
variable. The author has made these accounts as 
detailed as possible, within the limits posed by the 
existing data. There is a short text on identification, 
but more space is given to the ecology, biology and 
social behaviour of these birds. The accounts finish 
with a synopsis of status and conservation. 

A colour photograph of the male (both sexes in 
the case of the Great Argus) illustrates most of the 
species. As most of the specimens appear to be in a 
zoo-like setting the photographs and the birds are in 
top shape. A number of additional illustrations are 
taken from charming paintings by Joseph Wolfe, 
executed in the 1870s. In this case, both the male and 
female are shown. Display postures and details of 
sub-species are shown by black-and-white line draw- 
ings made by the author. 

I think it is important that someone does what this 
author does; that is he collates disperse and disparate 


The Biology of Plethodontid Salamanders 


Edited by Richard C. Bruce, Robert G. Jaeger, and Lynne 
D. Houck. 2000. Kluwer Academic/Plenum Publishers, 
New York. xiii + 485 pp., illus. US$195. 


Plethodontid (or lungless) salamanders make up 
the largest family of salmanders with some 350 or 
more species. These fascinating creatures have been 
the focus of many studies. The fourth “Conference 
on the Biology of Plethodontid Salamanders” was 
held in June 1998 at the Highlands Biological 
Station in North Carolina. Many of the papers first 
presented at that conference are collected together in 
this book. This is the first time that a formal pro- 
ceedings has been published as a result of the confer- 
ences. 

The book is divided into three sections. Part 1 
consists of papers presented at a symposium at the 
conference held in honour of Richard Highton on his 
retirement. Highton’s pioneering work on molecular 
systematics and speciation in plethodontid salaman- 
ders stretches back over 40 years. The symposium 
was entitled “Points of View on Defining and 
Naming Species of Plethodontid Salamanders.” Five 
invited papers tackle these themes in a variety of dif- 
ferent species groups. Highton himself has a paper 
on the Plethodon jordani and P. glutinosus complex- 
es which clearly illustrates the difficulty in delineat- 
ing species. As recently as the early 1980s both P. 
jJordani and P. glutinosus were each considered to be 
a single variable species, yet Highton provides 
detailed evidence on geographic protein variation 
that both “species” may actually consist of up to 14 
species in total. Part 1 concludes with a summary 
paper (not presented at the conference) on “Syste- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


data into a single coherent volume. I do not have 
access to the 500 references used by Jonhsgard, nor 
do I have the time to sift through and select the salient 
information. This author performs these tasks and 
does them well. I suspect it will not attract a huge 
audience. Pheasant breeders will want more depth 
(and will use the actual references) and birders will 
need less (and will be content with field guides). I 
hope there are enough readers in the middle who are 
willing to buy this book and encourage this prolific 
author to continue. Certainly it is a fine book for any- 
one interested in the birds themselves, and not just 
simply identifying them in the fieid. It is a good 
resource for students as well as a sound addition to 
any library. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario K1J 6K5 Canada 


matics at the turn of a century” by Stevan Arnold. It 
highlights some of the conceptual (and philosophi- 
cal) controversies in systematics and suggests direc- 
tions research may take in the 21st century to resolve 
some of these dilemmas. Debates on what constitutes 
a valid species are not new to systematics and new 
types of data, regarding allozymes and mtDNA 
sequences have not been able to resolve problems. 
Ultimately, it must be acknowledged that defining 
species and species boundaries 1s subjective, for 
species themselves are but “ephemeral fragments of 
a grand evolutionary continuum” (page 95). 

Part 2 — “Evolutionary and Comparative Biology 
of Plethodontids” — contains eight papers on a 
broad range of topics. A number of the papers tackle 
the entire family of plethodontid salamanders, or 
large subgroups. Sexual size dimorphism is critically 
reviewed as is courtship and its relationship to form, 
function and phylogeny. The life history evolution 
and adaptive radiation of the hemdactyliine salaman- 
ders is also examined. This small group of approxi- 
mately 25 species continues to offer puzzles to sys- 
temacists. Is the group monophyletic? If so it 
exhibits more life history variation than any other 
comparable vertebrate group. Other papers focus on 
single species, for example, maternal behaviour of 
Desmognathus ocoee and population cytogenetics of 
Eurycea wilderae. 

The third part of the book focuses on the 
“Behavioral Ecology of Small Plethodons.” Most of 
these eight papers deal with the Redback 
Salamander, Plethodon cinereus, undoubtedly one 
of the best studied vertebrate species. A variety of 


2001 


topics are explored including kin recognition, 
pheromonal attraction, and alternative mating strate- 
gies. 

Despite the title of this book, the focus is primari- 
ly phylogeny and evolutionary history. There is 
growing evidence that salamanders, as well as anu- 
rans, are a significant part of the global amphibian 
decline, yet reading this book one is struck by the 
lack of papers on conservation biology. The token 
exception is a paper on using artificial coverboards 
to monitor Redback Salamanders. 

The hefty price tag on this book makes one won- 
der whether publishing a conference proceedings 


Birds of Europe 


By Lars Svensson and Peter J. Grant, illustrations by 
Killian Mullarney and Dan Zetterstrom. 1999. Princeton 
Field Guides, Princeton University Press, Princeton, 
New Jersey. 400 pp., illus. Cloth U.S. $39.50; paper 
PS 5h29.95, 


Another bird guide for an area already blessed 
with several excellent volumes may seem hardly 
worthy of comment, let alone review. But this is a 
field guide with a difference: conceived by the late 
Peter Grant and Killian Mullarney, and sixteen years 
in the making, it is arguably the best field guide ever 
produced. It certainly is by far the best I have ever 
seen, and it now sets a standard that must be met by 
future guides on both sides of the Atlantic. 

What is so exceptional about it? I compared it 
with two leading guides, one from North America, 
the National Geographic Guide, and the other from 
Europe, Lars Jonsson’s excellent Birds of Europe. 
The latter has been my own choice for a European 
guide, and I used it extensively on a recent trip to 
Greece. Hence I had ample opportunity to compare it 
with the current volume, which proved to be the 
guide of choice with all my British companions! 

All three books are about the same size and for- 
mat, although the Princeton Guide [PFG] is the 
shortest of the three, the result of a conscious deci- 
sion by the authors. To cover 848 species in a book 
of this size demands both a small typeface and small 
images on the plates — I need reading glasses to use 
it — but these initial drawbacks are quickly over- 
come by the remarkable quality of both text and 
plates. 

Comparing the three volumes in terms of the 
length of each entry and the number of images, I 
selected some species that occur reasonably often on 
both sides of the Atlantic. In general, PFG has more 
text and more images for each species than either of 
the other two guides, and in some cases more than 
double. 

This, however, was just the beginning. Those 


BOOK REVIEWS 


tS 


was a good idea. The cost will surely prevent many 
university libraries, not to mention most individuals, 
from acquiring the book. The Highton symposium 
could have been published as a special section of any 
number of journals thus preserving the group 
dynamic. Nonetheless, the papers in this volume are 
first rate and it is an important contribution to our 
understanding of the lungless salamanders. 


DAVID SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, 
Oxford Mills, Ontario KOG 1S0 Canada 


small images are consistently superbly executed, 
with short notes appended throughout to emphasize 
key points. One feature I particularly enjoyed was 
the use of numerous small vignettes to illustrate 
key features of behaviour. Thus, a Northern Fulmar 
floating on a cliff updraft, and another series of 
three birds over the sea illustrating “gliding on 
long stiff wings”. The plate for this species also 
includes images of dark and light morph birds seat- 
ed on the water, three [light, intermediate, and 
dark] in flight, upper surfaces, and a single bird 
undersurface; plus another vignette of a bird nest- 
ing on a cliff-face. Not all species receive quite as 
extensive a series of illustrations, but this is fairly 
typical for the most widespread. As there are 
between two and six species featured on each 
plate, these inevitably appear cluttered — until one 
comes to use them, when one finds a significant 
degree of consistency in the layout of each page. 
This allows ready comparison of the species 
grouped there, and one finds neat hairlines separat- 
ing the images of different species, while dashed 
lines separate different races. 

The text itself is particularly concise and well- 
organized, with intelligent use of italic to highlight 
important features. One has the sense it is written by 
someone who really knows the species being dis- 
cussed: for example, under Short-eared Owl we are 
told that “In certain lights e.g. at dusk or overcast 
weather, can look surprisingly whitish”, while under 
Tawny Owl [a dark-eyed species] one is warned that 
“. . yellow-eyed species e.g. Long-eared Owl, can 
look dark-eyed at night in headlights owing to large 
pupils”. 

The introductory sections to various families or 
groups are another valuable innovation, additional to 
the usual section in the Introduction to the book. 
There is half a page of hints on “Watching Sea- 
birds”, a page on “Birds of Prey”, while shorebirds 
[“Waders”’] and Gulls each receive two pages with 


716 


useful information on molts, and the gulls section 
has an excellent plate illustrating the sequences of 
gull plumage over four years. On the other hand, the 
page for introduction for the Sylviidae Warblers cov- 
ers the differences between the five major genera, a 
potential source of considerable confusion to the 
beginner. 

In the past, one of the difficulties with field guides 
has been that there has been a dichotomy between 
the needs of the novice and those of the veteran. I do 
not think this is the case here: it is indeed most 
accessible to a beginner, but also answers the 
demands of the experienced birder. 

I could go on, but the reader will have the pic- 
ture. This is a superb guide, meticulously thought- 


Hoofed Mammals of British Columbia 


By David Shackleton. 1999. University of British 
Columbia Press and Royal British Columbia Museum, 
Vancouver. 268 pp., illus. $24.95. 


The Royal British Columbia Museum (RBCM) 
and the University of British Columbia (UBC) are 
co-publishing a series of six handbooks that replace 
the now out-of-print The Mammals of British 
Columbia (Cowan and Guiget 1965). Shackleton’s 
volume on our westernmost province’s hoofed mam- 
mals, or ungulates, is the third of this series. Since the 
first two handbooks on bats and on oppossums, 
shrews, and moles appeared in 1993 and 1996, 
respectively, it appears that will take almost another 
decade to see the series’ completion. However, after 
reading Shackleton’s treatment of British Columbia’s 
ungulates (a major component of any jurisdiction’s 
“charismatic megafauna’’) I’d suggest to those ama- 
teur naturalists with a keen interest in the mammals 
of western Canada that it will be worth the rather 
lengthy wait. 

In 1993, Brad Stelfox edited the excellent Hoofed 
Mammals of Alberta with chapters written by a col- 
lection of that province’s leading mammalogists 
each of whom addressed their own particular spe- 
cialty. In my office, Stelfox’s book remains a much- 
used reference for its well-researched chapters that 
examine all aspects of the ecology of ungulates. But, 
as a handbook to throw into a backpack or vehicle 
when heading out on a trip, it is not exactly suitable 
given its hard cover and size (22 X 29 cm). In con- 
trast, Hoofed Mammals of British Columbia is much 
more field friendly with its well-bound softcover and 
compact size (14 X 22 cm). It is also written for a 
quite different target audience with none of the dis- 
cussions on hunting, economics, and game ranching 
that are central to the Alberta book. The RBCM and 
‘UBC Press have obviously targeted this series at 
weekend to serious amateur naturalists who want a 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


out, clearly and knowledgeably written, with the 
most consistently satisfying plates of any guide I 
have ever used. True, there is a minor glitch here 
and there [and I find some of the sound descriptions 
hard to come to terms with] but overall it’s difficult 
to imagine how it could be improved! It will now 
be the essential guide for birding in Europe, and 
while I would not normally advocate such a book 
for North American use, perhaps this is an excep- 
tion. This book is sheer delight. Get it! 


CLIVE E. GOODWIN 


1 Queen Street Suite 401, Cobourg, Ontario K9A 1M8 
Canada 


well-written overview of the general biology of the 
particular mammal group with specific information 
on individual species and subspecies. 

The first 43 pages of this handbook cover General | 
Biology and include brief discussions on such topics 
as the evolution of ungulates, group living, food 
habits, mineral requirements, social behaviour, and 
reproduction. This is all done in a light and easy-to- 
read manner accompanied by 27 black-and-white 
pictures and diagrams. More serious students of nat- 
ural history will no doubt often find themselves 
wanting more information than Shackleton provides; 
however, I believe that he has done an admirable job 
in giving just enough to answer some of readers’ 
questions (and likely leading to a few more) without 
getting overly bogged down in detail. To assist the 
more adventurous he provides a few selected refer- 
ences that range from older “classics” by writers 
such as Ian McTaggart Cowan and Val Geist to 
recent investigations of the genetic relationships 
between subspecies of caribou as determined by 
DNA techniques. 

The next section looks at the province’s 19 ungu- 
lates in more general terms, including their presence 
or absence of each within each of British Columbia’s 
nine ecoprovinces and 14 biogeoclimatic zones. The 
inside back cover of the book includes a fold-out 
colour map of the biogeoclimatic zones. Unfor- 
tunately, readers have to discover this for themselves 
and I did not realize this until well after I had read 
through this section. The section also very briefly 
discusses exotic ungulates, a history and general sta- 
tus of management and conservation, and some of 
the techniques used by biologists to study ungulates 
in British Columbia. Also provided are a checklist, 
which includes extinct, non-native, and feral domes- 
tic species and subspecies, an identification key, and 
a key to skulls. The latter is particularly useful for 


2001 


use in the field when skulls or skull fragments are 
encountered during a wilderness expedition. 

The main portion of the book is devoted to indi- 
vidual accounts for the province’s eight native 
species of ungulates (with 18 subspecies) and one 
introduced species. Each account gives other com- 
mon names of the species or subspecies, a descrip- 
tion, information on its natural history, range, con- 
servation status, and traditional aboriginal use within 
the province. The accounts are well written and 
informative and [| particularly like the maps, which 
show relative density across the province (plentiful, 
moderate, few, absent) based on information provid- 
ed by regional wildlife biologists. The descriptions 
include body measurements (all in millimetres) 
based on collections within the province. For a book 
intended for a lay audience, particularly for older 
readers and those coming from the United States, it 
would have been beneficial if non-metric measure- 
ments had also been provided. 


Swallow Summer 


By Charles R. Brown. 1998. University of Nebraska Press, 
Lincoln, Nebraska. 371 pp., illus. U.S.$16.95. 


“This is a book about why I love to do research” are 
the first words of the preface, and Charles Brown’s 
love and enthusiasm for research permeate the book. 
For fifteen summers Dr. Brown, with his wife and 
graduate students, studied Cliff swallows in the area 
around Cedar Point Biological Station in western 
Nebraska. The station is close to the Platte River and 
descriptions of the countryside are glowing. His studies 
focused on nesting habits, life span and migration pat- 
terns, and resulted in the publication of “Coloniality in 
the Cliff Swallow: The Effect of Group Size on Social 
Behavior’, in 1996 by University of Chicago Press. 

Swallow Summer is written in the form of a day-to- 
day diary for the 1995 summer season, but includes 
some of the findings from all the years of research in a 
less academic format than the 1996 book. It gives an 
insight into the daily highs and lows of field research, 
the hard slog of routine data gathering, and the 
dynamics within the team involved in collecting, 
banding, and recording weights and measurements of 
up to 900 birds a day. The author describes the daily 
doings in an interesting way, and readers will look 
forward to reading about the next day’s activities. The 
Cliff Swallow colonies were almost all found in large 
culverts and most had been nesting colonies for at 
least 15 years. Working at the mouths of these cul- 
verts to set up mist nets was not pleasant — the 
stream waters were polluted with farm runoff. 

The summer climate in Nebraska produces 
extreme heat (100-plus deg. F.) and occasional vio- 
lent wind and thunder storms, all of them potentially 
dangerous and curtailing bird collection. Bad weather 


BOOK REVIEWS 


ely, 


The book ends with a series of three appendices 
of scientific names of plants and animals used, esti- 
mated numbers of each subspecies of ungulate in the 
province, and a summary of gestation period, weight 
at birth, and approximate date-of-birth for 10 
species. An excellent glossary, lists of general and 
specific references, and an index are also provided. 

Overall, this is a well researched, well written, 
and nicely produced book that fulfills its objective. I 
have no reservations in recommending it for every- 
one with an interest in the large mammals of western 
Canada and I will definitely have it along with me on 
my next visit to British Columbia. This book, along 
with the five others in the series, are and will be a 
significant contribution to Canadian mammalogy. 


ALASDAIR M. VEITCH 


P.O. Box 102, Norman Wells, Northwest Territories 
XOE OVO Canada 


was a hindrance, but at the end of the season, it was 
considered successful and knowledge about Cliff 
Swallow nesting and life histories had advanced. 
Three graduate students had also learned a great deal 
— about themselves, about working in a team and 
just what field research was all about. 

Some questions about colony size were not 
answered that summer. Questions such as why are 
some colonies smaller than others established in a 
similar space. Are breeding colonies formed to act 
only as information centres to communicate news 
about food sources? Are the “inspection tours” by 
non-breeding birds in late summer solely for picking 
out a nesting site for the following year? What is the 
effect of the parasitic swallow bug on breeding suc- 
cess? Among the discoveries were three colonies in 
new sites with birds from old colonies; four different 
birds were seen carrying an egg from one nest to 
another and a captured swallow had been banded 11 
years previously — the highest age recorded for a 
Cliff Swallow. One half-serious conclusion Dr. 
Brown draws 1s a parallel with human “colonies”: 
swallows can be competitive, co-operative, mean, 
insecure but very social birds. 

This is recommended reading for any naturalist, 
birdwatcher, and in particular for a biology student 
who might want to find out what field research 
entails. Judging by his enthusiasm and devotion to 
Cliff Swallows, Dr. Brown is probably still studying 
them every summer in Nebraska and taking more 
first-rate photographs. 


JANE E. ATKINSON 


255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 


718 


BOTANY 


Flora of New Brunswick 


By Harold R. Hinds. 2000. Second Edition. Department of 
Biology, Bag Service #45111, University of New Bruns- 
wick, Fredericton, New Brunswick E3B 6E1. 699 pp., 
illus. $50.00 + $8.00 shipping and handling. 


This volume is a tremendous jump ahead of the 
first edition which was published by the author in 
1986. In the introductory area is a map of New 
Brunswick depicting the counties and major rivers, a 
Foreward by T. G. Dilworth, Acknowledgements, a 
Glossary with many illustrations, an Introduction, 
How to use this book, Abbreviations, signs and sym- 
bols, Rarity rankings, Common Names (sources), 
Mi’kmagq and Mailisect names, Chromosome counts, 
Notes: folklore, toxicity and other information, 
Specific epithets, History of plant collecting by C. 
Mary Young (5 pages), and History, physical setting, 
and regional variation of the flora by S.R. Clayden (39 
pages). This contribution by Claden includes informa- 
tion on bedrock assembly, earliest plant life, modern- 
ization of the landscape and flora, glaciations and 
their aftermath, the present environment, ecoregions 
and their floras, acknowledgements, and references. 

The main text starts on page 75 which begins with 
a Key to Families. This is followed by the Ferns and 
Fern Allies, Division: Phiophyta (Gymnosperms), 
Flowering Plants (Dicotyledons), and Flowering 
Plants (Monocotyledons). All taxa are keyed with 
numbered unindented keys. The maps and illustra- 
tions which could be found with some difficulty at 
the end of the 1986 volume are now conveniently 
displayed on either side of the text of most individu- 
al species, although rare species are not mapped and 
some species are lacking an illustration. Text of 
some species that are of particular interest is high- 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


lighted. The work is completed by a Bibliography, 
An Appendix with additions to the Flora of New 
Brunswick since the publication of the 1986 volume, 
Exclusions from the Flora of New Brunswick, Plants 
believed to be Extirpated from New Brunswick, and 
Plants found in neighbouring provinces/states to 
Look for in New Brunswick, a Summary of Taxa, 
Nomenclatura! Innovations, and Addenda followed 
by the index. Also included were four sheets of 
paper with additional minor corrections dated 19 
April 2001. 

In the Acknowledgements thanks were given to 
George Argus (Salicaceae), Jim Goltz (Orchi- 
daceae), Jerrold Davis (Puccinellia), Tony Reznicek 
(Carex), Don Britton and Dan Brunton (Isoetes), 
Pierre Taschereau (Atriplex), the late Herb Wagner 
(Ferns and fern allies), and Arthur Haines and Tom 
Vining (for permission to rework various taxa from 
their Flora of Maine). Work for this revised edition 
was sponsored by the New Brunswick Department 
of Natural Resources Environmental Trust Fund, the 
New Brunswick Wildlife Trust Fund, Parks Canada, 
and several corporations and businesses as well as 
the commitment to time and labour by many individ- 
uals. Hal Hinds worked hard to complete this second 
edition and fortunately saw it in print before he 
passed away 9 May 2001. 


WILLIAM J. CODY 


Biological Resources Program, Eastern Cereal and Oilseed 
Research Centre, Agriculture and Agri-Food Canada, Wm. 
Saunders Building, Central Experimental Farm, Ottawa, 
Ontario K1A 0C6 Canada 


Flora of Florida, Volume 1, Pteridophytes and Gymnosperms 


By Richard P. Wunderlin and Bruce F. Hansen. 2000. 
University Press of Florida, Gainesville, Florida. xii + 
366 pp., illus. U.S. $49.95. 


This is the first volume of a series of eight that will 
encompass the flora of Florida: Pteridophytes and 
Gymnosperms (1), Dicot families (2 to 6) and Monocot 
families (7 and 8). Because it is the first of the series, in 
addition to Acknowledgements, there is an Intro- 
duction which includes the Historical Background, the 
Organization of the Flora, and Taxonomic Concepts. 
This is followed by a description of the Physical 
Setting (Physiography, Geology, Soils, Climate, and 
Fire) by Ronald L. Myers (10 pages) and Vegetation of 
Florida (Upland Communities, Wetland Communities, 
‘and Aquatic Communities) by Ronald L. Myers (15 
pages), and Botanical Exploration in Florida by 


Richard P. Wunderlin, Bruce F. Hansen, and John 
Beckner (66 pages). All of the above is most useful 
background information for anyone interested in the 
vegetation of this state which houses over 4000 taxa. 

A key to the Major Vascular Plant Groups: Pteri- 
dophytes, Gymnosperms, Dicotyledons, and Mono- 
cotyledons can be found on page 106. This is fol- 
lowed by Pteridophytes (Pteridophyta) authored by 
Nauman, Wunderlin and Bruce F. Hansen (pages 
101-300) and Gymnosperms authored by Wunderlin 
and Hansen (pages 301-328), an Appendix which 
presents a synopsis of the families (28), total genera 
(70), species (170), native genera (58), native species 
(127), endemic genera (0), and endemic species (1), 
Literature Cited, General Index, Index of Common 
Names, and Index of Scientific Names. 


2001 


Throughout this main part of the volume there is a 
wealth of information: excellent keys, scientific names 
(with an English translation of the Latin species 
name), common names, synonomy, publication data 
for all taxa, detailed descriptions of families, genera 
and species, habitats, known distribution in the state, 
overall distribution in North America, and if non- 
native, where it came from. In addition, interesting and 
useful comments are found throughout the text and an 
excellent page drawing is provided for one species of 


BOOK REVIEWS 


719 


each genus. Now we can hope that the remaining 
seven volumes will soon be available for anyone inter- 
ested in the vegetation of this fortunate state. 


WILLIAM J. Coby 


Biological Resources Program, Eastern Cereal and Oilseed 
Research Centre, Agriculture and Agri-Food Canada, Wm. 
Saunders Building, Central Experimental Farm, Ottawa, 
Ontario K1A 0C6 Canada 


Plant Life Histories: Ecology, Phylogeny and Evolution 


Edited by Jonathan Silvertown, Miguel Franco, and John L. 
Harper. 1997. Cambridge University Press, New York. 
xviii + 313 pp., illus. U.S. $29.95. 


“Comparative ecology” explores the relationships 
between species traits and environmental character- 
istics in order to generate hypotheses and evaluate 
theoretical expectations regarding the evolution of 
adaptive traits. Many biologists spend careers deter- 
mining which traits are correlated with one another, 
and then trying to decide if correlated traits are the 
result of common descent or environmental adapta- 
tion. But scientists need to beware of the risk of phy- 
logenetic constraints being interpreted as adaptive 
traits. This book (apparently the proceedings of a 
Royal Society Symposium) addresses a weakness in 
this field of comparative biology, in that standard 
statistical analysis of trait correlations may be com- 
promised because each species is not a truly inde- 
pendent observation. That is, species differ in the 
degree to which they are phylogenetically related, 
and hence this limits the inferences possible about 
ecological adaptation and evolutionary trade-offs. 

Chapters are grouped into five sections: phyloge- 
netic perspectives; reproductive traits; seed biology; 
recruitment and growth; and interspecific interac- 
tions. In the book’s preface, the editors point out that 
despite the weaknesses of earlier studies, some cor- 
relations and trade-offs in plant life histories seem to 
be robust across a variety of plant families. For 
example, plant longevity is negatively correlated 
with reproductive allocation and seed dormancy, and 
is positively correlated with outcrossing, genetic 
diversity, age at first reproduction, and seed mass. 
Seed mass, in turn, is negatively correlated with seed 
dormancy, specific leaf area, and relative growth 
rate, and is positivély correlated (at least among 
herbs) with the presence of vesicular-arbuscular 
mycorrhizae. Life form (whether a plant is herba- 
ceous or woody) is correlated with leaf chemistry, 
mating system, and ectomycorrhizae. 

Individual chapters highlight particular questions, 
approaches and findings in plant life history 


research. Highlights include Silvertown & Dodd’s 
(Chapter 1) suggestion to limit comparisons to con- 
specific or confamilial pairs, and Donoghue & 
Ackerly’s (Chapter 2) use of sensitivity analysis to 
explore the implications of uncertain phylogenies 
when testing for trait correlations. Crawley et al. 
(Chapter 3) explore correlates of exotic plant distri- 
bution in the British Isles, including phylogenetically 
controlled independent contrasts of native and alien 
species traits; they conclude that plants alien to 
Britain tend to be larger, have larger seeds, are more 
likely to flower very early or very late in the year, 
have long-lived seed banks and are more likely to be 
pollinated by insects than their native counterparts. 
Hamrick and Godt (Chapter 6) conclude that genet- 
ic diversity and its distribution are more closely asso- 
ciated with individual species’ life history traits than 
with their phylogenetic status; e.g., outcrossing 
species have less genetic diversity among populations, 
regardless of their ancestry or other traits. Rees 
(Chapter 7) concludes that large-seeded plants, long- 
lived plants, and those with wide lateral spread are 
less likely to have long-lived seed banks. Westoby et 
al. (Chapter 8) conclude that larger seed mass does 
convey benefits in seedling establishment under a 
wide variety of circumstances, but is correlated with 
several other plant attributes (including height, growth 
form, dispersal mode, specific leaf area, and potential 
growth rate). Chapter 10 (by van Groenendael et al.) 
explores the relationship between clonality and habitat 
wetness, noting that a positive correlation disappears 
when one considers that the majority of aquatic and 
semi-aquatic species are monocots. Franco and Silver- 
town (Chapter 11) use three different approaches to 
confirm that species with high risks of adult mortality 
tend to reproduce earlier. Sibly (Chapter 12) reviews 
what is known of life history evolution constrained by 
trade-offs in heterogeneous environments, confirming 
that a gradual cline in habitat frequency can still pro- 
duce an abrupt change in the optimal phenotype. 
Futuyama and Mitter (Chapter 13) discuss how diffi- 
cult it is to rigorously evaluate the co-evolution of 
herbivorous insects and their hosts. Fitter and 


720 


Moyersoen (Chapter 14) investigate the history of 
symbioses in plants, noting that revised classifications 
suggest a single ancestor for all species supporting 
nitrogen-fixing symbioses. They also observe that 
non-mycorrhizal plant species should be considered 
atypical habitat specialists (typically ruderal or wet- 
land species), having thinner roots, smaller seeds, and 
habitats with less extreme pH than mycorrhizal 
species. The final chapter by Goldberg (Chapter 15) 
confirms that competitive hierarchies between pairs of 
plant species are consistent, though her analysis (as 
she acknowledges) did not utilize the methods of 
independent contrasts employed elsewhere in the 
book to account for relatedness of species. 

Some chapters (4, 5, and 9) were disappointingly 
jargon-filled, poorly written and edited, or too the- 


Flora of Mount Rainier National Park 


By David Biek. 2000. Oregon State University Press, 
Corvallis. vi + 506 pp., illus. U.S. $29.95. 


Mount Rainier National Park which now covers 
235 612 acres or 368 square miles lies west of the 
Cascade Divide and is located in Pierce and Lewis 
Counties about 55 miles southeast of Seattle in the 
state of Washington. It was established on 2 March 
1899. Much of the park is surrounded by National 
Forests and Wilderness areas but a good portion of 
the western border is privately owned and has been 
clear cut. Mount Rainier which is the highest of the 
Cascade volcanoes (14 410 feet) occupies about one- 
third of the area of the park and one-quarter of the 
park is bare alpine rock, ice, and snowfields. Studies 
of the plants found in the park began in the late 
1900s and C. V. Piper published articles in 1901 and 
1902 entitled The Flora of Mount Rainier in which 
295 species were listed. Since then a number of indi- 
viduals have increased the knowledge of the flora of 
the park and published their results. 

The introduction to the volume describes the his- 
tory of the park, how to use the book, a description 
of how the information in the flora is provided (sci- 
entific names, authorities, common names, minimum 
number of technical terms, habitat and ecological 
information, where to look for each species, the use 
of metric units, and abbreviations found throughout 
the text), how to use the keys, information on plant 
names (the plant names in this book for genus, 
species, subspecies, or variety follow J. T. Kartesz 
(1994: A Synonymized Checklist of the Vascular 
Flora of the United States, Canada, and Greenland), 
information about the park and its climate. This is 
followed by information on forests and plant com- 
munities, plant geography and distribution, weeds, 
rare plants, and history. 

The 439 pages encompass the keys and descrip- 
tions to the major vascular plant groups: Ferns and 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


oretical; this book covers a narrow enough field of 
interest even without such irritations. I cannot rec- 
ommend this book for most field biologists or plant 
ecologists, because of its specialized subject mat- 
ter. Furthermore, editorial direction and inputs 
appear to have been minimal and incomplete. But 
for researchers in the fields of plant population 
genetics and evolutionary ecology, this volume 
provides an important synthesis of thinking on a 
multi-faceted problem being addressed by scien- 
tists with a wide range of backgrounds, perspec- 
tives, and techniques. 


PHILIP J. BURTON 


Box 3398, Smithers, British Columbia VOJ 2NO Canada 


Fern Allies, Gymnosperms, Dicots, and Monocots. 
The families, genera, and species are in alphabetical 
sequence to make the use of the flora easier for those 
not acquainted with the relationships in more techni- 
cal works. Family descriptions and generic descrip- 
tions are provided where there are more than one 
genus or species found in the park. Throughout the 
text the keys and descriptions of families, genera, 
and species are not in the technical format normally 
found in floras but are easy to read and be under- 
stood by those who have never taken a botanical 
course in university but who want to Jearn something 
about the vegetation found in this most interesting 
region. Following the description is useful informa- 
tion on habitat and where to look for a species and 
other interesting information. Line drawings which 
were previously published in the Vascular Plants of 
the Pacific Northwest by C. L. Hitchcock are found 
beside each species (reprinted by permission of 
University of Washington Press). In addition colour 
photographs of 64 species on eight pages can be 
found in the middle of the book. 

Following the main text are an appendix of nine 
pages on which information is provided regarding 79 
species which have recently been added to the flora 
of the park, plus a glossary, a bibliography, and an 
index of scientific and common names. 

The Flora of Mount Rainier National Park by 
David Biek will be an invaluable help to many of the 
thousands of individuals who visit the park every 
year who want to increase their knowledge of the 
plants of this most interesting region. 


WILLIAM J. CODY 


Biological Resources Program, Eastern Cereal and Oilseed 
Research Centre, Agriculture and Agri-Food Canada, Wm. 
Saunders Building, Central Experimental Farm, Ottawa, 
Ontario KIA 0C6 Canada 


2001 


BOOK REVIEWS 


721 


The Flora of Manitoulan Island and the Adjacent Islands of Lake Huron, 
Georgian Bay and the North Channel, Third Edition 


By J. K. Morton and J. M. Venn. 2000. University of 
Waterloo Biology Series Number Forty. 376 pp. Avail- 
able from University of Waterloo, Biology Department. 
Cloth $40.00 + $7.50 postage and handling + GST; Coil 
bound $27.50 + $7.50 postage and handling + GST. 


The first edition of the flora of this region was 
published in 1977 by J. K. Morton (University of 
Waterloo Biology Series Number fifteen, 62 pages). 
This work included a 12-page introduction, 42 pages 
listing the flora of 906 species (692 native and 214 
alien) with comments, plus literature cited, and 
indexes to English names, genera, and families. 

The second edition was published in 1984 by J. K. 
Morton and J.M. Venn (University of Waterloo 
Biology Series Number twenty-eight, 248 pages). 
This work included a 36-page introduction, 86 pages 
listing an expanded flora of 1167 species (831 native 
and 336 alien plus 6 subspecies and 30 hybrids) with 
frequently expanded comments, plus literature cited, 
index to scientific names, index to common names, 
16 pages with 124 excellent colour photographs, and 
an index of species found on 53 small adjacent 
islands. This is followed by 106 pages of distribution 
maps in the sequence of the text. 

The third edition is another great step forward. 
The introduction is expanded to 46 pages including 
information on geology, glacial and post glacial his- 
tory, climate, botanical exploration, vegetation in the 
various plant communities, conservation, composi- 
tion and origin of the flora, changes to the flora (15 
species appear to have been lost and 72 species, 
including 32 that are native, have been added since 
the second edition), statistics of the flora, arrange- 
ment of the flora, recording and mapping, rarity and 
abundance, excluded records, chromosome numbers 
of plants from the Manitoulin Region, abbreviations 
and symbols, acknowledgements, and a postscript. 


ENVIRONMENT 


Of particular interest in the above is the following 
breakdown of the taxa: 


Native Introduced Total 

Species 861 425 1286 
Subspecies + 4 8 
Varieties 9 3 12 
Hybrids 34 10 44 
908 442 1350 


Of the native taxa, 59 are rare in the Manitoulin 
Region although most are common in other parts of 
Ontario or in neighbouring parts of the continent. 
The area, which covers about 2800 sq. km. (about 
1090 square miles), contains about a third of the 
flora of Canada although it occupies only 0.028% of 
the country. It is thus, without question, one of the 
most interesting areas in our country. 

In the flora section, which now numbers 106 
pages, in addition to the new taxa, many of the taxa 
have additional information which will be most 
interesting to the reader. This, like the second edition 
is followed by the bibliography, index to scientific 
and common names, the list of illustrations, the 124 
colour photographs, the floras of the smaller islands, 
and distribution maps. 

This publication will be invaluable to anyone 
interested in the flora of Ontario and the authors are 
to be congratulated for the result of their continuing 
study of the region. 


WILLIAM J. Coby 


Biological Resources Program, Eastern Cereal and Oilseed 
Research Centre, Agriculture and Agri-Food Canada, Wm. 
Saunders Building, Central Experimental Farm, Ottawa, 
Ontario KIA 0C6 Canada. 


Food Webs and Container Habitats: The Natural History and Ecology of Phytotelmata 


By R.L. Kitching. 2000. Cambridge University Press, New 
York. xiii + 431 pp., illus. U.S. $100. 


To most Canadians, a pitcher plant (Sarracenia 
sp.) 18 unique because it is one of very few carnivo- 
rous plants. However, it is the life that thrives, not 
that which perishes, in these pitchers (and other 
water-filled plant cavities) that is the topic of this 
very thorough book. The over 100 years of research 
on phytotelmata (food webs in plant containers) is 
covered quite thoroughly. The author himself has 
been involved in this research for over 30 years, 


starting with his post-graduate work at Oxford. 
Studying most food webs in nature is very complex, 
there are no isolated food webs, though container 
habitats come close, and offer ecologists small, rela- 
tively simple systems to study. 

The information in this book is of the weird and 
wonderful sort that makes birders, plant enthusiasts, 
herpetophiles, and really all naturalists exclaim, 
“wow!” However, this work is not a coffee table 
book, nor is it targeted at the average naturalist 
(whatever that may be!). The book is a summary of 


W222 


many people’s research, and is directed towards 
ecologists and entomologists, though researchers of 
other phytotelm taxa will be interested as well. 

The book is organized into five parts, with the first 
one giving a description of the major types of con- 
tainer habitats (tree holes, bromeliads, etc.) and the 
ecological niches of the inhabitants (saprophages to 
predators); it is this section that will be most con- 
sumable by naturalists. The book culminates in an 
annex entitled “The Bestiary” which outlines the ani- 
mals known to make use of phytotelmata, from ver- 
tebrates to flatworms, though by far the majority of 
this section is devoted to the animals that dominate 
the world as a whole, the insects. The Bestiary is a 
straightforward systematic breakdown of the ani- 
mals, with very useful tables which direct the reader 
to published works of each taxon. 

The bulk of the book compares phytotelmata on 
many scales, from local to global, across time and 
habitats. The organisms from different areas and dif- 
ferent types of cavities are thoroughly described; the 
relationships among these phytotelm are explained 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


quite well. It is these sections that draw most heavily 
on the primary literature. 7 

The bad points? There are too few photographs, 
and those that are there are all dark. Almost all of the 
graphs and line drawings are pixelated. Pixelated! 
For hundreds of years, publishers have been able to 
make a straight line look like a straight line, and in 
this era of technology there is absolutely no excuse 
for such inferior quality, especially from such a fine 
publisher. The book is directed primarily at biolo- 
gists, and as such it is understandable that the bino- 
mial names of organisms are used; using common 
names occasionally (when such exist), or at least 
parenthetically, would colour the writing with a 
favourable hue for the non-specialist. 

Overall though, this is a well-written book on a 
subject that certainly deserves the attention. 


RANDY LAUFF 


Department of Biology, St. Francis Xavier University, 
Antigonish, Nova Scotia B2G 2W5 Canada 


The Friendship of Nature, A New England Chronicle of Birds and Flowers 


By Mabel Osgood Wright. Edited by Daniel J. Philippon. 
1999. John Hopkins University Press, Baltimore. 172 
pp., illus. Reissue of 1900 book. U.S. $16.95. 


Every so often a book comes along which revives 
a figure of the past. That is the case with The 
Friendship of Nature, written by Mabel Osgood 
Wright, founder of the Connecticut Audubon Society, 
author of 25 works of fiction and non-fiction, long- 
time associate editor of Bird-Lore magazine (now 
Audubon), accomplished photographer, and the 
organizing force behind one of the first privately 
owned bird sanctuaries in the United States. As editor 
Dan Philippon (University of Minnesota, Twin 
Cities) writes in his lengthy but excellent introduc- 
tion, Osgood Wright is “not only a neglected writer 
and illustrator but also a lost hero of the conservation 
movement”. 

The introduction provides intriguing background 
to this remarkable woman, who realized the need to 
integrate the personal and the regional, to extend 
herself through contact with nature. The Friendship 
of Nature, roughly organized according to the sea- 
sons, paints a loving portrait of the birds and flora 
Osgood Wright encourages and tends in her large 
garden, and the wildlife and seasonal transforma- 
tions she experiences in the surrounding countryside. 

Her keen observations and insights, expressed in 
meticulous prose and poetic descriptions typical of 
the period (1894), are a delight to read. I particu- 

_larly like her charming, and wholly accurate, 


sketches of the bobolink: “in the tufts of grass out 
in the open, hovering above a nest that is merely a 
heap of twigs, the bobolink calls in a perfect ecsta- 
sy: ‘Bobolink 0’ wadolink, winterseeble-see me-see 
me-see!’” (springtime), and “madcap bobolinks are 
now anxious to disperse their broods before the 
mowers lay bare their shabby nests; and half 
bewitched, they sing, and pausing, float with out- 
stretched wings, then soaring, pour out torrents of 
high notes” (summer). 

But it isn’t all pretty pictures and words. Osgood 
Wright also makes astute observations regarding 
institutions such as agriculture: “If you serve Nature, 
waiting her moods, taking what she yields unforced, 
giving her a love devoid of greed, she will be a regal 
mistress, and all she has to bestow will be yours. 
Exact and say to one little field: ‘This year you shall 
yield this crop or that,’ and it becomes a battle- 
ground, where Nature, well equipped, wages war 
with man.” 

The Friendship of Nature would make an excel- 
lent gift for naturalists, birders, and gardeners who 
love books. Not too long, divided into 11 neat and 
clearly written chapters, and illustrated with Osgood 
Wright’s own photographs, it is a quick and gratify- 
ing read. 


R. SANDER-REGIER 


RR5 Shawville, Quebec JOX 2YO Canada 


2001 


Earth Future: Stories from a Sustainable World 


By Guy Dauncey. 1999. New Society Publishers, Gabriola 
Island. xii + 161 pp. $17.95. 


In this day and age when the world's population is 
bombarded with negativity regarding the earth's 
future, the positive potential often appears lost. Guy 
Dauncey author, speaker, organizer, and consultant 
on post-industrial environmentally sustainable future 
provides some possible scenarios. In his short works 
of fiction he attempts to offer an environmentally 
sustainable positive vision. The majority of the ini- 
tiatives he notes are already happening. 

The 21 short stories, two poems and one pledge 
tell not of the completion but of the making of a 
vision. Subjects covered include the expected recy- 
cling, organic gardening, co-operatives, nature of 
consciousness, de-globalization to name a few. Most 
steries have concluding notes with lists provided of 


BOOK REVIEWS 


723 


contacts, many being internet URL’s. Dauncey does 
not aim to provide a balanced view. His intent is 
rather to provide a look at a positive possible future. 
There is recurring anti-hitech theme with a strong 
emphasis on low tech solutions. 

If one is looking for positive infusion for a possi- 
ble future, this book provides it. The book also has 
the potential, I feel, as a discussion starter. Guy 
Dauncey provides all in all not a bad read, with a 
strong emphasis on the positive future, which 
requires us all to create our positive visions of the 
future. 


M. P. SCHELLENBERG 


434 4th Avenue SE, Swift Current, Saskatchewan S9H 3M1 
Canada 


Vanishing Borders: Protecting the Planet in the Age of Globalization 


By Hilary French. 2000. W.W. Norton & Company Ltd., 
London. 257 pp., illus. U.S. $13.95; Pdf version 2 down- 
loads at U.S. $5.50 each. 


In today’s modern world with increased rapidity 
of communication and travel, borders no longer limit 
potential dangers of the outside world’s impact on a 
nation’s environmental resources. Hilary French pro- 
vides, in this Worldwatch publication, an overview 
of the threats as well as potential cures to the prob- 
lems of globalization. 

The reader becomes acquainted with the issues in 
the first section of the book entitled “The Ecology of 
Globalization.” In this section chapters with its 
descriptive titles; “Nature Under Siege,” “the Biotic 
Mixing Bowl,” “Global Grocers,” “the Export of 
Hazard”, and “Sharing the Air”; one is provided with 
an array of facts and analyses describing the plight 
of the world’s resources. Issues encountered includ- 
ed under funding, shortcomings of present eco- 
nomics, unsustainable agricultural progress, bioinva- 
sions to name a few. 

In the second section of the book, “Reforming 
Global Governance” policy problems and potential 
solutions are offered. Once again the reader is pro- 
vided insight into the chapter contents via chapter 
entitled; “Trade Wars”, “Greening the Financial 
Architecture”, “Strengthening Global Environmental 


Governance”, and “Partnerships for the Planet.” As 
the titles suggest impact of present globalization 
policies, and present economic theory are discussed 
in relation to their environmental impact. 

Overall, Hilary French has provided a book with a 
great deal of information summarized from a large 
number of sources in a fashion which will be famil- 
iar to those who are familiar with the World Watch 
Institute. A great deal of the referenced material 
originates in reports normally not available to the 
common reader. The summarization of the present 
day data continues to indicate a gloomy future for 
the world community as do many other publications 
on the future. The outlining of solutions makes this 
book somewhat unique. 

The book is meant for the general reader to pro- 
vide information to be used in the decision making 
process at all levels, personal or international. The 
material has been provided in a well organized and 
readable fashion. For those with an interest and stake 
in the future this will provide a good read and refer- 
ence. 


M. P. SCHELLENBERG 


434 4th Avenue SE, Swift Current, Saskatchewan S9H 3M1 
Canada 


724 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Pandora's Poison: Chlorine, Health, and a New Environmental Strategy 


Joe Thornton. 1999. MIT Press, Cambridge, Massachusetts. 
597 pp., illus. U.S.$34.95. 


This is a thick, formidable book. It presents a 
compelling, seemingly irrefutable position: that pro- 
duction, use, and disposal of chlorine gas and its 
chemicals, a phenomenon of the past century, is bad 
for us, bad for society, and bad for the environment. 
The author has achieved his aim of a readable essay, 
while his figures, tables, appendices, and about a 
thousand references, offer detailed documentation. 

Chlorine itself is an effective bleach and a disin- 
fectant. Chlorine gas does not occur naturally, but 
industry now produces 40 million tons of it annually. 
This man-made gas is the feedstock for 11 000 
organochlorine compounds including plastics, pesti- 
cides, and solvents. Chlorination decreases a chemi- 
cal’s stability, making it more reactive, and usually 
more toxic. And while some of these pollutants may 
degrade in less than a year in the temperate climates 
in which they are produced, they may last for cen- 
turies near the frigid poles, which become the ulti- 
mate global sinks. Organochlorines are now “ubiqui- 
tous on a global scale.” 

Sadly, scientists assigned to investigate the 
degree of toxicity of the commonest chlorine com- 
pounds have followed the Risk Paradigm. They 
tend to calculate only the Lowest Observed 
Adverse Effect Level (LOAEL), or the No 
Observed Adverse Effect Level (NOAEL). Each 
chemical is studied separately. Yet, any exposure to 
any organic chlorine compound carries some risk of 
mutations, of cancer, of endocrine disruption 
(including infertility), of neurotoxicity, and of sub- 
clinical effects, among others. 

Thornton, in striking contrast to industry, advo- 
cates the Ecological Paradigm, because, as Canadian 
ecologist Glen A. Fox has pointed out, “health dam- 
age occurs on a continuum.” The Ecological 
Paradigm, a holistic approach, views all organochlo- 
rines as “end products unified by a single history, a 
common economic dynamic, and a single root mate- 
rial.” There is no safe dose. A central tenet is the 
precautionary principle. For this, the first rule is 
Reverse Onus, which means erring on the side of 
caution. The second is Zero Discharge and the third 
is Clean Production. 


What dangers are known? Prostate cancer, non- 
Hodgkin’s lymphoma, and multiple myeloma have 
tripled in prevalence between 1950 and 1994 in the 
United States, and cancers of the testis, kidney, thy- 
roid, and liver have doubled. Perchloroethylene, used 
in dry-cleaning, causes leukemia in the laboratory, 
and trichloroethylene causes kidney cancer. The rate 
of non-Hodgkin’s lymphoma in domestic dogs dou- 
bles when the owner applies 2,4-D on the lawn four 
times during the summer. Dioxin deposition in Great 
Lakes sediments has increased 2000-fold since 1940. 
Thousands of animals died near Seveso, Italy, when 
chlorophenols and dioxins escaped from a factory 
manufacturing trichlorophenols. 

What can be done? The first principle of public 
health practice must be prevention. The ever- 
increasing production of dangerous chlorine prod- 
ucts must be checked; PVC or vinyl plastic is now 
the largest user of chlorine in the world. Phasing 
out chlorine represents a “major step on the road 
toward a sustainable economy based on clean mate- 
rials and production techniques.” When a new pulp 
and paper plant is to be built, totally chlorine-free 
bleaching methods (TCF) allow less expensive con- 
struction. Major companies have eliminated the use 
of chlorinated solvents and some have switched to 
propane, isobutane, and ammonia as refrigerants. In 
dry-cleaning, chlorine should be replaced with ‘wet 
cleaning’ as some establishments have already 
done. Plastics can be made from polypropylene 
rather than PVC (vinyl). Ozonation is preferred to 
chlorine in swimming pools. Sometimes the substi- 
tutes cost less, sometimes more, but jobs need not 
be lost. Thornton explains the political and social 
barriers and the details of the economic costs and 
benefits of each intervention. The environment is at 
stake. 

Industry opened Pandora’s box less than a century 
ago. Now it is time to shut it. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


2001 


NEw TITLES 


Zoology 


Animal behavior desk reference. 2001. By E. M. 
Barrows. 2nd edition. CRC Press, Boca Raton, Florida. 
936 pp. U.S. $129.95. 


*Bats of Papua New Guinea. 1998. By F.J. Bonaccorso. 
Conservation International (distributed by University of 
Chicago Press, Chicago). 492 pp., illus. U.S. $40. 


Bears of the world. 2000. By L. Craighead. Voyager, 
Stillwater, Minnesota. 132 pp., illus. U.S. $29.95. 


Biology of marine birds. 2001. Edited by E.A. 
Schrieber and J. Burger. CRC Press, Boca Raton, Florida. 
744 pp. U.S. $79.95. 


*Birds of Delaware. 2000. By G. K. Hess, R. L. West, 
M. V. Barnhill II, and L. M. Flemming. University of 
Pittsburgh Press, Pittsburgh. xv + 635 pp., illus. U.S. $65. 


*Birds of southern South America and Antarctica. 
2001. By M.R. de la Pena and M. Rumball. Originally 
published by Collins in 1998. Princeton University Press, 
Princeton. 304 pp., illus. U.S. $24.95. 


Canadian feathers: a loon-atics guide to anting, 
mimicry, and dump nesting. 2001. by P. E. Bumstead. 
Simply Wild Publications, Calgary. $29.99. 


Carnivore conservation. 2001. Edited by J.L. 
Gittleman, S. M. Funk, D. Macdonald, and R. K. Wayne. 
Cambridge University Press, New York. xii + 675 pp., 
illus. Cloth U.S. $130; paper U.S. $49.95. 


*Cetacean societies: field studies of dolphins and 
whales. 2000. Edited by J. Mann, R. C. Connor, P. L. 
Tyack, and H. Whitehead. University of Chicago Press, 
Chicago. xiv + 433 pp., illus. Cloth U.S. $85; paper U.S. 
$35. 


+Changing tracks: predators and politics in Mt. 

McKinley National Park. 2001. By T. Rawson. 
University of Alaska Press, Fairbanks. xiii + 317 pp., 
illus. Cloth U.S. $39.95; paper U.S. $24.95. 


*The complete guide to the wildlife of Britain and 
Europe. 2001. By R. Hume and P. Hayman. Mitchel 
Beazley, London, England. 288 pp., illus. £25. 


Ecotoxicology of wild mammals. 2001. Edited by R. F. 
Shore and B. A. Rattner. John Wiley and Sons, New York. 
U.S. $199.95. 


+A field guide to the reptiles of east Africa. 2002. By S. 
Spawls, K. Howell, R. Drewes, and J. Ashe. Academic 
Press, San Diego. 543 pp., illus. 


*Hummingbirds of North America; the photographic 
guide. 2001. By S. N. G. Howell. Academic Press, San 
Diego. ix + 219 pp., illus. U.S. $24.95. 


+In search of the golden frog. 2000. By M. Crump. 
University of Chicago Press, Chicago. xvi + 299 pp., illus. 
US!$27. 


BOOK REVIEWS 


725 


*Life underground: the biology of subterranean 
rodents. 2000. Edited by E. A. Lacey, J. L. Patton, and 
G. N. Cameron. University of Chicago Press, Chicago. x1 
+ 449 pp., illus. Cloth U.S. $65; paper U.S. $24. 


*Marine mammals of the Pacific northwest. 2001. By 
P. Folkens. Harbour Publishing, Madeira Park, British 
Columbia. Brochure $9.95. 


Neotropical tree boas: natural history of the Corallus 
hortulanus complex. 2001. By R. W. Henderson. 
Krieger Publishing, Melbourne, Florida. Illus. 


*The new birdwatcher’s pocket guide to Britain and 
Europe. 2002. By P. Hayman and R. Hume. Mitchell 
Beazley, London, England. 272 pp., Illus. £9.99. 


*On the move: how and why animals travel in groups. 
2000. Edited by S. Boinski and P. A. Garber. University 
of Chicago Press, Chicago. xi + 811 pp., illus. Cloth U.S. 
$95; paper U.S. $35. 


+Owls aren’t wise and bats aren’t blind: a naturalist 
debunks our favorite fallacies about wildlife. 2000. By 
W. Shedd. Three Rivers Press (Canadian distributor 
Random House, Mississauga). ix + 322 pp., illus. $21. 


+Primate conservation biology. 2000. By G. Cowlishaw 
and R. Dunbar. University of Chicago Press, Chicago. 498 
pp., illus. Cloth U.S. $75; paper U.S. $27. 


*Radio tracking and animal populations. 2001. Edited 
by J. J. Millspaugh and J. M. Marzluff. Academic Press, 
San Diego. xvi + 474 pp., illus. U.S. $69.95. 


*Red-tailed hawk. 2000. By C.R. Preston. Wild Bird 
Guides. Stackpole Books, Mechanicsburg, Pennsylvania. 
103 pp., illus. U.S. $19.95. 


+Resolving human-wildlife conflicts: the science of 

wildlife damage management. 2001. By M. Conover. 
Lewis Publishers, CRC Press, Boca Raton, Florida. 440 
pp. U-S. $69.95. 


Human dimensions of wildlife management in North 
America. 2001. Edited by D. J. Decker, T. L. Brown, and 
W.F. Siemer. The Wildlife Society, Bethesda, Maryland. 
464 pp. cU.S. $40. 


*+Sacred hunt: a portrait of the relationship between 
seals and Inuit. 2001. By D.F. Pelly. Greystone Books 
(Douglas and McIntyre), Vancouver. xv + 127 pp., illus. 
$34.95. 


*+Seabird bycatch: trends, roadblocks, and solutions. 
2001. Edited by E. F. Melvin and J. K. Parrish. University 
of Alaska Sea Grant, Fairbanks. Vii + 206 pp., illus. U.S. 
$20. 


*Survivors in armour: turtles, tortoises, and terrapins. 
2001. By R. Orenstein. Key Porter Books, Toronto. xi + 
308 pp., illus. $45. 


A whale biologist at work. 2001. By S. B. Collard, III. 
Watts, Danbury, Connecticut. 48 pp., illus. Cloth U.S. 
$22.50; paper U.S. $6.95. 


726 


Wolf: spirit of the wild: a celebration of wolves in 
word and image. 2000. Edited by D. Landau. Sterling, 
New York. xi + 182 pp., illus. U.S. $24.95. 


The world of frogs, toads, salamanders, and newts. 
2000. Edited by R. Hofrichter. Firefly, Willowdale, 
Ontario. 264 pp., illus. U.S. $49.95. 


*The world of humming birds. 2001. By R. Burton. 
Firefly Books, Willowdale, Ontario. 158 pp., illus. $40. 


Botany 


*The ecology of trees in the tropical rain forest. 2001. 
By I. M. Turner. Cambridge University Press, New York. 
xiv + 298 pp., illus. U.S. $80. 


*Ecological management of agricultural weeds. 2001. 
By M. Liebman, C.L. Mohler, and C. P. Staver. 
Cambridge University Press, New York. xi + 532 pp., 
illus. U. S. $120. 


*Guide to standard floras of the world. 2001. By D. G. 
Frodin. 2nd edition. Cambridge University Press. New 
York. xxiv + 1100 pp., U.S. $240. 


*Lichens of North America. 2001. By I. M Brodo, S. D. 
Sharnhoff, and S. Sharnhoff. Yale University Press, New 
Haven. 


Orchids and their conservation. 2001. By H. Koopowitz. 
Timber Press, Portland, Oregon. 177 pp., illus. U.S. 
$39.95. 


*Rare vascular plants of Alberta. 2001. Edited by L. 
Kershaw, J. Gould, D. Johnson, and J. Lancaster. 
University of Alberta Press, Edmonton. xliv + 484 pp., 
illus. $29.95 


+ Vegetation and the terrestrial carbon cycle; modelling 
the first 400 million years. 2001. By D. I. Beering and F. 
I. Woodward. Cambridge University Press, New York. x+ 
405 pp., illus. U.S. $150. 


Environment 


*+Bioconservation and sytematics. 2001. Edited by J. B. 
Phipps and P. M. Catling. Canadian Botanical Association 
(Dr. M. Fisher, Box 160, Aberdeen, Saskatchewan SOK 
OAO) iii + 101 pp., illus. $23 in Canada; U.S. $17 in 
U.S.A., U.S. $21 elsewhere. 


Biological diversity: balancing interests through adap- 
tive collaborative management. 2001. By L. E. Buck, 
C. C. Geisler, J. Schelhas, and E. Wollenberg. CRC Press, 
Boca Raton, Florida. 504 pp. U.S. $49.95. 


Biogeography of the West Indies: patterns and per- 
spectives. 2001. Edited by C. A. Woods and F. Sergile. 
2nd edition. CRC Press, Boca Raton, Florida. 608 pp., 
U.S. $139.95. 


{Biotic forest communities of Ontario. 2001. By N. D. 
Martin and N. M Martin. 3rd edition. Commonwealth 
Research, Belleville, Ontario. 195 pp., illus. $10. 


' *Bringing the biosphere home, learning to perceive 
global environmental change. 2001. By M. Thomashow. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


MIT Press, Cambridge, Massachusetts. xi + 244 pp., U.S. 
927.95. 


The earth’s biosphere. 2002. By V. Smil. MIT Press, 
Cambridge. 360 pp., illus. U.S. $32.95. | 


+Elements of mathematical ecology. 2001. By M. Kot. 
Cambridge University Press, New York. 453 pp., illus. 
Cloth U.S. $110; paper U.S. $39.95. 


Environmental careers, environmental employment, 
and environmental training. 2001. Edited by W. L. 
Filho. Peter Lang Scientific Publishers, New York. U.S. 
$31.95. 


+Conservation of an exploited species. 2002. Edited by 
J.D. Reynolds, G. M. Mace, K. H. Redford, and J. G. 
Robinson. Cambridge University Press, New York. xx + 
524 pp., illus. Cloth U.S. $120; paper U.S. $44.95. 


The Everglades, Florida Bay, and the coral reefs of the 
Florida Keys: an ecosystem sourcebook. 2001. By J. W. 
and K. G. Porter. CRC Press, Boca Raton, Florida. c1064 
pp., illus. U.S. $199.95. 


+Flammable Australia: The fire regimes and biodiversi- 
ty of a continent. 2002. Edited by R. A. Bradstock, J. E. 
Williams, and A. M. Gill. Cambridge University Press, 
New York. ix + 462 pp., illus. U.S. $130. 


Fungal conservation: issues and solutions. 2001. Edited 
by D. Moore, M. M. Nauta, S. E. Evans, and M. Rotheroe. 
Cambridge University Press, New York. x + 262 pp., 
illus. 


*+Globalization and environmental reform. 2001. By 
A. P. J. Mol. MIT Press, Cambridge, Massachusetts. x + 
273 pp., illus. U.S. $37.95. 


Hands on nature. 2000. Edited by J. Lingelbach and L. 
Purcell. Vermont Institute of Natural Science, Woodstock. 
329 pp., illus. U.S. $24.95. 


+The hiking trails of Florida’s national forests, parks, 
and preserves. 2001. By J. Molloy. University Press of 
Florida, Gainsville (distributed by Scholarly Book 
Services, Toronto). xv + 190 pp., illus. $27.95. 


+How green is the city? 2001. Edited by D. Devuyst. 
Columbia University Press, New York. xxi + 457 pp. 
Cloth U.S. $70; paper U.S. $30. 


Insects and gardens: in pursuit of a garden ecology. 
2001. By E. Grissell. Timber Press, Portland, Oregon. 348 
pp., illus. U.S. $29.95. 


+Lessons from Amazonia: the ecology and conservation 
of a fragmented forest. 2002. Edited by R.O. 
Bierregaard, Jr., C. Gascon, T. B. Lovejoy, andi: 
Mesquita. Yale University Press, New Haven. xv + 478 
pp., illus. U.S. $65. 


+A long look at nature: the North Carolina State 
Museum of Natural Sciences. 2001. By M. Martin. 
University of North Carolina Press, Chapel Hill (distribut- 


2001 


ed by Scholarly Book Services, Toronto). xi + 174 pp., 
illus. $32.95. 


*The love of nature and the end of the world: the 
unspoken dimensions of environmental concern. 2002. 
By S.W. Nicholsen. MIT Press, Cambridge, 
Massachusetts. ix + 216 pp. U.S. $27.95. 


*The natural history of an oil field: development and 
the biota. 2000. Edited by J. C. Truett and S. R. Johnson. 
Academic Press (Harcourt), Troy, Missouri. xvi + 422 pp., 
illus. 


+Norms of nature. 2001. By P.S. Davies. MIT Press, 
Cambridge, Massachusetts. xiv + 234 pp. 


*Political nature: environmentalism and the interpreta- 
tion of western thought. 2001. By J. M. Meyer. MIT 
Press, Cambridge, Massachusetts. xii + 210 pp. 


Reefscape: reflections on the Great Barrier Reef. 
2001. By. R. Love. Joseph Henry Press, Washington. 250 
pp. U.S. $24.95. 


*Scaling relations in experimental ecology. 2001. 
Edited by R. H. Gardner, W. M. Kemp, V.S. Kennedy, 
and J. E. Petersen. Columbia University Press, New York. 
Xxx + 373 pp., illus. Cloth U.S. $65; paper U.S. $30. 


Self-organization in biological systems. 2001. By S. 
Camazine, J.-L. Deneubourg, N. Franks, J. Sneyd, G. 
Theraulaz, and E. Bonabeau. Princeton University Press, 
Princeton. U.S. $65. 


7Solutions for an environment in peril. 2001. Edited by 
A.B. Wolbarst. Johns Hopkins University Press, 
Baltimore. xv + 214 pp. U.S. $22.50. 


Spatial processes and management of marine popula- 
tions. 2001. Edited by G. H. Kruse, N. Bez, A. Booth, 
M. W. Dorn, S. Hills, R. N. Lipcius, D. Pelletier, C. Roy, 
S.J. Smith, and D. Withrell. Alaska Sea Grant College 
Program, Fairbanks. 730 pp. U.S. $40 in Canada and 
U.S.A.; U.S. $60 elsewhere. 


*Thoreau’s country: journey through a transformed 
landscape. 2001. By D. Foster. Harvard University Press, 
Cambridge, Massachusetts. 270 pp., illus. U.S. $16. 


World resources 2000 — 2001 people and ecosystems: 
the fraying web of life. 2000. By World Resources 
Institute, Washington. ix + 389 pp., illus. Cloth U.S. $49; 
paper U.S. $27.nnn 


BOOK REVIEWS 


727 


Miscellaneous 


*Backcountry huts and lodges of the Rockies and 
Columbias. 2001. By J. Scott. Johnson Gorman, 
Calgary. 287 pp., illus. $24.95. 


*Cheltenham in Antarctic: the life of Edward Wilson. 
2001. By D. M. Wilson and D. B. Elder. Reardon 
Publishing, Leckhampton, England. 144 pp.. illus. £9.99. 


+Minutes of meetings, 1961 to 1969 of the McIlwraith 
Ornithological Club, London, Ontario, Canada. 2002. 
By W. W. Judd. Phelps Publishing, London. iii + 168 pp. 
$10. 


+Rediscovering the Great Plains: journeys by dog, 

canoe, and horse. 2001. By N. Henderson. Johns 
Hopkins University Press, Baltimore. Xv + 214 pp., illus. 
UWS.529.95. 


+Smithsonian Institute secretary, Charles Doolittle 

Walcott. 2001. By E.L. Yochelson. Kent State 
University Press, Kent, Ohio. xvi + 589 pp., illus. U.S. 
$55. 


Understanding soil change. 2001. By D. D. Richter, Jr. 
and D. Markewitz. Cambridge University Press, New 
York. xiv + 255 pp., illus. U.S. $69.95. 


Books for Young Naturalists 


Bears. 2001. By D. Fertl, M. Reddy, and E.D. Stoops. 
Sterling, New York. 80 pp., illus. U.S. $17.95 


Eagles of Devil Mountain. 2001. By M. J. Rauzon. 
Danbury, Watts, Connecticut. 48 pp., illus. Cloth U.S. 
$22.50; paper U.S. $6.95. 


Fireflies. 2001. By S. Walker. Lerner, Minneapolis. 48 
pp., illus. U.S. $22.60. 


Mountain gorillas. 2001. By K. Kane. Lerner, 
Minneapolis. 48 pp., illus. U.S. $21.95. 


Oceans. 2001. By S. H. Gray. Compass Point Books, 
Minneapolis. 48 pp., illus. U.S. $15.95. 


Songbirds: the language of song. 2001. By S.A. 
Johnson. Carolrhoda, Minneapolis. 48 pp., illus. U.S. 
$23.95. 


Tundra. 2001. By S.H. Gray. Compass Point Books, 
Minneapolis. 48 pp., illus. U.S. $15.95. 


tavailable for review 
*assigned for review 


The Ottawa Field-Naturalists’ Club Awards April 2001 


Each year, The Ottawa Field-Naturalists’ Club makes a 
number of awards to certain members and some non- 
members who have distinguished themselves by accom- 
plishments in the field of natural history and conservation, 
or by extraordinary activity within the Club. The follow- 
ing citations, in an abbreviated form, were read at the 
Club’s annual soirée held, as usual, at the Unitarian 
Church of Ottawa. 


HONORARY MEMBERSHIP: DONALD M. BRITTON 

Donald M. Britton is one of Canada’s most distinguished 
and accomplished botanists. He is recognized worldwide as 
an authority on fern taxonomy and systematics with more 
than 50 years of contributions to understanding the taxono- 
my and evolution of major groups including Dryopteris, 
Polypodium, Woodsia, Polystichum, Isoetes, and 
Lycopodium. He has been a leader in the use of cytological 
techniques in his studies, as well as more traditional 
approaches such as morphological, anatomical, develop- 
mental, and phytogeographic analyses, thereby helping to 
define biosystematics most appropriately. 

Dr. Britton was born in Toronto and received his bache- 
lor’s degree from the U of T. His doctoral work was done 
at the University of Virginia, where he received his Ph.D. 
in 1950. Although he did his early work on the Boragin- 
aceae (which includes viper’s bugloss and forget-me-nots) 
and the genus Rubus (blackberries), he gradually turned 
almost all of his attention to the ferns and fern-allies. After 
a few years in the Horticulture Department at the 
University of Maryland, Dr. Britton returned to Canada in 
1958 to take up a position in the Botany and Genetics 
Department at the University of Guelph, where he 
remained until his retirement, and where he continues to 
devote time to botanical studies. His activities in Guelph 
included continuous involvement in the Guelph Field- 
Naturalists’ Club, where he served for many years as 
Chairman of their Conservation Committee and led many 
field trips. 

Dr. Britton has been a prolific author (including co- 
authoring Ferns of Canada with William Cody). His 
involvement with the Flora North America project includes 
serving as regional reviewer for pteridophytes of eastern 
Canada as well as being an author of the treatment of 
Tsoetes. He has described numerous taxa new to science, 
including two quillworts from the Ottawa Valley. He is an 
exceptional field botanist with a careful and thorough 
approach to all his research, assuring the lasting value and 
reliability of his observations. 

Dr. Britton is an exceptionally generous scholar who has 
inspired professionals and amateurs alike across North 
America and Europe. Those lucky enough to have worked 
with him have been touched by his warmth, humour, and 
especially his equal accessibility to students, amateurs and 
scholars. 

In 1991, the Canadian Botanical Association presented 
Dr. Britton with its premier achievement award, the 
Lawson Medal, for outstanding scientific achievement over 
the period of a career. It is now our turn to honour him by 
welcoming Dr. Britton as an Honorary Member of the 
_ Ottawa Field-Naturalists’ Club. 


HONORARY MEMBERSHIP: JOHN (JACK) GILLETT 

John Montague Gillett, better known to his friends 
and colleagues as “Jack,” is one of the Ottawa Field- 
Naturalists’ Club’s most distinguished members. His 
career as a respected Canadian plant taxonomist, and his 
activities in the Club make him a most fitting recipient of 
Honorary Membership. 

Jack is a native Ottawan. After serving in the Royal 
Canadian Air Force during World War II, he returned to 
Canada and earned his B.A. from Queen’s University and 
Ph.D. at Washington University in St. Louis. In 1949, Jack 
began his botanical career in the herbarium at the Depart- 
ment of Agriculture and participated in many expeditions. 
He left the Plant Research Institute in 1972, to become 
curator of the herbarium at the National Museum of Natural 
Sciences [Canadian Museum of Nature]. Here he continued 
work on a flora of Gatineau Park and taxonomy of many 
plant groups, especially clovers (the genus Trifolium). He 
retired from the Museum in 1983. 

His work on Canadian botany includes monographs on 
Hypericum, Polygala, Bartonia-Obolaria, Gentianella- 
Gentiana, Trifolium, and others. Floristic works include 
studies of the Mealy Mountains in southern Labrador, the 
Madoc region of Ontario, the St. Lawrence Seaway and the 
Ottawa District. In the course of his career, he has 
described several new species of plants from Canada, not 
an insignificant thing in the second half of the 20th century. 

Many of Jack’s papers have been of great importance to 
plant taxonomy in North America. Among these are his 
papers on the economically important genera Agropyron 
and Trifolium. In recent years he has made a series of won- 
derful contributions to Trail & Landscape on the plants of 
the Ottawa district. These articles are useful identification 
aids backed by careful taxonomic evaluations and judge- 
ment, not merely lists of plants. They are of use to amateur 
and professional alike. His Index to the Transactions of the 
Ottawa Field-Naturalists’ Club and the Ottawa Naturalist 
was a tedious task, but a remarkable labour of great benefit. 
His Checklist of the Vascular Plants of the Ottawa-Hull 
Region, written with David White, is an indispensable tool 
for anyone working with the plants of the region. 

Jack has a long history of activity in the Club. From 1958 
to 1970 he served on the Council. From 1959 to 1961 he 
served in the arduous post of Treasurer, and for several 
years in the 1960s, he was Auditor. From 1966 to 1970 he 
served as Chair of the Publications Committee. As well as 
these administrative tasks Jack has also been prominent in 
Club activities with his many contributions to Trail & 
Landscape (16 papers and counting) and has been leading 
field trips and giving talks for more than half a century. 

In recognition of Jack Gillett’s outstanding botanical 
achievements and his considerable service to the OFNC, 
the Club is pleased to award him an Honorary 
Membership. 


HONORARY MEMBERSHIP: E. FRANKLIN POPE 

In the long history of the Ottawa Fieid-Naturalists’ Club, 
few people can match the years of steady dedicated com- 
mitment that Frank Pope has devoted to this organization. 
For the past twenty-two years, he has served continuously 
on Council in various capacities and has been active in 


728 


2001 


many committees. In 1980 he served on the Executive as 
Corresponding Secretary, followed by three years as 
Recording Secretary. He was elected Club President in 
1984, and again in 1985, a period when the Club was 
deeply involved in environmental advocacy, taking a coura- 
geous and public stand on what was then a controversial 
viewpoint. 

For five years, 1988 to 1991, he served as Chairman of 
the Finance Committee. In 1992, he was again elected 
President, a position which he held for the next four years. 
During this second term as President, he had the added 
responsibilities of overseeing all the numerous arrange- 
ments associated with the 1993 conference of the 
Federation of Ontario Naturalists, which was hosted by the 
OFNC. Always ready to offer his services where needed, in 
1999 he took over the vacant position of Club Treasurer, 
which he filled again in the year 2000. An impressive 
record of service! 

Repeatedly over the years, Frank has assumed responsi- 
bilities that made heavy demands on his time and energies. 
Most notable of these undertakings was his unflagging 
leadership in the effort to gain environmental protection for 
Alfred Bog. In this long drawn-out struggle, Frank’s com- 
mitment never wavered in the face of setbacks. A large sec- 
tion of Alfred Bog is now safe from commercial develop- 
ment due, in large part, to Frank’s steadfast leadership. 

Over a period of many years, Frank has been an ever- 
present strength on Council, adding his thoughtful and 
experienced contributions to deliberations. His long pres- 
ence on Council has provided the Club with important con- 
tinuity. In recognition of his years of remarkable and dedi- 
cated service, the Ottawa Field-Naturalists’ Club is pleased 
to confer Honorary Membership on E. Franklin Pope. 


HONORARY MEMBERSHIP: JOYCE and ALLAN REDDOCH 

Joyce and Allan Reddoch have been club members since 
1967. Both have served on Council and on various commit- 
tees, performing outstanding service on the conservation 
committee. They have contributed greatly both to the run- 
ning of the club and to its natural history goals. They have 
added significantly to the knowledge of Canadian natural 
history, with particular emphasis on the Ottawa area. Their 
fame as orchid experts extends far beyond the boundaries 
of the Ottawa region. 

Joyce’s ten-year tenure as editor for Trail & Land- 
scape has been a monumental contribution to the Ottawa 
Field-Naturalists’ Club. The attention to quality and 
detail that she brought to this demanding position ensured 
that the high standards set by the founding editor were 
maintained. 

The years of work that Allan and Joyce have spent on 
conservation issues, preparing reports, attending meetings, 
and hounding bureaucrats and politicians, demonstrate a 
dedicated labour seldom matched. The Club was always 
assured that, with the Reddochs representing us, our posi- 
tion would be presented with a measure of excellence and 
integrity that we could be proud of. 

Joyce and Allan’s work on orchids has been and is great- 
ly valued by naturalists and professionals alike. They have 
published papers on their work in both popular and scientif- 
ic venues culminating in the authoritative and thorough 
treatment of the orchids of the Ottawa District in The 
Canadian Field-Naturalist 111(1): 184 pages as a special 
issue in 1997. 

Over the years, Allan and Joyce have always operated as 


THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 


129 


a team, and as it is impossible to separate the activities of 
one from the other the logical decision was to give them a 
joint award. It is our pleasure to add Joyce and Allan 
Reddoch to our roster of Honorary Members. 


CONSERVATION AWARD — MEMBER: ALBERT DUGAL 

The cause of natural history conservation in the Ottawa 
area has been well served by OFNC members over the 
years. However, the road to habitat protection does not run 
smoothly and it takes individuals with a high degree of dili- 
gence and dedication to keep working on issues over the 
long term. Albert Dugal is one such individual. 

Most club members will be familiar with Albert’s long 
association with the Leitrim Wetlands. For over 20 years 
Albert has been working hard to save this ecologically sig- 
nificant area. His inventory (still ongoing) of Leitrim’s vas- 
cular plants has thus far recorded an astonishing 500 
species. Encouraged by Albert, local naturalists have 
recorded impressive lists of other organisms in the wetland 
complex. 

Albert has written extensively about the Leitrim Wet- 
lands (see Trail & Landscape 24(2), 26(3) and 27(4)) pro- 
viding in-depth information about a once little-known area. 
Whenever it seems that the issue has moved to the back of 
people’s consciousness, Albert returns with new informa- 
tion. In the last couple of years he has redoubled his efforts 
in the fight to save the remaining Leitrim wetlands from 
development. Working with the OFNC Conservation 
Committee as well as members of the Sierra Club, Albert 
has attended meetings, written letters, prepared briefs and 
given talks about the Leitrim site, raising people’s aware- 
ness of the issue and encouraging them to join with his 
efforts for preserving this area forever. 

Albert also continues working for protection for the for- 
mer South Gloucester Conservation Area, adding new 
species to his plant inventory, leading interested parties into 
the area, and providing expertise and background informa- 
tion for the Conservation Committee to aid in the commit- 
tee’s efforts. As well, he was a vital member of the 
Environmental Health Advisory Group (EHAG) under the 
Regional Municipality of Ottawa-Carleton. Albert has also 
acted as the Club’s liaison with other groups such as the 
Goulbourn Environmental Advisory Committee, where his 
botanical expertise was instrumental in persuading Goul- 
bourn Council to reject the residential subdivision plan for 
Westridge Phase III which encroached upon the Poole 
Creek Wetland. He also triggered the review which restored 
provincially significant status to the Fernbank Wetland. 

When the OFNC established the Conservation Award in 
1981 to recognize outstanding contributions to natural his- 
tory preservation in the Ottawa area, Albert Dugal was the 
first recipient. At that time his efforts in gaining protection 
for the Shaw Woods (successful) and for the South 
Gloucester Conservation Area (ongoing), as well as for his 
work in bringing attention to the unique natural values of 
Petrie Island were recognized. In 1991 Albert again won 
the Conservation Award, this time for his unstinting and 
persistent work on seeking protection for the Leitrim 
Wetlands. 

For the year 2000 Conservation Award for Members, we 
are delighted that once again, Albert Dugal is the recipient. 
His perseverance and tenacity over many, many years in 
working for the protection of our precious natural heritage, 
particularly the Leitrim Wetlands, is without doubt impres- 
sive. Albert Dugal is more than worthy of this award. 


730 


CONSERVATION AWARD — NONMEMBERS: DAVID MILLER 

From time to time, it is necessary to recognize outstand- 
ing contributions to the cause of natural history conserva- 
tion in the Ottawa Valley by outside governmental or non- 
governmental organizations, or individuals working within 
these organizations, and therefore not directly associated 
with the Ottawa Field-Naturalists’ Club. As originally con- 
ceived in 1992, the award for such organizations and indi- 
viduals was made only to non-members. We are making an 
exception in this case because, although David Miller has 
been a member of the Club since 1993, he has made his 
most significant contributions to conservation through his 
involvement with the Regional Municipality. 

For those who have worked on conservation concerns 
in the Ottawa area over the last decade, the name David 
Miller is a familiar one. David is an environmental plan- 
ner, formerly with the Regional Municipality of Ottawa- 
Carleton, now with the new city of Ottawa, and one of the 
more visible faces in the planning department. David is 
particularly skilful at translating ecological information 
and its implications into a form easily understood by 
bureaucrats and politicians. One recent example of this 
was his role in the acquisition of a significant portion of 
the South March Highlands, identified as a critical area 
both Provincially and Regionally. Because he was able to 
effectively demonstrate the tremendous conservation 
opportunity this presented, the powers-that-be acted 
quickly. The end result has been the largest single pur- 
chase of conservation lands in the region since at least the 
1960s. Public approval for this decision was overwhelm- 
ingly positive. 

Many club members will recall the Region’s Natural 
Environment Systems Strategy (NESS) project of the mid 
to late 1990’s which identified environmentally signifi- 
cant areas within the district. David’s participation in site 
investigations and supervision of this project was vital in 
ensuring that it received the appropriate attention and con- 
sideration. 

One of David’s most important qualities is his ability to 
listen to and understand the various positions on environ- 
mental issues. Representation of natural areas in the latest 
Regional Official Plan was substantially increased over 
previous versions, almost certainly due to David’s ability to 
present sound ecological concepts in such a way that they 
could be integrated within the planning process. 

David was also instrumental in setting up a Working 
Group to look at the feasibility of establishing a community 
based land trust organization, and to provide a forum to dis- 
cuss opportunities or partnership proposals prior to more 
formal land trust activities. The OFNC has been represent- 
ed on this committee from the beginning.Other local con- 
servation issues that have benefited from David’s participa- 
tion include the regional Wildlife Protocol and protection 
of Petrie Islands. It is therefore our pleasure to give to 
David Miller the club’s year 2000 Conservation Award for 
non-members. 


GEORGE MCGEE SERVICE AWARD: PHILIP MARTIN 

In organizations such as the Ottawa Field-Naturalists’ 
Club, run entirely by volunteers for the benefit of all mem- 
bers, there are always certain individuals who stand out by 
virtue of their special contribution of time and effort. The 
_ George McGee Service Award is given in recognition of 
the efforts of these individuals. This year, we are pleased to 
make that award to Philip Martin. 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Philip joined the OFNC in 1982 shortly after arriving in 
Ottawa and soon became an active participant in club 
affairs. Philip has been regularly leading walks since 1983. 
For many years, he led the spring flower walk in May, an 
astronomy outing in late August or early September, and a 
general autumn walk in Gatineau Park in early November. 
During the past few years, the number of walks in which he 
has participated as a leader has risen, and last year, he was 
involved in seven dealing with animals, shrubs, flowers, 
water plants, and even stars. Philip is always anxious to 
share his knowledge of plants and fungi with others, and 
his warm, genial manner has made him a favourite field trip 
leader. His interest in excursions led to his chairing the 
Excursions and Lectures Committee not once, but twice, 
from 1984-1986 and from 1997-2000. 

Upon leaving his post as Chair of the E&L Committee, 
Philip volunteered his time to the Conservation Committee, 
where he now a major player in the Club’s effort to save 
the Leitrim wetlands. 

Those of you who come out to these soirées know that 
Philip has been one of the organizers of the event and has 
overseen the photography and art exhibits for a number of 
years. 

Philip Martin is clearly a most fitting recipient of this 
year’s George McGee Service Award. 


ANNE HANES NATURAL HISTORY AWARD: ROBERT E. LEE 

Robert E. Lee has been a member of the Ottawa Field- 
Naturalists’ Club since 1981, having come up through the 
ranks of the Macoun Field Club, where his leadership con- 
tinues today to be an outstanding contribution to the natu- 
ralist community. 

Over the years he has built up an impressive store of 
knowledge of the natural history of the Ottawa District, 
much of it gained through independent investigation on his 
part. 

Rob possesses that combination of attributes that best 
exemplify a top-rate naturalist: a lively curiosity concern- 
ing the world of nature, the patience for careful observation 
and meticulous recording, and the tenacity to see a project 
through to its successful conclusion. He has demonstrated a 
fine talent for independent research, using imagination, 
resourcefulness and ingenuity to obtain and record biologi- 
cal data. 

His recent study of leopard frogs culminated in an excel- 
lent article published in Trail & Landscape. In this study, 
Rob produced new and hitherto unpublished information 
about the habits and movements of leopard frogs. 

Curious as to how many leopard frogs were summering 
on his land, Rob devised a method to distinguish one indi- 
vidual from another. Keen observation led him to the con- 
clusion that, as in fingerprints, no two frog patterns were 
precisely identical. From this basis, and working with the 
patterns of 175 different frogs, he cleverly devised a classi- 
fication system, with a key which permitted quick identifi- 
cation of individuals. Armed with this identification tool, 
he was able to follow the activities of individual frogs, and 
thus to document behaviour and migration patterns. His 
observations will surely lead to further research on these 
amphibians. 

Anne Hanes, the founding editor of Trail & Landscape, 
would be deeply satisfied with this well-conducted investi- 
gation by an amateur naturalist. His fellow naturalists are 
pleased to honour Robert E. Lee with the coveted Anne 
Hanes Natural History Award. 


2001 


MEMBER OF THE YEAR, 2000: SANDRA GARLAND 

Our club has a long history of fulfilling our goal of dis- 
seminating information relating to natural history through 
traditional means such as public lectures and print media. 
In recent times we have taken advantage of newly- 
developed electronic technologies to enhance our activities 
in this respect. 

In 1996, a notice posted in Trail and Landscape asked 
for volunteers to join a development team to produce a web 
site for the Ottawa Field-Naturalists’ Club. Sandra Garland 
was one of the individuals who stepped forward, although 
she had little previous experience with the Internet and was 
already devoting many volunteer hours to the Fletcher 
Wildlife Garden. She was duly appointed as Web Master. 
In the ensuing years, Sandy has spent countless hours 
developing original material, transforming information sub- 
mitted by the club’s committees and individual members 
into web pages, adding graphic images, and conducting 
maintenance on the site to keep it both fresh and timely. 

The site now has sections showcasing the activities of 
many of our club’s committees. For example, the club’s 
publications are highlighted, with on-line tables of contents 
and notes on the availability of books and special issues of 
our journals for purchase. Sandy has taken the time to dis- 
cuss the techniques for web page production with members 
of the Macoun Field Club, our junior naturalists, who now 
have their own sub-site complete with articles, photographs 
and images, and notes of their upcoming activities. 


THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 


731 


Extensive information is available on the Fletcher Wildlife 
Garden, denoting yet another of Sandy’s interests in the 
club, including an outstanding “virtual-tour” of the garden 
for web surfers who cannot visit Ottawa in person. 

Although regular web surfers may recognize pleasing 
page design, they probably do not appreciate the underlying 
skills which are required to produce the look-and-feel 
which they experience, nor how much time and effort is 
required to ensure that the page content is accurate and 
always current. This was acknowledged in the Autumn, 
2000 issue of Nature Canada, where Curt Schroeder, the 
Nature On-Line columnist, wrote: 

“Naturalist groups are getting better at using the 
power of the Internet to communicate to their mem- 
bers and the public. One of the best sites I have seen 
belongs to the Ottawa Field-Naturalists’ Club. The 
site... is well designed with excellent sub-section navi- 
gation... the Ottawa club has done a great job in pro- 
moting their activities on the web.” 

Such praise is well deserved by one very special individ- 
ual. The Ottawa Field-Naturalists’ Club wishes to express 
its Own appreciation for the exemplary dedication and 
enthusiasm of Sandy Garland, our Web Master, by present- 
ing her with the Member of the Year Award for 2000. 


AWARDS COMMITTEE IRWIN M. Bropo (Chair), 
CHRISTINE HANRAHAN, STEPHEN DARBISHIRE and 
SHEILA THOMSON 


News and Comment 


Froglog: Newsletter of the Declining Amphibian Populations Task Force (47, 48) 


Number 47, October 2001, contains: DAPTF seed grants 
— Endangered Chinhai Salamander colonising newly cre- 
ated breeding habitat (Max Spaareboom, Xie Feng and Fei 
Liang) — Amphibian decline in Venezuela: the state of 
knowledge (Cesar Luis Barrio Amoros) — Froglog Shorts 
— Publications of Interest. 

Number 48, December 2001, contains: Project Anuran 
(Emily Fitzherbert and Tony Gardner) — Disinfection of 
Ambystoma tigrinum virus (ATV) (Jesse Brunner and Tim 
Sesterhenn) — Is the canary singing? (A. Stanley Rand) — 
A hierarchical approach in studying the effects of an insecti- 
cide on amphibian communities (Michelle D. Boone, 
Christine M. Bridges, and Nathan E. Mills) — Report on the 
roundtable organized by DAPTF monitoring protocols work- 
ing group, 15 July 2001 — Assessing the conservation status 
of Australian frogs (IUCN — ASH Workshop 6-9 February 
2001) — Recent studies of European frogs reveal complexi- 
ties of the link with UV-B (Tim Halliday, DAPTF 


Marine Turtle Newsletter (94) 


The October 2001 issue, 32 pages, contains: EDITORIAL: 
MTN/MTM: Status Update (Brendan J. Godley and 
Annette C. Broderick — ARTICLES: Post-nesting move- 
ments of the green turtle, Chelonia mydas, nesting in the 
south of Bioko Island, Equatorial Guinea, West Africa (J. 
Tomas, A. Formia, J. Castroviejo, and J. A. Raga) — 
Commensal barnacles of sea turtles in Brazil (Leandro 
Bugoni, Ligia Krause, Alexandre Oliveira de Almeida and 
Alessandra Angelica de Padua Bueno — N Notes: Link 
between green turtles foraging in Brazil and nesting in 
Costa Rica? (Eduardo H. S. M. Lima and Sebastian 
Troeng) — Notes on the trade in marine turtle products in 
Bangladesh (M. Zahirul Islam) — An oft told story: man’s 
impact on green turtles in the Caribbean, , circa 1720 
(Michael G. Frick and Arnold Ross — Long distance trans- 
portation of turtle eggs from Sukabumi to Bali (Indonesia) 


Point Pelee Natural History News 1(3), 1(4) 


This newsletter for Point Pelee, Ontario, is edited by 
Alan Wormington (e-mail: wormington@juno.com). 
Editorial Assistants are M. Lea Martell and Matthew J. 
Smith. The wed site is www.wincom.net/~fopp/Natural_ 
History_News.htm 

1(3), Fall 2001, contains: ARTICLES: Biological Invasions 
of a Lake Erie (Joseph H. Leach) — Noteworthy Bird 
Records: September to November 2001 (Alan 
Worthington) — “Western” Solitary Sandpiper: New to 
Ontario (Alan Worthington) — IN THE FIELD 

1(4), Winter 2001, contains: ARTICLES: Wood Stork: 
New to Essex County (Alan Wormington) —- Noteworthy 
bird records: June to August 2001 (Alan Wormington) — 
Dainty Sulphur: New to Essex County (Henrietta T. 


International Director) — Farmland tree frog conservation 
project (Reprinted from the International Conservation 
Newsletter 9(2), 2001, of the Society for Wildlife and Nature 
(SWAN), Taiwan) — Froglog Shorts —-- Publications of 
Interest — Instructions to Contributors. 

Froglog is the bi-monthly newsletter of the Declining 
Amphibian Populations Task Force of The World 
Conservation Union (IUCN)/Species Survival Commission 
(SSC) and is supported by The Open University, The 
World Congress of Herpetology, The Smithsonian Insti- 
tution, and Harvard University. The newsletter is Edited by 
John W. Wilkinson, Department of Biological Sciences, 
The Open University, Walton Hall, Milton Keynes, MK7 
6AA, United Kingdom; e-mail: daptf@open.ac.uk. Funding 
for Froglog is underwritten by the Detroit Zoological 
Institute, P. O. Box 39, Michigan 48068-0039, USA. Frog- 
log can be accessed at http://www?2.open.ac.uk/biology/ 
froglog/ 


(Ismu Sutanto Suwelo, Saddon Silalahi, and Adang 
Gunawan — MEETING REPORTS: ANNOUNCEMENTS — BOOK 
REVIEWS — NeEws & LEGAL BRIEFS — RECENT 
PUBLICATIONS. 

The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, School of Biological Sciences, University of Wales, 
Swansea, SA2 8PP Wales, United Kingdom; e-mail 
MTN @swan.ac.uk; Fax +44 1792 295447. Subscriptions to 
the MTN and donations towards the production of MTN 
and its Spanish edition NTM [Noticiero de Tortugas 
Marinas] should be sent to Marine Turtle Newsletter c/o 
Chelonian Research Foundation, 168 Goodrich Street, 
Lunenburg, Massachusetts 01462 USA; e-mail 
RhodinCRF@aol.com; fax + 1 978 582 6279. MTN web- 
site is: <http://www.seaturtle.org/mtn/> 


O’ Neill) — Point Pelee Butterfly Count: August 11, 2001 
(Sarah Rupert) — Zabulon Skipper: New to Ontario and 
Canada (Jerry Ball, Thomas A. Hanrahan, and Paul R. 
Desjardins) — Clouded Skipper: New to Ontario and 
Canada (Henrietta O’ Neill and Alan Wormington) — I — 
IN THE FIELD — NEWS AND ANNOUNCEMENTS — UPCOMING 
EVENTS AND OUTINGS. 

Subscription rates are Canada: CAN $15 (one year) or 
$30 (two years); International: US $15 (one year) or $30 
(two years). Send payment (and e-mail address, optional) to 
The Friends of Point Pelee, 1118 Point Pelee Drive, Leam- 
ington, Ontario N8H 3V4. Issues will be mailed in March, 
June, September, and December, and back issues will be 
available for $15 per Volume/ $5 per issue (postage paid). 


32 


2001 


Canadian Species at Risk November 2001 


Issued by the Committee on the Status of Endangered 
Wildlife in Canada (COSEWIC), the list is 32 pages con- 
taining [1] About COSEWIC, (mandate, membership, defi- 
nitions); — [2] Summary Tables (COSEWIC species at 
risk, not at risk, and data deficient; results of November 
2001 COSEWIC meeting), [3] COSEWIC Lists (Explan- 
ation of symbols, Geographical occurrence and abbrevia- 
tions; List | Species designated in the five “risk” cate- 


NEWS AND COMMENT 


733 


gories, List 2. Species examined and designated in the NOT 
AT RISK category; List 3 Species examined and designated 
in the DATA DEFICIENT category) — [4] Record of Status 
Re-examinations — [5] List of name changes.s. 

It is available from COSEWIC Secretariat, Chief, 
Coleen Hyslop, c/o Canadian Wildlife Service, Environ- 
ment Canada, Ottawa, Ontario K1A 0H3. See Web site: 
http://www.cosewic.gc.ca 


Renew (Recovery of Nationally Endangered Wildlife) Report 11: 2000-2001 Annual Report 


Contents: Report from Co-Chairs — Highlights of 
2000-2001 |Of 107 Endangered and 76 Threatened 
species on the November 2000 COSEWIC list: 64 have 
recovery teams in place, 19 have approved plans, 6 more 
waiting approval, 25 have recovery plans or strategies in 
draft form, 68 are the focus of recovery efforts, 40 others 
are included in ecosystem recovery, 17 show stable or 
increasing population trends, $26.6 million expended on 
recovery (salaries + expenses), employment equivalent to 


about 129 salaried and 25 volunteer people working full- 
time, and 214 organizations have made financial contribu- 
tions] — Approaches to Recovery — Species at risk web 
site coverage [“Species search” web site: www. 
speciesatrisk.ga/Species/English/ SearchRequest. cfm — 
Status of recovery planning (summaries for 69 species and 
6 ecosystemstems/landscapes) — Financial contributors 
— Funding per species — Canadian Wildlife Directors 
Committee. 


Amphipacifica: Journal of Aquatic Systematic Biology 3(2) 15 November 2001 


CONTENTS: Donald E. McAllister (1934-2001): a tribute 
— The amphipod genus Allorchestes in the North Pacific 
region: systematics and distributional ecology (E. A. 
Hendrycks and E. L. Bousfield) — Systematics of the sub- 
terranean amphipod genus Stygobromus (Crangonyctidae) 
in western North America, with emphasis on the hubbsi 
group (Daqing Wang and John R. Holsinger). 

Amphipacifica is published by Amphipacifica Research 


Publications. Dr. E. L. Bousfield, Managing Editor, 
Ottawa; Dr. D. G. Cook, Technical Editor, Greely, Ontario. 
Subscriptions (4 numbers per volume) are renewable at $50 
(Can) or $40 (US) including surface postage. Author 
charges are $25 per printed page, subject to change. For 
further information please contact Dr. E. L. Bousfield, 
Managing Editor, 1710-1275 Richmond Road, Ottawa, 
Ontario, Canada K2B 8E3; e-mail: elbousf@magma.ca 


Conserving Borderline Species: A Partnership between the United States and Canada 


The Canadian Wildlife Service, Environment Canada, 
has published a 28-page booklet (including covers) on the 
“Framework for Cooperation between the U.S. Department 
of the Interior and Environment Canada in the Protection 
and Recovery of Wild Species at Risk” that was signed 7 
April 1997 “to facilitate cooperation in identifying and 
recovering shared species at risk”. 

“The goal of the Framework is to prevent populations of 
wild species shared by the United States and Canada form 
becoming extinct as a consequence of human activity, 
through the conservation of wildlife populations and ecosys- 
tems on which they depend.” The booklet text covers: 
Introduction (United States Fish and Wildlife Service and 
the Canadian Wildlife Service are responsible for imple- 


menting the Framework; since these have varying jurisdic- 
tions, the Framework does not consider issues involving 
marine mammals, fish, or sea turtles); Species accounts 
(Black-footed Ferret, Swift Fox, Woodland Caribou, 
Grizzly Bear, Whooping Crane, Piping Plover, Marbled 
Murrelet, Lake Erie Water Snake, Karner Blue Butterfly, 
Western Prairie Fringed Orchid); Table of Shared Species; 
What You Can Do (Report, Observe, Cooperate, Join, 
Inform, Do not disrupt, Ensure, Learn, Respect, Protect); 
For more information (in Canada contact Canadian Wildlife 
Service at 1-800-668-6767 or www. speciesatrisk.gc.ca; in 
United States contact U.S. Fish and Wildlife Service at 
1-800-344-WILD or visit [http://endangered.fws.gov]); Text 
of Framework. 


Index to Volume 115 


Compiled by Leslie Durocher 


Abalone, Northern, 555 
Pinto, 555 
Threaded, 555 
Abalone, Haliotis kamtschatkana, in Canada, Review of the 
Status of the Northern, 555 
Abies balsamea, 10,64,100,107,215,426,447 
grandis, 4 
lasiocarpa, 4,505 
Acantholumpenus mackayi, 116,566 
Accipiter cooperii, 61,346,415 
gentilis, 58,515 
striatus, 58,440 
Acer glabrum, 224 
macrophyllum, 224,455 
negundo, 257 
rubrum, 83,446 
saccharum, 40,64,446 
spicatum, 9 
Achillea millefolium, 2,467 
millefolium ssp. borealis, 96 
millefolium ssp. lanulosa, 96 
millefolium var. nigrescens, 96 
millefolium var. occidentalis, 96 
sibirica, 110 
Achmorphorus occidentalis, 433 
Achnanthes, 165 
Achnatherum hymenoides, 306 
Acipenser brevirostrum, 565 
fulvescens, 565 
medirostris, 565 
oxyrhynchus, 568 
transmontanus, 565 
Acremonium sp., 37 
Acrocheilus alutaceus, 565 
Actaea rubra, 93,110 
Actitis macularia, 272,434,440,488 
Adams, S. B. and C. A. Frissell. Thermal Habitat Use and 
Evidence of Seasonal Migration by Rocky Moun- 
tain Tailed Frogs, Ascaphus montanus, in Montana, 
251 
Adoxa moschatellina, 303,319 
orientalis, 319 
Aedes spp., 274 
communis, 280 
pionips, 274 
punctor, 280 
Aegolius funereus, 58,476 
Aegolius funereus, in Western Interior Alaska, Diets of 
Nesting Boreal Owls, 476 
Agastache utricifolia, 3 
Agelaius phoeniceus, 273,434,549 
Agropyron cristatum, 76,513 
smithii, 76 
trachycaulum, 406 
trachycaulum var. novae-angliae, 98 
trachycaulum var. trachycaulum, 98 
Agrostis alba, 76 
hyemalis, 218 


Aira praecox, 452,468 

Aix sponsa, 499,502 

Alaska, Diets of Nesting Boreal Owls, Aegolius funereus, 
in Western Interior, 476 

Alaska, First Record of an Anomalously White Killer 
Whale, Orcinus orca, Near St. Lawrence Island, 
Northern Bering Sea, 501 

Alaska, Three New Taxa and a Summary of the Mustard 
Family, Brassicaceae (Cruciferae), in Canada and, 
341 

Alasmidonta heterodon, 329 
undulata, 329 
varicosa, 329 

Alberta Border, A Significant New Record of the Pygmy 
Shrew, Sorex hoyi, on the Montana-, 513 

Alberta, Diet of the Prairie Rattlesnake, Crotalus viridis 
viridis, in Southeastern, 241 

Alberta, Hunting Methods and Success Rates of Gyr- 
falcons, Falco rusticolus, and Prairie Falcons, Falco 
mexicanus, Preying on Feral Pigeons (Rock Doves), 
Columba livia, in Edmonton, 395 

Alberta, Short-eared Owl, Asio flammeus, Attack on a 
Burrowing Owl, Athene cunicularia, in Suffield 
National Wildlife Area, 345 

Alberta, Tiger Salamander, Ambystoma tigrinum, Move- 
ments and Mortality on the Trans-Canada Highway 
in Southwestern, 199 

Alberta Wildlife Status Reports: (32 to 36), 542 

Alces alces, 174 

Alder, 9,58 
Red, 455 

Alectoria nigricans, 325 
ochroleuca, 325 

Alewife, 330,631 

Alfalfa, 317 

Alliaria, 341 

Allium acuminatum, 468 
cernuum, 451,467 

Allolumpenus hypochromus, 565 

Alnus spp., 9,58 
crispa, 476 
incana, 93 
rubra, 455 
rugosa, 93,110 

Alopecurus pratensis, 302,305 

Alopex lagopus, 22,515 

Alosa sp., 631 
aestivalis, 330,565 
pseudoharengus, 330,631 
sapidissima, 164,330,569 

Alternaria alternata, 37 

Alvo, R. and P. J. Blancher. Common Raven, Corvus 
corax, Observed Taking an Egg from a Common 
Loon, Gavia immer, Nest, 168 

Alyssum, 341 

Ambloplites rupestris, 150 

Amblystegium varium, 407 


734 


2001 


Ambystoma sp., 499 
tigrinum, 199,272,500 
tigrinum tigrinum, 202 

Ambystoma tigrinum, Movements and Mortality on the 
Trans-Canada Highway in Southwestern Alberta, 
Tiger Salamander, 199 

Ameirus nebulosus, 330,58 1,594,63 1,647 

Amelanchier spp., 9 
alnifolia, 110,257 
alnifolia ssp. compacta, 217 

American Birding Association Ludlow Griscom Award for 
Publications in Field Ornithology: W. Earl Godfrey, 
June 2000, 185 . 

American Society of Mammalogists, 82nd Annual Meeting 
of the, 542 

Americanus nebulosus, 155 

Amerorchis rotundifolia, 98 

Ammocrypta pellucida, 566 

Ammodramus caudacutus, 434 
leconteii, 273,434 

Ammodytes sp., 644 
americanus, 658,669 

Ampelomyces sp., 37 

Amphiagrion saucium, 402 

Amphipacifica: Journal of Aquatic Systematic Biology, 
542,733 

Amphipod, 644 

Anagallis arvensis, 232 

Anarhichas lupus, 570 
orientalis, 116,566 

Anas acuta, 272,434,499 
americana, 272,434 
bahamensis, 502 
castanea, 502 
clypeata, 434 
crecca, 434 
discors, 272,434 
gibberifrons, 502 
platyrhynchos, 272,434,499,502 
rubripes, 440,499 
strepera, 434 

Anas platyrhynchos, Feeding on a Wood Frog, Rana sylvat- 
ica, An Observation of a Mallard, 499 

Anas platyrhynchos, in Eastern South Dakota, Evidence for 
Double Brooding by a Mallard, 502 

Anderson, J. R., A. C. Nilssen and W. Hemmingsen. Use of 
Host-mimicking Trap Catches to Determine which 
Parasitic Flies Attack Reindeer, Rangifer tarandus, 
Under Different Climatic Conditions, 274 

Andromeda glaucophylla, 94 
polifolia ssp. glaucophylla, 94 
polifolia ssp. polifolia, 94 

Andropogon scoparium, 248 

Androsace septentrionalis, 94,301,318 

Anemone canadensis, 93,110 
mulifida, 93 
parviflora, 93 

Angelica lucida, 301,317 

Angelica, Seacoast, 317 

Anguilla rostrata, 330,631,647 

Anisakis spp., 122 

Anodonta implicata, 329 

Anquilla rostrata, 569 

Ant, Flying, 644 


INDEX TO VOLUME 115 


735 


Antennaria sp., 217 
densifolia, 301,320 
pulcherrima, 96,302,320 
rosea, 96 
Anthoxanthum odoratum, 451,461,467 
Anthus spragueti, 257,434 
Antilocapra americana, 257 
Anus acuta, 440 
Apeltes quadracus, 330 
Aphragmus, 341 
Apios americana, 446 
Apocynum androsamifolium, 110 
cannabinum, 216 
Apodemus sylvaticus, 473 
Apple, 54 
Aquila chrysaetos, 515 
Aquila chrysaetos, Attack on a Harlequin Duck, Histri- 
onicus histrionicus, in Northern Labrador, Obser- 
vation of a Golden Eagle, 515 
Aquilegea canadensis, 217 
Arabidopsis, 341 
Arabis, 341 
arenicola, 94 
boivinii, 301,312 
divaricarpa vat. divaricarpa, 313 
drummondii, 302,313 
holboellii, 110 
holboellii var. retrofracta, 302,313 
holboellii var. secunda, 302,313 
nuttallii, 302 
Aralia nudicaulis, 110,216 
Arbutus menziesii, 461 
Arbutus, 461 
Archibold, O.W., 106 
Archilochus colubris, 434 
Arctagrostis latifolia, 323 
Arctagrostis latifolia, and Arctic Lupine, Lupinus arcticus, 
in Upland Tundra on Herschel Island, Yukon 
Territory, Observations of Change in the Cover of 
Polargrass, 323 
Arctophila fulva, 302,305 
Arctostaphylos alpina var. alpina, 94 
alpina var. rubra, 94 
rubra, 94 
uva-ursi, 94,218 
uva-ursi ssp. adenotricha, 94 
uva-ursi Ssp. uva-ursi, 94 
Ardea herodias, 165,433,440,582,589,594 
Arenaria laterifolia, 93 
Argia fumipennis violacea, 404 
Armeria maritima, 467 
Armoracia, 341 
Arrow-grass, Seaside, 305 
Arrow-wood, Downy, 217 
Arrowhead, 77 
Artemisia abronatum, 110 
absinthium, 110 
borealis, 96 
campestris ssp. borealis, 96 
cana, 513 
canadensis, 96 
granatensis, 294 
michauxiana, 302,320 
tilesii, 90,302,320 


736 


tilesii ssp. elatior, 96 
tridentata, 2 
tripartita, 2 
Ascaphus montanus, 251 
truei, 251 
Ascaphus montanus, in Montana, Thermal Habitat Use and 
Evidence of Seasonal Migration by Rocky Moun- 
tain Tailed Frogs, 251 
Ascodichaena rugosa, 36 
Ascomycota, 35 
Ascomycotina, 36 
Asemichthys taylori, 565 
Ash, 248 
Black, 10 
Green, 257 
Mountain, 9 
Asio flammeus, 257,345,440,513 
otus, 345 
Asio flammeus, Attack on a Burrowing Owl, Athene cunic- 
ularia, in Suffield National Wildlife Area, Alberta, 
Short-eared Owl, 345 
Asodichaenetum, 36 
Aspen, 199 
Large-toothed, 446 
Quaking, 476 
Trembling, 2,9,100,107,216,224,257,413,426,446 
Asphodel, Common False, 310 
Northern False, 309 
Sticky False, 309 
Ass, Feral, 31 
Aster borealis, 96 
brachyactis, 96 
ciliolatus, 110,216 
conspicuus, 110 
ericoides, 110,407 
hesperius, 110 
Johannensis, 96 
laurentianus, 287 
longifolius, 96 
macrophyllus, 216 
pauciflorus, 409 
Aster, Ciliolate, 216 
Gulf of St. Lawrence, 287 
Large-leaved, 216 
Aster, in the Prince Edward Island National Park, Ger- 
mination Potential, Updated Population Surveys and 
Floral, Seed and Seedling Morphology of Symphyo- 
trichum laurentianum, the Gulf of St. Lawrence, 
287 
Asteromassaria, 38 
macrospora, 37 
Asterosporium, 38 
asterospermum, 37 
Astragalus adsurgens ssp. robustior, 302,316 
alpinus, 95,325 
cicer, 302,316 
miser, 2 
neglectus, 217 
umbellatus, 323 
Athene cunicularia, 257,345 
Athene cunicularia, in Suffield National Wildlife Area, 
Alberta, Short-eared Owl, Asio flammeus, Attack on 
a Burrowing Owl, 345 
Athya collaris, 434 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Athysanus, 341 
Atkinson, J. E., Reviews by, 521,711,717 
Atractylis arbuscula var. shizogynophylla, 294 
Atriplex cf. subspicata, 93 
patula, 410 
subspicata, 407 
Aureobasidium pululans, 37 
Avens, Yellow, 316 
Avian, 54 
Avocet, American, 434 
Aythya spp., 440 
affinis, 272,434 
ferina, 434 
valisineria, 434 


Bachelor’ s-button, 320 

Badger, 345 

Baetis, 142 

Baetisca, 142 

Baird, R. W. Status of Harbour Seals, Phoca vitulina, in 
Canada, 663 

Baird, R. W. Status of Killer Whales, Orcinus orca, in Can- 
ada, 676 

Balaena mysticetus, 567 

Balaenoptera acutorostrata, 118,568 
borealis, 568 
musculus, 567 
physalus, 116,121,567 

Balsamorhiza delioidea, 451,462 
sagittata, 451 

Balsamorhiza deltoidea (Asteraceae) in Canada, Status of 
the Deltoid Balsamroot, 451 

Balsamroot, Arrowleaf, 451 
Deltoid, 451,462 

Balsamroot, Balsamorhiza deltoidea (Asteraceae) in 
Canada, Status of the Deltoidm 451 

Banksiola, 142 

Barbarea, 341 

Barley, Meadow, 305 

Barnes, D. M. and A. U. Mallik. Effects of Beaver, Castor 
canadensis, Herbivory on Streamside Vegetation in 
a Northern Ontario Watershed, 9 

Barry, S. J., 257 

Barton, D. R., 68 

Bartramia longicauda, 272 

Bartsia alpina, 95 

Basil, Wild, 217 

Bass, 160 
Largemouth, 150 
Rock, 150 
Smallmouth, 150,330,620,629,650 
Striped, 330 

Bat, Big Brown, 421 
Evening, 421 
Hoary, 421 
Indiana, 420 
Little Brown, 421 
Northern Long-eared, 420 
Red, 205,420 
Silver-haired, 421 
Southeastern, 421 
Yellow-winged, 207 

Bat, Lasiurus borealis, in Mixed Mesophytic Forests of 
Kentucky, Possible Microclimate Benefits of Roost 
Site Selection in the Red, 205 


2001 


Bats in Southern Illinois, Timing of Pregnancy, Lactation, 
and Female Foraging Activity in Three Species of, 
420 

Bayberry, 447 

Beamish, R. J. Updated Status of the Vancouver Island 
Lake Lamprey, Lampetra macrostoma, in Canada, 
127 

Beamish, R. J., J. H. Youson, L. A. Chapman. Status of the 
Morrison Creek Western Brook Lamprey, Lampetra 
richardsoni, in Canada*, 573 

Bear, 249 
Black, 84,174,414 
Brown, 170 
Grizzly, 174,495 
Polar, 670 

Bear, Ursus arctos, Usurps Bison Calf, Bison bison, 
Captured by Wolves, Canis lupus, in Yellowstone 
National Park, Wyoming, Grizzly, 495 

Bearberry, Common, 218 

Beaver, 9,53 

Beaver, Castor canadensis, Herbivory on Streamside Vege- 
tation in a Northern Ontario Watershed, Effects of, 
= 

Bedstraw, Northern, 111 

Beech, American, 64,206,446 

Beetle, 644 

Bélanger, L., 75 

Bélanger, M., 99 

Beluga, 567 

Bennett, B., 301 

Benz, G. W., A. Kingman, and J. D. Borucinska. Gillnet 
Survival and Healing by a Porbeagle, Lamna nasus, 
506 

Berardius bairdi, 567 

Berry, Cloud, 275 

Berteroa, 341 

Betula alleghaniensis, 64,100,446 
glandulosa, 9 
nana, 275 
papyrifera, 9,57,107,219,426,446,476 
pubescens tortuosa, 275 
pumila var. glandulifera, 93 

Beyer, D. E., Jr., B. J. Roell, J. H. Hammill, and R. D. Earle. 
Records of Canada Lynx, Lynx canadensis, in the 
Upper Peninsula of Michigan, 1940-1997, 234 

Bib, 121 

Bilberry, 275 

Bindweed, Low, 217 

Birch, Dwarf, 275 
Mountain, 275 
Paper, 476 
River, 9 
White, 9,57,107,219,426,446 
Yellow, 64,100,446 

Bishop, C. A., 510 

Bison bison, 174,247,257 ,343,495 

Bison, 247,257,343,495 
Wood, 174 : 

Bison, Bison bison, Calf by a Wolf, Canis lupus, and Four 
Coyotes, Canis latrans, in Yellowstone National 
Park, Killing of a, 343 

Bison bison, Calf by a Wolf, Canis lupus, and Four Coy- 
otes, Canis latrans, in Yellowstone National Park, 
Killing of a Bison, 343 

Bison bison, Captured by Wolves, Canis lupus, in 


INDEX TO VOLUME 115 


737 


Yellowstone National Park, Wyoming, Grizzly 
Bear, Ursus arctos, Usurps Bison Calf, 495 
Bison Calf, Bison bison, Captured by Wolves, Canis lupus, 
in Yellowstone National Park, Wyoming, Grizzly 
Bear, Ursus arctos, Usurps, 495 
Bistort, 323 
Alpine, 311 
Bittern, American, 428 
Bittersweet, Climbing, 216 
Blackbird, Brewer’s, 273,435 
Red-winged, 273,434,549 
Rusty, 435 
Yellow-headed, 434,549 
Blackbirds, Xanthocephalus xanthocephalus, During 
Migration, Inability to Predict Geographic Origin of 
Yellow-headed, 549 
Blackfly, 644 
Bladderwort, Flat-leaved, 319 
Greater, 319 
Blancher, P. J., 168 
Blaney, C. S. and P. M. Kotanen. The Vascular Flora of 
Akimiski Island, Nanavut Territory, Canada, 88 
Blarina brevicauda, 53,248 
Bleier, W. J., 549 
Bloater, 565 
Blue-eyed Mary, Small-flowered, 3 
Bluebird, Mountain, 434 
Bluegrass, Cusick’s, 305 
Fowl, 2 
Kentucky, 76,461 
Sandberg’s, 3 
Bluestem, Little, 248 
Bluet, Bog, 404 
Boreal, 404 
Familiar, 404 
Hagen’s, 404 
Marsh, 404 
Northern, 404 
Orange, 404 
Rainbow, 404 
Spring Northern, 404 
Stream, 404 
Taiga, 404 
Tule, 404 
Vesper, 404 
Blush, Sea, 462 
Blysmus rufus, 97 
Boar, Wild, 248 
Bobcat, 235 
Bobolink, 273,432 
Bocaccio, 570 
Bog-laurel, 318 
Bombycilla cedrorum, 434 
Bonasa umbellus, 53,83,434 
Boreal Dip Net, The, 5(1), 186 
Borucinska, J.D., 506 
Bosmina, 134 
Botaurus lentiginosus, 433 
Botriosphaeria sp., 40 
Botritys cinerea, 37 
Botrychium lunaria, 92 
minganense, 92 
Botryosphaeria sp., 37 
Bouteloua gracilis, 513 
Bower, Virgin’s, 449 


738 


Bowman, J., C. V. Corkum, and G. J. Forbes. Spatial 
Scales of Trapping in Small-mammal Research, 472 

Bowman, J., G. J. Forbes and T. G. Dilworth. Distances 
Moved by Small Woodland Rodents within Large 
Trapping Grids, 64 

Boykinia richardsonii, 302,315 

Boykinia, Richardson’s, 315 

Bracken-fern, Eastern, 216 

Branta canadensis, 75,407.433,440,499 
canadensis canadensis, 76 
canadensis interior, 76 
canadensis maxima, 75 
canadensis moffitti, 75 

Brassica, 341 

Braya, 341 
humilis, 302,313 

Braya, Dwarf, 313 

Brazil, J..515 

Brisson, J., 34 

British Columbia, New Records of Land Snails from the 
Mountains of Northwestern, 223 

British Columbia, Predation on Nesting Woodpeckers in, 
413 

British Columbia, Status of the Stickleback Species Pair, 
Gasterosteus spp., in Hadley Lake, Lasqueti Island, 
579 

British Columbia, Status of the Stckleback Species Pair, 
Gasterosteus spp., in Paxton Lake, Texada Island, 
591 

British Columbia, Status of the Stickleback Species Pair, 
Gasterosteus spp., in the Vananda Creek watershed 
of Texada Island, 584 

Brochet d°’ Amérique, 597 
grand, 597 
maillé, 597 
vermiculé, 597 

Brochet d°’ Amérique, Esox americanus americanus, au 
Canada, Rapport sur la Situation du, 597 

Brome, 76,451.461 
Japanese, 2 
Rip-gut, 461.468 
Soft, 461.468 

Bromus spp., 451,461 
hordeaceus, 461.468 
inermis, 76 
japonicus, 2 
rigidus, 461,468 
tectorum, 3 

Broom, Scotch, 451,461 

Brown, R. S., 68 

Brownell, V. R., 402 

Bryant, J. E., 517 

Bubo virginianus, 58,345.354.434,440.480,5 13,515,543 

Bubo virginianus, Evidence of an Indirect Dispersal 
Pathway for Spotted Knapweed, Centaurea macu- 
losa, Seeds, via Deer Mice, Peromyscus manicula- 
tus, and Great Horned Owls, 354 

Bubo virginianus, Predation on Richardson’s Ground 
Squirrels, Spermophilus richardsonii, Great Horned 
Owl, 543 

Bucephala albeola, 434 
clangula, 434,440 

Buckthorn, Common, 218 
Glossy, 218 

Buckwheat, Parsnip-flowered, 2 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Budworm, Spruce, 58,99 
Buffalo, Bigmouth, 116,566 
Black, 116,566 
Buffaloberry, 257 
Bufflehead, 434 
Bufo americanus, 499 
boreas, 500 
bufo, 200 
Bug, True, 462 
Bullhead, Brown, 330,631,647 
Bulrush, Common Great, 309 
Great, 408 
Prairie, 407 
Bunchberry, 217 
Bunias, 341 
Bunker-Popma, K., 436 
Bunting, Lark, 244,273 
Snow, 440 
Bur-reed, Small, 304 
Giant, 77 
Burton, P. J., Review by, 719 
Bush, Antelope, 3 
Buteo jamaicensis, 346,434,481,515,546 
lagopus, 60 
lineatus, 61 
regalis, 257,346 
swainsoni, 257,346,513 
Buteo swainsoni, Prey and Reproduction in a 
Metapopulation Decline Among Swainson’s 
Hawks, 257 
Buttercup, Arctic, 312 
Butterwort, Common, 319 


Cactus, Brittle Prickly-Pear, 3 
Caddisfly, 142 
Cakile, 341 
Calamagrostis canadensis, 97 
stricta, 406 
stricta ssp. inexpansa, 98 
stricta ssp. stricta, 98 
Calamospiza, 244 
melanocoris, 273 
Calcarius ornatus, 273 
Calidris alba, 440 
alpina, 490 
bairdii, 272 
himantopus, 490 
maori, 408 
minutilla, 408.490 
pusilla, 490 
Callorhinus ursinus, 566 
Calopteryx, 404 
Caltha palustris, 93 
Calypso bulbosa var. americana, 98 
Calystegia spithamaea, 219 
spithamaea ssp. spithamaea, 217 
Camarosporium sp., 37 
Camas leitchtlinii. 461 
Camas, 452 
Common, 467 
Great, 461 
Meadow Death, 2 
Camassia spp., 452 
guamash, 467 
Camelina, 341 


2001 


Camnula pellucida, 268 

Campanula rotundifolia, 96,110,216 

Campbell, D., 510 

Campbell, R. R. Rare and Endangered Fishes and Marine 
Mammals of Canada: COSEWIC Fish and Marine 
Mammal Subcommittee Status Reports XIV, 564 

Campbell, R. R. Rare and Endangered Fishes and Marine 
Mammals of Canada: COSEWIC Fish and Marine 
Mammal Subcommittee Status Reports: XIII, 115 

Campostoma anomalum, 157,565 
anomalum michauxi, 157 
anomalum pullum, 157 
oligolepis, 158 

Campostoma anomalum, in Canada, Updated Status of the 
Central Stoneroller, 157 

Canada and Alaska, Three New Taxa and a Summary of 
the Mustard Family, Brassicaceae (Cruciferae), in, 
341 

Canada: COSEWIC Fish and Marine Mammal Subcom- 
mittee Status Reports XIV, Rare and Endangered 
Fishes and Marine Mammals of, 564 

Canada: COSEWIC Fish and Marine Mammal Subcom- 
mittee Status Reports: XIII, Rare and Endangered 
Fishes and Marine Mammals of, 115 

Canada: Insights from Long-term BBS Routes, Breeding 
Bird Declines in the Boreal Forest Fringe of 
Western, 425 

Canada Lynx, Lynx canadensis, in the Upper Peninsula of 
Michigan, 1940-1997, Records of, 234 

Canada, Meriam’s Shrew, Sorex merriami, and Preble’s 
Shrew, Sorex preblei: Two New Mammals for, | 

Canada, Rapport sur la Situation du Brochet d’ Amérique, 
Esox americanus americanus, au, 597 

Canada, Review of the Status of the Northern Abalone, 
Haliotis kamtschatkana, in, 555 

Canada, Status of Harbour Seals, Phoca vitulina, in, 663 

Canada, Status of Killer Whales, Orcinus orca, in, 676 

Canada, Status of Scouler’s Corydalis, Corydalis scouleri 
(Fumariaceae) in, 455 

Canada, Status of Snake-root Sanicle, Sanicula arctopoides 
(Apiaceae) in, 466 

Canada, Status of the Bluntnose Minnow, Pimephales nota- 
tus, in, 145 

Canada, Status of The Bridle Shiner, Notropis bifrenatus, 
in, 614 

Canada, Status of the Deltoid Balsamroot, Balsamorhiza 
deltoidea (Asteraceae) in, 451 

Canada, The Status of the Mira River Population of Lake 
Whitefish, Coregonus clupeaformis, in, 623 

Canada, Status of the Morrison Creek Western Brook 
Lamprey, Lampetra richardsoni, in, 573 

Canada, Status of the Purple Sanicle, Sanicula bipinnatifida 
(Apiaceae), in, 460 

Canada, Status of the Texada Stickleback Species Pair, 
Gasterosteus spp., in, 152 

Canada*, Status of the Weed Shiner, Notropis texanus, in, 
608 

Canada, Status of the White-Beaked Dolphin, Lagenor- 
hynchus albirostris, in, 118 

Canada, String and Net-Patterned Salt Marshes: Rare 
Landscape Elements of Boreal, 406 

Canada, The Vascular Flora of Akimiski Island, Nanavut 
Territory, 88 

Canada, Updated Status of the Central Stoneroller, 
Campostoma anomalum, in, 157 


INDEX TO VOLUME 115 


739 


Canada, Updated Status of the Northern Madtom, Noturus 
stigmosus, in, 138 
Canada, Updated Status of the Vancouver Island Lake 
Lamprey, Lampetra macrostoma, in, 127 
Canadian Species at Risk, 185, 541,733 
Canis aureus, 22,171 
familiaris, 483 
latrans, 19,31,99,170,343 
lupus, 19,22,174,179,235,343,495 
mesomelas, 22,171 
Canis latrans, in Yellowstone National Park, Killing of a 
Bison, Bison bison, Calf by a Wolf, Canis lupus, 
and Four Coyotes, 343 
Canis lupus, and Four Coyotes, Canis latrans, in Yellow- 
stone National Park, Killing of a Bison, Bison 
bison, Calf by a Wolf, 343 
Canis lupus, in Yellowstone National Park, Wyoming, 
Grizzly Bear, Ursus arctos, Usurps Bison Calf, 
Bison bison, Captured by Wolves, 495 
Canis lupus, “Standing Over” And “Hugging” in Wild 
Wolves, 179 
Canvasback, 434 
Capelin, 121,569,657 
Capreolus capreolus, 248 
Capsella, 341 
Caragana arborescens, 257,302,317 
Caragana, 257 
Common, 317 
Cardamine, 341 
pratensis ssp. angustifolia, 94 
Cardaria, 341 
Carduelis flammea, 440,477 
pinus, 435 
tristis, 435 
Carex sp., 217,275,325,505 
aenea, 302,306 
albo-nigra, 302,306 
aquatilis, 97 
aquatilis ssp. stans, 302,306 
atratiformis ssp. raymondii, 302,306 
atrofusca, 302,307 
aurea, 97,302,307 
bicolor, 302,307 
bigelowii, 97 
brunnescens, 97 
capillaris, 97 
capillaris ssp. capillaris, 302,307 
capillaris ssp. robustior, 302,307 
capitata, 97 
castanea, 218 
chordorrhiza, 97,302,307 
concinna, 97 
crawfordii, 302,307 
diandra, 97,302,307 
filifolia, 513 
flava, 97 
garberi, 97 
glareosa, 97 
gynocrates, 97 
inops, 461 
interior, 97 
intumescens, 217 
lachenalii, 302,307 
leptalea, 97 
limosa, 97 


740 


livida, 97,302,307 
mackenziei, 97,407 
magellanica, 97 
microglochin, 97 
norvegica, 97 
obtusata, 302,307 
oligosperma, 97 
paleacea, 97 
pellita, 218 
praticola, 97 
rariflora, 97,302,307 
richardsonii, 217 
rostrata, 97 
saxatilis, 97 
stylosa, 302,307 
subspathacea, 89 
tenuiflora, 302,308 
trisperma var. trisperma, 97 
utriculata, 97 
vaginata, 97 
viridula, 97 
Caribou, 50,174,274,326,495 
Barren-ground, 174 
Peary, i179 
Woodland, 174 
Carnegie, S. D., E. J. Urton, and D. L. Gummer. Short- 
eared Owl, Asio flammeus, Attack on a Burrowing 
Owl, Athene cunicularia, in Suffield National Wild- 
life Area, Alberta, 345 
Carp, Common, 336,484 
Carpodacus purpureus, 435 
Carroll, S. K., 420 
Garter i, C7420 
Carum carvi, 95 
Carya spp., 83,206,247 
Cassiope tetragona ssp. saximontana, 302,318 
Castilleja caudata, 302,318 
miniata, 302,319 
pallida ssp. candata, 319 
raupii, 95,302,319 
yukonis, 302 
Castor canadensis, 9,53 
Castor canadensis, Herbivory on Streamside Vegetation in 
a Northern Ontario Watershed, Effects of Beaver, 9 
Castostomus castostomus lacustris, 568 
Cat’s-ear, Hairy, 467 
Cataptrophorus semipalmatus, 272 
Catbird, Gray, 428,440 
Catfish, 155,160,581,594 
Flathead, 565 
Catharus fuscescens, 434 
guttatus, 434 
ustulatus, 434,477 
Catling, P.M. and V. R. Brownell. Biodiversity of Adult 
Damselflies (Zygoptera) at Eastern Ontario Gravel 
Pit Ponds, 402 
Catling, P. M., A. Sinclair, and D. Cuddy. Vascular Plants 
of a Successional Alvar Burn 100 Days After a 
Severe Fire and Their Mechanisms of Re-establish- 
ment, 214 
Catoptrophorus semipalmatus, 434 
Catostomus sp., 566 
commersoni, 330,336,63 1,647 
platyrhynchus, 565 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Cattail, 503 
Common, 304,408 
Narrow-leaved, 77,232 

Cedar, Eastern Red, 215. 
Eastern White, 100,219 

Celastrus scandens, 216 

Centaurea cyanus, 301,320 
diffusa, 3 
maculosa, 354 

Centaurea maculosa, Seeds, via Deer Mice, Peromyscus 
maniculatus, and Great Horned Owls, Bubo virgini- 
anus, Evidence of an Indirect Dispersal Pathway for 
Spotted Knapweed, 354 

Centraria nivalis, 325 

Centrocercus urophasianus, 212,257 

Cephenemyia trompe, 274 

Cerastium nutans, 301,311 

Cervus elaphus, 174,247,344,505 

Cervus elaphus, Droppings by Clark’s Nutcracker, Nuci- 
fraga columbiana, Opportunistic Foraging at Ameri- 
can Elk, 505 

Ceryle alcyon, 165,434,440 

Cestode, 150 

Cetraria cucullata, 325 
islandica, 325 

Chaenorrhinum minus, 218 

Chamaedaphne calyculata, 94 

Chapman, L.A., 573 

Char, Arctic, 568 

Chara, 141,582,587,593,619 

Charadrius melodus, 352,480 
semipalmatus, 490 
vociferus, 257,272,434,490 

Charadrius melodus, and Least Tern, Sterna antillarum, 
Productivity along the Missouri River in South 
Dakota, Influence of Predation on Piping Plover, 
480 

Charadrius melodus, from Natal Sites in Northwestern 
North Dakota, Initial Movements of Juvenile Piping 
Plovers, 352 

Cheatgrass, 3 

Chelydra serpentina, 182,510 

Chelydra serpentina, and Eggs in a Wood Chip Pile, 
Hyperthermia Induced Mortality of Gravid Snap- 
ping Turtles, 510 

Chelydra serpentina, Limb Mutilations in Snapping 
Turtles, 182 

Chenonetta jubata, 502 

Chenopodium capitatum, 93 
glaucum ssp. salinum, 93 
simplex, 216 

Cherry, Black, 83,248 
Choke, 9,217 
Pin, 9,217,447 

Chickadee, Black-capped, 417,434,440 
Boreal, 434,477 

Chicken, Domestic, 83 

Chickweed, 312 
Blinks, 311 
Nodding, 311 
Water, 311 

Chilipepper, 570 

Chinook, 687 3 

Chionoecetes opilio, 122 


2001 


Chipmunk, 547 
Eastern, 53,84 

Chiselmouth, 565 

Chlidonias niger, 434 

Chokecherry, 257 

Chondestes grammacus, 273 

Chordeiles minor, 434 

Chorispora, 341 

Choristoneura fumiferana, 58,99 

Chromagrion conditum, 402 

Chruszez, B., 199 

Chrysanthemum arcticum ssp. polare, 96 
leucanthemum, 90 

Chrysemys picta, 512 

Chrysocyon brachyurus, 22 

Chrysops nigripes, 277 

Chub, Creek, 164 
Gravel, 566 
Hornyhead, 157,565 
River, 157,565 
Silver, 565 

Chubbs, T. E., B. Mactavish, K. Oram, P. G. Trimper, K. 
Knox and R. I. Goudie. Unusual Harlequin Duck, 
Histrionicus histrionicus, Nest Site Discovered in 
Central Labrador, 177 

Chubsucker, Creek, 569 
Lake, 566 

Cicuta bulbifera, 95 
maculata, 95,110 
virosa, 95,302,318 

Cinquefoil, Norwegian, 316 

Circus cyaneus, 257,434,440,480,513 

Cirsium arvense, 110 
discolor, 220 
vulgare, 216 

Cisco, Bering, 565 
Blackfin, 116,566 
Deepwater, 566 
Least, 569 
Longjaw, 566 
Shortjaw, 116,566 
Shortnose, 116,566 
Spring, 565 

Cistothorus palustris, 434 

Cladina sp., 90,178,468,476 

Cladocera, 644 

Cladosporium sp., 36 
cladosporioides, 37 
herbarum, 37 

Clangula hyamalis, 440 

Clematis tangutica, 302,312 
virginiana, 449 

Clematis, Golden, 312 

Clemmys guttata, 182 
insculpta, 182 

Clethrionomys gapperi, 53,64,273,472 
rutilus, 477 ; 

Clevenger, A. P., M. McIvor, D. MclIvor, B. Chruszcz and 
K. Gunson. Tiger Salamander, Ambystoma tigrinum, 
Movements and Mortality on the Trans-Canada 
Highway in Southwestern Alberta, 199 

Cliff-brake, Slender, 304 

Clinopodium vulgare, 217 

Clinostomus elongatus, 116,565 


INDEX TO VOLUME 115 


741 


Cloudberry, 316 
Clover, 76,218 
Alsike, 232 
Bush, 248 
Sand, 468 
White, 218 
Club, Devil’s, 456 
Club-moss, Common, 303 
Clubrush, Tufted, 309 
Clupea harengus, 121,569,644,657,669 
pallasi, 356,569,668 
Cocconeis, 165 
Coccothrausters vespertinus, 435 
Coccyzus erythropthaimus, 434 
Cochlearia, 34} 
Cod, Atlantic, 115,121,565,669 
Cody, W. J., C. E. Kennedy and B. Bennett. New Records 
of Vascular Plants in the Yukon Territory III, 301 
Cody, W. J., Reviews by, 718,720,721 
Coeloglossum viride ssp. bracteatum, 302,310 
Coelomycetes, 35 
Coenagrion resolutum, 404 
Coho, 127 
Colaptes auratus, 413,434,440 
Coleoptera, 644 
Collinsia parviflora, 3,462 
Coltsfoot, 324 
Palmate, 456 
Sweet, 320 
Columba livia, 395 
Columba livia, in Edmonton, Alberta, Hunting Methods 
and Success Rates of Gyrfalcons, Falco rusticolus, 
and Prairie Falcons, Falco mexicanus, Preying on 
Feral Pigeons (Rock Doves), 395 
Columbine, Wild, 216, 
Columella edentula, 225 
Comandra livida, 95 
Condylura cristata, 53 
Coniothyrium fagi, 37 
Conringia, 341 
Conserving Borderline Species: A Partnership between the 
United States and Canada, 733 
Contropus borealis, 433 
sordidulus, 433 
Cook, F. R., Review by, 194 
Cooley, D. A., 323 
Coot, American, 272,428 
Copepod, 594 
Corallorhiza trifida, 98 
Coregonus sp., 565,623,643 
alpenae, 566 
artedi, 568 
clupeaformis, 568,623 ,636,669 
clupeaformis ssp., 566,623 
hoyi, 565 
huntsmani, 566,624,635 
Johannae, 566 
kiyi, 116,565 
laurettae, 565 
nasus, 569 
nigripinnis, 116,566 
reighardi, 116,566 
sardinella, 569 
zenithicus, 116,566 


742 


Coregonus clupeaformis, in Canada*, The Status of the 
Mira River Population of Lake Whitefish, 623 

Coregonus huntsmani*, Updated Status Report on the 
Endangered Atlantic Whitefish, 635 

Corkum, C. V., 472 

Cormorant, Double-crested, 176,433,440 
Great, 440 

Cornus canadensis, 95,110,217 
stolonifera, 9,174,217 

Coronopus, 341 

Corrigan, S. and A. W. Diamond. Northern Gannet, Morus 
bassanus, Nesting on Whitehorse Island, New 
Brunswick, 176 

Corvus brachyrhynchos, 84,168,273,346,434,440,483 
corax, 168,434,440,515 
corone, 168 

Corvus corax, Observed Taking an Egg from a Common 
Loon, Gavia immer, Nest, Common Raven, 168 

Corydalis aurea, 219,455 
aurea ssp. aurea, 216 
pauciflora, 455 
scouleri, 455 
sempervirens, 455 

Corydalis, Golden, 216 
Scouler’s, 455 

Corydalis, Corydalis scouleri (Fumariaceae) in Canada, 
Status of Scouler’s, 455 

Corydalis scouleri (Fumariaceae) in Canada, Status of 
Scouler’s Corydalis, 455 

Corylus cornuta, 9,109 

Coryphaenoides rupestris, 570 

Cotton-grass, 323 
Slender, 308 

Cottontail, Nuttall’s, 244,273 

Cottonwood, Narrow-Leaf, 543 

Cottus sp., 566 
bairdi, 142,568 
confusus, 566 
ricei, 565 

Coturnicops noveboracensis, 434 

Cowbird, Brown-headed, 273,427 

Coyote, 19,31,99,170,343,483,671 

Coyotes, Canis latrans, in Yellowstone National Park, 
Killing of a Bison, Bison bison, Calf by a Wolf, 
Canis lupus, and Four, 343 

Crab, 122 

Crambe, 341 

Cranberry, Red, 476 

Crane, Sandhill, 168,408,433 
Whooping, 406 

Crane’s-bill, Bicknell’s, 216 

Crappie, Black, 620 

Crataegus monogyna, 461 

Crayfish, 165 

Creeper, 329 

Crepis elegans, 302,320 
tectorum, 303,320 

Cress, Marsh Yellow, 315 

Créte, M., 99 

Crocuta crocuta, 497 

Crossbill spp., 440 

Crossman, E.J., 157,614 

Crotalus viridis viridis, 241 

Crotalus viridis viridis, in Southeastern Alberta, Diet of the 
Prairie Rattlesnake, 241 


THE CANADIAN FIELD-NATURALIST 


Vol. fi 


Crow, American, 84,168,273,346,434,440,483 
Hooded, 168 

Cryptogramma stelleri, 302,304 

Cuckcoo, Black-billed, 432 

Cuddy, D., 214 

Cumming, E. E., K. A. Hobson, and S. L. Van Wilgenburg. 
Breeding Bird Declines in the Boreal Forest Fringe 
of Western Canada: Insights from Long-term BBS 
Routes, 425 

Cuon alpinus, 31 

Curlew, Long-billed, 272 

Currant, Stink, 456 

Cyanocitta cristata, 84,434,440 

Cyclops, 134 

Cyclopterus lumpus, 570,657 

Cylindrocarpon destructans, 37 

Cylindrocladium sp., 37 

Cymbella, 165 

Cynomys, 543 
ludovicianus, 244 

Cynosurus echinatus, 452,461,468 

Cyprinella spiloptera, 615 

Cyprinus carpio, 336,484 

Cypripedium arietinum, 220 
passerinum, 98 

Cystophora cristata, 566 

Cytisus scoparius, 451,461,467 

Cytospora sp., 36 


Dace, Banff Longnose, 566 
Blacknose, 157 
Leopard, 565 
Nooksack, 566 
Pearl, 568 
Redside, 116,565 
Speckled, 565 
Umatilla, 116,565 
Dactylis arctica, 325 
glomerata, 452,461,468 
Damsel, Aurora, 404 
Eastern Red, 404 
Damselflies (Zygoptera) at Eastern Ontario Gravel Pit 
Ponds, Biodiversity of Adult, 402 
Dancer, Violet, 404 
Dandelion, Common, 216,321 
Danthonia californica, 461 
spicata, 216 
Daphnia, 134,594 
Darbyshire, S., 357 
Darter, Channel, 566 
Eastern Sand, 566 
Greenside, 566 
Least, 565 
Rainbow, 569 
River, 565 
Tesselated, 565 
Dasyscyphus sp., 37 
de Boer, D. H. and O. W. Archibold. Slumping Activity and 
Forest Vegetation Along the Northeastern Shore of 
Waskesiu Lake, Prince Albert National Park, 
Saskatchewan, 106 
Decapod, 644 
Decapoda, 644 
Deer, Black-Tailed, 283,468 
Coastal Black-tailed, 299 


2001 


Mule, 103,174,248 
Red, 248 
Roe, 248 
Sitka, 299 
White-tailed, 19,53,99,172,247,296 

Deer in the Yukon, Mule, Odocoileus hemionus, and 
White-tailed, O. virginianus, 296 

Deer, Odocoileus virginianus, in the Bas-Saint-Laurent 
region, Québec, Effects of Reopening Hunting on 
Survival of White-tailed, 99 

Deer, Odocoileus virginianus, in the Northwest Territories, 
An Unusual Record of a White-tailed, 172 

Deiluiis, G., 210 

Dekker, D. and J. Lange. Hunting Methods and Success 
Rates of Gyrfalcons, Falco rusticolus, and Prairie 
Falcons, Falco mexicanus, Preying on Feral Pigeons 
(Rock Doves), Columba livia, in Edmonton, 
Alberta, 395 

Delphinapterus leucas, 123,567 

Delphinium menziesii, 451 

Delphinus delphis, 122,567 

Dendroica coronata, 434,440,477 
magnolia, 434 
palmatum, 434 
penslyvanica, 434 
petechia, 434 

Dermacentor albipictus, 174 

Deschampsia cespitosa, 98 

Descurainia, 341 
incisa var. incisa, 301,313 
incisa var. viscosa, 341 
pinnata ssp. filipes, 314 
pinnata ssp. nelsonii, 302,314 
sophia, 314 
sophioides, 302,314 

Desert-parsley, Barestem, 467 
Fern-leaved, 3 

de Solla, S. R., D. Campbell, and C. A. Bishop. Hyper- 
thermia Induced Mortality of Gravid Snapping 
Turtles, Chelydra serpentina, and Eggs in a Wood 
Chip Pile, 510 

Deuteromycota, 35 

Devil, King, 446 

Dhole, 31 

Diamond, A.W., 176,349 

Diaptomous, 134 

Diarimella, 36 
laurentidae, 35 

Diatrypae sp., 36 

Diatrype disciformis, 37 
stigma, 37 

Dicrotendipes, 165 

Didelphis virginiana, 53 

Diervilla lonicera, 218 

Dilworth, T. G., 64 

Dinemasporium sp., 36 

Diospyros virginiana, 247 

Diplodia sp., 37 ; 

Diplotaxis, 341 

Dipodomys, 244,546 
ordii, 345 
stephensi, 64 

Diptera pupae, 644 

Discus whitneyi, 225 

Distichlis spicata, 447 


INDEX TO VOLUME 115 


743 


Dock, Willow, 311 

Dodecatheon hendersonii, 462 

Dog, African Wild, 22,171,497 
Domestic, 483 

Dogfish, Spiny, 569 

Dogtail, Hedgehog, 452,461,468 

Dogwood, Red-osier, 9,174,217 

Dolichonyx oryzivorus, 273,434 

Dolphin, Atlantic White-sided, 118,567 
Bottlenose, 122,567 
Common, 122,567 
Northern Right Whale, 567 
Pacific White-sided, 567 
Risso’s, 567 
Striped, 567 
White-Beaked, 118,567 
Whitenose, 118 

Dolphin, Lagenorhynchus albirostris, in Canada, Status of 
the White-Beaked, 118 

Donovan, M. T. and G. W. Douglas. Status of Snake-root 
Sanicle, Sanicula arctopoides (Apiaceae) in 
Canada, 466 

Douglas, G. W., 460,466 

Douglas, G. W. and J. A. Jamison. Status of Scouler’s 
Corydalis, Corydalis scouleri (Fumariaceae) in Can- 
ada, 455 

Douglas, G. W. and J. Pojar. Trillium ovatum Pursh variety 
hibbersonii (Taylor et Szczawinski) Douglas et 
Pojar, variety nova, 343 

Douglas, G. W. and M. Ryan. Status of the Deltoid 
Balsamroot, Balsamorhiza deltoidea (Asteraceae) in 
Canada, 451 

Douglas-fir, 1,461,467 
Interior, 413 

Dove, Mourning, 434 
Rock, 395 

Doves), Columba livia, in Edmonton, Alberta, Hunting 
Methods and Success Rates of Gyrfalcons, Falco 
rusticolus, and Prairie Falcons, Falco mexicanus, 
Preying on Feral Pigeons (Rock, 395 

Dowitcher, Short-billed, 487 

Draba spp., 25,341 
aurea, 94 
borealis, 302,314 
cinerea, 302,314 
densifolia, 301,314 
glabella, 94 
incana, 94 
lonchocarpa var. vestita, 301,314 
nemorosa Vat. leiocarpa, 302,314 
scotteri, 302,314 
stenopetala, 302,315 

Draba, Gray-leaved, 314 
Northern, 314 
Nuttall’s, 314 
Star-flower, 315 

Dracocephalum parviflorum, 216 

Dragonhead, Small-flowered, 216 

Dreissena polymorpha, 619 

Drepanocladus uncinatus, 325 

Driessena polymorpha, 334 

Dropseed, Sand, 3 

Drosera anglica, 93 
rotundifolia, 93 

Dryas integrifolia, 89,323 


744 


Dryas, 323 

Dryocopus pileatus, 413,434 

Duck, American Black, 440 
Australian Wood, 502 
Black, 499 
Harlequin, 177,440,515 
Pink-eared, 502 
Ring-necked, 434 
Ruddy, 428 
Wood, 499,502 ) 

Duck, Histrionicus histrionicus, in Northern Labrador, 
Observation of a Golden Eagle, Aquila chrysaetos, 
Attack on a Harlequin, 515 

Duck, Histrionicus histrionicus, Nest Site Discovered in 
Central Labrador, Unusual Harlequin, 177 

Dumetella carolinensis, 434,440 

Dumont, P., 614 

Dunlin, 487 

Dupontia fisheri ssp. psilosantha, 98 

Durella sp., 37 

Dyatrype spp., 36 


Eagle, Bald, 176,408,440,483,515 
Golden, 483,515 

Eagle, Aquila chrysaetos, Attack on a Harlequin Duck, 
Histrionicus histrionicus, in Northern Labrador, 
Observation of a Golden, 515 

Earle, R. D., 234 

Eaton, B. R. and Z. C. Eaton. An Observation of a Mallard, 
Anas platyrhynchos, Feeding on a Wood Frog, 
Rana sylvatica, 499 

Eaton, Z. C., 499 

Edge, T. A. and J. Gilhen. Updated Status Report on the 
Endangered Atlantic Whitefish, Coregonus hunts- 
mani*, 635 

Eel, 647 
American, 330,569,631 

Eelgrass, 644 

Eider, Common, 439 

Elderberry, Red, 224,456 

Eleocharis acicularis, 97 
palustris, 410 
smallii, 97 

Elk, 247,344 
American, 505 

Elk, Cervus elaphus, Droppings by Clark’s Nutcracker, 
Nucifraga columbiana, Opportunistic Foraging at 
American, 505 

Elliptio complanata, 331 

Elliptio, Eastern, 331 

Elm, American, 257 

Elodea canadensis, 603 

Elymus innovatus, 110 
mollis ssp. mollis, 98 
repens, 303,305 
spicata, 3 
trachycaulus, 447 
trachycaulus ssp. subsecundus, 302,305 
trachycaulus ssp. trachycaulus, 98 

Empetrum nigrum, 94 

Emphemerella, 142 

Empidonax alnorum, 433 
minimus, 433 

Emydoidea blandingii, 512 


THE CANADIAN FIELD-NATURALIST 


Vols 


Enallagma antennatum, 404 
aspersum, 402 
boreale, 404 
carunculatum, 404 
civile, 404 
cyathigerum, 403 
ebrium, 404 
exsulans, 403 
geminatum, 404 
hageni, 404 
signatum, 404 
vernale, 403 
vesperum, 402 
Endophragmiella sp., 37 
enfin , 597 
Enhydra lutris, 556,566 
épaulard, 676 
Ephemeroptera, 644 
Epicoccum nigrum, 37 
purpurascens, 37 
Epilobium angustifolium, 95 
ciliatum, 95 
palustre, 95 
Epinette noire, 44 
Epischura, 134 
Eptesicus fuscus, 421 
Equisetum arvense, 92,110,302,303 
fluviatile, 92 
hyemale, 110 
scirpoides, 218,302,303 
variegatum, 92 
variegatum ssp. variegatum, 302,304 
Equus africanus, 31 
Eremophila alpestris, 263,433 
Erethizon dorsatum, 53 
Erigeron acris ssp. politus, 302,320 
acris var. asteroides, 96 
elatus, 302,320 
lonchophyllus, 96 
philadelphicus, 110 
Erignathus barbatus, 566 
Erimystax x-punctata, 566 
Erimyzon oblongus, 569 
sucetta, 566 
Eriogonum heracleioides, 2 
Eriophorum angustifolium, 97 
gracile, 302,308 
russeolum, 97 
russeolum vat. albidum, 302,308 
tenellum, 97 
vaginatum, 97,323 
viridi-carinatum, 97 
Eruca, 341 
Erucastrum, 341 
Erysimum, 341 
cheiranthoides, 90 
Eschrichtius robustus, 567 
Esox americanus, 597,62 
americanus americanus, 115,564,597 
americanus vermiculatus, 568,597 
lucius, 597,620 
masquinongy, 597,620 
niger, 330,565,597,629,647,649 
Esox americanus americanus, au Canada*, Rapport sur la 
Situation du Brochet d’ Amérique, 597 


2001 


Etheostoma blennioides, 566 
caeruleum, 569 
microperca, 565 
olmstedi, 565 

Eubalaena glacialis, 567 

Euchalon, 569 

Euconulus fulvus, 225 

Eulachon, 355 

Eumetopias jubatus, 355,566,67 | 

Eumetopias jubatus, Cooperative Foraging by Steller Sea 
Lions, 355 

Euphagus carolinus, 435 
cyanocephalus, 273,435 

Euphrasia arctica, 95 
hudsoniana, 95 

Euthamia graminifolia, 232 

Eutrema, 341 

Eutypella sp., 37 

Exoglossum maxillingua, 565 


Fagus grandifolia, 40,64,206,446 

Fagus grandifolia dans une forét ancienne: bioindicateurs 
et structure mycosociologique, Biodiversité micro- 
fongique du, 34 

Fairy-candelabra, 318 

Falcipennis canadensis, 477 
canadensis franklinii, 84 

Falco columbarius, 58,346,395 ,434 
mexicanus, 346,395 
peregrinus, 346,395 
rusticolus, 395,515 
sparverius, 60,434,440,483 

Falco mexicanus, Preying on Feral Pigeons (Rock Doves), 
Columba livia, in Edmonton, Alberta, Hunting 
Methods and Success Rates of Gyrfalcons, Falco 
rusticolus, and Prairie Falcons, 395 

Falco rusticolus, and Prairie Falcons, Falco mexicanus, 
Preying on Feral Pigeons (Rock Doves), Columba 
livia, in Edmonton, Alberta, Hunting Methods and 
Success Rates of Gyrfalcons, 395 

Falcon, Peregrine, 346,395 
Prairie, 346,395 

Falcons, Falco mexicanus, Preying on Feral Pigeons (Rock 
Doves), Columba livia, in Edmonton, Alberta, 
Hunting Methods and Success Rates of Gyrfalcons, 
Falco rusticolus, and Prairie, 395 

Feldhamer, G. A., 247 

Feldhamer, G. A., T. C. Carter, and S. K. Carroll. Timing 
of Pregnancy, Lactation, and Female Foraging 
Activity in Three Species of Bats in Southern 
[llinois, 420 

Felis rufus, 235 

Fern, Braun’s Holly, 446 

Ferron, J., 43 

Fescue, Barren, 468 
Idaho, 3,461,468 
Red, 76,467 
Six-weeks, 3 

Festuca brachyphylla, 98 
idahoensis, 3,461,468 
rubra, 76,89,467 
rubra ssp. rubra, 98 
saximontana, 98 

Finch, Purple, 435 


INDEX TO VOLUME 115 


745 


Fir, Balsam, 10,57,64,100,107,219,426,447 
Grand, 4 
Subalpine, 4,505 
Fisher, 52 
Fishers, Martes pennanti, in Vermont, Fall Food Habits and 
Reproductive Condition of, 52 
Flake, L. D., 502 
Fleabane, Bitter, 320 
Flicker, Northern, 413,428,440 
Floater, Alewife, 329 
Brook, 329 
Eastern, 329 
Newfoundland, 329 
Triangle, 329 
Flook, D. R. and J. E. Bryant. A Remembrance of John 
Clifton Ward, 1921-1999, 517 
Flounder, Summer, 570 
Winter, 570 
Witch, 570 
Yellowtail, 570 
Fly, Gall, 354 
Muscoid, 274 
Nasopharyngeal Bot, 274 
Reindeer Warble, 274 
Flycatcher, Alder, 433 
Great-crested, 433 
Least, 428 
Olive-sided, 433 
Forbes, G. J., 64,472 
Forget-me-not, Blue, 3 
Forktail, Eastern, 404 
Forsyth, R. G. New Records of Land Snails from the Moun- 
tains of Northwestern British Columbia, 223 
Fox, Archie,22/515 
Blanford’s, 22 
Red, 22,77,170,480 
Silver, 170 
Fox, Vulpes vulpes, Family in Southern Ontario, Two 
Cases of Infanticide in a Red, 170 
Foxes, Vulpes vulpes, in Southern Ontario, Comparison of 
Parental Roles in Male and Female Red, 22 
Foxtail, Meadow, 305 
Fragaria vesca, 110 
virginiana, 94,110 
virginiana ssp. virginiana, 217 
Fragilaria, 165 
Fraxinus sp., 248 
nigra, 10,40 
pennsylvanica, 257 
Fringilla coelebs, 49 
Prissell, €)A., 251 
Frog, Boreal Chorus, 499 
Common, 500 
Leopard, 272 
Mink, 499 
Rocky Mountain Tailed, 251 
Tailed, 251 
Wood, 499 
Frog, Rana sylvatica, An Observation of a Mallard, Anas 
platyrhynchos, Feeding on a Wood, 499 
Froglog: Newsletter of the Declining Amphibian Popula- 
tions Task Force, 185,541,732 
Frogs, Ascaphus montanus, in Montana, Thermal Habitat 
Use and Evidence of Seasonal Migration by Rocky 
Mountain Tailed, 251 


746 


Fulica americana, 272,434 

Fundulus diaphanus, 116,336,565,644,647 
notatus, 565 

Fusarium oxysporum, 37 

Fusicoccum sp., 37 


Gadus morhua, 115,121,565,657,669 

Gadwall, 434 

Galbraith, D. A. Arboreal Courtship Behaviour by Eastern 
Garter Snakes, Thamnophis sirtalis sirtalis, in 
September in Bruce County, Ontario, 347 

Gale, Sweet, 311 

Galeopsis tetrahit ssp. bifida, 303,318 

Galium borealie, 110 
labradoricum, 96 
trifidum, 96 

Gallinago gallinago, 434,492 

Gallus gallus, 83 

Gammarus sp., 154 

Gannet, Morus bassanus, Nesting on Whitehorse Island, 
New Brunswick, Northern, 176 

Gannet, Northern, 176;440 

Gar, Spotted, 565 

Garbary, D., Review by, 526 

Garenne, 43 

Gasterosteus spp., 116,566,579,584,591 
aculeatus, 152,336,584 
aculeatus complex, 579,591 

Gasterosteus spp., in Canada, Status of the Texada Stickle- 
back Species Pair, 152 

Gasterosteus spp., in Hadley Lake, Lasqueti Island, British 
Columbia*, Status of the Stickleback Species Pair, 
579 

Gasterosteus spp., in Paxton Lake, Texada Island, British 
Columbia, Status of the Stickleback Species Pair, 
591 

Gasterosteus spp., in the Vananda Creek watershed of 
Texada Island, British Columbia*, Status of the 
Stickleback Species Pair, 584 

Gaultheria hispidula, 94 

Gavia immer, 168,433,440,582,589,594 
stellata, 440 

Gavia immer, Nest, Common Raven, Corvus corax, 
Observed Taking an Egg from a Common Loon, 
168 

Gawn, M., Reviews by, 189,381 

Geese in Southern Quebec, Establishment of a Breeding 
Population of Canada, 75 

Gende, S. M., J. N. Womble, M. F. Willson, and B. H. 
Marston. Cooperative Foraging by Steller Sea 
Lions, Eumetopias jubatus, 355 

Gentianella amarella ssp. acuta, 95 

Gentianopsis detonsa, 95 

Geocaulon lividum, 95 

Geochelone chilensis, 182 

Geoduck, 559 

Geothlysis trichas, 434 

Geranium bicknellii, 216 

Germander, American, 447 

Geum aleppicum ssp. strictum, 302,316 
macrophyllum vat. perincisium, 94 
triflorum, 3 

Gilhen, J., 635 

Ginns, J. and S. Darbyshire. A Tribute to Douglas Barton 
Osbourne Savile, 1909-2000, 357 


THE CANADIAN FIELD-NATURALIST 


Volk 105 


Giuliano, W.M., 52 
Giroux, J.-F., J. Lefebvre, L. Bélanger, J. Rodrigue and S. 
Lapointe. Establishment of a Breeding Population 
of Canada Geese in Southern Quebec, 75 
Glaucomys, 543 
sabrinus, 417 
Glaucomys volans, 248 
Glaux maritima, 89 
maritima ssp. obtusifolia, 94 
Gliocladium sp., 37 
Globicephala macrohynchus, 567 
malaena, 567 
Glyceria sp., 218 
striata, 98 
Glyptocephalus cynoglossus, 570 
Goby, Round, 142 
Godwit, Hudsonian, 487 
Marbled, 272 
Gold, Spring, 462,467 
Golden-Plover, American, 487 
Goldeneye, Common, 434,440 
Goldenrod, Early, 217 
Grass-leaved, 232 
Gray-stemmed, 217 
Hairy, 217 
Upland White, 217 
Zigzag, 449 
Goldfinch, American, 435 
Gomphonema, 165 
Goniodes spp., 212 
Goodchild, C. D. The Status of the Mira River Population 
of Lake Whitefish, Coregonus clupeaformis, in 
Canada, 623 
Goodwin, C. E., Reviews by, 708,715 
Goodyear, G., 515 
Goodyera repens, 98 
Goose, Canada, 75,89,407,433,440,499 
Giant Canada, 75 
Lesser Snow, 89 
Western Canada, 75 
Goosefoot, Maple-leaved, 216 
Gopher, Northern Pocket, 273 
Pocket, 245 
Goshawk, Northern, 58,515 
Gosse, J. W. and W. A. Montevecchi. Relative Abundances 
of Forest Birds of Prey in Western Newfoundland, 
Si) 
Goudie, R. I., 177 
Grackle, Common, 435,440 
Grampus griseus, 567 
Grant, P., Review by, 709 
Grass, 505 
Acuminate Panic, 217 
Colonial Bent, 218 
Glomerate Satin, 218 
Italian Rye, 305 
_ Manna, 218 
Narrow-leaved Panic, 217 
Needle-and-thread, 3 
Orchard, 452,461 
Poverty Oat, 216 
Quack, 305 
Reed Canary, 76,305 
Seashore Salt; 447 
Slender Wheat, 406,447 


2001 


Sweet Vernal, 461,467 
Wiry Panic, 218 
Witch, 232 
Wood Panic, 217 
Grebe, Eared, 433 
Horned, 272,433,440 
Pied-billed, 428 
Red-necked, 428,440 
Western, 433 
Grenadier, Rock, 570 
Grindelia integrifolia, 468 
Grosbeak, Evening, 435 
Rose-breasted, 434 
Groundnut, 446 
Groundsel, Balsam, 216 
Black-tipped, 321 
Boreal, 320 
Grouse spp., 440 
Ruffed, 53,83,434 
Sage, 212 
Sharp-tailed, 210,272,434 
Spruce, 84,477 
Grouse, Tympanuchus phasianellus, Ectoparasites in 
Lekking Sharp-tailed, 210 
Grus americana, 406 
canadensis, 168,408,433 
Grypus, 653 
Gull, Bonaparte’s, 434,440 
California, 272,434,480 
Franklin’s, 272,432 
Glaucous, 168 
Great Black-backed, 168,440 
Herring, 168,434,440 
Iceland, 440 
Lesser Black-backed, 168 
Ring-billed, 272,434,480 
Gum, Black, 83 
Gummer, D. L., 345 
Gumweed, Entire-leaved, 468 
Gunson, K., 199 
Gyrfalcon, 395,515 
Gyrfalcons, Falco rusticolus, and Prairie Falcons, Falco 
mexicanus, Preying on Feral Pigeons (Rock Doves), 
Columba livia, in Edmonton, Alberta, Hunting 
Methods and Success Rates of, 395 


Haddock, 121,570 
Haemaphysalis spp., 212 
Haematopota pluvialis, 277 
Hai, D: J.,.118 
Hairgrass, Early, 452,467 
Hake, Pacific, 668 
Red, 570 
Silver, 570,658 
Haliaeetus leucocephalus, 176,408,440,483,515 
Halibut, Atlantic, 570 
Greenland, 570 - 
Pacific, 570,687 
Halichoerus grypus, 566,653 
Halichoerus grypus, in the Northwest Atlantic*, The Status 
of the Grey Seal, 653 
Halimolobos, 341 
Haliotis kamtschatkana, 555 
kamtschatkana assimilis, 555 
rubra, 558 


INDEX TO VOLUME 115 


747 


Haliotis kamtschatkana, in Canada, Review of the Status of 
the Northern Abalone, 555 

Hammill, J. H., 234 

Hammill, M. O., 653 

Hanson, J. M. and A. Locke. Survey of Freshwater Mussels 
in the Petitcodiac River Drainage, New Brunswick, 
329 

Hare, 543 
Arctic, 179 
Snowshoe, 53,234,479,546 

Harebell, 216 

Harrier, Northern, 257,432,440,480,513 

Hatfield, T. Status of the Stickleback Species Pair, 
Gasterosteus spp., in Hadley Lake, Lasqueti Island, 
British Columbia*, 579 

Hatfield, T. Status of the Stickleback Species Pair, 
Gasterosteus spp., in the Vananda Creek watershed 
of Texada Island, British Columbia*, 584 

Hatfield, T. and J. Ptolemy. Status of the Stickleback 
Species Pair, Gasterosteus spp., in Paxton Lake, 
Texada Island, British Columbia*, 591 

Hawk, Cooper’s, 61,346,415 
Ferruginous, 257,346 
Red-shouldered, 61 
Red-tailed, 346,434,481,515,546 
Rough-legged, 60 
Sharp-shinned, 58,440 
Swainson’s, 257,272,346,513 

Hawk’ s-beard, Annual, 320 
Elegant, 320 

Hawks, Buteo swainsoni, Prey and Reproduction in a 
Metapopulation Decline Among Swainson’s, 257 

Hawthorn, Common, 461 

Hazel, 109 
Beaked, 9 

Heal-all, Common, 218 

Heath, J. P., G. Goodyear, and J. Brazil. Observation of a 
Golden Eagle, Aquila chrysaetos, Attack on a 
Harlequin Duck, Histrionicus histrionicus, in 
Northern Labrador, 515 

Hedysarum alpinum var. americanum, 95 
boreale ssp. mackenzii, 95 
mackenzii, 95 

Hemiptera, 644 

Hemipteran, 462 

Hemlock, Eastern, 85,446 
Western, 455 

Hemmingsen, W., 274 

Hemp, Indian, 216 

Hempnettle, 318 

Hendersonia sp., 37 

Hendricks, L. N., 505 

Hendricks, P. A Significant New Record of the Pygmy 
Shrew, Sorex hoyi, on the Montana-Alberta Border, 
513 

Hendricks, P. and L. N. Hendricks. Opportunistic Foraging 
at American Elk, Cervus elaphus, Droppings by 
Clark’s Nutcracker, Nucifraga columbiana, 505 

Heron, 582,589,594 
Great Blue, 165,433,440,483 

Herring, 121,356,657 
Atlantic, 569,644,669 
Blueback, 330,565 
Lake, 568 
Pacific, 569,668 


748 


Hesperian, Northwest, 226 

Hesperis, 341 

Hetaerina, 404 

Hexagenia, 142 
bilineata, 142 
limbata, 142 

Hicklin, P. and K. Bunker-Popma. The Spring and Fall 
Migrations of Scoters, Melanitta spp., at Con- 
federation Bridge in the Northumberland Strait 
between New Brunswick and Prince Edward Island, 
436 

Hickory, 83,206,247 

Hieracium piloselloides, 216,220,446 

Hierochloe odorata, 98 

Higgins, K. F., 480 

Hill, M. M. A., G. L. Powell, and A. P. Russell. Diet of the 
Prairie Rattlesnake, Crotalus viridis viridis, in 
Southeastern Alberta, 241 

Hippocampus stenolepis, 687 

Hippoglossoides platessoides, 570 

Hippoglossus hippoglossus, 570 
steolepis, 570 

Hippopotamus, 170 

Hippuris tetraphylla, 95 
vulgaris, 95 

Hirundo rustica, 433 

Histrionicus histrionicus, 177,440,515 

Histrionicus histrionicus, in Northern Labrador, Obser- 
vation of a Golden Eagle, Aquila chrysaetos, Attack 
on a Harlequin Duck, 515 

Histrionicus histrionicus, Nest Site Discovered in Central 
Labrador, Unusual Harlequin Duck, 177 

Hobson, K. A., 425 

Hoefs, M. Mule, Odocoileus hemionus, and White-tailed, 
O. virginianus, Deer in the Yukon, 296 

Holcus lanatus, 468 

Holly, False, 447 

Holm, E. and E. J. Crossman. Updated Status of the Central 
Stoneroller, Campostoma anomalum, in Canada, 
lsy7/ 

Holm, E. and N. E. Mandrak. Updated Status of the 
Northern Madtom, Noturus stigmosus, in Canada, 
138 

Holm, Es, P: Dumont J. Leclerc, (G- Rey-sandyEa se 
Crossman. Status of The Bridle Shiner, Notropis 
bifrenatus, in Canada, 614 

Holt, D. W. and W.N. Tiffney. A Yellow Wood Lily, 
Lilium philadelphicum, from Nantucket Island, 
Massachusetts, With Notes on its Occurrence in 
New England, 351 

Honeysuckle, 217 
Bush, 218 

Honkenya peploides, 93 

Hordeum brachyantherum, 303,305 
Jubatum, 98,407 

Horsetail, 111 

Horsetail, Field, 303 
Variegated, 304 

Houston, C. S., 257 

Houston, C. S., Reviews by, 192,193,195,382,387,388, 
523,524,527,703,724 

_ Houston, J. Status of the Bluntnose Minnow, Pimephales 
notatus, in Canada, 145 

Houston, J. Status of the Texada Stickleback Species Pair, 
Gasterosteus spp., in Canada, 152 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Houston, J. Status of the Weed Shiner, Notropis texanus, in 
Canada, 608 : 
Houstonia sp., 218 
Huggard, D. J., 1 
Hummingbird, Ruby-throated, 434 
Humpback, 122 
Hutchinsia, 341 
Hutchinson, J. T. and M. J. Lacki. Possible Microclimate 
Benefits of Roost Site Selection in the Red Bat, 
Lasiurus borealis, in Mixed Mesophytic Forests of 
Kentucky, 205 
Hyaena brunnea, 31 
vulgaris, 31 
Hyalella azteca, 165 
Hybognathus argyritis, 565 
hankinsoni, 157,569 
nuchalis regius, 565 
Hybomitra auripila, 277 
lundbecki, 277 
montana, 277 
nitidifrons confiformis, 277 
sexfasciata, 277 
Hydropsyche, 142 
scalaris, 142 
Hydrotaea sp., 274 
Hyena, Brown, 31 
Spotted, 497 
Striped, 31 
Hylocomium splendens, 325 
Hymenoptera, 644 
Hypericum sp., 216 
perforatum, 218, 
Hyperoodon ampullatus, 567 
Hyphomyceétes, 35 
Hypochaeris radicata, 467 
Hypoderma tarandi, 274 
Hypoxylon, spp., 36 
cohaerens, 37 
fragiforme, 37 
mammatum, 37 
rubiginosum, 37 
Hypoxylonetum, 36 
Hyppopotamus amphibius, 170 
Hyssop, Nettle-leaved Giant, 3 


Iagomorph, 53 
Iberis, 341 
Ichthyomyzon castaneus, 565 
fossor, 565 
Icterus galbula, 435 
Ictiobus cyprinellus, 116,566 
niger, 116,566 
Idahoa, 341 
Illex illecebrosus, 669 
Illinois, Timing of Pregnancy, Lactation, and Female For- 
_ aging Activity in Three Species of Bats in Southern, 
420 
Impatiens glandulifera, 446 
Indiangrass, 248 
Iris versicolor, 98 
Tsatis, 341 
Ischnura verticalis, 404 
Tsoetes braunii, 303 
echinospora, 303 
maritima, 301,303 


2001 


Ivy, Poison, 447 
Ixoreus naevius, 477 


Jackal, 22 
Golden, 31,171 
Silverbacked, 171 
Jackrabbit, White-tailed, 273 
Jacob’s Ladder, Annual, 3 
Jaeger, Long-tailed, 168 
Parasitic, 168 
Jamieson, G. S. Review of the Status of the Northern Aba- 
lone, Haliotis kamtschatkana, in Canada, 555 
Jamison, J. A., 455 
Jay, Blue, 84,434,440 
Gray, 434,477 
Jehl, J. R. Jr., and W. Lin. Population Status of Shorebirds 
Nesting at Churchill, Manitoba, 487 
John, R., Reviews by, 188,191,385,702,706,707,713 
Johnson, M. T. and C. J. Rothfels. The Establishment and 
Proliferation of the Rare Exotic Plant, Lythrum hys- 
sopifolia, Hyssop-leaved Loosestrife, at a Pond in 
Guelph, Ontario, 229 
Junco hyemalis, 440,434 
Junco, Dark-eyed, 434,440 
Juncus, 406,587,593 
alpinoarticulatus, 97 
alpinus, 97 
balticus var. littoralis, 97 
bufonius, 97,232,302,309 
castaneus, 97 
dudleyi, 232 
effusus, 232 
triglumis ssp. albescens, 97 
Juneberry, 217 
Junegrass, 3 
Juniper, 275,348 
Juniperus communis, 275,348 
communis var. depressa, 92 
communis var. montana, 92 
horizontalis, 92 
virginiana, 215 


Kalmia angustifolia, 178 
polifolia, 94,302,318 

Karagatzides, J. D., 210 

Karstenula sp., 37 

Kehoe, F. P. and A. W. Diamond. Increases and Expansion 
of the New Brunswick Breeding Population of 
Black-legged Kittiwakes, Rissa tridactyla, 349 

Kennedy, C. E., 301 

Kennedy, C. E., C. A. S. Smith, and D. A. Cooley. Obser- 
vations of Change in the Cover of Polargrass, 
Arctagrostis latifolia, and Arctic Lupine, Lupinus 
arcticus, in Upland Tundra on Herschel Island, 
Yukon Territory, 323 

Kentucky, Effects of Enclosed Large Ungulates on Small 
Mammals at Land Between The Lakes, 247 

Kentucky, Possible Microclimate Benefits of Roost Site 
Selection in the Red Bat, Lasiurus borealis, in 
Mixed Mesophytic Forests of, 205 

Kestrel, American, 60,434,440,483 

Killdeer, 257,272,432,488 

Killifish, Banded, 116,336,565,644,647 

King Devil, Glaucous, 216 

Kingbird, Eastern, 169,433 


INDEX TO VOLUME 115 


749 


Kingfisher, 582,589,594 
Belted, 165,434,440 

Kinglet, Ruby-crowned, 434 

Kingman, A., 506 

Kittiwake, Black-legged, 168,349 

Kittiwakes, Rissa tridactyla, Increases and Expansion of 
the New Brunswick Breeding Population of 
Black-legged, 349 

Kiyi, 116,565 

Knapweed, Diffuse, 3 
Spotted, 354 

Knapweed, Centaurea maculosa, Seeds, via Deer Mice, 
Peromyscus maniculatus, and Great Horned Owls, 
Bubo virginianus, Evidence of an Indirect Dispersal 
Pathway for Spotted, 354 

Knetter, J. M., R. K. Murphy, and R. S. Lutz. Initial 
Movements of Juvenile Piping Plovers, Charadrius 
melodus, from Natal Sites in Northwestern North 
Dakota, 352 

Knotweed, Eastern, 311 

Knox. Keo 077 

Kobresia simpliciuscula, 97 

Koeleria macrantha, 3,513 

Kogia breviceps, 567 
simus, 567 

Kotanen, P. M., 88 

Kruse, C. D., K. F. Higgins, and B. A. Vander Lee. Influ- 
ence of Predation on Piping Plover, Charadrius 
melodus, and Least Tern, Sterna antillarum, 
Productivity along the Missouri River in South 
Dakota, 480 


Labidesthes sicculus, 565 

Labrador, Observation of a Golden Eagle, Aguila chrysae- 
tos, Attack on a Harlequin Duck, Histrionicus his- 
trionicus, in Northern, 515 

Labrador, Unusual Harlequin Duck, Histrionicus histrioni- 
cus, Nest Site Discovered in Central, 177 

Lachance, S. Rapport sur la Situation du Brochet d’ Amé- 
rique, Esox americanus americanus, au Canada*, 
597 

Lacki, M. J., 205 

Lacroix, Cons 287 

Ladies’-tresses, Hooded, 310 

Lagenorhynchus acutus, 118,567 
albirostris, 118,567 
obliquidens, 567 

Lagenorhynchus albirostris, in Canada, Status of the 
White-Beaked Dolphin, 118 

Lagurus curtatus, 242,273 

Lambdina fiscellaria, 58 

Lamna nasus, 506 

Lamna nasus, Gillnet Survival and Healing by a Porbeagle, 
506 

Lamoureux, J., M. Créte and M. Bélanger. Effects of 
Reopening Hunting on Survival of White-tailed 
Deer, Odocoileus virginianus, in the Bas-Saint- 
Laurent region, Québec, 99 

Lampetra sp., 569 
ayresi, 576 
macrostoma, 127,565 
richardsoni, 116,128,573 
richardsoni marifuga, 566 
tridentata, 127,574 


750 


Lampetra macrostoma, in Canada, Updated Status of the 
Vancouver Island Lake Lamprey, 127 
Lampetra richardsoni, in Canada, Status of the Morrison 
Creek Western Brook Lamprey, 573 
Lampmussel, Eastern, 329 
Yellow, 329 
Lamprey, Chestnut, 565 
Copper River, 569 
Darktail, 565 
Lake, 565 
Morrison Creek, 116,566 
Morrison Creek Western Brook, 573 
Nass River, 569 
Northern Brook, 565 
Pacific, 127 
Sea, 128,330,632 
Vancouver Island Lake, 127 
Lamprey, Lampetra macrostoma, in Canada, Updated 
Status of the Vancouver Island Lake, 127 
Lamprey, Lampetra richardsoni, in Canada, Status of the 
Morrison Creek Western Brook, 573 
Lampsilis cariosa, 329 
radiata radiata, 329 
Lance, Sand, 658 
Lange, J.,395 
Langur, Hanuman, 170 
Lanius ludovicianus, 257 
Lapointe, S., 75 
Laportea canadensis, 449 
Larch, Eastern, 476 
Larix laricina, 89,476 
Lark, Horned, 263,432 
Larkspur, Menzies’, 451 
Larus argentatus, 168,434,440 
californicus, 272,434,480 
delawarensis, 272,434,480 
fuscus, 168 
glaucoides, 440 
hyperboreus, 168 
marinus, 168,440 
philadelphia, 434,440 
pipixcan, 272,434 
Lasionycteris noctivagans, 421 
Lasiosphaeria sp., 37 
Lasiurus borealis, 205,420 
cinereus, 421 
Lasiurus borealis, in Mixed Mesophytic Forests of Ken- 
tucky, Possible Microclimate Benefits of Roost Site 
Selection in the Red Bat, 205 
Lathyrus japonicus, 95,467 
japonicus var. maritimus, 461 
martimus var. pellitus, 95 
ochroleucus, 95,110 
palustris, 95 
venosus, 110 
Lauff, R., Review by, 721 
Laurel, Sheep, 178 
Lavia frons, 207 
Lebouria cooperi, 150 
Eeclere J 614 
Ledum groenlandicum, 94 
Leech, Smooth Turtle, 182 
| Lefebvre, J., 75 
Lemmiscus curtatus, 4 
Lemmus sibiricus, 477 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Lepidium, 341 
densiflorum var. densiflorum, 315 . 
densiflorum var. elongatum, 315 
densiflorum var. macrocarpum, 301,315 
ramosissimum, 303,315 

Lepisosteus oculatus, 565 

Lepomis spp., 150 
auritus, 116.566 
cyanellus, 565 
gibbosus, 155,336,594 
gulosus, 566 
humilis, 116,566 
megalotis, 565 

Leptodea ochracea, 329 

Leptodora, 134 

Leptographium sp., 37 

Lepus, 543 
americanus, 43,53,234,477,546 
arcticus, 43,179 
europaeus, 43 
timidus, 43 
townsendi, 273 

Lepus americanus, en semi-liberté, Influence de paramétres 
climatiques sur les patrons d’activité saisonniers et 
journaliers du liévre d’ Amérique, 43 

Lernaea cyprinacea, 150 

Lesage, V. and M. O. Hammill. The Status of the Grey 
Seal, Halichoerus grypus, in the Northwest Atlan- 
tic, 653 

Lespedeza sp., 248 

Lesquerella, 341 

Lestes congener, 404 
disjunctus disjunctus, 404 
dryas, 404 
eurinus, 404 
forcipatus, 404 
inaequalis, 404 
rectangularis, 404 
unguiculatus, 404 
vigilax, 404 

Lethenteron alaskense, 565 

Libertella faginea, 37 

Lichen, Caribou, 178 

Lien, J., D. Nelson, and D.J. Hai. Status of the White- 
Beaked Dolphin, Lagenorhynchus albirostris, in 
Canada, 118 

Ligula intestinalis, 150 

Ligusticum scoticum, 95 

Lilium philadelphicum, 217,351 
philadelphicum f. flaviflorum, 351 

Lilium philadelphicum, from Nantucket Isiand, Massachu- 
setts, With Notes on its Occurrence in New England, 
A Yellow Wood Lily, 351 

Lily-of-the-valley, Wild, 217 

Lily, Wood, 217 
Yellow Wood, 351 

Lily, Lilium philadelphicum, from Nantucket Island, 
Massachusetts, With Notes on its Occurrence in 
New England, A Yellow Wood, 351 

Lim, B. K., Review by, 525 

Limanda ferrugineus, 570 

Limnodromus griseus, 490 

Limosa fedoa, 272 
haemastica, 490 

Limosella aquatica, 302,319 


2001 
Lin, W., 487 
Lingcod, 570,668,687 


Linnaea borealis, 109 
borealis ssp. longiflora, 96 
borealis var. americana, 96 

Linum lewisii ssp. lepagei, 95 

Linz, G. M., 549 

Lion, 497 
Kalahari, 31 

Liriodendron tulipifera, 206 

Lissodelphis borealis, 567 

Listera borealis, 98 
cordata, 98 

Littorina littorea, 644 

Lobelia kalmii, 96 

Lobularia, 341 

Locke, A., 329 

Locoweed, Late Yellow, 317 
Pendant-pod, 317 
Scamman’s, 317 

Lolium spp., 232 
perenne, 462,468 
perenne ssp. multiflorum, 303,305 

Lomatium dissectum, 3 
nudicaule, 467 
utriculatum, 462,467 

Lomatogonium rotatum, 95 

Longspur, Chestnut-collared, 273 

Lonicera sp., 217 

Loon, 582,589,594 
Common, 168,433,440 
Red-throated, 440 

Loon, Gavia immer, Nest, Common Raven, Corvus corax, 
Observed Taking an Egg from a Common, 168 

Looper, Hemlock, 58 

Loosestrife, Hyssop-leaved, 229 
Purple, 232 
Winged, 232 

Loosestrife, at a Pond in Guelph, Ontario, The Estab- 
lishment and Proliferation of the Rare Exotic Plant, 
Lythrum hyssopifolia, Hyssop-leaved 229 

Lophius americanus, 570 

Lotus micranthus, 468 

Louse, 212 

Lousewort, Small-flowered, 319 
Whorled, 319 

Lucerne, Yellow, 317 

Lumpfish, 570,657 

Lunaria, 341 

Lundholm, J. T., Review by, 529 

Lupine, Arctic, 323 
Two-coloured, 468 

Lupine, Lupinus arcticus, in Upland Tundra on Herschel 
Island, Yukon Territory, Observations of Change in 
the Cover of Polargrass, Arctagrostis latifolia, and 
Arctic, 323 

Lupinus arcticus, 323. 
bicolor, 468 

Lupinus arcticus, in Upland Tundra on Herschel Island, 
Yukon Territory, Observations of Change in the 
Cover of Polargrass, Arctagrostis latifolia, and Arctic 
Lupine, 323 

eR S352 

Luxilis chyrsocephalus, 565 

Luzula multiflora, 461 


INDEX TO VOLUME 115 


751 


nivalis, 325 
parviflora, 97 
Lycaon pictus, 22,171,497 
Lycopodium clavatum var. monostachyon, 302,303 
Lymnatria dispar, 85 
Lynx canadensis, 234 
Lynx, Canada, 234 
Lynx canadensis, in the Upper Peninsula of Michigan, 
1940-1997, Records of Canada Lynx, 234 
Lynx, Lynx canadensis, in the Upper Peninsula of Michi- 
gan, 1940-1997, Records of Canada, 234 
Lythrum alatum, 232 
hyssopifolia, 229 
salicaria, 232 
Lythrum hyssopifolia, Hyssop-leaved Loosestrife, at a Pond 
in Guelph, Ontario, The Establishment and 
Proliferation of the Rare Exotic Plant, 229 
Lythrurus umbratilis, 565 


Mackerel, 658 
Atlantic, 570 

MacNulty, D.R., N. Varley, and D. W. Smith. Grizzly 
Bear, Ursus arctos, Usurps Bison Calf, Bison bison, 
Captured by Wolves, Canis lupus, in Yellowstone 
National Park, Wyoming, 495 

MacQuarrie, K. and H. Schaefer. New Plant Records for 
Prince Edward Island, 446 

Macrhybopsis storeriana, 565 

Mactavish, B., 177 

Madtom, Brindled, 138,566 
Carolina, 139 
Margined, 116,139,566 
Northern, 138,566 

Madtom, Noturus stigmosus, in Canada, Updated Status of 
the Northern, 138 

Magpie, Black-billed, 273,434 

Maianthemum canadense, 110,217 
trifolium, 98,302,309 

Malacoraja senta, 570 

Malacorhynchus membranaceus, 502 

Malaxis paludosa, 301,310 

Malcolmia, 341 

Mallard, 272,434,499,502 

Mallard, Anas platyrhynchos, Feeding on a Wood Frog, 
Rana sylvatica, An Observation of a, 499 

Mallard, Anas platyrhynchos, in Eastern South Dakota, 
Evidence for Double Brooding by a, 502 

Mallik, A. U.,9 

Mallotus villosus, 121,569,657 

Malus spp., 54 

Mammenga, P.W., 502 

Mandrak, N.E., 138 

Manitoba, Population Status of Shorebirds Nesting at 
Churchill, 487 

Maple, Bigleaf, 224,455 
Douglas, 224 
Manitoba, 257 
Mountain, 9 
Red, 83,446 
Sugar, 64,446 

Margariscus margarita, 568 

Margaritifera margaritifera, 329 

Marine Turtle Newsletter, 185,541,732 

Marmot, 31 
Alpine, 170 


{52 


Marmota spp., 31,543 
marmota, 170 

Marston, B.H., 355 

Marten, 235,515 
American, 62 

Martes americana, 235,515 
americana atrata, 62 
pennanti, 52 

Martes pennanti, in Vermont, Fall Food Habits and Repro- 
ductive Condition of Fishers, 52 

Martin, M. D., R. S. Brown, D. R. Barton, and G. Power. 
Abundance of Stream Invertebrates in Winter: 
Seasonal Changes and Effects of River Ice, 68 

Martin, Purple, 433 

Mary, Blue-eyed, 462 

maskinongé, 597 

Massachusetts, With Notes on its Occurrence in 
New England, A Yellow Wood Lily, Lilium phila- 
delphicum, from Nantucket Island, 351 

Massarina sp., 37 

Matricaria maritima ssp. phaeocephala, 96 

Matthiola, 341 

Mayfly, 142 

Mclvor, D., 199 

Mclvor, M., 199 

McNicholl, M. K., Review by, 187 

Meadow-sweet, Narrow-leaved, 218 

Meadowlark, Western, 257,434 

Meadow Rue, Alpine, 312 

Mech, L. D. “Standing Over” And “Hugging” in Wild 
Wolves, Canis lupus, 179 

Medicago falcata, 303,317 
lupulina, 216 
sativa, 303,317 

Medick, Black, 216 

Megaceryle alcyon, 582,589,594 

Megaptera novaeangliae, 116,122,567 

Melanitta fusca, 437 
nigra, 437 
perspicillata, 434,437 

Melanitta spp., at Confederation Bridge in the North- 
umberland Strait between New Brunswick and 
Prince Edward Island, The Spring and Fall Migra- 
tions of Scoters, 436 

Melanogrammus aeglefinus, 121,570 

Melanoplus sp., 268 

Meleagris gallopavo, 83 

Melilotus alba, 303,317 
officinalis, 76,303,317 

Melosira, 165 

Melospiza georgiana, 434 
lincolnii, 434 
melodia, 434,440 

Mentha arvensis, 110,302,318 

Menyanthes trifoliata var. minor, 95 

Mephitis mephitis, 53,84,480 

Merganser, Common, 440 
Red-breasted, 434,440 

Mergus merganser, 440 
serrator, 434,440 

Merlangius merlangius, 121 

Merlin, 58,346,395,434,440 

~ Merluccius bilinearis, 570,658 
productus, 668 

Mertensia maritima, 95 


THE CANADIAN FIELD-NATURALIST 


Vole aus 


Mesoplodon bidens, 116,567 
carlhubbsi, 567 
densirostris, 567 
mirus, 567 
stejnegeri, 567 
Mice, Peromyscus maniculatus, and Great Horned Owls, 
Bubo virginianus, Evidence of an Indirect Dispersal 
Pathway for Spotted Knapweed, Centaurea macu- 
losa, Seeds, via Deer, 354 
Michener, G.R. Great Horned Owl, Bubo virginianus, 
Predation on Richardson’s Ground Squirrels, Sper- 
mophilus richardsonii, 543 
Michigan, 1940-1997, Records of Canada Lynx, Lynx 
canadensis, in the Upper Peninsula of, 234 
Microchampignon, 35 
Microdiplodia sp., 36 
Microgadus tomcod, 330,570,647 
Microphysula cookei, 223 
ingersolli, 224 
Micropterus spp., 160 
dolomieu, 150,330,620,629,650 
salmoides, 150 
Microthyrium sp., 37 
Microtus sp., 244,273,354,543 
montanus, 4 
ochrogaster, 31 
pennsylvanicus, 53,242,477,513 
pinetorum, 248 
xanthognathus, 477 
Midshipman, Plainfin, 668 
Miller, E. H., 413 
Mink, 53,165,178,480,515 
Sea, 566 
Minnow, Bluntnose, 145,565,608 
Brassy, 157,569 
Bullhead, 145 
Cutlips, 565 
Eastern Silvery, 565 
Fathead, 145 
Pugnose, 565 
Slim, 145 
Western Silvery, 565 
Minnow, Pimephales notatus, in Canada, Status of the 
Bluntnose, 145 
Minotilta varia, 434 
Mint, Field, 318 
Minuartia biflora, 302,311 
dawsonensis, 93 
obtusiloba, 325 
Minytrema melanops, 566 
Mirounga angustirostris, 566,671 
Moehringia lateriflora, 93,302,31 
Mole, Star-nosed, 53 
Mollisia sp., 37 
Molothrus ater, 273,435 
Moneses uniflora, 94 
Monger, S., 343 
Monkfish, 570 
Monodon monoceros, 567 
Monolepis nuttalliana, 409 
Montana-Alberta Border, A Significant New Record of the 
Pygmy Shrew, Sorex hoyi, on the, 513 
Montana, Thermal Habitat Use and Evidence of Seasonal 
Migration by Rocky Mountain Tailed Frogs, Asca- 
phus montanus, in, 251 


2001 


Montevecchi, W. A., 57 
Montia fontana, 302,311 
Moose, 174 
Morone americana, 330,620,63 1,647 
saxatilis, 330 
Morus bassanus, 176 
Morus bassanus, Nesting on Whitehorse Island, New 
Brunswick, Northern Gannet, 176 
Moschatel, 319 
Mosquito, 274 
Moth, Gypsy, 85 
Mountain-heather, Four-angled, 318 
Mouse, 543 
Deer, 64,273,354,414,472 
Olive-Backed Pocket, 242 
Western Harvest, 64 
White-footed, 64,245,248,472 
Wood, 473 
Woodland Jumping, 53,64 
Moxostoma carinatum, 566 
dusquesnei, 116,566 
erythrurum, 565 
hubbsi, 566 
valenciennesi, 116,568 
Mucket, Tidewater, 329 
Mucor racemosus, 37 
Mudwort, Water, 319 
Mugwort, Aleutian, 320 
Michaux’s, 320 
Muhlenbergia glomerata, 218 
Mule, 296 
Mule, Odocoileus hemionus, and White-tailed, O. virgini- 
anus, Deer in the Yukon, 296 
Mullein, Common, 216 
Mulligan, G.A. Three New Taxa and a Summary of the 
Mustard Family, Brassicaceae (Cruciferae), in 
Canada and Alaska, 341 
Murphy, K.M., 343 
Murphy, R.K., 352 
Muskellunge, 620 
Muskox, 179,328 
Muskrat, 53,273,331,503 
Mussel, Zebra, 334,619 
Mussels in the Petitcodiac River Drainage, New 
Brunswick, Survey of Freshwater, 329 
Mustard, Tansy, 313 
Mustela sp., 273 
erminea, 416 
frenata, 415 
macrodon, 566 
vison, 53,165,178,480,515 
Myagrum, 341 
Myarchus crinitus, 433 
Myosotis alpestris, 325 
stricta, 3 
Myotis austroripareous, 421 
lucifugus, 421 
septentrionalis, 420 
sodalis, 420 
Myoxocephalus quadricornis, 116,566 
thompsoni, 566 
Myrica gale, 89,302,311 
pensylvanica, 447 
Myriophyllum, 619 
exalbescens, 603 


INDEX TO VOLUME 115 


753 


sibiricum, 95 
spicatum, 620 
verticillatum, 95 


Nagorsen, D. W., G. G. E. Scudder, D. J. Huggard, H. 
Stewart, and N. Panter. Meriam’s Shrew, Sorex 
merriami, and Preble’s Shrew, Sorex preblei: Two 
New Mammals for Canada, | 

Nanavut Territory, Canada, The Vascular Flora of Aki- 
miski Island, 88 

Napaeozapus insignis, 53,64 

Narwhal, 567 

Nasturtium, 341 

Navicula, 164 

Nectria sp., 36 
cinnabarina, 37 
coccinea, 37 
episphaeria, 37 
galligena, 37 

Needlegrass, Columbian, 2 

Nehalennia irene, 404 

Nelson, D., 118 

Nematode, 122,150,659 

Nemopanthus mucronata, 447 

Neogobius melanostomus, 142 

Neohendersonia kickxii, 37 

Neotoma, 244,546 

Neslia, 341 

Nesovitrea binneyana occidentalis, 225 

Nettle, Wood, 449 

New Brunswick and Prince Edward Island, The Spring and 
Fall Migrations of Scoters, Melanitta spp., at Con- 
federation Bridge in the Northumberland Strait 
between, 436 

New Brunswick Breeding Population of Black-legged 
Kittiwakes, Rissa tridactyla, Increases and Expan- 
sion of the, 349 

New Brunswick, Northern Gannet, Morus bassanus, 
Nesting on Whitehorse Island, 176 

New Brunswick, Status of the Sympatric Smelt (Genus 
Osmerus) Populations of Lake Utopia, 131 

New Brunswick, Survey of Freshwater Mussels in the 
Petitcodiac River Drainage, 329 

Newfoundland, Relative Abundances of Forest Birds of 
Prey in Western, 57 

Night-Heron, Black-crowned, 433 

Nighthawk, Common, 428 

Nilssen, A.C., 274 

Nitzschia, 164 

Nocomis, 162 
biguttatus, 157,565 
micropogon, 157,565 

North Dakota, Initial Movements of Juvenile Piping Plovers, 
Charadrius melodus, from Natal Sites in North- 
western, 352 

Northwest Territories, An Unusual Record of a 
White-tailed Deer, Odocoileus virginianus, in the, 
Ly2 

Notemigonus crysoleucas, 647 

Notorus miurus, 566 

Notropis anogenus, 565,618 
atherinoides, 569 
bifrenatus, 565,614 
buchanani, 565 
dorsalis, 565 


754 


heterodon, 565,608 
heterolepis, 608,617 
hudsonius, 569,608 
photogenis, 116,565 
rubellus, 565 
stramineus, 615 
texanus, 565,608 
volucellus, 142,615 
Notropis bifrenatus, in Canada, Status of The Bridle 
Shiner, 614 
Notropis texanus, in Canada, Status of the Weed Shiner, 
608 
Noturus eleutherus, 139 
flavater, 139 
flavipinnis, 139 
flavus, 139,569 
furiosus, 139 
gyrinus, 139 
insignis, 116,139,566 
miurus, 138,568 
munitus, 139 
placidus, 139 
stigmosus, 138,566 
Noturus stigmosus, in Canada, Updated Status of the 
Northern Madtom, 138 
Nucifraga caryocatactes, 505 
columbiana, 505 
Nucifraga columbiana, Opportunistic Foraging at 
American Elk, Cervus elaphus, Droppings by 
Clark’s Nutcracker, 505 
Numenius americanus, 272 
phaeopus, 490 
Nuphar, 587,593 
variegatum ou rubrodiscum, 603 
Nutcracker, Clark’s, 505 
European, 505 
Nutcracker, Nucifraga columbiana, Opportunistic Foraging 
at American Elk, Cervus elaphus, Droppings by 
Clark’s, 505 
Nuthatch, Red-breasted, 434 
Nyctea scandiaca, 168,345 
Nycticeius humeralis, 421 
Nycticorax nycticorax, 433 
Nymph, Dragonfly, 644 
Mayfly, 644 
Stonefly, 644 
Nymphea tuberosa, 603 
Nyssa sylvatica, 83 


O’ Neill, J., Review by, 386 
Oak, 247 
Bur, 215 
Chestnut, 83,206 
Garry, 451,461,467 
Northern Red, 83 
Red, 215 
White, 83,206 
Oatgrass, California, 461 
Oceanodroma leucorhoa, 440 
Ocella impi, 565 
Ochrotomys nuttalli, 248 
~ Octopus, 121 
Odobenus rosmarus, 655 
rosmarus rosmarus, 566 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Odocoileus hemionus, 103,174,248,296 
hemionus columbianus, 283 
hemionus sitkensis, 299 
hemionus ssp. columbianus, 468 
virginianus, 19,53,99,172,247,296 

Odocoileus hemionus, and White-tailed, O. virginianus, 
Deer in the Yukon, Mule, 296 

Odocoileus virginianus, Deer in the Yukon, Mule, Odo- 
coileus hemionus, and White-tailed, 296 

Odocoileus virginianus, in the Bas-Saint-Laurent region, 
Québec, Effects of Reopening Hunting on Survival 
of White-tailed Deer, 99 

Odocoileus virginianus, in the Northwest Territories, An 
Unusual Record of a White-tailed Deer, 172 

Odonata, 644 

Oldsquaw, 440 

Oncorhynchus spp., 165,569,668 
clarki, 128,154,582,589,591 
kisutch, 127,155,575,594 
mykiss, 165,631 
tshawytscha, 687 

Ondatra zebithicus, 53,273,331,503 

Onion, Hooker’s, 468 
Nodding, 451,467 

Ontario, Arboreal Courtship Behaviour by Eastern Garter 
Snakes, Thamnophis sirtalis sirtalis, in September 
in Bruce County, 347 

Ontario, Comparison of Parental Roles in Male and Female 
Red Foxes, Vulpes vulpes, in Southern, 22 

Ontario Natural Heritage Information Centre Newsletter 
6(1), 186 

Ontario, The Establishment and Proliferation of the Rare 
Exotic Plant, Lythrum hyssopifolia, Hyssop-leaved 
Loosestrife, at a Pond in Guelph, 229 

Ontario, Two Cases of Infanticide in a Red Fox, Vulpes 
vulpes, Family in Southern, 170 

Ontario Watershed, Effects of Beaver, Castor canadensis, 
Herbivory on Streamside Vegetation in a Northern, 
9 

Ophiodon elongatus, 570,668,687 

Oplopanax horridus, 456 

Oporornis agilis, 434 
philadelphia, 434 

Opossum, Virginia, 53 

Opsopoeodus emiliae, 565 

Opuntia fraglis, 3 

Oram, K., 177 

Orchid, 218 
Northern Bog, 310 
Northern Green, 310 

Orcinus, 679 
glacialis, 679 
nanus, 679 
orca, 123.501, 567 ,67,1;676 

Orcinus orca, in Canada*, Status of Killer Whales, 676 

Orcinus orca, Near St. Lawrence Island, Northern Bering 
Sea, Alaska, First Record of an Anomalously White 
Killer Whale, 501 

Orconectes, 165 

Oriole, Northern, 428 

Ortega, Y. K., 354 

Orthilia secunda, 94 | 

Oryctolagus cuniculus, 43 

Oryzopsis hymenoides, 306 


2001 


Oscillatoria, 165 

Osmerus sp., 566 
dentex, 132 
eperlanus, 132 
mordax, 131,330,631 
spectrum, 131,569 

Osmerus) Populations of Lake Utopia, New Brunswick, 
Status of the Sympatric Smelt (Genus, 131 

Osprey, 60,440 

Ottawa Field-Naturalists’ Club, April 27, 2001, Awards 
made at the Annual Soirée of the, 728 

Ottawa Field-Naturalists’ Club Committee Reports for 
2000, The, 535 

Ottawa Field-Naturalists’ Club, 9 January 2001, Minutes of 
the 122nd Annual Business Meeting of the, 534 

Otter, Sea, 556 

Ovenbird, 428 

Ovibos moschatus, 179,328 

Ovis canadensis, 3 

Owl, spp., 515 
Barred, 61 
Boreal, 58,476 
Burrowing, 257,345 
Great Horned, 58,345,354,434,440,480,5 13,515,543 
Long-eared, 345 
Short-eared, 257,345 ,432,440,513 
Snowy, 168,345 
Spotted, 61,547 

Owl, Asio flammeus, Attack on a Burrowing Owl, Athene 
cunicularia, in Suffield National Wildlife Area, 
Alberta, Short-eared, 345 

Owl, Athene cunicularia, in Suffield National Wildlife 
Area, Alberta, Short-eared Owl, Asio flammeus, 
Attack on a Burrowing, 345 

Owls, Aegolius funereus, in Western Interior Alaska, Diets 
of Nesting Boreal, 476 

Owls, Bubo virginianus, Evidence of an Indirect Dispersal 
Pathway for Spotted Knapweed, Centaurea macu- 
losa, Seeds, via Deer Mice, Peromyscus manicula- 
tus, and Great Horned, 354 

Owl, Bubo virginianus, Predation on Richardson’s Ground 
Squirrels, Spermophilus richardsonii, Great Horned, 
543 

Oxytropis campestris ssp. jordalii, 302,317 
deflexa ssp. foliolosa, 302,317 
deflexa ssp. sericea, 302,317 
nigrescens, 325 
scammaniana, 302,317 
viscida var. hudsonica, 95 

Oxyura jamaicensis, 434 


Paddlefish, 566 
Paecilomyces sp., 40 
Paintbrush, Common Red, 319 
Raup’s, 319 
Scarlet, 319 ; 
Pandion haliaetus, 60,440 
Panicum sp., 218 
acuminatum var. acuminatum, 217 
capillare, 232 
flexile, 218 
linearifolium, 217 
philadelphicum, 217 
virgatum, 248 


INDEX TO VOLUME 115 


755 


Panopea abrupta, 559 

Panter, N., | 

Panthera leo, 31,497 

Papaver spp., 325 

Paralichthys dentatus, 570 

Parascalops breweri, 53 

Parelaphostrongylus tenuis, 174 

Parnassia palustris, 94,110 
parviflora, 94 

Parrya, 341 
arctica, 302,315 
nudicaulis, 325 

Partridge, Gray, 272,434 

Parus spp., 416 
atricapillus, 434,440 
hudsonicus, 434 

Passer domesticus, 273,435 

Passerculus sandwichensis, 273,434,440 

Pea, Beach, 461,467 

Pearlshell, Eastern, 329 

Pearlwort, Arctic, 312 

Pearson, D. E. and Y. K. Ortega. Evidence of an Indirect 
Dispersal Pathway for Spotted Knapweed, Cen- 
taurea maculosa, Seeds, via Deer Mice, Pero- 
myscus maniculatus, and Great Horned Owls, Bubo 
virginianus, 354 

Pedicularis capitata, 325 
groenlandica, 96 
lanata, 325 
macrodonta, 302,319 
parviflora, 96 
sudetica, 96 
verticillata, 302,319 

Pelecanus erythrorhynchos, 408,433 

Pelican, American White, 408,433 

Pendantgrass, 305 

Penicillium sp., 37 

Penny, J. L. and G. W. Douglas. Status of the Purple 
Sanicle, Sanicula bipinnatifida (Apiaceae), in Can- 
ada, 460 

Pennycress, Field, 315 

Pennyroyal, False, 217 

Penstemon confertus, 3 

Penstemon, Yellow, 3 

Peppergrass, Branched, 315 

Perca flavescens, 150,336,620,63 1,647 

Perch, Pacific Ocean, 570 
White, 330,620,63 1,647 
Yellow, 150,336,620,63 1,647 

Percina copelandi, 566 
shumardi, 565 

Perdix perdix, 272,434 

Perisoreus canadensis, 434,477 

Periwinkle, 644 

Perognathus fasciatus, 245 
maniculatus, 242 

Peromyscus sp., 53,249,273,543 
leucopus, 64,248,472 
maniculatus, 64,244,354,414,472 

Peromyscus maniculatus, and Great Horned Owls, Bubo 
virginianus, Evidence of an Indirect Dispersal Path- 
way for Spotted Knapweed, Centaurea maculosa, 
Seeds, via Deer Mice, 354 

Persimmon, 247 


756 THE CANADIAN FIELD-NATURALIST Vol. 115 


Petasites spp., 324 Pike, Northern, 620 
frigidus, 216 Pimephales notatus, 145,565,608,615 
frigidus ssp. frigidus, 302,320 promelas, 145 
frigidus var. palmatus, 456 tenellus, 145 
sagittatus, 96 vigilax, 145 
Petrel, Leach’s Storm, 440 Pimephales notatus, in Canada, Status of the Bluntnose 
Petromyzon marinus, 128,330,632 Minnow, 145 
Phalacrocorax auritus, 176,440 Pimpernel, 232 
carbo, 440 Pine, 85,413 
Phalaris arundinacea, 76,302,305 Eastern White, 219 
Phalarope, Red-necked, 487 Jack, 10,215,347,426 
Wilson’s, 272 Lodgepole, 4,199,344,495,505 
Phalaropus lobatus, 492 Ponderosa, | 
tricolor, 272 Virginia, 248 
Phalocrocorax auritus, 433 White, 446 
Pheucticus ludovicianus, 434 Pinguicula vulgaris, 96 
Phialostromatinae, 36 vulgaris ssp. vulgaris, 302,319 
Philonotis fontana, 325 Pinson, 49 
Phleum pratense, 76 Pintail, 499 
Phoca groenlandica, 570 Northern, 272,432,440 
hispida, 567 White-cheeked, 502 
largha, 663 Pinus spp., 85 
vitulina, 568,656,663 banksiana, 10,92,215,347,426 
vitulina concolor, 566 contorta, 4,199,344,413,495,505 
vitulina mellonae, 566,663 ponderosa, 1,413 
vitulina richardsi, 566,663 strobus, 215,446 
Phoca vitulina, in Canada*, Status of Harbour Seals, 663 virginiana, 248 
Phocoena phocoena, 116,123,567 Pipilo erythrophthalmus, 85,434 
Phocoenoides dalli, 567 Pipistrelle, Eastern, 420 
Phoebe, Eastern, 433 Pipistrellus subflavus, 420 
Say’s, 433 Pipit, Sprague’s, 257,434 
Phoma sp., 37 i Placobdella parasitica, 182 
Phomopsis oblonga, 37 Plaice, American, 570 
Phragmites australis, 408 Plankton, 644 
Phryganea, 142 Plantago lanceolata, 462 
Phyllodoce X intermedia, 301,318 major, 90,232 
Phyllosticta sp., 37 maritima, 95,407,467 
Physaria, 341 Plantain, Common, 232 
Physella, 165 Narrow-leaved, 462 
Physeter macrocephalus, 567 Seaside, 467 
Pica pica, 273,434 Platanthera sp., 218 
Picea engelmannii, 4,505 aquilonis, 302,303,310 
engelmannii x glauca, 413 dilatata, 98 
glauca, 10,57,64,89,100,107,199,215,426,447,476 hyperborea, 98,302,310 
glauca var. glauca, 92 obtusata, 98,302,310 
glauca vat. porsildii, 92 Plecoptera, 644 
mariana, 10,44,57,64,89,178,426,446,476 Plectritis congesta var. congesta, 462 
rubens, 446 Plectrophenax novalis, 440 
sitchensis, 455 Pleospora sp., 37 
Pickerel, Chain, 330,565,629,641,647 Plover, Piping, 352,480 
Grass, 568,620 Semipalmated, 487 
Red-finned, 598 Plover, Charadrius melodus, and Least Tern, Sterna antil- 
Redfin, 115,564,565,620 larum, Productivity along the Missouri River in 
Picoides arcticus, 416 South Dakota, Influence of Predation on Piping, 
gossypinus, 417 480 
leucopus, 417 Plovers, Charadrius melodus, from Natal Sites in North- 
pubescens, 413,434 western North Dakota, Initial Movements of Juvenile 
villosus, 413,434 Piping, 352 
Pig, Sea, 653 Pluvialis dominica, 488 
Pigeon, Feral, 395 Poa spp., 325,505 
Pigeons (Rock Doves), Columba livia, in Edmonton, alpina, 98 
Alberta, Hunting Methods and Success Rates of arctica ssp. arctica, 98 
Gyrfalcons, Falco rusticolus, and Prairie Falcons, cusickii, 303,305 


Falco mexicanus, Preying on Feral, 395 glauca, 98 


2001 


palustris, 2,98 
pratensis, 76,461 
pratensis ssp. alpigena, 98 
secunda, 3,513 
Poacher, Pixy, 565 
Podiceps auritus, 272,433,440 
grisegena, 433,440 
nigricollis, 433 
Podilymbus podiceps, 433 
Poecile atricapilla, 417 
hudsonica, 477 
Pogonatum alpinum, 325 


Point Pelee Natural History News, 542,732 


Poison-ivy, 216 
Pojar, J., 343 
Polargrass, 323 


INDEX TO VOLUME 115 7 


gramineus, 97 
richardsonii, 97,302,304 
zosteriformis, 302,304 
Potamyia flava, 142 
Potato, 447 
Potemonium micranthum, 3 
Potentilla anserina var. anserina, 94 
anserina var. groenlandica, 94 
crantzii, 89 
fruticosa, 95 
multifida, 95 
nivea, 95 
norvegica, 90,302,316 
palustris, 95 
pensyvanica var. pectinata, 95 
pulchella, 95 


Polargrass, Arctagrostis latifolia, and Arctic Lupine, rubricaulis, 302,316 
Lupinus arcticus, in Upland Tundra on Herschel Powell, G.L., 241 
Island, Yukon Territory, Observations of Change in Pane. Gas 


the Cover of, 323 
Poigonum viviparum, 325 
Pollachius virens, 570,669 
Pollock, 570,669 
Polycentropus, 142 
Polygonum alaskanum, 302,311 

bistorta, 325 

buxiforme, 303,311 

fowleri, 93 

lapathifolium, 303,311 

viviparum, 93,302,311,323 
Polynema, 36 

muirii, 35 
Polyodon spathula, 566 
Polystichum braunii, 446 

munitum, 456 
Pomoxis nigromaculatus, 620 
Pondweed, 304 

Closed-leaved, 304 

Eel-grass, 304 

Richardson’s, 304 
Pooecetes gramineus, 273,434 
Poor-cod, 121 
Poplar, 257 

Balsam, 107,216,426,476 
Populus angustifolia, 543 

balsamifera, 93,107,426,476 


balsamifera ssp. balsamifera, 216 


Prairie-Chicken, Greater, 257 

Prairie Dog, Black-tailed, 244 

Presbytis entellus, 170 

Prickleback, Blackline, 116,566 

Primula egaliksensis, 94 
stricta, 94 

Prince Edward Island National Park, Germination Potential, 
Updated Population Surveys and Floral, Seed and 
Seedling Morphology of Symphyotrichum lauren- 
tianum, the Gulf of St. Lawrence Aster, in the, 287 

Prince Edward Island, New Plant Records for, 446 

Prince Edward Island, The Spring and Fall Migrations of 
Scoters, Melanitta spp., at Confederation Bridge in 
the Northumberland Strait between New Brunswick 
and, 436 

Pristiloma arcticum, 223 
arcticum arcticum, 225 
arcticum crateris, 225 
chersinella, 223 

Procyon lotor, 31,53,84,182,249,417,480 

Progne subis, 433 

Pronghorn, 257 

Prosopium sp., 569 
coulteri, 569 
cylindraceum, 568 

Protonotaria citrea, 417 

Prunella vulgaris ssp. vulgaris, 218 


deltoides, 481 Prunus pensylvanica, 9,217,447 
grandidentata, 446 serotina, 83,248 

tremuloides, 2,9,93,100,107,199,215,224,257,413, virginiana, 9,257 

426,446,476 virginiana ssp. virginiana, 217 


Porbeagle, 506 


Pseudacris triseriata maculata, 499 


Porbeagle, Lamna nasus, Gillnet Survival and Healing by —-Pseudolachea sp., 37 


a, 506 
Porcupine, 53 
Porichthys notatus, 668 
Porpoise, Dall’s, 567 - 

Harbour, 116,123,567 
Porzana carolina, 272,434 
Posthodiplostomum minimum, 150 
Potamogeton spp., 582,587,593,603 

alpinus, 96 


alpinus ssp. tenuifolius, 302,304 


filiformis, 96 
foliosus var. macellus, 302,304 


Pseudopleuronectes americanus, 570 
Pseudorca crassidens, 567 
Pseudoterranova decipiens, 659 
Pseudotsuga menziesii, 1,413,461,467 
Psiada coronopus, 294 
Pteridium aquilinum, 219 
aquilinum var. latiusculum, 216 
Ptolemy, J., 591 
Puccinellia andersonii, 302,305 
lucida, 98 
nuttalliana, 89,406 
phryganodes, 89 


758 


Pumpkinseed, 155,336,594 
Punctum randolphii, 223 
Pungitius pungitius, 330,569,647 
Purshia tridentata, 3 
Pussytoes, 217 
Dense-leaved, 320 
Showy, 320 
Pyganodon cataracta, 329 
fragilis, 329 
Pylodictis olivaris, 565 
Pyrola asarifolia, 94 
chlorantha, 94 
grandiflora, 94 
minor, 302,318 
rotundifolia ssp. grandiflora, 94 


Quaternaria quaternata, 37 

Québec, Effects of Reopening Hunting on Survival of 
White-tailed Deer, Odocoileus virginianus, in the 
Bas-Saint-Laurent region, 99 

Quebec, Establishment of a Breeding Population of Canada 
Geese in Southern, 75 

Quercus spp., 247 
alba, 83,206 
garryana, 451,461,467 
macrocarpa, 215 
prinus, 83,206 
rubra, 83,215 

Quillwort, Maritime, 303 

Quiscalus quiscula, 435,440 


Rabbit, 543 
Rabida, 139 
Raccoon, 31,53,182,249,417,480 
Common, 84 
Rail, Yellow, 434 
Raisin, Wild, 447 
Raja laevis, 569 
radiata, 569 
Rana pipiens, 272 
septentrionalis, 499 
sylvatica, 499 
temporaria, 500 
Rana sylvatica, An Observation of a Mallard, Anas 
platyrhynchos, Feeding on a Wood Frog, 499 
Rangifer tarandus, 50,174,274,326,495 
tarandus caribou, 174 
tarandus groenlandicus, 174 
tarandus pearyi, 179 
Rangifer tarandus, Under Different Climatic Conditions, 
Use of Host-mimicking Trap Catches to Determine 
which Parasitic Flies Attack, 274 
Ranunculus abortivus, 93 
cymbalaria, 93 
gmelinii, 93 
hyperboreus, 93,302,312 
macounii, 93 
pedatifidus, 93 
reptans, 93 
subrigidus, 93 
Raphanus, 341 
- Rapistrum, 341 
Raptor spp., 515 
Raspberry, 58,112 


THE CANADIAN FIELD-NATURALIST 


Vokimis 


Dwarf, 216,316 
Wild Red, 217 
Rat, African Pouched, 248 
Kangaroo, 64 
Norway, 273 
Ord’s Kangaroo, 345 
Rattlesnake, Prairie, 241 
Rattlesnake, Crotalus viridis viridis, in Southeastern 
Alberta, Diet of the Prairie, 241 
Rattus spp., 53 
norvegicus, 273 
Raven, 515 
Common, 168,427,440 
Raven, Corvus corax, Observed Taking an Egg from a 
Common Loon, Gavia immer, Nest, Common, 168 
Recovery, 186,541 
Recurvirostra americana, 434 
Redcedar, Western, 252,455 
Redhead, 434 
Redhorse, Black, 116,566 
Copper, 566 
Golden, 565 
Greater, 116,568 
River, 566 
Redpoll, Common, 440,477 
Redstart, American, 434 
Redtop, 76 
Reed, 408 
Regulus satrapa, 434 
Reindeer, 274 
Reindeer, Rangifer tarandus, Under Different Climatic 
Conditions, Use of Host-mimicking Trap Catches to 
Determine which Parasitic Flies Attack, 274 
Reinhardtius hippoglossoides.570 
Reithrodontomys, 244 
megalotis, 64 
Renew (Recovery of Nationally Endangered Wildlife) 
Report 11: 2000-2001 Annual Report, 733 
Rhabdochona cascadilla, 150 
Rhamnus cathartica, 218 
frangula, 218 
Rheum rhaponticum, 301,311 
Rhinanthus cristagalli, 96 
minor, 90 
Rhinichthys atratulus, 157 
cataractae ssp., 566 
cataractae smithi, 566 
falcatus, 565 
osculus, 565 
umatilla, 116,565 
Rhizopus sp., 37 
Rhodiola rosea ssp. integrifolia, 302,315 
Rhubarb, 311 
Rhus rydbergii, 216 
typhina, 216 
Rhynocladiella sp., 37 
Ribes americanum, 110 
bracteosum, 456 
oxyacanthoides, 94 
triste, 94 
Ricegrass, Indian, 306 
Ripara ripara, 433 
Rissa tridactyla, 168,349 
Rissa tridactyla, Increases and Expansion of the New 


2001 


Brunswick Breeding Population of Black-legged 


Kittiwakes, 349 
Robin, American, 427,440,477 
Rockcress, Drummond’s, 313 
Holboell’s, 313 
Rockfish, Aurora, 570 
Black, 570 
Blue, 570 
Canary, 570 
China, 570 
Copper, 570 
Golden, 570 
Quillback, 570 
Redbanded, 570 
Shortbelly, 570 
Silvergray, 570 
Splitnose, 570 
Sripetail, 570 
Vermillion, 570 
Widow, 570 
Yelloweye, 570 
Rodewald, A. D., 82 
Rodrigue, J., 75 
Roell, B. J., 234 
Rorippa, 341 
palustris, 302,315 
palustris ssp. fernaldiana, 94 
palustris ssp. glabra var. glabrata, 94 
Rosa spp., 447 
acicularis, 110,219,257 
acicularis ssp. sayi, 216 
blanda, 218 
Rose, 110 
Prickly, 111,216,257 
Smooth, 218 
Wild, 447 
Roseroot, 315 
Rothfels, C. J., 229 
Roy, G., 614 
Roy, J., Reviews by, 520,521 
Rubus spp., 58 
acaulis, 95 
chamaemorus, 90,275,302,316 
idaeus ssp. melanolasius, 217 
pubescens, 216,302,316 
spectabilis, 456 
strigosus, 110 
Rumex acetosa ssp. alpestris, 302,311 
acetosella, 468 
occidenalis, 93 
salicifolius ssp. triangulivalvis, 302,311 
Rush, 406 
Dudley’s, 232 
Soft, 232 
Three-Square, 410 
Toad, 232,309 . 
Russell, A. P., 241 
Ryan, M., 451 
Ryegrass, 232 
Perennial, 462,468 


Saccostomus mearnsi, 248 
Sage-grouse, Greater, 257 
Sage, Silver, 513 


INDEX TO VOLUME 115 759 


Sagebrush, Big, 2 
Threetip, 2 
Sagina nodosa, 93 
saginoides, 302,312 
Sagittaria spp., 77 
Salamander, 272,499 
Eastern Tiger, 202 
Tiger, 199,500 
Salamander, Ambystoma tigrinum, Movements and Mor- 
tality on the Trans-Canada Highway in South- 
western Alberta, Tiger, 199 
Salicornia borealis, 89 
europaea, 93,407 
Salix spp., 9,89,174,224,257,413 
alaxensis, 476 
alaxensis ssp. longistylis, 302,310 
arctica, 323 
arctophila, 93,302,310 
barrattiana, 302,310 
bebbiana, 93 
brachycarpa, 93 
candida, 93 
glauca, 275 
glauca ssp. callicarpaea, 94 
glauca var. callicarpaea, 94 
glaucophylloides, 94 
lapponum, 275 
myricoides, 94 
myrtillifolia, 94 
nigra, 348 
pedicellaris, 94,302,310 
planifolia, 94 
pyrifolia, 302,311 
reticulata, 94,323 
serissima, 94 
Salmo salar, 133,330,569,63 1 
trutta, 165,631,647 
Salmon, 668 
Atlantic, 133,330,569,631 
Coho, 155,575,594 
Pacific, 165 
Salmonberry, 456 
Salmonids, Pacific, 569 
Salvelinus alpinus, 568 
fontinalis, 135,165,251,330,631,650 
fontinalis timagamiensis, 116,566 
malma, 129 
namaycush, 631,669 
Sambucus racemosa, 224,456 
Sander-Regier, R., Reviews by, 384,386,528,707,712,722 
Sanderling, 440 
Sandlance, American, 669 
Sandpiper, Baird’s, 272 
Least, 408,487 
Semipalmated, 487 
Solitary, 488 
Spotted, 272,434,440,488 
Stilt, 487 
Upland, 272 
Western, 408 
Sandwort, Blunt-leaved, 312 
Mountain, 311 
Sanicle, Pacific, 460,466 
Purple, 460,466 


760 


Sierra, 462 
Snake-root, 466 
Sanicle, Sanicula arctopoides (Apiaceae) in Canada, Status 
of Snake-root, 466 
Sanicle, Sanicula bipinnatifida (Apiaceae), in Canada, 
Status of the Purple, 460 
Sanicula sp., 218 
arctopoides, 466 
bipinnatifida, 460,466 
crassicaulis, 460,466 
graveolens, 462 
Sanicula arctopoides (Apiaceae) in Canada, Status of 
Snake-root Sanicle, 466 
Sanicula bipinnatifida (Apiaceae), in Canada, Status of the 
Purple Sanicle, 460 
Sapsucker, Red-naped, 413 
Williamson’s, 413 
Yellow-bellied, 416,434 
Sardine, Pacific, 565 
Sardinops sagax, 565 
Sarracenia purpurea, 93 
Sarsaparilla, Wild, 111,216 
Saskatchewan, Slumping Activity and Forest Vegetation 
Along the Northeastern Shore of Waskesiu Lake, 
Prince Albert National Park, 106 
Saskatoon, 257 
Sassafras, 248 
Sassafras albidum, 248 
Saumure, R. A. Limb Mutilations in Snapping Turtles, 
Chelydra serpentina, 182 
Saussurea angustifolia, 325 
Savile, D.B.O. Evolution of a Naturalist, 365 
Savile, 1909-2000, A Tribute to Douglas Barton Osbourne, 
351), 
Saxifraga adscendens ssp. oregonensis, 302,315 
aizoides, 302,315 
bronchialis ssp. funstonii, 302,316 
hirculus, 94,323 
punctata, 325 
tricuspidata, 89 
Saxifrage, 323 
Spotted, 316 
Wedge-leaved, 315 
Yellow Mountain, 315 
Sayornis phoebe, 433 
saya, 433 
Scaup, 440 
Lesser, 272,434 
Schaefer, H., 446 
Schellenberg, M.P., Reviews by, 194,723 
Scheuchzeria palustris, 96 
palustris ssp. americana, 302,304 
Scheuchzeria, 304 
Schmutz, J. K., C. S. Houston, and S. J. Barry. Prey and 
Reproduction in a Metapopulation Decline Among 
Swainson’s Hawks, Buteo swainsoni, 257 
Schoenocrambe, 341 
Scirpus acutus, 301 
caespitosus, 97 
caespitosus ssp. austriacus, 302,309 
hudsonianus, 97 
paludosus, 407 
rollandii, 302,309 
rufus var. neogaeus, 97 
validus, 303,309 


THE CANADIAN FIELD-NATURALIST 


Vol ans 


Sciurid, 543 
Sciurus spp., 84,543 
carolinensis, 53,547 
Scolicosporium sp., 37 
Scomber scombrus, 570,658 
Scoter, Black, 437 
Surf, 437 
White-winged, 434,437 
Scoters, Melanitta spp., at Confederation Bridge in the 
Northumberland Strait between New Brunswick 
and Prince Edward Island, The Spring and Fall 
Migrations of, 436 
Scouring-rush, Dwarf, 218,303 
Scudder, G. G. E., 1 
Sculpin, Cultus Pygmy, 566 
Deepwater, 566 
Fourhorn, 116,566 
Great Lakes Deepwater, 569 
Mottled, 142,568 
Shorthead, 566 
Spinynose, 565 
Spoonhead, 565 
Scutellaria parvula, 218 
Seal, Bearded, 566 
Grey, 566,653 
Harbour, 566,655,663 
Harp, 570 
Hooded, 566 
Largha, 663 
Northern Elephant, 566,671 
Northern Fur, 566 
Ringed, 567 
Seal, Halichoerus grypus, in the Northwest Atlantic*, The 
Status of the Grey, 653 
Seals, Phoca vitulina, in Canada*, Status of Harbour, 663 
Sea Lion, California, 566 
Northern, 671 
Steller, 355,566 
Sea Lions, Eumetopias jubatus, Cooperative Foraging by 
Steller, 355 
Sea Wind: Bulletin of Ocean Voice International 14(4), 186 
Sebastes aurora, 570 
babcocki, 570 
brevispinis, 570 
caurinus, 570 
diploproa, 570 
entomelas, 570 
goodei, 570 
jordani, 570 
maliger, 570 
melanops, 570 
miniatus, 570 
mystinus, 570 
nebulosus, 570 
Sebastolobus alascanus, 570 
altivelis, 570 
Sebates alutus, 570 
norvegicus, 570 
paucispinis, 570 
pinniger, 570 
ruberrimus, 570 
saxicola, 570 
Seburn, D., Reviews by, 522,712,714 
Sedge, 2172755505 
Black, 306 


2001 


Bladder, 217 
Blunt, 307 
Bronze, 306 
Chestnut, 218 
Cordroot, 307 
Crawford’s, 307 
Golden, 307 
Hairlike, 307 
Lesser Panicled, 307 
Livid, 307 
Long-styled, 307 
Long-stoloned, 461 
Loose-flower Alpine, 307 
Richardson’s, 217 
Sparse-leaved, 308 
Two-coloured, 307 
Two-parted, 307 
Two-toned, 306 
Woolly, 218 
Sedum spathifolium, 451 
Seiurus aurocapilus, 434 
noveboracensis, 434 
Selaginella selaginoides, 302,303 
Selaginella, Northern, 303 
Semotilus atromaculatus, 164 
corporalis, 615 
Senecio congestus var. palustris, 96 
hadrosomus, 294 
hyperborealis, 302,320 
indecorus, 96 
lindstroemii, 325 
lugens, 302,321 
pauperculus, 96,216 
Serviceberry, 9 
Setophaga ruticilla, 434 
Shad, American, 164,330,569 
Shark, 671 
Sheep, Bighorn, 3 
Sheffield, G., 501 
Shepherdia argentea, 257 
canadensis, 95 
Shiner, Bigmouth, 565 
Blackchin, 565,608,617 
Blacknose, 608,617 
Bridle. 565,614 
Emerald, 569 
Ghost, 565 
Golden, 647 
Mimic, 142 
Pugnose, 565,618 
Redfin, 565 
Roseyface, 565 
Silver, 116,565 
Spottail, 569,608 
Striped, 565 


Weed, 565,608 - 

Shiner, Notropis bifrenatus, in Canada*, Status of The 
Bridle, 614 

Shiner, Notropis texanus, in Canada*, Status of the Weed, 
608 


Shootingstar, Henderson’s, 462 
Shoveler, Northern, 434 
Shrew, 54,354 

Common, 6 


INDEX TO VOLUME 115 


761 


Dusky, 3 
Dwarf, 1,514 
Meriam’s, | 
Preble’s, | 
Pygmy, 513 
Short-tailed, 53 
Soricidae, 273 
Vagrant, 3 
Shrew, Sorex hoyi, on the Montana-Alberta Border, A 
Significant New Record of the Pygmy, 513 
Shrew, Sorex merriami, and Preble’s Shrew, Sorex preblei: 
Two New Mammals for Canada, Meriam’s, | 
Shrew, Sorex preblei: Two New Mammals for Canada, 
Meriam’s Shrew, Sorex merriami, and Preble’s, | 
Shrike, Loggerhead, 257 
Shrimp, 644 
Sialia currucoides, 434 
Sigmodon, 546 
Silene uralensis ssp. ogilviensis, 302,312 
Silverside, Brook, 565 
Simuliidae, 644 
Simulium, 274 
Sinadoxa corydalifolia, 319 
Sinapis, 341 
Sinclair, A., 214 
Siskin, Pine, 435 
Sisymbrium, 341 
Sitta canadensis, 434 
Sium suave, 95 
Skate, Barndoor, 569 
Smooth, 570 
Thorny, 569 
Skua, Arctic, 515 
Skullcap, Small, 218 
Skunk, Striped, 53,84,480 
Smelowskia, 341 
borealis, 342 
calycina var. americana, 342 
calycina var. porsildii, 342 
Johnsonii, 341 
media, 341 
ovalis, 342 
pyriformis, 342 
Smelt, Arctic Rainbow, 132 
European, 132 
Lake Utopia Dwarf, 566 
Pygmy, 131,569 
Rainbow, 131,330,631 
Smelt (Genus Osmerus) Populations of Lake Utopia, New 
Brunswick, Status of the Sympatric, 131 
Smilacina trifolia, 98 
Smith, C.Ay S., 323 
Smith, D. W., 495 
Smith, D. W., K. M. Murphy, and S. Monger. Killing of a 
Bison, Bison bison, Calf by a Wolf, Canis lupus, 
and Four Coyotes, Canis latrans, in Yellowstone 
National Park, 343 
Snail, 165 
Vancouver, 223 
Snails from the Mountains of Northwestern British 
Columbia, New Records of Land, 223 - 
Snake, Eastern Garter, 347 
Plains Garter, 272 
Red-sided Garter, 347 


762 


Snakeroot, 218 
Snakes, Thamnophis sirtalis sirtalis, in September in Bruce 
County, Ontario, Arboreal Courtship Behaviour by 
Eastern Garter, 347 
Snapdragon, Lange Dwarf, 218 
Snipe, Common, 428,488 
Snowberry, 3,112,216,451 
Solanum tuberosum, 447 
Solidago candensis, 110 
flexicaulis, 449 
hispida var. hispida, 217 
juncea, 217 
multiradiata, 96 
nemoralis, 217 
ptarmicoides, 2\7 
uliginosa, 96 
Solomon’s Seal, Three-leaved, 309 
Two-leaved, 109 
Somateria mollissima, 439 
Sonchus arvensis, 90,110 
arvensis ssp. uliginosus, 96,303,321 
Sora, 272,428 
Sorbaria sorbifolia, 301,316 
Sorbus americana, 9 
Sorex sp., 53,354,477 
cinereus, 2 
hoyi, 513 
merriami, | 
monticolus, 3 
nanus, 1,514 
preblei, | 
vagrans, 3 
Sorex hoyi, on the Montana-Alberta Border, A Significant 
New Record of the Pygmy Shrew, 513 
Sorex merriami, and Preble’s Shrew, Sorex preblei: Two 
New Mammals for Canada, Meriam’s Shrew, | 
Sorex preblei: Two New Mammals for Canada, Meriam’s 
Shrew, Sorex merriami, and Preble’s Shrew, 1 
Sorghastrum nutans, 248 
Sorrel, Garden, 468 
Green, 311 
South Dakota, Evidence for Double Brooding by a Mallard, 
Anas platyrhynchos, in Eastern, 502 
South Dakota, Influence of Predation on Piping Plover, 
Charadrius melodus, and Least Tern, Sterna antil- 
larum, Productivity along the Missouri River in, 
480 
Sow-thistle, Perennial, 321 
Sparganium angustifolium, 98 
eurycarpum, 77 
minimum, 302,304 
multipedunculatum, 302,304 
natans, 98 
Sparrow, Chipping, 427 
Clay-coloured, 428 
House, 273,435 
Lark, 273 
Le Conte’s, 273,434 
Lincoln’s, 434 
Savanna(h), 273,427,440 
Sharp-tailed, 434 
Song, 428,440 
Swamp, 434 
Vesper, 273,434 
White-throated, 434 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Speckman, S. G. and G. Sheffield. First Record of an 
Anomalously White Killer Whale, Orcinus orca, 
Near St. Lawrence Island, Northern Bering Sea, 
Alaska, 501 
Speotyto cunicularia, 272 
Spergularia canadensis, 89,170 
marina, 407 
Spermophilus, 244,543 
niger, 547 
parryii, 546 
richardsonii, 242,257,543 
tridecemlineatus, 273 
Spermophilus richardsonii, Great Horned Owl, Bubo virgini- 
anus, Predation on Richardson’s Ground Squirrels, 
543 
Sphagnum, 323 
Sphyrapicus nuchalis, 413 
thyroideus, 413 
varius, 416,434 
Spike-Rush, Creeping, 410 
Spiraea alba, 218 
Spiraea, False, 316 
Spiranthes romanzoffiana, 98,302,310 
Spizella pallida, 434 
passerina, 434 
Sporobolus crytandrus, 3 
Spot, Conical, 224 
Spreadwing, Amber-winged, 404 
Common, 404 
Emerald, 404 
Lyre-tipped, 404 
Slender, 404 
Spotted, 404 
Sweetflag, 404 
Sprite, Sedge, 404 
Spruce, Black, 10,57,64,178,426,446,476 
Engelmann, 4,505 
Interior, 413 
Red, 446 
Sitka, 455 
White, 10,57,64,100,107,199,219,426,447,476 
Squalus acanthias, 569 
Squid, 121 
Short-finned, 669 
Squirrel, 84 
Arctic Ground, 546 
Flying, 543 
Fox, 547 
Gray, 53 
Ground, 170,244 
Northern Flying, 417 
Red, 53,417 
Richardson’s Ground, 242,257,543 
Thirteen-lined Ground, 273 
Squirrels, Spermophilus richardsonii, Great Horned Owl, 
Bubo virginianus, Predation on Richardson’s 
Ground, 543 
St. John’ s-wort, 216 
Common, 218 
Stachys palustris, 110 
Stafford, J. D., L. D. Flake, and P. W. Mammenga. Evidence 
for Double Brooding by a Mallard, Anas platyrhyn- 
chos, in Eastern South Dakota, 502 
Starling, European, 434,440 


2001 


Starwort, Long-stalked, 312 
Stellaria borealis ssp. borealis, 93 
calycantha, 93 
humifusa, 93 
longifolia, 93 
longipes, 93,302,312 
media, 303,312 
Stenella coeruleoalba, 567 
Stenonema, 142 
Stercorarius longicaudus, 168 
parasiticus, 168,515 
Sterna antillarum, 480 
caspia, 440 
hirundo, 434,440 
paradisaea, 492 
Sterna antillarum, Productivity along the Missouri River in 
South Dakota, Influence of Predation on Piping 
Plover, Charadrius melodus, and Least Tern, 480 
Sternotherus odoratus, 182 
Stewart, H., 1 
Stewart S. E. and C. R. Lacroix. Germination Potential, 
Updated Population Surveys and Floral, Seed and 
Seedling Morphology of Symphyotrichum lauren- 
tianum, the Gulf of St. Lawrence Aster, in the 
Prince Edward Island National Park, 287 
Stickleback, 579,584,591 
Balkwell Lake, 116 
Charlotte, 569 
Emily Lake, 116 
Enos Lake, 116,566 
Fourspine, 330 
Giant, 566 
Hadley Lake, 116,566 
Ninespine, 330,569,647 
Paxton Lake, 566 
Priest Lake, 116 
Texada, 152 
Threespine, 152,336,579,584,591 
Unarmoured, 566 
Vananda Creek, 566 
Stickleback Species Pair, Gasterosteus spp., in Canada, 
Status of the Texada, 152 
Stickleback Species Pair, Gasterosteus spp., in Hadley 
Lake, Lasqueti Island, British Columbia*, Status of 
the, 579 
Stickleback Species Pair, Gasterosteus spp., in Paxton 
Lake, Texada Island, British Columbia*, Status of 
the, 591 
Stickleback Species Pair, Gasterosteus spp., in the Vananda 
Creek watershed of Texada Island, British Colum- 
bia*, Status of the, 584 
Stictochironemus, 165 
Stipa acutus 408 
comata, 3,513 
hymenoides, 301,306 
nelsonii ssp. dorei, 302 
nelsonii var. dorei, 2 
pungens, 410 
Stizostedion vitreum, 160 
vitreum glaucum, 566 
Stonecat, 569 
Stonecrop, Broad-leaved, 451 
Stoneroller, Central, 157,565 
Largescale, 158 


INDEX TO VOLUME 115 


763 


Stoneroller, Campostoma anomalum, in Canada, Updated 
Status of the Central, 157 
Strawberry, Barren, 216 
Scarlet, 217 
Strix occidentalis, 61,547 
varia, 61 
Stronglyocentrotus spp., 556 
franciscanus, 559 
Strophitus undulatus, 329 
Sturgeon, Atlantic, 568 
Green, 565 
Lake, 565 
Shortnose, 565 
White, 565 
Sturna vulgaris, 434 
Sturnella neglecta, 257,434 
Sturnus vulgaris, 440 
Suaeda calceoliformis, 93 
Subularia, 341 
Sucker, Jasper Longnose, 568 
Mountain, 565 
Salish, 566 
Spotted, 566 
White, 330,631,647 
Sula bassanus, 440 
Sumac, Staghorn, 216 
Sunfish, 150 
Green, 565 
Longear, 565 
Orangespotted, 116,566 
Redbreast, 116,566 
Sus scrofa, 248 
Swallow, Bank, 433 
Barn, 433 
Tree, 433,440 
Sweet-clover, White, 317 
Yellow, 76,317 
Sweet-coltsfoot, 216 
Switchgrass, 248 
Swordfern, Western, 456 
Swordfish, 570 
Sylvilagus spp., 53,543 
floridanus, 43 
nuttallii, 244,273 
Symphoricarpos alba, 219 
albus, 3,110,216,451 
occidentalis, 110 
Symphyotrichum laurentianum, 287 
Symphyotrichum laurentianum, the Gulf of St. Lawrence 
Aster, in the Prince Edward Island National Park, 
Germination Potential, Updated Population Surveys 
and Floral, Seed and Seedling Morphology of, 287 
Synedra, 165 


Tabanus, 274 
Tabellaria, 165 
Tachycineta bicolor, 433,440 
Talbott, S. C., 82 
Tamias, 543 
striatus, 53,84 
Tamiasciurus, 543 
hudsonicus, 53,417,547 
Tansymustard, Northern, 314 
Western, 314 


764 


Taraxacum carneocoloratum, 301,321 
ceratophorum, 96 
lacerum, 96 
officinale, 90,216,303,321 
Taxidea taxus, 345 
Taylor, E. B. Status of the Sympatric Smelt (Genus Osmer- 
us) Populations of Lake Utopia, New Brunswick, 
131 
Teal, Blue-winged, 272,428,434 
Chestnut, 502 
Green-winged, 434 
Grey, 502 
Teesdalia, 341 
Teradoxa omeiensis, 319 
Tern, Arctic, 492 
Black, 432 
Caspian, 440,483 
Common, 434,440 
Least, 480 
Tern, Sterna antillarum, Productivity along the Missouri 
River in South Dakota, Influence of Predation on 
Piping Plover, Charadrius melodus, and Least, 480 
Testudo hermanni, 182 
Teucrium canadense, 447 
Thaleichthys pacificus, 355,569 
Thalictrum alpinum, 302,312 
venulosum, 93,110 
Thamnolia subuliformis, 325 
Thamnophis radix, 272 
sirtalis parietalis, 347 
sirtalis sirtalis, 347 
Thamnophis sirtalis sirtalis, in September in Bruce County, 
Ontario, Arboreal Courtship Behaviour by Eastern 
Garter Snakes, 347 
Théau, J. et J. Ferron. Influence de paramétres climatiques 
sur les patrons d’activité saisonniers et journaliers 
du hevre d’Amérique, Lepus americanus, en semi- 
liberté, 43 
Thelypodium, 341 
Thistle, Bull, 216 
Canada, 110 
Thlaspi, 341 
arvense, 303,315 
Thomomys, 546 
bottae, 245 
talpoides, 273 
Thornyhead, Longspine, 570 
Shortspine, 570 
Thrasher, Brown, 434 
Thrift, 467 
Thrush, Hermit, 434 
Swainson’s, 428,477 
Varied, 477 
Thuja occidentalis, 44,100,215 
plicata, 252,456 
Thunnus thynnus, 568 


Thuya, 44 
Thysanocarpus, 341 
Mek, 212 

Winter, 174 


Tiffney, W.N., 351 

- Tightcoil, Arctic, 225 
Tilia americana, 40 
Timmia austriaca, 325 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Timoney, K. P. String and Net-Patterned Salt Marshes: 
Rare Landscape Elements of Boreal Canada, 406 
Timothy, 76 
Toad, American, 499 
Boreal, 500 
Common, 200 
Tofieldia coccinea, 302,309 
glutinosa, 98 
glutinosa ssp. brevistyla, 302,309 
pusilla, 98,302,310 
Tomcod, 570,647 
Atlantic, 330 
Topminnow, Blackstripe, 565 
Touch-me-not, Glandular, 446 
Towhee, Eastern, 85 
Spotted, 434 
Toxicodendron rydbergii, 447 
Toxostoma rufum, 434 
Trefoil, Small-flowered Birds-foot, 468 
Trematodes, 150 
Trematosphaeria sp., 37 
Triaenodes aba, 142 
Trianga flavipes, 272 
Trichoderma, 36 
harizanum, 37 
viridae, 37 
Trichophorum pumilum, 309 
Trichostema brachiatum, 217 
Trientalis borealis, 94 
Trifolium spp., 76 
hybridum, 232 
repens, 218 
willdenowii, 468 
Triglochin maritimum, 96,302,305,407 
palustre, 96,407 
Trillium ovatum var. hibbersonii, 343 
Trillium ovatum Pursh variety hibbersonii (Taylor et Szcza- 
winsk1) Douglas et Pojar, variety nova, 343 
itimper, PGs 177. 
Tringa flavipes, 490 
solitaria, 488 
Tripleurospermum phaeocephalum, 96 
Trisetum spicatum vat. spicatum, 98 
Trisopterus luscus, 121 
minutus, 121 
Triticum aestivum, 513 
Troglodytes aedon, 414,434 
Trout, Aurora, 116,566 
Brook, 135,165,251,330,631,650 
Brown, 165,631,647 
Cutthroat, 128,154,582,589,592 
Lake, 631,669 
Rainbow, 165,631 
Tsuga canadensis, 40,85,446 
heterophylla, 455 
Tsuji, L. J. S., J. D. Karagatzides, and G. Deilutis. Ecto- 
-parasites in Lekking Sharp-tailed Grouse, Tympanu- 
chus phasianellus, 210 
Tuna, Bluefin, 568 
Turdus migratorius, 434,440,477 
Turkey, Wild, 83 
Tursiops truncatus, 122,567 
Turtle, Blanding, 512 
Musk, 182 
Painted, 512 


2001 


Snapping, 182,510 
Spotted, 182 
Wood, 182 

Turtles, Chelydra serpentina, and Eggs in a Wood Chip 
Pile, Hyperthermia Induced Mortality of Gravid 
Snapping, 510 

Turtles, Chelydra serpentina, Limb Mutilations in Snap- 
ping, 182 

Twedt, D. J., G. M. Linz, and W. J. Bleier. Inability to 
Predict Geographic Origin of Yellow-headed Black- 
birds, Xanthocephalus xanthocephalus, During 
Migration, 549 

Twinflower, 109 

Tympanuchus cupido, 257 
pallidicinctus, 434 
phasianellus, 210,272 

Tympanuchus phasianellus, Ectoparasites in Lekking 
Sharp-tailed Grouse, 210 

Typha spp., 503 
angustifolia, 77,232 
latifolia, 98,302,304,408 

Tyrannus tyrannus, 169,433 


Ulmus americana, 40,257 
Uncinula sp., 37 
Urchin, Red Sea, 559 
Sea, 556 
Urophora spp., 354 
Urophycis chuss, 570 
Ursus sp., 249 
americanus, 84,174,414 
arctos, 170,495 
arctos horribilis, 174 
maritimus, 670 
Ursus arctos, Usurps Bison Calf, Bison bison, Captured by 
Wolves, Canis lupus, in Yellowstone National Park, 
Wyoming, Grizzly Bear, 495 
Urtica dioica, 110 
Urton, E. J., 345 
Ustulina deusta, 36 
Ustulinetum, 38 
Utricularia, 582,587,593 
intermedia, 96,302,319 
vulgaris, 96 
vulgaris ssp. macrorhiza, 302,319 
Uvulifer sp., 164 
vancleavi, 150 


Vaccinium spp., 476 
myrtillus, 275 
oxycoccos, 94 
uliginosum, 94 
vitis-idaea, 476 
vitis-idaea ssp. minus, 94 
Valeriana dioica ssp. sylvatica, 96 
septentrionalis, 96 
Valsa spp., 36 
ambiens, 37 
Valsetum, 36 
Vander Lee, B. A., 480 
Van Why, K. R. and W. M. Giuliano. Fall Food Habits and 
Reproductive Condition of Fishers, Martes pen- 
nanti, in Vermont, 52 
Van Wilgenburg, S.L., 425 


INDEX TO VOLUME 115 


765 


Varden, Dolly, 129 
Varley, N., 495 
Veery, 428 
Veitch, A. M. An Unusual Record of a White-tailed Deer, 
Odocoileus virginianus, in the Northwest Terri- 
tories, 172 
Veitch, A.M., Reviews by, 704,716 
Velvet-grass, Common, 468 
Verbascum thapsus, 216 
Vergara, V. Comparison of Parental Roles in Male and 
Female Red Foxes, Vulpes vulpes, in Southern 
Ontario, 22 
Vergara, V. Two Cases of Infanticide in a Red Fox, Vulpes 
vulpes, Family in Southern Ontario, 170 
Vermivora celata, 434 
peregrina, 434 
Vermont, Fall Food Habits and Reproductive Condition of 
Fishers, Martes pennanti, in, 52 
Vernalgrass, Sweet, 451 
Verticillium sp., 36 
lecanii, 37 
Vertigo columbiana, 225 
modesta, 225 
modesta parietalis, 226 
Vertigo, Columbia, 225 
Vespericola columbianus, 223 
columbianus pilosus, 226 
Vetche2i7 
American, 111 
Chick-pea Milk, 316 
Milk, 323 
Neglected Milk, 217 
Standing Milk, 316 
Timber Milk, 2 
Tufted, 76,217,317 
Viburnum nudum, 447 
rafinesquianum, 217 
Vicia sp., 217 
americana, 110 
craccas (6.217.303 317 
Viola adunca, 217 
nephrophylla, 93,218 
praemorsa ssp. praemorsa, 451 
renifolia, 93,218 
Violet, Hooked-spur, 217 
Kidney-leaved, 218 
Northern Bog, 218 
Vireo gilvus, 434 
olivaceus, 434 
Philadelphicus, 434 
solitarius, 434 
Vireo, Philadelphia, 434 
Red-eyed, 427 
Solitary, 434 
Warbling, 428 
Vitrina pellucida, 225 
Vole, 53,354,543 
Gapper’s Red-backed, 273 
Meadow, 242,273,478,513 
Montane, 4 
Northern Red-backed, 478 
Prairie, 31 
Red-backed, 64,472 
Sagebrush, 4,242,273 


766 


Vujanovic, V. et J. Brisson. Biodiversité microfongique du 
Fagus grandifolia dans une forét ancienne: bioindi- 
cateurs et structure mycosociologique, 34 

Vulpes cana, 22 
vulpes, 22,77,170,480 

Vulpes vulpes, Family in Southern Ontario, Two Cases of 
Infanticide in a Red Fox, 170 

Vulpes vulpes, in Southern Ontario, Comparison of Parental 
Roles in Male and Female Red Foxes, 22 

Vulpia bromioides, 468 
octoflora, 3 


Waiser, B., Review by, 530 
Waldsteinia fragarioides, 216 
Walleye, 160 
Blue, 566 
Walrus, 655 
Atlantic, 566 
Walters, E. L. and E. H. Miller. Predation on Nesting Wood- 
peckers in British Columbia, 413 
Wapiti, 174 
Warbler, Black-and-White, 434 
Chestnut-sided, 434 
Connecticut, 428 
Magnolia, 434 
Mourning, 434 
Orange-crowned, 434 
Palm, 434 
Prothonotary, 417 
Tennessee, 428 
White-crowned, 477 
Yellow, 427 
Yellow-rumped, 434,440,477 
Ward, 1921-1999, A Remembrance of John Clifton, 517 
Warmouth, 566 
Water-Hemlock, European, 318 
Watermilfoil, 620 
Eurasian, 620 
Waterthrush, Northern, 434 
Waxwing, Cedar, 434 
Weasel, 273 
Long-tailed, 415 
Wedgemussel, Dwarf, 329 
Weed, Willow, 311 
Weickert, C. C., J. C. Whittaker, and G. A. Feldhamer. 
Effects of Enclosed Large Ungulates on Small 
Mammals at Land Between The Lakes, Kentucky, 
247 
Whale, Baird’s Beaked, 567 
Beluga, 122 
Blainville’s Beaked, 567 
Blue, 567 
Bowhead, 567 
Cuvier’s Beaked, 567 
Dwarf Sperm, 567 
False Killer, 567 
Fin, 116,121,567 
Grey, 567 
Hubbs’ Beaked, 567 
Humpback, 116,567 
Killer, 123,501,567,671,676 
Long-finned Pilot, 567 
Minke, 118,568 
Northern Bottlenose, 567 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Pygmy Sperm, 567 
Right, 567 
Sei, 568 
Short-finned Pilot, 567 
Sowerby’s Beaked, 116,567 
Sperm, 567 
Stejneger’s Beaked, 567 
True’s Beaked, 567 
Whale, Orcinus orca, Near St. Lawrence Island, Northern 
Bering Sea, Alaska, First Record of an Anomal- 
ously White Killer, 501 
Whales, Orcinus orca, in Canada*, Status of Killer, 676 
Wheat, 513 
Wheatgrass, Bluebunch, 3 
Crested, 76,513 
Western, 76 
Whimbrel, 487 
Whiskers, Old Man’s, 3 
Whitefish, 669 
Atlantic, 566,624,635 
Broad, 569 
Giant Pygmy, 569 
Lake, 568,623,636 
Lake Simcoe, 566,623 
Mira, 565 
Pygmy, 569 
Round, 568 
Squanga, 565,623 
Whitefish, Coregonus clupeaformis, in Canada, The Status 
of the Mira River Population of Lake, 623 
Whitefish, Coregonus huntsmani, Updated Status Report 
on the Endangered Atlantic, 635 
Whiting, 121 
Whitlow-grass, Scotter’s, 314 
Wood, 314 
Whitman, J. S. Diets of Nesting Boreal Owls, Aegolius 
funereus, in Western Interior Alaska, 476 
Whittaker, J. C., 247 
Widgeon, American, 272,434 
Willet, 272,434 
Willow, 9,174,224,257,275,413 
Arctic, 323 
Balsam, 311 
Barratt’s, 310 
Black, 348 
Bog, 310 
Feltleaf, 310 
Northern, 310 
Willson, M. F., 355 
Wintergreen, Lesser, 318 
Wolf, 19,22,179,343,495 
Gray, 174,235,497 
Maned, 22 
Wolf, Canis lupus, and Four Coyotes, Canis latrans, in 
Yellowstone National Park, Killing of a Bison, Bison 
- bison, Calf by a, 343 
Wolffish, 570 
Bering, 116,566 
Wolves, Canis lupus, in Yellowstone National Park, 
Wyoming, Grizzly Bear, Ursus arctos, Usurps 
Bison Calf, Bison bison, Captured by, 495 
Wolves, Canis lupus, “Standing Over” And “Hugging” in 
Wild, 179 
Womble, J. N., 355 


2001 


Wood-Pewee, Western, 433 

Woodpecker spp., 440 
Black-backed, 416 
Downy, 413,434 
Hairy, 413,434 
Pileated, 413,434 
Red-headed, 428 

Woodpeckers in British Columbia, Predation on Nesting, 
413 

Woodrush, Many-flowered, 461 

Woodsia glabella, 302,304 

Woodsia, Smooth, 304 

Worm, Meningeal, 174 

Wren, House, 414,434 
Marsh, 434 

Wyatt, V., Review by, 710 

Wyoming, Grizzly Bear, Ursus arctos, Usurps Bison Calf, 
Bison bison, Captured by Wolves, Canis lupus, in 
Yellowstone National Park, 495 


Xanthocephalus xanthocephalus, 434,549 

Xanthocephalus xanthocephalus, During Migration, In- 
ability to Predict Geographic Origin of Yellow- 
headed Blackbirds, 549 

Xiphias gladius, 570 

Xylaria, 38 
polymorpha, 37 


Y-Prickleback, 565 
Yahner, R. H., A. D. Rodewald, and S. C. Talbott. Edge- 
Related Nest Predation Associated With the 


Index to Book Reviews 


Botany 

Biek, D. Flora of Mount Rainier National Park, 720 

Graham, L E. and L. W. Wilcox. Algae, 526 

Hinds, H. R. Flora of New Brunswick, 718 

Lee, R. E. Phycology, 526 

Morton, J. K. and J. M. Venn. The Flora of Manitoulan 
Island and the adjacent islands of Lake Huron, 
Georgian Bay and the North Channel, Third 
Edition, 721 

Silvertown, J., M. Franco, and J. L. Harper. Plant Life 
Histories: Ecology, Phylogeny and Evolution, 719 

Wunderlin, R. P. and B. F. Hansen. Flora of Florida, Volume 
1, Pteridophytes and Gymnosperms, 718 


Environment 

Brownell, V.R. and J. L. Riley. The Alvars of Ontario: 
Significant Alvar Natural Areas in the Ontario 
Great Lakes Region, 529 

Copeland, G. Acts of Balance: Profits, People and Place, 
194 

Dauncey, G. Earth Future: Stories from a Sustainable 
World, 723 

French, H. Vanishing Borders: Protecting the Planet in the 
Age of Globalization, 723 

Herriot, T. River in a Dry Land: A Prairie Passage, 387 


INDEX TO VOLUME 115 


767 


Retention of Residual Trees in Harvested Hardwood 
Stands, 82 

Yarrow, 2,467 

Yellow-poplar, 206 

Yellowlegs, Lesser, 272,432,488 

Yellowthroat, Common, 434 

Youson, J. H., 573 

Yukon, Mule, Odocoileus hemionus, and White-tailed, O. 
virginianus, Deer in the, 296 

Yukon Territory, Observations of Change in the Cover of 
Polargrass, Arctagrostis latifolia, and Arctic Lupine, 
Lupinus arcticus, in Upland Tundra on Herschel 
Island, 323 

Yukon Territory III, New Records of Vascular Plants in 
the, 301 


Zalophus californianus, 566 
Zannichellia palustris, 97 
Zapus hudsonius, 477 
Zapus princeps, 245 
Zapus princeps, Mouse, Western Jumping, 242 
Zenaida macroura, 434 
Zigadenus venenosus, 2 
Ziphius cavirostris, 567 
Zonitoides arboreus, 225 
Zonotrichia albicollis, 434 
leucophrys, 477 
Zoogenetes harpa, 225 
Zostera marina, 97,644 
Zygomycota, 36 


Kitching, R. L. Food Webs and Container Habitats: The 
Natural History and Ecology of Phytotelmata, 721 

McNeill, J. R. Something New Under the Sun: An Environ- 
mental History of the Twentieth-Century World, 
527 

Osgood Wright, M. The Friendship of Nature, A New 
England Chronicle of Birds and Flowers, 722 

Shepard, F. R. Encounters With Nature: Essays by Paul 
Shepard, 386 

Suzuki, D. and K. Vanderlinden. You Are the Earth, From 
Dinosaur Breath to Pizza from Dirt, 528 

Thornton, J. Pandora’s Poison: Chlorine, Health, and a 
New Environmental Strategy, 724 


Miscellaneous 

Koerner, L. Linnaeus: Nature and Nation, 195 

Meine, C. and R. L. Knight. The Essential Aldo Leopold: 
Quotations and Commentaries, 530 

Merrill, M. D. Yellowstone and the Great West: Journals, 
Letters, and Images from the 1871 Hayden Expe- 
dition, 388 


Zoology 
Able, K. P. Gatherings of Angels: Migrating Birds and their 
Ecology, 523 


768 


Beletsky, L. Belize and Northern Guatemala: The Eco- 
travellers’ Wildlife Guide, 710 

Boitani, L. and T. K. Fuller. Research Techniques in Animal 
Ecology, Controversies and Consequences, 707 

Brown, C. R. Swallow Summer, 717 

Bruce, R. C., R. G. Jaeger, and L. D. Houck. The Biology 
of Plethodontid Salamanders, 714 

Campbell, R. W., N. K. Dawe, I. MacTaggart-Cowan, J. M. 
Cooper, G. W. Kaiser, M. C. E. McNall and G. E. J. 
Smith. The Birds of British Columbia Volume 3, 
187 

Conant, R. and J. T. Collins. A Field Guide to Reptiles and 
Amphibians of Eastern and Central North America, 
194 

Crichton, E. G. and P. H. Krutzsch. Reproductive Biology 
of Bats, 525 

Davies, N. B. Cuckoos, Cowbirds and Other Cheats, 520 

del Hoyo, J., A. Elliot, and J. Sargatal. Handbook of the 
Birds of the World: Volume 6 Mousebirds to Horn- 
bills, 706 

Desmarais, S. and P. R. Krausman. Ecology and 
Management of Large Mammals in North America, 
704 

Forsyth, A. Mammals of North America: Temperate and 
Arctic Regions, 384 

Gosling, L. M. and W. J. Sutherland. Behaviour and 
Conservation, Conservation Biology Series 2, 712 

Grimmet, R., C. Inskipp, and T. Inskipp. A Guide to the 
Birds of India, Pakistan, Nepal, Bangladesh, 
Bhutan, Sri Lanka, and the Maldives, 189 

Isenberg, A. C. The Destruction of the Bison, 703 

Johnsgard, P. A. Prairie Birds: Fragile Splendor in the 
Great Plains, 703 

Johnsgard, P. A. The Pheasants of the World: Biology and 
Natural History, 713 


THE CANADIAN FIELD-NATURALIST 


Vol. 115 


Johnsgard, P. A. Trogons and Quetzals of the World, 191 
Kaufman, K. Kingbird Highway: The Story of a Natural 
Obsession that Got a Little Out of Hand, 524 

Kiemens, M. W. Turtle Conservation, 712 

Konig, C., F. Weick, and J.-H. Becking. Owls: A Guide to 
the Owls of the World, 193 

MacKinnon, J. and K. Phillipps. A Field Guide to the Birds 
of China, 381 

Mason, A. The Nature of Spiders, Consummate Killers, 
386 

Morrison, M. L., L. S. Hall, S. K. Robinson, S. I. Rothstein, 
D. C. Hahn, and T. D. Rich. Research and Man- 
agement of the Brown-headed Cowbird in Western 
Landscapes, 192 

Parsons, H. The Nature of Frogs: Amphibians With Atti- 
tude, 522 

Pearson, D. L. and L. Beletsky. Ecotravellers’ Wildlife 
Guide to Ecuador and its Galapagos Islands, 709 

Raffeale, H., J. Wiley, O. Garrida, A. Keith, and J. 
Raffeale. A Guide to the Birds of the West Indies, 
188 

Rouxel, R. Snipes of the Western Palearctic, 385 

Shackleton, D. Hoofed Mammals of British Columbia, 716 

Sibley, D. A. The Sibley Guide to Birds, 708 

Snyder, N. and H. Snyder. The California Condor: A Saga 
of Natural History and Conservation, 382 

Stattersfield, A. and D. Capper. Threatened Birds of the 
World, 707 

Svensson, L. and P. J. Grant. Birds of Europe, 715 

Thurston, H. The Nature of Hummingbirds: Rainbows on 
Wings, 521 

Tickell, W. L. N. Albatrosses, 521 

Waldbauer, G. Millions of Monarchs, Bunches of Beetles: 
How Bugs find Strength in Numbers, 711 

Wauer, R. Birder’s Mexico, 702 


SPECIAL ISSUES 
THE CANADIAN FIELD-NATURALIST 


THE HISTORY OF THE EXPLORATION OF THE VASCULAR 
FLORA OF CANADA, SAINT-PIERRE ET MIQUELON, AND GREENLAND 
by James S. Pringle 
Volume 109(3) July - September 1995 


A LIFE WITH BIRDS: PERCY A. TAVERNER, 
CANADIAN ORNITHOLOGIST, 1875-1947 
by John L. Cranmer-Byng 
Volume 110(1) January - March 1996 


THE ORCHIDS IN THE OTTAWA DISTRICT 
by Joyce M. Reddoch and Allan H. Reddoch 
Volume 111(1) January - March 1997 


A PASSION FOR WILDLIFE: A HISTORY OF THE CANADIAN WILDLIFE SERVICE 
by J. Alexander Burnett and including selected publications from work 
by the Canadian Wildlife Service, by Anthony J. Erskine 
Volume 113(1) January - March 1999 


Copies of these important issues are available from: 


Business Manager 
Canadian Field-Naturalist 
Box 35069, Westgate P.O. 

OTTAWA, Ontario K1Z 1A2 


$10.00 plus $2.50 handling and mailing each. 


769 


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 
readers and other potential contributors are invited to submit 
for consideration their manuscripts meeting these criteria. 
The journal also publishes natural history news and com- 
ment items if judged by the Editor to be of interest to read- 
ers and subscribers, 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 encouraged to support the 
journal by becoming either members or subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
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 publi- 
cation of the data must have been obtained by the authors. 
Also authors are expected to have complied with all perti- 
nent legislation regarding the study, disturbance, or collec- 
tion of animals, plants or minerals. The place where voucher 
specimens have been deposited, and their catalogue num- 
bers, should be given. Latitude and longitude should be 
included for all individual localities where collections or 
observations have been made. 

Print the manuscript on standard-size paper, doubie- 
space throughout, leave generous margins to allow for 
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key words. Generally, words should not be abbreviated but 
use SI symbols for units of measure. The names of authors 
of scientific names should be omitted except in taxonomic 
manuscripts or other papers involving nomenclatural prob- 
lems. “Standard” common names (with initial letters capi- 
talized) should be used at least once for all species of high- 
er animals and plants; all should also be identified by scien- 
tific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports should not be cited 
here but placed in the text or 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 numer- 
als, and placed on a separate page. Mark in the margin of 
the text the places for the figures and tables. 

Check recent issues (particularly in 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. Photographic reproduction of line drawings, no 
larger than a standard page, are preferable to large origi- 
nals. Prepare line drawings with India ink on good quality 
paper and letter (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, chosen by the Editor. Authors are encouraged to sug- 
gest 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 revi- 
sion — sometimes extensive revision and reappraisal. The 
Editor makes the final decision on whether a manuscript 
is acceptable for publication, and in so doing aims to main- 
tain the scientific quality, content, overali high standards 
and consistency of style, of the journal. 


Special Charges — Please take note 


Authors must share in the cost of publication by pay- 
ing $80 for each page in excess of five journal pages, plus 
$15 for each illustration (any size up to a full page), and up 
to $80 per page for tables (depending on size). Repro- 
duction of color photos is extremely expensive; price quo- 
tations may be obtained from the Business Manager. 
Reprint order forms are included when galley proofs are 
sent to authors. If grant or institutional funds are available, 
we ask authors to defray a higher proportion of the cost of 
publishing, $80 per page for all published pages. Govern- 
ment institutions are expected to pay the full cost of publi- 
cation. Authors must also be charged for their changes in 
proofs. 

Limited journal funds are available to help offset publi- 
cation charges to authors with minimal financial resources. 
Requests for financial assistance should be made to the 
Business Manager when the manuscript is accepted. 


Reprints 


An order form for the purchase of reprints will accom- 
pany the galley proofs sent to the authors. 


FRANCIS R. Cook, Editor 


RR 3 North Augusta, Ontario KOG 1RO, Canada 


770 


otany: Flora of New Brunswick Second Edition — Flora of Florida, Volume | Pteridophytes and 
Gymnosperms — Plant Life Histories: Ecology, Phylogeny and Evolution — Flora of Mount Rainier 
National Park — The Flora of Manitoulan Island and the Adjacent Islands of Lake Huron, Georgian 
_ Bay and the North Channel, Third Edition 


vironment: Food Webs and Container Habitats: The Natural History and Ecology of Phytotelmata — 

The Friendship of Nature, A New England Chronicle of Birds and Flowers — Earth Future: Stories 
from a Sustainable World — Vanishing Borders: Protecting the Planet in the Age of Globalization 
_ —- Pandora’s Poison: Chlorine, Health, and a New Environmental Strategy 


; 
TABLE OF CONTENTS (concluded) 
| 


Jew Titles 
fhe Ottawa Field-Naturalists’ Club Awards April 2001 


News and Comment 


‘roglog: Newsletter of the Declining Amphibian Populations Task Force (47, 48) — Marine Turtle 
Newsletter (94) — Point Pelee Natural History News 1(3), 1(4) — Canadian Species at Risk 
November 2001 — Renew (Recovery of Nationally Endangered Wildlife) Report 11: 2000-2001 
Annual Report — Amphipacifica: Journal of Aquatic Systematic Biology 3(2) 15 November 2001 — 
Conserving Borderline Species: A Partnership between the United States and Canada 


ndex to Volume 115 Compiled by LESLIE DUROCHER 
\dvice to Contributors 


Mailing date of the previous issue 115(3): 30 April 2002 


718 


721 
725 


728 


732 
734 
766 


THE CANADIAN FIELD-NATURALIST Volume 115, Number 4 


Articles 


Great Horned Owl, Bubo virginianus, predation on Richardson’s Ground Squirrels, 
Spermophilus richardsonii GalL R. MICHENER 


Inability to predict geographic origin of Yellow-headed Blackbirds, 
Xanthocephalus xanthocephalus, during migration 
DANIEL J. TWEDT. GEORGE M. LINZ, and WILLIAM J. BLEIER 


Review of the status of the Northern Abalone, Haliotis kamtschatkana, 
in Canada G. S. JAMIESON 


Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports XIV R. R. CAMPBELL 


Status of the Morrison Creek Western Brook Lamprey, Lampetra richardsoni, in Canada 
R. J. BEAMISH, J. H. Youson, and L. A. CHAPMAN 


Status of stickleback species pair, Gasterosteus spp., in Hadley Lake, 
Hasqueti Island, British Columbia TODD HATFIELD 


Status of the stickleback species pair, Gasterosteus spp., in the Vananda Creek watershed 
of Texada Island, British Columbia TOM HATFIELD 


Status of the stickleback species pair, Gasterosteus spp., in Paxton Lake, 
Texada Island, British Columbia ToM HATFIELD and JAUNITA PTOLEMY 


Rapport sur la situation du Brochet d’amérique, Esox americanus americanus, 
au Canada STEPHANIE LACHANCE 


Status of the Weed Shiner, Notropis texanus, in Canada J. HOUSTON 


Status of the Bridle Shiner, Notropis bifrenatus, in Canada 
E. HoLm, P. DUMONT, J. LECLERC, G. Roy, and E. J. CROSSMAN 


Status of the Mira River population of Lake Whitefish, Coregonus clupeaformis, 
in Canada CHERYL D. GOODCHILD 


Updated status report on the endangered Atlantic Whitefish, Coregonus huntsmani 
THOMAS A. EDGE and JOHN GILHEN 


Status of the Grey Seal, Halichoerus grypus, in the Northwest Atlantic 
VERONIQUE LESAGE and MIKE O. HAMMILL 


Status of Harbour Seals, Phoca vitulina, in Canada ROBIN W. BAIRD 


Status of Killer Whales, Orcinus orca, in Canada ROBIN W. BAIRD 


Book Reviews 


Zoology: Birder’s Mexico — The Destruction of the Bison — Prairie Birds: Fragile Splendor in the 
Great Plains — Ecology and Management of Large Mammals in North America — Handbook of 
Birds of the World: Volume 6 Mousebirds to Hornbills — Threatened Birds of the World — 
Research Techniques in Animal Ecology, Controversies and Consequences — The Sibley Guide to 
Birds — Ecotravellers’ Wildlife Guide to Ecuador and its Galapagos Islands — Belize and Northern 
Guatemala: The Ecotravellers’ Wildlife Guide — Millions of Monarchs, Bunches of Beetles: How. 
Bugs find Strength in Numbers — Behaviour and Conservation Biology Series 2 — Turtie 
Conservation — The Pheasants of the World: Biology and Natural History Second Edition — The 
Biology of Plethodontid Salamanders — Birds of Europe — Hoofed Mammals of British Columbia 
— Swallow Summer 


(continued on inside back cover) 


ISSN 0008-3550 


2001 | 


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