eer ee erent _—— HAIR WARD) UN DV ERS Wy Ernst Mayr Library of the Museum of ¢ omparative Zoology The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada fut, a4 January—March 2006 Volume 120, Number 1 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Michaille Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- ments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Donald M. 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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: January—March 2006 (June 2007). Cover: Adult Le Conte’s Sparrow, Ammospiza leconteii, attending to the nest. 21 July 1999. Photo by Michael Patrikeev. See arti- cle New nesting records of the Le Conte’s Sparrow — pages 22-26. Mc UBRARy, UL 3 1 mp7 ARy UN; VERGO THE CANADIAN Ty FIELD-NATURALIST Volume 120 2006 Volume 122 The Ottawa Field-Naturalists’ Club Transactions Promoting the study and conservation of northern biodiversity since 1880 THE OTTAWA FIELD-NATURALISTS’ CLUB OTTAWA CANADA The Canadian Field-Naturalist January—March 2006 Volume 120, Number | Predicting Bird Oiling Events at Oil Sands Tailings Ponds and Assessing the Importance of Alternate Waterbodies for Waterfow!: a Preliminary Assessment ROBERT A. RONCONI Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada Present address: Department of Biology, University of Victoria, P.O. Box 3020 Station CSC, Victoria, British Columbia V8W 3N5 Canada; e-mail: rronconi@uvic.ca Ronconi, Robert A. 2006. Predicting bird oiling events at oil sands tailings ponds and assessing the importance of alternate waterbodies for waterfowl: a preliminary assessment. Canadian Field-Naturalist 120(1): 1-9. Tailings ponds are an integral part of oil sands mining development in northeastern Alberta, but waterfow! and shorebirds often land in these ponds during spring migration where they may become covered with oil. For decades, managers have developed and implemented methods for deterring birds from landing in these ponds, yet no deterrent strategy is fully effec- tive. Therefore, to enhance deterrence strategies, it will be important to understand the environmental conditions that influ- ence bird use of tailings ponds. This study quantified waterfowl flights over, and use of, tailings ponds and compared this use to waterfowl activity at natural waterbodies in the region over a single spring migration period. Results suggest that waterfowl are most likely to land on tailings ponds before lakes have thawed, after which migratory ducks appeared mainly to use natural waterbodies for migratory stopover sites. Very high numbers of waterfowl were observed on one waterbody, Kearl Lake, suggesting that this lake may be of greater importance to spring staging waterfowl than previously thought. A small sample of birds oiled at tailings ponds were examined in relation to spring weather conditions. Logistic regression analysis demonstrated that the probability of birds being oiled tended to increase with precipitation levels. Results of this study suggest that (1) preservation of natural waterbodies may play an important role in minimizing bird use of tailings ponds, and (2) future bird deterrence efforts should especially aim to deter birds during rainy weather conditions when birds may be more likely to become oiled. These results were from a small sample size, are preliminary in nature, and should be interpreted with caution. A concerted and careful effort to collect and thoroughly analyze long-term records of oiled birds may reveal important environmental effects predicting bird oiling events. Key Words: waterfowl, oil sands, mining, tailings ponds, lakes, In northeastern Alberta, oil sands development has grown considerably over the past several decades. This industry promises to grow into the 21‘' century since recent oil reserve estimates have placed Canada sec- ond on the world list of oil reserves with 175 billion barrels of oil in oil sands deposits (CAPP 2003*). Al- though most of these reserves are extractable with new drilling technologies, an estimated 20% are recoverable only with open pit mining practices (CAPP 2003*). One of the primary environmental concerns stemming from oil sands mining is the existence of tailings ponds which have oil floating on the surface. Birds landing on these ponds might become covered in oil and die. This problem has been recognized since the initial development of oil sands mining (SCL 1973; Schick and Ambrock 1974*; Ward et al. 1976*; Hennan and Munson 1979) and significant effort has been dedicat- ed to researching the conditions that promote and pre- vent birds from becoming oiled (Boag and Lewin 1980; Gulley 1980; Yonge et al. 1981*; Ronconi and St. Clair weather, migration, radar, conservation, Alberta. 2006; also see review by Golder Associates 2000*). Nevertheless, birds still continue to be oiled and oil sands companies carry on their search for better means of reducing these incidents. Previous oil sands research recommended the cre- ation of uncontaminated waterbodies in close proximity to tailings ponds, allowing birds alternate landing sites (Gulley 1980; Golder Associates 2000*). Moreover, research in other industrial contexts indicates that birds can be deterred from an area more effectively when alternate roosting sites (including ponds) are available (Martin and Martin 1984; Gosler et al. 1995; Stevens et al. 2000). Yet others have argued that such diversion- ary ponds would only draw more birds to the region (Allen 1990). Instead, the preservation of naturally occurring waterbodies may be an optimal solution to provide alternate landing sites for birds, particularly if these alternative bodies are already used by migrating birds in the region. Comprehensive surveys of water- fowl at the major waterbodies in the vicinity of Alber- 2 THE CANADIAN FIELD-NATURALIST ta’s oil sands were conducted prior to oil sands devel- opment (Hennan and Munson 1979), but the current status of these waterbodies and their use by migratory birds are unknown. Moreover, subsequent surveys of waterbird numbers at natural waterbodies (Shortt 1985*; Van Meer and Arner 1985*) did not compare levels of bird activity between these natural ponds and nearby tailings ponds. Such information would make it possible to assess the importance of alternative water- bodies as stopover sites during migration, and to assess temporal changes within seasons in the attractiveness of tailings ponds to waterfowl. In addition to information about the alternative stopover sites for migrating birds, researchers and man- agers have also sought to understand the conditions under which birds become oiled on tailings ponds. Over a three-year period, Gulley (1980) identified three main conditions as determinants of the numbers of birds that were killed each year. First, more birds were oiled when more birds were migrating (Gulley 1980), though the synchrony of migration is not fully predictable and tends to vary among both years and species (Bellrose 1976). Second, cold spring weather and delayed ice break-up correlated with bird oilings. Finally, greater than average seasonal rainfall has been correlated with higher rates of oiled birds. In contrast, other studies failed to observe any environmental (wind, tempera- ture, precipitation) effects on the probability of bird landings in tailings ponds (Ronconi and St. Clair 2006). Yet neither analysis examined the frequency of bird oiling events in relation to daily weather patterns and this information may be most likely to reveal the causes of variation. The study is situated at one of more than 10 oil sands tailings ponds operating in northeastern Alberta (Gold- er Associates 2000*), all of which are situated along a major migratory flyway for waterfowl traveling to the Peace-Athabasca Delta, an internationally signifi- cant staging area (Bellrose 1976; Hennan and Munson 1979). The objectives in this study were three-fold. First, | examined the seasonal abundance and direc- tion of migratory bird movements over tailings ponds. Second, I compared numbers of waterfowl that landed at tailings ponds with numbers of birds at natural ponds in a broader region and with historical data. Finally, I assessed potential correlations between daily weather conditions and the frequency of oiled birds found dur- ing April and May of 2003. Because data were available only for a single season, I present these as preliminary results and encourage future researchers to investigate these findings over longer time periods. Methods The study was undertaken at the Muskeg River Mine (MRM), Albian Sands Energy Ltd., located approxi- mately 75 km north of Fort McMurray, Alberta (Fig- ure 1). During April and May 2003, the mine was beginning initial oil production stages. The tailings Vol. 120 pond complex consists of three ponds totaling ~ 740 ha of surface area when full, but water coverage during this study period was approximately 530 ha. Natural waterbodies were selected for surveys based on acces- sibility, size, and proximity to the MRM tailings ponds. These included two lakes near the mine, four lakes south of the mine near other oil sands development, and eight lakes north of the mine that were isolated from development (Figure 1). As these lakes were sit- uated along a North-South axis relative to the MRM, all lakes were potentially usable by waterfowl] migrat- ing along this corridor. Airport radar systems have been used to study bird migration patterns in Alberta (Richardson and Gunn 1971), and marine radar has been used by ornitholo- gists to study various aspects of bird activity (Cooper et al. 1991; Burger 1997; Bertram et al. 1999; Gau- threaux and Belser 2003). In this study, marine type radar was used to assess spring migratory activity over the MRM tailings ponds between 3 and 29 May 2003. Migratory activity was measured by the groups of birds flying over the study area (groups rather than individ- uals are identifiable by radar) and the direction of flight. Radar observations were made from 0.5 hours before sunrise to 4 hours after sunrise. This period was con- sidered a high migratory period in northeastern Alberta (Richardson and Gunn 1971). Owing to some techni- cal difficulties, directional data were not recorded for the first four days of this period, and data collection was not possible on six other days because of heavy precipitation that obscured radar observations. Tailings pond and lake surveys were conducted from fixed observation points using binoculars and spot- ting scopes. Tailings ponds were surveyed regularly from 18 April to 29 May, and lakes were surveyed 9 to 28 May as most lakes were still completely or partially frozen during the first week of May. Observers counted individual ducks, gulls, coots, loons, and grebes on the water. No more than two or three geese were observed during any lake count; therefore, goose counts were omitted from the results. Because of the large size of McCleland Lake, all portions of it were not simulta- neously visible, thus counts were only a minimum esti- mate of bird numbers on that lake. Kearl Lake was surveyed more often than the other natural waterbod- ies because it was nearest to the MRM tailings ponds and thus most likely to offer an alternate landing site for migratory birds. During the mid-1970s Mildred Lake was dammed, converting most of the lake into a tailings pond and leaving only a portion of the lake in its semi-natural state. Surveys of Mildred Lake in- cluded only the semi-natural portion. When applicable, results were compared to previous surveys (see Hen- nan and Munson 1979 for detailed waterfowl surveys in the region). Reports of oiled birds on the tailings ponds were col- lected from 10 April to 29 May, inclusive. Although systematic searches for oiled birds were not conduct- 2006 ed, observers spent 4 to 8 hours observing waterfowl] at the tailings ponds on most days thus ensuring some consistency in effort throughout this period. Weather data for wind speed and direction, temperature, and rainfall were recorded from meteorological stations on site, and snowfall events were converted into rain- fall equivalency. Mean temperature is reported here as the daily mean of maximum and minimum tempera- tures. Wave conditions on the ponds and percent cloud cover were recorded by observers. All occurrences of oiled birds are reported (below); however, analyses of weather effects were conducted only on oiled birds that were still alive when found, an assumption that those birds became oiled either the day they were found or the previous night. For analysis by logistic regression, oiled birds were the dependent variable (1 = days with oiled birds; 0 = days with no oiled birds). Days when observers were not present on site were excluded from analyses. Weather data were the independent variables (wind, temperature, precipitation, wave conditions, % cloud cover). Except for wave conditions, analyses of weather effects were assessed for three time periods: date of oiled bird finding, one day prior (i.e. previous 24 hour conditions), and two days prior (i.e. previous 24-48 hour conditions). Wave condition was only analyzed for the first two time periods since it was considered likely that rough water effects would not persist for as long as two days. Logistic regression analysis (SPSS 11.5) was used to assess weather effects and to build a model for pre- dicting their effects on the probability of birds being oiled. Owing to small sample sizes of data from a single season, a liberal alpha value (0.10) was used for logis- tic regression analysis. Tests were conducted for inter- ance ®o xo) cs oa i — 80 2 3 ire £23 6 =o Oo Ta) [@) Oo 40 ®o DO =o oO 8 20 Oo — = © = as 0 05/05/03 42/05/03 19/05/03 Date RONCONI: PREDICTING BIRD OILING AT OIL SANDS PONDS ; ieee H xs @ / © Lakes Surveyed if = Ga Muskeg River Mine ) ‘a 05 @ Fort McMurray | \3 ~~ Rivers 2s ao in eee Major Lakes ___| Wood Buffalo N.P FIGURE 1. Map of study area showing the location of the Muskeg River Mine and surrounding natural water- bodies surveyed in 2003. Lake names are as follows: (1) Poplar, (2) Ruth and Crane, (3) Mildred, (4) Isa- dore’s, (5) Kearl, (6) McCleland, (7) Unknown Lakes, (8) Go Go, (9) Herb, (10) Six Lakes, (11) Larocque, and (12) Net. NW &N Flights NE & E Flights SE & E Flights SW & W Flights Bird Groups per Hour 26/05/03 > FiGurE 2. Migration patterns from radar data at the Muskeg River Mine tailings ponds, 3-29 May 2003. Bird groups were the unit of measure because group size could not be inferred from radar images. Gaps in the data set resulted from days of heavy precipitation which obscured radar observations. 4 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Numbers of waterfowl at the Muskeg River Mine (MRM) and surrounding natural waterbodies, 2003. Distance (km) Other Lake from Water Location Date Coordinates MRM Size! Gulls Ducks _ birds Muskeg River Mine Tailings Ponds 18 April 57°13'47"N, 111°35'10"W 0 0 B85 3 20 April 0 8 0 23 April 0 32 0 24 April 0 13 0 25 April 4 13 0 26 April 0 22 4 27 April 5) 55 0 03 April 0 86 0 04 April 0 90 0 05 April 0 16 0 06 May 0 166 58 07-29 May 0-14 < 28 <2 Near Muskeg River Mine Kearl Lake 09 May 57°18'08"N, 111°15'06"W 19 > 15 104 1] 15 May 300 2700 present 19 May 440 859 30 May 2 8 9 Isadore’s Lake 26 May 57°13'43"N, 111°36'29"W 1 = 66 102 2) North of Muskeg River Mine McCleland Lake? 23 May 57°30'35"N, 111°21'16"W 32 >> 12 645 5 Unknown Lake 23 May 57°50'25"N, 111°06'29"W V2 < 4 0 0 Unknown Lake 2 23 May 57°50'28"N, 111°06'46"W 72 < 0 4 2, Go Go Lake 23 May 57°52'48"N, 111°02'56"W Ty « 0) 0 2 Herb Lake 23 May 57°54'13"N, 111°02'11"W 80 = 0 5 1 Six Lakes 24 May 57°58'SO"N, 111°01'45"W 88 = 0 0 3 Net Lake 24 May 58°07'23"N, 110°46'55"W 108 = 0 5 3 Larocque Lake 24 May 58°06'S1"N, 110°49'11"W 107 = 3 0 1 South of Muskeg River Mine Mildred Lake 25 May = 57°02'45"N, 111°33'52"W 18 = 0 0 0 Crane Lake 25 May 56°59'45"N, 111933'12"W 24 0 158 present 28 May 0 106 5 Ruth Lake 25 May 56°59'22"N, 111°34'06"W Ds > 19 306 present 27 May 12 350 1 Poplar Reservoir Dil Maya y5 62553 20Ns ile29 500i 3p = 0 15 2 Relative to Muskeg River Mine tailings ponds = (approximately equal), > (larger), < (smaller). 2Tncludes coots, loons, grebes. 3 Entire lake was not observable, therefore counts are minimum numbers of birds. actions and confounding variables. For detailed descrip- tion of the model-building approach see Ronconi and St. Clair (2006) and Hosmer and Lemeshow (1989). Results Counts of bird groups by radar provided an indica- tion of overall bird activity relative to the numbers of birds landing at tailings ponds. The radar system was operational from 3 — 29 May, during which two peaks in migratory activity were observed (Figure 2). Peaks in migratory flights (NW & N flights) coincided with two peaks in bird activity (bird groups per hour). How- ever, a third peak in groups per hour on 27 May showed no corresponding peak in NW & N flights. Notwith- standing the three peaks, these data document high migratory volumes over most of May when counts of waterfowl on lakes were conducted. However, the num- bers of ducks that also landed on the tailings ponds (Table 1) were consistently low over this same period with no comparable peaks. Counts of birds at tailings ponds and natural water- bodies showed much variability and peaks occurred at different times for different lakes. Counts of ducks and other waterbirds at the MRM tailings ponds (Table 1) showed peaks in activity from 26 April through to 6 May, after which activity dropped considerably and remained low throughout May. Conversely, high counts of birds on surrounding lakes were observed in May (Table 1), though comparable counts from April were not possible because the natural waterbodies were still frozen at that time. In particular, Kearl Lake showed a peak around mid May with an estimated 2700 ducks 2006 and 300 gulls on 15 May. As numbers were low at Kearl Lake in early May, these data, in combination with low May counts at tailings ponds, suggest that bird use of tailings ponds may have shifted to natural waterbodies for much of May. North of MRM, counts showed very low numbers except for McCleland Lake with over 600 ducks on 23 May. This high count suggests that McCleland was still being used as a migratory stopover at this time, but lakes farther north were likely only occupied by small numbers of breeding pairs. Between 25 and 27 May, counts of lakes south of MRM had higher numbers than maximum counts at MRM tail- ings ponds throughout April or May, but by this date many of these birds may have been resident breeders. Although duck use of tailings ponds appeared low in comparison with natural waterbodies, overall duck numbers for this region were higher than historical sur- veys reported. Comparisons with Hennan and Mun- son’s (1979) surveys showed several changes (Table 2). Duck numbers at McCleland Lake fell within the range of historical counts and were higher than four of the five years surveyed between 1973-1977, even though McCleland was only censused once in 2003. The highest count at Kearl Lake in 2003 was 4.6 times the maximum recorded by once annual surveys be- tween 1973 and 1977. South of the MRM, Mildred Lake showed no use in 2003, in contrast to abundant use in previous surveys, whereas numbers were high- RONCONI: PREDICTING BIRD OILING AT OIL SANDS PONDS 5 er at Ruth Lake than were recorded during the 1970s. Due to inconsistencies in data collection methods and dates of surveys, we can not speculate on trends or changes in waterfowl abundance since the 1970s. However the dramatically high count of ducks on Kear! Lake suggest that this lake may be more heavily used as a migratory stopover site than might have been sus- pected from earlier surveys. Fifteen oiled birds were observed at the MRM tail- ings ponds during the study period (Table 3), nine of which were alive when found. Of these, four Lesser Scaup (Aythya affinis) were lightly oiled and able to fly, making it probable that these were non-fatal oiling events. The remaining live oiled birds included one goose, one shorebird, one gull and two ducks. Days when live oiled birds were found were plotted in rela- tion to weather variables (Figure 3). A logistic regres- sion analysis was conducted on 47 days of observa- tions, six of which had reported live oiled birds. This analysis found no effect of temperature, sea-state, wind direction, or cloud cover. Univariate analyses found date (P = 0.160) to be marginally significant, and rain- fall (P = 0.086), rainfall during the previous 24 hours (P = 0.066), and wind speed during the previous two days (P = 0.050) as significant variables affecting the probability of birds being oiled. However, when com- bined in a multivariate analysis no variables retained significance but the two rainfall variables were only TABLE 2. Comparisons of spring 2003 duck counts with historical counts from 1973-1977 (Hennan and Munson 1979). Maximum Number of Ducks Recorded During Spring Migration Location 1973 1974 MRM Tailings Ponds ~ = Kearl Lake 529 586 McCleland Lake! 194 1154 Mildred Lake? 246 313 Ruth Lake 113 20 Poplar Reservoir - ~ 1975 1976 1977 2003 - - - 166 24 37 140 2700 225 360 195 645 136 17 - 0 26 1] - 350 16 l 14 15 ' For 2003 surveys the entire lake was not observable; therefore, count is a minimum estimate of ducks ca. 1975 most of Mildred Lake was converted into a tailings pond. Surveys in 2003 checked only the portion of Mildred Lake that remains in its natural state. TABLE 3. Summary of oiled birds found at the Muskeg River Mine tailings ponds, April and May 2003. Date Common Name Scientific Name 26 April goose spp. unknown 03 May Common Goldeneye Bucephala clangula 19 May Lesser Scaup Aythya affinis 20 May Bonaparte’s Gull Larus philadelphia 21 May small shorebird Calidris spp. 21 May Lesser Yellowlegs Tringa flavipes 26 May Northern Shoveler Anas clypeata 26 May Northern Shoveler Anas clypeata 26 May small Shorebird unknown 26 May medium passerine unknown 26 May large shorebird unknown Group Size Status Oiled Status 1 Live heavy — >80% 1 Live heavy — 80% 4 Live light < 10%! ] Live heavy — 70% 1 Live heavy — 60% 1 Dead moderate — 30% ] Live heavy > 50% 1 Dead heavy — 100% 1 Dead heavy — 100% 1 Dead heavy — 90% ] Dead heavy — 100% ' Lightly oiled but able to fly. 6 THE CANADIAN FIELD-NATURALIST marginally non-significant (P = 0.107 and 0.105). To examine the apparent, though non-significant, effect of rainfall on birds being oiled, the cumulative rainfall was summed for the day of plus day before oiled birds were found and this had a significant effect (Wald = 4.973, d.f. = 1, P = 0.026) on the probability of birds being oiled. The positive coefficient of this variable indicated that the probability of a bird becoming oiled increased with higher rainfall. With the cumulative rainfall vari- able, the model was significant (-2LL = 30.264, model - = 5.636, df. = 1, P =0.018), but the model fit was weak (Nagelkerke’s r? = 0.212, Hosmer and Lemeshow test ¥° = 1.330, df. = 1, P= 0.095). Small sample size likely contributed to the marginal significance of vari- ables and the poor fit of the model, yet the persistence of rainfall effects indicate that precipitation was relat- ed to birds being oiled this year and this may be true more generally. Discussion Timing of Spring Migration Although the radar system was not operational in April, visual observations suggested that migration activity in the study area was fairly low in April and early May and that migration peaked in mid-May. The radar was also not in operation (due to weather) for six days in May near peak migration (Figure 2); there- fore, migration records presented here may have missed some critical portions of the migration. Nevertheless, the results on migration timing agree with other stud- ies in northeastern Alberta. Richardson (1969) reported peak spring migration over east-central Alberta in May. Aerial surveys of lakes in this region during 1972 and 1973 reported peak numbers on 15 May and 25 May, respectively (SCL 1973). Likewise, Schick and Am- brock (1974*) showed peak migration to occur in mid- dle and end of May in 1973. During aerial surveys in 1976/1977, Hennan and Munson (1979) reported peak spring migration for ducks in early May, with most abundant species being scaup (Aythya sp.), Ring- necked Ducks (Aythya collaris), Buffleheads (Buce- phala albeola), and to a lesser extent Mallards (Anas platyrhynchos). Annual variation in timing of spring migration is likely to vary with timing of ice break-up of the Athabasca River and also the Peace-Athabasca Delta (Schick and Ambrock 1974*). Although the timing of migration in northeastern Alberta is relatively well known, the composition and numbers of birds passing through the area are less well known. During the same time period of this study in 2003, Ronconi and St. Clair (2006) reported the fol- lowing numbers of birds flying over the tailings ponds: 1140 ducks, 2132 shorebirds, 11 339 geese, 218 swans, 965 gulls, 56 terns and 205 others (loons, grebes, cranes, herons, cormorants, coots). These numbers give some indication of guild composition during migration but should be used cautiously as such because these were visual, daytime observations and many birds in Alberta migrate at night and at high altitudes (Richard- Vol. 120 son 1971; Blokpoel 1973; Blokpoel and Burton 1973). Hennan (1972) reported hundreds of thousands of waterfowl using the Peace-Athabasca Delta as a spring staging area, thus providing some indication of the volume of birds that may pass over the tar sands area each year. Nevertheless, many individuals of some waterfowl species may also take alternate migration routes to Alaska or nest in southern Canada (Miller et al. 2005). Finally, in this study area we may also be observing resident as well as migratory waterfowl. This study makes no distinction between the two groups; howev- er, the high proportions of northwesterly and northerly flights (Figure 2) suggest that most of the birds ob- served were migrants. It is likely that the small num- bers of birds observed on the tailings ponds in May and on some lakes in late May were residents rather than stopover migrants. Surprisingly, the ecology of waterfowl at staging and stopover areas is fairly poorly understood (Arzel et al. 2006). Numbers of Birds on Ponds and Lakes Despite the apparent peak in migration around mid- May, the numbers of ducks on the tailings ponds peaked sooner than this. Working in a nearby area, Gulley (1980) found that duck dependence on tailings ponds was higher when spring ice break-up was de- layed. In 2003, ice break-up on the nearby Athabasca River began 23 April and was finished by 1 May, yet many of the nearby lakes remained at least partially frozen through the first week of May. Duck surveys at Kearl Lake showed low numbers on 9 May when this lake was still partially frozen, but numbers grew by 23 times only one week later when the lake was ice- free. These results are mainly qualitative, but they sup- port the hypothesis of others (Gulley 1980; D. Mar- tindale, personal communication) that duck use of tailings ponds is highly dependent on the availability of open water elsewhere. After 9 May, duck numbers at the MRM tailings ponds were seldom above 10 indi- viduals, while thousands of individuals were using Kearl Lake less than 20 km away. Nevertheless, these conclusions about seasonal change in dependence on tailings (early spring) to natural waterbodies (mid to late May) are tenuous due to several inconsistencies in survey methodology: (1) Kearl Lake was the only natural water body surveyed more than twice, (2) most natural waterbodies were surveyed only in late May and might have missed peak migration, and (3) lack of personnel did not allow for simultaneous surveys of tailings ponds and natural waterbodies. Bird move- ments between wetlands may have been a concern; however, the extremely low numbers of ducks at the tailings ponds after 7 May suggest little, if any, move- ment between artificial and natural ponds after this date. Finally, no data are available on wetland charac- teristics, which were beyond the scope of this project but may have had significant influence on waterfowl use of waterbodies. 2006 oe Mean Wind Speed cast sesees Mean Temperature 5 20 Daily Precipitation Ss = One or More & Live Oiled Bird Found E = 15 FEMS ae O wee le Se ur 2S aoa OO or - ® ns U \ ep) 12) & 5 : f -10 g g gf RONCONI: PREDICTING BIRD OILING AT O11. SANDS PONDS 7 . *, cer A) + gle i 1 a hee yo tN ae ia Ce eee | Reece o 7 & & se \ 9 IN a> VY FIGURE 3. Weather patterns with respect to dates when live oiled birds were found at the Muskeg River Mine tailings ponds, April and May 2003. Comparisons with census results from the 1970s sug- gest some changes in lake use. However, duck activity levels can vary widely within seasons, as was indicated in our results and other studies (e.g. SCL 1973; Gul- ley 1980; Van Meer and Arner 1985*; Shortt 1985*). The maximum counts between 1973 and 1977 were typically based on only one or two surveys during May, making them similarly prone to high variability. Al- though qualitative and tenuous, these comparisons rein- force the belief (SCL 1973; Schick and Ambrock 1974*; Ward et al. 1976*; Hennan and Munson 1979) that some lakes are important as stopover sites for migratory ducks, though not as major staging areas. As with our results, censuses from 1973 to 1977 ranked McCleland and Kearl lakes among the most important stopover sites for ducks (Hennan and Munson 1979). Other lakes in the region with high numbers included Little McCleland, Mildred, Horseshoe, and Saline lakes (Hennan and Munson 1979). The lack of ducks on Mil- dred Lake in 2003 is not surprising considering only a small portion of the lake remains in its natural state. The lack of ducks on remote lakes north of McCleland Lake was not surprising given the relatively small size of the lakes surveyed. Nonetheless, Hennan and Mun- son (1979) identified other remote lakes with high wat- erfowl densities, though these were not accessible for surveys by land in 2003. Generally the oil sands region is not recognized as an important staging area for migratory waterfowl (SCL 1973; Schick and Ambrock 1974*; Ward et al. 1976*; Hennan and Munson 1979), although some staging does occur nearby. The Peace-Athabasca Delta, north of oil sands regions, has been recognized as an internationally significant staging area for hundreds of thousands of waterfowl (Hennan 1972). Further south of Fort McMurray, Gordon Lake was previously rec- ognized as an important waterfowl staging area with spring numbers greater than 5600 in 1973 and 1974 (Hennan and Munson 1979). The count of 2700 water- fowl at Kearl Lake in 2003 suggest that this lake may bear greater importance as a staging area than might have been concluded from those earlier surveys, but further surveys may be needed to verify annual varia- tion in waterfowl use of this lake. Surveys of other nat- ural waterbodies also revealed that smaller ponds sup- port small populations of birds throughout the spring and summer (Shortt 1985*). Because availability of natural waterbodies may minimize duck dependence on tailings ponds, further investigations of waterfowl use of these waterbodies is merited. Should it continue to appear that they are important to waterfowl, future oil sands developers may consider conserving these locations and avoiding the establishment of tailings ponds in close proximity to them. 8 THE CANADIAN FIELD-NATURALIST Probability of Oiling Logistic regression analysis of several weather vari- ables found wind speed and precipitation to be the only variables that might predict the few observations of oiled birds in 2003. Strong winds may mix waters and spread oily films over wider areas, hence creating greater risk of oiling, but with consistent winds oily sheets may also be pushed to one shore of the pond. Thus any link between winds and oiling may be com- plicated. A two-day cumulative rainfall measure best predicted days with oiled birds, but rainfall the day of an oiling event and rainfall from the previous 24 hours showed some possible predictive ability. These uncer- tain results, along with small sample size, suggest only a potential effect of precipitation. Results of this study support a previous analysis across years (Gulley 1980) which showed that precipitation levels affected num- bers of waterfowl deaths. Although Ronconi and St. Clair (2006) found no correlation between weather and bird landings on tailings ponds, the more important factor may be whether or not birds become oiled when they land. If precipitation increases the frequency of birds be- ing oiled, as these results cautiously suggest, I specu- late that it may be caused by two possible mechanisms. First, poor weather conditions such as rain, strong winds, headwinds, and cold temperatures can delay migration, cause changes or a temporary reversal in migration direction, and cause migratory birds to be temporarily grounded (Richardson 1978; Elkins 1983). Waterfowl in particular appear to be erratic in their mig- ratory behaviour which may be influenced by break-up of ice cover (P6ysd 1996), temperatures (Austin et al. 2002), weather fronts (Custer et al. 1997), and espe- cially wind patterns (Bergman 1978; Liechti 1993; Bruderer 1994; Alerstam and Gudmundsson 1999). The significance of precipitation as a predictor of oil- ing may be due to correlations between rainfall and other weather changes, which may together be affect- ing bird movements and halting of migrations. The more frequently migratory waterfowl are forced to move and stop during migration, the greater the risk of encountering oil sands ponds and oil on the water sur- face in these ponds. Second, precipitation may additionally increase oil- ing probabilities through birds experiencing reduced visibility during landings under rainy and snowy con- ditions. Without clear views of the water, they may be more likely to land in patches of oil which might other- wise be avoided. Reduced visibility as a causal mech- anism may also explain the non-significance of wind speed, wind direction, and temperature since these variables should be less likely to affect the visibility of oil patches. Snow could affect in-flight visibility of birds, and on two of six days with oiled birds, the precipitation that occurred was a mix of rain and snow. Moreover, for three of the remaining days with oiling Vol. 120 events (19, 20, and 21 May), heavy snow had fallen for two days previously (17 and 18 May). Although fog was not measured in this study, fog may also limit vis- ibility and thus influence oiling events (Gulley 1980). Careful collection and thorough statistical analyses of long-term oiled bird records at tailings ponds may reveal important trends and weather association. These data could be valuable to improving the effectiveness of bird deterrent strategies, and I encourage oil sands companies to standardize collection protocols and pool their data for such an investigation. Conclusions As the development of oil sand mines continues in the 21*' century, tailings pond and bird oiling problems will persist, and findings from current and previous studies will become more relevant to managers seek- ing a solution to this issue. Findings from this study suggest the importance of maintaining natural water- bodies in the region, especially those that are already used as migratory stopover sites for waterfowl. The significance of rainfall affecting numbers of birds being oiled suggests that bird deterrence efforts could benefit from enhancement on rainy days, and future deterrent development should pursue strategies that are effective during rainy weather more so than during clear condi- tions. With decades of data on oiled birds, and 30 years of research on bird deterrence, the oil sands industry in northern Alberta is positioned to offer significant insight into waterfowl management issues for future develop- ment here and at other sites of industrial development. Acknowledgments Logistical support and funding was provided by Albian Sands Energy Ltd., with additional logistical support from the University of Alberta, Edmonton. I thank Colleen Cassady St. Clair for support and guid- ance throughout this study and for reviewing drafts of this manuscript. I am grateful for support from Darrell Martindale, excellent field assistance from Andrew Forrest and Sarah Wong, and technical support from Will Neagle and Peregrine Systems Inc. Field work for this project was made possible with the help of Albian Sands employees Christine Theriault, Maurice Kay, Bob McKenzie, Thor Einarson, Brian Kean, and others on the Tailings and Site Services Teams. 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Received 17 February 2004 Accepted 17 May 2006 A Test of Interspecific Effects of Introduced Eastern Grey Squirrels, Sciurus carolinensis, on Douglas’s Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia YEEN TEN Hwanc!* and SERGE LARIVIERE!” ‘Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada *Delta Waterfowl Foundation, R.R. #1, Box 1, Site 1, Portage La Prairie, Manitoba RIN 3A1 Canada >Corresponding author: Department of Biology, University of Western Ontario, 1151 Richmond Street N., London, Ontario N6A 5B7 Canada; e-mail: yeenten@ gmail.com Hwang, Yeen Ten, and Serge Lariviére. 2006. A test of interspecific effects of introduced Eastern Grey Squirrels, Sciurus caro- linensis, on Douglas’s Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia. Canadian Field-Naturalist 120(1): 10-14. We compared the effects of absence and presence introduced Eastern Grey Squirrels (Sciurus carolinensis) on the demography of native Douglas’s Squirrels (Jamiasciurus douglasii) in two urban parks in Vancouver, British Columbia: Ecological Reserve #74 in Pacific Spirit Regional Park (Douglas’s Squirrel only) and Stanley Park (Douglas’s and Eastern Grey squirrels). Based on the exploitative competition hypothesis, we predicted that in the presence of introduced Eastern Grey squirrels, Douglas’s Squirrels would occur at lower densities, have larger home ranges, lower body mass, and poorer reproduction. Using mark-recapture methods, we found no differences in density, home range, or body mass of Douglas’s Squirrel between parks. However, the proportion of breeding Douglas’s Squirrels was higher in Ecological Reserve #74 in the absence of Eastern Grey Squirrel, than in Stanley Park. We found no evidence that Eastern Grey Squirrels are displacing Douglas’s Squirrels in Stanley Park, but less conspicuous negative effects such as reduced breeding propensity may still reflect the competitive interactions of the two squirrel species. Key Words: Density, home range, body mass, reproduction, Eastern Grey Squirrel, Sciurus carolinensis, Douglas’s Squirrel, Tamiasciurus douglasii, exploitative competition, British Columbia. In western Canada, Eastern Grey Squirrels (Sciurus carolinensis) were introduced to Stanley Park in the city of Vancouver, British Columbia, shortly before 1914. The three to four pairs of introduced Eastern Grey Squirrels came from eastern Canada (Robinson and Cowan 1954). In the early 1980s, Eastern Grey Squirrels colonized the rest of the city of Vancouver, including many of its urban parks. Since then, Eastern Grey Squirrels have spread to many parts of the Greater Vancouver Regional District (Gonzales 1999, 2000). On the mainland in southwest British Columbia, the native tree squirrel is the Douglas’s Squirrel (Jamias- ciurus douglasii); it is the ecological counterpart of the Red Squirrel (7. hudsonicus) that is widespread in temperate forests of North America (Steele 1998, 1999). The Eastern Grey Squirrel is native in eastern North America (Hall 1981) and its geographical range over- laps broadly with that of the Red Squirrel. Although Red Squirrels are associated with conifer forests and Eastern Grey Squirrels with mast-bearing hardwood forests, the two species often coexist in mixed forests or urban parks (Riege 1991). Forty years after Eastern Grey Squirrels were introduced to southwestern British Columbia, Robinson and Cowan (1954) conducted a live-trapping and observational study on Eastern Grey Squirrels in Stanley Park. They noted that, in decidu- ous habitat in Stanley Park, Eastern Grey Squirrels lived at high densities together with lower densities of Douglas’s Squirrels. They suggested that the East- ern Grey Squirrels outcompete Douglas’s Squirrels for food, and thus restrict Douglas’s Squirrels to areas of predominately coniferous habitat (Robinson and Cowan 1954). Whether competitive interactions between the two species still exist today (> 80 years since the intro- duction of Eastern Grey Squirrels) remains unknown. In Great Britain, Eastern Grey Squirrels were intro- duced at the end of the 19" century, and have caused the decline of the smaller native European Red Squir- rel (Sciurus vulgaris) from its former range (Gurnell and Pepper 1993; Wauters and Gurnell 1999; Tean- gana et al. 2000). The mechanisms of replacement of European Red by Eastern Grey squirrels are not yet fully understood (Skelcher 1997). Juvenile recruitment of European Red Squirrels is typically lower when Eastern Grey Squirrels are present (Wauters et al. 2000, 2001). Also, diseases such as parapoxvirus have been hypothesized to facilitate the replacement of European Red Squirrels by Eastern Grey Squirrels in the United Kingdom (Tompkins et al. 2003). The Eastern Grey Squirrel averages 400-700 g in body mass (Koprowski 1984), and is larger than Doug- las’s Squirrel, which averages 250-300 g (Steele 1999). Douglas’s Squirrels feed primarily on coniferous seeds, new shoots of conifers, green vegetation, acorns, nuts, mushrooms, fruits, and berries. Also, Douglas’s Squir- rels actively defend food middens within their territory 10 2006 (Steele 1999). In its native range, the Eastern Grey Squirrel feeds heavily on hickory nuts, beechnuts, acorns, and walnuts (Koprowski 1984); however, intro- duced Eastern Grey Squirrels in British Columbia feed extensively on maple, oak, hazelnut, mushrooms, berries and pine cones (Robinson and Cowan 1954). Both species prefer seeds, and other plant foods pre- dominate in the diet only when seed crops are small (Moller 1983). Also, in Stanley Park, a few Eastern Grey Squirrels depend almost entirely on “hand-outs” from park visitors as an artificial food source. Eastern Grey Squirrels living in close proximity to humans often venture out of the forests and feed on artificial food sources. Eastern Grey Squirrels have a dominance hierarchy and are not territorial (Flyger 1955; Thompson 1977) whereas Douglas’s Squirrels are territorial year round (Steele 1999). Territorial behaviour exhibited by Doug- las’s Squirrels is mainly used against conspecifics sim- ilar to Red Squirrels (Smith 1981). Intruding Eastern Grey Squirrels are generally free to enter their terri- tory (Robinson and Cowan 1954). The interspecific effects of the introduced Eastern Grey Squirrels on the native Douglas’s Squirrels are unknown in southwestern British Columbia. We tested the hypothesis that Eastern Grey Squirrels negatively impact native Douglas’s Squirrels through exploita- tive competition for resources. With the introduction of Eastern Grey Squirrels, resources such as food and nest sites in areas where both squirrels occur would decrease, causing an increase in home range size of Douglas’s Squirrels because larger ranges are needed to collect more food (Schoener 1971). Larger home ranges would decrease the density of Douglas’s Squir- rels in the given area. Also, with increased home ranges, Douglas’s Squirrels would expend more energy forag- ing and defending the area and would have less ener- gy for reproduction and maintenance. Therefore, we predicted that in areas with Eastern Grey Squirrels, Douglas’s Squirrels would occur at lower densities, have larger home ranges and lower body mass, and that they would reproduce less well. Materials and Methods We worked at two study sites. The species co-occur in Stanley Park, which was the original site of Eastern Grey Squirrel introduction to western Canada, and of Robinson and Cowan’s (1954) study. The control study site was Ecological Reserve #74 (Pacific Spirit Region- al Park) where only Douglas’s Squirrels occur. Al- though Eastern Grey Squirrels do not occur in Eco- logical Reserve #74, they do occur in residential areas nearby and are abundant on the University of British Columbia campus only 1-2 km distant. Both study areas are in the Coastal Western Hem- lock biogeoclimatic zone. The dominant coniferous species in Stanley Park were Western Hemlock (Tsuga HWANG AND LARIVIERE: EFFECTS OF GREY ON DOUGLAS’S SQUIRRELS 1} heterophylla), Western Redcedar (Thuja plicata), and Douglas Fir (Pseudotsuga menziesii). Stanley Park is a mosaic of second- (> 50 years old) and old-growth (> 120 years old) conifer stands. Ecological Reserve #74 represented a mature conifer stand (> 90 years old) with Western Hemlock, Western Redcedar, Dou- glas Fir, and Sitka Spruce (Picea sitchensis) with scat- tered Red Alder (Alnus rubus) and Big Leaf Maple (Acer macrophyllum). Understory and ground cover in both study areas consisted mainly of Vine Maple (Acer circinatum), Salmonberry (Rubus spectabilis), Red Huckleberry (Vaccinium parvifolium), Salal (Gaulthe- ria shallon), leaf litter, ferns and mosses. In each study area, we established a 9-ha live-trap- ping grid consisting of 96 (6 by 16) stations located at 30 m intervals. We set Tomahawk live-traps (model 201, Tomahawk Live Trap Co., Tomahawk, Wisconsin) at alternate trap stations, resulting in 48 traps per live- trapping grid. We live-trapped squirrels for 2 days every 2 weeks from May to August 1997 (9 trap ses- sions) in both study areas. We baited the traps with sunflower seeds and set them in early morning and checked traps 4-6 hours later. Captured squirrels were ear-tagged with num- bered metal tags (Monel #1, National Band and Tag Co., NewPort, Kentucky), sexed and weighed (+ 5 g using a Pesola® spring balance). We noted breeding condition by palpation of male testes and female mam- maries (Sullivan and Moses 1986). We estimated population size, proportion of repro- ductive squirrels, body mass, and home range size. Population size and trappability were estimated for each trap session using the Jolly-Seber mark-recap- ture model (Jolly 1965; Seber 1965). We were not able to use radio-telemetry to estimate the territory size of Douglas’s Squirrels in this study; therefore we esti- mated home range size by using trapping location for animals that were captured = 2 times using the exclu- sive boundary strip method (Hayne 1949, 1950; Stickel 1954). Also, we only considered home ranges of fe- males because in late spring and early summer, males roam extensively while searching for potential mates. We used chi-square test to examine the trappability of squirrel populations between both sites. We used one-tailed t-test to compare densities of Douglas’s Squirrels between sites. Due to unequal variances for Eastern Grey Squirrel densities in Stanley Park, we used Mann-Whitney U test to compare differences in densities between both species in the two study areas. We used two-way Analysis of Variance (ANOVA) to examine the difference in body mass and home ranges of Douglas’s Squirrels between Ecological Reserve #74 and Stanley Park. The Z-test was used to test for significant differences in the proportion of squirrels in breeding condition (Sokal and Rohlf 1995). Since sample sizes of breeding individuals were small, the statistical power of the Z tests is very low in detecting 12 THE CANADIAN FIELD-NATURALIST Vol. 120 Table 1. Summary of demographic parameters for Douglas’s Squirrels (DS) and Eastern Grey Squirrels (EGS) in Ecological Reserve #74 and Stanley Park, Vancouver, British Columbia, Canada from May to August 1997. Values shown are mean#+ SE. Ecological Reserve #74 Stanley Park DS DS EGS Density (individuals/ha) 3.5+04 2.9+0.1 2.5+04 Home range (ha) 0.37 + 0.096 0.44+0.11 — Trappability (7%) Male 74.7 60.3 30.0 Female 80.9 74.0 30.0 Body mass (g) Male 181.8 + 2.69 184.5+3.81 608.6 + 11.91 Female 177.4 +4.62 181.1+4.40 603.7 + 15.59 Proportion of breeding females (7%) 20 5) 77 a significant effect; however, the Z-statistic was used as an index of differences between study areas. All val- ues are reported as mean + SE. Results Density and home range From May to August 1997, we captured 30 male and 24 female Douglas’s Squirrels in Pacific Spirit Park. In Stanley Park, we caught 19 male and 25 female Douglas’s Squirrels. Eastern Grey Squirrels were only found in Stanley Park, and we caught 22 males and 17 females. See Table 1 for a summary of demograph- ic parameters. Trappability of Douglas’s Squirrels in Pacific Spirit Park was similar between males and females, at 74.7% and 80.9%, respectively. Male and female Douglas’s Squirrels in Stanley Park also have relatively high trap- pability values, both ranging from 60.3% to 74.0%. On the other hand, Eastern Grey Squirrels had very low trappability, with males and females both having 30%. Trappability of Eastern Grey Squirrels was sig- nificantly lower in Stanley Park (y? = 14.54, df = 1, P <0.001) than for Douglas’s Squirrels. Densities of Douglas’s Squirrels did not differ Gia osridig—i2ene) — OMI) libetweent = colopical Reserve #74 (3.5 individuals/ha + 0.4) and Stanley Park (2.9 individuals/ha + 0.1). In Stanley Park, density also did not differ (Mann-Whitney T = 56, P = 0.71) between Eastern Grey Squirrel (2.5 individuals/ha + 0.4) and Douglas’s Squirrels (2.9 individuals/ha + 0.1). No effect of sex (F, ,, = 0.18, P = 0.67) or study area tes = 1.24, P = 0.27) on the home range of Douglas’s Squirrels, nor were significant interactions detected (Ban = 0.44, P=0.51). Home ranges aver- aged 0.31 + 0.05 ha (n = 96). Body mass and reproduction No effect of study area (F, 93 = 0.075, P = 0.78) on body mass of Douglas’s Squirrels was found, but there was a gender effect (F, ,,= 5.41, P = 0.022), with males (185.3 + 4.1 g, n = 48) being heavier than females (177.2 + 5.3 g,n = 49) in both parks. For comparison purposes, Eastern Grey Squirrels in Stanley Park averaged 606.4 + 19.1 g with no gen- der difference (t = 0.25, df = 37, P = 0.80). On average, male Eastern Grey Squirrels weighed 608.5 + 11.9 g, and females weighed 603.8 + 15.6 g. From 20 May to 28 August 1997, proportions of fe- male Douglas’s Squirrels in breeding condition differed significantly between sites (Z = 2.56, P < 0.001), with a higher proportion of breeding females in Ecological Reserve #74 compared to that in Stanley Park: 20% (15 breeding out of 77 individuals) and 5% (3/62), respectively. In Stanley Park, the proportion of fe- male Eastern Grey Squirrels in breeding condition (77%: 17/22) was higher than for Douglas’s Squirrels (5%: 3/62). However, the proportion of breeding male Douglas’s Squirrels at the two sites was similar and averaged 98.3% (59/60). Discussion We did not detect strong effects of exploitative com- petition between Eastern Grey Squirrels and Douglas’s Squirrels. There was no evidence of reduced density, large ranges, and lower mass in Douglas’s Squirrels where the two species co-occurred. However, the pro- portion of breeding Douglas’s Squirrels was significant- ly lower in Stanley Park compared to that in Ecologi- cal Reserve #74, suggesting possible negative effects of Eastern Grey Squirrels on the breeding propensity of Douglas’s Squirrels. Densities of Douglas’s Squirrels are limited by food availability (Sullivan and Sullivan 1982; Ransome and Sullivan 1997), and old-growth forests provide higher- quality habitat for Douglas’s Squirrels than younger forests due to greater and more reliable quantities of conifer seed (Buchanan et al. 1990). In our study, the densities of Douglas’s Squirrels were similar at both sites, suggesting that habitat suitability was similar, and further suggesting that reduced breeding propensity was caused by the presence of Eastern Grey Squirrels, and not simply an artifact of difference in food avail- ability among study areas. A possible explanation for weak effects of exploita- tive competition between the two species of squirrels could be due to the fact that Eastern Grey Squirrels often can exploit broadleaved and deciduous wood- 2006 land to increase breeding success by feeding on high- energy food (Skelcher 1997; Gurnell et al. 2001), but since both study areas consist of mixed woods with high conifer content, advantages in breeding success of Eastern Grey Squirrels were perhaps less pronounced (Teangana et al. 2000). Thus, Douglas’s and Eastern Grey Squirrels could coexist for many years although perhaps Douglas’s Squirrel would slowly decline due to lower reproductive outputs (Reynolds 1985; Gur- nell 1996). Our density estimates for Eastern Grey Squirrels (2.5 individuals/ha) also were slightly higher than those reported >45 years earlier (1.7 individuals/ha; Robin- son and Cowan 1954), suggesting that the population of Eastern Grey Squirrels in Stanley Park is relatively stable, even if their range elsewhere has expanded (Gonzales 1999, 2000). If the combined presence of both Douglas’s and Eastern Grey squirrels in Stanley Park accelerates the depletion of the autumn seed crop, this might force a greater dependence on alternative plant foods in the spring and summer, which Eastern Grey Squirrels are better able to use because of their larger body size (Ackerman and Weigl 1970; Steele and Weigl 1993). An investigation into the feeding behavior and space use of both species would provide a better understanding of how Douglas’s and Eastern Grey squirrels coexists in anthropogenic environments such as urban parks. This study highlights the need for more long-term population monitoring of the interac- tion of the two squirrel species in areas where East- ern Grey Squirrels were introduced. Acknowledgments Douglas Ransome and T. P. Sullivan helped design the study and provided valuable insight, statistical help and field equipment. Many thanks to Mike Mackintosh (Wildlife Manager, Vancouver Parks Board) and Greg Paris (Wildlife Manager, Greater Vancouver Regional District) for permission and help with study area design. J. N. M. Smith reviewed an earlier draft of this man- uscript. Literature Cited Ackerman, R., and P. D. Weigl. 1970. Dominance relations of red and grey squirrels. Ecology 51: 332-335. Buchanan, J. B., R. W. Lundquist, and K. B. Aubry. 1990. Winter populations of Douglas’s squirrels in differ- ent-aged Douglas-fir forests. Journal of Wildlife Man- agement 54: 577-581. Flyger, V. F. 1955. Implications of social behavior in grey squirrel management. Transactions of the North American Wildlife and Natural Resources Conference 20: 381-389. Gonzales, E. K. 1999. Eastern gray squirrels in BC: an intro- duction to an introduction. Discovery 28: 22-25. Gonzales, E. K. 2000. Distinguishing between modes of dispersal by introduced western grey squirrels (Sciurus carolinensis). M.Sc. thesis, University of Guelph, Guelph, Ontario. 98 pages. HWANG AND LARIVIERE: EFFECTS OF GREY ON DOUGLAS'S SQUIRRELS 13 Gurnell, J. 1996. Conserving the red squirrel in Thetford Forest Park: the ecology of a pine forest. Edited by P Ratcliffe and J. Calridge. Forestry Commission, Edin burgh. Gurnell, J., L. A. Wauters, D. Preatoni, and G. Tosi. 200! Spacing behaviour, kinship, and population dynamics of grey squirrels in a newly colonized broadleaf woodland in Italy. Canadian Journal of Zoology 79: 1533-1543 Gurnell, J., and H. Pepper. 1993. A critical look at consery ing the British Red Squirrel Sciurus vulgaris. Mammal Review 23: 127-137. Hall, E. R. 1981. The mammals of North America. Second edition. John Wiley & Sons, New York, New York. Hayne, D. W. 1949. Calculation of size of home range. Jour- nal of Mammalogy 30: 1-17. Hayne, D. W. 1950. Apparent home range of Microtus in rela- tion to distance between traps. Journal of Mammalogy 3) 26-39. Jolly, G. M. 1965. Explicit estimates from capture-recapture data with both death and immigration-stochastic model Biometrika 52: 225-247. Koprowski, J. L. 1984. Sciurus carolinensis. Mammalian Species 480: 1-9. Moller, H. 1983. Foods and foraging behaviour of red (Sci- urus vulgaris) and grey squirrels (Sciurus carolinensis). Mammal Review 13: 81-99. Ransome, D. B., and T. P. Sullivan. 1997. Food limitation and habitat preference of Glaucomys sabrinus and Tami- asciurus hudsonicus. Journal of Mammalogy 78: 538-549. Reynolds, J. C. 1985. Details of the geographic replacement of the red squirrel (Sciurus vulgaris) by the grey squirrel (Sciurus carolinensis) in eastern England. Journal of Ani- mal Ecology 54: 149-162. Riege, D. A. 1991. Habitat specialization and social factors in distribution of red and grey squirrels. Journal of Mam- malogy 72: 152-162. Robinson, D. J., and I. McT. Cowan. 1954. An introduced population of the grey squirrel (Sciurus carolinenesis Gmelin) in British Columbia. Canadian Journal of Zool- ogy 32: 261-282. Schoener, T. W. 1971. Theory of feeding strategies. Annual Review of Ecology and Systematics 2: 369-404. Seber, G. A. F. 1965. A note on the multiple-recapture census. Biometrika 52: 249-259. Skelcher, G. 1997. The ecological replacement of red by grey squirrels. Pages 67-78 in The conservation of red squirrels, Sciurus vulgaris L. Edited by J. Gurnell, and P. Lurz. Peo- ple’s Trust for Endangered Species, London. Smith, C. C. 1981. The indivisible niche of Tamiasciurus: An example of nonpartitioning of resources. Ecological Monographs 51: 343-363. Sokal, R. R., and F. J. Rohlf. 1995. Biometry: The principles and practice of statistics in biological research. 3™ edition. W. H. Freeman & Company, New York, New York. Steele, M. A. 1998. Tamiasciurus hudsonicus. Mammalian Species 586: 1-9. Steele, M. A. 1999. Tamiasciurus douglasiti. Mammalian Species 630: 1-8. Steele, M. A., and P. D. Weigl. 1993. The ecological signifi- cance of body size in fox squirrels (Sciurus niger) and gray squirrels (S. carolinensis). Pages 57-69 in Proceedings of the Second Symposium on Southeastern Fox Squirrels. Sciurus niger. Edited by N. D. Moncrief, J. W. Edwards. 14 THE CANADIAN FIELD-NATURALIST and P. A. Tappe. Virginia Museum of Natural History Special Publication Number 1. Stickel, L. F. 1954. A comparison of certain methods of measuring ranges of small mammals. Journal of Mammal- ogy 35: 1-15. Sullivan, T. P., and R. A. Moses. 1986. Red squirrel popu- lations in natural and managed stands of lodgepole pine. Journal of Wildlife Management 50: 595-601. Sullivan, T. P., and D. S. Sullivan. 1982. Population dynamics and regulation of the Douglas’s squirrel (Jamiasciurus douglasii) with supplemental food. Oecologia 53: 264-270. Teangana, D. O., S. Reilly, W. I. Montgomery, and J. Rochford. 2000. Distribution and status of the red squirrel (Sciurus vulgaris) and grey squirrel (Sciurus carolinensis) in Ireland. Mammal Review 30: 45-56. Tompkins, D. M., A. R. White, and M. Boots. 2003. Eco- logical replacement of native red squirrels by invasive greys driven by disease. Ecology Letters 6: 189. Vol. 120 Thompson, D. C. 1977. Social system of grey squirrel. Behaviour 64: 305-328. Wauters, L. A., and J. Gurnell. 1999. The mechanism of replacement of red squirrels by grey squirrels: a test of the interference competition hypothesis. Ethology 105: 1053- 1071. Wauters, L. A., J. Gurnell, A. Martinoli, and G. Tosi. 2001. Does interspecific competition with grey squirrels affect the foraging behavior and food choice of red squir- rels? Animal Behavior 61: 1079-1091. Wauters, L. A., P. W. W. Lurz, and J. Gurnell. 2000. Interspecific effects of grey squirrels (Sciurus carolinensis) on the space use and population demography of red squir- rels (Sciurus vulgaris) in conifer plantation. Ecological Research 15: 271-284. Received 22 July 2004 Accepted 22 September 2005 Responses of Syrphids, Elaterids and Bees to Single-tree Selection Harvesting in Algonquin Provincial Park, Ontario Erica Nov!, HUME DouGLas!*, and WILLIAM J. CRINS? 'Biology Department, Trent University, Peterborough, Ontario K9J 7B8 Canada *Present address: Biology Department, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6 Canada 3Planning and Research Section, Ontario Parks, Ontario Ministry of Natural Resources, 300 Water Street, Peterborough, Ontario K9J 8M5 Canada Nol, Erica, Hume Douglas, and William J. Crins. 2006. The response of syrphids, elaterids and bees to single-tree selection harvesting in Algonquin Provincial Park, Ontario. Canadian Field-Naturalist 120(1): 15-21. The species composition of hoverflies (Syrphidae), click beetles (Elateridae), and bees (Apoidea) was studied to determine whether there was a positive response in these flower-seeking insect groups to gaps in the canopy created through single- tree selection harvesting of Sugar Maple (Acer saccharum) and Yellow Birch (Betula alleghaniensis) in hardwood forests of the Great Lakes-St. Lawrence forest region of Algonquin Provincial Park, Ontario. There were significantly more hoverflies and bees collected in forest stands harvested within the previous five years than in wilderness zone (unharvested at least for 40 years) stands or stands harvested 15-20 years previously (old logged stands). Click beetles, especially Selatosomus pul- cher (LeConte), were collected most often in old logged stands. Bees and click beetles were collected significantly later in the season in logged than in wilderness zone stands. Malaise traps resulted in higher capture rates for syrphids than pan traps, and only with these higher capture rates did we detect a significant increase in species richness in recently logged stands over that in wilderness stands. Changes in the numbers and phenology of flower-visiting insects may impact on repro- ductive success of flowering plants of the forest understory and deserves further study. Key Words: single-tree selection logging, syrphids, hoverflies, Syrphidae, click beetles, Elateridae, bees, Apoidea, Algonquin Provincial Park, hardwood forests. Single-tree selection harvesting’ in shade-tolerant hardwood forests creates cyclic perforations in the canopy on a 20-25 year rotation and is hypothesized to simulate conditions found under gap-phase dynam- ics. This form of silviculture is practised in many parts of the Great Lakes-St. Lawrence Forest Region in Ontario and Quebec (Hunter 1990). Very little is known about the impact on non-target species of removing 25% of the basal area of the forest through selection cutting (Annand and Thompson 1997). Particular invertebrate groups are most likely to respond to changes in light conditions of the forest floor as a result of reductions in canopy cover. Many plants respond positively to increased light. As a result, insects with adult stages attracted to conspicuous flow- ers, also are expected to increase in abundance. These include bees (Hymenoptera: Apoidea), hoverflies (Dip- . tera: Syrphidae), a family of flies that contains impor- tant flower visitors and pollinators (Waldbauer 1983; Kula 1997; Goulson and Wright 1998; Gross 2001), and many click beetles (Coleoptera: Elateridae), a less well-known group of flower visitors (Johnson 2002). Changes in other components of the forest as a result of selection cutting may also benefit these insects. Larvae of hoverflies of the subfamily Syrphinae feed largely on aphids (Vockeroth 1992), and may benefit from increased herbaceous growth associated with canopy opening. Larvae of many elaterids (Johnson 2002) and some hoverflies of the subfamily Eristali- nae (Vockeroth 1992) inhabit decaying wood. Their response to logging may depend on the availability of amounts, sizes and decay states of the standing and fallen trees remaining after each phase of the logging cycle. Soil-inhabiting elaterid larvae and ground-nest- ing bees may be affected by changes in vegetation, litter composition and depth, microclimate, and soil disturbance (Penev 1992; Sugar et al. 1998). Distur- bance caused by construction of sand logging roads may result in greater colonization by psammophilous bees and elaterids in a previously unsuitable forest stand (Cane 1991). We studied the impact of selection cutting on these non-target invertebrates in Algonquin Park, Ontario. We predicted positive density responses for all three groups, either as a result of an increase in spring flora, or other changes associated with logging. We also pre- dicted that the presence of insects that visit flowers would extend later into the growing season in logged forests than in unlogged stands because of the greater amount of light penetrating to the forest floor through- out the spring and summer seasons. We tested these predictions by comparing these insect communities in Under selection harvesting, only some of the trees are cut in order to balance the volume of wood harvested with the amount produced through new growth and to enhance biodiversity and ecological functions. This system of silviculture differs from selective harvesting in which only the best timber trees are cut and the forest is gradually degraded. 15 16 THE CANADIAN FIELD-NATURALIST stands cut 6 months to 3 years before our sampling, 15-20 years before, and in stands that have not been subjected to logging for at least 40 years. Study Area and Methods The study was conducted in the summers of 1997 and 1998 in Algonquin Provincial Park, Ontario (42°32'N, 78°36'W), as part of a larger study that was examining impacts of logging on birds (Jobes et al. 2004) and ground beetles (Carabidae: Vance and Nol 2003). Forest stands were chosen randomly (within the constraint that they be within 500 m of road access) in wilderness designated zones (Algonquin Park Man- agement Plan 2000), last logged at least 40 years prior to survey), old logged (15-20 years post cutting), and recently logged (6 months — 3 years post cutting) areas. Canopy cover (estimated in July) ranged from 60- 70% in the wilderness zone to 25-35% in old cut and recently cut stands, and was significantly higher in the wilderness zone stands than in both the old logged and recently-cut stands (Jobes et al. 2004). Other veg- etation differences included a significant increase in vegetation under 2 m and percent logging residue cover (mostly tops of trees) in the recently cut stands over that in the other two treatments, and a signifi- cant increase in stem densities and saplings (2-5 m) and subcanopy (5-10 m) vegetation in the old logged treatments as compared to the other two (Jobes et al. 2004). In 1997, we used a single Malaise trap set on a ridge in one stand of each of the recently logged and wilder- ness zone treatments. It was emptied at 1-week inter- vals every other week from May to August. Thus, we had no replication of stands for this year. In 1998, we set yellow pan traps (21 cm? x 5 cm depth) filled with water to a depth of 2-3 cm, and a small amount of detergent in a total of 18 stands. These traps were set, within stands, at 100 m intervals at permanent mark- ers. We sampled eight stands in the wilderness zone, seven in the recently logged areas, and three in the old logged areas. Stands were separated by a minimum of 200 m, but most stands were separated by > 3 km, in stands consisting of at least 75% upland deciduous forest (Vance and Nol 2003) so we assumed that they were spatially independent. Each stand contained a minimum of three traps. The sampling was uneven in the treatments because of accessibility of stands and because two of the initial old logged treatments were logged in the winter of 1997-1998. These traps were set from early May to late August, and emptied weekly for three weeks in early, mid, and late summer periods, for a total of nine trapping weeks. There were 69 inci- dents of trap disturbance by mammals. Pitfall traps were used to obtain activity-density esti- mates for click beetles (Baars 1979). Pitfalls were con- structed using | litre plastic 11.5 cm diameter contain- ers lined with 500 mL plastic containers of the same diameter (Spence and Niemela 1994). The traps were dug into the ground so that the top of the upper contain- Vol. 120 er was level with or slightly below the surface of the soil. Each trap was filled with approximately 300 ml of water, with a few drops of dish soap added to reduce surface tension. Two pitfall traps were set at 4 m to each side of the permanent stand markers, at the same intervals as the pan traps. In 1997, pitfalls were set for one week in each of May through August for a total of four weeks. In 1998, pitfalls were set for the same periods as the pan traps. A small number of traps (5%) were disturbed in all treatments All invertebrates were sorted initially to class, and all but insects were discarded. Insects of the three target groups were then sorted and identified with the use of keys by Becker (1956, 1974); Brown (1934); Coovert and Thompson (1977); Curran (1921, 1922, 1925, 1934, 1941); Curran and Fluke (1926); Dietrich (1945); Fluke and Hull (1945); Fluke and Weems (1956); Hull and Fluke (1950); Michener et al. 1994; Mitchell (1960); Shannon (1939); Telford (1970); Vockeroth and Thompson (1981); and Vockeroth (1992). Voucher specimens are deposited in the Canadian National Collection of Insects and Arthropods, Agriculture and Agri-food Canada, Ottawa. As we began sampling in all treatments at the same time, we used the dates (1998 only) on which the tar- get groups were captured as independent samples, for our comparisons of phenology. These data were high- ly skewed so we used the Kruskal-Wallis ANOVA to test for significance among treatments. We were unable to use stands as sampling units for any statistical com- parisons because capture rates were too low. Thus, we tested for significant differences among treatments in species and numbers of individuals trapped, using chi- square tests, with proportion of total trapping effort among treatments used to calculate expected num- bers. We report standardized residuals ((Observed — Expected)/ VExpected) and their direction to show which treatment had greater or less than the numbers of individuals or species than expected based on our sampling effort (Quinn and Keough 2002). For pit- falls and pan traps we usually caught single individu- als of the target species and these were spread across a range of stands within that treatment, so that the as- sumption of independence for this statistical test was probably not violated (1.e., the samples are not biased by many individuals caught in one particular stand and treatment except in cases where noted below). Given the low capture rate for any of our target groups we did not calculate or analyse patterns of diversity. To assess spring ephemeral abundance, we counted, in | m* quadrats placed 5 m to the side of pitfall and pan trap locations, all visible stems of spring-blooming forb species during the first sampling period in May 1998, before canopy leaf emergence. Results Each of the three target groups had substantially higher captures per unit effort (combining sample across all stands per treatment because of low num- 2006 bers captured) in at least one logged treatment than in the wilderness zone treatment (Table 1). Syrphids. In both years, with both pan and malaise trap- ping methods, more species of syrphids were caught in logged than in unlogged landscapes (Table 2). In 1997, eight species were found in wilderness zone malaise traps and 25 species were trapped from the recently logged stands with three species in common between the two treatments. As Malaise traps were opened for an equal number of days in the two stands, we calculated expected numbers using the binomial distribution, based on P = 0.5. Significantly more NoL, DOUGLAS, AND CRINS: RESPONSES OF SYRPHIDS, ELATERIDS AND BEES TABLE |. Catch per unit effort (x100) for all Syrphidae, Elateri dae and Apoidea caught in pan traps in recently logged, old logged, and wilderness stands in 1998, and combined catch for pitfalls in 1997 and 1998. Recently Old Logged Logged Wilderness Total pan trap days 1894 1040 2961 Total pitfall days 4796 3088 7938 Syrphidae (pan) 0.68 2.31 0.17 Elateridae (pan) 1.37 6.82 3.7] Elateridae (pitfall) 0.27 1.49 0.30 Apoidea (pan) 3.22 2.11 1.28 TABLE 2. Species and numbers of individuals captured of Syrphidae from pan traps set in recently logged, old logged, and wilderness stands in Algonquin Provincial Park using Malaise traps in 1997 and pan traps in 1998. ¥° statistic provided for test of association between number of individuals and species and logging treatment. Standardized residuals give direction and strength of deviation (Quinn and Keough 2002) Species Year Sub-Family Syrphinae Baccha elongata (Fabricius) Dasysyrphus pauxillus (Williston) Epistrophe nitidicollis (Meigen) Eupeodes perplexus (Osburn) Eupeodes americanus (Wiedemann) or pomus (Curran) Melanostoma mellinum (Linnaeus) Meliscaeva cinctella (Zetterstedt) Parasyrphus semiinterruptus (Fluke) Parasyrphus sp. Platycheirus confusus (Curran) Platycheirus obscurus (Say) Sphaerophoria novaeangliae (Johnson) Syrphus rectus (Osten Sacken) Toxomerus geminatus (Say) Sub-Family Eristalinae Brachyopa notata (Osten Sacken) Brachypalpus oarus (Walker) Chalcosyrphus nemorum (Fabricius) Cheilosia rita (Curran) Cheilosia tristis (Loew) Eristalis dimidiatus (Wiedemann) Helophilus fasciatus (Walker) Lejota aerea (Loew) Lejota cyanea (Smith) Neoascia distincta (Williston) Pipiza femoralis (Loew) Rhingia nasica (Say) Sericomyia chrysotoxoides (Macquatt) Sphegina brachygaster (Hull) Sphegina flavomaculata (Malloch) Sphegina keeniana (Williston) Sphegina campanulata (Robertson) Temnostoma balyras (Walker) Volucella bombylans (Linnaeus) Xylota quadrimaculata (Loew) Xylota confusa (Shannon) Number of individuals ¥7= 45.9, P < 0.001 Standardized residuals, individuals (1998 data only) Number of species ¥7= 5.5, n.s. Standardized residuals, species (1998 data only) 1997 11 Wilderness 1998 Old logged 1998 Recently logged 1997 1998 i) bho vs) — Om W— ‘= et emt KD ND 042 18 THE CANADIAN FIELD-NATURALIST syrphid individuals and species were captured in the Malaise trap set in the recently logged than in the wilderness habitat (Binomial test, species: P < 0.0001, individuals: P < 0.0001). In 1998, using pan trap catch- es, two species, each consisting of one individual each, were found only in the wilderness zone, whereas nine species were found only in the recently logged land- scape. Only six species were found in both years using the two capture techniques. Significantly more individ- uals were caught in 1998, in the old logged stands than in either the recently logged or wilderness treatments (Table 2), but this result was entirely due to a capture of 17 individuals of the species Melanostoma mellinum on 14 May 1998 in the old logged treatment. In 1997, we did not begin sampling until late May, after leaf-out in the park. In 1998, there was a strong bias in the collection times for all syrphids, with 16 of 22 collection dates for syrphids (73%) in May, prior to leaf-out. There was, however, no significant differ- ence among treatments in the collection dates of syr- phids in 1998 (median sample dates (range): wilder- ness: 29 May (26 May-6 August), n = 5 trap sample dates; old logged: 21 May (14 May-2 July), n = 4; recently logged: 27 May (9 May-24 June), n = 13, Kruskal-Wallis 77 = 4.33, P =0.11). Bees. Twelve species of bees were identified, with five of these from wilderness and old logged stands and all twelve in the recently logged stands (Table 3). All but four species (excluding Andrena) were halictids. For both number of species and number of individu- als the recently logged stands had significantly more bees than expected based on catch effort (Table 3). All but 6 of the 57 dates on which we captured bees were in May. Bees were caught at a significantly later date in recently logged stands than in either old logged or wilderness stands (median collection dates (range): wilderness: 10 May (8-20 May), n = 18 trap sample dates; old logged: 9 May (9 May-24 June), n = 13; recently logged: 19 May (9 May-14 July), n = 26, Kruskal-Wallis 77 = 13.8, P < 0.001). Elateridae. In this group of insects, two species, Cteni- cera triundulata and C. vulnerata were found only in the wilderness stands, both in very low numbers (Table 4). The number of species among treatments did not deviate from expected using the pan trap data but was marginally higher than expected in the old logged treatment using the combined 1997/1998 pitfall data (combined years due to small sample sizes, y7 = 5.89, P <0.06). Numbers of individual click beetles were signifi- cantly higher in the old logged habitat, and significant- ly lower in the wilderness zone than expected based on catch effort, using pan traps and pitfall traps (Table 4). Elaterids were collected in 1998 from pan traps at significantly later dates in old logged and recently logged stands than in wilderness zone stands (median date (range); wilderness zone: 13 May (8 May-5 June), Vol. 120 ‘n = 9; old logged: 28 May (8 May-9 July), n = 23: recently logged: 19 May (9 May-2 July), n = 23; Krus- kal-Wallis y? = 7.6, P= 0.02). Using 1998 pitfall data, elaterids were sampled significantly later in recently logged stands than in the other two treatments (wilder- ness zone: 17 May (13 May-4 August), n = 20; old logged: 17 May (14 May-3 July, n = 22; recently logged: 24 June (12 May-24 June), n = 9; Kruskal- Wallis 7? = 6.34, P = 0.04). The 1997 elaterid data were too sparse for analysis. Spring ephemerals. We recorded 15 species of spring ephemerals in our quadrats (1998 only). The num- bers of ephemerals found in all of the stands were extremely variable (cv’s range from 156-165%). Although the mean number of stems in the recently logged stands was more than twice as large as this number in wilderness zones, there was no significant difference among treatments (recently logged: 59.7 + 16.7 / m’, n = 23 plots; old logged: 46.5 + 22.1 / m?, n= 13 plots; wilderness zone: 26.5 + 18.8 /m*,n=18 plots; IB sn = (0.88, n.s.). Discussion Our results suggest that all three of groups of flower- visiting insects were more abundant in one of the logged habitat types than in the wilderness zone. This suggests that these insect groups were positively affected by single-tree selection harvesting as it is practised recent- ly and/or how it was practised in the 1970s, when the selection cuts were heavier than is currently accept- able (Algonquin Forestry Authority, unpublished). This result is not explained simply by greater numbers of spring ephemerals, as, at least with our sampling methodology, we found no significant difference among treatments in the availability of flowers measured prior to leaf-out. Differences were apparent in number of individual insects, but, in general, not in insect species richness, except in the case of the Malaise traps set in 1997, where both numbers of individuals and species richness were higher in collections from the trap in the recently logged stand. Given that this result is for one trap only in each treatment, this result should be viewed cautiously. As many larval syrphids prey on Homoptera in herbaceous vegetation and many adults feed on nectar sources in this vegetation, the presence of much high- er percent cover of herbaceous or soft woody vegeta- tion (37 to 43% in old logged and wilderness stands respectively, to 78% in recently logged stands, partic- ularly Rubus idaeus L. var. strigosus (Michx), Jobes 1999) could explain the significant increase in syr- phid abundance that we found through Malaise trap- ping. An apparent increase also was noted in our pan trapping results but not established statistically because of the generally poor success of this method for cap- turing syrphids in any of our stands. A similar positive numerical response in syrphids (also not tested statis- tically) was found in Poland’s polluted spruce forests, 2006 NoL, DOUGLAS, AND CRINS: RESPONSES OF SYRPHIDS. ELATERIDS AND BEES 19 TABLE 3. Species and Genera of bees collected from pan traps and numbers of individuals and minimum number of specie from Algonquin Park, summer 1998, 122 individuals captured. Statistics and standardized residuals as in Table 2 Genus Wilderness Old logged Recent logged Andrena' Bombus perplexus Cresson Bombus vagans Smith Hylaeus elliptica (Kirby) Lasioglossum admirandus (Sandhouse) Lasioglossum divergens (Lovell) Lasioglossum laevissimus (Smith) Lasioglossum nymphaearum (Robertson) Lasioglossum quebecensis (Crawford) Lasioglossum rohweri (Ellis) Lasioglossum versans (Lovell) Lasioglossum zephyrus (Smith) Osmia atriventris Cresson Number of individuals x? = 21.2, P < 0.001 Standardized residuals, individuals Minimum number of species y= 20.4, P < 0.001 Standardized residuals, species 36 Wr o 20 | 21 — ~ o Nw — — WwW Ww to ' Not identified to species because of poor quality of specimens. TABLE 4. Species and number of individuals of Elateridae collected from wilderness, old logged stands, and recently logged stands in Algonquin Park in 1997 and 1998 using pan traps (individuals: y7= 175.1, P < 0.00; species: y? = 3.2. ns.) and pitfall traps (individuals: y= 90.7, P < 0.001, species: y?= 5.9, n.s.). Analysis for pitfalls from two years combined). Standardized residuals as in Table 2. Wilderness Old logged Recent logged Species 98 Pan 97 Pit 98 Pit 98 Pan 97Pit 98 Pit 98Pan 97Pit 98 Pit Agriotes collaris (LeConte) 2 | 5 6 l 3 Agriotes stabilis (LeConte) l Athous acanthus (Say) | Ctenicera hieroglyphica (Say) ((Say)) 1 2 Z, l Ctenicera insidiosa (LeConte) l Ctenicera resplendens (Eschscholtz) ] l Ctenicera spinosa (LeConte) | l Ctenicera triundulata (Randall) 2 Ctenicera vulnerata (LeConte) 1 Dalopius cognatus Brown l l Dalopius fuscipes Brown l | l Dalopius spp. (females) 2, 2 | 3 2 Limonius aeger LeConte 2 Limonius confusus LeConte > l l - l 2 9 a Melanotus castanipes (Paykull) (Paykull) l Neohypdonus tumescens (LeConte) | l 2 Selatosomus pulcher (LeConte) 2 10 59 2 28 10 Number of individuals 11 5 20 71 6 42 27 2 9 Standardized residuals, individuals -3.4 -2.6 11.8 8.6 -1.3 -3.] Number of species 5 3 9 5 5 8 7 2 - Standardized residuals, species =|? -0.8 1.1 22. 0.7 0.6 as an apparent result of drastic reductions in canopy cover and greater herbaceous and grass undergrowth (Kula 1997). Some species of at least two genera of hoverflies that were found in the wilderness zone (Melanostoma and Platycheirus) specialize on grami- noid and other anemophilous pollen, enabling them to be active later into the spring/summer when grami- noids typically flower (Vockeroth 1992; Gilbert 1993). However, none of the species captured is restricted to flying in the spring. Bees, like syrphids, were most abundant in the re- cently logged areas. Unlike syrphids, larval bees are fed pollen and thus are more wholly dependent on flo- ral resources. Some bees also may be affected by the 20 THE CANADIAN FIELD-NATURALIST availability of open flight paths present in the open understory of the recently logged treatments, to make pollen-gathering trips for brood provisioning, although the effects of forest habitat on bee movement is prob- ably species-specific, and for many species these move- ment patterns are unknown (Kreyer et al. 2004). Wood- land and edge ground nesting bees (the majority of the species caught in the recently logged stands) are thought to benefit from patches of disturbed habitat associated with habitat fragmentation (Cane 1991) and thus, may benefit from the building of sand logging roads. Halictid bees can be major components of pol- linator communities in other forest types (Pascarella 1997). Their role in northern hardwood forests has not been adequately assessed. Elaterids were most abundant in the old logged habi- tats, using two methods of sampling, indicating posi- tive responses to the much higher cover of the sapling (2-10 m) layers found in this treatment (Jobes et al. 2004). The most abundant species, Selatosomus pul- cher, has been trapped in open field habitats (Boiteau et al. 2000). This species is morphologically nearly indistinguishable from the European species C. cru- ciatus, a species whose larvae develop in the sandy and moist soils in forest and forest edge (Leseigneur 1972). The higher abundance of elaterids in the old logged treatment areas may be due largely to this species’ positive response to the edge-like conditions present in this treatment. The lower numbers of ela- terids in the recently logged stands may indicate the absence of colonization because of the short time frame. Analagous results are suggested in a study of herbaceous plants in recently logged White Pine (Pinus strobus) stands in Algonquin. Plants in those stands were most similar to plants in unlogged stands pre- sumably because there had been insufficient time for changes in community assemblages after the logging (Kingsley 1998). As little is known about the role of adult food avail- ability in the distribution of elaterids, the near signifi- cant positive effect on species richness may also be due to the greater availability of soil invertebrates or plant material suitable for larval food produced by a four-fold increase in the number of stumps in old logged versus either wilderness or recently logged stands (Jobes 1999). Of the two elaterids caught only in the wilderness treatment, only one may be an old growth dependant species. Ctenicera vulnerata (1 specimen caught) has rarely been collected, and has not been reported pre- viously from Ontario. The other, Crenicera triundulata (2 specimens) cannot be considered old growth-restrict- ed, as it has been collected in moderate numbers in an apparently healthy spruce (Picea sp.) in an old field by HD. We found very few bees, syrphids or click beetles in our samples from the wilderness zone stands, although we expected to find some, at least early in the season when the canopy had not yet leafed out or of species Vol. 120 that utilized downed woody debris (e.g., Chalcosyr- phus, Xylota, Brachyopa). The reason for their paucity in the wilderness zone is unclear but deserves further study. The numerical increase and changes in phenology of flower-associated insect species suggests the potential for greater pollinator efficiency in logged forests, and hence greater reproductive success, through greater seed set of flowers. A more detailed examination of species-specific patterns of habitat use by pollinators and their functional role in hardwood forests is worthy of future research (e.g., Sheffield et al. 2003; Kreyer et al. 2004). Acknowledgments We thank, in particular, Christine Vance, who tire- lessly emptied pan traps in 1998 for this project. We also thank Andrew Jobes for collecting some of the data on spring ephemeral abundance and providing the canopy cover information. We thank E. C. Becker for verifying elaterid identifications, and R. Tuckerman, J. Grixti and T. Romankova for identification of bees. D. Voigt and E. Addison provided the stimulus for this project through their project on disturbance of ecosys- tems in Algonquin Park. We thank them for their exten- sive logistic support. The Algonquin Forestry Authority (AFA) also provided details on the history of the stands. N. Quinn (Algonquin Park) kindly provided funding for our work on birds and beetles in the park. This proj- ect was also funded by NSERC (Canada) grants to E. Nol. Literature Cited Annand, E. M., and F. R. Thompson III. 1997. Forest bird response to regeneration practices in central hardwood forest. Journal of Wildlife Management 61: 159-171. Baars, M. 1979. Catches in pitfall densities in relation to mean densities of carabid beetles. Oecologia 41: 25-46. Becker, E. C. 1956. Revision of the nearctic species of Agri- otes (Coleoptera: Elateridae). Canadian Entomologist 88, Supplement |. 101 pages. Becker, E. C. 1974. Revision of the nearctic species of Athous (Coleoptera: Elateridae) east of the Rocky Mountains. Canadian Entomologist 106: 711-758. Boiteau, G., W. P. L. Osborn, X. Xiong, and Y. Bousquet. 2000. The stability of vertical distribution profiles of insects in air layers near the ground. Canadian Journal of Zoology 78: 2167-2173. Brown, W. J. 1934. The American species of Dalopius Esch. (Elateridae, Coleop.). Canadian Entomologist 66: 30-39, 66-72, 87-96, 102-110. Cane, J. H. 1991. Soils of ground-nesting bees (Hymenoptera: Apoidea): texture, moisture, cell depth and climate. Jour- nal of the Kansas Entomological Society 64: 406-413. Coovert, G. A., and F. C. Thompson. 1977. The Sphegina species of eastern North America (Diptera: Syrphidae). Proceedings of the Biological Society of Washington 90: 536-552. Curran, C. H. 1921. Revision of the Pipiza group of the fam- ily Syrphidae (flower-flies) from north of Mexico. Proceed- ings of the California Academy of Sciences, Fourth Series 11: 345-393. 2006 Curran, C. H. 1922. The syrphid genera Hammerschmidtia and Brachyopa in Canada. Annals of the Entomological Society of America 15; 239-255. Curran, C. H. 1925. Revision of the genus Neoascia Willis- ton (Diptera: Syrphidae). Proceedings of the Entomolog- ical Society of Washington 27: 51-62. Curran, C. H. 1934. Notes on the Syrphidae in the Slosson collection of Diptera. American Museum Novitates (724). Curran, C. H. 1941. New American Syrphidae. Bulletin of the American Museum of Natural History 78: 243-304. Curran, C. H., and C. L. Fluke. 1926. Revision of the nearc- tic species of Helophilus and allied genera. Transactions of the Wisconsin Academy of Sciences, Arts and Letters 22: 207-281. Dietrich, H. 1945. The Elateridae of New York State. Cornell University Agricultural Experimental Station Memoirs 269. 79 pages. Fluke, C. L., and F. M. Hull. 1945. The Cartosyrphus flies of North America (Syrphidae). Transactions of the Wiscon- sin Academy of Sciences, Arts and Letters 37: 221-263. Fluke, C. L., and H. V. Weems, Jr. 1956. The Myoleptini of the Americas (Diptera, Syrphidae). American Museum Novitates (1758). Gilbert, F. S. 1993. Hoverflies. Naturalists’ Handbooks S. Richmond Publishing Co. Ltd., Slough, England Goulson, D., and N. P. Wright. 1998. Flower constancy in the hoverflies Episyrphus balteatus (Degeer) and Syrphus ribesii (L.) (Syrphidae). Behavioral Ecology 9: 213-219. Gross, C. L. 2001. The effect of introduced honeybees on native bee visitation and fruit-set in Dillwynia juniperina (Fabaceae) in a fragmented ecosystem. Biological Con- servation 102: 89-95. Hull, F. M., and C. L. Fluke. 1950. The genus Cheilosia Meigen (Diptera, Syrphidae): The subgenera Cheilosia and Hiatomyia. Bulletin of the American Museum of Natural History 94: 303-401. Hunter, M. L., Jr. 1990. Wildlife, forests, and forestry: principles of managing forests for biological diversity. Prentice-Hall, Englewood Cliffs, New Jersey. Jobes, A. 1999. Effects of selection cutting on habitat struc- ture and bird communities in the tolerant hardwood forests of Algonquiin Provincial Park, Ontario. B.Sc. thesis. Trent University, Peterborough, Ontario. 60 pages. Jobes, A. P., E. Nol, and D. R. Voigt. 2004. Effects of selec- tion cutting on bird communities in contiguous eastern hardwood forests. Journal of Wildlife Management 68: 51-60. Johnson P. J. 2002. Elateridae. Pages 160-173 in American Beetles. Volume 2. Edited by R. H. Arnett, M. C. Thomas, P. E. Skelley, and J. H. Frank. CRC Press LLC. Boca Raton, Florida. Kingsley, A. L. 1998. Response of birds and vegetation to the first cut of the uniform shelterwood silvicultural sys- tem in the white pine forests of Algonquin Provincial Park, Ontario. Unpublished M.Sc. thesis, Trent University, Peter- borough, Ontario. 110 pages. Kreyer, D., A. Oed, K. Walther-Hellwig, and R. Frankl. 2004. Are forests potential landscape barriers for forag- ing bumblebees? Landscape scale experiments with Bom- NOL, DOUGLAS, AND CRINS: RESPONSES OF SYRPHIDS, ELATERIDS AND BEES bus terrestris agg. and Bombus pascuorum (Hymen« yptera, Apidae). Biological Conservation 116: 111-119 Kula, E. 1997. Hoverflies (Dipt.: Syrphidae) of spruce forest in different health condition. Entomophaga 42: 133-138 Leseigneur, L. 1972. Coléoptéres Elateridae de la Faune de France continentale et de Corse. Bulletin Mensuel de la So ciété Linnéenne de Lyon. Supplément. 379 pages Michener, C. D., R. M. McGinley, and B. N. Danforth. 1994. The bee genera of North and Central America (Hy menoptera: Apoidea). Smithsonian Institution, Washington 209 pages. Mitchell, T. B. 1960. Bees of the Eastern United States. Vol- ume |. North Carolina Agricultural Experiment Station, Raleigh, North Carolina. Pascarella, J. B. 1997. Pollination ecology of Ardisia escal- lonioides (Myrsinaceae). Castanea 62: 1-7. Peney, L. D. 1992. Qualitative and quantitative spatial varia- tion in soil-wire-worm assemblages in relation to climatic and habitat factors. Oikos 63: 181-192. Quinn, G. P., and M. J. Keough. 2002. Experimental design and data analysis for biologists. Cambridge University Press, Cambridge. Shannon, R. C. 1939. Temnostoma bombylans and related species (Syrphidae, Diptera). Proceedings of the Ento- mological Society of Washington 41: 215-224. Sheffield, C. S., P. G. Kevan, R. F. Smith, S. M. Rigby, and R. E. L. Rogers. 2003. Bee species of Nova Scotia, Canada, with new records and notes on bionomics and floral relations (Hymenoptera: Apoidea). Journal of the Kansas Entomological Society 76: 357-384. Spence J. R., and J. K. Niemela. 1994. Sampling carabid assemblages with pitfall traps: the madness and the meth- od. Canadian Entomologist 126: 881-94. Sugar, A., A. Finnamore, H. Goulet, J. Cumming, J. T. Kerr, and L. Packer. 1998. A preliminary survey of Sym- phytan and Aculeate Hymenoptera from oak savannahs in Southern Ontario. Proceedings of the Entomological Society of Ontario 129: 9-18. Telford, H. S. 1970. Eristalis (Diptera: Syrphidae) from America north of Mexico. Annals of the Entomological Society of America 63: 1201-1210. Vance, C., and E. Nol. 2003. Temporal effects of selection logging on ground beetle communities in northern hard- wood forests of eastern Canada. Ecoscience 10: 49-56. Vockeroth, J. R. 1992. The flower flies of the subfamily Syrphinae of Canada, Alaska, and Greenland. The Insects and Arachnids of Canada, Part 18. Research Branch, Agri- culture Canada, Publication 1867. Vockeroth, J. R., and F. C. Thompson. 1981. Syrphidae. Pages 713-743 in Manual of Nearctic Diptera, Volume 2. Edited by J. F. McAlpine, B. V. Peterson, G. E. Shewell. H. J. Teskey, J. R. Vockeroth, and D. M. Wood. Agriculture Canada, Research Branch Monograph (28). Waldbauer, G. P. 1983. Flower associations of mimetic Syr- phidae (Diptera) in northern Michigan. Great Lakes Ento- mologist 16: 79-85. Received 14 July 2004 Accepted 7 February 2006 New Nesting Records of the Le Conte’s Sparrow, Ammospiza leconteii, from Northeastern Ontario, with Some Notes on Nesting Behaviour MICHAEL PATRIKEEV Ontario Parks, 199 Larch Street, Suite 404, Sudbury, Ontario P3E 5P9 Canada Present address: 3 Helen Street, Dundas, Ontario L9H 1N2 Canada; e-mail: mpatrikeev@hotmail.com Patrikeev, Michael. 2006. New nesting records of the Le Conte’s Sparrow, Ammospiza leconteii, from northeastern Ontario, with some notes on nesting behaviour. Canadian Field-Naturalist 120(1): 22-26. Le Conte’s Sparrow is sparsely distributed through northeastern Ontario with no confirmed records from Algoma District and eastern Lake Superior. Two nests were found in open poor fen between Wawa and Hawk Junction in central Algoma District in 1999. Notes on behaviour, nests, nestling development and feeding effort were taken during 9 hours and 38 minutes at one nest over three days. Key Words: Le Conte’s Sparrow, Ammospiza leconteii, nesting, northeastern Ontario. Le Conte’s Sparrow (Ammospiza leconteii) is dis- tributed through grasslands and wetlands of central and southern Canada east to Ontario and Quebec, and in the north-central United States. “It is one of the least known North American sparrows and any new infor- mation on this species may be of interest” (Lowther 1996). A cryptic species, it nests in thick clumps of dead grass, and its nests are difficult to find. Only about 50 nests had been recorded range-wide by the early 1960s (Walkinshaw 1968) and few have been docu- mented since then (Lowther 1996). Only nine nests of this species had been found in Ontario prior to 1999 (Peck and James 1987; Sandilands and Campbell 1987; G. Peck, personal communication). In Ontario, Le Conte’s Sparrow has been reported in widely separated areas in the forest zone with the greatest concentrations in the James Bay Lowland and northwestern Ontario (Sandilands and Campbell 1987). During the first Ontario Breeding Bird Atlas (1981-1985) this species was found in only 20 of 137 UTM blocks (100 x 100 km) and nesting was confirmed in only four blocks (Sandilands and Campbell 1987). None of the first Atlas records were from Algoma District, although Le Conte’s Sparrow was considered a rare spring species in the vicinity of Wawa (Ontario Ministry of Natural Re- sources 1990). In 2001-2004, surveyors for the second Ontario Breeding Bird Atlas reported probable nesting in two blocks (Sault Ste. Marie and St. Joseph Island), and possible nesting in five additional blocks in the vicinity of Sault Ste. Marie and Thessalon at the south end of Algoma District (Ontario Breeding Bird Atlas website: http://www.birdsontario.org/atlas). Here I describe two new nests from central Algoma District, northeastern Ontario. A small group with 2-3 singing males was found in an upland wetland (5.8 ha, 48°02'42.24N", 084°34'01.30W") in Esquega Township east of Wawa and southwest of Hawk Junction (Fig- ure 1). The wetland (329 m above sea level) encom- passed elements of several wetland types gradually changing from sphagnum bog with few stunted Black Spruce (Picea mariana) to open poor fen and then to a meadow marsh (Harris et al. 1996). Le Conte’s Sparrows inhabited approximately 2 ha of open poor fen in this wetland (see Figure 2). The wetland was bordered by alder thickets (Speckled Alder, Alnus incana) with a few dead Balsam Poplars (Populus balsamifera) and by mixed forest dominat- ed by Trembling Aspen (P. tremuloides) and White Birch (Betula papyrifera). Water regime of the wet- land changed from late May through July 1999. The lower parts (mostly meadow marsh section) were flood- ed in late May (up to 0.5 m deep), but water gradually — receded by late June. The area remained damp, but no standing water remained (natural flow in this wetland was likely disrupted years ago when it was transsected by Highway 101). The area occupied by Le Conte’s Sparrows had a ground cover of sphagnum moss and was dominated by two species of sedge: Few-seeded Sedge (Carex oligosperma) and Aw]-fruited Sedge (C. stipata); Canada Bluejoint (Calamagrostis canaden- sis) was also present. Singing males were heard at the wetland on 21 May 1999, and a display flight observed on 27 May. A flight song is described by Murray (1969). I also observed a “spy flight’: a sparrow would rise from the grass and fly in a low arch quickly dropping back into the cover. Le Conte’s Sparrow likely uses this flight to deter- mine precise position of an approaching intruder. No song or audible call was uttered during such flight. Two nests were found in the wetland, on 13 June and 17 July 1999, respectively. The first nest contained five eggs (Figure 3), but was empty on 17 June. The eggs matched a description of Le Conte’s Sparrow eggs from Walkinshaw (1968). The second nest contained five newly-hatched young on 17 July and still contained _ five young on 23 July when the young would be seven days old. On day seven, the young were in cinnamon-_| coloured plumage with dark streaks on the back, dark | De 2006 _Manitoba omens OS eg 44 at PATRIKEEV: NESTING OF LE CONTE’S SPARROW IN NORTHEASTERN ONTARIO 23 Hudson Bay ( co a —S % a oe Quebec Poff Xe | Algoma District 7 | FIGURE |. Position of Wawa and Algoma District in Ontario. grey crowns and cheeks, tan median crown-stripe and black shafts. The median crown-stripe was seen in at least some chicks (Figure 4). Bill colour changed from fleshy on days two-three to grey-pink on day seven. Gape was bright flesh red on day seven. According to Gollop et al. (1966), the young leave the nest at the age of seven days. The two nests were approximately 150 m apart and might have belonged to the same pair. Both nests were built of dried stems, and lined with fine dried stems of sedge and grass. Outer rim of both nests was loosely built. The first nest was well hidden under and inside a clump of dried graminoids and could not be seen from above. The second nest was attached to dried and liv- ing sedge stems and a small Leatherleaf (Chamaeda- phne calyculata) under a loose cover of sedges and grasses. Both nests were 5-6 cm above the wet ground. Observations totaling 9 hours and 38 minutes were made at the second nest from a blind (through a tele- photo lens) during late afternoon-early evening hours on 19 July (1610-2045 hours), 21 July (1800-2110) and 22 July (1812-2005). All observations were conducted during calm fair weather. The pair at the nest under observation consisted of a slightly larger brighter male, and a scruffier duller female. Le Conte’s Sparrows nev- er flew directly to and from the nest, but landed some distance away and walked (hopped) to it, fed the young and walked away before taking on the wing. Also both birds were never present at the nest at the same time during the observation period. A total of 80 feeding visits was recorded. The birds were bringing food every 6.6 minutes on average (SD + 5.2, Standard error 0.6). Lowther (1996) reported | feeding/minute at a nest con- taining a cowbird. Gender of feeding birds was as- sumed on 55 occasions. The male fed the young on average every 14.1 minutes (SD + 13.5) and the female every 13.5 minutes (SD + 13.5). No differences be- tween sexes were detected in feeding efforts (P = 0.50). Intervals between feedings became shorter as the young were growing: every 7.9 minutes + 5.9 on 19 July, 5.7 + 4.6 on 21 July and 4.9 + 3.0 on 22 July (Kruskal- Wallis one way ANOVA: P = 0.07). 24 THE CANADIAN FIELD-NATURALIST Vol. 120 FIGURE 2. Habitat of Le Conte’s Sparrow in Esquega Township: this wetland encompassed elements of several wetland types gradually changing from a sphagnum bog to open poor fen and then to a meadow marsh. Le Conte’s Sparrows inhabited about 2 ha of open poor fen. Photo by Michael Patrikeev. FiGurE 3. Nest of Le Conte’s Sparrow with five eggs. The nest was built of dead stems, and lined with fine stems of sedge and grass. It was well hidden under and inside a clump of dead graminoids and could not be seen from above. Photo by Michael Patrikeev. 13 June 1999. 2006 PATRIKEEV: NESTING OF LE CONTE’S SPARROW IN NORTHEASTERN ONTARIO Nm Ww Ficure 4. Nestlings of Le Conte’s Sparrow seven days old. Photo by Michael Patrikeey. 23 July 1999. Lowther (1996) noted that hardly any information was available on this species’ diet. Although in many cases food brought to the young was too small to iden- tify through a telephoto lens, I observed small green caterpillars, a white moth, a spider, a small sphinx cater- pillar and two grasshopper abdomens. The last three items were brought by the female who tried to feed them to the small young. The content of the sphinx caterpillar was eventually squeezed into the young’s mouths, and one grasshopper abdomen was swallowed by the female itself after futile attempts to feed it to the young. I did not observe wetlands similar to the one des- cribed above in the Wawa area (Michipicoten and Es- quega townships, and the adjacent parts of Lake Super- ior Provincial Park) in 1999-2000 and no Le Conte’s Sparrows were detected in other visited wetlands. How- ever, this species likely occurs in low densities through- out Algoma District where suitable habitat is available. Acknowledgments I am grateful to Carol Dersch (Lake Superior Provincial Park) and Kim Taylor (North-Eastern Sci- ence and Technology, Ontario Ministry of Natural Re- sources) for help with plant identification, to George Peck (Royal Ontario Museum) who provided me with information on the number of confirmed nesting rec- ords of this species in Ontario, to Richard Knapton (University of Alberta, Edmonton), John C. Eitniear (San Antonio, Texas), A. J. Erskine and an anonymous reviewer for useful comments on this manuscript. Literature Cited Gollop, J. B., J. A. Slimmon, and R. V. Folker. 1966. Some 1965 bird records for the Saskatoon district. Blue Jay 24: 76-78. Harris, A. G., S. C. McMurray, P. W. C. Uhlig, J. K. Jeglum, R. F. Foster, and G. D. Racey. 1996. Field guide to the wetland ecosystem classification for northwestern Ontario. Ontario Ministry of Natural Resources, Northwest Science and Technology. Thunder Bay, Ontario. Ontario Field Guide FG-01. 74 pages + Appendices. Lowther, P. E. 1996. Le Conte’s Sparrow (Ammodramus leconteii). In The Birds of North America (224). Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadelphia, Pennsylvania, and The American Ornithol- ogists’ Union, Washington, D.C. Murray, B. G., Jr. 1969. A comparative study of Le Conte's and Sharp-tailed sparrows. Auk 86: 199-231. Ontario Ministry of Natural Resources. 1990. Checklist of Birds/Liste des oiseaux. Lake Superior Provincial Park and adjacent areas/Pare provincial du lac Supérieur et les régions environnantes. Ministry of Natural Resources/Min- istére des Richesses naturelles. MNR Number 3752. ISBN 0-7729-6809-8. 24 pages. Peck, G. K., and R. D. James. 1987. Breeding Birds of On- tario: Nidiology and Distribution. Volume 2: Passerines. Life Sciences Miscellaneous Publication, Royal Ontario Museum, Toronto, Ontario. xi + 387 pages. Sandilands, A., and C. Campbell. 1987. Le Conte’s Spar- row/Bruant de Le Conte Ammodramus leconteii. Pages 26 THE CANADIAN FIELD-NATURALIST Vol. 120 454-455 in Atlas of the Breeding Birds of Ontario. Com- piled by M. D. Cadman, P. F. J. Eagles, and F. M. Hellein- er. Federation of Ontario Naturalists and Long Point Bird Observatory. University of Waterloo Press. Walkinshaw, L. H. 1968. Passerherbulus caudacutus (Lath- am). Le Conte’s Sparrow. Pages 765— 776 in Life histories of North American cardinals, grosbeaks, buntings, towhees, finches, sparrows, and allies. Edited by O. L. Austin, Jr. United States National Museum Bulletin 237, Part 2. Received 21 October 2004 Accepted 14 September 2005 Conditions for Sexual Interactions RICHARD P. THIEL Canadian Field-Naturalist 120(1): 27—30. of the two species. Wolves and Coyotes can hybridize in captivity (Young and Goldman 1944; Silver and Silver 1969; Kolenosky 1971; Schmitz and Kolenosky 1985a, 1985b). Previously it was assumed that behavioral bar- riers prevented hybridization between Canis species (Wolf x Coyote) in the wild. Evidence of such crosses in the wild remained elusive until Lehman et al. (1991) reported on the presence of Coyote genes in Wolf mito- chondrial DNA material from the Upper Great Lakes region of North America probably from crosses be- tween female Coyotes and male Wolves. This stimulat- ed further investigations (Roy et al. 1994; Pilgrim et al. 1998; Boyd et al. 2001; Wayne et al. 1995; among oth- ers), confirming the introgression of Coyote genes in _ Wolf mtDNA in a region extending from the western _ Great Lakes east to Quebec. _ Confounding these findings, recent DNA studies _ among Canis in eastern Ontario and northeastern Unit- ed States hypothesize the existence of a unique and putative Eastern (Timber) Wolf (Canis lycaon (Wilson et al. 2000, 2003) while other researchers argue the possibility that the Eastern Wolf is the Red Wolf (Canis rufus), or some type of a Wolf (Canis lupus) x Coyote hybrid (Schmitz and Kolenosky 1985a; Wayne et al. 1995; Nowak 2003; Wayne and Vila 2003; Phillips et al. 2003; Sears et al. 2003; among others). It is speculated that the tumultuous ecological up- , heavals caused by Eurasian settlement of North Ameri- ca disrupted Canis communities creating conditions | facilitating hybridization (Wayne et al. 1995; among others). Forest fragmentation and heavy exploitation of i | i | Canis populations are believed responsible for current _ hybridization between various Canis communities in southeastern Ontario (Sears et al. 2003; Theberge and | Theberge 2004). _ The question remains under what environmental — . behavioral circumstances did (do) unions between Coyotes and Wolves occur, given the usual aggressive | 1 Between Wild Grey Wolves, Canis lupus, and Coyotes, Canis latrans Wisconsin Department of Natural Resources, Sandhill Wildlife Area, Box 156, Babcock, Wisconsin 54413 USA Thiel, Richard P. 2006. Conditions for sexual interactions between wild Grey Wolves, Canis lupus, and Coyotes, Canis latrans. | Genetic evidence for the hybridization of wild Grey Wolves and Coyotes was first reported by Lehmann et al (1991). Subse- quent genetic and landscape-environmental analyses have attempted to grasp the extent of Wolf-Coyote crosses in North America. Since Wolves are normally territorial and thus aggressive towards Coyotes, hybridization events remain rare, not withstanding the taxonomic debates regarding Canis in eastern Ontario. In this paper I report on amicable interactions between Wolves and Coyotes observed in Wisconsin in recent decades and discuss circumstances that may lead to pairing between individuals Key Words: Grey Wolf, Canis lupus, Coyote, Canis latrans, interactions, hybridization, Wisconsin, Upper Great Lakes region. tendencies of Wolves to Coyotes? Lehman et al. (1991) believed that initial hybridization most likely occurred when young, male Wolves dispersed into an area with an endemic Coyote population where female Wolves were rare. I report encounters between Coyotes and Wolves in Wisconsin that support this theory during a period when Wolves were re-colonizing landscapes occupied by Coyotes. Because of the ongoing debate over Canis affinities in the east, I confine my comments to the Upper Great Lakes region of North America. Methods It is generally accepted that two taxonomically dis- tinct wild canid species occur in Wisconsin: Grey Wolves (Canis lupus) and Coyotes (Canis latrans). Natural re-colonization of Wolves in Wisconsin began in the mid 1970s from expansion of the Minnesota population (Mech and Nowak 1981; Thiel and Welch 1981). Wolf monitoring has been conducted since 1979 in Wisconsin involving techniques that include a com- bination of winter track surveys, aerial surveillance of radio-collared Wolves, summer howl surveys, and col- lection of reports by government employees and citi- ~ zens (Wydeven et al. 1995). The following cases of Wolf-Coyote interactions are derived from visual obser- vations from both ground and aerial surveillance, and from interpretation of trails in snow. Results Florence County Trails, raise-leg urinations (RLUs) and radio telemetry data indicated that three lone male Wolves occupied home ranges near one another in western Flo- rence County (45°50'N, 88°50'W) in northeastern Wis- consin between 1982 and 1984. At that time 17 — 25 Wolves lived in 2-4 packs along the Minnesota-Wis- consin border (46°15'N, 92°00'W), and in two north- central Wisconsin packs, 150 km southeast of the border Deh 28 THE CANADIAN FIELD-NATURALIST area (45°30'N, 90°00'W). The Florence County Wolves were 100 km east of the two north-central packs. One of these Wolves, male 077, was captured and collared, and aerially radio-monitored about once-weekly for 250 days between 6 May 1983 and 11 January 1984, occupying a 44 km* home range. Several citizen and US Forest Service reports in- volved sightings of a Coyote trailing a lone Wolf in the areas occupied by the lone male Wolves. Wolf Pro- ject personnel also observed snow sign indicating that lone Wolves and Coyotes interacted. On 17 February 1982, Wolf Project technician, Larry Prenn, and I fol- lowed the trail of a Coyote and Wolf over 3 km. We were not able to ascertain the timing of each canid’s travel, but based on highly convoluted trail maneuvers, it appeared they were traveling together. The Wolf was likely a male (RLUing) and the Coyote was a female (squat urinations, including one with blood). On 6 December 1983, while homing in on the signal of Wolf 077 using a Cesna 180 fixed-wing aircraft, pilot Dan Doberstein and I saw a pair of canids lying within 15- 20 m on the ice of Halsey Lake. One appeared 2 or 3 times larger than the other. After determining that nei- ther was Wolf 077, we began a slow descent. When approximately 200 m above them, the smaller canid stood up and quickly ran off into thick lowland conifer cover. We identified it as a Coyote. As it disappeared into cover, a third, larger canid ran from shore to where the second canid was still lying. On our third pass we approached within 10 m of these canids; the second one was larger, and we agreed they were both Wolves. Shortly thereafter we located Wolf 077 male 9.6 km to the east. The following morning we attempted to inspect their trails, but high winds overnight and thin ice made inter- pretation challenging. We found outlines of two large tracks corresponding in size to Wolf, and one small Coyote-sized track. No evidence of a kill-site was found that might have attracted the two species. Wood County In each of seven winters between 1995 and 2002 high school students under my supervision followed the snow trails of a non-collared male Wolf, 501 that had escaped into the 36 km? Sandhill Wildlife Area (SWA) in Wood County (44°17'N, 90°10'W) (Thiel 2000; Thiel unpublished data) in May 1995. This is a deer research facility enclosed by 3 m tall x 29.5 km long fence. At that time, Wolves were colonizing this portion of west- central Wisconsin. In winter, 1996-1997, a territorial pair of Wolves colonized the area surrounding SWA. In January 1998, a pair of Wolves believed to be yearling siblings, entered SWA, and subsequent snowfall pre- vented them from digging out beneath the fence and leaving. Wolf 501 paired with one of these Wolves, a female based on bloody urine discovered on 20 Janu- ary 1998. The third Wolf thereafter roamed the area as a loner. Vol. 120 In March a 6.5 year-old radioed Coyote was killed by Wolves within SWA. Likely, Wolf 501 male was in- volved in its death. Wolf 501 was last observed with another Wolf in October 1998. In four additional win- ters of snow tracking, Wolf 501 was the only Wolf de- tected within SWA. On 22 January 2001, High School students reported trailing Wolf 501 male and encountered where the Wolf had physically interacted with a Coyote, resulting in an injury to the Wolf’s right front foot. On 5 February several students and I encountered Wolf 501 male’s trail where it intersected a Coyote’s trail, heading in the same direction. In backtracking we determined the Coyote was a proestrus female based on bloody urine. The mutual trail led toa 5 m x 5 m site completely padded down with Wolf and Coyote tracks. Numerous small droplets of blood and several tufts of canid fur were found in the padded-down site. Genetic testing to ascertain the species leaving the blood and fur was cost- prohibitive. By carefully back-trailing and following the tracks of both the Coyote and Wolf 501 male as they left the site we established the Wolf had been injured by the proestrus Coyote at the padded down site. The Coyote and Wolf walked on the same trail for approximately 1 km beyond the padded down site be- fore separating. Bloody Coyote urine was noted three times along this trail, and the Wolf inspected each. At one spot along the trail the Coyote and Wolf sat down within 1.5 m of each other. We followed the Wolf an additional 3.2 km after their trails separated. The Wolf RLU’d 8 times, and in each of five beds we found the imprint of his bloody limb. It was evident that both canids were present together and spent considerable time at the padded down site. Since we found no evidence of prey remains, and since Wolf 501 male did not react aggressively when injured by the proestrus Coyote we concluded that his injury was caused by unwanted sexual advances. At approximately 0830 (CDT) on 22 June 2001, amateur wildlife photographer, Rudi Van Stedum, saw male Wolf 501 male standing on rock talus in a forest in SWA at a distance of approximately 50 m. The Wolf was watching her as she idled her car along a woods trail. Simultaneously Van Stedum saw movement ob- liquely behind and left of the Wolf. A Coyote appro- ached but retreated when it became aware of her car. Meanwhile, the Wolf turned and left the rock, facing her. At that point, the Coyote again approached the Wolf from behind and thrust its muzzle forcefully into the left rump of the Wolf. A yelp from the Wolf was clearly audible while the Wolf’s gaze remained fixed on Van Stedum. The Coyote stepped back, and both stood still for a moment before the Wolf walked a ways off and turned momentarily to look at her. As the Wolf began slowly walking away, the Coyote approached it from behind and jabbed it a second time in its left hip with its muzzle. The Wolf yelped again. Both then trailed off into the undergrowth. 2006 THIEL: CONDITIONS FOR SEXUAL INTERACTIONS BETWEEN GREY WOLVES AND COYOTES 29 Wildlife biologist Wayne Hall and I inspected Van Stedum’s photographs, and while they were taken in poor light conditions and no single frame showed both the Wolf and Coyote, the position of trees in the background of all photographs attested to the accuracy of her observation. Wolf 501 male roamed SWA alone in winter 2001-2002 and disappeared sometime after October 2002. No evidence of hybrids materialized subsequent to the observations in 2001. Discussion Coyote mtDNA in Wolves was calculated as the result of six hybridization episodes between male Wolves and female Coyotes (Lehman et al. 1991); Wayne and Vila (2003) speculated that, “... female wolves and male coyotes are more closely matched in size, and thus may also be more likely to mate...”, but they further noted that no Coyotes sampled had Wolf- like haplotypes. Several researchers have speculated that male Wolf dispersers provide the potential pool for male Wolf x female Coyote unions (Lehman et al. 1991; Wayne and Vila 2003; among others). While probably true, this is not necessarily because male Wolves disperse beyond the edge of Wolf range, as is implied. During winter 1983-1984, Michigan and Wisconsin DNR biologists discovered several lone male Wolves living in a several thousand square kilometer area. Some of these Wolves, like Wolf 077, seemed to have home ranges, as evi- denced by RLU’ ing (R. Thiel and J. Hammill, unpub- lished notes). They likely originated from at least 100 km away where the nearest known packs then existed. But female Wolves also dispersed to this area. In sum- mer-fall 1986 radio-collared female Wolf 035 dispersed from Douglas County, Wisconsin (46°15'N, 92°00'W) 277 km to Iron County, Michigan (46°20'N, 88°59'W) and settled into a 174 km” home range, becoming the first known Wolf to reach upper Michigan since their demise in the late 1950s (Thiel 1988). The first breed- ing Wolf pack in upper Michigan would establish itself within this same area in 1990-1991 (J. Hammill, per- sonal communication, | March 2004), implying that other female Wolves also dispersed to this region. The male Wolf x female Coyote hybridization hypothesis is therefore not explained in our region by any superior dispersal advantage displayed by male Wolves. Because of their size, Wolves can easily overpower and kill Coyotes. Ballard et al. (2003: page 267, Table 10.4) summarized Wolf-Coyote interactions in Yellow- stone National Park and noted that 11 percent of inter- actions resulted in killed Coyotes. In fatal encounters, the number of Wolves exceeded the number of Coy- otes. Wisconsin biologists have also encountered aggres- Sive interactions between Wolves and Coyotes, includ- ing killings of Coyotes. Killings involved superior numbers of Wolves (R. Thiel, unpublished data). In contrast, the few observations of single Wolf and Coy- ote encounters are not known to have resulted in killings. In the fall of 1981 pilot Jim Dienst! and I watched dispersing lone male Wolf 023 chase a Coyote from a White-tailed Deer (Odocoileus virginianus) fawn kill, but made no attempt to catch or kill it. SWA male Wolf 501 likely killed at least one Coyote when paired with a female Wolf. Four years later, this same male Wolf - who remained a loner in those four years consorted with one or more Coyotes on at least three occasions; one case involving a proestrus female Coy- ote. As in the Wisconsin cases reported here, Michigan DNR biologist, J. Hammill (personal communication, 1 March 2004), witnessed two separate incidents where a single Wolf and Coyote apparently traveled and bed- ded down together during the early years of Wolf re- colonization. Within Wolf range, owing to the aggressive stance pack Wolves normally display towards foreign Wolves and Coyotes (Arjo and Pletscher 1999; Ballard et al. 2003), hybridization events must be exceedingly rare. Observations reported here indicate that single Wolves are capable of genial behavioral interactions with Coy- otes. Further, they suggest a potential for sexual ad- vances of male Wolves towards female Coyotes, when and where male Wolves lack access to female Wolves. Unions between male Wolves and female Coyotes probably remain localized (however, note Schmitz and Kolenosky 1985a; Sears et al. 2003; and Theberge and Theberge 2004), and likely occur in areas of exceed- ingly low Wolf densities where access to female Coy- otes by single male Wolves far exceeds access to female Wolves. Under such circumstances, hybridiza- tion is at least possible, and then depends on the behav- ioral nuances of these canids as individuals. Acknowledgments I thank L. D. Mech, United States Geological Sur- vey, and A. Wydeven, Wisconsin DNR, for comments on an earlier draft of this manuscript. Literature Cited Arjo, W. M., and D. H. Pletscher. 1999. Behavioral responses of coyotes to wolf recolonization in northwestern Montana. Canadian Journal of Zoology 77: 1919-1927. Ballard, W. B., L. N. Carbyn, and D. W. Smith. 2003. Wolf interactions with non-prey. Pages 259-271 in Wolves: Behavior, Ecology, and Conservation. Edited by L. D. Mech and L. Boitani. University of Chicago Press. Boyd, D. K., S. H. Forbes, D. H. Pletscher, and F. H. Allen- dorf. 2001. Identification of Rocky Mountain gray wolves. Wildlife Society Bulletin 29: 78-85. Kolenosky, G. B. 1971. Hybridization between wolf and coy- ote. Journal of Mammalogy 52: 446-449. Lehman, N. E., A. Eisenhawer, K. Hansen, L. D. Mech, R. O. Peterson, P. J. P Gogen, and R. K. Wayne. 1991. Intro- gression of coyote mitochondrial DNA into sympatric North American gray wolf populations. Evolution 45: 104-119. Mech, L. D., and R. M. Nowak. 1981. Return of the gray wolf to Wisconsin. American Midland Naturalist. 105: 408-409. 30 THE CANADIAN FIELD-NATURALIST Nowak, R. M. 2003. Wolf evolution and taxonomy. Pages 239 - 258 in Wolves: Behavior, ecology, and conservation. Edited by L.D. Mech and L. Boitani. University of Chicago Press. Phillips, M. K., V. G. Henry, and B. T. Kelly. 2003. Restora- tion of the red wolf. Pages 272-288 in Wolves: Behavior, ecology, and conservation. Edited by L. D. Mech and L. Boitani. University of Chicago Press. Pilgrim, K. L., D. K. Boyd, and S. H. Forbes. 1998. Testing for wolf-coyote hybridization in the Rocky Mountains using mitochondrial DNA. Journal of Wildlife Management 62: 683-689. Roy, M.S., E. Geffen, D. Smith, E. A. Ostrander, and R. K. Wayne. 1994. Patterns of differentiation and hybridization in North American wolflike canids, revealed by analysis of microsatellite loci. Molecular Biology and Evolution 11: 533-570. Sears, H. J., J. B. Theberge, M. T. Theberge, I. Thornton, and G. D. Campbell. 2003. Landscape influence on Canis morphological and ecological variation in a Coyote-Wolf C. lupus X latrans hybrid zone, southeastern Ontario. Canadian Field-Naturalist 117: 589-600. Schmitz, O. J., and G. B. Kolenosky. 1985a. Wolves and coy- otes in Ontario: morphological relationships and origins. Canadian Journal of Zoology 63: 1130-1137. Schmitz, O. J., and G. B. Kolenosky. 1985b. Hybridization between wolf and coyote in captivity. Journal of Mammalo- gy 66: 402-405. Silver, H., and W. T. Silver. 1969. Growth and behavior of the coyote-like canid of northern New England with obser- vations on canid hybrids. Wildlife Monograph (17). 41 pages. Theberge, J. B., and M. T. Theberge. 2004. The wolves of Algonquin Park: a 12 year ecological study. Department of Geography, Publication Series Number 56. University of Waterloo. Thiel, R. P. 1988. Dispersal of a Wisconsin wolf into upper Michigan. Jack-pine Warbler 66: 143-147. Vol. 120 Thiel, R. P. 2000. Successful release of a wild wolf, Canis lupus, following treatment of a leg injury. Canadian Field- Naturalist 114: 317-319. Thiel, R. P., and R. J. Welch. 1981. Evidence of recent breed- ing activity in Wisconsin wolves. American Midland Natu- ralist 106: 401-402. Wayne, R. H., N. Lehman, and T. K. Fuller. 1995. Conser- vation genetics of the gray wolf. Pages 399-407 in Ecol- ogy and conservation of Wolves in a changing world. Edit- ed by L.N. Carbyn, S. H. Fritts and D. R. Seip. Canadian Circumpolar Institute. Wayne, R. K., and C. Vila. 2003. Molecular genetic studies of wolves. Pages 218-238 in Wolves: Behavior, ecology, and conservation. Edited by L. D. Mech and L. Boitani. Univer- sity of Chicago Press. Wilson, P. J.,S. Grewal, I. D. Lawford, J. N. M. Heal, A. G. Granacki, D. Pennock, J. B. Theberge, M. T. Theberge, D. R. Voigt, W. Waddell, R. E. Chambers, P. C. Paquet, G. Goulet, D. Cluff, and N. B. White. 2000. DNA profiles of the eastern Canadian wolf and the red wolf provide evi- dence for a common evolutionary history independent of the gray wolf. Canadian Journal of Zoology 78: 2156-2166. Wilson, P. J.,S. Grewal, T. McFadden, R. C. Chambers, and B.N. White. 2003. Mitochondrial DNA extracted from east- em North American wolves killed in the 1800s is not of gray wolf origin. Canadian Journal of Zoology 81: 936-940. Wydeven, A. P., R. N. Scultz, and R. P. Thiel. 1995. Monitor- ing a recovering gray wolf population in Wisconsin, 1979- 1991. Pages 147-156 in Ecology and conservation of Wolves in a changing world. Edited by L. N. Carbyn, S. H. Fritts and D. R. Seip. Canadian Circumpolar Institute. Young, S. P., and E. A. Goldman. 1944. The wolves of North America. Dover, New York. / American Wildlife Institute. Washington, D.C. 636 pages. Received 22 March 2004 Accepted 7 November 2005 Multiple Scale Den Site Selection by Swift Foxes, Vulpes velox, in Southeastern Colorado ANN M. KiITCHEN!, Eric M. GESE?, and SARAH G. Lupts! ' Department of Forest, Range and Wildlife Sciences, Utah State University, Logan, Utah 84322-5230 USA. Present address National Zoological Park, Smithsonian Institution, Washington DC 20013-7012 USA. Corresponding author: Henderson An@si.edu 7 U.S. Department of Agriculture, Wildlife Services, National Wildlife Research Center, Department of Forest, Range and Wildlife Sciences, Utah State University, Logan, Utah 84322-5230 USA. ' Kitchen, Ann M.., Eric M. Gese, and Sarah G. Lupis. 2006. Multiple scale den site selection by Swift Foxes, Vulpes velox, in | southeastern Colorado. Canadian Field Naturalist 120(1): 31-38. Predation by Coyotes (Canis latrans) is a major source of mortality in Swift Fox (Vulpes velox) populations. Year-round den use by Swift Foxes is likely to be a predator avoidance strategy. Due to the importance of denning to Swift Fox ecology, we recorded den site selection of Swift Foxes in southeastern Colorado. Den site selection was recorded at two scales: micro- habitat characteristics at the den and den placement within the home range. The number of den entrances, height and width of each entrance, aspect, hill position, slope, percent rock in soil, vegetative cover, and horizontal foliar density of 42 Swift Fox dens were examined during December 1999 — April 2000. This was compared to the same microhabitat characteristics at 42 random sites within Swift Fox home ranges to determine if Swift Foxes were using site characteristics according to their availability. Our results indicated that Swift Foxes were not highly selective of den sites based on the microhabitat characteristics evaluated in this study, although Swift Foxes selected areas of intermediate rock percentages. In addition, Swift Foxes were radio-tracked throughout the sample period and the location and frequency of use of known dens were recorded. Within the core area of home ranges, Swift Foxes used more dens (mean = 3.51 + 1.70 (SD)), and had a higher frequency of use of dens (mean = 8.20 + 6.01) than in the mid-range area (number of dens, mean = 0.90 + 0.94; frequency, mean = 1.27 + 2.12) and the boundary area (number of dens, mean = 0.34 + 0.53; frequency, mean = 0.45 + 0.93) of the home range. We discuss our results in terms of the importance of dens in facilitating escape from Coyotes. These results illustrate the need for examining den site selection at multiple scales to determine all selection factors, and to provide infor- ‘mation useful for recovery and management efforts for this species. Key Words: Swift Fox, Vulpes velox, denning, habitat selection, home range use, Colorado. Introduction The Swift Fox (Vulpes velox) is native to the short and mid-grass prairies of North America. The current range of the Swift Fox is much reduced from the his- toric range, and the species was formerly a candidate for endangered species listing (United States Fish and Wildlife Service 1996). The Swift Fox has been local- ly extirpated in the northern prairies in Canada, North Dakota, and much of South Dakota (Scott-Brown et al. 1987). One of the principal causes of Swift Fox pop- ulation declines across much of the historical range is a reduction in suitable habitat (Hillman and Sharps 1978). __ The combined availability and distribution of suitable _den sites and escape holes is likely a habitat compo- nent for Swift Foxes (Egoscue 1979; Rongstad et al. 1989; Herrero et al. 1991). Swift Foxes were referred to as the “burrowing fox” by Lewis and Clark and are ‘ considered one of the most fossorial canids in North | America because they use dens throughout the year | (Kilgore 1969; Egoscue 1979). Swift Foxes spend the / majority of diurnal hours in dens or near den entrances, | often concurrently with a mate (Cutter 1958; Kitchen et al. 1999). Denning might enable Swift Foxes to maintain homeostasis by providing a cool, damp micro- habitat refuge. Dens offer shelter from the elements and facilitate escape from predators, especially Coyotes (Canis latrans), in an environment that offers little nat- ural cover (Egoscue 1979; Rongstad et al.1989; Her- rero et al. 1991; Pruss 1999). Kitchen et al. (1999) found all Coyote-killed Swift Foxes near the periphery or outside their home range boundary, a significant dis- tance from their nearest currently used den. Thus, knowledge of habitat requirements for den sites might be an important component of Swift Fox conservation efforts (Hillman and Sharps 1978; Pruss 1999): however, results of studies on Swift Fox den site ‘selection have varied. Swift Foxes may modify bur- rows dug by other animals such as American Badgers (Taxidea taxus), ground squirrels (Spermophilus spp.) or Black-tailed Prairie Dogs (Cynomys ludovicianus), or dig their own dens (Kilgore 1969; Hillman and Sharps 1978; Cameron 1984; Pruss 1999). Dens have been found in a variety of habitat types including short- and mid-grass prairie, grazed prairie, cultivated fields, fence rows, and rock outcrops (Cutter 1958; Hines 1980; Cameron 1984: Urésk and Sharps 1986; Rong- stad et al. 1989; Pruss 1999). Occasionally, dens have been associated with man-made structures such as cul- verts, buildings, and cemetery gravesites with a con- crete cap (Kilgore 1969; Hillman and Sharps 1978; Jackson and Choate 2000). Dens are frequently locat- Sil a2 THE CANADIAN FIELD-NATURALIST ed on or near tops of gently sloping hills, which are well drained and have a clear line of sight (Hillman and Sharps 1978; Cameron 1984; Uresk and Sharps 1986; Pruss 1999; Harrison 2003). However, dens also have been found in level, open areas and at the base of hills (Kilgore 1969; Hines 1980; Cameron 1984). Den entrance characteristics vary among seasons and individual dens. There are indications that natal dens typically have more entrances than non-natal dens (Kilgore 1969; Hillman and Sharps 1978). Hines (1980) reported that Swift Foxes seem to select for entrances with west and east exposures. Similarly, Uresk and Sharps (1986) documented a trend toward dens with easterly exposures. In contrast, Rongstad et al. (1989) and Pruss (1999) found entrance exposure to be random. An additional factor that might influence the selec- tion of dens is the position of the site within the home range. Previous studies investigating den use in Swift Foxes have not considered den placement within the range as a factor influencing selection. We contend, however, that due to the importance of dens for pred- ator escape (Herrero et al. 1991; Kitchen et al. 1999; Pruss 1999), placement relative to the home range core and boundary areas might be a driving force in selec- tion. It is likely den placement is correlated to overall space use patterns and dens are positioned to facili- tate a quick escape from an approaching predator. The objective of this study was to determine the fac- tors affecting den site selection for Swift Foxes on the Pinon Canyon Maneuver Site, in southeastern Col- orado. We examined den site placement and micro- habitat characteristics of active Swift Fox dens and compared these to sites not used by Swift Foxes. Methods Study Area We conducted the study on the 1040-km? Pinon Canyon Maneuver Site (PCMS) in Las Animas Coun- ty, Colorado. The dominant cover type on the PCMS is native grassland (60%), with Juniper (Juniperus monosperma) and Pinyon Pine (Pinus edulis) com- munities found in the hills and canyons (Shaw et al. 1989). Annual precipitation ranges from 26 to 38 cm (United States Department of Army 1980). Mean monthly temperatures range from —1°C in January to 23°C in July (Andersen and Rosenlund 1991). Preda- tors known to kill Swift Fox are Coyotes (Covell 1992; Kitchen et al. 1999) and raptors (Covell and Rongstad 1990). Swift Fox Capture Swift Foxes were captured with double-door box traps (80 x 25 x 25 cm) baited with chicken or mack- erel (Covell 1992). We deployed traps in the evening and checked them the following morning. Traps were not activated during periods when nighttime temper- atures dropped below -10°C. We attached a radiocollar Vol. 120 and ear tag to the Swift Fox and recorded the weight, sex and approximate age (using tooth wear) of the ani- mal (Rongstad et al. 1989). All Swift Foxes were re- leased at the site of capture. We followed telemetry procedures recommended by White and Garrott (1990). Locations were obtained by triangulating 2-3 bearings to the animal’s position within a 10-minute period. Triangulation angles were maintained between 20° and 160° (Gese et al. 1988). We used aerial telemetry (Mech 1983) to locate missing animals. Telemetry error was determined by comparing telemetry locations with actual locations of stationary reference transmitters. When Swift Foxes were located in a den, we record- ed the UTM coordinates of the den. We attempted to locate Swift Foxes approximately every 1-3 days with locations obtained throughout the 24-h period to reduce bias in home range estimates (used for den placement analysis). Den Site Characteristics Microhabitat characteristics of den sites were meas- ured at a randomly selected sub-sample of the known active dens used by Swift Foxes during the breeding season (defined on the basis of energetic demands due to climatic changes and prey abundance, and behay- ioral characteristics to be 15 December to 14 April). Active dens were found by tracking radio-collared Swift Foxes. Den site selection was not stratified by sex as Swift Foxes form pair bonds with males and females concurrently using the same dens. The following physical characteristics were meas- ured at each den site: number of entrances, height and width of each entrance, den aspect, den position on hills, slope of site, percent rock in soil, vegetative cover, and horizontal foliar density. These character- istics were selected based on field observations and published descriptions of potentially important habi- tat characteristics (Hines 1980; Cameron 1984; Ursek and Sharps 1986; Rongstad et al. 1989; Covell 1992; Pruss 1999). Den position on a hill was classified as bottom, mid- dle, top, or flat when there was no hill. We determined maximum slope outside the den using a hand-held cli- nometer (Hays et al. 1981). The percentage of rock in the soil was visually estimated from the presence of rock in the dirt mound runway leading to the den entrance. Vegetation cover was determined by estimat- ing the percent cover of grasses, forbs, shrubs, and bare soil in eight 1-m? plots randomly located within 10 m of the den. Percent soil and vegetation cover were esti- mated within the nearest 5% category. We indexed horizontal foliar density at 5 distances (10, 20, 30, 40, and 50 m) from the den in the four cardinal directions with a vegetation profile board and techniques for the fixed distance approach (Hays et al. 1981). The number of squares visible on a vertical board was counted by an observer lying down to approximate the visibility from the height of a Swift Fox. 2006 BRITISH COLUMBIA KITCHEN, GESE, AND LupPIS: DEN SITE SELECTION BY Swirr FOXES 33 Current distribution W@ Historic distribution Study site (PCMS) FicurE |. Swift Fox current and historic range and study area, Pinon Canyon Maneuver Site, Colorado, 2000. Den Site Selection To examine den site selection, we compared micro- habitat characteristics of dens to control sites that rep- resented available sites (Taylor et al. 1999). For each of the active den sites, we randomly selected one control site (not a den) within the Swift Foxes home range. We used the control site to compare the site characteristics of active den sites versus random sites within the Swift Foxes’ home range. The control site was located 500 m : away from the active den site in a random direction. This distance was arbitrary but allowed for selection of a random location within the home range. Because control sites were in the home range of the Swift Fox inhabiting the active den site, they were considered available as a possible denning location to the Swift Fox. The same den site characteristics were evaluated on the control sites, excluding variables associated with den entrances. Den Placement Dens used by Swift Foxes during the breeding sea- son were recorded using telemetry procedures to assess the placement and use of the dens within the home range. The Swift Foxes’ home ranges were described using a 95 % fixed kernel home-range estimator based on > 30 locations recorded within the breeding sea- son (Worton 1989) with Arcview 3.0 software (Envi- ronmental Systems Research Institute, Inc., Redlands, California). Areas within the home range were then defined as the core area, which was the area within the 50% isopleth, the mid-range area, between the 50% and 75% isopleths, and the boundary area, between the 75% and the 95% isopleths. The number of dens and the frequency of use of those dens in the core, mid- range, and boundary areas of the home range were compared. In addition, we compared the number of dens in each area to the number expected from the proportion of each area in the home range; i.e., 50% within the core area, 25% within the mid-range area, and 20% within the boundary area. Statistical Analysis Because the habitat measurements were categorical, chi-square tests were used to assess the difference between habitat characteristics at active dens versus control sites (Zar 1999). Whether den entrances were randomly distributed around a circle (i.e., no predom- inant direction) was tested using Rayleigh’s nonpara- metric Z test (Zar 1999). We attempted to predict class membership (i.e., active dens or control sites) by using a classification tree analysis with cost complexity prun- ing (CART: Verbyla 1987; S-Plus Version 6.0 1988- 34 THE CANADIAN FIELD-NATURALIST 2001.). Also called tree regression (Rejwan et al. 1999), this method repeatedly partitions the study sites into two groups (active dens and control sites) that are as similar as possible based upon the five variables com- mon to both groups (position, slope, horizontal foliar density, percent rock in soil, and percent vegetative cover). Each of the independent variables is used at each step in the analysis regardless of whether they were previously used in the tree. The tree that is cre- ated is hierarchically structured with the complete data set at the top (the root) and the binary splits, referred to as “nodes”, below to the final undivided “leaves” or groupings at the bottom of the tree. Analysis of variance with Tukey-adjusted post-hoc comparisons was used to assess the difference in the number of dens used and the number of times these dens were used by area (SAS Version 8.2 2001). Log transformations were used to achieve normality where deviations occurred. In cases where more than one den per Swift Fox was sampled, one den was randomly chosen for analysis to avoid pseudoreplication. Results Forty-two dens belonging to 42 Swift Foxes were located from December 1999 to April 2000 and used to evaluate den site characteristics. Most active dens had only one or two entrances and the maximum num- ber of den entrances was seven. The mean height of den entrances was 19.4 + 3.1 cm and the mean entrance width was 17.9 + 2.5 cm. The percentage of dens with entrances oriented to the north, south, east, and west were 29.9, 21.1, 21.9, and 27.1%, respectively. These aspects were randomly distributed around a circle (Z,, = 0.088; P = 0.54), i.e., dens were not oriented in a predominant direction. The mean maximum slope of active dens was 1.3 + 1.3 degrees. We found no difference between the slope at den sites and control sites (y? = 2.71, df = 4, P=0.61). Most dens (88.7%) and control sites (80.3%) were located on gentle slopes between 0 and 2 degrees. Four percent of control sites were located on slopes 60 50 - 40 - Percent Use (do) o>) I Vol. 120 that exceeded 5 degrees, however, no dens were found on slopes that steep. The majority of Swift Fox dens were positioned either on flat ground or in the middle of a hill, and relatively few dens were located at the top or bottom of a hill (Figure 1). Control sites were found in similar positions with no detected difference between site position of used and control sites (y* = 2.04, df = 3, P=0.59; Figure 1). We also found no difference in the pattern of visibility by distance from the den as com- pared to control sites (y* = 0.06, df = 4, P = 0.99). Visibility from both active den sites and control sites decreased uniformly in each direction as one moved away from the site. The vegetative cover of den sites and control dens was similar (y7 = 1.98, df = 3, P =0.598) with both dens and control sites occurring in areas mostly covered in grasses and soil (Figure 2). There was a significant difference in the percentage of rock at dens versus control sites (y7 = 8.91, df = 3, P = 0.03; Figure 3) with dens having higher propor- tions of intermediate percentages of rock and control sites having higher proportions of rock below 5% or above 20%. The classification tree used to characterize the habitat at dens versus control sites failed to pro- duce any biologically relevant pattern. The average number of dens used per Swift Fox during the breeding season was 5.1 + 2.1. The num- ber of dens used by Swift Foxes varied by area of the home range (Wa 26 = 37.53, P < 0.001; Figure 4a). Tukey’s adjusted post-hoc tests indicated that more dens were used in the core area than in the mid-range and boundary areas (mid-range: t,, = 6.29, P < 0.001; boundary: t= - 7.54, P < 0.001). There were also sig- nificantly more dens in the mid-range area than the boundary area (t,, = -2.20, P = 0.035). In addition, we found that the number of dens in each area varied from that expected from the proportion of the area, and thus the frequency of use of the area (y* = 38.41, df = 3, P < 0.0001). Overall, there was a difference in the frequency of use of dens in the different areas of the home range (F,.. = 44.94, P < 0.001; Figure 4b). The number of 2,33 w Den sites O Control sites | (0) 5; Bottom Middle Position Top Flat FIGURE 2. Position of Swift Fox den sites and control sites, Pinon Canyon Maneuver Site, Colorado, 2000. 2006 100 - ioe) i=) Percent Use (o>) Oo NO A Sy [> I i] | i — Shrubs Grasses KITCHEN, GESE, AND LupIs: DEN SITE SELECTION BY SWIFT FOXES Soil -) wv g Den sites 0 Control sites Forbs Vegetative Cover FIGURE 3. Vegetative cover at Swift Fox den sites and control sites, Pinon Canyon Maneuver Site, Colorado, 2000. times that dens in the core area were used was higher than in the mid-range or boundary areas (mid-range: t,,= 7.52, P < 0.001; boundary: t,= 7.65, P <0.001). There was no difference in the number of times dens in the mid-range and boundary areas were used (f,, = 1.55, P = 0.13). The pattern of den use was variable. For example, one Swift Fox was located in only one den for a period spanning 67 days, while another Swift Fox used at least four different dens in a five-day period. Discussion Our results indicated Swift Foxes on the PCMS did not select den sites based on the microhabitat charac- teristics that we evaluated. However, Swift Foxes ap- peared to select areas of intermediate rock percentages. The selection of areas with intermediate rock percent- ages may be related to den structure and strength. Foxes had no preference for particular positions or vegeta- tion at den sites despite other studies concluding that Swift Foxes preferentially selected den sites on hilltops (Uresk and Sharps 1986; Pruss 1999; Harrison 2003). The finding that Swift Foxes in our study area select den sites with habitat characteristics that do not differ from control sites was not unexpected considering the homogeneous prairie environment on the PCMS. Our result does differ from other Swift Fox studies which reported preferences for certain habitat characteristics, however. The importance of selection of hill position and vegetation characteristics might be determined by variation in topography and habitat in the geographic area. Habitat selection by Swift Foxes on the PCMS appears to operate at the landscape level instead of at the den site level as Swift Foxes were only trapped on the open prairie on the PCMS despite efforts to capture Swift Foxes in wooded areas (Schauster et al. 2002). Den site selection occurred at a larger scale; the placement of dens relative to areas within the home range was a significant factor in site selection. Foxes ® 27680): cob) B08. ‘m Den sit 3 40 m Den sites Oo 20 ‘oO Control sites 0 2) ee < o 9Q\o 9 9 9 ao. oa oY s\ oS io s a Percent Rock FicurE 4. Percentage of rock at Swift Fox den sites and control sites, Pinon Canyon Maneuver Site, Colorado, 2000. 36 THE CANADIAN FIELD-NATURALIST Vol. 120 6 - ane rane oe. @ 2 a = 0 hy Ne le Mid-range Boundary Area & 15 - Y” ® Ey 0] or 7) o 7 a4 2 E > 0 Core Mid-range Boundary Area FIGURE 5. Placement of Swift Fox dens within the home range (top) and frequency of Swift Fox den use by area of the home range (bottom), Pinon Canyon Maneuver Site, Colorado, 2000. used more dens in the core area of the home range as compared to the mid-range or boundary areas. In addi- tion, dens in the core area were used more frequently by radio-tracked Swift Foxes than dens in the mid- range or boundary area. Den placement within the home range may be designed to facilitate escape from Coyotes. Previous studies have indicated that Swift Foxes use dens as a mechanism of escape from pred- ators (Herrero et al. 1991; Kitchen et al. 1999; Pruss 1999) and that Coyote-caused mortality might be a key factor in limiting population growth in North America as well as the re-establishment of Swift Foxes on the Canadian prairies (Scott-Brown et al. 1987). Kitchen et al. (1999) found that all Coyote-caused Swift Fox mortalities occurred near or outside the Swift Fox’s home range boundary, a substantial distance from their nearest currently used den. Thus, Swift Foxes might maintain the majority of dens in the core area of the home range where they spend most of their diurnal hours (Schauster et al. 2002), and use dens only occasionally in the outer areas of their range to reduce encounters with and facilitate escape from predators. Previous studies have suggest- ed that other preferences for den sites also might be due to Coyote avoidance, including den locations that are negatively associated with water sources (Pruss 1999), and dens that are closer to roads than unoccu- pied sites (Pruss 1994; Harrison 2003). The importance of interspecific competition in deter- mining den site selection has also been noted in Kit Foxes (List and Macdonald 2003) and Arctic Foxes (Alopex lagopus; Frafjord 2003). Frafjord (2003) doc- umented that the distance to Red Fox (Vulpes vulpes) dens and the elevation from the tree line were the most important factors determining den use in Arctic Foxes. Kit Foxes in Mexico denned more often in grasslands and less in Prairie Dog (Cynomys ludovicianus) towns than expected (List and Macdonald 2003). These au- thors hypothesized that Kit Foxes reduced the amount of time they spent in Prairie Dog towns due to the activity of Coyotes. Thus, Swift Fox den site selection in homogenous environments might be unrelated to microhabitat char- acteristics and more dependent on interspecific inter- actions and predator avoidance. This study also illus- trates the importance of examining den site selection at multiple scales to determine all site selection fac- tors. Due to the importance of denning to Swift Fox 2006 ecology, a full understanding of den site selection in various environments is essential to recovery, reintro- duction, and management efforts. Acknowledgments Funding and assistance provided by the U.S. Army, Directorate of Environmental Compliance and Man- agement (DECAM), Fort Carson, Colorado, through the U.S. Fish and Wildlife Service (USFWS), Colorado Fish and Wildlife Assistance Office, Golden, Colorado, and the Utah Cooperative Fish and Wildlife Research Unit at Utah State University, Logan, Utah. Addition- al support provided by the U.S. Department of Agri- culture, Wildlife Services, National Wildlife Research Center, Logan Field Station at Utah State University, Logan, Utah. We thank T. Warren, G. Belew, and R. Bunn at DECAM, and B. Rosenlund at the USFWS for logistical support. We thank E. Bergman, E. Joyce, S. Karki, M. Klavetter, A. Kozlowski, H. Kratz, and L. Schutte for field assistance. Research protocols were approved by the Institutional Animal Care and Use Committees (IACUC) at the National Wildlife Research Center and Utah State University. Literature Cited Andersen, D. E., and B. D. Rosenlund. 1991. Fish and wildlife management recommendations: Pinon Canyon Maneuver Site, Las Animas County, Colorado. 1-84. United States Fish and Wildlife Service, Golden, Colorado, USA. Cameron, M. W. 1984. The Swift Fox (Vulpes velox) on the Pawnee National Grassland: its food habits, population dynamics and ecology. Unpublished MS thesis, University of Northern Colorado, Greeley, Colorado. Covell, D. F. 1992. Ecology of the Swift Fox (Vulpes velox) in southeastern Colorado. M.S. thesis, University of Wis- consin, Madison, Wisconsin. Covell, D. F., and O. J. Rongstad. 1990. Ecology of swift fox on the Pinon Canyon Maneuver Site. Annual Report to Fort Carson, Colorado. Cutter, W. L. 1958. Denning of the Swift Fox in Northern Texas. Journal of Mammalogy 39: 70-74. Egoscue, H. G. 1979. Vulpes velox. Mammalian Species 122: 1-5. Frafjord, K. 2003. Ecology and use of arctic fox Alopex lagopus dens in Norway: tradition overtaken by interspe- cific competition? Biological Conservation 111: 445-453. Gese, E. M. 1998. Response of neighboring Coyotes (Canis latrans) to social disruption in an adjacent pack. Canadian Journal of Zoology 76: 1960-1963. Gese, E. M., O. J. Rongstad, and W. R. Mytton. 1988. Home-range and habitat use of coyotes in southeastern Colorado. Journal of Wildlife Management 52: 640-646. Harrison, R. 2003. Swift Fox demography, movements, den- ning, and diet in New Mexico. Southwestern Naturalist 48: 261-273. Hays, R. L., C. Summers, and W. Seitz. 1981. Estimating wildlife habitat variables. U.S.D.I. Fish and Wildlife Ser- vice. FWS/OBS-8 1/47. Herrero, S., C. Mamo, L. Carbyn, and M. Scott-Brown. 1991. Swift Fox reintroduction into Canada. Provincial Museum of Alberta. Natural History Occasional Paper Number 15. KITCHEN, GESE, AND LuPIS: DEN SITE SELECTION BY SwWirt FOXES 37 Hillman, C.N., and J.C. Sharps. 1978. Return of Swift Fos to northern Great Plains. Proceedings of the South Dakota Academy of Science 57; 154-162 Hines, T. D. 1980. Home range and movements of Swift Fox (Vulpes velox) as determined by radio telemetry. Proceed ings of the Nebraska Academy of Science 90: 7 Jackson, V. L., and J. R. Choate. 2000. Dens and den sites of the Swift Fox, Vulpes velox. Southwestern Naturalist 45 212-220. Kilgore, D. L. 1969. An ecological study of the Swift Fox (Vulpes velox) in the Oklahoma Panhandle. American Mid land Naturalist 81: 512-533. Kitchen, A. M., E. M. Gese, and E. R. Schauster. 1999 Resource partitioning between Coyotes and Swift Foxes space, time, and diet. Canadian Journal of Zoology 77: 1645-1656. List, R., and D. W. Macdonald. 2003. Home range and habitat use of the kit fox (Vulpes macrotis) in a prairie dog (Cyno- mys ludovicianus) complex. Journal of Zoology 259: 1-5. Mech, L. D. 1983. Handbook of animal radiotracking. Uni- versity of Minnesota Press, Minneapolis. Pruss, S. D. 1994. An observational natal den study of wild Swift Fox (Vulpes velox) on the Canadian Prairies. Unpub- lished Master’s of Environmental Design thesis. Univer- sity of Calgary, Alberta. Pruss, S. D. 1999. Selection of natal dens by the Swift Fox (Vulpes velox) on the Canadian prairies. Canadian Journal! of Zoology 77: 646-652. Rejwan, C., N. C. Collins, L. J. Brunner, B. J. Shuter, and M.S. Ridgway. 1999. Tree regression analysis on the nest- ing habitat of smallmouth bass. Ecology 80: 341-348. Rongstad, O. J., T. R. Laurion, and D. E. Andersen. 1989. Ecology of Swift Fox on the Pinon Canyon Maneuver Site, Colorado. Final Report, DECAM, Ft. Carson, Colorado. S-Plus Version 6.0. 1988-2001. Insightful Corporation, New York, New York. SAS Version 8.2. 2001. SAS Institute, Cary, North Carolina. Schauster, E. R., E. M. Gese, and A. M. Kitchen. 2002. Population ecology of Swift Foxes (Vulpes velox) in south- eastern Colorado. Canadian Journal of Zoology 80: 307- 319. Scott-Brown, J. M., S. Herrero, and J. Reynolds. 1987. Swift Fox. Pages 432-441 in Wild furbearer management and conservation in North America. Edited by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. Ontario Min- istry of Natural Resources, Toronto. Shaw, R. B., S. L. Anderson, K. A. Schulz, and V. E. Diersing. 1989. Plant communities, ecological checklist, and species list for the U.S. Army Pinon Canyon Maneu- ver Site, Colorado. Colorado State University. Science Series 37. Taylor, J. S., K. E. Church, and D. H. Rusch. 1999. Micro- habitat selection by nesting and brood-rearing northern bobwhite in Kansas. Journal of Wildlife Management 63: 686-694. United States Departmeht of Army. 1980. Draft: environ- mental impact statement for acquisition of training land in Huerfano, Las Animas and Pueblo counties, Colorado. United States Department of Army, Fort Carson, Colorado. United States Fish and Wildlife Service. 1996. Endangered and threatened wildlife and plants; review of plant and ani- mal taxa that are candidates for listing as endangered or threatened species. Federal Register 61: 7596-7613. 38 THE CANADIAN FIELD-NATURALIST Uresk, D. W., and J. C. Sharps. 1986. Denning habitat and diet of the Swift Fox in western South Dakota. Great Basin Naturalist 46: 249-253. Verbyla, D. L. 1987. Classification trees: a new discrimina- tion tool. Canadian Journal of Forest Resources 17: 1150- IS. White, G. C., and R. A. Garrott. 1990. Analysis of radio- tracking data. Academic Press, New York, New York. Vol. 120 Worton, B. J. 1989. Kernel methods for estimating the uti- lization distribution in home range studies. Ecology 70: 164-168. Zar, J. H. 1999. Biostatistical analysis. Prentice-Hall, Inc. New Jersey. Received 8 December 2004 Accepted 15 November 2006 First Records of the Northern Long-eared Bat, Myotis septentrionalis, in the Yukon Territory THOMAS S. JuNG!, BRIAN G. SLOUGH”, DAVID W. NAGORSEN?, TANYA A. DEwEY?, and Topp Powe! 'Yukon Department of Environment, Fish and Wildlife Branch, Box 2703, Whitehorse, Yukon Territory Y1A 2C6 Canada; e-mail: thomas.jung@ gov.yk.ca 235 Cronkhite Road, Whitehorse, Yukon Y1A 5S9 Canada $Mammalia Biological Consulting, 4268 Metchosin Road, Victoria, British Columbia V9C 374 Canada 4Mammal Division, Museum of Zoology, University of Michigan, 1109 Geddes Road, Ann Arbor, Michigan 48109-1079 U.S.A. Jung, Thomas S., Brian G. Slough, David W. Nagorsen, Tanya A. Dewey, and Todd Powell. 2006. First record of the Northern Long-eared Bat, Myotis septentrionalis, in the Yukon Territory. Canadian Field-Naturalist 120(1): 39-42. Three adult male Northern Long-eared Bats, Myotis septentrionalis, were captured in mist nets in July 2004 in the LaBiche River Valley, southeastern Yukon. These are the first records of M. septentrionalis in the Yukon. Further survey work is needed to delineate the extent of the range and population structure of this and other species of bats in northwestern North America. Key Words: Northern Long-eared Bat, Myotis septentrionalis, distribution, Yukon. The diversity and distribution of bats in northwestern North America (e.g., =>55° N) is poorly documented. Few areas have been adequately surveyed, particularly for rare or cryptic species (sensu Nagorsen and Brigham 1993; van Zyll de Jong and Nagorsen 1994; Parker and Cook 1996; Parker et al. 1997). Youngman (1975) reported the Little Brown Bat (Myotis lucifu- gus) as the only species of bat in the Yukon. However, Youngman (1975) noted the possibility of other species occurring in the Yukon, such as the Big Brown Bat (Eptesicus fuscus) and Long-legged Bat (Myotis volans), based on single specimens collected near Fair- banks, Alaska, and Atlin, British Columbia, respective- ly. In the 30 years subsequent to Youngman (1975), no species of bats other than M. /ucifugus have been con- firmed from the Yukon. Slough (2000*), however, recorded a bat in southeastern Yukon that appeared to be a Big Brown Bat according to its echolocation call characteristics (C. Corben, personal communication), but it may have been a Silver-haired Bat (Lasionycteris noctivagans) and remains to be verified by a capture. We believe that species other than M. lucifugus may — extend northward into the Yukon, and that a lack of records of other species is largely a function of previous search effort. Based on a specimen collected in Nahanni National Park, Northwest Territories (van Zyll de Jong 1985), captures in recent surveys in adjacent British Columbia (Wilkinson et al. 1995*: Vonhof and Wilkinson 1997*), and an unidentified Myotis in a recording of an echolo- cation call (Slough 2000*), we suspected that the North- em Long-eared Bat (Myotis septentrionalis) and the Western Long-eared Bat (Myotis evotis) could be pres- ent in southeastern Yukon. In summer 2004, we under- took a preliminary field survey of the mammalian diversity in the boreal forest of southeastern Yukon, including a bat inventory. Here, we provide the first records of the Northern Long-eared Bat in the Yukon. Methods On 28 July — 2 August 2004, we used mist-nets to capture bats in the LaBiche River Valley (60.126°N, 124.064°W) of extreme southeastern Yukon. Mist-net- ting occurred at three sites; two sites were over small ponds (>150 m7’) within dense boreal forest, and anoth- er at an intersection of a roughly 3 m wide forest trail and a roughly 30 m wide forest access road. Nets were set across and bordering the ponds and perpendicular to the road and forest trail. Nets were opened 0.5 hr before sunset (about 23:00 hrs PDT) and closed 2—3 hr thereafter. A 6 m long mist net open for | hr was used as our definition of | mist-net hr. Captured bats were examined for external diagnostic characteristics according to van Zyll de Jong (1985) and Nagorsen (2002), including presence of dark shoulder patches in the pelage, ear length, number of hairs on the trailing edge of the uropatagium, presence of keeled calcar, and relative length and shape of the tragus. We measured forearm length, ear length, and mass, and determined age as per van Zyll de Jong (1985). No voucher specimens were procured; rather, we obtained morphological measurements and wing punches for mitochondrial DNA sequencing to verify our field identifications. Wing punches were stored in 70% ethanol until lab analysis. Genetic analyses were conducted at the University of Michigan’s Museum of Zoology (Ann Arbor, Michigan), and included an unweighted maximum parsimony analysis of 745 bp of the cytochrome b gene using 21 specimens of Myotis (Figure 1). 39 40 THE CANADIAN FIELD-NATURALIST Results and Discussion We amassed 42.1 mist-net hr over six nights of sam- pling; only six bats were captured. On 28 July 2004, we captured two adult male M. septentrionalis and one M. lucifugus at a small, shallow pond (ca. 108 m7). On 29 July 2004, a third adult male northern M. septentrion- alis and two M. lucifugus were captured at the same pond. Although three of our captures were consistent with the morphological traits of M. septentrionalis (Table 1), this species can be difficult to distinguish from other species of potentially sympatric long-eared Myotis (1.e., M. evotis and M. keenii), based on external characteristics alone (van Zyll de Jong 1979; Nagorsen 2002). DNA sequences, however, confirmed our tenta- tive species identifications: the three M. septentrionalis are included in the M. septentrionalis clade (Figure 1). The presence of M. septentrionalis in the Yukon is not surprising, considering that captures were reported from elsewhere in the Liard River Watershed by Wilkinson et al. (1995*), Bradbury et al. (1997*) and Vonhof and Wilkinson (1997*). The only voucher taken in those surveys to confirm identification, however, was a specimen of M. septentrionalis collected by Vonhof and Wilkinson (1997*) from the Fort Nelson River, British Columbia, near Highway 77 (Royal British Columbia Museum, RBCM 19516), about 140 km southeast of our captures. It is not known if there are breeding populations of M. septentrionalis in the Yukon, as no juveniles, or lactating or post-lactating females were captured. Breeding populations of the Northern Long-eared Bat, however, have been reported from nearby northeastern British Columbia (Wilkinson et al. 1995*, Vonhof and Wikinson 1997*). Similarly, it is not known if M. septentrionalis hibernates in the Yukon. No bat hibernacula have been found in the Yukon. Despite low capture rates, we suspect that M. septen- trionalis were locally common in our survey area, based on the number of captures relative to M. lucifu- gus. Moreover, we often detected what appeared to be M. septentrionalis at all of the mist-netting sites, based on low intensity echolocation calls (Faure et al. 1993) heard on narrowband bat detectors tuned to 40-50 kHz and observations of detection distances of passing bats. Our data, along with that of Wilkinson et al. (1995*), Bradbury et al. (1997*), and Vonhof and Wilkinson Vol. 120 (1997*), suggest that M. septentrionalis may be locally common and widely distributed throughout the Liard River Watershed in British Columbia and the Yukon. We suspect that M. septentrionalis may occur through- out the Liard River Watershed in the Yukon, from the LaBiche River west to the Little Rancheria River. Other species of bats (e.g., M. evotis, M. volans, Eptesicus fuscus, and Lasionycteris noctivagans) may also be present in southern Yukon, but not yet confirmed with a capture, voucher, or DNA sample. Perhaps the most intriguing question is whether M. septentrionalis is a long-established member of the Yukon mammalian fauna that has simply gone unde- tected, or, alternatively, is it a species that has recently (i.e. <100 years ago) colonized southeastern Yukon? Other mammals have recently colonized the Yukon, from the south, including: Mule Deer and White-tailed Deer (Odocoileus hemionus and O. virginianus; Hoefs 2001), Coyote (Canis latrans; Youngman 1975), and Cougar (Puma concolor; Jung et al. 2005). Moreover, Humphries et al. (2002) provided data from simulation models that suggested that the range of M. lucifugus, and presumably other species of bats in the boreal for- est, would substantially move northward in response to global warming. Regardless of the origin of M. septen- trionalis in the Yukon, these records underscore that our knowledge of the diversity and distribution, and population structure of bats in the Yukon is limited. Directed surveys are needed to address these informa- tion gaps and provide a basis for management and con- servation initiatives that include bats. Acknowledgments S. Cannings and A. Runck kindly provided com- ments on an earlier draft of this note. Additional tissue samples for DNA sequence comparison were kindly provided by T. Dewey, P. Myers, D. Burles, and R. Benedict. The staff of Devon Canada Corporation’s LaBiche River Gas Plant kindly went out of their way to accommodate us and provide logistical support. Funding was provided by the Yukon Department of Environment and NatureServe Yukon. Additional finan- cial support was provided by a Northern Research Endowment Grant from the Northern Research Insti- tute, Yukon College, to B. Slough. TABLE 1. Age, sex, and morphometrics of Myotis septentrionalis (n = 3) captured in the LaBiche River valley, southeastern Yukon, 28-29 July 2004. Specimen GenBank Forearm Ear Mass Label Accession Species! Sex Age Length (mm) Length (mm) g) LaBiche-1 AY883902 M. septentrionalis Male Adult 16 37.7 7.6 LaBiche-3 AY883904 WM. septentrionalis Male Adult 15 35.8 6.6 LaBiche-9 AY883905 M. septentrionalis Male Adult 14 37.6 6.9 ' As tentatively identified in the field and subsequently confirmed with genetic analyses. 2006 100 89 95 JUNG ET AL.: RECORDS OF THE NORTHERN LONG-EARED BAT IN THE YUKON 4] LaBiche9 M. septentrionalis (British Columbia, 5 individuals) Mh. septentrionalis (Nebraska) M. septentrionatis (South Dakota) LaBichel LaBiche3 M. septentrionalis (Michigan) M. auriculus LaBiche2 M. lucifugus (South Dakota) M. lucifugus (South Dakota) LaBiche10 fi. lucifugus (Michigan) M. evotis (South Dakota) M. evotis (Colorado) M. volans MM. yumanensis Eptesicus fuscus FIGuRE |. Placement of Myotis septentrionalis (LaBiche 1,3, and 9) and M. lucifugus (LaBiche 2 and 10) samples collected in the LaBiche River Valley, Yukon, based on an unweighted maximum parsimony analysis of cytochrome b (745 bp) for these samples compared to other Myotis. Eptesicus fuscus is used as an outgroup. Numbers above the branches represent bootstrap support values for those nodes. Documents Cited (marked * in the text) Bradbury, S. M., S. Morris, and S. McNally. 1997. Bat sur- vey of the Liard River watershed in British Columbia. Un- published Report. British Columbia Ministry of Environ- ment, Lands and Parks, Victoria, British Columbia. 29 pages. Slough, B. G. 2000. A survey of the bat fauna of the Yukon Territory: 1999 field studies. Unpublished Report. North- em Research Institute, Yukon College, Whitehorse, Yukon. 26 pages. Vonhof, M. J., and L. C. Wilkinson. 1997. Roosting habitat requirements of northern long-eared bats (Myotis septentri- onalis) in the boreal forest of northeastern British Colum- bia. Unpublished Report. British Columbia Ministry of Environment, Lands and Parks, Fort St. John, British Columbia. 88 pages. Wilkinson, L. C., P. F. J. Garcia, and R. M. R. Barclay. 1995. Bat survey of the Liard River watershed in northern British Columbia. Unpublished Report. British Columbia Ministry of Environment, Lands and Parks, Victoria, British Columbia. 39 pages. Literature Cited Faure, P. A., J. H. Fullard, and J. W. Dawson. 1993. The gleaning attacks of the northern long-eared bat, Myotis septentrionalis, are relatively inaudible to moths. Journal of Experimental Biology 178: 173-189. Hoefs, M. 2001. Mule, Odocoileus hemionus, and White- tailed, O. virginianus, deer in the Yukon. Canadian Field- Naturalist 115: 296-300. Humphries, M. M., D. W. Thomas, and J. R. Speakman. 2002. Climate-mediated energetic constraints on the distri- bution of hibernating mammals. Nature 418: 313-316. Jung, T. S., and P. J. Merchant. 2005. First confirmation of cougar, Puma concolor, in the Yukon. Canadian Field-Nat- uralist 119(4): 580-581. 42 THE CANADIAN FIELD-NATURALIST Nagorsen, D. W. 2002. An identification manual to the small mammals of British Columbia. Ministry of Sustainable Resource Management, Ministry of Water, Land and Air Protection, and Royal British Columbia Museum, Victoria, British Columbia. 153 pages. Nagorsen, D. W., and R. M. Brigham. 1993. Bats of British Columbia. Volume 1. The Mammals of British Columbia. UBC Press, Vancouver, British Columbia. 176 pages. Parker, D. I., and J. A. Cook. 1996. Keen’s long-eared bat, Myotis keenii, confirmed in southeast Alaska. Canadian Field-Naturalist 110: 611-614. Parker, D. 1I., B. E. Lawhead, and J. A. Cook. 1997. Distrib- utional limits of bats in Alaska. Arctic 50: 256-265. Vol. 120 van Zyll de Jong, C. G. 1979. Distribution and systematic relationships of the long-eared Myotis in western Canada. Canadian Journal of Zoology 57: 987-994. van Zyll de Jong, C. G. 1985. Handbook of Canadian mammals. Volume 2: Bats. National Museum of Natural Sciences, Ottawa, Ontario. 212 pages. van Zyll de Jong, C. G., and D. W. Nagorsen. 1994. A review of the distribution and taxonomy of Myotis keenii and Myotis evotis in British Columbia and the adjacent United States. Canadian Journal of Zoology 72: 1069-1078. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museums of Canada, Ottawa, Ontario. 192 pages. Received 4 February 2005 Accepted 12 September 2005 Recent Declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces ANTHONY J. ERSKINE 16 Richardson Street, Sackville, New Brunswick E4L 4H6 Canada Erskine, Anthony J. 2006. Recent declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces. Canadian Field-Naturalist 120(1): 43-49. House Sparrows, Passer domesticus, were introduced to North America after 1850, increased and spread up to 1920, and sta- bilized or decreased thereafter until 1960. In the Maritimes (and perhaps some other areas), a further decline set in after 1970, continuing to the present. Now the species is rare to absent in much of the Maritimes, except around farms with livestock. Decline here since 1970 probably approaches 90 per cent in most other areas of human settlement except south of 45°N Similar declines are known in the U.K., but seem poorly documented, if recognized, in North America outside of our region. Key Words: House Sparrows, Passer domesticus, declines, New Brunswick, Nova Scotia, Prince Edward Island. Fifty years ago, House Sparrows (Passer domesti- cus; HoSp hereafter — a standard abbreviation used in many reports of organized birding activities) were so nearly ubiquitous in and around human settlements in eastern Canada that one sometimes had to “tune them out” to hear other birds. Then, it would have been un- imaginable that this abundant species could become scarce in our lifetimes. HoSp, and their (urban) habi- tats, were virtually beneath notice both of birders and academics, so few serious studies of the species were made (see Kendeigh 1973), and none in Canada, dur- ing the rapid expansion of research on birds after World War 2. The result is a continuing scarcity of hard data on HoSp, precluding rigorous discussion of its popu- lation ecology. This study is a first look at what hap- _ pened in the recent past and what may still be investi- gated usefully, in the Maritime Provinces of Canada {New Brunswick, Nova Scotia, and Prince Edward Island (hereafter “the Maritimes”)]. HoSp were first introduced to North America around 1850, and became established in the Maritimes by the 1880s (Christie 1979; Lowther and Cink 1992). Num- _ bers and densities in the next century are known main- ly from anecdotal remarks in general accounts. As was general in eastern North America by the early 20" Cen- tury, HoSp were nearly ubiquitous and very common around human settlements, including cities, towns, vil- lages, and even isolated farms. HoSp thrived on grain Wastage, as well as using other food wastes. Nearly all _ human settlements, through the 1940s, harboured dom- | estic animals, especially livestock and poultry, which | were fed on small grains at least seasonally. Motor vehicles began replacing horses in transporta- i tion by the 1920s, and the resulting decrease in horse droppings and other waste grain sources led to a decline | in HoSp, documented mainly by general statements ' (e.g., Bent 1958). Motor vehicle use in the Maritimes | was much less during the Great Depression of 1930s, when most people could not afford cars, and in World War 2, when gasoline and tires were reserved for mil- itary use. Use of horses again became widespread in those decades, and that presumably slowed or halted the earlier decline of HoSp. After the war, motor vehi- cles again became available, use of horses declined rap- idly, and probably HoSp also declined. Very few bird counts were made in the Maritimes before 1960, when HoSp still were considered widespread and abundant (Squires 1952; Godfrey 1954; Tufts 1962). Winter bird counts at Wolfville, Nova Scotia (J. S. Erskine 1968) showed no obvious trend in HoSp numbers between 1948 and 1968. Christie (1979) summarized bird status changes in the Maritimes over the century to 1978, not- ing arrival, increase, and subsequent decline in HoSp, with numbers likely stable after 1930. Starting in 1977, winter plot counts through 1988 in Sackville, New Brunswick (Erskine 1992b), showed further decreases in HoSp — evidently nor related to decline of horses and other livestock, which already were absent in such towns (compare Erskine and McManus 2005). Also in the 1980s, Christie (1985 and following years) remarked on generally lower HoSp numbers on New Brunswick Christmas Bird Counts (CBCs hereafter). Some naturalists (personal communi- cations) remarked on similar decreases in Nova Scotia in the early 1990s. The cooperative Breeding Bird Sur- vey (BBS) documented HoSp declines across eastern Canada in summers 1967-1996 (Downes and Collins 2003), with decreases each decade (significant only in 1981-1990) in the Bird Conservation Region (BCR 14) including the Maritimes. HoSp continued to decline, and by 2000 were rare in Sackville, New Brunswick (personal observations), and in nearby towns. In the near-absence of even quasi-systematic counts of HoSp, CBCs provided the only widespread sam- pling of bird numbers in the Maritimes, with many more counts in more years than the BBS. Although CBC data are poorly standardized (e.g., Dunn et al. 2005), those surveys are extensive enough, both here and across 43 44 THE CANADIAN FIELD-NATURALIST Canada, to allow examination of HoSp occurrence in various habitats, in an attempt to identify factors in- volved in their recent declines. Methods Christmas Bird Counts are single-day annual counts in the period 13 December to 5 January, each restrict- ed to a 24-km diameter circle selected subjectively. Bird species richness and availability of observers are usual selection criteria. CBC data for HoSp in the Maritimes in 1960-2003 were extracted from N. B. Naturalist (formerly Nature News), Nova Scotia Birds (formerly N.S. Bird Society Newsletter), and Island Naturalist. Audubon publica- tions (American Birds, formerly Audubon Field Notes) included many fewer counts (“circles” hereafter) in the Maritimes, and were used only to fill gaps in regional publication series. All data were tabulated in year vs. location matrices. Data were selected by excluding circles surveyed in <=20 years and others with obvious “discontinu- ities” in coverage or effort. The selected circles were grouped by major non-urban habitat combinations, most including also some human settlements, thus (names in parentheses used in text and tables hereafter): e (Forest) inland, predominantly forested, little or no farming; e (Coastal) much forest or bog, farming insignifi- cant; e (Farm=forest) inland or coastal, farms and other open areas minor or unproductive but roughly equal in area to forests; e (Farm>forest) farming major (much exceeding forest), relatively productive, usually around a siz- able town; e (Major urban) cities or larger towns, other habi- tats minor. To allow for varying observer effort, between years or circles, published counts (“raw data”) were also con- verted to birds/10 party-hours (“adjusted data”; see also Discussion). Statistics on area changes in major habitats of the Maritimes over the study period (roughly 1960-2000) were assembled from Statscan websites, Canada Year Books, and some other directories. Extrapolation from CBC data to Maritimes HoSp populations was attempted. Counts from the selected circles were expanded across the five habitat groupings above. Further adjustment was made to cover two 1m- portant omissions in coverage: e not all suitable areas within active CBCs were surveyed, and ¢ not all HoSp in the areas surveyed were detected. Those factors could not be assessed rigorously, but “informed guesses” from personal experience (see Appendix) allowed tentative adjustment of extrapo- lated estimates. Vol. 120 Results HoSp counts (adjusted data) for each selected circle in the various habitat-groupings were summarized as decade-means (1960-69, 1970-79, etc.; Table 1). Circles lacking data for >1 decade were excluded, as were those lacking 1 decade and >=3 years in a 2"! decade. Mean data were estimated for circles missing one decade, using ratios of HoSp counts in other circles in that and the next decade(s). Some general patterns were evident: (i) a HoSp decline (from an earlier “plateau’), most often starting in the 1980s, was apparent nearly everywhere, in all habitats, and whether raw or adjusted data were compared; (ii) the recent decline was less pronounced in cir- cles where farmlands were important, or at least equal in area to forests, than elsewhere; the recent decline began earlier in cities and large towns than elsewhere. Observer effort in and around cities and towns in 1960-1979 increased so much that increased detec- tion of HoSp then sometimes balanced or exceeded declines, which were less apparent in raw data vs. adjusted data. Area changes in major habitats were substantial, as follows: (a) Forests remained the largest land cover type in New Brunswick and Nova Scotia throughout; forest cover increased in Prince Edward Island, but that was minor in the overall picture. HoSp were never found in or near forests except in association with human settlements, so changes in forest area are ignored in further discussion. (b) Among farmlands, “improved lands” (= crops, pasture, fallow; the main farmland types used by HoSp, aside from buildings) decreased in all provinces, by 35% in Prince Edward Island (where those remain dominant land-uses), and by 55% in New Brunswick and Nova Scotia, between 1961 and 1996. Published statistics available did not distinguish “commercial” and “marginal” farms; most decline in New Brunswick and Nova Scotia was in marginal farms, on which before 1960 farming was only one of several sources of income. Land-use on commercial farms became more intensive in 1961-1996 (personal observations), in the Maritimes as elsewhere, but that was not obvi- ous from the published statistics. (c) Urban area statistics allowed few generalizations. Larger communities mostly increased in human pop- ulation and area, whereas many smaller ones decreased, some dwindling to scattered houses. Cultural changes within urban areas were many and widespread, involv- ing sweeping changes in commerce, transportation, heating, garbage disposal, tolerance of livestock and poultry, recreation, etc. [see review in Erskine and McManus 2005]. Correlations with single factors in urban change could not be made rigorously so did not warrant statistical tests. (ii1) 2006 ERSKINE: RECENT DECLINES OF HOUSE SPARROWS 45 Table 1. Comparison of CBC HoSp means over decades, by habitats: (a) Raw data, (b) Adjusted data. Missing data added from ratios (of all groups) to next decade. Circles with no HoSp ever omitted. (a) Raw data 1960-69 1970-79 1980-89 1990-99 2000-02 Forest areas 7 circles 439 530 450 117 49 Coastal settlements, no farming 11 circles 875 1306 1023 404 138 Farming & woodlands nearly equal 11 circles 2120 2482 1823 949 735 Major farming areas, larger communities 8 circles 3306 4596 4953 2695 1296 Cities 4 circles 4017 3569 1664 374 157 Grand totals: 10757 12483 9913 4539 2375 (b) Adjusted data 1960-69 1970-79 1980-89 1990-99 2000-02 Forest areas 7 circles 599 476 331 58 25 Coastal settlements, no farming 11 circles 596 442 211 101 37 Farming & woodlands near equal 11 circles 1255 927 672 216 160 Major farming areas, larger communities 8 circles 2130 1840 1153 500 240 Cities 4 circles 1568 1001 283 39 19 Grand totals: 6148 4686 2650 914 48] Extrapolated HoSp populations by decades, expand- ed from surveyed circles across suitable habitats, and adjusted for incomplete coverage and detection, are shown in Table 2. That suggested that since the 1970s total HoSp num- bers in the Maritimes decreased greatly, probably by three-quarters or more, with urban numbers — in New Brunswick and Prince Edward Island — diminishing to less than 10% of levels prevailing before 1980. Farm- land HoSp numbers still may be one-half to two-thirds of those present 30-40 years earlier — more likely less. They now account for the vast majority of all HoSp remaining in New Brunswick, Prince Edward Island, _ and northern and eastern parts of Nova Scotia, where | winter conditions and especially snow cover limit sur- vival at that season. Numbers in western and southern Nova Scotia declined to a lesser degree. _ Discussion Johnston and Klitz (pages 15-52 in Pinowski and Kendeigh 1977) stated “The house sparrow seems » clearly a product of interaction between sparrows and » sedentary, agricultural man... This is a long-term rela- » tionship, sufficient to allow fine-grain adaptation of the birds to man’s way.” There seems no reason to dispute that generalization, on the long-term, broad-scale stage _ for which it was stated, but regional evidence suggest- ed that “man’s way” in the Maritimes recently changed more rapidly than did HoSp adaptations to it. Generalization from CBC data. Rigorous conclu- sions regarding the recent HoSp decline in the Mar- itimes (or elsewhere) will not emerge from CBCs alone, but improved understanding may result when various kinds of information are combined. The scarci- ty of hard data on this species, anywhere in North America, is embarrassing. In the Maritimes and adja- cent regions, the only multi-year study found that pro- vided HoSp density indices directly was the one in Sackville, New Brunswick (Erskine 1992b) that stim- ulated this investigation. Knowledge of habitats and diet here relied on casual observations, as no planned studies had been made — and HoSp numbers in most areas now are too low for economical study. CBC data are poorly standardized, and adjusting effort using party-hours addressed only a small part of variation in CBC data (compare Dunn et al. 2005). Regional data have not been examined rigorously, but it was obvious that the few — mostly urban — circles deploying large numbers of observers detected far fewer HoSp relative to effort than were found on most counts with fewer observers. Thus, declines in large urban areas may have been less drastic than indicated by the counts adjusted by party-hours, but they were impres- sive: compare means of raw data for 1966-1970 vs. 46 THE CANADIAN FIELD-NATURALIST Vol. 120 Table 2. Expansion of counts in Table 1 (raw data) to total Maritimes population estimates (explanation in Appendix). Habitats Extrapolated estimates 1960s 1970s 1980s 1990s 2000s Forest+urban 1694 2045 Wiss 452 189 Coastal+urban 16944 25290 19809 7824 2673 (Farm=forest)+urban 55506 64980 47730 24846 19242 (Farm>forest)+urban 78520 109160 117630 64010 30780 Large urban 120510 107070 49920 11220 4710 farming 67013 87070 115752 71085 45020 urban 206161 221475 121072 37267 12574 Grand totals: 273174 308545 236824 108352 57594 1996-2000 at Fredericton (3379 vs. 53), Moncton (1046 vs. 142), Saint John (S501 vs. 74). Also, CBC data often were less adequate for HoSp than for most other bird species. When HoSp were abundant and ubiquitous in urban areas, some CBCs did not actually count them during the “census”, mere- ly inserting round-number estimates as reported totals. Where such obvious guesses recurred in several years, those counts were excluded. A more frequent problem was that parts of many urban areas, where HoSp were the most expectable birds, were ignored as unproduc- tive (of more interesting species!) until HoSp decline was far advanced. HoSp declines in urban areas. That the recent HoSp decline in the Maritimes began earlier in cities (except perhaps “Greater Halifax’) than less urbanized situations suggested that influences formerly encour- aging HoSp to associate with human settlements are now less strong. Changes since 1960 in the regional urban milieu that might have been expected to affect HoSp numbers included: Favourable influences (i) winter bird-feeding increased greatly since 1960; however, small grains — preferred by HoSp (Bent 1958, also personal observations) — are now much less often presented (Ruther- ford 1984), feeding stations now offering most- ly sunflower and thistle seeds, rather than oats, cracked corn, or “chicken scratch feed” mix- tures; (11) shelter in urban hedges and other shrubs in- creased, presumably in response to increased disposable income of home-owners; Negative influences (a) food from human garbage, livestock wastage and manure decreased greatly, by disappearance of live- stock, especially horses, and improved garbage dispos- al, these factors all related to increasing “urbanization”; (b) nesting and wintering shelter within buildings decreased, through removal of dilapidated structures and renovation of others — especially closing of open- ings (personal observations); (c) transmittal of pathogens (e.g. Salmonella) may have been aided by increased transportation between urban areas (Erskine 1980b; a HoSp die-off at Char- lottetown, Prince Edward Island, had been reported in the preceding winter); (d) avian predators in towns (especially Merlin, Falco columbarius, and Sharp-shinned Hawk, Accip- iter striatus) increased, with Merlins breeding in urban areas here since the 1980s; domestic cats, Felis catus, also increased. The recent major decline of urban HoSp in the Mar- itimes indicated that urban areas no longer provide a favourable environment for this species, in winter or summer or both. Complete disappearance of HoSp in winter in many smaller human settlements suggested a “threshold effect’; below some population level, which likely varies with winter severity and availability of suitable habitats nearby, production in urban HoSp may no longer balance winter losses in this resident species. HoSp declines in farmlands. HoSp decline in farm- ing areas seemed better correlated with decreases in farms with livestock than with total numbers of farms or areas of “improved farmland”. Loss of invertebrate food through pesticide use, as in England (D. Sum- mers-Smith, letter), seems plausible but is unstudied here; chemical treatments in regional agriculture are obvious especially in potato- and fruit-producing areas, less so in grain production (mostly oats). Much Mar- itimes farmland area is inaccessible to HoSp in win- ter owing to snow cover. Livestock disappeared com- pletely from most urban areas, and also from many former farms now used as rural residences or “hobby farms”. Horses are scarce generally, and cattle now are mostly found in large feed-lot operations, where larger birds (gulls, pigeons, corvids, starlings) often out-com- pete HoSp — though a few of the latter persist (per- sonal observations). Concentrations of HoSp persist in productive farming areas, even when disappearing from nearby towns, presumably because livestock operations now are mainly in better farming areas. From the viewpoint of agriculture (and general econ-_| omy), the Maritimes are a “backwater”. Agriculture here ranks far behind forestry, tourism, fisheries, and “gen- 2006 eral service” on economic and employment indices, despite the local importance of potatoes and orchards. Snow cover and related temperatures are major fac- tors limiting agriculture (and HoSp) here. Soils and geology underlie the picture, with limestone areas very limited. Agricultural changes since World War II in- volved abandonment of former marginal agricultural lands rather than intensified use of better farmlands. HoSp responses to habitat changes. All CBC _ habitat groupings used herein involved combinations of human settlements, of varying size, with nearby habitats, among which only farmlands were regularly used by HoSp. Human settlements formerly provided _HoSp with both sheltered nesting places and nearby _ feeding opportunities. After nesting, urban HoSp often | withdrew from built-up areas to glean waste grain in agricultural lands within a radius of 2-3 km, return- ing to towns and villages with the approach of winter | (personal observations; Summers-Smith 1963). Where | little farmland exists near urban settlements, in forest- | ed regions (e.g., Miramichi, New Brunswick) or with- in larger cities, urban HoSp presumably depended | year-round on feeding opportunities within the city. Cultural changes altered urban areas more, as well | as in different ways, than farmlands. Thus, HoSp in areas that combined extensive farming with urban settlements were (and are) less restricted for feeding ' than those relying year-round on urban areas. Recent | trends suggest that HoSp might disappear within a few | decades from most urban habitats in New Brunswick and Prince Edward Island, where snow-cover limits ' their use of non-urban habitats in winter. They may persist longer in “Greater Halifax”, and in some towns in farming areas of western Nova Scotia, where snow cover is less and winter temperatures more moderate. | Such changes presuppose continuation of climates | similar to recent decades; if “global warming” brings \ served numbers across potential HoSp habitat to regional populations (Table 2) involved much “in- _ formed guesswork”, as hard data do not exist for most | needed adjustments. Valuable perspective may emerge from such simple “modeling”, even though it lacks | rigour and precision. | Modeling allowed comparison with the only other ' independent estimate of Maritimes HoSp populations, _ derived from abundance indices in the Maritimes breed- ing bird atlas (Erskine 1992a) in 1986-1990. The Atlas ) estimates used a general algorithm (C. Field and P. _Payzant, Appendix D in Erskine 1992a) for extrapo- lation to areas unsampled or lacking abundance indices; _ that algorithm may not have been fully appropriate for _ HoSp, of which density varied greatly between habi- tats. The Atlas estimate of HoSp (140 000 pairs) was ERSKINE: RECENT DECLINES OF HOUSE SPARROWS 47 of a similar order to that for the same period (1980s) in Table 2 (ca. 240 000 individuals). The HoSp situation elsewhere. No systematic stud- ies of HoSp were available in other parts of Canada. CBC data there — tabulated and adjusted for party- hours as in the Maritimes (unpublished MSS) — re- vealed no such drastic HoSp declines as seen here in 1960-2003. Continuing availability of small grains in both rural and urban areas of the Prairie Provinces south of the boreal forest were well-correlated with the highest HoSp CBC counts on the continent, with no suggestion of declines unrelated to count efforts. British Colum- bia CBC data were extremely variable, with generally low numbers, as was to be expected from the varied geography and relative scarcity of arable farming. No long-run counts suggested continuing declines in HoSp there. Very few long-run counts were available in north- ern, deep-snow areas of western Canada, where most showed no HoSp at any time. Quebec experiences the largest average snowfalls in Canada outside the western mountains. Most of the few long-run (>20 years) CBCs in Quebec suggested some decline in recent decades, from “adjusted counts”, as reported also from independent surveys (1970-1991) by Cyr et Larivée (1995); those declines were less ex- treme than in the Maritimes. In Ontario, where CBCs are more numerous than elsewhere in Canada, HoSp (raw) numbers remained generally stable over the decades, except in a few long- run urban circles near the northern limits of human set- tlement. Adjusted data, however, suggested declines, starting in 1980s, on nearly half the long-run counts; Sudbury showed the largest proportional decline, and counts at Sault Ste. Marie and Thunder Bay, also north- ern areas with much snow cover — and despite both being ports involved in grain shipping from the Prairies, also declined. In the United Kingdom, where the HoSp situation has been followed for decades, especially by Denis Summers-Smith (1963, 2003, and letters), declines evidently are of a similar order to those found in the Maritimes. Although research on HoSp is limited there too, much detailed information is available on habitat relations of HoSp in the U.K. Summers-Smith’s con- clusions (op. cit.) regarding causation agree well with tentative inferences suggested here for the Maritimes — despite the near-absence of snow cover as a limit- ing factor in the U.K. The intensity of agriculture there is much more comparable to that in southern Ontario or the Prairie Provinces than to the Maritimes. At this time it seems likely that many different factors, some general and widespread, others applying only region- ally, are involved in recent declines of HoSp in the wider picture. A recent discussion of HoSp decline in North America (Holder 2003) drew on wide-ranging sur- veys, including BBS and CBC, but reached few con- 48 THE CANADIAN FIELD-NATURALIST clusions except that competition with recent House Finch populations had not affected HoSp numbers. Use by Holder of only CBCs reported to Audubon limited his understanding of the Maritimes situation. His expressed suggestion (page 65) that the HoSp sit- uation in Canada might parallel that in the U.K. — where very different climatic and economic regimes prevail — seems unlikely. Acknowledgments Thanks to the many observers and compilers of CBCs whose data were used in this study, and also to publishers and editors of magazines (named in text) in which those reports were presented. Denis Summers- Smith (U.K.) and Jenny De Laet (Belgium) comment- ed very helpfully on an earlier version of the MS. Reviewers for this journal provided much-needed en- couragement as well as helpful suggestions for im- provement of the text. Literature Cited Bent, A. C. 1958. Life histories of North American black- birds, orioles, tanagers, and allies. United States National Museum Bulletin 211 [pages 1-24; Dover reprint con- sulted]. Christie, D. S. 1979. Changes in Maritime (sic) bird popu- lations, 1878-1978. Journal of the New Brunswick Muse- um 1979: 132-146. Christie, D. S. (1985 and following years). [general comments in annual “Christmas Counts” articles] in New Brunswick Naturalist. Cyr, A., et J. Larivée. 1995. Atlas saisonnier des oiseaux du Québec. Les Presses de |’ Université de Sherbrooke et La Société de Loisir Ornithologique de |’’Estrie, inc. [pages 660-661]. Downes, C. M., and B. T. Collins. 2002. The Canadian Breed- ing Bird Survey, 1967-2000. Canadian Wildlife Service Progress Notes Number 219. 39 pages. Dunn, E. H., C. M. Francis, P. J. Blancher, S. R. Drennan, M.A. Howe, D. Lepage, C.S. Robbins, K. V. Rosenberg, J. R. Sauer, and K. G. Smith. 2005. Enhancing the sci- entific value of the Christmas Bird Count. Auk 122: 338- 346. Vol. 120 Erskine, A. J. 1992a. Atlas of breeding birds of the Maritime Provinces. Nova Scotia Museum and Nimbus Publishing Ltd. 270 pages. Erskine, A. J. 1992b. A ten-year winter bird count in Sack- ville, New Brunswick. Canadian Field-Naturalist 106: 499- 506. A. J., and R. McManus, Jr. 2005. Bird status changes — and changes in environment — in the Chignecto isthmus region of Atlantic Canada. Canadian Wildlife Service Technical Report Series Number 430 Atlantic Region. vit+167 pages. Erskine, J.S. 1968. Winter birds of Wolfville, Nova Scotia: 1948-1968. Nova Scotia Museum Occasional Paper Num- ber 7, Science Series (5). 15 pages. Godfrey, W. E. 1954. Birds of Prince Edward Island. Nation- al Museum Canada Bulletin 132: 155-213. Holder, M. 2003. The decline of the House Sparrow. Birders Journal 12(2): 61-66. Kendeigh, S. C., Editor. 1973. A symposium on the House Sparrow (Passer domesticus) and European Tree Sparrow (P. montanus) in North America. Ornithological Mono- graphs (A.O.U.) 14. 121 pages. Lowther, P. E., and C. L. Cink. 1992. House Sparrow. Jn the Birds of North America, number 12. Edited by A. Poole, P. Stettenheim, and F. Gill. Philadelphia: Academy of Natural Sciences; Washington, D.C.: American Ornithol- ogists’ Union. 20 pages. Pinowski, J., and S. C. Kendeigh. Editors. 1977. Granivorous birds in ecosystems. Their evolution, populations, energet- ics, adaptations, impact and control. International Biologi- cal Program 12. Cambridge University Press, Cambridge, U.K. 431 pages. Rutherford, L. A. 1984. Effects of winter feeding on bird populations in Halifax, Nova Scotia. Unpublished Master — of Environmental Studies thesis, Dalhousie University, Halifax, Nova Scotia. 125 pages. Squires, W. A. 1952. The birds of New Brunswick. Mono- graph Series Number 2. Saint John, New Brunswick Muse- _ um. 164 pages. Summers-Smith, J. D. 1963. The House Sparrow. New Nat- | uralist Monograph 19. Collins, London. 269 pages. Summers-Smith, J. D. 2003. The decline of the House Spar- row: a review. British Birds 96: 439-446. Tufts, R. W. 1962. The birds of Nova Scotia. Halifax, Nova || Scotia Museum. 481 pages. Received 14 February 2005 Accepted 13 September 2005 2006 ERSKINE: RECENT DECLINES OF HOUSE SPARROWS 49 Appendix (i) Factors, by habitat groupings, for expansion from long-run CBC circles to total areas in Maritimes. Note: Forest areas support very few or no HoSp, so those areas were ignored, except where CBCs had been done. Habitats Number long-run Potential CBC circles CBC circles + other potential habitat | Forest + urban 7 add 8 | Coastal + urban 11 add 60 | (Farm = forest) + urban 1] add 37 (Farm > forest) + urban 8 add 1] | Large urban 4 add 4 (ii) Expansion factors, by habitat groupings, for incomplete coverage of suitable habitats in circles and incomplete detec- tion of HoSp in areas covered. | Habitats Coverage Detection | Forest + urban 1.2 1.5 » Coastal + urban 2.0 eS (Farm = forest) + urban 3.0 2.0 (Farm > forest) + urban 5.0 2.0 ) Large urban 5.0 3.0 (iii) Segregation of HoSp numbers among habitats, where appropriate. Forest and coastal habitats omitted as not used by HoSp; thus only 3 & 4" categories above required segregation, with proportions varying over time. Decades 1960s 1970s 1980s 1990s 2000s | Farm:urban ratio 50:50 50:50 70:30 80:20 90:10 First Record of the Southern Red-Backed Vole, Clethrionomys gapperi, in Newfoundland: Implications for the Endangered Newfoundland Marten, Martes americana atrata BRIAN J. HEARN, ! JOHN T. NEVILLE,! WILLIAM J. CURRAN,! and DEAN P. SNow? ‘Natural Resources Canada, Canadian Forest Service — Atlantic Forestry Centre, P.O. Box 960, Corner Brook, Newfoundland A2H 6J3 Canada > College of the North Atlantic, Fish and Wildlife Program, P.O. Box 822, Corner Brook, Newfoundland A2H 6H6 Canada Hearn, Brian J., John T. Neville, William J. Curran, and Dean P. Snow. 2006. First record of the Southern Red-Backed Vole, Clethrionomys gapperi, in Newfoundland: implications for the endangered Newfoundland Marten, Martes americana atrata. Canadian Field Naturalist 120(1): 50-56. We report on the first capture of the Southern Red-backed Vole (Clethrionomys gapperi), the eleventh non-native terrestrial mammal established on the island of Newfoundland over the last 150 years. Red-backed Voles may have been accidentally introduced by unknown sources in pulpwood imports or may have been deliberately introduced in an attempt to augment the depauperate small mammal fauna as a vigilante recovery effort for the endangered Newfoundland Marten (Martes americana atrata). We anticipate significant utilization of the Red-backed Vole as prey by both Newfoundland Marten and Red Fox (Vulpes vulpes) with associated demographic responses within and between these species. Red-backed Voles will likely change habitat utilization patterns for the endemic subspecies of Meadow Vole, Microtus pennsylvanicus terraenovae. Key Words: Southern Red-backed Vole, Clethrionomys gapperi, Marten, Martes americana atra, introduced species, New- foundland. The island of Newfoundland has only 13 resident native species of terrestrial mammals (Bangs 1913; Cameron 1958: Dodds 1983; Table 1). A fourteenth species, the Newfoundland Wolf (Canis lupus beothu- cus) was extinct by the early 20" century (Allen and Barbour 1937), and one of the three native mustelids, the Newfoundland Marten (Martes americana atrata) is currently listed by the Committee on the Status of Endangered Wildlife in Canada as endangered (Lemon 1996*). As well, the indigenous community of terres- trial mammals in Newfoundland has a skewed compo- sition, with a disproportionate number of predators and few prey species; historically, only one microtine, the native subspecies of Meadow Vole (Microtus pennsyl- vanicus terraenovae) occurred on the island (Dodds 1983). Moreover, Cameron (1958) considered 10 of the 14 native species of terrestrial mammals as endemic subspecies; recent genetic analysis has confirmed the subspecies classification of the Newfoundland Marten (Kyle and Strobeck 2003). However, over the last 150 years, an additional nine species of mammals have been either intentionally or accidentally introduced to the island (Dodds 1983; Gould and Pruitt 1969; Northcott 1974*; Northcott et al. 1974; Payne 1976; Table 1). A tenth species, the Eastern Coyote (Canis latrans) be- came established on the island of Newfoundland in 1985, likely after crossing on sea ice from mainland Nova Scotia (Parker 1995). Herein, we report the first record of the eleventh non-native mammal established on the island of Newfoundland, the Southern Red- backed Vole (Clethrionomys gapperi), and discuss the circumstances and implications of this recent introduc- tion, particularly with reference to the Newfoundland Marten. Field Sites and Sampling Between 2 and 5 November 1999, as part of a larger study investigating the demography and ecology of the Newfoundland Marten, we indexed small mammal pop- ulations on four sites inside the Pine Marten Study Area (PMSA) in southwestern Newfoundland (48°37'N, 57°53'W). The PMSA is a 2078-km* wildlife reserve, that was created in 1973 by the Newfoundland and La- brador Wildlife Division, for the protection of the New- foundland Marten (Snyder and Bissonette 1987). Sites 1-3 were overmature (81+ years), Balsam Fir (Abies balsamea) stands, whereas Site 4 was a regenerating (10-15 yrs) Balsam Fir stand that regenerated after an outbreak of Spruce Budworm (Choristoneura fumifer- ana) and Hemlock Looper (Lambdina fiscellaria fiscel- laria) in 1987. Site 4 contained a mix of young Balsam Fir and early successional species, such as White Birch (Betula papyrifera) and Pin Cherry (Prunus pensylvani- ca), had a significant number of dead Balsam Fir snags, and had no canopy. Each site was trapped with 100 snap traps on 3 consecutive nights. Traps were bait- ed with peanut butter, and placed in pairs at 50 trapping stations, spaced 15 m apart, yielding a 735-m transect, following the methods previously used by Thompson and Curran (1995). All captures were recorded and col- lected for further analysis. Forty small mammals were collected on the four sites over 1200 trap nights: 8 Meadow Voles, three Deer Mice (Peromyscus maniculatus), 10 Masked Shrews (Sorex cinereus), and 19 Southern Red-backed Voles. How- 50 2006 TABLE |. Native and non-native ‘? terrestrial mammals on the island of Newfoundland prior to the discovery of the Southern Red-backed Vole!. Order Insectivora Chiroptera Family Soricidae Vespertilionidae Lagomorpha Leporidae Rodentia Cricetidae Scuridae Castoridae Muridae Carnivora Mustelidae Felidae Canidae Ursidae Arctiodactyla Cervidae Scientific Name (Authority) Sorex cinereus acadicus (Gilpin) Myotis lucifugus lucifugus (Le Conte) Myotis septentrionalis (van Zyll de Jong) Lepus arcticus bangsti (Rhoads) Lepus americanus struthopus (Bangs) Microtus pennsylvanicus terraenovae (Bangs) Peromyscus maniculatus (Wagner) Ondatra zibethicus obscurus (Bangs) Tamiasciurus hudsonicus ungavensis (Anderson) Tamias striatus lysteri (Richardson) Castor canadensis caecator (Bangs) Rattus norvegicus norvegicus (Berkenhout) Mus musculus domesticus (Rutty) Mustela erminea richardsonii (Bonaparte) Mustela vison (Schreber) Martes americana atrata (Bangs) Lontra canadensis degener (Bangs) Lynx lynx susolanus (Bangs) Vulpes vulpes deletrix (Bangs) Canis lupus beothucus? (Allen and Barbour) Canis latrans var. (Lawrence and Bossert) Ursus americanus hamiltoni (Cameron) Rangifer tarandus terraenovae (Gmelin) Alces alces americana (Clinton) HEARN, NEVILLE, CURRAN, AND SNOW: RED-BACKED VOLE IN NEWFOUNDLAND Common Name Masked Shrew * Little Brown Bat Northern Long-eared Bat Arctic Hare Snowshoe Hare * Meadow Vole Deer Mouse * Muskrat Red Squirrel ° Eastern Chipmunk ~ Beaver Norway Rat * House Mouse * Short-tailed Weasel Mink * Newfoundland Marten? Otter Canada Lynx Red Fox Eastern Timber Wolf Eastern Coyote ~ American Black Bear Woodland Caribou Moose 'Several other species are either seasonal visitors (e.g., Arctic Fox (Alopex lagopus), Polar Bears (Ursus maritimus)) or have been introduced to offshore islands (e.g., Bison (Bison bison), Bank Vole (Clethrionomys glareolus suecicus)). Adapted from Cameron (1958) and Dodds (1983). *Subspecies designation confirmed via recent genetic analysis (Kyle and Strobeck 2003). 3Extinct ever, Red-backed Voles were captured on only two of the four sites (Sites 3 and 4). The two sites where Red- backed Voles were captured were approximately 6 km apart, but on opposite sides of Little Grand Lake. Move- ment between these two sites would require an over- land dispersal of at least 11 km, suggesting that the spe- cies is well established in the area. At least one of the collected females had obvious placental implantation scars on the uterine horns, indicating recent breeding. Species identification was confirmed by pelage and dental characteristics (Banfield 1974). Discussion Sites 1—3 had previously been snap trapped for small mammals for 4 years, between 1990 and 1993 (Thomp- son and Curran 1995). Additionally, Site 4, as well as the general area encompassing all four of our sampled sites, was extensively live trapped from 1993 to 1997, as part of a larger study investigating Marten, small mammal communities, and forest structure (Adair 2003). Neither of these two previous trapping efforts captured Red- backed Voles, suggesting that the species had recently become established, most likely sometime during or after 1998. Moreover, a more extensive small mammal survey, conducted two weeks before this survey in the Red Indian Lake area, approximately 35 km east of our four trap- ping sites, using identical methods but involving 16 sites and >4900 trap-nights, captured 116 small mam- mals, but no Red-backed Voles. Autumn (October) small-mammal surveys in the Red Indian Lake area, repeated annually since 1999, have subsequently doc- umented the arrival (2001) and eastward dispersal of the Red-backed Vole in southwestern and south-central Newfoundland (B. J. Hearn, unpublished data). Two explanations for this introduction are possible. Red-backed Voles may have been accidentally intro- duced in pulpwood or pulp chips imported through the port of Stephenville, (approximately 38 km SW of our sampling sites). Alternatively, the species may have been deliberately introduced by unknown persons to increase prey diversity and abundance for the Newfoundland Marten. Because Red-backed Voles are a common prey item in the diet of American Marten elsewhere (Mar- tin 1994), this management option was debated during preparation of the Newfoundland Marten Recovery Plan (Forsey et al. 1995*), resulting in a divergence of opinions among local stakeholder groups. Although ultimately rejected by the Newfoundland Marten Re- covery Team as an ethical recovery action, the limited prey base and the ecological consequences of introduc- ing the Red-backed Vole were publicly discussed in 1998, during a two-day symposium on issues concern- Sy THE CANADIAN FIELD-NATURALIST ing the Newfoundland Marten. Given the coincidence in timing (ca. 1998) and spatial distribution of this dis- covery (inside the PMSA), we suggest that it is likely that Red-backed Voles were deliberately introduced. Regardless of the source of this introduction, the direct and indirect effects of this new species on com- munity structure in general, and the Newfoundland Marten in particular, warrant investigation. As an exam- ple, Cameron (1958) reported that Newfoundland Meadow Voles are less selective in their habitat require- ments than mainland Meadow Voles, and atypically, occupy forested habitats — likely due to the historical absence of Red-backed Voles (Cameron 1958; Clough 1964; Cameron 1965; Morris 1969; Folinsbee et al. 1973). Consequently, the introduction of the Red- backed Vole will probably result in a niche contraction for the endemic Meadow Vole, and we predict that Meadow Voles will be displaced from forested areas and restricted to their preferred habitat (1.e., moist meadow-like areas (Folinsbee et al. 1973)). Preliminary analysis of our 1999-2006 small-mammal trapping data from the RIL area is documenting an irrupting population of Red-backed Voles (>3000 Red-backed Voles captured since 2001), and an essential elimina- tion of Meadow Voles from forested sites where we had previously captured Meadow Voles. Consequently, the positive effect of increased prey diversity and biomass for the Newfoundland Marten will be offset, to some degree, by a decrease in distri- bution and densities of Meadow Voles, which have been a historically important prey species (Bateman 1986; Drew 1995; Gosse and Hearn 2005; Tucker 1988). Further, indirect effects of the Red-backed Vole on Newfoundland Marten could occur as a result of the impact of this species on local Red Fox (Vulpes vulpes) populations. Given the depauperate prey base in New- foundland, Red Fox and Marten display considerable dietary overlap, and Red Fox are the most important natural predator of Newfoundland Marten (B. J. Hearn, unpublished data). Consequently, Marten may be nega- tively affected if the introduction of Red-backed Voles has a positive effect on Red Fox demography, leading to increased intraguild predation on Marten (Lindstr6m et al. 1995) or increased Marten—Fox competition for food (Kurki et al. 1997; Lindstrém et al. 1994; Marc- strom et al. 1988). Pine Marten (Martes martes) popu- lations in Scandinavia increased in the 1980s, follow- ing a decline of Red Fox due to a sarcoptic mange epidemic, as a direct result (hypothesized) of reduced predation by Red Fox (Lindstrom et al. 1994; Helldin 1998) or reduced competition for Microtus prey (Storch et al. 1990). Although counterintuitive to con- ventional competition theory, increases in food re- sources (prey availability) can actually increase intra- guild predation by supporting increased populations of the superior competitor. Parallel circumstances have been described for other threatened or endangered car- nivores elsewhere (e.g., Coyote predation on Swift Vol. 120 Foxes (Vulpes macrotis) (Sovada et al. 1998), Coyote predation on San Joaquin Kit Foxes (Vulpes macrotis) (Ralls and White 1995; White and Garrott 1997; Cypher and Spencer 1998), and Lion (Panthera leo) and Spotted Hyena (Crocuta croenta) predation on African Wild Dogs (Lycaon pictus) (Creel 2001; Creel et al. 2001)). African Wild Dog densities are actually lowest where density of their major prey is highest, due to intraspecific predation (Mills and Gorman 1997). It is noteworthy that Marten live-trapping captures on the Northern Peninsula of Newfoundland increased in 2005 following a recent (2002-2004) rabies outbreak (Whit- ney 2004*) that virtually eliminated Red Fox from the local landscape (M. McGrath, personal communication). Changes in prey availability clearly affect densities of carnivores, improving demographic performance (1.e., increased reproductive output and survival (Fuller and Sievert 2001). Furthermore, decreased food re- sources have been associated with reduced population density or trapper success, enlarged home ranges, and lower reproductive performance of American Marten (Bulmer 1974; Fryxell et al. 1999; Poole and Graf 1986; Simon et al. 1999; Thompson and Colgan 1987). The reduced prey base for Marten in Newfoundland compared with that available on the mainland portion of the province (Labrador) has been suggested as a likely explanation for the difference in the demograph- ic health of these two populations (Bissonette et al. 1988*). By comparison, Labrador has 17 small mam- mal species, including the Red-backed Vole (Tucker 1988) and Marten populations appear capable of sup- porting a commercial harvest; 500-800 Marten were harvested annually between 1995 and 1998 (Simon et al. 1999). However, an increase in vole biomass may not lead to an immediate increase in Marten densities if territo- ry size in Newfoundland is adjusted to the lowest (or historical) level of prey availability. Such appears to be the case for European Badger (Meles meles) in Scot- land (Kruuk and Parrish 1982), Kit Foxes in California (White and Ralls 1993), and some Coyote populations in the western United States (Mills and Knowlton 1991). Interestingly, Payer (1999) in the largest Ameri- can Marten study reported to date (n > 140 marten ranges), found no difference in mean home-range area for American Marten in Maine between 1995-1998, despite a three-fold decline and recovery in small mammal populations. Thompson and Colgan (1987, 1990) suggested that, based on energetics, Marten in Ontario cannot survive exclusively on small rodents during late winter, and further, take small mammals opportunistically as they hunt for larger prey (i.e., Snowshoe Hare). Moreover, Gosse and Hearn (2005) reported that Snowshoe Hare provide >90% of the caloric intake of Newfoundland Marten in winter, the most energetically stressful period annually (Buskirk et al. 1988). As cautioned by Fuller and Sievert (2001), | the correlation between food and density-related demo- | 2006 HEARN, NEVILLE, CURRAN, AND SNOW: RED-BACKED VOLE IN NEWFOUNDLAND 53 59°0'0"W 58°30'0"W 58°0'0"W 49°0'0"N 48°30'0"'N * Trap Sites 48°0'0"'N =a kilometers 0 5 10 20 30 58°30'0"W 58°0'0"W 57°30'0"W 57°0'0"W 49°0'0"N 48°30'0"N 48°0'0"'N 57°30'0"W 57°0'0"W Ficure |. Map of southwestern Newfoundland showing location of Southern Red-backed Vole (Clethrionomys gapperi) cap- tures in 1999, and capture sites relative to other areas mentioned in text; A indicates location of first Southern Red- backed Vole capture in Red Indian Lake area in 2001. Inset map shows the location of areas relative to the island of Newfoundland. graphic parameters is statistically broad, and many site- specific factors (e.g., prey availability, habitat patchi- ness, habitat selection under foraging versus predation- risk trade-offs), influence home-range characteristics and carnivore density. Newfoundland Marten have been geographically and reproductively isolated from mainland Marten populations for the last 7000 years (South 1983), resulting in a genetically distinct subspecies amidst a much more genetically homogenous Canadian main- 54 land population (Kyle and Strobeck 2003). Additional- ly, Newfoundland Marten are large (males = 1254 g,n= 122; B. J. Hearn, unpublished data), and have extreme- ly large home-range requirements (males = 30.8 km?, n= 43; B. J. Hearn, unpublished data) compared with nearby mainland populations in Quebec (males = 937 g, n= 67; 74 km’, n = 40; Potvin and Breton 1997) and Maine (males = 776 g, n = 23, Katnik 1992; 4.04 km”, n = 96; Payer 1999). Collectively, these population characteristics suggest that the Newfoundland Marten is a product of a unique ecological setting and evolu- tionary selective factors operating on a geographically and reproductively isolated island population. Thus, it seems unlikely, given the historical ecological set- ting, that Newfoundland Marten spatial requirements evolved to access or select habitat (Hearn et al. 2005*) based on utilizing small mammals as prey. We predict that the establishment of the Red-backed Vole on the island of Newfoundland will cause direct and indirect community-level effects in general and for Newfoundland Marten in particular (e.g., increased uti- lization of the Red-backed Vole as prey by Newfound- land Marten and Red Fox and associated demographic responses and interactions, more restricted habitat uti- lization by Newfoundland Meadow Voles, and chang- ing utilization and dispersal of seeds by introduced Red-backed Voles and consequently changing plant regeneration patterns). The direct and indirect effects of previous introductions of non-native species to the island of Newfoundland have been previously described for predator—prey interactions (Bergerud 1971, 1983), plant successional patterns (Thompson et al. 1992, Thompson and Curran 1993), and species interactions (Benkman 1993). Additional small mammal field surveys are being conducted to document the changing distribution of Red-backed Voles on the island and may offer addi- tional information about the mechanism by which this species was introduced. In this regard, genetic analysis may be helpful in determining the source population for this introduction. Future field studies, in areas where Newfoundland Marten spatial characteristics have been well documented (Hearn et al. 2005*) are being con- sidered to assess the effects of this new prey species on Newfoundland Marten population characteristics. Acknowledgments Funding for this work was provided by Natural Resources Canada, Canadian Forest Service; the New- foundland and Labrador Department of Environment and Conservation — Inland Fish and Wildlife Division; Newfoundland and Labrador Department of Forest Resources and Agrifoods — Forestry Division; Corner Brook Pulp and Paper; Abitibi Consolidated; and the Western Newfoundland Model Forest through support of a five-year research project concerning Newfound- land Marten. We thank D. J. Harrison, B. Krohn, J. Loo, and two anonymous reviewers for comments on this manuscript. THE CANADIAN FIELD-NATURALIST Vol. 120 Documents Cited (marked * in text) Bissonette, J. A., R. J. Frederickson, and B. J. Tucker. 1988. The effects of forest harvesting on Marten and small mammals in western Newfoundland. Report prepared for the Newfoundland and Labrador Wildlife Division and Corner Brook Pulp and Paper Limited, Utah Cooperative Fish & Wildlife Research Unit, Logan, Utah. Forsey, O., J. 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Forest Ecology and Management 47: 29-37. Vol. 120 Tucker, B. J. 1988. The effects of forest harvesting on small mammals in western Newfoundland and its significance to Marten. M.Sc. thesis. Utah State University, Logan, Utah. White, P. J., and R. A. Garrott. 1997. Factors regulating Kit Fox populations. Canadian Journal of Zoology 75: 1982— 1988. White, P. J., and K. Ralls. 1993. Reproduction and spacing patterns of Kit Foxes relative to change in prey availabil- ity. Journal of Wildlife Management 57: 861-867. Received 21 February 2005 Accepted 15 March 2006 Long-range Homing by an Adult Female Black Bear, Ursus americanus L. J. LANDRIAULT!, M. N. HALL?, J. HAMR?, and F. F. MALLoryY4 ‘Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada Present address: P.O. Box 521, Dowling, Ontario POM 1RO Canada 2Ontario Ministry of Natural Resources, Sudbury, Ontario P3E 1G7 Canada 3Cambrian College, Sudbury, Ontario P3A 3V8 Canada ‘Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada Landriault, L. J.,M.N. Hall, J. Hamr, and F. F. Mallory. 2006. Long-range homing by an adult female Black Bear, Ursus amer- icanus. Canadian Field-Naturalist 120(1): 57—60. An adult female Black Bear was repeatedly captured and relocated as a result of nuisance behaviour. The relocation distances ranged from 40 km to 389 km (mean = 152 km, n = 6). She homed successfully from all relocations, even when accompanied by young-of-the-year. Differential homing ability among bears may depend on first homing from a short relocation, facilitating subsequent responses to longer distance relocations. Key Words: Black Bear, Ursus americanus, homing, distance, relocation, translocation, Ontario. The ability of animals to return home when removed from familiar territory has been recorded for several different species including: pigeons, Columba spp. (Papi 1992); Deer Mice, Peromyscus maniculatus (Bovet 1968; Teferi and Millar 1993); Red Squirrels, Tamias- ciurus hudsonicus (Bovet 1995); Raccoons, Procyon lotor (Belant 1992); Wolves, Canis lupus (Fritts et al. 1984); and Brown Bears, Ursus arctos (Miller and Ballard 1982). The use of relocation as a management tool for nuisance bears has provided researchers with the opportunity to study the homing behaviour of a large mammal. Several studies conducted on Black Bears (Ursus americanus) have determined that a high proportion of relocated bears return to the capture area (Harger 1970; Alt et al. 1977; Rutherglen and Herbi- son 1977; McArthur 1981; Rogers 1986a; Shull 1994), with adult bears more likely to return to the capture area than juveniles (Harger 1970; Rogers 1986a; and Landriault 1998). During a four-year study of relocat- ed nuisance Black Bears in Sudbury, Ontario (46°N, 81°W), one bear was successful in homing over an exceptionally long distance (Landriault 1998). This report describes her relocation and homing history, and briefly discusses factors that may affect homing suc- cess in relocated bears. Animals were captured in large barrel traps or with the use of immobilization agents delivered via a dart gun or jab stick. In accordance with the study protocol, nuisance Black Bears were immobilized using a mix- ture of ketamine hydrochloride (Ketaset®) and xylazine hydrochloride (Rompun®) at a dosage of 4.4 mg/kg and ae mg/kg, respectively. Captured bears were weighed, various body measurements obtained, and the animals were ear-tagged (Kurl-lock metal tags, Ketchum Man- ufacturing Inc, Brockville, Ontario). A premolar was extracted for ageing purposes. Some animals were fitted with a very high frequency (VHF) radio-transmitter collar (148 to 152 MHz range, Lotek Inc., Newmarket, Ontario). If re-captured, most bears were immobilized again to assess changes in condition by means of growth rates for juveniles and changes in weight for adults. The majority of capture and release locations were estimat- ed using topographical maps or a hand-held global positioning system (GPS) unit; however, the only infor- mation available for the first and third captures des- cribed in this note was the township name. In these cases the center of the township was used to estimate release and homing distances. In the study area, town- ships were 10 x 10 km such that the center of the town- ship could only be a maximum of 7 km from the true location. Distances presented in this report are straight- line measurements and do not take topographical fea- tures into consideration. Returning within 20 km of the capture site was defined as successful homing (Lan- driault 1998). Adult female Black Bear LRO4 was captured as a nuisance animal and relocated a total of 6 times (Table | and Figure 1). In all cases she was captured because she was foraging in improperly stored refuse at resi- dences in suburban areas. She was initially captured as a nuisance animal in June of 1994. At this time, she was estimated to be 9 years of age based on the examination of premolar cementum annuli (Johnston et al. 1987). She was ear-tagged and relocated approx- imately 40 km south of the capture site. LRO4 was recaptured in June of 1995, an estimated 6 km from her first capture location. She was lactating; however, no cubs were found. She was fitted with a radio-collar and relocated a second time, 105 km to the north. LRO4 was subsequently recaptured in September of 1995, 7 km from her second capture location, and was accompanied by three cubs. The four bears were relo- cated as a family unit, approximately 112 km. LRO4 was captured again in June of 1996, within 17 km of her previous capture location, accompanied by three yearlings. LRO4 and two of the yearlings were relo- SY) 58 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Relocation history of nuisance Black Bear LR04, including relocation distance, bearing, and proximity of consecu- tive capture and release sites. All capture locations are in the suburban areas of the City of Sudbury, Ontario. Body weight information has been included as a means to assess the general condition of the animal. Capture Body Proximity of date weight capture site to (kg) previous capture site (km) 20 June 1994 65 14 June 1995 76 6 18 September 1995 147 7 8 June 1996 67 13 5 July 1996 82 10 1 October 1997 140 <1 cated 169 km. LRO4 was subsequently captured a fifth time in July of 1996, within 11 km of her previous cap- ture site. She was not accompanied by any offspring. On this fifth relocation she was transported 389 km to the northwest and was recaptured in October of 1997, within 7 km of her previous capture location. At this time she weighed approximately 140 kg and was re- located 100 km. From 1994 to 1997, adult female LRO4 was relo- cated a total of six times, with relocation distances ranging from 40 to 389 km. She homed successfully in all cases and the maximum distance between any two capture sites was 10 km. The longest homing dis- tance observed for this animal (389 km) appears to be a record distance for Black Bears. The true distance travelled by bear LR04 to return to her home range is expected to be much greater than 389 km, due to vari- ations in topography and the probable deviations from straight-line homing as a result of landscape features, human habitation, foraging, and interactions with other animals. Other observations of long-range homing in relocated Black Bears include animals returning from distances of 99 km (Rutherglen and Herbison 1977) and 229 km (Harger 1970) on the Upper Peninsula in Michigan, 103 km in the Rocky Mountains of British Columbia (Erickson and Petrides 1964), and 182 km in Central Ontario (Landriault 1998). Although all of these observations are from adult animals, in some in- stances juveniles can home from long distances. As part of the same study, a juvenile female homed from a relocation distance of 92 km (Landriault 1998). The differential homing ability of adults versus juveniles has been attributed to an established home range and to experience gained during seasonal foraging excursions (Anderson et al. 1977; Rogers 1986b, Rogers 1987a). Juvenile male Black Bears generally disperse be- tween 2 and 4 years of age, prior to establishing home ranges. In contrast, juvenile females generally take up residence in their maternal home range (Rogers 1987b; Schwartz and Franzmann 1992). It has been hypothe- sized that bears without established home ranges have limited homing success due to a lack of effort rather Proximity of Relocation release site to Bearing to distance previous release relocation (km) site (km) site 40 166° 105 141 B25 a i 6 825% 166 76 343° 389 222 330° 100 290 323% than ability (Anderson et al. 1977; Rogers 1986b). As predicted by this hypothesis, lower homing success has been observed in juvenile males than in adults and juvenile females (Harger 1970; Rogers 1986a; and Landriault 1998). Adult females would benefit from a strong desire to return to their established home range where they have been able to meet the nutritional re- quirements necessary for reproduction (Rogers 1976; LeCount 1982; Elowe and Dodge 1989; Kolenosky 1990) and have successfully avoided fatal conflicts for them and/or their young, with local conspecifics (Jonkel and Cowan 1971; Tietje et al. 1986). Experience would also be expected to increase an animal’s homing success. Many Black Bears cover ex- tensive area during yearly foraging excursions (Rogers 1987a) and these foraging excursions may allow bears to develop orientation and homing skills, resulting in increased homing success with age. Many young bears may not have gained enough experience to home suc- cessfully from significant distances. It is unknown whether bear LRO4 would have homed from 389 km had she been relocated that dis- tance after her initial capture. It is possible that repeat- ed relocations result in training individual bears to home from long distances. Black Bears that home suc- cessfully after the first relocation are likely to be suc- cessful after all subsequent relocations of increasing | distance (Blanchard and Knight 1995; Landriault' 1998), suggesting that homing ability in bears is en-- hanced with experience. Homing pigeons are common-- ly trained by gradually increasing the displacement| distance from the home loft, and young birds displaced! too far early in their training do not home successfully; (Papi 1992). The successful return from a relatively) short distance appears to enhance homing effort and/ or skill, and increases the chance of success from longer distances. The desire for adults to return to an established home) range, combined with experience acquired in navigat- ing over long distances, appears to explain why adult! bears are more adept at homing. The inadvertent train-) ing of bear LR04 by gradually increasing the reloca-)) 2006 100 0 EE _ LANDRIAULT, HALL, HAMR, AND MALLORY: LONG-RANGE HOMING 59 © Lake Huron a1 | 100 Kilometers SIGURE 1. Capture and release locations for nuisance Black Bear LRO4. Black circles represent the capture locations and grey circles represent release locations. The thin black lines on the map correspond to the two-lane paved provincial highways along the relocation routes, while the shaded squares are cities and small towns along these routes. Irregu- lar shaded areas are lakes. ion distances may explain how she was able to home successfully over a longer distance than any previous- y reported for a Black Bear. Acknowledgments Funding for this research was provided by The Fed- sration of Ontario Naturalists, the Ontario Ministry of Natural Resources, the Ontario Federation of Anglers d Hunters, the Bear Alliance-Canada, Laurentian Jniversity, and Cambrian College. Literature Cited t, G. L., G. J. Matula, F. W. Alt, and J. S. Lindzey. 1977. Movements of translocated nuisance black bears of north- eastern Pennsylvania. Transactions of the Northeast Fish and Wildlife Conference 34: 119-126. derson, P. K., G. E. Heinsohn, P. H. Whitney, and J. P. Huang. 1977. Mus musculus and Peromyscus maniculatus: homing ability in relation to habitat utilization. Canadian Journal of Zoology 55: 169-182. Belant, J. L. 1992. Homing of relocated raccoons, Procyon lotor. Canadian Field-Naturalist 106: 382-384. Blanchard, B. M., and R. R. Knight. 1995. Biological con- sequences of relocating grizzly bears in the Yellowstone ecosystem. Journal of Wildlife Management 59: 560-565. Bovet, J. 1968. Trails of deer mice (Peromyscus maniculatus) traveling on the snow while homing. Journal of Mam- malogy 49: 713-725. Bovet, J. 1995. Homing in red squirrels (Jamiasciurus hud- sonicus): the importance of going straight. Ethology 101: 1-9. Elowe, K. D., and W. E. Dodge. 1989. Factors affecting black bear reproductive success and cub survival. Journal of Wild- life Management 53: 962-968. Erickson, A. W., and G. A. Petrides. 1964. Population struc- ture, movements, and mortality of tagged black bears in Michigan. Pages 46-67 in The black bear in Michigan. 60 THE CANADIAN FIELD-NATURALIST Edited by A. W. Erickson, J. Nellor, and G. A. Petrides. Michigan Agriculture Experiment Station Research Bul- letin 4. Fritts, S. H., W. J. Paul, and L. D. Mech. 1984. Movements of translocated wolves in Minnesota. Journal of Wildlife Management 48: 709-721. Harger, E. M. 1970. A study of homing behavior of black bears. M.A. thesis, Northern Michigan University, Mar- quette. 81 pages. Jonkel, C. J., and I. McT. Cowan. 1971. The black bear in the spruce-fir forest. Wildlife Monographs 27: 1-57. Johnston, D. H., D. G. Joachim, P. Bachmann, K. V. Kar- dong, R. E. A. Stewart, L. M. Dix, M. A. Strickland, and I. D. Watt. 1987. Aging furbearers using tooth structure and biomarkers. Pages 228-243 in Wild furbearer manage- ment and conservation in North America. Edited by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. Ontario Trappers Association, North Bay, Ontario, Canada. Kolenosky, G. B. 1990. Reproductive biology of black bears in east-central Ontario. International Conference for Bear Research and Management 8: 385-392. Landriault, L. J. 1998. Nuisance black bear (Ursus ameri- canus) behavior in central Ontario. M.Sc. thesis, Laurent- ian University, Sudbury, Ontario, Canada. 95 pages. LeCount, A. L. 1982. Characteristics of a central Arizona black bear population. Journal of Wildlife Management 46: 861-868. McArthur, K. L. 1981. Factors contributing to the effective- ness of black bear transplants. Journal of Wildlife Manage- ment 45: 102-110. Miller, S. D., and W. B. Ballard. 1982. Homing of transplant- ed Alaskan brown bears. Journal of Wildlife Management 46: 869-876. Papi, F. 1992. Animal homing. Chapman and Hall Publish- ing, London. 390 pages. Vol. 120 Rogers, L. 1976. Effects of mast and berry crop failures on survival, growth, and reproductive success of black bears. Forty-First North American Wildlife Conference, pages 431-438. Rogers, L. L. 1986a. Homing by radio-collared black bears, Ursus americanus, in Minnesota. Canadian Field-Naturalist 100: 350-353. Rogers, L. L. 1986b. Effects of translocation distance on frequency of return by adult black bears. Wildlife Society Bulletin 14: 76-80. Rogers, L. L. 1987a. Navigation by adult black bears. Jour- nal of Mammalogy 68: 185-188. Rogers, L. L. 1987b. Effect of food supply and kinship on social behaviour, movements, and population growth of black bears in northeastern Minnesota. Wildlife Mono- graphs 97: 1-72. Rutherglen, R. A., and B. Herbison. 1977. Movements of nuisance black bears (Ursus americanus) in southeastern British Columbia. Canadian Field-Naturalist 91: 419-422. Schwartz, C. C., and A. W. Franzmann. 1992. Dispersal and survival of subadult black bears from the Kenai Peninsula, Alaska. Journal of Wildlife Management 56: 426-431. Shull, S. D. 1994. Management of nuisance black bears (Ursus americanus) in the Interior Highlands of Arkansas. M.S. thesis, University of Arkansas, Fayetteville. 101 pages. Teferi, T., and J. S. Millar. 1993. Long distance homing by the deer mouse, Peromyscus maniculatus. Canadian Field- Naturalist 107: 109-111. Tietje, W. D., B. O. Pelchat, and R. L. Ruff. 1986. Cannibal- ism of denned black bears. Journal of Mammalogy 67: 762-766. Received 23 February 2005 Accepted 26 September 2005 Early Ontogenetic Diet in Gray Wolves, Canis lupus, of Coastal British Columbia HEATHER M. BryAN!, Curis T. DARIMONT! 4, THOMAS E. REIMCHEN!, and PauL C. PagueT?= 'Department of Biology, University of Victoria, P.O. Box 3020, Station CSC, Victoria, British Columbia V8W 3N5 Canada *Raincoast Conservation Society, P.O. Box 26, Bella Bella, British Columbia VOT 1B0 Canada Faculty of Environmental Design, University of Calgary, Calgary, Alberta T2N 1N4 Canada ‘Corresponding author: cdarimon@uvic.ca Bryan, Heather M., Chris T. Darimont, Thomas E. Reimchen, and Paul C. Paquet. 2006. Early ontogenetic diet in Gray Wolves, Canis lupus, of coastal British Columbia. Canadian Field-Naturalist 120(1): 61-66. Within populations, different age classes often consume dissimilar resources, and provisioning of juveniles by adults is one mechanism by which this can occur. Although the diet of Gray Wolves (Canis lupus) has been studied extensively, the diet of pups is largely unknown. We examined faeces deposited by altricial pups and adult providers during the first two months following birth at two den sites over two years on the central coast of British Columbia, Canada. Pups and adult wolves con- sumed similar species, and Black-tailed Deer (Odocoileus hemionus) constituted most of the diet for both age groups. Pup and adult diet, however, diverged. Specifically, adult deer occurred significantly less frequently in the diet of pups than in the diet of adult wolves, which suggests that adults selectively provisioned pups. We speculate that this may relate to adaptive strategies of adult wolves to provide their offspring with food of optimal nutritional value or reduced parasitic burden, and/or logistic factors associated with provisioning such as prey transportability and availability. Key Words: Gray Wolf, Canis lupus, Sitka Black-tailed Deer, Odocoileus hemionus, provisioning, pups, diet, British Columbia. Different age classes within a species often consume dissimilar resources, increasing population niche width and decreasing intraspecific competition (Polis 1984; Bolnick et al. 2003). Provisioning of young by adults is one mechanism whereby differences in diet between young and adults may occur (Markman et al. 2002). Foragers that select foods to optimize growth and development of their offspring may have a selective advantage over those that provision their young oppor- tunistically (Krebs and Avery 1984; Wright et al. 1998). Providers may select resources that are easy to trans- port (Molsher et al. 2000) or that have nutritional qual- ities important for developing juveniles (Krebs and Avery 1984). Furthermore, providers may alter their behaviour while rearing young to reduce exposure of their offspring to parasites (Christe et al. 1994; Mer- ‘ila and Allander 1995; Tripet et al. 2002). ~ Young Gray Wolves (Canis lupus) rely predomi- nantly on food killed and delivered by closely related adult providers for many months (Paquet and Carbyn 2003). Although diets of adult wolves have been stud- ied extensively, the diet of juvenile wolves has not been well documented in any system (Paquet and Carbyn 2003). _ Wolves of coastal British Columbia (BC) have a potential niche that includes up to 15 mammal species (Darimont and Paquet 2000*, 2002; Darimont et al. 2004), which likely differ in transportability, nutritional characteristics, and parasite load. Herein, we investi- gated potential age class differences in diet by exam- ining faeces of pups and adults deposited at den sites ‘within the first two months following birth for two social groups over two breeding seasons. Our objectives were to describe the diet of adult and juvenile wolves during the denning season and examine whether the diet of pups differed from that of adults. Given the ubig- uity of evidence from a variety of other systems, we pre- dicted that adult wolves would selectively provision food resources to developing pups. Study Area We collected faeces from two home sites (i.e., repro- ductive areas) on the central coast of British Colum- bia in July of 2001 and 2002. The study area has been described well elsewhere (Darimont and Paquet 2002; Darimont et al. 2004). One site was located on Chat- field Island (area: 48 km?; distance from mainland: 1 km; centre island location: 52°16'N, 128°05'W) and the other on Yeo Island (98 km?; 0.25 km; 52°21'N, 128°08'W). Both sites were <100 m ASL (above sea level). During the study, social groups were composed of 3-7 pups and 3-8 adults. Potential prey species in- clude Sitka Black-tailed Deer (Odocoileus hemionus sitkensis), mustelids, ursids, birds, rodents, fish, and marine mammals and invertebrates (Darimont and Paquet 2000*, Darimont et al. 2004). Materials and Methods Scat collection and analysis Both sites were on or within 50 m from transects monitored for faeces during spring (April/May) before arrival of wolves or birth of pups, and summer (July) after wolves had left the areas. Thus, we are confident that faeces collected during July represent resources 61 62 acquired during May and June, at which time pups were approximately 2-8 weeks old. This period also corresponds to the birth pulse of many prey in the area, including deer (Shackleton 1999). In all cases, wolves had moved to another site by July, so we sampled all or most faeces that surround- ed the abandoned sites (within ~100 m). Based on diameter, we subjectively classified faeces (n = 479) as those deposited by pups or adults. In July, pup fae- ces are easy to distinguish from those of adults and there is no overlap in diameter (Weaver and Fritts 1979). The few faeces that were mostly liquid or suggestive of severe weathering (i.e., amorphous shape or irreg- ular surface; n ~ 20), which we felt could not be accu- rately classified, were discarded. Faeces were autoclaved, washed, and dried so that only macroscopic components such as hair, bone, teeth, and hooves remained (Cuicci et al. 1996). To identify mammalian prey items, we examined the entire sam- ple with the naked eye to detect hair clumps that dif- fered in colour, size and/or texture from others, and other clues such as teeth, claws and hooves. Subse- quently, we microscopically examined five sub-sam- ples, randomly selected via a grid system in the sample tray, which typically contained five to 10 hairs each. We identified species by comparing with a reference hair collection, as well as cuticular scale imprints using dissecting and compound microscopes and dichoto- mous keys (Mayer 1952; Adorjan and Kolenasky 1969). We differentiated between adult deer and fawn hair using a combination of diameter and colour charac- teristics (Scott and Shackleton 1979). Birds and inver- tebrates were identified by feathers, claws, shell frag- ments, and cuticles and were scored as present or absent. Using an 8 x 10 grid of 1-cm squares, we esti- mated the percentage of prey species in each faeces. All faeces were analyzed by one observer (HB) to eliminate inter-observer bias (Spaulding et al. 2000). Before faecal analysis, we tested observer accuracy with a reference collection and scored 92% (23 out of 25 correct). Following identification of prey remains, we also assessed precision and scored 100% (25/25) on classification of taxa. Finally, we investigated our precision in assessing adult/fawn categorization. We classified 77% (17/22) correctly. This bias, however, was directional: we scored five adults incorrectly as fawns. Data analysis We used two indices in analysis. Percentage of whole scat equivalents (pWSE) is the mean percentage of each prey species found per faeces (Angerbjorn et al. 1999; Elmhagen et al. 2002). For example, two fae- ces containing deer and seal in percentages of 40,60 and 60.40 respectively, would have pWSE of 50 for both prey species. Percent occurrence per item (O/T) is the number of occurrences of each prey species divid- ed by the total number of items identified, where an THE CANADIAN FIELD-NATURALIST Vol. 120 item is defined as an occurrence of a prey species in an individual faeces (Theberge et al. 1978). We used log-linear analysis on pWSE data to test for differences in diet between pups and adult wolves while simultaneously assessing influences of years and sites. We excluded birds and invertebrates from this analysis because conversion to pWSE was not possi- ble on presence/absence data. These items occurred infrequently and were never found alone in faeces. In this log-linear analysis, we also combined non-deer items into an “other” category due to infrequent occur- rence (see Results). To measure dietary breadth of both age classes, we used Levin’s Measure, standardized on a scale from 0 to 1 (Hurlbert 1978; Krebs 1999; Elmhagen et al. 2002). Levin’s measure of dietary breadth, B, is cal- culated using pWSE data as follows: [1] =| stato» where p,; is the proportion of resource 7 in a diet consisting of n resources. We also calculated dietary overlap between age classes using Schoener’s measure (p;,; Shoener 1970) and Horn’s index (R,; Horn 1966). Both calculations indicate food-niche overlap between populations j and k, where p;; and p;, are the proportions of resource 1 in diets, and n is the number of available resources. Although Shoener’s measure is frequently used and — therefore comparable to other literature, Horn’s index is less biased under changing numbers of resources, sample size, and resource evenness (Krebs 1999). Calculations were performed on pWSE data. [2] P; =| ¥ (minimum p,,, p, )/100 [3] 5 _ ©, + Pi) log (p; +p.) - Sp, logp , -Y pi. logp , 2 2log? Results Pup and adult wolves consumed a breadth of resources; however, Sitka Black-tailed Deer consti- tuted most of the diet for both age classes (Table 1). Deer represented 54.6 to 100% of whole scat equiva- lents, depending on site and year, and accounted for 50.0 to 98.4% of all prey items (Table 1). After deer, Harbour Seal (Phoca vitulina), invertebrates, and birds were the most common prey items, whereas Beaver (Castor canadensis), Short-tailed Weasel (Mustela ermina), Mink (Mustela vison), River Otter (Lutra can- adensis), Marten (Martes americana), and Fisher pups and adult wolves (Table 1). Pups and adults had | | | | (Martes pennati) occurred infrequently in the diets of © | 63 ES | WoL\ 2T IN aw De JET: EARL EN, AND PAQUET IMCHEN, T. REIM ONT, ARIM N,D BRYA 2006 srcent dict- 4 Average ae 719%. Tz ble «). , ylves ther (la It we hig / breadths and adu iderably his , 1e a « a cons similar foi pl ie ia tee ia 2). liet was ary overld Ps index of 0.96 (Ta and adult die oe a whereas eae shbitice stween pup aia of gape ed cl iia f erage die A ference : c age clas rtion of ¢ h Tia pa ajor diffe ’ different ropo “d wit Dein 1) A maj ion of di a lower p compare 1). ; Ds a ortic ned a fawn ; Table NE a nSm < he prop consun ion of 0.001; “ars 2 a oe ¥ (oe) fon n pe a << t ms Pups 2r proporti r=? Pc); across yea a1 £ ae BAe. inet 5s ZZ sumed. a higher p 7.77, df = 2 varied ac ze class 2 3 Za pone St | and a h Ne eee ion, age >. Epa it eilias we i | deer (Partial x f this differe en location, P=002: Mh Koa Liaal adults itude o a etwee =i,F= ; = Z S i “S = Hn | a chmcnens 36 oe “= Rete field Island = S Ame I The m ions (ass¢ artial Chatfi ord- 25 Bal ess 6 nn locations item; Part of seal on > 1). Acec a Loleas =] and lo and prey ite rtion of se (Table alysis, gy eu Se ine iia < > of wolf, ue high propo en g-linear ana volf, 5 ages Zz. 1). A hig 3-way as: m log- ass of w & A] é& ea] TAN venti | Table . d the 4 seal fro e class 0) 20. E icles es ee Spee Bim das 3 9 vt Salas in 200 we exc locati Partial x areas er are - 5 no) *"o. i) ingly, mee smo sigican Pan to other aller 42 3 inal dee sSOC as not : d, si ificantly sm S a — | he as ; Was foun , can d th RS s oon Lpelait ; rey item We then d a signifi ‘olves at bo Q» on | dp ). sume fe) 5 8 pape rec a P=0.65 consum id adult w 1) 5 = ~ © 5 ee i oa) re) ened df= 1, z that she than Fe 1, P<0.001). a ‘Ss = ae) ears, t =1, 5g E ~ Qn+e hone eon 7a dk propia etn 7 @ OD Va) ’ 10) ia F diet Eh pene aia a tee Gein ing wolf die E oq a Re lie thie loca ists regarding atterns 49 = ieee Ta) Pe . ture exists temporal E 2003) 3 2 S atl iscussion of litera atial and Cuicei 2003 2 a i Sola Re << ye large body ddresses spa terson and variability. ©) lap iu er te A lars ily a 3; Peters lation >on- a3 y Q i oI ZZ it primar n 2003; ee SUN IESE ° = | titp Carby f intrap ists 1 es sie 5 ere + bu and ts o ion ex 0 studi =o} % ieee (Paquet ther aspec informatio’ clude two F aint SES Sale l not oth dietary ions in diet o 205 =: | + and little Exceptions i the berge Balk he ively ng. Ey aon rere = 2 a oy Relat ° ing young differen / areas ever. Bees a raising y nt earby how eo ©°o xt of enifica Ean a study, e. gO iS B= |RES lac is t found sig den sites 6). Neither s s we do he > e va} mat nN . ~ 225 alo aed ien Za aha s between hko 198 adult diet as olves from vO E as) oq) Yori wolve 78: Meles pup and d adult w the prey 5 s = < QMlho mS et al. 19) d between juvenile an iches at h and 285 eae bie differentiate how that ju retary breadth an 3 ow Il 1 1 in. we § have s : dieta ) wolves 2 iy 5'o Ton ere, we ds imilar adult ny ane 5 2 ras) if . astal islan have ae) show that hemselves ! So a) No) oO CO S they ults s “1, to t 5. At iD a) a N S Slaw en (erent tw ree level, a These res Ss similar for pups. ¢ 2es Ie) ae Para eCcl erlap. items s Ses ; differ = F SS = ~ a Teel Z. sp te ove ith food y spec diets d é Set odera with food pu EEy SES at idesice As g fe r~ SAS Saat een nting eis a show = as owe N55 = t hu < veo fein cation an vee ete See a eae 3 = Lone ner res ; nd adu 3 ontrolling ved tha a = v a} Va) x lo | a fi nN pups a isioning. Cc sults shov deer tha oR ron ee is S It 28 é |x tela co aa eeu lective prov log-linear re ion of adu Rese WS [i 7 MOISES: her ea T prop ced by LS S| 20 et sue Ss year nee pres Id be E eeae ot = = c S & S F ee = B |nw 08 29 CI Senaene by adults. detected eee a bias in No ao N = : i : : . h = § a AlRSS aia SNES AER inition ae The . — | jj 62° N. latitude) in the Taiga Shield ecozone. Records show Coyotes have been seen since the 1960s. Prior to 2000, evidence of Coyotes breeding in the Taiga Shield has been anecdotal. In 2000, a Coyote was repeatedly seen at the Yellowknife airport and in 2001, a pair of Coyotes was observed with two pups. Since then, Coyote pups have been observed annually at the airport and adult Coyotes are seen regularly within the city of Yellowknife, an urban island within the Taiga Shield ecozone. Unlike in most regions occupied by Coyotes, medium-sized prey are rarely seen. Recently, Coyotes have become a poten- tial hazard to aircraft at the Yellowknife airport. Although Coyotes appear to have established themselves within the city of Yel- lowknife, maintaining a presence beyond the urbanized area remains uncertain. Key Words: Coyote, Canis latrans, distribution, range extension, reproduction, Northwest Territories. Coyotes (Canis latrans) have been uncommon in the Northwest Territories (NWT) north of Great Slave Lake (north of the 62" parallel). Banfield (1974) and Voigt and Berg (1987) suggested Coyote distribution was limited to the Taiga Plain ecozone and did not extend beyond the northwest shore of Great Slave Lake where the Taiga Shield ecozone begins. The Taiga Shield ecozone is characterized by a patchwork of wetlands, forests, meadows, and shrublands situated on top of glaciated bedrock with numerous lakes and rock outcrops. Summers are short and winters are long and cold. Although sightings of Coyotes beyond cur- rent range limits may be expected, occurrences of re- production beyond the Taiga Plain and into the Taiga Mica would be less likely. Bekoff (1977) and Bekoff and Gese (2003) estimated the range of Coyotes to be further northward but their delineation appears based B the northern limit of trees (“tree-line”). This paper en on recorded sightings and breeding of Coyotes ‘in the Taiga Shield ecozone in northwestern Canada. lidentify current conflicts and management responses for Coyotes as their numbers have increased in a north- em urban area. IMethods | Irecorded and investigated Coyote sightings in the ellowknife area since summer 2001. I solicited sight- ‘ing and harvest information from wildlife officers, hunt- ‘ers, and trappers in the Northwest Territories and west- m Nunavut. I searched historical records for Coyotes ong fur harvest records, Wolf (C. /upus) bounty ‘teceipts, and annual reports. Carcasses of Coyotes ob- ained since 2001 were examined whenever possible. Results and Discussion The earliest recorded sighting of Coyotes in the Taiga Shield ecozone occurred in the Yellowknife area in the early 1960s. In winter 1974-1975, five Coyotes were observed in the Long Lake area near the Yellow- knife airport (J. Hordal, Hay River, Northwest Territo- ries, personal communication). Prior to 2000, three other sightings occurred outside (>40 km) of the Yellowknife area (Figure 1) and in the late 1960s, a trapper shot a Coyote on Stark Lake, east from Lutsel K’e (formerly Snowdrift) (A. Boucher, Lutsel K’e, Northwest Term- tories, personal communication). One Coyote was oc- casionally sighted at the Yellowknife airport in 2000 and in spring 2001, two adult Coyotes were often re- ported near the airport; two pups were observed later that summer, the first documented occurrence of Coy- otes breeding this far north in the Northwest Territories. There have been two oral reports of Coyotes at Kug- luktuk (67°49'N, 115°06'W), Nunavut, near the north- ern coast of mainland Canada. Although exact dates and written reports are absent, the sightings are note- worthy as they occurred on the tundra 40-60 kilome- tres beyond the tree-line. The first was by a hunter who shot a Coyote 70 km west of Kugluktuk around 1988. The second was circa 1997 when two hunters saw a Coyote southwest of Bloody Falls on the Coppermine River (A. Niptanatiak, Kugluktuk, Nunavut, personal communication). The presence of Coyotes at the Yellowknife airport is not surprising because the area is characterized by open grassy habitat in contrast to the surrounding boreal forest. Rodents, birds, hares, and other wildlife on the airport property serve as food. The availability of food 67 68 THE CANADIAN FIELD-NATURALIST scraps near the city of Yellowknife proper and at the city dump should contribute to a continued presence by Coyotes within the urbanized area. It is possible that the Coyotes observed near the Yellowknife airport in win- ter of 1974-1975 had bred in the area as three of the five Coyotes did not appear adult size (J. Hordal, personal communication). Coyotes in northern urban areas The existence and colonization of Coyotes in and around Yellowknife is of concern to residents and air- port authorities. Residents are primarily concerned for the safety of their pets. Airport authorities are concerned as Coyotes and their pups occupy the Yellowknife air- port property and represent an air safety risk. In 2002, 2003 and 2004, 2,7, and 4 pups were observed on the airstrip area respectively (S. Loutitt, Yellowknife air- port, personal communication). Coyotes have not been limited to the airport. Seven were observed together on Back Bay in Yellowknife on 9 December 2004 (J. Bastedo, Yellowknife, personal communication) repre- senting the largest number of Coyotes observed together within Yellowknife to date. In addition, a Coyote pup was found dead on 30 August 2004 (J. Bastedo, per- sonal communication) and another Coyote pup was road-killed on 29 October 2004, both within Yellow- knife city limits. Furthermore, on 21 October 2004, a Coyote pup was killed on the highway about 60 kilo- metres northwest of Yellowknife and may represent a separate litter (Figure 1). The origin of this newest incursion of Coyotes is unknown, although they may have followed the single highway corridor connecting Yellowknife to the south. Coyotes have been expanding their range elsewhere over recent decades and human-induced habitat ch- anges are likely contributory (Moore and Parker 1992; Chubbs and Phillips 2002; Chubbs and Phillips 2005). Factors that might discourage Coyotes from colonizing northward include morphology, behaviour, and repro- duction (Phillips 1982). Although not morphologically equipped for travel in areas with deep snow, Coyotes might overcome this limitation by using trails and areas with hard snow (Murray and Boutin 1991; Murray and Lariviere 2002). Wolves often kill coyotes where their ranges overlap (Krefting 1969; Paquet 1989; Crabtree 1998), further discouraging northern Coyote range expansion. Air-strike risk Coyotes at the Yellowknife airport present a public safety risk because of aborted take-off and landing attempts and potential for collisions causing property damage and personal injury (B. Webber, Yellowknife airport, personal communication). Although the airport perimeter is fenced, Coyotes can access the area through holes in the chain-link or by digging under- neath. Continual removal efforts would be required to be effective over the long-term. Vol. 120 Although Coyotes have the potential to cause signif- icant damage to small planes, those struck at southern airports are often hit by undercarriage and do little overall damage to the aircraft (Dolbeer et al. 2000; Cleary et al. 2004). The greater risk is from aborted take-off and landings (Dolbeer et al. 2000), as pilots tend to see Coyotes on runways before collisions occur. There is anecdotal evidence that Coyotes may dis- perse other wildlife at airports such as Sandhill Cranes (Grus canadensis), Red Fox (Vulpes vulpes), and deer (Odocoileus sp.), and thus reduce the overall risk of wildlife strikes by aircraft (R. Dolbeer, United States Department of Agriculture, personal communication; S. Loutitt, personal communication). Coyotes are known to predate on Red Fox in some southern regions (Voigt and Earle 1983; Sargeant et al. 1987; Harrison et al. 1989), although fox predation by Coyotes has not yet been reported at the Yellowknife airport. There are few site-specific wildlife studies that address relative abun- dance and spatial patterns of wildlife at airports (Hoff- man et al. 1996; Dolbeer et al. 2000). The dilemma at the Yellowknife airport is one of risk management. Air- port management staff believe co-existence is possible and have chosen not to eliminate Coyotes but to moni- tor their presence and reduce numbers when required. Coyote persistence Whether Coyotes remain in the Yellowknife region is uncertain, although they occupy similar habitat else- where (e.g., Alaska; Labrador). Coyotes expanded their range northwest to Alaska in the late 1800s by scav- enging along trails left by the gold rush (Gier 1975) and arrived in the Yukon by 1910 (Parker 1995). Coy- otes appeared in the Northwest Territories fur harvest in the early 1930s when records were first kept. Sup- pressants to early colonization of Coyotes in the Yel- lowknife region include hunting, trapping, predation by Wolves, and incidental mortality at poison baits during Wolf control in the early 1960s (Heard 1983; Cluff and Murray 1995). Wolves are occasionally observed at the Yellowknife airport, and in 2004 a pack of five were reported within city limits. To sustain a Coyote population a reasonable prey base must exist. Birds, Snowshoe Hare (Lepus ameri- canus) and other small mammals would be the main source of food for forest-dwelling Coyotes (Créte et al. 2001). White-tailed Deer (O. virginianus) are only rarely sighted in the Taiga Shield (unpublished data; Veitch 2001). Wood Bison (Bison bison athabascae) have expanded their range into the area since 1999 but this may adversely impact Coyotes if Wolf numbers also increase. Garbage and domesticated animals in the Yellowknife area would augment the available food supply. The Yellowknife city dump, electrified only in summer against Black Bears (Ursus americanus), is not a barrier to Coyotes. Cabins and homes scattered on the landscape outside of Yellowknife may also supply 2006 CLUFF: COYOTE BREEDING RANGE IN NORTHWEST TERRITORIES 69 Area of Detail \ asp Ficure |. Taiga Plain and Taiga Shield ecozones bordering Great Slave Lake in the Northwest Territories showing records of Coyotes since the 1960s documented in text. Circles represent sites prior to 2000, squares in 2003 and 2004. food. Consequently, Coyote persistence in Taiga Shield of the Northwest Territories may be strongly linked to a suburban environment, especially if an inadequate prey base exists elsewhere. How the public and wildlife management authorities choose to deal with Coyotes as urban carnivores will be a key factor for Coyote persist- ence in the Yellowknife region. Acknowledgments I thank the trappers, wildlife officers, and biologists in the Northwest Territories and Nunavut for their assis- tance in documenting and mapping sightings of Coy- Otes in the two jurisdictions. I also appreciate the efforts of airport management staff to document their sight- ings of Coyotes and to facilitate logistics of my investi- gations. E. Campbell was especially helpful in referring me to trappers and providing earlier personal sightings of Coyotes. Comments from A. Gunn, G. Elliott, M. Créte, and two anonymous referees helped improve an earlier version of the manuscript. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. University of Toronto Press, Toronto, Ontario. Bekoff, M. 1977. Canis latrans. Mammalian Species Number 79. Bekoff, M., and E. M. Gese. 2003. Coyote (Canis /atrans). Pages 467-481 in Wild mammals of North America: Biolo- gy, management and conservation. 2™ edition. Edited by G. A. Feldhamer, B. C. Thompson and J. A. Chapman. Johns Hopkins University Press, Baltimore, Maryland. Chubbs, T. E., and F. R. Phillips. 2002. First record of an eastern Coyote, Canis latrans, in Labrador. Canadian Field- Naturalist 116: 127-129. Chubbs, T. E., and F. R. Phillips. 2005. Evidence of range expansion of eastern Coyotes, Canis latrans, in Labrador. Canadian Field-Naturalist 119(3): 381-384. Cleary, E. C., R.A. Dolbeer, and S. E. Wright. 2004. Wildlife strikes to civil aircraft in the United States 1990-2003. Fed- eral Aviation Administration National Wildlife Strike Data- base Serial Report Number 10. Report of the Associate Administrator of Airports, Office of Airport Safety and Stan- dards, Washington, D.C. 70 THE CANADIAN FIELD-NATURALIST Cluff, H. D., and D. L. Murray. 1995. Review of wolf control methods in North America. Pages 491-504 in Ecology and conservation of wolves in a changing world. Edited by L.N. Carbyn, S. H. Fritts, and D. R. Seip. Canadian Circumpolar Institute, University of Alberta, Edmonton, Alberta. Crabtree, R. L. 1998. The total impact. The Yellowstone wolf tracker 1: 14-17. Créte, M., J. P. Ouellet, J. P. Tremblay, and R. Arsenault. 2001. Suitability of the forest landscape for coyotes in north- eastern North America and its implications for coexistence with other carnivores. Ecoscience 8: 311-319. Dolbeer, R. A., S. E. Wright, and E. C. Cleary. 2000. Ranking the hazard level of wildlife species to aviation. Wildlife Society Bulletin 28: 372-378. Gier, H. T. 1975. Ecology and behaviour of the coyote (Canis latrans). Pages 247-262 in The wild canids. Edited by M. W. Fox. Van Nostrand Reinhold, New York, New York. Harrison, D. J., J. A. Sherburne, and J. A. Bissonette. 1989. Spatial relationships between coyotes and red fox in eastern Maine. Journal of Wildlife Management 53: 181-185. Heard, D.C. 1983. Historical and present status of wolves in the Northwest Territories. Pages 44-47 in Wolves in Canada and Alaska; their status, biology, and management. Edited by L. N. Carbyn. Report Series Number 45, Canadian Wild- life Service, Edmonton, Alberta. Hoffman, T. R., R. F. Krischke, and J. A Brent. 1996. A bio- logical assessment of Portland International Airport with rec- ommendations to alleviate wildlife hazards to aircraft opera- tions. United States Department of Agriculture — Port of Portland, Portland, Oregon. Krefting, L. W. 1969. The rise and fall of the coyote on Isle Royale. Naturalist (Leeds) 20: 24-32. Moore, G. C., and G. R. Parker. 1992. Colonization by the eastern coyote (Canis latrans). Pages 23-37 in Ecology and management of the eastern coyote. Edited by A. H. Boer. Vol. 120 Wildlife Research Unit, University of New Brunswick, Fredericton, New Brunswick. Murray, D. L., and S. Boutin. 1991. The influence of snow on lynx and coyote movements: does morphology affect behaviour? Oecologia 88: 463-469. Murray, D. L., and S. Lariviére. 2002. The relationship between foot size of wild canids and regional snow condi- tions: evidence for selection against a high footload? Journal of Zoology (London) 256: 289-299. Paquet, P. C. 1989. Behavioral ecology of sympatric wolves (Canis lupus) and coyotes (C. latrans) in Riding Mountain National Park, Manitoba. Dissertation, University of Alberta, Edmonton, Alberta. Parker, G. 1995. Eastern coyote. The story of its success. Nim- bus Publishing Ltd., Halifax, Nova Scotia. 256 pages. Phillips, M. K. 1982. Factors influencing coyote distribution. Page 51 in Proceedings of the 33"! Alaska Science Confer- ence, Science in the North, 16-18 September 1982, Fair- banks, Alaska. Sargeant, A. B., S. H. Allen, and J. O. Hastings. 1987. Spatial relations between sympatric coyotes and red foxes in North Dakota. Journal of Wildlife Management 51: 285-293. Veitch, A. M. 2001. An unusual record of a White-tailed deer (Odocoileus virginianus) in the Northwest Territories. Cana- dian Field-Naturalist 115: 172-175. Voigt, D. R., and W. E. Berg. 1987. Coyote. Pages 345-357 in Wild furbearer management and conservation in North America. Edited by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. Ontario Trappers Association, North Bay, Ontario. Voigt, D. R., and B. D. Earle. 1983. Avoidance of coyotes by red fox families. Journal of Wildlife Management 47: 852-857. Received 7 March 2005 Accepted 2 November 2005 Lake Sturgeon, Acipenser fulvescens, Movements in Rainy Lake, Minnesota and Ontario W. EUGENE ADAMS, Jr.!*, LARRY W. KALLEMEYN?, and Davip W. WILLIs!4 'Department of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, South Dakota 57007 USA *B-mail: Geno.Adams@staff.sd.us 3U.S. Geological Survey, 3131 Hwy 53, International Falls, Minnesota, 56649, USA. E-mail: larry_kallemeyn@usgs.gov 4E-mail: David.Willis@sdstate.edu, corresponding author. Adams, W. Eugene, Jr., Larry W. Kallemeyn, and David W. Willis. 2006. Lake Sturgeon, Acipenser fulvescens, movements in Rainy Lake, Minnesota and Ontario. Canadian Field-Naturalist 120(1): 71-82. Rainy Lake, Minnesota-Ontario, contains a native population of Lake Sturgeon (Acipenser fulvescens) that has gone largely unstudied. The objective of this descriptive study was to summarize generalized Lake Sturgeon movement patterns through the use of biotelemetry. Telemetry data reinforced the high utilization of the Squirrel Falls geographic location by Lake Stur- geon, with 37% of the re-locations occurring in that area. Other spring aggregations occurred in areas associated with Kettle Falls, the Pipestone River, and the Rat River, which could indicate spawning activity. Movement of Lake Sturgeon between the Seine River and the South Arm of Rainy Lake indicates the likelihood of one integrated population on the east end of the South Arm. The lack of re-locations in the Seine River during the months of September and October may have been due to Lake Sturgeon moving into deeper water areas of the Seine River and out of the range of radio telemetry gear or simply moy- ing back into the South Arm. Due to the movements between Minnesota and Ontario, coordination of management efforts among provincial, state, and federal agencies will be important. Key Words: Lake Sturgeon, Acipenser fulvescens, biotelemetry, Rainy Lake, Ontario, Minnesota. Movement patterns of Lake Sturgeon (Acipenser ful- vescens) were relatively unstudied through the early 1970s (Scott and Crossman 1973). However, in the past decade, Lake Sturgeon movements have been studied at an increasing rate throughout much of the species’ range (Rusak and Mosindy 1997; McKinley et al. 1998; Auer 1999; Borkholder et al. 2002; Knights et al. 2002). Lake Sturgeon have been found to move large distances for purposes such as spawning (Threader and Brosseau 1986; Fortin et al. 1993; Rusak and Mosindy 1997; Auer 1999) while maintaining smaller ranges in other systems (Bassett 1982*). Such movement pat- ‘terns make Lake Sturgeon management difficult, par- ticularly for multiple agencies dealing with border waters of substantial size. Rainy Lake, a border water shared by Minnesota and Ontario, contains a Lake Sturgeon population that has gone largely unstudied. Lake Sturgeon in this system have been segregated from upstream Namakan Lake and downstream Lake of the Woods/Rainy River sys- i populations by dams constructed on the outlets /of Rainy and Namakan lakes early in the 20" century. |The Lake Sturgeon is a Minnesota state-listed species | of special concern (Minnesota Department of Natural i Resources 2003*), while The Committee on the Sta- tus of Endangered Wildlife in Canada considers west- em populations of Lake Sturgeon to be endangered is the Rainy River-Lake of the Woods populations to be of special concern (Environment Canada 2005*). | Management of the Lake Sturgeon population in Rainy Lake is a cooperative effort among the Ontario Ministry of Natural Resources, the Minnesota Depart- ment of Natural Resources, and the United States fed- eral government. Movement of Lake Sturgeon across international borders leads to an increase in impor- tance of appropriate harvest regulations and an under- standing of movement patterns and spawning locations. Such information will allow managers to protect areas that are important for Lake Sturgeon spawning. The objective of this descriptive study was to deter- mine generalized Lake Sturgeon movement patterns in Rainy Lake through the use of biotelemetry. Move- ment data obtained from tagged Lake Sturgeon were organized using a geographic information system (GIS) and used to help identify seasonal movement patterns and potential spawning locations. Study Site Rainy Lake, located on the Minnesota-Ontario bor- der, consists of three main basins: the North Arm, Redgut Bay, and the South Arm, all of which are part of the Winnipeg-Nelson drainage system in the Lake Winnipeg primary watershed (Figure 1). The North Arm and Redgut Bay are both located entirely in Cana- da. Rainy Lake has a total surface area of 92 000 ha, a maximum depth of 49.1 m, and a mean depth of 9.9 m. Of the three main lake basins, the South Arm has the greatest surface area at 49 200 ha, with 27 300 ha in Ontario and 21 900 ha in Minnesota. The South Arm is also the deepest basin with a mean depth of 9.9 m and a maximum depth of 49.1 m. The South Arm ex- tends for 56 km along the border of the United States TA 2 THE CANADIAN FIELD-NATURALIST and Canada. The South Arm of Rainy Lake, along with a 40-km section of the Seine River, was the area of focus for this study. The watershed associated with Rainy Lake encom- passes over 37 500 km? and can be divided into two sub-basins, including a 19 270-km? area above the outlet of Namakan Reservoir at Kettle Falls and Squir- rel Falls and a 19 320-km? area below Kettle Falls and Squirrel Falls that drains directly to Rainy Lake. Overflows between Namakan Reservoir and Rainy Lake also occur at Bear Portage and Gold Portage. Below Kettle Falls and Squirrel Falls there are two principal tributaries that enter Rainy Lake, the Turtle River with a mean discharge of 37 m*/sec and the Seine River with a mean discharge of 48 m?/sec. Smaller, ungauged tributaries into the South Arm of Rainy Lake include the Rat and Pipestone rivers, both of which may be used by Lake Sturgeon for spawning. Changes in flow initiated at the headwaters of the watershed take about 21 days to reach the outlet of Rainy Lake under normal flow conditions with the water drop- ping about 135 m in the 338 km between these two points (International Rainy Lake Board of Control/ International Lake of the Woods Control Board 1984*). Long-term flow records indicate that approximately 8.3 billion m? of water move through the Rainy Lake watershed annually (Ericson et al. 1976*). Methods Biotelemetry was utilized in the identification and characterization of Lake Sturgeon seasonal movement patterns and likely spawning locations in Rainy Lake. Lake Sturgeon were captured for transmitter implan- tation in October of 2002 (n = 13), May and June of 2003 (n = 20), and October of 2003 (n = 8) at the mouths of tributaries where they were known to concentrate (tagging locations are noted in Table 1). Large mesh multifilament gill nets with mesh sizes of 103, 114, 127, 152, and 178 mm (bar measure; i.e., distance from knot to adjacent knot) were used to capture fish. Net lengths varied from 30 to 91 m, with all nets having a height of 1.83 m. Most nets were fished for approxi- mately 24 hours, but actual times varied among sam- pling crews. Captured Lake Sturgeon were measured, weighed, sex was noted if gonads were identifiable during surgery, and a pectoral fin ray section was re- moved for aging (Adams et al. 2006). Lotek Wireless combined acoustic/radio transmit- ters (CART) were surgically implanted into 41 Lake Sturgeon (>8 kg) throughout the duration of this study. Thus, only a single tag was implanted and used for both radio and acoustic tracking, with the radio portion of the tag allowing tracking of fish in shallow or tur- bulent habitats, while the ultrasonic portion of the tag was needed for tracking of Lake Sturgeon in deeper habitats (Winter 1996). Surgical implantation proce- dures followed guidelines set by Hart and Summerfelt (1975). A 5- to 8-cm incision was made on the ventral Vol. 120 fish surface approximately 3 cm off the midline and approximately 3 cm from the anterior end of the pelvic girdle. A 0.5-cm exit hole for the whip antenna was started with a scalpel on the midline approximately 3 cm posterior to the incision. A curved, 12-gauge catheter needle was passed through the hole and out through the incision with care taken to not penetrate or cut the viscera. The whip antenna projecting from the CART tag was then threaded though the end of the catheter needle that was protruding through the incision with the antenna being passed out of the peritoneal cavity through the antenna hole. The CART tag was then inserted into the peritoneal cavity with minimal pressure exerted on the internal organs. Fish were placed in a holding pen following implantation, if nec- essary. When full equilibrium was regained, fish were placed in the water at the site of capture and monitored until they vacated the area. All fish handling and sur- gery followed guidelines for use of wild fishes estab- lished by Nickum et al. (2004). Lotek CART 16-2 series tags were implanted in 33 Lake Sturgeon with the remaining eight fish receiving CART 16-1 series tags. Lotek CART 16-2 tags were 16 mm X 85 mm with a weight in air of 36 g, while Lotek CART 16-1 tags were 16 mm Xx 60 mm with a weight in air of 25.3 g. Implanted CART tags did not exceed 2% of the total body weight of any given fish (Gallep and Magnuson 1972; McCleave and Stred 1975; Stasko and Pincock 1977; Moser et al. 1990). Aerial tracking of Lake Sturgeon occurred at least once per week throughout the spring spawning season and summer months when the airplane was opera- tional. Transects were flown covering the eastern half of the South Arm, with every flight including a 40-km stretch of the Seine River that flows into Rainy Lake. Transects were chosen to maximize area covered with the least amount of flight time. The remaining portion of the South Arm was flown at least once every two flights, time and weather permitting. The North Arm of Rainy Lake was flown once in the summer of 2003 in an attempt to locate a fish that could not be located in the South Arm. Radio tracking by boat took place on average two to three times per week depending upon weather con- ditions. Tracking was also dependent on time availabil- ity, with partial and full days devoted to boat tracking when possible. Locations highly used by Lake Stur- geon were sampled by boat during May and June be- cause recognition of individual signals was difficult from the airplane. Distributional patterns of Lake Sturgeon were sum- marized by month during the spawning period (May and June) and by two-month periods throughout the rest of the summer (July/August and September/Oc- tober). Movement rates were calculated as the linear distance between successive fixes divided by the num- | ber of elapsed days. Lake Sturgeon home ranges were | calculated using the Animal Movement extension in | ——————— = 2 ey a 2006 ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 73 Kilometers : Seine River IRS Pipestone River ——= 5a eo te | Kettle Falls ; Squirrel Falls’ Sok » 7 Canadian Channel Ca a’ x= — FIGURE 1. Rainy Lake, Minnesota and Ontario. The line across the South Arm indicates the border between Canada and the United States. _ ArcView 3.2 (Hooge et al. 1999*), with home ranges estimated by using minimum convex polygons (White and Garrett 1990). Home ranges included only water area; land masses were excluded. Lake Sturgeon eggs were sampled during 2004 in an attempt to confirm spawning locations. Egg col- lectors were set weekly from 7 June through 1 July in three areas of high Lake Sturgeon use. Collectors consisted of an 18-m mainline of vinyl-coated wire cable (4.8 x 6.4 mm) with cement blocks (203 x 406 x 102 mm) attached at 3-m intervals and wrapped in furnace filter material. Floats were attached to each end of the mainline and egg collectors were set in shallow water areas of 0.3 to 4.6-m depth. Egg collectors were checked once weekly during the entire time period in which they were deployed. A sample of all egg types present on the collectors was removed and transport- ed back to laboratory facilities to be hatched and the fry subsequently identified. Results and Discussion Lake Sturgeon tagging and survival Forty-one Lake Sturgeon were implanted with CART tags during 2002 and 2003 (Table 1). Individual Lake Sturgeon were re-located between 0 and 36 times. No mortalities were positively identified. However, fish 99, implanted in October of 2003, was not located during the remainder of the study. Fish 30 was not located for 12 months after it was tagged and released, but then was subsequently located 11 times throughout the re- mainder of the study. Fish 106 was found at the same location, adjacent to the Seine River First Nations vil- lage on the Seine River, weekly for 11 weeks. We later learned that fish 106 may have perished in a gill net used in the Seine River First Nations subsistence fish- ery that had been left on the bottom of the Seine River due to entanglement with the substrate. All other fish were located at least once throughout the study. Lake Sturgeon implanted with CART tags had a mean fork length of 1217.4 mm (range 951-1505 mm), girth of 502.4mm (range 372-639 mm), mass of 15.35 kg (range 7.5-30 kg), and age of 25.9 (range 15-55) (Table 1). These ranges were representative of the Lake Sturgeon population in Rainy Lake at the time of this study (Adams et al. 2006). Transmitter detection Acoustic equipment was tested during the spring of 2003. Tags were lowered at 2-m intervals between 74 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Fish description, transmitter code, and capture location at time of tagging for Lake Sturgeon implanted with Lotek CART transmitters in Rainy Lake, 2002-2003. Ages were determined from pectoral fin ray sections. NA = not available. Tagging Total Fork Code date length length Girth number Month/day/year (mm) (mm) (mm) 1 10/2/02 NA 1100 444 yD 5/21/03 1506 1433 635 3 10/9/02 NA 1210 556 4 10/9/02 NA 1275 615 5 10/9/02 NA 994 372 6 10/9/02 NA 1050 425 i] 10/9/02 NA 951 395 8 10/8/02 NA 1212 485 9 10/8/02 NA 1313 582 10 10/8/02 NA 1386 614 11 10/8/02 NA 1447 639 12 10/8/02 NA 1388 585 13 10/8/02 NA 1200 445 14 10/8/02 NA 1125 407 15 5/28/03 1372 1260 495 16 5/20/03 1400 1270 503 17 5/28/03 1404 1300 520 18 5/28/03 1410 1285 475 19 6/18/03 1270 1140 410 20 5/28/03 1200 1060 415 21 6/18/03 1425 1280 570 22 5/28/03 1505 1365 545 23 6/18/03 1120 1000 460 24 5/21/03 1460 1315 578 25 5/20/03 1379 1260 525 ay 5/23/03 1640 1505 585 30 5/29/03 1444 1291 595 39 5/23/03 1555 1429 580 43 5/29/03 1335 1205 520 Sil 5/29/03 1180 1063 405 56 5/23/03 1575 1443 589 63 5/23/03 1135 1025 435 69 5/28/03 1280 1140 410 Ve 10/8/03 1449 1325 568 82 10/7/03 1400 1275 486 87 10/8/03 1095 997 431 93 10/8/03 1265 1134 442 99 10/8/03 1217 1094 450 106 10/8/03 1288 1186 461 111 10/8/03 1272 1138 467 119 10/8/03 1160 1045 480 Weight Tagging (kg) Sex” Age location 11.5 FI 22 Canadian Channel 30 F 5p) Brule Narrows 19 FM 26 Stokes Bay Des) FM Di Canadian Channel 8 MM 15 Stokes Bay 95 Unk 19 Stokes Bay eS) MM 17 Squirrel Falls 16 Unk 28 Squirrel Falls 20.5 FM 32 Canadian Channel DiS es) FM 28 Canadian Channel 30 FM 31 Stokes Bay DORMS FM 26 Kettle Falls IS). Unk 18 Squirrel Falls 11.5 Unk 19 Squirrel Falls 14.5 Unk 21 Kettle Falls 16.5 Unk NA Brule Narrows 16.75 MM 21 Kettle Falls IS) MM 35 Kettle Falls 9.5 M 32 Kettle Falls 9 MM 20 Kettle Falls 20 FM 30 Squirrel Falls 1935 FM 27 Kettle Falls 11 Unk 21 Kettle Falls 20.75 FM 22 Brule Narrows 18 Unk 26 Brule Narrows 19.25 FM 47 Seine River VES le 29 Squirrel Falls 22 FM 47 Stokes Bay 16.5 FI 22 Stokes Bay 8.5 Unk 19 Seine River 16 Unk 44 Seine River 8 Unk 18 Seine River 10 Ip 23 Rat River Bay Mouth 21 F NA Seine River 5) Unk Bi Brule Narrows 8 Unk 18 Seine River 10.5 M 16 Seine River 9.5 Unk 16 Brule Narrows 12 M 18 Seine River eS M Ds) Seine River 10.5 Unk 19 Seine River “F = female, maturity unknown; FM = female, mature; FI = female, immature; M = male, maturity unknown; MM = male, mature; Unk = sex unknown 2 and 20 m. The highest maximum detectable dis- tance from the boat was 0.95 km when the CART tag was lowered to 6 m. The lowest maximum detectable distance of 0.53 km occurred at a tag depth of 18 m. The mean maximum detectable distance was 0.65 km with a standard error of 0.04. Limitations in the per- formance of the acoustic portion of the tags led to the discontinuation of acoustic telemetry from the study beginning during July of 2003. No fish locations were recorded with the acoustic equipment despite attempts on 10 dates between 15 May and 15 June 2003. The complexity of the bottom contours of Rainy Lake may have been the reason for the lack of reception of the directional acoustic signal. Radio telemetry equipment was also tested. The highest maximum detectable distance. from the boat was 0.47 km when the CART tag was lowered to 2 m. The lowest maximum detectable distance of 0.017 km occurred at a tag depth of 20 m. The mean maximum detectable distance was 0.21 km with a standard error of 0.05. The maximum detectable distance by airplane with the tag lowered to 8.77 m was 0.55 km with a standard error of 38.5. 2006 Canadian Channel -92.964° -92.823° -92.701° ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 75 _ 4 Kilometers -92.580° -92.458° -92.336°W FIGURE 2. Distribution of Lake Sturgeon locations in Rainy Lake during May and June, 2003 and 2004. Lake Sturgeon locations The 41 implanted Lake Sturgeon were located 587 times throughout the study, with fish being tracked through September 2004. In 2003 Lake Sturgeon were located 285 times and in 2004 Lake Sturgeon were located 302 times. Lake Sturgeon utilized Squirrel Falls/Kettle Falls and adjacent areas (Canadian Channel, Hale Bay) ex- tensively throughout the study (37% of total locations, including both spawning and non-spawning seasons). A rock shelf below the Squirrel Falls Dam is one of the areas thought to be a spawning site for Lake Sturgeon in Rainy Lake. Egg samplers were placed below the Squirrel Falls Dam, in the Canadian Channel, and below the falls in the Pipestone River in 2004. Lake Sturgeon spawning, verified by hatching eggs and iden- tifying larvae, was confirmed only at the Squirrel Falls Dam and occurred between 14 June and 17 June of 2004. Thus, we cannot exclude the possibility of spawn- ing at other locations, but were able to verify spawning below the Squirrel Falls Dam. Other sites with Lake Sturgeon spawning habitat characteristics as listed by Scott and Crossman (1973) (e.g., 0.6-4.6 m depth, swift current, below falls) include the Kettle Falls Dam, Pipestone River, Rat River, and the Seine River (Crilly Dam and Highway 11 Bridge). Lake Sturgeon were located at all of these sites throughout the study, in- cluding the spawning season. The majority of May locations (89%) for both 2003 (85%) and 2004 (90%) were associated with the cur- rent area directly below Kettle Falls, Squirrel Falls, and in the Canadian Channel (Figure 2), which suggests pre-spawning aggregations. Four of the other six May locations were associated with the Brule Narrows, also an area with consistent current. Fewer Lake Sturgeon were located in the Kettle Falls/Squirrel Falls/Canadian Channel area as June progressed in both years (Adams 2004*). We suspect that Lake Sturgeon gradually left the spawning areas at Kettle Falls and Squirrel Falls, and dispersed pri- marily to the main basin east of Brule Narrows. Dur- ing this month, Lake Sturgeon locations in the Seine River were spread throughout Seine Bay and 36 km upriver. The increased number of Seine River locations from 2003 to 2004 was due to the increased number of tagged Lake Sturgeon. Only four fish from the Seine River were tagged for 2003 tracking, while six addi- tional fish were tagged for 2004 tracking. 76 THE CANADIAN FIELD-NATURALIST Vol. 120 2 4 6 Kilometers -93.270° -93.067° -92.963° -92.701° -92.580° -92.458°W FIGURE 3. Distribution of Lake Sturgeon in Rainy Lake during July and August, 2003 and 2004. During the July/August period, the majority of Lake Sturgeon were found from the Brule Narrows east- ward and in the Seine Bay/Seine River system, with only five locations found west of the Brule Narrows (Figure 3). Some Lake Sturgeon continued to be locat- ed in the Kettle Falls/Squirrel Falls area during this period in both years (15 locations and 35 locations, respectively). Locations in the main body of the South Arm were associated with the north shore more than the south shore during this time period (96% in 2003 and 84% in 2004). There are substantial habitat dif- ferences between the two areas with the south shore consisting of a sharp drop-off into depths of over 30 m. Conversely, the north shore consists of rock reefs and depths shallower than 15 m. Either the fish were less likely to use the deeper water, or the lack of locations was due to limitations of the telemetry equipment with radio signals detected from depths no greater than 18 m. September locations were obtained during both 2003 and 2004 with October locations obtained only in 2003. Locations during this time period were dis- tributed throughout the South Arm (Figure 4). Loca- tions in the Seine River were few, with zero in 2003 and two in 2004. Fish 75 was tagged in the Seine River but located in the South Arm during this time period. Lake Sturgeon were located on the south shore of the South Arm for the first time in September. One poten- tial explanation is that Lake Sturgeon were staging at locations in the vicinity of winter habitats. Rusak and Mosindy (1997) found that Lake Sturgeon in the Lake of the Woods/Rainy River system demonstrated con- sistent preferences for specific areas in the main basin of Lake of the Woods during the winter; these areas were adjacent to the mouth of the Rainy River where the Lake Sturgeon would later spawn in the spring. Lake Sturgeon movement rates Mean movement rates of Lake Sturgeon were cal- culated by month for descriptive purposes (Figure 5). Movement rates increased from May to June during 2003 and 2004, with a maximum movement rate of | 0.80 km/day in June of 2004 and a minimum move- | ment rate of 0.17 km/day in May of 2003. Movement | rates in five of the 10 months exceeded the mean spring 2006 4 Kilometers | -92.963° -92.823° -92.701° ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 77 -92.580° -92.458°W Ficure 4. Distribution of Lake Sturgeon locations in Rainy Lake during September and October 2003 and September 2004. movement rate of 0.50 km/day observed by Knights et al. (2002) for Lake Sturgeon in the upper Missis- sippi River system. Lake Sturgeon in Rainy Lake like- ly were dispersing into the lake away from spawning sites during the post-spawn period in June. However, movement rate patterns were dissimilar for the same months between years. Further research thus will be needed to ascertain seasonal movement patterns and explanations for those patterns. | Movement of fish between the Seine River and the main body of the South Arm did occur but was mini- mal throughout the study. Rusak and Mosindy (1997) found that in the Lake of the Woods/Rainy River sys- _ tem, there was a separation in the population based | primarily on winter habitat use. “River” fish spawned in the Rainy River and remained there throughout the _ winter months. “Lake” fish spawned in the Rainy River | but moved into the main basin of Lake of the Woods | during the winter months. If a similar dichotomy exists / in the Rainy Lake/Seine River population, there may | be greater movement of Lake Sturgeon between these a at two areas during the time period in which we did not attempt to locate fish (i.e., late fall and winter). Thus, Lake Sturgeon may move from Rainy Lake into the Seine River during this time period, but further inves- tigation will be needed. Movement patterns varied for individual Lake Stur- geon in Rainy Lake. Some fish did not exhibit dis- cernable movement patterns, such as fish 21, which was located throughout the east end of the South Arm from Stokes Bay to the Brule Narrows and also in Seine Bay and at the mouth of Seine Bay (Figure 6). Locations in 2003 began in the east and moved west with the last location east of the Brule Narrows. The first locations in 2004 were in the Seine Bay/Seine River area with the remainder of locations further east. One factor contributing to the differential patterns between years may be the spawning interval exhibited by Lake Sturgeon, which ranges between | and 3 years for males and 4 and 6 years for females (Magnin 1966). Telemetry results from Rainy Lake suggest that some Lake Sturgeon remained in the same area for 78 THE CANADIAN FIELD-NATURALIST Vol. 120 May June July August September Month FIGURE 5. Mean monthly movement rate (+ standard error) of Lake Sturgeon in Rainy Lake during 2003 and 2004. 2006 ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 79 4 Kilometers Tagging Location | 6/18/03 FiGureE 6. Distribution of locations for Lake Sturgeon number 21 in Rainy Lake, 2003 and 2004. extended periods. For example, fish 15 was tagged at the Squirrel Falls Dam on 28 May 2003 and was next located in the Brule Narrows area (Figure 7). Subse- quent locations were all in the area of the Brule Nar- rows until 26 August 2003. The next locations were in the Squirrel Falls Dam area beginning on 28 Sep- tember 2003. The fish was then located in the Squirrel Falls Dam area on 18 May 2004 and was not located in the Brule Narrows area until 30 June 2004. Lake Sturgeon home ranges _ Home ranges for Lake Sturgeon in Rainy Lake var- ied substantially, with a maximum home range size of } 14 844 ha, a minimum size (excluding fish 106, pos- ‘sible mortality) of 84 ha, and a mean home range size \of 4 625 ha (standard error = 642). Lyons and Kemp- inger (1992*) reported that most Lake Sturgeon in the Lake Winnebago system had consistent movement pat- terns, while others demonstrated variable movement patterns and apparently did not establish home ranges. They also found that Lake Sturgeon in Lake Win- nebago did not remain in any particular location for long periods of time. Conclusions This research should provide biologists from mul- tiple agencies insight into general movement patterns of Lake Sturgeon inhabiting Rainy Lake. Telemetry data indicated high utilization of the Squirrel Falls area, with 37% of total locations occurring at that area. Squirrel Falls was the only site at which spawning was confirmed by collection of Lake Sturgeon eggs, although other aggregations in areas associated with Kettle Falls, the Pipestone River, and the Rat River also could indicate spawning activity. Movement of Lake Sturgeon between the Seine River and the South Arm of Rainy Lake indicates the likelihood of one integrated population on the east end of the South Arm. The lack of locations in the Seine River during the months of September and October might result from Lake Sturgeon moving into deeper water areas of the Seine River and out of the range of telemetry gear or simply moving back into the South Arm. Further re- search will be needed to determine the winter range of Lake Sturgeon in Rainy Lake. The Lake Sturgeon has historically been a resource of both cultural and economic importance to the Rainy Lake area. Due to 80 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 2. Home ranges and mean movement rates (SD) by year for Lake Sturgeon in Rainy Lake. Missing values (dots) indicat that a fish was located zero or one times that year and thus movement could not be calculated. Fish number Total Home range Mean movement rate Mean movement rate locations (ha) 2003 (km/d) 2004 (km/d) 1 Ds 2164 0.320 (0.22) 0.265 (0.31) 2 12 7758 0.170 (0.14) 0.564 (0.26) 3 7) 2985 0.267 (0.20) 0.372 (0.33) 4 7 14844 0.619 (0.50) 2 5 13 3779 0.285 (0.33) 0.510 (0.36) 6 15 5416 0.448 (0.70) 0.663 (0.72) 7 23 3558 0.289 (0.28) 1.065 (0.74) 8 20 VAY 0.076 (0.05) 0.554 (1.11) 9 11 8600 0.156 (0.20) i 10 29 11894 0.808 (0.67) 0.977 (1.29) 11 17 10501 0.309 (0.23) 0.382 (0.23) 12 12 578 0.233 (0.25) 0.261 (0.43) 13 31 4404 0.420 (0.54) 0.120 (0.23) 14 12 3806 1.121 (1.00) 0.195 (0.21) 15 34 7141 0.352 (0.42) 0.714 (0.92) 16 19 2434 0.324 (0.25) : 17 29 10245 0.565 (0.53) 0.552 (0.41) 18 28 Sly) 0.279 (0.28) 0.513 (0.46) 19 14 6434 0.396 (0.39) 1.091 (1.74) 20 34 1859 0.299 (0.51) 0.870 (1.30) 21 15 10017 0.316 (0.11) 0.817 (0.61) 22 26 8492 0.459 (0.36) 0.511 (0.38) 23 19 4095 0.684 (0.80) 0.187 (0.20) 24 19 9411 0.587 (0.94) 0.236 (0.31) 25 7 7677 : 1.713 (1.56) 27 10 1921 0.324 (0.19) 0.200 (0.20) 30 10 1196 : 0.527 (0.66) >I 18 1049 0.131 (0.06) 0.262 (0.38) 56 6 1499 : 0.651 (0.81) 69 10 2448 0.267 (0.36) 0.960 (1.26) TS 14 159 1.160 (2.77) 82 6 641 0.213 (0.26) 87 5 433 0.397 (0.15) 93 8 2355 0.995 (0.97) 106 14 71 0.099 (0.04) ita 3 84 1.114 (0.35) 119 6 380 0.583 (0.90) the movements between Minnesota and Ontario, coor- dination of future management efforts among provin- cial, state, and federal agencies will be important. Acknowledgments Technical review was provided by Trent Sutton, Daniel Hubbard, and three anonymous reviewers. Par- tial funding for this project was provided by the Na- tional Park Service through the Great Plains Cooper- ative Ecosystem Studies Unit Cooperative Agreement Modification Number J6820031002 (administered through the University of Nebraska), the U.S. Geolog- ical Survey, the Minnesota Department of Natural Re- sources, and the Ontario Ministry of Natural Resources. Field assistance was provided by Voyageurs National Park employees. This manuscript was approved for publications by the South Dakota Agricultural Exper- iment Station as Journal Series Number 3472. Documents Cited (marked * in text) Adams, W. E., Jr. 2004: Lake sturgeon biology in Rainy Lake Minnesota and Ontario. M. Sc. thesis, Department of Wild- life and Fisheries Sciences, South Dakota State University Brookings. Bassett, C. 1982. Management plan for lake sturgeon (Acr- penser fulvescens) in the Indian River and Indian Lake Alger and Schoolcraft counties, Michigan. U.S. Fores! Service, Manistique Ranger District, Hiawatha Nationa! Forest, in cooperation with the Michigan Department of! Natural Resources, Manistique. Environment Canada. 2005. Nature and wildlife. Avail: able: http://www.ec.gc.ca/wild_e.html (September 2005) Ericson, D. W., G. F. Lindholm, and J. O. Helgesen. 1976 Water resources of the Rainy Lake watershed, northeastern )06 4 Kilometers ADAMS, KALLEMEYN, AND WILLIS: LAKE STURGEON MOVEMENTS 8 | | Tagging Location|. 4 bs SORNB ya ae 2003 ry @ 2004 Ficure 7. Distribution of locations for Lake Sturgeon number 15 in Rainy Lake, 2003 and 2004. Minnesota. U. S. Geological Survey, Hydrologic Investi- gations Atlas HA-556, Washington, D.C. ooge, P. N., W. Eichenlaub, and E. Solomon. 1999. The animal movement program. U.S. Geological Survey, Alas- ka Biological Science Center, Anchorage. ternational Rainy Lake Board of Control/International Lake of the Woods Control Board (IRLBC/ILWCB). 1984. Briefing paper submitted to the International Joint Committee, St. Paul, Minnesota. yons, J., and J. J. Kempinger. 1992. Movements of adult lake sturgeon in the Lake Winnebago System. Wisconsin Department of Natural Resources, Research Report No. 156, Madison. linnesota Department of Natural Resources (MN DNR). 2003. Minnesota’s list of endangered, threatened, and Special concern species. Minnesota Rules, Chapter 6134. Available: http://www.dnr.state.mn.us/ets/index html (Sep- tember 2005). iterature Cited dams, W. E., Jr., L. W. Kallemeyn, and D. W. Willis. 2006. Lake sturgeon population characteristics in Rainy Lake, Minnesota and Ontario. Journal of Applied Ichthy- ology 22: 97-102. uer, N. A. 1999. Population characteristics and movements of lake sturgeon in the Sturgeon River and Lake Superior. Journal of Great Lakes Research 25: 282-293. Borkholder, B. D., S. D. Morse, H. T. Weaver, R. A. Hugill, A. T. Linder, L. M. Schwarzkopf, T. E. Perrault, M. J. Zacher, and J. A. Frank. 2002. Evidence of a year- round resident population of lake sturgeon in the Kettle River, Minnesota, based on radio telemetry and tagging. North American Journal of Fisheries Management 22: 888- 894. Fortin, R., J.-R. Mongeau, G. Desjardins, and P. Dumont. 1993. Movements and biological statistics of lake sturgeon (Acipenser fulvescens) populations from the St. Lawrence and Ottawa River system, Quebec. Canadian Journal of Zoology 71: 638-650. Gallep, G. W., and J. J. Magnuson. 1972. Effects of negative buoyancy on the behavior of the bluegill, Lepomis macro- chirus Rafinesque. Transactions of the American Fisheries Society 101: 507-512. Hart, L. G., and R. C. Summerfelt. 1975. Surgical proce- dures for implanting ultrasonic transmitters into flathead catfish (Pylodictis olivaris). Transactions of the American Fisheries Society 104: 56-59. Knights, B. C., J. M. Vallazza, S. J. Zigler, and M. R. Dewey. 2002. Habitat and movement of lake sturgeon in the upper Mississippi River System, USA. Transactions of the American Fisheries Society 131: 507-522. Magnin, E. 1966. Recherches sur les cycles de reproduction des sturgeons Acipenser fulvescens Rafinesque. Annales de la Station Centrale d’Hydrobiologie Appliquee 9: 8-242. 82 THE CANADIAN FIELD-NATURALIST McCleave, J. D., and K. A. Stred. 1975. Effect of dummy telemetry transmitters on stamina of Atlantic salmon (Salmo salar) smolts. Journal of the Fisheries Research Board of Canada 32: 559-563. McKinley, S., G. V. Der Kraak, and G. Power. 1998. Sea- sonal migrations and reproductive patterns in the lake stur- geon, Acipenser fulvescens, in the vicinity of hydroelectric stations in northern Ontario. Environmental Biology of Fishes 51: 245-256. Moser, M. L., A. F. Olson, and T. P. Quinn. 1990. Effects of dummy ultrasonic transmitters on juvenile coho salmon. American Fisheries Society Symposium 7: 353-356. Nickum, J. G., H. L. Bart, Jr, P. R. Bowser, I. E. Greer, C. Hubbs, J. A. Jenkins, J. R. MacMillan, J. W. Rachlin, J. D. Rose, P. W. Sorensen, and J. R. Tomasso. 2004. Guidelines for the use of fishes in research. American Fish- eries Society, American Institute of Fishery Research Bio- logists, and American Society of Ichthyologists and Her- petologists, Bethesda, Maryland. Rusak, J. A., and T. Mosindy. 1997. Seasonal movements of lake sturgeon in Lake of the Woods and the Rainy River, Ontario. Canadian Journal of Zoology 74: 383-395. Vol. 12 Scott, W. B., and E. J. Crossman. 1973. Freshwater fishe of Canada. Fisheries Research Board of Canada, Bulleti: Number 184, Ottawa. Stasko, A. B., and D. G. Pincock. 1977. Review of under water biotelemetry, with emphasis on ultrasonic techniques Journal of the Fisheries Research Board of Canada 34 1261-1285. Threader, R. W., and C. S. Brosseau. 1986. Biology an management of the lake sturgeon in the Moose River, On tario. North American Journal of Fisheries Managemen 6: 383-390. White, G. C., and R. A. Garrett. 1990. Analysis of wildlif radio-tracking data. Academic Press, San Diego, California Winter, J. 1996. Advances in underwater biotelemetry. Page 555-590 in Fisheries techniques, second edition. Edite: by B. R. Murphy and D. W. Willis. American Fisherie Society, Bethesda, Maryland. Received 8 March 2005 Accepted 11 October 2005 lorthern Range Expansion and Invasion by the Common Carp, 'yprinus carpio, of the Churchill River System in Manitoba .SCAL H. J. BApiou! and L. GORDON GOLDSBOROUGH! lepartment of Botany, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada; e-mail: pbadiou@shaw.ca; Present address: 48 DesMeurons Street, Winnipeg, Manitoba, RZH 2M1 Canada lelta Marsh Field Station, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada diou, Pascal H.J., and L. Gordon Goldsborough. 2006. Northern range expansion and invasion by the Common Carp, Cyprinus carpio, of the Churchill River system in Manitoba. Canadian Field-Naturalist 120(1): 83-86 cent fisheries data from northern Manitoba indicates that the Common Carp (Cyprinus carpio) has extended the northern lit of its range. Additionally, it also appears that carp have invaded and established viable populations in the Manitoba rtion of the Churchill River. Habitat degradation and altered flow regimes as result of hydroelectric development in north- 1 Manitoba may have facilitated the expansion of carp in the region. y Words: Common Carp, Cyprinus carpio, exotic species, range expansion, invasion, habitat disturbance, Manitoba. The Common Carp (Cyprinus carpio) was intro- ced intentionally to Manitoba, Canada, when fish ym Minnesota were stocked in the Assiniboine River itershed southeast of Brandon in 1886 (Stewart and atkinson 2004). These early attempts at introduc- n failed, and carp only became established after pop- itions from the northern U.S. entered the province ‘ough the Red River (Atton 1959). In fact, the Com- mn Carp was largely unknown in Manitoba until 1938, 1en it was positively identified in the Red River near ckport (Hinks 1943). Today, the Common Carp is und throughout most of the southern and central Zions of the province, but particularly in the Red and siniboine rivers, and lakes Manitoba, Winnipeg, and innipegosis. Common Carp do not appear to have ex- nded into the eastern portion of the province, likely e to the fact that they cannot easily migrate upstream this region because of the many obstructions they juld encounter (i.e., Beaver dams). Additionally, lakes this portion of the province are characterized by cky substrates and clear deep waters typical of the ecambrian Shield and do not provide suitable habi- for viable populations of the Common Carp. Benthivorous fish such as the Common Carp can ve profound effects in aquatic ecosystems by increas- 2 turbidity through sediment resuspension while for- ing and spawning (Breukelaar et al. 1994; Zambra- yet al. 2001). Carp are known to uproot submerged acrophytes during spawning and accidentally con- me them while foraging for benthic invertebrates olterman 1990; Roberts et al. 1995; Lougheed et al. 98; Zambrano and Hinojosa 1999). Sediment resus- nsion and excretion by carp can increase water col- nn nutrient concentrations, thereby causing phyto- ankton to flourish (Lamarra 1975; Breukelaar et al. 94: King et al. 1997). A shading effect results from ese blooms, further suppressing submerged macro- phytes and creating an ecological feedback mechanism that perpetuates the turbid state (Scheffer 1998). Carp physically disturb the sediment-water interface and the algae associated with sediment. These algae play a major role in the stabilization of the bottom sediments (Taylor et al. 1998) and the regulation of nutrient fluxes across the sediment-water interface (Goldsborough and Robinson 1985; Carlton and Wetzel 1988; Woodruff et al. 1999). It is thought that many of the once clear, shallow lakes and wetlands in southern Manitoba have switched to the turbid state due to the loss of sub- merged macrophyte cover which, in turn, was caused by the proliferation of Common Carp (Badiou 2005). Before 1940, carp were found only in the waters of the Red River (Figure 1a). After invading the province from the south, the Common Carp spread quickly throughout the Red and Assiniboine River watersheds prior to the 1950s and established themselves in the south basins of lakes Winnipeg and Manitoba (Figure 1b). However, the greatest range expansion occurred between 1950 and 1970 when carp were easily able to access the northern and western portions of the province after successfully invading Lake Winnipeg, Lake Man- itoba, and Lake Winnipegosis, the largest lakes in the province (Figure Ic). Swain (1979) indicated that by 1976 the Common Carp was found in most large lakes and virtually all major rivers in Manitoba, with the exception of the Churchill River system. A specimen collected from Split Lake (Figure Id) in 1963 was considered to be the most northerly record for the distribution of carp in the world (McCrimmon 1968). These original reports stated that carp caught from Split Lake appeared emaciated and unhealthy. According to Atton (1959), cold summer temperatures prevented the carp from invading the Churchill River system. However, as shown in Figure 1d, the Common Carp has migrated northward through the Nelson River 83 84 THE CANADIAN FIELD-NATURALIST system and, in the last decade, has become established in the Churchill River system according to carp pro- duction data spanning the period from 1970 to 2004. On the Nelson River system, carp have become established in Split Lake where commercial catches (3-74 kg per year) have been reported starting in 1996. Fish monitoring studies for the Limestone Generating Station (C. Barth, personal communication) and the pro- posed Gull Generating Station (D. J. Pisiak, unpub- lished data) documented Common Carp in Stephens Lake and in the forebay of the Limestone Generating Station (Figure 1d). Furthermore, during the course of fish studies for the proposed Conawapa Generating Station, a large, approximately 4 kg, Common Carp was caught in a backwater area near the lower Lime- stone Rapids (P. Nelson, personal communication). This is the first report of Common Carp downstream of the Limestone Generating Station and the most northerly location documented for the Nelson River. More important than the northward expansion of carp on the Nelson River system is the invasion of the Churchill River system in the last decade. Recent fish- eries records for northern Manitoba provided by the Freshwater Fish Marketing Corporation indicate that the Common Carp is now established in and around the Churchill River system with commercial catches report- ed from Britton, Guthrie, Highrock, Loon, Sisipuk, and Southern Indian lakes (Figure 1d). Commercial catches in these lakes were much higher (range 8—1184 kg per year) relative to those from Split Lake on the Nelson River system. Carp were likely introduced into the Churchill River system from the Saskatchewan River system, either by fishers using them as bait or by birds that may have inoculated carp eggs from one system into the other. A thorough search of the primary liter- ature and government documents revealed that Com- mon Carp have not been reported in the Saskatchewan portion of the Churchill River. The fact that carp have not been reported in the Churchill River in Saskatche- wan indicates that the transfer of the Common Carp between the Saskatchewan River and Churchill River systems must have occurred within the province of Manitoba in the vast wetland area that occurs be- tween these two watersheds along the Manitoba/Sas- katchewan border. Based on the fact that carp were found in Split Lake as early as 1963, it is not surprising that they have con- tinued to migrate northward through the Nelson River system. However, given the poor condition of carp in Split Lake in 1963, a change must have occurred, which has allowed them to persist in this system. The two basic habitat requirements necessary to sustain carp populations are: (1) a shallow marsh environment with abundant aquatic vegetation for spawning and, (2) an area of deep water where carp can overwinter (Mc- Crimmon 1968). We hypothesize that the construction of the Kettle Generating Station in 1974 at the outlet of Stephens Lake on the Nelson River provided the neces- sary habitat requirements to allow the establishment and Vol. 120 sustain viable populations of Common Carp in Split Lake and Stephens Lake. Construction of this generating sta- tion increased the size of Stephens Lake by 242 km? and resulted in the flooding of 192 km? of land (Envi- ronment Canada / Fisheries and Oceans 1992a). This created an extensive area of shallow marsh-like habitat where carp could spawn in the summer. This was the case in Ontario where Swee and McCrimmon (1966) reported that Lake St. Lawrence near Cornwall, formed as the result of flooding during hydroelectric develop- ment on the St. Lawrence River, provided ideal spawn- ing habitat for the Common Carp. However, it was also noted that fluctuations in lake level associated with hydroelectric activities caused many of the eggs at- tached to vegetation to be exposed and destroyed. More importantly, the construction of the Kettle Generating Station, with a forebay head of 30 meters (Environment Canada / Fisheries and Oceans 1992a), created a deep basin with warmer profundal zone waters to which carp can escape during cold winter months. The completion of the Churchill River Diversion in 1976 raised the water level in Southern Indian Lake by approximately 3 m and flooded 187 km? of surround- ing land (Environment Canada / Fisheries and Oceans 1992a). Like the flooding experienced on Stephens Lake after the construction of the Kettle Generating Station, flooding on Southern Indian Lake would have also provided ideal spawning habitat for carp. The extensive flooding on Southern Indian Lake increased shoreline erosion and drastically increased suspend- ed sediment concentrations in the lake from 5 mg-L"!, pre-impoundment, to 30-50 mg-L"', post-impoundment (Environment Canada / Fisheries and Oceans 1992b). This anthropogenic increase in suspended sediments would have conferred a competitive advantage on carp which can feed more efficiently in highly turbid waters due to their superior olfactory senses, relative to sight- feeders (Panek 1987). Water flows on the Churchill and Nelson river sys- tems have been altered dramatically as a result of the Churchill River Diversion, which has diverted 80% of the flow from the Churchill River at Southern Indian Lake into the Nelson River system. Carp are found in large, slow-moving rivers as well as fast flowing streams (Panek 1987). Due to the ability of carp to inhabit wa- ters of varying flow, the increased flow on the Nelson and decreased flow on the Churchill, as well as the in- creased variability in flows in both systems as a result of hydroelectric development, would also have con- ferred an advantage on the Common Carp relative to native fish that may be less tolerant of variations in flow regime. Overall, it is not surprising that Common Carp are invading northern Manitoba in step with hydroelec- tric development, as many authors have demonstrated the link between anthropogenic habitat alteration and invadability of ecosystems (Moyle and Light 1996; Keith and James 1999; Byers 2002). Keith and James (1999) showed a positive correlation between the num- 2006 BADIOU AND GOLDBOROUGH: CARP OF THE CHURCHILL RIVER SYSTEM 85 ex ats pen (Wax a Ls raz uae RZ ip oA Hudson Bay Ficure |. Range expansion of the Common Carp (Cyprinus carpio) in Manitoba (A) prior to 1940, (B) prior to 1950, (C) prior to 1970, and (D) prior to 2000. Carp distributions prior to 1970 were estimated based on date of first occur- rences reported in Swain (1979), while the range expansion between 1970 and 2000 was estimated from carp pro- duction data provided by Manitoba Conservation and the Freshwater Water Fish Marketing Corporation for the period from 1970 to 2000. Numbers in D indicate locations discussed in the text: (1) Split Lake, (2) Stephens Lake. (3) Limestone Generating Station, (4) Britton Lake, (5) Sisipuk Lake, (6) Guthrie Lake, (7) Highrock Lake, (8) Southern Indian Lake, (9) Loon Lake, (10) Churchill River Estuary, and (11) Nelson River Estuary. 86 THE CANADIAN FIELD-NATURALIST ber of reservoirs and the number of introduced species in North American drainages. Given that the Common Carp has already been reported in subarctic regions (Lukin 1999) and is tolerant of a wide range of salin- ity levels (Lam and Sharma 1985) it is a matter of time before the populations of carp established on the Nel- son and Churchill river systems reach the Churchill River and Nelson river estuaries. Acknowledgments We thank Ellen Smith of the Freshwater Fish Mar- keting Corporation for providing us with carp produc- tion data from Manitoba lakes. We also thank Patrick Nelson, Paul Graveline and Cam Barth of North/South Consultants Inc., in Winnipeg for helpful comments. This is publication number 311 from the Delta Marsh Field Station (University of Manitoba). Literature Cited Atton, F. M. 1959. The invasion of Manitoba and Saskatche- wan by carp. Transactions of the American Fisheries Soci- ety 88: 203-205. Badiou, P. H. J. 2005. Ecological impacts of an exotic ben- thivorous fish in wetlands: a comparison between common carp (Cyprinus carpio L.) additions in large experimental wetlands and small mesocosms in Delta Marsh, Manitoba. Ph.D. dissertation, Department of Botany, University of Manitoba, Winnipeg, Manitoba. Breukelaar, A. W., E. H. R. R. Lammens, J. G. P. Klein Breteler, and I. Tatrai. 1994. Effects of benthivorous bream (Abramis brama) and carp (Cyprinus carpio) on sed- iment resuspension and concentrations of nutrients and chlorophyll a. Freshwater Biology 32: 113-121. Byers, J. E. 2002. Impact of non-indigenous species on na- tives enhanced by anthropogenic alteration of selection regimes. Oikos 97: 449-458. Carlton, R. G., and R. G. Wetzel. 1988. Phosphorus flux from lake sediments: Effects of epipelic algal oxygen pro- duction. Limnology and Oceanography 33: 562-570. Environment Canada / Fisheries and Oceans. 1992a. Fed- eral Ecological Monitoring Program (FEMP) Final Report. Volume 2. Environment Canada / Fisheries and Oceans. 1992b. Fed- eral Ecological Monitoring Program (FEMP) Final Report. Volume 1. Goldsborough, L. G., and G. G. C. Robinson. 1985. Effect of an aquatic herbicide on sediment nutrient flux in a fresh- water marsh. Hydrobiologia 122: 121-128. Hinks, D. 1943. The Fishes of Manitoba. Department of Mines and Natural Resources, Winnipeg. Keith, B. G., and H. B. James. 1999. Invasion of North Amer- ican drainages by alien fish species. Freshwater Biology 42: 387-399. King, A. J., A. I. Robertson, and M. R. Healey. 1997. Experimental manipulations of the biomass of introduced carp (Cyprinus carpio) in billabongs. I. Impacts on water- column properties. Marine and Freshwater Research 48: 435-443. Vol. 120 Kolterman, B. F. 1990. Effects of common carp and black bullheads on sago pondweed. M.Sc. dissertation, South Dakota State University. Lam, T. J., and R. Sharma. 1985. Effects of salinity and thyroxine on larval survival, growth and development in the carp, Cyprinus carpio. Aquaculture 44: 201-212. Lamarra, V. A. 1975. Digestive activities of carp as a major contributor to the nutrient loading of lakes. Verhandlungen Internationale Vereinigung fiir theoretische und angewandte Limnologie 19: 2461-2468. Lougheed, V. L., B. Crosbie, and P. Chow-Fraser. 1998. Predictions on the effect of common carp (Cyprinus carpio) exclusion on water quality, zoopiankton, and submergent macrophytes in a Great Lakes wetland. Canadian Journal of Fisheries and Aquatic Sciences 55: 1189-1197. Lukin, A. A. 1999. Occasional introduction of carp Cyprinus carpio into a subarctic water body. Journal of Ichthyology 39: 419-420. McCrimmon, H. R. 1968. Carp in Canada. Fisheries Re- search Board of Canada. Bulletin 165. Moyle, P. B., and T. Light. 1996. Fish invasions in California: do abiotic factors determine success. Ecology 77: 1666- 1670. Panek, F. M. 1987. Biology and ecology of carp. Pages 1-16 in Carp in North America. Edited by E. L. Cooper. Amer- ican Fisheries Society, Bethesda, Maryland. Roberts, J., A. Chick, L. Oswald, and P. Thompson. 1995. Effect of carp, Cyprinus carpio L., an exotic benthivorous fish, on aquatic plants and water quality in experimental ponds. Marine and Freshwater Research 46: 1171-1180. Scheffer, M. 1998. Ecology of shallow lakes. Chapman & Hall, Toronto. Stewart K. W., and D. A. Watkinson. 2004. The freshwater fishes of Manitoba. University of Manitoba Press, Win- nipeg. Swain D. P. 1979. Biology of the carp (Cyprinus carpio L.) in North America and its distribution in Manitoba, North Dakota and neighbouring U.S. waters. Manuscript Report Number 79-73. Manitoba Department of Mines, Natural Resources and Environment, Winnipeg. Swee, U. B., and H. R. McCrimmon. 1966. Reproductive biology of the carp, Cyprinus carpio L., in Lake St. Law- rence, Ontario. Transactions of the American Fisheries Society 95: 372-380. Taylor, I. S., D. M. Paterson, and A. Mehlert. 1998. The quantitative variability and monosaccharide composition of sediment carbohydrates associated with intertidal diatom assemblages. Biogeochemistry 45(3): 303-327 Woodruff, S. L., W. A. House, M. E. Callow, and S. C. Leadbeater. 1999. The effects of biofilms on chemical processes in surficial sediments. Freshwater Biology 41: 73-89. Zambrano, L., and D. Hinojosa. 1999. Direct and indirect effects of carp (Cyprinus carpio L.) on macrophyte and benthic communities in experimental shallow ponds in central Mexico. Hydrobiologia 408/409: 131-138. Zambrano, L., M. Scheffer, and M. Martinez-Ramos. 2001. Catastrophic response of lakes to benthivorous fish introduction. Oikos 94: 344-350. Received 21 March 2005 Accepted 30 January 2006 Diversity and Range of Amphibians of the Yukon Territory BRIAN G. SLOUGH! and R. LEE MENNELL? ' 35 Cronkhite Road, Whitehorse, Yukon Territory Y1A 5S9 Canada 2 P.O. Box 10002, Whitehorse, Yukon Territory Y1A 7A1 Canada Slough, Brian G., and R. Lee Mennell. 2006. Diversity and range of amphibians of the Yukon Territory. Canadian Field-Naturalist 120(1): 87-92. Four amphibian species occur in the Yukon: Western Toad (Bufo boreas; first verified record in 1961), the Boreal Chorus Frog (Pseudacris maculata, first record in 1995), the Columbia Spotted Frog (Rana luteiventris, first record in 1993), and the Wood Frog (Rana sylvatica; first record in 1933). The Western Toad is restricted to the Liard Basin in five geographically separated areas. Breeding sites have been located on the lower Coal River and vicinity. The Boreal Chorus Frog has been documented from a small area of the lower La Biche River valley near the Yukon-British Columbia-Northwest Territories border. The Colum- bia Spotted Frog occurs in two widely separated areas, at Bennett Lake in the southwest and in the Liard Basin in the southeast The Wood Frog is widespread below treeline. Limited survey efforts continue to hinder our knowledge of amphibian distribution. Key Words: Bufo boreas, Western Toad, Pseudacris maculata, Boreal Chorus Frog, Rana luteiventris, Columbia Spotted Frog, Rana sylvatica, Wood Frog, amphibian, Yukon, distribution. Biological surveys began in the interior of the Yukon Territory in 1899 (Osgood and Bishop 1900); however, these were often directed at mammals and birds, and did not report amphibian observations. The earliest known amphibian records for the region are from Atlin (Slevin 1928; Western Toad and Wood Frog) and Bennett Lake, British Columbia (Logier 1932; Colum- bia Spotted Frog, formerly Western Spotted Frog). The first published report of amphibians in the Yukon was by Loomis and Jones (1953), who reported a Wood Frog collected at Dezadeash Lake in 1948 by the University of Kansas Museum of Natural History. Museum collections from the Canadian Museum of Nature (CMN) and the American Museum of Natural History (AMNH) date to 1933 and 1936, respectively. Logier and Toner (1961) report only the single locali- ty record of Loomis and Jones (1953). Using an ex- panded database, including museum specimens and inferences from locality records in Alaska, the North- west Territories and British Columbia, Martof (1970) shows an approximate range for the Wood Frog; although a large area with no data existed in the east- ern and northern Yukon. The amphibian range maps in Stebbins (2003) were modified with information provided by BGS. The Western Toad (Bufo boreas) was first observed in 1961 in the southeast Yukon (Cook 1977). The Columbia Spotted Frog was first observed in 1993 (Mennell 1997) in the southwest Yukon. There are no published records of the Boreal Chorus Frog or other species for the Yukon. All species of amphibians have been underrepresented in records due to the remote- ness of breeding sites and lack of directed surveys. Several amphibian surveys and expert observations have been made since 1993 by the authors and others, and the data have remained unpublished. We typical- ly used visual encounter surveys and dipnetting of terrestrial, semi-terrestrial (wetland) and aquatic habi- tats (Thoms et al. 1997). We review unpublished data, unpublished reports and museum collections to pres- ent an update on the amphibian species of the Yukon and their ranges. Anecdotal information on phenology and habitats is provided. Scientific and common names follow Collins and Taggart (2002). Species Accounts WESTERN TOAD (Bufo boreas) Western Toads occur from south-coastal Alaska, through western Canada and the western United States to Baja California. Cook (1977) reported two museum specimens of the Western Toad from the Yukon. The Whitehorse specimen, collected in 1948, was verified in 1997 by F. Cook, who also verified the label in 2003 as collected by William Mason July 1-10. Mason was an entomologist and the date notation suggests a spec- imen taken in a pan or similar trap set for that period and contents preserved on the last date (F. Cook, per- sonal communication). This specimen is here consid- ered to be either an extra-limital stray or erroneously labelled, since the nearest confirmed breeding records of Western Toads are 100 km to the south in British Columbia on Bennett Lake (Slough 2004*; 17 km from the British Columbia-Yukon border) and on Tagish Lake (Mennell and Slough 1998*; 40 km from the border). A single adult was found in non- breeding habitat on Tagish Lake 9 km from the border (Mennell and Slough 1998*). All other records are from the Liard Basin in the southeast Yukon, where the Western Toad is now known from at least 11 popu- lations in five geographically separated areas (Figure 1). These are summarized from west to east. Two adult Western Toads were observed near a geo- thermal spring on the Meister River (60°17'N, 130°07'W) in 1988 (D. Mossop, personal communi- 87 88 THE CANADIAN FIELD-NATURALIST cation). RLM searched this area in 2004 but did not observe Western Toads. A single adult was observed near Upper Liard (60°05’°N, 128°58’W) in 2004 (C. Eckert, personal communication; verified from pho- tographs). At least six breeding populations of Western Toads occur in close proximity (within 16 km) on the lower Coal River and wetlands to the west. The most visited population and breeding site is the Coal River Springs (60°09'N, 127°26'W). G. Scotter collected two speci- mens in 1977 (CMN 17798-1 and -2), and at least six Western Toad records of adults and metamorphs were documented between 1982 and 2004 (B. Slough and L. Mennell, unpublished data; Slough 2005a*). Two breeding sites exist in spring marshes below the upper and lower springs. The temperature of the main source pool is 11°C, likely maintaining frost-free hibernating sites nearby. Other breeding sites occur in riparian floodplain backwater channels and ponds above the springs (60°11'N, 127°29'W), below the springs on an island in the Coal River (60°07'N, 127°24'W), and fur- ther downstream on the Coal River in a bedrock pool [60°08'N, 127°14'W] (J. Staniforth and J. Meikle, per- sonal communication; L. Mennell, unpublished data; Slough 2005a*). Juvenile toads have also been ob- served in the chain of lakes southwest of the springs (60°14'N, 127°29'W) [B. Slough, unpublished data; Slough 2005a*] and in a tributary of the lakes (60°06'N, 127°32'W) [Slough 2005b*], suggesting the presence of breeding populations. Cook (1977) reported a specimen collected in 1961 from North Toobally Lake (60°20'N, 126°15'W). An adult Western Toad was observed 23 km to the south near South Toobally Lake (60°08’N, 126°19’W) in 1996 (C. Eckert, H. Grunberg, P. Sinclair, personal communication; verified from photograph). A single adult Western Toad was observed on the lower Beaver River [60°02’N, 124°32’W] in 1997 (B. Bennett, V. Loewen, J. Staniforth, personal communication; veri- fied from photograph). All of the Western Toad local- ities are low elevation sites with early deep snow accu- mulation. The snow cover likely prevents deep frost penetration and permits safe hibernation, as speculated by Cook (1977). Western Toad breeding sites are predominantly shal- low, silty areas on ponds, lakes and rivers. Adults are often encountered far from water in forests or mead- ows. Breeding dates in the Yukon are unknown; how- ever, populations in northwestern British Columbia breed in late May (B. Slough and L. Mennell, unpub- lished data). A population in the Atlin Warm Springs, 20 km south of Atlin, breeds in late February-early March. Tadpoles have been observed on the Coal River in July, and tadpoles and metamorphs were observed there on 10 August 2004. BOREAL CHORUS FROG Pseudacris maculata While Pseudacris maculata is currently considered a distinct species following Platz (1989), it was formerly Vol. 120 called P. triseriata maculata (and earlier Pseudacris triseriata septentrionalis). Recent phylogeographic studies of North American chorus frogs have shown that range limits and taxonomy of the triseriata com- plex may need revision (Moriarty and Cannatella 2004). The Boreal Chorus Frog ranges across much of cen- tral North America east of the continental divide, from the Northwest Territories to southern James Bay and south to Arizona and New Mexico. We present the first records of the Boreal Chorus Frog from the Yukon. Boreal Chorus Frog calls were identified at five natural wetlands and roadside/airstrip ditches along a 10 km section of road and airstrip in the lower La Biche River valley in the southeast Yukon (60°03'N 123°58'W to 60°08'N 124°03'W) between 8 and 17 June in 1995, 1997 and 1999 (Figure 1) [B. Bennett, C. Eckert, V. Loewen, P. Sinclair, J. Staniforth, personal communi- cation; B. Slough, unpublished data]. BGS also iden- tified Boreal Chorus Frog tadpoles there on 10 June 1999. Extensive surveys by the authors in southeastern Yukon in 2004 did not yield any further observations of the species. Matsuda et al. (2006) show the species 160 km to the southeast at Fort Nelson, British Colum- bia, and it occurs 125 km northeast at the junction of the South Nahanni and Liard rivers (Fournier 1998*). COLUMBIA SPOTTED FROG (Rana luteiventris) The Columbia Spotted Frog occurs between the Rocky Mountains and Coast Ranges from the Yukon to Utah and Nevada in the south. Mennell (1997) located two populations of Columbia Spotted Frogs adjacent to the West Arm of Bennett Lake in 1993, in a pond (60°05'N 135°01'W) and a beaver pond on a creek (60°O1'N 135°12'W) [Figure 1]. Slough (2002) obtained single voucher specimens and preserved toe- tips (for phylogeographic studies, n=10 and 11, respec- tively) from the two ponds in 2001 (Slough et al. 2002*). A third population was found in a beaver pond in the same area (Slough 2002) [60°03’N 134°58’W] where a voucher specimen and six toe-tips were obtained; specimens are held at the Redpath Museum, McGill University [RM 3382, 3379 and 3381 for the three populations, respectively]. Additional Columbia Spot- ted Frog adults (n=7 in 4 sites) were observed within 2.5 km of Site 2 in apparent non-breeding habitats (i.e., early life-stages not observed) along the Partridge River to Bennett Lake (R. McClure, personal communication, verified from photograph; B. Slough and L. Mennell, unpublished data). These three breeding populations are the most northerly of a series of populations that extends into northwestern British Columbia (Matsuda et al. 2006, Mennell 1997; Mennell and Slough 1998*; Slough et al. 2002*). In 2003, two adult Columbia Spotted Frogs were ob- served in Irons Creek Lake in the southeastern Yukon (60°00’N 127°05’W; P. Hovingh, personal communi- cation to Yukon Fish and Wildlife Branch) (Figure 1). Several adults, juveniles and one egg mass were ob- served in 2004 and 2005 (Slough 2005b*). The long- 2006 SLOUGH AND MENNELL: AMPHIBIANS OF THE YUKON 89 Key to Map Symbols Boreal Chorus Frog Columbia Spotted Frog Westem Toad FiGuRE 2. Distribution of amphibians in the Yukon. Refer to text for data sources. Shaded areas are mountain ranges. 90 THE CANADIAN FIELD-NATURALIST term prospects of this population are unknown as this small lake has regularly been stocked with Rainbow Trout (Oncorhynchus mykiss) since 1990, with the lat- est stocking occurring in 2003 (D. Toews, personal communication). Rainbow trout are known to prey on Columbia Spotted Frog tadpoles and juveniles, which overwinter in lakes (Pilliod and Peterson 2000). Fur- thermore the heavily forested shoreline habitat is a- typical for Columbia Spotted Frogs (Ovaska 1999*). The species has been observed only on human-altered shoreline which has been cleared of trees and graded into Irons Creek Lake, creating shallows. In 2004, RLM observed an adult Columbia Spotted Frog 15 km west of this site on an ox-bow channel of the Hyland River (60°O1'N 127°12'W). The Columbia Spotted Frog localities in the Liard Basin and Coast Mountain Cordillera are in areas with early deep snow accumulation which likely prevents thick ice formation and permits safe aquatic hibernation. Columbia Spotted Frogs were noticeably abundant in and around pond outlets, where they were invariably perched at the waters edge. Breeding and transforma- tion dates in the Yukon are unknown; however popu- lations in nearby northwestern British Columbia have been observed extensively (Mennell 1998*; B. Slough, unpublished data). Breeding takes place on a single warm day from 18-22 May (six years of observations) in traditional shallow basins along the margins of ponds. Some of the ponds may be partially ice-covered with melting snow surrounding the ponds. They breed com- munally with up to 30 individuals congregating in a few square metres. Males begin calling above water over a larger area and finally call underwater as the aggre- gation consolidates and begins amplexis, leaving a mass of egg clutches unattached in shallow water. Breeding sites are occasionally compromised by falling water levels, leaving egg masses landlocked and stranded. Tadpoles are free-swimming in 14-16 days and meta- morphs appear on pond edges throughout August (Men- nell 1998*). Metamorphs have also been observed in late May, and tadpoles have been observed in mid- September at higher elevation ponds, suggesting that some tadpoles overwinter. Woop FROG (Rana sylvatica) The Wood Frog ranges from northern Alaska and the Yukon through most of Canada below treeline east to the Atlantic and south to the Appalachian Mountains. Disjunct populations occur in several states south of this range. Numerous collections and observations reveal that Wood Frogs are common and abundant in wet- lands across the Yukon below the arctic and alpine tree- line (B. Slough, unpublished data) [Figure 1]. The most northerly location is Ney Khwi Vun (“Frogs in the Lake’), on the Old Crow Flats (68°01'N 140°05'W; Gray and Alt 2000*). Museum collections exist in the University of Kansas Museum of Natural History (n = 1), the American Museum of Natural History (n Vol. 120 = 16), the Canadian Museum of Nature (n=56) and the Auke Bay Laboratory, Alaska Fisheries Science Center (Hodge 1976; n = 2). One of the museum spec- imens from mouth of Coal Creek (AMNH 44751) was originally misidentified as Pseudacris septentrionalis (now P. maculata) until verification as a Wood Frog in 1995 by L. S. Ford Curatorial Associate, AMNH (F. Cook, personal communication to P. Milligan). In July 2000, BGS obtained Wood Frog samples for phy- logeographic studies (J. Irwin and J. Lee- Yaw, unpub- lished data); eight juveniles from Nogold Creek, west of Mayo (63°26'N 135°06'W), and 10 juveniles from south of Whitehorse (60°35’N 134°58’W). The CMN also has tissue from 11 adults and 25 juveniles from “Paddy’s Pond”, Whitehorse, July and August 1992 by Gerry Whitley. The Wood Frog specimens and observations (Fig- ure 1) document an extensive range for the species in the Yukon. Most observations are from road and river access corridors, and gaps in the range exist in inac- cessible areas. We believe that the species occupies suitable habitats below treeline within these gaps. Fur- thermore, the distribution is continuous in areas of ex- tensive wetlands such as the Whitehorse area, where only a small sample of locations was plotted. Yukon Wood Frogs were assigned to the Alaskan phenotype by Martof and Humphries (1959). While we have no data on most visible characters used to differentiate the types, there is a north-south gradient in the frequency occurrence of the mid-dorsal stripe, from approximately 100% in the north and central Yukon to 50% or less in the south. Wood frogs breed in the Yukon as soon as shorelines become ice-free in late April-early May. They breed over a period of up to two weeks, they are frequently solitary breeders, and they do not necessarily use tra- ditional breeding sites, but use any site with shallow water and sparse emergent grass or sedge vegetation. Egg masses are attached singly to grass or sedge stalks. Metamorphs move onto land throughout July. The most common Wood Frog breeding habitats are shallow, permanent and fishless ponds. Neighbouring Species Three species of salamanders occur near the south- western Yukon, but have yet to be recorded there. The Long-toed Salamander (Ambystoma macrodactylum) ranges as far north as the Taku River (Hodge 1976; P. Milligan, personal communication; Mennell and Slough 1998*) and upstream to the junction of the Nakina and Sloko rivers (K. Heinemeyer, personal communication), 110 km from the British Columbia- Yukon border (observation not shown by Matsuda et al. 2006). The Northwestern Salamander (Ambystoma gracile) and Roughskin Newt (Taricha granulosa) oc- cur in Southeast Alaska (MacDonald 2003), but they occupy Pacific coastal habitats not present in the Yukon. 2006 Future Surveys Dedicated amphibian surveys and expert observa- tions have greatly increased our knowledge of the species and their ranges in the Yukon. We presented significant new observations of Western Toads, Colum- bia Spotted Frogs and Wood Frogs for the Yukon, and the first records of Boreal Chorus Frogs. Continued efforts should identify additional populations, their degree of isolation and habitat requirements; all nec- essary for conservation planning. The ranges of the Western Toad, Columbia Spotted Frog and Boreal Cho- rus Frog in the Yukon imply data deficiencies for these species in northern British Columbia. The Columbia Spotted Frog is common on Atlin Lake, south of Atlin (Slough et al. 2002*; not shown by Matsuda et al. 2006). Acknowledgments Funding to BGS and RLM for field surveys was provided by NatureServe Yukon, Yukon Department of Environment; and by Northern Research Endow- ment Grants from the Northern Research Institute, Yukon College. Canadian Parks and Wilderness Soci- ety- Yukon Chapter kindly supported BGS on several wilderness trips. We are grateful to the many individ- uals who reported amphibian sightings. Significant Wood Frog sightings were reported by D. Mossop, J. Meikle, L. Karnis, M. Gill, R. Lamb and G. Whitley. E. Wind and T. Jung reviewed earlier versions of the manuscript. G. Perrier of Dragonfly Maps provided digital cartographic design of the distribution map. We thank P. Milligan for his work as the founding Yukon DAPCAN coordinator. Documents Cited [marked * in text citations] Fournier, M. 1998. Amphibians & reptiles in the Northwest Territories. A brochure prepared for Ducks Unlimited, Environment Canada, Indian and Northern Affairs Canada, and Northwest Territories Resources, Wildlife and Eco- nomic Development, Yellowknife, Northwest Territories. Ecology North, Yellowknife. Gray, D. R., and B. T. Alt. 2000. Resource description and analysis of Vuntut National Park of Canada. Western Cana- da Service Centre, Parks Canada, Vancouver, British Colum- bia. 650 pages. Mennell, R. L. 1998. The ecology and population status of the Columbia spotted frog (Rana luteiventris) at their northern range limit. Prepared for the Northern Research Institute, Yukon College, Whitehorse. 21 pages. Mennell, R. L., and B. G. Slough. 1998. Amphibian and biodiversity inventories of ecoregions in northwestern British Columbia. Habitat Conservation Trust Fund Pro- ject TF28056. Ministry of Environment, Lands and Parks, Victoria, British Columbia. 86 pages. Ovaska, K. E. 1999. Status report on the Columbia spotted frog, Rana luteiventris, in Canada. Committee on the Sta- tus of Endangered Wildlife in Canada. 55 pages. Slough, B. G. 2004. Western toad inventory in the Chilkoot Trail National Historic Site, July-August 2004. Parks Cana- da Species at Risk Inventory Fund Project SARINV04-30. 54 pages. SLOUGH AND MENNELL: AMPHIBIANS OF THE YUKON 9) Slough, B. G. 2005a. Western toad, Bufo boreas, stewardship in the Yukon. NatureServe Yukon, Whitehorse. 26 pages Slough, B. G. 2005b. Assessment of the Impact of Rainbow Trout Introductions on Columbia Spotted Frogs in Lrons Creek Lake, Yukon. Fish and Wildlife Branch, Yukon Department of Environment, Whitehorse. iv + 17 pages Slough, B. G., J. T. Irwin, and D. M. Green. 2002. Post glacial Colonization and Genetic Diversity of the Columbia Spotted Frog at its Northern Range Limit. Progress report, prepared for the Northern Research Institute, Yukon Col lege, Whitehorse. 12 pages. Literature Cited Collins, J. T., and T. W. Taggart. 2002. Standard common and current scientific names for North American amphib- ians, turtles, reptiles and crocodilians, fifth edition. Cen- ter for North American Herpetology. 45 pages. Cook, F. R. 1977. Records of the boreal toad from the Yukon and northern British Columbia. Canadian Field- Naturalist 91: 185-186. Hodge, R. P. 1976. Amphibians and reptiles in Alaska, the Yukon, and Northwest Territories. Alaska Northwest Pub- lishing Co., Anchorage, Alaska. 89 pages. Logier, E. B. S. 1932. Some accounts of the amphibians and reptiles of British Columbia. Contribution Number 5, Royal Ontario Museum of Zoology. Reprinted from Trans- actions of the Royal Canadian Institute, Volume XVIII, Part 2: 311-338. Logier, E. B. S., and G. C. 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A field guide to western reptiles and Agriculture, Odgen, Utah. amphibians. Third edition. Houghton Mifflin Company, Platz, J. E. 1989. Speciation within the chorus frog, Pseu- Boston, New York. xiii + 533 pages. dacris triseriata: morphometric and mating call analyses of Thoms, C., C. C. Corkran, and D. H. Olson. 1997. Basic the boreal and western subspecies. Copeia 1989: 704-712. amphibian survey for inventory and monitoring in lentic Slevin, J. R. 1928. The amphibians of western North America. habitats. Pages 35-46 in Sampling amphibians in lentic Occasional Papers of the California Academy of Sciences habitats. Edited by D. H. Olson, W. P. Leonard and R. B. XVI. 152 pages. Bury. Northwest Fauna Number 4: 1-134. Society for Slough, B. G. 2002. Geographic distribution: Rana luteiven- Northwestern Vertebrate Biology, Olympia, Washington. tris. Herpetological Review 33: 146. Received 10 March 2005 Accepted 30 November 2005 Temporal Variability of Cetaceans near Halifax, Nova Scotia PETER SIMARD!:2, JENNIFER L. LAWLOR?, and SHANNON GowaAns! 4 ' Blind Bay Cetacean Studies, 144 Victoria Road, Dartmouth, Nova Scotia B3A 1V7 Canada + University of South Florida, College of Marine Science, 140 7" Avenue South, St. Petersburg, Florida 33701 USA; e-mail psimard@ marine.usf.edu. ‘Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada ‘Eckerd College, 4200 54" Avenue South, St. Petersburg, Florida 33711 USA Simard, Peter, Jennifer L. Lawlor, and Shannon Gowans. 2006. Temporal variability of cetaceans near Halifax, Nova Scotia. Canadian Field-Naturalist 120(1): 93-99. Annual and seasonal trends in sightings of coastal cetaceans near Halifax, Nova Scotia, were studied using observations from whale watching and dedicated research vessels from late spring to early fall of 1996 to 2005. Four species of cetaceans rou- tinely used the area during the summer: White-beaked and Atlantic White-sided dolphins (Lagenorhynchus albirostris and L. acutus), Harbour Porpoises (Phocoena phocoena), and Minke Whales (Balaenoptera acutorostrata). The dolphin species were temporally separated, with White-beaked Dolphins being common earlier in the summer than White-sided Dolphins. White-sided Dolphins were unusually abundant in 1997, and were found in larger groups (mean = 46.5 + 46.19 sd) than White- beaked Dolphins (mean = 9.1 + 5.19 SD). The area also appears to be an important habitat for dolphin calves and juveniles of both species. Fin Whales (B. physalus) were commonly observed in relatively large groups in 1997, but were uncommon or absent in other years. Humpback Whales (Megaptera novaeangliae), Blue Whales (B. musculus) and North Atlantic Right Whales (Eubalaena glacialis) were uncommon in the area, although Humpback Whales were sighted frequently in 1997. Increased numbers of White-sided Dolphins, Fin and Humpback whales in 1997 may be explained by increased prey abundance and decreased sea-surface temperatures. Key Words: White-beaked Dolphin, Lagenorhynchus albirostris, Atlantic White-sided Dolphin, Lagenorhynchus acutus, Har- bour Porpoise, Phocoena phocoena, Minke Whale, Balaenoptera acutorostrata, Fin Whale, Balaenoptera physalus, Humpback Whale, Megaptera novaeangliae, seasonality, group size, calves, Nova Scotia. In temperate ecosystems, it is common for cetaceans to exhibit seasonal and annual fluctuations in their dis- tribution and abundance (e.g., Hooker et al. 1999). These fluctuations are thought to be a reflection of seasonal changes in the environment, such as sea sur- face temperature (e.g., Hooker et al. 1999) and prey distribution (e.g., Kenney et al. 1996). Monitoring highly mobile, top level predators such as cetaceans serves not only to identify their habitat use patterns and track their population sizes, but also to monitor the health of the overall marine ecosystem. The area between Halifax and Mahone Bay, Nova Scotia, is habitat for several species of cetaceans. It is used extensively by the commercial fishing industry, commercial shipping (especially near Halifax Harbour) and recreational boaters, and it is being used increas- ingly for whale watching. Given the high potential for anthropogenic influences in the Halifax area, there is good reason to study and monitor cetaceans there. Methods Field Methods Data were collected from 1996 to 2005, primarily from 1 June to 30 September. From 1996 to 1999, 12.6 m and 14 m whale watching vessels were used to make three 3-hour trips daily, weather permitting. In 2000 and 2001, an 8.5 m auxiliary sailing vessel was used, and in 2002 to 2005 a 4.2 m rigid inflatable boat was used. The latter two vessels were used only for research, and cruises in 2000 to 2005 were generally conducted only in weather conditions suitable for photo-identification. All surveys were limited to day- light hours and the area bounded by approximately 44°19' to 44°38'N latitude and 63°15' to 64°10'W lon- gitude (Figure 1). The path of all vessels was oppor- tunistic, based on where cetaceans were previously sighted. In 1996, only date, time, species, group size and the presence of calves and juveniles were recorded. In subsequent years, latitude and longitude positions were recorded (using GPS) for hourly search effort and species positions and photo-identification data were col- lected. Only those sightings identified to species level were included in analysis. Animals were classified as adults, juveniles or calves. For odontocetes, first year calves were identified by having a length of <50% of adult body length, while juveniles were identified by having 50% — 70% of adult body length (Reeves et al. 1999; Lien et al. 2001: Bjorge and Tolley 2002). The identification of calves was often facilitated by their position near an adult, with younger calves displaying “chin-slap”™ surfacings, and in the case of neonatal calves, by fetal folds and bent-over dorsal fins (Weinrich et al. 2001). For mys- ticetes, calves were identified by having a length of 94 THE CANADIAN FIELD-NATURALIST <40%, and juveniles by having a length of 40% — 60% of adult body length (Aguilar 2002; Clapham 2002; Perrin and Brownell 2002). Over the ten-year study period, changes in method- ology introduced several biases to our data. Data col- lected from the whale watching boats (1996-1999) were particularly problematic as the boat returned sev- eral times per day with new passengers to the same area where cetaceans had previously been sighted, potentially causing groups to be double-sampled and introducing a positive bias to calculations of hourly sighting rates. Cetacean encounters on the whale watch- ing vessels were typically shorter in duration, and there- fore a larger proportion of time was spent searching for new groups, introducing another potentially positive bias. However, the whale watching vessels were more likely to operate in inclement weather than the re- search vessels, which should negatively bias the sighting data. Larger groups were preferred on whale watching cruises, positively biasing group sizes. Ves- sel cruising speed also varied greatly in the nine year study period (ranging from 5 to 15 knots). These dif- ferences in survey speed changed our spatial sampling greatly, and therefore likely affected the sighting data. The height of observers above the sea surface also var- ied greatly, from a minimum 1.25 m to a maximum 5 m. Search effort varied greatly between different years and months, and as a result time periods with low effort may have had unreliable sighting rates (see Table la and Ib). Calculation of Sighting Indices Because of these biases, regular group sighting rates (sightings per hour) would not be directly comparable. Therefore, the data were restricted in order to calcu- late a “sighting index” for each species per year and per month. Potential positive biases from repeated sampling of the same cetaceans throughout the day were eliminated by daily scoring each species as pres- ent or absent. This restriction also reduces biases intro- duced from differences in vessel speed, and time spent searching vs. time spent with groups of cetaceans. Cal- culated sighting indices could range from a maximum of 1.0 (sightings occurring every day of field effort) to 0.0 (no sightings). In order to determine if weather significantly affected sighting rates, the sighting rates during good weather and bad weather were subjected to a chi-squared test. Good weather was defined as visibility = 1 km, wind < Beaufort 3 (< = 19 km/hr), wave height < 0.75 m, and swells < 1.5 m. Results from this test indicate that sighting rates were signifi- cantly lower in poor weather (y? = 0.9999, df = 5, 0.025 75%. Coniferous Crown closure 2 10%, coniferous species > 75%. Mixedwood Crown closure 2 10%, neither type > 75% Habitat type Description Immature-Pole Young Mature-old 0-40 years old stands. Open areas and new stands. Pole stands are thick stands of trees (7.5 to 12.4 cm diameter at breast height), usually with little understory. 40-80 year-old forests. Achievement of dominance by some trees and death of other trees leads to reduced competition that allows understory plants to become established. The forest canopy has begun differentiation into distinct layers. Vigorous growth and a more open and multi-storied stand than in the pole stage. > 81 years old late successional stands consisting mostly of mature stands with even canopy of trees, with or without coarse woody debris down and leaning logs. A few old stands with tall and large canopy trees, canopy gaps, large snags, large downed woody debris, and developed understories. A second cycle of shade tolerant trees may have become established. to identify animals. When encountering animals, foot- prints and trails were studied to ascertain criteria used to differentiate American Marten from Fisher. Autocorrelation may occur during analysis of track survey data because of the uncertainty in whether one or more animals have made the tracks being counted (Proulx and O’Doherty 2006). It is sometimes difficult to confirm that a series of tracks along a transect be- longs to the same animal (de Vos 1951), as home ranges overlap (Buskirk and Ruggiero 1994), and winter dis- persal movements are known to occur (Clark and Campbell 1976). On the basis of track characteristics, Proulx et al. (2005) deduced that tracks of two different animals could be as close as 100 m from each other along the same transect. To minimize spatial autocor- relation, only tracks >100 m apart within the same for- est stand were recorded (Bowman and Robitaille 1997; Proulx et al. 2006). In western North America, American Martens ap- pear to be associated with late successional stands. Consequently, I hypothesized that American Marten tracks would not be distributed at random among habi- tat types, but that they would be more frequent in mature and old-growth stands. My hypothesis was declared before examining the track survey data. The proportion of inventory transects within each habitat type was used to determine the expected frequency of track intersects/habitat type (i.e., availability) if tracks were distributed randomly with respect to habitat types (Proulx et al. 2006). For each survey, I used a one- tailed Fisher Probability test (Siegel 1956) to compare the proportion of American Marten tracks in the habi- tat growth types, i.e., mature/old vs. immature/young. Data from the three inventories were pooled to increase sample size. Chi-square statistics with Yates correc- tion (Zar 1999) were used to compare observed to expected frequencies of track intersects among habi- tat type. If the chi-square analysis suggested an over- all significant difference between the distribution of observed and expected frequencies, comparisons of observed to expected frequencies for each habitat class were conducted using the G test for correlated proportions (Sokal and Rohlf 1981). Probability values <0).05 were considered statistically significant. Results Temperatures ranged from -21°C to 0°C and snow depths exceeded 45 cm during each inventory. Inven- tories were conducted < 24 h after snowfalls or flurries, thus assuring that tracks were fresh. In 1999, 41.1 km of transects were inventoried: 27.8 km (67.6%) in immature/young stands, and 13.3 km (32.4%) in mature/old stands. Eight tracks were recorded: 3 (37.5%) in young stands, and 5 (62.5%) in mature-old forests. The observed distribution of tracks was not significantly (P = 0.31) different from a random distribution of tracks among habitat types. In 2002, 36.0 km of transects were inventoried: 16.6 km (46%) in immature/young stands, and 19.4 km (54%) in mature/old stands. Ten tracks were recorded: 1 (10%) in a young stand, and 9 (90%) in mature/old forests. The observed distribution of tracks was not sig- nificantly (P = 0.07) different from a random distribu- tion of tracks among habitat types. In 2005, 51.1 km of transects were inventoried: 27.1 km (53%) in immature/young stands, and 24 km (47%) in mature/old forests. Twenty-six tracks were recorded: 6 (23%) in young stands, and 20 (77%) in 006 PROULX: WINTER-HABITAT USE BY AMERICAN MARTEN 103 20 18 218 5 14 = ee E10 eo §8 = 6 = 5 4 z= 2 0 1 ~” aw} n Mo} ~ SS no) o 3 =| = 3 = = 2 Pee a, PS Ove ni pote. (Qive ae & D c z as = Eas 1 z “eee. ie) 5 Ga 3 ov 2 o& 2s 23 & jam > ‘S omm.s a oO = owt =) oO = tS Sap Aes ee = 2 ee Habitat types IGURE 2. Observed and expected number of American Marten tracks per habitat type in Weyerhaeuser’s Grande Prairie Forest Management Area. jature-old forests. The observed distribution of tracks vas significantly (P = 0.002) different from a random istribution of tracks among habitat types. A total of 128.2 km of transects were inventoried rom 1999 to 2005: 39.1 km (30.5%) in immature/pole tands, 32.5 km (16.9 % in coniferous, 5.8% in mixed oniferous-deciduous, and 2.7% in deciduous) in young tands, and 44.2 km (18.8% in coniferous, 13.5% in nixed coniferous-deciduous, and 11.9% in deciduous) n mature/old forests. Forty-four tracks were recorded: one in immature/pole stands, 10 tracks (22.7%) in oung coniferous forests, and 34 tracks (77.3%) in nature/old stands (Figure 2). The observed distribution f American Marten tracks was significantly different rom a random distribution of tracks among habitat ypes (x7 = 39, df: 4, P < 0.001) (Figure 2). Tracks were ignificantly less frequent than expected in immature/ ole stands (G = 17.2, df: 1, P < 0.001), and more requent than expected in mature/old mixedwood stands G = 6.42, df: 1, P < 0.02). The distribution of tracks uggested that American Martens used young forests, nd mature/old coniferous and deciduous stands, ac- ording to their availability. Jiscussion In 1999 and 2002, because of small sample sizes, no ignificant difference was found in the distribution of \merican Marten tracks among habitat types. Howev- r, during both years, most tracks were in mature-old orests. Pooling data increased sample size and statisti- al power, and showed that American Marten tracks were definitely more frequent in late-successional stands. This finding is in agreement with previous stud- ies conducted in western coniferous forests (Spencer et al. 1983; Wilbert 1992; Koehler et al. 1990; Therrien 2002; Proulx et al. 2006). American Martens prefer mature and old-growth forests over young and imma- ture forests because this is where they maximize for- aging efficiency and finding rest sites that can protect them from harsh environmental conditions Powell et al. 2003). Although young forests are believed to be subopti- mal due to a reduction of necessary structure and diver- sity compared to mature and decadent forests (Thomp- son and Harestad 1994), in this study, nearly a quarter of American Marten tracks were located in young coniferous stands. As late/successional stands are decreasing in importance in Weyerhaeuser’s FMA due to timber harvest, and oil and gas exploitation, Ameri- can Martens may have to make greater use of young forests. However, previous studies showed that Ameri- can Martens used young forests in winter where suffi- cient physical structure (Bowman and Robitaille 1997; Poole et al. 2004) and prey (Potvin et al. 2000) were present. More research on the characteristics of young forests used by American Martens is required to better understand their importance as winter habitat for this species in boreal ecosystems. The fact that no tracks were recorded in immature and pole stands is in agreement with previous studies that have generally shown that American Martens make little or no use of early/successional stages \C.£., 104 (Steventon and Major 1982; Thompson 1994; Thomp- son and Harestad 1994; Chapin et al. 1998; Heinemey- er 2002; Poole et al. 2004). Previous studies have also shown that American Martens appear intolerant of habi- tat fragmentation, and they would not tolerate more than 30-35% cutovers in their home range (Snyder and Bissonette 1987; Hargis and Bissonette 1997; Chapin et al. 1998; Potvin et al. 2000; Poole et al. 2004). In Weyerhaeuser’s FMA, mature and old forests may be highly fragmented due to extensive clearcuts, and an important network of seismic lines and roads. Un- doubtedly, these habitat modifications would have an impact on the distribution of American Martens, and cause shifts in home range boundaries (Poole et al. 2004). Forest development plans taking into consid- eration the winter habitat requirements of American Marten should therefore be developed locally accord- ing to the distribution of late/successional forests, cut blocks, and access networks (e.g., Proulx 2001*). Acknowledgments I thank Luigi Morgantini, Wendy Crosina, Dan Beil- man, and Darren Lapp from Weyerhaeuser Company Ltd., and Pauline Feldstein from Alpha Wildlife, for helping with the project logistics. I also thank Vivian Banci, Valeria Vergara, and Jamie Farkvam for field assistance, and two anonymous referees for helpful comments. Documents Cited (marked * in text) Alberta Government. 1999. Alberta natural regions. Alberta Environment Protection, Edmonton, Alberta. Proulx, G. 2001. Characteristics and management of Amer- ican marten habitat at stand and landscape levels. Alpha Wildlife Research & Management Ltd. report prepared for British Columbia Ministry of Forests, Prince George Forest District, Prince George, British Columbia. Literature Cited Bowman, J. C., and J-F. Robitaille. 1997. Winter habitat use of American Martens Martes americana within second- growth forest in Ontario, Canada. Wildlife Biology 3: 97-105. Buskirk, S. W., and L. F. Ruggiero. 1994. American Marten. Pages 7-37 in American marten, fisher, lynx, and wolverine in the Western United States. Edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. J. Lyon, and W. J. Zielinski. United States Department of Agriculture, Forest Service, General Technical Report RM-254, USA. Campbell, T. M. 1979. Short-term effects of timber harvests on pine marten ecology. M.Sc. thesis, Colorado State Uni- versity, Fort Collins, Colorado. Chapin, T. G., D. J. Harrison, and D. D. Katnik. 1998. Influence of landscape pattern on habitat use by American marten in an industrial forest. Conservation Biology 12: 1327-1337. Clark, T. W., and T. M. Campbell. 1976. Population organi- zation and regulatory mechanisms of pine Martens in Grand Teton National Park, Wyoming. Pages 293-295 in Conference on scientific research in national parks. Volume 1. Edited by R. M. Linn, National Park Service Transac- tions Proceedings Series 5, Washington, D.C. THE CANADIAN FIELD-NATURALIST Vol. 120 de Vos, A. 1951. Tracking of fisher and marten. Sylva 7: 15-19. Halfpenny, J. C., R. W. Thompson, S. C. Morse, T. Holden, and P. Rezendes. 1995. Snow tracking. Pages 91-163 in American marten, fisher, lynx and wolverine: survey meth- ods for their detection. Edited by W. J. Zielinski and T. E. Kucera. United States Department of Agriculture, Forest Service Pacific Southwest Research Station General Tech- nical Report PSW-GTR-157. Hargis, C. D., and J. A. Bissonette. 1997. Effects of forest fragmentation on populations of American marten in the intermountain west. 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Budd’s flora of the Canadi- an Prairie Provinces. Agriculture Canada, Research Branch. Publication 1662. Murie, O. J. 1975. A field guide to animal tracks. The Peterson’s field guide series. Houghton Mifflin Co., Boston. Massachusetts, USA. Poole, K. G., A. D. Porter, A. deVries, C. Maundrell, S. D. Grindal, and C. C. St. Clair. 2004. Suitability of a young deciduous-dominated forest for American marten and the effects of forest removal. Canadian Journal of Zoology 82: 423-435. Potvin, F., L. Bélanger, and K. Lowell. 2000. Marten habitat selection in a clearcut boreal landscape. Conservation Biol- ogy 14: 844-857. Powell, R. A.,S. W. Buskirk, and W. J. Zielinski. 2003. Fisher and marten. Pages 635-649 in Wild mammals of North America. 2" edition. Edited by G. A. Feldhamer, B. C. Thompson, and J. A. Chapman.The Johns Hopkins Univer- sity Press, Baltimore, USA. Proulx, G., K. B. Aubry, J. Birks, S. W. Buskirk, C. Fortin, H. C. Frost, W. B. Krohn, L. Mayo, V. Monakhoy, D. Payer, M. 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Biostatistical analysis. 4" edition, Prentice Hall, New Jersey. Received 9 June 2005 Accepted 17 October 2005 Notes Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine J.C. A. BuRCHSTED! and FRED BURCHSTED2 ' Department of Biology, Salem State College, Salem, Massachusetts 01970 USA * Research Services, Widener Library, Harvard University, Cambridge, Massachusetts 02138 USA Burchsted, J.C. A., and Fred Burchsted. 2006. Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine. Canadian Field-Naturalist 120(1): 106-108. The Lady Crab (Ovalipes ocellatus), mainly found south of Cape Cod and in the southern Gulf of St. Lawrence, is reported from an ocean beach on the north shore of Massachusetts Bay (42°28'60"N, 70°46'20"W) in the Gulf of Maine. All previ- ously known Gulf of Maine populations north of Cape Cod Bay are estuarine and thought to be relicts of a continuous range during the Hypsithermal. The population reported here is likely a recent local habitat expansion. Key Words: Lady Crab, Ovalipes ocellatus, Gulf of Maine, distribution. The Lady Crab (Ovalipes ocellatus) is a common member of the sand beach fauna south of Cape Cod. Like many other members of the Virginian faunal province (between Cape Cod and Cape Hatteras), it has a disjunct population in the southern Gulf of St. Lawrence (Ganong 1890). The Lady Crab is of consid- erable ecological importance as a consumer of mac- robenthic invertebrates, and is of biogeographic inter- est as a member of a large disjunct fauna. However, the distribution of the Lady Crab in the Gulf of Maine is unclear from the literature. Here we gather and dis- cuss available reports of the Lady Crab in the Gulf of Maine and report an additional site that may represent a recent habitat expansion. This paper is intended as a basis for further work on Lady Crab distribution and for monitoring changes. Lady Crabs have not generally been reported from the Gulf of Maine. Field guides (Gosner 1978; Mein- koth 1981; Miner 1950) and a review of tidal flats (Whitlatch 1982) give their northern limit as Cape Cod except for the disjunct Gulf of St. Lawrence popula- tion. Recent distributional studies of Gulf of Maine crabs show Lady Crabs on the south shore of Cape Cod and onto Georges Bank but no further north (Williams and Wigley 1977; Stehlik et al. 1991). This confirms the older work of Rathbun (1930). Williams (1984), and the update by Nizinski (2003), give the range briefly as Prince Edward Island to Georgia, but in this they may elide the northern New England gap. Pol- lock (1998) states the range as Virginian and Acadi- an, but again the range between the southern Gulf of St. Lawrence and Cape Cod is unspecified. The Lady Crab is not listed in Bay of Fundy, Maine, and New Hampshire checklists (Croker 1972; Brinkhurst et al. no date; Linkletter et al. 1977; Perkins and Larsen 1975) and distributional studies of the fauna of Maine sand beaches (Larsen and Doggett 1990) and mud flats (Larsen and Doggett 1991). Lady Crabs were not found in intensive local studies of western Cape Cod Bay (Davis and McGrath 1984) or Ipswich Bay (Dexter 1944). Berrick (1986) reports Lady Crabs as common on Cape Cod Bay sand flats (which commonly reach 20°C in summer). The few other published records of Lady Crabs in the Gulf of Maine are from warm, estuarine habitats. Bousfield and Thomas (1975) identify a group of three warm water species which tend to co-occur (Balanus improvisus, Urosalpinx cinera, and Ovalipes ocellatus) but do not specify locations for the Lady Crab. Members of this group occurred in 1963 at: Cape Cod Bay, Duxbury and Cape Ann (Massachusetts); Cape Neddick and Boothbay Harbor (southern Maine); and the Bay of Fundy. Bousfield found Lady Crabs at Sesuit Harbor, East Dennis, Massachusetts, in 1964 (Judith Price, personal communication). Lady Crabs are reported in Salem Sound, Massachusetts, in 1997 by Chase et al. (2002). Although Dexter (1947) found no Lady Crabs in an intensive study of a tidal inlet at Annisquam (Gloucester, Massachusetts) in 1933-1940, he later (Dexter 1985) found them there during what he terms a period of warm water from 1952 to 1964. We have found Lady Crab shells washed up on an ocean beach adjacent to the Essex and Ipswich river estuaries (Crane Beach, Ipswich, Massachusetts, 42°41'30"N, 70°46'20"W; 1995 and 1996). Fefer and Schettig (1980) mention, but do not specify, one or more southern Maine (south of Saco) locations from unpublished data by Doggett, Larsen, and Sykes. They were not found in the 1873 U.S. Fisheries Bureau investigation of Casco Bay which discovered several other warm water species (Verrill 1874). There are no records for the northern Maine cold water region (north of the Sheepscot River estuary) identified by Bousfield and Laubitz (1972) and confirmed by Larsen and 106 2006 ss Doggett (1990). In the northern Gulf of Maine, there are several reports of Lady Crabs in the Bay of Fundy: Bousfield and Leim (1960) report Lady Crabs in the Minas Basin and Channel in 1922 (Bass River and Scotman Bay, noted also in Rathbun 1930) and again in 1959 (Bass River, Diligent River, and a cheliped on beach at Kingsport). Bousfield and Thomas (1975) state that their Gulf of Maine records of Lady Crabs are from warm (circa 20°C summer temperatures — similar to their range south of Cape Cod) brackish water pockets or from estuaries with two-layer circulations where endemic warm-water populations could be maintained as des- cribed by Bousfield (1955) and Sponaugle et al. (2002). Bousfield and Thomas (1975) suggest these estuarine populations are relicts from the Hypsithermal warm era (9500-3000 years Before Present). The many Vir- ginian invertebrates and marine algae now extending north to Cape Cod with additional disjunct populations in the southern Gulf of St. Lawrence had continuous ranges at that time. The short abstract by Dexter (1985) provides no details, but his 1952-1964 warm period may correspond to the 1951-1954 warming reported by Loder et al. (2001), with the crabs becoming local- ly extinct during the succeeding cold period (1959- 1967). Dexter’s observations suggest the possibility that the estuarine distribution is dynamic, populations becoming extinct and reestablished with changing conditions. For the ten years we have been keeping records, Lady Crabs have been commonly found living on an ocean beach, called Preston Beach in Marblehead and Phillips Beach contiguously to the south in Swamp- scott, on the north shore of Massachusetts Bay (42°28'60"N, 70°46'20"W). There is no nearby estu- arine habitat and the beach is exposed to heavy surf (unlike the Cape Cod Bay flats) [Leo et al. 2003]. Salem Sound is about six coastline miles to the north around the headlands of Marblehead/Marblehead Neck. This is the first reported exposed ocean beach location for Lady Crabs in the Gulf of Maine. Phillips Beach is in an area of wind-driven summer upwelling that keeps Summer temperatures around 15-16°C (Geyer et al. 1992). Lady Crabs were commonly found swimming in shallow water and buried in sand just above low tide line. The habitat is thus very like that of the Lady Crab south of Cape Cod. Most of the living animals were found on the southern portion of the beach. This area is influenced by groundwater seepage on the high beach. Old faunal lists for Swampscott and vicinity yield no records of Lady Crabs: Grabau (1898) for Nahant, Revere, and Phillips Beach; Pearse (1913) for Nahant. This suggests our Massachusetts population may mark a local habitat expansion. Discovery of other ocean beach populations and determination of the status of the estuarine populations would help clarify the distribution and habitat associ- ations of Lady Crabs in the Gulf of Maine. NOTES 107 Acknowledgments We thank E. L. Bousfield for information on his col- lections and Judith C. Price of the Canadian Museum of Nature for searching the collections for Lady Crab records. Literature Cited Berrick, S. 1986. Crabs of Cape Cod. Cape Cod Museum of Natural History, Brewster, Massachusetts. 76 pages. Bousfield, E. L. 1955. Ecological control of the occurrence of barnacles in the Miramichi estuary. National Museum of Canada, Bulletin, Biological Series 137.65 pages. Bousfield, E. L., and D. R. Laubitz. 1972. Station lists and new distributional records of littoral marine invertebrates of the Canadian Atlantic and New England Regions. National Museums of Canada, Publications in Biological Oceanography (5). 51 pages. Bousfield, E. L., and A. H. Leim. 1960. The fauna of Minas Basin and Minas Channel. National Museum of Canada Bulletin 166. 30 pages. Bousfield, E. L., and M. L. H. Thomas. 1975. Postglacial changes in distribution of littoral marine invertebrates in the Canadian Atlantic region. Pages 47-60 in Environ- mental change in the Maritimes. Proceedings of the Nova Scotia Institute of Science 27 (Supplement 3). Brinkhurst, R. O., L. E. Linkletter, E. I. Lord, S. A. Con- nors, and M. J. Dadswell. No date. A preliminary guide to the littoral and sublittoral marine invertebrates of Pas- samaquoddy Bay. Environment Canada Biological Station, St. Andrews, New Brunswick. 166 pages. [Published after 1975] Chase, B. C., J. H. Plouff, and W. M. Castonguay. 2002. The marine resources of Salem Sound, 1997. Massachusetts Division of Marine Fisheries Technical Report TR-6. 143 pages. Croker, R. A. 1972. Checklist with habitat notes, of some common intertidal, estuarine, and nearshore invertebrate animals of New Hampshire and Southern Maine. Jackson Estuarine Laboratory, University of New Hampshire, Dur- ham, New Hampshire. 25 pages. Davis, J. D., and R. A. McGrath. 1984. Some aspects of nearshore benthic macrofauna in western Cape Cod Bay. Pages 77-102 in Observations on the ecology and biolo- gy of western Cape Cod Bay, Massachusetts. Edited by J. D. Davis and D. Merriman. (Lecture notes on coastal and estuarine studies 11). Springer-Verlag, Berlin, New York. 289 pages. Dexter, R. W. 1944. The bottom community of Ipswich Bay, Massachusetts. Ecology 25: 352-359. Dexter, R. W. 1947. The marine communities of a tidal inlet at Cape Ann, Massachusetts: A study in bio-ecology. Eco- logical Monographs 17: 261-294. Dexter, R. W. 1985. Invasions of southern marine fauna into Cape Ann, Massachusetts, during periods of warmer sea-water. American Zoologist 25 (4): A64. Fefer, S. I., and P. A. Schettig. 1980. An ecological charac- terization of coastal Maine. Volume 4: Appendices. Pub- lication FWS/OBS-80/29, Biological Services Program, US. Fish and Wildlife Service. Ganong, W. F. 1890. Southern invertebrates on the shores of Acadia. Transactions of the Royal Society of Canada 8: 167-185. Geyer, W. R., G. B. Gardner, W. S. Brown, J. Irish, B. Butman, T. Loder, and R. Signell. 1992. Physical ocean- 108 ographic investigation of Massachusetts and Cape Cod Bays. Massachusetts Bays Program, Boston, Massachu- setts. 497 pages. Gosner, K. L. 1978. A field guide to the Atlantic seashore. Houghton Mifflin, Boston, Massachusetts. 329 pages. Grabau, A. W. 1898. Zoology: Marine invertebrates. Pages 67-94 in Guide to localities illustrating the geology, marine zoology and botany of the vicinity of Boston. Edited by A. W. Grabau and J. E. Woodman. American Association for the Advancement of Science Fiftieth Anniversary Meet- ing, Boston, August, 1898. Larsen, P. F., and L. F. Doggett. 1990. Sand beach macro- fauna of the Gulf of Maine with inference on the role of oceanic fronts in determining community composition. Journal of Coastal Research 6: 913-926. Larsen, P. F., and L. F. Doggett. 1991. The macroinvertebrate fauna associated with the mud flats of the Gulf of Maine. Journal of Coastal Research 7: 365-375. Leo, W., R. Geyer, and M. Mickelson. 2003. Physical and biological oceanography of Massachusetts Bay. Pages 4- 1 — 4-22 in Briefing for OMSAP Workshop on Ambient Monitoring Revisions, June 18-19, 2003. Report ENQUAD ms-085, Massachusetts Water Resources Authority, Boston, Massachusetts. 250 pages. Linkletter, L. E., E. 1. Lord, and M. J. Dadswell. 1977. A checklist of the marine fauna and flora of the Bay of Fundy. Huntsman Marine Laboratory, St. Andrews, New Brunswick. 68 pages. Loder J. W., J. A. Shore, C. G. Hannah, and B. D. Petrie. 2001. Decadal-scale hydrographic and circulation variabil- ity in the Scotia-Maine region. Deep-Sea Research Part II- Topical Studies in Oceanography 48: 3-35. Meinkoth, N. A. 1981. The Audubon Society Field Guide to North American seashore creatures. Knopf, New York. 799 pages. Miner, R. W. 1950. Field book of seashore life. G. P. Putnam’s Sons, New York. 888 pages. Nizinski, M. S. 2003. Annotated checklist of decapod crus- taceans of Atlantic coastal and continental shelf waters of the United States. Proceedings of the Biological Soci- ety of Washington 116: 96-157. THE CANADIAN FIELD-NATURALIST Vol. 120 Pearse, A. S. 1913. Observations on the fauna of the rock beaches at Nahant, Massachusetts. Bulletin of the Wiscon- sin Natural History Society (n.s.) 11: 8-34; 12: 72-80. Perkins, L. F., and P. F. Larsen. 1975. A preliminary check- list of the marine and estuarine invertebrates of Maine. TRIGOM Publication No. 10. 37 pages. Pollock, L. W. 1998. A practical guide to the marine animals of northeastern North America. Rutgers University Press, New Brunswick, New Jersey. 367 pages. Rathbun, M. J. 1930. The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. U.S. National Museum Bulletin 152. 609 pages. Sponaugle, S., R. K. Cowen, A. Shanks, S. G. Morgan, J. M. Leis, J. S. Pineda, G. W. Boehlert, M. J. Kingsford, K. C. Lindeman, C. Grimes, and J. L. Munro. 2002. Predicting self-recruitment in marine populations: Bio- physical correlates and mechanisms. Bulletin of Marine Science, 70 (Supplement S.): 341-375. Stehlik, L. L., C. L. MacKenzie, Jr., and W. W. Morse. 1991. Distribution and abundance of four brachyuran crabs on the northwest Atlantic shelf. Fishery Bulletin 89: 473- 492. Verrill, A. E. 1874. Explorations of Casco Bay in 1873. Pro- ceedings of the American Association for the Advance- ment of Science (Portland Meeting), 22: 340-395. Whitlatch, R. B. 1982. The ecology of New England tidal flats: a community profile. Publication FWS/OBS-81/01, Office of Biological Services, Fish and Wildlife Service, Washington, D.C. 125 pages. Williams, A. B. 1984 Shrimps, lobsters, and crabs of the Atlantic coast of the Eastern United States, Maine to Flori- da. Smithsonian Institution Press, Washington, D.C., 550 pages. Williams, A. B., and R. L. Wigley. 1977. Distribution of decapod crustacea off northeastern United States based on specimens at the Northeast Fisheries Center, Woods Hole, Massachusetts. NOAA Technical Report NMFS Circular 407. 44 pages. Received 5 August 2004 Accepted 8 February 2006 2006 NOTES 109 Ring-billed Gull, Larus delawerensis, Food Piracy on Diving Ducks WILLIAM J. WALLEY 222 Bossons Avenue, Dauphin, Manitoba R7N OR2 Canada Walley, William J. 2006. Ring-billed Gull, Larus delawerensis, food piracy on diving ducks. Canadian Field-Naturalist 120(1) 109-110. One of four Ring-billed Gulls observed 26 April 2002 at Dauphin Lake, Manitoba attacked a group of diving ducks and took abandoned food items. Key Words: Ring-billed Gull, Larus delawerensis, Common Goldeneye, Bucephala clangula, kleptoparasitism, Manitoba. Kleptoparasitism or piracy is the forceful taking of food by one species of bird from another (Terres 1980). According to Bent (1921) Ring-billed Gulls (Larus delawerensis) have been observed to hover over feed- ing ducks, notably Red-breasted Mergansers (Mergus serrator) darting at them as they re-surfaced apparent- ly to steal their prey although success is not reported. Although Ring-billed Gulls do not usually pirate other gull species (Ryder 1993) they have been observed pirating food from European Starlings, Sturnus vul- garis (Burger and Gochfeld 1981). Clapp et al. (1983) state that a variety of species are pirated by the Ring- billed Gull. Jarvis and Southern (1976), in their study of the food habits of this gull in the Great Lakes region, made no mention of piracy or attempted pira- cy during the breeding season. Crescent Cove at the south end of Dauphin Lake (51°10'N; 99°53'W) in west central Manitoba was windless and sunny but cold (-11°C) at mid morning on 26 April 2002. There, 15 m from shore, a stretch of water 70 m in length and 30 to 50 m in width had opened up parallel to a sandy beach. Small numbers of grebes and diving ducks, about 5 and 25 respectively, swam about on the calm surface, some diving for food. Ring-billed Gulls were also on the water, stood on the edge of the ice, or flew around in the area. Upon my arrival on the nearby ridge above the beach 2 Red-necked (Podiceps grisegena), 3 Horned grebes (Podiceps auritus), 3 Redheads (Aythya americana), and 5 Canvasbacks (Aythya valisneria), and unhurriedly swam away leaving a half dozen Common Goldeneyes (Bucephala clangula) and 3 Greater Scaup (Aythya marila). With the ducks diving and returning to the surface, a Ring-billed Gull, only one of four gulls believed to actually be an attacker, typically took wing off the surface from among the waterfowl and flew low toward a duck up to 15 or 20 m away where it had just surfaced. Upon reaching the targeted bird, the gull flew up and hovered just above and behind its head as described by Bent (1921). The duck plunged out of sight and the gull, using its bill, snatched the food item surrendered. One attack was initiated from as close as 1.5 m. Often a duck surfaced without prey and with sev- eral ducks diving it was remarkable how quickly and efficiently the gull identified a successful duck from the others and attacked it. Two prey items were iden- tified: crayfish (Decapoda) and small fish which, when plucked from the water, were eaten on the spot after the gull alighted on the surface nearby or flew to a shelf of ice where the item was tossed down, re-grasped, then swallowed. During the forty minute observation, attacks on the diving ducks were initiated 20 — 25 times. With one exception — a female scaup — all attacks targeted the goldeneyes. Other than one instance where four male goldeneyes directed a threat display involving the thrusting of their heads low over the water toward a nearby gull, little aggression was shown by the ducks. Avoidance of contact and hence injury from the gull attacks appeared to be paramount for the ducks. Frus- tration on the part of the ducks was not in evidence, but determination to capture more prey was. In April at Dauphin Lake, carrion, notably discard- ed coarse fish from ice fishing activities, would seem to be an important source of food for early arriving carnivorous or omnivorous birds from the south such as gulls, Bald Eagles (Haliaeetus leucocephalus), American Crows (Corvus brachyrhyncos) and some species of hawks, in addition to resident populations of the Common Raven (Corvus corax). A possible shortage of fish and/or other sources of food in the Crescent Cove area of the lake and, perhaps, on the lake ice generally due to competition, may have been a factor in the gulls’ attacks on the goldeneyes. Ob- servations of the lake from shore at Crescent Cove with the use of a 20 power spotting scope that day revealed only one such food item, the carcass of a Canada Goose (Branta canadensis) lying on the ice 250 m out. In immediate succession at this source of carrion, includ- ed: a Northern Harrier (Circus cyaneus) with two crows nearby; an adult Bald Eagle that causing the harrier to fly off and three immature Bald Eagles that displaced the adult eagle at the carcass; all to the ex- clusion of any gulls, suggesting intensive competition for food. This competition may have resulted in the unusual method of securing food observed in at least one Ring-billed Gull. Acknowledgments I thank Ken Kingdon of Parks Canada, Riding Mountain National Park, for literature concerning the 110 Ring-billed Gull — Red-breasted Merganser record. Appreciation is also extended to A. J. Erskine and other reviewers whose constructive comments were well received. Literature Cited Bent, A. C. 1921. Life histories of North American gulls and terns. U.S. National Museum Bulletin 113. Washing- ton, D.C. Burger, J., and M. Gochfeld. 1981. Age-related differences in piracy behavior of four species of gulls, Larus. Behav- ior 77: 242-267. Clapp, R. B., D. Morgan-Jacobs, and R. C. Banks. 1983. Marine birds of the southeastern United States and Gulf of Mexico. Part 3: Charadriiformes. U.S. Fish and Wildlife THE CANADIAN FIELD-NATURALIST Vol. 120 Service, Biological Service, FWS-OBS-83/30. Washing- ton, D.C. Jarvis, W. L., and W. E. Southern. 1976. Food habits of ring-billed gulls breeding in the Great Lakes region. Wil- son Bulletin 88: 621-631. Ryder, J. P. 1993. Ring-billed Gull. The Birds of North Ameri- ca. Number 33. The American Ornithologists’ Union, Wash- ington, D.C. 22 pages Terres, J. K. 1980. The Audubon society encyclopedia of North American birds. Alfred A. Knopf, New York. 1109 pages. Received 28 February 2005 Accepted 2 January 2006 Probable Black Bear, Ursus americana, Retrieval of an Elk, Cervus elaphus, Carcass from a Small Lake in Riding Mountain National Park, Manitoba WILLIAM J. WALLEY 222 Bossons Avenue, Dauphin, Manitoba R7N OR2 Canada Walley, William J. 2006. Probable Black Bear, Ursus americana, retrieval of an Elk, Cervus elaphus, carcass from a small lake in Riding Mountain National Park, Manitoba. Canadian Field-Naturalist 120(1): 110-112. Strong circumstantial evidence indicated that a sow Black Bear (Ursus americana) retrieved an Elk (Cervus elaphus) carcass from near the middle of a small lake to feed herself and three cubs. Key Words: Black Bear, Ursus americana, Elk, Cervus elaphus, carcass, small lake, Riding Mountain National Park. Accounts of observations of Black Bears (Ursus americana) retrieving carcasses from bodies of water are difficult to locate, even those based on circumstan- tial evidence. Studies of food habits of Black Bears by Hatler (1972) and Machutchon (1989) cite no such instances. Herein is provided a description of a series of observations that lead to the conclusion that an Elk (Cervus elaphus) carcass was brought to shore from mid-lake by a female Black Bear. The observation reported was made at Grayling Lake (50°47"N, 100°W) in Riding Mountain Nation- al Park, Manitoba. This is a kettle lake (Lang 1974) and as such is deep (9 m) for its small surface of approximately 25 hectares. It is situated well below surrounding rolling hills in mixed forest about 9 km north of Clear Lake. With the exception of an 80-m gap to the NW and the lake outlet through sedge and grass meadows to the north, tall spruce (Picea) trees encircle and shelter Grayling Lake. At 0815 on 2 August 2004 under full sunshine a large object was seen floating just west of the middle of the lake by the author and two companions from north-south orientated Highway 10 that passes just to the west of the lake (Figure 1). Identification of the brown-gray object was not made even with 8 x 40 binoculars. At 12:15 that day the lake was re-visited. Though air movement was Table 1. Wind direction and speed in km/hr in Riding Moun- tain National Park 2 August 2004* Time At Wasagaming At old Kippen’s mill site 08:00 NW 6 SW 3.6 09:00 NW 4 NW 63 10:00 NW 9 NW 8.1 11:00 NW 7 WNW 7.8 12:00 NW 11 NW 8.6 *Recorded by Environment Canada (personal communica- tion, K. Kingdon, 2005) almost negligible, the object had been moved close to the east shore, where two Coyotes (Canis latrans) fed on it. Some 50 m to the north, a Black Bear was un- hurriedly walking north away from the carcass and the Coyotes. It then disappeared into heavy coniferous forest. Shortly after, the Coyotes moved off and a bear appeared at a small partial clearing above the shore and the carcass. In subsequent days the bear, from this point, repeatedly chased off Coyotes but tolerated the intensive food collecting activity of a pair of Common Ravens (Corvus corax). On 3 August a sow led her three cubs to the carcass, where they fed. In the next few days the remains of the carcass were identified as cow Elk (Cervus elaphus) by Ken Kingdon 2006 AN WIND OUT OF NW I NOTES 11] observation points COW ELK CARCASS STRANGE "OBJECT" | 1215 A 3 815 oO 50° ' dock / HWY 10 GRAYLING LAKE /_—_______ 200 m i 3 = Expected location of carcass if blown by wind FIGURE |. Elk carcass locations relative to wind direction, Grayling Lake, 2 August 2004. of Parks Canada (personal communication) and inde- pendently by the author. It was hypothesized that the bear had retrieved the carcass from the lake and had to swim out at least 200 m to get it. Veteran Parks Canada Warden Pat Rousseau (personal communication), an expert on Black Bears, concurred. In his experience he had not actually seen bears drag Elk carcasses ashore but, from an aircraft, had seen a Black Bear swimming halfway out toward an Elk carcass 40 m from the shore of a lake. On another occasion he had seen Black Bear and Gray Wolf (Canis lupus) tracks associated with an Elk car- cass on the shore of a remote lake in the park. In the mid-1990s a party of hikers reported an animal with a black head behind a carcass in deep water of Bead Lake Four less than 3 km NW of Grayling Lake. The only scavenger other than a bear that it might have been mistaken for would be the much rarer Wolverine (Gulo gulo) (Pat Rousseau, personal communication). Environment Canada wind data in the critical time period that day (08:00—12:00) showed only very light breezes primarily from the NW which almost certainly eliminated the wind as being a factor in the carcass being blown to the east shore (Table 1) (Figure 1). On 112 a similarly calm day, 13 August, blocks of wood were placed in the middle of the lake to see if water currents would carry them to the east shore. No definitive proof was obtained that the bear swam out and hauled the carcass a minimum of 200 m to shore. However, the negative results of the water cur- rents experiment coupled with the data on wind direc- tion and speed that day provided a strong suggestion that this is what happened. Still, it is possible that anoth- er species of scavenger (e.g., Wolverine) brought the carcass to shore, and then abandoned it. Further, al- though none was seen before 08:15 or after 12:15 there is a possibility that, while the carcass was unobserved by the author between these times that day, one or more people kayaking or canoeing on the lake took the car- cass to the east shore. Interviews with such recreation- ists at the lake on subsequent days revealed no knowl- edge of the carcass being moved. Hence, both of these scenarios seem very unlikely. In retrospect, the bloated condition of the carcass near the middle of Grayling Lake on 2 August doubt- less produced a powerful olfactory stimulus for the bear and her cubs. THE CANADIAN FIELD-NATURALIST Vol. 120 Acknowledgments I thank Parks Canada Warden Patrick Rousseau for his accounts of Black Bears and Elk and Warden Service Communications Officer of Riding Mountain National Park, Ken Kingdon, for providing informa- tion on wind speed and direction. Appreciation is also extended to the reviewers of the manuscript who made constructive comments and gave suggestions. Documents Cited (marked * in text) Parks Canada. 2004. Riding Mountain National Park esti- mated elk and moose populations from aerial survey 1976- 2004. Literature Cited Hatler, D. F. 1972. Food habits of black bears in interior Alaska. Canadian Field-Naturalist 86: 17-31. Lang, A. H. 1974. A guide to the geology of Riding Mountain National Park. The Geological Survey of Canada. Miscel- laneous Report 20, Department of Energy, Mines and Resources, Ottawa. 68 pages. Machutchon, A. G. 1989. Spring and summer food habits of black bears in the Pelly River Valley, Yukon. Northwest Science 63(3): 116-118. Received 28 February 2005 Accepted 2 January 2006 Book Reviews ZOOLOGY rhe Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi By Terry Stevenson and John Fanshawe. 2006. Princeton University Press: 41 William Street, Princeton, New Jer- sey, 08540 USA. 640 pages U.S. $35.00 Paper. Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, and Burundi is one of a number of bird guide books on this region. Indeed, this publication is a paper- back version of the award-winning 2001 hardback edi- ‘ion. It has not been updated. The first of these books I bought was Birds of East Africa by Williams and Arlott, published in 1963. At he time I was very pleased at the style and quality of his work. Some 665 species were illustrated. In 1999 his was superseded by Birds of Kenya and Northern Tanzania by Zimmerman, Turner and Pearson. This 900k was a giant leap forward with 1084 species illus- rated (even though this only covered 40% of the area of East Africa.) The illustrations increased by 419 spe- >ies despite the smaller area covered. Also the quality of the artwork and printing was far better and was a sloser match to European and North American guides. Now we have the new Birds of East Africa with Norm Arlott again providing much of the artwork. This vol- ume illustrates 1388 species and is another, although smaller leap forward. Birds of East Africa covers the resident, migrant, and vagrant bird species of Kenya, Tanzania, Uganda and the two smaller countries of Rwanda and Burundi (the area of these two together is less than 3% of the region, but adds a surprising number of species to the list). The text is high quality and faces the colour plate page (a more convenient format). The species accounts de- scribe identification, status, range, habits, and voice. The colour distribution maps are small, but clear. The question now arises, which is the better book for the visitor going on typical trips to the major parks in Kenya and northern Tanzania? Birds of Kenya and Northern Tanzania is slightly smaller (around 5%) and covers less territory. While this loses about 20% of the species, most of these are specialist species con- fined to Uganda, Rwanda and Burundi (about 225 species that are wholly or primarily confined to those countries ). The remainder are species predominantly from southern Tanzania. To my eye the artwork in The Birds of East Africa is noticeably better both for the colourful species and the LBJ’s (“Little Brown Jobs”: warblers and Cisti- colas), but this not a huge difference. I checked sev- eral species from my slide collection and found Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi was generally more accurate. The user will find it very difficult to separate Long-billed from Tawny pipit using Birds of Kenya and Northern Tanzania, but will find only a small improvement with Birds of East Africa as these birds are difficult to separate anyway. There is not much to choose between the texts (al- though it is more extensive in Birds of Kenya and Northern Tanzania) and the coverage of endemics is very similar. It is easier to have the text, illustration and map side by side (Birds of East Africa ). The biggest difference lies in the range maps. The Birds of East Africa shows only a single range designation, covering winter, summer and migratory ranges. In comparison, Birds of Kenya and Northern Tanzania shows summer and winter ranges separately. Just as important, Birds of Kenya and Northern Tanzania more clearly defines where the birds are located. For example waterside birds like bitterns and herons are show in localities where water occurs. Similarly plains species are shown in the plains. The Birds of East Africa maps blanket the entire area that water birds can occur giving the impres- sion they can be anywhere, even in dry savannah. Birds of Kenya and Northern Tanzania includes a map of region inside its front cover that shows the key locations mentioned in the text. Although The Birds of East Africa has maps that show features (like for- est cover), they do not depict the location of national parks, cities, etc. Neither book gives alternative names, which is a bit irritating when making comparisons with park checklists, older books, trip reports etc. (e.g. Schoenicola brevirostris is called the Fan-tailed Grass- bird or Broad-tailed Warbler and not to be confused with Cisticola juncidis called the Fan- tailed Warbler but now known as the Zitting Cisticola.) If you are planning a trip that includes Kenya, Tan- zania, Uganda, Rwanda or Burundi, The Birds of East Africa is the more valuable book to take with you. When I go to Kenya and Tanzania later this year, the extra information given by the range maps would prove beneficial (These more detailed range maps will be particularly useful for the first time visitor.), but | am taking Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi because of ease of use and higher accuracy of the illustrations. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada HS 114 The Birding Sites of Nova Scotia By B. Maybank. 2005. Nimbus Publishing Ltd., 3731 Mackintosh St., Box 9166, Halifax, Nova Scotia B3K 5MB8 Canada. 654 pages, $35. There is a path that runs from a popular beach in southern Nova Scotia. If you know exactly where, you can leave this path and cross several hundred metres to the start of a more secret path. This second path runs through a spruce grove along a cliff. If you know exactly which spruce to slip behind, you will find a hidden rope that will allow you to drop several metres down a cliff face. You will be in a very secluded cove with a golden beach and blue water, sheltered from any cold wind. This secret spot must be the only place in Nova Scotia that Blake Maybank has not visited! His new book is an incredibly detailed set of directions to (almost) every tiny nook of this beautiful province. The main purpose of this book is to give sufficient directions for the visitor or native Nova Scotian to find the numerous coves and special places that dot the province. This is not as easy as it sounds as some of the roads and tracks are obscure and hard to distinguish. For example, I know native-born people who did not know of the track to Cherry Hill Beach, even though they had driven past it many times. This is because it is tucked between houses and looks like a driveway. Very sensibly Maybank has given the GPS coordinates of this location and others where someone might have trouble. At each locality the author describes the habitat and comments on the birds typically to be found there. He also includes cultural highlights such as churches, lighthouses, museums and other tourist attractions. Although the title suggests this book is only about birds, Maybank includes similar information on mammals, butterflies, reptiles and amphibians. Where noticeable he comments on the wildflowers but this is not a focus. Each section has icons representing the types of birds, plants or mammals found and other key features like washrooms to be found on the site. Giving directions can, of necessity, be dry reading even (or especially?) when they are given in great detail. The Complete Fauna of Iran By Eskandar Firouz. 2005. I. B. Tauris. London, New York. Xiv + 322 pages. US $90. This book covers the vertebrate fauna of Iran, mam- mals, birds, reptiles, amphibians and freshwater fish- es. It is a revised, English language version of a 2000 publication by the same author in Farsi (reviewed by Coad and Keivany 2002). Those fluent in Farsi might prefer that version at the bargain price of $5.63! Iran is a country that often appears in the political news for its great economic and strategic geographi- cal importance. This is mirrored in its vertebrate THE CANADIAN FIELD-NATURALIST Vol. 120 The writer’s style is much more upbeat and entertain- ing in the other sections. I was amused to read (Meat Cove) is “beyond the radar ... its delights are shared by a privileged minority” (Meat Cove, despite its name, is indeed a beautiful place of seabirds, orchids and gor- geous scenery; naturalists should certainly go there). Maybank does not give a detailed account of all the birds likely to be seen at a given locality. He uses gener- ic terms such as gulls, shorebirds and warblers. Howey- er he has set up a website where you can access bird lists and information on the key localities. I assume he plans to keep this site current, so it is much more useful then putting a list in the book (which would soon be dated). There are a few items that I think need changing. Page 243B has lost part of its text, an obvious printing problem. The author states “There are no deer ticks ... hence no Lyme disease.” It is not this simple. Dog ticks (Dermacentor variabilis) also carry the Lyme disease bacteria, Borrelia burgdorferi (technically they are “infected”, but maybe not “infective”). Whether or not the D. variabilis tick transmits Lyme disease is not yet resolved, suggesting caution. I was dismayed to see he did not mention Grassy Island (and hence Roseate Terns) in his account of Pearl Island (perhaps the last pair has been extirpated). The book has no index, mean- ing you will need a map to get you oriented. (Find the county you want and use the index to get you to the cor- rect chapter.) Any traveler who has gone to the expense of travel- ing to Nova Scotia needs this book; it will be invalu- able. They will get to more places and see things they would have missed. Nova Scotians will also benefit. I doubt if anyone has traveled so widely and could pro- vide such a precise record as Maybank. I will be using this book for my upcoming trip to Ile Madame in Cape Breton. As my last trip was in winter, I look forward to more informed and interesting birding this time. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada fauna, which includes such famous economic species as the caviar-bearing sturgeons and many diverse species of interest to naturalists close to its location at the intersection of the Palaearctic, Ethiopian and Oriental realms. : The Introductory sections of the book are particu- larly valuable for readers not familiar with this part of the world, and encompass 48 pages. A transliteration system is explained for the Farsi common names of organisms (a local language name list is always use- ful as scientific names may be more of a barrier than a 2006 help outside academia), the distribution of Lranian prov- inces is mapped, major freshwater drainage basins outlined, and topography summarized in colour. A sum- mary of zoological work in Iran follows with accounts of the transition between hunting and falconry to conservation and protection. [ran now has an impres- sive list of 16 National Parks, 13 National Nature Mon- uments, 33 Wildlife Refuges and 92 Protected Areas. Summaries of physical geography, climate, vegeta- tion, zoogeography, threatened species and some fos- sils follow, illustrated by colour habitat photographs. Appendix I lists all the parks, their area, year estab- lished and a grid reference for a map in the Introduc- tory section. Appendix II describes the Pardisan proj- ect (from the Old Persian for paradise), a 270-ha park in Tehran intended to replicate Iranian environments as an innovative educational project and centre for re- search in all fields related to the natural world. Con- ceived by Firouz, Pardisan was meant to be an academy of sciences, a zoo, herbarium, aquarium and museum. Hopefully it will come to fruition. The Bibliography encompasses 7 pages and is a general introduction to the literature on Iran. This literature is surprisingly diverse and could not be covered in this book — that on freshwater fishes, for example, numbers over 2900 items (see www.briancoad.com). The book ends with indices to Latin and English names of vertebrates and a gen- eral index. The mammals occupy 60 pages, birds 88, reptiles 54, amphibians 10 and freshwater fishes 34. The num- ber of mammal species referred to is 168, birds 514, reptiles 199, amphibians 20, and freshwater fishes 173, for a total of 1074. Because of space limitation, not all species are described. Introductory sections give an overview of work on each group and an update from the 2000 edition in Farsi; e.g., six species of mammals have been added to the fauna. Iranian scientists became very active in the 1990s and subsequently, as evidenced by their publications in such journals as Zoology in the Middle East. Comments are made on conservation and the destruction of habitats that threaten so many species, here as in the rest of the world. As an example, the first family dealt with, the hedge- hogs, number four species which are listed by English name, Farsi name in English script, Latin name and a numbered listing of the provinces in which they occur. This requires the reader to find the map of provinces at the front of the book, and these maps are very small (15 by 10 mm), and would require a knowledge of Iran- ian geography to interpret easily. Any illustrated species is indicated by a diamond next to the English name. A general review of hedgehog biology precedes this list. No mention is made of the absence of hedges in Iran (where hedgehogs are found in burrows, bushes and rock crevices), and the disconcerting habit shown by some species of hissing loudly when disturbed is also omitted, an attribute not noted in the more sedate English hedgehogs. BooK REVIEWS Mammals of note, and evidence of the land bridge that Iran forms between Europe, Asia and Africa, in clude lions and tigers (both now, sadly, extinct), Egypt- ian fruit bats, wolves, red foxes and jackals, cheetahs, leopards and lynx, hyenas and badgers, deer and gaz- elles, and jerboas and hares. The plan to re-introduce lions from India to southern Iran is noted but not whether this actually took place. Rumours in the 1970s had them introduced, but quickly wiped out by local villagers who were not ardent conservationists when faced with a potential predator on their sheep and goats, not to mention sleepy goatherds and shepherds. The illustrations are a mix of photographs and colour drawings. The birds are illustrated mainly from colour draw- ings provided by Birdlife International and are a good summary of that fauna. Necessarily, this section in par- ticular must be quite abbreviated in its text, given the bird diversity, but books devoted to birds are widely available for the fauna of the Middle East. Status sym- bols are given for each species rather than provincial distributions as with mammals; e.g., R = common res- ident, r = scarce resident, again a system that requires reference back to the page listing 16 possibilities. These could certainly have been included with each species’ name in an abbreviate form. The reptiles are well-illustrated with colour photo- graphs. The fauna is varied and interesting but not evi- dent without effort in field situations. I seldom saw snakes and lizards while living in Iran although small geckos could be found on house walls (and had a fero- cious bite). One of the largest lizards readily encoun- tered is the Bengal Monitor which reaches 2.75 m. Chasing one of these rapid lizards across the desert could be quite tiring if they did not try to hide behind a bush much too small to conceal their length. Marsh crocodiles live in the southeast of Iran, much decimat- ed by drought in the late 1990s. They are here reported as not attacking people, a fact I wish I had known in the 1970s when I might have sampled their home streams for fish less nervously. Five species of marine turtles are known from the Persian Gulf, all badly in need of protection. The amphibian fauna is small in species although toads and frogs are common in and near water. Tree frogs are found on banks of streams, trees being rare in much of desert Iran. Extensive critical comment on the freshwater fishes would be inappropriate given the input by this review- er. Only freshwater fishes are dealt with, although this includes Caspian Sea fishes where salinities are about one-third of sea water. The diverse fauna of the Per- sian Gulf and Sea of Oman are not covered—a recent summary work is Assadi and Dehqani Posterudi (1997). One loss of information is that space does not always allow for a full explication of evocative Farsi names. The enormous sturgeon, Huso huso, reaching 1.5 ton- nes and 6 m in length, is known as fil mahi which 116 THE CANADIAN FIELD-NATURALIST translates appropriately as elephant fish while the pike, Esox lucius is ordak mahi, duck fish from its duck- like snout. The author is eminently qualified to write on this topic having been the Director of the Department of the Environment and vice-president of IUCN. He is regarded as the father of Iran’s environmental move- ment. The Farsi version of this book won the Iranian Publisher’s Association top award. This book is highly recommended as a well-writ- ten overview of a fascinating fauna. Fishes of the World By Joseph S. Nelson. 2006. John Wiley & Sons, Hoboken, New Jersey. Fourth Edition. xvii + 601 pages. U.S. $125. This book covers all the 515 families of fishes in the world in 62 orders and 5 extant classes. Fossil families are more briefly treated. Each class, subclass, order, suborder, family and subfamily receives a brief descrip- tion. In this way the reader is treated to a tour of the world’s fishes through their higher classification and relationships. This being the fourth edition, 30 years after the first, proves the need, and success, of such a work. Scientists and students look to this work for an overview of fish classification and, with an estimated 28 400 species at the end of 2006 compared to about 26 730 tetrapods (mammals, birds, reptiles and amphib- ians), the fishes need this structure. At the individual species level, constantly changing as new ones are described or old ones revised, the online “Catalog of Fishes” from the California Academy of Sciences is the update source (www.calacademy.org/research/ich thyology/catalog/index.htm). A website, http://members.shaw.ca/fishesoftheworld/ index.htm contains errata, a list of new families post- 1958, student exercises and links to relevant websites. Changes in the numbers of taxa between the four edi- tions of the book are listed in the table below. Variation in the number of families is mostly due to lumping and splitting as Nelson points out at the website mentioned above. Discovery of new families is a rarity in fishes. However, the estimated number of known fish species for the end of 2006 cited above is almost 10 000 more than in 1976, showing that this field of endeavour has been most active. The text has continued to be expanded over the ear- liest editions, with families once without a description now provided with one. Much necessarily remains the same from the last edition as relationships, anatomy and species content remain unchanged in certain families. Vol. 120 Literature Cited Assadi, H., and R. Dehqani Posterudi. 1997. Atlas-e Mahian-e Khalij-e Fars 0 Dary-ye Oman/Atlas of the Persian Gulf & the Sea of Oman Fishes. Iranian Fisheries Research and Training Organization, Tehran. 10 + 226 + 23 pages. In Farsi and English. Coad, Brian W., and Yazdan Keivany. 2002. Review of “A Guide to the Fauna of Iran.” E. Firouz. 2000. Iran University Press (Uni- versity Publication Centre), Tehran. ISBN: 964-01-0956-8. vi + 491 pages. 45,000 Rls ($5.63) (hardbound)”. Copeia 2002(4): 1164-1166. BRIAN W. COAD Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada Not all genera are listed within each family, although a couple listing would be a great convenience, and not all families have a line drawing (about 74% do). Some families have undergone significant changes, with new information on species numbers, ecology and fossil members added, including the new coelacanth found in a fish market. Several groups (e.g., Characiformes, Osphronemidae) have been rearranged based on new systematic studies, and Nelson makes a concerted ef- fort to follow a cladistic framework. However, as Nel- son notes in his preface, it is naive to accept the latest proposals as being the best. Nelson has used his own considerable judgment in determining which groups are sufficiently systematically stable to include. This has also resulted in some families (e.g. Cichlidae) los- ing some structure, as new studies have shown past taxonomic groupings to be non-monophyletic. The literature on fish classification is extensive, sometimes arcane in its arguments, and scattered in journals worldwide. Having an author mull over this lit- erature and provide an interpretation and digest for con- sideration is a great service to students and to those pro- fessionals who need to dip into these waters at intervals. In this new edition, Nelson has included many more references, providing the most recent literature. The Bibliography is comprehensive and covers 54 pages. Some authors may be disappointed that not all their papers are cited, but Nelson has generally included good summary papers that will provide, within their own references, many titles of more specific articles. The Index at 63 pages is also most useful in locating particular taxa but certain genera will not be found (as noted above) and the reader will have to resort to Cat- alog of Fishes. Additionally, the previous edition used the names of orders as the running header throughout the text. The use of the more inclusive class level as the running header in the new edition is less helpful in flipping through to find the desired pages. Marine and Year Species Freshwater Diadromous* Families Orders Classes 1976 18,818 6851 11,967 450 46 + 1984 21,450 8411 IBS 445 50 4 1994 24,618 9966 14,652 482 57 5 2005 ZU UT 11,952 16,025 515 62 5 2006 The single disappointment of the new edition is the loss of the chart of categories and relationships of fish- es from the inside front cover. It is now buried on (un- numbered) pages xvi-xvii, between the Acknowledg- ments and the Introduction, making it much harder to find. Owners of the book may want to place a perma- nent bookmark in the chart for quick reference. The new chart, while showing better resolution of groups in this edition, has also lost all the names between Class and Order. While some of these names were for non-cladistic grades, their presence on the previous edition’s chart was of great benefit, particularly for stu- dents, to determine exactly what is a “teleost” or “acti- nopterygian”, or any of the other names that are com- monly used by ichthyologists. At the end of the list of errata on the website given above, and on page 9 of the new edition, Nelson draws Insects: Their Natural History and Diversity: North America By Stephen A. Marshall. 2006. Firefly Books Ltd., 66 Leek Crescent Richmond Hill, Ontario L4B 1H1 Canada. 720 pages. Can $95. Hardcover. It has always been said that you can’t judge a book by its cover, but after reading /nsects: Their Natural History and Diversity lve learned that this well-worn axiom isn’t always true. When this book first crossed my desk, to say that I was instantly enamoured would be an understatement. It was so beautiful, the cover adorned with a stunning jewel-toned dogbane beetle (Chrysochus auratus). 1 almost didn’t want to crack the binding. However my curiosity finally got the better of me and I’m glad it did because once I started read- ing I couldn’t put the book down. Visually stunning, with over 4000 color photographs of insects in their natural habitats, nsects: Their Nat- ural History and Diversity has the look and feel of a glossy coffee table book while still being full of accu- rate, well researched information. As its title implies Insects: Their Natural History and Diversity focuses on the diversity and natural his- tory of common families of northeastern North Amer- ican insects. The book opens with a brief synopsis of basic insect anatomy and morphology. This is fol- lowed by chapters covering the diversity of all insect orders, including all the major families, along with two chapters on non-insect arthropods and methods for observing, collecting and photographing insects. The book’s last 50 pages are dedicated to illustrated keys to order and family as well as a key to the most commonly encountered insect larvae. These keys are designed to facilitate ease of use and therefore empha- size morphological characters visible to the naked eye or easily seen with a hand lens. Also peppered through- out the book are helpful suggestions on where to look for and find various insect orders/families. For exam- ple “Depending on your inclination and the weather, BooK REVIEWS 117 attention to the need for ichthyologists, and the work that still remains for future generations. He urges us to support the replacement of retiring ichthyologists to continue the work. The Introduction of Fishes of the World provides a sampling of all the rich areas of ich- thyology to be studied and highlights the importance of fishes to all of us. Perhaps if the Introduction were required reading for everyone, Nelson’s enthusiasm for fishes would be passed on to all, and the worth of ichthyological research would be clearly visible to governments and society. This book continues to be a seminal work, finding an essential place in libraries and on the bookshelves of anyone interested in fishes. BRIAN W. CoaD and ALISON M. MURRAY Canadian Museum of Nature, Ottawa, Ontario KIP 6P4 Canada With a photographic guide to insects of eastern a good place to start looking for assassin bugs would be in your kitchen light fixture. Unless you are much more fastidious than most, the odds are that among the crispy critters accumulated there you will find a large black assassin bug called the Masked Bed Bug Hunter (Reduvius personatus).” Considering that Insects: Their Natural History and Diversity is priced so as not to be cost prohibitive and is clearly written in plain language makes it highly accessible to a broad audience including naturalists, amateur entomologists as well as seasoned profession- als. The author has also included a dollop of humour and wit throughout the text. For example this passage describing the appearance of springtails: “Some are covered with scales, like those of a butterfly, many are brilliantly colored and all are morphologically bizarre, starting from the long, forked tail used to make Her- culean leaps, and ending with the deeply pocketed mouth that makes springtails look like they have lost their dentures and then sucked on a bunch of lemons.” This book would make a great textbook for a natu- ral history or general entomology course. Especially when you consider that the impetus for this book cen- ters on materials originally gathered in support of the author’s third-year course “The Natural History of Insects” at the University of Guelph. With its depth of scope and true-to-life color photographs /nsects: Their Natural History and Diversity would be indispensable in the field; however, due to its size it would be a bit unruly to have to lug around. I thoroughly enjoyed this book and would recom- mend it whole heartedly to anyone who has an interest in entomology, natural history or a simple curiosity about the six-legged world that surrounds us. GINA PENNY Department of Biology, St. Francis Xavier University, Anti- gonish, Nova Scotia B2G 2W5 Canada 118 THE CANADIAN FIELD-NATURALIST Vol. 120 Secret Weapons — Defenses of Insects, Spiders, Scorpions, and Other Many-Legged Creatures By Thomas Eisner, Maria Eisner, Melody Siegler. Belknap Press of Harvard University Press, Cambridge. 2005. 372 pages. This book is dominantly about chemical defences of terrestrial arthropods, though other types of defences (camouflage, catapulting, sticky traps) are also men- tioned. Chapters are short, and typically feature the defences of one organism, or at most a small group of related organisms. One or more colour photos, in them- selves almost worth the cost of the book, enhance both the aesthetics of the book and the ability to drive home each chapter’s message. Chemical formulae are illus- trated in those chapters focusing on chemical defences. As the subtitle alludes, an array of terrestrial arthro- pods is dealt with, but by far, the majority of this book is about insect defences. However, scorpions, centi- pedes and other non-insects together account for about 15% of the book’s material. Examples include the well-known spray of the Bombardier Beetles, the less well-known vomit attacks of noctuid moth caterpillars and immobilizing wax of oleander aphids. This book will appeal to a wide array of readers, the caveat being that those with no (or little) training in chemistry (like me) will miss out on some of the nuances of the chemicals themselves; this did not give Rodents and Lagomorphs of British Columbia By David W. Nagorsen. 2005. Royal British Columbia Muse- um, Victoria, British Columbia. I read this book both with a general interest in mam- mals overall, and, as a raptor biologist, a desire to more completely understand the creatures which are the prey of my birds. I was not disappointed on either account. From a simply presentational point of view, the art- work sticks out as being very aesthetically pleasing. Each species is illustrated with a pencil drawing of the complete animal — not in a standard three-quarter pose or profile, but in varying postures that show snapshots of each animal’s character. Additionally, the skull of each species is presented in lateral, dorsal and palatal views — perhaps this is not useful to the general reader, but I certainly benefited from them; the illustrators (Michael Hames, Donald Gunn and Bill Adams) must be commended. The book has three keys — one each for whole ani- mals, skulls and chipmunk genital bones. Anyone who has used an unillustrated key understands the frustra- tion of trying to interpret the author’s meaning (what is meant by “relatively large?’”’). A fully illustrated key has diagrams for each couplet — the keys in this book are not fully illustrated, but there are plenty of dia- grams that make the process of keying out an organ- ism rather straightforward. An introductory chapter describes the biogeography of British Columbia, along with conservation strate- gies, winter survival and several other topics which link these species together. Maps and good photographs are featured here. rise to any problems at all for my enjoyment of this book. The other caveat being that those not liking those long, tongue-tripping scientific names of organisms will have to gloss over them periodically as they read; even where common names exist for the organism of the chapter’s focus, the scientific name is typically used throughout the text. Grade-school children will be able to use this book though clearly not as completely as older folk; entomology instructors could easily incor- porate this material into lectures. Truly, this book can meet the needs of many. There is an abundant amount of fascinating mate- rial that simply makes this well-written book a joy to read; I have been equally satisfied after having read Waldbauer’s books, including What Good are Bugs? (also from Harvard Press). Unlike the latter book, the chapter bibliographies of Secret Weapons are replete with papers of the book’s authors, particularly the Eisners. Their work in chemical ecology, translated by them from their papers to this book, is a grand exam- ple of making science more consumable by the non- specialist. RANDY LAUFF St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada The species accounts dominate this book. Each runs about six pages and describes physical aspects of the species, its natural history, geography, and conserva- tion status. An occurrence map accompanies each ac- count; I believe British Columbia residents and non- residents alike would have benefited from place names being included on the maps — especially those which are only mentioned once or twice in the whole book. Similarly, even the introductory maps do not include all the major sites mentioned in the book. There are a small number of mistakes, some edito- rial (e.g. “supraorbital bone” is used instead of “supra- orbital process”), some factual (e.g. “[The muskrat has] rooted cheek teeth.”). In the introduction to the lago- morphs, I think it would have been appropriate to differentiate rabbits from hares. I take particular exception to the oft-used expression that some introduced animal (in this book, the East- ern Cottontail) “seems to be filling an ecological niche.” This is at best an empty comment (what introduced animal would not have a different ecology than the natives) and at worst could be interpreted as an excuse to tolerate and perhaps enhance introductions. Those few shortcomings aside, this is‘a very good book which serves its purpose admirably. I will refer to it frequently. RANDY LAUFF St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada 2006 Book REVIEWS 119 Tanagers, Cardinals, and Finches of the United States and Canada By D. Beadle and J. Rising. 2006. Princeton University Press, 41 William Street, Princeton, New Jersey 08540 USA 196 pages, U.S.$29.95 Paper. The authors use 200 selected photographs to com- plement text covering the biology, identification, molts, song and distribution of the tanagers, cardinals, and finches found north of Mexico. In all, forty-six species are covered. Of these, one is an introduced species and 31 are native residents of some part of North America. The remaining 14 are vagrants with five as Alaska spe- cials and seven creeping north over the southern border. I have just spent a couple of hours resolving an iden- tification issue using about 20 photographs off the Inter- net (out of the 200 I found there) of a single species. Most of those photos were of limited use. Normally I find such photographs less useful than artwork because they are so influenced by light, angle and the condition of the specimen. However, my jaundiced eye is very impressed with the photographs in this book. I do be- lieve that I would be able to separate female House and Purple finches using the photographs and text given in this book. Indeed I used this book to confirm that I had correctly identified a Hoary Redpoll I saw recently in the Arctic. The typical resident species are represent- ed by several photographs taken at different angles thus giving the reader a better understanding of the bird. The 31 residents each have a range map giving sum- mer, winter, and permanent ranges. These are about 5.5 cm square and very readable and useful. There is a written description of range too, which adds more detail. I had no difficulty deciding I was at the northern edge of my Hoary Redpoll’s range. The text is clear and provides great detail without wasted words. The vagrants are similarly treated, but do not have a range map. Under distribution their normal range (outside of North America) is described, followed by a summary of their occurrence in North America, with additional details, such as precise locations and dates, where warranted. This is a very useful supplementary guide (to the classical all-species field guides) for providing addi- tional information on this group of birds. It is conve- niently sized to carry in the field and will be especially useful in sorting out look-alikes such as Hoary and Common redpolls. It provides carefully organized and well-presented information to those who wish to learn more about these birds than is given in the typical field guide. This is the type of book that makes a most wel- come present! Roy JOHN 2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada Wheatears of Palaearctic: Ecology, Behaviour and Evolution of the Genus Oenanthe By E. N. Panov. Pensoft, Sofia, Moscow. 439 pages, U.S. $167.50 Wheatears are the quintessential Afro-Eurasian bird. Found anywhere there is open ground, the more deso- late the better, wheatears make ideal subjects for orni- thological study; indeed, according to the author they are among the most studied groups of passerines. They are easy to see and follow, they exhibit interesting be- haviours, and, as a group, pose many riddles. How can up to four species with seemingly identical foraging behaviours co-exist in a relatively inhospitable habi- tat? Why in some species is there high sexual dimor- phism and in others virtually none? What light can this group renowned for intra-species variation shed on the nature of polymorphism and its evolutionary ori- gins? What are the advantages of a contrasty black and white plumage? How did one species of this desert loving family break through the oh so inhospitable for wheatears boreal forest to successfully colonize much of the Arctic including not once but twice in North America? These and other questions are copiously answered in this book by E. N. Panov, a Russian scientist with a passion for this intriguing group. The book is divid- ed into two parts, the first dealing with general fea- tures of wheatear biology, the latter providing detailed monographs on individual species or superspecies found in Eurasia. Between them, these sections cover every conceivable angle on wheatears, from the geo- graphic distribution and origins of the group in its entirety through habitats, movements, social behav- iour, reproductive behaviour to predators and parasites. Each monograph follows a template covering system- atics, distribution, behaviour, nesting, and movements. The book is extensively illustrated with black and white sketches illustrating everything from behaviours, nest sites to plumage variations and is amply provided with sonograms, family trees, graphs and maps illus- trating key points. There is also a section of 40 colour plates showcasing the array of arid habitats so beloved of the genus, portraits of selected species, and at least one picture of one being devoured by a snake. The book closes with a 14-page bibliography and an index help- fully divided into a general subject index as well as an index of animal names. There are about twenty species of wheatear, with the exact number hard to pinpoint because of blurry lines with other turdidae such as saxicola (“stone- chats’’) and uncertainty about the taxonomy of several closely related taxa. Taxonomic and genetic evidence suggests that the genus had its origins in Africa, where it may have emerged from an ancestor shared with redstarts. The large amount of variation and frequent contact zones within the genus suggest that there is an ongoing evolutionary process. The English name, which comes from the old English for white rump, 120 refers to the bold white “T” pattern which is character- istic of many members of the genus. Its Russian name, Kamenka, refers to the stony habitat they so love; their ability to cope with extreme conditions is witnessed by the fact for some 160 000 square kilometres in the Sahara the White-crowned Black Wheatear is the only breeding passerine, and the fact that the Northern Wheatear has successfully colonized arctic tundra. Given the lack of culinary choice their dietary habitats are pretty well catch as catch can; recorded food items include scorpions and bumblebees. The Northern Wheatear, the only member of the genus in North America, is remarkable. Each year birds from Alaska, in the words of the author, “stubbornly make for their remote African homeland”, an epic voy- age that would exhaust an Arctic Tern. Birds nesting in Eastern Canada head in the opposite direction, and are the only Canadian passerine that winters in Africa. Being found in places where lots of ornithologists live; the Northern Wheatear is one of the better known members of the group. From these studies, we know that they maintain a frenetic lifestyle during the breed- ing season that can, in the far north, involve a 24/7 rou- tine of zealously patrolling its borders while attending the needs of two females in widely separated territo- ries. According to Panov, Wheatears are highly aggres- sive with a “low threshold for territorial aggression”. BOTANY Giant Trees of Western America and the World By Al Carder. 2006. Harbour Publishing Co., Madeira Park, British Columbia. 138 pages. $18.05. Al Carder has been researching giant trees for a lot of his long career, collecting stories of large trees in the Pacific forest of North America, his home and main study area. This book is a sequel to his earlier work trying to record the dimensions of many of the world’s largest trees now including more facts and more tree species. The book is ordered by continents and tree families to give the reader perspective of the variety of trees which have achieved huge dimensions, but many of which have disappeared in the past century. A short anecdotal essay at the beginning puts Carder’s agenda forward quickly. He has always been awestruck by the beauty and perspective of being among the lar- gest trees and wants to preserve the memory of those trees for future ages. Sadly the largest specimens of any tree species are gone from our world due to com- mercial logging in the last two centuries and when sin- gle giant trees survived the logging, these succumbed to windstorms, having been left without the surround- ing forests which protected and supported their growth over the centuries. In North America, Australia and New Zealand, the story of logging is the history of large trees and our most impressive forests, a history THE CANADIAN FIELD-NATURALIST Vol. 120 To communicate with one another they have three dif- ferent song groups and a remarkably varied repertoire of physical signals involving different postures and movements. The author also usefully informs us that they have an unpleasant taste, evidenced by experi- ments in which hornets were offered wheatear “meat”. It is disappointing that the author, demonstrably a competent illustrator and photographer, did not choose to include a series of systemic plates illustrating the various taxa and their variations. Another disappoint- ment is that the author focused the species accounts on Eurasian taxa; one suspects that this reflects the author’s research interests but also the fact that African taxa are relatively less well known. In the review copy the hatching on many of the maps was blurry, making them hard to figure out. Nonetheless, there are few errors in the book, although the fact that the title con- tains a grammatical error did not inspire much up front confidence. This book is a scientific monograph primarily aimed at ornithologists. It poses, and proposes answers to a number of interesting ecological and behavioural ques- tions and will be of great interest to anyone intrigued by evolutionary biology. MARK GAWN Permanent Mission of Canada, 5 avenue de |’ Ariana, Gene- va, Switzerland. which has more to do with destruction of forests than the story of humans interacting with their environment. The book is short with only one hundred and thirty pages devoted to describing the trees. Each tree des- cription has an accompanying page with a diagram of the tree at its largest reported size and shape, each species being unique in shape. Fifty of those pages are devoted to the trees of North America with the other pages listing and describing the outstanding trees of other continents. North America is obviously Carder’s first priority but his love of trees encompasses the entire world. Many trees in England and Europe have been avail- able for study to written history over the centuries and Carder has researched these written accounts to give us historical anecdotes from their long life. A Tule tree in Mexico with an opening in the trunk large enough to accommodate twelve horsemen when report- ed 375 years ago, and the Sweet Chestnut in Sicily whose spread of branches sheltered the Queen of Ara- gon and her escort of 100 horsemen over 600 years ago are two of the anecdotes which describe trees still standing. Whether the tree has records of growing to over 400 feet as the Douglas Fir or whether its claim is to have lived over 5000 years as the English Yew, the grandeur of the trees is given to us by Carder to 2006 be considered, enjoyed and lamented as phenomena which have existed but sometimes passed us by. I enjoyed reading about present and past giant trees and was entertained different times looking for inter- esting facts and diagrams. It is not a book to be read at one sitting, even though short, because the tree species stand alone and reading several is like reading a list which can go on too long. A short reading of one or two tree descriptions at each visit is sufficient with another few minutes reading the next descriptions at ENVIRONMENT BOoK REVIEWS 121 another time. Soon the book becomes a reference for tree species and musings of how much we have missed and what we might visit. | have shown it to my students and they spent time with it as a picture and description book to browse and marvel. Al Carder has given us a little gem of knowledge and history for entertainment based on good science and research. Jim O’ NEILL 28718 Five Mile Road, Livonia, Michigan, 48154 USA Fire and Avian Ecology in North America — Studies in Avian Biology Number 30 Edited by Victoria A. Saab and Hugh D.W. Powell. 2005. Cooper Ornithological Society. 193 pages. This technical work is a collection of 11 papers, ten of which address the role and effect of fire in one or more ecosystems (e.g., Boreal Forest, Oak Woodlands, Interior Chaparral); the first chapter is a cogent sum- mary of the ten others. The chapters more or less follow a logical pattern in their layout. Each paper shows a map of the distri- bution of the habitat(s) discussed, and normally one or two other figures. A table of pertinent literature is presented in each chapter, which also includes respons- es to fire of a number of bird species. The papers des- cribe historic fire regimes, including the use of fire by aboriginals, as well as the effect of fire suppression on birds (and by default, the plants making up the habi- tat for them); sections on conservation strategies, in- cluding the use of prescribed burns and withholding of fire suppression end each chapter. The only irritant to me was lumping all the refer- ences at the end of the book; normally, each paper should be concluded with its own collection of refer- ences. Should someone want to photocopy one chapter, they would have to photocopy the entire compilation of references, instead of just the pertinent ones. None- the-less, it is a solid work. RANDY LAUFF St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada Fisheries Assessment and Management in Data-Limited Situations Edited by G. H. Kruse, V. F. Gallucci, D. E. Hay, R. I. Perry, R. M. Peterman, T. C. Shirley, P. D. Spencer, B. Wilson, and, D. Woodby. 2005. Alaska Sea Grant, Fairbanks, Alas- ka. Publication Number: AK-SG-05-02. 958 pages. Price: $50.00 US, ISBN:156612-093-4. Maintaining sustainable fisheries resources requires the combined efforts of scientists, fisheries managers and policy makers. Balancing fisheries growth and conservation is a difficult task, particularly in situations Where available information is limited. Data-limited fisheries are often plagued by a lack of long-term data on the basic biology, ecology and productivity of the species that are relevant to these resources. A number of successful fisheries have been developed by com- bining a scientific background with robust policy man- agement to successfully develop sustainable fisheries, even in situations where data is limited. However, newly developing and small-scale fisheries often oper- ate without sufficient data to develop production mod- els, assessment techniques and sustainable manage- ment strategies. To address these challenges, a symposium entitled “Assessment and Management of New and Developed Fisheries in Data-limited Situations” was held in Anchorage, Alaska, in 2003. The purpose of this sym- posium was to share knowledge, research and manage- ment strategies for newly developing and small-scale fisheries by bringing together fisheries scientists, man- agers and policy makers from all over the world. The result is a proceedings book titled “Fisheries Assess- ment and Management in Data-limited Situations.” Specifically, this book is a collection of forty-six peer- reviewed research papers that provide case studies and management considerations for fisheries with limited data. The book is divided into seven sections: (1) case studies of fishery failures and successes; (2) indica- tors of stock health and productivity from limited sam- pling programs; (3) involvement of fishermen and use of local knowledge; (4) multi-species and ecosystem indicators and models; (5) precautionary management approaches; (6) stock assessment models; and (7) stock assessment surveys and applications. The first two sec- tions focus on science-based assessments of fishery case studies while the final five sections identify and dis- cuss management and assessment strategies. Through- out many of the case studies in this book, several key management themes emerge that are highly relevant to fisheries managers in data-limited situations. Section one provides insight into the failures and successes of the world’s fisheries. Several case studies on a variety of species are described, and management implications are discussed. A key paper in this sec- 122 tion provides a general overview of the world status of data-limited fisheries using landings statistics with a Bayesian approach (Vasconcellos and Cochrane). Each of the subsequent case studies describes tech- niques for managing both struggling and successful fisheries. These case studies focus on the development of new or developed fisheries that have limited data (Bechtol and Trowbridge; Bureau and Hand; Therri- ault et al.) and the recovery and management of strug- gling resources (Bargmann et al.; Kruse et al.; Stephen- son and Jackson). Despite their geographically and species-specific nature, the general knowledge con- tributed by these case studies may be highly relevant to similar fisheries in data-limited situations in other parts of the world. The second section focuses on developing indicators of stock health and productivity from limited data sets. Each paper describes a case study where early warning signs can be detected in order to predict the state of fisheries resources. The indicators that are described include biological indicators (LeBlanc et al.; Tribuzio et al.), performance indicators and stock dynamics (Hay et al.; Orensanz; Raid et al.; Ziegler et al.) and management indicators (Howland and Tallman; Scan- dol; Weyl et al.). A key paper from this section is a simulated study of Yellowfin Bream (Acanthopagrus australis) where empirical stock-status indicators were assessed using quality control methods (Scandol). The third section focuses on the integration of local fishermen and traditional ecological knowledge in fish- eries management. A key paper in this section by Freire describes the use of data provided by anglers in the assessment of Brazil’s growing recreational fisheries industry. This paper represents the first data of their kind from South America and illustrates the potential value of recreational anglers as a key data source for improving fisheries management in data-limited situa- tions. The fourth section contains papers that are geared towards the assessment of the effects of fisheries at the ecosystem level by using ecosystem indicators (Stob- berup et al.) or models (Ainsworth and Pitcher; Che- ung and Pitcher; Morato and Pitcher). This section contains an interesting study by Cheung and Pitcher that outlines possible strategies for evaluating fish- eries management policies that aim to conserve bio- diversity. The fifth section contains a number of stud- ies that take a precautionary approach to data-limited fisheries management. The majority of the studies in this section focus on fisheries uncertainty (A’mar and Punt; Campbell and Dowling; Punt and Methot; Welch; Zheng). One key paper involves an evaluation of a multi-stage approach to data-limited fisheries manage- ment. Perry et al. assesses the four-stage approach to new and developing fisheries (i.e., data collection, de- sign of key studies, implementation of selected strate- gies and establishment of the new fishery) designed by Fisheries and Oceans Canada. One of the most chal- lenging factors in this staged framework, as with the management of many of the world’s fisheries, is the THE CANADIAN FIELD-NATURALIST Vol. 120 contradiction between the demands of access to fish- eries and the conservation of these resources. This paper addresses some key points that are relevant to balancing these challenges. The sixth section contains several papers that address the use of models in stock assessment. Stock assess- ment bias and sensitivity are addressed by two key papers in this section (Ernst and Valero; Mesnil). Also, Catch-Survey Analysis (CSA) is discussed by two pa- pers to estimate stock abundance (Collie et al.) and to address concerns over sensitivity and bias (Mesnil). Similarly, the seventh section addresses stock assess- ment methods and surveys. In this section, several key methods are described, including population density assessments using scuba (Byerly and Bechtol) and the applications of acoustical estimations (Kaljuste et al.). This section also addresses sampling designs for esti- mating bycatch (Menon et al.), methods for account- ing for climate variability in models (Shotwell) and assessments to address concerns over recreational over- harvesting (Therriault and Hay). A case study discussing the modeling of climate variability in relation to fore- casting Pacific salmon spawner-recruit dynamics (Shot- well et al.) is particularly relevant to data-limited fish- eries that require the input of variable environmental data into forecast models. This book contains an excellent assemblage of well- written, highly relevant research papers on data-limit- ed fisheries. Several management themes emerge from this book, including (1) the development of key rapid assessment tools, (2) the integration of traditional knowl- edge from fishermen and local ecological knowledge, (3) the adoption of a precautionary approach to fish- eries management, (4) the evaluation of sensitivity and bias in stock assessment models, and (5) the develop- ment of robust stock assessment methods. These key strategies recur throughout many of these studies and are relevant to fisheries worldwide. Further, several novel strategies for managing data-limited fisheries are discussed, including the development of key modeling and assessment tools. This book offers one of the first attempts to assemble key literature on data-limited fish- eries into a single volume. This is an invaluable resource for the management of new or developing fisheries. The intention of this collection is to provide scien- tific background to new and developing fisheries. Each research paper attempts to suggest applications of their case study to other similar data-limited fisheries resources. However, the seven sections contained in the book are organized with little or no preface or inter- pretation of this research. Throughout the book, the links between the results of each case study to a broad- er application are often tenuous. For most proceedings books in this style, it is common to omit interpretations by the authors or editors in each section. However, the nature of this research demands a synopsis in each sec- tion to highlight the applications and extensions of this research. Managers require the interpretation of fish- eries experts who are experienced with data-limited 2006 situations to identify how the methods described in this book can be applied to their own data-limited situations. In summary, the management themes that emerge from this collection are highly relevant to all fisheries man- agers, policy makers and scientists working with data- limited fisheries. However, this book would be consid- erably more useful to these stakeholders if the context Lapland — A Natural History By D. Redcliff. 2006. Yale University Press, P.O. Box 209040, New Haven, Connecticut 06520-9040. 352 pages, U.S. $60. Cloth. Lapland: a Natural History by the late Derek Red- cliff, is a remarkable account of the birds and higher plants of northern Scandinavia. This book begins with a general description in land use and the geolog- ical background terms followed by a skillful history of the earlier naturalist visitors. The book is mainly given over to a detailed evaluation of the various habi- tats found in this area. These include Boreal Forests, Forest Peatlands, Lakes and Rivers, Coastlands, Tun- dra and Man-influenced habitats. Each of these chap- ters is divided into vegetation and flora and birds. The birds section is subdivided into key groups such as wildfowl, waders (shorebirds), birds of prey, songbirds and birds that are of particular importance in that habi- tat (e.g., wildfowl or grouse). The author writes in a style reminiscent of a profes- sor giving a lecture to keen students and as a result the book is easy and pleasant to read. The accounts are packed with information drawn from many research sources, backed up by the author’s own observations. The text does contain references, but not in sufficient number to impede the flow of ideas. Every few pages there are four half-page photos. These depict habitats, plants, birds and bird nests. Particularly impressive are the almost two dozen photos of shorebird nests (have you ever tried to find a shorebird nest?). These photos, most by the author, are of very fine quality. Interspersed with the text are some engaging black-and-white illustrations by Mike Unwin, reminiscent of those in the journal, British Birds. I particularly enjoyed the way Redcliff linked the occurrence of a species with the other components of the habitat. He does a wonderful job of showing the inter-dependance of each constituent of a habitat and the consequences of disruptions to the balance, both natural and man-induced. Even though I found this a treasure trove of infor- mation (and wisdom) I did have several problems. I hoped, while doing this review, I could compare the plant life of Northern Canada, Svalbard and Lapland. The author’s inconsistent use of scientific names be- came a source of frustration for making this compari- son. The worst case was his discussions of Dwarf Aza- lea. As he does not include the scientific name it is hard to be sure which plant he is referring to. Lapland Rose- Book REVIEWS 123 of each section was clearly described in a one to two page preface and synopsis summary that outlines how this knowledge could be applied in practice. M. R. DONALDSON Ottawa-Carleton Institute of Biology, Carleton University, Ottawa, Ontario KIS 5B6 Canada bay (Canada) or Lapland Rhododendron (Europe) is Rhododendron lapponica, but the author quotes this plant separately. Indeed, he includes a beautiful photo of this lovely plant. He also includes a photo of “Dwarf Azalea’, but it is a very un-Rhododendron-like plant (more like a saxifrage or even Moss Campion). After much searching and consultation with Bill Cody (author of several books etc. on northern plants) I was still un- clear until I found a reference to Loiseleuria procum- bens as Trailing Azalea (and several other English names) while identifying plants from Cambridge Bay, Nunavut. Similarly I was never sure if the Finnmark Primrose was the same as Primula erects or nutans or if it was a new species or subspecies. Also the book only covers the higher plants well. There is relatively scant mention of the lichens and other more primitive plants. I am sure that there must be well over 500 species of lichens in this area, yet the book refers to only a handful. Other groups are even more poorly represented. On the animal side, birds are well covered. Mammals get an introduction and other groups (reptiles, amphibians, butterflies and dragon- flies) get an honourable mention. Other wildlife such as beetles and bugs do not get any attention. I also had problems with many of Redcliff’s terms. To start with, he does not really define what he means by Lapland, Fennoscandia, Fennoscandinavia and Scandinavia. As he uses terms like “Southern Lapland”, I felt it important to understand the region’s precise location. His map showing the location is vague so I thought I would check further. Lapland is somewhat well defined (northern Norway, Sweden, Finland and Russia — or the home of the Sami). Actually I have found Finnmark, Nordland, Nord-Trondelag and Troms [counties of Norway], Lapland [a province of Finland], Jamtlands Lan, Norrbottens Lan and Vasterbottens Lan [counties of Sweden], and Murmansk [an oblast — or “province” — of Russia] form Lapland. Knowing at least where the county of Finnmark is located is most useful when reading the text. However, the other terms vary from source to source. They could include some or all of Norway, Sweden, Denmark, Finland, Iceland and the Faeroes. Similarly I had trouble with the terms flark, fell, strang, patterned fen and mire. Eventually I did get them sorted out but a glossary would have been so helpful. I have several problems with the index. First, it does not reference any scientific names, only English ones. None of the photos are listed. Some of the entries are 124 listed forward (e.g., Lapland Rhododendron and Alpine Lady Fern) and some entries are listed backward (e.g., Primrose, Finnmark and Birch, Downy) making it dif- ficult to decide which letter to use. The author includes range maps from other sources. The maps for plants use the scientific names (If you did not know Silene acaulis is a Moss Campion [as opposed to the synonyms: Silene bryoides, Silene exs- capa] you would be a little lost. The lovely photo of Parmelia centrifuga is not identified as a concentric ring lichen). The map captions for birds use English names. The maps are not in the index. The Three-toed Woodpecker was split in 2003 into the American Three-toed (Picoides dorsalis) and Eura- sian Three-toed woodpeckers (Picoides tridactylus) using differences in mitochondrial DNA sequences. The author has not mentioned this or any other recent splits. Perhaps this is not as significant a issue in a book focused on ecology rather than identification. Finally, the book contains two photos of people; one is the author, although the caption omits this detail and the other is Jeanette. The book is dedicated to this lady, but her presence is never explained (she is Mrs. Red- cliff). THE CANADIAN FIELD-NATURALIST Vol. 120 This is a great book for anyone interested in the far north or who wants to understand the complexities of life. It is wonderful and fascinating reading if you are passionate about natural history, but is of more limited value for research. Derek Redcliffe was the first to establish the role of agricultural pesticides with the decline of birds of prey. He was chief scientist of the Nature Conservan- cy Council in the UK where he acted as a scientist and lobbyist for nature conservation. He fearlessly attacked and changed agricultural and forestry inter- ests for the damage to wildlife value caused by their policies and practices in place at the time. Not an easy or popular task in Maggie Thatcher’s world. Despite his legendary contribution to UK conserva- tion he was denied any honour by the state — in a country that knights pop stars. Redcliff was known an extraordinary naturalist, good company and friend. His last book 1s a fitting legacy. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Fish, Fur and Feathers: Fish and Wildlife Conservation in Alberta, 1905-2005 Fish and Wildlife Historical Society. Federation of Alberta Naturalists (FAN), Edmonton. $39.50, 418 pages. This sumptuous, well-illustrated book is an eclectic mix of articles, reminiscences and interviews. The main text is on white paper, and the other contributions on yellowish paper. Only rarely does this confuse the read- er but the white paper portion of chapter four ends in the middle of an incomplete sentence. As expected with any product of a committee, some chapters are stronger than others. We often know who wrote a particular piece, but sometimes we don’t. No- where, not even at the end of the two-page Editor’s Note, are we told the name of the chief editor (Bill Wishart, as I learned from a special enquiry to Edmon- ton). At the end of each chapter are references and a useful chronological list of the main happenings. In Alberta, as elsewhere, the responsibility for fish and wildlife management grew from a handful of vol- untary game guardians to a diverse group of biologists and enforcement officers within professional, academ- ic, and conservation communities. Yet for the better part of a century the emphasis was on fishing and hunt- ing. In 1967 the first female biologist (now the presi- dent of the Canadian Medical Association) was hired, and only in 1987 was the first non-game biologist hired. Now there are 14 recovery teams developing and imple- menting plans for threatened and endangered species. Some accounts are superb. The account of the change in upland game numbers associated with harvesting practices, from threshing machines, grain separators, and wheat stooks to combine harvesters, is among the best I have encountered. Alberta has had some unique successes. The Pere- grine Falcon restoration program, under the direction of Richard Fyfe, was immensely important. One week in the 1960s, biologists taught a three- or four-evening short course in wildlife biology and management, spe- cially designed for Members of the Legislative Assem- bly; more than half of them attended! A M.Sc. thesis on the fish and wildlife damage inherent in the planned Highwood Dam helped to influence the shelving of that project. Biographical information is generally excellent, giv- ing credit to world leaders in biology, especially Wil- liam Rowan. Even though this book is co-sponsored by the FAN, amateurs are somewhat slighted, except for Frank Farley, Cam and Joy Finlay, Elsie McAllis- ter, Myrna Pearman, Dewey Soper, and Kerry Wood. Edgar Jones is barely mentioned and Otto H6hn, Robert Lister and Al Oeming are omitted. Informa- tion resulting from bird banding is provided only as regards waterfowl. The Beaverhills Bird Observatory is not indexed. The outstanding studies of Golden Eagle migration west of Calgary by Peter Sherrington, the landmark studies of Ferruginous Hawk biology by Josef Schmutz, and the northern owl banding by the Edmonton group are not mentioned. The compilers are frank about some of the bureau- cratic bungles (“bureaucracies are often much too short- sighted;’ page 207) and failures, such as legislation 2006 drafted, but never proclaimed. The “biggest mistakes of the century” are identified as the transfer of diseased bison to Wood Buffalo Park, game farming, predator control attempts, “too many fishermen,’ and loss of habitat. There are a few errors of fact and interpretation. The two-page account of the Prairie Farm Rehabilita- tion Administration is unusually misleading when it speaks of 87 community pastures by 1999, omitting the fact that only one (in three blocks within the Suffield Military Range) of these PFRA pastures is in Alberta. The American White Pelican was threatened, never endangered (page 341). Gary Pelchat was born in Pon- teix, not the never-existent Phoenix, Saskatchewan. David Munro is misspelled “Munroe”. The first two Franklin Arctic expeditions were hoping to find an arc- tic sea route, not an overland route to Asia. The index is inadequate, failing to index names of any organism The Gulf of Alaska: Biology and Oceanography By P. Mundy 2005. Alaska Sea Grant College Program, Uni- versity of Alaska-Fairbanks, Edited by Exxon Valdez Oil Spill Trustee Council. Alaska is perceived “...as a place of wilderness, beauty and a special way of life”. This book describes the Gulf of Alaska (GOA), a region with many world records. Besides covering a huge area, having the third largest permanent ice field in the world and being a central place, crossroads, for almost all of the Pacific salmon, GOA also is among the most productive oceans in the world. The introduction is a chapter on general oceanog- raphy for the lay public; the classic concept of the ‘Conveyor Belt’ is presented. Due to its ecological im- portance, I like the human footprint statement such as “Human uses of the GOA are extensive” and“... it is rare to walk the intertidal zone anywhere in GOA without seeing evidence of human activity”. The “...marine pollution and floating refuse from as far away as Asia, or originating from deliberate deep-ocean dumping or accidents at sea, can be swept north and westward around the shelf edge in the GOA. Trash from the international fishing industry operating 200 miles offshore is commonly found on beaches”. Further, one will read that GOA has arguably the best known rocky tidal zone on earth, and this small tidal zone is cru- cial for the ocean. This book admits climate change but leaves it somewhat undefined whether it is a man- made issue, or not. The book contributors emphasize that to fully under- stand the ecology of GOA first one needs to make an effort to identify of all relevant species, e.g., plankton. Shrimp, once among the dominant benthic epifauna in Lower Cook Inlet, Kodiak and along the Alaskan Peninsula, declined after maximum harvests in 1977, and today its fishery is virtually nonexistent. Red King Crab crashed in 1980, and Tanner Crab fisheries are Book REVIEWS 125 (e.g., pronghorn, ring-necked pheasant, and liver fluke). For consistency, the book title might better have been Fin, Fur and Feathers. With pressure from an increasing urban human population, highways, and mechanization of farming, mining, oil extraction, forestry, dams and irrigation, it will be difficult to maintain biodiversity in the decades to come. Some hope may be gained from looking at both the successes and failures of the past. A large book, it is too heavy for bedside reading. Caveats aside, this is a wonderful collection of infor- mation that should be in all major Canadian libraries, and read by everyone interested in the history of fish and wildlife conservation and management. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Canada down as well. The clam fishery in Cordova never recov- ered from damages brought by an earthquake. Despite its fishery of global importance, the two short para- graphs for the Pacific Cod section are rather uninfor- mative. The fisheries for Walleye Pollock have been down since 1982. The Herring loss story for Seldovia Bay and Lower Cook Inlet is reported on as well. Salmon from Japan, Russia, British Columbia, all of Alaska and the Pacific North West spend some part of their life cycle in the GOA. The canned salmon industry collapsed in 1959, the year when Alaska be- came a U.S. state. Despite all of this overwhelming evidences of limit- ed ocean supplies, some book contributors still claim in their chapters the traditional views of “unlimited re- sources”. Other sections blame the “stocks” and anglers for their impacts (rather than their managers with a governmental mandate for safeguarding the public resource). Fortunately, sea floor dredging effects are explained, since fishery impacts to the soft-bottom benthic community are a possible driver of commu- nity change. It is suggested in the book text that this affects Steller Sea Lions as well. Throughout, the book shows that for non-commer- cial species we lack management information. These species compose the bulk of the biomass. By-catch of the juvenile cohort occurs in fisheries like Herring, Pollock, and Salmon. However, for many of the non- commercial species the magnitude of this phenomenon is not really known, nor how much it affects these stocks. Even apart from all of these severe marine issues mentioned in the book, the described terrestrial situa- tion is equally puzzling: The book states that in the GOA, 24% of the water bodies listed are on the state’s list of polluted sites attributed to some aspect of log- ging. 126 The book makes clear that Alaskan “Subsistence Harvest” is a much-debated issue, a major political player in Alaska and beyond, and thus, well supported by the government. However, only 2200 people in the Exxon Valdez Oil Spill (EVOS) region actually still practice this form of life style. This must be among the most subsidized lifestyles in the world. The gov- ernment uses undercover agents to report on how much natural resources are taken. How much do Alaskans value their nature? This book provides an answer: Anchorage operates under a Clean Water Act Section waiver, only completing pri- mary treatment of the sewage. The classic cocktail of PCB. DDE, DDT and Dioxin gets named several times, but there are many more pollutants in the Arctic. The complex toxic pollution of the Copper River is men- tioned though. I think that most readers will get excited that the Alaskan government does not choose to moni- tor for environmental pollutants. A seabird chapter deals with nesting species, not with its pelagic, non-breeding populations. Cliff nesting murtres and Black Kittiwakes are the most well studied: underground and nocturnal species are hardly known. The statements “There is no evidence that seabirds in the GOA have been directly affected by commercial fisheries” and “ ...do not compete with commercial fisheries for biomass” give the wrong impression. The book publishers need to be chastised for this misleading statement as it sets the stage for inappro- priate environmental management policies in favour of economic activities. Seabird starvations in GOA are mentioned several times: e.g., related to climatic ENSO (El Ninio/South- ern Oscillation) events. There is nice documentation of increased gull predation e.g., Glaucous-winged Gulls on Black-legged Kittiwakes, due to increased gull populations. The given Marbled Murrelet estimate of 200 000 nesting birds is likely low when compared with the authoritative literature. The provided nesting seabird population table is misleading since non-breed- ers occur in the region and thus, populations are higher. The Minerals Management Services, EVOS, NMFS (National Marine Fisheries Service), and the Alyeska Consortium did extensive benthos survey and moni- toring efforts in GOA in the 1970s. The reader is left wondering where these raw data are and this book does not help us to learn where to find them. One simply has to trust the “experts” blindly. This must set the stage for conflict. The contaminant issue I find not well handled in this book either, considering that EVOS was a GOA event of global attention and recognition. Since this book is an EVOS publication one has to ask, what did we really learn from EVOS? There is no chapter devoted to toxicology or sections contributed by a professional Alaskan toxicologist. One will have a hard time to find much oil spill literature. At least we know, in 2000, 604 spills were reported in the EVOS GOA THE CANADIAN FIELD-NATURALIST Vol. 120 region alone, proving we do have an oil and hazardous substances problem. The book notes that there is a conflict between in- creasing development and loss of environmental qual- ity in the GOA region (as found anywhere else in the world). Rats, ground squirrels and voles introduced in most of the Aleutians between 1700 and the 1900s make for a great example. Other examples include fox farms starting 1894 onwards, followed by canneries, the Klondike Goldrush of 1897, a copper mine operated from 1905 and the Northwestern railway built in 1911. Marine mammals are elaborated on in greater detail: seal decline in 1973-2000 and the sea lion decline of approximately 83% (maybe >90%). The federal fund- ing for investigating this sea lion decline is between 40 to 60 Million $$; a record in federal spending on one species. Was it a success? The infamous Killer Whale story and their blame for the seal decline gets featured as well: e.g.. in Prince William Sound. Killer Whales consume 400 seals each year (somebody has to eat the individuals of poor health: sharks were suggested as well). Killer Whales are believed to have suffered through EVOS. Ecologists might be happy to see that the textbook example of how Sea Otters and their over-harvest links with Sea Urchin and kelp populations is mentioned. The book reports that lead in Sea Otters can now be traced directly to industrial sources, namely from North America and Asia, which makes for a great example for the need of international management in GOA (none of this is demanded in the book text though). Overall. it remains unclear to me what the exact focus of this book is, and why it was published. Readers must see it as a major oversight that no professional contacts and addresses are provided for the 13 contrib- utors or even editors. I found the content editing some- times poor and not well structured. This becomes obvi- ous in the multiple mentioning of the study area and its repetitive definitions and maps throughout chapters. Some of the chapter sub-headings are virtually similar to the chapter heading, other sub-headings and themes re-occur in other chapters. The quality and amount of citations varies throughout chapters. I like the abbre- viation and acronym section because it also includes websites and their URLs. Some of the figures are stun- ningly made, but unfortunately in black and white. It becomes quickly obvious that in concert with the many other signs and evidences there is something wrong with the natural state of GOA, and subsequently how it is managed. This book is a great example how unpublished and non-peer reviewed grey literature can rather add to confusion and is not achieving its goals for sustainable management. FALK HUETTMANN Biology and Wildlife Department, Institute of Arctic Biolo- gy. 419 Irving I, University of Alaska, Fairbanks, Alaska 99775-7000 USA 2006 MISCELLANEOUS Fossil Legends of the First Americans By Adrienne Mayor. 2005 Princeton University Press, Prince- ton, New Jersey, USA. 446 pages. illustrated. The lure of fossils is not a recent venture. For cen- turies, by accident or pursuit, “sports of nature” as they once were called have fascinated our cultures. Yet, while fossils rose from “sports” to conceptions of the organic past, this was essentially a western perspective, shaped by culture and scientific advancements. It is a little surprise, however, that the North American abo- riginals or First Nations people were also drawn to these pieces of the past. The reasons varied, to be sure, but hidden to us were concepts of evolution and a sense of deep time. It is a great fortune that author Adri- enne Mayor has investigated and recorded this diver- sity of cultures, much of it based on the oral, not written, tradition, and to some extent existing in vari- ous artifacts. Mayor’s first foray into fossil lore and history was The First Fossils Hunters, Paleontology in Greek and Roman Times (2000) which examined Greek and Roman period and place with respect to fossils. Of course these cultures are rich in textual artifacts and documentation, and were in essence at the root of western civilization. Mayor’s recent study, Fossil Leg- ends of the First Americans, is somewhat more of a challenge. As part of paleontological lore, aboriginal communities certainly had contact with fossils but to what extent? Lacking written tradition, painted in broad strokes, and compounded by our interpretation of their awareness, what were their concepts of “deep time”, extinction, or even a notion of evolution? Mayor argues that aboriginals had more than a cur- sory view of fossils, and in many respects, and with a much finer, multi-colored strokes particular to specific groups, their views are much richer than anyone expect- ed. By reviewing cultures from south to north, each North American culture ebbed and flowed territorially, originating or modifying concepts of their surround- ings and to some degree of the fossil bones and stones encountered. Some Navajos in the southwest United States, for example, believe that it is horrific to disturb buried bones. “The Navajos envisioned a series of past worlds that were destroyed before this world” says Mayor, each with good and evil monsters. By digging them up, the fear emerges of bringing these past mon- Sters to our world. The large fossils found in the Nava- jo territory bespeak of life of great size in the form of petrified logs from the Petrified Forest National Park in Arizona, and of other possibilities in the rich, pale- ontological nest that they could have come across. This is more than mere tales of monsters. Fossils were not always considered dire by those who came across them. “Rock medicine of the Crows and their traditional enemies, the Sioux, often included unusual fossils, especially ammonites and baculites” Book REVIEWS 127 (page 268). They believed that these fossils carried with them mystical powers. In Alberta, medicine pouches of the Blackfeet often carried “buffalo-calling stones”, which are the segmented portions of a Cretaceous bac- ulite that look the shape, in cross section, of a buffalo. Aboriginals even attach some significance to the esthet- ics of fossil-bearing rock, in one case a point made of Pennsylvanian-age rock containing the ornate designs of the single- celled marine fusulinids from Kansas. Throughout Mayor’s work she references the revered paleontologist the late George Gaylord Simpson (1902- 1984). Like the few earlier records of ethnology and paleontology, Simpson is often taken to task for his forays into the history of fossil collecting in North America. His definition of “discovery” was much nar- rower than Mayor’s. He believed that “merely seeing a fossil bone or picking up an idle curiosity is hardly discovery” (1943, page 26). Therefore, Simpson gave little weight to beliefs and customs of First Nations people with regard to fossils in his reviews. He knew that there was contact with fossils, as in the case with the abundant fossils from the “Big Bone Lick” of the Ohio Valley, popular in the lore of American history as well as paleontology, but that’s about it. Despite the fact that Simpson’s views were reflective of 1940s American culture, Mayor’s stance is that the discoveries “were not scientific in the modern sense, of course, but there is something akin to scientific inquiry in these activities” (page 29-30). Many of the aboriginal oral traditions were handed down from one generation to another, thus becoming a permanent record in their culture. This is one of the requisites of scientific tra- dition. The 19" century American expansionism included vast paleontological initiatives like those seeded by Joseph Leidy (1823-1891), O. C. Marsh (1831-1899), and E. D. Cope (1840-1897). Ethnological initiatives also followed, but for some reason, cultural interpre- tation and fossils never met. Even when aboriginals were hired as scouts for the paleontologists, the “pale- ontologists ... [left] folklore to the ethnologists, so the ethnologists left paleontological topics out of their conversations with Native Americans” (page 195). As such, paleontological/ethnological information gathered today, some if recorded for the first time by Mayor, is tainted with over a 150 years of invading culture. A number of these aboriginal cultures have substituted their original, monstrous, interpretations of fossils with post-1840s terminology like “dinosaurs”. Where Mayor falls short, however, is in the extent of parallelism between aboriginal and western views of fossils and their meaning. There are a lot of “may have[s]’”, “could appear[s]” between the two views of scientific and cultural interpretation, and thus some of the interpretations seem somewhat forced. For exam- ple, an Assinoboine tale has “a great river monster” succumbing to a “violent death by the impact of a 128 “thunder stone”, a black projectile that came whistling out of the west” at great speed and sound, “a scenario that seems akin to the modern theory of an asteroid impact 65 million years ago” (page 289; emphasis added). The implied connection is not tenable. Likely an interpretation of a meteor, the connection between the extinction of a group of species by an extraterres- trial object is a contemporary idea. Fossil Legends of North American should have been written a century ago when contact with the First Nations people was still fresh. But the exploration of the cultural/science intertwining is still a relatively recent phenomenon. In this case, only a few other re- cent titles address fossils in a cultural context (see Cohen 2002; and to a lesser extent Semonin 2000). Still, if it were not for Mayor’s effort, from document- THE CANADIAN FIELD-NATURALIST Vol. 120 ing the oral traditions, the searching of archives, muse- ums and journals, we would be left with a culturally insensitive, simpleton view of rich and diverse cul- tures and their varied, unique surroundings. Literature Cited Cohen, C. 2002. The fate of the mammoth: Fossils, myth, and history. University of Chicago Press, Chicago. 297 pages. Mayor, A. 2000. The First Fossil Hunters. Princeton University Press, Princeton, New Jersey. 361 pages. Semonin, P. 2000. American monster: How the Nations first prehis- toric creature became a symbol of National Identity. New York University Press, New York. 482 pages. Simpson, G. G. 1943. The discovery of fossil vertebrates in North America. Journal of Paleontology, 17: 26-38. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s Drive of Vancouver By Rick Hudson, 2006. Volume 2 Revised Edition. Harbour Publishing Co. Ltd., P.O. Box 219, Madeira Park, British Columbia VON 2HO0 Canada. 368 pages $26.95 Paper. Rick Hudson has succeeded in expressing his pas- sion for mineral collecting through his field guide book. On a second level he also tries to promote a sense of adventure by teasing the reader to go out and explore or prospect for nature’s buried treasures. The author structured the book so the reader can eas- ily cross-reference required information in the lengthy location lists. A “How to use this Book” and a “Quick Reference Guide” helps to quickly locate items of interest throughout the book. The book cover illustrates the hobby of mineral col- lecting as a family activity. Hudson educates the reader from first principles on how to become a rockhound. To accomplish this, he describes how to prepare your- self by choosing proper clothing, safety gear, collect- ing tools and maps. He warns the future mineral col- lector about natural hazards they may encounter such as rattlesnakes, grizzly bears and ticks. Ethics, access rights and mineral rights are also discussed to keep the collector within acceptable collecting practices and to NEw TITLES +Available for review * Assigned ZOOLOGY The Eastern Palaearctic Species of Agnetina (Plecoptera: Perlidae). Scopolia 56. Edited by I. Sivec, L. Zhiltzova and B. Stark. 2005. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 21 pages. EUR 15.00 Paper. Discovering Amphibians. By J. Himmelman. 2006. Nim- bus Publishing Ltd., P.O. Box 9166, Halifax, Nova Scotia B3K 5M8 Canada. 208 pages. $23.95 Paper. A Field Guide to the Amphibians and Reptiles of Bali. By J. Lindley McKay. 2006. Krieger Publishing, P.O. Box 9542, Melbourne, Florida USA. US$39.50 Cloth. avoid unintentional trespassing. Mineral collecting is attractive to many people be- cause as you find an interesting mineral specimen, you want to learn how nature created it and or its geological history. The author has included short geological descriptions partitioned by collecting areas as well as mining histories of local economic importance within the collecting area. Colour plates illustrate the minerals found in some of the collecting sites and detailed occurrence maps are included in all the collecting area sections through- out the book. The colour plates represent minerals that are possible to find in the collecting sites. Government mineral index files were included in the locality infor- mation along with detailed road log descriptions. The author has made a successful merger of person- al knowledge and technical data to compile an inform- ative volume which can provide years of enjoyment for many future rockhounds and their families and friends. ROGER POULIN President, Sudbury Rock and Lapidary Society, 3171 Romeo Street, Val Caron, Ottawa P3N 1G5 Canada The Amphibians of Belarus. Advances in Amphibian Research, Volume 10. By S. Drobenkov, R. Novitsky, L. Kosova, K. Ryzhevich, and M. Pikulik. 2006. Pensoft Pub- lishers, Acad. G. Bonchev Street Bl. 6, 1113 Sofia, Bulgar- ia. 176 pages. EUR 34.00 Paper. The Amphibians and Reptiles of the Honduran Mosquita. By J. McCranie, J. Townsend and L. Wilson. Krieger Publish- ing P.O. Box 9542, Melbourne, Florida 32902-9542 USA. 302 pages. $63.50 US. Cloth. Bolton’s Catalogue of Ants of the World, 1758 — 2005. By B. Bolton, G. Alpert, P. Ward and P. Naskrecki. 2006. Har- 2006 vard University Press, 79 Garden Street, Cambridge, Mass- achusetts 02138 USA. US$49.95. The Beetles of the Galapagos Islands, Ecuador. By Stew- art B. Peck. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 SXN United Kingdom. £26.50 Paper. [Available in Canada from publisher Monograph orders NRC Resarch Press M-SS, National Research Coun- cil Canada, Ottawa, Ontario KIA OR6 Canada. $39.95 + GST 7%). Bird Coloration — Volumes I and II. Edited by G. Hill and K. McGraw. 2006. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 589 and 477 pages. US$95.00. * Gibson’s Guide to Birdwatching and Conservation. By Merritt Gibson. 2006. Nimbus Publishing Ltd., 3731 Mackin- tosh Street, Box 9166, Halifax, Nova Scotia B3K 5M8 Cana- da. 224 pages. $24.95. * Birds of East Africa — Kenya, Tanzania, Uganda, Rwan- da and Burindi. By T. Stevenson and J. Fanshawe. 2006. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 604 pages. US$35 Paper. Atlas — Birds of Moscow City and the Moscow Region. Pensoft Series Faunistica 54. By M. Kalyakin and V. Voltzit. 2006. (In English and Russian (bilingual). Pensoft Publishers, Geo Miley Street 13a, 1111 Sofia, Bulgaria. 372 pages. EUR 59.90. The Birds of Malawi — An Atlas and Handbook. By Fran- coise Dowsett-Lemaire and Robert Dowsett. 2006. Tauraco Press, Jupille-Liége, Belgium. 556 pages. £25.00 Paper. Birds of Mexico and Central America. By Ber van Perlo. 2006. Princeton University Press, 41 William Street, Prince- ton, New Jersey 08540 USA. 336 pages. $US29.95 Paper. Birds of Moscow City and the Moscow Region. By M. Kalyakin and O.V. Voltzit. 2006. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. In English and Russ- ian (bilingual). 372 pages. EURO 58.00 Cloth. + Birds of Prey Rescue. By Pamela Hickman. 2006. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 66 pages. $9.95 Paper. New Holland Field Guide to the Birds of South-East Asia. By Craig Robson. 2006. New Holland Publishers (UK) Ltd., Garfield House, 86-88 Edgware Road, London W2 2EA United Kingdom. £19.99 Paper. Birds, Mammals and Reptiles of the Galapagos Islands: An identification guide. 2"¢ Edition. By Andy Swash and Robert Still. 2006. Yale University Press, P.O. Box 209040, New Haven, Connecticut US$25 Paper. Birds — A Visual Guide. By Joanna Burger. 2006. Firefly Books Ltd. 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 304 pages. $29.95. Birds of the World. By Les Beletsky. 2006. UBC Press, 2029 West Mall, Vancouver, British Columbia V6T 1Z2 Canada. 628 pages. $55 Cloth. The Black Bear Almanac. By D. Smith. 2006. Globe Pequot Press, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut 0643 USA. US$29.95. Cloth. Fauna De Burpestidae De La Penisula Iberica Y Baleares (Coleoptera). Edited by A Verdugo. [In Spanish.] 2005. Book REVIEWS 129 Pensoft Publishers, Geo Miley Street 13a, 1111 Sofia, Bul garia. 350 pages. EUR 89.00. A Catalogue of Butterflies of the Ex-USSR, with Remarks on Systematics and Nomenclature. By S. Korb. 2005 Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bul garia. 158 pages. EUR 34.00 Paper. Butterflies of Vietnam. Volume 1: Nymphalidae: Satyrinae By A. Monastyrskii. 2005. Pensoft Publishers, Geo Mile Street 13a, 1111 Sofia, Bulgaria. 198 pages. EUR 45.00 Paper Butterflies of the World, Part 22, Nymphalidae X, Charax- es I. By B. Turlin. 2005. Pensoft Publishers, Geo Miley Street 13a, 1111 Sofia, Bulgaria. 40 pages. EUR 38.00 Cloth. Butterflies of the World, Part 23, Papilionidae XII, Par- nassius Apollo III Text. By E. Moehn. {In German] 2005. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bul- garia. 34 pages. EUR 25.00 Cloth. World Butterflies. By Bernard d’ Abrera. 2006. Hill House, Publishers, 491 Illington Road, Ossining, New York 10562 USA. 272 pages. £18.50 Paper. Dodo: The Bird Behind the Legend. By Alan Grihault. 2005. Imprimerie & Papeterie Commerciale Ltee. Port-Louis, Mauritius. 171 pages. £16.99 Paper. * Fishes of the World. 4" edition. J. Nelson. 2006. John Wiley & Sons Canada Ltd. 6045 Freemont Boulevard, Mis- sissauga, Ontario L5R 4J3 Canada. 601 pages. US$125.00, $162.99 Can. Grizzlies and Grizzled Old Men : A Tribute to the Men Who Fought to Save the Great Bear. By Mike Lapinski. 2006, FalconGuide, 246 Goose Lane, P.O. Box 480, Guilford, Connecticutt 06437 USA. 186 pages. US$14.95 Paper. Harvestmen. By R. Pinto-de-rocha, G. Machado and G. Giri- bet. 2006. Harvard University Press, 79 Garden Street, Cam- bridge, Massachusetts 02138 USA. 516 pages. US$125. Lemurs of Madagascar. By R. A. Mittermeier, W. R. Kon- stant, Frank Hawkins, Edward E. Louis et al. 2006. Conser- vation International ,1919 M Street, NW Suite 600, Washing- ton, District of Columbia 20036 USA. 520 pages. USS$25. Paper 2™ edition revised from the original 1994 edition. Field Guide to North Atlantic Wildlife. By N. Proctor and P. Lynch. 2006. Yale University Press, P.O. Box 209040, New Haven, Connecticut USA. 221 pages. US$19.95. * Birding Sites of Nova Scotia —- A Comprehensive, Year- Round Guide for Birders and Other Nature Lovers. By B. Maybank. 2006. Nimbus Publishing Ltd., P.O. Box 9166, Halifax, Nova Scotia B3K 5M8 Canada. 555 pages. $35. The World of the Polar Bear. By Norbert Rosing. 2006. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 204 pages. $45. World Catalogue of the Family Sepsidae (Insecta: Diptera). Zoologicheskie Issledovania (Zoological Studies), Num- ber 8. By A. Ozerov. 2005 Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 76 pages. EUR 29.00 Paper. * Tanagers, Cardinals, and Finches of the United States and Canada. By D. Beadle and J. Rising. 2006. Princeton University Press, 41 William Street, Princeton, New Jersey 08540 USA. 196 pages. US$29.95. Paper. Fauna de Tenebrionidae de La Peninsula Iberica y Balear- es. Volume 1 Lagriniinae y Pimeliinae. Edited by A. Vinolas. 130 C. Cartagena [in Spanish]. 2005. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 428 pages. EUR 110.00. Whalewatcher — A global guide to watching whales, dol- phins and porpoises in the wild. By T. Day. 2006. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 160 pages. $19.95. The Urban Whale. Edited by S. Kraus and R. Rolland. 2006. Harvard University Press, 79 Garden Street, Cambridge, Mass- achusetts 02138 USA. 514 pages. US$55. New Xixuthrina from Indo-Australian Region (Coleoptera, Cerambycidae, Prioninae). By G. Marazzi, V. Marazzi and Z. Komiya. 2006 Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 48 pages. EUR 35.00. BOTANY Aquatic and Wetland Plants of Eastern North America — Volumes I and II. By G. Crow and C. Hellquist. 2006. (paper- back edition) University of Wisconsin Press, 1930 Monroe Street, Madison, Wisconsin 53711 USA. U.S.$45/volume Emily Dickinson’s Herbarium (Facsimile Edition) Intro- duction. By R. Sewall. 2006. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 224 pages. US$135. * Flowers — how they changed the world. By W. Burgur. 2006. Prometheus Books, 59 John Glenn Street, Amherst, New York, USA. 337 pages. US$23. Cloth. Plants on Islands Diversity and Dynamics on a Conti- nental Archipelago. By Martin L Cody. 2006. University of California Press, 2120 Berkeley Way, Berkeley, Califor- nia 94720 USA. 315 pages. US$49.95. Cloth. North American Mushrooms: A Field Guide to Edible and Inedible Fungi. By Dr. Orson Miller and Hope Miller. 2006. Globe Pequot Press, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut 06437 USA. U.S.$25.95 The Tree of Life. By G. Lecointre and H. Le Guyader. 2006. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138. 560 pages. U.S. $39.95. ENVIRONMENT A Complete Guide to Arctic Wildlife. By R. Sale. 2006. Fire- fly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 400 pages. $49.95. Wild Borneo. By Nick Garbutt and Cede Prudente. 2006. New Holland Publishers (UK) Ltd., Garfield House, 86 — 88 Edgware Road, London,W2 2EA United Kingdom. £29.99 Cloth. * Guide to Deserts. Edited by A. Warren and T. Allan. 2006. Firefly Books Ltd., 66 Leek Crescent Richmond Hill, Ontario L4B 1H1 Canada. 240 pages. $19.95. Ecological Nationalisms — Nature, Livelihoods, and Iden- tities in South Asia. Edited by Gunnel Cederlof and K. Sivaramakrishnan. 2006. University of Washington Press, 1326 Fifth Avenue, Suite 555 Seattle, Washington 98101-2604 USA. US$50. Cloth. The Emirates: A Natural History. Edited by Peter Hellyer and Simon Aspinall. Trident Press. 175 Piccadilly, Mayfair, London WIJ 9TB United Kingdom. 428 pages. £65.00 Cloth. THE CANADIAN FIELD-NATURALIST Vol. 120 Endangered — Wildlife on the brink of extinction. By G. McGavin and D. Burnie. 2006. Firefly Books Ltd. 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 192 pages. $35. Evolutionary Dynamics. By M. Nowak. 2006. Harvard Uni- versity Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 312 pages. U.S.$35. The Revenge of Gaia — Why the Earth is fighting back — and how we can still save humanity. By James Lovelock. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN United Kingdom. 177 pages. £16.99 Cloth. Galapagos: A Natural History. By John Kricher. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN United Kingdom. US$19.95, £12.95. Paper. Galapagos Islands Explorer. By Nigel Sitwell. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN United Kingdom. £9.99. Field Guide to Gold Gemstones and Mineral Sites of BC — Volume 2 Revised Edition (Sites within a Day’s Drive to Vancouver). By Rick Hudson. 2006. Harbour Publishing, 4437 Rondeview Road, Madeira Park, British Columbia VON 2HO Canada. 368 pages. $26.95 * A Paddler’s Guide to Killarney and the French River. By K. Callan. 2006. The Boston Mills Press, 132 Main Street, Erin, Ontario NOB 1T0 Canada. 168 pages. $19.95. * Lapland — A Natural History. By D. Redcliff. 2006. Yale University Press, P.O. Box 209040, New Haven, Connecticut 06520-9040 USA. 352 pages. US$60 Cloth. The Man-Eaters of Eden: Life and Death in Kruger Na- tional Park. By Robert Frump. 2006. The Lyons Press, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut 06437 USA. 272 pages. US$24.95 Cloth. Western Newfoundland and Gros Morne National Park. By B. Bursey. 2006. Nimbus Publishing Ltd., 3731 Mackin- tosh Street, Box 9166, Halifax, Nova Scotia B3K 5M8 Cana- da. 88 pages. $22.95. New Zealand — a natural history. By Tui De Roy and Mark Jones. 2006. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 160 pages. $40. A View of the River. By L. Leopold. 2006. Harvard Uni- versity Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 298 pages. US$19.95 + States of Nature. By Tina Loo. 2006. UBC Press, 2029 West Mall, Vancouver, British Columbia V6T 1Z2 Canada. 320 pages. $85 Cloth. Southeast Asian Biodiversity in Crisis. By Navjot S Sodhi and Barry W Brook. 2006. Cambridge University Press, The Edinburgh Building, Shaftesbury Road, Cambridge, CB2 2RU United Kingdom. £65.00 Cloth. The Green Tiger — The Costs of Ecological Decline in the Philippines. By Barbara Goldoftas. 2006. Oxford Universi- ty Press, 198 Madison Avenue, New York, New York 10016 USA. 264 pages. US$74 Cloth, US$24.95 Paper. Vietnam: A Natural History. By Eleanor Jane Sterling, Martha Maud Hurley and Le Duc Minh. 2006. Yale Universi- ty Press P.O. Box 209040, New Haven, Connecticut 06520- 9040 USA. 448 pages. US$40 Cloth. 2006 Book REVIEWS 13] Waterfowl Ecology and Management — 2" Edition. By G. Baldassarre and E. Bolen, 2006. Krieger Publishing P.O. Box 9542, Melbourne, Florida 32902-9542 USA. 580 pages. US$112.50. + Walk around the Pond. By G. Walbauer. 2006. Harvard University Press, 79 Garden Street, Cambridge, Massachu- setts 02138 USA. 286 pages. US$22.95. Wilderness Predators of the Rockies — The Bond Between Predator and Prey. By Mike Lapinski. 2006. Falcon Guide, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut 06437 USA. 208 pages. U.S. $17.95 Paperback The World’s Wild Places. By C. Prior. 2006. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1HI Canada. 224 pages. $49.95, News and Comment Ontario Natural Heritage Information Centre Science and Information Newsletter 11(1) Winter 2006 Contents of this 20-page issue: FEATURE ARTICLE: Trans- ferring conservation science to practitioners — 2005 PRO- GRAM HIGHLIGHTS: Plant communities of Ontario upper great lake marshes — NHIC to track rare disjunct arctic- alpine plants — 2005 species at risk field survey highlights — Rare plant surveys of Long Point region — Natural spaces program — Mapping species at risk with biotics — Two major dragonfly meetings in Ontario — 2005 PROJECT HIGH- LIGHTS: NHIC launches Ontario Odonata atlas website — Aquatic species database — Great Lakes islands conservation blueprint — Lichen biodiversity project — NEWS AND Marine Turtle Newsletter (111) January 2006. 28 pages: EDITORIAL: Does the Mediter- ranean Green Turtle exist? (N. Mrosovsky) — Comment on the guest editorial by Paul J. Ferraro (P.C.H. Pritchard) — Response to Comment by Peter C. H. Pritchard (P. J. Ferraro — ARTICLES: Report on long-term transmitter harness retention by a Leatherback Turtle (S. Dowling, R. Solano, A. Diaz-Merry, J. Ordonez, J. Taylor, D. R. Evans, D. Godfrey, D. Bagley, L. Ehrhart, and S. Eckert — Marine Turtles on the Southern Coast of Bioko Island (Gulf of Guinea, Africa), 2001-2005 (H. Rader, M. A. E. Mba, W. Morra, and G. Hearn — Incidental capture of marine turtles in Marine fisheries of southern Spain (J.C. Baez, J. A. Camidax, and L. Rueda — First evidence of Leatherback movement from Africa to South America (A. Billes, J. Fretey B. Verhage, B. Huijbregts, B. Giffoni, L. Prosdocimi, D. A. Albareda, J.-Y Georges, and M. Tiwari — NOTES: Charybdlis hellerii, a non-indigenous portunid crab from the gastrointestinal contents of a Kemp’s Ridley Sea Turtle (Lepidochelys Kempi) in Georgia, USA (M. Frick and K. Williams) — An update on Eritrea’s marine Wildlife Afield 2(1,2) A British Columbia semi-annual publication (ISSN 1712- 2880) published by The Biodiversity Centre for Wildlife Studies twice a year. Additional information at www.wild lifebc.org. Volume 2, Number 1, contains FROM THE EDITOR: Spinning a web — On the covers — FEATURE ARTICLES: Migratory occurrence and status of select shorebirds in the vicinity of Fort St. John, British Columbia (Chris Siddle) — Survey of Painted Turtles in the Creston Valley, British Columbia, in 2003 and 2004 (Linda M.Van Damme) — NOTES: Changing status of Ring-necked Ducks wintering on Salt Spring Island, British Columbia, 1978-2004 (Robert B. Weeden — Barred Owl feeds on Ban-tailed Pigeon (David C. Schultz) — First winter occurrences of the Cape May Warbler for British Columbia (Phil Ranson) — Burdock entrapment of a Ruby-crowned Kinglet in the Creston Valley — First nest and eggs the Canada Warbler for British Col- umbia (R. Wayne Campbell) — Pinyon Jay (Gymnorhinus cyanocephalus): A new species for British Columbia (Kim Asquith and Malcolm Dennington) — Incidences of Gopher NOTES: Summer experience program at the NHIC — Ontario eastern habitat joint venture — GIS Project — Eastern region Natureserve conference — NHIC data supports the assess- ment of species of global conservation concern in Canada — NHIC joins new biodiversity section of MNR — Publication notices — NHIC staff information. Mailing address for Natural Heritage Information Centre, Ontario Ministry of Natural Resources, 300 Water Street, 2" Floor, North Tower, P.O. Box 7000, Peterborough, Ontario K9J 8M5, Canada; www.nhic.mnr.gov.on.ca/MNR/nhic.cfm. turtle programme and first record of Olive Ridley turtle nesting in the Red Sea (n. Pilcher, S. Mahmud, S. Howe, Y. Tecle- mariam, S. Weldeyohannes, T.Menggsu, and M. Giotom — Epibionts associated with Chelonia mydas from northern Brazil (S. Pereira, E.H. S. M. Lima. L. Ernesto, H. Mathews, and A. Ventura — Small Leatherback found in Dominica (R. Byrne and S. A. Eckert) —MTSG UPDATE — ANNOUNCE- MENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICA- TIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, Centre for Ecology and Conservation, University of Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 263700. Subscriptions and donations towards the production of the MTN can be made online at or postal mail to Michael Coyne (online Editor) Marine Turtle Newsletter, 1 Southampton Place, Durham, North Carolina 27705 USA (e-mail: mcoyne@seaturtle.org). Snake predation in bluebird nest boxes — Unsuccessful nesting attempt of Black-necked Stilt in British Columbia (Ryan Tomlinson) — Status of the Rock Pigeon (Columba livia) in Nelson, British Columbia (Janice E. Arndt) — WILDLIFE DATA CENTRE: Featured species — Thinhorn Sheep (Fred L. Bunnell) — Report of the Wildlife Data Centre: | January 2005 to 30 June 2005 (Michael I. Pres- ton) — BRITISH COLUMBIA ROUND-UP: News of Friends — From the Archives — Final Flight — Publications of In- terest (Compiled and edited by Chris Siddle) — Announce- ments and Meetings. Volume 2, Number 2, contains: FROM THE EDITOR: A turn of the times —On the covers — FEATURE ARTICLES: Field Survey of Red-eared Sliders (Trachemys scripta ele- gans) in the Lower Fraser Valley, British Columbia, in 2005 (Cory G. Bunnell — Origin and current status of the Pacific Pond Turtle (Actinemys marmorata) in British Columbia Francis R. Cook, R. Wayne Campbell and Glenn R. Ryder — Recent range expansion of Sandhill Cranes (Grus canadensus tabida) in southeastern British Columbia (Doug - 132 2004 las Leighton — Diet of Great Horned Owl in the Creston Valley, British Columbia, 1998-2005 (Linda M. Van Damme — NOTES: Anti-predator behaviour by a Mule Deer to Coyotes in Vernon, British Columbia (Chris Siddle) — Attempted capture of a Red-legged Frog by an American Robin (Michael I. Preston) — Evidence of an unusual prey item in a Barred Owl pellet (Christian W. Gronau) — Northernmost nest record of the Flammulated Ow! (Otus flammeolus) in Canada (Karen L. Wiebe — First nesting record of Say’s Phoebe (Sayornis saya) in the East Kootenay, British Columbia (Robert S. Ferguson and Mark Dascher) — Current status of the Black- throated Blue Warbler in British Columbia (Gary S. Davidson — Field observations of Gray Flycatchers breeding at Sum- merland, British Columbia, 1988-2005 (I. Laurie Rockwell) — American Crow predation in an Eared Grebe nesting colony at Rawlings Lake, British Columbia (Chris Siddle) Errata The Canadian Field-Naturalist 119(3) Catling, Paul M. 2005 [2006]. Additions to the flora of the continental Northwest Territories from the Great Slave area. Canadian Field-Naturalist 119(3): 437-440. (page 438) Actaea rubra f. neglecta : This form had previous- ly been reported by Cody 1961. Canadian Field-Naturalist 75(2): 62 from Fort Simpson. Michael Oldham (Ontario Ministry of Natural Resources) reports confirming its pre- sence there in August 2007. NEWS AND COMMENT 133 — SPECIES OF REGIONAL SIGNIFICANCE: Canvasback Fort St. John (Chris Siddle) — Bobolink — Nakusp to Edge- wood (Gary S. Davidson) WILDLIFE DATA CENTRE Featured species — Turkey Vulture (R. Wayne Campbell, Michael I. Preston, Linda M. Van Damme, and Diann Mac- Rae — BRITISH COLUMBIA ROUND-UP: Field notes and caring people — News of Friends — From the archives — Final flight — Publications of interest [Compiled and edited by Chris Siddle}] — Announcements and meetings. Annual memberships in the Biodiversity Centre for Wildlife Studies (which included Wildlife Afield are $30 (individual), $40 (family). $20 (student), $500 (life). Biodiversity Centre for Wildlife Studies, PO Box 6218, Station C, Victoria, British Columbia V9P 5LS Canada. Tel/Fax: 250-477-0465; e-mail: editor@ wildlifebc.org (page 449) Sonchus arvennsis subspecies uliginosus : Thiert reported this in 1963 Sida 1(3): 169 as S. arvensisvar. glabrescens from the Yellowknife Highway (Hwy 3). My appreciation to Michael Oldham for bringing these to my attention. PAUL M. CATLING Agriculture and Agri-Food Canada Ottawa, Ontario 134 THE CANADIAN FIELD-NATURALIST Vol. 119 Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit, 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 publication of the data must have been obtained by the authors. Also authors are ex- pected to have complied with all pertinent legislation regard- ing the study, disturbance, or collection of animals, plants or minerals. The place where voucher specimens have been de- posited, and their catalogue numbers, should be given. Lati- tude and longitude should be included for all individual local- ities where collections or observations have been made. Print the manuscript on standard-size paper, doublespace throughout, leave generous margins to allow for copy mark- ing, and number each page. For Articles and Notes provide a bibliographic strip, an abstract and a list of key words. Generally, words should not be abbreviated but use SI sym- bols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manuscripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in the text or in a sepa- rate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “British” or “American” spellings are acceptable in English but should be consistent within one manuscript. The Oxford English Dictionary, Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Illustrations Photographs should have a glossy finish and show sharp contrasts. Electronic versions should be high resolution. Photo- graphic reproduction of line drawings, no larger than a standard page, are preferable to large originals. Prepare line drawings with India ink on good quality paper and 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, cho- sen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—sometimes extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for pub- lication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by paying $80 for each page, plus $15 for each illustration (any size up to a full page), and up to $80 per page for tables (depending on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Business Manager. If grant or institutional funds are not available, club members and subscribers may apply for a waiver of charges for the first five pages. Limited joumal funds are available to help offset publi- cation charges to authors with minimal financial resources. Requests for financial assistance should be made to the Editor when the manuscript is accepted. Reprints An order form for the purchase of reprints or PDF will accompany the galley proofs sent to the authors, and an invoice will be sent after publication. Payment should be by cheque or money order paid when the invoice is received. FRANCIS R. COOK, Editor TABLE OF CONTENTS (concluded) Volume 120 Number 1 Notes Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine A.C. A. BURCHSTED and FRED BURCHSTED Ring-billed Gull, Larus delawerensis, food piracy on diving ducks WILLIAM J. WALLEY Probable Black Bear, Ursus americanus, retrieval of an Elk, Cervus elaphus, carcass from a small lake in Riding Mountain National Park, Manitoba WILLIAM J. WALLEY Book Reviews ZooLoGyY: The Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi — The Birding Sites of Nova Scotia — The Camplete Fauna of Iran — Fishes of the World — Insects: Their Natural History and Diversity: With a Photographic Guide to Insects of Eastern North America — Secret Weapons: Defenses of Insects, Spiders, Scorpions, and Other Many-legged Creatures — Rodents and Lagomorphs of British Columbia — Tanagers, Cardinals, and Finches of the United States and Canada — Wheatears of Palaeartic: Ecology, Behaviour and Evolution of the Genus Oenanthe Borany: Giant Trees of Western North America and the World ENVIRONMENT: Fire and Avian Ecology in North America: Studies in Avian Biology Number 50 — Fisheries Assessment and Management in Data-limited Situations — Lapland: A Natural History — Fish, Fur and Feathers: Fish and Wildlife Conservation in Alberta, 1905-2005 — The Gulf of Alaska: Biology and Oceanography MISCELLANEOUS: Fossil Legends of the First Americans — A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites within a Day’s Drive of Vancouver New TITLES News and Comment | Ontario Natural Heritage Information Centre Science and Information Newsletter 11(1) Winter 2006 — Marine Turtle Newsletter (111) — Wildlife Afield 2 (1,2) — Errata The Canadian Field-Naturalist 119(3) Advice to Contributors ! | Mailing date of the previous issue 119(4): 25 April 2007 2006 106 109 110 113 120 121 127 128 132 134 THE CANADIAN FIELD-NATURALIST Volume 120 Number 1 Articles Predicting bird oiling events at oil sands tailings ponds and assessing the importance of alternate waterbodies for waterfowl: a preliminary assessment ROBERT A. RONCONI A test of interspecific effects of introduced Grey Squirrels, Sciurus carolinensis, on Douglas’s Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia YEEN TEN HWANG and SERGE LARIVIERE Reponses of syrphids, elaterids, and bees to single-tree selection harvesting in Algonquin Provincial Park, Ontario ERICA NOL, HUME DOUGLAS, and WILLIAM J. CRINS New nesting records of the Le Conte’s Sparrow, Ammospiza leconteii, from northeastern Ontario with some notes on nesting behaviour MICHAEL PATRIKEEV Conditions for sexual interactions between wild Wolves, Canis lupus, and Coyotes, Canis latrans RICHARD P. THIEL Multiple scale den selection by Swift Foxes, Vulpes velox, in southeastern Colorado ANN KITCHEN, ERIC M. GESE, and SARAH G. KuPIS First records of the Northern Long-eared Bat, Myotis septentrionalis, in the Yukon Territory THOMAS S. JUNG, BRIAN G. SLOUGH, DAVID W. NAGORSEN, TANYA A. DEWEY, and TODD POWELL Recent declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces ANTHONY J. ERSKINE First record of the Southern Red-backed Vole, Clethrionomys gapperi, in Newfoundland: Implications for the endangered Newfoundland Marten, Martes americana atrata BRIAN J. HERN, JOHN T. NEVILLE, WILLIAM J. CURRAN, and DEAN P. SNow Long-range homing by an adult female Black Bear, Ursus americanus L. J. LANDRIAULT, M. N. HALL, J. HAme, and F. F. MALLORY Early ontogenetic diet in Wolves, Canis lupus, of coastal British Columbia HEATHER M. BRYAN, CHRIS T. DARIMONT, THOMAS E. REIMCHEN, and PAUL C. PAQUET Extension of Coyote, Canis latrans, breeding range in the Northwest Territories, Canada H. DEAN CLUFF Lake Sturgeon, Acipenser fulvescens, movements in Rainy Lake, Minnesota and Ontario W. EUGENE ADAMS JR., LARRY W. KALLEMEYN, and DAVID W. WILLIS Northern range expansion and invasion by the Common Carp, Cyprinus carpio, of the Churchill River system in Manitoba PASCAL H. J. BADIOU and L. GORDON GOLDSBOROUGH Diversity and range of amphibians of the Yukon Territory BRIAN G. SLOUGH and R. LEE MENNELL Temporal variability of cetaceans near Halifax, Nova Scotia PETER SIMARD, JENIFER L. LAWLOR, and SHANNON GOWANS Winter habitat use by American Marten, Martes americana, in western Alberta boreal forests GILBERT PROULX 2006 39 43 93 100 (continued on inside back cover) ISSN 0008-3550 84 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 120, Number 2 April—June 2006 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patron Her Excellency The Right Honourable Michaille Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- ments of high quality for living things. Honorary Members Edward L. Bousfield Ellaine Dickson George F. Ledingham E. Franklin Pope Charley D. Bird Bruce Di Labio Stewart D. MacDonald William O. Pruitt, Jr. Donald M. Britton R. Yorke Edwards Hue N. MacKenzie Joyce and Allan Reddoch Irwin M. Brodo Anthony J. Erskine Theodore Mosquin Dan Strickland William J. Cody John M. Gillett Eugene G. Munroe John B. Theberge Francis R. Cook C. Stuart Houston Robert W. Nero Sheila Thomson 2006 Council President: Mike Murphy Ronald E. Bedford —_ Diane Kitching Dan Millar Vice-President: Ken Alison Fenja Brodo Karen McLachalan Hamilton Justin Peter Recording Secretary: Susan Laurie-Bourque Julia Cipriani David Hobden Stanley Rosenbaum Treasurer: Frank Pope William J. Cody Diane Lepage Henry Steger Past President: Gary McNulty Francis R. Cook Anne MacKenzie Chris Traynor Susan Howell Gillian Marston Eleanor Zurbrigg To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc@achilles.net. For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. PAP Registration Number 9477. Canada We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward our mailing costs. Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca Copy Editor: Elizabeth Morton Honorary Business Manager: W. J. Cody; Acting Business Manager: E. Franklin Pope, P.O. Box 35069, Westgate P.O., Ottawa, Ontario K1Z 1A2 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwegsc.gc.ca Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen Charles D. Bird Brian W. Coad Donald F. McAlpine Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. Subscriptions and Membership Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 per! year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) andi), $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA), must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available tc 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 Pos’ Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: April-June 2005 (October 2006). Cover: Elk, Cervus elaphus, part of herd near Hartsmore, Ontario. Photographed 1 February 2005 by R. Rosatte. See paper on late! born calf — pages 188-191. Volume 120, Number 2 The Canadian Field-Naturalist —“'BRARY NOV 28 2007 AR VA UNE iD April-June 2006 A Tribute to George Wayne Douglas 1938 — 2005 ener L. PENNY! -'Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, P.O. Box 9993 Stn Prov Govt, | Victoria, British Columbia V8W 9R7 Canada. Penny, Jenifer L. 2006. A tribute to George Wayne Douglas 1938-2005. Canadian Field-Naturalist 120(2): 135-146 Coming from humble beginnings, George Wayne Douglas, with his determination and strong spirit, estab- lished himself as one of British Columbia’s most res- pected botanists. I first met George in 1995 when | began working with him at the British Columbia Con- servation Data Centre (CDC). In time, I came to know him as an adept field botanist, a knowledgeable ecolo- gist, an accomplished author, a practical taxonomist, a cunning businessman, a conservationist at heart, and a generous mentor. During our numerous field trips, George often talked about writing his memoirs and enjoyed recounting the stories and adventures that would go into the chapters. He had a lot of different experiences throughout his life that would have result- ed in an interesting read. He was a born leader: he had a strong character, held his ground on issues, and had a critical, but practical approach. George had a vision for botany in British Columbia and he brought that vision to fruition. _ George (known to family and close friends as Wayne) was born in the Royal Columbian hospital in New Westminster on 22 June 1938. He spent his early years exploring the bushes around the base of Burna- by Mountain near Vancouver, British Columbia, this may have been the root of his inspiration to study botany and ecology later in his life. He grew up in a large family with six brothers and sisters: Bonnie, Mar- ‘ilyn, and David who currently live in British Columbia, ‘and Robert, who lives in South Africa (siblings David ‘Charles and Judy predeceased him). His family was ‘not affluent, so George always aspired to do well for himself, which he certainly accomplished. As a youth, he excelled at sports but wasn’t that jinterested in academics. In high school (Figure 1), he ‘ran record sprints and was an excellent basketball play- ler, as well as an accomplished golfer. He would con- ‘tinue to enjoy golfing throughout his life, playing at least two to three times a week. When he first started, vat about 12 years of age, he used to sneak onto the Van- couver Golf Course, and golf his way to school, hide ' FIGURE |. George at Como Lake High School (circa 1956). his clubs and then golf his way home after school. Later he was on the high school golfing team and was an amateur champion. He always knew how to have fun, especially back in high school, but he also worked hard. One of his favourite jobs at that time was at Labatt’s Brewery where his Friday afternoons were spent “sampling” the products. 136 Despite his lack of interest in scholarly pursuits in high school, George ended up at the University of Washington (College of Forest Resources) in Seattle, Washington, on a track and field, basketball and golf scholarship. George continued to work on his athlet- ics; however, much to his surprise, he became very keen on his studies and excelled in his field. Al- though he never finished high school, during his edu- cational career he ended up receiving three post-grad- uate degrees! George had a scholarship but he still had to work part-time jobs to finance going to university. Amongst his occupations were working in a mill, a cannery and a slaughterhouse. He was also an ambulance and taxi driver. Later in his schooling he got work that better suited his career goals like as a forestry consultant, where he supervised forest inventory, mapping and re- forestation projects. He also worked as an ecological consultant while attending University of Washington and completed inventories in the North Cascades Na- tional Park focussing not only vegetation, but also wildlife, soils and geology. During his bachelor studies, George received tute- lage from two famous Pacific Northwest botanists, Dr. C. Leo Hitchcock and Dr. Art Kruckeberg. In later years, George dedicated one of his books to Dr. Kruck- eberg (Mountain Plants of the Pacific Northwest), in which he wrote that Dr. Kruckeberg had inspired him during his first plant identification attempts and start- ed him on the road to his lifelong career in botany. Somehow, George even ended up getting bestowed with Dr. Hitchcock’s plant press, a true honour for a botanist! As a result of considerable effort during his bachelor degree, he was accepted into graduate school and studied under Dr. Ronald J. Taylor. George re- ceived a Bachelor of Science degree in forestry in 1968 and a Master’s of Science degree in 1970 from Uni- versity of Washington. George’s master’s thesis involved a vegetation study in the subalpine zone of the western North Cascades, Washington, which was published in Arctic and Alpine Research (Douglas 1970). Following his Master’s de- gree, he moved back to Canada to embark on his Ph.D. at the University of Alberta (Department of Botany) in Edmonton, Alberta, with Dr. Larry Bliss. It was a good time to return to Canada since the United States Army was rallying young troops for Vietnam. His doctorate was a study of alpine and high subalpine plant com- munities of the North Cascades Range, in Washington and British Columbia, and he published the research in Ecological Monographs (Douglas and Bliss 1977). In 1974, following his doctorate at University of Alber- ta, he received a Postdoctoral Fellowship in the Depart- ment of Botany to study lichens of the North Cascade Ranges of Washington (Douglas 1974). His graduate work took him to the places he loved — the mountains of the Cascade Range. In fact, there is a special place on Mount Baker called Grouse Ridge where he wanted to have his ashes spread, but alterna- THE CANADIAN FIELD-NATURALIST Vol. 120 tively he thought that Mount Prevost overlooking his home in Duncan would be a fine place too, which is where his family ultimately scattered his ashes. After finishing his graduate studies, George took courses in landscape architecture at Pennsylvania State University, which later enhanced his immense enjoy- ment of gardening at home, but at the time, made him more marketable as a consultant. Then he focussed entirely on developing his consulting firm, Douglas Ecological Consultants Ltd., based out of Victoria, Bri- tish Columbia. His consulting work was to take him all over British Columbia and to the Yukon Territory and resulted in valuable vegetation interpretations and guidance to land managers on the issues of environ- mental impacts and biodiversity monitoring. In 1973 and 1974, he started working in the Yukon Territory, where the alpine was full of new species and breath-taking landscapes. His time there began when he was contracted by the Canadian Wildlife Service to examine the vegetation and soils of Kluane National Park and their vulnerability to land use within the park. George ended up spending eight summers working in the park, a good time for the business, but also some of the best memories of his life. Parks Canada also hired him to work on many im- portant projects in the 1970s. In fact, he was awarded some of Parks Canada’s largest contracts ever. Dur- ing this time he employed numerous prestigious sci- entists and colleagues to provide expertise on his proj- ects. Among other projects, he was asked to evaluate the use and ecological impacts of chemical fire retar- dants in North America and to make recommenda- tions on their future use, to assess the environmental impact of recreational use (mainly human and horse trampling) on different vegetation types in Waterton Lakes National Park, Alberta, and to assess the envi- ronmental impact of five potential transportation cor- ridors in Kluane National Park. He also carried out a four-year biophysical inventory project in Kluane Na- tional Park, which was a multidisciplinary study that included ten professional scientists and reported on landforms, surficial deposits, soils and vegetation. Douglas Ecological Consultants also worked on projects involving vegetation reconnaissance of large tracts of land in British Columbia, examining spruce hybridization in the province, evaluating the British Columbia ecological reserve system, air quality mon- itoring projects using lichens in northern Alberta, and a variety of environmental impact studies. In the early 1980s, George started working on con- tracts for the British Columbia Ministry of Environ- ment (MOB), for which he would work until his retire- ment in 2002. He started out at MOE as a consultant, designing and coordinating a biological monitoring net- work using lichens in northeastern British Columbia. He also investigated the occurrence of alpine plant communities in British Columbia for the Ecological Reserves unit of MOE, and suggested where new eco- logical reserves should be established. 2006 PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS 137 eo: , . ~* a re : he . be Pea oe Pa Se ee alt Se . ’ FIGURE 2. George (left) and Lloyd Freese above the Tatshenshini River. 1992. Photo credit unknown. At that time, he also became a Royal British Colum- bia Museum research associate and started working on a complete taxonomical and ecological treatment of approximately 360 taxa in the Asteraceae (sunflower family) in British Columbia. Unfortunately, only two of the planned three volumes were ever published due to funding difficulties. He was also involved in numer- ous articles and government documents in the 1980s, and he edited and contributed to an attractive and informative coffee table book on Kluane Park, Yukon, called Kluane, Pinnacle of the Yukon (Theberge and Douglas Editors 1980). His work in the Yukon also resulted in the Rare Vascular Plants of the Yukon (Dou- glas et al. 1981). George also established a strong relationship with ecologists in the British Columbia Ministry of Forests (MOF) in the 1980s as well as the 1990s. He worked on floristic manuals and bibliographies for them, and annotated several thousand sheets of British Colum- bia plants. George and his colleagues at MOF (Del Meidinger, Andy Mackinnon and Jim Pojar) felt that it was of utmost importance to have a flora for British Columbia to provide a foundation for all inventory, floristic and conservation work going on within the province. Therefore, he collaborated with them on the very first flora for British Columbia and became the chief editor of the The Vascular Plants of British Col- umbia (Douglas et al. 1991-1994). For the first time ever, botanists had a flora for British Columbia all in one work. Previously, botanists had to carry a vari- able number of floras depending on what part of the province they were working in. In the 1990s, George was also involved in the fol- lowing very popular field guides: Plants of Northern British Columbia (MacKinnon et al. 1992), Plants of the Southern Interior of British Columbia (Parish et al. 1996), and the second, expanded edition of the Moun- tain Plants of the Pacific Northwest (Taylor and Dou- glas 1995). In total, he contributed approximately 45 publications during that decade that greatly enhanced the knowledge of vascular plants in British Columbia and the Pacific Northwest. In addition to his valuable contributions to the botan- ical literature, George also added thousands of valuable collections to major herbaria in Canada. During his career, he collected 13893 specimens, many of which represented new taxa for the province, new range exten- sions for known taxa, rare or at-risk species, and other interesting finds. From his collections in the Cascade Mountains he and John Packer named a new Erigeron (fleabane) for the Pacific Northwest, Erigeron salishii (Salish Daisy) (Douglas and Packer 1988). He also made the first collections in British Columbia of species like Orthocarpus barbatus (Grand Coulee Owl-clover), Erigeron leibergii (Leiberg’s Fleabane), and Tonella tenella (Small-flowered Tonella). 138 In 1991, George started working for the British Columbia government as the Program Botanist at the CDC. This position gave him the opportunity to fur- ther contribute to floristics in British Columbia through the CDC mission of identifying, tracking, and monitor- ing the rarest or most at-risk plants in the province. To accomplish this goal, George continued to collect and amass important collections. In addition, through inventory work, George travelled throughout the prov- ince chasing the plants that were on the CDC’s track- ing list and surveying a wide variety of habitats, includ- ing range lands, wetlands and mountain tops. The year that I joined the CDC, we toured the peaks of the south- em Rocky Mountains for five days via helicopter escort. Because of his background in alpine ecology, mountain excursions were definitely a focus for George, but also critical was our inventory work in the other habitats for rare plants, such as the sand dunes of the west coast of Vancouver Island, the hot springs in the East Koote- nays, and the beaches of Osoyoos Lake. These trips sound like a series of holidays but that was where our work took us to confirm the conservation status of rare and at-risk plants in British Columbia. In order to accurately survey and track the species, George not only saw the need for meticulous inspec- tion of the herbaria collections and for inventory work in poorly surveyed areas as well as in regions under high development pressure, but he also saw the need for an updated flora. Therefore, in the late 1990s he started focusing his efforts on the production of the Illustrated Flora of British Columbia (Douglas et al. 1998-2002). At that time, he also co-authored the Rare Native Vascular Plants of British Columbia (Douglas et al. 2002) so that land use planners and field workers had an easy-to-use reference to the CDC-listed plants. The Illustrated Flora of British Columbia (Douglas et al. 1998-2002) dominated George’s life from the late 1990s to 2002. Since he was so dedicated to its com- pletion, he worked on it at home as much as he did in the office. The production of the flora not only involved coordinating the contributors’ schedules, keeping them on timelines and reviewing the products, but it also involved compiling high quality illustrations (some bor- rowed, others newly commissioned). The maps were a whole other project (published in Volume 8 of the series) — the first complete set of distribution maps for British Columbia plants ever done. It was an enormous effort but with enormous benefits. The inventory work that George was involved with at the CDC not only allowed for more accurate prov- incial listing of rare and at-risk taxa, but also allowed him to prepare numerous status reports on the local southeast Vancouver Island and Okanagan Valley En- dangered and Threatened plant taxa. As a result of these studies, he authored and co-authored 40 Commit- tee on the Status of Endangered Wildlife in Canada (COSEWIC) status reports and update status reports, more than any other contributor of reports to the nation- al species assessment team! THE CANADIAN FIELD-NATURALIST Vol. 120 Along the way, George encouraged many of the in- dividuals that he mentored to co-author status reports. He often invited enthusiastic students and other pro- tégés to join him on field outings where he was very forthcoming about sharing his botanical knowledge. He also spent time helping the Co-operative Educa- tion students that worked at the Royal British Colum- bia Museum and many others that sought his advice. He encouraged his staff with regular bouts of appre- ciation. Every year, at Christmas his botanical support- ers at the CDC were invited to lunch at his favourite spot, The Royal Colwood golf course clubhouse, where patrons have a great view of the lovely grounds with its ancient Douglas-firs and Garry Oaks, manicured greens and ponds. Through his work it was obvious that George was a conservationist at heart. He may not have portrayed the image of one in the typical sense, but he always extolled the virtues of preserving natural areas. In the early 1990s, he participated in the Sierra Club expe- dition down the Tatshenshini River, northern British Columbia, part of a successful campaign to have a pro- tected area established in there (Figure 2). He became a member of the Garry Oak Ecosystem Recovery Team Rare Plant Recovery Implementation Group, and contributed valuable input to the mem- bership discussions. He was also on the Adiantum capillus-veneris (Maidenhair Fern) recovery team and performed an advisory role to the South Okanagan- Similkameen Conservation Program. In his consulting work, he contributed to conservation through research and inventory, but also through work with golf clubs advising them on how to create environmentally friend- ly courses. He also supported an important Nature Conservan- cy of Canada campaign to purchase one of the last remaining sizeable stands of Garry Oak in sem1-pris- tine condition left on southeastern Vancouver Island, a property owned by the Elkington family, located east of Duncan. He did early inventories in the Garry Oak ecosystem, and advised the Nature Conservancy staff about the biodiversity values of the property during the raising of funds to purchase it. In 1999, George had the opportunity to conduct the most intensive and objective study of a Garry Oak woodland ever done in British Columbia on this very property, later coined the “Cow- ichan Garry Oak Preserve”. The study examined the composition, phenology, stand structure, and soils of the Garry Oak woodland. Phenological data was ob- tained on a weekly basis from March until September and soils were also examined in detail from three soil pits. In 2004, Douglas Ecological Consultants Ltd. repeated the vegetation component of the study to mon- itor changes after five years (Smith and Douglas 2005). During his government career, George reached out to many different groups to communicate his message to protect rare and at-risk plants. In particular, he estab- lished a relationship with Osoyoos Indian Band, and befriended Environmental Officer Ron Hall. He spent 2006 many summers doing inventories on Osoyoos band lands monitoring the special habitats that they have conserved over the years in the region, while all sur- rounding properties have been converted to vineyards, orchards, golf courses, and residential and commercial developments. Even after he retired from government in 2002, he continued to work with the band helping them with invasive species control in sensitive rare plant habitats. When George retired from government after 11 years of service in early 2002, he started up Douglas Eco- logical Consultants Ltd. full-time once again. He was very excited about getting out in a consulting capacity like the “old days” on some field trips since the inven- tory work had been scant in recent years with the gov- ernment. In his capacity as a consultant to government, he became active in a variety of interesting botanical and ecological projects and produced many reports. In the last three years of his life, he worked on numer- ous status reports, four species at risk recovery strate- gies, and one multi-species recovery strategy. Sadly, his life and work were cut short by a long battle with cancer. He died on 10 February, 2005 at Cowichan District Hospital, Duncan, British Columbia. He left behind wife Sylvia, son Geordie, daughter-in-law Mary McPhail, and grandchildren Mackenzie and Ryder, his four siblings and numerous nieces and nephews. He has, however, left a truly inspiring legacy. His botanical contributions are used often by many plant lovers, botanists and ecologists in British Columbia and beyond. He will be remembered as a leader, not a fol- lower, who made things happen. He was hard-working and passionate about his work. His lengthy bibliogra- phy and professional experience, which follows, will attest to this assertion. Other tributes to George W. Douglas appeared in Botanical Electronic News (BEN), the first, an obitu- ary written by his MOF colleagues, Del Meidinger, Jim Pojar and Andy MacKinnon, and the second, a tribute to George’s immense bibliography (http://www. ou.edu/cas/botany-micro/ben/ben344.html; http://www. ou.edu/cas/botany-micro/ben/ben351.html). The bib- liography found in this article is an updated version of the one that was published in October of 2005 in “BEN # 351” and is found at http://www.ou.edu/cas/ botany-micro/ben/suppl/35 1/gwdouglas.html. An obituary also appeared in the local Duncan and Victoria papers that featured one of George’s favourite pictures with his beloved cat, Garfield, where the two of them are reading a Czech satirical, antimilitaristic novel from the First World War given to him many years ago by a valued colleague (Figure 3; http://www. obituariestoday.com/Obituaries/ObitSearchList.cfm). Acknowledgments Thanks to Sylvia Douglas for important insights into an era in George’s life that pre-dates me and for editing help. Thanks also to Brenda Costanzo, Adolf Ceska, PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS 139 esvesebere the: 7 ee. FIGURE 3. George and his beloved cat, Garfield. Circa 1993. Photo by Sylvia Douglas. Sharon Hartwell, Diana Demarchi, Frank Lomer, Marie Malo (née Fontaine), Gail Harcombe, Marta Donovan, and Del Meidinger who graciously provided review comments. George would have appreciated that they contributed to the article. Positions in Scientific and Professional Societies Member: Northwest Scientific Association; Ottawa Field-Naturalists’ Club; Rare Plant Consortium; Asso- ciation of Professional Biologists Research Associate: Royal British Columbia Muse- um (1991-2004). Dissertations and Post-doctoral Fellowships Douglas, G. W. 1970. A vegetation study in the subalpine zone of the western North Cascades, Washington. M.Sc. thesis, University of Washington, Seattle, Washington. 293 pages. Douglas, G. W. 1973. Alpine plant communities of the North Cascades Range, Washington and British Columbia. Ph.D. thesis, University of Alberta, Edmonton, Alberta. 145 pages. Postdoctoral Fellowship: 1973-1974, Department of Botany, University of Alberta, Edmonton, Alberta. Publications Douglas, G. W., and R. J. Taylor. 1970. Contributions to the flora of Washington. Rhodora 72: 496-501. 140 Franklin, J. F., W. H. Moir, G. W. Douglas, and C. Wiberg. 1971. Invasion of subalpine meadows by trees in the Cas- cades Range, Washington and Oregon. Arctic and Alpine Research 3: 215-224. Douglas, G. W., and T. M. Ballard. 1971. Effects of fire on alpine plant communities in the North Cascades, Wash- ington. Ecology 52: 1058-1064. Douglas, G. W. 1972. Subalpine plant communities of the western North Cascades, Washington. Arctic and Alpine Research 4: 147-166. Douglas, G. W., and R. J. Taylor. 1972. The biosystematics, chemotaxonomy, and ecology of Claytonia lanceolata Pursh. in western Washington. Canadian Journal of Botany 50: 2177-2187. Douglas, G. W., D. B. Naas, and R. W. Naas. 1973. New plant records and ranges for Washington. Northwest Sci- ence 47: 105-108. Douglas, G. W., and J. Ramsden. 1973. Representation of leaf orientation data using equal-area projection techniques. Canadian Journal of Botany 51: 1081-1088. Taylor, R. J., G. W. Douglas, and L. M. Sunquist. 1973. Contributions to the flora of Washington: IT. Northwest Sci- ence 47: 169-179. Douglas, G. W. 1974. Lichens of the North Cascades Range, Washington. The Bryologist 77: 582-592. Douglas, G. W. 1974. Montane zone vegetation of the Alsek River region, southwestern Yukon. Canadian Journal of Botany 52: 2505-2532. Taylor, R. J., and G. W. Douglas. 1975. Mountain wildflow- ers of the Pacific Northwest. Binford and Mort, Portland, Oregon. 176 pages. Douglas, G. W., and D. H. Vitt. 1976. Moss-lichen flora of St. Elias-Kluane Ranges, southwestern Yukon. The Bry- ologist 79: 437-456. Douglas, G. W., K. E. Denford, and I. Karas. 1977. A con- tribution to the taxonomy of Antennaria alpina var. media, A. microphylla, and A. umbrinella in western North Amer- ica. Canadian Journal of Botany 55: 925-933. Douglas, G. W., and L. C. Bliss. 1977. Alpine and high sub- alpine plant communities of the North Cascades Range, Washington and British Columbia. Ecological Monographs 47: 113-150. Douglas, G. W., and G. Ruyle-Douglas. 1978. Contributions to the floras of British Columbia and the Yukon Territory I. Vascular Plants. Canadian Journal of Botany 56: 2296- 2302. Douglas, G. W., and G. Ruyle-Douglas. 1978. Nomenclatur- al changes in the Asteraceae of British Columbia. I. Sene- cioneae. Canadian Journal of Botany 56: 1710-1711. Taylor, R. J., D. B. Naas, R. B. Naas, and G. W. Douglas. 1978. Contributions to the flora of Washington. III. North- west Science 52: 220-225. Taylor, R. J., and G. W. Douglas. 1978. Plant ecology and natural history of Chowder Ridge, Mt. Baker: A potential alpine research natural area in the western North Cascades. Northwest Science 52: 35-50. Douglas, G. W. 1980a. The flora of the Haines Road region, northwestern British Columbia. British Columbia Provin- cial Museum Occasional Paper. Douglas, G. W. 1980b. Vegetation. Jn Biophysical inventory studies of Kluane National Park. Parks Canada, Winnipeg. The flora of the Haines Road region, Northwestern British Columbia. British Columbia Provincial Museum Occa- sional Paper. THE CANADIAN FIELD-NATURALIST Vol. 120 Douglas, G. W., and W. L. Peterson. 1980. Contributions to the floras of British Columbia and the Yukon Territory. II. Mosses and Lichens. Canadian Journal of Botany 58: 2145-2147. Murray, D. F., and G. W. Douglas. 1980. The Green Man- tle. Pages 52-63 in Kluane, Pinnacle of the Yukon. Edited by J. Theberge and G. W. Douglas. Doubleday Canada Ltd., Toronto, Ontario. 175 pages. Theberge, J., and G. W. Douglas. 1980. Kluane, Pinnacle of the Yukon. Doubleday Canada Ltd. Toronto, Ontario. 175 pages. Douglas, G. W., W. L. Peterson, and A. C. Skorepa. 1981. A preliminary checklist of mosses and lichens in the Fort McMurray area, Alberta. Canadian Journal of Botany 59: 1456-1464. Douglas, G. W., G. W. Argus, L. Dickenson, and D. Brun- ton. 1981. Rare vascular plants of the Yukon. National Museum of Canada Syllogeus 28. 61 pages. Douglas, G. W., and M. J. Ratcliffe. 1981. Some rare plant collections, including three new records for Canada, from Cathedral Provincial Park, southern British Columbia. Canadian Journal of Botany 59: 1537-1538. Douglas, G. W. 1982. The sunflower family (Asteraceae) of British Columbia. Volume I. Senecioneae. British Columbia Provincial Museum Occasional Paper Series 23. 180 pages. Douglas, G. W., A. Ceska, and G. G. Ruyle. 1983. A floris- tic bibliography for British Columbia. Land Management Report Number 15. British Columbia Ministry of Forests, Victoria, British Columbia. 143 pages. Straley, G. B., R. L. Taylor, and G. W. Douglas. 1985. Rare Vascular Plants of British Columbia. Canada National Museum Syllogeus 59. 165 pages. Douglas, G. W. 1986. Nomenclatural changes in the Aster- aceae of British Columbia. II Astereae, Eupatoreae, Inuleae, Lactuceae and Heliantheae. Canadian Journal of Botany 64: 2726-2727. Douglas, G. W., and J. G. Packer. 1988. Erigeron salishii, a new Asteraceae for British Columbia. Canadian Journal of Botany 66: 414-416. Douglas, G. W., G. B. Straley, and D. Meidinger. 1989. The vascular plants of British Columbia. Part 1 - Gymnosperms and Dicotyledons (Aceraceae through Cucurbitaceae). Spe- cial Report Series Number 1. British Columbia Ministry of Forests, Victoria, British Columbia. 208 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1990. The Vascular Plants of British Columbia. Part 2 — Dicotyledons (Diapensiaceae through Portulaceae). Special Report Series Number 2. British Columbia Ministry of Forests. Victoria, British Columbia. 155 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1991. The Vascular Plants of British Columbia. Part 3 — Dicotyledons (Primulaceae through Zygophyllaceae) and Pteridophytes. Special Report Series Number 3. British Columbia Min- istry of Forests, Victoria, British Columbia. 177 pages. Douglas, G. W. 1992. Composites, Ferns and Allies. Pages 95-120 in Plants of Northern British Columbia. Edited by A. Mackinnon, J. Pojar and R. Coupé. Lone Pine Publish- ing, Edmonton, Alberta. 351 pages. Douglas, G. W. 1993. Tracking rare British Columbian vascu- lar plants with a computerized data bank. Iiamna 2(2): 1. Douglas, G. W., G. B. Straley, and D. Meidinger. 1994. The Vascular Plants of British Columbia. Part 4 — Monocotyle- dons. British Columbia Ministry of Forests, Victoria, Bri- tish Columbia. 255 pages. 2006 Douglas, G. W., J. Pojar, D. Meidinger, and K. McKeown. 1994. Rare vascular plant collections from the St. Elias Mountains, northwestern British Columbia. Canadian Field-Naturalist 108(4): 391-396. Straley, G. B., and G. W. Douglas. 1994. Rare and endan- gered vascular plants — an update. Pages 95-112 in Bio- diversity in British Columbia Our Changing Environment Edited by L. E. Harding and E. McCullum. Environment Canada, Canadian Wildlife Service, Pacific and Yukon Region. Vancouver, British Columbia. 425 pages. Douglas G. W. 1995. The sunflower family (Asteraceae) of British Columbia. Volume II. Astereae, Anthemdieae, Eupa- toreae and Inuleae. Royal British Columbia Museum, Vic- toria. 393 pages. Taylor, R. J., and G. W. Douglas. 1995. Mountain plants of the Pacific Northwest: a field guide to Washington, west- ern British Columbia and southeastern Alaska. Mountain Press Publishing Company, Missoula, Montana. 437 pages. Douglas, G. W. 1996. Asteraceae: sunflower family. Pages 106-145 in Plants of the Southern Interior of British Colum- bia. Edited by R. Parish, R. Coupé and D. Lloyd. Lone Pine Publishing, Edmonton, Alberta. 463 pages. Douglas, G. W. 1996. Endemic vascular plants of British Columbia and immediately adjacent regions. Canadian Field-Naturalist 110(3): 387-391. Douglas, G. W., and J. M. Illingworth. 1997. Status of the white-top aster, Aster curtus (Asteraceae), in Canada. Cana- dian Field-Naturalist 111(4): 622-627. Douglas, G. W., F. Lomer, and H. L. Roemer. 1998. New or rediscovered native vascular plant species in British Columbia. Canadian Field-Naturalist 1 12(2): 276-279. Douglas, G. W., G. B. Straley, and D. V. Meidinger. 1998. Rare Native Vascular Plants of British Columbia. British Columbia Ministry of Environment, Lands and Parks, Vic- toria, British Columbia. 423 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998. Illustrated Flora of British Columbia. Volume 1: Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998. Illustrated Flora of British Columbia. Volume 2: Dicotyledons (Balsaminaceae through Cucurbitaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 401 pages. Douglas, G. W., and J. M. Illingworth. 1998. Status of the Water-plantain Buttercup, Ranunculus alismifolius var. alismifolius (Ranunculaceae), in Canada. Canadian Field- Naturalist 112(2): 280-283. Douglas, G. W., and M. Ryan. 1998. Status of the Yellow Montane Violet, Viola praemorsa ssp. praemorsa (Vio- laceae) in Canada. Canadian Field-Naturalist 112(3): 491- 495. Jamison, J. A., and G. W. Douglas. 1998. Status of the Coastal Wood Fern, Dryopteris arguta (Dryopteridaceae), in Canada. Canadian Field-Naturalist 112(2): 284-288. Penny, J. L., G. W. Douglas, and G. A. Allen. 1998. Status of the Bearded Owl-clover, Triphysaria versicolor ssp. ver- sicolor (Scrophulariaceae), in Canada. Canadian Field- Naturalist 112(3): 481-485. Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illustrat- ed Flora of British Columbia. Volume 3: Dicotyledons (Diapensiaceae through Onagraceae). British Columbia PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS 14] Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia 401 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illus trated Flora of British Columbia. Volume 4: Dicotyledons (Orobanchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Col- umbia Ministry of Forests. Victoria, British Columbia 401 pages. Douglas, G. W., and M. Ryan. 1999. Status of the Golden Paintbrush, Castilleja levisecta (Scrophulariaceae), in Can- ada. Canadian Field-Naturalist 113(2): 299-301. Lomer, F., and G. W. Douglas. 1999. Additions to the flora of the Queen Charlotte Islands, British Columbia. Cana- dian Field-Naturalist 113(2): 235-240. Miller, M. T., and G. W. Douglas. 1999. Status of Lyall’s Mariposa Lily, Calochortus lyalli (Liliaceae), in Canada. Canadian Field-Naturalist 1 13(4): 652-658. Penny, J. L., and G. W. Douglas. 1999. Status of the Tall Bugbane, Cimicifuga elata (Ranunculaceae), in Canada. Canadian Field-Naturalist 113(3): 461-465. Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illustrat- ed Flora of British Columbia. Volume 5: Dicotyledons (Salicaceae through Zygophyllaceae and Pteridophytes). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 389 pages. Douglas, G. W., and J. A. Jamison. 2001. Status of Scouler’s Corydalis, Corydalis scouleri (Fumariaceae), in Canada. Canadian Field-Naturalist 1 15(3): 455-459. Douglas, G. W. 2001. Abronia umbellata ssp. acutalata: Rarest plant on the planet or the rarest plant in Canada? Menziesia 6: 4-5. Donovan, M. T., and G. W. Douglas. 2001. Status of snake- root, Sanicula arctopoides (Apiaceae), in Canada. Canadian Field-Naturalist 115(3): 466-471. Douglas G. W., D. Meidinger, and J. Pojar. 2001. Illustrated Flora of British Columbia. Volume 6: Monocotyledons (Acoraceae to Najadaceae). British Columbia Ministry of Environment, Lands and Parks, British Columbia Min- istry of Forests, Victoria, British Columbia. 361 pages. Douglas G. W., D. Meidinger, and J. Pojar. 2001. Illustrated Flora of British Columbia. Volume 7: Monocotyledons (Orchidaceae to Zosteraceae). Ministry of Sustainable Resource Management and British Columbia Ministry of Forests. Victoria, British Columbia. 379 pages. Douglas, G. W., and J. Pojar. 2001. Trillium ovatum Pursh variety hibbersonii (Taylor et Szczawinski) Douglas ef Pojar, variety nova. Canadian Field-Naturalist 115(2): 343. Douglas, G. W., and M.W. Ryan. 2001. Status of Deltoid Bal- samroot, Balsamorhiza deltoidea (Asteraceae), in Canada. Canadian Field-Naturalist 115(3): 451-454. Penny, J. L., and G. W. Douglas. 2001. Status of Purple Sanicle, Sanicula bipinnatifida (Apiaceae), in Canada. Canadian Field-Naturalist 1 15(3): 460-465. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare Native Vascular Plants of British Columbia. Second edi- tion. Province of British Columbia, Victoria, British Colum- bia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2002. Ilustrat- ed Flora of British Columbia. Volume 8: General Sum- mary, Maps and Keys. British Columbia Ministry of Sus- tainable Resource Management and British Columbia Ministry of Forests. Victoria, British Columbia. 457 pages. 142 Douglas, G. W. 2005. Conservation evaluation of Lemmon’s Holly Fern, Polystichum lemmonii (Dryopteridaceae), a Threatened fern in Canada. Canadian Field-Naturalist 118(2): 164-168. Douglas, G. W., and J. L. Penny. 2005. Conservation eval- uation of Howell’s Triteleia, Triteleia howellii (Liliaceae), an Endangered lily in Canada. Canadian Field-Naturalist 118(2): 174-178. Douglas, G. W., and J. M. Illingworth. 2005. Conservation evaluation of the Pacific population of Tall Woolly-heads, Psilocarphus elatior (Asteraceae), an Endangered herb in Canada. Canadian Field-Naturalist 118(2): 169-173. Smith, T., G. W. Douglas, and A. Harris. 2005. Conserva- tion evaluation of Small-flowered Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada. Canadian Field-Nat- uralist 118(2): 179-184. Douglas, G. W., and J. Desrosiers. 2006. Conservation eval- uation of Pacific Rhododendron (Rhododendron macro- phyllum), in Canada. Canadian Field-Naturalist 120(2): 169-174. Douglas, G. W., and J. L. Penny. 2006. Conservation eval- uation of Slender Collomia, Collomia tenella, in Canada. Canadian Field-Naturalist 120(2): 175-178. Douglas, G. W., and Jenifer L. Penny. 2006. Conservation evaluation of the Small-flowered Tonella, Tonella tenella, in Canada. Canadian Field-Naturalist 120(2): 179-182. Douglas, G. W., J. L. Penny, and K. Barton. 2006. Conser- vation evaluation of the Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada. Canadian Field- Naturalist 120(2): 163-168. Douglas, G. W., J. L. Penny, and K. Barton. 2006. Conser- vation evaluation of the Stoloniferous Pussytoes, Antennar- ia flagellaris, in Canada. Canadian Field-Naturalist 120(2): 183-187. Douglas, G. W., and M. W. Ryan. 2006. Conservation eval- uation of Prairie Lupine, Lupinus lepidus var. lepidus, in Canada. Canadian Field-Naturalist 120(2): 147-152. Douglas, G. W., and M. W. Ryan. 2006. Conservation eval- uation of the Seaside Bird’s-foot Trefoil, Lotus formosis- simus, in Canada. Canadian Field-Naturalist 120(2): 153- 156. Douglas, G. W., and M. J. Oldham. Conservation evaluation of Scarlet Ammannia, Ammania robusta, in Canada. Cana- dian Field-Naturalist (submitted). Douglas, G. W., and M. J. Oldham. Conservation evaluation of Toothcup, Rotala ramosior, in Canada. Canadian Field- Naturalist (submitted). Unpublished Reports Douglas, G. W. 1971. An ecological survey of potential nat- ural areas in the North Cascades National Park Complex. Unpublished report submitted to the Intercampus Educa- tional and Scientific Preserves Committee. 137 pages. Douglas, G. W. 1974. A reconnaissance survey of the vegeta- tion of Kluane National Park. Report submitted to Cana- dian Wildlife Service, Edmonton, Alberta. 219 pages. Douglas, G. W. 1974. Ecological impact of chemical fire retardants: a review. Report submitted to Northern Forestry Research Centre, Environment Canada, Edmonton, Alber- ta. NOR-X-109. 33 pages. Douglas, G. W., and L. J. Knapik. 1974. Montane zone soil characteristics in Kluane National Park. Report submitted to Canadian Wildlife Service, Edmonton, Alberta. THE CANADIAN FIELD-NATURALIST Vol. 120 Blood, D. A., G. W. Douglas, and L. J. Knapik. 1975. Soil, vegetation and wildlife resources of five potential transportation corridors in Kluane National Park, Yukon. Report submitted to Donald A. Blood & Associates Ltd., Nanaimo, British Columbia. 460 pages. Douglas, G. W. 1975. An evaluation of the examination and selection of ecological reserves in Priority Area one, west- central British Columbia. Report submitted to British Col- umbia Ministry of Forests, Victoria, British Columbia. 14 pages. Douglas, G. W. 1975. Spruce (Picea) hybridization in west- central British Columbia. Report submitted to British Columbia Ministry of Forests, Victoria, British Columbia. 15 pages. Douglas, G. W., J. A. Nagy, and G. W. Scotter. 1975. Effects of human and horse trampling on natural vegetation, Wat- erton Lakes National Park. Report submitted to Canadian Wildlife Service, Edmonton, Alberta. 135 pages. Douglas, G. W. 1977. Vegetation. Jn Environmental impact statement, Shakwak Highway improvement, British Colum- bia and Yukon, Canada. Two volumes. Report submitted to Department of Public Works Canada, and United States Department of Transportation. Douglas, G. W. 1978. The status of some rare plants in the Haines Junction area, Yukon Territory. Report submitted to Department of Public Works, Canada, Whitehorse, Yukon Territory. 10 pages. Douglas, G. W. 1979. An initial environmental impact assess- ment of the vegetation and flora along the Watson Lake- Haines Junction section of the Alaska Highway. Report submitted to Department of Public Works, Canada, Van- couver, British Columbia. 34 pages. Douglas, G. W. 1979. Vegetation. In Shakwak Highway proj- ect reclamation program formulation. I. Technical report of soil and plant community types and reclamation materials and methods suitable for use in segments one to eight. Report submitted to Department of Public Works, Canada, Whitehorse, Yukon Territory. Douglas, G. W. 1980. Alpine plant communities of British Columbia and their occurrence in existing or proposed Eco- logical Reserves. Report submitted to Ecological Reserves Unit, Ministry of the Environment, Victoria, British Colum- bia. Douglas, G. W. 1980. Vegetation. Jn Biophysical inventory studies of Kluane National Park. Report submitted to Parks Canada, Winnipeg, Manitoba. Blood, D.A., E. A. Harding, M. E. Walmsley, and G. W. Douglas. 1987. A preliminary assessment of resources in the Strathcona Recreation Area. Report submitted to British Columbia Parks and Outdoor Recreation Division, North Vancouver, British Columbia. Lea, E. C., and G. W. Douglas. 1991. Endangered and Threatened Plant species of the southern interior of Bri- tish Columbia — Protecting the insignificant. Pages 59-66 in Community Action for Endangered Species — A public symposium on British Columbia Threatened and Endan- gered Species and Their Habitats. Edited by S. Rautio. Federation of British Columbia Naturalists and Northwest Wildlife Preservation Society, Vancouver, British Colum- bia. 238 pages. Douglas, G. W. 1995. Status Report on the Rabbit-brush goldenweed (Ericameria bloomeri [A. Gray] J. F. Mc- Bride). Unpublished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 10 pages. 2006 Illingworth, J. M., and G. W. Douglas. 1995a. Status report on the slender woolly-heads, Psilocarphus tenellus Nutt. var. tenellus. Unpublished report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 19 pages. Illingworth, J. M., and G. W. Douglas. 1995b. Status report on the tall woolly-heads, Psilocarphus elatior (A. Gray) A. Gray. Unpublished report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 19 pages. Ryan, M., and G. W. Douglas. 1995a. Status report on the golden paintbrush, Castilleja levisecta, in Canada. Unpub- lished report submitted to Committee on the Status of En- dangered Wildlife in Canada, Ottawa, Ontario. 26 pages. Ryan, M., and G. W. Douglas. 1995b. Status report on the yellow montane violet, Viola praemorsa var. praemorsa, in Canada. Unpublished report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 22 pages. Desrosiers, J., and G. W. Douglas. 1997. Status Report on Pacific rhododendron, Rhododendron macrophyllum D. Don ex Don (Ericaceae), in Canada. Committee on the Sta- tus of Endangered Wildlife in Canada, Ottawa, Ontario. 22 pages. Douglas, G. W., and J. M. Illingworth. 1996. Status report on the white-top aster, Aster curtus Cronq. In Canada. Un- published report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 26 pages. Illingworth, J. M., and G. W. Douglas. 1996. Status report on the water-plantain buttercup, Ranunculus alismaefolius Geyer ex Benth. var. alismaefolius. Unpublished report submitted to Committee on the Status of Endangered Wild- life in Canada, Ottawa, Ontario. 24 pages. Ryan, M. W., and G. W. Douglas. 1996a. Status report on the deltoid balsamroot, Balsamorhiza deltoidea Nutt. in Canada. Unpublished report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 30 pages. Ryan, M., and G. W. Douglas. 1996b. Status report on the seaside birds-foot lotus, Lotus formosissimus in Canada. Unpublished report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 25 pages. Ryan, M., and G. W. Douglas. 1996c. Status report on the prairie lupine, Lupinus lepidus var. lepidus, in Canada. Un- published report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 26 pages. Penny, J. L., and G. W. Douglas. 1997. Status report on bearded owl-clover, Triphysaria versicolor ssp. versicolor, in Canada. Unpublished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 29 pages. Jamison, J. A., and G. W. Douglas. 1998. Status report on coastal wood fern, Dryopteris arguta, in Canada. Unpub- lished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 23 pages. White, D. J., and G. W. Douglas. 1998. Update COSEWIC status report on giant helleborine orchid (Epipactis gigan- tea). Unpublished report submitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 15 pages. White, D. J., and G. W. Douglas. 1998. Update COSEWIC status report on Southern Maidenhair Fern (Adiantum capil- lus-veneris). Unpublished report submitted to Committee PENNY: TRIBUTE TO GEORGE WAYNE DOUGLAS 143 on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 14 pages. Douglas, G. W., and M. J. Oldham. 1999. Status report on scarlet ammannia, Ammannia robusta, in Canada. Unpub lished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 26 pages Douglas, G. W., and M. J. Oldham. 1999. Status report on toothcup, Rotala ramosior, in Canada. Unpublished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 25 pages. Fontaine, M., and G. W. Douglas. 1999. Rare Grassland Plants. Species and Ecosystems at Risk in British Colum- bia series. British Columbia Ministry of Environment, Lands and Parks, Victoria, British Columbia. 4 pages. Fontaine, M., and G. W. Douglas. 1999. Rare Forest Plants of Southern British Columbia. Species and Ecosystems at Risk in British Columbia series. British Columbia Min- istry of Environment, Lands and Parks, Victoria, British Columbia. 4 pages. Donovan, M., and G. W. Douglas. 2000. Status report on snake-root sanicle, Sanicula arctopoides, in Canada. Un- published report submitted to the Committee on the Sta- tus of Endangered Wildlife in Canada, Ottawa, Ontario. 20 pages. Douglas, G. W., and J. A. Jamison. 2000. Status report on Scouler’s corydalis, Corydalis scouleri, in Canada. Unpub- lished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 18 pages. Douglas, G. W., and J. L. Penny. 2000. New vascular plant species to the British Columbia flora. Menziesia 5(4): 5. Douglas. G. W., J. L. Penny, and M. Donovan. 2000. Sta- tus reports on two Sanicula species. Menziesia 5(4): 4-S. Miller, M., and G. W. Douglas. 2000. Status report on Lyall’s mariposa lily, Calochortus lyallii, in Canada. Unpublished report submitted to the Committee on the Status of En- dangered Wildlife in Canada, Ottawa, Ontario. 32 pages. Penny, J. L., and G. W. Douglas. 2000. Status report on purple sanicle, Sanicula bipinnatifida, in Canada. Unpub- lished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 21 pages. Penny, J. L., and G. W. Douglas. 2000. Status report on tall bugbane, Cimicifuga elata, in Canada. Unpublished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 22 pages. Penny, J. L., and G. W. Douglas. 2000. Tracking rare native vascular plants in British Columbia. Pages 75-77 in Pro- ceedings of a conference on the biology and manage- ment of species and habitats at risk. Edited by L. M. Dar- ling. British Columbia Ministry of Environment, Lands and Parks, Victoria, British Columbia. Douglas, G. W., J. L. Penny, and R. E. Maxwell. 2001. Composition, phenology, stand structure and soils of a Quercus garryana (Garry Oak) woodland community at Cowichan Garry Oaks Preserve, Quamichan Lake, Van- couver Island, British Columbia, Nature Conservancy of Canada. Victoria, British Columbia. 53 pages. Smith, T. W., G. W. Douglas, and A. Harris. 2001. Update COSEWIC Status Report on Small-Flowered Lipocarpha, Lipocarpha micrantha, in Canada. Unpublished report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 23 pages. Douglas, G. W. 2003. COSEWIC Status Report on Baikal Sedge, Carex sabulosa, in Canada. Unpublished report sub- mitted to Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 22 pages. 144 Douglas, G. W. 2003. COSEWIC Status Report on Lem- mon’s holly fern, Polystichum lemmonii, in Canada. In COSEWIC Assessment and Status Report on Lemmon’s Holly Fern, Polystichum lemmonii, in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 15 pages (Available online at: http://www.sara registry.gc.ca/status/status_e.cfm). Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status report on Howell’s Triteleia, Triteleia howellii, in Canada. In COSEWIC Assessment and Status report on Howell’s Triteleia, Triteleia howellii, in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 21 pages (Available online at: http://www.sararegistry.gc. ca/status/status_e.cfm). Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status report on Slender Collomia, Collomia tenella, in Canada. In COSEWIC Assessment and Status report on Slender Collomia, Collomia tenella, in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 15 pages (Available online at: http://www.sararegistry.gc. ca/status/status_e.cfm). Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status report on the Small-flowered Tonella, Tonella tenella, in Canada. In COSEWIC Assessment and Status report on the Small-flowered Tonella, Tonella tenella, in Canada. Committee on the Status of Endangered Wildlife in Cana- da, Ottawa, Ontario. 16 pages (Available online at: http:// Wwww-.Ssararegistry.gc.ca/status/status_e.cfm). Douglas, G. W., J. L. Penny, and K. Barton. 2003. COSEWIC Status Report on Dwarf Woolly-heads, Psilocar- phus brevissimus var. brevissimus, in Canada. In COSEWIC Status Assessment and Report on Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada. Com- mittee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 21 pages (Available online at: http://www. sararegistry.gc.ca/status/status_e.cfm). Douglas, G. W., J. L. Penny, and K. Barton. 2003. COSEWIC Status Report on Stoloniferous Pussytoes, Antennaria flagellaris, in Canada. In COSEWIC Assess- ment and Status Report on Stoloniferous Pussytoes, Antennaria flagellaris, in Canada. Committee on the Sta- tus of Endangered Wildlife in Canada Ottawa, Ontario. 19 pages (Available online at: http://www.sararegistry.gc. ca/status/status_e.cfm). Douglas, G. W., J. Pojar, P. Sheppard, and S. Walker. 2003. Gwaii Haanas Rare Vascular Plant Survey — Field Report. Unpublished report submitted to Parks Canada, Pacific and Yukon Region, Vancouver, British Columbia. 1 page. Douglas, G. W., and S. J. Smith. 2003. Stewardship account for foxtail muhly (Muhlenbergia andina). Unpublished report submitted to Ministry of Water, Land and Air Pro- tection, Biodiversity Branch. 17 pages. Douglas, G. W., and S. J. Smith. 2003. Stewardship account for pale bulrush (Scirpus pallidus). Unpublished report submitted to Ministry of Water, Land and Air Protection, Biodiversity Branch. 18 pages. Douglas, G. W., and S. J. Smith. 2003. Stewardship account for Southern Maidenhair-fern (Adiantum capillus-vener- is). Unpublished report submitted to Ministry of Water, Land and Air Protection, Biodiversity Branch. 18 pages. Brownell, V. R., and G. W. Douglas. 2004. Draft National Recovery Strategy for Toothcup, Rotala ramosior (L.) Koehne. Unpublished report submitted to Recovery of Na- tionally Endangered Wildlife (RENEW), Ottawa, Ontario. 39 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Douglas, G. W. 2004. A Survey of the Rare Native Vascular Plants of the Southern Okanagan Grasslands Protected Area. Unpublished report submitted to Parks Canada, Pacific and Yukon Region, Vancouver, British Columbia. 7 pages. Douglas, G. W. 2004. COSEWIC status report on the pink sand-verbena, Abronia unbellata in Canada. In COSEWIC assessment and status report on the pink sand-verbena, Abronia umbellata, in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 19 pages. Douglas, G. W. 2004. Draft National Recovery Strategy for Lemmon’s Holly Fern (Polystichum lemmonii). Unpub- lished report submitted to British Columbia Ministry of Water, Land, and Air Protection, Biodiversity Branch, Vic- toria, British Columbia. 15 pages. Douglas, G. W., and S. J. Smith. 2004. Draft COSEWIC Status Report on California Hedge-parsley, Yabea micro- carpa (Apiaceae) in Canada. Unpublished report submit- ted to British Columbia Ministry of Water, Lands and Air Protection, Biodiversity Branch, Victoria, British Colum- bia. 21 pages. Douglas, G. W., and S. J. Smith. 2004. Draft COSEWIC Status Report on Lindley’s Microseris, Microseris lind- leyi (Asteraceae), in Canada. Unpublished report submit- ted to British Columbia Ministry of Water, Land, and Air Protection, Biodiversity Branch, Victoria, British Colum- bia. 24 pages. Douglas, G. W., and S. J. Smith. 2004. Draft COSEWIC Status Report on Slender Popcorn Flower, Plagiobothrys tenellus (Boraginaceae) in Canada. Unpublished report submitted to British Columbia Ministry of Water, Land, and Air Protection, Biodiversity Branch, Victoria, British Columbia. 22 pages. Douglas, G. W., and S. J. Smith. 2004. Draft Update COSEWIC Status Report on Scouler’s Corydalis, Cory- dalis scouleri (Fumariaceae), in Canada. Unpublished report submitted to British Columbia Ministry of Water, Land, and Air Protection, Biodiversity Branch, Victoria, British Columbia. 18 pages. Douglas, G. W., and S. J. Smith. 2004. Draft National Multi- species Recovery Strategy for plants at risk in Garry Oak woodlands. Unpublished report submitted to British Col- umbia Ministry of Water, Land, and Air Protection, Bio- diversity Branch, Victoria, British Columbia. 10 pages. Douglas, G. W., and S. J. Smith. 2004. Draft National Recovery Strategy for Scarlet Ammannia (Ammannia robusta Heer & Regel). Unpublished report submitted to British Columbia Ministry of Water, Land, and Air Pro- tection, Biodiversity Branch, Victoria, British Columbia. 23 pages. Douglas, G. W., and S. J. Smith. 2004. Rare Native Vascu- lar Plants of the Southern Okanagan Grasslands Protect- ed Area Field Report. Unpublished report submitted to The Nature Trust of British Columbia, North Vancouver, British Columbia. 10 pages. Harris, A. G., R. F. Foster, G. W. Douglas, and S. J. Smith. 2004. National Recovery Strategy for Small-Flowered Lipocarpha (Lipocarpha micrantha) in Canada. Unpub- lished report submitted to the Ontario Ministry of Natural Resources, Northwest Region, Thunder Bay, Ontario, and British Columbia Ministry of Water, Land, and Air Pro- tection, Biodiversity Branch, Victoria, British Columbia. 25 pages. *COSEWIC. 2005a. COSEWIC assessment and status report on the Branched Phacelia Phacelia ramosissima in Can- ada. Committee on the Status of Endangered Wildlife in - 2006 Canada, Ottawa, Ontario. vi + 18 pages. (Available online at: http://www.sararegistry.gc.ca/status/status_e.cfm). *COSEWIC. 2005b. COSEWIC assessment and status report on the Cliff Paintbrush Castilleja rupicola in Canada. Committee on the Status of Endangered Wildlife in Cana- da, Ottawa, Ontario. vii + 18 pages. (Available online at: http://www.sararegistry.gc.ca/status/status_e.cfm). *COSEWIC. 2005c. COSEWIC assessment and status report on the Grand Coulee Owl-clover Orthocarpus barbatus in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. vi + 17 pages. (Available on- line at: http://www.sararegistry.gc.ca/status/status_e.cfm). “COSEWIC. 2005d. COSEWIC Assessment and Status report on the Mountain Holly Fern, Polystichum scopuli- num (Dryopteridaceae), in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. vii + 18 pages (Available online at: http://www.sarareg istry.gc.ca/status/status_e.cfm). “COSEWIC. 2005e. COSEWIC assessment and status report on the Spalding’s Campion, Silene spaldingii, in Canada. Committee on the Status of Endangered Wildlife in Cana- da, Ottawa, Ontario. vi + 18 pages. (Available online at: http://www.sararegistry.gc.ca/status/status_e.cfm). Smith, S., and G. W. Douglas. 2005. Cowichan Garry Oak Preserve Monitoring Report: Assessing changes to plant community composition 1999-2004. Douglas Ecological Consultants Ltd. Unpublished report submitted to the Nature Conservancy of Canada, Victoria, British Colum- bia. 22 pages + appendices. Professional Experience Summary 1966 — 1967: Forester and Consultant with Rayonier, Cana- da Ltd., Jeune Landing, BC and Mills, and Potter and Mills, Vancouver, British Columbia, where he supervised forest inventory, forest type mapping, and reforestation projects on northern Vancouver Island, British Columbia. Ecological Consultant to the Intercampus Edu- cation and Scientific Preserves Committee (Washington Universities) at Western Wash- ington State College, Bellingham, Washington, hired to examine and quantitatively sample more than twenty-five potential natural areas throughout the State of Washington, and rec- ommend which ones should be protected. Ecological Consultant to the United States National Park Service in Sedro Woolley, Wash- ington, where he initially provided a biological inventory (vegetation, wildlife, soils and geol- ogy) of the newly formed North Cascades National Park, Washington. He also conducted vegetation analysis of the entire park (18000 km?) and provided advice and research with respect to vegetation problems. Ecological Consultant to Canadian Wildlife Service, Edmonton, Alberta, and Parks Canada, Winnipeg, Manitoba. Project co-ordinator and chief ecologist examining vegetation and soils of Kluane National Park, Yukon Territory, and their relation to fragility and land use within the park. This study was also co-ordinated with small mammal and bird studies. 1968 — 1970: 1969 — 1974: 1973 — 1974: PENNY: TRIBUTE TO GEORGE WAYNE 1973 — 1974: 1974 — 1975: 1974 — 1975: 1974 — 1975: 1975 — 1977: 1975 — 2004: 1975 — 1979: 1977 — 1978: DOUGLAS 145 Ecological Consultant to Canadian Forestry Ser- vice, Edmonton, Alberta, reviewing and evalu ating the ecological impact and use of chem ical fire retardants in North America. He also made recommendations pertaining to the fu- ture use of chemical fire retardants in Canada. Ecological Consultant to Canadian Wildlife Service, Edmonton, Alberta, and Parks Cana- da, Calgary, Alberta. Project co-ordinator and chief plant ecologist for a study designed to assess the environmental impact of recreation- al use (mainly human and horse trampling) on different vegetation types in Waterton Lakes National Park, Alberta. Ecological Secretariat, Department of Lands and Forests, Victoria, British Columbia. Chief plant ecologist on a three-faceted study in west-central British Columbia. The study in- cluded a vegetation reconnaissance of a 1500 square mile area, examination of the spruce hybridization problem in the region, and an evaluation of the ecological reserve system and selection in British Columbia. Ecological Consultant with Don Blood & Associates Ltd., Lantzville, British Columbia, for Parks Canada, Winnipeg, Manitoba. Chief plant ecologist with a biological and geolog- ical team assessing the environmental impact of five potential transportation corridors in Kluane National Park, Yukon Territory. Ecological Consultant to Syncrude Canada Ltd., Edmonton, Alberta. This project involved setting up a continuous biological monitoring program for air quality in the tar sands refin- ery area near Fort McMurray, Alberta. Using lichens as an indicator, the affect of effluents (especially SO,) and their environmental im- pacts will be continuously evaluated. Research Associate in Biological Collections, Herbarium, Royal British Columbia Museum, Victoria, British Columbia. Author of a flora on the Asteraceae (Sunflower family) of Bri- tish Columbia. This work includes a complete taxonomical and ecological treatment of ap- proximately 350 taxa. Ecological Consultant to Parks Canada, Win- nipeg, Manitoba. Project co-ordinator and chief plant ecologist on four-year Biophysical Inventory Project in Kluane National Park, Yukon Territory. This multidisciplinary study included landforms, surficial deposits, vegeta- tion, and soils. A total of ten professional sci- entists were involved in this study. Ecological Consultant to Thurber Consultants Ltd., Victoria, British Columbia, with Depart- ment of Public Works Canada, Vancouver, British Columbia. Chief plant ecologist on environmental impact study team. This multi- disciplinary study included all aspects from engineering and social to environmental im- pacts with respect to the paving and relocation of a large section of the Alaska Highway. “In the production note, COSEWIC acknowledges the late George W. Douglas and Ms. Shyanne J. Smith for writing the status report. “In the production note, COSEWIC acknowledges the late George W. Douglas and Jacques Labreque for writing the status report. 146 1978 — 1979: 1978 — 1980: 1979 — 1980: 1980: 1981 — 1986: 1985: 1985 — 1986: 1986 — 1995: 1987: THE CANADIAN FIELD-NATURALIST Ecological Consultant to Department of Pub- lic Works, Canada — Whitehorse, Yukon Terri- tory. Chief plant ecologist on environmental impact study. Assessed the environmental im- pact of Alaska Highway relocation and paving between Watson Lake and Haines Junction. Ecological Consultant with Hardy and Asso- ciates Ltd. for Department of Public Works, Canada in Whitehorse, Yukon Territory. Chief plant ecologist on revegetation project. This multidisciplinary study examined vegetation and soils and recommended reclamation pro- cedures for a 175 mile segment of the Alaska Highway. Ecological Consultant to British Columbia Ministry of Environment, Land and Parks, Victoria, British Columbia. Designed and co- ordinated a biological monitoring network (using lichens) in northeastern British Colum- bia. Ecological Consultant to Ecological Reserves Unit, British Columbia Ministry of Environ- ment, Victoria, British Columbia. Provided an assessment of alpine plant communities in British Columbia and adjacent regions and their occurrence in existing or proposed eco- logical reserves. Taxonomical Consultant to British Columbia Ministry of Forests, Victoria, British Colum- bia. Provided editorial assistance for floristic manuals and prepared several reports, includ- ing a floristic bibliography. Identified and annotated several thousand sheets of British Columbia plant material. Taxonomical Consultant to National Museums of Canada, Ottawa, and World Wildlife Fund. Prepared detailed status reports on four En- dangered vascular plants of British Columbia. Interpretive Consultant to British Columbia Provincial Parks, Liard Hotsprings, and Mt. Robson Provincial Parks. Provided interpre- tive services for Provincial Parks summer interpretive programs. Taxonomical Consultant to British Columbia Ministry of Forests, Victoria, British Colum- bia. Chief editor and major contributor for the four-volume Vascular Plants of British Colum- bia. Ecological and Taxonomical Consultant for British Columbia Provincial Parks, Strathcona Provincial Park, with Donald A. Blood & Accepted 11 November 2006 1988 — 1989: 1991 — 2002: 1992: 2001 — 2004: 2002: 2002: 2003: 2003: 2003: 2003 — 2004: 2004: 1974 — 2004: Vol. 120 Associates, Nanaimo, British Columbia. Pro- vided ecological and floristic expertise with respect to a preliminary assessment of natu- ral resources. Ecological and Taxonomical Consultant to British Columbia Ministry of Parks, Ecological Reserves Unit, Victoria, British Columbia. Chief co-ordinator for study team (including Blood & Associates and Westland Resources Group) formulating a new ecological reserve system for British Columbia. Program botanist at British Columbia Con- servation Data Centre, British Columbia Min- istry of Environment, Lands and Parks, Vic- toria, British Columbia. Responsible for the tracking and inventorying vascular plants of conservation concern in British Columbia. Botanist, Sierra Club expedition down the Tat- shenshini River, northern British Columbia; part of the campaign to have a protected area established in the region. Recovery team member and scientific advisor, Garry Oak Ecosystem Recovery Team Plants at Risk Recovery Implementation Group. Recovery team member and scientific advisor, Adiantum capillus-veneris Recovery Team. Ecological Consultant to Parks Canada, Pacif- ic Rim National Park. Chief botanist involved in inventory and invasive species management. Botanical consultant to British Columbia Min- istry of Environment. Chief botanist on inven- tory projects in South Okanagan Grasslands Protected Area and adjacent parcels. Botanical Consultant to British Columbia Min- istry of Environment. Chief Botanist hired to inventory for Silene spaldingii to fill informa- tion gaps in its current status in the province. Botanical Consultant to Parks Canada. One of chief botanists hired to inventory rare plants on the Queen Charlotte Islands. Botanical Consultant to Osoyoos Indian Band. Botanist and Project Coordinator managing invasive species in sensitive habitats on band lands. Also trained band members in manag- ing the vegetation. Botanical Consultant to British Columbia Ministry of Environment. Chief botanist hired to inventory Azolla mexicana to fill informa- tion gaps in its current status in the province. Principal of Douglas Ecological Consultants Ltd. Botanist, Ecologist and Project Director. Conservation Evaluation of the Prairie Lupine, Lupinus lepidus var. lepidus, in Canada* GEORGE W. DouGLas! and MICHAEL RYAN2 ' Deceased Present address: 801 Frayne Road, RR # 1, Mill Bay, British Columbia VOR 1LO Canada Douglas, George W., and Michael Ryan. 2006. Conservation evaluation of the Prairie Lupine, Lupinus lepidus var. lepidus, in Canada. Canadian Field-Naturalist 120(2); 147-152. In Canada, Prairie Lupine, Lupinus lepidus var. lepidus, is restricted to southeastern Vancouver Island. Of the nine sites where it has been collected, five are extirpated and the status of two of the populations is uncertain. There are two extant popula- tions; some of the other sites may contain the species in the seed bank. Some of the sites are protected to a certain extent from direct habitat destruction by their remote location, although introduced herbaceous species may pose a serious threat by preventing the establishment of the species at other sites. Fire suppression or the lack of other types of disturbance also likely plays a role in discouraging emergence of Lupinus lepidus. Key Words: Prairie Lupine, Lupinus lepidus, endangered, distribution, population size, British Columbia. The Prairie Lupine, Lupinus lepidus Dougl. ex Lindl. var. lepidus* is a member of a genus of over 100 spe- cies. Most of these occur in North America but the - genus occurs on all continents except Australia (Hitch- cock et al. 1961). It is one of 20 species of Lupinus _ occurring in British Columbia (Pojar 1999) and of about 27 species in Canada (Scoggan 1978). The genus is of some economic importance in the horticultural trade. The Lupinus lepidus complex has been a source of difficulty to researchers over the years. Hitchcock et al. (1961) state that although the L. /epidus complex is distinct, the variation within this group has resulted in a large number of proposed subspecies and varieties _ of which some are considered to be distinct species by others. According to Phillips (1955), the nomenclature | is even more confusing, and the number of synonyms _ is greater for L. /epidus than for any other Lupinus spe- | cies. The work of Detling (1951), Phillips (1955) and Dunn and Gillett (1966) has improved the nomenclature and delineation of Lupinus species but many problems remain. The type specimen of L. lepidus var. lepidus was collected by David Douglas from “... Fort Vancouver ' to the Great Falls of the Columbia”. Subsequently, Hitchcock et al. (1961) recognized five varieties of L. | lepidus two of which, var. lepidus and var. lobbii (A. \ Gray) C. L. Hitche., are reported to occur in British Columbia. The latter variety is considered to be a sep- arate species (L. lyallii A. Gray) in more recent floras of Canada and British Columbia (eg., Dunn and Gillett 1966; Taylor 1974; Scoggan 1978; Douglas 1990; Pojar 1999) and occurs on well-drained soils in subalpine and alpine areas (Douglas 1990; Douglas and Bliss 1977). Lupinus lepidus var. lepidus as recognized by Hitch- cock et al. (1961), Douglas (1990), Douglas et al. (1998a) and Pojar (1999), included L. minimus Doug]. in synonomy but they failed to account for the loca- tion of the type specimen (Rocky Mountains, Kettle Falls in the upper Columbia River drainage) or the range of L. minimus in the western Cordillera east of the Coast-Cascade Mountains. Lupinus minimus is con- sidered by other authors to be a separate species (e.g., Cox 1973a, 1973b; Dunn and Gillett 1966; Scoggan 1978; Taylor 1974). Douglas et al. (2002a, 2002b) also now treat L. minimus as a distinct entity. The lat- ter has been reported to occur in the Columbia River drainage of southern British Columbia (Dunn and Gillett 1966; Taylor 1974), but no specimens have been located. It is also reported to occur in the Crowsnest area of Alberta (Dunn and Gillett 1966) and, more recently, in and adjacent to Waterton Lakes National Park (Argus and White 1978; Kuijt 1982) and at six locations near the southern Alberta border (Packer 1983). Both Kuijt (1982) and Packer (1983) consider L. minimus to be synonymous with L. lepidus. Lupinus lepidus is a multi-stemmed perennial herb 20-45 cm tall (Figure 1; Pojar 1999). The stems are slender with long, soft hairs. The plant has oblanceo- late palmate compound leaves that are 5 to 9 foliate and copiously silky-hairy. The leaves are mostly basal with only | to 4 leaves on the stem. The petioles are 2-5 times as long as the blade. The racemose, pea-like “The original field work for the Lupinus lepidus project was funded by the British Columbia Conservation Data Centre. The results appear in the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a Committee on the Status of Endangered Wildlife in Canada status report (Ryan and Douglas 1996*) and the subsequent assessment of | endangered (COSEWIC 2000’). This paper includes more recent information that did not appear in the original status report. ‘Taxonomy and nomenclature follow Douglas et al. (1998b, 1999, 200 1a, b) and Pojar (1999). ————————— } | 147 148 THE CANADIAN FIELD-NATURALIST blue flowers are similar to those of other Fabaceae species. The petals are 8-13 mm long, and the banner is often lighter or darker and reflexed below the mid- point. The wings are glabrous, the keel ciliate, and the calyx silky-hairy. Both lips are 6-7 mm long, the upper ones of which are bifid for at least half their length. Peduncles are 9-10 cm long, pedicels are 2 mm long and petals are 8-13 mm long. The hairy pods are 1-2 cm long and contain three to five large seeds. In British Columbia, it is separated from three similar looking species by its distinct range in the coastal lowlands on southeastern Vancouver Island. It is distinguished from most other species of Lupinus by its small stature, the white appressed hairs covering the stems and leaves and the thick caudex at the base of the plant. Distribution Lupinus lepidus extends from southwestern British Columbia to north-western Oregon, west of the Cas- cade Mountains (Douglas et al. 1990; Pojar 1999). In Canada it is rare on southeastern Vancouver Island, British Columbia (Figure 2; Douglas et al. 2002a; Pojar IO). Habitat Lupinus lepidus inhabits xeric sites ranging from grass-dominated meadows to steep rocky slopes where the vegetation is comprised of scattered clumps of Douglas-fir (Pseudotsuga menziesii), Arbutus (Arbu- tus menziesii), and Lodgepole Pine (Pinus contorta var. contorta). These sites are restricted to southeastern Vancouver Island, where rainfall is low compared to other coastal areas in British Columbia, and during the summer, they often experience an extended period of drought. Associated species include Scotch Broom (Cytisus scoparius), Hedgehog Dogtail (Cynosurus echinatus), Sweet Vernalgrass (Anthoxanthum odora- tum), several species of Brome (Bromus), and Common Camas (Camassia quamash). The habitat at the Some- nos site was unnatural, consisting of gravel deposited during railroad or highway construction. TABLE |. Populations of Lupinus lepidus in Canada. Collection Last Site Observation Langford Plains (Victoria) 1908 Observatory Hill (Victoria) 1915 Koksilah River Valley (Duncan) 1960 Cattle Point (Victoria) 1973 Beacon Hill (Victoria) 1993 Somenos Lake (Duncan) 1994 Mount Wells (Victoria) 2006 (2 plants/1 m? in 2003) Mount Braden (Victoria) 2005 Mount McDonald (Victoria) 2005 Vol. 120 Ficure 1. Illustration of Lupinus lepidus. (Line drawing from Pojar 1999) Collector/ Population data Observer (plants/area in m7?) Macoun Extirpated Newcombe Extirpated Hardy Unknown Brayshaw Extirpated Brayshaw Extirpated Douglas Extirpated Roemer No plants Roemer 2 plants Roemer 113 plants/75 m? 2006 Biology Little is known regarding the biology of Lupinus lepidus. However, many species of Lupinus share com- mon traits, some of which likely characterize L. lep- idus. It is likely to be associated with nitrogen-fixing Rhizobium, which may allow it to invade sites where soils are low in nitrogen. It is suspected that L. lep- idus is a short-lived colonizer of recently disturbed sites in Xeric, open, exposed areas where it persists for sev- eral years. It eventually declines, possibly from compe- tition with more aggressive species that either directly compete for resources or prevent the establishment of seedlings, or from lack of vigour in mature plants. The seeds may not have any dormancy requirements but like other legumes, L. /epidus seeds have a hard seed coat which may delay germination for several months or years. Nothing is known regarding pollination, seed set, and survival rates. Population Attributes Of the nine Canadian sites from which Lupinus lep- idus has been collected (Table 1, Figure 2), five pop- ulations are extirpated. Another two sites (Koksilah River and Mount Wells) are assumed extant but their status is unknown. Little is known regarding popula- tion trends for this species. Monitoring this taxon is difficult because populations may only be comprised of a few plants and therefore, easily overlooked, or they may remain in a seed bank for an unknown period before reappearing. However, four years of data col- lected at Somenos Lake (extirpated) showed a decline before it was destroyed at the site: 250, 1,0, and | in 1991, 1992, 1993 and 1994, respectively. Other sites with limited data include Mount Wells, which con- tained nine plants in 2001, only a few plants in 2002, two plants in 2003, and no plants in 2004, 2005 and 2006 (H. Roemer, personal communication). The steady decline at this site is attributed to the re-growth of shrubs following a fire. In addition, in 2005, two plants were observed on Mount Braden (up from sin- gle plant in 1996) and 113 plants were observed on Mount McDonald (where they had only previous been known from 1915). In 2006, a single plant was seen on Mount Braden and 54 plants on Mount McDonald, but a thorough search of all the locations from 2005 was not done and therefore, the population should not be considered in decline. Provincial, National, and Global Ranks The British Columbia Conservation Data Centre has ranked Lupinus lepidus as S1 and placed it on the British Columbia Ministry of Environment Red-list (Douglas et al. 2002a). This is the most critical cate- gory for imperiled rare native vascular plants in British Columbia. A rank of S1 is considered “critically imper- iled because of extreme rarity (5 or fewer occurrences or very few remaining individuals) or because of some factors making it especially vulnerable to extirpation or DOUGLAS AND RYAN: PRAIRIE LUPINE 149 FIGURE 2. The status and location of Lupinus lepidus in British Columbia. @ recently confirmed; ® present status unknown: O extirpated extinction” (Douglas et al. 2002a). Lupinus lepidus is on the Canadian Species At Risk Act legal list and the Committee on the Status of Endangered Wildlife in Canada has also assigned this species to the Endangered category (COSEWIC 2000’). Globally, it is ranked G5 or common and secure in its range (NatureServe 2006°). Threats and Protection The most direct threat to Lupinus lepidus is habitat destruction. Grass-dominated meadows, often associ- 150 ated with Garry Oak-Brome (Quercus garryana- Bromus) stands, commonly occurred on gentle slopes on the most climatically favourable coastal areas on south-eastern Vancouver Island and some of the Gulf Islands prior to European colonization. Since coloniza- tion both types of vegetation have been subjected to ex- tensive agricultural and residential development. Their destruction has continued to the present, resulting in the elimination of almost all sites occurring outside parks or ecological reserves. The loss of these habi- tats, estimated at 95% (Lea 2002), severely reduces the availability of sites for the establishment of L. lepidus var. lepidus and imposes severe limitations on the long- term survival of this species in Canada. Introduced species including Cytisus scoparius, Cynosurus echinatus and Anthoxanthum odoratum now dominate L. /epidus habitats, further threatening the survival of this species. The introduction of aggressive European species has resulted in substantial changes to the grass-dominated meadows associated with Quer- cus garryana, and rocky xeric sites in the Victoria area. Cytisus scoparius, in particular, has been one of the most devastating species. It has become a dominant shrub on xeric, exposed sites throughout much of south- eastern Vancouver Island and the Gulf Islands. Furthermore, the suppression of natural and unnat- ural periodic fires appears to have resulted in changes to the vegetation of many sites where this species would be expected to occur. In the past, aboriginal peoples may have set fire to these sites to maintain them as an important habitat for wildlife (Roemer 1972). This would have destroyed much of the competing vegeta- tion and created habitats where L. lepidus would have had the opportunity to become established. However, since that time, these sites have experienced little dis- turbance, resulting in invasion and expansion by trees, shrubs, and especially introductions, which effectively eliminate many herbaceous species. In Canada, some populations of L. /epidus are pro- tected to a certain extent by their location on public property. This was not the case, however, for the plant in Beacon Hill Park which was destroyed by mainte- nance activites along a roadbank in 1994, or for the population on the highway right-of-way at Duncan, which was also destroyed and covered in 1994 by heavy equipment used in a North Cowichan Munici- pality sewer line project. Extirpation is also a probable fate at Cattle Point and Observatory Hill. Even though Cattle Point is located in a municipal park, it is not protected from trampling by pedestrians and mountain bike enthusiasts. During the summer months, tour buses arrive at this park many times each day resulting in large numbers of people trampling the vegetation along the shoreline and adjacent rock outcrops. As a result of these activities, much of the vegetation has been degraded to the point where the number of potential- ly suitable habitats for the establishment of L. lepidus are far fewer today than in the past. The population THE CANADIAN FIELD-NATURALIST Vol. 120 could also have been overcome if it occurred in areas now dominated by shrubs. Observatory Hill is owned by the federal government and is home to the Dominion Astrophysical Observa- tory. Much of the vegetation has been replaced by exotic species but access is limited to a single road. Although further development on the hill is unlikely to occur, many of the suitable habitats in which L. lep- idus would likely be found are infested with Cytisus scoparius . Populations at Mount McDonald and Mount Braden are protected by their location within either the Greater Victoria Watershed District (which supplies drinking water to Victoria and adjacent communities and pro- hibits public access) or the CRD Sooke Hills Wilder- ness, respectively. Since these areas are dominated by steep rocky terrain and since there are few trails, veg- etation has not been degraded to the same extent as it has in more accessible areas near Victoria. The specific site at which L. lepidus was collected in the Koksilah River Valley is not known, thus it is not possible to provide any information on the extent to which this population is protected. However, if it is located on a steep, rocky, xeric sites similar to those found on Mount McDonald, Mount Braden, and Mount Wall, it is unlikely to be threatened in the near future by development or habitat destruction. The site on Mount Wells is on private land but is not easily accessed at the present time. Nearby mountains, however, have been subject to housing developments in recent years thus the site of L. lepidus is not neces- sarily secure. Lupinus lepidus is not formally protected in British Columbia, however, it could be in the future since this species is a potential candidate for listing under the provincial Wildlife Amendment Act (2004). As part of its commitment to the National Accord (National Accord for the Protection of Species at Risk), the province is required to take measures to protect this species. It is on Schedule 1 of the federal Species At Risk Act and a recovery strategy (which treats Lupi- nus lepidus in addition to several other species-at-risk) has been proposed (Parks Canada Agency 2005"). The recovery strategy is a planning document that identi- fies what needs to be done to arrest or reverse the de- cline of a species. Detailed planning is done at the action plan stage. However, an action plan is not cur- rently available, nor are there any stewardship activi- ties that are currently being implemented. Evaluation The Committee on the Status of Endangered Wildlife in Canada and the British Columbia Conservation Data Centre consider Lupinus lepidus to be endangered in Canada (Douglas et al. 2002a). It has only been col- lected nine sites in Canada, only three of which have recently been confirmed, all confined to southeastern Vancouver Island. It is believed to be extirpated at five 2006 of these sites and has not been confirmed recently at two sites. The prognosis for this species is not good considering the threats posed by aggressive competi- tive species. The loss of suitable habitats at other sites to residential development severely limits the potential of this species to become established at new sites. Fur- thermore, fire, which may have been an important agent in the continued existence of this species in British Columbia, is now actively suppressed. With so little information known on the ecology of L. lepidus, and on its relationship to the environment and other species, successful management of the species is hampered. Acknowledgments Special thanks to Hans Roemer, whose persistent searches during recent years continue to reveal Lupi- nus lepidus sites. Thanks also to Jenifer Penny for office assistance. Documents Cited (marked * in text) COSEWIC. 2000. Canadian Species at Risk. Committee on the Status of Endangered Wildlife in Canada. www. cosewic.gc.ca, Ottawa, Ontario. Available Online at: http:// Www.sararegistry.gc.ca/species/speciesDetails_e.cfm?sid =187. NatureServe Explorer. 2006. NatureServe Explorer: An online encyclopedia of life. Version 1.7. Arlington, Vir- ginia, USA. Web site: http://www.natureserve.org/explorer/ [Accessed 22 March 2006]. Parks Canada Agency. 2005. Recovery Strategy for Multi- species at Risk in Maritime Meadows Associated with Garry Oak Ecosystems in Canada (proposed). Jn Species at Risk Act Recovery Strategy Series. Edited by Parks Canada Agency. Ottawa, Ontario. 98 pages. Ryan, M., and G. W. Douglas. 1996. Status Report on the Prairie Lupine, Lupinus lepidus var. lepidus, in Canada. Committee on the Status of Endangered Wildlife in Canada. 25 pages. Literature Cited Argus, G. W., and D. J. White. 1978. The rare vascular plants of Alberta. National Museums Canada, Syllogeus 17, Ottawa, Ontario. Cox, B. J. 1973a. A chemosystematic comparison of the Lupi- nus lepidus-L. caespitosus complex. Bulletin of the Torrey Botanical Club 100: 12-17. Cox, B. J. 1973b. Protein relationships among the perennial caespitose lupines. Bulletin of the Torrey Botanical Club 100: 153-158. Detling, L. E. 1951. The caespitose lupines of western North America. American Midland Naturalist 45: 474-499. Douglas, G. W. 1990. Fabaceae. Pages 21-45 in The Vascular Plants of British Columbia. Part 2 — Dicotyledons (Dia- pensiaceae through Portulacaceae). Edited by G. W. Dou- glas, G. B. Straley, and D. Meidinger. British Columbia Ministry of Forests, Victoria, British Columbia. 158 pages. Douglas, G. W., and L. C. Bliss. 1977. Alpine and subalpine plant communities of the North Cascades Range, Washing- ton and British Columbia. Ecological Monographs 47: 1 13- 150. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare Native Vascular Plants of British Columbia. Second edi- DOUGLAS AND RYAN: PRAIRIE LUPINE 151 tion. British Columbia Ministry of Sustainable Resource Management, Victoria, British Columbia. 359 pages Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Ulustrated Flora of British Columbia. Volume 3. Dicotyledons (Dia- pensiaceae through Onagraceae). British Columbia Min- istry of Environment, Lands and Parks and British Colum- bia Ministry of Forests, Victoria, British Columbia. 423 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 200/a. Ilus- trated flora of British Columbia. Volume 6. Monocotyle- dons (Acoraceae through Najadaceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 361 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2001b. Ilustrat- ed flora of British Columbia. Volume 7. Monocotyledons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas G. W., D. Meidinger, and J. Pojar. 2002b. Illus- trated Flora of British Columbia. Volume 8 — General Summary, Maps and Keys. British Columbia Ministry of Sustainable Resource Management, British Columbia Min- istry of Forests, Victoria, British Columbia. 457 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1990. The Vascular Plants of British Columbia. Part 2 — Dico- tyledons (Diapengiaceae through Portulaceae). Special Report Series Number 2, British Columbia Ministry of Forests, Victoria, British Columbia. 155 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. Rare Native Vascular Plants of British Columbia. British Columbia Ministry of Environment, Lands and Parks, Vic- toria, British Columbia. 423 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998b. Illustrated flora of British Columbia. Volume 1. Gymnosperms and Dicotyledons. (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Dunn, D. B., and J. M. Gillett. 1966. The lupines of Canada and Alaska. Monograph Number 2. Canada Department of Agriculture, Ottawa, Ontario. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1961. Vascular plants of the Pacific Northwest. Part 3: Saxifragaceae to Ericaceae. University of Washing- ton Press, Seattle, Washington. 614 pages. Kuijt, J. 1982. A flora of Waterton Lakes National Park. The University of Alberta Press, Edmonton, Alberta. 684 pages. Lea, Ted. 2002. Historical Garry oak ecosystems of Greater Victoria and Saanich Peninsula. 1:20,000 Map. Terrestrial Information Branch, British Columbia Ministry of Sustain- able Resource Management. Victoria, British Columbia. Available online at: http://www.goert.ca/maps/go-020202. pdf. Packer, J. G. 1983. Flora of Alberta. Second edition. Univer- sity of Toronto Press, Toronto, Ontario. 687 pages. Phillips, L. L. 1955. A revision of the perennial species of Lupinus of North America exclusive of southwestern United States and Mexico. Research Studies of the State College of Washington 23: 161-201. Pojar, J. 1999. Fabaceae. Pages 64-180 in Illustrated Flora of British Columbia. Volume 3 — Dicotyledons (Diapen- siaceae through Onagraceae). Edited by G. W. Douglas. D. Meidinger, and J. Pojar. British Columbia Ministry or Sy THE CANADIAN FIELD-NATURALIST Vol. 120 Environment, Lands and Parks, British Columbia Ministry Taylor, T. M. C. 1974. The Pea Family (Leguminosae) of of Forests, Victoria, British Columbia. 423 pages. British Columbia. Handbook Number 12. British Colum- Roemer, H. L. 1972. Forest vegetation and environments of bia Provincial Museum, Victoria, British Columbia. 251 the Saanich Peninsula, Vancouver Island. Ph.D. thesis, Uni- pages. versity of Victoria, Victoria, British Columbia. 405 pages. Scoggan, H. J. 1978. The flora of Canada. Part 3. National Museum of Natural Sciences Publications in Botany. Num- _— Received 29 April 2002 ber 7. National Museum of Natural Sciences, Ottawa, Accepted 24 July 2006 Ontario 568 pages. Conservation Evaluation of the Seaside Birds-foot Trefoil. Lotus . . . * formosissimus, in Canada GEORGE W. DouGLas! and MICHAEL RYAN? 'Deceased. Present address: 801 Frayne Road, RR #1, Mill Bay, British Columbia VOR 1LO Canada. Douglas, George W., and Michael Ryan. 2006. Status of the Seaside Birds-foot Trefoil, Lotus formosissimus, in Canada Canadian Field-Naturalist 120(2): 153-156. - In Canada, Seaside Birds-foot Trefoil, Lotus formosissimus, is restricted to the Victoria area on southeastern Vancouver Island and two adjacent islands. Populations at five sites are extant and have been recently inventoried, whereas populations at two or more sites in Victoria are extirpated. Existing populations represent the northern range limits of L. formosissimus. Although known populations are protected to a certain extent from direct habitat destruction, introduced herbaceous species may pose | a serious threat to the continued existence of most populations and may prevent the establishment of L. formosissimus at other sites. Fire suppression, grazing, encroachment and low survivorship are also limiting factors. Key Words: Seaside Birds-foot Trefoil, Lotus formosissimus, endangered, distribution, population size, British Columbia. Seaside Birds-foot Trefoil, Lotus formosissimus Greene, is a member of a genus of about 200 species occurring throughout the world, primarily in temper- ate zones. It is one of nine species occurring in British - Columbia and Canada, four of which have been intro- duced from the Old World (Pojar 1999; Scoggan 1978). _ Lotus formosissimus is a sprawling or ascending, | multi-stemmed perennial that grows from a short root- | stock and thick taproot to 20-50 cm tall (Figure 1; ' Pojar 1999). It has compound, pinnate leaves 4-8 cm | long with 3 to 7, ovate to obovate leaflets. The three to five yellow and pink pealike flowers are 10-15 mm long and arranged in umbels. The umbels are borne on axil- lary peduncles 2-6.5 cm long and are usually subtend- ed by trifoliate bracts. The wings of the flowers are pinkish to purplish tinged, the keel is purple tipped, and the banner is yellow and 11-15 mm long. The wings and clawed keel exceed the calyx considerably. Pods are 27-36 mm long by 1.5-2 mm wide and en- close 7 to 15, dark brown to black seeds. Lotus for- mosissimus is distinguished from other Lotus species _ by its perennial habit, membranous stipules, trifoliate ' bracts (sometimes unifoliate), and pink and yellow _ flowers (Ryan and Douglas 1994*, 1996*). Bog Bird’s- _ foot Trefoil (L. pinnatus) also has yellow but never pink corollas. Ficure 1. Illustration of Lotus formosissimus. (Line drawing | from Pojar 1999). | Distribution Lotus formosissimus is restricted to the west coast | of North America, from southeastern Vancouver Island (S central California (Isely 1993; Pojar 1999). In Cana- and nearby islands (Figure 2; Dunn and Gillet 1966; da, it is restricted to southeastern Vancouver Island Pojar 1999; Douglas et al. 2002a, 2002b). OF Pe an 1 | ‘The original field work for Lotus formosissimus was funded by the British Columbia Conservation Data Centre. The results appear in the | British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a | Committee on the Status of Endangered Wildlife in Canada status report (Douglas and Ryan 1996*) and the subsequent assessment of Endan- | gered. This paper includes more recent information that did not appear in the original status report. 153 154 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Populations of Lotus formosissimus in Canada (in Victoria and west of Victoria at Department of National Defense — Rocky Point ). Collection Site Last Observation Oak Bay, Victoria 1896 Foul Bay, Victoria 1912 William Head, W of Victoria 2002 Bentinck Island, Rocky Point 2004 Rocky Point, E side 2005 Rocky Point, Church Point 2004 Trial Island, Victoria 2004 Collector Population Status (number/area) Anderson Extirpated Henry Extirpated Fairbarns 19/8 m? Fairbarns 278/110 m? Fairbarns 36-53/75.5 m? Fairbarns 209-327/137 m2 Fairbarns 383/5000 m? Habitat Lotus formosissimus grows in various coastal mead- ow and woodland habitats, ranging from open exposed grass-dominated meadows to exposed steep rocky sites inhabited by Garry Oak (Quercus garryana), to shad- ed Garry Oak-Brome (Quercus garryana-Bromus) forests. Associated species, which typically occur in the understory of Quercus garryana stands, include Lance-leaved Stonecrop (Sedum spathifolium), Sea Blush (Plectritis congesta), Nodding Onion (Allium cernuum), Menzies Larkspur (Delphinium menziesii), Orchard Grass (Dactylis glomerata), Sweet Vernalgrass (Anthoxanthum odoratum), Hairy Cat’s-ear (Hypo- chaeris radicata), Common Velvet-grass (Holcus lana- tus), Early Hairgrass (Aira praecox), Ribwort Plantain (Plantago lanceolata), and several species of brome (Bromus). Lotus formosissimus also occurs in vernal seeps. It is suspected that L. formosissimus is a poor competitor with a number of perennial grasses because it is absent in a number of grass-dominated micro-sites that are otherwise apparently suitable habitats. Biology Little is known regarding the biology of Lotus for- mosissimus. It appears that L. formosissimus shoots emerge in spring followed by flowering between May and the end of June when moist soil conditions and warm temperatures are prevalent in Quercus garryana stands and meadows. By July, when drought conditions are prevalent, seed maturation and dispersal begin, followed soon thereafter by die-back of the stems to the rootstock, which is located several centimetres below the soil surface. In his demographic studies of Lotus formosissimus, M. Fairbarns (personal communication) discovered, based on ex situ experiments, that seeds may remain in the seed bank for at least two years. He postulated that this was a likely explanation for his in situ observations of higher numbers of seedlings than numbers of seeds set in the previous year. Flowers likely require cross-pollination to set viable seed although pollinators have not yet been identified (Zandstra and Grant 1968). Seeds are believed to be easy to germinate (Hitchcock et al. 1961) but like other members of Fabaceae, they may have a hard seed coat, delaying the germination of seeds for several months or years. Mature plants may be long-lived and recruit- ment from germinating seeds is likely a rare event. It is believed that L. formosissimus, like other le- gumes, is associated with nitrogen-fixing Rhizobium that provides the plants with an independent source of nitrogen from that in the soil. Population Attributes Lotus formosissimus has been collected at seven sites in Canada on southern Vancouver Island or adjacent islands (Table 1, Figure 2). Populations at two of these sites are considered extirpated. Other older collections not listed in Table 1 may or may not represent differ- ent locations (e.g., Victoria, and Oak Bay, Victoria). The population on Trial Island has been known since the 1950s and surveyors reported 28 plants in 1994 and 380 plants in 2004. Likewise, the population at Rocky Point has also been observed for over 20 years and was represented by 165 plants in 1993 and 245-380 plants in 2005. The increase, however, is likely attributable to different observers and different sampling methods. It is very unlikely that the populations are actually in- creasing significantly. The remaining sites have not been monitored for any length of time and thus, no infor- mation is available on population trends. Provincial, National and Global Ranks The British Columbia Conservation Data Centre has ranked this species as S1 and placed it on the Ministry of Environment Red list. This is the most critical cat- egory for imperiled rare native vascular plants in the province. A rank of S1 is considered “critically imper- iled because of extreme rarity (five or fewer occurrences or very few remaining individuals) or because of some factors making it especially vulnerable to extirpation or extinction”. Lotus formosissimus is on the Canadian Species At Risk Act legal list (COSEWIC 2000*) and globally, it is G5 or common and secure in its range (NatureServe Explorer 2006'). Threats and Protection In the past, the most direct threat to Lotus formo- sissimus was that of habitat destruction. Grass-domi- nated meadows, often associated with Quercus gar- ryana-Bromus stands, commonly occurred on gentle slopes on the southeastern side of Vancouver Island and some of the Gulf Islands prior to European colo- nization. Since colonization, both types of vegetation have been subjected to extensive agricultural and res- 2006 idential development and have been essentially elimi- nated outside parks or ecological reserves. Direct habi- tat destruction, however, is no longer the most impor- tant threat. It is the loss of suitable sites outside the current occupied area of L. formosissimus that severely limits the potential for this species to spread into new areas and therefore, threatens its long-term survival in Canada. Less direct factors which threaten known popula- tions include the introduction of aggressive European species. These species have resulted in substantial changes to the grass-dominated meadows associated with Quercus garryana and the rocky xeric sites in the Victoria area. One of the most devastating species is Scotch Broom (Cytisus scoparius). This species has become a dominant shrub on xeric, exposed sites throughout much of southeastern Vancouver Island and the Gulf Islands and either competes with native plants such as L. formosissimus for resources or pre- vents the establishment of their seedlings. Furthermore, the suppression of natural and unnat- ural periodic fires may have resulted in changes to the vegetation of many sites where this species would be expected to occur. In the past, aboriginal peoples prob- ably set fire to many of these sites to maintain them as an important habitat for food plants and wildlife (Roe- mer 1972). During the past century these sites have experienced little disturbance resulting in increased domination of some sites by trees and shrubs, especially introductions, that effectively eliminate many herba- ceous species. Another limiting factor is grazing. M. Fairbarns (personal communication) observed that grazing prior to seed set had greatly limited seed production and caused reduced vigour among established plants on Bentinck Island. Finally, demographic constraints also play a role in the viability of populations. Fairbarns (personal com- munication) found low rates of juvenile survivorship and low levels of seed production. He found that they were related primarily to summer drought and grazing. All extant populations of L. formosissimus in British Columbia are protected from heavy recreational pres- sure and disturbance caused by the general public. Al- though these sites contain a large number of introduced species, the latter were not nearly as abundant at Rocky Point, Bentinck Island, and Trial Island as they are at other public sites. It is very likely that restricted public access and lack of disturbance are the primary reasons for the continued existence of L. formosissimus at these sites. The populations at Rocky Point and Bentinck Island are owned by the Department of National Defense. The Department of National Defence is aware of the loca- tion of rare species at Rocky Point and there are no plans to develop those areas in which L. formosissimus occurs. Bentinck Island is used by DND to detonate unused ammunition, but such activities do not coincide with DOUGLAS AND RYAN: SEASIDE BiIRDS-FOOT TREFOIL 155 VANCOUVER ISLAND FIGURE 2. The status and location of Lotus formosissimus sites in British Columbia. @ — recently confirmed sites, O — extirpated site. locations of L. formosissimus on the island. These pop- ulations are also protected by the Canadian Species at Risk Act by their occurrence on land under federal jurisdiction and therefore, are not likely to be directly harmed. The Canadian Forest Service is also involved in a number of projects with the goal of managing ap- propriately for species at risk on federal lands, includ- ing Scotch Broom removal from sites. The Lotus formosissimus population in Trial Island Ecological Reserve is also provided with a high degree of protection. Despite its proximity to Victoria, it has 156 very limited public access because it is only accessi- ble by boat, thus preventing excessive disturbance. A recovery strategy (which treats Lotus formosis- simus in addition to several other species-at-risk) has been proposed (Parks Canada Agency 2005"). The recovery strategy is a planning document that identi- fies what needs to be done to arrest or reverse the decline of a species. Detailed planning is done at the action plan stage. In the absence of an action plan, no attempts have been made to introduce L. formosissimus to suitable habitats or to increase the number of indi- viduals at current locations. However, Scotch Broom removal at the Bentinck Island site has revealed addi- tional Lotus plants (M. Fairbarns, personal communi- cation). The site now offers improved conditions for growth and reproduction. Evaluation of Status The British Columbia Conservation Data Centre con- siders Lotus formosissimus to be endangered in Cana- da (Douglas et al. 2002a). The Committee on the Sta- tus of Endangered Wildlife in Canada has also assigned this species to the Endangered category (COSEWIC 2000"). Between 925 and 1060 plants are known from five sites on southeastern Vancouver Island and adja- cent islands. The long-term survival of this species in Canada is dubious as a result of a number of limiting factors: shading by aggressive introduced species, poor survivorship, grazing, fire suppression and encroach- ment. Even though known populations are protected from excessive disturbance on federal lands and in the ecological reserve, the loss of suitable habitats at other sites severely limits the potential of this species to become established at new sites. Acknowledgments We thank Matt Fairbarns for population updates for all extant locations and Jenifer L. Penny for assistance in the office. Documents Cited (marked * in text) COSEWIC. 2000. COSEWIC assessment and status report on the Seaside birds-foot lotus Lotus formosissimus in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa, Ontario. vi + 18 pages. NatureServe Explorer. 2006. NatureServe Explorer: An online encyclopedia of life. Version 1.7. Arlington, Virgi- nia, USA. Web site: http://www.natureserve.org/explorer [Accessed 22 March 2006]. THE CANADIAN FIELD-NATURALIST Vol. 120 Parks Canada Agency. 2005. Recovery Strategy for Multi- species at Risk in Maritime Meadows Associated with Garry Oak Ecosystems in Canada (proposed) in Species at Risk Act Recovery Strategy Series. Edited by Parks Canada Agency. Ottawa, Ontario. 98 pages. Ryan, M., and G. W. Douglas. 1994. Status report on the Seaside birds-foot lotus, Lotus formosissimus Greene. Un- published report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa. British Colum- bia Conservation Data Centre, Victoria, British Columbia. 25 pages. Ryan, M., and G. W. Douglas. 1996. COSEWIC status report on seaside birds-foot lotus Lotus formosissimus in Canada. Committee on the Status of Endangered Wildlife in Cana- da. Ottawa, Ontario. 1-18 pages. Literature Cited Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare native vascular plants of British Columbia. Second edition. British Columbia Ministry of Sustainable Resource Management, Victoria, British Columbia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2002b. Illus- trated flora of British Columbia. Volume 8. General sum- mary, maps and keys. Ministry of Sustainable Resource Management, Ministry of Forests, Victoria, British Colum- bia. 458 pages. Dunn, D.B., and J. M. Gillett. 1966. The lupines of Canada and Alaska. Monograph number 2. Canada Department of Agriculture, Ottawa, Ontario. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1961. Vascular plants of the Pacific Northwest. Part 3: Saxifragaceae to Ericaceae. University of Washing- ton Press, Seattle, Washington. 614 pages. Isley, D. 1993. Lotus. Pages 616-622 in The Jepson manual: higher plants of California. Edited by J. C. Hickman. Uni- versity of California Press, Los Angeles, California. 1400 pages. Pojar, J. 1999. Fabaceae. Pages 64-180 in Illustrated Flora of British Columbia. Volume 3. Dicotyledons (Diapensi- aceae through Onagraceae). Edited by G. W. Douglas, D. Meidinger, and J. Pojar. British Columbia Ministry of Environment, Lands and Parks, British Columbia Ministry of Forests, Victoria, British Columbia. 423 pages. Roemer, H. L. 1972. Forest vegetation and environments of the Saanich Peninsula, Vancouver Island. Ph.D. thesis, Uni- versity of Victoria, Victoria, British Columbia. 405 pages. Scoggan, H. J. 1978. The flora of Canada. Parts 1-4. Nation- al Museum of Natural Sciences Publications in Botany 7: 1-1626. Zandstra, I. I., and W. F. Grant. 1968. The biosystematics of the genus Lotus (Leguminosae) in Canada. I. Cytotax- onomy. Canadian Journal of Botany 46: 557-583. Received 29 April 2002 Accepted 24 July 2006 Conservation Evaluation of the Bog Bird’s-foot Trefoil, Lotus pinnatus, . * in Canada ~ Marta Donovan! Victoria British Columbia V8W 9R7 Canada. Donovan, Marta. 2006. Conservation evaluation of the Bog Bird’s-foot Trefoil, Lotus pinnatus, in Canada. Canadian Field- ' Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, P.O. Box 9993 St~ Prov Gove. ) Naturalist 120(2): 157-162. In Canada, Bog Bird’s-foot Trefoil, Lotus pinnatus, is confined to vernally wet meadows, creek margins, ditches and seepages, where underground water comes to the surface. Seven localities, within a small area on the east coast of Vancouver Island near Nanaimo, British Columbia, have been confirmed in recent years. At two others it is considered extirpated. The lack of protection on private land, alterations to hydrology, and erosion arising from intensive recreational use of primary habitat by off-road vehicles as well as the appearance of invasive, species are the main concerns for British Columbia populations. Key Words: Bog Bird’s-foot Trefoil, Lotus pinnatus, endangered species, distribution, conservation, British Columbia. Bog Bird’s-foot Trefoil, Lotus pinnatus Hook’, is a low-growing, short-lived perennial of the pea family _ (Fabaceae) with many erect to spreading stems ranging from 15 to 60 cm long (Figure 1). The pinnately com- pound leaves are 4-8 cm long, with 5-9 elliptic, oblong or narrowly egg-shaped leaflets. The inflorescence is | a compact, stalked, axillary umbel of 3 to 12 flowers. The corollas are 10-15 mm long with a yellow ban- ner and keel and creamy-white wings. The keel petals are fused along one edge that is elongated into a well- _ defined beak towards the outside of the inflorescence. The calyx is tubular and lobed, with two upper lobes that are joined most of their length. The linear seed pods are 3-6 cm long and 1.5-2 mm wide and contain 5-20 cylindrical, glossy, dark-coloured seeds (Pojar 1999). In the field, several species superficially resem- ble L. pinnatus. Vetches (Vicia species) are often pres- “ent in similar habitats, but the leaves are smaller than those of L. pinnatus and the terminal leaflet in Vicia spp. _is a tendril. Seaside Birds-foot Trefoil (Lotus formo- sissimus) looks similar to L. pinnatus except the wing petals of L. formosissimus are pinkish-purple, while ‘those of L. pinnatus are cream-colored. In addition, |though a trifoliate bract usually subtends the flowers of L. formosissimus, bracts were sometimes absent in field specimens or only a unifoliate bract was present (Ryan and Douglas 1994*). Lotus pinnatus and L. for- |mosissimus could be confused if the plants are imma- ‘ture or not in flower, but the two species do not over- lap in their distribution in British Columbia. Lotus formosissimus is known only from the Victoria area and nearby islands, where it occurs in various xeric habitats, ranging from open grass-dominated meadows | i FiGure 1. Illustration of Lotus pinnatus (line drawing in Hitchcock et al. 1961; Pojar 1999). to exposed steep rocky sites with Garry Oak (Quercus garryana) (Ryan and Douglas 1994*). Lotus pinnatus is found only in the Nanaimo area on Vancouver Island, where it occurs in moist soil on exposed, coastal low- land areas. Bird’s-foot Trefoil (Lotus corniculatus) could also be confused for L. pinnatus as it occasion- ally grows in wet places, but it is usually found in drier, disturbed sites. Also, L. corniculatus has completely yellow flowers and its leaflets are smaller and more blunt than the other trefoils. '* The information, gathered during field surveys in 2003 formed the basis for a Committee on the Status of Endangered Wildlife in Canada |Status report (Donovan 2004*) and the subsequent assessment of Endangered (COSEWIC 2004*). This paper includes more recent information ie did not appear in the original status report. |’ Taxonomy and nomenclature follow Douglas et al. (1999). lSy7/ 158 Distribution Lotus pinnatus occurs in western North America from Vancouver Island, south to northwest Washing- ton, western Oregon and the Columbia River Gorge, to central California and sporadically eastward to Idaho (Figure 2; Isely 1993; Pojar 1999). In Canada, Lotus pinnatus is known from seven extant populations with- in a small area on the east coast of Vancouver Island near Nanaimo (Donovan 2004’). Habitat The British Columbia populations of L. pinnatus occur within the moist maritime subzone of the Coast- al Douglas-fir (CDF) biogeoclimatic zone (Meidinger and Pojar 1991) in the Nanaimo Lowlands Ecosection of the Georgia Depression Ecoprovince (Demarchi 1996’). In this region, the Olympic Mountains in Washington State to the south and the Insular Mountains on Vancou- ver Island to the west produce a rain shadow effect resulting in a Mediterranean-type climate with warm, dry summers and mild, wet winters. Most of the rain- fall occurs during the winter months, and limited pre- cipitation and high temperatures during the summer months result in pronounced summer moisture deficits (Meidinger and Pojar 1991). Lotus pinnatus grows in open, springy meadows, along the margins of creeks, or in seepages, where underground water comes to the surface and the plants are in close physical contact with cool, flowing water (H. Roemer, personal communication). In all cases, the soils are shallow (< 15 cm), over gently sloping sand- stone or conglomerate bedrock with abundant mois- ture during the growing and blooming period. Lotus pinnatus is most commonly associated with Yellow Monkey-flower (Mimulus guttatus), Sea Blush (Plec- tritis congesta), White Triteleia (Triteleia hyacinthina), Small-leaved Montia (Montia parvifolia), Scouler’s Popcornflower (Plagiobothrys scouleri) and American Speedwell (Veronica beccabunga ssp. americana). Stands of Douglas-fir (Pseudotsuga menziesii) and dense thickets of Nootka Rose (Rosa nutkana), Ocean Spray (Holodiscus discolor), Nine-bark (Physocarpus capitatus) and willow (Salix spp.) exist on the margin of some seepages but Lotus pinnatus does not occur in the shaded understory of these sites, suggesting that the species is shade-intolerant. Prolonged moisture, edge habitat along streams and meadows and shallow soils derived from sedimentary rock are key habitat compo- nents. Other factors, such as slope and aspect are vari- able and do not appear to be critical in defining suit- able habitat. In British Columbia, the elevation for this species ranges between 40 m and 150 m. Biology Zandstra and Grant (1968) studied the biosystem- atics of native and introduced Lotus species in Cana- da and reported that although many of the Old World species are polyploids, none of the North American THE CANADIAN FIELD-NATURALIST Vol. 120 FIGURE 2. Distribution of Lotus pinnatus in North America (from Donovan 2004*). species examined exhibited the same trait. North Ameri- can Lotus species have a base chromosome number of 6 or 7. Of the species native to Canada, L. pinnatus and L. formosissimus are considered to be the most closely related, based on morphological, habitat, and cytological characteristics. A chemotaxonomic study using thin-layer chromatography further supported the general relationships among Canadian Lotus species with L. formosissimus and L. pinnatus demonstrating the greatest degree of similarity (Grant and Zandstra 1968). These species differed from other native Cana- dian species in that both were perennial outcrossers with large flowers on long pedunculate umbels. The other three native species were self-fertile annuals with small flowers. Evolution in angiosperm reproduction has frequently occurred with a decrease in basic chro- mosome number, a shift from an outcrossing (cross- pollinating) to an inbreeding (self-pollinating) repro- ductive system and a shift from a perennial to an annual habit (Stebbins 1957). On this basis, Zandstra and Grant (1968) considered L. pinnatus and L. formo- sissimus to be more primitive than the annual species. Lotus pinnatus germinates in the late winter or early spring when soil moisture and temperature conditions are most favourable, and seedling survival appears to be dependent on continuous surface moisture during this period. Though the specific timing of events will depend on local habitat conditions, flowering in L. pin- natus generally occurs between May and the end of June. 2006 The pea-like, bisexual flowers are borne at the end of a long peduncle arising from the leaf axil, and the umbellate inflorescences are produced sequentially throughout the growing season. As the flowers senesce, some become progressively more reddish (Isely 1981). During July, when drought conditions are prevalent, seed maturation and dispersal begin, followed by die- back of the stems to the rootstock, which is located sev- eral centimetres below the soil surface. The seeds lack any strong adaptations for long-distance dispersal by wind or animal vectors. Most seeds are gravity-dis- persed and generally land in the immediate vicinity of the parent plant. The plant’s habitat along stream channels may permit the legumes and seeds to be trans- ferred by water during times of seasonal flooding. However, germination and seedling survival appear to depend upon continuous surface moisture, and the species does not appear to be a strong competitor with native shrub species or with invasive alien plants. The chances of a healthy population returning to Canada if local populations become extirpated are highly unlike- ly. The nearest population in Washington State, from which collections were made in 1940, is 240 km away in Bremerton. Whether this population is extant is un- known. As with other perennial species in the genus Lotus, Zandstra and Grant (1968) suggested that the flowers of L. pinnatus required cross-pollination to set viable seed. Given the bright yellow appearance of the flower and the wide keel that serves as a suitable landing plat- form, flowers of L. pinnatus appear to be adapted to bee pollination. Although the importance of the bum- blebee to the flower (and vice versa) is yet to be specif- ically determined, bumblebees were observed visiting flowers of L. pinnatus at Harewood Plains during field observations made in 2003. Although bees are capable of dispersing pollen over short distances, cross-polli- nation between most populations in Canada is unlikely in view of the distances involved. Like many other legumes, L. pinnatus appears to be associated with nitrogen-fixing Rhizobium bacteria that occupy root nodules and provide the plants with a source of reduced nitrogen in exchange for a supply of carbon and other nutrients. Specific strains of Rhi- zobium bacteria are required for effective nodulation of Lotus species grown as forage crops, such as L. corniculatus and L. tenuis. To maximize establishment in areas that have never produced Lotus, inoculation of seeds with the appropriate rhizobia is necessary (Blumenthal and McGraw 1999). An examination of root nodules collected from L. pinnatus at Harewood Plains indicated the presence of bacteria, most likely of genus Rhizobium (S. Berch, personal communication). Population Attributes Lotus pinnatus has been reported from nine sites in British Columbia (Figure 3; Table 1), all in the Nanai- DONOVAN: BOG BiIRD’S-FOOT TREFOIL 159 BRITISH , COLUMBIA = Ta Ke ree VICTORIA FiGurE 3. Distribution of Lotus pinnatus in British Colum- bia (from Donovan 2004*) @ confirmed: a uncon- firmed. mo area on Vancouver Island and on nearby Gabriola Island. Of the nine reported records, seven populations were verified in recent years. The remaining two sites have been extensively disturbed and it is unlikely that these populations currently exist. Between 1500 and 2000 individual plants were counted in Canada in 2003 over an area of 650 square metres. Relatively little is known of demographic characteristics and population trends of L. pinnatus due to a lack of long-term mon- itoring. Estimates of population trends in L. pinnatus are currently unreliable as the clumped, sprawling habit makes it difficult to identify separate individuals and counting methods have varied with different investiga- tors. Until counting methods are standardized, popula- tion numbers should be considered rough estimates. Provincial, National and Global Ranks The British Columbia Conservation Data Centre has ranked L. pinnatus as “S1” and placed it on the “Red list”. A rank of “S1” indicates that the species is “critically imperiled because of extreme rarity (often 160 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Locations and Population Sizes for Lotus pinnatus in British Columbia. Populations Observation Collector/ Number of Date Observer Individual Plants Apparent Status Harewood Plains North’ 7 July 2003 M. Donovan 1500 in 500 m? Extant Harewood Plains South 1 12 July 2003 M. Donovan 25 in 4 m? Extant Harewood Plains South 2 12 July 2003 M. Donovan 10 in 2 m Extant Woodley Range 1 June 2003 M. Donovan 120-140 in 90 m? Extant Gabriola Island, off Peterson Road, on Perry Road 25 May 2003 Donovan/Penny 65-70 in 10.5 m? Extant Nanaimo, west of Cinnabar Valley 15 June 2003 M. Donovan 40 in 25 m? Extant Nanaimo, south of Extension 23 June 2003 M. Donovan 30 to 45 in 17 m2 Extant, but declining Waddington & Departure Bay Roads Nanaimo 2 June 1939 J.W. Eastham N/A Extirpated Departure Bay Road and Island Highway, Nanaimo 20 June 1965 K. Beamish N/A Extirpated “The collection by W. R. Carter at Mt. Benson in | June 1918 is included with this population as Harewood Plains may be considered the foothills of Mt. Benson. 5 or fewer occurrences) or because of factors making it especially vulnerable to extirpation.” This is the most critical status that can be applied to a species at the provincial level. Though it currently lacks formal pro- tection in British Columbia, Lotus pinnatus 1s a poten- tial candidate for listing under the provincial Wildlife Amendment Act (2004). At the national level, Lotus pinnatus has been as- sessed as Endangered by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC 2004") and appears on Schedule 1 of the federal Species At Risk Act (Environment Canada 2006’). Globally, Lotus pinnatus has a rank of G4G5 which indicates that the plant is apparently secure in most of its range (NatureServe Explorer 2006’). Threats and Protection Habitat loss presents a serious and urgent threat to Lotus pinnatus in Canada. Over 90% of the extant pop- ulation of L. pinnatus is located on land that is privately owned, making this species vulnerable to habitat loss as a result of urban expansion and residential devel- opment. In addition to potential commercial and residential development, the population of L. pinnatus at Hare- wood Plains is under direct and immediate threat of habitat degradation and fragmentation resulting from intensive, unauthorized use of recreational off-road vehicles; including ATVs, 4 x 4s and dirt bikes (C. Thir- kill, personal communication). In some areas at this site, the thin and fragile soils have been rutted down to bedrock and plants have been dislodged on to bare rock where they cannot re-establish (Figure 4). There is high potential for further impacts as vehicles con- tinue to access the site, despite efforts by the former landowner (Weyerhaueser Canada) to block access to off-road vehicles with boulder placements, ditches and the installation of several cement barriers along the access road (C. Thirkill, personal communication). In an effort to minimize the disturbance of sites desig- nated as “Environmentally Sensitive Areas” the City of Nanaimo has established a Development Permit Area on a portion of the property at Harewood Plains (Rob Lawrance, personal communication). As the habitat at Harewood Plains supports several other species at risk such as Dense-spike Evening Primrose (Epilobium den- siflorum), Green-sheathed Sedge (Carex feta), Foothill Sedge (Carex tumulicola), Slim-leaf Onion (Allium amplectens) and Nuttall’s Quillwort (/soetes nuttallii), a working group comprised of representatives from industry, non-governmental organizations and local and provincial governments has been established in order to develop proactive communication with the different land owners, managers and stakeholders at this location. At the site south of Extension, near Nanaimo, at least some portions of the population have been reported extirpated due to recreational off-road vehicle use and residential development (A. Ceska, personal commu- nication). Other than habitat destruction, competitive exclu- sion from native and non-native vegetation represents the most significant ongoing threat to L. pinnatus at all sites. Encroachment of native shrub species in poten- tial habitats may prevent this species from occupying new sites. Invasive alien grass species that threaten the persistence of L. pinnatus include Sweet Vernalgrass (Anthoxanthum odoratum), Orchard Grass (Dactylis glomerata), Kentucky Bluegrass (Poa pratensis) and Barren Brome (Bromus sterilis). Scotch Broom (Cytisus scoparius) is the most dominant invasive alien shrub. Logging operations close to (within 50 m of) popula- tions of L. pinnatus at Harewood Plains increase the potential for the spread of aggressive native and non- native plant species. By disturbing soils and carrying seeds, off-road vehicles are also a major factor in the spread of invasive non-native plants in sensitive areas. The population located in the Woodley Range Eco- logical Reserve is protected, to a certain extent, by its 2006 DONOVAN: BOG Birpd’s-FOOT TREFOIL 16] a ~ Ss N ay he if FiGure 4. Destruction of habitat caused by all-terrain vehicles at Harewood Plains (Photo George Douglas, 2003). location within an ecological reserve, but this repre- sents only 7% of the total Canadian population. In addition, plants at this location are vulnerable to adja- cent land uses including all-terrain vehicle use, log- ging and other land clearing activities which could impact on the hydrological features that provide seep- age to the location. A national multi-species strategy that addresses the recovery of Lotus pinnatus and five other endangered plant species of vernal pools and other ephemeral wet areas has been completed. This strategy outlines recov- ery goals for each species, suggests strategic approach- es for meeting the recovery goals and recommends that a draft recovery action plan (RAP) be completed by October 2009, which includes proposing Critical Habi- tat for this species under the Species At Risk Act (Parks Canada Agency 2005*). Evaluation The British Columbia Conservation Data Centre considers Lotus pinnatus to be endangered in British Columbia (Donovan 2004*) and the Committee on the Status of Endangered Wildlife in Canada has assessed the species as Endangered (COSEWIC 2004"). Only a few small fragmented populations remain, and these are restricted to wet meadows of limited occurrence that are considerably disjunct from the main range of the species in the north-western United States. The populations are at risk from continued habitat loss, en- croachment of invasive species and recreational off- road vehicle use. Development of the habitat supporting the only sizeable remaining population could result in significant losses to the population (COSEWIC 2004 ). Acknowledgments Thanks to Jenifer Penny, Charles Thirkill and Don Piggott for help with field surveys. Hans Roemer and Shannon Berch provided information and technical assistance. Matt Fairbarns provided strong support throughout this study and Ted Lea reviewed the man- uscript. I am indebted to the late George Douglas for nurturing my interest in plants. Documents Cited (marked * in text) COSEWIC. 2004. COSEWIC assessment and status report on the bog bird’s-foot trefoil Lotus pinnatus in Canada. Committee on the Status of Endangered Wildlife in Cana- da. Ottawa, Ontario. 33 pages. www.cosewic.gc.ca Demarchi, Dennis A. 1996. An Introduction to the Ecore- gions of British Columbia. Wildlife Branch, Ministry of Environment, Lands and Parks, Victoria. (Available at http: srmwww.gov.be.ca/ecology/ecoregions/title_author.html [Accessed 22 June 2006]) Donovan, M. 2004. COSEWIC status report on the bog bird’s-foot trefoil Lotus pinnatus in Canada. Jn COSEWIC assessment and status report on the bog bird’s-foot trefoil Lotus pinnatus in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa, Ontario. 33 pages. 162 Environment Canada, Schedule 1: List of species at risk. Website: http://www.sararegistry.gc.ca/species/schedules_e. cfm?id=1 [Accessed 22 June 2006]. NatureServe Explorer. 2006. NatureServe Explorer: An on- line encyclopedia of life. Version 1.7. Arlington, Virginia, USA. Web site: http://www.natureserve.org/explorer [Ac- cessed 22 March 2006]. Parks Canada Agency. 2005. Recovery Strategy for Multi- species at Risk in Maritime Meadows Associated with Garry Oak Ecosystems in Canada (proposed) in Species at Risk Act Recovery Strategy Series. Edited by Parks Canada Agency. Ottawa, Ontario. 98 pages. Literature Cited Blumenthal, M. J., and R. L. McGraw. 1999. Lotus adap- tation, use and management. Pages 97-119 in Trefoil: The science and technology of Lotus. Edited by P. R. Beuse- linck. Crop Science Society and the American Society of Agronomy. Madison, Wisconsin. Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illus- trated Flora of British Columbia. Volume 3. Dicoteyle- dons (Diapensiaceae through Onagraceae). British Colum- bia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 423 pages. Grant, W. F., and I. I. Zandstra. 1968. The biosystematics of the genus Lotus (Leguminosae) in Canada. II. Numer- ical cytotaxonomy. Canadian Journal of Botany 46: 585- 589. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1961. Vascular Plants of the Pacific Northwest. Part 3: Saxifragaceae to Ericaceae. University of Washing- ton Press, Seattle, Washington. 614 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Isely, D. 1981. Leguminosae of the United States. III. Sub- family Papilionoideae: tribes Sophoreae, Podalyrieae, and Loteae. In: Memoirs of the New York Botanical Garden. 25(3): 181. Isely, D. 1993. Lotus. Pages 616-622 in The Jepson Manual: Higher Plants of California. Edited by J.C. Hickman. Uni- versity of California Press, Los Angeles, California. Meidinger, D., and J. Pojar (editors). 1991. Ecosystems of British Columbia. Special Report, Series 6, British Colum- bia Ministry of Forests, Victoria, British Columbia. 330 pages. Pojar, J. 1999. Fabaceae. Pages 64-180 in Illustrated Flora of British Columbia. Volume 3. Dicotyledons (Diapensi- aceae through Onagraceae). Edited by G. W. Douglas, D. Meidinger, and J. Pojar, British Columbia Ministry of Environment, Lands and Parks, British Columbia Ministry of Forests, Victoria, British Columbia. Ryan, M., and G. W. Douglas. 1994. Status report on the Seaside birds-foot lotus, Lotus formosissimus Greene. Un- published report submitted to the Committee on the Status of Endangered Wildlife in Canada, Ottawa. British Colum- bia Conservation Data Centre, Victoria, British Columbia. 25 pages. Stebbins, G. L. 1957. Self-fertilization and population vari- ability in higher plants. American Naturalist 91: 337-353. Zandstra, I. I., and W. F. Grant. 1968. The biosystematics of the genus Lotus (Leguminosae) in Canada. J. Cytotax- onomy. Canadian Journal of Botany 46: 557-583. Received 24 July 2006 Accepted 24 July 2006 Conservation Evaluation of the Pacific Population of Dwarf Woolly- heads, Psilocarphus brevissimus var. brevissimus, in Canada’ GEORGE W. DouGLas'!, JENIFER L. PENNY’, and KSENIA BARTON? ' Deceased. 2 Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, PO Box 9993 Stn Prov Govt, Victoria British Columbia V8W 9R7 Canada 4Current address: 204-4272 Albert Street, Burnaby, British Columbia V5C 2E8 Canada Douglas, George W., Jenifer L. Penny, and Ksenia Barton. 2006, Conservation evaluation of the Pacific Population of Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada. Canadian Field-Naturalist 120(2): 163-168 In Canada, Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, is restricted to the Similkameen River valley, south of Princeton in southwestern British Columbia and the extreme southeast and southwest corners of Alberta and Saskatchewan, respectively. This paper deals with the three British Columbia populations which represent the northwestern limit of the species which ranges from south-central British Columbia, southward in the western United States to Montana, Idaho, Washington, Oregon, Nevada, Utah, Wyoming , California and Baja California, Mexico. In British Columbia, P. bre- vissimus is associated with calcareous vernal pools and ephemeral pond edges in large forest openings. This habitat is rare in the area the few existing populations could easily be extirpated or degraded through slight changes in groundwater levels, -coalbed methane gas drilling, housing development or recreational vehicles. Key Words: Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus, British Columbia, endangered, distribution, population size. | Dwarf Woolly-heads, Psilocarphus brevissimus vat. (\ _ Nutt.', is a member of a genus of five Q®W species which occur in the Americas (Cronquist 1950). It is one of three species occurring in British Columbia es Fen Ne Yy and Canada (Douglas 1998). Two varieties of P. bre- =e - _vissimus, var. brevissimus and var. multiflorus, are rec- a Ce . 7, “4 ognized. The var. multiflorus occurs only in California ae, UB SS ‘(Morefield 1993). Psilocarphus brevissimus var. bre- aS _vissimus was first recorded in Canada in 1997 (Dou- _gias et al. 1998a). __ Psilocarphus brevissimus var. brevissimus is a pros- trate, matted annual herb with a short taproot (Figure 1; ‘Douglas 1998). The plants are from 8 to 20 cm long. ‘The few leaves are restricted to the stems and are oppo- site, lance-linear to lance-oblong or lance-triangular, 14-15 mm long and white woolly-hairy. The flower heads | jare disciform, lack a pappus, and are usually solitary ‘in the leaf axils or at the tips of the branches and lack involucres. The receptacular bracts are 2.4-4 mm long, ‘hooded and balloonlike. The achenes are more or less cylindric, glabrous, nerveless and tipped with a small, _ 1-2 mm long, offset style. | In British Columbia herbaria, specimens of P. bre- \vissimus var. brevissimus may be distinguished from [" elatior by their prostrate and matted habit and much — Fygure 1. Illustration of Psilocarphus brevissimus var. brevis- narrower leaves. It is separated from P. tenellus var. simus (Line drawing by Elizabeth J. Steven in Douglas tenellus by its longer (2.5-4 mm versus 1.3-2.5 mm et al. 1998; 2002a). } } | The field work for Psilocarphus brevissimus was funded by the British Columbia Conservation Data Centre. The results appear in the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002). This information formed the basis for a Commit- f tee on the Status of Endangered Wildlife in Canada status report (Douglas et al. 2003") and the subsequent assessment of Endangered \(COSEWIC 2003°). This paper includes more recent information that did not appear in the original status report. ‘Taxonomy and nomenclature follows Douglas et al. (1998b; 1998c; 1999; 2001). 163 164 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Locations and population sizes for Psilocarphus brevissimus var. brevissimus south of Princeton, British Columbia. Collection site Observation Collector/Observer | Number of plants/area Princeton, Tracey Lake, ca. 0.75 km SE of Lake 1996 Lomer 30/unknown Princeton, Tracey Lake, ca. 1.3 km ESE of Lake 2002 Douglas and Penny 450/392 m? 2004 Douglas and Smith —7200+500/100 m? Princeton, Stevenson Lake, ca. 0.75 km SE of 2002 Douglas and Penny 900 000+100 000/570 m2 2004 Douglas and Smith —_11 775/155 m? long) receptacular bracts and its much narrower leaves. For many years in Alberta and Saskatchewan, this species was called P. elatior (Packer 1983; Kershaw 2001; Saskatchewan Conservation Data Centre 2004’). In 2004, Alberta taxonomists examined material from both provinces and reidentified them as Psilocarphus brevissimus var. brevissimus (J. Gould, personal com- munication). It is quite likely that the “species of con- cern” ranking given by COSEWIC (2004°) for the Alberta/Sackatchewan populations of P. elatior will now be applied to the P. brevissimus populations. North American and Provincial Ranges Psilocarphus brevissimus var. brevissimus ranges from south-central British Columbia, southward in the western United States to Montana, Idaho, Washington, Oregon, Nevada, Utah, Wyoming, California and Baja California, Mexico (Morefield 1993; Cronquist 1994; Douglas 1998). There are also disjunct populations in Chile and Argentina (Cronquist 1950, 1955). In Cana- da, the species is known only from along 2.6 km of Highway #3 in the Similkameen River valley south of Princeton in south-central British Columbia (Fig- ure 2; Douglas 1998; Douglas et al. 2002) and in the south-eastern and south-western corners of Alberta and Saskatchewan, respectively. Habitat Psilocarphus brevissimus var. brevissimus occurs in the southern interior of British Columbia in the lower montane zone in the Interior Douglas-fir biogeocli- matic zone (Hope et al. 1991). Climatic conditions are continental, characterized by hot, dry summers, a fairly long growing season and cool winters. A rain- shadow effect prevails in this area due to the pres- ence of the Coast-Cascade Mountains to the west. Within this zone, the sites occur in a matrix of level to gently sloping Big Sagebrush (Artemisia tridentata) shrub/grassland with scattered Ponderosa Pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii) [Nomenclature follows Douglas et al. 1998, 1998c, 1999 and 2001]. The area is at the western edge of the distribution of open shrub/grassland at that elevation. Specifically, P. brevissimus var. brevissimus occurs in calcareous clay soils in vernal pools in large forest openings dominated by Scouler’s Popcornflower (Pla- giobothrys scouleri) and Close-flowered Knotweed (Polygonum polygaloides ssp. confertiflorum (Figure 3). Associates in these sites include One-spike Oat- grass (Danthonia unispicata), Tiny Mousetail (Myosu- rus minimus), Carolina Meadow-foxtail (Alopecurus carolinianus), Common knotweed (Polygonum avic- ulare), Lowland Cudweed (Gnaphalium palustre), and Annual Hairgrass (Deschampsia danthonioides). Psilo- carphus brevissimus var. brevissimus also occurs at the edge of ephemeral ponds in the same type of sub- stratum (Figure 4). The vernal pools and ephemeral pond edges are wet in spring and dry and compacted in the summer. Psilocarphus brevissimus var. brevissimus is consid- ered to be a vernal pool specialist (Schlising and San- ders 1982; Keeley and Zedler 1998; Bauder 2000). Keeley and Zedler (1998) define vernal pools as “‘pre- cipitation-filled seasonal wetlands inundated during periods when temperature is sufficient for plant growth, followed by a brief waterlogged-terrestrial stage and culminating in extreme desiccating soil conditions of extended duration”. The species is able to outcompete grassland species due to its tolerance of inundation, and aquatic/wetland species due to its tolerance of soil dessication and heat during summer drought (Bauder 2000). Biology Little information is available on the biology of Psilocarphus brevissimus var. brevissimus in British Columbia. Only basic facts about its life cycle and reproduction are known. Keeley and Zedler (1998) recognize four stages in the annual vernal pool cycle: (1) a wetting phase; (2) an aquatic or inundation phase; (3) a waterlogged-terres- trial phase; and (4) the drought phase. In vernal pool Species, germination is typically initiated during the wetting or inundation phases. Flowering is usually ini- tiated during the transition to the waterlogged-terrestrial phase. In the Princeton area of south-central British Columbia, this phase occurs during June and fruiting follows in July during the drought period. Psilocarphus brevissimus var. brevissimus is an an- nual herb that likely reproduces primarily by self-pol- lination. The assumption of a selfing breeding system is based on the taxon’s lack of pollinator attracting struc- tures and the proximity of the stigmas to the central pollen-producing flowers. In addition, the abundance of woolly hairs and lack of achene pappus appear to limit pollen dispersal by wind. Another possibility is that plants set seed without pollination, through asex- ual reproduction (Cronquist 1950). 2006 Animals may be the primary vectors of seed disper- sal for Psilocarphus brevissimus var. brevissimus. Birds may disperse seeds over small and large geographic areas. The use of vernal pools by various migrating avian species, particularly waterfowl and shorebirds, is well documented in other regions (Silveira 1998). Finally, cottontail rabbits (Sy/vilagus spp.) are poten- tial agents of dispersal of P. brevissimus var. brevis- simus seeds. A study in California found that P. bre- vissimus Var. brevissimus seeds were commonly found in cottontail rabbit pellets. The excreted seeds germi- nated easily, suggesting that the lagomorphs may be important agents of seed dispersal between vernal pools on a small scale (Zedler and Black 1992). British Columbia sites for P. brevissimus var. brevissimus may provide habitat for Nuttall’s Cottontail (Sy/vila- gus nuttallii), a species that frequents grasslands. Population Attributes The three populations of Psilocarphus brevissimus var. brevissimus were discovered in 1997 in the Simil- kameen River valley, west of Princeton, British Colum- bia (Table 1; Douglas et al. 1998a). These populations are between approximately 0.8 and 2.6 km apart (Fig- ure 2). During the period from 2002 to 2004, two of the populations decreased in size by about 25%. Plant numbers increased at one site from 450 to about 7200 plants and decreased at the other from about 900 000 to about 12000 plants. Trends for these populations are not well known but it is obvious that they can be expected to vary marked- ly due to differences in yearly seed germination and seedling success. Population sizes of this annual herb are strongly tied to annual precipitation patterns (Baud- er 2000) and are typical of vernal pool plants (Griggs and Jain 1983). Provincial, National and Global Ranks Globally, Psilocarphus brevissimus var. brevissimus has a rank of G4T4 indicating that in most of its range the plant is secure and frequent to common. Since the species is restricted to British Columbia it has a nation- al rank in Canada of N1. Provincially, P. brevissimus var. brevissimus has been ranked S1 by the Conserva- tion Data Centre and appears on the British Columbia Ministry of Sustainable Resource Management red list (Douglas et al. 2002). This is the most critical rank that can be applied to species at the provincial level and indicates that the species is “critically imperiled be- cause of extreme rarity (typically five or fewer occur- rences or very few remaining individuals) or because of some factor(s) making it especially vulnerable to extirpation or extinction”. Threats and Protection The major threats to P. brevissimus var. brevissimus are through drilling for coalbed methane gas, housing development and off-road recreational vehicles. In 2002 DOUGLAS, PENNY, AND BARTON: DWARF WOOLLY-HEADS 165 ™m— Ly» \ \ \ a BRITISH COLUMBIA PRINCETON FIGURE 2. Distribution of Psilocarphus brevissimus var. bre- vissimus in British Columbia. three test holes, targeting coalbed methane gas deposits, were drilled in the Similkameen valley. If brought into production the drilling, access roads and pumping sta- tions could cause major degradation in the area. Other potential threats result from activities that are permitted in the Agricultural Land Reserve. The private property where P. brevissimus var. brevissimus occurs is on the Agricultural Land Reserve, where pri- mary land use is agriculture. For instance, some types of fill/soil removal are allowable without application to the Agricultural Land Commission (Provincial Agri- 166 THE CANADIAN FIELD-NATURALIST Vol. 120 FIGURE 3. Calcareous vernal pool habitat of Psilocarphus brevissimus var. brevissimus south of Princeton, British Columbia. Colorado Rush (Juncus confusus), another rare species in British Columbia, is conspicuous on both sides of the dried- up vernal pool. The several large plants in the habitat are the introduced grass, American Sloughgrass (Beckmannia syzigachne). cultural Land Commission 2003°). On these lands, development pressures do not appear to be an imme- diate issue at this time. The Agricultural Land Reserve status may prevent subdivision development, but does allow other activities that could also potentially threaten the populations. In recent years many tracts of Agri- cultural Land Reserve land in southern British Colum- bia has been converted to housing developments, shop- ping malls and golf courses, either by decisions of the Agricultural Land Commission or very rarely by an “order in council” by the sitting provincial legislature. Another threat likely to have a negative impact is recreational off-road vehicle use. The authors observed evidence of all-terrain vehicles and dirt bike use in the area of the sites in 2004. The vernal pools and ephemer- al pond edges are wet in spring and are susceptible to permanent changes if disturbed at this. An additional threat to the population of P. brevis- simus var. brevissimus is its vulnerability to extirpation due to the extremely small occupied area and popula- tion size. Such small populations are at risk of inbreed- ing depression (Primack 1998). Due to lack of genetic variation, these small populations are vulnerable to demographic and environmental variation. Further- more, suitable habitats for P. brevissimus var. brevis- simus in the southern interior of British Columbia are extremely limited and thus opportunities for coloni- sation are also limited. Psilocarphus brevissimus vat. brevissimus 1s not for- mally protected in British Columbia, however, it could be in the future since this species is a potential candi- date for listing under the provincial Wildlife Amend- ment Act (2004). As part of its commitment to the National Accord (National Accord for the Protection of Species at Risk), the province is required to take measures to protect this species. It is on Schedule 1 of the federal Species At Risk Act and a recovery strategy is expected to be completed by the summer of 2006. Evaluation The British Columbia Conservation Data Centre considers P. brevissimus var. brevissimus to be threat- ened/endangered in British Columbia (Douglas et al. 2002) and the Committee on the Status of Endangered Wildlife in Canada has assessed the species as endan- gered (COSEWIC 2003*). The three populations are small and vulnerable to a number of threats. With lim- ited knowledge of the plant’s biological and ecological requirements, this species is vulnerable to extirpation in British Columbia. Without research on growth re- quirements and further demographic information, the stability of the present populations will remain un- known. The limited number of plants also reduces the potential for genetic variation, which may be necessary to respond to environmental changes in the future. 2006 Acknowledgments We thank Frank Lomer for providing information on his original discovery of Psilocarphus brevissimus var. brevissimus in 1997 and for his assistance in re- locating the sites in 2002. Shyanne J. Smith assisted with field work in 2004. Documents Cited (marked * in text) COSEWIC., 2003. COSEWIC assessment and status report on Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus in Canada, Committee on the Status of Endan- gered Wildlife in Canada. Ottawa, Ontario. 19 pages. www. cosewic.gc.ca. COSEWIC., 2004. Canadian Species at Risk. Committee on the Status of Endangered Wildlife in Canada. Ottawa, Ontario. 13 pages. www.cosewic.gc.ca. Douglas, G. W., J. L. Penny and K. Barton. 2003. COSE- WIC Status report on Dwarf Woolly-heads, Psilocarphus brevissimus var. brevissimus in Canada. In COSEWIC as- sessment and status report on Dwarf Woolly-heads, Psilo- carphus brevissimus var. brevissimus in Canada. Commit- tee on the Status of Endangered Wildlife in Canada. Ottawa, Ontario. 1-19 pages. www.cosewic.gc.ca. Provincial Agricultural Land Commission. 2003. Agricul- tural Land Reserve — Frequently Asked Questions. Provin- cial Agricultural Land Commission. 133-4940 Canada Way, Burnaby, British Columbia V5G 4K6 Canada. http://www. alc.goy.be.ca/faq/faq_alr.htm. Saskatchewan Conservation Data Centre. 2004. Tracked species list for vascular plants. Saskatchewan Conservation Data Centre, Fish and Wildlife Branch, Saskatchewan Environment and Resource Management. Regina, Saskat- chewan. Literature Cited Bauder, E. T. 2000. Inundation effects on small-scale plant distributions in San Diego, California vernal pools. Aquatic Ecology 34: 43-61. Cronquist, A. 1950. A review of the genus Psilocarphus. Research Studies of the State College of Washington 18: 71-89. Cronquist, A. 1955. Vascular plants of the Pacific Northwest. Part 5: Compositae. University of Washington Press, Seat- tle, Washington. 343 pages. Cronquist, A. 1994. Intermountain flora-vascular plants of the Intermountain West, U.S.A. Volume 5: Asterales. New York Botanical Garden, New York, New York. 496 pages. Douglas, G. W. 1998. Asteraceae. Pages 96-392 in Illustrat- ed flora of British Columbia. Volume 1. Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). Edited by G. W. Douglas, G. B. Straley, and D. Meidinger. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., F. Lomer, and Hans L. Roemer. 1998a. New or rediscovered native vascular plant species in British Columbia. Canadian Field-Naturalist 112: 276-279. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare native vascular plants of British Columbia. Second edi- tion. Province of British Columbia, Victoria, British Colum- bia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illustrated flora of British Columbia. Volume 4. Dicotyledons (Oroban- DOUGLAS, PENNY, AND BARTON: DWARF WOOLLY-HEADS 167 oa or . ; vy. - aint of, * | a : a i FIGURE 4. The ephemeral pond habitat of Psilocarphus bre- vissimus Var. brevissimus south of Princeton, British Columbia. Myosurus minimus, Plagiobothrys scouleri and Polygonum aviculare are also prominent species on the calcareous clay soils. The adjacent bands of vegetation consist mainly of Meadow Barley (Hordewmn brachyantherum) in the foreground and Common Cat- tail (7ypha latifolia) in the background. chaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2001. Illustrat- ed flora of British Columbia. Volume 7. Monocotyledons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Colum- bia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1998b. Illustrated flora of British Columbia. Volume 1. Gymno- sperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998c. Illustrated flora of British Columbia. Volume 2. Dicotyledons. (Balsaminaceae through Cuscutaceae). Bri- tish Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 401 pages. 168 Griggs, F. T., and S. K. Jain. 1983. Conservation of vernal pool plants in California, I]. Population biology of a rare and unique grass genus Orcuttia. Biological Conserva- tion 27: 171-193. Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson, W. L. Harper, and B. M. Wikeem. 1991. Interior Douglas- fir zone. Pages 153-166 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Ministry of Forests Special Report Series Number 6, Vic- toria, British Columbia. 330 pages. Keeley, J. E., and P. H. Zedler. 1998. Characterization and global distribution of vernal pools. Jn Ecology, conserva- tion, and management of vernal pool ecosystems: Proceed- ings from a 1996 Conference. Edited by C. W. Witham, E. T. Bauder, D. Belk, W. R. Ferren Jr., and R. Ornduff. California Native Plant Society, Sacramento, California. Kershaw, L. 2001. Rare vascular plants of Alberta. The Uni- versity of Alberta Press, Edmonton, Alberta. 484 pages. Morefield, J. D. 1993. Psilocarphus. Page 329 in The Jep- son manual: Higher plants of California. Edited by J.C. Hickman. University of California Press. Berkeley, Cali- fornia. 1400 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Packer, J. G. 1983. Flora of Alberta. Second edition. Univer- sity of Toronto Press, Toronto, Ontario. 687 pages. Primack, R. B. 1998. Essentials of conservation biology. Sinauer Associates Inc., Sunderland, Massachusetts. Schlising, R. A., and E. L. Sanders. 1982. Quantitative analy- sis of vegetation at the Richvale vernal pools, California. American Journal of Botany 69: 734-742. Silveira, J. G. 1998. Avian uses of vernal pools and implica- tions for conservation practice. Jn Ecology, conservation, and management of vernal pool ecosystems. Proceedings from a 1996 Conference. Edited by C. W. Witham, E. T. Bauder, D. Belk, W. R. Ferren Jr., and R. Omduff. Califor- nia Native Plant Society, Sacramento, California. Zedler, P. H., and C. Black. 1992. Seed dispersal by a gen- eralized herbivore: rabbits as dispersal vectors in a semi- arid California vernal pool landscape. American Midland Naturalist 128: 1-10. Received 8 October 2002 Accepted 20 March 2006 macrophyllum, in Canada* GEORGE W. DouG-Las! and JULIE DEsROSIERS2 ' Deceased. in Canada at this time. The Pacific Rhododendron, Rhododendron macro- phyllum D. Don ex G. Don’, is a member of a genus of almost 1000 species occurring mainly in north- eastern Asia (Wallace 1993). It is one of three species | occurring in British Columbia (Meidinger 1999) and four occurring in Canada (Scoggan 1979). Rhododendron macrophyllum was first reported by naturalist Archibald Menzies in 1792 in his “Journal of Vancouver’s Voyage”, and subsequently described by David Don in 1834. Joseph Hooker also described plants collected during Vancouver’s voyages, and in 1855 he published a description of this rhododendron under the species name californicum. This name was in common use by botanists for close to a century. Rhododendron macrophyllum is an erect to spread- ing, branched shrub, 1-8 m tall (Meidinger 1999; Fig- ure 1). The evergreen leaves are alternate, often appear- |ing whorled, ovate or elliptical, 8-20 cm long, 3-6 cm |wide, leathery, dark green above, paler beneath and glabrous, or nearly so. The pale-pink to deep rose- purplish corollas are tubular bell-shaped, spotted red- |brown and 2.5-4 cm long and up to 5 cm across. The fruits consist of woody capsules 1.5-2 cm long with numerous minute seeds. It is easily separated from the other two Rhododendron species in British Columbia. Lapland Rosebay (Rhododendron lapponicum) is a northern plant with much shorter habit and smaller leaves, while White-flowered Rhododendron (R. albi- florum) is a higher elevation species with white flowers. Conservation Evaluation of Pacific Rhododendron. Rhododendron * 4254 Garden Drive, Burnaby, British Columbia VSG 4G6 Canada Douglas, George W., and Julie Desrosiers. 2006. Conservation evaluation of Pacific Rhododendron, Rhododendron macrophyllum, in Canada. Canadian Field-Naturalist 120(2); 169-174. In Canada, Pacific Rhododendron (Rhododendron macrophyllum) is restricted to the Skagit River drainage and southeastern _ Vancouver Island in southwestern British Columbia. These populations represent the northern limits of the species, which ranges south to northern California. In British Columbia, Rhododendron macrophyllum is usually associated with coniferous forests. Until recently this species was considered to be rare. Results of this study indicate that the populations in the Skagit River watershed are large, extensive and mostly contained in protected areas, thus the Species is not considered to be at risk Key Words: Pacific Rhododendron, Rhododendron macrophyllum, distribution, conservation evaluation, British Columbia North American and Provincial Ranges Rhododendron macrophyllum is restricted to coastal habitats from northern California north through Oregon and Washington to southwestern British Columbia (Hitchcock et al. 1959). In Canada, it occurs in south- western British Columbia in the western Cascade Moun- tains (Skagit River valley) and on southeastern Van- couver Island (Shawnigan and Rhododendron lakes) [Meidinger 1999; Douglas et al. 2002; Figure 2}. Habitat Skagit River Valley The Rhododendron macrophyllum populations in the Skagit River valley occur on well-drained, coarse- textured, gravelly, dry soils overlain by fibremors. Ele- vations range from 500 to 870 metres. About half of the Rhododendron populations in the Skagit River valley are within the Interior Douglas-fir (IDF) Zone while the remaining are within the Coastal Western Hemlock (CWH) Zone (Hope et al. 1991; Pojar et al. 1991). Summaries of all plant communities within these zones are taken from Desrosiers and Cad- rin (1993). The four plant communities within the Interior Douglas-fir (IDF) Zone are described below: (a) Douglas-fir/Falsebox/Feathermoss (Pseudotsuga menziesii Paxistima myrsinites/Pleurozium schreberi) community. Sites for this community are located in the dry, mid- dle part of the Skagit River Valley, on the flats of the The field work for the Rhododendron macrophyllum project was funded by the British Columbia Conservation Data Centre and The Skagit avironmental Endowment Commission. The results appear in the British Columbia Conservation Data Centre database and a Skagit Envi- onmental Endowment Commission report (Desrosiers and Cadrin 1993"). This information formed the basis for a Committee on the Status of Endangered Wildlife in Canada status report (Desrosiers and Douglas 1997") and the current assessment of nor at risk (COSEWIC 2006’). Taxonomy and nomenclature follows Douglas et al. (1998a, 1998b, 1999a, 1999b, 2000, 2001) and Schofield (1992). 169 170 THE CANADIAN FIELD-NATURALIST Vol. 120 FiGurE 1. illustration of Rhododendron macrophyllum. (Line drawing by Lora May Richards) valley floor. These are the driest IDF sites and were mid-range in terms of their nutrient regime. The 80- 115 year old stands are dominated by Pseudotsuga menziesii, Paxistima myrsinites, and Pleurozium schre- beri. Other important species include Rhododendron macrophyllum, Lodgepole Pine (Pinus contorta), White Hawkweed (Hieracium albiflorum), Heart-leaved Tway- blade (Listera cordata) and Cow-wheat (Melampyrum lineare var. lineare). Rhododendron macrophyllum had an average cover of 27% in this community. (b) Douglas-fir — Western Redcedar/Beaked Hazelnut (Pseu- dotsuga menziesii-Thuja plicata/Corylus cornuta) com- munity. The lowest elevations in the lower part of the valley contained these sites. They tended to be intermediate, both environmentally and floristically, for the IDF in this area. Pseudotsuga menziesii, Thuja plicata, and Corylus cornuta were dominants in these 80-100 year old stands. Cover values for Thuja plicata, Western Hemlock (Tsuga heterophylla), Paxistima myrsinites 1 | ' n 2006 BRITISH COLUMBIA Koksilah River VICTORIA DOUGLAS AND DESROSIERS: PACIFIC RHODODENDRON 171] -=-- as ly ‘MANNING \. PROVINCIAL \ PARK BB Ecological Reserve ---- Boundary =—— Road FIGURE 2. The location of Rhododendron macropyhilum sites in British Columbia. and Dull Oregon-grape (Mahonia nervosa) were high- est in these sites. Average cover of Rhododendron macrophyllum was 3%. (c) Douglas-fir — Western Redcedar/Vine Maple (Pseudot- suga menziesii-Thuja plicata/Acer circinatum) community. This community is found at a higher elevation than other IDF sites, on the east-facing slopes of the Rhodo- dendron Ecological Reserve. The sites were mid-range in term of moisture regime and were nutrient-poor. Pseudotsuga menziesii, Thuja plicata, Acer circinatum and Step Moss (Hylocomium splendens) were the dom- inant species. Douglas Maple (Acer glabrum var. dou- glasii), Thimbleberry (Rubus parviflorus var. parvi- florus), Oceanspray (Holodiscus discolor), Hooker's Fairybells (Prosartes hookeri var. oregana), Fireweed (Epilobium angustifolium), and Lanky Moss (Rhy- tidiadelphus loreus) were also important. Rhododen- dron macrophyllum had an average cover of 3%. (d) Pacific Rhododendron/Westerm Tea-berry/Cladonia (Rho- dodendron macrophyllum/Gaultheria ovatifolia/Cladonia) community. This Rhododendron community occurs on the east- ern section of the Rhododendron Ecological Reserve as well as on the west side of the Skagit River. These sites were among the driest and most nutrient-poor of all sites sampled. The stands were 105-115 years old. 172 Tree cover in these plots was extremely low. Pseudo- tsuga menziesii had an average cover of 8% while West- em White Pine (Pinus monticola) had an average cover of 7%. Average cover of Rhododendron macrophyl- lum was the highest of all plots at 70%. Species of importance in this community included Soopolallie (Shepherdia Canadensis), Paxistima myrsinites, Twin- flower (Linnaea borealis), Mahonia nervosa, Kinnikin- nick (Arctostaphylos uva-ursi) and species of lichen (Cladonia spp.). All of the plots sampled in the northern half of the study area fell within a single community type in the Coastal Western Hemlock (CWH) Zone. This com- munity may be summarized as follows: Douglas-fir — Western Hemlock/Falsebox (Pseudotsuga men- ziesti-Tsuga heterophylla/Paxistima myrsinites ) community. Compared to IDF sites the CWH sites have higher moisture regimes, lower nutrient levels and higher tree cover values. In addition to Pseudotsuga menziesii, Tsuga heterophylla and Paxistima myrsinites other important species include Amabilis Fir (Abies ama- bilis), Grouseberry (Vaccinium scoparium) and Hylo- comium splendens. The stands ranged from 80-150 years old. Average cover of Rhododendron macro- phyllum was 18%. Southeastern Vancouver Island The Rhododendron sites on southeastern Vancouver Island are within the Coastal Douglas-fir Zone (Neuz- dorfer et al. 1991). The populations east of Shawnigan Lake were found in the following plant community: Western Redcedar/Pacific Rhododendron-Alaskan Blueberry/ Step Moss (Thuja plicata/Rhododendron macrophyllum- Vaccinium alaskense/Hylocomium splendens) community. This community was located on a moderate, south- west slope at an elevation of 490 m approximately 16 km west of Shawnigan Lake. The small (550 m7) area on which Rhododendron occurred was dominated by 40 m tall Thuja plicata. The shrub layer was well- developed with Rhododendron macrophyllum, Vac- cinium alaskense, Red Huckleberry (V. parvifolium) and Salal (Gaultheria shallon) dominating. A dense layer of mosses and lichens was dominated by Hylo- comium splendens. At Rhododendron Lake three markedly different communities occur in bands around Rhododendron Lake. A very wet band of vegetation (about 3-5 m wide), dominated mainly by Sweet Gale (Myrica gale), occurs around the immediate edge of the lake. Adja- cent to the latter band is another, slightly drier but still very moist 3-10 m band of vegetation containing Myri- ca gale and Rhododendron macrophyllum. This satu- rated habitat is atypical for Rhododendron macrophyl- lum. A wider band (5-25 m wide) of forested vegetation occurs upslope from the lakeside communities. Approx- imately nine ha was covered by Rhododendron plants in the latter two vegetation bands. The two communi- ties containing Rhododendron may be summarized as follows: THE CANADIAN FIELD-NATURALIST Vol. 120 (a) Pacific Rhododendron — Sweet Gale/Salal (Rhododendron macrophyllum-Myrica gale/Gaultheria shallon) community. This community occurs in a 3-10 m wide band that extends for about 300 m along the south side of the lake. Soils are constantly wet with the presence of Skunk Cabbage (Lysichitum americanum) and Sphag- num sp. indicating the hydric nature of the site. This site had been logged many years earlier. It appears all trees cut were Yellow-cedar (Chamaecyparis nootka- tensis) and would have had about a 40-60% average cover. At the present time, Rhododendron macrophyl- lum has an average cover of 47% with only Gaultheria shallon being more abundant (average cover of 57%). Bracken Fern (Preridium aquilinum), Labrador Tea (Ledum groenlandicum) and Myrica gale were also important species in the community. (b) Western Hemlock/Pacific Rhododendron/Salal (Tsuga heterophylla/Rhododendron macrophyllum/Gaultheria shal- lon) community. Upslope from the previous community is a forested community dominated by Tsuga heterophylla. The site has been partially logged and was previously dom- inated by Thuja plicata. Other coniferous species pres- ent, but in low numbers, include Chamaecyparis nootka- tensis, Thuja plicata, Pinus monticola and Pseudotsuga menziesii. Important understory species include Gaul- theria shallon, Rhododendron macrophyllum and Hylocomium splendens. Rhododendron has an aver- age cover of 61%. Biology and Ecology Research on the biology and ecology of Rhododen- dron macrophyllum has been limited. The majority (60%) of populations in the Skagit Valley showed good to very good vigour. One in three populations had young genets less than one metre in height. These appear, for the most part, to have germinated on rot- ten logs or thick moss mats. Asexual reproduction, by layering, appears to be much more common than sexual reproduction. Many sites showed evidence of fire on the domi- nant trees and charcoal layers were often found in the soil profile. This history of fire and its importance to the maintenance of Rhododendron macrophyllum pop- ulations was also noted in an Olympic National For- est study (Henderson et al. 1989). Population Attributes Approximately 200 populations of Rhododendron macrophyllum have been observed at 15 ecosites in the Skagit River valley over a distance of 25 km (Fig- ure 2). Of these, 92 were examined in detail at 13 eco- sites for this study (Table 1). Populations ranged in size from 4 m? to about 200 000 m°’. The latter area (part of the Rhododendron Ecological Reserve) had an extrapolated count of approximately a million canes. Two populations on Vancouver Island were also exam- ined. Counts at one of them (Rhododendron Lake) revealed 400 plants and approximately 9000 canes. 2006 DOUGLAS AND DESROSIERS: PACIFIC RHODODENDRON 173 TABLE |. Locations and population sizes of Rhododendron macrophyllum in south-western British Columbia Number of Populations Map Location Number(s)/ Ecosite Name Manning Provincial Park 1. Cayuse Flats | 2. Silverdaisy Trail 3 3. Rhododendron Flats 4 Skagit Valley Provincial Park 4. Upper Skagit Area 3 5. Klesilkwa River 6 6. Valley Floor North 7. Pinus contorta Terrace North 8. Pinus contorta Terrace South 9. Shawatum Area 10. Rhododendron Ecological Reserve, East 11. Rhododendron Ecological Reserve, West 12. Nepopekum area 13. Galene Lakes Trail Vancouver Island 1. Rhododendron Lake 2. Shawnigan Lake | n WAwWe LK WY Approximate Area of Populations (m*) Approximate Total Number of Canes 300 1 000 250 1 750 150 2 400 12 600 360 600 400 2 100 3 450 no count 920 4 000 140 300 200 000 1 000 000 40 550 60 1 000 100 1 250 90 000 9 000 550 no count ' Detailed counts of canes per plant were made for 50 plants at both Cayuse Flats and Rhododendron Lake. Numbers of canes per plant ranged from 18 to 24 at Cayuse Flats and 20 to 25 at Rhododendron Lake (Desrosiers and Douglas 1997*). The intensive inventory work conducted by the British Columbia Conservation Data Centre resulted in the documentation of thousands of plants in the Skagit Valley (Table 1). Prior to this work the species had been reported as a rare plant in British Columbia (Straley et al 1985; Argus and Prior 1990). Locations of Rhododendron macrophyllum in the nearby Chilliwack River valley were mentioned by Slaney (1971*); however, no references or supporting information were provided in that document. It is pos- sible that these populations could be extirpated due to development. In addition, Rhododendron macro- phyllum was reported from Lightning Lake in Man- ning Park (Carl et al. 1952); this population was not verified by the authors or others working in that area. Provincial, National and Global Ranks Provincially R. macrophyllum is ranked S4 (Douglas et al. 2002) which indicates that it is “rare or uncommon (typically 21 to 100 occurrences); may be susceptible to large-scale disturbances; e.g., may have lost extensive peripheral populations”. Since the species is restricted to British Columbia, the National rank is N1. Globally, Rhododendron macrophyllum is ranked GS. This rank- ing indicates that, on a global scale, the plant is con- sidered “frequent to common or very common; demon- strably secure and essentially ineradicable under present conditions”. Threats and Protection Habitat destruction, due to logging and road build- ing, likely eliminated some populations of Rhododen- dron macrophyllum, especially in the Skagit River area. The establishment of Manning Provincial Park and the more recent Skagit Valley Provincial Park has eliminated most threats to R. macrophyllum. On Vancouver Island, where the two populations are locat- ed within timber harvesting areas, small informal pro- tected areas have been established. In the Skagit River valley it was evident that fire had previously swept through most of the ecosites (Des- rosiers and Cadrin 1993"). Whenever possible, natural fires are quickly extinguished in the Cascades Moun- tains. If this control is successful over long periods of time, fuel loads may build up resulting in extremely hazardous conditions. It is, therefore, possible that pop- ulation sizes may be temporarily reduced from time to time after large burns. Rhododendron macrophyllum receives protection since all but the Vancouver Island populations occur in provincial parks. The Vancouver Island populations have informal protection set up by the timber compa- nies that own the properties Until recently there was formal protection under the Dogwood, Rhododendron and Trillium Protection Act. This act was repealed in 2002 due to our greater knowl- edge of the abundance of the species involved and the fact that sufficiently large numbers of their populations occur in protected areas. Evaluation Rhododendron macrophyllum was considered rare in Canada for many years (Straley et al. 1985; Argus and Pryor 1990). The present study, however, indicates that the species is not rare and is much more abun- dant than previously thought. Most of the extant sites 174 are secure since they are contained either within E. C. Manning Provincial Park, the adjacent Skagit Val- ley Provincial Park or in several ecological reserves. Acknowledgments We thank Shane Ford for locating and examining the Shawnigan Lake population and Jenifer L. Penny for office assistance. Documents Cited (marked * in text) COSEWIC. 2004. Canadian Species at Risk. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. www.cosewic.gc.ca. Desrosiers, J., and C. M. Cadrin. 1993. The Pacific Rho- dodendron (Rhododendron macrophyllum D. Don ex G. Don) communities of the Skagit River watershed, British Columbia. Unpublished Report, The Skagit Environmen- tal Endowment Commission. 27 pages. Desrosiers, J., and G. W. Douglas. 1997. Status Report on the Pacific Rhododendron, Rhododendron macrophyllum D. Don ex G. Don (Ericaceae), in Canada. Ottawa, Ontario. 22 pages. Slaney, F. F. 1971. Rhododendrons in the Skagit Valley. The Canadian Skagit Environmental Newsletter Volume 7, 4 pages. Literature Cited Argus, G. W., and K. M. Pryer. 1990. Rare vascular plants in Canada — our natural heritage. Canadian. Museum of Nature, Ottawa, Ontario. 191 pages. Carl, G. C., C. J. Guiguet, and G. A. Hardy. 1952. A natural history survey of the Manning Park area, British Columbia. British Columbia Provincial Museum Occasional Paper 9. Victoria, British Columbia. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare native vascular plants of British Columbia. Second edition. British Columbia Ministry of Sustainable Resource Management, Victoria, British Columbia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- trated flora of British Columbia. Volume 3. Dicotyledons (Diapensiaceae through Onagraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 423 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- trated flora of British Columbia. Volume 4. Dicotyledons (Orobanchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illustrated flora of British Columbia. Volume 5. Dicotyledons (Sali- caceae to Zygophyllaceae) and Pteridophytes. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 389 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2001. Illustrat- ed flora of British Columbia. Volume 7. Monocotyledons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Douglas, G. W., G. B. Straley, D. Meidinger and J. Pojar. 1998a. Illustrated flora of British Columbia. Volume 1. Gymnosperms and Dicotyledons. (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998b. Illustrated flora of British Columbia. Volume 2. Dicotyledons. (Balsaminaceae through Cuscutaceae). Bri- tish Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, Bri- tish Columbia. 401 pages. Henderson, J. A., D. H. Peter, R. D. Lesher, and D. C. Shaw. 1989. Forested plant associations of the Olympic National Forest. United States Department of Agriculture, Forest Service, Pacific Northwest Region, Ecological Tech- nical Paper Report 001-88, Portland, Oregon. 502 pages. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1959. Vascular plants of the Pacific Northwest. Part 4: Ericaceae through Campanulaceae. University of Washington Press. Seattle, Washington. 510 pages. Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson, W. L. Harper, and B. M. Wikeem. 1991. Interior Dou- glas-fir zone. Pages 153-166 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Ministry of Forests Special Report Series (6) Victoria, British Columbia. 330 pages. Meidinger, D. 1999. Ericaceae. Pages 14-55 in Illustrated Flora of British Columbia. Volume 3 — Dicotyledons (Dia- pensiaceae through Onagraceae). Edited by G. W Douglas, D. Meidinger and J. Pojar. British Columbia Ministry or Environment, Lands and Parks, British Columbia Ministry of Forests, Victoria, British Columbia. 423 pages. Nuszdorfer, F. C., K. Klinka, and D. A. Demarchi. 1991. Coastal Douglas-fir zone. Pages 95-112 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Ministry of Forests Special Report Series (6), Victoria British Columbia. 330 pages. Pojar, J.. K. Klinka, and D. A. Demarchi. 1991. Coastal Western Hemlock Zone. Pages 95-111 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Ministry of Forests Special Report Series (6), Victoria, British Columbia. 330 pages. Schofield, W. B. 1992. Some common mosses of British Columbia. Royal British Columbia Museum. 394 pages. Scoggan, H. J. 1979. The flora of Canada. Part 4 — Dicotyle- doneae (Loasaceae to Compositae). National Museum of Natural Sciences Publications in Botany (7). Ottawa, On- tario. Pages 1117-1626. Straley, G. B., R. L. Taylor, and G. W. Douglas. 1985. The : rare vascular plants of British Columbia. National Muse- - um of Natural Sciences. Syllogeus (59), Ottawa, Ontario. . 165 pages. Wallace, G. D. 1993. Ericaceae. Pages 544-567 in The Jep- son Manual — Higher plants of California. Edited by J.C. Hickman. University of California Press, Berkeley, Cali- fornia. 1400 pages. Received 14 October 2002 Accepted 20 March 2006 Conservation Evaluation of Slender Collomia, Collomia tenella. in Canada* GEORGE W. DouG-as'! and JENIFER L. PENNY? ' Deceased. 2 Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, PO Box 9993 Stn Prov Govr, Victoria, British Columbia V8W 9R7 Douglas, George W., and Jenifer L. Penny. 2006. Conservation evaluation of Slender Collomia, Collomia tenella, in Canada Canadian Field-Naturalist 120(2): 175-178. In Canada, Slender Collomia, Collomia tenella, is restricted to the Princeton area in southwestern British Columbia. The single population represents the northern limits of the species, which ranges from southwestern British Columbia, south in the western United States to Wyoming, Utah, Nevada and Oregon. In British Columbia, C. tenella is associated with an eroded section of a steeply sloping, southeast-facing sandy ridge. Population numbers fluctuate and in some years plants fail to appear. The major threats to C. tenella are through drilling for coalbed methane gas, sand removal for road construction, housing development and off-road recreational vehicles. Key Words: Slender Collomia, Collomia tenella, endangered, distribution, population size, British Columbia. Slender Collomia, Collomia tenella A. Gray’ (Pole- - moniaceae), is member of a genus of about 13 species | found in North and South America (Hitchcock et al. 1959). Four species occur in British Columbia and Canada (Scoggan 1979; Pojar 1999). Collomia tenella _ was first recorded in Canada by Douglas et al. (1998a). _ Collomia tenella is an ascending to spreading, freely branched, annual, tap-rooted herb up to 15 cm tall | (Figure 1; Pojar 1999). The leaves are alternate, lin- | ear, entire, 1-5 cm long and 1.5 mm wide. Flowers are single or in pairs at the branch tips, in the leaf axils or at the forks of the branches. The pinkish to white corollas are five-lobed. The calyces, which bow out and often form purplish knobs at the sinuses, have 1- _ 2mm long, triangular teeth. The fruits consist of cap- | sules with 1-seeded chambers; the seeds become sticky _ when moistened. } | North American and Provincial Ranges Collomia tenella ranges from southwestern British Columbia, south in the western United States to Wyo- /ming, Utah, Nevada and Oregon (Hitchcock et al. ' h| 1959). In Canada, C. tenella is known only from the | Princeton area in British Columbia (Figure 2; Pojar | 1999; Douglas 2002a). Ficure |. Illustration of Collomia tenella (line drawing from Douglas et al. 1999b). ‘ Habitat | Collomia tenella occurs in the southwestern interi- (Hope et al. 1991). Climatic conditions in this region \ai of British Columbia, along the Similkameen River are continental, characterized by hot, dry summers, a _| valley in the Interior Douglas-fir biogeoclimatic zone fairly long growing season, and cool winters. A rain- | 1 i] { 1 ) * The field work for the Collomia tenella project was funded by the British Columbia Conservation Data Centre. The results appear in the | British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis fora Committee on the Status of Endangered Wildlife in Canada status report (Douglas and Penny 2003*) and the subsequent assessment of "ae (COSEWIC 2003). This paper includes more recent information that did not appear in the original status report. ‘" Taxonomy and nomenclature follow Douglas et al. (1998b, 1999a, 1999b, 2000, 2001). iS) 176 shadow effect prevails in this area due to the pres- ence of the Coast-Cascade Mountains to the west. Within this region, Collomia tenella occurs on an eroded, steeply-sloped, southeast-facing section of a sandy ridge (Figure 3). The sandy ridge, formed by flu- vial processes during recent glaciation, consists of fine- textured sands. The eroded sections of the slopes are sparsely vegetated with about 20% cover. The vegeta- tion consists of a variety of herbs and shrubs including Arrow-leaved Balsamroot (Balsamorhiza sagittata), Timber Milk-vetch (Astragalus miser), Narrow-leaved Collomia (Collomia linearis), Thread-leaved Phacelia (Phacelia linearis), Silky Lupine (Lupinus sericeus), Saskatoon (Amelanchier alnifolia), Dalmation Toad- flax (Linaria genistifolia ssp. dalmatica), Cheatgrass (Bromus tectorum), and Bluebunch Wheatgrass (Pseu- doroegneria spicata). Scattered Douglas-fir (Pseudot- suga menziesii) and Ponderosa Pine (Pinus ponderosa) also occur on the ridge (Figure 3). Biology There is no information available on the biology and ecology of Collomia tenella in the literature. Since it is an annual plant it appears to be absent in some years if environmental conditions for seed germination and/ or seedling survival are not favourable. Other annual members of the genus are self-compatible and self- pollinating (Wilken 1993). Seeds may be dispersed by animals since they are sticky when moistened. Population Attributes The single population of Collomia tenella, covering an area of less than 60 m?, was discovered in 1997 in the Similkameen River valley, west of Princeton, British Columbia (Figure 2; Douglas et al. 1998a). Counts for the population in 1997, 2000, 2002, 2003 and 2004 were 10, 1,0, 127 and 0, respectively. In 2003, the area occupied was ca. 56 square metres, with plants in four groups of 56, 67, 1 and 3 plants. These fluctuations indicate that seed germination and/or seedling success of this annual species is highly variable and thus, trends are difficult to determine. A fairly thorough search in adjacent areas in southern British Columbia have not revealed similar habitats or other populations of this species. Provincial, National and Global Ranks The British Columbia Conservation Data Centre has ranked Collomia tenella as S1 and placed it on the British Columbia Ministry of Environment Red list (Douglas et al. 2002a). This is the most critical cate- gory for imperilled rare native vascular plants in British Columbia. A rank of S1 is considered “critically imper- illed because of extreme rarity (5 or fewer occurrences or very few remaining individuals) or because of some factors making it especially vulnerable to extirpation or extinction” (Douglas et al. 2002a). Since the species is restricted in Canada to British Columbia, the Nation- THE CANADIAN FIELD-NATURALIST Vol. 120 } BRITISH ¢ — COLUMBIA FIGURE 2. Distribution of Collomia tenella in British Columbia. al rank is N1. Globally, C. tenella is ranked “G4?” or apparently secure but with a range of uncertainty which includes the possibility it is more vulnerable or conversely that is more widespread, abundant, and secure. Threats and Protection The major threats to Collomia tenella are through drilling for coalbed methane gas, sand removal for road construction, housing development and off-road recre- ational vehicles. In 2002, three test holes targeting coal- bed methane gas deposits were drilled in the Simil- 2006 DOUGLAS AND PENNY: SLENDER COLLOMIA 177 FIGURE 3. Collomia tenella habitat occurs on the eroded section near the top of this sandy ridge south of Princeton, British Columbia. Prominent trees on the ridge are Pseudotsuga menziesii and Pinus ponderosa. kameen area. If brought into production, the drilling, access roads and pumping stations could cause exten- sive degradation to the natural habitat in the area. Other potential threats result from activities that are permitted in the Agriculture Land Reserve. The private property where C. tenella occurs is in the Agricultural Land Reserve, where primary land use is agriculture. For instance, some types of fill/soil removal are allowable without application to the Agricultural Land Commission (Provincial Agricultural Land Com- mission 2003"). On these lands, development pressures may not appear to be an issue at this time. The Agri- cultural Land Reserve status may prevent subdivision development, but does allow other activities that could also potentially threaten the populations. In recent years many tracts of Agricultural Land Reserve land in southern British Columbia has been converted to housing developments, shopping malls and golf cours- es, either by decisions of the Agricultural Land Com- mission or very rarely by an ‘order in council’ by the sitting provincial legislature. Another threat likely to have a negative impact is recreational off-road vehicle use. The authors observed evidence of all-terrain vehicles and dirt bike use in the area of the sites in 2002. The steep and relatively unsta- ble slopes where C. tenella occurs are highly suscep- tible to disturbance but at the same time, present a desirable challenge for off-roaders. Although weeds are not a major threat to C. tenella populations, weed control activities do constitute a minor potential threat to populations. Under the Weed Control Act, an occupier must control noxious weeds growing or located on land and premises. Marginally specific chemical weed control compounds that kill broad-leaved plant species would likely also kill C. tenella. Noxious weeds such as Dalmation Toadflax (Linaria genistifolia var. dalmatica) and Cheatgrass (Bromus tectorum) occur in the C. tenella habitat and could increase. Currently the habitat is not highly in- vaded, but a variety of other introductions could also eventually take root there as they have in adjacent habi- tats in the surrounding landscape. Therefore, the eco- logical integrity of the C. tenella habitat could even- tually become as comprised as some of these other surrounding habitats are. An additional threat to the population of C. renella is its vulnerability to extirpation due to the extremely small occupied area and population size. Such small populations are at risk of inbreeding depression (Pri- mack 1998). Due to lack of genetic variation, these small populations are vulnerable to demographic and environmental variation. Furthermore, suitable habitats for C. tenella in the southern interior of British Colum- bia are extremely limited and thus, opportunities for colonisation are also limited. 178 Collomia tenella is on Schedule 1 of the federal Species at Risk Act. As part of its commitment to the National Accord (National Accord for the Protection of Species at Risk), the province is required to take measures to protect this species. A recovery strategy will be required under this commitment. Additionally, C. tenella is a potential candidate for listing under the provincial Wildlife Amendment Act. Evaluation The British Columbia Conservation Data Centre con- siders Collomia tenella to be Endangered in British Columbia (Douglas et al. 2002a) and the Committee on the Status of Endangered Wildlife in Canada has also assessed the species as Endangered (COSEWIC 2003*; Douglas 2005"). The single population is small and vulnerable to a number of threats. The small num- ber of plants also reduces the potential for genetic vari- ation which may be necessary to respond to environ- mental changes in the future. Without research on growth requirements and more demographic informa- tion, the stability of the present populations will remain unknown, but the population appears to be quite vul- nerable. Acknowledgments We thank Frank Lomer for providing information on his original discovery of Collomia tenella in 1997 and for his help in finding the site in 2002. Thanks also to Shyanne J. Smith who assisted with fieldwork in 2004. Documents Cited (marked * in text) COSEWIC. 2003. COSEWIC assessment and status report on Slender Collomia, Collomia tenella in Canada. Com- mittee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 13 pages. www.cosewic.gc.ca. Douglas, G. W., and J. L. Penny. 2003. COSEWIC Status report on Slender Collomia, Collomia tenella in Canada, in COSEWIC assessment and status report on Slender Collomia, Collomia tenella in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 13 pages. www.cosewic.gc.ca. Provincial Agricultural Land Commission. 2003. Agricul- tural Land Reserve — Frequently Asked Questions. Provin- cial Agricultural Land Commission, 133-4940 Canada Way, Burnaby, British Columbia V5G 4K6 Canada (Available at: http://www.alc.gov.bc.ca/faq/faq_alr.htm). Literature Cited Douglas, G. W., F. Lomer, and Hans L. Roemer. 1998a. New or rediscovered native vascular plant species in British Columbia. Canadian Field-Naturalist 112: 276-279. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare native vascular plants of British Columbia. Second edition. THE CANADIAN FIELD-NATURALIST Vol. 120 Province of British Columbia, Victoria, British Columbia. 358 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- trated flora of British Columbia. Volume 3. Dicotyledons (Diapensiaceae through Onagraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 423 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- trated flora of British Columbia. Volume 4. Dicotyledons (Orobanchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illus- trated flora of British Columbia. Volume 5. Dicotyledons (Salicaceae to Zygophyllaceae) and Pteridophytes. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 389 pages. Douglas, G.W., D. Meidinger, and J. Pojar 2001. Illustrat- ed flora of British Columbia. Volume 7. Monocotyledons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998b. Illustrated flora of British Columbia. Volume 1. Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1959. Vascular plants of the Pacific Northwest. Part 4: Ericaceae through Campanulaceae. University of Washington Press. Seattle, Washington. 510 pages. Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson, W. L. Harper, and B. M. Wikeem. 1991. Interior Dou- glas-fir zone. Pages 153-166 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Ministry of Forests Special Report Series (6), Victoria, British Columbia. 330 pages. Pojar, J. 1999. Polemoniaceae. Pages 36-60 in Illustrated flora of British Columbia.Volume 4. Dicotyledons (Oro- banchaceae through Rubiaceae). Edited by G. W. Douglas, D. Meidinger; and J. Pojar. British Columbia Ministry of Environment, Lands and Parks and British Columbia Min- istry of Forests, Victoria, British Columbia. 427 pages. Primack, R. B. 1998. Essentials of conservation biology. Sinauer Associates Inc., Sunderland, Massachusetts. Scoggan, H. J. 1979. The flora of Canada. Part 4. Dicotyle- doneae (Loasaceae to Compositae). National Museum of Natural Sciences Publications in Botany 7 (4): 1117-1710. Wilken, D. A. 1993. Collomia. Pages 825-826 in The Jepson manual: Higher plants of California. Edited by J. C. Hick- man. University of California Press, Berkeley, California. 1400 pages. Received 4 November 2002 Accepted 10 April 2006 Conservation Evaluation of the Small-flowered Tonella, Jonella tenella, in Canada’ GEORGE W. DouGLAs! AND JENIFER L. PENNY? ' Deceased. 2 Conservation Data Centre, Ministry of Environment, Ecosystems Branch, P.O. BOX 9993 St~ Prov Govt, Victoria British Columbia V8W 9R7 Canada Douglas, George W., and Jenifer L. Penny. 2006. Status of the Small-flowered Tonella, Jonella tenella (Scrophulariaceae) in Canada. Canadian Field-Naturalist 120(2): 179-182. In Canada, the Small-flowered Tonella, Tonella tenella, is restricted to the west side of Saltspring Island in the Gulf Islands of southwestern British Columbia. This population represents the northern limits of the species which is disjunct from its main range in southern Washington (Columbia River gorge), through Oregon to central California. In British Columbia, Jonella tenella is associated with rock outcrops and dry, steep, sparsely forested talus slopes at elevations of 50 to 300 m. The population on Saltspring Island is on private property and not directly imperilled at this time. There is, however, a potential for housing development in the future on this waterfront site, thus the authors consider the species endangered. Key Words: Small-flowered Tonella, Zonella tenella, endangered, distribution, population size, British Columbia. The Small-flowered Tonella, Tonella tenella (Benth.) Heller’, is a member of a genus of only two species occurring in western North America (Hitchcock et al. 1959). It is the only species found in British Columbia and Canada (Pojar 2000). Tonella tenella was first re- corded in Canada by Douglas and Ruyle-Douglas (1978). Tonella tenella is a slender, ascending to prostrate, annual herb from a delicate taproot (Figure 1; Pojar 2000). The smooth, often branched stems are 5-25 cm long with opposite leaves 1-2 cm long. The leaves are stalked, simple, ovate to round, and few-toothed or lobed below and unstalked and deeply 2-3 lobed above. The blue or white flowers are small, slightly zygomor- phic and long-stalked with deeply five-lobed calyces. The corollas are also five-lobed. There are four, epi- petalous stamens and one pistil with 2 ovules. Anthers are 0.4 mm long. The fruits are obtuse to globe-shaped capsules containing 2 to 4, 1-1.5 cm long, wingless seeds. Distribution The southwestern British Columbia occurrence of Tonella tenella is disjunct from its main range in south- ern Washington (Columbia River gorge), through Ore- gon to central California (Wetherwax 1993; Pojar 2000). In Canada, T. tenella is known only from the west side of Saltspring Island in the Gulf Islands of southwestern British Columbia (Figure 2; Pojar 2000; Douglas et al. 2002a, 2002b). . FiGureE 1. Illustration of Tonella tenella (Line drawing from Douglas et al. 2000) The field work for Tonella tenella was funded by the British Columbia Conservation Data Centre. The results appear in the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a Committee on the Status of Endangered Wildlife in Canada status report (Douglas and Penny 2003”) and the subsequent assessment of Endangered (COSEWIC 2003"). This paper includes more recent information that did not appear in the original status report. "Taxonomy and nomenclature follows Douglas et al. (1998a, 1998b, 1999a, 1999b, 2000, 2001) and Schofield (1992). 179 180 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Locations and sizes of Tonella tenella subpopulations on Saltspring Island, British Columbia. Collection site Last Observation 1- Upper slope, south 1976 2- Upper slope, north 2002 3- Mid-slope 2002 4-Lower slope 2002 5- ca. 10 m uphill from beach 2002 Habitat Tonella tenella sites in British Columbia are found in a climatic and floristic anomaly in coastal British Columbia, the dry Coastal Douglas-fir (Pseudotsuga menziesii) Zone (Nuszdorfer et al. 1991) of southeast- ern Vancouver Island. This area is in a rainshadow belt created by the Olympic Mountains to the south, result- ing in a relatively warm and dry Mediterranean climate. Within this region, Jonella tenella occurs on west- facing slopes on stable talus. This talus occurs in open Big-leaf Maple (Acer macrophyllum)-Arbutus (Arbutus menziesii) forests, where associates include Cleavers (Galium aparine), Little Western Bitter-cress (Car- damine oligosperma), Miner’s-lettuce (Claytonia per- foliata), Common Vetch (Vicia sativa), Barren Brome (Bromus sterilis), and Large-flowered Blue-eyed Mary (Collinsia grandiflora) or in open Douglas-fir — Arbu- tus — Garry Oak forests with Oregon Beaked Moss (Kindbergia oregana), heron’s-bill moss (Dicranum species), Harford’s Melic (Melica harfordii), Galium aparine, Upright Hedge-parsley (Torilis japonica) and Barren Brome (Bromus sterilis). Biology Species of Tonella, along with its sister genus, Blue- eyed Mary (Collinsia) of the tribe Collinsieae, are self- compatible annuals (Armbruster et al. 2002). Tonella tenella is considered a small-flowered type and accord- ing to Armbruster et al. (2002), large- and small-flow- ered taxa appear to have differences in timing of self- pollination. Large-flowered taxa maintain herkogamy (spatial separation of anthers and stigmas) early in anthesis by differential elongation of staminal filaments, while small-flowered taxa do not show this elongation pattern. As a result, large-flowered taxa experience a delay in self-pollination whereas in small-flowered taxa, anther-stigma contact and self-pollination occur early. Furthermore, the stigmas are receptive to pollen- tube growth early in Tonella species. Small-flowered populations are almost certainly autogamous (Arm- bruster et al. 2002). Low elevation pollinators for the tribe include the insects Bombus, Anthophora, Empho- ropsis, Synhalonia and Osmia (Armbruster et al. 2002). Population Attributes Four small subpopulations of Tonella tenella were recently confirmed in 2002 at the Saltspring Island site. These subpopulations, plus an earlier (1976) collection Collector Number of plants/area (m7) Douglas unknown Lomer 56+ /40.18 Lomer 100-150/15 Douglas 65+/-15/6 Lomer 30/1 record, occur in a narrow band extending up the moun- tainside for about 425 m. The subpopulations consist- ed of 6 to 356 plants with areas of 1 to about 40 m? (Table 1). Since the plant is inconspicuous and extreme- ly difficult to detect, it is quite likely other subpopu- lations occur on the slope. Provincial, National and Global Ranks Globally, Tonella tenella has a rank of G5 indicating that in most of its range the plant is common. Since the species is restricted to British Columbia it has a national rank of N1. Provincially, 7. tenella has been ranked as S1 by the Conservation Data Centre and appears on the British Columbia Ministry of Environ- ment Red List (Douglas et al. 2002a). This is the most critical rank that can be applied to species at the provin- cial level and indicates that the species is “critically imperiled because of extreme rarity (typically five or fewer occurrences or very few remaining individuals) or because of some factor(s) making it especially vul- nerable to extirpation or extinction”. Threats and Protection The most immediate threat to Jonella tenella in British Columbia is habitat destruction through a hous- ing development on waterfront private property. Sup- pression of both natural and human-induced fires in the last century may also have had an effect on the sur- vival of 7. tenella. The vegetation in this region would naturally be maintained by fires; however, in their absence, high fuel loads build and catastrophic fires could result. This may result in unsuitable conditions for T. tenella. Introduced species are also a problem. The vegetation characterizing the 7. tenella site has been altered by the introduction of European species. Although a large number of native forbs occur in these areas, much of the vegetation is dominated to a large extent by introduced species, particularly grasses, in- cluding a number of species of Bromus. The Tonella tenella site occurs on private land. Tonella tenella is not formally protected in British Columbia; however, it could be in the future since this species is a potential candidate for listing under the provincial Wildlife Amendment Act (2004). As part of its commitment to the National Accord (National Accord for the Protection of Species at Risk), the province is required to take measures to protect this species. It is on Schedule 1 of the federal Species At Risk Act and a recovery strategy will be required. 2006 Evaluation of Status The British Columbia Conservation Data Centre con- siders T. tonella to be Endangered in British Columbia (Douglas et al. 2002a). The Committee on the Status of Endangered Wildlife in Canada has also assigned this species to the Endangered category (COSEWIC 2003"). Only one extant population of Tonella tenella is known in British Columbia; thus, the loss of the one site would result in the loss of the species from British Columbia and Canada. Potential housing developments threaten 7. tenella in Canada since the one site occurs on private oceanside property. Acknowledgments We thank Frank Lomer for his assistance with field work and Marilyn Lambert for providing water trans- portation. Documents Cited (marked * in text) COSEWIC 2003. COSEWIC assessment and status report on the small-flowered tonella Tonella tenella in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. vii + 14 pages. Available at: www.sararegistry.gc. ca/status/status_e.cfm. Douglas, G. W., and J. L. Penny. 2003. COSEWIC status report on the small-flowered tonella Tonella tenella in Canada in COSEWIC assessment and status report on the small-flowered tonella Tonella tenella in Canada. Com- mittee on the Status of Endangered Wildlife in Canada. Ottawa. 1-14 pages. Literature Cited Armbruster, W. S., C. P. H. Mulder, B. G. Baldwin, S. Kalisz, B. Wessa, and H. Nute. 2002. Comparative analy- sis of late floral development and mating-system evolution in tribe Collinsieae (Scrophulariaceae s.1.). American Jour- nal of Botany 89: 37-49. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare native vascular plants of British Columbia. Second edition. Province of British Columbia, Victoria, British Columbia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- trated flora of British Columbia. Volume 3. Dicotyledons (Diapensiaceae through Onagraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 423 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- trated flora of British Columbia. Volume 4. Dicotyledons (Orobanchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Colum- bia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illustrated flora of British Columbia. Volume 5. Dicotyledons (Sali- caceae to Zygophyllaceae) and Pteridophytes. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 389 pages. Douglas, G. W., D. Meidinger, and J. Pojar 2001. Illustrat- ed flora of British Columbia. Volume 7. Monocotyledons (Orchidaceae to Zosteraceae). British Columbia Ministry DOUGLAS AND PENNY: SMALL-FLOWERED TONELLA 18] a) BRITISH ‘e COLUMBIA eS | ~™*©_f VICTORIA FIGURE 2. Distribution of Jonella tenella in British Columbia. of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas G. W., D. Meidinger, and J. Pojar. 2002b. Illustrated flora of British Columbia. Volume 8 — General Summary, Maps and Keys. British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 457 pages. Douglas, G. W, and G. Ruyle-Douglas. 1978. Contributions to the flora of British Columbia and the Yukon Territory I. Vascular plants. Canadian Journal of Botany 56: 2296-2302. Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. Illustrated flora of British Columbia. Volume 1. Gymno- sperms and Dicotyledons. (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998b. Illustrated flora of British Columbia. Volume 2. Dicotyledons. (Balsaminaceae through Cuscutaceae). Bni- tish Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 401 pages. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1959. Vascular plants of the Pacific North- west — Part 4: Ericaceae through Campanulaceae. Universi- ty of Washington Press, Seattle, Washington. 510 pages. 182 Nuszdorfer, F. C., K. Klinka, and D. A. Demarchi. 1991. Coastal Douglas-fir zone. Pages 95-112 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Ministry of Forests Special Report Series (6), Victoria, British Columbia. 330 pages. Pojar, J. 2000. Scrophulariaceae. Pages 114-218 in Illus- trated flora of British Columbia.Volume 5. Dicotyledons (Salicaceae through Zygophyllaceae) and Pteridophytes. Edited by G. W. Douglas, D. Meidinger, and J. Pojar. Bri- tish Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 427 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Schofield, W. B. 1992. Some common mosses of British Columbia. Royal British Columbia Museum Handbook. Royal British Columbia Museum, Victoria, British Colum- bia. 394 pages. Wetherwax, M. 1993. Tonella. Page 1063 in The Jepson manual: Higher plants of California. Edited by J.C. Hick- man. University of California Press, Berkeley, California. 1400 pages. Received 26 November 2002 Accepted 20 March 2006 | | | | | ) ] | ‘ Conservation Evaluation of Stoloniferous Pussytoes, Antennaria . . * flagellaris, in Canada GEORGE W. DouGLas!, JENIFER L. PENNY2, and KSENIA BARTON? ' Deceased. 2 Conservation Data Centre, British Columbia Ministry of Environment, Ecosystems Branch, P.O. Box 9993, Stn Prov Govt, Victoria, British Columbia V8W 9R7 Canada 3Current address: 204-4272 Albert Street, Burnaby, British Columbia V5C 2E8 Canada Douglas, George W., Jenifer L. Penny, and Ksenia Barton. 2006. Conservation evaluation of Stoloniferous Pussytoes, Anten- naria flagellaris, in Canada. Canadian Field-Naturalist 120(2): 183-187. In Canada, Stoloniferous Pussytoes, Antennaria flagellaris, is restricted to the Similkameen River valley south of Princeton, in south-western British Columbia. The three populations represent the northern limits of the species which ranges from south- western British Columbia, south in the western United States to Idaho, Wyoming, Nevada and California. In British Colum- bia, Antennaria flagellaris is associated with eroded, unstable, calcareous clay seepage slopes on open, southerly aspects. This habitat is infrequent in the Similkameen River area and the few existing plant populations could easily be extirpated through slight changes in drainage through drilling for coalbed methane gas, road-building, or housing development. Key Words: Stoloniferous Pussytoes, Antennaria flagellaris, endangered, distribution, population size, British Columbia. Stoloniferous Pussytoes, Antennaria flagellaris (A. Gray) A. Gray’ is a member of a genus of approximate- ly 35 species mainly of North America but also found in South America (Cronquist 1994). Fifteen species occur in British Columbia (Douglas 1998) and approx- imately 18 in Canada (Kartesz 1999). Antennaria fla- gellaris was first recorded in Canada by Douglas et al. (1998a). Antennaria flagellaris is a stoloniferous, short-lived perennial from a fibrous root (Figure 1; Douglas 1998). The stolons are up to 10 cm long while the few central flowering stems are up to 3.5 cm tall. The numerous basal leaves and the few stem leaves are linear to linear- wy oblanceolate, silky woolly-hairy and 1-3 cm long. The FS flower heads are terminal with involucres of two types. The female involucres are 7-13 mm tall bracts that are tinged brown or reddish-brown, thinly woolly-hairy below. The male involucres are 4-7 mm tall translucent bracts that are brownish at the tips. The female flowers are 5-7 mm tall while the male flowers are 3-4.5 mm tall. é The single-seeded achenes are elliptic, warty and 2-3 mm a long with a 6-8 mm tall pappus that has white, hairlike a (\ bristles. VAN A hl INIA 4 Distribution Antennaria flagellaris ranges from southwestern British Columbia, south in the western United States to Ficure 1. Illustration of Antennaria flagellaris (Line drawing Idaho, Wyoming, Nevada and California (Cronquist by Elizabeth J. Steven in Douglas et al. 1998, 2002b). “The field work for Antennaria flagellaris was funded by the British Columbia Conservation Data Centre. The results appear in the British | Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002a). This information formed the basis for a Com- 4 } | } ; ' | | ] Mittee on the Status of Endangered Wildlife in Canada status report (Douglas and Penny 2004") and the subsequent assessment of Endan- gered (COSEWIC 2004’). This paper includes more recent information that did not appear in the original status report. ‘Taxonomy and nomenclature follow Douglas et al. (1998b, 1999, 2001). 183 184 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Locations and Population Sizes for Antennaria flagellaris near Princeton, British Columbia. Collection site Last Observation 1. Stevenson Creek 1997 2. Whipsaw Creek, 2 km N of 1997 3. Stevenson Lake, 300 m NE of 2002 4. Tracey Lake 2003 1994). In Canada, A. flagellaris is known only from along a 3.2-km length of Highway #3 in the Similka- meen River valley south of Princeton in southwestern British Columbia (Figure 2; Douglas 1998; Douglas et al. 1998a, 2002a, b). Habitat Antennaria flagellaris occurs in the southern inte- rior of British Columbia along the Similkameen River valley in the lower montane zone in the Interior Dou- glas-fir biogeoclimatic zone (Hope et al. 1991). Cli- matic conditions in this region are continental, char- acterized by hot, dry summers, a fairly long growing season, and cool winters. A rainshadow effect prevails in this area due to the presence of the Coast-Cascade Mountains to the west. Within this zone, A. flagellaris occurs in calcareous, gravelly-clay soils or gravelly sands. The sites are characterized by a unique hydrol- ogy. The slopes, which are moderate (20-30%) with open southerly aspects, are eroded, unstable, and char- acterized by ephemeral seepage. These moist microsites are saturated by underground water in the winter and dry up in the summer. The slow down-slope movement that occurs as a result of the unusual hydrology is unique in the region and excludes many taxa which are not able to tolerate the conditions. As a result, the slope is disturbed and sparsely vegetated with A. flagellaris being the dominant component (Figure 3). Vegetation in the vicinity of the A. flagellaris sites is characterized by Big Sagebrush (Artemisia tridentata) shrub/grassland with scattered Ponderosa Pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii) [Nomenclature follows Douglas et al. 1998b, 1998c, 1999 and 2001]. The sites occur at the western edge of the distribution of open shrub/grassland at that ele- vation. Associates of A. flagellaris include Cushion Buckwheat (Eriogonum ovalifolium var. nivale), Par- snip-flowered Buckwheat (E. heracleoides var. angus- tifolium), Swale Desert-parsley (Lomatium ambiguum), Thread-leaved Sandwort (Arenaria capillaris ssp. amer- icana), Slender Wheatgrass (Elymus trachycaulus), One-spike Oatgrass (Danthonia unispicata), Common Rabbit-brush (Ericameria nauseosus var. speciosa), and Cut-leaved Daisy (Erigeron compositus var. glabra- tus). No non-native species were found in association with Antennaria flagellaris. Biology Little information is available on the biology of A. flagellaris in British Columbia. Only basic reproductive facts have been compiled about the species. Collector Number of plants/area (m7) Lomer 50 +3 or 4 Lomer Unknown Douglas and Penny 1 400 000 + 100 000/ 2100 Lomer 4-500/84 Antennaria flagellaris is a short-lived perennial, dioe- cious (male and female structures on separate plants) and wind-pollinated. This species produces seeds sex- ually by outcrossing (Bayer 1996). The fruits consist of single-seeded achenes. The numerous hair-like bristles of the mature achene facilitate its dispersal by wind. Antennaria flagellaris also reproduces vegetatively by producing stolons that terminate in plantlets. Ini- tially, the mother plant provides the plantlet with nutri- ents via the stolon. The genetically identical plantlets eventually become independent plants as stolons are severed. This mode of reproduction results in very re- stricted dispersal, as stolons only grow up to 10 cm long (Douglas 1998). Population Attributes There are three recently surveyed populations of Antennaria flagellaris in the Similkameen River valley, south of Princeton, British Columbia (Table 1). These populations occur along a 3.2-km length of Highway #3 (Figure 2). The populations range from a couple of square metres to over 2100 m/ in size and number from 50 to approximately 1 400 000 + 100 000 plants. Short- and long-term trends for these populations are unknown but can be expected to vary markedly due to differences in yearly plantlet survival, seed germi- nation and seedling success. Growing conditions on the ephemerally moist, clay slopes may vary from year to year according to normal climatic fluctuations affect- ing site hydrology. Provincial, National and Global Ranks Globally, Antennaria flagellaris has a rank of G5? indicating that in most of its range the plant is appar- ently secure to demonstrably widespread, abundant, and secure. Since the species is restricted to British Colum- bia it has a national rank in Canada of N1. Provincial- ly, Antennaria flagellaris has been ranked as S1 by the Conservation Data Centre and appears on the British Columbia Ministry of Environment Red list (Douglas et al. 2002a). This is the most critical rank that can be applied to species at the provincial level and indicates that the species is “critically imperiled because of ex- treme rarity (typically five or fewer occurrences or very few remaining individuals) or because of some factor(s) making it especially vulnerable to extirpation or extinc- tion”. Threats and Protection The major threat to populations of Antennaria fla- gellaris are through drilling for coalbed methane gas, 2006 road construction, housing developments and off-road recreational vehicles. In 2002 three test holes, target- ing coalbed methane gas deposits, were drilled in the Similkameen valley. If brought into production the drilling, access roads and pumping stations could cause major degradation in the area. The southern interior of British Columbia has recent- ly seen an increase in housing projects and the Prince- ton area is no exception. Any developments which even slightly change groundwater levels could be devastat- ing to this taxon. Furthermore, suitable habitats for A. flagellaris are extremely restricted in Canada, only occurring in the Princeton area, thus opportunities for colonisation are extremely limited. Although weeds are not a major threat to A. flagel- laris populations, weed control activities do constitute a minor potential threat to populations. Under the Weed Control Act, an occupier must control noxious weeds growing or located on land and premises; thus margin- ally specific chemical weed control compounds, that kill broad-leaved plant species, would likely be fatal to A. flagellaris. Recreational off-road vehicle use could also pres- ent a threat due to the nature of the steep, clay slopes where A. flagellaris occurs. When moist, this habitat would be highly susceptible to disturbance and at the same time, be at its most desirable to off-road users. Overuse of the habitat in this way could disturb the sites enough to radically alter the hydrological regime, and promote the establishment of non-native species. Other potential threats result from activities that are permitted in the Agriculture Land Reserve. The private property where A. flagellaris occurs is on the Agricul- tural Land Reserve, where primary land use is agri- culture. For instance, some types of fill/soil removal are allowable without application to the Agricultural Land Commission (Provincial Agricultural Land Com- mission 2003"). On these lands, development pressures may not appear to be an issue at this time. The Agri- cultural Land Reserve status may prevent subdivision development, but does allow other activities that could also potentially threaten the populations. In recent years many tracts of Agricultural Land Reserve land in south- ern British Columbia have been converted to housing developments, shopping malls, and golf courses, either by decisions of the Agricultural Land Commission or, very rarely, by an “order in council” by the sitting provincial legislature. An additional threat to the population of A. flagel- laris is its vulnerability to extirpation due to the ex- tremely small occupied area and population size. Such small populations are at risk of inbreeding depression (Primack 1998). Due to lack of genetic variation, these small populations are vulnerable to demographic and environmental variation. Furthermore, suitable habi- tats for C. flagellaris in the southern interior of British Columbia are extremely limited and thus opportunities for colonisation are also limited. DOUGLAS, PENNY, AND BARTON: STOLONIFEROUS PUSSYTOES 185 BRITISH COLUMBIA FiGuRE 2. Distribution of Antennaria flagellaris in British Columbia. Antennaria flagellaris is not formally protected in British Columbia, however, it could be in the future since this species is a potential candidate for listing under the provincial Wildlife Amendment Act (2004). As part of its commitment to the National Accord (National Accord for the Protection of Species at Risk), the province is required to take measures to protect this species. It is on Schedule | of the federal Species At Risk Act and a recovery strategy is expected to be com- pleted by the summer of 2006. 186 THE CANADIAN FIELD-NATURALIST Vol. 120 FIGURE 3. Open habitat of Antennaria flagellaris in the Princeton area. Antennaria flagellaris is abundant in the open areas between clumps of Eriogonum heracleoides var. angustifolium and Ericameria nauseosus Vat. speciosa. Evaluation The British Columbia Conservation Data Centre con- siders A. flagellaris to be Endangered in British Colum- bia (Douglas et al. 2002a). The Committee on the Sta- tus of Endangered Wildlife in Canada has also assigned this species to the Endangered category (COSEWIC 2004*). The three populations are small and vulnerable to a number of threats. With limited knowledge of the plants, biological and ecological requirements, this species is vulnerable to extirpation in British Columbia. Without research on growth requirements and further demographic information, the stability of the present populations will remain unknown. The limited number of plants also reduces the potential for genetic variation which may be necessary to respond to environmental changes in the future. Acknowledgments We thank Frank Lomer for providing information on his original discovery of A. flagellaris and for his assis- tance in relocating the sites in 2002 and 2003. Documents Cited (marked * in text) COSEWIC. 2004. COSEWIC assessment and status report on Stoloniferous Pussytoes, Antennaria flagellaris in Cana- da. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. Available at: http://www.sarareg istry.gc.ca/virtual_sara/files/cosewic/as%5Fstoloniferous% 5Fpussytoes%5Fe%2Epdf. Douglas, G. W., and J. L. Penny. 2004. COSEWIC Status report on Stoloniferous Pussytoes, Antennaria flagellaris, in Canada, in COSEWIC assessment and status report on Stoloniferous Pussytoes, Antennaria flagellaris in Cana- da. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 13 pages. Available at: http://www. sararegistry.gc.ca/virtual_sara/files/cosewic/sr%5Fstolon iferous%5Fpussytoes%5Fe%2Epdf. Provincial Agricultural Land Commission. 2003. Agricultur- al Land Reserve — Frequently Asked Questions. Provincial Agricultural Land Commission. 133-4940 Canada Way, Burnaby, British Columbia V5G 4K6 Canada. Available at: http://www.alc.gov.bc.ca/faq/faq_alr.htm. Literature Cited Bayer, R. J. 1996. Phylogenetic inferences in Antennaria (Asteraceae: Inuleae: Gnaphaliinae) based on sequences from the nuclear ribosomal DNA internal transcribed spac- ers (ITS). American Journal of Botany 83: 516-527. Cronquist, A. 1994. Intermountain flora — Vascular plants of the Intermountain West, U.S.A. Volume five: Asterales. New York Botanical Garden, New York, New York. 496 pages. Douglas, G. W. 1998. Asteraceae. Pages 96-392 in Illustrated flora of British Columbia. Volume 1. Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). Edited by G. W. Douglas, G. B. Straley, and D. Meidinger. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., F. Lomer, and H. L. Roemer. 1998a. New or rediscovered native vascular plant species in British Columbia. Canadian Field-Naturalist 112: 276-279. « 2006 Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare native vascular plants of British Columbia, Second edition. Province of British Columbia, Victoria, British Columbia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999. Illustrated flora of British Columbia. Volume 4. Dicotyledons (Oro- banchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar 2001. Illustrat- ed flora of British Columbia. Volume 7. Monocotyledons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas G. W., D. Meidinger, and J. Pojar. 2002b. Illus- trated flora of British Columbia. Volume 8 — General Sum- mary, Maps and Keys. British Columbia Ministry of Sus- tainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 457 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1998b. Illustrated flora of British Columbia. Volume 1. Gymno- sperms and Dicotyledons. (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. DOUGLAS, PENNY, AND BARTON: STOLONIFEROUS PUSSYTOES 187 Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998c, Illustrated flora of British Columbia. Volume 2 Dicotyledons. (Balsaminaceae through Cuscutaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 401 pages. Hope, G. D., W. R. Mitchell, D. A. Lloyd, W. R. Erickson, W. L. Harper, and B. M. Wikeem. 1991. Interior Douglas- fir zone. Pages 153-166 in Ecosystems of British Columbia. Edited by D. Meidinger and J. Pojar. British Columbia Min- istry of Forests Special Report Series Number 6, Victoria, British Columbia. 330 pages. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First Edition. /n Synthesis of the North American flora, Version 1.0. Edited by J.T. Kartesz and C. A. Meacham. North Carolina Botanical Gar- den, Chapel Hill, North Carolina. Primack, R. B. 1998. Essentials of conservation biology. Sin- aur Associates Inc., Sunderland, Massachusetts. Received 30 October 2002 Accepted 20 March 2006 Late-born Elk, Cervus elaphus, Calf Observed Near Bancroft, Ontario Rick C. Rosatte!? AND JOE NEUHOLD? ‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, DNA Complex, 2140 East Bank Drive, Peterborough, Ontario K9J 7B8 Canada 2 825 Wentworth St., Peterborough, Ontario K9J 8R7 Canada » 3Corresponding author: e-mail: rick.rosatte@ontario.ca Rosatte, Rick, and Joe Neuhold. 2006. Late-born Elk, Cervus elaphus, calf observed near Bancroft, Ontario. Canadian Field- Naturalist 120(2): 188-191. A recently born male Elk (Cervus elaphus) calf was observed and photographed approximately 30 km east of Bancroft, Ontario, on 5 October 2004. Based on the gestation period for Elk, the estimated conception date was between late Decem- ber 2003 and early January 2004. This is unusual as the majority of calves are born between late May and early June with conception having occurred in September/October. This reproductive strategy allows for an optimum growth period for calves to occur before the onset of winter which increases the probability of survival. An eleven-year-old cow Elk was iden- tified as the mother of the late born calf. The age of the cow, and her inability to meet the annual energy requirements neces- sary for normal reproduction, may have contributed to the late conception and parturition dates. Key Words: Elk, Cervus elaphus, Elk calf, restoration, late-born, Ontario. In 1998, the Ontario Ministry of Natural Resources (OMNR) and partner organizations implemented a plan to restore Elk (Cervus elaphus) to Ontario, Cana- da (Bellhouse and Broadfoot 1996). Between 1998 and 2001, 443 elk acquired from Elk Island National Park (EINP), Alberta, were released in four different areas of Ontario; Lake of the Woods, Lake Huron/North Shore (near Blind River), French River/Burwash, and Bancroft/North Hastings (Rosatte et al. 2002). In total, 120 Elk were released during 2000/2001 in the Ban- croft/North Hastings area at a location near Gin Lake (UTMC = 304000E 4990000N). Each summer, Elk calves have been observed and each winter, Elk calf surveys were conducted using OMNR helicopters to determine Elk calf survival and estimate the size of the herd. On 5 October 2004, a male Elk calf with spots, esti- mated to be about three weeks of age, was observed and video taped near Hartsmere, Ontario (UTMC= 300000E 4995000N), approximately 25 km east of Bancroft, Ontario (Figure 1). The calf weighed about 25-30 kg based on body size; average birth weights for Elk calves are 15-22 kg (Hudson et al. 2002). On 1 November 2004, the calf was again observed nurs- ing from a 11-year-old cow Elk which had been ear- tagged (All-Flex ear-tag # 347), weighed (238 kg), and radio-collared (Lotek GPS Collar- 148.115 Mhz) at EINP, Alberta, and released in the Bancroft/North Hastings, Ontario, area during April 2001. The calf was observed nursing from the cow during December 2004 and January 2005. The calf, still with spots, was pho- tographed again on | February 2005, near Hartsmere, Ontario (Figure 2). It was with a group of 38 Elk com- prised of 13 bulls {4 mature (>Syr), 4 yearlings, 5 im- mature (2-4 yrs), 8 calves, and 17 cows (which includ- ed the calf’s mother — Figure 3)}. Based on observation during January/February 2005 the late-born calf with spots was estimated to be 25%-35% smaller than the other calves (Figure 2). Those calves were of a size — comparable to Elk calves handled at EINP, Alberta, where calf weights averaged 120 kg (264 Ib) (n=16) during January 2001. These weights are comparable to 8-9 mo old Elk calves in other studies that were born during early spring and summer (Cook 2002; Peek 2003). Unfortunately, the spotted calf was killed on 25 February 2005. Based on tracks in the fresh snow, injuries to the calf, and blood on the antlers of a mature bull Elk, the calf was likely gored to death. Gestation periods for Elk range between 247-262 days with a mean of about 255 days (Bubenik 1982; Hudson et al. 2002). As this calf was first observed on 5 October 2004, conception must have occurred dur- — ing late December 2003 or early January 2004. Wishart (1981) reported a cow Elk in Alberta that was shot in September 1979 and had a fully developed fetus. In this case, the conception date was estimated to be Jan- uary 1979 and the cow was estimated to be 10 years of age. In Elk, reproductive success generally declines after 7 years of age (Raedeke et al. 2002). One adaptive strategy of Elk is the production of highly developed, rapid growing, large young, so that calves quickly reach a size that maximizes survival during winter (Geist 2002). The high energy require- ments for rapid growth of Elk calves is met through the energy acquired from maternal milk. Elk calves | acquire most of their nutritional requirements during | their first month of life from milk; however, after this ; neonatal period, requirements are greater and the inges- - tion of solid forage is required. That is why parturition throughout most of Elk range is early spring and sum- mer (late May/early June) when forage is high in pro- - tein and nutritious (Hudson et al. 2002). The timing | 188 2006 ROSATTE AND NEUHOLD: LATE-BORN ELK CALF 189 FiGurE 1. Late born elk calf with spots photographed by J. Neuhold near Hartsmere, Ontario, on October 5, 2004. of calving for Elk is such that inclement winter/spring weather is avoided but is early enough (e.g. June) so that calves can attain sufficient weight and size to sur- vive the winter (Hudson et al. 2002). Juvenile Elk have evolved to reduce or cease growth during fall and win- ter, when the nutritive value of forage declines. Cook et al. (1996) reported that calf growth in mid-Novem- ber was 50% of that found in September and hypoth- esized that the nutritional strategy of Elk calves during the late fall/early winter was primarily to maintain con- dition rather than maximizing growth. Inevitably, ener- gy acquisition for rapid growth becomes a problem for late-born calves, as not only is forage less nutri- tious during the fall and winter, but milk production in lactating cow Elk also declines during that period due to poor quality forage (Cook et al. 1996; Cook 2002). In addition, survival was found to be lower for late-born Elk calves in Yellowstone National Park than for earlier-born calves (Singer et al. 1997), and calf mortality during winter was primarily due to mal- nutrition. The calf in this study was observed nursing in mid-January 2005 and supplemental feeding by peo- ple in the area may have provided sufficient energy for the cow to continue milk production during the winter period. Nutrition and lactation can affect the timing of es- trous in Elk and subsequently the parturition date (Cook 2002). Generally, cows that are suckling a calf during the rut come into estrous later than cows that do not have a calf (Hudson et al. 2002). During the early part of rut, estrous approximates 21 days but is variable (19-25 days) (Bubenik 1982) and estrous length in- creases and becomes irregular during the later stages of the breeding period (Hudson et al. 2002). Three to four estrous periods during one breeding season are possible with a refractory period of 62 days between sequential estrous cycles (Bubenik 1982) and Elk can be in heat as late as February. Older aged ungulates ~ often cannot support production or reproduce (Cameron 1994; Cook 2002). Generally, reproductive success for elk 11 years of age is low (Raedeke et al. 2002). The age of the cow in this study, and her inability to meet the annual energy requirements necessary for normal reproduction, may have contributed to the late concep- tion and parturition dates. Acknowledgments The authors wish to thank the numerous partners of the Ontario Elk Restoration and Research Program: the Provincial Elk Technical Team, the Local Imple- mentation Committees, Cambrian College, the Rocky Mountain Elk Foundation, Elk Island National Park, the Ontario Federation of Anglers and Hunters, Trent University, Lakehead University, Laurentian University, 190 THE CANADIAN FIELD-NATURALIST Vol. 120 ai 1 1 PAl uf FIGURE 2. Late born elk calf with spots was about 25% smaller than other calves when photographed by R. Rosatte near Hartsmere, Ontario, 1 February 2005. = - : : = a fe Sil 20 EL ag pes TMT ES ea ay ee at ai ee ee a FIGURE 3. Elk calf with spots with 11-year old mother (on right) (photo by R. Rosatte near Hartsmere on 1 February 2005) ————E—————__ sr t—— a a a eS a 2006 University of Guelph, Sault College, Safari Club Inter- national (Ontario Chapter), Northern Ontario Tourist Outfitters, French River Resorts Association, Parks Canada, Northern Ontario Heritage Foundation, On- tario Fur Managers Federation, and the Ontario Minis- try of Natural Resources. C. Davies, manager, OMNR, Wildlife Research & Development Section, reviewed the manuscript and provided useful comments. Literature Cited Bellhouse, T., and J. Broadfoot. 1996. Plan for the restora- tion of elk in Ontario. Ontario Ministry of Natural Re- sources internal report, Ontario, Canada, 55 pages. Bubenik, A. 1982. Physiology. Pages 125-179 in Elk of North America: ecology and management. Edited by J. W. Thomas and D. Toweill. Stackpole Books, Harrisburg, Pennsylvania. Cameron, R. 1994. Reproductive pauses by female caribou. Journal of Mammalogy 75: 10-13. Cook, J. 2002. Nutrition and food. Pages 259-349 in North American elk: ecology and management. Edited by D. Toweill and J. W. Thomas. Smithsonian Institution Press, Washington, D.C. Cook, J., L. Quinlan, L. Irwin, L. Bryant, R. Riggs, and J. Thomas. 1996. Nutrition-growth relations of elk calves during late summer and fall. Journal of Wildlife Manage- ment 60: 528-541. ROSATTE AND NEUHOLD: LATE-BORN ELK CALF 19] Geist, V. 2002. Adaptive behavioral strategies. Pages 389-433 in North American elk: ecology and management. Edited by D. Toweill and J. W. Thomas. Smithsonian Institution Press, Washington, D.C. Hudson, R., J. Haigh, and A. Bubenik. 2002. Physical and physiological adaptations. Pages 199-257 in North Ameri- can elk: ecology and management. Edited by D. Toweill and J. W. Thomas. Smithsonian Institution Press, Washing- ton, D.C, Peek, J. 2003. Wapiti. Pages 877-888 in Wild Mammals of North America: Biology, Management and Conservation. Edited by G. Feldhamer, B. Thompson and J. Chapman. The Johns Hopkins University Press, Baltimore, Maryland. Raedeke, K., J. Millspaugh, and P. Clark. 2002. Population characteristics. Pages 449-491 in North American elk: ecology and management. Edited by D. Toweill and J. W. Thomas. Smithsonian Institution Press, Washington, D.C. Rosatte, R., J. Hamr, B. Ranta, J. Young, and N. Cool. 2002. Elk restoration in Ontario, Canada: Infectious dis- ease management strategy, 1998-2001. Annals of the New York Academy of Sciences 969: 358-363. Singer, F., A. Harting, K. Symonds, and M. Coughenour. 1997. Density dependence, compensation, and environ- mental effects on elk calf mortality in Yellowstone National Park. Journal of Wildlife Management 61: 12-25. Wishart, W. 1981. January conception in an elk in Alberta. Journal of Wildlife Management 45: 544. Received 17 February 2005 Accepted 7 October 2005 Annual Variation in Habitat Use by White-footed Mice, Peromyscus leucopus: The Effects of Forest Patch Size, Edge and Surrounding Vegetation Type CHRISTINE S. ANDERSON!, DOUGLAS B. MEIKLE’, ALAN B. Capy”, and ROBERT L. SCHAEFER? ' Present address: Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, Columbus, Ohio 43210 USA. e-mail: anderson.1309@osu.edu; formerly C. Schandorsky; corresponding author. "Department of Zoology, Miami University, Oxford, Ohio 45056 USA 3Department of Mathematics and Statistics, Miami University, Oxford, Ohio 45056 USA Anderson, Christine S., Douglas B. Meikle, Alan B. Cady, and Robert L. Schaefer. 2006. Annual variation in habitat use by White- footed Mice, Peromyscus leucopus: The effects of forest patch size, edge and surrounding vegetation type. Canadian Field-Naturalist 120(2): 192-198. White-footed Mice (Peromyscus leucopus) were trapped for two years in the exterior matrix, edge, and interior forest habitat sections of six forests patches in a fragmented agricultural landscape. We used data on the capture locations of P. leucopus individuals from the two years, which differed in rainfall (1.e., summer of 2000 with 50% more rain than summer of 1999), to assess how patch size, edge habitat, and surrounding habitat type influence habitat use and movements in populations of this forest habitat generalist. We found that the proportion of individuals subsequently captured in the forest edge from the exterior was 16 times greater in the wet year than in the dry year and approximately twice as many P. leucopus were not subsequently recaptured from the exterior matrix in the dry year compared to the wet year. For each year, captures between habitats did not differ in relation to patch size, edge forest habitat, or exterior matrix type. These results illustrate the generalist habitat pref- erences of P. leucopus, but emphasize annual variation in their behavior and distribution. Key Words: White-footed Mice, Peromyscus leucopus, forest patches, habitat preference, edge habitat, exterior matrix, Ohio, Indiana. Human activities have led to a reduction and frag- mentation of secondary-growth deciduous forests in North America, as well as changes in the quality of the remaining habitat (Yahner 1988; Fahrig 1997). The effects of fragmentation include those related to loss of connectivity among remnant patches, and increases in edge habitat, which both can cause local changes in abundance, distribution, and species’ persistence (An- dren 1994, 1997; Murcia 1995). In addition, the frag- mentation of forested habitat results in the creation of forest patches that vary in size. Studies have shown that there are taxonomic differences in the relationship be- tween animal population densities and patch area. Insects and birds generally show a positive relationship, likely due to an avoidance of edge habitat, which is greater in proportion in smaller patches (Connor et al. 2000). The majority of mammal species show fairly constant population densities with patch area, this follows the theory of island biogeography (Bowers and Matter 1997). In contrast, investigators have found a consistent negative relationship between density of the White-foot- ed Mouse (Peromyscus leucopus Rafinesque), a forest habitat generalist, and forest patch area (Nupp and Swihart 1996, 1998; Krohne and Hoch 1999; Mossman and Waser 2001; Schmid-Holmes and Drickamer 2001; Anderson et al. 2003; Anderson and Meikle 2006). Although resource availability may influence popula- tion dynamics of small mammals (e.g., Nupp and Swi- hart 1996), there are other factors that may be respon- sible for the negative density-area relationship. One possibility is that dispersal is inhibited from smaller forest patches due to lower habitat quality in the matrix than under continuous forest cover (Brown and Lit- vaitis 1995; Nupp and Swihart 1998; Krohne and Hoch 1999). We hypothesize in the current study that if inhib- ited dispersal results in the negative density-area rela- tionship, then fewer P. leucopus should tend to move from the forest edge to the agricultural fields surround- ing smaller compared to those surrounding larger for- est patches. In addition to patch area, another consequence of forest fragmentation is an increase in the proportion of edge habitat in the landscape since smaller forest patches have a greater edge: interior ratio than larger forest patches. Forest edges usually exhibit differences in microclimate, species composition, and vegetation structure compared to forest interiors (Harris 1988; Saunders et al. 1991). These difference may influence abundance and distribution of animal species. Studies have found that some forest animals may prefer for- est edges while others may avoid those areas (Harris 1988; Saunders et al. 1991; Murcia 1995). However, there is debate as to whether P. /eucopus prefers for- est edge to forest interior habitat. Some investigators have reported greater densities of Peromyscus spp. near edges (Cummings and Vessey 1994; Boyne and Hobson 1998; Manson et al. 1999), possibly because 192 = 2006 vegetation in edge habitat has been shown to be struc- turally more complex compared to interior habitat (Cummings and Vessey 1994; Anderson et al. 2003). In contrast, others have found fewer mice and lower maternal survival and litter success in edge habitats, presumably because parasitism, predation and inter- specific competition can be higher along edges (Mat- ter et al. 1996; Morris and Davidson 2000; Wolf and Batzli 2001, 2004). Since there is no evidence to date of higher rates of predation on mice in forest edges, then we predict that mice may prefer the high quality vege- tation in forest edges such that there would be higher recapture rates in forest edges than in forest interiors. Another consequence of forest fragmentation is that forest patches are surrounded by a matrix of agriculture or residential development that may inhibit the move- ment of organisms (Fitzgibbon 1997; Collinge 1996). Even though the agricultural matrix surrounding forest patches may be considered “hostile” habitat for some species (Collinge 1996), generalists such as P. leuco- pus (Gottschang 1981) may disperse relatively easily (Lackey 1978) through row crop fields (Mossman and Waser 2001). However, it is not well understood whe- ther matrix types (e.g., corn or pasture) differentially affect capture rates of White-footed Mice in the exterior matrix. M’Closkey and Lajoie’s (1975) work suggests that P. leucopus may prefer corn over pasture because mice were absent from grasslands. Even though grass- es may provide cover for mice from aerial predators, Getz (1961) also noted that individual Peromyscus avoided grassy sites under forest canopy (also see M’Closkey and Lajoie 1975) likely because it rustles as the mouse moves (Barnum et al. 1992). If surrounding habitat type influences movements of mice through- out the landscape, we predict that they may stay in or move into the surrounding matrix habitat at greater rates when the matrix is comprised of corn rather than pasture. The purpose of this study was to use data on the first two captures of P. leucopus individuals in differ- ent habitats (i.e., exterior matrix, forest edge, and forest interior habitat) in small and large forest patch- es for two summers to test the hypotheses related to patch size, edge habitat, and surrounding habitat type as described above. In addition, we compared captures among habitats between two summers differing in rainfall amounts (i.e., one dry year and one wet year) because temporal variation in precipitation regulates rodent population dynamics (Lewellen and Vessey 1998). Utilizing data on habitat use instead of overall abundance, we assumed that if mice tended to prefer one habitat over another, then, on average, they would move into (or stay in) the preferred habitats in rela- tively high proportions. Since much of the work on the effects of forest fragmentation has been focused on birds, this study contributes to our understanding of how human-dominated landscapes influence habitat use and movements of populations of small mammals. ANDERSON, MEIKLE, CADY, and SCHAEFER: WHITE-FOOTED MICE 193 Methods Study Area Peromyscus leucopus were trapped in both 1999 and 2000 in the same three “small” (~S-9 ha) and three “large” (~1 10-150 ha) forest patches in a row-crop agri- cultural landscape covering approximately 144 km* in SW Ohio and SE Indiana, USA (study area, 39°S1'N, 84°74'W; Anderson and Meikle 2006; Table 1). We analyzed data from four transects of six Sherman™ live traps (7.5 x 7.5 x 25.5 cm) that were placed per- pendicular to the center of one edge of each patch, with each transect beginning 20 m exterior to and ex- tending 30 m into the patch (Figure |). The transects and the traps in each transect were spaced 10 m apart, resulting in a 30 m by 50 m grid with a total of 24 traps at each forest patch. The eight traps in the agricultural field outside of each forest patch were considered “ex- terior” and the next eight traps were considered “edge”. The eight traps inside the forest patch were consid- ered “interior” traps. Based on our previous findings (Anderson et al. 2003) and corroborated by Burke and Nol (1998), the width of the edge habitat was defined based on typical edge floristic features. The surround- ing habitat type in the exterior matrix was either pas- ture or corn fields (Table 1). Data collection Traps were set on 26 nights in each forest patch be- tween June and September in 1999 and 2000 for a total of 7488 trap-nights. When a mouse was captured ini- tially it was lightly anesthetized with halothane and a 12 mm AVID® microchip (PIT tag) was injected under the scruff of its neck. To distinguish between P. leu- copus (primarily a woodland species) and P. manicu- latus (primarily a grassland species), which are mor- phologically similar and overlap in the nonforested habitat within the study area, saliva samples were ana- lyzed for the migration of the amylase enzyme with polyacrylamide gel electrophoresis (Aquadro and Pat- ton 1980; Bruseo et al. 1999). We followed the proto- col of Bruseo et al. (1999) with minor modifications. Each patch was trapped for two consecutive nights every 5-7 d and we recorded the date, identification number and location of every individual at every trap- ping event. Based on trap location we determined which habitat (i.e., exterior, edge, or interior) each individ- ual was located in for its first capture and then where it was subsequently captured for its second capture event. It is possible that the length of time between any two capture events for a particular individual could be 24 hours apart or more. A fourth category for the sub- sequent capture was “not subsequently recaptured”, which was used if the individual was not recaptured again after tagging. Only the first two capture events for each individual were analyzed so that each individ- ual was included only once in the analysis to avoid pseudoreplication. In order to verify whether a mouse moving into a particular habitat reflects fidelity to this THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Description of study sites and their location, size, and surrounding habitat type (i.e., exterior matrix habitat) in 1999 and 2000. Patch names Location Small patches ERC (Ecology Research Station) Oxford, Ohio Indiana Bath, Indiana Jericho Oxford, Ohio Large patches Bachelor Woods and Game Preserve Oxford, Ohio Hueston Woods State Park Reily Township Reily, Ohio Preble/Butler Counties, Ohio Surrounding habitat type Size (ha) 1999 2000 5 pasture pasture 8 com com 9 com com 150 pasture pasture 200 com pasture 110 corm soybeans* *This patch was not included for tests involving surrounding habitat type since it was the only patch with soybeans. habitat shift, we also analyzed the data for the first year using all captures of all mice instead of the first two capture events only (i.e., 188 mice captured a total of 901 times) and found similar results (not presented), suggesting a permanence of habitat shifts. Although over winter survival was very low (i.e., four known individuals) between the two years of this study (An- derson 2004), we did not include those four individuals in the data set for the second year. Rainfall amounts from the Ohio Agricultural Research & Development Center (OARDC) weather station on Miami Universi- ty’s Oxford campus (http://jrscience.wcp.muohio.edu/ van tagepro/latestweather.html) were totaled for each summer month (June — August) in 1999 and 2000. The OARDC is located within the center of the study area. Statistical analyses Data on the proportion of mice trapped for their sec- ond capture in the exterior, edge, or interior habitats, or not subsequently recaptured, based on the three initial capture habitats (i.e., exterior, edge, or interior) were compiled (see Table 2). The data set was analyzed with PROC CATMOD using chi-square tests to determine whether captures among habitats differed in relation to patch size, surrounding matrix type, and year (SAS 1999-2000). Subsets of the data were analyzed with PROC FREQ using either chi-square tests or Fisher’s exact tests when sample size was small (n < 50 or cells Agricultural field Forest Interior Ficure |. The trapping grid design for each of the forest patch- es in this study. The letter “O” denotes each outside (1.e., exterior) trap, the letter “E” represents the eight edge traps, and the “I” denotes the interior trap loca- tions. with values <5; SAS 1999-2000). Data from each year were analyzed separately (see Results; Table 2). In post hoc analyses, Fisher’s exact tests and chi-square tests were used where appropriate (Moore and McCabe 1993) with a Bonferroni adjustment to the level of sig- TABLE 2. The habitat where each Peromyscus leucopus was trapped for its second capture based on where it was first cap- tured in 1999 and 2000. Values are pooled across patch sizes and number of individuals and row percentages are given in parentheses. Habitat where first captured Habitat where subsequently (i.e., second) trapped 1999 Exterior Edge Interior Not subsequently recaptured Exterior 9 (28%) tm GZ) 2 (6%) 20 (63%) Edge 6 (14%) 19 (43%) 10 (23%) 9 (21%) Interior 2 (4%) 8 (15%) 17 61%) 27 (50%) 2000 Exterior Edge Interior Not subsequently recaptured Exterior 1 (8%) 6 (50%) 1 (8%) 4 (33%) Edge ily 1G) 18 (46%) 10 (26%) 10 (26%) Interior 0 (0%) 7 (13%) 23 (41%) 26 (46%) | | : | } ' | | | 2006 nificance at & = 0.05 (Sokal and Rohlf 1981). An un- paired t-test was used to test for a difference in mean monthly rainfall in the summer (June — August) of 1999 compared to 2000 (Sokal and Rohlf 1981). Results Patch size did not influence captures of P. leucopus among habitats in either 1999 or 2000 (1999: x7, = 0.9, P = 0.8; 2000: y°, = 4.7, P = 0.2). Hence, there was no difference in captures of mice among different habi- tats (i.e., exterior matrix, edge, or interior habitats, or not subsequently recaptured) for those individuals that had been initially trapped in edge habitat in small com- pared to large patches in either year (1999: Fisher’s exact test, n = 44, P = 0.5; 2000: Fisher’s exact test, n= 39, P=0.7). Data were pooled across patch sizes within each year for all other analyses (Table 2). The proportion of P. leucopus recaptured in edge habitat (i.e., both the first and second captures occurred in edge habitat) compared to the proportion recaptured in inte- rior habitat did not differ in either year (1999: oa = 1.4) P=0.2; 2000: x7, = 0.2, P = 0.6), suggesting that mice did not show a preference for edge or interior forest habitat. Surrounding habitat type (i.e., corn or pasture) did not influence the proportion of mice subsequently captured across different habitats that had been first captured in the edge in either year (1999: Fisher’s exact test, nm = 44, P = 0.5; 2000: Fisher’s exact test, n = 33, P = 0.2). Likewise, surrounding habitat type did not affect subsequent captures across habitats for mice that had been first captured in the exterior matrix habitat in either year (1999: Fisher’s exact test, n = 32, P=0.3; 2000: Fisher’s exact test, n = 10, P=0.3). When captures between habitats in 1999 and 2000 were compared, P. /Jeucopus first captured in the exte- rior were subsequently trapped in different proportions among habitats in 1999 compared to 2000 (Fisher’s exact test, = 44, P = 0.002); specifically, the propor- tion of individuals subsequently captured in the forest edge from the exterior was 16 times greater in 2000 (6/12 = 50%) than in 1999 (1/32 = 3%; Fisher’s exact test,n = 44, P=0.0008; Table 2, Figure 2a). Likewise, a greater proportion of mice stayed in the exterior matrix compared to moving into the forest (edge and interior combined) in 1999 (9/12) than in 2000 (1/7; Fisher’s exact test, n = 20, P = 0.02). Approximately twice as many P. leucopus were not subsequently re- captured after their first capture in the exterior matrix in 1999 (20/32 = 63%) compared to 2000 (4/12 = 33%), although the test was not significant (Fisher’s exact test,n = 44, P=0.1; Figure 2b). Mean monthly rain- fall amounts were over 50% higher in the summer (June-August) of 2000 (mean + SE = 11.8 + 0.9 cm) compared to 1999 (5.3 + 1.0 cm; t,= 4.7, P = 0.009). Discussion Patch size Studies have shown that populations of P. leuco- pus, unlike birds, insects, and most other mammals, ANDERSON, MEIKLE, CADY, and SCHAEFER: WHITE-FOOTED MICE 195 100 @ Subsequently 80 trapped in exterior interior, of not subsequently 60 recaptured 40 @ Subsequently | trapped in edge 20 0 b 100 gw Subsequently trapped in extenor edge, or interior @ Not subsequently | recaptured Percentage of mice initially captured in exterior matrix 0 aa [hoo 199 2000 Year FIGURE 2. Percentage of mice (a) subsequently captured in forest edge habitat in 1999 compared to 2000 and (b) not subsequently recaptured in 1999 compared to 2000 (see Table 2 for sample sizes) that had been initially trapped in the exterior matrix habitat. have greater densities in smaller forest patches (Nupp and Swihart 1998; Krohne and Hoch 1999; Anderson et al. 2003). One proposed mechanism to account for this negative density-area relationship is that habitat quality is lower in the exterior matrix than in woodland sites (Nupp and Swihart 1996; Krohne and Hoch 1999). Predation pressure may be elevated in the agricultural fields surrounding smaller patches because some pred- ators may attain higher densities in a landscape with a variety of land uses (Brown and Litvaitis 1995). We predicted that fewer mice would move from the forest edge to the agricultural fields surrounding smaller than larger forest patches, but our results did not support the inhibited dispersal hypothesis. Diaz et al. (1999) found no difference in movements of Apodemus syl- vaticus to the exterior in relation to patch size, where- as Krohne and Hoch (1999) found that P. leucopus populations in Indiana had lower dispersal rates from smaller patches, a factor that may have resulted in higher densities in those patches (Adler and Levins 1994). Nupp and Swihart (1996) concluded that both reduced dispersal and greater abundances of food con- tributed to the high densities of P. leucopus they ob- served in small forest woodlots. 196 It is unclear why densities of P. leucopus tend to be higher in small forest patches (Anderson and Meikle 2006). One possibility is that immigration rates are in fact greater into smaller forest patches, resulting in higher densities in those patches (Bowman et al. 2002). However, we did not find that movements from exterior to the edge or interior habitats were greater in smaller than larger patches in this study. If smaller and larger forest patches do not vary in emigration and immigra- tion rates, as our data suggest, then a second possibility is that other factors, such as greater food abundance or cover from predators, may have resulted in greater abun- dances of mice in smaller patches or in particular the edges of small patches (Nupp and Swihart 1996; Diaz et al. 1999: Anderson et al. 2003; Anderson and Meikle 2006). A third possibility is that lower densities in larg- er patches may be the result of mice dispersing to low density regions within the heterogeneous forest patch to alleviate high densities, with fewer sites likely pres- ent in smaller forest patches (Krohne and Burgin 1990; Krohne and Hoch 1999). Edge habitat Differences in species composition and vegetation structure in forest edges compared to forest interiors, regardless of patch size; may influence the distribution of small mammals within forest fragments (Matlack 1994; Collinge 1996). Currently, there are conflicting reports about the effects of edge habitat on the abun- dance and distribution of White-footed Mice (e.g., Cummings and Vessey 1994; Anderson et al. 2003; Wolf and Batzli 2004). We found that P. leucopus were recaptured in the same proportion in forest edge com- pared to forest interior habitat in either year. Our data on initial and second capture habitat locations support those studies that reported no difference in the relative abundance of mice in those two habitats (Heske 1995; Nupp and Swihart 1996, 1998). This result is somewhat surprising since forest edge habitats have been shown to have greater structural complexity in the understory vegetation compared to edge habitat (Anderson et al. 2003) and White-footed Mice have been shown to prefer areas with more plant cover (Drickamer 1990). One explanation could be that mice are not moving into edge habitat but are already present in high abun- dances with small territory sizes. Perhaps once territo- ries are established in early spring, when movements among high (i.e., edge) and low quality (i.e., interior) habitats are negligible due to competitive exclusion. Surrounding habitat type Types of vegetation in the surrounding matrix habi- tat may differentially affect the population dynamics of rodent species in a fragmented landscape. For exam- ple, Wolf and Batzli (2004) found that predation risk for P. leucopus was higher at edges adjacent to re- stored prairies than at edges adjacent to agricultural fields. Our findings showed that P. leucopus moved in similar proportions from the forest edge into two dif- ferent matrix habitat types (i.e., corn and pasture). This THE CANADIAN FIELD-NATURALIST Vol. 120 is in contrast to studies that have found that P. leucopus tend to avoid grassy habitats and can be found in corn fields at densities as high as those in woodlots (Getz 1961; M’Closkey and Lajoie 1975; Wegner and Mer- riam 1979, 1990). Mice in our study sites may have simply responded to the seasonal presence of cover and not to the type of cover (Cummings and Vessey 1994). Henein et al. (1998) designed a simulation model illus- trating flexible and opportunistic responses of P. leu- copus to landscape structure and connectivity via an expansion of their use of corn and small grain fields. Similar results have been found for A. sylvaticus, a mouse species also described as a habitat generalist (Garcia et al. 1998; Kozakiewicz et al. 1999). Annual variation Rainfall may impact rodent populations directly by influencing their reproductive status and survival, or indirectly in relation to changes in food sources and vegetative cover from predators (Kaufman et al. 1995; Kesner and Linzey 1997). Monthly rainfall data showed that totals in the summer of 2000 were almost 50% greater than in 1999. For mice initially captured in the exterior matrix habitat, we found that a greater propor- tion moved to the edge habitat and more were subse- quently recaptured in the exterior matrix in the wet year compared to the dry year. If rainfall influenced patterns of capture among habitats, one explanation is that in 1999 (i.e., the dry summer) a higher proportion of mice were not subsequently recaptured from the ex- terior matrix habitat because they died due to poor con- ditions, or moved throughout the landscape in search of other better quality habitats (Kesner and Linzey 1997; Lidicker and Stenseth 1992). In 2000, the shrub and herbaceous layers in the forest habitat may have had more or higher-quality food sources compared to the exterior matrix as a consequence of rainfall which caused an influx of mice to the forest habitat (Parmenter and MacMahon 1983; Bowers and Dooley 1999; Hut- chinson et al. 1999). Our results illustrate the gener- alist habitat preferences of P. leucopus but emphasize annual variation in their distribution within the forest and surrounding habitats. However, year-to-year dif- ferences in patterns of habitat use may be influenced by a number of variables, and a longer-term study would help to elucidate those factors (Kesner and Linzey 1997): Acknowledgments We thank the landowners for permission to use study sites, in addition to the Ohio Department of Natural Resources, Hueston Woods State Park, Miami University’s Natural Areas Committee and Miami Uni- versity’s Ecology Research Center. We thank J. Bruseo, M. Spritzer, S. Vessey, and three anonymous reviewers for their valuable advice. K. Fielitz, J. Gliha, C. Har- rison, R. Lim, A. Lohrey, D. Markert, M. Tarbox, K. Wolfe and especially R. Young provided valuable sup- port in the laboratory and in the field. 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Changes in wildlife communities near edges. Conservation Biology 2: 333-339. Received 13 May 2005 Accepted 12 December 2006 Response of the Sea Louse Lepeophtheirus salmonis Infestation Levels on Juvenile Wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, Salmon to Arrival of Parasitized Wild Adult Pink Salmon ALEXANDRA Morton! and RoB WILLIAMS2 ' Raincoast Research Society, 13 Cramer Pass, Simoom Sound, British Columbia VOP 1SO Canada email: wildorca@island.net 2 Pearse Island, Box 193, Alert Bay, British Columbia VON 1A0 Canada Morton, Alexandra, and Rob Williams. 2006. Response of the sea louse Lepeophtheirus salmonis infestation levels on juve- nile wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, salmon to arrival of parasitized wild adult Pink Salmon. Canadian Field-Naturalist 120(2): 199-204. Recent recurring infestations of Sea Lice, Lepeophtheirus salmonis, on juvenile Pacific salmon (Oncorhynchus spp.) and subsequent annual declines of these stocks have made it imperative to identify the source of Sea Lice. While several studies now identify farm salmon populations as sources of Sea Louse larvae, it is unclear to what extent wild salmonid hosts also contribute Sea Lice. We measured Sea Louse numbers on adult Pink Salmon (Oncorhynchus gorbuscha) migrating inshore. We also measured Sea Louse numbers on wild juvenile Pink and Chum salmon (Oncorhynchus keta) migrating to sea before the adults returned, and as the two age cohorts mingled. Adult Pink Salmon carried an average of 9.89 (SE 0.90) gravid lice per fish, and thus were capable of infecting the adjacent juveniles. Salinity and temperature remained favourable to Sea Louse reproduction throughout the study. However, all accepted measures of Sea Louse infestation failed to show significant increase on the juvenile salmon, either in overall abundance of Sea Lice or of the initial infective-stage juvenile lice, while the adult wild salmon were present in the study area. This study suggests that even during periods of peak inter- action, wild adult salmon are not the primary source of the recent and unprecedented infestations of Sea Lice on juvenile Pacific Pink and Chum salmon in the inshore waters of British Columbia. Key Words: Sea Lice, Lepeophtheirus salmonis Pink Salmon, Oncorhynchus gorbuscha Chum Salmon, O. keta, salmon farm, British Columbia The sea louse Lepeophtheirus salmonis is a common salmon-specific parasite (Nagasawa et al. 1993; Kaba- ta 1973). While their preferred Pacific host is the Pink Salmon (Oncorhynchus gorbuscha) (Nagasawa 1987; Nagasawa et al. 1993), L. salmonis is found on all species of Pacific salmonids, as well as Atlantic Salmon (Salmo salar), Sea Trout (Salmo trutta trutta), Rain- bow Trout (Salmo gairdneri) and Artic Charr (Salveli- nus alpinus). Once considered benign (Boxshall 1974; Nagasawa 1987; Berland 1993), L. salmonis is now a significant problem species on farmed salmon (Pike and Wadsworth 1999; Johnson et al. 2004). Salmon farms are a series of floating net pens attached to the shoreline or anchored in the nearshore. Salmon arrive from hatcheries and are fed in these pens for 18-24 months, then harvested and sold as food. All farm salmon in our study area were Atlantic Salmon. Optimal salinity for L. salmonis survivorship and reproduction is 30%o (Johnson and Albright 1991b), with copepodids tolerating salinities as low as 15%c (Pike and Wadsworth 1999). L. salmonis hatch into a free-swimming naupliar stage directly from egg strings attached to the female’s body (Johnson and Albright 1991a). The interval from hatching to infective capa- bility (copepodid stage) is approximately 4 d at 10°C and 2d at 15°C (Johnson and Albright 1991b). The copepodid stage is the first attached stage, thus abun- dant presence of this stage signals a local source of gravid Sea Lice and a host salmonid population. When wild salmon enter freshwater to spawn, the attached L. salmonis experience decay of life-processes leading to death (Hahnenkamp and Fyhn 1985; McLean et al. 1990; Johnson and Albright 1991b). As one would expect from an obligate parasite, there have been no reports of L. salmonis over-wintering off-host in the nearshore marine environment. While early studies on juvenile wild Atlantic sal- monids Salmo salar and Salmo trutta make no mention of parasitism by L. salmonis (Heuch and Mo 2001): Sea Lice are now reported in high numbers on wild European salmonids adjacent to farms (Tully et al. 1993; Birkeland 1996). Historically, researchers did not look specifically for L. salmonis on juvenile Pacific salmon in the inshore environment. However, the sem- inal works on juvenile Pink and Chum salmon (Healey 1978, 1982; Parker 1965; Parker and Vanstone 1966) did note the presence of the smaller sea louse, Caligus clemensi (Parker and Margolis 1964). C. clemensi is a generalist non salmon-specific parasite that has been reported on 13 species of fish (Jones and Nemec 2004’). It seems reasonable that if Caligus was noted, then the larger, more conspicuous L. salmonis would also have been reported had it been historically present on juve- nile Pink salmon. Today, Sea Lice are reported on juvenile Pink and Chum salmon in some areas of coastal British Colum- IP) 200 bia and appear associated with salmon farming (Mor- ton and Williams 2003; Morton et al. 2004; Morton et al. 2005; Krkosek et al. 2005; Krkosek et al. 2006). The impact of Sea Lice is host-size dependant (Grimnes and Jakobsen 1996; Bjgrn and Finstad 1997) and this is critical to Pink and Chum salmon as they enter the marine environment at the smallest size of any salmon- id. Pink and Chum Salmon weigh less than one gram in the study area (Broughton Archipelago) throughout March and April (Morton et al. 2005), and are without protective scales (A.M. personal observations). Juve- nile Pink and Chum salmon exhibit prolonged inshore residency from March to September (Healey 1978). If some adult wild salmonids occur inshore year round, their numbers are low. The largest wild host popula- tion for L. salmonis are the mature Pink Salmon that enter coastal waters in July and run through Septem- ber (Heard 1991). Thus, in a natural setting, a system has evolved that isolates returning adult salmon from the most juvenile stage conspecifics. Juvenile Pink and Chum salmon do not share habitat with a significant L. salmonis host population until July, by which time they have grown to weigh several grams. Krkosek et al. (2005) report that after initial infection by salmon farms, juvenile salmon schools will re-infect themselves some weeks later as lice mature and being reproducing. Salmon farming has altered this arrangement by sit- ing 26 salmon farms in the nearshore environment of the Broughton Archipelago (Figure 1), each site hold- ing potentially 1 million Atlantic Salmon (Naylor et al. 2003), a known host for L. salmonis. In July of 2003 there were approximately 6 million Atlantic Salmon among the 17 farm sites leased by Stolt Sea Farms with another nine sites leased by another company (Her- itage) that did not report numbers (Orr in press). Orr (in press) reports an average of 2.2 — 9.2 gravid Sea Lice per salmon on the 6 million farm Atlantic Salmon for 2003-2004. While salmon farms are now a recognized source of Sea Lice, this study further illuminates this subject by measuring the response in Sea Louse numbers on wild juvenile salmon to the arrival of a large popula- tion of this Sea Louse’s preferred host, the adult Pink Salmon. We hypothesise that if wild adult salmon were a substantial source of Sea Lice on juvenile salmon in the study area, then the number of copepodid L. sal- monis should rise in response to the arrival of in-migrat- ing adult Pink Salmon. To test this, we first measured lice levels on juvenile Pink Salmon, then looked for evidence of copepodid Sea Louse recruitment subse- quent to wild adult Pink Salmon arrival in the area. Understanding the mechanisms of L. salmonis infes- tation of inshore juvenile wild salmon and the relative importance of both wild and farm lice sources has become crucial to both wild and farm salmon manage- ment. This study provides analysis on a previously THE CANADIAN FIELD-NATURALIST Vol. 120 unreported aspect of the dynamic between adult and juvenile Pacific salmon and Sea Lice. Methods This study took place in the 400 km’ Broughton Archipelago in British Columbia. There are seven major Pink Salmon rivers and 26 Atlantic Salmon farm sites between the rivers and the open waters of Queen Charlotte Strait (Figure 1). The Broughton Archipel- ago’s 200 km of inlets and 90 km of passages and small bays makes it ideal marine rearing habitat for juvenile Pink and Chum salmon (Healey 1978). With- in this region, Fife Sound (Figure 1) through Tribune Channel is considered a primary route used by return- ing adult Pink Salmon (G. Neidrauer, Fisheries and Oceans Canada, Fisheries Patrolman, General Deliv- ery, Simoom Sound, BC VOP 1S0 personal commu- nication). Juvenile Pink and Chum salmon were sampled once each week at the same sampling location from 1 June 2003 through 19 September 2003 (Figure 1). The sal- mon farm at this site was fallow, and without any farm salmon. A 50' x 4' beach seine of %" mesh was used in June and a 120' x 8' beach seine with 4" mesh and a %" bunt was used for the remainder of the study. Sample size ranged from 20 to 30 fish per week. The juvenile salmon were placed individually in Whirlpak™ bags immediately after capture, laid on ice and then frozen. They were later examined under 30x magnification and the number of Sea Lice, their age-class and sex were determined and recorded using published taxonomic keys (Johnson and Albright 1991a). Adult female lice were classified as either virgin or gravid as per the Norwegian method of counting sea lice (Karin Box- aspen, Researcher, Institute of Marine Research, Ber- gen, Norway). Results are reported only for the salmon- specific Sea Louse, L. salmonis. We used three independent methods to estimate the timing of the wild adult Pink Salmon return to the study area. First, we used measured Pink Salmon catch-per- unit directly at our juvenile salmon sampling site. We used two fishing rods with lures known to attract Pink Salmon in this area (A.M. personal observations). We fished for 166 hook-hours between 4 June and 19 Sep- tember. The second method compared our catch-per-unit- effort analysis with the timing of the adult return re- ported by Fisheries and Oceans Canada (DFO) as determined by their enumeration program on the known Pink Salmon-bearing streams nearby (Figure 1). The Fisheries and Oceans Pink Salmon enumeration flights began in the second week of August and continued through the end of October. Finally, a tourism operator in the Glendale River watershed (Figure 1) had guides on the river for 3 days in May, 11 days in June, 23 days in July, 28 days in August, and reported to us when Pink Salmon were first sighted. 2006 Wakeman River J Embly River 50° oO Queen Charlotte Strait MORTON AND WILLIAMS: SEA LOUSE INFESTATION LEVELS 20) 128 Kingcome River Ahnuhati River Ahta River Kakwetan Active farm site @ Fallow farm site @ Sample site FiGurE 1. Study area, the Broughton Archipelago, showing major Pink Salmon rivers, salmon farm sites and status, and study site. The adult salmon caught in this study were lifted from the water without a net and kept free from con- tact with any surface in order to reduce loss of lice. They were examined immediately for Sea Louse counts, stages and species identification using a hand lens. We estimated the infective capability of the return- ing adult salmon by counting gravid female sea lice per fish: a standard measure used to trigger de-lousing treatment of farm salmon to reduce infective capabil- ity and lower sea lice numbers on adjacent juvenile wild salmon (Heuch and Mo 2001). Louse counts were not expected to follow a nor- mal distribution. Rather than transforming the data, we made comparisons using Mann-Whitney U tests (MWU; the non-parametric equivalent of a two-sample t-test) in GraphPad version 3.05 by InStat (GraphPad Software Inc., 5575 Oberlin Drive, #110, San Diego, California 92121 USA). We used the following standard measures of louse infestation rates (Margolis et al. 1982): prevalence = the proportion of fish infested with lice; intensity = the mean number of lice on each infected fish; and abun- dance = the mean number of lice on the entire sample. Results We fished for 47 “hook-hours” 4 June through July, without catching one adult Pink Salmon. We fished for 119 hook-hours in August until 19 Septem- ber, and beginning on 4 August, caught 50 adult Pink Salmon at an average of 0.78 Pink Salmon/hook-hour (SE = 0.10, n = 17 fishing sessions, range 0-1.4 Pink Salmon/ hook-hour). Given that there was no variance in the 12 June/July fishing sessions (all 12 failed ses- sions to catch a fish), a Wilcoxon signed-rank test was conducted to assess whether the median catch-per-unit- effort in August and September differed significantly from the observed zero catch-per-unit-effort in June and July. The sum of positive ranks was 136, indicating that the median value was significantly greater than 0 (P < 0.0001). Fisheries and Oceans Canada estimated that 188 730 Pink Salmon entered the Broughton Archipelago rivers from mid-August through September 2003 (G. McEachen, Fisheries and Oceans Canada, chief of conservation management for the central coast, Central Coast Area Office, 315-940 Alder Street, Campbell River, British Columbia VOW 2P8, personal commu- 202 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Mass, length, and sea lice infection of Pink and Chum salmon collected during the two sampling periods with average sea surface temperature and salinity for both time periods. “Total” signifies all L. salmonis and “cope” signifies copepodite L. salmonis only. Period n Grams Sizecm Salt % Temp °C Prevalence % Abundance Intensity (SE) (SE) (SE) (SE) (SE) Range (SE) Total Cope Total Cope Total Cope June — July 623 6.94 8.47 29.1 12.1 ASO ez 1.20 0.079 2.79 1.09 (0.15) (0.03) (0.03) (0.01) (0.12) (0.012) (0.26) (0.018) 0-42 0-2 August — VAS a2 el Gy eel 223.8 30.8 11.1 400 48 1.00 0.048 2.50 1 19 September (0.57) (0.10) (0.11) (0.05) (0.25) (0.018) (058) (0) 0-33 0-1 nication). This would indicate that Pink Salmon were present seaward of the rivers and within the study area for 2-3 weeks prior to the end of the second week of August, when they were first identified in the rivers, through September. We cannot be certain how many of these fish passed the sampling site, but Fife Sound is considered the primary migration corridor for adult Pink Salmon (Neidrauer, personal communication). The tour operator and his guides looked for Pink Salmon on each of their tours, but did not see Pink Salmon in the largest Pink Salmon river in the study area, the Glendale River, until 20 August (H. Pattinson, owner/operator, Tide Rip Tours, 1660 Robb Avenue, Comox, British Columbia V9M 2W7 Canada, person- al communication). Again, assuming that adults took two to three weeks to migrate through the Broughton Archipelago to reach the Glendale watershed, one would expect that the adults reached our sampling location in early August. In fact, these three methods show very good agreement, and all point to a best esti- mate return date for large numbers of wild adult Pink Salmon in the first days of August. Consequently, we estimate that juvenile Pink and Chum salmon were introduced to louse-infected adults around the first day of August. We divided the pre- and post-exposure stages into two sampling periods: 1 June to 31 July, and 1 August to 19 September. As one would expect from a rapidly growing cohort, the juvenile salmon increased in both length and weight throughout the study (Table 1). The average seawater temperature and salinity varied slightly between the two sampling periods, but re- mained at all times within optimal range for L. salmo- nis (Table 1). The mean number of gravid lice on return- ing adult Pink Salmon was 9.31 (SE 0.93). No measure of total Sea Louse occurrence varied between the sampling periods (Table 1) (abundance — MWU statistic: 43186, P = 0.4088; intensity - MWU statistic: 7153.5, P = 0.3412). As well, no measure of copepodite Sea Lice varied significantly (abundance —MWU 44072; P = 0.6448). Given that there was no variance in the observed intensity of copepodid infection on juvenile salmon in the later sampling period (that is, all seven fish infected with copepodites carried a single copepodid louse), a Wilcoxon signed-rank test was used to assess whether the median early-season intensity differed from the observed intensity of 1 louse/fish in the later sampling season. The mean intensity of infection on the 45 fish infected with copepodites in the earlier sampling sea- son was 1.09 (SE 0.043, median 1.00, range 1-2). This was not significantly greater than the | louse/infected fish observed in the later sampling season (sum of all signed ranks = 10, after correcting for ties; P = 0.13). Discussion The number of returning adult Pink Salmon carry- ing gravid Sea Lice was on the order of two hundred thousand, with approximately nine gravid Sea Lice per fish and thus we estimate that the juvenile Pink and Chum salmon in Fife Sound were exposed to the infec- tive capability of close to two million gravid female Sea Lice. However no marked increase above the pre- existing background infection levels could be detected on juvenile salmon after the adult fish arrived in the area. Even if the total number of Sea Lice failed to in- crease on juvenile salmon after the return of adult Pink Salmon, an increase in the proportion of lice at the copepodid stage would have indicated arrival of a source of Sea Lice that exceeded ambient levels. How- ever, no increased copepodid settlement was observed. In fact, prevalence of copepodids was slightly lower after the adult Pink Salmon returned, which is as one would expect from an epizootic living out the course of its infestation. In the early, i.e., pre-exposure sam- pling period, 7.2% of juveniles sampled carried at least one copepodid, and after the adult Pink Salmon re- turned, 4.8% of juveniles sampled carried at least one copepodid. The juvenile salmon were traveling west out of an area where gravid Sea Lice occurred on farm salmon. The decline we recorded in copepodite Sea Lice is contrary to what Krkosek et al. (2005) found where infected juvenile salmon re-infected each other as their farm-origin Sea Lice matured. The difference in 2006 findings might be a result of geography. Krkosek et al. (2005) was conducted within the confined water- way of Tribune Channel, whereas this work occurred in an area of much greater flush at the archipelago edge. There are no juvenile Pink and Chum salmon in Tribune Channel in August. The juvenile Pacific salmon out-migrations and adult salmon in-migrations are generally timed such that interaction between the two cohorts occurs in open waters. Ocean temperature and salinity were unlikely to have hindered parasite growth or transfer during August and September. Mean sea-surface temperature fell 1°C dur- ing the adult spawning in-migration, which had the potential to slow reproduction, but salinity increased, which favoured reproduction. Both values, however, were well within optimal range necessary for Sea Lice reproduction (Johnson and Albright 1991b; Boxaspen and Naess 2000; Tucker et al. 2000). Consequently, we conclude that the juvenile hosts in our study simply failed to respond measurably to exposure to a large influx of infested wild adult salmon. We fully expected louse loads to increase when the adult Pink Salmon returned. We know that Sea Lice must be transmitted among wild salmon. The fact that Louse numbers did not increase in our study was sur- prising, and leads us to conclude that, compared to ambient levels of farm-origin parasitism in the Brough- ton Archipelago, the contribution of wild-source lice was negligible. We suspect the higher numbers of salmon-specific Sea Lice on juvenile Pink and Chum salmon in June and July originated from the popula- tion of several million, farmed Atlantic Salmon in the Broughton Archipelago in high-density open-net pens. Orr (in press) reports an average of several gravid sea lice per each of 6 million farm salmon during our study period, which translates into a very large number of copepodite sea lice occurring to the east of our study site area, but not at our study site itself which was adja- cent to an empty salmon farm. Why we did not find an increase in Sea Lice in August and September is perplexing. The mandatory 2-4 day dispersal term in the plankton layer likely limits Sea Louse recruitment rates to allow both para- site and host to thrive. Perhaps there were not enough wild adult Pink Salmon in the Broughton Archipelago for measurable Sea Lice transmission to the juvenile cohort in the face of such dispersal. Perhaps Sea Louse recruitment is an infrequent event between free-ranging salmon, and our sample size was too small to capture it. Perhaps the duration of exposure to adult Pink Salmon was not long enough or the dynamic of juvenile and adult salmon travelling in opposite directions with juve- niles along the shoreline and adults mid-channel sup- pressed copepodid recruitment. Perhaps some small increase in parasite load did occur, but natural variabil- ity and small sample size left us with insufficient sta- tistical power to detect this small effect above the ambi- ent levels in the Broughton Archipelago. Morton et al. MORTON AND WILLIAMS: SEA LOUSE INFESTATION LEVELS 203 (2004) reported that in areas without salmon farms juvenile Pink Salmon were not infested with L. salmo- nis. Therefore, it is possible that in the absence of farm salmon the 4.8% prevalence we found on juvenile sal- mon in August and September could have been entirely or partially in response to arrival of the wild adult Pink Salmon. Likely our inability to detect response in Sea Louse populations to returning adult Salmon was some com- bination of these. The lack of observed recruitment of new Sea Lice suggests that some mechanism has evolved to suppress transfer of lice from returning adults to out-migrating smolts in a naturally function- ing ecosystem. Because Sea Louse pathogenicity is host-size dependant, protecting the smaller salmon from L. salmonis infestation would have clear evolutionary benefits, particularly for the Pink and Chum salmon that go to sea smaller than any other salmonid. If exposure to abundant Sea Lice infested wild adults did not trigger a measurable increase in Sea Lice loads on juvenile Pink and Chum salmon, then this leaves the other, less abundant (if at all present), over- wintering inshore wild salmon species poor candidates for explaining the enormous and recent spring L. salmo- nis outbreaks in the Broughton Archipelago. Clearly, more extensive research is urgently needed. In the meantime, our study suggests that wild Pink Salmon do not appear to be the primary source of the high Sea Lice infestations reported on wild juvenile Pink and Chum salmon in the Broughton Archipelago. They may, however, be the primary source infecting the farm salmon and thus pose a risk to the industry. We conclude the most obvious source of L. salmonis on juvenile Pink and Chum salmon in the Broughton Archipelago is the stationary population of several million, farmed Atlantic Salmon. Acknowledgments This research was supported in part by Tides Canada. RW is supported by the Jane Marcher Foundation. Our thanks to Ron Turner for assistance in beach seining. Documents Cited (marked * in text) Jones, S., and A. Nemec. 2004. Pink Salmon Action Plan: sea lice on juvenile salmon and on some non-salmonid species in the Broughton Archipelago in 2003. Canadian Science Advisory Secretariat Research Document. Available from: http://www.dfo-mpo.ge.ca/csas/Csas/publications ResDocsDocRech/2004/2004_105_e.htm. Literature Cited Berland, B. 1993. Salmon lice on wild salmon (Salmo salar L.) in western Norway. Pages 179-187 in Pathogens of wild and farmed fish: Sea Lice. Edited by G. A. Boxshall and D. Defaye. Ellis Horwood, London, United Kingdom. Birkeland, K. 1996. Consequences of premature return by sea trout (Salmo trutta) infested with the salmon louse (Lepeophtheirus salmonis Kroyer); migration, growth and mortality. Canadian Journal of Fisheries and Aquatic Sci- ence 53: 2808-2813. 204 Bjorn, P. A., and B. Finstad. 1997. The physiological effects of salmon lice infection on sea trout post smolts. Nordic Journal of Freshwater Research 73: 60-72. Boxaspen, K., and T. Naess. 2000. Development of eggs and the Planktonic stages of salmon lice (Lepeophtheirus sal- monis) at low temperatures. Contributions to Zoology 69: 51-55. Boxshall, G. A. 1974. Infections with parasitic copepods in North Sea marine fishes. Journal of Marine Biology Asso- ciation. UK 54: 355-372. Grimnes, A., and P. J. Jakobsen. 1996. The physiological effects of salmon lice infection on post-smolt of Atlantic salmon. Journal of Fish Biology 48: 1179-1194. Hahnenkamp, L., and H. J. Fyhn. 1985. The osmotic res- ponse of salmon louse, Lepeophtheirus salmonis (Cope- poda: Caligidae), during transition from sea water to fresh water. Journal of Comparative Physiology B 155: 357-365. Healey, M. C. 1978. The distribution, abundance and feeding habits of juvenile Pacific salmon in Georgia Strait, Bri- tish Columbia. Fisheries Marine Service (Canada) Tech- nical Report 78. 49 pages. Healey, M. C. 1982. The distribution and residency of juve- nile Pacific salmon in the Strait of Georgia, British Colum- bia, in relation to foraging success. Edited by B. R. Melteff and R. A. Neve. Proceedings of the North Pacific Aqua- culture Symposium Alaska Sea Grant Report 82: 61-69. Heard, W. 1991. Life history of pink salmon. Pages 121- 230 in Pacific salmon life histories. Edited by C. Groot and L. Margolis. University of British Columbia Press, Van- couver, British Columbia. Heuch, P. A., and T. A. Mo. 2001. A model of salmon louse production in Norway: effects of increasing salmon pro- duction and public management measures. Diseases of Aquatic Organisms 45: 145-152. Johnson, S. C., J. W. Treasurer, S. Bravo, K. Nagasawa, and Z. Kabata. 2004. A review of the impact of parasitic copepods on marine aquaculture. Zoological studies 43: 229-243. Johnson, S. C., and L. J. Albright. 1991a. The develop- ment stages of Lepeophtheirus salmonis (Kr@yer, 1837) (Copepoda: Caligidae). Canadian Journal of Zoology 69: 929-950. Johnson, S. C., and L. J. Albright. 1991b. Development, growth and survival of Lepeophtheirus salmonis (Cope- poda: Caligidae) under laboratory conditions. Journal of Marine Biology Association U.K. 71: 425-436. Kabata, Z. 1973. The species of Lepeophtheirus (Copepoda: Caligidae), a parasite of fishes from the Pacific coast of North America. Journal of Fisheries Research Board of Canada 30: 729-759. Krkosek, M., M. A. Lewis, and J. P. Volpe. 2005. Trans- mission dynamics of parasitic sea lice from farm to wild salmon Proceedings of the Royal Society, Series B doi: 10.1098/rspb.2004.3027. Krkosek, M., M. A. Lewis, A. B. Morton, L. N. Frazer, and J. P. Volpe. 2006. Epizootics of wild fish induced by farm fish. Proceedings of the National Academy of Sci- ences103: 15506-15510. Margolis, L., G. W. Esch, J. C. Holmes, A. M. Kuris, and G. A. Schad. 1982. The use of ecological terms in para- sitology (report of an ad hoc committee of the American Society of Parasitologists). Journal of Parasitology 68: 131- 133. THE CANADIAN FIELD-NATURALIST Vol. 120 McLean, P. H., G. W. Smith, and M. J. Wilson. 1990. Residence time of sea louse, Lepeophtheirus salmonis K.., on the Atlantic salmon, Salmo salar L., after immersion in fresh water. Journal of Fish Biology 37: 311-314. Morton, A., and R. Williams. 2003. First report of the Sea Louse, Lepeophtheirus salmonis, infestation on juvenile Pink Salmon, Oncorhynchus gorbuscha, in nearshore habi- tat. Canadian Field-Naturalist 117: 634-641. Morton, A., R. Routledge, C. Peet, and A. Ladwig. 2004. Sea Lice Lepeophtheirus salmonis infection rates on juve- nile pink Oncorhynchus gorbuscha and chum salmon Oncorhynchus keta in the nearshore marine environment of British Columbia, Canada. Canadian Journal of Fish- eries and Aquatic Science 61: 147-157. Morton, A. B., R. Routledge, and R. Williams. 2005. Tem- poral patterns of sea lice infestation on wild Pacific salmon in relation to the fallowing of Atlantic salmon farms. North American Journal of Fisheries Management 25: 811-821. Nagasawa, K. 1987. Prevalence and abundance of Lepeoph- theirus salmonis (Copepod: Caligidae) on high-sea salmon and trout in the North Pacific Ocean. Bulletin of the Japan- ese Society of Scientific Fisheries 53: 2151-2156. Nagasawa, K., Y. Ishida, M. Ogura, K. Tadokora, and K. Hiramatsu. 1993. The abundance and distribution of Lep- eophtheirus salmonis (Copepoda: Caligidae) on six species of Pacific salmon in offshore waters of the North Pacific Ocean and Bering Sea. Pages 166-178 in Pathogens of wild and farmed fish: Sea Lice. Edited by G. A. Boxshall and D. D. Defaye. Ellis Horwood, New York. Naylor, R. L., J. Eagle, and W. L. Smith. 2003. Salmon aquaculture in the Pacific Northwest: a global industry with local impacts. Environment 45: 18-39. Orr, C. (in press). Estimated sea louse egg production from Marine Harvest Canada (Stolt) farmed salmon, Broughton Archipelago, British, Columbia, Canada, 2003-2004. North American Journal of Fisheries Management. Parker, R. R. 1965. Estimation of sea mortality rates for the 1961 brood-year pink salmon of the Bella Coola area, British Columbia. Journal of Fisheries Research Board of Canada 22: 1523-1554. Parker, R. R., and L. Margolis. 1964. A new species of parasitic copepod, Caligus clemensi sp. Nov. (Caligoida: Caligidae), from pelagic fishes in the coastal waters of British Columbia. Journal of the Fisheries Research Board of Canada 21: 873-889. Parker, R. R., and W. E. Vanstone. 1966. Changes in chemi- cal composition of central British Columbia Pink Salmon during early sea life. Journal of Fisheries Research Board Canada 23: 1353-1384. Pike, A. W., and S. L. Wadsworth. 1999. Sea lice on salmonids: Their biology and control. Advanced Parasitol- ogy 44: 234-337. Tucker, C. S., C. Sommerville, and R. Wooten. 2000. The effect of temperature and salinity on the settlement and survival of copepodids of Lepeophtheirus salmonis (Kr@yer, 1837) on Atlantic salmon Salmo salar L. Journal of Fish Diseases 23: 309. Tully, O., W. R. Poole, and K. F. Whelan. 1993. Infestation parameters of Lepeophtheirus salmonis (Kroyer) (Cope- pod: Caligidae) parasitic on sea trout, Salmo trutta L., off the west coast of Ireland during 1990 and 1991. Aquacul- ture and Fisheries Management 24: 545-555. Received 17 April 2005 Accepted 9 February 2007 Physical Characteristics, Hematology, and Serum Chemistry of Free- ranging Gray Wolves, Canis lupus, in Southcentral Alaska MATTHEW J. BUTLER!, WARREN B. BALLARD!, and HEATHER A. WHITLAW2 'Department of Natural Resources Management, Texas Tech University, Box 42125, Lubbock, Texas 79409-2125 USA 29305 Winston Avenue, Lubbock, Texas 79424 USA Butler, Matthew J., Warren B. Ballard, and Heather A. Whitlaw. 2006. Physical characteristics, hematology, and serum chemistry of free-ranging Gray Wolves, Canis lupus, southcentral Alaska. Canadian Field-Naturalist 120(2): 205-212. Examination of morphometric characteristics and blood parameters has become a widely used tool for assessing the physio- logical and nutritional status of wild and captive animals. During 1976 through 1984, 155 Gray Wolves (Canis lupus) were chemically immobilized in south-central Alaska. Of those, we obtained physical measurements from 132 and blood samples from 121 individuals. Also, 208 carcasses of harvested and dead radiocollared Wolves were weighed and measured. We obtained blood samples from three of the fresh carcasses. We measured age, body weight, skull length and width, and upper and lower canine length. We analyzed blood serum for Ca, P, Fe, chlorides, creatinine, glucose, lactic dehydrogenase, alkaline phosphatase, glutamic oxalic transaminase, triglyceride, beta globulin, serum urea nitrogen, and uric acid. We obtained packed cell volume and hemoglobin values from whole blood. We classified samples by season, sex, and age. Seasonal differences were observed for physical measurements, packed cell volume, alkaline phosphatase, and serum urea nitrogen. Age differences were observed for physical measurements, hemoglobin, packed cell volume, alkaline phosphatase, P, Ca, creatinine, serum urea nitrogen, and percent femur bone marrow fat. However, differences among sexes were observed for physical measurements only. These data provide a baseline for physical condition, hematology, and serum chemistry for free-ranging Gray Wolves. Key Words: Gray Wolf, Canis lupus, blood, chemistry, hematology, measurements, physical, serum, Alaska. Examination of morphometric characteristics and blood parameters has become a widely used tool for assessing the physiological and nutritional status of wild and captive animals (e.g., LeResche et al. 1974; Seal and Mech 1983; DelGiudice et al. 1987; Franz- mann and Schwartz 1988). Although baseline hema- tology and serum chemistry values for many species of captive animals exist, baseline values for free-rang- ing animals are also necessary to properly interpret data. It is also important to understand how factors such as age, sex, and season influence hematology and serum chemistry values. Hematology and serum chemistry of captive Gray Wolves (Canis lupus) has been studied and described by Fox and Andrews (1973) and Constable (1998) in Alaska, Seal and Mech (1983) and DelGiudice et al. (1987) in Minnesota, Pospisil et al. (1987) in the Czech Republic, and Drag (1995) in Missouri. Seal et al. (1975) and DelGiudice et al. (1991) in Minnesota, Messier (1987) in Quebec, and Constable et al. (1998) in Alaska studied and described hematology and serum chemistry of free-ranging Gray Wolves. Mech et al. (1984) reported changes in hematology and serum chemistry of a female Gray Wolf recuperating from near starvation in Minnesota. However, little baseline data exist for age, sex, and season for free-ranging Gray Wolf populations. The purpose of our study was _ to describe physical characteristics and report base- line hematology and serum chemistry values for age, sex, and season in free-ranging Gray Wolves of south- central Alaska. Study Area The study was conducted in an area of approxi- mately 45 000 km’, known as Alaska Game Manage- ment Unit 13, located approximately 125 km north- east of Anchorage, Alaska. The area was bisected by three major river systems including the Copper, Nel- china, and Susitna rivers. Primary prey species were Moose (Alces alces) and Barren-ground Caribou (Ran- gifer tarandus granti). Alternate prey species includ- ed Dall Sheep (Ovis dalli), Beaver (Castor canaden- sis), and Snowshoe Hare (Lepus americanus). Climate, topography, and other important aspects of the study area were described by Ballard et al. (1987, 1991b). Gray Wolf densities during this study ranged from 10.3/1000 km? in autumn 1975 to 2.6/1000 km? in spring 1982 (Ballard et al. 1987). Mean annual litter size in November ranged from 3.7 to 7.3 pups. The Wolf population was heavily exploited by humans (i.e., >40% of population) during the course of this study (Ballard et al. 1987). The reference values we present here occurred dur- ing a period when both Moose and Caribou popula- tions were increasing. Moose densities were highly variable in the study area and averaged about 603 to 741/1000 km? (Ballard et al. 1991b) and the popula- tion increased at a finite rate of increase of 1.03 to 1.06 annually (Ballard et al. 1986). Caribou popula- tions during this time period increased from approxi- mately 10 000 in 1976 to about 25 000 in 1984 (Ber- gerud and Ballard 1988). Winter conditions as reflected by snow depths were relatively moderate during this 205 206 time period except during 1978-1979 which was classi- fied as severe (Ballard et al. 1991b). Methods We captured Gray Wolves for radiocollaring by dart- ing them from a helicopter during 1976 through 1984 using methods described by Ballard et al. (1982, 1991a). We immobilized Wolves with phencyclidine hydrochloride, etorphine hydrochloride, or a combina- tion of zolzepam hydrochloride and tiletamine hydro- chloride following dosages and methods described by Ballard et al. (1982, 1991a). Induction times averaged 7.2 minutes (Ballard et al. 1991a). After immobiliza- tion each Wolf was radiocollared, weighed, sexed, ear- tagged, bled, and a number of physical measurements were recorded on standardized forms. Physical meas- urements included skull length (SL) and width (SW), and upper (UCL) and lower canine length (LCL). We measured canine length from the edge of the gum line to the tip of the tooth. Skull and tooth measurements were taken with calipers and recorded to the nearest mm. Hunters and trappers were paid $10 (U.S.) per skinned Wolf carcass. Carcasses were weighed (CW) and then kept frozen until examined. Skulls were not boiled and cleaned before measuring to maintain sim- ilarity with live Wolf measurements. Also, the effects of freezing carcasses on physical measurements were likely minimal. We determined percent femur bone marrow fat (MF) by the dry weight method (Neiland 1970). Because this Wolf population was heavily exploited (Ballard et al. 1987) we were able to do com- parisons with many individuals that had been radio- . collared and harvested within the same year. We collected blood samples from saphenous veins using the B-D Vacutainer system (Becton-Dickenson Co., Rutherford, New Jersey, USA). Generally blood was taken within 15 minutes of induction time and serum was separated each evening. Only three blood samples were collected from carcasses. Those samples were obtained within 15 minutes of harvest. We used one EDTA tube to preserve whole blood. Packed cell volume (PCV) was determined with uncoagulated whole blood using a microhematocrit centrifuge (Triac, Clay-Adams Co., Parisppany, New Jersey, USA) while hemoglobin (Hb) was estimated with an AO Hb-Meter (American Optical, Buffalo, New York, USA). Clot- ted blood was centrifuged within 36 h of collection and serum was then stored frozen at -17°C. We sent frozen serum to Pathologist Central Laboratory, Seattle, Wash- ington, USA, where it was analyzed for the following via an auto-analyzer: calcium (Ca), phosphorous (P), iron (Fe), chlorides (CHL), creatinine (CRE), glucose (GLU), lactic dehydrogenase (LDH), alkaline phos- phatase (SAP), glutamic oxalic transaminase (SGOT), triglyceride (TRI), serum urea nitrogen (SUN), and uric acid (UA). Protein fractions were determined by stan- THE CANADIAN FIELD-NATURALIST Vol. 120 dard protein electrophoresis for beta globulin (BEG). Because of unbalanced cells in our analyses, we pooled years, sorted blood and physical measurements by sex, age, and season, and conducted multiple analy- ses of variance. We conducted post-hoc comparisons with a Scheffe test (Sokal and Rohlf 1995). We deter- mined correlations among variables by Spearman’s rank correlation (Conover 1980). We classified ages as pup (<12 months), yearling (13 to 23 months), and adult (>24 months), while seasons were divided into autumn (September through November), winter (De- cember through February), spring (March through May), and summer (June through August). Differences were considered significant when P < 0.05 unless oth- erwise specified. Results We immobilized 155 individual Gray Wolves dur- ing 1976 through 1984. From that total, we obtained 121 blood samples (n = 121 serum samples, n = 117 whole blood samples) and 132 live body weights (BW). Due to low sample sizes, we excluded yearling and summer data from the analyses. In addition, we weighed and measured a total of 208 carcasses. We obtained three blood samples from fresh carcasses (n = 3 serum samples, n = 2 whole blood samples). Physical Characteristics Mean BW differed by season (F = 26.121, 2 df, P <0.001), sex (F = 37.084, 1 df, P < 0.001), and age (F = 147.890, 1 df, P < 0.001), and there were signifi- cant interactions between season and age (F = 15.665, 2 df, P < 0.001), and among season, sex, and age (F = 3.231, 2 df, P=0.043). Lowest BW occurred for pups during autumn, while highest BW occurred for adult males during autumn and winter (Table 1). Adult males had higher BW than male pups during autumn and winter; adult females had higher BW than female pups during autumn and spring (Table 1). Body weights were not different between adult males and adult fe- males among seasons (Table 1). Body weights between male and female pups was only significantly differ- ent during spring (i.e., males were heavier; Table 1). Body weights of male pups increased with seasons (i.e., over the course of the biological year); both win- ter and spring BW were higher than autumn weights, although winter and spring were not different (Table 1). In addition, BW of female pups increased with season though female pups were significantly heavier in the spring than during autumn (Table 1). Mean CW exhibited similar trends by sex, age, and season as BW although sample sizes were too small to evaluate post-hoc differences (Table 1). Season (F = 4.849, 2 df, P = 0.009), sex (F = 6.665, 1 df, P=0.011), and age (F = 24.849, 1 df, P < 0.001) each had a significant effect on CW and there were signifi- cant interactions between sex and age (F = 3.950, 2 df, P = 0.046), and among season, sex, and age 2006 TABLE 1. Mean physical measurements + | standard error by season, sex, and age for free-ranging Gray Wolves in south central Alaska during 1976 through 1984. Sample size denoted in parentheses. BUTLER, BALLARD, and WHITLAW: WOLVES IN SOUTHCENTRAL ALASKA 207 Physical Season measurements Age Sex Autumn Winter Spring Live-body weight (kg) Pup Male 26.2+2.3 (8) 37.9 + 14 (10) 413+ 14(16) Female 23.5+2.0 (7) 33.3'4+.25 (5) 33.34 0.9(20) Adult Male 46.8+1.5 (8) 48.1 + 1.3 (11) 46.2+ 0.9(17) Female 38.9+1.4(13) 41.141.2 (3) 424+ 1.0(14) Carcass weight (kg) Pup Male 19.7+2.7 (3) 30.7 + 0.9 (51) 299+ 25 (8) Female 25.0424 (3) 27.3 + 0.6 (51) 253+ 10 (7) Adult Male 39.1 (1) 39.8 + 0.9 (36) 335+ 18 (8) Female 24.8 +13.8 (2) 32.8 + 0.7 (32) 338+ 1.1 (6) Skull length (cm) Pup Male 24.5+0.8 (3) 26.2 + 0.2 (58) 27.1+ 0.3(18) Female 24.5+0.7 (5) 25.1 + 0.2 (52) 25.5+ 0.3(14) Adult Male 27.7+40.3 (3) 27.8 + 0.2 (39) 27.0+ 04(12) Female 24.8+1.0 (3) 26.5 + 0.2 (31) 26.4+ 0.4(12) Skull width (cm) Pup Male 12.34+0.5 (3) 13.5+0.1 (58) 14.44 0.2(18) Female 124+04 (5) 13.1 + 0.2 (52) 14.0+ 0.7(14) Adult Male 16.0+0.6 (3) 14.8 + 0.1 (35) 14.9+ 0.3(12) Female 13.4+05 (6) 14.5 + 0.3 (30) 14.34 0.1(11) Upper canine length (cm) Pup Male 2.1402 (2) 2.5 + 0.1 (38) 2.6+ 0.1(11) Female 1940.1 (5) 2.340.1 (45) 2.34 0.1(10) Adult Male 2.8 + 0.04 (6) 2.9+0.1 (32) 3.0+ 0.1 (7) Female 23-033) 2.5+0.1 (25) 25+ 0.04(6) (F = 4.105, 1 df, P=0.018). Lowest CW occurred for pups during autumn, while highest weights were for adult males during autumn and winter (Table 1). There were significant differences in average SL by season (F = 5.008, 2 df , P = 0.007), sex (F = 23.574, 1 df ,P <0.001), and age (F = 22.331, 1 df ,P < 0.001), and there were significant interactions between season and age (F = 3.769, 2 df , P = 0.024) and among sea- son, sex, and age (F = 3.874, 2 df , P = 0.022). Low- est mean SL was for pups during autumn and highest mean SL was for adult males during autumn and win- ter (Table 1). Males had higher mean SL than females during each season (Table 1). Mean pup SL increased from autumn through spring (Table 1). Mean SW differed by season (F = 3.831, 2 df , P=0.023), sex (F = 7.692, 1 df , P= 0.006), and age (F = 32.160, 1 df , P < 0.001), and there was a signif- icant interaction between season and age (F = 4.582, 2 df ,P =0.011). Mean male SW was greater than fe- male SW for all seasons and ages (Table 1). There were no differences in average adult SW among seasons (Table 1). Also, mean pup SW increased by season dur- ing the first year of life (Table 1). Mean UCL differed by season (F = 5.257, 2 df , P = 0.006), sex (F = 19.804, 1 df , P < 0.001), and age (F = 26.872, 1 df , P < 0.001), and there were no significant interactions. Mean UCL was lower during autumn than during either winter or spring (Table 1). Mean male UCL was greater than that of females (Table 1). Mean UCL for adults was greater than that for pups (Table 1). There was no significant difference in mean LCL by season (F = 0.002, 1 df , P = 0.964), sex (F = 1.707, 1 df , P=0.218), or age (F = 0.828, 1 df, P = (0.382), and there were no significant interactions. Mean LCL was 2.3 + 0.07 cm (n =18). Body weight was also correlated with the other physical character- istics, MF, and PCV (Table 2). Hematology Mean Hb concentration differed by age (F = 7.671, 1 df ,P =0.007) and there was a significant interaction between age and season (F = 3.876, 2 df , P =0.024): differences were between pups and adults during win- ter and autumn (Table 3). Mean pup Hb concentrations increased throughout the year, whereas adult Hb con- centrations remained relatively constant during the year (Table 3). Average PCV differed by season (F = 6.015, 2 df, P =0.003) and age (F = 29.083, 1 df, P < 0.001), and there was a significant interaction between season and age (F = 10.375, 1 df, P < 0.001). Adults during autumn had higher mean PCY than all other groups by season and age (Table 3). Mean PCV did not differ between adults and pups in winter and spring (Table 3). Packed cell volume was also correlated with Hb and BW (Table 2). Serum Chemistry Mean SAP concentration differed by season (F = 6.557, 2 df, P= 0.002) and age (F = 19.621, 1 df, P< 0.001), and there was a significant interaction between age and season (F = 5.189, 2 df, P = 0.007). Mean SAP con- centration appeared greater in pups than adults dur- ing autumn, winter, and spring (Table 3). However, only during autumn were SAP concentrations signifi- cantly different between pups and adults (Table 3). Among pups, autumn and winter SAP concentrations were higher than spring, but only the autumn—spring comparison was significant. Among adults, seasonal 208 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 2. Correlations among physical characteristics, hematology, and serum chemistry values for free-ranging Gray Wolves in south-central Alaska during 1976 through 1984. Characteristics Body weight Carcass weight Skull length Skull width Upper canine length Marrow fat Packed cell volume Hemoglobin Carcass weight Skull length Skull width Upper canine length Marrow fat Serum alkaline phosphatase Age Phosphorous Calcium Beta globulin Lactic dehydrogenase Packed cell volume Hemoglobin iP n P 0.954 23 <0.001 0.665 67 <0.001 0.639 62 <0.001 0.583 47 <0.001 0.725 14 0.003 0.271 97 0.007 0.362 82 0.001 0.365 108 <0.001 0.281 108 0.003 0.494 108 <0.001 0.316 79 0.005 —0.516 91 <0.001 0.600 140 <0.001 0.511 142 <0.001 0.500 91 <0.001 0.527 102 <0.001 TABLE 3. Mean blood characteristics + | standard error by season and age for free-ranging Gray Wolves in south-central Alaska during 1976 through 1984. Sample size denoted in parentheses. Blood characteristics Age Autumn Alkaline phosphatase (IU/L) Pup 177.8 + 33.1 Adult 493+ 6.1 Urea nitrogen (mg/dL) Pup 262+ 2.5 Adult 454+ 42 Hemoglobin (g/dL) Pup 179+ 04 Adult 199+ 0.0 Packed cell volume (%) Pup 478+ 2.2 Adult 63.6+ 2.0 SAP concentrations were not significantly different (Table 3). Serum alkaline phosphatase was also cor- related to age, P, and Ca (Table 2). Mean SUN concentration differed by season (F = 6.746, 2 df, P = 0.002) and age (F = 10.382, 1 df, P = 0.002), and there were no significant interactions. Mean SUN concentration was higher during spring than in winter; autumn SUN concentration was not different from spring or winter (Table 3). Mean adult SUN concentration was higher than that of pups, regardless of season or sex (Table 3). Mean Pi — 6595s lidine 0.012) ia (i 6.036: 1 df, P=0.016), and CRE (F = 4.189, 1 df, P = 0.043) concentrations differed by age with no significant inter- actions. Pup P concentrations were higher than adults, regardless of season or sex, as were Ca concentrations (Table 4). Mean adult CRE concentration was higher than that of pups, regardless of season or sex (Table 4). There were no significant differences in mean val- ues of BEG, CHL, GLU, Fe, LDH, SGOT, TRI, or UA by season, sex, or age, and there were no significant interactions (Table 5). Measures of BEG were also correlated with LDH concentrations (Table 2). Season Winter Spring (10) 141.9+ 32.0 (15) 65.0+ 6.9 (29) (21) 90.0 + 38.8 (12) 49.8+ 6.7 (37) (10) 254+ 60 (15) 46.1+ 4.1 (29) (21) 376+ 74 (12) 52.9+ 34 (37) (8) 18.0+ 0.8 (10) 19.14 0.2 (26) 7 (21) 189+ 0.6 (11) 190+ 0.3 (29) (7) 442+ 1.2 (12) 490+ 1.1 (32) (23) 55.14 1.8 (11) 497+ 13 (34) Bone Marrow Fat Mean MF differed by age (F = 5.062, 1 df, P=0.027). Mean pup MF was less than adult MF (Table 4). There were no differences in mean MF by season (F =0.351, 1 df, P = 0.555) or sex (F = 0.000, | df, P = 0.989) and there were no significant interactions. Percent bone marrow fat was also correlated to BW, CW, SL, SW, and UCL (Table 2). Discussion Average BW of free-ranging Gray Wolves in south- central Alaska was greater than captive Gray Wolves in Minnesota (VanBallenberghe and Mech 1975; Seal and Mech 1983; Mech 2006) probably reflecting sub- species differences (Mech 1970). The relationship be- tween CW and BW may allow managers to gain reli- able biological data from harvested individuals. Also, the relationship between BW and MF may allow biol- ogists to infer condition based upon BW. Pup and adult Hb concentrations were in the upper range of values reported for free-ranging (Seal et al. 1975; Messier 1987; DelGiudice et al. 1991) and cap- tive (Seal and Mech 1983; DelGiudice et al. 1987; 2006 Pospisil et al. 1987) Gray Wolves. In addition, we ob- served a correlation between Hb and BW and it has been suggested Hb concentration is a good indicator of physical condition (Franzmann and LeResche 1978; Messier 1987; Franzmann and Schwartz 1988). How- ever, Harlow and Seal (1981) observed little change in Hb concentrations in food-deprived captive Bad- gers (Taxidea taxus). Perhaps, lower Hb concentrations in pups are indicative of a dietary difference due to social status. Seal and Mech (1983) observed low Hb concentrations during July and August in captive Gray Wolves. They also noted reduced BW during this peri- od. Also, lower Hb concentrations in winter than in spring were reported by Thomas and Kittrell (1966) for German shepherds. Changes in Hb concentrations by season have also been observed in female Black Bears (Ursus americanus) (Franzmann and Schwartz 1988) and female White-tailed Deer (Odocoileus vir- ginianus) (DelGiudice et al. 1992). However, our Hb data were unaffected by season and we observed com- parable Hb concentrations in summer (18.6 + 1.4 g/dL, n= 5). Franzmann and LeResche (1978) did not ob- serve a seasonal effect on Hb concentrations in Moose. Perhaps, the reduced BW and Hb concentrations ob- served by Seal and Mech (1983) during summer can be attributed to a physiological response to greater sum- mer heat in Minnesota. However, the elevation of Alas- ka Game Management Unit 13 lies above 1,220 m and the research conducted by Thomas and Kittrell (1966) suggested increased elevation resulted in increased Hb concentrations in German shepherds. Thus, per- haps, high Hb concentrations in south-central Alaskan Gray Wolves may have been due to higher elevation. Thomas and Kittrell (1966) also observed increased elevation resulted in increased PCV levels in German shepherds. Perhaps, this explains why mean PCV in south-central Alaskan Wolves was greater than values reported by DelGiudice et al. (1991) in free-ranging Minnesota Wolves. Similar PCV values have been ob- served in free-ranging (Smith and Rongstad 1980) and captive (Gates and Goering 1976; Rich and Gates 1979) Coyotes (Canis latrans). Franzmann and LeResche (1978) and Franzmann and Schwartz (1988) suggested PCV was a good indi- cator of physical condition. We also observed a sea- sonal effect on PCV. Autumn PCV (59.9 + 2.0% ,n = 30) was greater than spring (49.4 + 0.8%, n = 66) and winter (49.4 + 1.6%, n = 23). This suggests Wolves in Alaska Game Management Unit 13 were in their best physical condition in autumn. However, differ- ent seasonal patterns were observed by Thomas and Kittrell (1966) in German shepherds: lower PCV in winter than spring. Seasonal changes in PCV have also been observed in female Black Bears (Franzmann and Schwartz 1988), female White-tailed Deer (Bahnak et al. 1979; DelGiudice et al. 1992), and Moose (Franz- mann and LeResche 1978). Female White-tailed Deer in Minnesota exhibited two peaks (February-March BUTLER, BALLARD, and WHITLAW: WOLVES IN SOUTHCENTRAL ALASKA 209 TABLE 4. Mean characteristics + | standard error by age for free-ranging Gray Wolves in south-central Alaska during 1976 through 1984. Sample size denoted in parentheses Characteristics Age Mean + standard error (n) Phosphorous (mg/dL) Pup 58+ 03 (54) Adult 44+ 04 (69) Calcium (mg/dL) Pup 96+ 03 (54) Adult 89+ 03 (70) Creatinine (mg/dL) Pup 09+ 0.04 (54) Adult 11+ 0.1 (70) Marrow fat (%) Pup 79.6+16.9 (64) Adult 87.0+ 9.2 (44) TABLE 5. Mean serum chemistry values + | standard error for free-ranging Gray Wolves (season, sex, and age com- bined) in south-central Alaska during 1976 through 1984. Sample size denoted in parentheses. Serum chemistry values Mean + standard error (71) 474+ 45 (71) 161.1+ 4.0 (124) 118.7+ 7.4 (122) 306.7 + 22.7 (123) Iron (umol/L) Chlorides (mmol/L) Glucose (mg/dL) Lactic dehydrogenase (IU/L) Glutamic oxalic transaminase (IU/L) 226.5 + 23.8 (120) Triglyceride (mg/dL) 518+ 5.8 (124) Beta gobulin (g/dL) 10+ 0.06 (93) Uric acid (mg/dL) 13+ 0.09(114) and October-November) in BW and PCV (DelGiudice et al. 1992). However, PCV levels in Michigan female White-tailed Deer reached their lowest levels in July through September when those animals reached slight- ly lower weights (Buhnak et al. 1979). Female Black Bear exhibited a decline in PCV during summer and, though not significant, an increase during autumn (Franzmann and Schwartz 1988). Franzmann and Le- Resche (1978) observed greater PCV during June through October in Moose. Our SUN data was greater than or in the upper range of values reported for free-ranging Wolves (Seal et al. 1975; Messier 1987), captive Wolves (Seal and Mech 1983; Drag 1995), captive Coyotes (Rich and Gates 1979; Dunbar and Giordano 2002), and free-ranging Coyotes (Smith and Rongstad 1980). However, other studies have reported concentrations similar to this study. DelGiudice et al. (1987) reported average SUN concentrations of well-fed captive Gray Wolves be- tween 37.5 and 44.4 mg/dL. However, during fasting those concentrations were reduced to a range of 12.0 to 19.8 mg/dL. Another study conducted on free-ranging Wolves (n = 11) in the Yukon-Charley Rivers National Preserve, Alaska (Constable et al. 1998), reported SUN concentrations (46.2 mg/dL) similar to our observa- tions. Higher dietary intake of protein is indicated by high- er SUN concentrations as described for Dogs (Lane 210 and Robinson 1970; Bressani and Braham 1977), White-tailed Deer (Seal et al. 1972; Bahnak et al. 1979), and Pronghorn (Antilocapra americana) (Seal and Hoskinson 1978). We observed lower SUN con- centrations in pups, suggesting a dietary difference, perhaps, due to social status. We also observed season- al differences in SUN concentrations. Urea nitrogen was greatest during spring, intermediate in autumn, and lowest in winter, suggesting greatest intake of pro- tein during spring. This may be indicative of increased Moose vulnerability due to reduced Moose group size and increased number of calves (Ballard et al. 1991b). Though sample size was low (n = 5), SUN concentra- tion during summer was relatively high (41.8 + 6.1 mg/ dL) which was comparable to well-fed captive Wolves (DelGiudice et al. 1987). Seasonal variation has also been observed in Black Bears (Franzmann and Schwartz 1988), female White-tailed Deer (Bahnak et al. 1979; DelGiudice et al. 1992), Santa Cruz Island Spotted Skunks (Spilogale gracilis amphiala) (Crooks et al. 2003), Santa Cruz Island Foxes (Urocyon littoralis) (Crooks et al. 2000), and Son Joaquin Kit Foxes (Vul- pes macrotis mutica) (McCue and O’Farrell 1992). For each of these species, peak SUN concentrations ap- peared to coincide with the most productive seasons. Phosphorus concentrations were similar to those reported for free-ranging (Seal et al. 1975; Messier 1987; Constable et al. 1998) and captive (Drag 1995; Constable et al. 1998) Wolves. Our analyses suggest- ed pup P concentrations were higher than adults. Messier (1987) observed lower P concentrations in adults as well. Calcium concentrations were similar to reported values (DelGiudice et al. 1987 in addition to those cited above). However, no other Wolf study reported pup Ca values but we observed pup Ca con- centrations were higher than adults. We also observed higher SAP concentrations in pups than adults. This has been observed in Gray Wolves (Messier 1987), Coyotes (Smith and Rongstad 1980), and Dogs (Pick- rell et al. 1974). High SAP concentration is indicative of bone formation and osteoblast differentiation (Searcy 1969). Serum alkaline phosphatase was also correlated to age, P, and Ca. Interestingly, of adult SAP values, 11.4% (all male and sampled in different seasons) were above the lower 95% confidence interval for pup SAP values. High SAP concentration has been asso- ciated with mammary tumors in dogs (Karayannopou- lou et al. 2003), mast cell disease in humans (Pardanani et al. 2002), pregnancy in animals (Stockham and Scott 2002; Bain 2003), healing of broken bones (Searcy 1969), and other diseases in Dogs (Cornelius 1980). Thus, SAP concentration may be used to identify in- jured or diseased individuals. Conclusions New patterns in physical condition, hematology, and serum Chemistry were identified for Gray Wolves. Pups appeared to be under greater nutritional stress than adults as suggested by reduced MF, PCV, and SUN and THE CANADIAN FIELD-NATURALIST Vol. 120 Hb concentrations in pups. We also confirmed estab- lished differences in SAP concentrations between adults and pups. However, abnormally high SAP concentra- tions were observed in a portion of the adult male Wolves, perhaps, identifying injured or diseased indi- viduals. High PCV in autumn suggested Gray Wolves of this population were in their best physical condition during autumn. High SUN concentrations in spring suggested these Wolves were under little nutritional stress during spring and appeared to be on a good nutritional plane during summer, as suggested by rel- atively high summer SUN concentrations. Also, high Hb concentrations and PCV in this population were probably a result of higher elevation. These data, classified by season, sex, and age, pro- vide a baseline physical condition, hematology, and serum chemistry for free-ranging Gray Wolves with a relatively high and increasing prey base in south-cen- tral Alaska. However, more information on yearlings and summer is needed to properly understand physical condition, hematology, and serum chemistry of this Gray Wolf population. Although we found several cor- relations among BW, Hb, PCV, MF, and several other variables, a number of measurements should be used to assess physical condition. Acknowledgments The study was funded by several Alaska Federal Aid to Wildlife Restoration Projects and the Alaska Power Authority. Publication costs were paid by the Department of Natural Resources Management, Texas Tech University. We gratefully acknowledge L. Au- miller, T. Balland, A. W. Franzmann, C. L. Gardner, P. Hessing, L. Metz, R. 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Journal of Mammalogy 56: 44-63. Received 20 September 2005 Accepted 28 November 2006 Are Lesser Snow Geese, Chen caerulescens caerulescens, Exceeding the Carrying Capacity of the Fraser River Delta’s Brackish Marshes? MIKE W. DEMARCHI LGL Limited, environmental research associates, 9768 Second Street, Sidney, British Columbia V8L 3Y8 Canada; e-mail demarchi@1g1.com Demarchi, Mike W. 2006. Are Lesser Snow Geese, Chen caerulescens caerulescens exceeding the carrying capacity of the Fraser River delta’s brackish marshes? Canadian Field-Naturalist 120(2): 213-224. Brackish marshes of the Fraser River delta provide important habitats for such high-profile animals as White Sturgeon (Acipenser transmontanus), Pacific Eulachon (Thaleichthys pacificus), Pacific salmon (Oncorhynchus spp.), Western Sand- piper (Calidris mauri), and Lesser Snow Goose (Chen caerulescens caerulescens), the latter comprising the “Fraser-Skagit™ segment of the Wrangel Island (Russia) population. This study assessed whether the current numbers of Snow Geese are exceeding the carrying capacity of brackish marshes in the Fraser River delta. Simulation modelling predicts that those marshes are presently capable of supporting ~17 500 Snow Geese—a value that is greatly exceeded by the numbers of geese that have over-wintered there in recent years (~80 000 in 2004-2005). The Pacific Flyway Council's target 3-y aver- age population and segment sizes of 120 000 and 50 000 - 70 000, respectively, were set without considering the carrying capacity of natural wintering habitats, the potential impacts of too many geese on upland agriculture, or implications for hazards to civilian aircraft at Vancouver International Airport. The modelled results of the present study suggest that the Fraser River delta can sustain the current numbers of Snow Geese that stage or winter there only if those birds also forage in agricultural and refuge fields—a relatively recent phenomenon that likely bolstered the Snow Goose population. Over-use by Snow Geese can degrade the productivity and habitat quality of marshes. There is documented evidence that some key plant species (e.g., Scirpus americanus) of the brackish marshes of the Fraser River delta are well below their biomass potential (~15%), primarily because of grubbing by Snow Geese. Other species that depend on this brackish environment as well as human interests in the Fraser River delta may be adversely affected by an overabundance of Snow Geese. The future effectiveness of hunting as a primary means of population regulation is questioned. Key Words: Lesser Snow Goose, Chen caerulescens caerulescens, American Three-square Bulrush, Scirpus americanus, brack- ish marsh, carrying capacity, Fraser River delta, bird strike, British Columbia. The recent dramatic increase in numbers of the mid- continent (North America) Lesser Snow Goose (Chen caerulescens caerulescens) has attracted considerable attention— mainly because of the habitat degradation it has caused (e.g., Kerbes et al. 1990; Williams et al. 1993; Ankney 1996). Consumption of agricultural foods (which may be more abundant and contain high- er fractions of metabolizable energy than native brack- ish-marsh plants) and a network of refugia are thought to have contributed to substantial increases in some North American populations of Lesser Snow Geese by improving overwinter survival (Ankney 1996; Jefferies 1997). Although the bird community that winters on the Fraser River delta contains many species of graz- ing waterfowl, the large amount of rhizome biomass consumed annually by Lesser Snow Goose (Boyd 1995) means it is potentially a keystone species there as it is elsewhere (e.g., Kerbes et al. 1990). Many Snow Geese that breed on Wrangel Island (Russia) use the Fraser River delta as staging or win- tering habitat between October and April (Subcom- mittee on White Geese 1992*; Boyd 1995; Pacific Fly- way Council 2006*). They forage in brackish marshes of the Fraser River delta, primarily on rhizomes of American Three-square Bulrush (Scirpus americanus) and shoots of Lyngbei Sedge (Carex lyngbyei). J. P. Hatfield (personal communication, cited in Campbell et al. 1990) noted that Snow Geese began foraging in agricultural fields on the Fraser River delta in 1977. The number of Snow Geese using those fields rose from 2000 in the early 1980s to between 20 000 and 30 000 in the 1990s (Boyd 1995; Canadian Wildlife Service et al. 1999*). The Canadian Wildlife Service (CWS) and local farmers actively enhance the Alaksen National Wildlife Area (ANWA) on Westham and Reifel islands in the Fraser River delta for wintering Snow Geese and other waterfowl. Fields within the ANWA are planted with crops to provide the birds with supplemental feed (Pacific Flyway Council 2006*). Because Snow Geese and other grazing waterfowl! can affect private agricultural fields outside the ANWA by consuming grass, crops, or crop residue such as corn and potatoes, and by altering surface-water drain- age, the “Greenfields Program” administered by the Delta Farmland and Wildlife Trust (DFWT) was estab- lished in 1990. The goal of that program is to spread waterfowl impacts over a larger area, thereby lessen- ing their severity in any one given area (Smith 1996*). The program encourages farmers to plant cover crops for soil conservation and provide forage for over-win- o>) 214 tering waterfowl. The costs of these cropping practices are shared among farmers, conservation societies, and government bodies. Snow Geese that overwinter on the Fraser River delta are managed according to the Pacific Flyway Council’s 2006 management plan (Pacific Flyway Council 2006*) which is a revised version of the 1992 plan (Subcom- mittee on White Geese 1992*). Although the overall Wrangel Island population target of 120 000 birds (3- year average) remains the same between the 1992 and 2006 plans, the target size of the Fraser-Skagit seg- ment (also referred to as “subpopulation” or “flock’”’) target was increased to a range of 50 000-70 000 birds in the 2006 plan. According to the 1992 plan, the seg- ment target was a minimum 3-year average of 35 000 birds. To achieve that, the ideal segment size had been identified to be between 30 000 and 60 000 birds (S. Boyd, personal communication). The historical size of the Wrangel Island population was considered when the goal was set, but some key factors were not: the carrying capacity of the Fraser and Skagit deltas, the possible impacts of Snow Geese on habitat quality and ecosystem function of the Fraser River delta, nor the possible impacts of Snow Geese on agriculture or air- safety. Under the 2006 plan, regulation of the Wrangel Island population is to occur by way of sport and sub- sistence hunting. Burton (1977) concluded 30 years ago that the brack- ish marshes of the Fraser River delta would be damaged if the Fraser-Skagit segment exceeded 20 000 Snow Geese. The purpose of this study was to evaluate, by way of simulation modelling, recent numbers of Snow Geese in light of the carrying capacity of the brackish marshes in the Fraser River delta. Study Area The Fraser River sustains the largest delta on Can- ada’s Pacific coast. The delta covers ~680 km? of aquat- ic and terrestrial areas, stretching 30 km from New Westminster westward to Sand Heads Lighthouse and from Point Grey southward to the international bound- ary (Butler and Campbell 1987). Brackish marshes occupy ~3% of the delta. The Fraser River delta is a critical link in a series of migratory bird habitats along the Pacific Flyway between the arctic breeding grounds of North America and northeast Asia and wintering areas in southwestern North America, Central America, and South America. The delta supports some of the highest densities of wintering waterfowl, shorebirds, and raptors in Cana- da (Butler and Campbell 1987). Brackish marshes of the Fraser River delta have evolved as a result of the interactions between the maritime conditions of the Pacific Ocean and the fresh water and sediment load of the Fraser River (Hutch- inson 1982). Local tides are predominantly semidiurnal with a maximum range of ~5 m. The Fraser River is the largest river in British Columbia, with a mean an- THE CANADIAN FIELD-NATURALIST Vol. 120 nual flow of 3500 m?/s. Eighty percent of the annual flow occurs during March to July (Hoos and Packman 1974, cited in Hutchinson 1982). Hutchinson (1982) divided the present brackish marshes of Lulu Island into three elevational zones: (1) low marsh—dominat- ed by Scirpus americanus and S. maritimus; (2) middle marsh—dominated by Carex lyngbyei, Triglochin mar- itimum, and S. maritimus; and (3) high marsh—a com- munity of Agrostis exerata, Potentilla pacifica, Dis- tichlis spicata, and Typha latifolia. Prior to dyking, the low elevations of much of the land near the western limit of the delta were subject to periodic flooding during spring freshet. Such flood- ing would have affected the delta’s vegetation commu- nities. Habitats of this area have been, and are heavily, modified by human activities. Although much of the Fraser River delta is used for agriculture, residential, commercial, and industrial developments are rapidly increasing at the expense of agricultural land-uses. Commercial greenhouse operations are increasingly replacing traditional field-based agriculture. Much of the upland and foreshore areas of Reifel Island and Westham Island make up a wildlife sanc- tuary that is managed by the CWS. The sanctuary com- prises 586 ha, of which 300 ha includes the federally owned ANWA that was designated a Ramsar site in May 1982 (Wetlands International, no date*). Approx- imately 410 ha of the sanctuary are in cultivated farmland. The adjacent George C. Reifel Refuge, ini- tiated in the 1960s, consists of 400 ha of Crown land designated as a Migratory Bird Sanctuary under the Migratory Birds Convention Act of 1917. Nearly 6000 ha in the municipality of Delta is used to produce vegetable crops and livestock. Of the crop- land, nearly two-thirds is planted in vegetables such as potatoes, peas, beans, corn, pumpkins, cabbage, rutaba- gas. Hay fields, pasture, and corn silage represent more than one-quarter of the cropland on the delta. The re- mainder of the cropland on the delta is allocated to cereals and berries. Methods A map of the brackish marshes was prepared by digitally scanning colour air photos (1:24 000 scale) at high resolution and ortho-rectifying each photo by tying control points to those on a TRIM base-map (1:20 000 scale). Air photos from 18 September 1991 were deemed the best in terms of resolution, scale, date, season, and tide height. I. Hutchinson (personal communication) indicated that the brackish marsh profile has likely undergone very minor net changes since that time. Habitat polygons were digitized man- ually on-screen while viewing habitat features at var- ious magnifications. The digital map upon which habi- tat calculations were made had a minimum resolution of 1:10 000. Small patches of Scirpus spp. near south- western Sea Island were amalgamated because it was believed that the distribution of those plants had in- 2006 creased since the photos were taken (S. Boyd, per- sonal communication). Interpretations of habitat type and zonation in the Fraser River delta were aided by information from Yamanaka (1975), Moody (1978), and Hutchinson (1982). Although changes in the marsh communities have undoubtedly occurred since maps in those reports were produced, the visual patterns of intertidal plant zonation were expected to have changed insignificantly. All mapping was done using the GIS programs MapInfo and ArcView. Estimates of Snow Goose numbers since 1987-1988 were obtained from Boyd (1995) and Pacific Flyway Council (2006*). The numbers of Snow Geese using agricultural fields were obtained from Boyd (1995) for the period 1987-1992, and from Canadian Wildlife Ser- vice et al. (1999*) for the period 1995-1998. Data on the Greenfields Program were obtained from Smith (1996*). Because of variation and error inherently associated with parameters used in the analyses, a stochastic sim- ulation model was developed. The model, run in MS Excel 98, incorporated published and unpublished data from several sources. Data in Imperial units (e.g., kcal) were converted to SI units (e.g., kJ). Stochasticity was incorporated in the model by randomly selecting val- ues that were within the 95% confidence limits, or hypothesized range of some parameters. The model in Appendix | was used to estimate the carrying capacity of the brackish marshes in the first half of the overwintering period (“autumn”, versus the second half of the overwintering period: “spring”). Because the Fraser River delta experiences substan- tially more Snow Goose-days during autumn than in spring (Boyd 1995) and has less autumn habitat than spring habitat (as mapped in this study), I assumed that any habitat-imposed limitations on the geese would occur in autumn. Each simulation first calculated the metabolizable caloric value of the available S. ameri- canus. From that value, the number of supportable Snow Goose-days was determined. In turn, Snow Goose-days (y) were used to estimate the size of the Fraser-Skagit segment (x) by using the inverse of the linear relation: y = 47.764(x) (P = 0.028; r = 0.89; df = 5; y-intercept = 0) between “autumn” Snow Goose-days on the Fraser River delta and the size of the Fraser-Skagit segment as calculated from data presented in Boyd (1995). The y-intercept of this equa- tion was set at 0 to account for segment sizes lower than those presented in Boyd (1995). Five thousand simulations were run. A lack of data on two issues may have implications for the accuracy of the model’s results, but because the parameters represented by these issues would not affect the model in a unidirectional manner, their impli- cations are not viewed as necessarily troublesome. First, Burton et al. (1979) suggested that Snow Geese feed almost exclusively on rhizomes of S. americanus during autumn. In fact, however, Burton (1977, page DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA 17) stated that Snow Geese strongly prefer both S. americanus and §, paludosus (the latter also known as. S. maritimus). Unfortunately, as observed using giz- zard contents, Burton (1977) could not differentiate between the cuticle reticulations of the rhizomes of the two species, thereby casting doubt on the accuracy of his conclusion about a dual food preference. Abun- dance of S. maritimus is about one-third that of S. americanus on the Fraser River delta (Yamanaka 1975). Second, the model assumes that all biomass grubbed by the Snow Geese is consumed by them. MclIlhenny (1932; cited in Burton 1977) guessed that over-win- tering Snow Geese in the southern United States re- jected up to 10 times as much grubbed vegetation as they consumed. Burton (1977) felt that although some food rejection occurred on the Fraser River delta it did not likely approach those proportions. Results and Discussion Distribution, Abundance, and Carrying Capacity of Brackish Marshes in the Fraser River Delta Autumn marsh habitat that is suitable for Snow Geese occurs in a narrow band along the western edge of the Fraser River delta’s westernmost islands and near Brunswick Point (Figure 1). Most spring marsh habitat is similarly distributed. According to habitat mapping, the Fraser River delta contains 917.1 ha of “low marsh” habitat and 946.3 ha of “mid-high marsh” habitat in areas used by Snow Geese. Low marsh habi- tat is used by Snow Geese for “grubbing” bulrush rhizomes in the autumn period and mid-high marsh habitat is used in the spring for foraging on emergent sedges (Boyd 1995). The results of autumn carrying capacity simulations of the Fraser River delta’s brackish marshes are sum- marized in Figure 2. During autumn, brackish marshes can presently support a number of Snow Goose-days that is commensurate with a mean Fraser-Skagit seg- ment size of 17 600 geese (range 7400-35 000). Num- bers larger than this will exceed the calculated carry- ing capacity of brackish marshes during most years. According to the model there was only a 1.4% chance that >30 000 geese could be sustained by the brackish marshes (Figure 2). Those observations, and the conclu- sion that §. americanus biomass in the Fraser River delta is only ~15% of what it might support in the ab- sence of Snow Goose grubbing (Boyd 1995), suggest that brackish marshes are either already over-grubbed or are at considerable risk of being over-grubbed. Distribution and Abundance of Snow Geese on the Fraser River Delta Numbers of Snow Geese in the Fraser-Skagit seg- ment have been estimated from aerial photos of geese on the Fraser and Skagit river deltas (Subcommittee on White Geese 1992*; Boyd 1995; Pacific Flyway Council 2006*). Most use of the Fraser River delta oc- curs during early October to mid-January (“autumn”) and from late February to mid-April (“spring”). Al- 216 THE CANADIAN FIELD-NATURALIST Vol. 120 : Mainland Ser i 130° } 125° 120° 115° : LOE NANO = ‘ ss (Vancouver) | \af. a Point, Grey WS8\ British (6 km) : pe. 55° , Columbia | ajperta | | I Sea Island a YVR Rich d) ZIG (Richmon t e(Richmoend Rete aS Low Marsh Lulu Island 4 Mid-High Marsh (Richmond) yued uoebinis 2 ; 4 a ————— Sand : q Kilometres Heads ae ~ 9% Lighthouse Woodward Island re ae ; Complex Westham Island (Delta) Mainland International Y Brunswick Point (Delta) Boundary (~7 km) — Figure 1. Distribution of intertidal marsh habitats used by Snow Geese during the autumn and spring periods on the Fraser River delta, British Columbia. Runways of the Vancouver international Airport (Y VR) are indicated. 2006 DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA 217 Frequency S Fraser-Skagit Sub-Population Size (birds) FiGuRE 2. Simulation model results of the size of the Fraser-Skagit segment that could be supported by the brackish marshes of the Fraser River delta for Snow Geese during the autumn. 95% confidence interval on mean: 17 606 +131 (n = 5000). 120000: _——_<_—__—$_——————____—__———_____“_- 100 000 ° 0 ® 30000 , o se ® P : oO : 2 el © 60000 - b—0 —_ ro) iS) 5 > 40 000 - 20 000 - fo} o N a \o co fom) N wv \o oo o N vT ~ io) co co foe) oo fo aD oa a a —) So = aN fon) fon) lon an an fo fo) a an nN —) o oC ee Maa) ap eh oet AE aad i = = = a A Q Year /_o— Wrangel Island Population oe Fraser-Skagit Segment | FiGurE 3. Estimates of the Wrangel Island Lesser Snow Goose Population size (total, spring) and size of the Fraser-Skagit segment (overwintering period beginning in year shown). Source: Pacific Flyway Council (2006*). 218 6 000 000 - 5 000 000 4 000 000 3 000 000 Goose Days 2 000 000 | ~ 1 000 000 measured 1987-88 1988-89 1969-90 1990-91 1991-92 1992-93 1993-94 THE CANADIAN FIELD-NATURALIST 1994-95 - 1995-96 1996-9? Vol. 120 O Autumn & Spring predicted 1997-98 - 1998-99 1999-00 - 2000-01 2001-02 2002-03 - 2003-04 2004-05 Year FiGuRE 4. Stacked area chart showing measured and predicted number of Snow Goose-days on the Fraser River delta since 1987-1998. Data from 1987-1988 through 1991-1992 were measured by Boyd (1995). The remaining data were pre- dicted using linear regression equations developed from data for the 1987-1988 through 1991-1992 period (Boyd 1995), and using population estimates (x values) from Pacific Flyway Council (2006*). Regression equations are as follows: (1) Autumn: y = 36.4(x) + 448 977 where y = autumn goose-days, and x = annual estimate of Fraser-Skagit segment (P =0.016, r = 0.94, df = 4). (2) Spring: y = 22.7(x) - 221 987 where y = spring goose-days, and x = annual estimate of Fraser-Skagit segment (P = 0.116, 7 = 0.78, df = 4). though geese occur on both deltas simultaneously during autumn and spring, most or all geese use the Skagit River delta for the four to five weeks between these periods. Since the winter of 1978-1979 (when the first photo counts on the Fraser and Skagit deltas were done) the Fraser-Skagit segment grew from ~27 000 birds (Pacific Flyway Council 2006*). Owing to high juvenile production and survival rates of Snow Geese, the Fraser-Skagit segment increased 3- to 4-fold in the late 1970s (Boyd 1995). Numbers of Snow Geese fluctuated in the 1980s then increased thereafter (Fig- ure 3). Figure 3 shows an upward trend in the size of the Fraser-Skagit segment from 1978 through spring 2005 (P < 0.001; r = 0.76; df = 26). Recent surveys indicate that during the winter of 2004-2005 the seg- ment was the highest ever recorded at ~80 000 birds (Pacific Flyway Council 2006*). From 1987-2005 the Fraser River delta sustained an estimated 2-5 mil- lion goose-days annually (Figure 4). Farm fields apparently play an increasingly impor- tant role in the over-wintering ecology of the Fraser- Skagit segment of Snow Geese. The percentage of Snow Goose days spent on upland fields between 1987- 1988 and 1991-1992 varied among years (Figure 5), but autumn-use, ranging between 7% and 30%, was typically much greater than spring use, which ranged from 0% to 20% (Boyd 1995). Although surveys of use of agricultural fields on Westham Island were conduct- ed via ground counts from autumn 1995 through spring 1998 (cf. photo counts up to that point), the use of farm fields has increased in recent years. From 1995-1996 through 1997-1998, the use of farm fields on Westham Island (representing ~90% of all use of upland fields (S. Boyd, personal communication) ranged between 17% and 46% of the total number of goose days spent on the Fraser River delta in the autumn, and increased from 13% to 70% of such days in the spring (Figure 5). If foods in brackish marshes were being fully uti- lized, such a pattern would be expected if more goose- days were spent on the Fraser River delta. Indeed, Figure 4 shows a predicted increase in goose days over time and the linear relation between annual goose days on agricultural fields and size of the Fraser-Skagit segment is significant (P = 0.002, r = 0.83, df = 7); the lowest proportion of field-use occurred when the least number of goose days were spent on the Fraser River delta (1989-1990; Figures 4 and 5). Snow Geese and the Brackish Marsh The brackish marshes of the Fraser River delta are probably being over-utilized by Snow Geese. Model results indicate that the Fraser-Skagit segment exists at current levels only because the geese can also for- age in upland fields. Maintaining numbers of Snow Goose above the carrying capacity of brackish marshes by way of providing supplemental feed on refuges and agricultural fields may degrade or continue to suppress the productivity of the brackish marshes. Boyd (1995) suspected that grubbing pressure by Snow Geese on 2006 photo counts Percent of Season Use BS S =! 1987- 1988- 1989- 88 89 90 9] DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA bhi al 1990- 1991- 219 ground counts @ Autumn 0 Spring 1995- 1996- 1997- G2 796 SOR WOR Year FiGurE 5. The estimated percentage of total Snow Goose-days on the Fraser River delta spent on agricultural fields during the “autumn” and “spring” seasons. Source: 1987-1992, Boyd (1995); 1995-1998, Canadian Wildlife Service et al. (1999*). +Spring use in 1995-1997 is likely underestimated because surveys ended several weeks before spring migration (departure). parts of the Fraser Delta was contributing to a reversal in marsh succession. Boyd (1995) also speculated that the reason that production in the brackish marshes of the Skagit River delta was even lower (i.e., 10%) than the 15% figure reported for the Fraser River delta was because the Skagit River delta supported about twice as many goose-days as did the Fraser River delta dur- ing 1987-1992. Further, there is some concern that in recent years a greater proportion of the Wrangel Island population has been overwintering on the Fraser and Skagit deltas (Pacific Flyway Council 2006*). If Snow Geese were excluded from all agricultural fields, including the sanctuary on Reifel and Westham islands, the segment would likely decline to a habitat- limited maximum size of ~15 000-20 000 birds. Al- though that scenario may be unrealistic, it is possible that changing land-uses and agricultural practices throughout the Wrangel Island population’s winter range, in addition to changes in farmers’ tolerance of Snow Geese in their fields, could direct even more for- aging pressure onto brackish marshes. Regardless of farm-field availability, brackish marshes will always be vital for Snow Geese because during extended periods of freezing, the marshes are used exclusively (Boyd 1995). Use of marshes during cold weather likely reflects difficulties geese experience in obtaining food from the frozen or snow-covered soil of agricul- tural fields. Snow Geese and Agricultural Fields Forage from agricultural fields has contributed to the improved survivorship and subsequent population growth of Snow Geese in other parts of North America (Ankney 1996; Jeffries 1997). Possible reasons for the onset of, and increased use of, upland fields by Snow Geese on the Fraser River delta are not known with certainty, but it is plausible that rapid increases in the Snow Goose population during the late 1970s were largely responsible (Figure 3). Burton (1977) predicted that if the Fraser-Skagit Snow Goose segment expanded beyond ~20 000 birds, the brackish marshes of the Fraser River delta would be adversely affected. Although estimates of S. amer- icanus biomass in the mid 1970s (Burton 1977) and in the early 1990s (Boyd 1995) are not directly com- parable due to different sampling strategies, a general review of the datasets suggests that rhizome densities during the early 1990s likely approximated those dur- ing the mid 1970s. This suggests that the brackish marshes were already being used to capacity by Snow Geese prior to the onset of farm-field use. Unfortunate- ly, there are no data documenting trends in S. ameri- canus biomass immediately following the rise in Snow 220 Goose numbers and onset of field use in the late 1970s and early 1980s so it is not known whether S. ameri- canus biomass declined under more foraging pressure then merely rebounded as incremental foraging pres- sure on the delta shifted to upland fields. The increased use of agricultural fields by the geese supports the hypothesis that the brackish marshes alone could not sustain the increased foraging pressure that a rising Fraser-Skagit segment was putting on the Fraser River delta. If the geese could not meet their energetic demands in brackish marshes, they would have been forced to explore alternate food sources such as those in agricultural fields. Several lines of evidence support this hypothesis. First, from the perspective of Snow Goose forag- ing habitat, agricultural production suitable for Snow Geese in the local area has probably declined since the 1970s, both in areal extent and crop type (Boyd 1995). So, contrary to what might be expected, little or no use occurred when agricultural habitats were better for Snow Geese. According to W. Temple (DWFT, person- al communication), there is no evidence that agricul- tural changes on the Fraser River delta influenced the initial use of upland habitats by Snow Geese in the late 1970s and early 1980s. W. Temple (personal communi- cation) indicated that crops that Snow Geese presently use were grown for many years before the birds started using them. In 1993, farming of corn, peas, and beans by commercial processors on the Fraser River delta was largely halted (W. Temple, personal communica- tion); thereby further diminishing the area of agricul- tural fields suitable for foraging Snow Geese. This like- ly put increased foraging pressure on other fields, and perhaps, on the brackish marshes. Second, S. americanus stands in the Fraser River delta are at about 15% of their biomass potential in the absence of grubbing by Snow Geese (Boyd 1995). This suggests that the brackish marshes are not pro- ducing vast amounts of unutilized rhizomatous foods. Assuming a “steady state” between rhizome produc- tion and consumption, as reported by Boyd (1995), brackish marshes are being fully utilized (even over- utilized) by consumers such as Snow Geese. Third, Snow Geese on the Fraser River delta exhibit a high degree of site fidelity (Burton 1977; Boyd 1995). Thus, the full or partial abandonment of an area could indicate decreased habitat suitability. A lack of food in the traditionally used brackish marshes is a plausible reason why Snow Geese explored upland feeding areas. Such exploratory behaviour was predicted by Burton (1977) before the segment increased and before agri- cultural fields were used by the geese. Finally, speculation that changes to hunting seasons and hunting areas in the Fraser River delta forced Snow Geese to use upland fields beginning in the late 1970s is not supported. The existence of a waterfowl refuge on Reifel Island has conceivably provided Snow Geese with upland foraging opportunities free of hunting pressure since the 1960s. Further evidence that hunting THE CANADIAN FIELD-NATURALIST Vol. 120 did not precipitate field use by Snow Geese is provided by Trumpeter Swans (Cygnus buccinator); a protected (i.e., unhunted) species since the 1930s. Trumpeter Swans and Snow Geese have similar diets during win- ter (Boyd 1995; Carter 1997). Trumpeter Swans have also experienced substantial increases in over-winter- ing populations in recent years. Carter (1997) noted that as numbers of Trumpeter Swans wintering near southwestern British Columbia have increased since the early 1970s, there has been a recent shift in habitat use from brackish marshes to agricultural areas. G. Fowler (CVWMBP, personal communication) indicated that the use of agricultural fields by Trumpeter Swans in the Comox Valley on Vancouver Island, British Columbia, began in the mid-1970s. The onset of swan use of upland habitats was unlikely a response to any refuge effects provided by fields, as hunting for other waterfowl continues to this day on the same farm fields used by swans. Further, if swans were seeking refuge, the Courtenay River estuary (their traditional brackish- marsh wintering ground that has been largely aban- doned as a foraging area in favour of upland sites) should still be used for foraging because it has been a no-shooting area since 1994. The swans roost in the estuary, but minimal feeding occurs there (G. Fowler, personal communication). Conclusions Currently, the increasing numbers of Snow Geese wintering on the Fraser River delta (~80 000 in 2004- 2005; Pacific Flyway Council 2006*) exceed the mod- elled autumn carrying capacity of the area’s brackish marshes (~17 500) by a large margin. As a result, the geese are highly dependent on forage provided in near- by agricultural and refuge fields. Despite compelling evidence that Snow Geese were grubbing S. ameri- canus biomass to ~15% of its biomass potential since at least the early 1990s (Boyd 1995) and a doubling of the Fraser-Skagit segment from 1995-2005, the 2006 management plan (Pacific Flyway Council 2006*) only states that increasing numbers of Snow Geese may be reducing the biomass and extent of S. americanus. The Pacific Flyway Council (2006* page 13) recom- mends researching the “trends in biomass and growth dynamics of American Bulrush, impacts of grubbing by Snow Geese on intertidal marshes, and the carrying capacity of the Fraser-Skagit.” The current plan relies on hunting to regulate the Wrangel Island population once it exceeds a 3-y average of 160 000 birds (Pacific Flyway Council 2006* page 32). As society’s values and interests have shifted, sales of migratory bird hunt- ing permits and participation in waterfowl hunting in British Columbia have declined steadily from 1974- 2003 (Environment Canada 2005*). Hunting oppor- tunity on the Fraser River delta has also diminished over time. Consequently, it is questionable that man- agers will be able to rely solely on hunting as a tool to effectively manage Snow Geese on the Fraser delta in the future. 2006 Given that the brackish marshes are currently being used to (present) carrying capacity, any further increas- es in the Fraser-Skagit segment will likely require: (1) a substantial increase in the production of S. amer- icanus rhizomes, (2) a concomitant increase in the use of agricultural fields by the geese, (3) natural selec- tion for lower adult body weights, and thus lower gross energetic demands, or (4) a combination of these. Recognizing the reduced production of the brackish marsh, Boyd (1995) recommended that consideration be given to increasing the productivity of S. ameri- canus by applying fertilizer. However, unless fertiliza- tion results in a yield of rhizomes exceeding that which Snow Geese could potentially consume, there is no rea- son to believe that Snow Geese would not consume fertilizer-induced growth back down to the “steady- state”. The current management plan (Pacific Flyway Council 2006*) does not identify a need to research ways of increasing the carrying capacity of the marsh as was recommended in the 1992 plan (Subcommittee on White Geese 1992*). By serving as food for herbivores and detritivores and as structural habitat (cover) for fish and inverte- brates, healthy brackish marshes in the Fraser River delta are integral to an ecosystem that supports such key animals as White Sturgeon, Pacific Eulachon, Pacific salmon, shorebirds, and Snow Geese among many others. Large numbers of Snow Goose have neg- ative implications not only for the brackish marsh eco- system, but also for human safety. Bird hazard evalua- tions have concluded that Snow Geese pose an extreme hazard to civilian aircraft at the Vancouver Interna- tional Airport on Sea Island (Demarchi and Searing 1995*). Consequently, an aggressive Snow Goose con- trol program is necessary each winter near Sea Island to mitigate that hazard. A prudent approach to managing the Wrangel Island population of Lesser Snow Geese would, at a mini- mum, be sensitive to Snow Goose conservation, poten- tial adverse effects of Snow Geese on the brackish marsh ecosystem, and the interactions between Snow Geese and agriculture, air-traffic safety, and other land uses in the Fraser River delta. Acknowledgments Many people made important contributions to this study and manuscript. Their assistance is greatly appre- ciated, but such assistance does not necessarily connote full agreement with the manuscript’s conclusions. S. Boyd, Canadian Wildlife Service (CWS) provided important data and reviewed drafts of this manuscript. K. Moore and M. Porter, CWS, also provided impor- tant data (Canadian Wildlife Service et al. 1999). S. Smith, Delta Farmland and Wildlife Trust (DFWT), provided data. W. Temple, DFWT, provided informa- tion about agriculture. C. Levings, Department of Fish- eries and Oceans and I. Hutchinson, Simon Fraser Uni- versity, provided information and advice on brackish DEMARCHI: SNOW GEESE OF THE FRASER RIVER DELTA marshes in the Fraser River delta. R. Alisauskas, CWS, provided information on the current status of bioen ergetic studies of Lesser Snow Geese and gave con- structive feedback on the simulation model. R. Trost, U.S. Fish and Wildlife Service, provided information about the Pacific Flyway Management Plan. G. Fowler, Comox Valley Waterfowl Management Project, provid- ed information about Trumpeter Swans in the Comox Valley. D. Kraege, U. S. Fish and Wildlife Service, reviewed drafts of this manuscript. S. Johnson and G. Searing, LGL Limited, contributed to the design of this assessment and reviewed a draft of this manu- script. R. Tamasi, LGL Limited, assisted with GIS and mapping. G. Glova, LGL Limited, reviewed a draft of this manuscript as did A. Erskine and an anony- mous reviewer. Documents Cited [marked * in text citations] Canadian Wildlife Service, Ducks Unlimited, and the British Columbia Waterfowl Society. 1999. Westham Island winter waterfowl surveys 1995-1998. Unpublished data. Delta, British Columbia. Demarchi, M. W., and G. F. Searing. 1995. Avian ecology and air-traffic safety at Vancouver International Airport. Unpublished report for Transport Canada, Ottawa, Ontario. 100 pages. Available from University of British Columbia Library. Environment Canada. 2005. Data accessed via the INTER- NET. http://www.ecoinfo.org/env_ind/region/ducks/ducks_ data_e.cfm#Graph2 Last updated 14 June 2005. Pacific Flyway Council. 2006. Pacific Flyway management plan for the Wrangel Island population of lesser snow geese. Revised Draft. White Goose Subcommittee, Pacific Flyway Study Committee. [c/o USFWS], 911 N.E. 11" Avenue, Portland, Oregon. Accessed via the INTERNET. http: pacificflyway.gov/Documents/Wilsg_plan.pdf Smith, S. 1996. The Greenfields project 1995/96 historical database on cover crops. /n The Greenfields Project 1995 *96: an investigation of cover crops and waterfow! use in the Boundary Bay area. Edited by S. Smith. Unpublished report by the Delta Farmland and Wildlife Trust, Suite 205-4882 Delta Street Delta, British Columbia V4K 2T8. Subcommittee on White Geese. 1992. Pacific Flyway man- agement plan for the Wrangel Island population of Lesser Snow Geese. Pacific Flyway Study Committee, Portland, Oregon. 25 pages. Wetlands International. No date. A directory of wetlands of international importance: Canada Ramsar Site 243. Infor- mation accessed via the INTERNET. http://www.wetlands. org/RSDB/_COP9Directory/Directory/4C A009 html Literature Cited Ankney, C. D. 1996. An embarrassment of riches: too many geese. Journal of Wildlife Management 60: 217-223. Boyd, W. S. 1995. Lesser Snow Geese (Anser C. caerules- cens) and American Three-square Bulrush (Scirpus amer- icanus) on the Fraser and Skagit River deltas. Ph. D. dis- sertation, Simon Fraser University, Burnaby, British Columbia. 154 pages. Burton, B. A. 1977. Some aspects of the ecology of Lesser Snow Geese wintering on the Fraser River delta tidal marshes. M.Sc. thesis, University of British Columbia, Vancouver, British Columbia. 173 pages. DD Burton, B. A., R. J. Hudson, and D. D. Bragg. 1979. Effi- ciency of utilization of bulrush rhizomes by Lesser Snow Geese. Journal of Wildlife Management 43: 728-735. Butler, R. W. and R. W. Campbell. 1987. The birds of the Fraser River delta: populations, ecology and international significance. Occasional Paper 65. Canadian Wildlife Ser- vice, Ottawa, Ontario. 73 pages. Campbell R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. Kaiser, M. C. E. MecNall. Editors. 1990. The birds of British Columbia. Volume Two. Royal British Columbia Museum, Victoria, British Columbia. 636 pages. Carter, B. P. 1997. Winter habitat use by Trumpeter Swans (Cygnus buccinator) in the Fraser River delta, British Columbia. M.Sc. thesis, Simon Fraser University, Burnaby, British Columbia. 80 pages. Frederick, R. B., and E. E. Klaas. 1982. Resource use and behaviour of migrating Snow Geese. Journal of Wildlife Management 46: 601-614. Frederick, R. B., W. R. Clark, and E. E. Klaas. 1987. Be- haviour, energetics, and management of refuging water- fowl: a simulation model. Wildlife Monographs 96. Hoos L. M., and G. A. Packman. 1974. The Fraser River Estuary: status of environmental knowledge to 1974. Fish- eries and Marine Service, Pacific Environment Institute. Special Estuary Series, Number. 1. Hutchinson, I. 1982. Vegetation-environment relations in a brackish marsh, Lulu Island, Richmond, B.C. Canadian Journal of Botany 60: 452-462. Jeffries, R. L. 1997. Long-term damage to sub-arctic coastal ecosystems by geese: ecological indicators and measures of ecosystem dysfunction. Pages 151-165 in Disturbance and recovery in arctic lands. Edited by R. M. M. Craw- ford. Kluwer Academic Publishers, Netherlands. THE CANADIAN FIELD-NATURALIST Vol. 120 Kerbes, R. H., P. M. Kotanen, and R. L. Jefferies. 1990. Destruction of wetland habitats by Lesser Snow Geese: a keystone species on the west coast of Hudson Bay. Jour- nal of Applied Ecology 27: 242-258. King, J. R. 1974. Seasonal allocation of time and energy resources in birds. Pages 4-70 in Avian energetics. Edited by R.A. Paynter. Nutall Ornithological Club, Cambridge. Mcllhenny, E. A. 1932. The blue goose in its winter habitat. Auk 49:279-306. McKelvey R., M. Bousfield, A. Reed, V. V. Baranyuk, and R. Canniff. 1989. Preliminary results of the Lesser Snow Goose collaring program on the Alaksen National Wildlife Area 1986 and 1987. Canadian Wildlife Service Progress Note Number 183, Ottawa, Ontario. 5 pages. Moody, A. I. 1978. Growth and distribution of the vegeta- tion of a southern Fraser River delta marsh. M.Sc. thesis, University of British Columbia, Vancouver, British Colum- bia. 153 pages. Sedinger, J. S., R. G. White, and J. Hupp. 1995. Metabo- lizability and partitioning of energy and protein in green plants by yearling Lesser Snow Geese. Condor 97: 116- 12? Williams, T. D., E. G. Cooch, R. L. Jeffries, and F. Cooke. 1993. Environmental degradation, food limitation and re- productive output: juvenile survival in Lesser Snow Geese. Journal of Animal Ecology 62: 766-777. Yamanaka, K. 1975. Primary productivity of the Fraser River delta foreshore: yield estimates of emergent vegetation. M.Sc. thesis, University of British Columbia, Vancouver, British Columbia. 133 pages. Received 10 November 2005 Accepted 20 September 2006 OF THE FRASER RIVER DELTA JEESE : SNOW MARCHI DE 2006 Os oO=2 -¢ =Jp ‘9 = 1daouaqut-A ‘(6661 ) pPAOG Woy Req jJUdUBas WFYS-osei4, POL Lp = Josey ur sAep 28008 UUININE [UOISsausar Suisn payeypnsyey pareynoed uonezipnn aja[dwoo sauinsse | Jo anyea y paienoyea Apmis stqy, (S661) PAog paeynoyea paeynoyea (6261) ‘[e 19 uoLNg (uonRoTuNUWOS jeuosiad *pAog *S) B]]ap JOATY Jase oy) UO 2SOY] UPY) 2ANDe OW aq O} Peaatjaq ae (ZR6]) SERN puke youapaiy Aq parpnys asaas ‘PION “(TR61) SEELY pue YUapasy JO / ae WOJJ anyeA URI ay) pue APATIOR JO [2AQ] _ MO], B UQAMNIOG SANyRA JO IsuvI se UdyR]L (S661) ‘Te 19 Ja8uIpag (S661) “[e 19 JasuIpag (LQ61 ‘Te 19 YOUepaty Ul Pard) pL6T Sury (6861) “Te 19 Aaapayo (6861) “Te 19 Aaalaypyy (S661) PAog JUSWIWIOD/201No0g (POL LE/(89%((S9+99)4(ZO- 1D) +(L9%T79)))/91D))) = S1O*60%P LO = OT OO000I*E1O*«71D = VL16 0001/(9E' 7I)NAAMLAPAGNVYU = 60/LO = 60/9) = (00L9'098S)NAAM.LAPGNVAa = OIALT TDNAFMLAPAUNVa = bOx((1L8' SSh)NFAMLAGGNV®) = SO-€9x((1L8°SSp) NSAMLAPAGNVY) = 60¢ ceO'C LECT OIG ONFEMLAPGNVY = (D uUINJOS) onyeA spsiq MI JUDIOYJAO9 pexy A/3y By A/,Wl/3 Avp/3¥ Aep/3¥ 34/0 JUDIOYJOOS a[qQuLreA P/F ynpe/sytusant s}uQ) uoneindod asoor Moug 13eBYS J0se14 PISIA SNUDILLAUD *§ JO ANIRA ILIOTRO B[qeZI[OqRIsI\\ pouinsuoo SSRWOIG SNUDILIAUD *§ paqqnis Jo uOHIOdOIg PISIA SNUDIIAAUUD *§ BIJAP JOATY JOse1, [BIOL (YSIBUI MO]) SOYSILU SNUDILAAUUD *§ JO BAIR [BIO], Path snupoisaup *§ ayeis Apeays DOURUDIUIRIY S[IUSANL :pouunsuod snUuDpI1IAUD ‘¢ JOURUDIUILIY I[NPY :Pauinsuod snUDILIAUD *§ SOWOZIYI SnUuDILAAUD SNd1IG JO KB1BUS I{QIZI[OgrIDIN| J0ud}SIX9 dAogeR JUoWIAIOUT AWANOY afusane uvoyy :— 0 Ie Adios BOUN}SIXY ynpy ues ‘OD 0. 1e Adious doUN]sIxq A310uU9 Q0U9d)STxo ][Npe pur sjluaanf usamjoq sdUdIOIFIG 1y310Mm Apog urou o[TuoANe Wy31em Apog uvow inpy (2[qeLIeA) soptucant uonsodoig Jojoureleg MOY ne OOOO —————————————————E—EEEEEEEEeeeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeeeee__ nnn "86 [99XH SIN WIM poulloy -Jad aia SUONR[NOTRS [apoyy “UWININe ay) SuLINp assay MOUS Jasse] JOJ BaP JOATY JosesJ oy) JO soyssew ysryoesq ay} Jo Aytoedeo SurAsed oy} JO} [OPOU UONR[NUITS *] XIGNAddy 224 ADDENDA Last Flight? My third, and supposed final, flight of 10 Novem- ber 2005 was in a De Havilland Dash 8 that was to take me home from a journey that began a few hours earlier in a 6-seat Piper Navaho that departed from a snow- covered gravel strip in northern B.C., then continued in a Bombardier CRJ from Prince George to Vancouver (YVR). The westbound departure from YVR’s runway 26L would take us over the marsh along the foreshore of Sea Island. It was like so many others I’ve been on since 1994 when I began studying bird-hazards to air- craft at YVR and helped develop their 24/7 wildlife control program to mitigate those hazards—many of which I had witnessed first-hand. There I was, wonder- ing what critters might be flying around in the flight path this time. For night-time flights such as this, my usual thoughts turn to some poor, unsuspecting barn owl out patrolling the airfield in search of a plump vole. But this time a far more serious hazard was pon- dered: snow geese that breed in the Russian arctic and winter here by the tens of thousands. By chance, I had just submitted a manuscript for publication earlier in the week, addressing adverse effects of a burgeoning snow goose population on the Fraser Delta. And two days earlier on final approach I had spied a flock of geese loafing in the marsh near the flight path. I had heard recently that the wintering flock this year was bigger than ever. With plenty of runway lights still below us, we levi- tated steeply into the darkness. By the time we were over the marsh, we’d reached an altitude that, based on my numerous hours of field-observations from a ground station near the runway’s end, led me to believe that we had cleanly escaped the main bird-hazard zone yet again. Still, I peered out the right side at the faintly illuminated propeller hauling us into the blackness. At some 300+ km/h in the dark there was no chance for more than a glimpse of three or four whitish blurs, accompanied by subtle impact vibrations so transient that I can’t remember if they were heard, felt, or both. Snow geese? Gulls? “Shit!” seemed like the right thing to say, so I did as I sat there with a helpless feeling wondering what, if any, damage had been done, and on tenterhooks bracing for more white blurs that, luckily, didn’t appear. THE CANADIAN FIELD-NATURALIST Vol. 120 As the craft began its usual roll toward Victoria I said to myself, “Okay, maybe I’m making this out to be a bigger deal than it is,’ thinking that my height- ened awareness of bird hazards probably causes me to fret more than I should. Except this time, the turn continued a bit longer so that instead of heading over the Strait of Georgia, we were heading southeast over the lights of Delta. I knew something was amiss. Then the captain announced that we’d hit some birds and that we were going back to YVR for an “inspection”. Well, at least the plane sounded okay and I couldn’t see any evidence of severe damage like fire or smoke. Touch- down, taxi, stop, shutdown, back home—sort of. Sitting on the apron and watching an aircraft mechanic inspect the wing, cowling, and what looked to be a bloodstained propeller (I surmised that blood does not show up so clearly on a black background) some questions arose. Can his flashlight look-see really pronounce the plane safe? Should I just grab my gear, walk off this plane and board another? Is it too late to catch a ferry? I sat there knowing that bird-strikes are not that uncommon and that by choice or not, the sim- ple fact that I choose to fly means that I must have some faith in our air industry’s standards and its technicians and mechanics. Besides, what could I know of the con- dition of any other aircraft in the fleet? When the cap- tain announced that the plane was damaged and that we would have to deplane and board another, well, let’s just say that I was not put out. As deftly handled by the gate crew on what was already a very busy evening for regional air-travel out of Gate C38, within an half an hour 37 of the original 50 of us were lining up to board a shorter Dash 8. The others might seek the calming benefit of a pint or two before catching their ride. On the way out of the cov- ered walkway I engaged the ground crewman’s humour about this time we should avoid the “shredded tweet”; taking it as a reference to the fact that this mustn’t be a particularly unique event if it’s got its own pun. Yet I must say that his humour was tempered somewhat by my second-hand knowledge of the deadly havoc a few birds can wreak when they strike the “wrong” parts of a fully loaded, outbound plane. I’m happy to report the most exciting thing about the fourth of my supposed three plane rides that day was the “free” candy. Home safe (but a little shaken) in time for dinner. My next trip to YVR will be on Thurs- day. Faith be with me. MIKE W. DEMARCHI 12 November 2005 ——————————— LL Lt s—S Predicting Raccoon, Procyon lotor, Occurrence Through the Use of Microhabitat Variables RoGeR A. BALDWIN! 4, ALLAN E. HousSTON?, MICHAEL L. KENNEDY!, and Pin SHUO Liv? ' Department of Biology, The University of Memphis, Memphis, Tennessee 38152 USA * Ames Plantation, The University of Tennessee, P.O. Box 389, Grand Junction, Tennessee 38039 USA § Department of Environmental Science and Geography, William Patterson University, Wayne, New Jersey 07470 USA * Current address: Department of Fishery and Wildlife Sciences, P.O. Box 30003 MSC 4901, New Mexico State University, Las Cruces, New Mexico 88003 USA Baldwin, Roger A., Allan E. Houston, Michael L. Kennedy, and Pin Shuo Liu. 2006. Predicting Raccoon, Procyon lotor, occur- rence through the use of microhabitat variables. Canadian Field-Naturalist 120(2): 225-231. Recent increases in Raccoon (Procyon lotor) abundance have been implicated for decreased nesting success of songbirds and transmission of rabies. Understanding the relationship between occurrence and microhabitat factors should be helpful in managing this species, though our current understanding of this relationship is inadequate. Therefore, we conducted a study in western Tennessee during 2000-2002 to determine this association. Occurrence (capture) data were assessed from results of live trapping at 176 and 112 trap sites during winter and summer, respectively, at three sites. A maximum of 26 habitat variables were measured at each trap location; all grids were combined for statistical analyses to account for varying relationships between occurrence and microhabitat factors across different landscapes. Univariate and stepwise logistic-regression analyses were used to assess associations among microhabitat variables and occurrence. Resulting models were validated through the jackknife procedure. Predictive equations were constructed from logistic-regression models to compute capture probabilities. Univariate analyses yielded numerous significant variables with those representing forest characteristics and proximity to water generally the most significant. Strong concordance was observed between winter and summer seasons for most variables though several differed (number of large hardwood snags, ground dens, and plant food species, distance to potential water and roads). Such temporal variability was expected due to seasonal differences in habitat components and biological needs of Raccoons. Variables included in derived models were similar to those scoring highest in univariate analyses; classification rates for models (winter = 72%; summer = 78%) were among the highest recorded for generalist species. By accounting for landscape attributes and replicating across sites, more accurate and useful models were developed. Such models should provide the information required to effectively manage this species. Key Words: habitat partitioning, logistic regression, mesopredator, microhabitat, Procyon lotor, Raccoon, western Tennessee. Raccoons (Procyon lotor) and their associated habi- | Kennedy 1992; Pedlar et al. 1997) indicate various asso- tats have been the subject of several biological investi- gations (see Lotze and Anderson 1979; Kaufmann 1982; Broadfoot et al. 2001; Gehrt 2003). At a landscape level, this medium-sized predator (mesopredator) is thought to be most abundant in aquatic associated habitats (Johnson 1970; Minser and Pelton 1982). However, Raccoons can thrive in an array of habitats although | seeming to depend on particular features to reach high _and stable populations (Broadfoot et al. 2001; Zeveloff / 2002). Several investigators (i.e., Morris 1987; Pedlar vet al. 1997; Chamberlain et al. 2002) have noted that knowledge of spatial scale and landscape composition i is integral to understanding habitat associations of spe- ‘cies, though at present, much of the understanding of ‘the relationship of Raccoons to their habitat is based on assumptions drawn from investigations (e.g., Sanderson 1987; Minser and Pelton 1982; Oehler and Litvaitis 1996) conducted at the landscape level (macrohabitat). The relationship between the occurrence (defined as presence at a location) of Raccoons and microhabitat ‘factors has not been studied at multiple sites collec- tively and findings of previous investigations (Leberg and Kennedy 1988; Kennedy et al. 1991; Kissell and ————— i) ciations depending on season and location. Currently the relationship between the occurrence of Raccoons and habitat variables appears to be unclear and in need of additional study (Kissell and Kennedy 1992; Cham- berlain et al. 2003), and models allowing for the pre- diction of occurrence based on microhabitat factors are lacking from the published literature. Previous studies (Noss et al. 1996; Oehler and Lit- vaitis 1996; Rogers and Caro 1998) have indicated that due to a combination of factors (e.g., removal of top predators, altered land use, reduced hunting), abun- dance of mesopredators has increased in recent years. Estes (1996) noted that increased mesopredator popu- lations could influence numerous aspects of ecosystems. For example, high densities can have negative effects on populations of ground-nesting birds (Crabtree and Wolf 1988; Vickery et al. 1992; Schmidt 2003) and, by enhancing the spread of diseases, can impact negative- ly the health of populations (Carey and McLean 1983; Hill et al. 1993; Schuburt et al. 1998; Rosatte 2000). Predators play an important role in structuring biologi- cal communities (Meffe et al. 1997), and an increase in abundance of these species requires strong management 25 226 and conservation planning to limit the negative impact of these taxa on ecosystems. However, sound management and conservation plans are difficult to derive without a clear understanding of specific habitat factors critical to target species. At this time, managers of natural resources are faced with increasing popula- tions of Raccoons but limited habitat information on which to make management decisions (see Rosatte 2000 for the consideration of habitat factors in con- trolling rabies). In particular, models allowing for the prediction of Raccoon occurrence are lacking. For example, certain factors are important components in determining Raccoon occurrence (e.g., den sites and aquatic habitats, Pedlar et al. 1997; Broadfoot et al. 2001; Henner et al. 2004). However, the exact rela- tionship among these components is not known, there- by requiring individuals to make management deci- sions without the necessary information (i.e., containing rabies outbreaks, Rosatte et al. 2001). Therefore, the purpose of this study was to assess at a microhabitat scale the relationship of occurrence of Raccoons (based on capture frequency) with selected habitat factors from multiple sites representing a mosaic of habitat types. Specifically, the following predictions were assessed: (1) there is an association between occur- rence (based on capture) and selected (individual) habi- tat variables; and (2) selected habitat variables can be used to construct models predictive of species occur- rence. Study Area This study was conducted in temperate deciduous forest in western Tennessee characterized by a frag- mented landscape consisting of various levels of for- est, early successional and agricultural fields, residen- tial buildings, and road systems at three sites. Site 1 was located at the Edward J. Meeman Biological Station (Meeman), which was located approximately 20 km north of Memphis, Tennessee (35°33'N, 90°09'W). This location was comprised primarily of hardwood forests with old-field and Kudzu (Pueraria lobata) habitats interspersed throughout. Old-fields were dominated by goldenrod (Solidago spp.), fescue (Festuca spp.) and Switch Grass (Panicum virgatum). Topography of the site was characterized by numerous drainages that resulted in a gently rolling terrain throughout the area. Upland and bottomland forests included various oaks (Quercus spp.), hickories (Carya spp.), and maples (Acer spp.) as well as Tulip Poplar (Liriodendron tulipifera) and Sweet Gum (Liguidambar styraciflua; Maris 1998). Several smali ponds and intermittent streams occurred throughout the site. Sites 2-3 were located at the Ames Plantation (Ames; 35°06'N, 89°12'W), which was a 7462 ha farm located northwest of Grand Junction in Fayette and Hardeman counties, and was operated by The Hobart Ames Foundation in cooperation with the University THE CANADIAN FIELD-NATURALIST Vol. 119 of Tennessee. Ames was located approximately 79 km southeast of Meeman. Site 2 was located adjacent to the North Fork of the Wolf River. It was comprised pri- marily of agricultural fields and bottomland hardwood forest with some upland forest present. Site 3 was composed of upland and bottomland forest, cropland, and old-field habitats. In general, agricultural crops included Soybeans (Glycine max), Corn (Zea mays), and cotton (Gossypium spp.). Typical upland tree spe- cies were Loblolly Pine (Pinus taeda), oaks, and hick- ories; typical bottomland species included oaks, maples, Cottonwood (Populus deltoides), and Sweet Gum (Gabor 1993). Old-field habitats included native warm season grasses and were similar to those at Meeman. Topography of upland forest sites was characterized by gently rolling slopes, whereas bottomland forest, old-field, and agricultural areas constituted a flatter topography. Ponds and intermittent streams were num- erous, and drainages were interspersed throughout all of these sites. Methods Trapping grids were established at each site. Site 1 followed a 5 x 10 trap configuration with traps locat- ed approximately 150 m apart. Sites 2—3 followed an 8 X 8 trap configuration with traps located approxi- mately 230 m apart though two trap locations were excluded from analysis for site 2 as habitat factors were not measured. Difference in trap configuration for site 1 was due to the limited size of the area. Collectively, the three sites represented most habitat types occurring in western Tennessee. The association of microhabitat variables and occur- rence of Raccoons was assessed during two periods that coincided with times when leaves were present or absent (winter and summer, respectively) on most woody and herbaceous vegetation (Kolowski and Woolf 2002). During winter, sites were operated on selected nights from 3 November—7 April, 2000-2002; sum- mer trapping included only sites 1-2 and were oper- ated on selected nights from 7 May—12 October, 2001— 2002. An approximate total of 2000 trap nights (one trap night = one trap set for one night) were observed for each site during both seasons. Raccoon-size Tomahawk (Number 108; Tomahawk Live Trap Co., Tomahawk, Wisconsin, USA) and Havahart (Woodstream Corporation, Lititz, Pennsyl- - vania, USA) live traps were used. Traps were baited with canned cat food in winter and a combination of cat food, dog food, and doughnuts in summer. Initial- ly captured individuals were anesthetized with a mix- ture of ketamine hydrochloride (Ketaset; Bristol Lab- oratories, Syracuse, New York, USA) and acepromazine maleate (PromAce; Ayerst Laboratories, New York, New York, USA) at a 10:1 ratio with 0.1 cc of ketamine hydrochloride used per estimated kg of live weight (Bigler and Hoff 1974). Raccoons were tagged in both | 2005 ears with Number 3 Monel (National Band and Tag Company; Newport, Kentucky, USA) ear tags to deter- mine recaptures. For the winter season, 19 habitat variables were measured to determine the influence of habitat at the micro-scale on raccoon occurrence. “Slope” represent- ed the average percentage slope for a 32 m radius around the trap site as measured by a clinometer. “Total basal area” [25 cm diameter at breast height (dbh)], “basal area of small trees” (5-35 cm dbh) and “basal area of large trees” (>35 cm dbh) represented the amount of area (m*) covered by trees per hectare and were determined through use of a prism sweep (10 _ basal area factor prism) conducted at the trap site and at two additional sites 11.4 m in two random cardinal ' directions. Heights were recorded for each tree meas- _ ured during the basal area estimation using a haga alti- + meter (Forestry Suppliers, Inc., Jackson, Mississippi, _ USA) with the mean recorded as “average height”. _ “Number of fallen logs” (£10 cm in diameter) “number of total snags” (210 cm dbh), “number of small hard- _ wood snags” (10-35 cm dbh), “number of large hard- _ wood snags” (>35 cm dbh), “number of pine snags” (210 cm dbh), “number of ground dens”, “number of , tree dens”, and “number of total dens” were counted within a 32 m radius of trap site. An opening of 25 cm ' in diameter was required to be considered a potential _ den. The “number of plant food species” represented _ the number of plant species present within a 32 m _ radius of the trap site that could be utilized as food + sources. The “number of stems” 1—5 cm in diameter - was counted for a 3.2 m radius around the trap site. _ The procedure was repeated in two random cardinal _ directions 11.4 m from the trap site and the average used. “Distance to potential water” represented the Nearest distance to a water source that held water _ 230 days a year. “Distance to permanent water” was 1 _ the minimum distance to a water source that held water /211 months a year. “Distance to road” was measured to the nearest road or man-made vehicular trail, while | ieee to open area” represented the nearest dis- | ba to a non-forested patch. All distance measure- ments were in meters and were measured using digital i _ orthophoto quarter-quadrangles georeferenced in Arc- | View software. ) Seven additional variables were assessed during the Bsepiner season yielding a total of 26. “Depth of leaf j litter” was averaged from 10 random measurements jtaken within a 32 m radius around the trap site. “Verti- | Wal cover’ was assessed through the use of a2 mx 0.2m | ‘cover board checkered with 0.1 x 0.1 m black and . ‘white squares. The percentage of board uncovered was i pggorded at the trap site in all four cardinal directions. ‘The percentage of squares uncovered on a spherical ‘densiometer (Forestry Suppliers, Inc., Jackson, Missis- ; ! sippi, USA) was recorded to determine * ‘canopy cover’. ‘Readings were taken in all 4 cardinal directions at the trap site. “Grass cover”, “woody cover”, “forb cover” , | BALDWIN, HOUSTON, KENNEDY, and LIU: PREDICTING RACCOON OCCURRENCE nN nN ~ and “bare soil cover” were visually estimated for a 3.2 m radius around the trap site. All procedures for measuring summer variables (except “depth of leaf lit- ter”) were repeated in two random cardinal directions 11.4 m from trap site and the average used. Because the purpose of this investigation was to assess at a microhabitat scale the relationship of occur- rence of Raccoons from multiple sites, statistical analy- ses were conducted only on data from all sites com- bined for both winter and summer seasons. A natural log transformation was applied to all continuous vari- ables; percentage variables were arcsine transformed to approximate a normal distribution (Zar 1999). Uni- variate logistic regression was used to assess associa- tions between single habitat variables and Raccoon occurrence with a non-adjusted @ = 0.05 to indicate significance following suggestions by Moran (2003). In stepwise logistic-regression analyses, an @ = 0.15 was used as a minimum threshold for inclusion into the regression function to reduce the data set (Hosmer and Lemeshow 2000). Multicollinearity effects be- tween two significant variables were addressed by assessing correlations among habitat variables. If two significant variables were correlated at r > 0.70, only the more significant variable of the pair was included in further analyses to reduce redundant variables (Ag- resti 1996). Variables remaining after univariate logistic regres- sion were included in a backward stepwise logistic- regression function and were removed from the model at P > 0.15. Subsequent models often contained a large number of variables. Therefore, for practicality and management purposes, these models were reduced further by forcing exclusion of variables with lowest t ratios resulting in a minimum variable model. The ¢ ratio represents the ratio of each regression coefficient to its standard error. Relative importance of variables included in the final models was ascertained through t ratios, with maximal f ratios reflecting the best vari- able to predict occurrence (Hacker and Coblentz 1993; Hosmer and Lemeshow 2000; Kolowski and Woolf 2002). Percentage correct classification of trap sites was determined using logistic regression models. Accuracy of these models was determined using the jackknife procedure as a pseudo-validation technique (Morri- son 1976; Kolowski and Woolf 2002). This procedure tested percentage correct classification by removing one trap site at a time and then classified that site based on the model built from all other sites combined, result- ing in a less-biased percentage classification (Hacker and Coblentz 1993; Kolowski and Woolf 2002). All statistical procedures were conducted using SYSTAT 10.0 (SPSS 2000). Results Trapping resulted in 209 total captures of 112 indi- vidual Raccoons obtained from 176 trap sites in winter 228 THE CANADIAN FIELD-NATURALIST Vol. 119 TABLE 1. Resulting ¢ ratios and P values from univariate logistic regression of captures of Raccoons (Procyon lotor) com- pared to habitat variables at three sites during winter and two sites during summer 2000-2002 in western Tennessee. All variables are considered significant at P < 0.05. Basal area measurements are per ha. Height and distance measurements are in m. See text for explanation of variables. Winter Summer (n = 12454) (n = 8406) Variable t ratio P value t ratio P value Slope 2.517 0.012 1.878 0.060* Total basal area 4.986 <0.001 3.593 <0.001? Basal area of small trees 4.252 <0.001 3.400 0.001 Basal area of large trees 5.516 <0.001 3.447 0.001° Average height 4.052 <0.001 2.433 0.015 Number of fallen logs 4.548 <0.001? 2.653 0.008? Number of total snags 2.677 0.007 1.753 0.080? Number of small hardwood snags 1.506 0.132 1.647 0.100 Number of large hardwood snags 2.786 0.005? -0.205 0.838 Number of pine snags 2.579 0.010 1.494 0.135 Number of ground dens 1.947 0.052 0.412 0.680 Number of tree dens 3.399 0.0014 2.556 0.011? Number of total dens 3.410 0.001? 2.265 0.024 Number of plant food species 4411 <0.001? -0.046 0.963 Number of stems 2.657 0.008? 2.140 0.032? Distance to potential water -6.045 <0.001° -0.170 0.865 Distance to permanent water -2.045 0.041° -3.158 0.002 Distance to road 0.908 0.364 2.645 0.008* Distance to open area 3.669 <0.001° 2.023 0.043 Depth of leaf litter 0.699 0.485 Vertical cover -0.843 0.399 Canopy cover -3.018 0.003? Grass cover 0.182 0.855 Woody cover 1.353 0.176 Forb cover -0.742 0.458 Bare soil cover 3.052 0.002 “Variable included in stepwise logistic-regression function. (site 1 = 100 total captures of 49 individuals; site 2 = 40 total captures of 26 individuals; site 3 = 69 total cap- tures of 37 individuals) and 173 total captures of 117 individuals from 112 trap sites in summer (site 1 = 93 total captures of 52 individuals; site 2 = 80 total cap- tures of 65 individuals). Winter.— Sixteen habitat variables were significant statistically when compared to Raccoon occurrence using univariate logistic regression; greatest positive associations occurred for basal area of large trees and total basal area, whereas the strongest negative asso- ciation was for distance to potential water (Table 1). The only habitat variable not closely related to Rac- coon occurrence was distance to road. Through step- wise procedures and model construction, six variables (four positive, basal area of large trees, number of plant food species, number of tree dens, basal area of small trees; two negative, distance to potential water, number of total dens) were selected (Table 2). Per- centage of trap sites correctly classified was 72%. Summer.—Thirteen habitat variables were signifi- cantly associated to Raccoon occurrence based on univariate logistic regression; strongest positive asso- ciations were for total basal area, basal area of large trees, basal area of small trees, and bare soil, whereas distance to permanent water and canopy cover had the strongest negative associations (Table 1). In general, variables associated to basal area, height of stand, num- ber of logs, den sites, canopy cover, number of stems, . bare soil, and distances to permanent water, roads, and | open areas were associated to occurrence; variables not associated to occurrence included snags, leaf lit- ter, food species, vertical density, vegetative ground | cover, and potential water sources. Three significant | variables (two positive, total basal area, distance to ) road; one negative, distance to permanent water) were ° selected through subsequent stepwise procedures and | model construction resulting in the correct classifica- tion of 78% of trap sites (Table 2). Discussion Univariate analyses yielded numerous significant variables indicating those selected for the current in- vestigation influenced Raccoon occurrence. These vari- ables were similar to those correlated in previous inves- | tigations (e.g., association with forest characteristics, | | | 2005 BALDWIN, HOUSTON, KENNEDY, and LIU: PREDICTING RACCOON OCCURRENCE TABLE 2. Regression coefficients, ¢ ratios, and P values (P) for Raccoons (Procyon lotor) derived from logistic-regression functions, as well as percentage correct classifications using the jackknife procedure. Data included in analyses were as fol lows: winter = three sites totaling 12454 trap nights; summer = two sites totaling 8406 trap nights. See text for explanation of variables and sampling locations. Logistic regression Season Variable Coeff Winter Constant -4.605 Distance to potential water -0.251 Basal area of large trees 0.194 Number of plant food species 0.491 Number of total dens -0.389 Number of tree dens 0.381 Basal area of small trees 0.149 Summer Constant -4.605 Distance to permanent water -0.338 Total basal area 0.355 Distance to road 0.230 * Percentage correct classification of sites with no captures. » Percentage correct classification of sites with captures. © Percentage correct classification of all sites combined. Leberg and Kennedy 1988; Kennedy et al. 1991; Ped- lar et al. 1997; Dijak and Thompson 2000; associa- tion with water sources, Leberg and Kennedy 1988; _Dijak and Thompson 2000) and suggest that a strong component of forested areas with large trees and close proximity to water sources are important factors in _ predicting Raccoon occurrence. Additional variables significant in the present investigation were not sig- nificant in other studies (e.g., average height, slope, _and distance to roads — Leberg and Kennedy 1988; Kissell and Kennedy 1992). This lack of concordance may be attributed to differences in landscape compo- sition and a lack of replication (Maurer 1986; Temple and Wilcox 1986; Oehler and Litvaitis 1996). Bald- win (2003) found similar results when addressing in- dividual sites for Virginia Opossums (Didelphis vir- 'giniana) in western Tennessee. However, by assessing multiple sites, noticeable trends were observed. The lack of concordance for individual sites was attributed to differing landscape compositions. By assessing mul- tiple sites of similar landscape composition, the dif- ference i in significant habitat factors dissipated. Simi- lar results were observed for Striped Skunks (Mephitis jmephitis — Baldwin et al. 2004) and suggest that land- ‘scape composition is an important factor when con- seetng models predictive of occurrence (Temple and ) Wilcox 1986; Dijak and Thompson 2000). Differential use of habitat by Raccoons across sea- sons has been documented (e.g., Lotze and Anderson 1979; Kaufmann 1982; Chamberlain et al. 2002), al- though quantitative assessments of these shifts among seasons have received little study. Nevertheless, con- Le nce of significant variables across seasons was generally high although a few exceptions were noted. For example, large hardwood snags and ground dens Were important factors during winter but were non- i i \\ Jackknife classification t ratio P No capture* Capture” “Combined -10.656 <0.001 70 We 72 -4.671 <0.001 2.732 0.006 2.583 0.010 -2.249 0.024 2.081 0.037 2.046 0.041 -7.034 <0.001 57 88 78 -4.177 <0.001 3.322 0.001 2.656 0.008 significant during summer. Both variables provide ther- mal shelter for Raccoons during winter (Stains 1961; Sanderson 1987) but may be less important during warmer seasons. Likewise, number of plant food species and distance to roads were significant factors during winter but not in summer. The non-signifi- cance of these variables may be related to the abun- dance of food and the lack of hunter use of roads dur- ing summer. This non-use of roads as travel corridors may have resulted in increased use of stream banks during winter (Hilty and Merenlender 2004), thus yielding different results between seasons for dis- tance to potential water. Ultimately, temporal vari- ability should be expected due to seasonality in vari- ables measured as well as in the biological needs of Raccoons and should be considered when modeling Raccoon occurrence. Most previous investigations have attempted to deter- mine those factors most important in influencing Rac- coon occurrence at the micro-scale but did not include modeling techniques in their analyses (Leberg and Kennedy 1988; Kennedy et al. 1991; Kissell and Ken- nedy 1992; but see Pedlar et al. 1997 for different mod- eling strategy). Such modeling strategies allow for the prediction of heavy-use areas by Raccoons and are particularly useful to managers of natural resources. Therefore, such procedures were incorporated into the current investigation and yielded models that gener- ally included the strongest variables from univariate analyses (i.e., forest and water characteristics). Result- ing models maintained high classification rates while allowing managers to focus only on those variables most important for predicting Raccoon occurrence. Such techniques have yielded slightly higher classifi- cation rates for habitat specialists [Fisher (Martes pen- nanti) = 79% — Carroll et al. 1999; Iberian Lynx (Lynx 230 pardinus) = 83% — Palma et al. 1999] likely due to their need for more specific habitat components. None- theless, classification rates observed in the current study are generally higher than those reported for other hab- itat generalists [Bobcat (Lynx rufus) = 59-70% — Kolowski and Woolf 2002; Striped Skunk = 56-75% — Baldwin et al. 2004]. Therefore, models that uti- lize replicated sites and account for differences in land- scape composition could serve as a blueprint for future investigations involving habitat generalists ultimately resulting in models more useful for wildlife managers. Unfortunately, construction of such models can be expensive. Care must be taken to develop an appro- priate sampling strategy to maximize results. Acknowledgments Partial funding for this project was provided by the Tennessee Wildlife Resources Agency, the Ames Plan- tation, and The University of Memphis. Thanks are extended to S. B. Franklin for critical review of the manuscript, and to M. Biernacki, S. B. Franklin, and T. A. Wasklevicz for assistance with statistical analy- sis and GIS application. Special thanks go to B. D. Carver, J. R. Hisey, J. B. Jennings, S. J. Mahady, and numerous students from The University of Memphis, as well as several individuals from the University of Tennessee and the Ames Plantation for their assis- tance in data collection. Literature Cited Agresti, A. 1996. An introduction to categorical data analy- sis. John Wiley & Sons, New York. Baldwin, R. A. 2003. An assessment of microhabitat vari- ables and capture success of selected mammalian meso- predators. M.S. thesis, The University of Memphis, Mem- phis, Tennessee. Baldwin, R. A., A. E. Houston, M. L. Kennedy, and P. S. Liu. 2004. 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Raccoons: a natural history. Smithsonian Institution Press, Washington D.C., USA. Received 3 August 2005 Accepted 3 October 2005 Notes Scoter, Melanitta spp., Migrations Interrupted by Confederation Bridge: An Update KATHERINE BUNKER-POPMA 40 Weldon Street, Sackville, New Brunswick E4L 4N4 Canada Bunker-Popma, Katherine. 2006. Scoter, Melanitta spp., migrations Interrupted by Confederation Bridge: An update. Canadian Field-Naturalist 120(2): 232-233. Continued monitoring over 10 years of scoter migrations through Northumberland Strait confirmed that these birds continue to perceive the Confederation Bridge (completed in 1997) as an obstacle. Such problems — and alternatives not causing them — need serious consideration when “strait crossings” are contemplated elsewhere. Key Words: Scoters, Melanitta spp., migrations, Northumberland Strait, bridge, New Brunswick. While the Confederation Bridge (46°15'N, 63°40'W) linking Prince Edward Island to New Brunswick was being completed in 1997, Hicklin and Bunker-Popma (2001) examined the possible influence of the bridge upon migration by marine birds. They concluded that migrating scoters (Melanitta spp., seaducks) were un- willing to fly under the (40 metre high) bridge and often reluctant to fly over it. Earlier information, recently supported by several studies (e.g. Wilson et al. 2003*), suggested that, in recent years at least, most migrating scoters passing through the Maritime Provinces in spring cross the Chignecto Isthmus 40-50 km. further west and do not approach the Confederation Bridge. Much less is known about their fall migrations. TABLE 1. Ten years of scoter movement at Confederation Bridge, New Brunswick/Prince Edward Island. Spring (S) and fall (F) totals shown separately for each year. Year Observation Scoters seen to Scoters seen to pass time (hr) approach bridge over bridge* (%) 1997 S 54 731 243 (33) F 102 876 338 (39) 1998 S 15 70 29 (41) F 49 923 388 (42) 1999 S 4 16 0 F 78 1495 881 (59) 2000 S 35 389 67 (17) F 74 1675 859 (51) 2001 S 16 212 115 (54) F 72 906 431 (48) 2002 S 18 461 150 (33) F 70 1368 725 (53) 2003 S 27 604 129 (21) F 50 1824 1274 (70) 2004 S 28 SS) 275 (53) F 47 1253 696 (56) 2005 S 11 46 6 (13) F 22 435 156 (36) 2006 S 14 124 68 (55) F 40 1038 547 (53) Totals S 222 3168 1082 (34) F 604 11793 6295 (53) Grand Totals 826 14961 7377 (49) * No scoters were seen to pass under the bridge May, 2006 I monitored seaduck migrations at Confederation Bridge each year since 1997, using procedures of our original study (Hicklin and Bunker-Popma 2001) to see if these birds habituated to the perceived obstacle. Results of these observations, summarized in Table 1, suggest that the majority of scoters still, after 10 years, have not learned a simple, general, way of passing the bridge. Nearly half of those seen (34% in spring, 53% in fall) circled upwards and passed extremely high over the bridge without landing. Others landed on the water, and remained there, often for hours. Still oth- ers followed the bridge towards the opposite shore, and sometimes were lost to sight while still flying. As my observations do not suggest that birds in the two latter categories stay near the bridge indefinitely, pre- sumably they continue their migrations later, but tim- ing of their departures and the routes followed remain unknown. Given the small fraction of regional scoter popula- tions that approach it, this bridge cannot be considered a major obstacle to their migrations, at a flyway level. Nevertheless, the continuing problem posed to the birds that encounter this bridge needs to be borne in mind whenever and wherever bridges are proposed in loca- NOTES tions traversed by major portions of migratory species populations. Acknowledgments A. J. Erskine edited my original draft and encour- aged publication of this update. Document Cited (marked * in text) Saint John Naturalists Club, Inc. 2003. Mystery migra- tion: Secret of the Black Scoter (bilingual). Video (23:45 English, 24:15 French); filmed and edited by Rowan Ridge Productions Ltd. (This video refers to several unpublished studies on scoter migrations in the Maritime Provinces). Literature Cited Hicklin, P. W., and K. Bunker-Popma. 2001. The spring and fall migrations of scoters, Melanitta spp., at Confederation Bridge in the Northumberland Strait between New Bruns- wick and Prince Edward Island. Canadian Field-Naturalist 115: 436-445. MacKinnon, C., and A. Kennedy. 2006. An observation of the spring 2006 migration of Black Scoter, Melanitta nigra, in Northumberland Strait interrupted by the Confederation Bridge. Canadian Field-Naturalist 120(2): 233-234. Received 14 December 2004 Accepted 26 February 2007 An Observation of the Spring 2006 Migration of Black Scoter Melanitta nigra, in Northumberland Strait, Interrupted by the Confederation Bridge, New Brunswick — Prince Edward Island COLIN MACKINNON and ANDREW KENNEDY Canadian Wildlife Service, Atlantic Region, P.O. Box 6227, Sackville, New Brunswick E4L 1G6 Canada MacKinnon, Colin, and Andrew Kennedy 2006. An observation of the spring 2006 migration of Black Scoter Melanitta nigra, in Northumberland Strait, interrupted by the Confederation Bridge, New Brunswick — Prince Edward Island. Canadian Field-Naturalist 120(2): 233-234. An observation from the bridge of a flock of Black Scoters suggests that, nine years after construction, the 12.9 km Confed- eration Bridge may still be a partial barrier to bird migration. Only 3 of 18 scoters (16.7%) crossed over the structure during the observation period. Key Words: Black Scoter, Melanitta nigra, seaducks, migration, Confederation Bridge, Northumberland Strait, New Brunswick, Prince Edward Island, Cape Jourimain National Wildlife Area. Construction of the Confederation Bridge began in 1995 and was completed in 1997. This 12.9 km con- crete structure spans the Northumberland Strait at its narrowest point between the Cape Jourimain Nation- al Wildlife Area, New Brunswick, and Borden, Prince Edward Island. As part of collecting pre-construction baseline data in 1990, MacKinnon et al. (1991) mon- itored seabird and seaduck movement through the Northumberland Strait. Twenty-two hours of surveys, during spring and fall, produced 5214 observations on 25 species. Of these, 2041 (39.1%) were seaducks, 1400 (26.9%) were three species of scoters and 641 (12.3%) were Common Eider, Somateria mollisima (MacKinnon et al. 1991). During the early 1990s, concerns were expressed over possible effects the Confederation Bridge may have on seabird migration. The closest comparable existing structure was the Canso Causeway, connect- ing Cape Breton Island with mainland Nova Scotia. Erskine and Smith (1986) reported that, although the causeway was a comparatively low and short structure, Common Eider were believed to have treated the struc- ture as a barrier to flight. A flock of eiders flew parallel to the causeway, gained altitude, and eventually crossed over. The question was, would seaducks do the same with the Confederation Bridge? In 1997, Hicklin and Bunker-Popma (2001) con- ducted surveys at Confederation Bridge during spring 234 migration, similar to those reported by MacKinnon et al. (1991). The Confederation Bridge was then opera- tional, and their observations suggested that it formed a partial barrier to scoter migration. Of 3986 and 1441 scoters observed by them during respective spring and fall migration, only 12.8% of scoters crossed over the bridge during the spring and slightly more, 21.9%, in the fall. Bunker-Popma (2006) subsequently reported additional years of observation. The Confederation Bridge utilizes a multi-span con- crete box girder structure and is composed of three main components: the approaches (east and west), a 40 m- high deck and 60 m-high ship-navigation span. The east approach is 0.6 km over 7 piers and the west approach is 1.3 km over 14 piers. The main bridge is 11.0 km (44 piers) in length. In the middle of the bridge is a 2.50 km navigation section. The typical span, between piers, is 250 m. The Confederation Bridge has been in operation for nine vears. During the morning of 13 April 2006, we observed the behaviour of a flock of Black Scoters, Melanitta nigra, during their northward migration (46°13'N, 63°44'W). This observation proved more interesting as the authors were travelling across the bridge, by vehicle, at the time and were able to track the flock’s activities over most of the crossing. At the time of observation, visibility was excellent, no clouds, +6°C and wind light SSW. Shortly after we entered the Confederation Bridge from Borden, Prince Edward Island, at 08:10, a flock of 18 Black Scoters was observed flying parallel with and approximately 100 m south of the Bridge at a height of approximately 40 m above the water. The flock was observed to rise and approach the bridge deck on three occasions. On two occasions, the flock approached the deck and then veered back to its origi- nal flight path, parallel with the Bridge. On the third occasion, the flock gained altitude and, similar to the previous occasions, aborted its attempted flight over the bridge after coming within approximately 50 m of the structure. However, this time the flock completed a clockwise circle, towards the bridge, in order to regain their original flight path. At the instant in which the flock began this circle, three birds broke from the main group and promptly crossed over the Bridge, behind our vehicle, at a height of approximately 10 m above the bridge deck. This crossing occurred approximate- ly 4.5 km from the Prince Edward Island side of the THE CANADIAN FIELD-NATURALIST Vol. 120 bridge. The velocity of the flock was constant with our driving speeds of about 80 km/h, until it could no longer be viewed in the vicinity of the navigation channel. While rising over the navigational channel, the scoters remained out of sight below us. We main- tained a constant speed of 80 km/hr and the flock was immediately re-sighted once we descended from the elevated navigation span. Upon re-locating the flock, we noted that the group had decreased in size, while out of sight, from 15 to 8 birds. We did not observe any scoters coming into view on the north side of the Bridge and assume the missing seven birds did not cross the structure. The remaining eight birds eventually landed in the water approximately 100 m south of the bridge and 500 m from the New Brunswick coastline (46°10'N, 63°49'W). Observations ceased at 08:25. It is notewor- thy that the Confederation Bridge is built in a general “S” curve, thus the flight of the scoter flock was fol- lowing the contours of the bridge and not a straight line. This observation demonstrates that nine years after the Confederation Bridge was built, some seaducks, such as Black Scoters, still may perceive the structure as a partial barrier to their northward migration through the Northumberland Strait. Acknowledgments We thank P. Hicklin, K. Davidson and A. J. Erskine for helpful reviews of the manuscript. Literature Cited Bunke-Pompa, K. 2006. Scoter, Melanitta spp., migrations interrupted by Confederation Bridge: an update. Canadi- an Field-Naturalist 120(2): 232-233. Erskine, A. J., and A. D. Smith. 1986. Status and movements of Common Eiders in the Maritime Provinces. Pages 20-29 in Eider Ducks in Canada. Edited by A. Reed. Canadian Wildlife Service, Report Series Number 47. 177 pages. Hicklin, P., and K. Bunker-Popma. 2001. The spring and fall migrations of scoters Melanitta spp., at Confederation Bridge in the Northumberland Strait between New Bruns- wick and Prince Edward Island. Canadian Field-Naturalist 115: 436-445. MacKinnon, C. M., R. W. Daury, and R. J. Hicks. 1991. Seabird and seaduck movement through the Northumber- land Strait, 1990. Technical Report Number 130. Canadian Wildlife Service — Atlantic Region. 86 pages. Received 25 April 2006 Accepted 6 February 2007 2006 NOTES First Record of a River Otter, Lontra canadensis, Captured on the Northeastern Coast of Alaska SHAWN P. HASKELL! 'LGL Alaska Research Associates, Inc., 1101 E 76th Avenue, Suite B, Anchorage, Alaska 99518 USA; e-mail: shawn haskell @ ttu.edu Haskell, Shawn P. 2006. First record of a River Otter, Lutra canadensis, captured on the northeastern coast of Alaska. Canadian Field-Naturalist 120(2): 235-236. A River Otter (Lontra canadensis) was captured on the Beaufort Sea coast of northeastern Alaska during a coastal fish study in late July 2001. The otter was dead upon retrieval and of adult size. River Otter distribution is not known to extend north of Alaska’s Brooks Range. This occurrence may have represented a one-time spring dispersal event. However, fluviomor- phological features of some river systems may provide suitable habitat for an overwintering otter in this region of Alaska’s North Slope. Key Words: River Otter, Beaufort Sea, Lutra canadensis, fyke net, distribution, Alaska. In late July 2001, a River Otter (Lontra canaden- sis) of adult size was retrieved from a shoreline fyke net in Lions Bay, a barrier island lagoon system on the Beaufort Sea coast of northeastern Alaska. The live-capture fishing net was located at 70°10.024'N, 146°12.557'W, approximately 82 km east of Prudhoe Bay and 11 km west of the Canning River; the nearest non-industrial areas of human habitation are Barter Island, 100 km to the east, and the Colville River delta, 170 km to the west. A westerly wind event had caused coastal water levels to rise, completely submerging the trap-end of the net in about 1.5 m of water, drowning the otter. Physical samples or measures were not taken of the specimen. The animal was discarded to the water at the capture site. About two weeks after the capture, the author retrieved the carcass for brief examination. Marine arthropods of the family Mysidae had entered all openings in the otter’s body, expediting decompo- sition. River Otter distribution is commonly noted as ex- tending far north up the Mackenzie River drainage of northwestern Canada but then falling south of the Alaskan Brooks Range to the west (Rue 1981; Whitak- er 1996; Forsyth 1999; Wilson and Ruff 1999). Bee and Hall (1956) reported a River Otter sighting in the cen- tral Brooks Range at Kanayut Lake and tracks observed near the Beaufort Sea coast at the Kuparuk River. Manville and Young (1965) also displayed a point loca- tion near the mouth of the Kuparuk River, but given Survey methods, that point may stem from the same observation of tracks documented by Bee and Hall (1956). A commercial winter fishery operated in the Colville River Delta since the early 1950s has never captured a River Otter (James Helmericks, homestead- er and naturalist, personal communication). There are only two state sealing records for River Otters in Alas- ka’s North Slope region; one was caught near Wain- wright in northwestern Alaska in 1990 and the other near Anaktuvuk Pass in the central Brooks Range in 1985 (Geoff Carroll, Alaska Department of Fish and Game Area Biologist, personal communication). Biol- ogists have reported seeing small numbers (i.e., one or two) and tracks of otters north of Anaktuvuk Pass but south of the Arctic Coastal Plain on the Anaktuvuk and Chandler rivers, both tributaries to the north-flowing Colville River, where freshwater upwellings provide open water throughout winter and habitat for fishes such as Arctic char (Salvelinus alpinus; Geoff Carroll, Alaska Department of Fish and Game Area Biologist, personal communication). It is unknown whether the 2001 otter capture in Lions Bay signified an anomalous extralimital occur- rence or a case of regular periodic use by otters of this area seldom occupied by humans. Leffingwell (1919) noted that larger rivers in the region supported popu- lations of fish year-round. Relatively small catches of juvenile amphidromous fishes during the summers of 1999 and 2001 in Lions Bay indicated that these fish, and likely some adults of the same species, did over- winter in the area (Griffiths et al. 2002"). Craig and McCart (1974°) identified groundwater springs in the nearby Kavik River and Canning River drainage, as well as a field of aufeis in the Canning River Delta, that supported overwintering fish. Some of these areas could potentially provide suitable habitat for an over- wintering otter on Alaska’s North Slope. Acknowledgments This capture occurred during the daily operation of a remote coastal fish study operated by LGL Alaska Research Associates, Inc. The study was funded by the Exxon Owners Group of Point Thomson, admin- istered and reviewed by the Environmental Studies Group of BP Exploration (Alaska) Inc. Documents Cited (marked ° in text) Craig, P. C., and P. McCart. 1974. Fall spawning and over- wintering areas of fish populations along routes of the pro- posed pipeline between Prudhoe Bay and the Mackenzie Delta. Chapter 3 in Fisheries research associated with pro- posed gas pipeline routes in Alaska, Yukon, and Northwest 236 Territories. Edited by P. J. McCart. Arctic Gas Biological Report Series 15(3). 36 pages. Griffiths, W. B., L. R. Martin, S. P. Haskell, W. J. Wilson, and R. G. Fechhelm. 2002. Nearshore Beaufort Sea fish studies in the Point Thomson area, 2001. Report for BP Exploration (Alaska) Inc. by LGL Alaska Research Asso- ciates, Inc., Anchorage, Alaska. 55 pages + appendices. Literature Cited Bee, J. W., and E. R. Hall. 1956. Mammals of northern Alas- ka on the Arctic Slope. University of Kansas, Lawrence, Kansas. Forsyth, A. 1999. Mammals of North America: temperate and arctic regions. Firefly Books Ltd., Willowdale, Ontario. Leffingwell, E. K. 1919. The Canning River region, northern Alaska. Government Printing Office, Washington D.C. THE CANADIAN FIELD-NATURALIST Vol. 120 Manville, R. H., and S. P. Young. 1965. Distribution of Alaskan mammals. Bureau of Sport Fisheries and Wildlife. Circular 211. U.S. Government Printing Office, Washing- ton, D.C. Rue, L. L., IM. 1981. Furbearing animals of North America. Crown Publishers, Inc., New York. Whitaker, J. O., Jr. 1996. National Audobon Society field guide to North American mammals. Alfred A. Knopf, Inc., New York. Wilson, D. E., and S. Ruff. 1999. The Smithsonian book of North American mammals. UBC Press, Vancouver/Toronto. Received 4 August 2002 Accepted 24 December 2004 Osprey, Pandion haliaetus, Depredates Common Eider, Somateria mollissima, Duckling BRENDA M. BLINN!, VICKY VIOLETTE!~, and ANTONY W. DIAMOND! ' Atlantic Cooperative Wildlife Ecology Research Network, University of New Brunswick, P.O. Box 45111, Fredericton, New Brunswick E3B 6E1 Canada > Present address: Department of Fish and Game, Bay-Delta Region, P.O. Box 47, Yountville, California 94599 USA; e- mail: bblinn@dfg.ca.gov 3 Mailing address: 80, Notre-Dame Nord, Louiseville, Québec JSV 1X5 Canada Blinn, Brenda M., Vicky Violette, and Antony W. Diamond. 2006. Osprey, Pandion haliaetus, depredates Common Eider, Somateria mollissima, duckling. Canadian Field-Naturalist 120(2): 236-237. To our knowledge, we present the first published account of a successful attack on a Common Eider (Somateria mollissima) duckling by an Osprey. Ospreys were sighted periodically during 940 hours of Common Eider observations in June-August, 2001 on Grand Manan Island, New Brunswick. One duckling (<1 week old) was observed depredated by an Osprey. Vulner- ability of ducklings to predation may promote such opportunistic behaviour. The rarity of observations of such incidents suggests depredation on ducklings by Osprey is uncommon. Key Words: Common Eider, Somateria mollissima, Osprey, Pandion haliaetus, predation, Grand Manan Island, New Brunswick. The diet of Ospreys (Pandion haliaetus) consists al- most exclusively of live fish (Poole et al. 2002). Coastal populations may switch prey species depending on the pattern and timing of fish migration (Greene et al. 1983). Ospreys are known to feed occasionally on rodents, birds, reptiles, amphibians, and invertebrates (see Wiley and Lohrer 1973 for literature review). Birds reported include grebes, gulls, ducks, including a 2-week old Wood Duck (Aix sponsa; Wiley and Lohrer 1973), and passerines (Chubbs and Trimper 1998). From 7 June to 3 August 2001, a study of Common Eiders (Somateria mollissima) was conducted on Grand Manan Island located 35 km from the New Brunswick south shore. It is the largest (25 km long, 11 km wide) of 17 other islands in the archipelago. A total of 940 hours of observations were made at eight 1 km‘? sites, typically between 06:00 and 21:00 h. Duckling num- bers and ages were recorded at 20-minute intervals using scan sampling (Altman 1974). Predatory attacks were recorded as they occurred. Eider broods occupied the intertidal zone which was covered with brown fucoid algae (Ascophyllum nodosum and Fucus vesi- culosus) with some rock, sand or mud. A mixed forest of Balsam Fir (Abies balsama), spruce (Picea sp.) and alder (Alnus sp.), as well as shrubs (Rubus sp.) and var- ious grasses (Graminae) were usually present within 10-50 m of the high tideline. A vertical tidal range of up to 8.4 m can be expected in this area. Approxi- mately 3370 Common Eider pairs (Ronconi and Wong 2002) and 15 other Anatidae species breed in the archi- pelago (Dalzell 1991). On 4 July 2001, an Osprey of unknown breeding sta- tus captured and carried off a 0-7 day-old eider duck- ling approximately 100 m offshore. The researcher (V. V.) was situated near the waterfowl sanctuary of Cas- talia Marsh (44°42'N, 66°48'W). Just prior to the attack, an eider créche of 25 adult females and 62 ducklings was observed in the area dabbling for invertebrates. Duckling ages and counts (number) in this group were 0-7 days old (18), 8-13 days old (38), and 14-21 days old (6). At 20:00, the créche stopped foraging and became alert when an Osprey flew nearby before leav- 2006 ing the site. It returned at 20:03, swooped down, took an eider duckling without any defensive behaviour by the hens, and flew out of sight. The presence of the duckling in the raptor’s talons was verified using a Bushnell Spacemaster 45-60 x 200 spotting scope. Ospreys were not observed at Castalia for the remainder of the evening. Ospreys were recorded at Castalia and at five other sites during visits both before and after the attack. Only lone Ospreys were seen at any given time, and no nests were present within any of the study sites. Bald Eagles (Haliaeetus leucocephalus) also took three ducklings at Castalia and two ducklings at another site approximately 10 km away. One of the reasons suggested to explain Ospreys tak- ing non-fish prey is reduced visibility caused by turbid water (Chubbs and Trimper 1998). Machmer and Yden- berg (1990) found that capture rate by Ospreys de- creased greatly when winds were above 25.2 km/h. On the evening of the attack, moderate winds (~13 km/h), medium-sized waves and clear skies probably did not contribute to poor fishing conditions, although incom- ing tide and floating algae could have made fish less detectable. Ospreys may take easily-captured (usually weak or captive) birds when available (Dement’ev and Glad- kov 1951). Eider ducklings <3 weeks old have poorly developed diving skills (Hamilton 2001). This makes them vulnerable to predation, especially in the first 10 days of life, with reported mortalities exceeding 90% from gull attacks (Mawhinney and Diamond 1999). Because Ospreys cannot dive deeper than one meter into water (Prevost 1982), they forage in shallow- water environments (Poole et al. 2002). Shallow-water areas are often near brood-rearing habitat of eiders (McAloney 1973). In spite of many hours of observation of eider créch- es in waters surrounding Grand Manan Island, we ob- served only one instance of duckling depredation by an Osprey. Ducklings less than one week old, which are vulnerable to predation, were observed during five out of eight observation periods (63%) and during 32 out of 309 scans (10%) at Castalia. Therefore, it seems un- likely that this Osprey was a “specialist”; i.e., fed more frequently on ducklings than the average rate for the species. Predation on eider ducklings by Ospreys is evidently a rare event everywhere. Acknowledgments Financial, logistical and collaborative support that made this observation possible was provided by the Atlantic Cooperative Wildlife Ecology Research Net- NOTES 237 work (ACWERN) at the University of New Brunswick, the New Brunswick Wildlife Trust Fund, Acadian Sea- plants Ltd. and the New Brunswick Department of Natural Resources and Energy. Our sincere gratitude goes to Greg McGuire for his assistance in the field. André Breton provided valuable comments on earlier versions of the manuscript. This is ACWERN Publi- cation Number UNB-59. Literature Cited Altmann, J. 1974. Observational study of behavior: sampling methods. Behaviour 49: 227-267. Chubbs, T. E., and P. G. Trimper. 1998. The diet of nesting Ospreys, Pandion haliaetus, in Labrador. Canadian Field- Naturalist 112: 502-505. Dalzell, B. 1991. Grand Manan Birds..A checklist with occurrence graphs and a site guide. Third Edition. Brian Dalzell, Grand Manon, New Brunswick. Dement’evy, G. P., and N. A. Gladkovy. Editors. 1951. Birds of the Soviet Union. Volume |. Sovietskaya Nauka, Moscow. Greene, E. P., A. E. Greene, and B. Freedman. 1983. For- aging behaviour and prey selection by Ospreys in coastal habitats in Nova Scotia, Canada. Pages 257-267 in Biology and management of Bald Eagles and Ospreys. Edited by D.M. Bird. Harpell Press, Ste. Anne de Bellevue, Quebec. Hamilton, D. J. 2001. Feeding behaviour of Common Eider ducklings in relation to availability of rockweed habitat and ducklings age. Waterbirds 24(2): 233-241. Machmer, M. M., and R. C. Ydenberg. 1990. Weather and Osprey foraging energetics. Canadian Journal of Zoology 68: 40-43. Mawhinney, K., and A. W. Diamond. 1999. Using radio- transmitters to improve estimates of gull predation on Com- mon Eider ducklings. Condor 101: 824-831. McAloney, R. K. 1973. Brood ecology of the Common Eider (Somateria mollissima dresseri) in the Liscombe area of Nova Scotia. M.Sc. thesis, Acadia University, Wolfville, Nova Scotia. Poole, A. F., R. O. Bierregaard, and M.S. Martell. 2002. Osprey (Pandion haliaetus). In The Birds of North Amer- ica, Number 683. Edited by A. Poole and F. Gill. The Birds of North America, Inc., Philadelphia, Pennsylvania. Prevost, Y. A. 1982. The wintering ecology of Ospreys in Senegambia. Unpublished Ph.D. dissertation, University of Edinburgh, U.K. Ronconi, R. A., and S. N. P. Wong. 2002. Seabird colonies of the Grand Manan Archipelago: 2001 census results and guidelines for surveys and future monitoring. Grand Manan Whale and Seabird Research Station. Bulletin Number 4. Wiley, J. W., and F. E. Lohrer. 1973. Additional records of non-fish prey taken by Ospreys. Wilson Bulletin 85: 468- 470. Received 14 October 2005 Accepted 23 April 2007 238 THE CANADIAN FIELD-NATURALIST Vol. 120 Extension de |’aire de distribution de 1 Umbre de vase, Umbra limi, dans le nord-est du Québec JEAN-FRANCOIS DESROCHES! 'Cégep de Sherbrooke, Département des Techniques de bioécologie, 475 rue du Cégep, Sherbrooke, Québec JIE 4K1 Canada; e-mail: Jean-Francois.Desroches @cegepsherbrooke.qc.ca Desroches, Jean-Frangois. 2006. Extension de |’aire de distribution de ’Umbre de vase, Umbra limi, dans le nord-est du Québec. Canadian Field-Naturalist 120(2): 238-239. Le 26 juillet 2005, plusieurs Umbres de vase (Umbra limi) ont été capturés au secteur Les Buissons de Pointe-aux-Outardes, sur la Céte-Nord au Québec. Ces mentions élargissent l’aire de distribution de ce poisson de 300 km vers le nord-est. On 26 July 2005, we caught many Central Mudminnows (Umbra limi) at the Les Buissons sector of Pointe-aux-Outardes, on the Céte-Nord region of Québec. These records extend the known distribution of this fish 300 km to the northeast. Mots-Clés: Umbre de vase, Mudminnow, Umbra limi, distribution, aire de distribution, Québec L’Umbre de vase (Umbra limi) se retrouve dans le centre de I’ Amérique du Nord, du centre-sud des Etats- Unis jusqu’au sud du Canada, vers le nord (Page et Burr 1991). Au Canada, il est réparti d’ouest en est depuis |’ Alberta jusqu’au Québec (Scott et Crossman 1974). Au Québec, l’espéce est confinée au sud-ouest de la province, se rendant a l’est jusque dans la région de la ville de Québec selon les données actuellement disponibles (Scott et Crossman 1974; Bergeron et Brousseau 1981; Bernatchez et Giroux 2000). En juillet 2005, dans le but de capturer des amphi- biens aquatiques, deux nasses a ménés ont été placées dans un étang a castors situé dans le secteur Les Buis- sons de Pointe-aux-Outardes (49°7'47"N; 68°21'36"0). Les nasses ont été relevées le 26 juillet 2005. Dans la premiere nasse, il n’y avait aucune capture, mais quel- ques Umbres de vase ont été capturés en fouillant dans l'eau avec une épuisette. Neuf spécimens, soit quatre jeunes (longueur totale 27,0-30,7 mm) et cing adultes (longueur totale 54,5-62,7 mm) ont été gardés et sont conservés dans la collection de poissons du Musée canadien de la nature (CMNFI 2005-0032). Dans la seconde nasse se trouvaient une jeune Anguille d’ Amé- rique (Anguilla rostrata) vivante et les restes de deux Umbres de vase, probablement dévorés partiellement ou régurgités par l’anguille. Des fouilles effectuées avec une épuisette ailleurs dans |’étang, la méme journée, ont permis la capture de quelques autres Umbres de vase et de tétards de grenouilles. Cet étang a castors est relié a un lac plus gros, le lac Lagacé, qui a une superficie d’environ 6 ha (850 m x 85 m). Le contour de l’étang est irrégulier, l’eau est teintée de brun, et on y retrouve quelques arbres morts. Il présente certains méandres vaseux ot |’on retrouve un peu de végétation aquatique submergée. La forét bordant l’étang est mixte et composée notamment d’épinettes (Picea sp.), de Sapin baumier (Abies bal- samea) et de Bouleau blanc (Betula papyrifera). Des aulnes (Alnus sp.) bordent I’étang. L’Umbre de vase est une espéce tolérante reconnue pour vivre dans des eaux chaudes et riches en végéta- tion, généralement peu oxygénées (Scott et Crossman 1974). Dans le sud-ouest du Québec, on le trouve dans les baies riches en végétation des lacs et riviéres lentes, dans les étangs 4 castors et les étangs de tour- biéres, et dans les fossés (observation personnelle). Les plans d’eau retrouvés sur la Cote-Nord sont en général des lacs clairs typiques a l1’Omble de fontaine (Salve- linus fontinalis), habitat peu propice a l’?Umbre de vase. On y retrouve également des étangs a castors et des tourbiéres, lesquels sont plus favorables a l’espéce. L’Umbre de vase pourrait donc se retrouver ailleurs sur la Céte-Nord. La découverte de |’;Umbre de vase sur la C6te- Nord au Québec, a 300 km au nord-est de sa distribu- tion actuellement connue, revét un intérét particulier. Il apparait probable que ce poisson soit plus répandu qu’on le croyait au Québec et qu’il ait été sous-inven- torié. Il s’agit d’une petite espéce ne présentant aucun intérét pour la péche sportive ou commerciale et vivant le plus souvent dans des milieux peu propices aux poissons convoités pour la péche. Ces facteurs con- tribuent sans doute a la méconnaissance de sa distribu- tion dans le nord du Québec, et possiblement sur la rive sud du Saint-Laurent vers l’est. Il est également pos- sible que cette population d’Umbres de vase soit le fruit d’une introduction volontaire ou accidentelle par des pécheurs les ayant prélevés plus au sud, dans le but de s’en servir comme appats de péche. Des inventaires mieux ciblés permettraient de préciser la distribution de cette espéce au Québec. Remerciements L’auteur remercie Julie et Félix Soucy pour leur aide sur le terrain. Littérature citée Bergeron, J. F., et J. Brousseau. 1981. Guide des poissons d’eau douce du Québec. Ministére du Loisir, de la Chas- se et de la Péche, Québec. 217 pages. Bernatchez, L., et M. Giroux. 2000. Les poissons d’eau douce du Québec et leur répartition dans |’est du Canada. Broquet inc., Boucherville, Québec. 350 pages. 2006 NOTES 239 Page, L. M., et B. M. Burr. 1991. A field guide to freshwa- — Scott, W. B., et E. J. Crossman. 1974. Poissons d'eau douce ter fishes of North America, North of Mexico. Houghton du Canada. Ministére de l'Environnement, Service des Mifflin Company, Boston et New York. 432 pages. péches et des sciences de la mer, Ottawa. 1026 pages Regu 24 aoit 2005 Acceptée 19 février 2007 The Mink Frog, Rana septentrionalis, in Southeastern Labrador JEAN-FRANCOIS DESROCHES!, ISABELLE PICARD, and JOHN E. MAUNDER? ' Cégep de Sherbrooke, Département des Techniques de bioécologie, 475 du Cégep, Sherbrooke, Québec JIE 4K 1 Canada; e-mail: jean-francois.desroches@ cegepsherbrooke.qc.ca * P.O. Box 250, Pouch Cove, Newfoundland and Labrador AOA 3L0 Canada Desroches, Jean-Francois, Isabelle Picard, and John E. Maunder. 2006. The Mink Frog, Rana septentrionalis, in Southeastern Labrador. Canadian Field-Naturalist 120(2): 239-240. Mink Frogs (Rana septentrionalis) were collected at three new localities in southeastern Labrador in 2003 and 2005, extend- ing the known range of the species about 200 km to the east, to the limits of the coast. La Grenouille du Nord (Rana septentrionalis) a été trouvée dans trois nouvelles localités dans le sud-est du Labrador en 2003 et 2005. Ces mentions élargissent |’aire de répartition connue de |’espéce d’environ 200 km vers lest, aux limites de la c6te de l’Océan Atlantique. Key Words: Mink Frog, Rana septentrionalis, range, Labrador. The Mink Frog (Rana septentrionalis) has the most northerly southern limit of any North American anuran (Hedeen 1986). This southern limit ranges from south- ern Manitoba and northern Minnesota, south and east through Wisconsin, northern Michigan and southern Ontario, then northeast through northern New York, northern New England, New Brunswick, and Nova Scotia (Conant and Collins 1998; Cook 1984; Hedeen 1986). The species has recently been introduced on the island of Newfoundland (Warkentin et al. 2003). Its northern limit remains poorly known, largely because the species is particularly aquatic (Hedeen 1986) and its call and appearance are unfamiliar to many observers. In Labrador, the Mink Frog has been documented from several scattered localities, though not from the extreme north or southeast (Maunder 1983, 1997; Cook 1984; Desroches and Rodrigue 2004). In 2003, during a primarily botanical and malaco- logical field trip for the Provincial Museum of New- foundland and Labrador (JEM), and in 2005, during a vacation trip (J-FD and IP), we surveyed ponds and brooks along all of the main roads and smaller town access roads of southeastern Labrador, as far to the northeast as the town of Cartwright. FicurE |. The Mink Frog in Labrador. Previous records: dots On 3 August 2003 a single adult interface round (Maunder 1983) and squares (Maunder 1997). Question =) o° mark: Questionable record (see Maunder 1983: 42, 44). Gray zone: proposed distribution based on previous records (Bleakney 1958; Cook 1984; Desroches and freshly dead from unknown causes, was collected (NFM HE-250) from the shoreline of a medium-sized pond, located beside the access highway running be- Rodrigue 2004). Stars: New records in southeastern tween the Trans-Labrador Highway and the town of Labrador (1: southwest of Paradise River, 2: Charlot- Cartwright, about 39.5 km southwest of the road tetown, 3: St. Lewis, 4: east of Churchill Falls). Bro- junction to the town of Paradise River (53°09'52"N; ken line: Labrador/Québec border. 57°31'S6"W). This locality was, at the time, about 100 km east of the known range of the species (num- ber | in Figure 1). The water of the pond was clear. 240, The dominant emergent plants were Yellow Pondlily (Nuphar variegata), Narrowleaf Bur-reed (Spargani- um angustifolium), and Water Lobelia (Lobelia dort- manna). The shoreline of the pond was mainly gravel- ly, over a boulder substrate, and was well-vegetated. The surrounding habitat was Kalmia scrub heath, with patches and strips of low Black Spruce (Picea mari- ana) woods in more sheltered areas. In 2005, Mink Frogs were observed at two addition- al localities in southeastern Labrador, about 200 km east of the former known range of the species (num- bers 2 and 3 in Figure 1). On 11 July, a subadult Mink Frog was collected (CMN-35805) from a small pond located beside a gravel road in the town of Charlotte- town (52°46'09"N; 56°06'59"W). The pond was about 60 x 15 m in size, and had clear water, a muddy bot- tom, and grassy and rocky shores. Trees at the site were mostly Black Spruce. On 18 July, two Mink Frog tad- poles were collected (CMN-35810) by dipnetting at Blue Hill Pond, St. Lewis (52°24'55"N; 55°55'45"W). The pond had clear water, a sandy and rocky bottom, and some emergent plants near the shore. It was bor- dered by Sweet Gale (Myrica gale) and alder (Alnus sp.), and the adjacent forest was dominated by Black Spruce. Two Threespine Sticklebacks (Gasterosteus aculeatus) were caught in the same habitat, but re- leased. In 2004, an additional observation of the Mink Frog was recorded at St. Lewis by Christian Fortin (Fortin 2005) but no voucher specimen was collected (C. Fortin, personal communication). In 2005, Mink Frogs were also observed in west- ern Labrador, near the site of an old record, along the Trans-Labrador Highway, east of the town of Churchill Falls (53°29'29"N; 63°40'06"W) (number 4 in Figure 1). Two adults and one subadult were collected (CMN- 35798). Although this observation is not a range exten- sion it is included here because it is a new collection record. During our searches for amphibians in southeastern Labrador, the only other species observed was Amer- ican Toad (Bufo americanus). Tadpoles of this species were seen and/or collected at five locations along the roads. THE CANADIAN FIELD-NATURALIST Vol. 120 The new Mink Frog collections recorded in this paper show that the species is more widespread in southeastern Labrador than has been previously doc- umented. Further Labrador surveys should emphasize areas north of the presently known distribution of the species. Acknowledgments The authors thank Frederick W. Schueler for com- ments on the manuscript before submission, and Francis R. Cook for useful comments on the submit- ted manuscript. Literature Cited Bleakney, J.S. 1958. A zoogeographical study of the amphib- ians and reptiles of eastern Canada. National Museum of Canada Bulletin 155. 119 pages. Conant, R., and J. T. Collins. 1998. A field guide to the rep- tiles and amphibians, Eastern and Central North America. Revised third edition. The Peterson Field Guide Series, Houghton Mifflin Company. Boston and New York. 616 pages. Cook, F. R. 1984. Introduction to amphibians and reptiles of Canada. National Museum of Natural Sciences, and Nation- al Museums of Canada, Ottawa. 211 pages. Desroches, J.-F., and D. Rodrigue. 2004. Amphibiens et reptiles du Québec et des Maritimes. Editions Michel Quintin, Waterloo. Québec. 288 pages. Fortin, C. 2005. Sortie herpétologique sur la Cote-Nord et au Labrador: de Natashquan a St. Lewis. Le Naturaliste can- adien 129: 50-56. Hedeen, S. E. 1986. The southern geographic limit of the Mink Frog, Rana septentrionalis. Copeia 1986: 239-244. Maunder, J. E. 1983. Amphibians of the province of New- foundland. Canadian Field-Naturalist 97: 33-46. Maunder, J. E. 1997. Amphibians of Newfoundland and Labrador: status changes since 1983. Herpetological Con- servation 1: 93-99. Warkentin, I. G., C. E. Campbell, K. G. Powell, and T. D. Leonard. 2003. First record of Mink Frog, Rana septen- trionalis, from insular Newfoundland. Canadian Field- Naturalist 117: 477-478. Received 15 August 2005 Accepted 15 March 2007 2006 NOTES 24 First Record of the Great Barracuda, Sphyraena barracuda from Canada DARLENE BALKWILL!, BRIAN W. Coap!, ISMAEL GALVEZ2, and JOHN GILHEN ‘Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada 21219 Cromwell Road, Halifax, Nova Scotia B3H 4L1 Canada 4Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada Balkwill, Darlene, Brian W. Coad, Ismael Galvez, and John Gilhen. 2006. First record of the Great Barracuda Spn Vraena i barracuda, from Canada. Canadian Field-Naturalist 120(2); 241-242. We report the collection, and provide a description, of the first Great Barracuda, Sphyraena barracuda, in Canadian water based on a partial skeleton from Country Island, Nova Scotia. Key Words: Great Barracuda, Sphyraena barracuda, new record, fish fauna, Nova Scotia, Canada. The cranium, lower jaw and attached trunk verte- brae of a Great Barracuda, Sphyraeana barracuda (Family Sphyraenidae), were found dried on a beach on Country Island, Guysborough County, Nova Scotia at 45°06'00"N, 61°32'37"W in May 1999 by Ismael Galvez. This is the first record for this species from Canada (Scott and Scott 1988; Coad et al. 1995). The specimen is catalogued in the Nova Scotia Museum of Natural History, Halifax, under NSM 85385. Description Three species of barracuda are reported from the western Atlantic Ocean north of Florida, Sphyraena barracuda (Walbaum, 1792), S. borealis DeKay, 1842, and §. guachancho Cuvier in Cuvier and Valenciennes, 1829. Only S. borealis has been recorded from Cana- da, a single specimen from Halifax Harbour, Nova Scotia, caught on 5 October 1928 (Scott and Scott 1988). Carpenter (2002) records S. barracuda and § guachancho from Massachusetts (rare) southwards. Our specimen is neither S. borealis nor S. guachan- cho on the basis of length, these two species general- ly reach maximums of 46-50 cm, (commonly) 35 cm and 50-60 cm, respectively (De Sylva 1984; Robins and Ray 1986; Carpenter 2002). Murdy et al. (1997) give | m for the latter species. Our specimen is close in size to a disarticulated specimen of S. barracuda in the Royal Ontario Museum (ROM R6281; pur- FicureE |. Photograph of skeletal remains of a Great Barracuda from Country Island, Guysborough County, Nova Scotia (NSM 85385)(bottom) with skull of ROM R6281 above. 242 THE CANADIAN FIELD-NATURALIST Vol. 120 1 2 3 CENTIMETRES FIGURE 2. Photograph of skeletal remains of a Great Barracuda from Country Island, Guysborough County, Nova Scotia (NSM 85385) flanked by the upper and lower jaws of ROM R6281. chased from Kensington Fish Market, Toronto, and thought to be from the Caribbean area) that has a re- corded total length of 86.0 cm. The length of the dor- sal skull surface (from the anterior tip excluding the upper jaw to the origin of the supraoccipital crest) is 135.1 mm in NSM 85385 and 116.8 mm in ROM R6281. These measurements make the Nova Scotian specimen ca. 99.5 cm total length. NSM 85385 has a head length of ca. 181.7 mm (the opercular bone is somewhat curved) and FishBase (www.fishbase.org) states that head length is 17.7% total length, giving a total length for the Nova Scotian specimen of 1.03 m. These two forms of measurement agree, making the Nova Scotian specimen about 1 m long in life. This unequivocally eliminates S. borealis and probably S. guachancho. Other, osteological, characters also strongly sug- gest the Nova Scotian specimen is a Great Barracuda. The specimen has the area between eyes flat (convex in S. guachancho), teeth are contiguous, vertically flat- tened and erect (conical and widely spaced in S. bore- alis, angled backwards in S. guachancho), and the tip of the maxilla reaches past the anterior orbit margin (not reaching the orbit in S. borealis) (De Sylva 1984; Murdy et al. 1997; Carpenter 2002; comparative mate- rial of S. guachancho ROM R1837 and R2706). We consider it unlikely that this specimen came from a local fish market and was imported. One of us (JG) has not seen this species on sale locally. The shore of Country Island is remote from commercial sites. It probably represents a stray that died in Canadian waters and was washed ashore. Acknowledgments We are indebted to Kevin Seymour, Department of Palaeobiology, Royal Ontario Museum, Toronto, for the loan of comparative material. Photographs were taken by Alison Murray, Canadian Museum of Nature, Ottawa, who also provided osteological advice. Literature Cited Carpenter, K. E. Editor. 2002. The Living Marine Resources of the Western Central Atlantic. Volume 3. Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. Food and Agriculture Organization, Rome. vi + 1375-2127. Coad, B. W., with H. Waszczuk, and I. Labignan. 1995. Encyclopedia of Canadian Fishes. Canadian Museum of Nature, Ottawa, and Canadian Sportfishing Productions, Waterdown, Ontario. viii + 928 pages, 128 colour plates. De Sylva, D. P. 1984. Sphyraenoidei: development and rela- tionships. Pages 534-540 in Ontogeny and Systematics of Fishes. Special Publication Number 1, American Society of Ichthyologists and Herpetologists, Lawrence, Kansas. ix + 760 pages. Murdy, E. O., R. S. Birdsong, and J. A. Musick. 1997. Fishes of Chesapeake Bay. Smithsonian Institution Press, Washington. xi + 324 pages. Robins, C. R., and G. C. Ray. 1986. A Field Guide to the Atlantic Coast Fishes of North America. Houghton Mif- flin Co., Boston. xi + 354 pages. Scott, W. B., and M. G. Scott. 1988. Atlantic Fishes of Canada. Canadian Bulletin of Fisheries and Aquatic Sci- ences 219: xxx + 731 pages. Received 18 July 2005 Accepted 16 January 2007 A oncmoms See I Book Reviews ZOOLOGY Animal Skulls — A Guide to North American Species By Mark Elbroch. 2006. Stackpole Books, 5067 Ritter Rd., Mechanicsburg, PA 17055. 727 pages, $34.65 Paper. As an instructor of comparative vertebrate anatomy, and a self-proclaimed enthusiast of skulls, | very much looked forward to reading and reviewing this book. I learned a fair amount from this book, mostly regard- ing animals that don’t live in my area, so overall, this book was for me, a valuable read. There is an enormous amount of material synthe- sized in this book. There are the expected figures illustrating the skull bones from a number of species; neighbouring figures illustrate the measurements most commonly used. However, the figure for reptiles has several errors, and those for the birds and mammals also have a few...not a good start for fundamental material near the beginning of the book. Skulls of each species are illustrated in three views within the species accounts. Elsewhere, montages of dorsal views (mammalian and herp crania) and lateral views (mammalian mandibles, birds) are quite useful, and where practical, are illustrated at life-size. Even though the skulls in these montages show less detail than in the species accounts, I’ve always found this set-up useful (as in the Golden Field Guide to Birds, where warblers and sparrows are illustrated in a simi- lar manner). There are a number of issues that I found annoying. _ Why, for instance, is the Mantled Howler Monkey des- cribed (and illustrated along with two other species of non-human primate)? They are native to Central and South America; Are they feral in North America? We are never told. We are never told of the distributions of any of the animals, a shortcoming if someone is try- ing to identify a skull they’ve found based on this book. Secondly, the author is somewhat free with his use of the word bone. “Occipital condyles”, for instance, are processes of bones, not bones unto themselves, though that is how they’re treated. In a book that thankfully retains appropriate terminology, this oversight will be undetected by the novice, but irritating to the experi- enced. I now need to shout...““Molars are not flat!” I have to remind my students of this constantly. In Eng- lish, we have the two words, “flat” and “broad”, and they are not interchangeable. Molars adapted for grind- ing vegetation are broad...they are anything but flat as erroneously described numerous times in this book. There are a few other annoying errors in this vein. Given the number of prominent, pre-publication re- viewers mentioned in the acknowledgements, one must wonder about their diligence to this project. The title is awkward, Animal Skulls (as opposed to part two in the series, Plant Skulls?)...and mislead- ing...a guide to North American Species: fish are not covered at all, amphibians and reptiles are given a token amount of space (12 species, fewer than 3% of those in North America), birds (71 species, 10% of breeders) are allotted only a fraction of the coverage of mammals (143 species, one-third of known species), this despite the fact that the number of species of birds alone outnumbers those of mammals in North Ameri- ca. I do understand that there is very little variation in skulls amongst warblers or sparrows, for example, and illustrating each species could quickly become re- dundant. However, the mammalian bias is still over- whelming. The mammalian dominance in this book is also reflected in the amount of text devoted to each species account — one line for each herp, one paragraph for each bird, four or more paragraphs for mammals. About 100 pages of measurement data are provided for mammals (which is wonderful), though none is pro- vided for the others. One is forced to ask, “Why bother with the birds and herps at all?” The same amount of space could have been used to make a more complete book focussing on mammal skulls alone. Then, subse- quent volumes addressing other taxa could complete the series. I particularly liked the early chapters on interpret- ing form, marks on skulls (from gnaw marks to builet holes) and preparing skulls. These chapters were in- formative and well-written. For me, a large part of the worthiness of this book was in these early chapters. That is not to diminish the worthiness of the mam- malian species accounts, they too are thorough and instructive. RANDY LAUFF Department of Biology, St. Francis Xavier University, Antigo- nish, Nova Scotia Canada B2G 2W5 244 A Field Guide to North Atlantic Wildlife By N. Proctor and P. Lynch. 2006. Yale University Press P.O. Box 209040, New Haven, Connecticut 06520-9040 USA 221 pages. U.S. $19.95 Paper. This book arrived on the morning I left for Svalbard in the Norwegian arctic. While the area covered by this book, offshore from North Carolina to Newfoundland, is some distance from Norway, the two areas do share a lot of wildlife. So I took the book along to make direct field comparisons. I like the author’s concept for this guide. He has selected the most likely species you have some possi- bility of seeing on a pelagic trip or ferry crossing in the northwestern Atlantic. This includes the large “plank- ton’ and algae, fish, turtles, whales, seals and seabirds. Only the more common species are included, enabling the authors to keep the book small [18 x 11 x 1.5 cm] and portable. This meant I was able to identify Beroe’s Comb Jelly, Fin Whale and Ivory Gull carrying only one guide. The guide does not include coastal species such as crabs, molluscs and inshore fish. The descriptions are well written. For example, the text for Fin Whale is a version of the classic descrip- tions found in many guides. I compared this to a whale that circled the ship and the information given was accurate down to the details of the back chevrons. I had far more problems with the illustrations. First there is no attempt to show scale. A 10-inch flying fish is shown as the same size as an 18-inch species. This is a critical characteristic for look-alike species such as Glaucus and Iceland gulls. You need to check the text to see that there is a 5-inch difference in size. Similarly the Northern Seal plate shows the tiny 4 foot Ringed Seal as the same size as the 9 foot Hooded Seal. I found this disconcerting. Some of the species shown are really cold water arctic animals. The North Atlantic is the southern fringe of their range. This is not mentioned for animals like Beluga, Dovekie or Harp Seal. I think giving their typ- A Complete Guide to Arctic Wildlife By Richard Sale. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 400 pages. $49.95 Paper. This is a comprehensive field guide to the birds and mammals of the Arctic. Author Sale describes the ecology and people of the region along with some Arctic history. He cannot avoid discussing both cli- mate change and the threat of pollution. Short chap- ters on geology, geography and biogeography set the background. These are followed by the main text: a field-guide style section on the Arctic’s mammals and birds. The information provided covers identification, size, voice, distribution, diet, breeding and taxonomy. The guide is illustrated with photographs supported THE CANADIAN FIELD-NATURALIST Vol. 120 ical range would add perspective and could be said in few words. The non-bird illustrations were very good. The artist has captured the short face of the Ringed Seal and the longer-nosed head of the Harp Seal very well. The fish portraits from the warm-water Dolphin [the fish] to the cold-water Cod are accurate. The bird illustrations are another story. For example, the illustration of the Arctic Tern looks almost identi- cal to that of the Common Tern. The Arctic has a short- er head and a longer tail giving it a different fore-and- aft look from the Common. The wing pattern is also quite different, having less black and giving the Arctic a much paler appearance. I had similar issues with other confusing pairs [Leach’s and Wilson’s storm-petrels, Glaucous and Iceland gulls, Audubon’s and Manx shearwaters, etc.] Many other species lacked the sub- tle plumage differences that help confirm identification. For example, the two-toned grey on the wing of a Kit- tiwake and the armpit marks on the murres. Many birds are shown in winter plumage only. This is a reason- able approach as many birds disappear inland or north to breed. However, the winter Atlantic Puffin is very odd and the loon illustrations do not capture these birds at all well. It appeared that the artist did not have field experience with these species. This is, however, a useful little book. If you are a novice I would recommend you take this book plus a modern bird guide on your pelagic trips. For more ex- perienced bird watchers this book will suffice to remind you of the key points for what you may see and will be useful for all the non-bird species. I thank Dr. Tom Smith, a fellow traveller, for sharing his immense knowledge of arctic wildlife, and thereby contributing to my review. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada by some field-guide style plates. Each species has a range map showing their circumpolar ranges. I am always delighted to find a book that shows birds and mammals from a bio-regional perspective. Last summer I saw eight Snow Geese on the west coast of Svalbard — an unusual sighting. When you look at the distribution of the birds it is not hard to see how their expanding population could spill over the now unfrozen ocean a short distance eastward. This kind of perspec- tive is difficult to reach when you use guides based on a political boundary (birds of Europe, Russia etc.) The author’s first problem was the difficultly of de- fining the boundary of the Arctic. He uses, in my opin- ion, the most sane choice. He has taken the 10°C July 2006 isotherm, modified by extreme winter temperatures. This still leaves out some areas that I think of as “Arc- tic’ such as James Bay and its Polar Bears and the northwest coast of Alaska with Belugas and Yellow- billed Loons. This, perhaps, proves that nature will never entirely fit into our human need for definitions. Following his definition, the author then chose the species to be included as arctic birds and mammals. Here | am a little perplexed. He has, for example in- cluded Carrion Crow but not Black-billed Magpie (which I have seen at 63°N). He has only an introduc- tory mention of the Gray Jay, which ranges to the shores of the Arctic ocean. Tree Swallow is given an individual account while the more northerly Cliff Swallow (I have seen it at 73°N) is merely mentioned as a rarity. | am also confused by the species he has decided to split and those he leaves as sub-species. For, example he has Iceland and Thayer’s gulls as separate species but not Mew and Common gulls. There seems to be better evi- dence to split the latter. Similarly you can question hav- ing separate entries for Black and Common scoters. These are minor points and not as significant to my two major concerns. The species descriptions are de- tailed and clear and are some of the best I have read. While they are first rate, the author gives exceedingly spartan information on confusing species. This is in contrast to the descriptions, which often allude to an important field mark. For example, he states the three central tail feathers of the jaegers are “distinctive.” In the field separating a Parasitic Jaeger with a 9 cm tail extension from a Long-tailed with a 12 cm extension is not that easy. I well remember a controversial bird The Black Flies (Simuliidae) of North America By P. Adler, D. Currie, and D. Monty Wood. 2004. Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario MSS 2C6 Canada, and Cornell University Press. 941 pages. $99.95 US. This is a huge book. It is huge in size, filling 941 pages. It is huge in scope, treating the biology, man- agement and identification of the entire black fly fauna of Canada, the United States, and Greenland. And it is huge intellectually, the culmination of many decades of work by an enthusiastic international community of researchers, among whom the authors of this book are prominent members. In the preface, Peter Adler states that the book is “geared to a general readership.’ I would not have guessed that this was true, at first glance, but after some perusal it seemed to me that the book would be useful from about the undergraduate level on up. The writing is clear and precise, but the vocabulary will be daunting for most people without biological training. I suspect that the book will find a place in almost all scholarly libraries, and that the specialists who will use it most probably ordered a copy before it came into print. But there is a wider audience for the book as well, and in BooK REVIEWS 245 that needed the late great Earl Godfrey's talents to re- solve as a Long-tailed Jaeger. Similarly the informa- tion on the Common and Hoary redpolls is not enough for field separation. There is no mention, for example, of the stubby bill of the Hoary — very evident in the nice accompanying photo. My second point is that this is not a complete guide to wildlife. Unlike such books as Les Beletsky and Dennis Paulson’s Ecotraveller's Wildlife Guide to Alas- ka it does not contain any information on plants, insects, marine invertebrates and fish or amphibians. Admit- tedly Beletsky only includes the commoner, more vis- ible species although his title does say it is a complete guide, but he does include a broad spectrum of the wild- life, other than birds and mammals, visible to the naked eye. While this in not meant as a coffee table book the photographs are really eye-catching. The bird plates are quite good, but I was less taken with the mammal plates. My chief reservation about all these illustrations is they frequently show only adult males. I will take this book with me on my upcoming trip to the Pribiloffs. For the most part I will be seeing wildlife that I am familiar with and I will not carry a conventional field guide to the birds or mammals. This book will provide me with more encyclopaedia-style information about the creatures I am seeing. | will also take Beletsky and Paulsen’s book and a flower guide. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada this regard I predict that it will be a required reference for many decades to come. The authors mention its rel- evance to medical and veterinary entomologists, aquat- ic biologists, environmental consultants, systematists, naturalists, pest management specialists and students, but I suspect the real list will be longer than that. The text is arranged in four parts, covering back- ground information, biology of black flies, economic concerns, and systematics and taxonomy. The first part provides an overview of the subject, a very readable history of the study of black flies (with lovely historical photographs) and a chapter on techniques for collecting and curating black fly specimens. As an entomologist (with no particular interest in black flies, 1] admit) I nonetheless found this section very interesting. The second part of the book deals with the biology of black flies, and consists of a fine treatment of the structure of these insects (and their immature stages), as well as a well-written overview of their cytology. Black fly cytology was pioneered by the late Klaus Rothfels, a man deeply missed by the authors, and who they refer to as “friend, mentor, and phenomenon.” A chapter summarizing the life history and behaviour 246 of black flies is also offered, along with an exhaustive summary of blood feeding host records, and records of black fly diseases and parasites. For most readers, it is likely that the third part of the book, which treats the management of black fly pests, will be the most interesting. I can see this part of the book providing fodder for any number of term papers, research paper introductions, and consulting and government reports. I was surprised, however, to find that the chapter on management (Chapter 8) was largely historical in nature, without a summary of con- trol recommendations. On careful reading, however, I could see the reason for this approach. Oil-based pes- ticides, DDT, and other chemical controls for black flies have not stood the test of time. The authors sum- marize the use of the bacterial toxin Bi, but give it only a qualified nod, with the caution that it too might lose its effectiveness over time. The chapter ends in an inter- esting discussion of repellants, and repellant clothing. The fourth and final part of the book is clearly the section closest to the authors’ own interests. It treats the 254 species of “North American” black flies, thor- oughly. The focus is not simply on identification, it is also deeply phylogenetic, and includes a very careful evolutionary justification for each and every level of the classification. This is followed by a superb species by species treatment of the entire fauna. Significantly, there is additional information on the economic sig- nificance (if any) of each and every species, adding greatly to the summary in part three of the book. As a non-specialist, I have to admit that the most impressive aspects of the book to me were the illus- THE CANADIAN FIELD-NATURALIST Vol. 120 trations. They begin on page 436 (well before the halfway mark) and they are so masterfully executed that I couldn’t help but come away thinking that there was something deeply beautiful and elegant about black flies, their larvae and pupae, and the fine details of their anatomy. More full-body illustrations of the adults would have been nice, but the overall sameness of the other life stages illustrated convinced me that side-by-side comparisons of the adults might not be all that useful. Maps follow the illustrations, and treat the United States and Canada on a county level, but they do not show Greenland. And, as one might expect in a book of this nature, the reference section and the indices are exhaustive. This is a magnum opus, and a tremendous labour of love. It ranks, in my opinion, right up there with Holldobler and Wilson’s The Ants. Having said that, it seems to me a shame that the book appears to have been presented as a somewhat impenetrable scholar- ly tome, without a showy slip cover, and with little or no fanfare for the “general readership” to which it was addressed. Don’t be scared off by the size, or the complexity of this book. It is a masterwork, and if any aspect of your interests overlaps with the subject of black fly biology, do consider adding this fine vol- ume to your library. JOHN ACORN University of Alberta, Edmonton, Alberta TST 5L7 Canada Literature Cited: Hollidobler, Bert, and E. O. Wilson. 1990. The ants. Belknap Press (Harvard University Press), Cambridge, Massachusetts. 732 pages. Insects: Their Natural History and Diversity: With a Photographic Guide to Insects of Eastern North America By Stephen A. Marshall. 2006. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 720 pages. CDN $95. Hardcover It has always been said that you can’t judge a book by its cover, but after reading Insects: Their Natural History and Diversity I’ve learned that this well-worn idiom isn’t always true. When this book first crossed my desk, to say that I was instantly enamoured would be an under-statement. It was so beautiful, the cover adorned with a stunning jewel-toned dogbane beetle (Chrysochus auratus). 1 almost didn’t want to crack the binding. However, my curiosity finally got the better of me and I’m glad it did because once I started read- ing I couldn’t put the book down. Visually stunning, with over 4000 colour photo- graphs of insects in their natural habitats, Insects: Their Natural History and Diversity has the look and feel of a glossy coffee table book while still being full of accurate, well researched information. As its title implies, Insects: Their Natural History and Diversity focuses on the diversity and natural his- tory of common families of northeastern North Ameri- can insects. The book opens with a brief synopsis of basic insect anatomy and morphology. This is followed by chapters covering the diversity of all insect orders, including all the major families, along with two chap- ters on non-insect arthropods and methods for observ- ing, collecting and photographing insects. The book’s last 50 pages are dedicated to illustrated keys to order and family as well as a key to the most commonly en- countered insect larvae. These keys are designed to facilitate ease of use and therefore emphasize morpho- logical characters visible to the naked eye or easily seen with a hand lens. Also peppered throughout the book are helpful suggestions on where to look for and. find various insect orders/families. For example, “Depending on your inclination and the weather, a good place to start looking for assassin bugs would be in your kitchen light fixture. Unless you are much more 2006 fastidious than most, the odds are that among the crispy critters accumulated there you will find a large black assassin bug called the Masked Bed Bug Hunter (Redu- vius personatus).” Considering that Insects: Their Natural History and Diversity is priced so as not to be cost prohibitive and is clearly written in plain language, it will be highly accessible to a broad audience, including naturalists, amateur entomologists as well as seasoned profession- als. The author has also included a dollop of humour and wit throughout the text. For example, this passage describing the appearance of springtails: “Some are covered with scales, like those of a butterfly, many are brilliantly colored and all are morphologically bizarre, starting from the long, forked tail used to make Her- culean leaps, and ending with the deeply pocketed mouth that makes springtails look like they have lost their dentures and then sucked on a bunch of lemons.” Book REVIEWS 247 This book would make a great textbook for a natu- ral history or general entomology course. Especially when you consider that the impetus for this book cen- tres on materials originally gathered in support of the author’s third-year course “The Natural History of Insects” at the University of Guelph. With its depth of scope and true to life colour photographs /nsects: Their Natural History and Diversity would be indispensable in the field; however due to its size, it would be a bit unruly to have to lug around. I thoroughly enjoyed this book and would recom- mend it whole-heartedly to anyone who has an interest in entomology, natural history or a simple curiosity about the six-legged world that surrounds us. GINA PENNY Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada The Northern Goshawk: A Technical Assessment of its Status, Ecology and Management By Micheal L. Morrison. 2006. Cooper Ornithological Soci- ety, Pennsylvannia. The Northern Goshawk: A Technical Assessment of Its Status, Ecology and Management, is just as the title suggests, a highly technical assessment of the goshawk. I do not recommend it to those looking for a light read. Micheal L. Morris, editor, has compiled 22 recent journal articles that study the Northern Gos- hawk, focussing on ecology and population. Morris outlines two objectives for the assessment; the first is to summarize existing knowledge of the goshawk into one book, and the other is try to provide a framework for the development of future regional conservation and management strategies for the species. Due to the legal issues surrounding the Northern Goshawk and its proposed listing as a Species at Risk, particularly in the United States, this book is a useful step for establishing what we do and what we do not know about this Accipiter. Unfortunately, it appears that the latter predominates. Morris structures this assessment in an appropriate way, Starting with an article that extensively covers the ecology of the goshawk, which helps provide the read- er with a general knowledge of the bird. After this first article, the volume is divided into three distinct sec- tions: Regional, Ecology, and Management, each of which contains a number of independent studies that Morris has gathered from several sources. Each of these studies take a highly regionally specific look at certain aspects of the Northern Goshawk. The goshawk has an extensive range over North America, Europe and Asia. As a result of this huge regional coverage there is extensive continental/regional variation amongst populations of these birds. This makes it difficult as a reader to distil any major trends between the articles. I found it hard to link many of the articles together because of the different styles by which the studies were carried out, as well as some differences of opin- ion. For example, Squires and Kennedy suggest that “(impacts from forest management] can either enhance or degrade goshawk habitat depending on type and extent of habitat alterations” (page 61). This is not so much a weakness of the assessment but rather the real- ity of goshawk management, and emphasizes the need for regional management strategies instead of an all encompassing global strategy, which is one of Morris’s initial goals for the assessment. Overall, this publication is a building block. It estab- lishes where we have been in terms of goshawk man- agement and conservation and makes clear where we need to head in the future. Morris has done a good job of selecting appropriate articles that clearly depict the challenges faced by researchers and the Northern Goshawk. I would recommend this book only to those who want to critically study goshawk ecology. JARED B. CLARKE 323 Habkirk Drive, Regina, Saskatchewan S4S 6A9 Canada 248 Turtles of the World By Franck Bonin, Bernard Devaux, and Alain Dupré (Trans- lated by Peter C.H. Pritchard). 2006. Johns Hopkins Uni- versity Press. 416 pages. Illus., $50 (US). First published in 1996 in French, this guide to the turtles of the world has been translated into English by eminent turtle biologist Peter Pritchard. In the trans- lator’s note, Pritchard acknowledges that he has also made some corrections and updates to the text. As such, the book includes recently discovered species such as Pelusios cupulatta, only described in 2003. The book is divided into two sections: General Biology and Identification. General Biology provides a brief (nine pages) overview of the evolutionary his- tory of turtles, their biology and conservation issues. Identification provides information on the approxi- mately 330 recognized species of turtles. The species are grouped by families within the two suborders Pleu- rodira and Cryptodira. In general, each species account follows the same standardized format: distribution, description, natural history, protection. The species accounts vary from about half a page to three pages in length. Each species also has a small regional map showing which part of the world the species occurs in and a larger, close-up map showing the distribution as shad- ed areas. These distribution maps also show major river systems, as well as some political boundaries. In addi- tion, almost every species has one or more colour pho- tographs. Any attempt to list the names of all the species in the world of a certain group is a daunting task of sort- ing out sometimes conflicting and constantly chang- ing taxonomy. The authors have accepted the recent splitting of the genus Clemmys and duly note this change in the appropriate species accounts. In con- trast, the entire genus Sternotherus has been subsumed within Kinosternon without any explanation. While there has been debate over the validity of Sternotherus as a distinct genus, lumping the two genera goes against the current standard North American taxonomy. The authors are also somewhat inconsistent in assigning species status. They do not provide species accounts for Chelodina mccordi or C. pritchardi (although they are mentioned under the species account for C. no- vaeguineae), yet both these species are widely accept- ed. On the other hand, they elevate Hardella indi, a subspecies of H. thurjii, to full species status without any justification. This may explain why Hardella indi has the shortest species account in the book — three sentences. THE CANADIAN FIELD-NATURALIST Vol. 120 There are some strange oversights in this book. The IUCN Red List of Threatened Species is the standard international ranking system for species at risk of be- coming extinct. It designates status for these species from Lower Risk up to Critically Endangered. Turtles of the World occasionally mentions if a species has been “red listed” but not consistently, even within the same genus. For example, all four species within the genus Acanthochelys are on the Red List, yet the spe- cies accounts mention this for only one of the species. The book says one of the other species is on the “IUCN priority list” and the “IUCN is worried” about another. None of the species accounts provides the IUCN status, so the reader will not learn that A. pal- lidipectoris is considered the most endangered of the four species. A thorough reference section is also lacking. The references for this book do not even fill two pages and almost all of the references are to other books on tur- tles. Accordingly, most of the information in the book is presented without any reference to a specific primary source. However, occasionally a statement is referenced and in almost every case that reference is absent from the reference section. Of the Canadian species, the map for Graptemys geographica (Common Map Turtle) excludes the Ot- tawa area and all of the Quebec part of the range, and maps for both Glyptemys insculpta (Wood Turtle) and Emydoidea blandingii (Blanding’s Turtle) omit Nova Scotia. Terrapene carolina (Eastern Box Turtle), a species never confirmed to have been historically pres- ent in Canada at all, is shown as ranging across all of southern Ontario as far north as Toronto. The range is also over-extended in Michigan, Ohio, Pennsylvania and New York. Despite these criticisms this is a significant book in turtle biology. The photographs are often excellent. The natural history section, while sometimes too short, pro- vides insight into an incredible variety of life history strategies. For many species, the contents of the protec- tion section provide important information on threats, as well as a wide diversity of conservation measures. This volume is not the definitive statement on any of these 300+ species but it is a good introduction to the diversity of turtles. DAVID SEBURN Seburn Ecological Services, 2710 Clarenda Street, Ottawa, Ontario K2B 7S5 Canada 2006 Book REVIEWS 249 The Wolves of Algonquin Park: A 12-year Ecological Study By John B. Theberge and Mary T. Theberge. Department of Geography, University of Waterloo, Waterloo, Ontario N2L 3G1 Canada. 163 pages. Available from bkevans@ fes.uwaterloo.ca. $23.50 Canadian, $20 U.S. John and Mary Theberge, a husband and wife team, have put together information available on wolves and their prey in one of North America’s great ecosystems. Research results from the twelve year period (1987- 1999) was used as a core for the publication. Student projects were as follows: | Ph.D. thesis, 7 MSc. theses, 11 BSc. theses). Overall, scientific publication of 55 papers is impressive. In the end, it was the Theberge’s that pulled the material together into this single mono- graph. For that they need to be congratulated. The results of the study are also based on previous work begun in the 1960s. The Theberges reached back in time and reanalyzed and re-evaluated some of the data collected earlier. A major portion of the review centred on the taxonomic study of Wolves in the park. They concluded that these canids were larger than the Coyote-Wolf hybrids to the southwest, smaller than wolves to the northwest. It reinforced earlier conclu- sions of the existence of “Algonquin type”, “tweed type” and “boreal type” Wolves. New is the interpre- tation that these Wolves are taxonomically connected to the Red Wolf taxon, once more common on the con- tinent to the southeast. In the “canid soup of genet- ics” these are many theories and assertions. So far there have been no clear indications of the “ultimate” posi- tion. Maybe none will ever emerge that will satisfy everyone who is interested in the subject. The The- berges have presented a convincing case. Other portions of the monograph are equally well presented. There is much in the way of natural history — basic insight into the ecology of the predator within the prey dynamics dictated by geographical and botan- ical (habitat) setting. In an age of computers, abstract BOTANY Flowers — How They Changed the World By William C. Burger. 2006. Prometheus Books, 59 John Glen Drive, Amherst, New York. 14228-2197 USA. 337 pages. US $23.00. Cloth. Why are flowers important? They are not just for the viewing pleasure of humans, although they are spec- tacular — but why? This book will answer these ques- tions. “Without the gift of flowers ... man might still be a nocturnal insectivore gnawing a roach in the dark.” It is often true that the more we know about some- thing the more we appreciate it and the more we enjoy it. A book that leads to enjoyment is worthwhile, but this book also provides a much needed perspective on the flowering plants at a time when the world faces a biodiversity crisis. It contributes to an improved analysis and spurious mathematical modelling, this work stands out as a major contribution to science. It combines solid field work with a realistic quantitative assessment of the results, and | would recommend this as a case study for first-year graduate students who are contemplating a career in wildlife research. Beyond that, the study has had a real impact on conservation. In 2001, a total year-round Wolf killing ban in areas bordering Algonquin Park was announced by provin- cial authorities. Political responses were based on re- search results. It is a rare occurrence that this happens and it is a response to the well executed research. By any standard, that is a remarkable outcome to many years of dedicated work by scientists whose meticulous research and commitment to science were also cou- pled to a dedication to conservation and to bioethics. This is a scholarly monograph that likely will not be surpassed for many years to come. At the outset the Theberges posed a series of ques- tions that they wanted to find answers to. Likely some were listed after the fact, when intensive research results came in from the field work and data analysis. That too has strength. I cannot see much that has been left out. Appropriate photos and illustrations are found throughout the text. Some questions arose in my mind as to alternate explanations for some of the conclusions reached. Such results are inevitable and provide a use- ful basis for future scholars. This monograph is an exceptional document and has set a very high standard. I highly recommend it to anyone just starting field research on mammals, or for those veterans, who have spent much of their time in the pursuit of knowledge in the field of wildlife science. Lu CARBYN 137 Wolf Willow Crescent, Edmonton, Alberta TST 1T1 Canada understanding of the natural world and a concern for the future. It is a very valuable contribution as well as a very enjoyable book that will appeal to anyone as a result of the friendly writing style. William Burger, curator emeritus in the Department of Botany at Chicago’s Field Museum, takes us on a journey to the discovery of how flowering plants have transformed the world. He begins with “What exactly is a flower?” which is a delightful and comprehensive introduction to flower morphology or what makes a flower a flower. Floral parts and their role are described in different kinds of flowers. It comes as a surprise to many people that the four bright white petals of a dog- wood flower are not petals and the flower is actually an 250 aggregation of many flowers, that a dandelion flower is made up of dozens of flowers, that grasses do have very special flowers with floral parts analogous to those of tulips. The book is full of fascinating anecdotes. Why are Wolves friends of plants? By 1926 Wolves had been extirpated in Yellowstone and Yogi, Booboo, and the other bears were of course more interested in picnic baskets than controlling the Elk so there were soon too many Elk and the vegetation began to change. Seven- ty-five thousand Elk were removed between 1926 and 1968 but it was not enough to get things back to the way they were — although it did maintain some plant diversity and kept the park green! Finally, in 1995 a human brain triumphed and thirty-one Wolves were reintroduced to Yellowstone and that quickly got things back to the way they were. There was less browsing pressure as “fear factor” came into play with Elk avoid- ing places where they were more likely to be ambushed by Wolves. Browsing was not only reduced, it was spa- tially variable, creating more niches for plants. The entire Chapter 2 is a very entertaining voyage though the annals of ecology indicating how plants are assist- ed and influenced by other organisms. Flowering plants have a vast array of enemies, the herbivores, which are everything that eats plants, but they have developed chemical and mechanical de- fences. This is the subject of Chapter 3 “flowers and their enemies.” It is full of the interesting stories that are known well to biologists but not to others — such as the central American Acacias that have a police force of stinging ants that emerge from the hollow thorns to attack intruders but also destroy the surrounding veg- etation that competes with the Acacia. This is a mutu- ally beneficial relationship. As well as a home in hollow thorns the ants are supplied protein by specialized leaf glands and sugar from other glands on the Acacia stems. Chapter 5 “how are flowering plants distinguished” outlines the evolution of the distinctive features includ- ing tissues adapted to prevent water loss and to allow transport of water and nutrients and particularly the seed; which allowed land plants to achieve reproduc- tion without external water, unlike ferns and frogs. Did you know that the simple, 120 million-year-old flower of Archaefructus has cast some suspicion on the evolution of flowers by reduction from Magnolia- like ancestors (contrary to what we told and regurgi- tated for the final exam a few years ago). The classifica- tion system, double fertilization and the evolution of fruit and many other complex phenomena are intro- duced in a clear and very interesting way. There is a wealth of information in this chapter and although it may be the most tedious chapter in the book, if you want to know what a plant is all about, you will not find a better general presentation of this information. Chapter 6, “what makes flowering plants so special?” points out that flowering plants were not present in THE CANADIAN FIELD-NATURALIST Vol. 120 the forests of 300 million years ago (that made coal) but appeared 120 million years ago, were apparently little affected by the great extinction that wiped out the dinosaurs and other lineages, and became 87% of all plants today. The estimated 100-fold increase in the number of plant species since the Carboniferous period is mostly a result of the proliferation of flow- ering plants. The chapter explains why they are so suc- cessful and special in sustaining 99% of life on the planet. It may seem strange to find an explanation of photosynthesis in the same chapter as an explanation for the similarity of the floras of eastern North Amer- ica and Japan, but the reader is definitely left with an idea of how special flowering plants are. With the excellent introduction in these chapters the reader has developed a comprehensive knowledge of botany and ecology which provides a basis for the last two chapters, which indicate how flowering plants changed the world. Chapter 7, “primates, people and the flowering plants” explains how flowering plants created a world in which humans could live. The nec- tar, pollen, foliage and fruits together set the stage for a proliferation of insects and this led to the evolution of many new groups of insect eaters including the earliest primates, — but how did flatter faces, dextrous digits, upright stance and larger brains develop? It is all here. The chapter concludes with an explanation of how flowering plants and people came together to develop agriculture and how major civilizations rely upon flow- ering plants. Of course flowering plants did not just create a world for humans, they also changed the world in many other ways and this is dealt with in Chapter 8, “how flow- ers changed the world.” As they vastly increased in numbers they increased overall biodiversity. Biochem- ical specialization is explained with fascinating exam- ples. Effect on the weather is considered including gradual cooling through carbon sequestration and the effect of grassland expansion. The chapter concludes with an outline of the effect of human actions on flow- ering plants and their impact on the world. An epilogue outlines the responsibility of humans with regard to protecting and managing plant diversity and preserv- ing the world. Following the epilogue is a series of notes 17 pages in length providing additional information sources for each chapter. This is an extremely useful section, vast- ly improving the educational value of the work, and it serves as an indication of the very extensive knowl- edge of the author. The book also has a-colour insert of 12 attractive plates. It concludes with a very useful glossary that defines terms such as carpel, caryopsis and cleistogamous. The index is very helpful. Burger’s book is definitely a journey. It pulls togeth- er a broad array of concepts into a developing theme of how flowering plants got us to where we are today. It offers a biological perspective including the basics 2006 of botany, ecology and evolution combined into a picture of plants and people and ends with an evalua- tion of where we are and where we are going. If every- one read and understood this book — if everyone had the knowledge and the respect for life and the inquir- ing and sensitive mind of its author — we would be in a better position to save the world. This book is an example of outstanding communi- cation. It provides biologists with what they need to know to teach, but it is not just for the biologist, nor just for Americans who spend nineteen billion dollars ENVIRONMENT Guide to Deserts By Andrew Warren and Tony Allan (General Editors). 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 240 pages. $19.95 Paper. After reading this book I realised I have never been to “see” a desert. I searched for Scimitar-horned Oryx (northern Sahara). I marvelled at the marine life off the Peruvian-Chilean coast (Sechura-Atacama desert). I have visited the impressive Abu Simbel temples (Libyan desert). But I have not really looked at the desert for its own sake. This is a mistake I will not repeat. While I have not been completely unobservant there are many subtleties I have missed. In future I will look for zengens, desert pavement, inselbergs and yar- dangs. Deserts is a neat, small book in which the author defines deserts and describes the climatic conditions that shape their landscapes. He gives examples of the different types of desert and illustrates the impact wind and — yes — water have had on the land and its inhabi- tants. There are six chapters on aspects of deserts and an atlas of the major desert areas. The chapters cover wildlife such as plants, invertebrates and mammals and how they are adapted to thrive in an arid ecosystem. The people of the desert, their food, clothes, homes, customs and religions are described. This is an impor- tant chapter with the current focus of news from the deserts of Badiyat Ash Sham (Iraq) and Dasht-E-Mar- gow (Afghanistan). The authors explain the riches of the desert. Not just oil, but farming and mining. They also look at conservation issues and the damage done by human activity. After reading this material I am more convinced we need to re-assess the use of water for irrigation. Book REVIEWS 251 annually on leisure-time activities related to flowers and plants — it is for anyone. Behind the provocative title is a beautifully written book describing what flow- ering plants are, how they have influenced the evolu- tion of life, contributed to the origin of humans and enabled humans to become the masters of our planet with the responsibility to manage its resources prop- erly. It is so far from being just another book! PAUL CATLING 170 Sanford Avenue, Ottawa, Ontario K2C OE9 Canada The atlas is really a field guide of the world’s 14 major arid zones [Canada’s minuscule desert in British Columbia is not mentioned]. In addition to a map there are descriptions of each area’s key characteristics. This includes some history, the people and key issues as well as a physical description. The authors have writ- ten a special chapter on visiting deserts to encourage people to see these special places. There are plenty of photos and illustrations. These have been chosen not so much as pretty pictures, but to illustrate the points in the text. They range from the stark beauty of Monument Valley, through the rock- cut ruins at Petra to simple depictions of dunes. They show the faces and costumes of Tuareg, Berber and Bedouin (but not Navajo or Apache). The author has not missed mine sites, luxury hotels or aerial views in making his selection. One photo supposedly contain- ing Giraffe, Ostriches and Eland shows six Oryx, some distant gazelles (presumably Springbock) but no Eland. This book is very interesting to read just because it puts deserts in perspective. For someone like me who goes to places looking for something special, this book gives added value to a trip. For all of us it gives a won- derful background on where desert’s ecosystem fits in the world’s biosphere. The authors have convinced me that deserts are important and I will pay more atten- tion in the future. Buy this book for yourself. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Day THE CANADIAN FIELD-NATURALIST Vol. 120 Endangered: Wildlife on the Brink of Extinction By George C. McGavin. 2006. Firefly Books Ltd. 192 pages. U.S. $35.00 Cloth An ecologist could not help but be drawn to a book with the title Endangered. Wildlife on the brink of Extinction. seeking an update on the current state of the world’s species at risk. Unfortunately McGavin’s book does not provide a succinct analysis of current status, the impacts we should be most concerned about, geographic localities of concern and the likely future trends. Rather this coffee table book provides a piece- meal of “information bites” for fifty animal groups ranging from dogs and cats to salmon and songbirds. The lengthy (50 of 192 pages) two chapter intro- duction on the “Nature of Extinction” and “The Sixth Extinction” could be a grade nine biology course cov- ering the basics of evolution and human development from our hunter-gather roots, through our agrarian and industrial revolutions to an assessment of supply and demand harvesting of nature. Instead of a discussion of the ecological implications of the loss of species to ecosystem integrity, extinction is considered more in terms of the extent of human mourning and the likely curiosity of future generations about species that once existed. The “Lost and the Last” chapter constitutes the cen- tral text starting with Monotremes and Marsupials. Typ- ical treatment of species groups include general state- ments such as: “Platypus eggs are eaten by introduced red foxes and rats as well as by native predators such as lizards. Many echidna eggs suffer the same fate.” This text provides no meaningful information on the status of even these most unique of creatures for which clear trends must surely be known. No consistent for- mat is used to cover each animal group, for example within the sub-section Insectivores Insectivora the sub- titles used could have been chosen at random: Sense of Smell; Desmans in Danger; Desert Moles; Rare and Restricted; and Solendon Capture. Often the informa- tion presented begs the question; What has this to do with wildlife on the brink of extinction? If general eco- logical information is to be presented it would have been better in a text box or sidebar as a special case study. MISCELLANEOUS Much more information could have been conveyed by following a consistent format for each animal group with sub-headings that allowed the reader to move from one group to another building an understanding of the most threatened species, the number of species threatened within each group, the ecological implica- tions of species loss, the key causes of species decline, likely future trends and actions required to reverse neg- ative trends. Summary graphs, tables or pictorial rep- resentations are virtually absent, yet they too could have been used to convey large amounts of information in a simple repeatable format for each species group. The most impressive aspect of Endangered is sure- ly the figures but, while there are captions, the text makes no direct reference to the figures and text and figures are often unrelated. In the Prosimians group the text states: “The strangest prosimian is surely the aye-aye. Its wide eyes, pointed ears and thin bony fin- gers lend it a goblin-like appearance”. No picture of the aye-aye is included, but pictures of the sifaka and loris are included with no information on their status. There is considerable variability in the treatment of each species group with some written in a more author- itative style, Bats Chiroptera is one example where there is a logical organization of sub-sections and more information on extinction. Variability is also seen in how comprehensively each species group is covered and this appears unrelated to the extent of information likely available. For example, the monkeys species group includes less than one page of text while the amphibians group has four pages of text. In summary, Extinction provides an enormous amount of anecdotal information that is interesting to read, but which is likely forgotten soon after having read it. The book leaves the reader with a sense of disaster for all creatures as a result of human activity (which may or may not be far off the truth) and attempts to deal with all the ills of humankind from switching off light bulbs to buying local produce. Unfortunately readers that might be inclined to read Endangered will likely come away knowing little more than they start- ed with. BRENT TEGLER North-South Environmental Inc., 35 Crawford Crescent, Unit U5, P.O. Box 518, Campbellville, Ontario LOP 1B0 Canada The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His Theory of Evolution By David Quammen. 2006. Atlas Books/ W. W. Norton & Company, New York. 304 pages. $30.00 Cloth. As naturalists, we never cease to be amazed by the myriad wonderful life forms in nature, be it the flow- ering plants that suddenly appear following snowmelt, the wide variety of birds that return north every spring, or the colourful mushrooms found each fall in our forests. However, the source of this very diversity may easily be perplexing: why are there so many species and how did they originate? The theory of evolution has gone a long way towards explaining these and related questions by means of a wealth of empirical and the- oretical studies. 2006 Evolution by natural selection as [formulated] pro- posed by Charles Darwin is one of the most impor- tant and far-reaching ideas ever formulated, as it pro- vides the basic mechanism whereby evolution has created the enormous diversity of life. David Quam- men’s objectives with his book were to offer a con- cise yet insightful overview of the background, birth and development of Darwin’s idea of natural selec- tion, and to consider the reasons for Darwin’s reluc- tance in making his theory public. Indeed, there were many reasons why Darwin might have wanted to keep his evolutionary views to himself. These included the fear of rejection from a science corps whose natural history was based largely on teleological ideals with foundations in religious agendas, as well as his person- al relationship with his beloved wife Emma, a pious Christian. In the end, it was Darwin’s receipt of a man- uscript from Alfred Russel Wallace, who at the time was collecting beautiful bird and insect specimens some- where in the Malay Archipelago, which prompted the eventual publication of On The Origin of Species in 1859. Quammen outlines this momentous piece of histo- ry concisely and does not attempt to compete with other more exhaustive biographies that provide a more detailed and scholarly account of Darwin’s idea. The result is a very readable book that, despite the fact that Quammen is not an academic scientist, is based on thorough research of original background material. Hence, this book follows in the tradition of Quammen’s previous successes, most notably The Song of the Dodo (1996), his outstanding and award Book REVIEWS 253 winning volume on island biogeography. Several interesting aspects of Darwin's life are also high- lighted, such as his mysterious illness, characterized by dizziness, stomach pain, and bouts of illness, often flaring up under stressful conditions. Darwin spent weeks on end at health clinics, surrendering his body to coldwater treatments, hoping to rid himself of the persistent ailment. The book also describes a number of the small natural history experiments that Darwin performed to understand the mechanisms of evolution, such as submerging common vegetable seeds in brine solution to investigate the possibility of over water dispersal, or dangling duck feet in water to see if gastropods would climb aboard on a flight destined for other ponds. With these sorts of amusing passages highlighting the development of the most important idea in biology, The Reluctant Mr. Darwin should be of great interest to naturalists regardless of whether they are deeply aware of evolu- tionary theory or simply familiar with Darwin’s work. The good quality of the material in this vol- ume, along with Quammen’s command of the Eng- lish language, makes this book well worth the read. Literature Cited Quammen, D. 1996. The Song of the Dodo — Island Biogeography in an Age of Extinctions. Hutchinson/Pimlico/Random House of Canada Limited, Mississauga, Ontario. JOHAN LINDELL 35 Charles Street West, Apartment 1408, Toronto, Ontario M4Y 1R6 Canada A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s Drive of Vancouver By Rick Hudson. 2006. Volume 2. Revised Edition. Harbour Publishing Co. Ltd., P.O. Box 219, Madeira Park, British Columbia VON 2HO Canada. 368 pages. 8 illustrations $16.95 CAD Rick Hudson has succeeded in expressing his pas- sion for mineral collecting through his field guide book. On a second level he also tries to promote a sense of adventure by teasing the reader to go out and explore or prospect for nature’s buried treasures. The author structured the book so the reader can easily cross-reference required information in the lengthy location lists. A “How to Use This Book” and a “Quick Reference Guide” help to quickly locate items of interest throughout the book. The book cover illustrates the hobby of mineral collecting as a family activity. Hudson educates the reader from first principles on how to become a Rock- hound. To accomplish this, he describes how to pre- pare yourself by choosing proper clothing, safety gear, collecting tools and maps. He warns the future min- eral collector about natural hazards they may encounter such as; rattlesnakes, grizzly bears and ticks. Ethics, access rights and mineral rights are also discussed to keep the collector within acceptable collecting prac- tices and to avoid unintentional trespassing. Mineral collecting is attractive to many people be- cause as you find an interesting mineral specimen, you want to learn how nature created it and or its geologi- cal history. The author included short geological des- criptions partitioned by collecting areas as well as min- ing histories of local economic importance within the collecting area. Colour plates illustrate the minerals found in some of the collecting sites and detailed occurrence maps are included in all the collecting area sections through- out the book. The colour plates represent minerals that are possible to find in the collecting sites. Government mineral index files were included in the locality infor- mation along with detailed road log descriptions. The author has made a successful merger of person- al knowledge and technical data to compile an inform- ative volume which can provide years of enjoyment for many future rockhounds and their families and friends. ROGER POULIN President, Sudbury Rock and Lapidary Society, 3171 Romeo Street, Val Caron, Ontario P3N 1G5 Canada 254 History of the Russian Zapovedniks 1985-1995 By F. Shtilmark. (Translated from Russian by G. H. Harper). 2003. Russian Nature Press (www.rusnatpress.org.uk). 307 pages, £30.95 This book makes a major contribution to the envi- ronmental history of Russia, its influence zone and also to the global nature heritage. The Zapovedniks - Russia’s protected nature reserve system — have with- stood the test of time, although it has been distorted by social and political factors. Due to the great Eng- lish translation by G. H. Harper, the Russian Nature Press offers us new views on conservation, as well as solutions to the current problems with protected areas world-wide. The original text is written by one of the leading Russian environmental writers and fascinat- ingly energetic activists providing for superb inside knowledge unknown to the western world. His visions are guided by principles such as “we have a duty to make our zapovedniks and national parks the pride of Russia...”. In this great book the reader will learn about Rus- sia’s nature, and foremost, about how to protect it dur- ing such fascinating and complex times like Tsarism, the rules of Lenin and Stalin, and Gorbachovs’ glas- nost and perestroika (the book manuscript was fin- ished 1996). Throughout the history of the Zapovedniks (in short called Zap’s) the role of the Academy of Sci- ence proves very significant. The composition and atti- tude of its board members towards protection vs. exploitation play a crucial role for Zap-Management (and species survival). Most readers will likely be surprised to learn that the Russian Zapovednik system actually acted as a model for setting up the global network of biosphere reserves. But to this very day there exists a funda- mental confusion in the western world regarding the terminology and meaning of Russian protected nature reserves. As described over the 307 pages, western- style National Parks represent only one component in the complex Russian system of Zapodniks, Zapoved- nosts, Zapovednayes and Zakazniks. A critical view of the author refers to National Parks simply as pseu- do-zapovedniks because “An absolute Zapovednik is an area of land or water that together with all products of nature on and in it (plants, animals, soils, rocks, min- erals) is preserved in perpetuity and in a state of invi- olability”. Many Zapovednys were originally used for the Tsars or as a prince’s hunting estate, e.g., the famous Kuban Hunting Reserve. So originally, these systems go back to land grants from the aristocrats. But the author takes great care in explaining that Qualified Zapovedniks have a weaker definition in their protection status, allowing for human use. A Zapovednost implies the complete withdrawal of an area of land from any eco- nomic use. The biological theory of Zapovednosts rests on the idea of an ecological equilibrium, and the capac- THE CANADIAN FIELD-NATURALIST Vol. 120 ity of organisms to regulate themselves in the absence of direct human influence. Zapovedaniye means a with- drawal of a certain area of nature from the sphere of ordinary economic activity for the attainment of par- ticular non-traditional economic, social and ecological purposes. It represents samples of “virgin nature select- ed for comparison with exploited land”. Zapovedny Management relates to a method of managing protect- ed territories and water bodies. Further, one can find Zapovednik Museums, as well as Game Zapovedniks and even Spa-Zapovedniks. Only the Zakazniks are areas set aside for hunting. Finally, the Etalons are pro- tected areas, model examples, representative for a spe- cific ecosystem type and completely free from human influence. Several black/white maps, portraits, tables and dia- grams are presented, and including with some paint- ed Russian scenery, turn this publication into a pre- cious book to own. The book makes a nice case that fur-bearing ani- mals were considered as Russia’s “Soft Gold”; they were one of the chief resources for foreign income. This wealth created the influential Moscow Fur Insti- tute. Besides the earlier over-exploitation of Russian beaver resources, the 20" century showed also a sharp decline of sable, and a subsequent nation-wide harvest moratorium was not really effective. Therefore, sable reserves were requested, and a Sable Zakaznik was set up by local people in Kamchatka as early as 1882. These Game Zapovedniks were called Superior Zap- ovedniks in which hunting was not allowed, and breed- ing programs were initiated. As elsewhere in the world, the tensions around pro- tected areas were intense. Borders of Zapovedniks were often not followed or defined even. Poaching was com- mon in taiga Zapovedniks or where “old believers”, Chinese bandits, and fur-traders lived. In Crimea and elsewhere, the designated Zap-guards and zoo-staff were tortured or even executed. Besides the many day to day struggles in Zapoved- niks, Lenin’s revolution then had a massive restruc- turing in place for them, resulting in the “Socialisa- tion of the Land”. It converted the entire country in a gigantic “national park”; ownership rights were removed with one stroke. Lenin was a hunter himself and had a great interest in nature affairs. That proved to be for- tunate for Russia’s nature, as well as for some aspects of the global environment. From that point on, the Russian Soviet Federal Socialist Republic (RSFSR, founded 25" October 1917) played a crucial role in the protected areas, and started its well known enthu- siasm for regulations. Some Russian Zapovednik administrators used the guiding principle “Nature is not a cathedral, but a work- shop”. And typical Zapovednik tasks dealt with the introduction (“Acclimatisation”) and control of animal populations. Culling was part of most Zapovedniks, 2006 and the culled game meat was contributed to the social food-supply or else was sold to Zapovednik staff. The elimination of crows, and cormorants as fish-eating birds took place Russia-wide. Bears, marten, gulls, skuas, crows, hawks and eagles got killed en masse. Extensive campaigns against wolves by poison and air were carried out for almost a century shaping the Rus- sian culture against predators to this very day. The text reads like a crime story. As felt in many other places in the world, the Stalinist revolution fur- ther led to more and major changes. Namely, a shift from nature protection to exploitation in the interest of the socialist construction. Protecting nature for the sake of nature was not en vogue anymore. Further, “Bourgeois” professors who comprised a large pro- portion of the conservation’s cadre in the Soviet state in the 1930s were eliminated, and many members of the Academy of Science got arrested. From then on, Zapovedniks were explored for minerals, coal and oil deposits, timber resources and fur bearing resources, culminating in slogans like “We must take the Urals apart’. The fetish of inviolability as a bourgeois theory got sacrified. Protection was now defined as exploita- tion. Under comrade Stalin, Zapovedniks turned into multi-purpose economic enterprises. It led to the view that planned exploitation of Natural Resources would be the main pre-condition of Nature Protection. From the 1960s onwards the Zapovednik system experienced a modernization. A. G. Bannikov and G. P. Dementey started to participate in the IUCN. Russian- Japanese international migratory bird conventions got signed. In 1970-71 the UNESCO biosphere reserves “Man and Biosphere” were established. In 1972 a U.S.- Russian agreement on protecting the natural environ- ment got accomplished, and the USSR Red Data book came into place. As early as 1975 multidisciplinary monitoring was demanded in Russia, a theme that is very relevant for today even, and not achieved by most countries yet. When cheap fuel got provided for that purpose to the Zapovedniks by the state, they got quickly swamped with motorboats, snow tractors and motorcycles. Heli- copters can be added to the list as well since flying was very common by then. The author explains that cater- pillars tracked and wheeled cross-country, and vehi- A Field Guide to the Identification of Pebbles By E. Van der Flier-Keller. 2007. Harbour Publishing, Box 219 Madeira Park, British Columbia VON 2HO Canada. 9 inch by 8 inch pamphlet, $7.95 Professor Van der Flier-Keller has produced a neat pebble guide for children. She has condensed funda- mental pebble geology into a single page. Then she portrays 34 types of rock as they appear when they are transformed into pebbles. She adds in a few man- made objects that also appear on beaches, such as glass. The pamphlet is a single long sheet of plasticized paper Book REVIEWS 25 a) cles ploughed up the tundra on Vrangel and Taymyr. In the 1990s, the U.S. American Soros and Mac- Arthur Foundations with their foreign funds gained considerable influence over Russia’s nature; WWF, Evraziya, the Japanese Society for the Protection of Birds, the Global Ecological Facility, and the Interna- tional Bank for Reconstruction & Development were other international players accounting for as much as one quarter of Western money on the expenditure for protecting Russia’s biodiversity. This foreign influence made many Russian politicians extremely uneasy. However, despite all of these gigantic efforts, and similar to the case in North America, nothing remains of the many original habitats such as the steppe. “We now need to introduce the specially-protected areas system almost forcibly to counterbalance irrational natural-resource-use, and to prevent deterioration of landscapes and valuable sites”. It becomes clear from the text that Russia and its Zapovedniks etc. are not set up for the information society at all, and thus Russian high quality online (raw) data are hard to come by still. Much crucial biodiver- sity information is published though in hardcopy and journals, or exists in filing cabinets, private ownerships and cabinets, and as expert knowledge. Otherwise, check the Russian Conservation News found online WWwW.russianconservation.org. One cannot find a better and centralized overview of Russian conservation history issues than this book; its detailed Appendices are further assets consisting of current and former Zapovednik lists, a Kyrillic ref- erence list and in-depth annotations for each chapter. I find that this book is a little too friendly on the wide- spread view in Russia that traditional Zapovednik man- agement caters primarily to the Russian race but leaves natives and other ethnical groups out. Marine protect- ed areas were apparently not really part of Russia's nature protection system either. However, “Come back in hundred years, then let’s talk” (And in the meantime, I highly suggest buying and reading this important book). FALK HUETTMANN Institute of Arctic Biology, Biology & Wildlife Department, University of Alaska-Fairbanks, Alaska 99775 USA with clear, “life” size photographs of pebbles. It is easy to use and is ideal for children as it does not get over- complicated and therefore overwhelming. For example, it only shows two forms of granite — pink and white. This is quite sufficient for an 8-year old to absorb. While the author has reduced the text to a simple ex- planation she does use the correct technical terms throughout. Younger children may need help with some of the terms (plutonic) or concepts (glacial deposits), but then this is what parents are for. The pamphlet folds 256 to 4.75 inches by 9 inches and can readily slip into a pocket or glove compartment. Although aimed at children this is a useful tool for the non-geologist adult who does not have the time (or inclination) to master the complex technology of geological science. I have a habit of picking up small NEw TITLES Prepared by Roy John + Available for review * Assigned ZOOLOGY The Amphibians of Belarus. By S. Drobenkov, R. Novitsky, L. Kosova, K. Ryzhevich, M. Pikulik. 2006. Pensoft Pub- lishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 176 pages. EURO 34.00 Paper. The Amphibians and Reptiles in Bulgaria. By V. Beshkov and K. Nanev. 2006. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 120 pages. EURO 38.90 Cloth. * Animal Skulls, A Guide to North American Species. By Mark Elbroch. 2006. Stackpole Books, 5067 Ritter Road, Mechanicsburg, Pennsylvania 17055 USA. 448 pages. $44.95 PB 740 pages. Atlas of Bird Migration. By J. Elphick (Editor) 2006. Fire- fly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 176 pages. $35 Cloth. Birder’s Companion. By S. Moss. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 192 pages, $16.95 Paper. * Bird Conservation Implementation and Integration in the Americas: Proceedings of the Third International Part- ners in Flight Conference [on CD]. By C. J. Ralph and T. Rich. 2007. USDA, Redwood Sciences Laboratory, 1700 Bayview Drive, Arcata, California 955231 USA. Birds of the Dominican Republic and Haiti. By S. Latta, C. Rimmer, A. Keith, J. Wiley, H. Raffaele, K. McFarland and E. Fernandez. 2006. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 360 pages. US. $35. Birds: A Visual Guide. By J. Burger. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 304 pages. $29.95 Cloth. Birds of the World. Edited by A. Gossler. 2006. Firefly Books, 66 Leek crescent, Richmond Hill, Ontario L4B 1H1 Canada. 384 pages. $39.95 Cloth. A Birdwatcher’s Companion to North American Birdlife. By C. Leahy. 2006. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 1072 pages. US. $19.95 Southern African Birdfinder. Callan Cohen, Claire Spot- tiswoode and Jonathan Rossouw. 2006. New Holland Pub- lishers (UK) Ltd., Garfield House, 86-88 Edgware Road, London W2 2EA, U.K. 456 pages, £19.99 Paper 100 Butterflies and Moths — Portraits from the tropical forests of Costa Rica. By J. Miller, D. Janzen and W. THE CANADIAN FIELD-NATURALIST Vol. 120 rocks and pebbles from the places I visit. I have not been consistent in identifying them, but I am sure this pamphlet will help me improve. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Hallwachs. Harvard University Press, 79 Garden St., Cam- bridge, Massachusetts 02138, USA. 260 pages. U.S. $39.95. *A Field Guide to North Atlantic Wildlife. By N. Proctor and P. Lynch. 2006. Yale University Press, P.O. Box 209040, New Haven, Connecticut 06520-9040 USA. 221 pages. U.S. $19.95. + Conservation of the Black-tailed Prairie Dog. Edited by J. Hoogland. 2006. Island Press, 1718 Connecticut Avenue, NW, Suite 300, Washington D.C. 20009. 368 pages. U.S. $35 Paper. + The Northern Goshawk: A technical Assessment of its Status, Ecology and management. Edited by M. Morrison. 2006. Cooper Ornithological Society, c/o Western Founda- tion of Vertebrate Zoology, 439 Calle San Pablo, Camarillo California 93012-8506, USA. 369 pages. U.S. $23 Paper. * Migrating Raptors of the World — Their Ecology and Conservation. By Keith L. Bildstein. 2006. Cornell Universi- ty Press, 512 East State Street, Ithaca, New York 14850 USA. 344 pages. $35.00 Cloth. Rattlesnake Adventures: Hunting with Oldtimers. By J. Kemnitzer, Jr. [Editor]. 2006. Krieger Publishing Company, P.O. Box 9542, Melbourne, Florida 32902-9542. 236 pages. US. $32.50. Atlas of the Millipedes (Diplopoda) of Britain and Ire- land. By P. Lee. 2006. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 216 pages. EURO 32.00 Cloth. An Atlas of the Reptiles of North Eurasia. Taxonomic Diversity, Distribution, Conservation Status. By N. Anan- jeva, N. Orlov, N. Khalikov, R. Darevsky, I. Ryabov, I. Bara- banov. 2006. Pensoft Publishers, Sofia-Moscow. 250 pages. EURO 75.00 Cloth. * Songbird Journeys: Four Seasons in the Lives of Migra- tory Birds. By Mary Chu. 2006. Walker & Company, 104 Fifth Avenue, New York, New York 10011 USA. 312 pages. U.S. $23.00 Cloth. Snake Venoms and Evenomations. By J-P. Chippaux. 2006 English Edition. Krieger Publishing, P.O. Box 9542, Mel- bourne, Florida 32902 USA. 300 pages. U.S. $58.50. + Tracks. By D. Jackson. 2006. University Press of Missis- sippi, 3825 Ridgewood Road, Jackson, Mississippi 39211 USA. $36.25 279pages. Cloth. The Alligator Snapping Turtle. By P. Pritchard. 2006 re- print. Krieger Publishing, P.O. Box 9542, Melbourne, Flori- da 32902 USA. 152 pages. U.S. $42. 2006 * Turtles of the World. By F Bonin, B Devaux and A Dupre. 2006. The Johns Hopkins University Press, 2715 North Charles Street, Baltimore, Maryland 21218-4363, USA. 416 pages. U.S. $50 Cloth. Whales, Dolphins and Seals — A Field Guide to the Marine Mammals of the World. By Hadoram Shirihai and Brett Jarrett. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN, U.K. 384 pages, £12.99 Cloth. BOTANY Botanical Riches — Stories of Botanical Exploration. By R. Aitken. 2007. Ashgate Publishing, Box 2225, Williston Vermont 05495. 256 pages. U.S. $ 50 Cloth. Wildflowers of the Rocky Mountains. By G. Scotter. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 256 pages. $29.95 Paper. * Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region. By Paul Martin Brown. 2006. Uni- versity Press of Florida, 15 Northwest 15" Street, Gaines- ville, Florida 32611-2079 USA. 366 pages. U.S. $ 29.95 * Wild Orchids of the Pacific Northwest and Canadian Rockies. By Paul Martin Brown. 2006. University Press of Florida, 15 Northwest 15" Street, Gainesville, Florida 326] 1- 2079 USA. 304 pages. U.S. $ 29.95 ENVIRONMENT Algonquin Souvenir. By M. Runtz. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 96 pages. $19.95 Cloth. An Artist’s and Photographer’s Guide to Wild Ontario. By R. Stimson and C. Thompson. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 224 pages. $29.95 Paper. The Enchantment of Birds. By R. Cannings. 2007. Grey- stone Books, Suite 201, Quebec Street, Vancouver, British Columbia VST 4S7 Canada. 168 pages. $29.95 Cloth. The International Politics of Bird Conservation. By R. Boardman. 2006. Edward Elgar Publishing, 9 Dewey Court, Northampton, Massachusetts 01060 USA. 288 pages. U.S. $110 Cloth. * The Atlas of Climate Change. By Kirstin Dow and Thomas E. Downing. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 SXN, U.K. £12.99, Paper. Principles of Population Genetics — 4" edition. By D. Hart and A. Clark. Sinauer Associates, Box 407, Sunderland, Massachusetts 01375 USA. 565 pages. U.S. $93.95. An Inconvenient Truth. By Al Gore. 2006. NHBS Environ- ment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN, UK. £14.99. The Rough Guide to Climate Change. By Robert Henson. 2006. NHBS Environment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN. U.K. £9.99. Book REVIEWS * The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His Theory of Evolu- tion. By David Quammen. 2006. Atlas Books, 10 East 53rd Street, 36th Floor, New York, New York 10022, USA. 304 pages. U.S. $22.95. + Marshes. By W. Burt. 2007. Yale University Press. P.O. Box 209040, New Haven, Connecticut 06520-9040 USA. 192 pages. U.S. $35 Paper. Nature’s Engraver — A life of Thomas Bewick. 2007. Far- rar Straus Giroux, North Point Press, 19 Union Square West, New York, New York 10003. 480 pages. $27.50 Cloth. Wild Nova Scotia. By L.Wagg. 2006. Nimbus Publishing, Box 9166, Halifax, Nova Scotia B3K 5M8 Canada. 96 pages $29.95. People, Fish and Whales — The Vancouver Aquarium Story. By Dr. Murray Newman. 2006. Harbour Publishing, P.O. Box 219, Madeira Park, British Columbia VON 2HO Canada. 120 pages. $19.95 Paper. * Pilgrimage to Vallombrosa — From Vermont to Italy in the Footsteps of George Perkins Marsh. By John Elder. 2006. University of Virginia Press, 210 Sprigg Lane, Char- lottesville, Virginia 22903 USA. U.S. $33.99 Cloth. Sarcophaginae et Paramacronychiinae du Proche Orient (Insecta, Diptera, Sarcophagidae). By A. Lehrer. 2006. Pen- soft Publishers, Sofia-Moscow. [In French.}, 264 pages. EURO 64.00 Cloth. Tales from the Thébaide- Reflections of a Turtleman. By P. Pritchard. 2006. Krieger Publishing Company, P.O. Box 9542, Melbourne, Florida 32902-9542 USA. 340 pages. U.S. $44.50. Wild Borneo. By N. Garbutt and J. Prudente. 2006. The MIT Press, Five Cambridge Center, 4" Floor, Cambridge, Mass- achusetts 02142-1493 USA. 176 pages. $34.95 US. The End of the Wild. By S. Meyer. 2006. The MIT Press, Five Cambridge Center, 4“ Floor, Cambridge, Massachusetts 02142-1493 USA. 96 pages. U.S. $14.95. + Wildfire in the Wilderness. By C. Czajkowski. 2006. Har- bour Publishing, P.O. Box 219, Madeira Park, British Colum- bia VON 2HO Canada. 224 pages. $19.95. Endangered Wildlife on the Brink of Extinction. By G. McGavin. 2006. Firefly Books Ltd., 66 Lark Crescent, Rich- mond Hill, Ontario L4B 1H1 Canada. 142 pages. $35. Cloth. YOUNG NATURALISTS Tale of a Great White Fish — A sturgeon story. 2007. Grey- stone Books, Suite 201 Quebec Street, Vancouver, British Columbia VST 4S7 Canada. 48 pages. $10.95 Cloth. SPECIAL NOTE Sale of the Herpetological Library of Adrian Crane (800 items including antiques). PDF catalogue from Breck @herplit.com News and Comment Marine Turtle Newsletter (112) April 2006. 28 pages: ARTICLES: Nesting of Green Turtles in Saint Leu, Reunion Island (S. Ciccione and J. Bourjea) — Sea Turtles and Fishery Interactions in Brazil: Identifying and mitigating potential conflicts (M. A. Marcovaldi, G. Sales, J. C. A. Thome, A. C. C. Dias da Silva, B. M. G. Gallo, E. H. S. M. Lima, E. P. Lima, C. Bellini) — First Report of Leatherback Turtle Entanglement in Trap Lines in the Uraguayan Continental Shelf (M. Laporta, P. Miller, S. Horta, and G. Riestra) — Loggerhead Turtle Nesting Activity in Kuriat Islands (Tunisia): Assessment of Nine Years Monitoring (I, Jribi, M. N. Bradai and A. Bouain) — Hawksbill Turtles on the Pacific Coast of Costa Rica (A. Gaos, R. Arauz and I. Yanez) — NOTES: Oceanic Movement of Benthic Foraging Juvenile Hawksbill Turtle from The Cocos (Keeling) Islands (S. D. Whiting and A. U. Koch) — Use of Marine Turtles in Zootherapy in Northeast Brazil (R. Romeu da N. Alves) — MEETING REPORTS — ANNOUNCE- MENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICA- TIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, Centre for Ecology and Conservation, University of Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 263700. Subscriptions and donations towards the pro- duction of the MTN can be made online at or postal mail to Michael Coyne (online Editor) Marine Turtle Newsletter, 1 Southampton Place, Durham, North Carolina 27705 USA (e-mail: mcoyne@ seaturtle.org). Herpetological Survey 2006 Atlas of Amphibians and Reptiles of Quebec Instructions for participants in the 19" herpetological sur- vey for the Quebec Atlas of amphibians and reptiles, coordi- nated by the St. Lawrence Natural History Society in col- laboration with the Ministére des Ressouces naturelles de la Faune du Québec. This provincial inventory is done every year with the important cooperation of volunteers. Observation cards to fill out and details of information needed can be ob- tained from David Rodrique, Director, Research and Con- servation, Program Coordinator, or Mathieu Oulette, Research and conservation, St. Lawrence Valley Natural History Society, Ecomuseum, 21 125 chemin Saint-Marie, Saint-Anne-de- Bellevue, Quebec H9X 3Y7; phone 514-457-9449, extension 105; fax 514-457-0769; e-mail: info@herpetofaune.org. Cur- rent information on The Atlas of Amphibians and Reptiles of Quebec is now available online at the St. Lawrence Natural History Society website: www.herpetofaune.org. Bill Cody Receives Yukon Biodiversity Awareness Award From 1946 to 1987 Bill (W.J.) Cody was employed with Agriculture and Agri-Food Canada as plant tax- onomist and curator of Canada’s largest collection of plants. From 1988 to 2006 he has continued to come in to work every day as an Honourary Research Associate and he has made a series of remarkable contributions to our knowledge of the plants of northwestern North America. His 643 page textbook on the Flora of the Yukon was published in 1996. It included current information on status, distribution, ecology, classifica- tion and identification. In 1997 this outstanding work was recognized with the prestigious Lawson Medal awarded by the Canadian Botanical Association. An updated second edition of the book was published by the National Research Council in 2000. Since his retirement Bill has published 14 articles on the flora of the Yukon in scientific journals. He has provided an enormous amount of information on plants to the agricultural sector, natural resources staff, wildlife biol- ogists, native people and landscape planners. Always willing to help, he has identified a thousand plants from the Yukon every year. It is therefore not a sur- prise that Bill has been recognized for his outstanding service with the Yukon Biodiversity Awareness Award. The award honours those who have made major con- tributions to educating people about biodiversity and its importance. 258 ) 2004 Bill started as an assistant with the Canada Depart- ment of Agriculture in Ottawa. Twenty-one years later in 1967 he was made a Research Scientist. This clas- sification was generally reserved for people who had a Ph.D., but Bill’s outstanding accomplishments at that time were judged by his colleagues and the sci- ence arm of the federal civil service to warrant treat- ment at the Ph.D. level. The awards that Bill has received since then are numerous, but one in partic- uar draws attention his broad contribution. In No- vember 2002 he received a Queen’s Golden Jubilee Commerorative Medal. These medals were awarded to a limited number of people who have made a sig- nificant contribution to Canada, in this case “especial- ly for his work on The Canadian Field-Naturalist, Canada’s foremost scientific journal for field biology.” Bill has served as the business manager, article and book review contributor, and manuscript reviewer for this journal for 60 years, and his influence on its devel- opment, support, improvement and content is beyond question. Bill also served as curator of the largest dried plant collection in Canada from 1959 to 1988 and was large- ly responsible for its development. This collection of now over | million specimens has become a major tool of Agriculture and Agri-food Canada for plant identi- fication. This collection provides a wealth of informa- Errata The Canadian Field-Naturalist 120(1) NEWS AND COMMENT 259 tion that is needed to implement Agriculture and Agri- food Canada’s Biodiversity Strategy and to respond to the International Convention on Biodiversity. It is particularly valued as a tool for improved plant clas- sification studies used by researchers in other institu- tions both in Canada and worldwide. It is also impor- tant in protecting Canada’s borders and enforcing federal regulations. Many thousands of specimens that Bill collected in the northern wilderness under extreme and dangerous conditions are part of this collection and serve as vouchers for his numerous publications and books. Bill was born in Hamilton on 2 December 1922. His father was a doctor and his mother a nurse at Hamilton General. He grew up in Hamilton and received his B.A. from McMaster University in 1946. The Yukon Biodiversity Awareness Award plaque that Bill received included a photo of one of Yukon’s rarest plants, McBride’s Phacelia (Phacelia mollis) which is a Beringian endemic (confined to the ungla- ciated area of Alaska and Yukon). Text slightly modified from that provided courtesy of Paul Catling, Canada Agriculture and Agri-food, Ottawa. Photograph of the award plaque courtesy Stephen Daryshire, Canada Agricuture and Agri-food, Ottawa. Book Review. Lapland a Natural History, pages 123-124: replace “Redcliff’ and Redcliffe” with Ratcliffe, throughout. Articles. Diversity and range of amphibians and reptiles of the Yukon Territory. Brian G. Slough and R. Lee Menell in Literature Cited page 91 “Matsurla” should read Matsuda. Recent Declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces. Anthony J. Erskine. Page 48 insert following Dunn et al. Erskine, A. J. 1980. A House Sparrow die-off. Nova Scotia Bird Society Newsletter 22: 183-184. For reference following Erskine 1992b insert Erskine before initials. Editor’s Report for Volume 119 (2005) Mailing dates for issues in volume 119: (1) 6 June 2006, (2) 13 June 2006, (3) 30 November 2006, (4) 17 April 2007. A summary of membership and sub- scriber totals 2004 is given in Table 1. The number of articles and notes in volume 119 is summarized in Table 2 by topic; totals for Book Reviews and New Titles are given in Table 3, and the distribution of con- tent by page totals per issue in Table 4. Council continued to authorize 40% of membership dues for publication of the journal and all of subscrip- tions (both individual and institutional). Council also has allocated 80% of the annual interest from the Man- ning Fund and other capital funds to The Canadian Field-Naturalist. Use of the Manning fund portion was broadened on an issue-by-issue basis to offset the publi- cation costs where author and institutional contributions were insufficient to cover page charges. The journal was printed at Gilmore Printers, Ottawa, and thanks are due business representative Tom Smith, customer representative Chuck Graham for overseeing production, and Wendy Cotie for typesetting and cor- rections. Leslie Cody prepared the Index for volume 119; Elizabeth Morton proofed the galleys. Long-time (60 years) Business Manager for the journal, Bill Cody, gradually relinquished much of his active role during the year and some of his duties were shouldered by treasurer Frank Pope and assistant treasurer Jim Ward. Roy John processed books sent by publishers, assigned reviewers, edited book reviews and prepared the New Titles listing. Sandra Garland the Ottawa Field- Naturalists’ Club webmaster, posted contents and ab- stracts from each issue as it appeared, and sent pdfs of articles and notes ordered by authors. Manuscripts (excluding book reviews, notices, and reports) submitted to The Canadian Field-Naturalist totalled 75 in 2005, equalling the total for 2004. The following reviewed papers submitted in 2005 (with number of manuscripts re- viewed in parentheses if more than one): TABLE 2. Number of articles and notes published in The Canadian Field-Naturalist Volume 119 (2005) by major field of study. Subject Articles Notes Total Mammals 19 17 36 Birds 14 5 19 Amphibians + reptiles 2 1 3 Fish 6 1 7 Invertebrates 7 2 9 Plants 13 1 14 Other Ie 0) 1 Totals 62 Dl 89 Associate Editors: R. Anderson, Canadian Museum of Nature, Ottawa, On- tario (2); C. D. Bird, Erskine, Alberta (9); R. R. Campbell, St. Albert, Ontario (3); P. M. Catling, Agriculture and Agri- food Canada, Ottawa (7); B. W. Coad, Canadian Museum of Nature, Ottawa (8); A. J. Erskine, Sackville, New Brunswick || (12); D. F. McAlpine, New Brunswick Museum, Saint John, New Brunswick (9); D. W. Nagorsen, Mammalia Biological Consulting, Victoria, British Columbia (9); W. O. Pruitt, Jr., University of Manitoba, Winnipeg (16). Additional Reviewers: W. B. Ballard, Texas Tech University, Lubbock, Texas (3); L. Bernatchez, Laval University, Quebec, Quebec; J. R. Bider, Baie o’ Urfe, Quebec; G. Blouin-Demers, University of Ottawa, Ontario; D. Boyd, Helena, Montana; M. Brigham, University of Regina, Saskatchewan; L. Carbyn. Canadian Wildlife Service, Edmonton, Alberta (3); J. Cayou- ette, Agriculture and Agri-Food Canada, Ottawa; J. Cebek. Trent University, Peterborough, Ontario; T. Chubbs, Happy Valley — Goose Bay, Labrador, Newfoundland & Labrado1 (3); H. D. Cluff, Resources, Wildlife & Economic Develop- ment, Yellowknife, Northwest Territories (2); M. Crete, Mini- stre de l’Environment, Quebec, Quebec; A. W. Diamond University of New Brunswick, Fredericton, New Brunswick (2); L. C. Dickamer, Northern Arizona University, Flagstaff Arizona; P. Dobson, School of Veterinary Medicine, Phila-) delphia, Pennsylvania; J. Eger, Royal Ontario Museum, Tor: TABLE 1. The 2005 circulation of The Canadian Field-Naturalist (2004 in parenthesis). Compiled by Robin Murphy, bmrgroup Ottawa. Canada USA Other Totals Memberships Family & individual 624 (717) 26 (26) 6 (6) 656 (749) Subscriptions Individuals 130 (165) 63 (63) 4 (5) O33) Institutions 156 (164) 229 (246) 34 = (30) 419 (440) TOTALS 910 (1046) 318 (335) 44 (41) 1272 (1422) Note: 22 countries are included under “Other” (outside Canada and United States): Austria, Belgium, Brazil, Denmark (2), Unite Kingdom (9: including 1 to Scotland), Finland (2), France (3: including 1 to St. Pierre & Miquelon), Germany (2), Icelanc Ireland, Japan, Mexico, Netherlands (3), New Zealand, Norway (4), Poland, Russia, South Africa, Spain (3), Sweden (2 | Switzerland (2), Trinidad and Tobago. 260 2005 TABLE 3. Number of reviews and new titles published in Book Review section of The Canadian Field-Naturalist Volume 119 by topic. ——— — — _ _ _— —_—_ — — EEEEEns=gQTeaeaEs«q@nanana=anaaa——— Reviews New Titles Zoology 33 104 Botany 5 19 Environment 17 10 Miscellaneous 6 20 Young Naturalists 0 2 Totals 61 lis) onto, Ontario; W. Foissner, Institut fur Zoologie, Salzburg, Austria; B. Freedman, Dalhousie University, Halifax, Nova Scotia; W. A. Fuller, Athabasca, Alberta; D. M. Green, Red- path Museum, McGill University, Montreal, Quebec (4); P. T. Gregory, University of Victoria, British Columbia (3); E. Haber, National Botanical Services, Ottawa, Ontario; F. H. Harrington, Mt. St. Vincent University, Halifax, Nova Scotia; -Y.L. Harms, University of Saskatchewan, Saskatoon; T. Hax- ton, Ministry of Natural Resources, Kemptville, Ontario (2); | S. Herrero, University of Calgary, Alberta (2); G. Holroyd, Can- cadian Wildlife Service, Edmonton, Alberta; E. Holm, Royal Ontario Museum, Toronto; S. Hooker, University of St. An- ‘drews, Fife, United Kingdom; C. S. Houston, Saskatoon, Sas- katchewan; R. James, Sutherland, Ontario (2); M. Johnson, Game & Fish Department, Bismarck, North Dakota; T. S. Jung, ‘Environment, Fish and Wildlife Branch, Whitehorse, Yukon (2); R. W. Knapton, Edmonton, Alberta; K. Larsen, University ‘College of the Cariboo, Kamloops, British Columbia; J. Lien, ‘Memorial University of Newfoundland, St. John’s; R. Mac- ‘Culloch, Royal Ontario Museum, Toronto, Ontario; F. F. Mal- ‘lory, Laurentian University, Sudbury, Ontario (2); J. Maunder, ‘Pond Cove, Newfoundland (3); L. D. Mech, U.S. Geological Survey, The Raptor Center, University of Minnesota, St. ‘Paul (3); R. Meisterfeld, Institute for Biology II (Zoology), ‘Department of General Biology, Aachen; D. Naughton, Cana- ‘tian Museum of Nature, Ottawa; J. S. Nelson, University of Alberta, Edmonton; J-P. Ouellet, Universite du Quebec a Ri- ‘nouski; P. Paquet, Meacham, Saskatchewan (3); D. Paulson, Seattle, Washington; G. Parker, Canadian Wildlife Service, sackville, New Brunswick; S. Petrie, Bird Studies Canada, ! Port Rowan, Ontario; G. Proulx, Alpha Research & Manage- ment Ltd., Sherwood Park, Alberta (2); Raine, Golder Asso- viates Ltd., Calgary, Alberta; R. Reeves, Okapi Wildlife Asso- Epiror’s REPORT 26) TABLE 4, Number of pages per section published in The Cana dian Field-Naturalist Volume 119 (2005) by issue (1) (2) (4) Total Articles 128 132 118 95 473 Notes 12 K 21 16 57 Book Reviews’ 15 20 1] 19 65 CFN/OFNC Reports” 0 2 x 4 14 News and Comment 2 2 | | 6 Index 0 0 0 32 32 Advice to Contributors | 0 | | Totals 158 164 160 168 650 * Total pages for book review section include both reviews and new titles listings. ™ Includes CFN Editor's report in (2), OFNC Annual Business Meeting (3) and OFNC Awards (4). ciates, Hudson, Quebec; J. D. Rising, University of Toronto, Ontario; L. Rogers, Ely, Minnesota; R. Rosatte, Ministry of Natural Resources, Trent University, Peterborough, Ontario; A. Russell, University of Calgary, Alberta; S. Sampson, Utah Museum of Natural History, Sault Lake City; Fred Scott, Acadia University, Wolfville, Nova Scotia; F. W. Schueler, Bishops Mills Natural History Center, Oxford Mills, Ontario (6); F. E. Schwab, College of the North Atlantic, Labrador City, Newfoundland and Labrador; D. Smith, Southern Connecticut State College, New Haven; K. W. Stewart, University of Manitoba, Winnipeg, Manitoba; J. Theberge, Oliver, British Columbia (3); I. Thompson, Canadian Forest Service, Sault Ste. Marie, Ontario (2); P. M. Youngman, Ottawa, Ontario. I am also indebted again to the President of the Ot- tawa Field-Naturalists’ Club Gary McNulty and the Club Council for continuing support of the journal; Chair- man Ron Bedford and the Publications Committee of the OFNC for editorial encouragement and support, the finance committee for input and encouragement to speed things up, the Canadian Museum of Nature for access to its library and the facilities at the Natural Heritage Building, 1740 Pink Road, Aylmer, Quebec, and to Joyce for continuing support. FRANCIS R. COOK Editor Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit by post to the Editor, in either English or French, three complete manuscripts written in the journal style. Manuscripts may also be submitted (one copy) by e- mail. The research reported should be original. It is recom- mended that authors ask qualified persons to appraise the paper before it is submitted. All authors should have read and approved it. Institutional or contract approval for the publica- tion of the data must have been obtained by the authors. Also authors are expected to have complied with all pertinent leg- islation regarding the study, disturbance, or collection of ani- mals, plants or minerals. The place where voucher specimens have been deposited, and their catalogue numbers, should be given. Latitude and longitude should be included for all indi- vidual localities where collections or observations have been made. Manuscripts should be printed on standard-size paper, dou- blespaced throughout, generous margins to allow for copy marking, and each page numbered. For Articles and Notes provide a bibliographic (citation) strip, an abstract, and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manu- scripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in a separate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “British” or “American” spellings are acceptable in English but should be consistent within one manuscript. The Oxford English Dictionary, Webster’s ’ New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Illustrations Photographs should have a glossy finish and show sharp contrasts. Electronic versions should be high resolution. | Photographic reproduction of line drawings, no larger than a standard page, are preferable to large originals. Prepare line drawings with India ink on good quality paper and let- ter (don’t type) descriptive matter. Write author’s name, title of paper, and figure number on the lower left corner or on © the back of each illustration. Reviewing Policy Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor (who | reviews it or asks another qualified person to do so), and at | least one other reviewer, who is a specialist in the field, cho- sen by the Editor. Authors are encouraged to suggest names | of suitable referees. Reviewers are asked to give a general | appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—some- times extensive revision and reappraisal. The Editor makes | the final decision on whether a manuscript is acceptable for — publication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by pay- ing $90 for each page, plus $20 for each illustration (any size up to a full page), and up to $90 per page for tables (depend- ing on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Editor. If grant or institutional funds are not available, club members and sub- scribers may apply for a waiver of charges for the first five — pages. Limited joumal funds are available to help offset publi- | cation charges to authors without grants or institutional sup- | port. Requests for financial assistance should be made to the | Editor when the manuscript is submitted. An order form for the purchase of repents will accompa- ny the galley proofs sent to the authors. Invoices for publi- | Reprints | cation costs will be sent when the submission is published. | FRANCcIs R. Cook, Editor |) RR 3 North Augusta, Ontario KOG IRO Canada || 262 TABLE OF CONTENTS (concluded) Volume 120 Number 2 Osprey, Pandion haliaetus, depredates Common Eider, Somateria mollissima, duckling BRENDA M. BLINN, VICKY VIOLETTE, and ANTONY W. DIAMOND Extension de l|’aire de distribution de vase, Umbra limi, dans le nord-est du Québec JEAN-FRANCOIS DESROCHES The Mink Frog, Rana septentrionalis, in southeastern Labrador JEAN-FRANCOIS DESROCHERS, ISABELLE PICARD, and JOHN E. MAUNDER First record of the Great Barracuda, Sphyraena barracuda, from Canada DARLENE BALKWILL, BRIAN W. COAD, ISMAEL GALVEZ, and JOHN GILHEN Book Reviews Zoo.LoGy: Animal Skulls: A Guide to North American Species — A Field Guide to North Atlantic Wildlife — A Complete Guide to Arctic Wildlife — The Black Flies (Simultidae) of North America — Insects: Their Natural History and Diversity: With a Photographic Guide to Insects of Eastern North America — The Northern Goshawk: A Technical Assessment of its Status, Ecology and Management — Turtles of the World — The Wolves of Algonquin Park: A 12-Year Ecological Study Borany: Flowers: How They Changed the World ENVIRONMENT: Guide to Deserts — Endangered: Wildlife on the Brink of Extinction MISCELLANEOUS: The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His Theory of Evolution — A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s Drive of Vancouver — History of the Russian Zapovedniks 1985-1995 — A Field | Guide to the Identification of Pebbles NEW TITLES News and Comment Marine Turtle Newsletter (112) — Herpetological Survey 2006 Atlas of Amphibians and Reptiles of Quebec — Bill Cody Receives Yukon Biodiversity Awareness Award — Errata: The Canadian Field- Naturalist 120(1) Editor’s Report for Volume 119 (2005) po to Contributors i Mailing date of the previous issue 120(1): 16 July 2007 2006 258 260 262 THE CANADIAN FIELD-NATURALIST Volume 120 Number 2 Articles George Wayne Douglas 1938-2005 JENIFER L. PENNY Conservation evaluation the Prairie Lupine, Lupinus lepidus var. lepidus, in Canada GEORGE W. DOUGLAS and MICHAEL RYAN Conservation evaluation of Seaside Bird’s-foot Trefoil, Lotus formosissimus, in Canada GEORGE W. DOUGLAS and MICHAEL RYAN Conservation evaluation of Bog Bird’s-foot Trefoil, Lotus pinnatus, in Canada MARTA DONOVAN Conservation evaluation of Dwarf Wolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada GEORGE W. DOUGLAS, JENIFER L. PENNY, and KSENIA BARTON Conservation evaluation of Pacific Rhododendron, Rhododendron macrophyllum, in Canada GEORGE W. DOUGLAS and JULIE DESROSIERS Conservation evaluation of Slender Collomia, Collomia tenella, in Canada GEORGE W. DOUGLAS and JENIFER L. PENNY Conservation evaluation of Small-flowered Tonella, Zonella tenella, in Canada GEORGE W. DOUGLAS and JENIFER L. PENNY Conservation evaluation of Stoloniferous Pussytoes, Antennaria flagellaris, in Canada GEORGE W. DOUGLAS, JENIFER L. PENNY, and KSENIA BARTON Late-born Elk, Cervus elaphus, calf with spots observed near Bancroft, Ontario Rick ROSATTE and JOE NEUHOLD Annual variation in habitat use by White-footed Mice, Peromyscus leucopus, in forest fragments: the effects of forest patch size, edge, and surrounding vegetation type CHRISTINE S. ANDERSON, DOUGLAS B. MEIKLE, ALAN B. CApDy, and ROBERT L. SHAEFER Response of the Sea Louse Lepeophtheirus salmonis infestation levels on juvenile wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, salmon, to arrival of parasitized wild adult salmon ALEXANDRA Morton and Ros WILLIAMS Physical characteristics, hematology, and serum chemistry of free-ranging Gray Wolves, Canis lupus, in southcentral Alaska MATTHEW J. BUTLER, WARREN B. BALLARD, and HEATHER A. WHITLAW Are Lesser Snow Geese, Chen caerulescens caerulescens, exceeding the carrying capacity of the Fraser River Delta’s brackish marshes? MIKE W. DEMARCHI Predicting Raccoon, Procyon loter, occurrence through the use of microhabitat variables ROGER A. BALDWIN, ALLAN E. HousTON, MICHAEL L. KENNEDY, and PIN SHUO LIU Notes Scoter, Melanitta spp., interrupted migrations by Confederation Bridge: An update KATHERINE BUNKER-POPMA An observation of the spring 2006 migration of Black Scoter, Melanitta nigra, in Northumberland Straight, interrupted by the Confederation Bridge, New Brunswick—Prince Edward Island CoLIN MACKINNON and ANDREW KENNEDY First record of a River Otter, Lutra canadensis, captured on the northern coast of Alaska SHAWN P. HASKELL 2006 133 147 153 158 163 169 le 179 183 188 192 192 235, (continued on inside back cover) ISSN 0008-3550 (7) 1S aH The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 120, Number 3 July-September 2006 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Michaélle Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- ments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson 2006 Council President: Mike Murphy Ronald E. Bedford = Diane Kitching Dan Millar Vice-President: Ken Allison Fenja Brodo Karen McLachalan Hamilton Justin Peter Recording Secretary: Susan Laurie-Bourque Julia Cipriani David Hobden Stanley Rosenbaum Treasurer: Frank Pope William J. Cody Diane Lepage Henry Steger Past President: Gary McNulty Francis R. Cook Ann Mackenzie Chris Traynor Susan Howell Gillian Marston Eleanor Zurbrigg To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc@achilles.net. For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward our mailing costs. PAP Registration Number 9477. Canada Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca Copy Editor: Elizabeth Morton Honorary Business Manager: William J. Cody Acting Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwgsc.gc.ca Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen Charles D. Bird Brian W. Coad Donald F. McAlpine Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail | or e-mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. Subscriptions and Membership | Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per year | (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must add an addi- tional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: July— August 2006 (November 2007). Cover: A Screech-Owl peers down from a perch at the McGill Bird Observatory in the Stoneycraft Wildlife Area, Ste-Anne-de- Bellevue, Quebec. Photographed 11 October 2004 by Marcel Gahbauer; e-mail: marcel @migrationresearch.org. See article | by Richards, Mineau, Bird, Wery, Larivée and Duffe, pages 289-297. LIBRA™® wr AVARD The Canadian Field-Naturalist{,\veRs'” Volume 120, Number 3 July-September 2006 An Ethogram Developed on Captive Eastern Coyotes Canis latrans JONATHAN G. Way!, DEAN-LORENZ M. SZUMYLO?, and Eric G. STRAUSS? 'Boston College, Environmental Studies Program and Lynch School of Education, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA; e-mail: jw9802@yahoo.com Urban Ecology Institute and Environmental Studies Program, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA 3Boston College, Biology Department and Urban Ecology Institute, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA; e-mail: strausse@be.edu 4Present address: 64 Cranberry Road, Marstons Mills, Massachusetts 02648 USA; Corresponding author: e-mail: szumylo@ be.edu Way, Jonathan G., Dean-Lorenz M. Szumylo, and Eric G. Strauss. 2006. An ethogram developed on captive eastern Coyotes Canis latrans. Canadian Field-Naturalist 120(3): 263-288. We studied capture Eastern Coyotes (Canis latrans) from 27-585 days of age and compiled an ethogram on them. A total of 72247 15-sec samples were taken, amounting to 301 h of field time varying between 59.4—61.3 h per Coyote. A total of 540 behavioral patterns was observed amongst the 16 behaviour categories ranging from 9 (miscellaneous) to 72 (explore investigate) action patterns per parent category. The 16 parent categories that we believed best described and appropriately sorted the behavioural actions were resting, sitting, sitting], sitting2, standing, traveling, explore/investigating, hunting, feed- ing, infantile, greeting, self play, play initiating, playing, agonistic, and miscellaneous. Exploring accounted for >31% of all of the behaviours observed with resting and sitting (combined), standing, traveling, and play as categories decreasing in order of most to least frequent. Despite some omissions in our ethogram and drift associated with its ongoing development, we believe that the large amount of data collected made it rigorous enough to be a useful guide for the species. We argue that although future research will no doubt add to and/or modify components of it, its ease of use in the field (in captivity or in the wild) and it being the first complete ethogram described for the species, make it a useful tool for future researchers. Key Words: canid, Canis latrans, eastern Coyote, ethogram, behavioural study. The concept of an ethogram, also called action system (Makkink 1936), behavioural inventory (Bekoff 1972a, 1978), or behaviour pattern (Scott and Fuller 1965), is a durable tool for analyzing vertebrate be- haviour and dates back to Jennings (1906), Makkink (1936), Tinbergen (1959), and Lorenz (1971). Although there have been direct observation behavioural studies of Coyotes (Canis latrans) in captivity (Snow 1967; Silver and Silver 1969: Bekoff 1972a, 1978: Parks 1979; Ryon 1986) and in the wild (Camenzind 1978; Bekoff and Wells 1981; Gese et al. 1996), there has yet to be a standardized and complete ethogram published for the species. Except for Goodmann and Klingham- mer’s (1990) non-peer reviewed but detailed manual which presents a Wolf (Canis lupus) ethogram and Scott and Fuller’s (1965) ethogram on Domestic Dogs (Canis familiaris) (which they compared with literature on Wolves, Coyotes and Red Foxes [Vulpes vulpes]), we could not find any other study that described a full behavioural repertoire of any other of the approximate- ly 35 canid species (Ewer 1973; Wozencraft 1989). Most studies examining behavioural data have focused on specific time periods (e.g., young pups/behaviour- al development — Bekoff 1972a, 1978, 1989), specific actions (e.g., social interactions — Fox 1970; Bekoff 1974; Zimen 1982), lumping behaviours into general categories (Silver and Silver 1969; Parks 1979; Gese et al. 1996), or describing many behavioural patterns but providing no summary or chart showing a complete ethogram (Mech 1970; Lehner 1978). Schleidt et al. (1984) termed these “partial ethograms” and stated the importance of publishing complete ethograms on as many species as possible. In addition, Macdonald et al. (2000) underscored the significance of developing ethograms on different species by stating that without one, meaning is not fixed. As such, ethograms are vital reference material for assessing a study’s conclusions (Macdonald et al. 2000). The objective of this paper is to describe, in detail, the full range of behavioural acts that we have ob- served in a captive group of five Eastern Coyotes. This ethogram is intended to be: (1) reader-friendly in for- mat and applied and used in future direct observation behavioural studies of wild and captive Coyotes: and (2) compared with ethograms of closely related species (e.g., Wolves, Jackals [Canis sp.] and foxes) and taxa (e.g., ursids, procyonids, and mustelids: Ewer 1973). 263 Methods On 12 April 2002, five (2 M; 3 F) of an original lit- ter of nine (3 M; 6 F) estimated 25 day old (Parks 1979), wild-born sibling Coyote pups were removed from under a shed in a residential area in Falmouth (Cape Cod), Massachusetts, and were reared at JGW’s house (note: the other four pups [1M; 3F] were placed back underneath the shed for the wild parents to raise). The five Coyotes taken into captivity were contained within a 53 m? area both inside and outside of JGW’s house until 13 May 2002, when the Coyotes were trans- ferred to a 16 m? quarantine facility for one month at the Franklin Park Zoo, Boston, Massachusetts. On 13 June 2002 the five Coyotes were moved to their perma- nent 403 m? public viewable exhibit at the Stone Zoo located in Stoneham, Massachusetts, and remained there for the duration of this study until late October 2003 — pup age 585 days). We chose the endpoint of our study as the last day of data collection prior to the Coyotes being permanently separated due to intrapack aggression (Late and Trans [See Table 1] were re- moved). The area of their exhibit was described by Frank (1987) as a suitable size for a long-term study of canid behaviour. Although the Coyotes were hand- raised, we made no attempt to interfere with or disci- pline their activities and therefore gave the Coyotes free access, at all times, to their exhibit/living facilities. The Coyotes were provided with puppy milk (Esbilac, PetAg, Inc., Hampshire, Illinois) via bottle or bowl up until 15 May and were given access to water and dry dog chow (commercially available dog foods until March 2003 then Mazuri exotic canine chow/diet [PMI Nutrition International, LLC., Brentwood, Missouri] thereafter) at all times. They were group fed (i.e., all five at once) 0.75 — 1.6 kg (varying with their age) of Nebraska Brand chopped frozen canine meat (Central Nebraska Packing Co., North Platte, Nebraska) mixed with dog chow on a daily basis and were given frozen or thawed laboratory rats, guinea pigs and mice (do- nated from a rodent breeding facility), and/or bones 2-4 times per week. Within their exhibit, the Coyotes commonly hunted (at least | prey item 2-3 times daily) and captured (about 1-2 prey items per week) Starlings (Sturnus vulgaris), House Sparrows (Passer domesti- cus), Chipmunks (Jamias striatus), Brown/Norway Rats (Rattus norvegicus), Meadow Voles (Microtus pen- nsylvanicus), and Gray Squirrels (Sciurus carolinen- sis), but did not appear to eat much or any of these prey items. Care and use of animal subjects was approved by Zoo New England’s Institutional Animal Use and Care Committee (letter dated 23 January 2002 to JGW), by Boston College’s Institutional Animal Care and Use Committee Protocol Number 01-03, and by the Massa- chusetts Division of Fisheries and Wildlife Permit #052. 02LP. We developed an ethogram that noted basic behav- ioural patterns (Scott and Fuller 1965; Bekoff 1972a, 1972b, 1978; Parks 1979). To avoid influencing Coy- THE CANADIAN FIELD-NATURALIST Vol. 120 ote behaviour with different observers only JGW inter- acted with and conducted activity budgets on them. Us- ing this technique, we acknowledge that we did not have interobserver reliability as described by Bekoff (1974) but the large amount of data collected attempt- ed to offset intraobserver reliability. We used focal indi- vidual animal sampling (i.e., one Coyote per 30 min bout of data collection), took instantaneous point or scan samples (Martin and Bateson 1986; Macdonald et al. 2000) every 15 sec for 30 min on the target Coy- ote, recorded the date, time, and weather before each observation bout took place, and noted important con- textual information in between each 15 sec sampling period. By using frequent (i.e., every 15 sec) instanta- neous samples, we tried to obtain an accurate approxi- mation from continuous recording (Martin and Bateson 1986). Thus, 120 samples per 30 min behavioural bout on a Coyote were ideally recovered. However, there were two reasons why we occasionally did not obtain 120 samples per bout: (1) a Coyote was momentarily out of sight during a particular 15 sec sampling period (usually this happened at most, once per 30 min bout); and (2) we had to stop a sampling bout earlier (e.g., rain, darkness, or some kind of disturbance that forced JGW to abandon an activity budget). We randomly chose which Coyote to conduct observations on before entering the field but attempted to evenly sample all Coyotes (1.e., each Coyote was scored every fifth time) during the study. We typically recorded behavioural data about 5 days/week and took between | and 4 (us- ually 1-2) 30 min behavioural bouts/day during day- light hours. Behaviours were also recorded on digital still and videocameras and 35-mm film cameras week- ly for the duration of the juvenile period of the pups (Parks 1979) then ad lib after pups reached adulthood (one yr of age — Bekoff and Jamieson 1975). We con- ducted most of the behavioural bouts during early to mid-morning (ca. 07:00-11:00 h) or between late-after- noon to early-evening hours (ca. 16:00-19:30 h); these were the times with the fewest number of people around the zoo (the zoo opened at 10:00 and closed at 16:30- 1800 h depending on the time of year). Because the Coyotes were habituated to and did not react negative- ly to JGWs presence, he followed them around the exhibit similar to the description of Goodall’s (1986) “follows” of wild Chimpanzees (Pan troglodytes) in Gombe, Tanzania. JGW made an effort not to influ- ence the movements and/or behaviour of Coyotes by minimizing his movement in the exhibit. This was es- pecially important because the Coyotes would follow him around the exhibit when he interacted with them but would generally ignore him when he was stand- ing erect and writing on a clipboard (J. Way, unpub- lished data). “Following” was a necessary technique to use on the Coyotes because there was not a single ob- servation spot outside the exhibit where the Coyotes could reliably be seen all of the time (i.e., many times trees and shrubs concealed the Coyote under observa- 2006 tion). Besides the senior author’s presence, other human contact was kept to a minimum before and after each sampling period to avoid humans affecting Coyote behaviour. Context was a critical variable with respect to the Coyotes’ response to human behaviour. For ex- ample, they generally ignored people (unless very loud) on the public path but would intently watch and/or bark at staff members that were behind (i.e., the opposite side of the public path that was off-limits to non-employees) or approaching their exhibit. Accordingly, zookeepers did not enter the exhibit to feed the Coyotes when JGW interacted with them. Despite these precautions there were no doubt many instances where Coyotes changed their activity in response to a person’s (public or staff) presence — sometimes even when a person simply walked by their exhibit area. Similar instances of Coy- otes shifting their behaviour because of the presence of people have been documented in areas where wild Coyotes inhabit urbanized areas (Way 2001). We at- tempted to mitigate these factors by increasing the total size of the pool of sample bouts. We entered all of the data into an excel spreadsheet. First, we entered the raw data into the spreadsheet. Then we alphabetized that data in order to group sim- ilar behaviours for each budget. Next we summed the frequency of each distinct behaviour observed. Finally we entered those summed values into a separate file (for each Coyote) that had our developing ethogram. Due to the large amount of data and behaviour se- quences in our ethogram (Table 1), properly entering and compiling data were critical parts of this ethogram’s creation. Behavioural actions or patterns (denoted as subcategories in Table 1), as defined by Scott and Ful- ler (1965), were grouped into “parent” categories, or behavioural complexes as described by Schleidt et al. (1984), or behavioural systems, as noted by Scott and Fuller (1965), based on motivational context and activ- ity observed (Parks 1979). In our ethogram, we organized the categories and the behavioural actions within each of the complexes generally on a scale from less to more active or intense an activity, then from solitary to group-oriented activ- ities. Notable exceptions were feeding, which was grouped next to hunting for obvious reasons, and self play, which was grouped close to social play but was of a solitary endeavor. The 16 parent categories that we believed best described and appropriately sorted the behavioural actions were resting, sitting, sitting], sit- ting2, standing, traveling, explore/investigating, hunt- ing, feeding, infantile, greeting, self play, play initiat- ing, playing, agonistic, and miscellaneous. There were instances (especially early in our study when we were working on differentiating different behavioural ac- tions) where a Coyote was engaged in an activity but we could not describe, other than in a general category, what the Coyote was doing. When this occurred (e.g., when a Coyote was behind trees and was only par- tially observed) we simply classified the behaviour in its parent category, so that we could at least provide a Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES 265 coarse description of a Coyote’s behaviour al a given 15 sec interval (Table 1). We calculated the percentage that each behaviour pattern was observed and added all within a parent cat- egory to examine the relative frequency that each behavy- ioural complex/system occurred (Table 1). In addition, we used the ages (in days) of the Coyotes to denote the first time that each behaviour was observed, with day 27 (14 April 2002) being the beginning point —i.c., the date we first did an ethogram on a Coyote. We want to stress here that although we report both the percent- age of each behaviour observed and the age that each behaviour was first recorded for comparison purposes (Table 1), the purpose of this manuscript is to describe the ethogram that we developed. We will more thor- oughly analyze these data in future manuscripts. Results and Discussion The ethogram that we developed on the Eastern Coyote is presented as Table 1. A total of 72247 15-sec samples were taken amounting to 301 h of field time, varying between 59.4-61.3 h per Coyote. A total of 540 behavioural patterns were observed amongst the 16 behaviour categories ranging from 9 (Miscellaneous) to 72 (Explore and Investigate) acts per parent category (Table 1; Figures 1-25). Resting (RE) occurred when a Coyote was lying down with its head on the ground. Sitting (SIT) was classified as sitting down either on rear or all four legs; this category was subsequently divided into sit] (SIT1) and 2 (SIT2). SIT1 was noted when a Coyote was sitting down on all four legs; in other words, it was almost ly- ing down but it had its head up. SIT2 was when a Coy- ote sat on its rear legs but its front legs were up in standing-like position. It was difficult to always reli- ably determine what the motivation was for a Coyote in the aforementioned positions. For example, a Coy- ote frequently sat in SIT1 position to sniff the ground. Because the animal was in a resting-like position we believed that it was important to separate these activi- ties from explore/investigate activities where the ani- mal was active (1.e., at least in a standing position). Similarly, it was sometimes difficult to understand the motivational context when an animal was just stand- ing (ST) and performing routine activities. Thus, we developed a category for just standing activities. Any of these categories could always be lumped together or put into different or new categories if the need arose — but the opposite cannot occur if the be- haviours are not split into discrete actions at the outset (Bekoff 1972b). For example, SIT1 Alt and SIT1 Alt Pt were very similar except for the panting involved. Al- though we could have easily combined these categories we thought that keeping them separated would allow us to potentially analyze this difference in the future (e.g., the level of panting on hot versus cold days). Resting and sitting behaviours were observed quite frequently and accounted for >23% of all of the etho- gram despite the Coyotes generally being much more Vol. 120 THE CANADIAN FIELD-NATURALIST 266 FOO O10) CO GOO ae 89) 19 hs) yh i 10 3=L9 Sus o[IYM Joalgo Suyysiug us OO COO COO OOO COO 1 © OW @ © 000 +6 SsuryyAue UO pasnsoj jou nq uado are sade ay SuNUe uado | jueg CAD FEW. OS) SEO OD OO OW. 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Traveling T was defined as a Coyote moving with purpose from one place to another (i.e., from point A to point B). A Coyote, when traveling, would move in a straight line and would waste little energy investigat- ing other objects until it reached its destination. This differed from explore/investigate E/I activities, where animals were characterized by actively sniffing objects or other Coyotes, digging, and moving around in a non-direct way in order to explore their surroundings. E/I behaviour is typically not described in the litera- ture for canids (but see Scott and Fuller 1965) yet was an important activity that our Coyotes engaged in, by far the most frequently of any parent category (Table 1; 32% of activity). The difference between explore/inves- tigate and traveling activities would be interesting to compare between wild and captive canids. The captive Coyotes that we studied were very active but did more exploring then traveling because of the relatively small area that they lived in. Conversely, Coyotes in the wild routinely have to traverse their home ranges (Camen- zind 1978; Gese et al. 1996; Way 2000) and potential- ly might do more traveling than exploring. Hunting (HU) behaviour was classified when a Coy- ote was actively trying to catch/kill another object. Many small mammals and birds entered their enclosure, giv- ing the Coyotes the opportunity to hunt (although we occasionally gave them live chicken chicks, we never scored the Coyotes when they were killing them). Ad- ditionally, when a Coyote was attempting to ambush a conspecific, we also classified this as hunting because its intentions were very similar to hunting prey: L.e., body very alert and focused, in a crouched position, and rushing at its intended target. Furthermore, differ- ent results occurred after an ambush; e.g., no social inter- actions occurred (i.e., both individuals walked away from one another), a play initiation was given once the ambusher got within close range of the Coyote, play immediately started, or fighting or agonistic behaviour ensued. Parks (1979) classified “ambushing” as a sep- arate category in her thesis but perhaps the larger size of the exhibit in which we studied our Coyotes allowed us to detect the similarity to hunting. Feeding was simply associated with ingesting some- thing. Although we documented our Coyotes feeding (FE) we attempted to avoid conducting behaviours around any type of feeding time (e.g., we did not do a behavioural bout when dog chow was just placed out or when meat, rats or mice were just given to them) to avoid biasing the data. Infantile (IN) behaviours were associated with neo- natal contact as described by Parks (1979), but we also included licking/grooming one another. Because the Coyotes were in contact with other Coyotes, we believed that this was an important division for a par- ent category even though other behavioural complex- es, such as resting, sit] and sit 2, are included in this category. These behaviours were intermittently seen THE CANADIAN FIELD-NATURALIST Vol. 120 me : 2 bt S FiGurE 1. JGW (first author) conducting a focal animal sam- pling bout on the Coyote to the right of him. nt FIGURES 2 and 3. Coyotes sleeping in rest position. FiGure 4. Coyote in Sit! Alert position. 2006 Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES FIGURE 9. Coyote in Stand Alert position. FIGURE 10. Coyote Standing and Howlin .¢ \ipee : % rs p in sit2 Alert position in JGW’s house. FiGuRE 11. Coyotes Standing and Group Howling. <—" Se Sis < ut . Rud FiGuRE 8. Coyote on left in Sit] Alert Pant position; Coyote © FIGURE 12. Coyote on the right and in the back are Traveling on right in Sit2 Alert position. and Trot Following the Coyote in the front. 284 THE CANADIAN FIELD-NATURALIST Vol. 120 Signi i pe is bs E 3 : Wrle . she US core "9 adi naar ac == FIGURE 16. Coyote in Hunting mode, chasing and outing a 4 bird. FiGuRE 15. Coyote in foreground in Hunt Ambush Crouch (Sitl) position. Coyote in background in Ambush- rush Conspecific position. This behavior can quickly turn into an Approach (Travel trot towards) when the Coyote is not in a crouched hunting mode and is moving towards another Coyote. FIGURE 18. Coyote Greeting Person (JGW). 2006 Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES 285 € FIGURE 19. Coyote in a true Greeting with JGW. FIGURE 24. Coyote trying to Play Initiate with JGW by Lying , on back and Pawing. FiGurE 25. Agonistic display. Coyote on top is in an Agonistic Dominance Display while the Coyote pinned to the ground is in Agonistic SuP (passive submission). throughout the 18 months of observation when Coy- otes would occasionally sleep on one another (espe- cially on rainy days) or one Coyote would lick anoth- er but was not greeting it. A licking Coyote appeared to be grooming the recipient, which made it hard to place into any other category (i.e., it was doing more FIGURE 22. Coyotes Play Group Wrestling. than simply greeting another Coyote). 286 Greetings (GR) were classified when non-agonistic and/or non-play interactions occurred between =2 Coy- otes. Zimen (1982) classified many of these behaviours as neutral social contacts but we separated Zimen’s descriptions of sniffing other Coyotes and left them in the explore/investigate category because the motiva- tion was not always clear (e.g., was sniffing another Coyote a social contact or just a Coyote curious to smell another one?). Simply put, when both Coyotes licked and/or had muzzle contact with each other and/ or wagged their tails in excitement to see one another, then we classified this as a greeting. We used Bekoff’s (1972a,b) definition of “play” to define our category, where play was the behaviour that was performed during social interactions in which there was a decrease in social distance between the interac- tants, and no evidence of social investigation or of ago- nistic or passive-submissive behaviours on the part of the members playing, although these actions may occur as derived acts during play (e.g., passive submission during a play bout). Bekoff (1974) also classified play as: (1) incorporating various contexts into unpredictable temporal sequences; (2) preceded by a metacommu- nicative signal (which we have a separate category for: i.e., play intitiation); (3) certain actions are repeated and performed in an exaggerated manner; and (4) the activity appears pleasurable to the participants: a play face (wide open jaws and eyes) is apparent. We also included the numerous role reversals in this category where more dominant Coyotes allowed lower ranked Coyotes to pin them to the ground; this was never documented during agonistic displays. Bekoff (1974) includes a description of this occurring amongst western Coyotes but does not delineate it as part of clas- sifying play. Additionally, we separated play initiation from play because the attempter was not always suc- cessful in initiating play during an attempt; 1.e., they appeared to be two separate categories. We observed many play initiation-like (e.g., approach/withdraw and general movements) behaviours occurring during play, but because they happened during play we kept them in this category; i.e., play was already initiated. Self play (PS) was similar to play but when a Coyote was by itself. Altogether, play-like sequences accounted for ca. 10% of all observations (Table 1). Agonistic (AG) behaviours were classified as aggres- sive acts associated with conflict where there were clear acts of dominance and submission or where growling or fighting was observed (Fox 1969; Scott and Fuller 1965; Zimen 1982). About 2% of all activities were aggressive, especially towards the end of the study when Coyotes were separated. Finally, a miscellaneous (MISC) category was one where we could not classify the behaviour pattern into any of the other parent sys- tems; these were very infrequent (Table 1). Despite the large number of behavioural patterns described herein, there are some important omissions in our ethogram. First, because the Coyotes were all siblings and were young adults (18 months) when the THE CANADIAN FIELD-NATURALIST Vol. 120 study ended, we did not notice any sexual/reproductive behaviours (although it was witnessed during their sec- ond winter: J. Way, unpublished data). Future etho- grams should add this behaviour as a separate (par- ent) category much as Scott and Fuller (1965), Zimen (1982), and Goodmann and Klinghammer (1990) did with their respective ethograms. Second, because the Coyotes were taken from the wild at 25 days of age, we missed many basic neonatal behaviours, such as crawling and nursing (Scott and Fuller 1965; Scott 1967; Parks 1979). Third, because the Coyotes were all siblings and of the same age, we could not document any adult-pup interactions such as epimeletic behay- iours, i.e., the giving of care (Scott and Fuller 1965). However, many times we did document et-epimeletic behaviours where the Coyotes were soliciting or call- ing for care where they would whine, wag their tails for a greeting (with JGW and/or another Coyote), and lick one another. Fourth, the full range of hunting be- haviours (e.g., killing/biting prey) was not documented mainly because of the rarity of observing a kill — espe- cially during the 15 sec intervals when we recorded data. Finally, there were some different behavioural actions that we could have added but simply did not because of the sheer volume of our ethogram. For ex- ample, under infantile behaviour we have sit] and sit2 although we do not note if their eyes are open or closed. Because both examples were rarely observed, this was a minor detail in the overall scheme of Table 1, but it does illustrate the arbitrarily defined nature of any ethogram (Schleidt et al. 1984), no matter how simple (i.e., the parent categories) or complex (i.e., the behav- ioural actions) it is. Also, we thought that sit] and sit2 were logical separations due to the noticeably different body positions; however, we theoretically could have had many more sitting positions based on a combina- tion of head, body, and leg positions. Although we attempted to be consistent with our behavioural classifications, there was no doubt that drift occurred in our categorizations (MacDonald et al. 2001). For example, many of the behaviours occurred well before first noted on Table 1. Although usually very similar to other categories (for example E/I wk sn and E/I st sn versus E/I step sn; Table 1) a new cat- egory randomly created during the study certainly introduces a bias to the finally tally of percentages in Table 1. However, the goal of this study was to pro- duce an end product (i.e., the ethogram) even at the expense of being consistent throughout — 1.e., there was no available model of a typical ethogram on wild canids. Also, a number of simple, but newly recorded, behaviours occurred after the study concluded. For example, E/I step sniff became much more frequent no doubt because of the new category described not because a Coyote just started doing that action pattern. Also, stand step alert and stand alert move head were described soon after this ethogram was finalized (J. Way, unpublished data). Although these were com- monly recorded after the Coyotes were separated they 2006 were not described herein. That being said, we believe that the ethogram is largely complete and that these slight nuances are offset by our rationale of produc- ing the ethogram when all five Coyotes were housed together (i.e., before they were permanently separat- ed) and the fact that the main categories would stay virtually identical even with these changes. The categories that we created are intended for ease of use in the field where a researcher can simply note the major behavioural system observed and the behay- ioural pattern associated with it. One advantage of this ethogram is that it is in discrete units (i.e., the behav- ioural patterns). However, data could always be lumped together (i.e., into general behavioural systems or =2 behavioural patterns could potentially be combined into one action) if the data/list are unmanageably large (e.g., when studying wild Coyotes). The disadvantage of merging behaviours into more broad groupings is that parent categories or lumped behavioural patterns can never be split back into more distinct behavioural patterns if not done at the outset; as Bekoff (1972b) indicated, this would cause data to be irreversibly lost. The previous studies that have described canid be- haviours were quite variable (Scott and Fuller 1965; Silver and Silver 1969; Bekoff 1972a; Parks 1979: Zimen 1982) so we basically combined data/categories from all of these studies and made modifications or additions as needed in order to try to create an accurate yet practical guide for researchers studying canids in the field. Future research should try to generate similar ethograms for other species and should continue to modify and/or expand our list if new behavioural ac- tions are found with Coyotes. Analyzing ethograms of closely related species may be one technique to effec- tively discriminate between them (e.g., Western Coy- otes, Eastern Coyotes and Wolves — Silver and Silver 1969; Bekoff 1978). Acknowledgments This study would not have been possible without the support from the staff at Zoo New England, including the keepers, management, veterinarians and their tech- nicians, and security. In-kind benefits from the Way family were invaluable. Peter Auger and the Biology Department and Urban Ecology Institute at Boston College provided support. Katie-Jo Glover and Kelly Holland assisted with data entry. The Town of Fal- mouth’s Animal Control and Department of Natural Resources were instrumental in helping JGW obtain the wild-born pups. Literature Cited Bekoff, M. 1972a. An ethological study of the development of social interactions in canids: a dyadic analysis. Ph.D. thesis, Washington University, St. Louis, Missouri. Bekoff, M. 1972b. The development of social interaction, play, and metacommunication in mammals: an ethological perspective. Quarterly Review of Biology 47: 412-434. Way, SZUMYLO, and STRAUSS: AN ETHOGRAM ON CAPTIVE EASTERN COYOTES 287 Bekoff, M. 1974. Social play and play-soliciting by infant canids. American Zoologist 14: 323-340 Bekoff, M. 1978. Behavioral development in coyotes and eastern coyotes. Pages 97-126 in Coyotes: biology, behav- ior and management. Edited by M. Bekoff, Reprint 2001, The Blackburn Press, Caldwell, New Jersey. Bekoff, M. 1989. Behavioral development of terrestrial car- nivores. Pages 267-294 in Carnivore behavior, ecology, and evolution. Edited by J. L. Gittheman, Volume 1, Comstock Publishing Associates, Cornell University Press, Ithaca, New York. Bekoff, M., and R. Jamieson. 1975. Physical development in coyotes (Canis latrans) with a comparison to other canids Journal of Mammalogy 56: 685-692. Bekoff, M., and M. C. Wells. 1981. Behavioural budgeting by wild coyotes: the influence of food resources and social organization. Animal Behaviour 29: 794-801. Camenzind, F. J. 1978. Behavioral ecology of coyotes on the national elk refuge, Jackson, Wyoming. Pages 267-294 in Coyotes: biology, behavior and management. Edited by M. Bekoff, Reprint 2001, The Blackburn Press, Caldwell, New Jersey. Ewer, R. F. 1973. The carnivores. Cornell University Press, Ithaca, New York. Fox, M. W. 1969. The anatomy of aggression and its ritual- ization in canidae: a developmental and comparative study. Behaviour 35: 242-258. Fox, M. W. 1970. A comparative study of the development of facial expressions in canids; wolf, coyote and foxes. Behaviour 36: 49-73. Frank, H. 1987. Man and wolf: advances, issues and problems in captive wolf research. Dr. W. Junk, Boston, Massachu- setts. Gese, E. M., R. L. Ruff, and R. L. Crabtree. 1996. Forag- ing ecology of coyotes (Canis latrans): the influence of extrinsic factors and a dominance hierarchy. Canadian Journal of Zoology 74: 769-783. Goodall, J. 1986. The chimpanzees of Gombe: patterns of behavior. The Belknap Press of Harvard University Press, Cambridge, Massachusetts. Goodmann, P. A., and E. Klinghammer. 1990. Wolf ethogram. Revised edition. Ethology Series Number 3. North American Wildlife Park Foundation (Wolf Park), Battle Ground, Indiana. Jennings, H. S. 1906. Behavior of the lower organisms. Columbia University Press, New York, New York. Lehner, P. N. 1978. Coyote communication. Pages 127-162 in Coyotes: biology, behavior and management. Edited by M. Bekoff. Reprint 2001, The Blackburn Press, Caldwell, New Jersey. Lorenz, K. 1971. Studies in animal and human behaviour. Volume 2. Harvard University Press, Cambridge, Massa- chusetts. Macdonald, D. W., P. D. Stewart, P. Stopka, and N. Yam- aguchi. 2000. Measuring the dynamics of mammalian societies: an ecologist’s guide to ethological methods. Pages 332-388 in Research techniques in animal ecology: con- troversies and consequences. Edited by L. Boitani and T. K. Fuller. Columbia University Press, New York, New York. Makkink, G. F. 1936. An attempt at an ethogram of the Euro- pean avocet (Recurvirostra avosetta L.) with ethological and psychological remarks. Ardea 25: 1-60. Martin, P., and P. Bateson. 1986. Measuring behaviour. Cam- bridge University Press, New York, New York. 288 Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. Reprint 1995, University of Min- nesota Press, Minneapolis, Minnesota. Parks, M. B. 1979. Physical and behavioral development of captive eastern coyote pups. M.S. thesis, University of Maine, Orono. Ryon, J. 1986. Den digging and pup care in captive coyotes (Canis latrans). Canadian Journal of Zoology 64: 1582- 1585. Schleidt, W. M., G. Yakalis, M. Donnelly, and J. McGarry. 1984. A proposal for a standard ethogram, exemplified by an ethogram of the bluebreasted quail (Coturnix chinensis). Zeitschrift fur Tierpsychologie 64: 193-220. Scott, J. P. 1967. The evolution of social behavior in dogs and wolves. American Zoologist 7: 373-381. Scott, J. P., and J. L. Fuller. 1965. Genetics and the social behavior of the dog. The University of Chicago Press, Chicago, Illinois. Silver, H., and W. T. Silver. 1969. Growth and behavior of the coyote-like canid of northern New England with obser- vations of canid hybrids. Wildlife Monographs 17: 1-41. Snow, C. J. 1967. Some observations on the behavioral and morphological development of coyote pups. American Zoologist 7: 353-355. Tinbergen, N. 1959. Comparative studies of the behavior of gulls (Laridae): a progress report. Behaviour 15: 1-70. THE CANADIAN FIELD-NATURALIST Vol. 120 Way, J. G. 2000. Ecology of Cape Cod coyotes (Canis latrans var.). M.S. thesis, University of Connecticut, Storrs. Way, J. G. 2001. The eastern coyote: documenting the habits of one of Cape Cod’s newest residents. Conservation Per- spectives, the online journal of the New England Chapter of the Society for Conservation Biology. URL: http://www. nescb.org/epublications/spring2001/coyotes.html. Wozencraft, W. C. 1989. Appendix: classification of the recent Carnivora. Pages 569-593 in Carnivore behavior, ecology, and evolution. Edited by J. L. Gittleman. Volume 1. Comstock Publishing Associates, Cornell University Press, Ithaca, New York. Zimen, E. 1982. A wolf pack sociogram. Pages 282-322 in Wolves of the world: perspectives of behavior, ecology, and conservation. Edited by F. H. Harrington and P. C. Paquet. Noyes Publications, Park Ridge, New Jersey. Received 7 March 2005 Accepted 30 January 2007 Editor’s Note A description of these Coyotes “in layman’s language” is now available. Way, J. G. 2007. Suburban howls: tracking the eastern Coy- ote in Urban Massachusetts. Dog Ear Publishing, Indiana- polis, Indiana. http://www.easterncoyoteresearch.com First Observations of an Eastern Screech-Owl, Megascops asio, Population in an Apple-Producing Region of Southern Quebec NGAIo L. RICHARDS!, PIERRE MINEAU2, DAVID M. Birp*, PIERRE WERY*, JACQUES LARIVEE®, and JASON DUFFE® ‘Anglia Ruskin University, East Road, Cambridge CB1 IPT United Kingdom; e-mail: n.richards@anglia.ac.uk Canadian Wildlife Service / National Wildlife Research Centre, Environment Canada, 2 Raven Road, Carleton University, Ottawa, Ontario KIA OH3 Canada ‘Avian Science and Conservation Centre, Department of Natural Resource Science, McGill University, Macdonald Campus, 21, 111 Lakeshore Road, Ste-Anne-de-Bellevue, Québec H9X 3V9 Canada 4Parc national du Mont-Saint-Bruno, 330 rang des 25 Est, Saint-Bruno-de-Montarville, Québec J3V 4P6 Canada Etude des populations d’ oiseaux du Québec, 194 Ouellet Street, Rimouski, Québec GSL 4R5 Canada °Cégep de Rimouski, 60 Evéché ouest, Rimouski, Québec GSL 4H6 Canada Richards, Ngaio L., Pierre Mineau, David M. Bird, Pierre Wery, Jacques Larivée, and Jason Duffe. 2006. First observations of an Eastern Screech-Owl, Megascops asio, population in an apple-producing region of southern Quebec. Canadian Field-Naturalist 120(3): 289-297. Baseline information was collected on a local Eastern Screech-Owl population found in the apple-producing region of Saint-Hilaire and Rougemont, Quebec, as part of a larger study on pesticide exposure conducted between 2000 and 2003. Screech-Owls visited or occupied 41 of 89 nest boxes installed in 12 orchards and 2 control locations. The mean height of occupied nest boxes was 3.83 m (2.00 — 5.80 m). Squirrels, Sciurus and Tamiasciurius sp., and chipmunks, Tamias striatus, Northern Flickers, Colaptes auratus, and wasps were the owls’ primary competitors for the boxes. Intact Screech-Ow] pel- lets retrieved from nest boxes (1 = 82) had a mean length and width of 3.57 and 1.44 cm, respectively, and weighed a mean of 1.77 g. Screech-Owls in the study area consumed a variety of small mammal, avian, insect and aquatic prey. Of these, Meadow Voles, Microtus pennsylvanicus, were the predominant prey item identified in pellets, and Mourning Doves, Zenai- da macroura, were the primary avian prey found in nest boxes. In Canada, the owl’s trend status remains largely unknown. The species is currently listed as “Not at Risk”, based on an assessment conducted for COSEWIC in 1986. Given that a lim- ited amount of information exists on the natural history and ecology of the species in Quebec, we also generated a map of the owl’s distribution in the province, using data from ornithological databases and rehabilitation facilities. Potential risks to the species within the province, particularly pesticide exposure and habitat loss, are briefly addressed and follow-up studies are discussed. Key Words: Eastern Screech-Owl, Megascops asio, natural history, ecology, apple orchard, Quebec, range, distribution, census, pel- let analysis, prey inventory, nest box, wing chord, population management, pesticide exposure, habitat loss, develop- ment, conservation. The Eastern Screech-Owl, Megascops asio, has been extensively studied in the United States (i.e., VanCamp and Henny 1975; Gehlbach 1994a). However, it has received considerably less attention north of the bor- der, where it occupies a much smaller area. Within Quebec, the Screech-Owl’s range is vaguely delineated and general knowledge about the species is limited. As part of a study assessing the exposure of the Screech- Owl to pesticides currently and previously used in apple orchards of southern Quebec (see Richards et al. 2004, 2005), we collected baseline information on the species. Here, we present observations on a popu- lation of Eastern Screech-Owls (Figures 1, 2, 3 that tesides in one of the primary apple-producing regions of Quebec. Methods Mapping the range of the Eastern Screech-Owl in Quebec We obtained case files from the Clinique des oiseaux de proie (COP), based in Saint-Hyacinthe, and the Cen- ter for the Rehabilitation of Injured Birds of the Mon- térégie (CRIB), based in Otterburn Park, which admin- ister care to the majority of injured or orphaned owls in the province. The locations from which adult and juvenile Screech-Owls were admitted were tabulated and mapped. These data harbour certain biases and so were used only to illustrate the overall, rather than density, distribution of the species within Quebec. For example, the number of owls admitted from an area is positively correlated with its proximity to the near- est rehabilitation facility (Richards et al., unpublished data). Visibility and predisposition to capture can arti- ficially inflate admission numbers; in this case facili- ties tend to receive an influx of orphaned Screech-Ow] babies and fledglings in the late spring and summer, largely from suburban settings, where they are highly conspicuous (Richards et al., unpublished data). Thus, each admission location was given one data point, and multiple admissions from any given area were not considered an indication of a large Screech-Ow] population or that it was especially suitable for the species. 289 290 We also obtained Screech-Owl locations from the Etudes des populations d’oiseaux du Québec (EPOQ) database, which is managed and continually updated by members of ornithological clubs and societies belong- ing to the Association québécoise des groupes d’ orni- thologues (AGQO). Finally, we contacted personnel from the Ontario Breeding Bird Atlas project (OBBA) and from the New Brunswick division of Bird Studies Canada to ensure that our map represented the species’ range at the eastern and western borders of Quebec as precisely as possible. Site selection and censusing All fieldwork was carried out in orchards of Rouge- mont and Saint-Hilaire (45:28"73°W to 4532"73°N), two adjacent apple-producing regions of southern Que- bec. The study region was first surveyed in the sum- mer of 2000 for areas containing habitat requirements for Screech-Owls; i.e., mature and mixed forest with old cavity-ridden trees and conifers for shelter, as well as running water. Potential orchards and control sites were then censused between November and March 2000 and 2002. By November, most first-year individ- uals should have dispersed from their natal habitat and established a territory. Thus, owls that respond are likely to be established, rather than transient, individ- uals. This provides a “true sense” of occupancy. We have found, based on our experience censusing Screech- Owls in southern Quebec, that this timeframe also corresponds to the period when individuals are rea- sonably responsive to broadcast calls. We walked or snowshoed between the edge of the orchard and the adjacent forest, broadcasting a com- bination of “bounce” (territorial) and “whinny* (defen- sive) calls into the forest from a small, hand-held tape player. Between 3 and 7 minutes of silence followed, and then the calls were repeated. If a response was not obtained, we then proceeded around the perimeter and repeated the process roughly 10 minutes later. When a response was obtained, the presence of an individual was visually confirmed with a headlamp or flashlight and its colour phase was noted. Two individuals ob- served in the immediate vicinity of one another (i.e., perched in the same tree) were considered a pair. We also noted responses from Barred Owls, Strix varia, or Great Horned Owls, Bubo virginianus. The pres- ence of these potential predators in an area may cause Screech-Owls to suppress vocalizations (Fuller and Mosher 1981), restrict the size of their territory, or deter them from establishing themselves altogether (Gehl- bach 1994a). The locations were then re-censused every winter to verify whether or not they were still occupied. The 12 orchard and 2 control sites where Screech-Owls were observed in the winter were censused again in June and July of 2001 and 2002. Given that Screech- Owls are not always particularly cooperative even under optimal conditions, locations were repeatedly censused (at least twice) if a response was not obtained THE CANADIAN FIELD-NATURALIST Vol. 120 during the first censusing effort. We did not census when wind speed exceeded 15 km/h, or during rain or snow, to ensure that censusing was carried out under the best possible conditions. Nest box installation and inspection Between 2000 and 2002, 89 nest boxes were in- stalled in the forests where Screech-Owls responded during our initial censusing efforts. All orchards were assigned a letter code to maintain orchard-owner con- fidentiality. Boxes were constructed of pine, Pinus sp., or Eastern Hemlock, Tsuga canadensis, with dimen- sions for American Kestrel, Falco sparverius, as per Henderson (1992). Material does not appear to influ- ence nest box selection by Screech-Owls (Gehlbach 1994b). One caveat is that boxes made of hemlock must be stored under conditions of relatively uniform heat or they will crack and warp. Pine boxes are heavier and more challenging to transport but much more resistant to temperature fluctuations, both inside during storage and in the field. Between 1.0 and 2.0 inches of dust-free wood chips were placed in the bottom of each box. The distances from the base of the box to the ground and from the box to the nearest orchard edge were recorded. Nest box covers were secured on each side with spring- loaded latches to facilitate inspections while minimiz- ing entry by mammalian predators, particularly Rac- coons, Procyon lotor. Though most locations were fairly remote, boxes installed in areas regularly ac- cessed by the public were painted with a brown, non- toxic paint, and placed a greater distance from the ground to reduce visibility and the possibility of distur- bance or vandalism (as described in Gehlbach 1994b). Boxes were initially inspected for pellets and prey remains 3 to 4 times per year: in the spring, in mid to late summer, in the late autumn and in the early winter, depending on ease of accessibility to each site espe- cially during periods of heavy snowfall. However, we soon stopped inspecting in the spring and summer be- cause of the disturbance posed to other species occupy- ing the boxes; unfortunately, we unwittingly caused nuthatch, Sitta sp., and European Starling, Sturnus vul- garis, parents to abandon their nests. From an ethical and logistical perspective, the late autumn and early winter were deemed the best times to inspect nest box- es. Thus, between 2003 and 2004, boxes were inspect- ed once or twice per year in the late autumn and/or early winter. When prey items were found, boxes were emptied and replenished with wood chips to prevent duplication of results on subsequent visits. Recovered wood chips were carefully sifted for pellets or prey items. Pellets, prey remains and prey inventory Pellets were air-dried prior to dissection for prey identification. Length, width and weight of intact pel- lets were recorded. Small mammal species were iden- tified by lower mandible (Cahn and Kemp 1930; Rac- 2006 zynski and Ruprecht 1974), using a reference collection of known skulls and mandibles. Vole species were fur- ther distinguished as Clethrionomys or Microtus sp. using Maisonneuve et al. (1997). Avian prey were iden- tified to species from feather remains with the help of two local wildlife rehabilitators, and referring to El- broch and Marks (2001). An inventory was then drawn up for each of the locations where prey items were found, Screech-Owl captures Screech-Owls were initially captured between June and August of 2001 and 2002 as part of an unsuccess- ful attempt to radio-track them and evaluate their use of orchards in relation to pesticide applications. We set up a mist net and a bal-chatri trap baited with a white mouse within orchards but near adjacent forest (as per permits no. 2001-05-07-103-16-SF and 2002-06-10- 120-16-SF). We then broadcast defensive and territori- al calls into the forest to attract owls. Capture attempts were initiated just prior to or soon after dusk. All captured adults and young were processed on- site. First, they were placed headfirst into a clean Pringles® chips canister and weighed with a 300 g spring-loaded Pesola scale. After the owl was removed from the canister, the keel region was gently palpated and a keel index was assigned on a scale of 0 to 5, with 1 being “emaciated”, 3 being of “medium” weight, and 5 being “obese”. The wing chord was also measured. Owls were aged as adults based on the presence of blood feathers and on other evidence of moulting. It was not possible to sex individuals. All owls were band- ed and released. Results Range map One hundred and sixty-nine locations were plotted: 20 from the COP (records spanning 1998 to 2001), 17 from the CRIB (records spanning 1988 to 2003), 7 from the OBBA (2001 to 2004 data) and 125 from the EPOQ database (1940 to 2004). According to the personnel at the New Brunswick division of Bird Stud- ies Canada, the species is incidental at best in the Mar- itime provinces, and they were unable to provide us with definitive sightings. Censusing Between the winters of 2000 and 2002, we censused 27 orchards and 3 potential control sites. We received a response from Screech-Owls in 12 orchards and 2 control sites, one a wooded area near a trailer camp- ground and the other in the research sector of the Mont- Saint-Hilaire Biosphere Reserve. Four pairs and 7 indi- viduals were observed during censusing and all were grey-phased. Great Horned Owls or Barred Owls were heard or seen in the vicinity of 6 of the 14 locations occupied by Screech-Owls and 6 of the 16 remaining locations. It is likely that some or all of the locations where we did not obtain a response are frequented by one or both RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC 29) ppg //~ Brunswick ~ / FIGURE 1. Distribution of the Eastern Screech-Owl (Meegas- cops asio) in Quebec. species; the habitat observed is suitable and they are all either quite close to the Mont-Saint-Hilaire Bios- phere Reserve or to Mont Rougemont, both of which have Great Horned Owl and Barred Ow! populations (C. Cormier, personal communication 2003). Nest box occupation A nest box was considered occupied if pellets, prey remains or an owl were observed in it during inspec- tions (Table 1). To determine whether height of nest box or distance from box to nearest habitat/forest edge influenced its selection or use, we first considered the height and distance of those boxes occupied ar least once by a Screech-Owl. Then, we further refined our analysis to nest boxes for which we had evidence of multiple uses, defined as three visits or more (Table 2): either evidence of occupancy on at least three sep- arate nest box inspections or discovery of at least three pellets or prey remains, given that Screech-Owls gen- erally regurgitate one pellet per resting period. We measured the distance from the tree on which nest boxes were installed to the nearest “true’ edge”, an important distinction as a network of paths criss- crossed most of the forests in which boxes were in- stalled. True edge was defined as an expanse of open area immediately adjacent to, rather than contained within, the forest itself, such as a field or an orchard. Box distance ranged from 0 to 68.0 m from edge. TABLE 1. General nest box occupancy of Eastern Screech- Owls, by location, in Saint-Hilaire and Rougemont, Quebec areas (2000 — 2004). Location Number of Percent boxes installed occupation Campground 9 89 Biosphere Reserve 6 100 A/AI 7 14 B 10 30 K 8 25 Cc 5) 20 D/F 10 100 G 7 14 I 11 0 M 7 29 N/N1 9 78 Total 89 A variety of other species occupied the nest boxes (see Table 3). To determine the overall frequency of nest box use, we evaluated the total number of times boxes were occupied by Screech-Owls at each site over the course of the study relative to other species. Table 4 shows occupation by the most commonly observed species in relation to that by Screech-Owls. Pellets and prey remains Nest boxes were installed at the 12 orchards and 2 control locations where Screech-Owls responded to censusing. However, in some of these locations, there was little or no evidence (1.e., pellets or prey remains) that Screech-Owls ever occupied these boxes. Only one pellet was found at Orchards A/A1, C and M, so these were excluded from the prey inventory analysis (Table 5). Intact pellets were retrieved from 8 of the orchards, including C and M, which were included in the analysis of pellet dimensions. TABLE 3. Nest box occupancy by other species. Mammal occupant Eastern chipmunk (Tamius striatus) Southern Flying squirrel (Glaucomys volans) Grey squirrel (Sciurus carolinensis) THE CANADIAN FIELD-NATURALIST Avian occupants Great Crested Flycatcher (Myiarchus crinitus) Downy Woodpecker (Picoides pubescens) European Starling (Sturnus vulgaris) Vol. 120 TABLE 2. Mean height from ground of nest boxes (n = 41) occupied by Eastern Screech-Owls in Saint-Hilaire and Rougemont areas, Quebec (2000 — 2004). Height from ground (m) Range Unoccupied boxes (n = 48) 4.22 2.40 — 6.82 Occupied boxes (n = 41) 3.83 2.00 — 5.80 “High-use”’ nest boxes (n = 33) 3.67 2.00 — 4.72 Screech-Owl captures Despite repeated censusing, a response was never obtained in Orchards G or M. These locations were therefore excluded from capture attempts. We did ob- tain a response in Orchard C, but the ow! never deigned to approach our traps. Nine individuals were captured in the summers of 2001 and 2002. On occasion, Screech-Owls began to respond to broadcast calls while some daylight re- mained. However, owls were always captured under cover of darkness, between 21:15 to 03:10. Though we were not always able to capture all the individuals observed, we did record their presence (see Table 6). Of the nine captured individuals and six observed, but not captured, only one was red-phased; the rest were grey. Discussion Eastern Screech-Owl range in Quebec The Eastern Screech-Owl’s northernmost North American range coincides with the southern tip of Quebec (Gauthier and Aubry 1996). To the west of Quebec, the species is commonly reported along the southern edge of Ontario in the Carolinian zone along Lake Erie and Ontario. More local and uncommon populations extend up another 100 to 200 kilometres north. In eastern Ontario, there appears to have been Other Hymenoptera: ants, bees*, wasps Eupplexoptera: Earwigs Unknown snake** Jumping Mouse Hairy Woodpecker Arachnid: Zapodidae (Picoides villosus) Spiders Red squirrel Northern Flicker Coleoptera: (Tamiasciurus hudsonicus) (Colaptes auratus) Unknown beetles White-breasted Nuthatch (Sitta carolinensis) *could also have been cached prey item **snakeskin found, could also have been part of nesting material 2006 an overall expansion of the range in the last 20 years (M. Gahbauer, personal communication, 2004). To the east of Quebec, the range may extend into the extreme southwestern part of New Brunswick (Gauthier and Aubry 1996); however, there are no confirmed breed- ing records for the species there. If Screech-Owls do in fact breed in New Brunswick, it is likely to be in very low numbers (B. Whittam, personal communication, 2004). Screech-Owls have also been observed in Nova Scotia and in Prince Edward Island, but their presence there is thought to be incidental (ibid). Elsewhere in Canada, the range is defined as extending from south- western Saskatchewan to southern Manitoba (Panak 1988*). All of the above suggests that an important proportion of the species’ range in Canada occurs in Quebec and in Ontario. Censusing For our purpose, which was to confirm the presence of resident Screech-Owls at a given location, we found the optimum census period to be between November and February. There are several logistical advantages to censusing towards the beginning of this period. During the early winter months, pairs jointly defend their territory and are very responsive to broadcast calls. As a result, there is a greater likelihood of obtaining a response from both the male and the female, which eliminates the need for speculation as to whether or not a location is occupied by a pair. Later on, heavy snowfall can lengthen the time required to census each location considerably, and render some virtually inac- cessible. Censusing can have adverse effects. In late Febru- ary and early March, females select a suitable nesting location, then remain in or near their cavity. Broadcast calls may elicit a response from territorial males, but RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC Ig? FIGURE 2: A grey-phase Screech-Owl discovered during a mid-winter census at the Réserve écologique Marcel- Raymond, Henryville, Quebec, near the Richelieu River. Photographed by Léopold Gaudreau, Assistant Deputy Minister, Sustainable Development Branch, Ministére du Développement durable, de |’Environ- nement et des Parcs. 9 December 2006. TABLE 4. Frequency of nest box occupation in orchards and control sites of Saint-Hilaire and Rougemont areas, Quebec (2000 — 2004) Location Number of CH/ boxes inspected** ESOW SQ NOFL WASP OTHER NO Biosphere Reserve 38 26 2 l 0 0 9 Campground 116 48 26 3 5 5 29 A/A1* 60 8 4 5 | 7 35 B/K* 173 16 39 9 i) 9 93 (c 47 4 10 6 l | 25 D/F* 103 20 33 2 3 4 4] G 51 5 3 2 l 5 25 I 67 2 29 0 l 9 26 M 39 4 22 0 2 0 1] N/N1* 59 13 24 l 3 7 1] Total 753 146 202 29 24 47 305 *Orchards are side by side or adjacent **Indicates number of boxes inspected at Iccation over duration of the study ESOW: number of boxes occupied by Screech-Owl CH/SQ: number of boxes occupied by chipmunk or squirrel NOFL: number of boxes occupied by a Northern Flicker OTHER: number of boxes occupied by other inhabitants (see Appendix 1) NO: number of unoccupied boxes THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 5. Eastern Screech-Ow! prey inventory based on pellets and prey remains found in nest boxes at 6 orchards and 2 control sites in Saint-Hilaire and Rougemont areas, Quebec (2001 — 2004) Scientific name Blarina brevicauda Microtus pennsylvanicus Peromyscus sp. Sorex sp.° Zapodidae® Aegolius acadicus Bombycilla cedrorum Cardinalis cardinalis Carduellis tristis Colaptes auratus Columba livia Cyanocitta cristata Junco hyemalis Molothrus ater Poecile atricapillus Picoides villosus Quiscalus quiscula Sayornis phoebe Sturnus vulgaris Toxostoma rufum Turdus migratorius Zenaida macroura Sittae Turdidae Tyrannidae Unidentified bird* Miscellaneous artifacts and prey Apis mellifera Rana clamitans Unidentified membrane! Corn, seeds Eggshell fragments * data spans 2001-2003 only > White-footed or Deer Mouse ° Masked or Smoky Shrew Common name Short-tailed Shrew Meadow Vole Northern Saw-whet Owl Cedar Waxwing Northern Cardinal American Goldfinch Northern Flicker Rock Dove Blue Jay Dark-eyed Junco Brown-headed Cowbird Black-capped Chickadee Hairy Woodpecker Common Grackle Eastern Phoebe European Starling Brown Thrasher American Robin Mourning Dove unidentified nuthatch unidentified thrush unidentified flycatcher Honey Bee Green Frog 4 Meadow or Woodland Jumping Mouse © down remains unidentifiable ‘ may be the remains of a frog D\F* SN = tS B\K N\NI fs 3 5 31 1 3 1 1 1 1 1 5) D 1 Biosphere Reserve Campground 3 5 53 19 5 1 13 1 1 2 1 1 3 1 1 1 1 1 1 1 1 8 1 1 12 2 3 1 2 1 1 2 TABLE 6. Eastern Screech-Owls (n = 9) captured and banded in one control and six orchard locations of Saint-Hilaire and Rougemont areas Quebec (2001-2002), by month of capture. See methods for keel index categories. Location Orchard I Campground Control Orchard K> Orchard A/A1°4 Orchard D/F** Capture date 29 August 2002 22 July 2001 1 adult* 13 July 2002 1 adult 3 juvenile 31 July 2002 1 adult 9 August 2002 1 adult | juvenile 1 juvenile Captured individual Keel index Wing chord Weight (g) (0-5) (mm) 2.5 to 3.0 170 78.0 3 177 78.0, 94.0, 96.0 52h 182,184,194 160 3 176 159 + 176 155 3 185 171 4 178 * red-phased individual > | adult and 1 juvenile observed but not captured at B in June 2002 “ orchards are adjacent to one another and considered to comprise one territory * 1 other juvenile observed but not captured * | adult and 2 juveniles observed in June 2001 but not captured 2006 RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC 295 FiGureE 3. An Eastern Screech-Owl peers sleepily from a cavity at Pare Summit on the Island of Montreal. As development within the province increases, so too does the importance of the parks and protected areas within the species range. Photo by Jean-Sébastien Guénette, 17 April 2005. given the necessity of devoting their energy to provi- sioning their mate and themselves, and to defending their territory against real rather than perceived intrud- ers, censusing at this time can impose an unnecessary and potentially harmful stress. Predators of the Screech- Owl can also be attracted to the broadcast calls, and to respondents, especially if illuminated by a flash- light or headlamp. We are aware of one instance, for example, where a Long-eared Owl, Asio otus, was drawn to a Screech-Owl during censusing and was illu- minated in the process of closing in on it. Since this study, a “minimal-disturbance” censusing protocol has been developed for the Screech-Owl and for other owl Species (see Balej 2006). Nest Box inspections Nest boxes (” = 89) were installed in the vicinity of where Screech-Owls responded during censusing. Ordi- narily, an owl’s presence in an area would go largely undetected because they are very inconspicuous and occupy natural cavities. As the study progressed, our primary concern became to place the nest boxes at a height that would reduce the possibility of drawing untoward human attention to them. Overall nest box occupation at each location varied from 0% to 100%. Forty-one boxes were occupied at least once by an owl, and these boxes were a mean of 3.8 m (2.0 to 5.8 m) from the ground (Table 2). Gehlbach (1994b) found owls nesting, on average, 3.7 m from the ground in natural cavities (m = 23). A number of species may compete with Screech- Owls for nest boxes. The four most frequent occu- pants were squirrels, Sciurus and Tamiasciurius sp., and chipmunks, Jamias striatus, Northern Flickers, Colaptes auratus, and Hymenoptera, especially wasps. During pre-snowfall inspections in October and No- vember, we frequently removed nesting material left by other species, and during inspections the following spring found Screech-Owl pellets and prey remains. Occasionally, Screech-Owls deposed chipmunks and squirrels, judging from pellets or prey remnants deposit- ed over nesting material. Northern Flickers have been reported to destroy Screech-Owl eggs (Penak 1986*). However, the Screech-Owls in our region infrequently (22 of the 146 recorded times) occupied nest boxes in the summer and early autumn months, when Flickers occupied them. Late autumn inspections enabled us to remove nesting material left by other species (or by Screech-Owls during the winter). We also could replen- ish the boxes with wood chips during these visits with- out having to disturb any of the species that had pre- viously occupied the boxes and make any necessary repairs. 296 Pellets The intact pellets retrieved from nest boxes (n = 82) ranged widely in length from 1.0 to 8.5 cm and from 0.8 to 3.2 cm in width, with a mean weight of 1.77 g. Elbroch and Marks (2001) recovered Screech-Ow1 pel- lets (n = 18) from nearby Vermont and New Hampshire which ranged from 1.6 to 3.8 cm length and 0.9 to 1.4 cm width. Fifty-six of our pellets exceeded the up- per length and width reported by Elbroch and Marks. Most of the intact pellets we measured were retrieved from nest boxes, where they were protected from the elements and from breaking. This, and our larger sam- ple size, may explain the discrepancy in our upper reported length. Prey Consumption Meadow voles were the predominant intact prey item found in our nest boxes. Screech-Owls are also highly insectivorous (Ritchison and Cavanagh 1992), and although we repeatedly found evidence of insect consumption, we were unable to fully assess or quan- tify it. Only insects with hard parts, such as beetles, Coleoptera, can be distinguished in pellets. For the most part, we found only a dust of legs and carapaces intermingled with the wood chips and with other arti- facts. Mourning Doves were the avian prey most fre- quently found in nest boxes, but few avian prey remains were found. Screech-Owls may simply prefer to pluck feathers on a perch or in a deeper tree cavity rather than in a nest box which feathers could potentially fill quite quickly. Hence, our prey inventory underestimates avian and insect consumption. As well, several cached frogs and shrews, and a decapitated vole, were also found in nest boxes at the Campground control site. On a number of occasions, remnants of crayfish and fish scales were recovered from this site, as well as pieces of a greenish membrane, which may have been a remnant from the frogs. The greatest diversity in prey items seems to have been consumed at the Biosphere Reserve. Conclusion The Screech-Owl is found in suburban, rural and agricultural habitats. It is considered to be one of the most ubiquitous owls in North America (Gehlbach 1995). There is, however, very little information avail- able about the species’ natural history and ecology, or about its density and overall distribution in Canada, where its status was last evaluated for the Committee on the Status of Endangered Wildlife in Canada over 20 years ago (by Penak 1986*). Paradoxically, the species is currently designated as “Not at Risk”, even though there are no current data on its population trend status (http://www.cws-scf.ec.gc.ca/mgbe/trends/ index .cfm?lang=e& go=info.bird&speciesid=3730). An important part of the species’ range in Canada occurs in southern Quebec, where it coincides with an agricultural belt and with the province’s major apple- producing region. Screech-Owls are known to favour THE CANADIAN FIELD-NATURALIST Vol. 120 orchard habitats (Gauthier and Aubry 1996) where pesticides are intensively applied. Our study evaluating the exposure of Screech-Owls to pesticides currently and previously used in one of the largest apple-pro- ducing regions of southern Quebec provided an oppor- tunity to collect information on the species’ regional diet, record physiological parameters, evaluate occupa- tion response to nest boxes and gain a sense of the den- sity and distribution of the local population. As far as we know this is the first time that these types of data have been collected on the Screech-Owls of the area. More baseline information of this type must be gath- ered for future monitoring and population management efforts throughout the owl’s Canadian range. Changes in development and land use patterns must also be examined. While we found that exposure to DDE and organophosphorus pesticides warranted further inves- tigation (see Richards et al. 2004, 2005), we also ob- served a marked increase in clearing and development of forested lands and orchards over the course of the study. We believe that loss of habitat and overall de- crease in habitat quality currently pose a far greater threat to the local population than exposure to pesti- cides. Our colleagues at the Quebec environment ministry (Ministere du Développement durable, de |’Environ- nement et des Parcs) have been conducting autumn and winter censuses in ecological reserves within the known range of the species since 2005. This is part of an ongoing study initiated to evaluate the relative risks that pesticide exposure and habitat loss/deterioration pose to Screech-Owls in southern Quebec (if interest- ed in collaborating on this study, please contact N. Richards). Even if the North American Screech-Owl popula- tion may not be at risk as a whole, we are concerned that habitat loss and modification may be adversely affecting regional populations. At the very least, a re- examination of the Screech-Owl’s distribution and pop- ulation status throughout its Canadian range, espe- cially in Quebec and Ontario, is long overdue. If this adaptable and opportunistic species is being adverse- ly affected, how does this bode for the more vulnera- ble species which occupy the same habitats? Acknowledgments We are grateful to the orchard owners of the study area, without whose collaboration this study could not have taken place. We thank Terry Ryan and Jean Gou- geon for building industrial quantities of nest boxes. Doctors Guy Fitzgerald and Claude Lacasse of the Clinique des oiseaux de proie and Annemarie Roth of the Centre for the Rehabilitation of Injured Birds of the Montérégie provided us with Screech-Owl ad- mission records. Stéphane Deshaies and Annemarie Roth helped identify avian prey remains. Ambroise Lycke, Christine Bourque, Patrick MacFarlane, Anke Roth, Gavin McMorrow, Jessus Karst, Hugo Gee, Mar- 2006 cel Gahbauer, Tina Newbury, Amy Stevenson, Mar- isha Futer, Jean-Sébastien Guénette, Joanna Coleman and Anneli Jokela all provided invaluable assistance and/or training in the field. We thank Nicole Kopysh at the Ontario Bird Banding Atlas project and Becky Whittam at the New Brunswick division of Bird Stud- ies Canada for information related to the Screech- Owl’s distribution in these provinces. Caroline Cormi- er of Nature-Action, Québec Inc. and Gina Rossini provided information on the whereabouts of birds of prey in Rougemont and Saint-Hilaire. We are pleased to acknowledge the collaboration and hard work of Rodolph Balej and Marcelle Ruest at the Ministére de Development durable de l’Environnement et des parcs. Funding for this study was provided to N. L. Richards by the Fonds Québécois de la recherche sur la nature et les technologies and by the Canadian Wildlife Ser- vice, Environment Canada. Additional funding was provided through a Natural Sciences and Engineering Research Council of Canada (NSERC) grant to P. Mineau. Initial stages of the project were funded by Environment Canada’s Science Horizons for Youth programme, in collaboration with World Wildlife Fund Canada. In particular, we gratefully acknowledge the continued funding and support provided by Bird Pro- tection Quebec (BPQ) (formerly the Province of Que- bec Society for the Protection of Birds — PQSPB). Documents Cited (marked * in text) Penak, B. L. 1986. COSEWIC Status report on the Eastern Screech-Owl Otus asio in Canada. Ottawa, Ontario. Literature Cited Balej, R. 2006. Census of nocturnal raptors in the ecologi- cal reserves of the province of Quebec. Winter 2006-2007. Service des aires protégées, Direction du patrimoine éco- logique et des parcs, Ministére du Développement durable, de l'Environnement et des Parcs. 15 pages. Cahn, A. R., and J. T. Kemp. 1930. On the food of certain owls in east-central Illinois. Auk 47: 323-328. Elbroch, M., and E. Marks. 2001. Bird Tracks and Sign. Stackpole Books, Mechanicsburg, Pennsylvania. Fuller, M. R., and J. A. Mosher. 1981. Methods of detecting and counting raptors: a review. Pages 235-246 in Estimat- ing the numbers of terrestrial birds. Edited by C. J. Ralph and J. M. Scott. Studies in Avian Biology 6. RICHARDS ET AL.: EASTERN SCREECH-OWL IN SOUTHERN QUEBEC 297 Gauthier, J., and Y. Aubry. 1996. Atlas of the breeding birds of southern Quebec, L’ Association québécoise des groupes ornithologues, Province of Quebec Society for the Pro tection of Birds, the Canadian Wildlife Service, Quebec region, Montreal. Gehlbach, F. R. 1994a. The Eastern Screech-Ow!: life history, ecology, and behaviour in the suburbs and countryside Texas A&M University Press, College Station, Texas Gehlbach, F. R. 1994b. Nest-box versus natural-cavity nests of the Eastern Screech-Owl: an exploratory study. Journal! of Raptor Research 28: 154-157. Gehlbach, F. R. 1995. Eastern Screech-Owl, Orus asio. Pages 1-24 in The birds of North America (165). Edited by A Poole and F. Gill. The Academy of Natural Sciences, Philadelphia, and The American Ornithologists’ Union, Washington, D.C. Henderson, C. 1992. Woodworking for wildlife: Homes for birds and mammals. Minnesota Department of Natural Resources. Maisonneuve, C., R. MecNicoll, S. St-Onge, and A. Desrosiers. 1997. Clé d’identification des micromam- miféres du Québec. Ministére de |’Environnement et de la Faune. Québec, Canada. Raczynski, J., and A. L. Ruprecht. 1974. The effect of diges- tion on the osteological composition of ow! pellets. Acta Ornithologica 84: 25-36. Richards, N. L., P. Mineau, and D. M. Bird. 2004. Exposure of the Eastern Screech-Owl to organophosphorus insecti- cides and anticoagulant rodenticides in apple orchards of southern Québec, Canada. Pages 389-408 in Raptors World- wide. Edited by R. C. Chancellor and B.-U. Meyburg. Pro- ceedings of the Sixth World Conference on Birds of Prey and Owls. May 18-23, Budapest, Hungary. Richards, N. L., P. Mineau, and D. M. Bird. 2005. A risk assessment approach to DDE exposure based on the case of the Eastern Screech-Owl (Megascops asio) in apple orchards of southern Québec, Canada. Archives of Envi- ronmental Contamination and Toxicology 49: 403-409. Ritchison, G., and P. Cavanagh. 1992. Prey use by Eastern Screech-Owls: seasonal variation in central Kentucky and a review of previous studies. Journal of Raptor Research 26: 66-73. VanCamp, L. F., and C. J. Henny. 1975. The Eastern Screech-Owl: its life history and population ecology in northern Ohio. North American Fauna 71: 1-65. Received 26 July 2005 Accepted 4 June 2007 Effects of Timber Harvesting and Plantation Development on Cavity-nesting Birds in New Brunswick STEPHEN J. WOODLEY!, GREG JOHNSON”, BILL FREEDMAN?, and Davip A. KirK* ‘Ecological Integrity Branch, Parks Canada, 25 Eddy Street, Hull, Québec K1A 0M5 Canada "Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada; Present address: Jacques-Whitford Environment Ltd., Fredericton, New Brunswick E3B 5C2 Canada >Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada *Aquila Applied Ecologists, C.P. 87, Carlsbad Springs, Ontario KOA 1KO Canada Woodley, Stephen J., Greg Johnson, Bill Freedman, and David A. Kirk. 2006 Effects of timber harvesting and plantation development on cavity-nesting birds in New Brunswick. Canadian Field-Naturalist 120(3): 298-306. We studied the abundance of cavity-nesting birds in forestry-related habitats in a region of Acadian forest in New Brunswick. We examined five reference stands of natural forest, a chronosequence of conifer plantations up to 19 years old (the oldest in the study area), two selectively harvested stands, and a 30-year-old naturally regenerated clear-cut. The species richness and abundance of cavity-nesting birds were higher in reference forest (average 10.0 species per stand; 5.3 territories per 10 ha) than in plantations (2.3/stand; 1.0/10 ha), selectively harvested stands (7.0/stand; 3.8/10 ha), or the naturally regenerated clear-cut (5.0/stand; 2.5/10 ha). A cluster analysis segregated the “community” of cavity-nesting birds of natural forest from those of other treatments. Of the various harvested stands and plantations, five with a relatively large number of residual snags clustered similarly in the cluster analysis, while those with no or very few snags also clustered together. We used arrays of nest boxes (12 per stand) to examine whether the availability of cavities was limiting the use of habitats otherwise suitable for foraging by cavity-dependent species. Nest-box use for nesting and roosting was much higher in the seven plan- tations examined (average 4.0/10 ha for nesting and 2.9/10 ha for roosting) than in three reference stands (each 0.3/10 ha), suggesting that the plantations were deficient in this critical-habitat element. Our results suggest that certain mitigations, such as leaving residual snags and living cavity-trees, would help maintain populations of some cavity-dependent birds in clear-cuts and plantations. However, some cavity-dependent species might not be accommodated by these mitigations and are potentially at risk in intensively managed areas, unless landscape-scale management plans ensure the survival of suffi- cient areas of older mixed-wood forest. Key Words: Acadian forest, cavity-nesting birds, conifer plantations, critical habitat, forestry, management recommendations, nest boxes, New Brunswick. Freedman et al. 1996; Hobson and Schieck 1999; Schieck and Hobson 2000). Intensive forestry prac- tices generally decrease the richness and abundance of cavity-dependent birds by causing the following habitat changes to occur (Niemi et al. 1998; Hobson and Bayne 2000; Flemming and Freedman 1998; Imbeau et al. 2001; Kirk and Machtans 2004; McRae et al. 2001): e the conversion of natural forest having complex biological and physical structure (i.e., multi- species stand composition and intricate spectra of age and size) into silvicultural plantations domi- nated by a monospecific cohort of similar-sized and -aged trees e the reduction in quantity of cavity trees, dead snags, and coarse-woody debris by intensive man- The effects of forestry on biodiversity range over a wide continuum, depending largely on the intensity of harvesting and subsequent management (Hunter 1990; Freedman et al. 1994; Angelstam and Milkusinski 1994; Freedman 1995; Hagan et al. 1997; Niemi et al. 1998; Imbeau et al. 2001; McRae et al. 2001; Kimmins 2003). For instance, clear-cutting followed by intensive management to develop an even-aged monocultural plantation has relatively extreme effects, while the effects associated with selective-harvesting and natural regeneration are much smaller. In boreal Europe, for example, natural pine (Pinus sylvestris) forest has been converted extensively into commercially preferred, short-rotation conifer plantations (Esseen et al. 1997), resulting in large declines of birds dependent on older natural forest (Virkkala 1991). Similar changes are occurring in temperate- and boreal-forest regions of Canada, where older mixed-species forest is being con- verted extensively into conifer plantations (Niemi et al. 1998; Imbeau et al. 2001; McRae et al. 2001). Cavity-dependent birds in temperate and boreal forest are generally most abundant and species-rich in older, uneven-aged, mixed-species stands (Hunter 1990; Schreiber and Decalesta 1992; Newton 1994; agement practices (in Canada and elsewhere snag felling may even be required under health and safety regulations; e.g., Naylor et al. 1999) e the truncation of stand age-at-maturity by short- rotation management (typically 40-60 years), which precludes the regeneration of large cavi- ty-trees and coarse-woody debris 298 2006 Moreover, at the level of landscape, forestry typi- cally decreases the average age and complexity of habitat “patches,” while isolating remnants of older natural forest (Hunter 1990; Freedman et al. 1994; Hagan et al. 1997; McRae et al. 2001). Landscape- scale forest management must accommodate species with extensive and specialized habitat needs, includ- ing some cavity-nesting birds dependent on older forest (Renken and Wiggers 1989, 1993; Bull and Holthausen 1993; Freedman et al. 1996; Bonar 2000). Because of the specialized need of cavity-users for critical habitat, they are often considered indicators of the ecological sustainability of forest management (Angelstam and Mikusinski 1997). It is particularly important to maintain the habitat used by “keystone” primary excavators, such as older heart-rotten trees and snags, because a diverse group of secondary users depends on the cavities they create (Freedman et al. 1996; Martin and Eadie 1999). The Pileated Woodpeck- er (Dryocopus pileatus) is one example of a keystone excavator whose abandoned cavities are used by other species for nesting or roosting (Bull and Jackson 1995; Bonar 2000). The effects of forestry on cavity-users are relatively wel known for forest types of the eastern and north- western United States (Raphael and White 1984; DeGraaf and Shigo 1985; Zarnowitz and Manuwal 1985; Renken and Wiggers 1989; Connor et al. 1994). However, much less information is available on which to base management decisions in the north-temperate and boreal regions of eastern Canada. This deficiency is important because habitat needs of cavity-nesters vary geographically and are likely to be different in boreal and Acadian forest than in other regions and biomes (Parker et al. 1999). In recognition of this dearth of information, studies were initiated in the mid-1990s of cavity-nesting birds in eastern Canada, with the in- tent of informing management guidelines to conserve their populations (Doyon et al. 1999; Naylor et al. 1999; Bonar 2000; Flemming et al. 2000). This advice is greatly needed, because of the increasing areas of natural forest that are being intensively managed for timber production in eastern Canada. Within this context, our study examines effects of forestry on cavity-dependent birds in a region of Aca- dian forest in southern New Brunswick. We assessed the degree to which cavity-nesting birds use conifer plantations and less-intensively managed stands, com- pared with natural forest. This was done by conducting breeding surveys of these habitats and by comparing occupancy of nest boxes as an indicator of limitation by natural cavities. Study Area The study area is in southeastern New Brunswick, Canada. It is located in the Atlantic Maritime Ecozone (Ecological Stratification Working Group 1995), with- in the Fundy Plateau Ecodistrict of the Acadian For- WOODLEY, JOHNSON, FREEDMAN, KIRK: EFFECTS ON CAVITY-NESTING BIRDS 299 est Region (Loucks 1962; Rowe 1972). The climate is humid temperate and the natural forest is dominated by mixed-species stands of Red Spruce (Picea rubens), White Spruce (P. glauca), Balsam Fir (Abies bal- samea), Sugar Maple (Acer saccharum), Red Maple (A. rubrum), Yellow Birch (Betula alleghaniensis), White Birch (B. papyrifera), and Mountain Birch (B. cordifolia). Extensive natural disturbances, including irruptions of native Spruce Budworm (Choristoneura fumiferana), have affected the natural forest, which tends to have a mixed-species canopy with gap-phase regenerating patches. Superimposed on the natural dis- turbance regime is a complex of anthropogenic influ- ences. This began with the selective harvesting of large trees for lumber and ship-building, followed by exten- sive deforestation for agricultural development, then abandonment of most poorer-quality farmland begin- ning in the 1920s, and more recently the widespread establishment of conifer plantations. Our study area is within the Greater Fundy Ecosys- tem (GFE), a region consisting of Fundy National Park and its surrounding area. The GFE was defined to study ecological consequences of the insularization of Fundy National Park, a 204 km? protected area embed- ded within a landscape whose matrix is being trans- formed by the conversion of natural forest into conifer plantations (Woodley et al. 1993, 1998; Freedman et al. 1994). The GFE itself is within the Fundy Model Forest (FMF), one of ten “model forests” established to demonstrate “sustainable forestry” in Canada. One declared indicator of sustainability in the FMF is that, at the landscape level, native biodiversity must not be compromised by forest management (Parker et al. 1999). Methods Stand selection We selected 18 stands for study (see Table 1), includ- ing representative natural (or “reference”) forest and habitats resulting from various intensities of manage- ment, including: ¢ five reference stands of unmanaged mature for- est, including three mixed woods (dominated by Red Spruce (Picea rubens), Balsam Fir (Abies balsamea), and White Birch (B. papyrifera), and Mountain Birch (B. cordifolia) and two soft- wood dominated by Red Spruce: ¢ ten stands that had been clear-cut and then in- tensively managed to develop conifer plantations ranging from 5 to 19 years in age (the oldest avail- able in the study area); eight were planted with Black Spruce (Picea mariana), one with Jack Pine (Pinus banksiana), and one with Norway Spruce (Picea abies): * two selectively harvested stands (1-year-old and 12-years-old); * one 30-year-old naturally regenerated clear-cut. 300 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. General habitat characteristics of stands surveyed for cavity-nesting birds. Survey Area Stand Site Year (ha) description Reference mixedwood forest Ra 1994 10 mature/mixedwood forest; Red Spruce, Balsam Fir, White Birch Rb 1995 20 mature/mixedwood forest; Red Spruce, Balsam Fir, White Birch Re 1994 10 mature/mixedwood forest; Red Spruce, Balsam Fir, White Birch Rd 1994 10 mature/Red Spruce-dominated forest Re 1995 20 mature/Red Spruce-dominated forest Plantations P5a 1994 10 5-year-old Black Spruce plantation P5b 1994 10 5-year-old Norway Spruce plantation 1Py/ 1995 20 7-year-old Black Spruce plantation with residual snags P8 1994 10 8-year-old Black Spruce plantation P9a 1994 10 9-year-old Black Spruce plantation P9b 1994 10 9-year-old Jack Pine plantation IIS) 1995 8 15-year-old Black Spruce plantation with residual snags P17 1994 10 17-year-old Black Spruce plantation P18 1995 10 18-year-old Black Spruce plantation P19 1994 10 19-year-old Black Spruce plantation Selectively harvested stands $12 1995 20 12-year-old selection harvested stand S1 1995 20 1-year-old selection harvested stand Naturally regenerated clear-cut NR30 1995 20 30-year-old naturally regenerated clear-cut with residual snags Note that selected study stands were not always paired because of the lack of availability of suitable stands that were given the same management. The study plots affected by forestry were essentially opera- tional stands embedded in a dissimilar habitat matrix generally of mixed-wood forests, and they were sur- veyed in their entirety. The reference plots were of a size and shape comparable to forestry-affected plots. The study stands were no more than 20 km from each other. Fieldwork was done during 1994 and 1995. Habitat description Trees (>5 cm diameter at breast height (DBH) and >1.5 m tall) and snags (standing dead trees) were sam- pled in twelve quadrats (20 m x 20 m) arranged as a grid across the study plot. However, in plantations 15 years and older, smaller quadrats (10 m x 10 m) were used because of the uniform structure of tree- sized vegetation. Shrubs (<5 cm DBH) were measured for diameter at 25 cm in two 5 m x 5 m sub-quadrats in opposite corners of each tree-quadrat. Length and diameter of coarse-woody debris (CWD; >5 cm diam- eter) were also measured in tree-quadrats. The field data were used to calculate the average DBH and basal area by species, including snags, and the volume of CWD. The cover of species of ground vegetation was deter- mined in 30 randomly located 1 m? quadrats per site. The cover of overhead canopy was estimated as the percentage area obscured by foliage when sighting upward through a 4.2-cm diameter cylinder. Here we present summary information; detailed data on woody habitat are in Flemming and Freedman (1998) and on ground vegetation are in Veinotte et al. (2004). Abundance of birds Cavity-nesting birds were surveyed by modified spot-mapping to identify territories (Bibby et al. 1999). Habitat use was designated as “nesting territory” if we found evidence of a nest, and as “present” if a nest site could not be located. The surveys were conducted in plots with areas of 8 to 20 ha (Table 1). Ten stands were surveyed in 1994 and eight in 1995, and each was examined seven times from early June through to early July. The locations of calling or singing male birds were mapped in the study plots and also in adjacent habitat, and territorial boundaries were estimated using standard spot-mapping procedures (Bibby et al. 1999). Where possible, we recorded the sex, age (adult/juve- nile), and relevant activity (nesting, feeding young, foraging, drumming, singing, calling) of individuals. This information contributed to the assignment of breeding territories. Nest-box survey To indicate whether cavities were a limiting factor for dependent species, we installed arrays of 12 nest boxes in each of seven plantations and three reference stands. The nest boxes were made of sections of White Cedar (Thuja occidentalis) with natural heart rot and were divided into three sizes: small (3.5-cm diameter entrance hole; ca. 6.5-cm internal width by 16 cm height), medium (6-cm hole, 11 x 34 cm), and large (9-cm hole; 15 x 48 cm). Because the site location and orientation of the entrance hole can affect occu- pancy (Rendell and Robertson 1994), these factors were standardized. Each nest box was mounted on an aspen pole supported by three steel cables fixed to 2006 WOODLEY, JOHNSON, FREEDMAN, KIRK: EFFECTS rebar pegs, and was placed at a predetermined height (small cavities at 2.5 m, medium at 3 m, and large at 5 m) with the entrance hole facing south. A total of 12 nest boxes was placed in each stand; two of each of size were positioned at 30 m and again at 100 m from a stand edge. The nest boxes were surveyed seven times during each of the 1994 and 1995 breeding sea- sons, and once late in the summer. We recorded the numbers of eggs, hatchlings, and fledglings of any species occupying the boxes, as well as roosting birds and non-avian species. Data analysis Because the study plots were unique habitats in terms of location, site and habitat attributes, and dis- turbance history, they were not treated as true statisti- cal “replicates” (Hurlbert 1984; Heffner et al. 1996). Moreover, many of our data did not meet assumptions of normality or homegeneity of variance. Consequent- ly, we restricted our analyses to simple comparisons among Classes of habitat types. Relationships among stands were examined using multivariate analyses, with data input being matrices of species abundance by site (Kovach 1995). A cluster analysis was used to identify groupings of stands (or “communities”), using an un- weighted pair-group procedure with arithmetic aver- ages (UPGMA,; Sokal and Rohlf 1995). Chi-square contingency tests were used to test for differences in occupancy rate of nest boxes between (a) unmanaged and managed stands and (b) boxes located at 30 m or 100 m from a forest edge. Results and Discussion Habitat The stands of natural (reference) forest had a large basal area (an indicator of biomass) of trees compared with the other habitats studied (Table 2). Although a high tree density occurred in older plantations and the naturally regenerated site, these were smaller trees than in natural forest. The species composition of the nat- ural forest was mixed, whereas the plantations are more strongly dominated by the planted conifer species. The natural forest also had relatively abundant snags. Of the various managed stands, only the 7-year-old and 15-year-old plantations (P7 and P15, respectively) had many snags, because the pre-harvest snags had not all been removed or felled during the clear-cut, and a sub- sequent herbicide treatment killed surviving hardwood trees. The 30-year-old naturally regenerated clear-cut also had abundant snags, as did the 12-year-old selec- tively harvested site. Coarse-woody debris was abun- dant in the stands of reference forest, where it origi- nated as natural deadfall. Coarse-woody debris was also abundant in the plantations and selectively har- vested stands, where it mostly originated as logging slash and, in older plantations, trees cut and left dur- ing a non-commercial thin. Similar but more detailed observations have been made by other studies of nat- ural forest and plantations in our study area (Flem- ming and Freedman 1998). TABLE 2. Summary of key habitat data (presented as mean area). ON CAVITY-NESTING BIRDS a c ° p=) SS = c Ss oO Reference Stands Sl Rd Re Rb Ra Tree Basal Area (m2/ha) Conifers 18.3 19.0 30.3 30.2 18.3 10.9 10.7 6.4 11.4 10.9 10.9 Hardwoods Snags 5.0 Sal 3.6 6.1 Tree Density (10°*/ha) Conifers 0.9 0.5 0.6 1.0 0.9 0.6 0.5 Hardwoods Snags ala —~olama ANS ome os SO) o oaa aoc arse = —i— A~ va oO res oi ~aAr wo = Cln “oro —— ¥ vy vu ~~ -_ fc ey — Y aac = oc °o S 0.5 0.5 0.7 0.2 ().2 Shrub Basal Area (m2/ha) 301 an ro) Vol. 120 302 THE CANADIAN FIELD-NATURALIST ssouypry saieds SITAOPAIIT, [EO], UdIA\ IOUT, Jadaaig uMmoig yoyeynN paisvaiq-ayty AA, YoyeuINN poaysvoig-poy dapeyolyD jeoog sapeyoiyD paddeo-yoryg MOT[EMS OIL, Joyoadpoon, paeayid TOYS UTSYION Joyoadpoon, payorq-yorlgq Jayoodpoom, Alle Jaysadpoom, Aumoqd Jayonsdes pal[aq-Mol]an [MO JO4M-MES [MO pened [ensoy abe I C0 CIS Is 6Id 8Id LId SId 6d "6d 8d Ld Sd Cd ‘PoSAAIVY SUA JSOLOJ [LUISLIO ay] BOUTS SIVIA JO IAQUINU SU} Po}LOIPUL SOPOd-dUILU JIS YJIM PoJeLOOsseK ssoquINN “JNd-1eI[9 payesouasar A][e.NjeU P[O-1VIK-(Y¢E B O} SIBJII NAOAA LVN ‘paysoarey ATOATIOIIAS SIOM JeY} SOUS 0} SIOJOI NOLLOATAS ‘suonejuryd sayruoo poseuvur ATOATSUIJUT 0} S:IQJOI NOLLV.LNV Id {38010} Sorseds-poxtuy ‘amjeu ‘Pe.njeU 0} sJo}a1 FONAUAATa “AUIS ay] UO Juasaid seM satoads ay) sayeorput g ‘AIOILLIO} G'Q> Sa}voIpUT + ‘vy Q] Jed SaLiojLL19} se poziprepur}s o1e ve ‘Sp1g SuNsou-AjIAvS JO ssouYyoL satoads pue souepunqy ‘¢ ATAVL OcdN satoadg 2006 180 160 140 120 100 80 Linkage Distance 60 40 20 P8 P9a P5Sa P19 WOODLEY, JOHNSON, FREEDMAN, KIRK: EFFECTS ON CAVITY-NESTING BIRDS 303 FIGURE 1. Cluster analysis of stands based on their species and abundances of cavity-nesting birds. See Table | for explana- tion of site codes. The abundance and cover of shrubs and ground veg- etation were highly variable among stands (Table 3). In general, these vegetation elements were more abun- dant in the plantations and other harvested stands, which were in younger stages of secondary succes- sion than the more mature natural forest. Similar, but more detailed observations of forest and plantations in our study area have been made by Veinotte et al. (2004). Cavity-nesting Birds A total of 16 species of cavity-nesting birds was observed (Table 3). The most abundant species were the Black-capped Chickadee (24% of the territories; see Appendix | for avian binomials), Boreal Chick- adee (23%), Red-breasted Nuthatch (16%), and Win- ter Wren (14%). The species richness was higher in the natural forest (average of 10.0 species per stand; total of 14 species present in the habitat type) than in plan- tations (2.3/stand; 8 species present), selectively har- vested stands (7.0/stand; 9 species present), or the nat- urally regenerated clear-cut (5.0/stand). Abundance was also higher in the reference forest (average 5.3 terri- tories/10 ha) than in plantations (1.0/10 ha), selective- ly harvested stands (3.8/10 ha) or the naturally regen- erated clear-cut (2.5/10 ha). Among the ten plantations studied, the highest density of cavity-nesting birds was in P15 (4.0 territories/10 ha), which also had a rela- tively large number of snags (2.1 m?/ha compared with an average of 0.4 m’/ha among the other nine planta- tions). A cluster analysis segregated the “community” of cavity-nesting birds of the natural forest from those of the other treatments (Figure 1). In the cluster analy- sis, the five reference stands were arranged together on the right-hand side. The selectively harvested stands (S1 and $12) and naturally regenerated stand (NR30) had relatively abundant cavity-nesting birds and clus- TABLE 4. Cavity-using species encountered in this study. American Kestrel Barred Owl Northern Saw-whet Owl Northern Flicker Yellow-bellied Sapsucker Downy Woodpecker Hairy Woodpecker Black-backed Woodpecker Pileated Woodpecker Tree Swallow Black-capped Chickadee Boreal Chickadee Brown Creeper White-breasted Nuthatch Red-breasted Nuthatch Wood Duck Winter Wren Falco sparverius Strix varia Aegolius acadicus Colaptes auratus Sphyrapicus varius Picoides pubescens Picoides villosus Picoides arcticus Dryocopus pileatus Tachycineta bicolor Poecile atricapillas Poecile hudsonicas Certhia americana Sitta carolinensis Sitta canadensis AIx sponsa Troglodytes troglodytes 304 tered close to the reference stands. Of the plantations, those with relatively abundant and species-rich cavity- nesting birds clustered together (1.e., P7, P15, P19, and P17), while the most depauperate ones formed another cluster (i.e., P5b, P8, P9a, P9b, P5a). The observation that cavity-nesters are more abun- dant in mature mixedwood forest than in harvested stands or plantations is consistent with research else- where, although the specifics vary depending on the forest region and species of birds present (e.g., Raphael and White 1984; DeGraaf and Shigo 1985; Zarnowitz and Manuwal 1985; Hansen et al. 1995; Kirk and Nay- lor 1995; Niemi et al. 1998; Hobson and Bayne 2000). Of the 16 species of cavity-nesting birds recorded during our study, eight were not recorded in any of the plantations: Saw-whet Owl, Barred Owl, Yellow-bellied Sapsucker, Downy Woodpecker, Black-backed Wood- pecker, Pileated Woodpecker, White-breasted Nuthatch and Winter Wren. One species, the Tree-Swallow, was only recorded as breeding in plantations in which nest boxes were present. Nest-box Occupancy Nest boxes were used by five species for nesting and by four for roosting (Table 4). Tree Swallow ac- counted for 41% of the nests, American Kestrel 31%, and Boreal Chickadee 17%. The most frequently roost- ing species were Tree Swallow (48%) and Northern Flicker (33%). The use of nest-boxes for nesting was much greater in plantations (average 4.0/10 ha) than in the natural forest (0.3/10 ha), and similarly for roost- ing (average 2.9/10 ha in plantations and 0.3/10 ha in reference stands). Overall, 33% of the nest boxes in plantations were used for nesting or roosting (28 of 84 boxes with nests), compared with only 3% in nat- ural forest (1 of 36 boxes with nests). These observa- tions support the notion that suitable cavities are a lim- iting factor in plantations in the study area, compared with the natural forest. The plantations we studied THE CANADIAN FIELD-NATURALIST Vol. 120 appear to provide suitable habitat for foraging and other purposes, but can only be used for nesting and roosting by cavity-dependent birds if their need for cavities is met. There were no recorded cases of a failed nest in the nest boxes. Of the 28 nest boxes used for nesting in plantations, 7.5% were located 30 m from an intact forest edge and 15.8% were 100 m from such an edge (marginally significant difference; y* = 3.78, 1 df; >0.05 P <0.1). The apparent reluctance to use nest-boxes located clos- er to a habitat edge may be associated with a greater risk of predation (Paton 1994; Derochers and Hannon 199M): Implications for Management It appears that conifer plantations in our study region can recover bird populations rather quickly after estab- lishment, but the species composition is different from that of natural forest, and cavity-dependent species are present in low abundance (Freedman et al. 1994; Freed- man and Johnson 1999; Johnson and Freedman 2002). Moreover, observations and models of stand develop- ment suggest that intensively managed plantations are depauperate in snags, cavity-trees, and coarse-woody debris and are likely to remain so over subsequent rota- tions because the harvesting removes all large-dimen- sion tree biomass (Flemming and Freedman 1998). These changes do not augur well for cavity-depend- ent species in stands and landscapes extensively con- verted into forestry plantations. Clearly, the extensive conversion of mixedwood Acadian forest into conifer plantations will result in a decline in the abundance and species richness of cavity- dependent birds at the scales of both stand and land- scape. Certain site mitigations might help some species, particularly the retention of some snags and large liv- ing trees within plantations, or much less preferably, the provision of nest boxes to provide local nesting and roosting habitat (Welsh and Capen 1992; Newton TABLE 5. Use of nest boxes in plantations and reference forest during two study years. Each site had an array of 12 nest boxes, each made of a section of a hollow log of White Cedar. All plots were 10 ha in area. Note some stands from Table 3 had no nest boxes installed, so the number of stands differs between Table 3 and 4. Reference Species Nesting Wood Duck 1 American Kestrel 2; Northern Flicker Tree Swallow D) Boreal Chickadee 1 Total nests I 0 5 Roosting American Kestrel Pileated Woodpecker Northern Flicker Tree Swallow Total roosting Plantation 2 2 2 1 1 1 3 3 1 2 2 6 6 5 1 3 2 2 1 3 2 1 2 5 3 4 2006 1994; Flemming et al. 2000). It is likely, however, that these site mitigations will not be sufficient to maintain some cavity-dependent species, particularly those with large home ranges that encompass a mosaic of habitat patches, such as Pileated Woodpecker (Bonar 2000; Flemming et al. 2000). These species will require large protected areas of unmanaged forest to function as population centres. Without sympathetic management of this sort, it is likely that the population viability of some cavity- dependent birds in regions extensively managed for forestry will become compromised. Some jurisdictions have established guidelines to help these species — in New Brunswick, for example, timber companies oper- ating on Crown Land are to required to retain at least 10% of conifer-dominated forest in mature or older suc- cessional stages, in patches of 500 or more ha (Sulli- van 1996). However, these requirements may not be adequate to maintain critical habitat for all species of cavity-nesting birds, particularly keystone primary exca- vators (Flemming et al. 2000). It is essential that these conservation guidelines be monitored to ensure they achieve their intended ecological purpose, including the maintenance of viable populations of all cavity- dependent species. In the meantime, a precautionary approach to ecologically sustainable forestry would require that additional large areas of natural forest be set aside from intensive economic use, as parks or other kinds of protected areas. 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Pages 4-7 in Using Population Viability Analysis in Ecosystem Management at Fundy National Park. Edited by S. P. Flemming. Parks Canada Ecosystem Science Review Report 1, Atlantic Region, Halifax, Nova Scotia. Veinotte, C., B. Freedman, W. Maass, and F. Kirstein. 2004. Comparison of the ground vegetation in silvicultural plantations and natural, mixed-species forest in the Greater Fundy Ecosystem, New Brunswick, Canada. Canadian Field-Naturalist (in press) Virkkala, R. 1991. Population trends of forest birds in a Finnish Lapland landscape of large habitat blocks: conse- quences of stochastic environmental variation or regional habitat alteration? Biological Conservation 56: 223-240. Welsh, C. J. E., and D. E. Capen. 1992. Availability of nest- ing sites as a limit to woodpecker populations. Forest Ecol- ogy and Management 48: 31-41. Woodley, S., G. Forbes, and A. Skibicki. 1998. State of the Greater Fundy Ecosystem. Greater Fundy Ecosystem Re- search Group, University of New Brunswick, Fredericton, New Brunswick. Woodley, S., J. Kay, and G. Francis. 1993. Ecological integri- ty and the management of ecosystems. Heritage Resource Centre, University of Waterloo, Ontario. Zarnowitz, J. E., and D. A. Manuwal. 1985. The effects of forest management on cavity-nesting birds in northwest- ern Washington. Journal of Wildlife Management 49: 255-263. Received 12 April 2005 Accepted 5 May 2007 Recent Invasion, Current Status and Invasion Pathway of European Common Reed, Phragmites australis subspecies australis, in the Southern Ottawa District PAUL M. CATLING! and SUSAN CARBYN? 'Biodiversity, National Program on Environmental Health, Agriculture and Agri-food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1 A 0C6 Canada; e-mail: catlingp@agr.ge.ca Biodiversity, National Program on Environmental Health, Agriculture and Agri-food Canada, 32 Main Street, Kentville, Nova Scotia B4N IJS Canada; e-mail: carbyns@agr.gce.ca Catling, Paul M., and Susan Carbyn. 2006. Recent invasion, current status and invasion pathway of European Common Reed, Phragmites australis subspecies australis, in the southern Ottawa district. Canadian Field-Naturalist 120(3): 207-312. A survey of populations of native North American Common Reed (Phragmites australis subsp. americanus) and the introduced invasive alien, European Common Reed (Phragmites australis subsp. australis) was conducted in four eastern Ontario town- ships in 2003. The historical status of the two taxa in the region was evaluated through reference to herbarium specimens collected over the past century. The introduced subspecies appears to have entered the district in the 1970s and subsequently assumed dominance but was not recognized as an alien until 2003. It is now widespread in the four townships where ninety-five populations were recorded during the 2003 survey. Of these, 25 were referable to the native subspecies which was localized and 70 were referable to the introduced subspecies which was widespread. The native subspecies occurs in natural wetlands and also spreads to roadsides, but the introduced subspecies is much more strongly associated with roads, where the rhizomes extend onto gravel shoulders and are broken and transported by construction equipment, graders, ploughs, mowers, and in the treads of many kinds of vehicles. Sensitive wetlands should have buffer zones exceeding 1000 m to prevent colonization of subsp. australis. Monitoring of the two subspecies will be essential to the protection of native biodiversity, since early detection of the alien subspecies in a sensitive wetland will offer the best opportunity for control. Key Words: North American Common Reed, Phragmites australis subsp. americanus, European Common Reed, Phragmites australis subsp. australis, invasion pathway, alien, invasive species, Ottawa, Canada. One of the most productive plants in temperate regions, Common Reed, Phragmites australis (Cav.) Trin. ex Steud. has economic and agricultural values and is under-exploited (Small and Catling 2001). Domestication and introduction of non-native races however, pose some potential risks to both agricul- ture and environment. It is a major pest of irrigation and flood control channels in some parts of the world and, being able to grow in water 2 m deep or on dry ground, it is not readily controlled by drawdown. It was only recently realized that both native and introduced races existed in North America (Catling et al. 2003). The native plants have been designated sub- species americanus Saltonstall, P.M. Peterson & Soreng (Saltonstall et al. 2004) and the introduced plants iden- tified as subspecies australis (Catling 2006, 2007, Cat- ling et al. 2007). It is believed that the introduced sub- species australis originated from southern Europe. It has been recently recognized as a major threat to North American biodiversity (Chambers et al. 1999; Meyer- son et al. 2000; Catling et al. 2003; Robichaud and Cat- ling 2003; Catling 2005). In a recent survey of the St. Lawrence River wetlands it was found to have a greater impact on native plant diversity than most other inva- sive aliens including Purple Loosestrife, Lythrum sali- caria, (Lavoie et al. 2003). The subspecies australis is currently spreading in Canada from areas of concen- tration in southern Ontario and southern Quebec and has only recently been reported from western Canada (Martin 2003; Schueler et al. 2003) and the maritime provinces (Catling et al. 2004). In eastern Ontario the alien subspecies australis was first recognized in 2003, but it apparently invaded the area a few decades earlier. Local botanists noticed the rapid spread of dense stands of Phragmites beginning in the 1970s and the rapid change from the status of a locally rare plant (Dore 1959) created suspicion about the possibility of an introduced race. Preliminary obser- vations suggested that it spread into the region along roads very recently and became widespread. Here we analyse its apparent recent spread into the region along roads by providing information on its history of inva- sion, current status and invasion pathway in the south- ern part of the Ottawa district based on identification of herbarium specimens and a current survey of popu- lations in four adjacent townships. This work will also provide benchmark data for analysis of future changes in status of native and introduced subspecies, thus con- tributing information needed to predict rate of spread and impact. Methods A recent study of plants of Phragmites australis sen- su lato in eastern Ontario suggested that plants with lower glumes exceeding 4.2 mm long were referable to the native subspecies americanus (Robichaud and Cat- 307 308 subsp. australis subsp. americanus Ficure |. Glumes of Phragmites australis. The smaller on the right of each attached pair is the lower glume, whereas the longer glume on the left of each pair is the upper glume. The subspecies australis on the left has short- er lower glumes as represented by three examples from an inflorescence of a specimen from Dundas County (Catling and Robichaud, 10 Oct. 2002, DAO). On the right are three pairs from an inflorescence of subspecies americanus from Leeds and Grenville County (Catling and Kostiuk, 30 Aug. 2003, DAO) showing its characteristically longer lower glumes. ling 2003). More recent work based on correlation with basal stem color and date of collection (Saltonstall et al. 2004; Catling and Mitrow in press) has supported the separation using 4.2 mm for length of longer glumes, this separation being correct (1.e. correlated with other diagnostic features) 97.3% of the time (Figure 1). Most recently (Catling 2006) the following key has been rec- ommended for distinguishing the two subspecies: Basal internodes red or reddish-purple; longer lower glumes Sirf Molen MON ooo ecpe ease uc subspecies americanus Basal internodes pale yellow; longer lower glumes 2.6-4.2 (Giss})) oo ONS gh odudovogec vee subspecies australis For the study reported here undertaken in 2003, we used a length of 4.2 mm or more to identify the native subspecies americanus and 4.1 mm or less to identify the introduced subspecies australis in a survey of four adjacent geographic townships (Figure 2): Rus- sell (45.2833°N, 75.2833°W), Osgoode (45.2333°N, 75.5000°W), North Gower (45.1500°N, 75.6833°W) and Marlborough (45.0666°N, 75.8166°W). Hybrid- like or intermediate clones were not observed during this study but several were located subsequently in townships to the north. The survey was conducted in May 2003 at which time the persistent flowering stalks of the previous year were readily visible on the landscape and the persist- ing inflorescences were not so damaged as to prevent measurement of first glume lengths. The survey in- cluded driving all roads in each township, checking known locations and searching larger open wetlands in Rideau Township on foot. All small wetlands visi- ble from roads were examined with binoculars. Loca- tion of populations was recorded with GPS and size of THE CANADIAN FIELD-NATURALIST Vol. 120 patches and number of flowering stalks and distance from road were also recorded. Populations were defined as less than 500 m apart or 500 m in length. A voucher inflorescence from one of the tallest flowering stems was collected from each population and from these vouchers the length of the longest first glume of 25 examined was recorded. Voucher specimens for some the collections were deposited in the AAFC [Agricul- ture and Agri-Food Canada] vascular plant collection (acronym DAO [Department of Agriculture, Ottawa”’]). All flowering culms within individual stands were sim- ilar and were assumed to represent single clones. Collections in Canadian herbaria from Ottawa area and Carleton and Prescott counties were identified to race (using criteria described above) in order to deter- mine status of the two subspecies in the region in the past. The historical information was compared to maps of the occurrence of the native and introduced Eastern Ontario Townships surveyed 1. Marlborough 2. North Gower 3. Osgoode 4. Russell FiGurRE 2. Location of four adjacent townships in eastern Ontario where native and introduced subspecies of Phragmites australis were surveyed and mapped in May 2003. Inset map above shows eastern Ontario and Great Lakes with Eastern Ontario region shown in black. 2006 20 km CATLING AND CARBYN: EUROPEAN COMMON REED IN OTTAWA DISITRICT 309 Phragmites australis subsp. australis Phragmites australis subsp. americanus FiGurE 3. Distribution of native and introduced subspecies of Common Reed Phragmites australis in four eastern Ontario town- ships based on a survey in May 2003. Aove, introduced Eurasian subspecies australis. Below, native North American subspecies americanus. subspecies in the four townships based on the results of the 2003 survey. The relationship to roads was analysed by compar- ing the distribution of both subspecies by plotting on township maps showing roads using histograms show- ing frequency of various distances from roads. In order to further elucidate the history of spread, the size of patches was compared on major and minor roads, all at least 20 years old, in Russell township. Results History of Invasion In Canadian herbaria there are 45 specimens of Phragmites australis collected in the southern portion of the Ottawa district (Ottawa-Carleton Regional Muni- cipality and Prescott-Russell counties). Of these, 21 collected prior to 1970 are referable to the native sub- species. The earliest collection of the introduced sub- species was in 1976 at Manotick (Hanes, DAO). Of the 24 specimens collected after 1970, only five are referable to the native subspecies, the remainder hav- ing the relatively short first glumes of the introduced subspecies. Thus the introduced subspecies appears to have entered the district in the 1970s and subsequently assumed dominance but was not recognized as an alien until 2003 (Catling et al. 2003). Present Status Ninety-five populations were recorded in the four townships during the 2003 survey. Of these, 25 were referable to the native subspecies and 70 were refer- able to the introduced subspecies. Both subspecies 310 THE CANADIAN FIELD-NATURALIST Vol. 120 Ficure 4. Patch of pale stems of P. australis subsp. australis on gravel portion of a roadside and extending in a large clone in the adjacent ditch and swamp. Highway 417, Russell Township, May 2003. occurred in all four townships, but the native sub- species was much more localized, with most popula- tions in a single wetland (Figure 2). Invasion Pathway As seen in the distribution maps (e.g. Figure 3) there is a strong association of the introduced subspecies with roads (Figure 4). This is confirmed in the histogram (Figure 5) where the vast majority of the populations of the introduced subspecies are within 100 m of a road. In fact 52% were in the roadside gravel (Figure 4) and 84% were within 3 m of the boundary between vege- tation and gravel shoulder. Populations of the native subspecies occurred at greater distances from a road (Figure 5) with 36% on the shoulder and 60% within 3 m. Although occurrence of the native subspecies on roadsides was much less, it is clearly spreading from natural habitats to roadsides. Since both subspecies may come into close contact on roadsides, there is an opportunity for the subspecies to hybridize. The intro- duced subspecies was most frequent in the eastern townships that have the most roads (Figure 3). The native subspecies was most frequent in the far western township which has the least extensive road network and the most extensive natural wetlands. Major concentrations of the introduced subspecies occurred along Highway 417 (upper right in Figure 3), the busiest and oldest major highway in the region, and the largest patches in Russell township occurred along this road, suggesting that this is one of the places where it first entered the region (Figures 3 and 6). The asso- ciation of the introduced subspecies with roads sug- gested spread by dispersal of rhizomes by road vehi- cles rather than wind-blown seed which would have resulted in more occurrences at greater distances from roads. The tough rhizomes extend into the gravel shoul- ders and even onto the asphalt (Figure 4). They are readily broken and transported by ploughs, graders and in the tire treads of many kinds of vehicles (personal observation). This mechanism of dispersal is believed to be largely responsible for the rapid spread and pres- ent strong association with roads. There is a possibility that spread along roads is facilitated by relatively high sodium salt (NaCl) con- centrations, as noted for many other vascular plants spreading along roadsides (e.g. Catling and McKay 1980, 1981; Reznicek and Catling 1987). Salt concen- trations may reduce competition by eliminating other species, thus promoting salt-tolerant species. Although the alien subspecies may be tolerant of relatively high salt concentrations, reliance on salty habitats is prob- ably much less than for other rapidly spreading road- side plants such as Carex praegracilis (e.g., Reznicek and Catling 1987). Some patches of the alien sub- species of Common Reed do occur at great distances from roads and some roadside patches extend up to 100 m from roads beyond the influence of sodium salt. 2006 80 60 40 | Phragmites australis subsp. americanus 20 > 6) i 8 WW > 0 CG — 1 1 1 as ee a | wa ~ 80 Ww 60 F 40 Phragmites australis subsp. australis 20 ~The sn a 0) 500 1000 1500 2000 DISTANCE FROM ROAD (m) FiGurE 5. Histograms showing frequency of distances from major and minor roads of native and introduced sub- species of australis in four eastern Ontario townships. Further evidence for a lack of strong dependence on sodium salt is the observation that the alien subspecies of Common Reed occurred along many minor and gravel roads that do not receive applications of de- icing salt. The introduced subspecies has colonized two natu- ral wetlands in the study area without spreading from a roadside. One of these stands is 500 m from a road along a pond shore used extensively as a staging area by waterfowl. This colonization may have been the result of transport of rhizomes by the waterfowl, which occurs with many aquatic plants, but is unusual. Once in a wetland system the rhizomes are likely to be trans- ported further by aquatic mammals and birds for build- ing habitations, platforms and nests. Successful ger- mination of wind-dispersed seed is also a possibility, but strong association with roads suggests that it hap- pens infrequently. Both native and introduced popula- tions produce apparently viable seed with quantities varying between poplulations (personal observation, CATLING AND CARBYN: EUROPEAN COMMON REED IN OTTAWA DISITRICT 31) 10 } 8 | | 6 | Phragmites australis } ssp. australis i 4 minor roads 9 | > a5 UF Z | PE | Of yess. ee | is} = r uw 8 5 6 Phragmites australis ssp. australis 4 hwy 417 | | OF I i fy a at a fo Se 4000 Tt T 0 1000 2000 3000 POPULATION SIZE (square m) FIGURE 6. Histograms showing frequency of clone sizes of the introduced Phragmites australis subspecies aus- tralis in relation to major and minor roads in Russell Township. Catling). As alien Common Reed increases on the landscape the frequency of unusual dispersal mecha- nisms is likely to increase, and spread through wet- lands may increase substantially. Future Spread Considering that it is being spread by road vehi- cles along corridors of essentially continuous habitat, including new and disturbed habitats, the rate of spread can be anticipated to be rapid. Carex praegracilis W. Boott is believed to have spread the same way through- out much of eastern North America from the midwest in only 30 years (e.g., Reznicek et al. 1976; Reznicek and Catling 1987). Since the introduced subspecies of Phragmites australis has apparently spread from road- sides to distances of up to 500 m from roads (Figure 5), it appears that it is not confined to the near road environment. Although it seems less likely that the introduced subspecies will invade sensitive wetlands through introduction by waterfowl, the chances will increase as it becomes more abundant. There are also a2 opportunities for spread into isolated wetlands through transport along tracks of off-road vehicles including motorcycles, ATVs and snowmobiles. The introduced subspecies has extensively spread into both saline and non-saline wetlands and along both dry and wet roadsides in parts of the United States. Within another 10-20 years it is expected to line many major roadways in eastern Canada and it will have entered many wetlands from adjacent or nearby roads. As it spreads it is most likely to become established first along major roads, providing an early warning of future impact on the nearby landscape. The introduction of invasive aliens into wetlands by roads provides another reason to have extensive buffer zones around wetlands with high protection priority. Considering that the vegetative spread of P. australis can be very rapid, and mammals and birds transport rhizome fragments over substantial distances for nest- ing, buffers exceeding 1000 m are desirable. Monitor- ing of the two subspecies will be essential to the pro- tection of native biodiversity, since early detection of the introduced subspecies in a sensitive wetland will offer the best opportunity for control. Acknowledgments J. Cayouette, W. J. Cody and F. W. Schueler pro- vided useful comments. Literature Cited Catling, P. M. 2005. New “top of the list” invasive plants of natural habitats in Canada. Botanical Electronic News 345: 1-5. http://www.ou.edu/cas/botany-micro/ben/ben345. html#1. Catling, P. M. 2006. Notes on the lectotypification of Phragmites berlandieri and identification of North Amer- ican Phragmites. Botanical Eclectronic News 366: 5-8. http://www.ou.edu/cas/botany-micro/ben/ben366.html#2. Catling, P. M. 2007. Additional notes on the identification of Phragmites in Canada. Botanical Eclectronic News 366: 1-3. http://www.ou.edu/cas/botany-micro/ben/ben370 html. Catling, P. M., and S. M. McKay. 1980. Halophytic plants in southern Ontario. Canadian Field-Naturalist 94: 248-258. Catling, P. M., and S. M. McKay. 1981. A review of the occurrence of halophytes in the eastern Great Lakes region. Michigan Botanist 20: 167-180. Catling, P. M., G. Mitrow, L. Black, and S. Carbyn. 2004. Status of the alien race of common reed in the Canadian maritime provinces. Botanical Electronic News 324: 1-3. http://www.ou.edu/cas/botany-micro/ben/ben324.html. THE CANADIAN FIELD-NATURALIST Vol. 120 Catling, P. M., G. Mitrow, and L. Black. 2007. Analysis of stem colour and correlated morphological characters for grouping Phragmites (Poaceae) taxa in eastern Ontario. Rhodora. Jn press. Catling, P. M., F. W. Schueler. L. Robichaud, and B. Blossey. 2003. More on Phragmites — native and introduced races. Canadian Botanical Association Bulletin 36(1): 4-7. Chambers, R. M., L. A. Meyerson, and K. Saltonstall. 1999. Expansion of Phragmites australis into tidal wet- lands of North America. Aquatic Botany 64: 261-273. Dore, W. G. 1959. Grasses of the Ottawa district. Publica- tion 1049. Research Branch, Department of Agriculture, Ottawa. 101 pages. Lavoie, C., M. Jean, F. Delisle, and G. Létourneau. 2003. Exotic plant species of the St. Lawrence River wetland: a spatial and historical analysis. Journal of Biogeography 30: 537-549. Martin, M. 2003. Common Reed (Phragmites australis) in the Okanagan Valley, British Columbia, Canada. Botani- cal Electronic News 318: 1. http://www.ou.edu/cas/botany- micro/ben/ben318 html Meyerson, L. A., K. Saltonstall, L. Windham, E. Kiviat, and S. Findlay. 2000. A comparison of Phragmites aus- tralis in freshwater and brackish marsh environments in North America. Wetlands Ecology and Management 8: 89- 103. Reznicek, A. A., and P. M. Catling. 1987. Carex praegra- cilis (Cyperaceae) in eastern North America: a remark- able case of rapid invasion. Rhodora 89 (858): 205-216. Reznicek, A. A., P. M. Catling, and S. M. McKay. 1976. Carex praegracilis W. Boott, recently adventive in south- ern Ontario. Canadian Field-Naturalist 90: 180-183. Robichaud, L., and P. M. Catling. 2003. Potential value of first glume length in differentiating native and alien races of common reed, Phragmites australis. Botanical Elec- tronic News 310: 1-3. http: www.ou.edu/cas/botany-micro/ ben/ben310.html. Saltonstall, K., P. M. Peterson, and R. J. Soreng. 2004. Recognition of Phragmites australis (Poaceae) Arundi- noideae) in North America: evidence from morphologi- cal and genetic analysis. Sida 21(2): 683-692. Schueler, F. W., A. Karstad, and J. H. Schueler. 2003. Non-native Phragmites communis in British Columbia. Botanical Electronic News 315: 1 http://www.ou.edu/cas/ botany-micro/ben/ben315.html. Small, E., and P. M. Catling. 2001. Poorly known econom- ic plants of Canada - 29. Common reed, Phragmites aus- tralis (Cav.) Trin. ex Steud. Canadian Botanical Associa- tion Bulletin 34(2): 21-26. Received 26 April 2005 Accepted 17 February 2007 Pack Size of Wolves, Canis lupus, on Caribou, Rangifer tarandus, Winter Ranges in Westcentral Alberta GERALD W. Kuzyk!“, JEFF KNETEMAN2, AND FIONA K. A. SCHMIEGELOW! ' Department of Renewable Resources, University of Alberta, Edmonton, Alberta, T6G 2H1 Canada * Alberta Sustainable Resource Development, Fish and Wildlife Division, Hinton Alberta, T7V 2E6 Canada 3 Present address: Ministry of Environment, Wildlife Science Section, P.O. Box 9338, Provincial Government, Victoria, British Columbia V8W 9M1 Canada; e-mail: Gerald. Kuzyk@gov.be.ca Kuzyk, Gerald W., Jeff Kneteman, and Fiona K.A. Schmiegelow. 2006. Pack size of Wolves, Canis lupus, on Caribou, Rangifer tarandus, winter ranges in westcentral Alberta. Canadian Field-Naturalist 120(3): 131-318. We studied pack size of Wolves (Canis lupus) on Woodland Caribou (Rangifer tarandus caribou) winter ranges in westcentral Alberta. These Caribou winter ranges are experiencing increasing pressure from resource extraction industries (forestry, energy sector) and concerns have been raised regarding increased Wolf predation pressure on Caribou in conjunction with landscape change. Thirty-one Wolves, from eight Wolf packs, were fitted with radiocollars on two Caribou winter ranges in the Rocky Mountain foothills, near Grande Cache, Alberta (2000-2001). There was a mean of 8.2 Wolves/pack and between 30 and 39 Wolves on each of the RedRock/Prairie Creek and Little Smoky Caribou ranges. The average pack size of Wolves in this region does not appear to have increased over that recorded historically, but the range (5-18) in the number of Wolves per pack varied considerably over our study area. Wolves preyed predominately on Moose (Alces alces), averaging one Moose kill every three to five days. There was some indication that pack size was related to prey size, with the smallest pack preying on Deer (Odo- coileus spp.). It was clear that Caribou could not be the primary prey for Wolves, due to their low numbers, and relative to the pack size and Wolf kills we observed. Key Words: Wolf, Canis lupus, Caribou, Rangifer tarandus, Moose, Alces alces, White-tailed Deer, Odocoileus virginianus, predation, pack size, colour phase, kills, Alberta. Pack size of Wolves (Canis lupus) can provide an important indication of ungulate abundance (Fuller 1989; Schmidt and Mech 1997; Hayes et al. 2003) and human impacts on the landscape (Mech 1995). The number of Wolves in each pack can be related to prey availability (Fuller 1989) and may be regulated by dispersal, pup survival and disease prevalence (Mech 1970; Mech et al. 1998). The amount of food available to each Wolf has been inversely related to pack size (Thurber and Peterson 1993; Schmidt and Mech 1997), with smaller pack sizes found in areas of low ungulate density (Messier 1985). In contrast, pack size may re- main stable when the amount of food per Wolf decreas- es, possibly due to a genetic advantage for the breeding pair when sharing limited food with their offspring (Schmidt and Mech 1997). Maximum Wolf pack size may be regulated by behaviors within the pack, such as social dominance, which can facilitate dispersal. Pack size has also been related to prey size. Average pack size of Wolves hunting White-tailed Deer (Odocoileus virginianus) is generally less than seven (Mech 1970; Fuller 1989), whereas packs preying on Moose (Alces alces) are generally larger, having ten or more Wolves per pack (Peterson et al. 1984; Mech et al. 1998; Hayes et al. 2003). Vucetich et al. (2004) demonstrated that larger packs are better able to compete with ravens and do accrue a foraging advantage. Reports of large Wolf Packs with over 20 members have been documented in regions where there is little Wolf harvest (Hayes and Harestad 2000) or in protected areas (Carbyn et al. 1993; Peterson 1995; Mech et al. 1998; Smith et al. 2004). Predators may negatively affect endangered prey species when landscapes become impacted by human disturbance (Schneider 2001). Landscapes in west- central Alberta are facing increased pressures from forest harvesting and the energy sector (oil and gas). Forest harvesting may alter the movements and distri- bution of Wolves (Kuzyk et al. 2004) and ungulates (Smith et al. 2000), and in combination with linear cor- ridors (roads, seismic lines) from oil and gas activities, can enhance human access to once remote areas and possibly increase human-caused Wolf mortality from shooting and trapping. Enhanced predation by Wolves, facilitated by landscape change such as increased travel efficiency on linear corridors (James and Stuart-Smith 2000) and associated increases in alternate prey pop- ulations (James et al. 2004), is thought to be the pri- mary factor for Caribou declines in Alberta (Edmonds 1988; McLoughlin et al. 2003). In Alberta, Woodland Caribou (Rangifer tarandus caribou) are classed as a threatened species under the provincial Wildlife Act and on the Species at Rist Act Public Registry [SARA, http://www.sararegistry.gc.ca/species/Species Details_e.cfm?sid=636] and special management con- siderations are necessary to maintain numbers and habitat (Edmonds 1998; Dzus 2001). More informa- tion is required on Wolf populations, such as pack sizes, to be better able to predict long-term changes in the dynamics of Wolves and Caribou associated with land- Sule) 314 scape change (Weclaw and Hudson 2004). The objec- tive of this paper is to present current information on pack size and associated predation by Wolves that can be used in decision-making processes for long-term Caribou conservation. Study Area The study area is approximately 5000 square kilo- metres, located in the foothills of west-central Alberta, near the town of Grande Cache (54°N 119°W). The area is classed into subalpine and boreal natural sub- regions (Beckingham and Archibald 1996), and con- tains several main rivers but lakes are scarce. Elevations range from 1300-1800 metres, and the climate is sub- arctic, with short wet summers and long cold winters. Temperatures average 16°C in July and -13.5°C in December (Beckingham and Archibald 1996). The forests are primarily Lodgepole Pine (Pinus contorta) and some White Spruce (Picea glauca). The wetlands support mostly Black Spruce (Picea mariana) and some Tamarack (Larix laricina). Some south facing slopes have Aspen (Populus tremuloides) and willow (Salix sp.). The study area contains three major Caribou herds: the Red Rock/Prairie Creek and Al La Peche are moun- tain herds and represent a migratory ecotype, while the Little Smoky herd is boreal or sedentary ecotype (Edmonds 1988). Population size for the mountain herds is estimated at 600-750 Caribou (Edmonds 1998), and the Little Smoky herd is estimated at fewer than 100 animals (Smith 2004). The study area also sup- ports a high diversity of other large mammals. Moose are found at densities ranging from 0.12 to 0.25 Moose per square kilometer (Alberta Fish and Wildlife unpub- lished data). Other ungulates occurring in the area are Elk (Cervus elaphus), Mule Deer (Odocoileus hemi- nous), White-tailed Deer, Bighorn Sheep (Ovis cana- densis), Mountain Goats (Oreamnos americanus) and Wild Horses (Equus cabalus). Wolves (Canis lupus), Coyotes (Canis latrans), Grizzly Bears (Ursus arctos), Black Bears (Ursus americanus) and Cougars (Felis concolor) also exist throughout the study area. Wolf Captures and Monitoring Wolf captures were conducted on two Caribou win- ter ranges in the winters of 2000 and 2001. Four Wolf packs were located on the winter range of a migratory mountain Caribou population (Red Rock/Prairie Creek herd) and four packs located on the winter range of a sedentary boreal Caribou population (Little Smoky herd) (Figure 1). Ungulate baits were placed in strate- gic locations seven to ten days prior to the initiation of the Wolf capture operation, to enhance detection of Wolves. Wolves were located by following trails in the snow from a fixed-wing aircraft (Mech 1966; Mech et al. 1998). All Wolf handling was approved by the Faculty of Agriculture, Forestry and Home Economics THE CANADIAN FIELD-NATURALIST Vol. 120 Animal Care Policy (Number 96-99D), subject to the protocols of the Canadian Council of Animal Welfare. Wolf captures were accomplished by either helicopter darting (Ballard et al. 1991) or netgunning, then phys- ically restraining the Wolf with restraining forks, and hand-injecting 1-2 mls of telazol at 200mg/ml (Kuzyk 2002). Adults were distinguished from pups by tooth eruption patterns (Van Ballenberghe and Mech 1975) whereas yearlings were classed on subjective physical criteria (size), as there is no definitive method to cat- egorize yearling Wolves (Mech et al. 1998). Adult Wolves were fitted with store-aboard GPS (Global Positioning System) collars (Lotek Engineering Sys- tems, Newmarket, Ontario or Televilt GPS-Simplex, Lindesberg, Sweden) or VHF (Very High Frequency) radiocollars (Lotek Engineering). Pups were fitted with VHF collars only. All Lotek GPS collars were equipped with remote dropoff units, intended to release when signaled from the air. All radiocollared Wolves were relocated by aerial radiotracking (Mech 1974) within one to four days of capture to determine if they had rejoined their packs. Color phases of Wolves were recorded as gray, black, or white following Dekker (1986), and adding a category of blue. Wolf pack size was recorded during aerial radio- tracking in February and March of each year, when pack size would be at a minimum (Mech 1970), thus yielding a conservative estimate. Most packs were intensively monitored in March of 2000, increasing the chances of a good count. The best estimates of Wolf pack size were made when the Wolves were traveling in single file on a linear corridor or river. Lone Wolves were assumed to account for 10% of the total popula- tion (Fuller 1989). We did not extrapolate our pack size data to determine Wolf density, due to insufficient temporal data required to adequately determine annual territory sizes. In addition, our broader project objec- tives entailed studying Wolves in late winter, corre- sponding with winter range occupancy by migratory Caribou. It is the winter range of these caribou that is currently undergoing industrial development, and thus of primary conservation concern. Wolf kills were determined during a two-week peri- od in March 2000 by aerially locating radiocollared Wolf packs and finding their ungulate kills (Mech 1974). Flights were conducted twice daily in hopes of detecting Wolf-killed deer (Fuller 1989). When a Wolf pack was located, Wolves were counted and the area searched for ungulate carcasses. A kill was as- sumed to be caused by Wolves if there were blood- trails indicating a successful chase and a disarticulated carcass (Hayes et al. 2000). At each kill site, the number and behavior of Wolves were recorded (Mech 1966). All Wolf-killed ungulates that could not be identified to species, gender and age (adult-calf) from the air were later ground inspected. 2006 7 eee ES Alberta * Grande Prairie LYNX CREEK PACK Grande Cache Wolf locations (GPS) Wolf Locations (VHF) Wolf pack territory Preliminary Wolf Pack Territroy | Caribou Management Zones Major Waterways Town of Grande Cache KUZYK, KNETEMAN, AND SCHMIEGELOW: PACK SIZE OF WOLVES ilometers | Ficure 1. Distribution of eight Wolf packs on two Caribou winter ranges in west-central Alberta during 2000 and 2001. TABLE 1. Sex ratio, age class and color phase of Wolves (n=31) captured in west-central Alberta during 22 January — 13 March, 2000 and 15—17 February, 2001. Sex Ratio Age Class Color Phases (n=31) (n=31) (n=31) 14 Males 15 adults 19 black (61%) 17 Females 8 yearlings 8 gray (26%) 8 pups 3 white (11%) 1 blue (2%) Results Thirty-one Wolves were captured on two Caribou ranges: 18 captured in 2000 and 13 in 2001 (Table 1). Nine Wolves were aerially darted and 24 were netted (two recaptures). There were no capture-related Wolf mortalities during this study. A total of 16 GPS (14 Lotek and 2 Televilt) and 17 VHF radiocollars were de- ployed. There were seven known mortalities of radio- collared Wolves (23%) over fifteen months: two were shot and five died of natural or unknown causes. Colors phases of the radiocollared Wolves were 61% black, 26% gray, 11% white and 2% blue. Eight Wolf packs were located on the two Caribou ranges (Tables | and 2; Figure 1). There were between 54 and 77 (mean = 66) Wolves on the Little Smoky and Red Rock/Prairie Creek Caribou ranges, equating TABLE 2. Pack size of Wolves on the Red/Rock Prairie Creek (RPC) and Little Smoky (LSM) Caribou winter ranges in west-central Alberta in late winters of 2000 and 2001. Wolf Pack Estimated pack size Caribou herd Lynx Creek 12-18 RPC Cutbank 7-8 RPC Prairie Creek 5-6 RPC Sheep Creek 6-7 RPC Simonette 7-11 LSM Little Smoky 7 LSM Deep Valley 9 LSM Berland 8-11 LSM Total (Range) 54-77 Total (Mean) 65.5 3) io) Mean TABLE 3. Ungulate kills by Wolf packs in west-central Alber- ta in March 2000. Wolf Days Moose Deer Ungulate Pack Monitored kills kills __ kills/day Lynx Creek 9 (Mar 2-10) 2 1/45 Cutbank 14 (Mar 2-15) 3 1/4.7 Simonette 14 (Mar 2-15) 4 2 1/18 Note: Prairie Creek pack not entered as only one Elk kill was located. 316 to 8.2 Wolves/pack. Each Caribou range had between 30-39 Wolves with Wolf pack sizes ranging from 5-18 (Table 2). Twelve ungulate kills were recorded from four Wolf packs during 14 days of monitoring in March 2000 (Table 3). Ungulate kills consisted of seven cow and two calf Moose, two deer (unknown species) and one cow Elk. Wolves preyed predominately on Moose, averaging one Moose-kill every three to five days. Discussion We recorded a mean pack size of 8.2 Wolves/pack for eight Wolf packs on the RedRock/Prairie Creek and Little Smoky Caribou ranges. This is similar to the 8.7 Wolves/pack averaged over five other Alberta Wolf studies conducted between 1975 and 1985 (Gun- son 1992) and marginally higher than the 6.8 Wolves per pack recorded in January 1995 in west-central Alberta during the Yellowstone-Idaho Wolf relocation (Kneteman 1995*). Pack size of Wolves in our study area varied considerably, ranging from 5 to 18 Wolves/ pack. The larger Wolf packs (Lynx Creek n = 18; Si- monette n = 11) preyed primarily on Moose, whereas the smallest pack (Prairie Creek n = 5) preyed pre- dominately on deer, consistent with results from other nearby studies (Carbyn 1974; Weaver 1994). A con- founding factor in estimating numbers of Wolf-killed ungulates in west-central Alberta is the difficulty of detecting Wolf-killed deer (Kuzyk 2002; Kuzyk et al. 2005), due to the small size and cryptic color of deer, and the short time required for Wolves to consume deer carcasses (Fuller 1989). All Wolf packs in this study were observed either hunting deer or at deer kills. During intensive monitoring in March 2000, the Prairie Creek pack was seen hunting deer, and was thought to have made deer kills, but those were never detected from the aircraft. This resulted in somewhat ambiguous information, as data indicate this pack killed only one Elk in nine days of monitoring. But further analysis of GPS data collected during the nine- day monitoring period suggested the Prairie Creek pack had made a minimum of three deer kills (Franke et al. 2006). The importance of deer to Wolves in this study area should not be underestimated. Kuzyk et al. (2005) found Wolves in this study area traveled 4.2 times less when near ungulate carcasses then when away from them, and suggested that Wolf packs preying primarily on deer, as opposed to Moose, may pose a greater pre- dation risk to Caribou due to associated increase travel and encounter rates. Further research to quantify the importance of deer in this Wolf-ungulate system should be initiated such as measuring Wolf encounter rates with deer (Kunkel et al. 2004). No Caribou kills were detected during this study, probably due to the low numbers of Caribou in the region (Smith 2004) and the short time it takes Wolves to consume a carcass (Hayes et al. 2000). Nevertheless, Caribou could not be the primary prey for Wolves in westcentral Alberta, as the numbers of Caribou could THE CANADIAN FIELD-NATURALIST Vol. 120 not support the numbers of Wolves we recorded. For example, there are fewer than 100 Caribou in the Lit- tle Smoky herd (Smith 2004), and four Wolf packs overlap their range, with each pack killing an esti- mated 40-85 ungulates each winter. Humans may impact Wolf pack size when access to remote areas increases, and human-caused mortality to Wolves increases from shooting, trapping and road collisions (Mech 1995). Landscapes in west-central Alberta are being altered by resource extraction indus- tries which have increased human access to previously remote areas. We recorded 23% mortality for radio- collared Wolves over fifteen months, similar to annual mortality rates (20-27%) found in other studies (Plet- scher et al. 1997; Mech et al. 1998). This level of mortal- ity is thought to be low and would not influence over- all Wolf population size due to the high reproductive and dispersal rates of Wolves (Mech et al. 1998; Hayes et al. 2003). Forest harvesting can increase human ac- cess and change the amount and spatial distribution of habitats for Wolves and their prey. Wolf packs in our study area had a seven-fold difference in the amount of forest harvested within their territories (Kuzyk et al. 2004). The Cutbank pack (n = 7-8) had 36% of the forest in their territory harvested whereas the Lynx Creek (n = 12-18) pack had 7% harvested. The lowest level of forest removal corresponded with the largest pack size of 18 Wolves. In 2001, this pack contained 12 black and 6 gray Wolves and was observed traveling on a road in an open forest cutblock (Kuzyk 2001). This pack size exceeds the largest recorded from nearby studies: 12 Wolves in the Simonette River area (Bjorge and Gunson 1989) and 10 Wolves in northern Jasper National Park (Weaver 1994). The color phase of a Wolf may influence its detec- tion by humans (Mech et al. 1998) and subsequent mor- tality. Radiocollared Wolves in this study were pre- dominantly black (61%), similar to those recorded in nearby Jasper National Park (53% black) (Dekker 1986). It is noteworthy that black Wolves may change to a lighter color possibly due to aging or physiologi- cal stress (Gipson et al. 2002). Black Wolves may be more easily detected by hunters, especially during win- ter and on fragmented landscapes. The two Wolves shot during our study were black, and of the seven total mortalities (2 shot, 5 natural or unknown), five Wolves were black (71%). Lone and small groups of Wolves are also an im- portant component of this population. During the two years of our monitoring, several Wolves dispersed from their natal territories, as individuals or in small groups (<3 Wolves), consistent with other research (Gese and Mech 1991). These lone Wolves, or small groups, could represent 10-30% of the Wolf population (Fuller 1989; Mech et al. 1998) and may be an important factor when assessing predation risk to Caribou. Such Wolves would be travelling great distances to establish new territories (Mech 1970), and may use human trails as travel routes 2006 (Kuzyk and Kuzyk 2002), thereby increasing their chances of encountering Caribou. In addition, pairs of Wolves may have proportionately higher kill rates than larger packs (Hayes et al. 2000). Further, if Wolf packs generally avoid Caribou habitats due to a lack of Moose (James et al. 2004), dispersing Wolves may select these habitats to avoid being killed by resident pack Wolves defending their territories, a primary cause of natural Wolf mortality (Mech 1994). Conducting research on single or small groups of Wolves would be logistically difficult but resulting information could lead to impor- tant insights into Caribou predation risk from Wolves. Documenting current pack size and prey relations of Wolves in our study area is an important step in understanding the potential implications of landscape change and resultant alteration of predator/prey sys- tems for Caribou in the area. However, as pack size could theoretically remain constant while overall num- bers increase, determining Wolf density is necessary. In combination with additional information on kill rates and prey availability, this would permit evaluation of the numerical and functional responses of Wolves in this system to landscape alteration, and provide a foun- dation for examining alternative management strategies aimed at long-term Caribou conservation. Acknowledgments Funding for this research was provided by the West- Central Alberta Caribou Standing Committee; the Al- berta Sport, Recreation, Parks and Wildlife Foundation; a University of Alberta Challenge Grant in Biodiversity (supported by the Alberta Conservation Association); and the Foothill Model Forest. We acknowledge the safe piloting conducted by C. Wilson from Bighorn Helicopters and D. Dennison from Coyote Air during Wolf captures and monitoring. S. Shirkoff and K. Lisgo provided logistical and data management support. The spatial data on the figure is published with permis- sion of Alberta Sustainable Resource Development, Natural Resource Canada and the Spatial Data Ware- house. Thanks to K. Smith for helpful comments on earlier drafts of this paper. Documents Cited (marked * in text) Kneteman, J. 1995. Summary report of gray wolf relocation from Alberta to Yellowstone National Park and central Idaho. 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Ecological Modelling 177: 75-94 Received 3 May 2005 Accepted 5 March 2007 Bur Buttercup, Ranunculus testiculatus, New to Eastern Canada MICHAEL J. OLDHAM!, CLIVE E. Goopwin2, and SEAN BLANEY? ' Ontario Natural Heritage Information Centre, Ministry of Natural Resources, 300 Water Street, P.O. Box 7000, Peterbor- ough, Ontario K9L 1C8 Canada * | Queen Street, Suite 405, Cobourg, Ontario K9A 1M8 Canada 4 Atlantic Canada Conservation Data Centre, P.O. Box 6416, Sackville, New Brunswick E4L 1C6 Canada Oldham, Michael J., Clive E. Goodwin, and Sean Blaney. 2006. Bur Buttercup, Ranunculus testiculatus, new to eastern Canada. Canadian Field-Naturalist 120(3): 319-322. Bur Buttercup (Ranunculus testiculatus) is newly reported for eastern Canada based on two collections from campgrounds in southern Ontario. This vernal, annual, Eurasian weed is widespread in western North America and is expanding its range in the east; it should be expected elsewhere in eastern Canada. Bur Buttercup is known to be toxic to livestock. Key Words: Bur Buttercup, Ranunculus testiculatus, Ranunculaceae, first records, weed, range expansion, campground, Ontario, eastern Canada. On 17 April 2004, Blaney found an unusual butter- cup growing on a campsite in Long Point Provincial Park, on the north shore of Lake Erie, Norfolk County, Ontario (42.58°North, 80.41° West). A few days later on 22 April 2004, Goodwin found the same species growing in a municipal campground in Cobourg, on the north shore of Lake Ontario, Northumberland Coun- ty, Ontario (43.96°North, 78.16° West). This distinctive plant, called Bur Buttercup (Ranunculus testiculatus Crantz), has not previously been reported from Ontario (Morton and Venn 1990; Newmaster et al. 1998) or eastern Canada (Scoggan 1978; Gleason and Cron- quist 1991; Whittemore 1997; Kartesz 1999). Ranunculus testiculatus is a small scapose, some- what tomentose, annual in the buttercup family (Ranun- culaceae) which flowers early in the season. The leaves are entirely basal and are 1.5 to 4 cm long, ternate to biternate, with narrow linear divisions. The several leaf- less peduncles are 2 to 8 cm tall, and each bears a sin- gle, inconspicuous pale yellow flower (Figure 1). The fruiting heads are bur-like when dry and mature, giving the plant its common name, Bur Buttercup. Superfi- cially the plant is very different in appearance from other buttercups and is often placed in a separate genus as Ceratocephalus testiculatus (Crantz) Roth by some authors (e.g., Cody 1988; Cusick 1989) or Cerato- cephala testiculata (e.g., Tutin et al. 1993; Mitchell and Tucker 1997; Kartesz 1999). However, most recent North American floras now include it within Ranuncu- lus (e.g., Gleason and Cronquist 1991; Whittemore 1997). A related species, R. falcatus L., has been report- 1 \ ed from North America, but all reports are apparently \ based on misidentified material of R. testiculatus (Whit- _ temore 1997). Common names used for R. testicula- tus in North America include Bur Buttercup or Bur- ' buttercup (e.g., Barkworth 1982; Pohl 1984; Cusick _ 1989: Gleason and Cronquist 1991; Swink and Wilhelm _ 1994), Hornseed Buttercup (e.g., Cody 1988; Douglas et al. 1999), Sage Buttercup (e.g., Mitchell and Tucker 1997), and Curve-seed-butterwort (e.g., Kartesz 1999). The first North American report of Ranunculus tes- ticulatus was from Utah in 1932, followed by discover- ies in other western states soon after: Oregon in 1938, Washington in 1940, and Colorado in 1948 (Barkworth 1982). It continues to spread in western North America (Hitchcock and Cronquist 1973; Taylor 1983; Weber 1990; Whittemore 1997). In Canada, Bur Buttercup has been known from southern British Columbia for some time (Boivin 1966; Scoggan 1978), where it is infrequent in south-central and south-eastern parts of the province in dry disturbed clearings and sagebrush slopes in the steppe zone (Douglas et al. 1999). Cody (1988) report- ed the species new to Saskatchewan based on a collec- tion from Assiniboia in 1987. Cody (1988) also mapped the Canadian distribution of R. testiculatus. Cusick (1989) was the first to report Bur Buttercup in eastern North America, based on a 1977 collection from South Bass Island in western Lake Erie, and pre- dicted it would be found elsewhere in the Great Lakes region. In New York State it was first collected in 1990 from the parking lot of a campground near Har- riman in Orange County by D. J. DeLaubenfels (speci- men in the New York State Museum, NYS A18183, personal communication C. Sheviak and T. Weldy). The first Chicago area collection was made in 1991 and with concerted searching it was found in 12 additional Chicago region counties in 1992 and 1993 (Swink and Wilhelm 1994). Other recent new state records are from Nebraska in 1970 (Macgrath and Weedon 1974), Iowa in 1984 (Pohl 1984), and Missouri in 1987 (Ladd and Schuette 1990). Bur Buttercup is considered a significant toxic weed of grazing lands in the western United States and has been responsible for the death of sheep (Olsen et al. 1982, 1983). Several authors have noted the affinity of this species for campgrounds, particularly in eastern North Amer- 319 320 THE CANADIAN FIELD-NATURALIST Vol. 120 Ficure |. Bur Buttercup (Ranunculus testiculatus) plants. Long Point Provincial Park, Ontario, 6 May 2004. Photograph by Michael Oldham. ica, for example the first reports from Iowa, Missouri, New York, Ohio, and the Chicago Region are all from campgrounds (Pohl 1984; Cusick 1989; Ladd and Schuette 1990; Swink and Wilhelm 1994; C. Sheviak personal communication). It seems likely that this species is dispersed by camping vehicles and associ- ated equipment. The bur-like fruiting heads of the buttercup may readily attach to blankets and tents, and are thus carried to campsites that appear to provide ideal disturbed conditions. Other vernal, ephemeral Eurasian weedy species have found a niche in camp- grounds and are presumed to have a similar dispersal mechanism, such as Aira caryophyllea (Silvery Hair- grass), Aira praecox (Early Hairgrass), Apera interrup- ta (Interrupted Windgrass), Cerastium pumilum (Cur- tis’ Mouse-ear Chickweed), Cerastium semidecandrum (Small Mouse-ear Chickweed), Erophila verna (Spring Whitlow-grass), Myosotis stricta (Blue Scorpion-grass), Poa bulbosa (Bulbous Bluegrass), and Veronica verna (Spring Speedwell) (Crins et al. 1987; Oldham et al. 1995; Goltz 1996; Oldham and Zinck 1997). Both Ontario records are from campgrounds, a municipal campground in Cobourg, Northumberland County, and a provincial park campground at Long Point, Norfolk County. At both Ontario locations Bur Buttercup is well established but quite local. At Long Point Provin- cial Park at least 500 plants occur, but they were found on only a single campsite, possibly suggesting a rela- tively recent introduction. In addition, the Long Point peninsula has been relatively well botanized in the past (Reznicek and Catling 1989). At Cobourg, Bur Buttercup grew in open, disturbed ground on at least four campsites with Erophila verna and Taraxacum officinale (Common Dandelion), while at Long Point it was found in sandy soil beneath planted Scots Pine (Pinus sylvestris) with Cerastium semidecandrum, Stel- laria pallida (Lesser Chickweed), Poa annua (Annual Bluegrass), Myosotis stricta, and Taraxacum offici- nale (Figure 2). Collection data for the Ontario records are as follows: Ontario, Norfolk County, Long Point Provincial Park, 42.58°North, 80.41°West, 6 May 2004, M. J. Oldham 29983 (DAO, MICH, NHIC, TRTE, UWO). Northumberland County, Cobourg Municipal Campground, 43.96°North, -78.16°West, 9 May 2004, M. J. Oldham 29996 (DAO, MICH, NHIC, TRTE). Acknowledgments Troy Weldy and Charles Sheviak provided infor- mation on Bur Buttercup in New York state. The comments of two anonymous reviewers were helpful in improving the manuscript. 2006 OLDHAM, GOODWIN, and BLANEY: BUR BUTTERCUP 32) FIGURE 2. Bur Buttercup (Ranunculus testiculatus) habitat at Long Point Provincial Park, Ontario, on 6 May 2004. Bur But- tercup plants appear as a light coloured area in front of the closest Scots Pine tree in the foreground. Photograph by Michael Oldham. Literature Cited Barkworth, M. 1982. Bur buttercup: a weedy immigrant. Utah Science 43: 6-9. Boivin, B. 1966. Enumération des plantes du Canada. II — Herbidées, 1° partie. 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Flora Europaea. Volume 1. Psilotaceae to Platanaceae. Cambridge Univer- sity Press, Cambridge, U.K. 581 pages. Weber, W. A. 1990. Colorado flora: eastern slope. Colorado Associated University Press, Niwot, Colorado. Whittemore, A. T. 1997. Ranunculus L. Pages 88-135 in Flora of North America North of Mexico. Vol. 3. Magno- liophyta: Magnoliidae and Hamamelidae. Edited by Flora of North America Editorial Committee. Oxford Universi- ty Press, New York. Received 16 May 2005 Accepted 8 February 2007 I Assessment of Effects of an Oil Pipeline on Caribou, Rangifer tarandus granti, Use of Riparian Habitats in Arctic Alaska, 2001—2003 LYNN E. Nogv!, MATTHEW K. BUTCHER2, MATTHEW A. CRONIN! “, and BILL STREEVER* 'ENTRIX, Inc., 1600 A Street, Suite 304, Anchorage, Alaska 99501 USA; e-mail: Inoel|@entrix.com *ENTRIX, Inc., 2701 First Avenue, Suite 500, Seattle, Washington 98121 USA 4Present address: University of Alaska Fairbanks, Palmer Research Center, 533 East Fireweed Avenue, Palmer, Alaska 99645 USA 4BP Exploration (Alaska) Inc., 900 East Benson Boulevard, Anchorage, Alaska 99519 USA Noel, Lynn E., Matthew K. Butcher, Matthew A. Cronin, and Bill Streever. 2006. Assessment of effects of an oil pipeline on Caribou, Rangifer tarandus granti, use of riparian habitats in arctic Alaska, 2001-2003. Canadian Field-Naturalist 120(3): 325-330. Elevated oil field pipelines may alter Caribou (Rangifer tarandus granti) movements and delay or prevent access to insect relief habitat. In an attempt to determine if the 40-km elevated Badami pipeline in northern Alaska changed Caribou use of riparian habitats at the three river crossings where the pipeline is buried, we quantified Caribou habitat use at all three crossings using time-lapse video cameras and aerial distribution surveys over three summers. We compared habitat use, behavior and duration of observations among pipeline and non-pipeline sites. We used a block experimental design with cameras at four sites at the three river crossings to evaluate differences in numbers of Caribou per day at pipeline and non-pipeline sites. At each crossing, four cameras were positioned, with one pair of cameras next to the pipeline (pipeline sites) and one pair of cameras |.8-3.2 km upstream from the pipeline (non-pipeline sites); where cameras monitored the river bank and channel (river habitat) and the tundra within about 200 m of the river (tundra habitat). Peak numbers of Caribou per day occurred during early July 2003 and mid-July 2001 and 2002. Large numbers of Caribou recorded north of the pipeline during aerial surveys did not usually correspond with increased number per day recorded by cameras suggesting Caribou probably also crossed the pipeline outside of the riparian areas. We assessed local changes in riparian habitat use by comparing the numbers of Caribou per day in river and tundra habitats at pipeline and non-pipeline sites and found no difference. We assessed regional changes in riparian habitat use by comparing numbers of Caribou per day at pipeline sites and at non-pipeline sites and found no difference. Caribou groups spent an average of | minute longer at tundra pipeline sites and groups spent 30 seconds longer feeding and trotting at pipeline sites, but these differences were not significant. Key Words: Caribou, Rangifer tarandus granti, aerial survey, insects, Badami pipeline riparian habitat, summer, time-lapse video, Alaska. In northern Alaska, Caribou (Rangifer tarandus granti) regularly use insect relief habitats during the summer (Cameron and Whitten 1979; Murphy and Curatolo 1987; Walsh et al. 1992; Pollard et al. 1996a, b: Young and McCabe 1998). This frequently involves north-south movements from inland feeding habitats to coastal insect relief habitats (Cameron and Whitten 1979; Pollard et al. 1996b). Oil transport pipelines tra- verse open tundra and riparian corridors, often in an east- west direction, so there is the potential for obstruction of movements to and from insect relief habitats (Nation- al Research Council 2003). Previous work has shown that pipelines associated with roads can have an impact Caribou movements (Murphy and Lawhead 2000); however, impacts from pipelines not associated with roads and elevated = 1.5 m above the tundra, a height designed as a mitigation to allow Caribou passage (Cronin et al. 1994"), have not been documented. One such pipeline on Alaska’s north slope runs 40 km from the Badami oil production site to the oil fields at Prudhoe Bay. Most of the Badami pipeline is elevated about 1.5 m above the tundra, but it is buried under- neath the three rivers it crosses (Figure | and 2). Cari- bou are common in this area during the summer, and aerial surveys document animals on the north and south sides of the pipeline, indicating that they cross the pipe- line (Jensen and Noel 2002°: Jensen et al. 2003°; Noel and Cunningham 2003"). However, the elevated sections of this pipeline could alter Caribou movement and delay access to coastal habitats (National Research Council 2003). If this occurs, Caribou should be more likely to cross the pipeline in the riparian zones (including the river and adjacent tundra) where the pipeline is buried. We recorded Caribou by using time-lapse video and aerial distribution surveys over three years to assess whether Caribou use of riparian habitats was influ- enced by the pipeline. Methods Study Area A 40-km elevated pipeline extends from the Badami facility (70°9'2.71"N, 147°1'25.93"W) across undevel- oped terrain to the Endicott pipeline near the Prudhoe Bay oil field (70°15'18.25"N, 148°1'32.75"W, Figure 323 Vol. 120 10 —— ; ? = jap Area ---- Pipelines Led — Roads i Bl Gravel pads and mines (5) Aerial Survey Area THE CANADIAN FIELD-NATURALIST FIGURE 1. Study area, North Slope, Alaska. Arrows indicate study sites. 1). The Badami pipeline is elevated = 1.5 m above the tundra surface for > 99% of its length. This pipeline is buried beneath the East Channel of the Sagavanirk- tok, Shaviovik, and Kadleroshilik rivers. At the Saga- vanirktok and Shaviovik rivers, the pipeline is buried for about 1000 m; at the Kadleroshilik River the pipeline is buried for about 400 m. The Sagavanirktok River crossing is split into two channels by a large vegetat- ed river bar; requiring two sites to be monitored. The tundra between these river channels is a gently rolling thaw-lake plain landscape with elevation rises of 6 to 8 m (Walker and Acevedo 1987). Time-Lapse Videography We used time-lapse video camera assemblies to monitor Caribou movements between 19 June and 26 August in 2001, 2002, and 2003. Each camera assem- bly consisted of aGYYR™ TLC1800-DC time-lapse videocassette recorder and a Panasonic™ WV-CL 322 color CCTV digital camera equipped with a Compu- tar™ APC auto-iris 8.5 mm semi wide-angle lens (Pol- lard and Noel 1994"; Noel et al. 1998). Each assembly was powered by four 12-volt, 80-amp sealed lead acid batteries, charged by four Solarex™ SX-56 photovolta- ic panels. The video recorder, camera, and batteries were housed in insulated aluminum casings to protect them from weather and animals. The time-lapse video recorders were set to record at 34 second intervals. Markers were placed 100 m from each camera to assist kilometers Badami Pipeline y a MIKKELSEN BAY BADAMI in determining visibility and to standardize the area sampled. Cameras had a 72° view angle resulting in a 7260-m*? field of view out to 100 m (i.e., the sampling area). Videotapes were changed at 10—13 day intervals. The cameras were arranged as follows at four river crossings (Figure 1): (1) the east side of the buried East Channel Sagavanirktok crossing (E Sag), (2) the west side of the buried East Channel Sagavanirktok cross- ing (W Sag), (3) the east side of the Shaviovik River crossing (Shav), and (4) the east side of the Kadlero- shilik River crossing (Kad). At each crossing, four cam- eras were positioned, with one pair of cameras next to the pipeline (pipeline sites) and one pair of cameras 1.8-3.2 km upstream from the pipeline (non-pipeline sites) (Figure 2). One of each pair of cameras moni- tored the river bank and channel (river habitat), while the other monitored the tundra within about 200 m of the river (tundra habitat). Non-pipeline sites with river channel width and con- figuration and tundra habitats similar to those of the corresponding pipeline sites were selected to reduce variability. This arrangement allowed us to collect Cari- bou data in four settings at each river crossing (Fig- ure 2): (1) along the pipeline corridor where the pipe- line was buried under the river (buried pipeline site/river habitat), (2) along the pipeline corridor where the pipe- line was elevated above the tundra (elevated pipeline site/tundra habitat), (3) 1.8-3.2 km upstream from the 2006 pipeline at the river’s edge (non-pipeline site/river habi- tat), and (4) upstream from the pipeline on the tundra (non-pipeline site/tundra habitat). Videotapes were viewed and the number of Caribou groups seen each day by each of the cameras was re- corded. Caribou groups were defined by lapses of 15- 30 seconds of tape with no Caribou observations. For each group observed within 100 m of the camera, data for the number of individuals, sex/age category, pre- dominant behavior, and direction of movement were recorded. Caribou group behaviors included: feeding, resting (laying), standing, walking, trotting, running, and swimming. Aerial Surveys Eleven systematic aerial strip-transect surveys were completed from a Cessna 206 fixed-wing aircraft (Caughley 1977) to document the number of Caribou within a 1043 km? area surrounding the Badami pipe- line between 25 June and | August 2001-2003 (Fig- ure 1) (Jensen and Noel 2002*; Jensen et al. 2003*; Noel and Cunningham 2003*). Transect centerlines were spaced at 1.6-km intervals, oriented north-south, and centered on township and section lines from | : 63360 scale U.S. Geological Survey (USGS) topogra- phic maps. Transects were flown at 90 m altitude and 130-180 km/h. Two observers, each searching an 800- m wide area on their side of the transect centerline, provided 100% coverage of the survey area. Aircraft wing struts were marked to enable visual control of transect strip width and estimation of distance between Caribou groups and the survey aircraft (Pennycuick and Western 1972). Global positioning system (GPS) receivers were used for navigating the aircraft along transects and for estimating the location of the aircraft when animals were observed. The locations of ani- mals were recorded using a GPS receiver linked to a notebook computer. For each sighting, species, group size, group composition, and perpendicular distance from the aircraft were recorded. Coordinates of animal sightings were later calculated using the visual esti- mates of distance from the aircraft to offset the GPS aircraft positions. Geographic Information System (GIS) software was used to complete geographic summaries. Weather Data An automated weather station was established near the East Channel Sagavanirktok River. Temperature (T100 probe with radiation shield) and wind (Gill 3- cup anemometer, Gill low threshold vane) sensors trans- mitted readings at 5-min intervals to a data recorder which averaged and stored values as mean hourly wind speed and mean hourly air temperature data (Dryden R2® data logger, Anchorage, Alaska). Data Analysis The cameras were arranged to create a complete block experimental design (Steel and Torrie 1980). Each river location (E Sag, W Sag, Kad, or Shav) was a complete block containing all four settings: buried NOEL ET AL: EFFECTS OF OIL PIPELINE ON CARIBOL 325 FIGURE 2. Top: An elevated pipeline on Alaska’s North Slope. Bottom: Schematic showing the positions of four cameras relative to a river and the Badami Pipeline. As shown, the pipeline is buried at river crossings. (Draw- ing by Daniel King not to scale.) pipeline site/river habitat, elevated pipeline site/tundra habitat, non-pipeline site/river habitat, and non-pipe- line site/tundra habitat, resulting in two cameras at the pipeline site and two cameras at the non-pipeline site (Figure 2). The potential for the pipeline to change Caribou habitat use was assessed by comparing mean numbers of Caribou per day (averaged for each year) among the four settings, and by evaluating the direc- tion, duration and behavior of Caribou observed by the time-lapse cameras. We summarize the aerial survey data to compare the numbers of Caribou recorded us- ing time-lapse cameras to the numbers of Caribou with- in the survey area, north of the pipeline and within ripar- ian habitats. Riparian habitats were defined as a 200 m area surrounding the Sagavanirktok, Kadleroshilik, and Shaviovik rivers (Figure 1). Observation duration, direction of movement, behavior, temperature, and wind speed were compared among habitats and with and without the pipeline using a general linear model and analysis of variance (ANOVA). The block design allowed for the evaluation of local (within the riparian zone) and regional changes in Caribou use of riparian habitats. At a local scale, if Caribou changed habitat use due to the elevated pipe- line within the riparian zone (i.e., within 200 m of the river bank), we would expect more Caribou to use river habitat at the pipeline sites compared to river habitat at the non-pipeline sites, as Caribou would avoid the elevated portion of the pipeline by crossing at the river where the pipeline was buried. At a regional scale, if Caribou moved toward the riparian corridors along the pipeline corridor, we would expect to see more Caribou at the pipeline sites (both river and tundra habi- tats) than at the comparable non-pipeline sites (both river and tundra habitats), as blockage of direct north- south movements by the elevated pipeline would lead to east-west Caribou movements along its length. We used complete block ANOVA to assess local and regional changes in Caribou use of riparian habitats. Because of the lack of independence between counts of Caribou on consecutive days, we averaged the num- ber of Caribou per day across each year, and used this mean as the response variable, giving an overall sam- ple size of n=47. The ANOVA model was: Caribou per day = Constant + Location + Year + Pipeline + Habitat + (Pipeline x Habitat), where: Constant = overall mean Caribou per day Location = a block for each river crossing (W Sag, E Sag, Kad, or Shav), Year = each year of the study (2001, 2002, and 2003) Pipeline = pipeline site or non-pipeline site, and Habitat = river or tundra. All statistical analyses were conducted with SYS- TAT® Version 10.2 (SYSTAT® Software Inc., Rich- mond, California). Results and Discussion Peak numbers of Caribou per day were recorded dur- ing early July in 2003 and mid July in 2001 and 2002 (Figure 3). Peaks in mean daily temperature coincided with increased numbers of Caribou per day recorded by cameras during late June and early July, but trends were not consistent. During the 11 aerial surveys be- tween 25 June and | August 2001-2003, 36% of the Caribou were distributed north of the pipeline within 13% of survey area and 17% of the Caribou were with- in riparian habitats representing 9% of survey area (Table 1). Generally, when few Caribou were record- ed within the survey area, no Caribou were recorded by the time-lapse cameras (Table 1, Figure 3). The sex-age distribution of Caribou occuring within the survey area was dominated by adult cows (61%-— 70%) based on the aerial survey data (Table 1). These sex-age distributions are estimates and likely under- represent calves based on the results of fall composi- tion surveys for this herd of 28% bulls, 42% cows and THE CANADIAN FIELD-NATURALIST Vol. 120 30% calves (Lenart 2003*). Sex-age distributions based on time-lapsed video were more similar to fall com- position results (Lenart 2003*), but nearly half of the individuals were unclassified (Table 1). Mean annual Caribou per day by camera ranged from 0.3—20.4 across habitats, pipeline configurations, river locations, and years (Table 2). Least squares means from the ANOVA show that Caribou were more abun- dant in tundra habitats (mean = 8.9 Caribou per day) than in river habitats (mean = 3.3 Caribou per day) (Table 3). This difference between habitats was the only significant factor (P = 0.02, Table 4). If local habitat use changed because of the presence of the elevated pipeline, more Caribou would have been seen in river habitat than in tundra habitat at the pipeline sites. This local effect was reflected in the ANOVA Habitat x Pipeline interaction term, which was not significant (P = 0.64, Table 4). The difference between mean Caribou per day in tundra and river habitats (7.6 and 3.2 Caribou per day, respectively) was smaller for pipeline sites than for non-pipeline sites (10.4 and 3.5 Caribou per day, respectively). Because we detected no effect on numbers of Cari- bou per day, we evaluated the duration that Caribou were recorded at pipeline and non-pipeline sites on the tundra (Table 5). Blockage of northward or southward movements across the pipeline corridor could result in delays at pipeline sites, which would be reflected in duration and behavior at these sites. Groups moving north appeared to spend an average of one minute longer at pipeline sites in tundra habitats, while groups mov- ing south spent about two minutes longer at non-pipe- line sites (Table 5). A general linear model was used to test whether the presence of the pipeline affected observation duration for Caribou moving northward potentially seeking refuge from insects. Neither the presence of the pipeline nor the explanatory variables habitat (river versus tundra), temperature, and wind speed (atmospheric conditions may have an effect on the severity of insect harrassment) had statistically significant (P = 0.05) effects on the observation dura- tion for Caribou moving northward. Duration of group behaviors indicated that Caribou spent about 30 sec- onds longer feeding and trotting on the tundra at pipe- line sites than non-pipeline sites (Table 6). Separate ANOVAs were applied to the duration of feeding, trot- ting, and walking activities in different habitats (river and tundra) and in the presence or absence of the pipe- line. For trotting and walking, neither habitat nor pipe- line or their interaction were significant explanatory variables (P > 0.2). For feeding Caribou, the duration within tundra (mean = 00:01:09) was significantly longer than within river (mean = 00:06:35, P = 0.002). If the pipeline caused regional changes in Caribou riparian habitat use, we would expect to see more Cari- bou at the pipeline sites than at the non-pipeline sites. However, the difference between mean Caribou per day at the pipeline and non-pipeline sites (5.4 and 6.8 Caribou per day, respectively, Table 3) was not signifi- 2006 NOEL ET AL: EFFECTS OF OIL PIPELINE ON CARIBOI 327 - Caribou per day 3 @ North of Pipeine £ : 5 Ps wt fh rr r) Tas 2 3 ; = oa =} z Soe Sue ae ols S = = & a oS BS = — = — = — —N N —N N N N —N N 1200 - 18 16 Cag ve & 800 12 £ oe 10 8 2 400 6 § ® E ra 3 4 = 200 > 0 0 1200 s 1000 Fe 2 = 800 2 o S be 600 ® 2 a don = Ee e = = 200+ 0 - nt RBs ses 8s 2228 8 8 8 i — - — os — N N N N N N N N Day of Year FiGureE 3. Comparison of numbers of Caribou recorded per day for all time-lapse cameras, numbers of Caribou recorded in riparian habitats north of the Badami pipeline corridor during aerial surveys (Figure 1), and mean daily temperature (°C), 19 June (day 170) 26 August (day 238) 2001-2003, North Slope, Alaska. 328 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Summary of Caribou sex-age classes, numbers, and distribution within the aerial survey area and observed in riparian areas at pipeline and non-pipeline sites using time-lapse video 19 June—26 August 2001—2003, North Slope, Alaska. For bulls, cows, and calves % is per cent of classified. Unclassified % is per cent of total. Total Total Percent Bulls Cows Calves Unclassified Caribou Groups _ of Total Aerial Surveys Survey Area (1043 km?) 2846 12% 16001 65% 5775 23% 5798 19% 30420 7715 North of Pipeline (136 km?) 1479 14% 7492 70% 1807 17% 287 3% 11065 134 36% Riparian Zone (98 km?) 811 21% 2392 61% 722 18% 1222 24% 5147 111 17% Time-Lapse Video All Sites (1293 days) 1352 36% 1352 36% 1005 27% 4165 53% 7874 635 Pipeline Sites (660 days) 664 37% 659 37% 457 26% 1816 51% 3596 352 46% Non-Pipeline Sites (633 days) 688 36% 693 36% 548 28% 2349 55% 4278 283 54% TABLE 2. Total Caribou, days of camera operation, and mean Caribou per day by camera with standard deviation (SD) for all three years pooled, 19 June—26 August 2001—2003, North Slope, Alaska. Camera Site Grand Total Habitat Tundra River Tundra River River Tundra River Tundra River Tundra River Tundra River Tundra River Tundra Pipeline Elevated Buried Non-pipeline Non-pipeline Buried Elevated Non-pipeline Non-pipeline Buried Elevated Non-pipeline Non-pipeline Buried Elevated Non-pipeline Non-pipeline Total Days of camera Caribou per Caribou operation day (SD) 562 89.88 5.8 (35.32) DD 87.62 0.2 (1.47) 488 86.88 Sli@5e70) 8 88.67 0.1 (0.82) 243 78.60 2.8 (14.56) 1122 77.88 12.9 (63.03) 320 81.23 3.6 (19.21) 596 77.78 6.7 (29.63) 151 79.67 1.6 (7.19) 234 88.42 2.5 (13.54) 591 84.98 6.5 (36.29) 1222 59.81 19.1 (77.81) 629 80.20 7.1 (27.53) 633 78.00 13 (C2) DD 76.32 2.6 (14.29) 828 76.93 9.5 (48.67) 7874 1292.87 TABLE 3. Caribou per day by pipeline and habitat with sites pooled (buried pipeline site/river habitat, elevated pipeline site/tundra habitat, non-pipeline site/river habitat, non-pipeline site/tundra habitat), 19 June—26 August 2001-2003, North Slope, Alaska. Habitat Pipeline River Tundra Total Pipeline 3.2 7.6 SD) Non-pipeline 3.5 10.4 6.8 Total Shes) 8.9 6.1 cant (P = 0.57, Table 4). Peaks in the number of Cari- bou north of the pipeline in 2002 and 2003 did not coincide with large numbers of Caribou recorded at cameras, suggesting that Caribou crossed the pipeline outside of riparian areas (Figure 3). The results of this study support the conclusion that pipelines elevated =1.5 m above the tundra did not cause changes in Caribou riparian habitat use at buried river crossings or delay north-south movements to and from coastal insect relief habitats (Cronin et al. 1994*, Murphy and Lawhead 2000). TABLE 4. Results of ANOVA for Caribou per day along three rivers crossed by the Badami pipeline during 19 June—26 August 2001-2003, North Slope, Alaska. Source Sum-of-Squares df Mean-Square F-ratio IP Pipeline (Present, Absent) 22.08 1.00 22.08 0.33 0.57 Habitat (River, Tundra) 415.86 1.00 415.86 6.29 0.02 Location (i.e., Block) (E Sag, W Sag, Kad, Shav) 170.90 3.00 56.97 0.86 0.47 Year (2001, 2002, 2003) 29.50 2.00 14.75 0.22 0.80 Interaction (Pipeline x Habitat) 14.40 1.00 14.40 0.22 0.64 Error 2512.10 38.00 66.11 329 ARIBOI FFECTS OF OIL PIPELINE ON ( i y) NOEL ET AL: 2006 PTO E100 LO:CET Il 1c:L0:0 PE:TO:0 60:1S:7I OOF 8F:90:0 = ST :90:0 OM:6P'Sl CSI [BIOL IV 9110/0 FO:10°0 OF-60:1 g9 ChC0:0 POCO:;0 ONC L“P CCl ££:90:0 80:90:0 C0:80:9 09 [PRIOL, 6F:00°0 SF:00°0 chEt-0 cr PO:€0:0 €1:c0'0 8OS:S0'E = #8 €v:90':0 = 8€:90:0 CO'BS:S = PS vApUNnL OF 10:0 Pes 10°0 8S:St-0 tC Se:10:0 SPI0‘0 62:90'l 8€ lelO:0 OF 10:0 (01010) 0 ee) TOATY ourjadid-uoyy TS 10:0 81100 LETT 9¢ 91:60:0 $S:20:0 =OS:8E:8 = BI 65:90:06 1:90:0 wIt'6 C6 [PIOL IS:10:0 T1100 I1:L¢:0 I€ Oe-T0:0 00:c0:0 OES: LII €1:40:0 — ZO:L0:0 LI:€c:6 = 08 eipuny 98-100 ST: 10:0 9eSe:0 ST €C:S1:0 Iv:70:0 9PSPbr 19 cCO:10:0 TE 10:0 Le81:0 =I TOATY ourpodid as uray [RIOL u ds urd [RIOL u ds urd [PIOL u SsuMoly SULTRY SUIpoa] ‘(W) SUOHRAJasgo Jo Jaquinu (GS) SUONeIAap prepurjs UyIM pajudsoid sonyea uray “eyseyy ‘edo[sg YON ‘EQOT-1OOT IsNsny 9Z—-ouNL 6] BuULINp 3uNI0N 10 SUPE “Surpaay sdnoss 10J says ourjedid-uou pue ourjadid ye syeyqey JOA pue espuny UTYIIM sUONRAIJASgO NOLIeD OaptA asdr]-auIN 10J SUONeINp UPd pure UONRANp [PIO], 9 AIAV], (LOTTE (CH) OL [e=70:0 66700 LP Li-0l 617 (91) 87 (tb) 06 £¢:€0:0 91°70:0 PSPS: LSI [RIOL, IV (SDE (6) 89 €1:90°0 [S:€0:0 Cr LE: 98 (C1) 97 (66) 8 8¢:70:0 €S: 10:0 61:SS:7 £6 [FIOL, (6DTE (QE) E9 8S:90:0 Le-F0:0 OF ESP c9 (QI) LZ (6'€) 0'8 CO:€0:0 T1700 PE:607 6S eipuny, (TUD9T (CH) £8 TS: 10°0 8F: 10:0 9¢:LE:0 I7 (SaDIGEG (6'¢) 16 6£:10:0 17: 10:0 CrSt:0 VE Joary — autfadid-uoyy (Dee (CH EL F700 60:70°0 COOP eel (LD OE (O'S) 96 e¢:P0:0 8:70:0 CE‘6S:7 v9 [BIO], (9DOE (OF) SL £0°€0°0 61°70:0 CTES:E 101 (31) 67 (6p) 88 60:S0:0 L700 ELSE:7 cr vApUuny, (6D Tr (Cr) 78 81:10:0 6£:10:0 LE:TS:0 CE (PDOE (Cr) SII F0:70:0 LI10:0 CC'VT0 61 JOATY autjadid PULA dway ds ural [BIOL u (GS) pul (qs) dway ds uBall [FIOL u yinos YON uonoa1g ‘(U) SUOTJBAIOSGO JO Joquinu (qs) SUONPLAANP PIepURIS YIM pojuasaid sanyea uray “eyse[y ‘adojs YON ‘EQOZ-1O0T IsnsNy 9Z—-oUNL 6] BuLINp yWNOs pue YOU SurAOUT sdnos Joy sayts sutfadid-uou pure ourjadid ye s}e} -IQPY JOALI PUP BIPUN) UIYIIA (PUOdas Jad siajaW) paads pum usu pure (D,) cMPIadua) URAL SUONPAIASGO NOgLIeD OaplA asde]-suI} 10} UONINP UvdLU PUe UOT}eAINP [eIOL “¢ ATV], 330 Documents Cited [marked * in text citations] Cronin, M. A., W. B. Ballard, J. Truett, and R. Pollard. 1994. Mitigation of the effects of oil field development and transportation corridors on caribou. Final Report to the Alaska Caribou Steering Committee by LGL Alaska Research Associates, Inc., Anchorage, Alaska. Jensen, P. G., and L. E. Noel. 2002. Caribou distribution in the range of the Central Arctic Herd. Part A: Aerial surveys in the Milne Point Unit, Prudhoe Bay Oilfield, Badami, and Bullen Point to Staines River study areas, summer 2001. Chapter 2A in Arctic Coastal Plain caribou distribution, summer 2001. Edited by M. A. Cronin. Unpublished report for BP Exploration (Alaska) Inc. by LGL Alaska Research Associates, Inc., Anchorage, Alaska. Pages 2-1-86. Jensen, P. G., L. E. Noel, and W. B. Ballard. 2003. Caribou distribution in the Badami and Bullen Point to Staines River study areas, Alaska, summer 2002. Chapter | in Caribou distribution in the range of the Central Arctic Herd, summer 2002. Report for BP Exploration (Alaska) Inc. by LGL Alaska Research Associates, Inc. (LGL Report P662) 42 pages plus appendices. Lenart, E. A. 2003. Unit 26A and B Caribou management report. Pages 304-326 in Caribou management report of survey and inventory activities 1 July 2000-30 June 2002. Edited by C. Healy. Alaska Department of Fish and Game, Juneau, Alaska. Noel, L. E., and EK. E. Cunningham. 2003. Caribou distribu- tion in the Badami and Bullen Point to Staines River sur- vey areas, Alaska, Summer 2003. Report for BP Explo- ration (Alaska) Inc. by ENTRIX, Inc., Anchorage, Alaska. 58 pages. Pollard, R. H., and L. E. Noel. 1994. Caribou distribution and parasitic insect abundance in the Prudhoe Bay oil field, summer 1993. Report to BP Exploration (Alaska) Inc. by LGL Alaska Research Associates, Inc., Anchorage, Alaska. 70 pages. Literature Cited Caughley, G. 1977. Sampling in aerial survey. Journal of Wildlife Management 41: 605-615. Cameron, R. D., and K. R. Whitten. 1979. Seasonal move- ments and sexual segregation of caribou determined by aerial survey. Journal of Wildlife Management 43: 626- 633. Murphy, S. M., and J. A. Curatolo. 1987. Activity budgets and movement rates of caribou encountering pipelines, THE CANADIAN FIELD-NATURALIST Vol. 120 roads, and traffic in northern Alaska. Canadian Journal of Zoology 65: 2483-2490. Murphy, S. M., and B. E. Lawhead. 2000. Caribou. Pages 59-84 in The Natural History of an Arctic Oil Field: Dev- elopment and the Biota. Edited by J. C. Truett and S.R. Johnson. Academic Press, San Diego, California. National Research Council. 2003. Cumulative environmen- tal effects of oil and gas activities on Alaska’s North Slope. The National Academies Press. Washington, D.C. 452 pages. Noel, L. E., R. H. Pollard, W. B. Ballard, and M. A. Cronin. 1998. Activity and use of active gravel pads and tundra by Caribou, Rangifer tarandus granti, within the Prudhoe Bay oil field, Alaska. Canadian Field-Naturalist 112: 400-409. Pennycuick, C. J., and D. Western. 1972. An investigation of some sources of bias in aerial transect sampling of large mammal populations. East African Wildlife Journal 10: 175-191. Pollard, R. H., W. B. Ballard, L. E. Noel, and M. A. Cronin. 1996a. Parasitic insect abundance and microclimate of gravel pads and tundra within the Prudhoe Bay oil field, Alaska, in relation to use by Caribou, Rangifer tarandus granti. Canadian Field-Naturalist 110: 649-658. Pollard, R. H., W. B. Ballard, L. E. Noel, and M. A. Cronin. 1996b. Summer distribution of Caribou, Rangifer taran- dus granti, in the area of the Prudhoe Bay oil field, Alas- ka, 1990-1994. Canadian Field-Naturalist 110: 659-674. Steel, R. G. D., and J. H. Torrie. 1980. Principles and pro- cedures of statistics: A biometrical approach. McGraw- Hill Book Company, New York. 674 pages. Walker, D. A., and W. Acevedo. 1987. Vegetation and a Landsat-derived land cover map of the Beechey Point quadrangle, Arctic Coastal Plain, Alaska. CRREL Report 87-5. U.S. Army Corps of Engineers, Cold Regions Re- search and Engineering Laboratory, Hanover, New Hamp- shire, USA. Walsh, N. E., S. G. Fancy, T. R. McCabe, and L. F. Pank. 1992. Habitat use by the Porcupine Caribou Herd during predicted insect harassment. Journal of Wildlife Manage- ment 56: 465-473. Young Jr., D. D., and T. R. McCabe. 1998. Grizzly bears and calving caribou: What is the relation with river corri- dors? Journal of Wildlife Management 62: 255-261. Received 17 August 2005 Accepted 19 April 2007 First Records of the Southern Red-backed Vole, Myodes gapperi, in the Yukon THOMAS S. JUNG!, AMY M. RUNCK?2, DAvip W. NAGORSEN?, BRIAN G. SLOUGH*, and Topp PoweLL>” 'Yukon Department of Environment, Fish and Wildlife Branch, P.O. Box 2703, Whitehorse, Yukon Y!1A 2C6 Canada; e-mail: thomas.jung@ gov.yk.ca Department of Biological Sciences, Idaho State University, 650 Memorial Drive, Pocatello, Idaho 83209-8007 USA 4Mammialia Biological Consulting, 4268 Metchosin Road, Victoria, British Columbia V9C 324 Canada 435 Cronkhite Road, Whitehorse, Yukon Y1A 5S9 Canada ‘Yukon Department of Environment, Fish and Wildlife Branch, P.O. Box 194, Watson Lake, Yukon YOA 1CO Canada °Current address: Alberta Department of Sustainable Resource Development, Fish and Wildlife Division, P.O. Box 9915, Fort McMurray, Alberta T9H 2K4 Canada Jung, Thomas S., Amy M. Runck, David W. Nagorsen, Brian G. Slough, and Todd Powell. 2006. First records of the Southern Red-backed Vole, Myodes gapperi, in the Yukon. Canadian Field-Naturalist 120(3): 331-334. Twenty Southern Red-backed Voles, Myodes gapperi, were collected in July 2004 in the LaBiche River valley of southeast- ern Yukon. Specimens were identified using morphological characteristics and analysis of cytochrome b gene sequences. These are the first records of this species in the Yukon. No Northern Red-backed Voles, M. rutilus, were collected and it is not known whether the two species are sympatric or parapatric in the Yukon. Further survey work is needed in southeastern Yukon to better delineate the extent of the northwestern range of this species and the extent, if any, of introgression with M. rutilus. Key Words: Southern Red-backed Vole, Myodes gapperi, distribution, Yukon. The Southern Red-backed Vole (Myodes [formerly Clethrionomys see Musser et al. 2005] gapperi) is broadly distributed across the boreal, montane, Pacific coastal, and other conifer-dominated forests of North America (Banfield 1974; Merritt 1981; Batzli 1999). Youngman (1975) did not report M. gapperi from the Yukon, in part, because he was of the opinion that M. gapperi and the Northern Red-backed Vole (M. rutilus) were conspecific. Cook et al. (2004), however, provided molecular data that reinforced the separate status of M. gapperi and M. rutilus. North American species of Myodes, including M. gapperi and M. rutilus, are cryp- tic and field identification is highly problematic where congeners come into contact (Merritt 1981; McPhee 1984; Runck 2001, Nagorsen 2002, 2005; Cook et al. 2004). Therefore, we considered that M. gapperi may have been overlooked in the Yukon due to the difficulty in distinguishing specimens from M. rutilus in the field, and a general paucity of mammalian diversity surveys in most of the Yukon. Based on the distribution of M. gapperi in adjacent northeastern British Columbia and southwestern Northwest Territories (Banfield 1974; Nagorsen 2005) we suspected the greatest likelihood of finding the species in the Yukon was in the Liard River Watershed. In summer 2004, we undertook a brief field survey of mammalian diversity in the bore- al forest of southeastern Yukon, partially with an aim to procuring specimens of M. gapperi. Here, we pro- vide the first records of the Southern Red-backed Vole in the Yukon. Methods On 28 July — 2 August 2004, we used pitfall and snap traps to collect shrews and rodents in the LaBiche River Valley (60.126°N, 124.064°W) of southeastern Yukon. Seven habitats were sampled: wet meadows and shrub thickets adjacent to Beaver (Castor canadensis) ponds; xeric grassy meadows; riparian old-growth White Spruce (Picea glauca) forest; lowland Black Spruce (Picea mariana) forest; subalpine old-growth Spruce-fir (Abies lasiocarpa) forest; second-growth mixedwood forest; and regenerating clearcut forest. All habitats were sampled with snap traps. We estab- lished 14 variable length traplines (ca. 100 m— 250 m) in each of the seven habitat types. Trap stations were set 10 m apart on the traplines. One Museum Special snap trap and one Victor snap trap (Woodstream, Lititz, Pennsylvannia) were set at each trapping station. Traps were baited with oats and peanut butter, run for 2-3 days and checked each morning. In addition to the snap traps, we established three traplines of metal conical pitfall traps in the wet meadow habitats, primarly to capture soricids. Pitfall traps were 25 cm in height and 15 cm in diameter at the top. Pitfall traps were also arranged in traplines with one trapline in each of the wet meadows sampled. Traps were spaced about 10 m apart, with 15 or 20 traps per trapline, and installed flush with the ground surface. Pitfall traps were not baited and they were left open for 3-5 days. Each mom- ing we checked the traps and processed the captures. 331 332 Captures were tentatively identified to species, mea- sured, aged, weighed, sexed, and examined for repro- ductive condition. Whole carcasses were frozen on site. Species were tentatively identified in the field us- ing morphological, pelage, and dental characteristics, following Nagorsen (2002) and van Zyll de Jong (1983). Because of the difficulty in distinguishing M. rutilus from M. gapperi in the field, we undertook sub- sequent laboratory investigations at Idaho State Uni- versity (Pocatello, Idaho) to confirm our field identifi- cations. All skulls were examined for fusion of the post-palatal bridge (Merritt 1981; Nagorsen 2002) and the first 600 base pairs (bp) of mitochondrial cyto- chrome b gene were examined to confirm species iden- tification. Restriction enzyme screening and sequencing of cytochrome b followed established protocols for Myodes (Runck and Cook 2005). All individuals un- derwent restriction enzyme digestion using the restric- tion enzyme ALU I to determine the presence of the restriction site AGCT which is found in the 600 bp frag- ment of the cytochrome b gene in M. rutilus but not M. gapperi (Runck 2001). Five random individuals were sequenced for the first 600 bp of the cytochrome b gene and were used in a phylogenetic reconstruction using published sequences of Myodes (Cook et al. 2004; Runck and Cook 2005), representing M. gapperi (n= 10), M. rutilus (n = 2), M. californicus (n= 1), M. glareolus (n= 1), and M. rufocanus (n = 1). Sequences for specimens not collected in LaBiche (Figure 1) were taken from GenBank. Phylogeographic reconstruction was conducted using neighbor-joining framework in PAUP b10 (Swofford 2002) using the Tamura and Nei (1993) model of sequence evolution as determined through Modeltest (Posada and Crandall 2000). Results and Discussion We captured 20 Myodes during sampling. Field identification, based on morphological (tail and pelage) characteristics, suggested that all captures were of M. gapperi. Further, 15 of the 20 specimens had complete post-palatal bridges, which is a primary diagnostic characteristic of M. gapperi (Merritt 1981; Nagorsen 2002). Two specimens had incomplete post-palatal bridges and the bridges of a further three were broken by the traps and not usable as a diagnostic character. Genetic analyses using restriction enzyme screening and sequencing of cytochrome b gene confirmed that all 20 specimens of Myodes collected in the LaBiche River Valley were M. gapperi (Figure 1). The Southern Red-backed Vole is now added to the list of mammals in the Yukon. This is the first addition- al rodent found in the Yukon since Youngman (1975). These 20 specimens were deposited at the Museum of Southwestern Biology at the University of New Mex- ico (Albuquerque, New Mexico). Capture rates were 0.28 per 100 trap nights (TN) and 1.32 per 100 TN in pitfall and snap traps, respec- THE CANADIAN FIELD-NATURALIST Vol. 120 tively. Captures of M. gapperi represented the range of sex and age classes (5 adult females, 7 adult males, 5 juvenile females, and 3 juvenile males), confirming that a breeding population was present. M. gapperi were taken in all of the habitats sampled. Species caught in association with M. gapperi in the LaBiche River Val- ley included: Meadow Voles (Microtus pennsylvani- cus, n = 14), Deer Mice (Peromyscus maniculatus, n = 13), Meadow Jumping Mice (Zapus hudsonius, n= 10), Eastern Heather Vole (Phenacomys ungava, n= 1), Masked Shrew (Sorex cinereus,n = 1), Pygmy Shrew (Sorex hoyi, n = 1), and Dusky Shrews (Sorex monticolus, n = 2). We do not know the extent of the range of the South- ern Red-backed Vole in the Yukon. They have been collected, however, near Fort Liard, Northwest Terri- tories (S. Carriére, unpublished data; Runck and Cook 2005). Specimens from near Watson Lake, Yukon (ca. 270 km W of the present study; n = 43), were also analyzed in the lab using skull and genetic characters and they were determined to be M. rutilus (T. S. Jung and A. M. Runck, unpublished data), suggesting that M. gapperi may not be widely distributed in southern Yukon. Perhaps M. gapperi is restricted in the Yukon to the extension of the Boreal Plain ecoregion in the extreme southeast. Further surveys from nearby river valleys, and subsequent DNA analysis, are needed to better document the ranges and contact zone of M. gap- peri and M. rutilus in southeastern Yukon and adjacent British Columbia and Northwest Territories. Our dis- covery of M. gapperi in extreme southeastern Yukon, along with that of the Yukon’s only records of Northern Long-eared Bats (Myotis septentrionalis; Jung et al. 2005) and other taxa (e.g., some species of neotropi- cal migrant passerines: Eckert et al. 1997), suggests that this ecoregion may be a zoogeographically unique part of the Yukon. Acknowledgments The staff of Devon Canada Corporation’s LaBiche River Gas Plant kindly went out of their way to accom- modate us and provide logistical support. Funding was provided by the Yukon Department of Environment, NatureServe Yukon, and a Northern Research Endow- ment Grant from the Northern Research Institute, Yukon College, to B. Slough. Thoughtful comments from S. Cannings, and five anonymous reviewers improved the manuscript. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. University of Toronto Press, Toronto, Ontario. 438 pages. Batzli, G. O. 1999. Northern red-backed vole, Clethrionomys rutilus. Pages 616-617 in The Smithsonian Book of North American Mammals. Edited by D. E. Wilson and S. Ruff. UBC Press, Vancouver, British Columbia. 750 pages. Cook, J. A., A. M. Runck, and M. J. Conroy. 2004. His- torical biogeography at the crossroads of the northern 2006 JUNG ET AL: SOUTHERN RED-BACKED VOLE IN THE YUKON 333 LaBiche DQ152248 Fort Liard, Northwest Territories AY952147 Swan Lake, British Columbia AY952158 85 LaBiche DQ152249 LaBiche DQ152250 LaBiche DQ152251 Fort Liard, Northwest Territories AY952146 High Latitude 98 Swan Lake, British Columbia AY952159 C. gapperi Duck Point, Alaska AY952195 Duck Point, Alaska AY952194 LaBiche DQ152252 93 Brown County, Minnesota AY952173 98 Brown County, Minnesota AY952174 ‘60 Lewis County, Washington AY952193 Lewis County, Washington AY952194 M. rufocanus Japan AY309418 M. californicus California AY309422 M. glareolus Finland AY119272 400 M. rutilus Alaska AY309426 M. rutilus Alaska AY309427 — 0.01 substitutions/site FiGurE 1. Placement of five specimens of Myodes gapperi (LaBiche DQ152248-DQ152252) captured in the LaBiche River Valley, Yukon, based a neighbor-joining analysis of cytochrome b gene sequences (600 bp) of these specimens and compared to sequences of other Myodes. GenBank accession numbers for sequences used are included for each specimen. Numbers above the branches represent bootstrap support values for those nodes. 334 continents: molecular phylogenetics of red-backed voles (Rodentia: Arvicolinae). Molecular Phylogenetics and Evolution 30: 767-777. Eckert, C. D., P. H. Sinclair, and W. A. Nixon. 1997. Breed- ing bird communities in the forests of the Liard River Valley, Yukon. Technical Report Series Number 297. Cana- dian Wildlife Service, Delta, British Columbia. Jung, T. S., B. G. Slough, D. W. Nagorsen, T. A. Dewey, and T. Powell. 2006. First records of the northern long- eared bat, Myotis septentrionalis, in the Yukon Territory. Canadian Field-Naturalist 120: 39-42. McPhee, E. C. 1984. Ethological aspects of mutual exclusion in the parapatric species of Clethrionomys gapperi and C. rutilus. Acta Zoologica Fennica 172: 71-73. Merritt, J. F. 1981. Clethrionomys gapperi. Mammalian Species 146: 1-9. Musser, G. G., and M. D. Carleton. 2005. Superfamily Muroidea. Jn Mammal Species of the world. Edited by D.E. Wilson and D. M. Reeder. John Hopkins University Press, Baltimore, Maryland. Nagorsen, D. W. 2002. An Identification Manual to the Small Mammals of British Columbia. Ministry of Sustainable Resource Management, Ministry of Water, Land and Air Protection and Royal British Columbia Museum, Victoria, British Columbia. 153 pages. Nagorsen, D. W. 2005. Rodents and Lagomorphs of British Columbia. Vol. 4: The Mammals of British Columbia. THE CANADIAN FIELD-NATURALIST Vol. 120 Royal British Columbia Museum, Victoria, British Colum- bia. 410 pages. Posada, D., and K. A. Crandall. 2000. Modeltest: testing the model of DNA substitution. Bioinformatics 14: 817- 818. Runck, A. M. 2001. Molecular and morphological perspec- tives in post-glaciation of Clethrionomys rutilus and Cleth- rionomys gapperi in southeast Alaska. M.S. thesis, Univer- sity of Alaska, Fairbanks, Alaska. 89 pages. Runck, A. M., and J. A. Cook. 2005. Post-glacial coloniza- tion of the southern red-backed vole (Clethrionomys gap- peri) in North America. Molecular Ecology 14: 1445-1456. Swofford, D. L. 2002. PAUP*. Phylogenetic analysis using parsimony, Version b10. Sinauer Associates, Sunderland, Massachusetts. Tamura, K., and M. Nei. 1993. Estimation of the number of nucleotide substitutions in the control region of mitochon- drial DNA in humans and chimpanzees. Molecular Biol- ogy and Evolution 10: 512-526. van Zyll de Jong, C. G. 1983. Handbook of Canadian Mam- mals. 1: Marsupials and Insectivores. National Museums of Canada, Ottawa, Ontario. 210 pages. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museums of Canada, Ottawa, Ontario. 192 pages. Received 12 September 2005 Accepted 4 September 2007 ee A Conservation Evaluation of Smooth Goosefoot, Chenopodium subglabrum (Chenopodiaceae), in Canada DIANA BIZECK!I ROBSON The Manitoba Museum, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2 Canada Robson, Diana Bizecki. 2006. A conservation evaluation of the Smooth Goosefoot, Chenopodium subglabrum (Chenopodi- aceae), in Canada. Canadian Field-Naturalist 120(3): 335-341. Smooth Goosefoot (Chenopodium subglabrum) is restricted to North America and reaches its northern distribution limit in Alberta, Saskatchewan and Manitoba. The habitat of Chenopodium subglabrum contains some element of active sand. It is commonly found on the stabilizing edges of active dunes as well as dune blowouts, and occasionally on bare or recently dis- turbed sand plains. Chenopodium subglabrum is a sexually reproducing annual species with seeds that may remain in the seed bank for several years, waiting for the conditions it favours for germination. Germination of this species is erratic, pos- sibly in response to moisture, making the overall population trend difficult to determine. The estimated population in Cana- da is likely between 5000 and 10000 individuals. There has been considerable loss of habitat as dunes become vegetated. The processes of dune stabilization and fire control threaten survival of this species. Key Words: Chenopodium subglabrum, Smooth Goosefoot, distribution, population size, rare, Alberta, Saskatchewan, Manitoba. Chenopodium subglabrum (S. Wats.) A. Nels., com- monly called Smooth Goosefoot, is a rare, shallow- rooted annual plant (Figure 1) found in the prairie provinces of Canada. Prior to 1990 only small num- bers of this species had been observed at 20 sites in 4 Canada (Smith and Bradley 1990*). Search efforts Jf 2 OP \ (/ ' ( Za a from the mid-1990s on resulted in the discovery of new populations in five Sand Hill regions in Saskat- chewan and Manitoba. In 2004, the highest number of individuals ever recorded in Canada was observed, suggesting that this species has a strong temporal com- ponent to its rarity. The objective of this paper is to doc- ument these recent findings and determine their con- servation implications. a Habitat Chenopodium subglabrum populations occur in the Aspen Parkland, Moist Mixed Grassland, and Mixed Grassland Ecoregions of the Prairie Ecozone (Acton et al. 1998). Within this ecological area, C. subglabrum is found in Mixed Grasslands with sandy soils. The populations in Saskatchewan and Alberta are isolated from each other as the land in between them is largely cultivated. The Manitoba populations are particularly isolated, being approximately 500 km from the near- est C. subglabrum population in Saskatchewan and about 350 km from the nearest population in North Dakota (Schmoller 2002*). This fragmentation is nat- ural since C. subglabrum does not generally grow on the silty and clayey soils that occur in between the var- 4 ious sand hills. The populations that occur along the \ South Saskatchewan River are less isolated than the ones growing in the sand hills to the south since habi- 2 tat along the river is largely undisturbed. , Chenopodium subglabrum is an early successional — Ficure 1. Illustration of Chenopodium subglabrum. Line habitat specialist growing typically in the stabilizing drawing by D. B. Robson. 335 336 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 1. Nationally and provincially rare plants associated with Chenopodium subglabrum. Associated Species Provincially rare * Bur Ragweed (Ambrosia acanthicarpa) Annual Skeleton-weed (Shinnersoseris rostrata) Hairy Prairie-clover (Dalea villosa var. villosa) Sand-dune Wheatgrass (Elymus lanceolatus Sand Nut-sedge (Cyperus schweinitziii) Ambrosia acanthicarpa Wingless Bugseed (Corispermum villosum) Shinnersoseris rostrata Ambrosia acanthicarpa Corispermum villosum Shinnersoseris rostrata Indian Rice-grass (Achnatherum hymenoides) Cyperus schweinitziti Purple Ball Cactus (Escobaria vivipara) Louisiana Broomrape (Orobanche ludoviciana) Shinnersoseris rostrata Winged Pigweed Province Sand Hill Nationally rare ! Alberta Dominion Nodding Umbrella-plant (Eriogonum cernuum) Sand-verbena (Tripterocalyx micranthus) Saskatchewan Dundurn ssp. psammophilus) Elbow Western Spiderwort (Tradescantia occidentalis) Great Eriogonum cernuum Pelican Lake Dalea villosa var. villosa Seward Elymus lanceolatus ssp. psammophilus Manitoba Routledge Sand Bluestem (Andropogon hallii) Tradescantia occidentalis Brandon Dalea villosa var. villosa (Carberry) 1According to Argus and Pryer 1990 2 According to provincial Conservation Data Centres sand at the edges of active dunes and blowouts, and along eroding, sandy river banks and coulees (Smith and Bradley 1990*; Robson 1997a; Lamont and Gerry 1998*). It is generally found on south- or west-facing actively eroding slopes, at the edge of stabilizing sand and sometimes in dune slacks. Populations tend to occur in discrete units clustered around active dunes and are typically highest in areas of finer and more compacted sand. Given that C. subglabrum was found historically along the South Saskatchewan River, it is possible that much of this species’ habitat disappeared when Gardiner Dam was built on the South Saskatche- wan River in the late 1960s, flooding the sand bars and terraces. Dynamic factors such as grazing, erosion and fire may aid in destabilizing sand, resulting in more habitat for C. subglabrum (Robson 1999). Commonly associated species include the following: Indian Rice-grass (Achnatherum hymenoides), Sand Grass (Calamovilfa longifolia), Narrow-leaved Goose- foot (Chenopodium pratericola), Canada Wild Rye (Elymus canadensis), Northern Wheat-grass (Elymus lanceolatus ssp. lanceolatus), Prairie Sunflower (Heli- anthus petiolaris ssp. petiolaris), Hairy Golden-aster (Heterotheca villosa var. villosa), Skeleton-weed (Ly- godesmia juncea) and Lance-leaved Psoralea (Psora- (Cycloloma atriplicifolium) Achnatherum hymenoides lidium lanceolatum) (Smith and Bradley 1990*; Rob- son 1997a). Shrub cover often includes: Common Bearberry (Arctostaphylos uva-ursi), Hoary Sagebrush (Artemisia cana ssp. cana), Creeping Juniper (Junipe- rus horizontalis) and Sandbar Willow (Salix exigua ssp. interior) (Smith and Bradley 1990*; Robson 1997a). Chenopodium subglabrum is also associated with six nationally rare and eight provincially rare plants in various locations (Table 1). Chenopodium subglabrum is an annual plant with bisexual flowers (Clements and Mosyakin 2004). Flowering occurs from June to August and seed pro- duction in August and September (Wallis and Wershler 1988). Although no studies on C. subglabrum pollina- tion have been performed, other species in this genus are wind- or self-pollinated (Johnson and Ward 1993; Royer and Dickinson, 1999). The distances that C. sub- glabrum pollen travels is unknown. Since the wind- pollinated C. pratericola often occurs in the same habi- tat with C. subglabrum, hybridization may be possible, although it has not been observed (Bassett and Cromp- ton 1982). The fruits and seeds of C. subglabrum lack structures that would aid in wind (1.e., hairs) or animal (i.e., burs or fleshy fruits) dispersal although the seeds may ad- 2006 here to wet fur. The seeds likely fall close to the par- ent plant and may be buried by shifting sand, or travel short distances in the winter over snow crust. This means that the main exchange of genetic material be- tween populations would occur via pollination. Lam- ont and Gerry (1998*) asserted that dry weather could limit its germination. The dramatic increase in the num- ber of individuals observed at most sites in Saskatche- wan in 2004 coupled with the observation that late sum- mer was unusually wet and cool suggests that maximum seed germination occurs under moist conditions. The seeds can likely remain dormant for several years (Rob- son 1999). This dormancy can last at least eight years; at the Beaver Creek site plants were observed in 1996 and 2004 but not in the years in between. There may be dormant seeds present in stabilized areas that would germinate under appropriate climatic conditions if the area were denuded of its vegetative cover. No attempts to propagate the species for seed pro- duction have been attempted. If seeds were harvested from the wild and grown in a greenhouse the seeds from those plants could be introduced back into the wild. However, given the species’ sensitivity to climate conditions, the seeds may not germinate in the wild the following year making assessment of the introduction success difficult to ascertain. Chenopodium subglabrum is both a halophyte (i.e., salt-loving plant) and a xerophyte (1.e., dry-lov- ing plant) making it a stress-tolerant species (Grime 1979). Plants in the Chenopodiaceae are generally non-mycorrhizal (Mukerji et al. 2000). Since C. sub- glabrum grows in areas of active sand where mycor- rhiza are usually not abundant, it is probably normally non-mycorrhizal. At several sites in Alberta and Saskatchewan there were specimens of C. subglabrum that had been grazed. Grazing may have been by cattle or wildlife. Some grazed plants produced side shoots to compensate for stem loss. A report on Ord’s Kangaroo Rat (a nation- ally rare mammal) ecology noted that there were seeds of C. subglabrum found in their pouches and food caches so seed predation occurs (Lamont and Gerry 1998*). The growth of several plants observed ap- peared to be inhibited by an unidentified purplish fun- gus infecting the leaves. The Grassland Natural Region is one of the most threatened natural regions in Canada. Over two-thirds of the Mixed Grassland has been lost to cultivation or other development (Wallis 1987). Although some sandy plains have been cultivated, the hilliest areas have not been. The main limiting factors affecting Chenopodi- um subglabrum are its natural narrow preference for unstabilized sites within dune fields and loss of natu- ral habitat through succession. Large areas of once active sand have become stabilized over the last fifty years. The active sand surface of some dunes in the Milk River Sand Dunes of Alberta has been reduced by 50% to 75% over 40 years (Wallis 1987). In the Middle ROBSON: SMOOTH GOOSEFOOT IN CANADA 337 Sand Hills of Alberta, 90% of the sand that was active in 1950 is now stabilized (Wallis 1987). Hugenholtz and Wolfe (2005) estimate that stabilization rates in the Great Sand Hills of Saskatchewan have ranged from 7.2 to 10.5 ha/yr in the northern portion and |.3-1.4 ha/yr in the southern portion since 1946. Sta- bilization rates from 1944 to 1996 in the Elbow Sand Hills have ranged from 0.4 to 1.9 ha/yr (Hugenholtz and Wolfe 2005). In the Seward Sand Hills stabiliza- tion rates from 1970 to 1991 were estimated to be 1.2 to 3.8 ha/yr (Hugenholtz and Wolfe 2005). The Tun- stall Sand Hills stabilization rates from 1969 to 199] were estimated to be 0.6 to 3.6 ha/yr (Hugenholtz and Wolfe 2005). In Manitoba only one of the six sand hills still has active dunes (Wolfe 2001). The Brandon (Car- berry) Sand Hills stabilization rate from 1928 to 1990 has ranged from 1.8 to 17.7 ha/yr (Hugenholtz and Wolfe 2005; Wolfe et al. 2000). While the exact mechanisms are unclear, a combi- nation of drought conditions and land use appears to be influential in dune stabilization (Hugenholtz and Wolfe 2005; Wolfe et al. 2001, 2002). If current trends continue, rare native plants that now have dangerously low populations could be eliminated entirely. How- ever, Wolfe and Thorpe (2005) speculate that climate change may result in a potential increase in the suscep- tibility of sand hills to erosion, possibly reversing this stabilization trend. Exotic invasive weeds are becoming more common in dune areas and have the potential to usurp C. subgla- brum habitat (Robson 1997a). Exotic plants observed in the same habitats with C. subglabrum include: Crested Wheat-grass (Agropyron cristatum), Lamb’s-quarters (Chenopodium album), Leafy Spurge (Euphorbia esula var. esula), sweet-clover (Melilotus spp.) and Russian- thistle (Salsola tragus) (Wallis and Wershler 1988; Schmoller 2002*; Robson et al. 2005). Oil and gas ex- ploration and extraction, and to a lesser extent recre- ation are also potential threats (Wallis 1987; Robson 1997a; Lamont and Gerry 1998*). Distribution Chenopodium subglabrum is found in the prairie provinces of Canada, south to Utah and Colorado, west to Washington and Nevada, and east to the Dakotas, Nebraska and Iowa (Figure 2) (Clements and Mosya- kin 2004). The populations in Manitoba, Washington, Nevada, Utah, Colorado and Iowa appear to be dis- junct from the species’ main range (Clements and Mosyakin 2004). In Alberta there are populations of C. subglabrum in five different regions of sand hills: Grassy Lake, Middle, Rolling Hills Lake, Dominion and Pakowki Lake Sand Hills. In Saskatchewan, C. subglabrum has been collected in 11 different sand hill regions: Big- stick-Crane Lakes, Birsay, Burstall, Cramersburg, Dun- dur, Elbow, Great, Seward, Pelican Lake, Piapot and Tunstall Sand Hills. In southwestern Manitoba Che- eS) eS) oo nopodium subglabrum populations are found in the Routledge (White and Johnson 1980; Robson et al. 2005) and Brandon (Carberry) Sand Hills. Due to the annual nature of this species, the popula- tion numbers fluctuate widely. Data collected in 2004- 2005 supports this hypothesis. At several Saskatche- wan locations fewer than 50 individuals were found in the late 1990s but several hundred were observed in 2004. In the Routledge Sand Hills of Manitoba, 19 plants were observed in 2004 but 68 were seen in 2005. Population fluctuations, possibly in response to cli- mate, make it difficult to accurately estimate the num- ber of plants and the vulnerability of populations. In 2004 there were an estimated 8400 plants growing in Canada; in other years only a few hundred to several thousand plants were observed. However, even though plants may not be growing at a site in a given year, seeds are likely present in the seed bank. The largest populations occur in the Great and Seward Sand Hills of Saskatchewan, and in the Grassy Lake Sand Hills of Alberta. Based on the observed fluctuations in population size and recognition that at any given year there are some C. subglabrum seeds dormant in the seed bank, there are likely between 5 000 and 10 000 plants in Canada. The following is an overview of populations in the three provinces of occurrence. Alberta The only Alberta site visited recently (in 2004) was Lost River. Most population research on C. subglabrum in Alberta was conducted in the late 1980s (Table 2). Since such a long time has passed since these sites were visited, the dunes may have stabilized further, reducing available habitat. Saskatchewan Extensive C. subglabrum survey work was done in the mid to late 1990s in the sand hills of Saskatchewan (Robson 1997a; Lamont and Gerry 1998*). Since the THE CANADIAN FIELD-NATURALIST Vol. 120 FiGurE 2. Distribution of Chenopodium subglabrum in North America. first status report was written in 1990 (Smith and Bradley 1990*) new populations have been discov- ered in the Bigstick-Crane Lakes, Seward, Great and Tunstall Sand Hills (Robson 1997b; Nelson Dynes & Associates 1998*; Lamont and Gerry 1998*) (Table 3). This is balanced somewhat by the apparent loss of five populations where no plants have been observed for at least four decades. The Dundurn, Elbow, Great and Seward Sand Hills locations were visited as recently as 2004. TABLE 2. Locations and population data for Chenopodium subglabrum in Alberta. Location Population Last observation! Population size Dominion Sand Hills Lost River 2004 5-50 Grassy Lake Sand Hills Barnwell 1988 8 Purple Springs 1988 30 Turin 1986 100-200 Medicine Lodge Coulee Medicine Lodge Coulee 1995 ? Middle Sand Hills Cavendish 1987 ? Hilda 1988 3 Pakowki Lake Sand Hills Pakowki Lake North 1980 4 Rolling Hills Lake Sand Hills Lonesome Lake 1988 1 Estimated size of the population in Alberta 7 200-300 ' Data obtained from Smith and Bradley (1990*) * Calculated by adding the minimum number of plants observed or the maximum number of plants estimated at each site and rounding up or down. 2006 ROBSON: SMOOTH GOOSEFOOT IN CANADA 339 TABLE 3. Locations and population data for Chenopodium subglabrum in Saskatchewan. Location Population Last observation’ Population size Bigstick-Crane Lakes Sands Hills Bigstick Sand Hills 1998 21-80 Crane-Lake Sand Hills 1997 311 Tompkins 1997 21 Birsay Sand Hills Dunblane 1961 4 Broderick Broderick 1989 ) Burstall Sand Hills Empress 1985 ) Burstall 1997 202 Cramersburg Sand Hills Cramersburg 1997 179 Dundurn Sand Hills Beaver Creek 2004 1-4 Beaver Creek East 1952 ? CFB Dundurn 1997 6 Elbow Sand Hills Bridgeford 1968 ? Elbow 2004 288-500 Head of Qu’appelle 1879 ? Great Sand Hills Sandhill Stockman’s Association 2004 2018-37967 Diamond Ranch 1997 435 Heck Stockman’s Association 2004 18-3307 Signal Stockman’s Association 2004 36-3307 Watson’s 2004 156 + 3307 McMahon McMahon 1949 ? Pelican Lake Sand Hills Caron 2002 <10 Piapot Sand Hills Piapot 1983 ? Patience Lake Patience Lake 1986 ? Seward Sand Hills SSH 1 1996 55-5407 SSH 2 2004 39-1000 SSH 3 2004 13-330 SSH 4 2001 90-5407 SSH 5 2001 10-540? SSH 6 2001 2 SSH7 2001 l Tunstall Sand Hills Bitter Lake 1997 1] Estimated population in Saskatchewan + 5000-9700 ' Data obtained from Harms (1990); Smith and Bradley (1990*); J. & W. Resource Management Consulting (1997*); Robson (1997a); Lamont and Gerry (1998*); Nelson Dynes & Associates (1998*). ? Estimated number of plants in 2004; determined by multiplying the number of dune fields present in the area by the aver- age number of plants found at other dunes in the same area. 3 Calculated by adding the minimum number of plants observed or the maximum number of plants estimated at each site and rounding up or down. TABLE 4. Locations and population data for Chenopodium subglabrum in Manitoba. Location Population Last observation Population size Routledge Sand Hills Routledge 2005 68-75 Oak Lake 1959 ? Brandon (Carberry) Sand Hills Spruce Woods 2005 9-25 Estimated size of the population in Manitoba ! 77-100 ' Calculated by adding the minimum number of plants observed or the maximum number of plants estimated at each site and rounding up or down. Manitoba Although the species was found near Oak Lake in 1959 it was not observed again until 2004 when it was rediscovered (The Manitoba Museum voucher speci- men catalogue # 37136) near Routledge (Robson et al. 2005) (Table 4). The active dunes in the Brandon (Carberry) Sand Hills were also searched for C. sub- glabrum in 2004 but no plants were found. However, in 2005 a small population of nine plants (The Mani- toba Museum voucher specimen catalogue # 37859) was located. Two plants were found in the same area in 2006. This is the first record ever for this location and represents a range extension for this species of approximately 100 km east. 340 Evaluation Significance Chenopodium subglabrum plays a role in dune sta- bilization, being one of the few plants that can colonize active sand. Several related species are important eco- nomically, including Quinoa (C. quinoa), grown for its seeds, and Good King Henry (C. bonus-henericus), grown for its leaves (Everett 1981). The aboriginal peoples of the western United States consumed seeds of Chenopodium pratericola and Fremont’s Goosefoot (C. fremontii) (Smith and Bradley 1990*). Rarity Status Chenopodium subglabrum was listed by the Com- mittee on the Status of Endangered Wildlife in Canada in 2006 as being a threatened species in Canada. Legal protection of this species via the Species at Risk Act, 2003 will likely occur in the next few years. This spe- cies is considered rare in all three prairie provinces (Packer and Bradley 1984; Maher et al. 1979; White and Johnson 1980) but is not yet protected by any provincial legislation. This species is not protected by any legislation in the United States. The NatureServe ranks are Global G3G4 (globally vulnerable to apparently secure), Canada N2 (nation- ally imperiled), Alberta S1 (provincially critically im- periled), Saskatchewan S2 (provincially imperiled), and Manitoba S1 (provincially critically imperiled). The United States ranks are: Montana S1 (critically imperiled in the state), North Dakota S1 (critically imperiled in the state), Nebraska S3 (vulnerable in the state) and Wyoming S3 (vulnerable in the state) (NatureServe 2005*). In the remainder of the states where it occurs it is has either not been ranked (e.g., Colorado, Idaho, Kansas, Nevada, Oregon, South Da- kota, Utah and Washington) or is considered unrank- able due to a lack of information (e.g. Michigan) (NatureServe 2005*). Future Outlook Chenopodium subglabrum is restricted to sandy habitats in the prairie provinces that are geographical- ly isolated from one another. Due to C. subglabrum’s annual nature, it experiences large fluctuations in pop- ulation size from year to year. Dune stabilization has been occurring at a high rate in all of the sand hill habitats where this species occurs. Increased pressure on these ecosystems from the oil and gas industry, and encroachment of exotic weeds further threaten this species. Acknowledgments The Canadian Wildlife Service (Environment Cana- da), Manitoba Conservation, and the Manitoba Museum Foundation Inc., provided funding for this research. Some of the data in this article were obtained from the original status report on this species prepared by Bonnie Smith and Cheryl Bradley. Thanks are extended to the many scientists, particularly Candace Elchuck and Joyce THE CANADIAN FIELD-NATURALIST Vol. 120 Gould, who provided information for this report, the landowners for allowing access to their land, and Janis Klapecki, Cary Hamel and Cathy Foster for participat- ing in the field surveys. Documents Cited (marked * in text) Harms, V. L. 1990. The vascular plant flora of the Beaver Creek Conservation Area, near Saskatoon, Saskatchewan. The W.P. Fraser Herbarium, and the Department of Biol- ogy, University of Saskatchewan, Saskatoon, Saskatche- wan. J. & W. Resource Management Consulting. 1997. Rare plant survey of Dundurn Military Training Reserve, Sas- katoon, Saskatchewan. Lamont, S., and A. Gerry. 1998. Species status report Chenopodium subglabrum (S. Wats.) A. Nels. Saskatche- wan Conservation Data Centre, Regina. NatureServe. 2005. NatureServe Explorer: An Online Ency- clopaedia of Life. Web site: http://www.natureserve.org/ explorer. [accessed June 2005]. Nelson Dynes & Associates Inc. 1998. Beverley-Cantuar comprehensive environmental protection plan. Saskatoon, Saskatchewan. Schmoller, D. 2002. Chenopodium subglabrum surveys 2002 field season. Yellowfield Biological Surveys, North Dakota. Smith, B., and C. Bradley. 1990. Status report on the Smooth Goosefoot Chenopodium subglabrum (S. Wats.) A. Nels. Committee on the Status of Endangered Wildlife in Cana- da. Ottawa. 27 pages. Literature Cited Acton, D. F., G. A. Padbury, and C. T. Stushnoff. 1998. The Ecoregions of Saskatchewan. Canadian Plains Re- search Center, University of Regina, Regina. 205 pages. Argus, G. W., and K. M. Pryer. 1990. Rare vascular plants in Canada. Canadian Museum of Nature, Ottawa. 274 pages. Bassett, I. J., and C. W. Crompton. 1982. The genus Cheno- podium in Canada. Canadian Journal of Botany 60: 586- 610. Clements, S. E., and S. L. Mosyakin. 2004. Flora of North America Volume 4: Magnoliophyta: Caryophyllidae, Part 1, Chenopodiaceae, Chenopodium. Oxford University Press, Toronto. Everett, T. H. 1981. Illustrated encyclopedia of horticulture. Volume 3. Garland Publishing Inc., New York. Grime, J. P. 1979. Plant strategies and vegetation processes. John Wiley & Sons, Toronto. Hugenholtz, C. H., and S. A. Wolfe. 2005. Recent stabiliza- tion of active sand dunes on the Canadian prairies and rela- tion to recent climate variations. Geomorphology 68: 131- 147. Johnson, D. L., and S. M. Ward. 1993. Quinoa. Pages 219- 221 in New Crops. Edited by J. Janick and J. E. Simon. John Wiley & Sons, New York. Maher, R. V., G. W. Argus, V. L. Harms, and J. H. Hudson. 1979. The rare vascular plants of Saskatchewan. National Museum of Natural Sciences Syllogeus 20. National Muse- ums of Canada, Ottawa. 81 pages. Mukerji, K. G., B. P. Chamola, and J. Singh. 2000. Myc- orrhizal biology. Kluwer Academic, New York. Packer, J. G., and C. E. Bradley. 1984. A checklist of the rare vascular plants in Alberta. Provincial Museum of Al- berta Natural History Occasional Paper 5. Alberta Culture, Edmonton. 112 pages. 2006 Robson, D. B. 1997a. Ecology of rare vascular plants in southwestern Saskatchewan. M.Sc. dissertation, University of Saskatchewan, Saskatoon, Saskatchewan. 128 pages. Robson, D. B. 1997b. Smooth Goosefoot discovered in the Webb sandhills. Blue Jay 55: 106-108. Robson, D. B. 1999. Reasons for prairie plant rarity. Pages 92-95 in Proceedings of the Fifth Prairie Conservation and Endangered Species Conference, Saskatoon, Saskat- chewan. Edited by J. Thorpe, T. A. Steeves, and M. Gollop. Provincial Museum of Alberta, Edmonton. Robson, D. B., J. Greenall, C. Hamel, and C. Foster. 2005. Smooth Goosefoot rediscovered in Manitoba. Blue Jay 63: 81-84. Royer, F., and R. Dickinson. 1999. Weeds of Canada. Lone Pine Publishing, Edmonton. Wallis, C. A. 1987. Critical, threatened and endangered habitats in Alberta. Pages 49-63 in Proceedings of the Workshop on Endangered Species in the Prairie Provinces, Edmonton, Alberta. Edited by G. L. Holroyd et al. Provin- cial Museum of Alberta Natural History Occasional Paper 9. Wallis, C. A., and C. Wershler. 1988. Rare wildlife and plant conservation habitats of southern Alberta. Albert Forestry, Lands and Wildlife; Alberta Recreation and Parks; World Wildlife Fund Canada, Edmonton. ROBSON: SMOOTH GOOSEFOOT IN CANADA 34] White, D., and K. Johnson. 1980. The rare vascular plants of Manitoba. National Museum of Natural Sciences Syllo- geus 27. National Museums of Canada, Ottawa. 77 pages. Wolfe, S. A. 2001. Eolian deposits in the Prairie Provinces of Canada. Natural Resources Canada, Ottawa. Wolfe, S.A., D. J. Huntley, P. P. David, J. Ollerhead, D. J. Sauchyn, and G. M. MacDonald. 2001. Late 18 century drought-induced sand dune activity, Great Sand Hills, Sas- katchewan. Canadian Journal of Earth Science 38: 105- 117. Wolfe, S. A., D. R. Muhs, P. P. David, and J. P. McGeehin. 2000. Chronology and geochemistry of late Holocene eolian deposits in the Brandon Sand Hills, Manitoba, Canada. Quaternary International 67: 61-74. Wolfe, S.A., J. Ollerhead, and O. B. Lian. 2002. Holocene eolian activity in south-central Saskatchewan and the south- ern Canadian prairies. Geographie physique et Quaternaire 56 (2-3): 215-227. Wolfe, S. A., and J. Thorpe. 2005. Shifting sands: Climate change impacts on sand hills in the Canadian prairies and implications for land use management. Prairie Forum 30: 123-142. Received 14 September 2005 Accepted 5 February 2007 Summer Movements and Impact of Individual Striped Skunks, Mephitis mephitis, on Duck Nests in Saskatchewan SERGE LARIVIERE!*, LYLE R. WALTON’, and FRANCOIS MESSIER? 'Delta Waterfowl Foundation, R.R. 1, Box 1, Site 1, Portage La Prairie, Manitoba RIN 3A1 Canada. Present address: Cree Hunters and Trappers Income Security Board, 2700 boulevard Laurier, Champlain # 1110, Québec (Québec) G1V 4K5 Canada *Wildlife Research and Development Section, Ontario Ministry of Natural Resources, Ontario Government Complex, Hwy 101 E., P.O. Bag 3020, South Porcupine, Ontario PON 1HO Canada > Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada Lariviére, Serge, Lyle R. Walton, and Frangois Messier. 2006. Summer movements and impact of individual Striped Skunks, Mephitis mephitis on duck nests in Saskatchewan. Canadian Field-Naturalist 120(3): 342-346. Striped Skunks (Mephitis mephitis) are important predators of duck eggs in the Canadian prairies. We estimated the impact of individual Striped Skunks on duck nests by intensive observations of foraging movements and depredation of duck nests in southern Saskatchewan, 1993-1994. Nightly, skunk movements were variable (range 0-20 km per night), and did not dif- fer among seasons for females, or between males and females during the parturition/rearing season. Overall, nightly move- ment of Striped Skunks averaged 7.4 km for females (SD = 5.9 km, n = 20 females) and 6.7 km for males (SD = 3.2 km,n=5 males). During 1 873 h of radio-tracking, we observed depredation of 10 duck nests by 8 skunks (7 F, 1 M). Using our observed estimate of one depredation for every 187 h, and averaged nightly activity of 8-10 h per night, we estimated that indi- vidual skunks find one duck nest every 19-23 nights. Thus, during the 60-day nesting season for ducks (mid-May to mid-July), individual skunks probably find 2-3 duck nests. These observations reinforce the growing evidence that, at normal duck nest densities (<2.5 nests/ha), depredation of eggs by Striped Skunks is opportunistic, and the impact of Striped Skunks on duck nests is a direct function of Striped Skunk abundance. Key Words: Striped Skunk, Mephitis mephitis, duck, movement, nest predation, prairie, predator ecology, Thickwood Hills, Saskatchewan. Predation on eggs is an important limiting factor for many bird populations (Ricklefs 1969; Martin 1995). In the grasslands of North America, population decline of many species of ground nesting birds is attributed to increased predation on eggs during the breeding sea- son (BOhning-Gaese et al. 1993). Of these birds, ducks (Anatidae) nesting in the prairies of Canada and the United States experience high predation rates on their nests. The primary mammalian predators are Striped Skunks (Mephitis mephitis), Red Foxes (Vulpes vulpes), Raccoons (Procyon lotor), American Badgers (Taxidea taxus) and Coyotes (Canis latrans). The impact of these predators on duck nest success is well recognized (Klett et al. 1988; Johnson et al. 1989; Pasitschniak-Arts and Messier 1995). However, the contribution of individ- ual animals is poorly understood, yet is critical for management and mitigation of nest predation. In the parklands of Saskatchewan, the Striped Skunk is an important predator of duck eggs (Pasitschniak- Arts and Messier 1995), yet it depredates duck nests opportunistically (Lariviére and Messier 1997a, 1998a). This suggests that the impact of skunks on duck nest success is the result of “chance” encounters with duck nests as skunks forage for other foods such as small mammals and insects (Greenwood et al. 1999). There- fore, the impact of Striped Skunks on duck nests should be a function of (a) habitat selection patterns (do skunks use habitats where ducks nest?), (b) movement rates (the longer the movements by skunks, the greater the chances of discovering a duck nest), and (c) density of skunks (the more skunks, the greater the impact on duck nests). A recent experiment suggested that indi- vidual predators were unlikely to key on duck nests at densities <2.5 nests/ha (Lariviére and Messier 1998a). Furthermore, partial depredation of duck nests by Striped Skunks commonly occurs (Lariviére and Mes- sier 1997a; Ackerman 2002), suggesting that duck eggs may not be a preferred food of skunks or that their importance may be marginal to the overall nutrition of Striped Skunks. An estimate of the impact of individual Striped Skunks on duck nests would help further our understanding of the role that duck eggs play in the for- aging ecology of this carnivore. Herein, we estimate the impact of individual Striped Skunks on duck nests by combining intensive radio-tracking and observations of free-ranging Striped Skunks in the Prairie Pothole Region of Saskatchewan, Canada. Methods We conducted this study ca. 80 km northwest of Saskatoon in the Thickwood Hills of southcentral Sas- katchewan, Canada (52°45'N, 107°08'W). In 1993, the area (ca. 220 km?) was dominated by farmland and interspersed with many wetlands and stands of trem- 342 2006 bling aspen (Populus tremuloides). Cropland repre- sented 66% of the area, and was primarily used for the production of grain (e.g., wheat, barley, rye, and oats), and oil crops (e.g., canola and flax). Other less common crops included peas, lentils, buckwheat, and canary seed. Woodland and areas managed for nesting waterfowl] (mostly dense nesting cover) occupied 11% and 9% of the landscape, respectively. Managed nesting areas consisted mostly of dense nesting cover, hay land, and idle pastures. Fields of dense nesting cover were sown with various tame and wild grasses to create prime habitat for nesting waterfowl. Small or linear cover types such as wetlands, rights-of-way, farmsteads, and miscellaneous habitats represented 8%, 4%, 1%, and 1% of the land available, respectively. Topography was gently rolling, and an extensive network of roads divid- ed the land. In 1993, Striped Skunks were live-trapped using opaque-sided livetraps (Dura-Poly livetraps, Ram Sup- plies, Birtle, Manitoba). Captured skunks were anes- thetized with halothane and Telazol® (Lariviére and Messier 1996a, 1996b), and equipped with motion- sensitive radio-collars (Telonics Inc., Mesa, Arizona, USA). Skunks were handled and released at site of capture. Radio-collared animals were located and tracked by a single observer on foot from 18:00 to 06:00 h, dur- ing the period of greatest activity (Lariviére and Mes- sier 1997b). During tracking, a single observer fol- lowed the foraging skunk while remaining downwind, and usually within 15-50 m of the animal (Lariviére and Messier 1998b). This distance allowed precise location of focal animals every 15 min, and allowed observation of foraging behavior. For short movements (<100 m), distance from previous location was esti- mated by observer. Locations immediately following an observer-induced defensive posture (n = 198: Lar- iviére and Messier 1996c) were discarded from analy- ses because of observer disturbance (Lariviére and Messier 1998b). At dark, we used night-vision equip- ment (AN-PVS 5C, Bill’s Electronics, Mildmay, Ontario) to continue observations (Lariviére and Mes- sier 1998b). Encounters with duck nests typically were easy to detect because of the flushing hen (Lariviére and Messier 1997a) or repeated visits to the nest (Lar- iviére and Walton 1998). Depredated nests were in- vestigated immediately following departure of the skunk, and usually reexamined the following day. For each location, we recorded position of the focal skunk using a Global Positioning System (GPS) unit. We estimated accuracy of our GPS as + 250 m. To overcome biases associated with the inaccuracy of our GPS on short movements, we only considered the total distance traveled by each skunk during the 12-h track- ing period (sum of all 15-min movements). We also only considered complete 12-h blocks for which onset and cessation of activity were known. Missing values (typically caused by delays of the GPS unit to obtain LARIVIERE, WALTON, MESSIER: IMPACT OF SKUNKS ON DUCK NESTS 343 fixes in forested habitats) for <2 intervals per night were approximated by the nightly mean of 15-min trav- el distances. Sessions with >2 missing values were dis- carded from analyses. Biological seasons for Striped Skunks were identified as pre-parturition (1 April-14 May), parturition/rearing (15 May- 30 June), pre-dis- persal (1-31 July), and dispersal (1-31 August). Each summer (May-August), >50% of radio-col- lared animals died or dispersed outside the study area (Lariviére and Messier 1998c). Thus, the resulting data set was not balanced, and precluded use of repeated- measures analyses, and comparison among seasons. Therefore, we used a one-way analysis of variance to test for effects of season on movements. To avoid bias- es caused by animals tracked >1 night in a given sea- son, we used the mean distance traveled for each skunk in each season (weighted mean, only one value per animal per season). Because some skunks were tracked during more than one season and they may not have been independent, we used an alpha value of 0.01 to remain conservative and reduce the chance of a Type 1 error. Results During 1993 and 1994, we obtained complete move- ment information for 63 12-h tracking sessions (11 in 1993, and 52 in 1994) on 25 Striped Skunks (5 M, 20 F). Because no skunks were radio-tracked during both years, we pooled data from 1993 and 1994 for analyses. Nightly movements of female Striped Skunks during pre-parturition (mean = 4.2 km, SD = 4.9 km, n = 4), parturition/rearing (mean = 7.2 km, SD = 6.1 km, n = 19), pre-dispersal (mean = 5.6 km, SD = 3.0 km, n= 8), and dispersal (mean = 3.0 km, SD = 3.1, = 2) were not statistically different Ga 0:70; P:=056): Distance traveled during 12-h tracking sessions were variable and ranged from 0.0 to 20.3 km per night (mean = 6.2 km, SD = 5.2 km, n = 33 female-season). When we considered only one value for each female (weighted mean; average of all seasons), nightly move- ments averaged 7.4 km (SD = 5.9 km, n = 20 females). During the parturition/rearing season, movements of males (mean = 6.7 km, SD = 3.2 km, n = 5) did not differ (t-test, df = 22, P = 0.84) from movement of females (mean = 7.2, SD = 6.1 km, n = 19). Move- ments of all skunks during the parturition/rearing peri- od averaged 7.1 km (SD = 5.6 km, n = 24 skunks). During spring and summer 1993 and 1994, we radio- tracked Striped Skunks for 1873 h (858 h in 1993, and 1015 h in 1994). During radio-tracking, we observed depredation of 10 duck nests (4 in 1993, 6 in 1994), thus averaging one depredation event for every 187 h of radio-tracking (1 for 215 h of radio-tracking in 1993, and | for 169 h in 1994). Observed depredation events were caused by 8 Striped Skunks (7 females, 1 male). Observed depredation events occurred between 9 June and 9 July in 1993, and between 15 May and 20 July in 1994. 344 From our observations, we extrapolated the number of nests depredated by a skunk during the duck nest- ing season. First, using our rate of discovery of | nest every 187 h of tracking, and allowing an average 8- 10 h of foraging movements per night (Lariviere and Messier 1997b), we estimated that individual skunks found 1 duck nest for every 19-23 nights of foraging. Considering the duck nesting season lasts approxi- mately 60 d (mid-May until mid-July), then individual skunks may find 2-3 duck nests per season. However, because skunks foraging in dense vegetation may have depredated duck nests without us noticing, this esti- mate represents a minimum number, but our close- range observations (typically <50 m) and use of night- vision equipment suggests that we did not miss many depredation events. Discussion Our results clearly show that the foraging behavior of Striped Skunks consists of highly variable foraging movements and opportunistic, but infrequent, depreda- tion of duck nests. The variable movements we observed within and among seasons, or between sexes, preclud- ed detection of statistical differences. However, the trend suggested that movements were shorter during the pre-parturition season, higher during parturition season, and declined temporally thereafter. Interesting- ly, the decline in skunk movements over time coin- cides with the steady increase in availability of both small mammals and insects over time (Lariviére and Messier 2000). Although we could not statistically link movements of skunks and food abundance because of only four sampling periods, we nonetheless observed apparently reduced skunk movements over time as prey abundance increased. An experiment done with sim- ulated nests in our study area in 1995 indicated that increases in availability of food (chicken eggs) caused a reduction of activity and home ranges in female Striped Skunks (Lariviére and Messier 2001a). Movement rates observed in our study are the largest reported for this species. However, most previous stud- ies of skunk movements relied on distance between consecutive den sites (Shirer and Fitch 1970), maxi- mum distance from the den during any given night (Verts 1967), or distance between recapture sites (Bjorge et al. 1981). Our study is one of three that reported actual distance moved from telemetry loca- tions. In Illinois, daily movements of Striped Skunks averaged 1.7 km for males (n = 4) and 1.4 km for females (n = 9) (Storm 1972). In North Dakota, night- ly movements ranged from 0.0 to 9.3 km for 26 females (mean = 2.7 km), and from 0.4 to 10.5 km for males (mean = 3.3 km) (Greenwood et al. 1985). We believe our values are higher because we collected a more accurate description of movement paths by collecting locations at closer intervals (i.e., every 15 min) where- as animals were located every 2 h in North Dakota (Greenwood et al. 1985), and at intervals varying THE CANADIAN FIELD-NATURALIST Vol. 120 between 10 and 120 minutes in Illinois (Storm 1972). Two additional explanations may explain the long movements of Striped Skunks in our study area. The first may be the composition of the landscape. Our study area was primarily cropland (66% of study area), a habitat that provides low abundance and diversity of foods for skunks (Larivieére and Messier 2000). Not surprisingly, Striped Skunks in our study area avoided cropland when foraging (Lariviere and Mes- sier 2000). Thus, large amounts of cropland may con- tribute to increased movements as skunks were forced to travel across large fields to access prime habitats embedded therein. Nightly movements of Striped Skunks may be lower in large patches of continuous foraging habitats, a phenomenon that would help ex- plain the typically higher nest success of ground-nest- ing birds in large patches of non-fragmented habitat (Pasitschniak-Arts and Messier 1996; Ball et al. 1995). Another explanation may be related to the amount of food in the landscape. Possibly, food abundance may have been lower in our study area compared to previous studies, thus contributing to longer move- ments. Food supplementation experiments on Striped Skunks and other species typically yield lower activi- ty, smaller ranges, and shorter foraging bouts (Boutin 1990; Lariviére and Messier 1998a). However, because we have no way of comparing food abundance among studies, we can only speculate on whether differences existed between our study area and those used in pre- vious studies. Managers attempting to mitigate predation on duck nests need to know whether high predation rates occur because of specialized foraging by a few individuals depredating numerous nests, or simply by a high abun- dance of individuals each depredating only a few nests. Previous observations of skunk foraging behavior sug- gested that predation is incidental to foraging for other foods (Vickery et al. 1992; Lariviére and Messier 1997a), and that greater predation rates are explained by a greater abundance of nest predators (Lariviere and Messier 1998a). Our study suggests that individual Striped Skunks in the Canadian prairies find between 0 and 3 duck nests per year. In fact, we observed only two skunks depredate more than one duck nest in any given year (both cases of females depredating two nests). Because our study area was located where nesting cover was common (managed nesting areas represented 9% of the study area), we believe our estimates probably are high because most areas of the Canadian prairies do not have 10% of the land as cover available for ducks, and hence probably hold fewer duck nests. Nonethe- less, this suggests that availability of duck nests in the landscape is not high enough to warrant special for- aging efforts; this reinforces the notion that nest depre- dation by Striped Skunks is strictly incidental (Lariv- iére and Messier 1997a, 1998a, 2001b; Vickery et al. 1992). Finally, the rate of encounter suggests that the 2006 impact of skunks on duck nests is directly related to skunk abundance, as was previously suggested (Lar- iviere and Messier 1998a). The impact of individual Striped Skunks on duck nests provides insight into the low success of manage- ment techniques that aim at educating individual pred- ators. For example, conditioned-taste aversion had been unsuccessful under field conditions when mammals such as skunks and Raccoons were the main predators (Clark et al. 1996). We believe our data suggest that encounter rates with duck nests by individual preda- tors are too low for such techniques to be effective, notwithstanding problems in causing aversion (Clark et al. 1996). At normal nest densities (<1 nest/ha), predators cannot subsist on eating duck eggs alone, and duck eggs only represent a marginal food item (Greenwood et al. 1999). Not surprisingly, foraging activity of Striped Skunks is directed at habitats where base prey such as insects and small mammals are most abundant, and not in areas managed for nesting ducks (Lariviere and Messier 2000). Thus, management tech- niques that aim to increase nest success should be directed at reducing predator abundance locally (Gar- retson and Rohwer 2001), or reducing overall habitat suitability for predators with implementation of large- scale habitat programs through agricultural policies (e.g., Conservation Reserve Program in the United States, Reynolds et al. 2001). Acknowledgments This study was supported by the Prairie Habitat Joint Venture through a research grant from Ducks Un- limited (Institute for Wetland and Waterfowl Research), the Canadian Wildlife Service, as well as support from the Delta Waterfowl Foundation, the National Science and Engineering Research Council through and oper- ating grant to SL, as well as postgraduate and post- doctoral fellowships from the Fonds pour la Forma- tion de Chercheurs et |’ Aide a la Recherche (FCAR), Québec. Field and laboratory assistance was provid- ed by B. Dinter, E. Howard, G. Poon, B. Fry, K. Eade, H. Royer, P. Toner, S. 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Received 18 August 2005 Accepted 23 April 2007 Life History Phenology and Sediment Size Association of the Dragonfly Cordulegaster dorsalis (Odonata: Cordulegastridae) in an Ephemeral Habitat in Southwestern British Columbia LAURIE B. MARCZAK'!, JOHN S. RICHARDSON!, and MARIE-CLAIRE CLASSEN2 ' Department of Forest Sciences, University of British Columbia, 3041-2424 Main Mall, Vancouver, British Columbia V6T 174 Canada; Corresponding author: e-mail: laurie@interchange.ubc.ca ? Department of Biological Sciences, University of Alberta, CW405, Biological Sciences Centre, University of Alberta, Edmonton, Alberta T6G 2E9 Canada Marezak, Laurie B., John S. Richardson, and Marie-Claire Classen. 2006. Life history phenology and sediment size associ- ation of the dragonfly Cordulegaster dorsalis (Odonata: Cordulegastridae) in an ephemeral habitat in southwestern British Columbia. Canadian Field Naturalist 120(3): 347-350. The life cycle of the dragonfly Cordulegaster dorsalis was studied over one year by systematic sampling of larvae in three intermittent headwater streams in southwestern British Columbia. We determined that larvae normally take three years to reach maturity, emerging throughout July and August. There is limited evidence suggesting a split cohort development, with early emergence after two years. Additionally, we tested whether larval instars were distributed randomly or if they occupied different sediment microhabitats. Smaller animals tend to be associated with smaller grained organic sediments, although there was high variation between the streams. Key Words: Odonata, dragonfly, Cordulegaster dorsalis, ephemeral stream, life history, phenology, British Columbia. Dragonfly larvae are often conspicuous and impor- tant predators in stream ecosystems. In British Colum- bia, the family Cordulegastridae is represented by a single species, Cordulegaster dorsalis Hagen. Cord- ulegastrids occur in lotic waters, usually in small streams, where larvae live as shallow burrowers in sedi- ment (Cannings 2002). In southwestern coastal British Columbia, larvae of C. dorsalis are often found in small headwater streams that are subject to extreme episodic drying. In the interior of the province, the species is less common and is mainly associated with small, spring-fed streams. Adults of C. dorsalis nor- mally fly from mid-May to early September, peaking in July (Cannings 2002). Published accounts of the life history, habitat associations or ecological requirements of C. dorsalis in its larval stage are sparse. Kennedy (1917) estimated C. dorsalis larvae in cen- tral California may require as many as four years to develop. The larval development of cordulegastrids in Europe ranges from 2 to 5 years and may include split cohorts (Ferreras-Romero and Corbet 1999). Split cohort timing has been documented in other species in the genus (C. boltonii [Donovan] in Ferreras-Romero and Corbet 1999). Corbet (1999: 230) maintains it is _ possible to identify a winter critical size, below which larvae in the first year of development will enter dia- _ pause during the subsequent summer period, requiring _ another year’s growth to reach metamorphosis. Such : differential growth rates within a single cohort may be produced by staggered egg-laying times, food limita- _ tion or variability in the environment. Given the tran- sitory nature of the streams used by C. dorsalis, it is possible that it exhibits plasticity in growth rates and emergence times in response to continual shrinking and rewetting of the streams. Dragonfly larvae increase in size by three to four orders of magnitude as they develop. Microhabitat selection may have much to do with the physical con- straints of size. Different species of burrowing odo- nates prefer certain sediment-size categories (Corbet 1999). C. boltonii has sediment-size preferences that vary with specific larval size classes (Prodon 1976 unpublished dissertation referenced in Corbet 1999). In contrast, C. maculata Sélys in Virginia is a habitat generalist, occuring in equal densities in both silts and sands (Burcher and Smock 2002). In general, small larvae of C. boltonii prefer much finer sediments than do larger larvae — larvae tend to select a particle size that offers the least obstruction to burrowing (Corbet 1999). We report life-history observations (morphological measurements) of larval C. dorsalis over a one year period in three small headwater streams subject to late summer drying. We observed that large and small lar- vae did not occur together within pools. At the same time, we observed surprisingly high densities of odo- nate larvae in small, shrinking pools during late sum- mer. We tested whether small and large larval instars occupy different sediment microhabitats and report life- history observations (morphological measurements) of larval C. dorsalis over a one-year period in three small headwater streams subject to late summer drying. 347 348 Head width (mm): THE CANADIAN FIELD-NATURALIST Vol. 120 Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul BR 100% FiGurE |. Life history patterns of Cordulegaster dorsalis shown for 2002-2003 — in G Creek (n = 175). Continuous lines ascending from left to right indicate inferred boundaries between successive hatching cohorts. The emergence period during late summer is shown (dotted line). Size ranges of the last three instars are indicated. The broken line indicates an alternate interpretation and represents the inferred boundary between the two components that could result from cohort-splitting in the two-year-old hatching cohort, thus distinguishing larvae that grow fast (above line) and slowly (below line). Numbers below columns indicate sample sizes for each sampling date. Widths of black bars for each interval of headwidth on a given date represent the relative fraction of the total sample that fell in that interval; total possible width for any given sampling date is represented by the black legend box labelled 100%. Methods Study Site This study took place in the 5157 ha Malcolm Knapp Research Forest, located in Maple Ridge, British Columbia. The forest is located in the Pacific coastal rain forest, with a maritime climate of cool dry summers and warm wet winters (Naiman et al. 2000). We sampled Cordulegaster dorsalis larvae for life history characteristics and sediment associations in three streams within the forest (I, F and G streams; 49°18'40"N, 122°32'40"W), which were part of an on- going riparian management experiment (see Kiffney et al. (2003) for site descriptions). Both F and G streams retained a 10 m forest buffer while I stream flowed through a 5-year-old clearcut. The dominant vegetation surrounding these creeks included Red Alder (Alnus rubra) and Vine Maple (Acer circinatum) with a canopy composed largely of Western Hemlock (Tsuga hetero- phylla), Western Redcedar (Thuja plicata) and Dou- glas-fir (Pseudotsuga menziesii). Vegetation adjacent to I stream consisted of shrubs, particularly Salmon- berry (Rubus spectabilis) and Huckleberry (Vaccinium ovatum). Life-history characteristics We surveyed approximately 200 m each of I, F, and G streams in August, October and December of 2002 and April, July and August of 2003, collecting larvae using dip nets and hand sorting. We measured head width (Hdw max. distance between the lateral margins of the compound eyes), total body length (Tbl; max. distance between the mouthparts and the end of the cerci measured along the dorsal surface) and wing- pad length to the nearest 0.1 mm with vernier-scale callipers. Using regression analysis we determined the scaling coefficient (reported as the slope of the line) for head width and body length. Size ranges of late instar classes were determined from the relationship between wingpad length and head width. Year class separation was analysed using wingpad to head width relation- ships together with visual assessment of size frequen- cy distributions (after Ferreras-Romero and Corbet 1999). The level of significance was set at P=0.05. Sediments The habitat associations of different size classes of larvae were characterized from August 2002 through April 2003. Sediment samples were collected at each point of larval capture and sieved into six size frac- tions (63 tm, 250 Um, 500 um, | mm, 2 mm, 4 mm). Each sediment fraction was air-dried and weighed, then ashed at 550°C for 2 h and reweighed to obtain weights of both inorganic and organic fractions. Cumu- lative percent curves were used to determine the medi- an particle size of each sample. We used linear regres- sion to investigate the data for relationships between larval head width, median sediment size and organic fraction. 2006 head width (mm) 0 10 20 30 40 50 % organic matter MARKZAK, RICHARDSON, AND CLASSEN: DRAGONFLY IN EPHEMERAL HABITAT 349 0 | nN we = median particle size (mm) FIGURE 2. Relationship between larval head width of Cordulegaster dorsalis and sediment characteristics — (A) percentage of organic matter; and (B) median particle size. Regressions were significant for F Creek (closed circles) but not for G Creek (open squares) or I Creek (open triangles). Results and Discussion During August 2003 when sample sizes were lar- gest, head width of F-0 (7.5 mm — 9.2 mm) and F-1 (6 mm — 7.8 mm) larvae overlapped, but wingpad lengths were discrete (3 mm — 5.5 mm in F-2, 6.9 mm — 10 mm in F-1 and 11.9 mm — 15.5 mm in F-0). Total body length and head width were correlated (R? = 0.86, P<0.001, TBL = 4.49 HW — 0.59). In G Creek, larvae of Cordulegaster dorsalis — de- rived from eggs that likely hatched in early summer — grew through late summer and early autumn (Figure 1). At the beginning of their second year most larvae had a head width between 2 mm and 5 mm. At the end of their second year of growth, larvae achieved head- widths between 5 mm and 8 mm. Emergence occurred at head widths greater than ca. 8 mm during July and August of the third year. The pattern of development that we observed in G Creek (where sample size was greatest, n = 180) indi- cated that, in general, C. dorsalis larvae require three years to achieve metamorphosis. It is possible to inter- pret these data as evidence of a split-cohort in the sec- ond year with a “fast” group of larvae emerging fol- lowing accelerated late summer to early fall growth at the end of their first year (Figure 1, dashed line). This interpretation would suggest a winter critical size of ca. 6 mm (Hdw; Figure 1). Sample sizes were too small in I Creek (n = 30) and F Creek (n = 21) to bol- Ster a split-cohort interpretation; the Hdw of collected larvae generally support a three-year development. Many Odonata retain a synchronized emergence period through the use of photoperiod cues (e.g. Aoki 1999; Burcher and Smock 2002). Some species of odonates appear to have both “slow” and “fast” com- ponents of the same cohort (Martin et al. 1991; John- son et al. 1995; Ferreras-Romero and Corbet 1999). We present two possible development schedules for C. dorsalis: a split cohort with a “fast” component emerging after two years and a “slow” component re- quiring three years to complete development. Cordule- gastridae are noted for their protracted larval develop- ment, a pattern that might be anticipated of shallow burrowers occupying small, cool streams (Corbet 1999). Two years would represent a remarkably short development time within this family, particularly given the known four-year development time for cordule- gastrids in northern California (Kennedy 1917). A more complete record of larval growth rates is required before evidence of a slow and fast development in C. dorsalis can be dismissed; in the meantime, the data most strongly support a simple three-year development schedule. During our surveys for larval development, we ob- served that small pools seldom contained both large and small larvae. Larvae were most abundant at medi- an particle sizes between 1.5 mm and 2.5 mm. The head width of larvae was negatively associated with percent total organic content in F Creek (Figure 2a: P <0.01; R? = 0.451) but no relationship was evident in either G or I creeks. Similarly, the head width of larvae was positively associated with median particle size in F Creek (Figure 2b; P < 0.05; R? = 0.446) while no relationship was evident in either G or I creeks. It should be noted that median particle and percent 350 organic matter are not independent measures; there is a negative correlation (r = -0.699). Post-hoc power analysis suggested that these results must be interpreted cautiously (power = 0.5134 organic; 0.5072 median particle). All odonate larvae are opportunistic predators that will eat almost any type of prey they can detect and subdue (Burcher and Smock 2002). Since larger lar- vae will eat smaller larvae when they encounter them, it seems plausible that some form of habitat partition- ing occurs to minimize intraspecific competition and cannibalism. How larvae are distributed at the micro- habitat scale will largely determine the frequency of their interactions with conspecifics, affecting the prob- ability of agonistic interactions. The presence or ab- sence of other predators, such as fish, may also affect the microdistribution of larvae. Suhling (1999) demon- strated that small larvae of the gomphid, Onychogom- phus uncatus (Charpentier), were significantly more abundant in sand where fish predation was also lowest while medium-sized and large larvae were most abun- dant in gravel and stones, respectively. In our study, smaller larvae appeared to be associ- ated with finer sediments with a higher organic frac- tion. There was a trend supporting larval size sorting relative to sediment microhabitats; however, these re- sults must be interpreted conservatively since this pattern was evident in only one of the three streams surveyed and overall sample sizes were low. In previ- ous habitat surveys of this genus, larvae of C. biden- tata Sélys were most abundant at a mean grain size of 2.04 mm while C. heros Theischinger were most abundant at 2.79 mm (Lang et al. 2001). This is con- sistent with our finding that larvae of C. dorsalis were most abundant within medium sand sediments with a median grain size between 1.5 mm and 2.5 mm. Although C. dorsalis larvae prefer smaller sediments as a group, our evidence is inconclusive regarding microhabitat partitioning amongst larval size classes. There are several alternative explanations for the size sorting we observed between small pools. Larvae may become trapped in these shrinking habitats in a nearly random fashion with respect to size, and the apparent sorting occurs post hoc through intraspecific predation. Smaller larvae would only survive in pools that have a low proportion of late instar larva when these habitats become isolated. Alternatively, the observed size sort- ing of large and small larvae may relate to a factor that is strongly correlated with sediment grain sizes such as Current speed or oxygen levels. THE CANADIAN FIELD-NATURALIST Vol. 120 Acknowledgments Robert Cannings provided helpful commentary and review. The authors would like to thank Trent Hoover and Conan Phelan for assistance with fieldwork, and members of the Stream and Riparian Research (StaRR) lab at the University of British Columbia for comment. The authors acknowledge funding support from NSERC and the Forest Investment Initiative Account. Literature Cited Aoki, T. 1999. Larval development, emergence and seasonal regulation in Asiagomphus pryeri (Selys) (Odonata: Gom- phidae). Hydrobiologia 394: 179-192. Burcher, C. L., and L. A. Smock. 2002. Habitat distribution, dietary composition and life history characteristics of odonate nymphs in a blackwater coastal plain stream. American Midland Naturalist 148: 75-89. Cannings, R. A. 2002. Introducing the dragonflies of British Columbia and the Yukon. Royal British Columbia Muse- um, Victoria. Corbet, P. S. 1999. Dragonflies: behaviour and ecology of Odonata. Cornell University Press, New York. Ferreras-Romero, M., and P. S. Corbet. 1999. The life cycle of Cordulegaster boltonii (Donovan, 1807) (Odona- ta: Cordulegastridae) in the Sierra Morena Mountains (southern Spain). Hydrobiologia 405: 39-48. Johnson D. M., T. H. Martin, M. Mahato, L. B. Crowder, and P. H. Crowley. 1995. Predation, density dependence, and life histories of dragonflies: A field experiment in a freshwater community. Journal of the North American Benthological Society 14(4): 547-562. Kennedy, C. H. 1917. Notes on the life history and ecology of the dragonflies (Odonata) of central California and Nevada. Proceedings of the U.S. National Museum 52: 483-635. Kiffney, P. M., J. S. Richardson, and J. P. Bull. 2003. Responses of periphyton and insects to experimental manipulation of riparian buffer width along forest streams. Journal of Applied Ecology 40: 1060-1076. Lang, C., H. Miiller, and J. A. Waringer. 2001. Larval habi- tats and longitudinal distribution patterns of Cordulegaster heros Theischinger and C. bidentata Sélys in an Austrian forest stream (Anisoptera: Cordulegastridae). Odonatolog- ica 30: 395-409. Martin, T. H., D. M. Johnson, and R. D. Moore. 1991. Fish- mediated alternative life-history strategies in the dragonfly Epitheca cynosura. Journal of the North American Ben- thological Society 10: 271-279. Naiman, R. J., R. E. Bilby, and P. A. Bisson. 2000. Riparian ecology and management in the Pacific coastal rain forest. BioScience 50: 996-1011. Suhling, F. 1999. Effects of fish on the microdistribution of different larval size groups of Onychogomphus uncatus (Odonata: Gomphidae). Archiv fiir Hydrobiologie 144: 229-244. Received 14 September 2005 Accepted 15 March 2007 An Analysis of the Vascular Flora of Annapolis Heathlands, Nova Scotia S. CARBYN!, P. M. CATLING2, S. P. VANDER KLOET®, and S. BASQUILL4 ‘Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, 32 Main Street, Kentville, Nova Scotia B4N 1J5 ? Agriculture and AgriFood Canada, Environmental Health, Biodiversity, Saunders Bldg., Central Experimental Farm, Ottawa, Ontario KIA 0C6 Canada; e-mail: catlingp@agr.ge.ca *Department of Biology, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada ‘Atlantic Canada Conservation Data Centre, PO Box 6416, Sackville, New Brunswick E4L 1G6 Canada; e-mail: sbasquill@ mta.ca Carbyn, S., P. M. Catling, S. P. Vander Kloet, and S. Basquill. 2006. An analysis of the vascular flora of Annapolis Heath- lands, Nova Scotia. Canadian Field-Naturalist 120(3): 351-362. A description and analysis of the vascular plant composition of heathlands in the Annapolis valley were undertaken to pro- vide a basis for biodiversity preservation within a system of protected sites. Species presence and abundance were recorded at 23 remnant sites identified using topographic maps, air photos, and Nova Scotia Department of Natural Resources records. A total of 126 species was recorded, of which 94 were native and 31 introduced. The Annapolis heathland remnants are strongly dominated by Corema conradii with Comptonia peregrina, Vaccinium angustifolium and Pteridium aquilinum var. latiusculum. A number of species, including Solidago bicolor, Carex tonsa var. rugosperma, Dichanthelium depauperatum, Lechea intermedia, Melampyrum lineare, and Rubus hispidus, were characteristic of heathland remnants, although they usu- ally contributed little to the total cover. The most frequent alien species were Hieracium pilosella and Festuca filiformis, but Pinus sylvestris, present at 7 of 18 sites, appeared to have the greatest impact in displacing native species. Species listed as at risk and sensitive in Nova Scotia, including Helianthemun canadense, Hudsonia ericoides and Viola sagittata var. ovata, occur in open disturbed sand in the Corema heathlands. Distinctive patterns of variation occur in several species and varia- tion in crop relatives is noted with particular reference to the genera Rubus (blackberries), Amelanchier (Juneberries, Saska- toon) and Vaccinium (Blueberries). The available evidence suggests that the heathlands and sandy barrens in the Annapolis valley differ from those further west in Canada and from anthropogenic and coastal heathlands of Nova Scotia in their species composition including particularly the presence of Corema conradii, Hudsonia ericoides and Amelanchier lucida. The need to protect representative examples is supported. Key Words: Heathlands, Annapolis valley, barrens, vascular plants, flora, Corema, Nova Scotia, Canada. In 1921, legendary Harvard botanist Merritt Lyndon accords. The only descriptions of Annapolis heathlands Fernald visited the Annapolis valley. He found exten- currently available (Fernald 1921; Roland 1946; Catling sive open heathlands. He noted: “near Berwick and from there to Wilmot were vast uncultivated plains carpeted, wherever dry enough, with a close growth of the New Jersey Pine barren Corema conradii, and, ... remnants of them near Middleton” (Fernald 1921). As recently as the 1960s open heathlands with scat- tered Red Pines (Pinus resinosa) occurred for many miles along the Evangeline Trail (Figure 1). It has been estimated that in pre-settlement times the actual area of heathland encompassed approximately 200 km”. Today less than 3% of the original heathland vegeta- tion remains in the Annapolis Valley (Catling et al. 2004), and even that is threatened by loss of natural ecological processes, invasive species and conversion of the landscape (Catling et al. 2004; Catling and Carbyn 2004). Protection of this ecosystem is impor- tant for the protection of (1) insect pollinators of adja- cent crops; (2) protection of wild relatives of crops for crop improvement; (3) benchmark research examples; (4) teaching examples; (5) nature-related recreational opportunities; and (6) protection of biodiversity gen- erally in connection with national and international et al. 2004) are brief, non-quantitative, and insufficient as a basis for protection of biodiversity. Here we pro- vide a description of remnants of natural heathland in the Annapolis Valley along with an indication of dom- inant species, rare and significant species, variation between sites and relationship to other eastern Cana- dian sand barrens. This is designed to provide a basis for further study and for the establishment of a system of protected sites. Methods Sites and data The study area consists of 23 sites in the Annapolis Valley (Table 1, Figure 2). Sites are defined as areas surveyed separated by at least 0.5 km. Information from the Nova Scotia Department of Natural Resources and topographical maps were used to determine the most probable locations of heathland vegetation occur- ring on sandy soil. At sites 1-19 abundance of vascular plants was recorded. Areas surveyed varied from approximately 0.561 hectares in extent. The majority of these 19 sites 351 B52 THE CANADIAN FIELD-NATURALIST Vol. 120 FiGurE |. Open heathland dominated by Corema conradii (Broom Crowberry) with Pinus resinosa (Red Pine). Photo by P. M. Catling in 2004 at site 8. were visited on at least two occasions in 2003, with one visit in early summer and another in autumn. Ap- proximately two hours were spent at each site on each visit. Species lists were made at each site (Table 2). An abundance value ranging from 0-5 was assigned to each species (1 = rare, 2 = uncommon, 3 = com- mon, 4 = frequent and locally dominant, 5 = dominant) based on consensus of two or three observers. Although some larger sites were visited more often, almost all species recorded were recorded in the first hour of two visits. Time spent at sites was therefore considered to be adequate and the lists are thought to be essentially TABLE 1. Site number and location of heathland study sites in the Annapolis Valley, Nova Scotia. Site Number, Name Evangeline Heathland E of 201 Evangeline Heathland W of 201 Trail Heathland Evangeline Main Heathland Brooklyn Street Wildlife Sanctuary Caribou Bog Heathland remnant Trailer Park Heathland Exit 17 Heathland, NE corner Exit 17 Heathland, NW corner 4 km W of Middleton W side of Middleton Evangeline Cemetary 13 E side of Greenwood Military Base Aldershot Military Base behind shooting range Greenwood section Exit 17 far east Greenwood Military Base main site Greenwood Military Base satellite 201 East near bog Plot — W of Whitman Road Plot — E of route 201 Plot — N of route 201 23 Plot — S of Ward Road BSR K OMNAAUNHKWN Ne Oo a OMADNFE ~~) ie) (=) i) N Latitude ° Longitude ° 45.0059 —64.8882 45.0048 —64.8930 45.0005 —64.9004 45.0094 —64.8705 45.0872 —64.5245 45.0377 —64.783 45.0219 —64.8255 44.9992 —64.9411 44.9962 —64.9509 44.9204 —65.1048 44.9607 —64.9999 44.9675 —64.9866 44.9896 —64.8955 45.0966 —64.5333 44.9833 —64.8873 44.9974 —64.9470 44.9801 —64.9381 44.9853 —64.9308 44.9791 —64.9366 45.0109 —64.8404 44.9992 —64.8854 44.9472 —65.0203 44.9681 —64.9367 ree a ——————eeeesss 2006 Heathland Sites Surveyed in CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS 353 Annapolis and Kings Counties Nova Scotia 20 Kilometres 65°0'W 64°50'W Legend A Sites Inventoried (19) @ Plots Evaluated (4) Major Roads ——— Minor Roads —™- County Boundary DW sandy soil 64°40'W 64°30'W 64°20'W FiGuRE 2. Heathland sites surveyed in the Annapolis Valley in 2004 (solid triangles) and location of plots evaluated for cover in 2005 (dots). The sandy soils of the valley (shaded) suggest the maximum extent of heathland. complete. Voucher specimens collected at various sites were deposited in the vascular plant herbarium at Agriculture Canada in Ottawa (DAO). Status of plant species was determined by using Maher et al. (1978), the recent listing of species at risk (Nova Scotia De- partment of Natural Resources 2001*) and informa- tion and lists on file at the Atlantic Conservation Data Centre in Sackville. At sites 20—23, plant association sampling methods were employed following provincial plot standards (Quigley et al. 2005*) and Canadian vegetation clas- sification conventions (CNVC Technical Committee 2004*). In May of 2005, at each of the four sites, a four hundred square meter plot was subjectively placed in a homogeneous tract of heathland vegetation, where can- opy tree cover exceeded 10 percent. The sampled asso- ciation was the open woodland stage of the Annapolis Valley sand plain. Species and cover values, represent- ing percentage of plot area within a vegetation layer, were visually assessed for five life forms (Table 3). Lichen names follow Esslinger (1997*), bryophyte names follow Anderson et al. (1990), and vascular plant names are from Kartesz and Meachum (1999). Although many voucher specimens were collected (and deposited at DAO), a sufficient number of speci- mens of the genus Aronia were not collected to enable a determination of which species (or hybrid) was pres- ent at a particular site. Thus only the genus name ap- pears in Table 2. Although Rosa carolina and Rosa vir- giniana were noted, both may have been present at any site where either was recorded, and some intermediates were noted. Rubus hispidus may be over-represented in the survey and Rubus arenicola may be under-rep- resented due to difficulties in distinguishing these, and hybrids between them may also have been present. Results and Discussion. Native species and limitations of the vegetation description Although 104 native species are recorded from the 23 sites surveyed, and although these sites covered a rather extensive area, they were mostly drier examples of dry, open sandy habitats (Figure 1). The relatively short-lived wetter examples and those associated with natural disturbances, such as fire, are now much less common than the drier examples. Thus as a reconstruc- Vol. 120 ia! 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Cc ANDMMNMNEAMNAMUOAMATAANAErAOARMH AMAA mAANNO TOC CO e— A ee A | 81 LI I Ol v LAN NAN onc mANN eo N Lao l(Q.0} r=GN! maa | Ama M (oe ioe) SI trot tl el NaN Cl IT Ol ioe) om "Jey Spasog syouaIAy dAOT CV (7) wnopisayy] wnyoiokyducs x] DAUIWUIDAS DIADIJAIS “Jey Déaov] Sayupsids loyng vqjp vavsids yur] vipuijoad puysvds "J 40]091g OSvpyjos ‘qsl[es (]) pyofiunupss osppijos ‘WV Sypsouau oOsvpiyos "| SisuapduDdd OsDpI]OS ‘Wy vaounl osppyos ‘WNN Vjynsagnd osppyos 4 7 DUulYyAUD auapls 4] adap winpas Ysieyy SiMuny X1d¢ x] DIJasojaow XAUNY 3 REA PIPED] “your|g vjooiuasiw snqny “YOuR|g SMUDJUOULIAA SNQNY “"T snpidsiy snqny IQIOg Sisuaiuaysayy snqny ‘| DUIIOADI DSOY ‘THA Dupiuis.1iA DSOY “MOL (J) asuappuvs vospuapopoyy 4 qT unasiuiydos snupydvy "J DANA SNIAANO “MIDPUL) (ASA) WHNISNYY] “eA UY] (TY) Wnuapnby wwipisalg ‘J 7] voiupajAsuad snunig “YIU DUIOIAS SNUNAG “] puplUis.slA SNUNA “XY Xajduns ppjyuajog x] DI9Ad DIYUajog “TT Sisuapoupds DII1WUAIOg x. DalvasID DIJYUA Og “XYOI] Saplojniuas snjndog “XY Vipjuapipuvss snjndog ‘ds wnuosKjog 4 7] DSSAadUO) DOJ “7 sisuajpid DOg DURA] IYUSIIS “IDAOO [P}O}] = DI ‘SAIS [210] = SL psonponyy = y. ‘JURUTUIOp = ¢ ‘JUeUTWOp AT[ROO] pue JUONbayy = p ‘UOWWOD = ¢ ‘UOWOOUN = Z ‘areI = | “AaT[eA stjodeuuy oy) Ur sojIs pueyyeoy 6] IV sNyeIs puke doURpUNge saloedg (panuiUo) *Z ATAVL 2006 CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS 357 Tne a eee tion of the Annapolis sand barren flora, the mesic and LE 3 5 — - successional stages of dry barrens are to a large ex- a AN Ee eae tent omitted. This may explain the lack of some species 8g = = = in the survey, such as Agalinis neoscotica (Greene) a Sale cence Si-estel | {ual Fern. (Middleton False Foxglove), which was des- Mt; cribed from Annapolis heathlands near Middleton, i Ren ete!) fy ule=te| and also Bartonia virginica (L.) BSP., Carex atlantica, 5 Polygala sanguinea L. and Sisyrinchium fuscatum be Sia) Pad be tw Bickn. All of these species occur on heathlands near | Middleton (personal observation), but were not seen s are tet. Tec aril during the present survey. Their abundance in the area q in the past (Fernald 1921, page 138) suggests that the e SMT ott [last evo | more mesic and naturally disturbed areas were much 3. x more prevalent only a short time ago. Fernald (1921) & al mureretaworce, hel (lah referred to these as the “the damper Polytrichim-car- At lc Ae ere peted areas.” He noted that “Bartonia virginica was | ‘i everywhere,” but it was not recorded in any of the bar- é SAE IP cont cle legen rens in our survey and only our site 10 approached this S damper sand barrens habitat. i Praline gh edt Te Me Lipeile? 4) In addition to the lack of mesic sites it is of interest a Mra ety cca. eee that in two days in 1920 Fernald (1921) recorded two S species from the drier heathlands near Middleton that : NOM iaetor lh cetier re | were not seen by us. These were Potentilla tridentata 2 || 2 and Pyrola rotundifolia. Although our data indicate the general and dominant composition of the Annapolis Sees ACO ily lett aly wilieel heathlands, the differences with Fernald’s brief survey suggest that it may never be possible to have a com- ah ee plete knowledge of their former floriste composition. ase bet eit (Cea ah T SN lal val Dominant vascular plants Relatively few of the 126 species (94 were native le de et and 31 introduced — Table 2) present in the 19 com- ME od Pasian gett es pletely surveyed sites were dominant in the vegetation. Species present at all sites included Corema conradii, Qu} Sp th tssh st Danthonia spicata, Pinus strobus and Solidago bicolor. Species present at 18 of the 19 sites were Amelanchier laevis, Carex tonsa var. rugosperma, Comptonia peregi- na, Dichanthelium depauperatum, Pteridium aquilinum var. latiusculum, Vaccinium angustifolium and Populus tremuloides. Species with the highest overall cover values included Corema conradii with 91 followed by Comptonia peregrina with 59, Preridium acquilinum var. latiusculum with 57, and Vaccinium angustifolium with 56. A number of species, including Solidago bi- TABLE 2. (continued) Species abundance and status at 19 heathland sites in the Annapolis Valley. 1 = rare, 2 = ie) tal 2 ¥ : color, Carex tonsa var. rugosperma, Dichanthelium = ue S depauperatum, Lechea intermedia, Melampyrum lin- iS) S & = eare and Rubus hispidus, were characteristic of heath- M 2 eS > land remnants, although they were rarely dominant. 2 = aS 5 The most frequent alien species were Hieracium pilo- g See e & = S sella and Festuca filiformis, both at 17 and 16 (respec- ‘@ oe 5 3 S Ws tively) of 19 sites. The most serious invasive of the = igs 2 SE oS Se SND heathlands in terms of displacing native species was 1 cae SS Sa PaaS Pinus sylvestris, present at 8 sites (Catling and Car- a Beeess2 ss byn 2004). 3|}/2| S888 = a8 "s S “s 3 The more wooded plot sites (20-23) gave a very sim- 3 > S 3 S SESES SS 25 ilar picture of the Corema-dominated heathland, but eels 5 s g§ : gs Soe : with scattered trees, mostly Pinus resinosa (Table 3, 15 || SRS RSS SSO) iss Figure 3). Three species more often associated with Tee = g S S § Sees S85 woodland than with barrens (Clintonia borealis, Lyco- *HalRRSSSSSSS5 BAe = 358 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 3. Cover values for vegetation layers in single 400 square meter plots at each of 4 sites in the Annapolis valley, based on data collected by Sean Basquill in May 2005. * = introduced, + = not listed in Table 2. Species Site 20 Site 21 Site 22 Site 23 Trees Amelanchier laevis Wieg. 2 = 2 = Betula populifolia Marsh 3 2 5 5 Picea mariana (P. Mill.) B.S.P. = ES 5 £ Pinus resinosa Ait. 20 25 20 10 Pinus strobus L. — = 5 8 Pinus sylvestris L. * = = = 10 Prunus pensylvanica L. f. - 1 = = Quercus rubra L. = 4 0.1 Es Shrubs and small trees Amelanchier laevis Wieg. = ~ : Betula populifolia Marsh 3 2 Sal 3 Comptonia peregrina (L.) Coult. 1 1 Juniperus communis L. var. depressa Pursh ~ - _ 0.1 Picea mariana (P. Mill.) B.S.P. - 0.1 - Pinus resinosa Ait. 4 + 1 Pinus strobus L. 0.5 1 Pinus sylvestris L.* - - - Quercus rubra L. — 1 - _ Vaccinium angustifolium Ait. 0.1 0.5 3 6 Herbs and dwarf shrubs Arctostaphylos uva-ursi (L.) Spreng. 30 5) 10 8 Betula populifolia Marsh = Carex tonsa (Fern.) Bickn. (sensu lato) 0.01 0.1 1 1 Clintonia borealis (Ait.) Raf. + - = 0.01 - Corema conradii (Torr.) Torr. ex Loud. 58 40 60 60 Deschampsia flexuosa (L.) Trin. — - 1 1 Dichanthelium depauperatum (Muhl.) Gould - - 5 0.1 - Epigaea repens L. — - 0.01 0.1 Festuca filiformis Pourret * 0.1 - Festuca trachyphylla (Hackel) Krajina * - 0.001 - - Gaultheria procumbens L. 0.5 0.01 - Hieracium piloselloides Vill. * - - 0.01 - Hudsonia ericoides L. 0.01 0.1 - 0.3 Kalmia angustifolia L. 0.1 0.01 - = Lechea intermedia Leggett 0.001 - = = Lycopodium obscurum L. + = = Lycopodium tristachyum Pursh 0.1 0.01 - Maianthemum canadense Desf. - - 0.01 - Oryzopsis asperifolia Michx. + — 0.1 0.2 = Poa pratensis L. - 0.1 0.1 0.1 Potentilla simplex Michx. - = - 0.2 Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw. 10 1 0.02 - Rubus hispidus L. - 0.001 - 0.01 Solidago bicolor L. - 0.2 = = Solidago nemoralis Ait. = 0.001 0.2 0.1 Taraxacum officinale G. H. Weber ex Wiggers - Trientalis borealis Raf. 0.01 = 0.01 Vaccinium angustifolium Ait. = Mosses Ceratodon purpureus (Hedw.) Brid.+ - 0.001 — 2 Dicranum scoparium Hedw.+ ~ 1 — = Pleurozium schreberi (Brid.) Mitt.+ 0.5 0.1 0.5 = Polytrichum commune Hedw.+ = — 0.1 1 Polytrichum juniperinum Hedw.+ - 0.01 0.3 3 Lichens Cladina rangiferina (L.) Nyl.+ 18 40 i 10 Cladina stellaris (Opiz) Brodo + 2 10 3 a a ————————————————————E 2006 Se ae nr % CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS 359 FIGURE 3. Open wooded heathland dominated by Corema conradii and Cladina rangiferina with scattered Pinus resinosa. Although there is much open area, the site exceeds the threshold 10% tree cover required for classification as a treed association under Canadian vegetation classification conventions (CNVC Technical Committee 2004*). Photo by S. Basquill on 25 May 2005 at site 20. podium obscurum and Oryzopsis asperifolia) were present a these more wooded sites. Although vascular plants were the focus of the descriptive work, the plot data included lichens and bryophytes, suggesting the former (Cladina rangiferina and C. stellaris) to be a significant component. This suggestion is supported by general observations at the 19 other sites where quan- titative data on bryophytes and lichens was not ob- tained. Although Roland (1946) described the Annapolis sand barrens and heathlands 60 years ago, his descrip- tion, which recorded 35 species, corresponds very closely to the composition seen in the relicts that re- main today. The successional processes are possibly also the same as described generally by Roland, but succession may have been much more rapid in some areas than in others where barrens and heathlands may have existed as a subclimax lasting for centuries, even without fire. Roland’s article was written at a time when sand barrens, heathlands and savanna were regarded as wasteland rather than as special places for native biodiversity, but his strong forestry theme is accompa- nied by many astute observations regarding the native flora. At the time of early settlement the barrens and heathlands may have expanded due to cutting and burn- ing of pine forests and abandonment of sandy land cleared for agriculture as suggested by Roland. How- ever there is little doubt that they also existed in pre- settlement times. Rare and significant species The only species considered to be at risk in Nova Scotia that occurs in the Annapolis heathlands is Helianthemum canadense (Long-Branch Frostweed, Figure 4). It was found at 6 of the sites, always in dis- turbed habitats with some bare sand. Both Viola sagit- tata var. ovata and Hudsonia ericoides are listed as sensitive (Nova Scotia Department of Natural Re- sources 2001*), and these also occurred in disturbed sandy areas (Figure 5). The provincially rare (Maher et al. 1978) Sisyrinchium fuscatum Bickn. (Coastal- Plain Blue-Eyed-Grass) was reported from “sandy areas near Middleton” (sub Sisyrinchium arenicola Bickn., Roland and Smith 1969), but was not seen during the present survey. In addition to the rare and threatened species there is a suggestion of some distinctive patterns of varia- tion due to taxonomic recognition of several variants. Among these is a sand barren ecotype of Amelanchier lucida (personal observation). Taxa of blackberries (Rubus particeps, R. arenicola), although not recog- nized as discrete in some recent literature, have been reported from the Annapolis heathlands suggesting at 360 least the presence of distinctive genetic variants with- in broadly defined species. Further study may indi- cate both of these to be worthy of taxonomic recog- nition. A restricted glabrous variant of Viola sagittata var. ovata (f. glabrata) occurs on the dry open sand with the more typical form. Three taxa have been described from Annapolis heathlands: Amelanchier lucida Fer- nald, Dichanthelium (sub Panicum) depauperatum vat. psilophyllum f. cryptostachys Fernald, and Rubus par- ticeps Bailey. The heathland ecosystem is particularly valuable as a reservoir of genetic variation in crops and crop rel- atives. Fernald (1921) commented on the remarkable variation in wild blueberries (Vaccinium angustifolium and V. myrtilloides) in Annapolis heathland remnants near Middleton. In addition to blueberries (Table 2) there are potential genotypes of Aronia (Aronia sp.), of blackberries (Rubus hispidus and others), juneber- ries (Amelanchier lucida and A. laevis), huckleberries (Gaylussacia baccata and G. dumosa), cherries (Pru- nus pensylvanica, P. virginiana, and P. serotina) and a strawberry (Fragaria virginiana). Variation between sites Although all sites shared dominant species (Table 2) they varied in diversity from 26 to 57 native species. Much of the variation between sites appeared to be attributable to disturbance and soil moisture. Those sites with lower lying and periodic moist areas and open sand had the highest vascular plant diversity. Site 10 for example was the most unlike the other sites in native species composition, possibly a result of parts of it having a relatively high water table and disturbed areas of periodically moist sand where dry ground species such as Carex tonsa var. rugosperma and Danthonia spicata occurred with wetland species such as Drosera intermedia. Characteristic native species and “at risk” or “sen- sitive” species were present at some sites but absent from others (Table 2). Sites also varied in the extent to which alien species were present and dominant (Table 2). For example parts of sites 1 and 4 had extensive and spreading stands of Pinus sylvestris, but displace- ment of native vegetation by this introduced tree was either less extensive or not observed at other sites. Festuca filiformis was a co-dominant at site 5 but not elsewhere. Species presence, diversity and extent of impacts all require consideration in selecting sites for protec- tion. The variation between sites in composition and abundance, including that of rare and/or characteristic species, suggests that protection of a number of sites will be necessary to protect representative ecosystem components. Relationship to other heathlands and barrens In Canada, Corema-dominated barrens are charac- teristic of the maritime region. Corema conradii has a restricted distribution extending from the Gulf of THE CANADIAN FIELD-NATURALIST Vol. 120 ~ af FIGURE 4. Helianthemum canadense (Long-Branch Frost- weed), a native plant at risk in Nova Scotia, with its main Nova Scotian occurrence on the Annapolis valley heathlands. Photo by P. M. Catling in 2004 at site 8. St. Lawrence region south to New Jersey. It does not occur in sand barrens further to the west in Canada, for example in the Ottawa valley, where Vaccinium angus- tifolium is the dominant heath shrub accompanied by other shrubs such as Prunus susquehanae and Comp- tonia peregina (Carbyn and Catling 1995). Other spe- cies present in the Annapolis heathlands but absent in Ottawa valley sites were Amelanchier lucida, Des- champsia flexuosa, Rubus hispidus and Solidago bi- color. Among the prevalent species in Ottawa valley sand barrens but absent in the Annapolis sites were Carex siccata, Carex lucorum, Dichanthelium sabu- lorum var. thinium, Polygonella articulata and Prunus susquehanae (Carbyn and Catling 1995). Within Nova Scotia, “barrens” with heath vegeta- tion occupied a large part of the western portion of the province (Strang 1972). Some of these barrens are a short-lived successional stage following fire, whereas in other cases they are long persisting. Those of short duration have in some cases been produced by cut- ting and then maintained by fires. Two such barrens were described by Hall and Aalders (1968). Such bar- rens, produced by human activities, have been consid- ered a degraded landscape of little economic value. The existence and importance of apparently natural bar- rens have only recently become apparent (Catling et al. 2004). The sites described by Hall and Aalders (1968) differ from those in the Annapolis Valley in lacking Corema conradii, the dominant of the Annapolis bar- rens. They were also without several other species, including Hudsonia ericoides. On this basis the dif- ferences between man-made and natural barrens seem 2006 CARBYN, CATLING, VANDER KLOET, and BASQUILL: ANNAPOLIS HEALTHLANDS 36) FIGURE 5. Open sandy ground in a disturbed area of heathland with Comptonia peregrina (Sweet Fern) and Viola sagittata var. ovata (Arrow-leaved Violet). Photo by P. M. Catling in 2004 at site 8. pronounced, but with only two anthropogenic sites, a more detailed comparison is unnecessary. The apparently longer persisting natural barrens in Nova Scotia are readily divided into two major vegeta- tion types based on either granitic or sandy substrates (personal observation). Sandy heathlands like those of the Annapolis Valley were also well developed in the interior of Yarmouth County and near Debert, Spring- hill, Parrsboro, and on coastal dunes in Guysborough and Kings counties and on Sable Island. The coastal heathlands on sand differ from the interior sites in both species composition and species abundance, and the differences suggest adaptation to different conditions. Differences also exist at the infraspecific level. For example Lechea intermedia vat. intermedia occurs in interior heathlands while L. intermedia var. juniperina occurs on the coastal sandy heathlands. The heathlands on Sable Island appear distinct (Catling et al. 1985). The interior sandy heathlands also appear to vary re- gionally, presumably due to differences in climate. The climate in southern Yarmouth and Shelburne differs from that of the Annapolis Valley (personal observa- tion). The data suggest that the Annapolis heathlands are distinctive in their floristic composition. Acknowledgments Mark F. Elderkin, Species at Risk biologist with the Nova Scotia Department of Natural Resources, assisted with current information on status ranks. Lawrence Benjamin and Randy Milton, also of the Nova Scotia Department of Natural Resources, provided informa- tion on site locations for rare plant species in the An- napolis Valley. Steve Javorek and M. Grant assisted with field studies (site 1). Geoff Mercer assisted with the inventory of sites on military bases at Aldershot (site 14) and Greenwood (sites 17 and 18). Documents Cited (marked * in text) CNVC Technical Committee. 2004. Plant association con- cept for the Canadian National Vegetation Classification. NatureServe Canada. 5 pages. http://www.cnve-cnvc.ca. Esslinger, T. L. 1997. A cumulative checklist for the lichen- forming, lichenicolous and allied fungi of the continental United States and Canada. North Dakota State University: http://www.ndsu.nodak.edu/instruct/esslinge/cheklst cheklst7.htm Nova Scotia Department of Natural Resources. 2001. General status ranks of wild species. Vascular plants. http: www.gov.ns.ca/natr/wildlife/genstatus/specieslist.asp Quigley, E., P. Neily, K. Keys, and B. Stewart. 2005. Nova Scotia Department of Natural Resources Forest Ecosys- tem Classification; Methods for Plot Establishment and Vegetation Inventory. Unpublished Internal Document. Literature Cited Anderson, L. E., H. A. Crum, and W. R. Buck. 1990. List of the mosses of North America north of Mexico. Bryol- ogist 93: 448-499. 362 Carbyn, S. E., and P. M. Catling. 1995. Vascular flora of sand barrens in the middle Ottawa valley. Canadian Field- Naturalist 109: 242-250. Catling, P. M., and S. Carbyn. 2004. Invasive Scots Pine (Pinus sylvestris) replacing Corema heathland in the Anna- polis valley. Canadian Field-Naturalist 119(2): 237-244. Catling, P. M.,S. Carbyn, S. P. Vander Kloet, K. MacKen- zie, S. Javorek, and M. Grant. 2004. Saving Annapolis Heathlands. Canadian Botanical Association Bulletin 37(1): 12-14. Catling, P. M., B. Freedman, and Z. Lucas. 1985. Vegeta- tion and phytogeography of Sable Island, Nova Scotia. Proceedings of the Nova Scotia Institute of Science 24: 181-248. Fernald, M. L. 1921. The Gray Herbarium expedition to Nova Scotia. Rhodora 23: 130-171. Hall, I. V., and L. E. Aalders. 1968. The botanical compo- sition of two barrens in Nova Scotia. Le Naturaliste cana- dien 95: 393-396. THE CANADIAN FIELD-NATURALIST Vol. 120 Kartesz, J. T., and C. A. Meachum. 1999. Synthesis of the North American Flora, version 1.0. North Carolina Botan- ical Garden, Chapel Hill, North Carolina. Maher, R. V., D. J. White, G. W. Argus, and P. Keddy. 1978. The rare vascular plants of Nova Scotia. Syllogeus (National Museums of Canada, Ottawa) 18.37 pages. Roland. A. 1946. The vegetation of the Annapolis valley I. — well drained sand areas. Acadian Naturalist 2(7): 1-20. Roland, A. E., and E. C. Smith. 1969. The flora of Nova Scotia. Nova Scotia Museum. Reprinted from Proceedings of the Nova Scotian Institute of Science 26: 1-238, 277- 743. Strang, R. M. 1972. Ecology and land use of the barrens of western Nova Scotia. Canadian Journal of Forest Research 2: 276-290. Received 25 September 2005 Accepted 17 February 2007 Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and Scavenger Activity on Tethered Carrion in Subtidal Benthic Communities off Western Vancouver Island SARAH Davies!, ALI GriFFiTHs!, and T. E. REIMCHEN? 'Bamfield Marine Sciences Centre, Bamfield, British Columbia VOR 1BO Canada *Department of Biology, P.O. Box 3020, University of Victoria, Victoria, British Columbia V8W 3N5 Canada; e-mail: reimchen@ uvic.ca (corresponding author) Davies, Sarah, Ali Griffiths, and T. E. Reimchen. 2006. Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and scavenger activity on tethered carrion in subtidal benthic communities off Western Vancouver Island. Canadian Field-Naturalist 120(3): 363-366. The influence of pelagic carrion food falls on marine benthic scavenging communities was investigated at two depths (10 m, 50 m) in Barkley Sound, west Vancouver Island, British Columbia from 12 May to 4 June, 2003. A remotely operated vehicle (ROV) equipped with video cameras was used to monitor anchored carrion (15 kg pig leg) during daylight and darkness. The videos were subsequently analyzed for species diversity, abundance and the intensity of scavenging. At 10 m, Redrock Crab (Cancer productus) and Kelp Greenling (Hexagrammos decagrammus) dominated, while at 50 m, Spot Shrimp (Pandalus platyceros), Spotted Ratfish (Hydrolagus colliei) and Pacific Hagfish (Eptatretus stoutii) were the dominant species, most of which were nocturnal. Hagfish were the major consumers of the carrion and after 23 days, no soft tissues remained at 50 m while 40% remained at 10 m. Within 24 hours of the carrion deployment, two of eleven ratfish succumbed, probably due to the direct clogging effects of hagfish mucus on the respiratory apparatus of the ratfish. These field observations are consistent with laboratory results suggesting high efficacy of hagfish mucus in competitive interactions. Key Words: Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, Spot Shrimp, Pandalus platyceros, Redrock Crab, Cancer productus, Kelp Greenling, Hexagrammos decagrammus, marine scavengers, carrion, remotely operated vehicle (ROV), nutrient cycling, Vancouver Island, British Columbia. Vertebrate carcasses provide an episodic but highly _ important food source for benthic organisms, particu- - larly at depths beyond the zones of primary produc- tivity (Lalli and Parsons 2001). Such carcasses have _ recently become of interest due to increasing discards _ in fisheries as this may shift benthic community struc- ture (Collins et al. 1999). Species such as ratfish and _ benthic sharks as well as crustaceans may benefit from such food falls (Collins et al. 1999; Tamburri and Barry _ 1999). There are few descriptions of the scavenging community or the interactions among species around _ these carcasses. We anchored carrion at two depths (10 m, 50 m) in _ Barkley Sound, west Vancouver Island, British Colum- bia and monitored the carrion with a remotely-operated vehicle (ROV). We provide here an examination of ' scavenger diversity, abundance and interactions among » species during daylight and darkness. \ ‘Methods Two soft-substrate sites (10 m, 50 m: 48°50'N, -125°10'W) in Mackenzie Anchorage, Barkley Sound, British Columbia were chosen and single 15 kg pig legs (frozen with intact skin) were anchored at each ‘site. Initial deployment occurred at 1030 h on 12 May 2003. An ROV (Deep Ocean Engineering Phantom HD2+) equipped with 5mW 635 nm lasers (10 cm Standard for scaling) and 250W Tungsten-Halogen lights were used to monitor the carrion. The ROV was stationed beside the carrion for 30 minutes during night trials and 15 minutes (due to less activity) during day trials. We switched the ROV lights off for 1-2 minutes on several occasions during the night monitoring phase but observed no changes in activity. Before surfacing with the ROV, areas within 5 m of carrion were scanned for individuals. Day trials of the ROV were carried out near 1400 h and replicated on three occasions (13, 14, 16 May) while night trials occurred near 2230 h and were replicated on four occasions (12, 13, 14, 23 May). The experiment was terminated on 4 June and the two carcasses returned to the surface and relative tissues were estimated. Twelve hours of videotape were ana- lyzed. Percent of remaining carrion and sizes of scav- engers were estimated from the laser standards. Species were identified and all individuals within several metres of the carrion were counted. General behavior of the species was noted. Paired sampled t-tests were used to test for significant differences for species and abun- dance between sites and within sites. Results Species Abundance Carrion anchored at the two depths attracted a dif- ferent assemblage of species (Figure 1). At 10 m, Redrock Crab (Cancer productus) (7-20 cm), and Kelp Greenling (Hexagrammos decagrammus) (11-23 cm) 363 364 Cumulative Abundance: Depth 10m Cumulative Abundance Species THE CANADIAN FIELD-NATURALIST Vol. 120 Cumulative Abundance: Depth 50m Cumulative Abundance Species FIGURE 1. Cumulative species abundances of scavengers observed at carrion anchored at 10 m and 50 m in Barkley Sound, Vancouver Island. were the dominant species, while at 50 m, Long-rayed Star (Stylasterias forriei) (20-60 cm), Spot Shrimp (Pandalus platyceros) (12-20 cm), Spotted Ratfish (Hydrolagus colliei) (18-25 cm), Pacific Hagfish (35- 65 cm) and Rocksole (Lepidosetta bilineata) were dominant. Only two of 16 species (Spotted Ratfish, Kelp Greenling) were common to both sites (Figure 2). Although all species were in direct contact or imme- diately adjacent to the carrion, only six species active- ly foraged off the carrion (Pacific Hagfish, Redrock Crab, Dungeness Crab (Cancer magister), Sunflower Star (Pycnopodia helianthoides), Long-rayed Star (Orthasterias koehleri), and Spot Shrimp). Hagfish undertook extensive burrowing and excavations and were the major consumers of tissues. Ratfish swam commonly in close vicinity to the carrion but we observed no foraging activity. Daily consumption rates at 10 m were initially high and then decreased while at the 50 m site, the rate increased throughout the duration of sampling. On the last ROV survey (Day 23), 40% of the soft tissues remained at the 10 m site while only bone remained at the 50 m site. Diel variations The scavenger community differed during daylight and darkness (Figure 3). At both sites there were in- creased numbers of species observed during darkness (10 m Day, 5 , Night, 11; 50 m Day, 5, Night, 9). At 10 m depth, nocturnal scavengers were Redrock Crab, Spotted Ratfish, Roughback Sculpin (Chitonotus pugetensis), and Dungeness Crab. At 50 m, Long rayed Star was present in both day and night trials and was seen continually in all surveys. Northern Ronquil (Ron- quilus jordani), Rocksole and Kelp Greenling were seen only during day trials. Pacific Hagfish, Spotted Ratfish, and Spot Shrimp were dominant at night. Behavioural interactions within and among scavenger species Diverse interactions occurred among species. Red- rock Crabs defended the carrion by direct attacks against conspecifics as well as against Kelp Green- ling. Pacific Hagfish excavated burrows within the carrion by rasping, full-body spinning and occasional knotting. Spotted Ratfish swam in close vicinity to the carrion and in several instances directly contacted the hagfish but we saw no evidence of agonistic behaviour. However, during the second ROV flight, we observed two freshly deceased ratfish lying with- in several meters of the carrion. There was no evi- dence of external damage to either ratfish but there was extensive subsurface haemorrhaging around the mouth on one ratfish and strands of mucus on the second and we infer rapid respiratory failure from the hagfish mucus that was prevalent in the water column around the carrion. During the ROV survey, hagfish did not feed on dead ratfish. Strands of mucus that separated from hagfish when feeding on the carrion were very adhesive as the strands disabled thruster motors of the ROV. Discussion High productivity and the recycling of nutrients sus- tain large populations of benthic organisms in the waters of the continental shelf (Nybakken 2001). We examined an experimental food fall at two (10 m, 50 m) natural habitats in coastal British Columbia and found that species assemblages differed between depths and with diel variation. The intensity of scavenging was greater at increased depth and increased during night, which corroborates well with other observations on scavenging communities (Collins et al. 1999). Shifts in diel patterns to nocturnal activity are common in marine 2006 DAVIES, GRIFFITHS, and REIMCHEN: SCAVENGER ACTIVITY 365 !Om/ 50m Abundance Difference FiGuRE 2. Relational abundance of scavenger species at carrion anchored at 10 m and 50 m depth (number of individuals at 10 m — number of individuals at 50 m depth), Barkley Sound. Positive values show greater occurrence at 10 m depth. Negative values show greater occurrence at 50 m depth. 10m Day/Night Abundance Difference Species FiGuRE 3. Relational abundance of scavenger species due to diel variation (number of individuals during daylight — number of individuals during night). Positive values show greater occurrence during daylight. Negative values show greater occurrence during night. benthic habitats (Lalli and Parsons 2001) and may reflect predator avoidance or reduction in competition. The agonistic encounters observed among crabs and fish indicate that carrion represents significant local energy enrichment to the benthic community (Stock- ton and DeLaca 1982). 50m —s Day/ Night Abundance Difference Species Spotted Hagfish were the major scavengers at 50 m and were largely nocturnal, consistent with the find- ings of Fernholm (1974). These fish play an important role in the turnover and cycling of nutrients (Lesser et al. 1996: Collins et al. 1999). We observed a much higher rate of tissue loss at the 50 m site where hagfish 366 were prevalent. Since hagfish are dependent on food falls, specialized adaptations to minimize competition are present. Knotting is considered to be an important feeding behavior in which the hagfish attaches to prey with tooth plates and passes a knot from the caudal region anterior towards the head allowing tissues to be ripped off (Collins et al. 1999). We also observed this behavior, but only on a single occasion. Rather, hagfish primarily used full body spinning to pull pieces of tis- sue from the carrion. Such full body rotation is also present in sharks feeding on large prey and may allow an increase in their effective gape (Helfman et al. 1997). Hagfish produce copious amounts of slime when feeding, agitated or threatened (Strahan 1963; Tam- burri and Barry 1999). There are probably diverse functions to this slime including burrowing, defense and competitive exclusion (Collins et al. 1999). Hag- fish burrowed into the carcass up to % of their total body length. While the mucus may facilitate entry into the carrion, the burrowing would also cover the gills and this would limit gas exchange of the hagfish. This suggests the potential for posterior cutaneous respira- tion. Lesser et al. (1996) noted that hagfish skin was well-vascularized and hagfish are therefore able to sur- vive with plugged nostrils and no respiratory current across the gills. Slime produced by hagfish has high strength (Fudge et al. 2003) and can also obstruct mouthparts and gills of other species attempting to feed at food falls (Tamburri and Barry 1999; Collins et al. 1999). Our ROV observations are consistent with these suggestions. Two of the ratfish swimming in close prox- imity to the foraging hagfish quickly succumbed, pre- sumably from loss of respiratory ability. The close prox- imity suggests high efficacy and rapid effects of the hagfish mucus in natural conditions even under high dilution effects. If these observations are representa- tive, it suggests that hagfish may not only have a sub- stantial competitive advantage over other fish species in these episodic carrion fallouts but also operate as predators through the lethal influences of mucus on other scavengers. THE CANADIAN FIELD-NATURALIST Vol. 120 Acknowledgments We thank Nimrod Levy for discussion, James Mor- timer, John Richards and Janice Pierce for ROV and vessel support, J. Yakimishyn and J. P. Danko for lab and technical support, and the Bamfield Marine Sci- ences Center for funding this project. TER acknowl- edges support (A2354) from the Natural Sciences and Research Council of Canada. Literature Cited Collins, M. A., C. Yau, C. P. Nolan, P. M. Bagley, and I. G. Priede. 1999. Behavioral observations on the scavenging fauna of the Patagonian Slope. Journal of the Marine Bio- logical Association of the United Kingdom 79: 963-970. Fernholm, B. 1974. Diurnal variation in the behavior of the hagfish Eptatretus burgeri. Marine Biology 27: 351-356. Fudge, D.S., K. H. Gardner, V. T. Forsyth, C. Riekel, and J. M. Gosline. 2003. The mechanical properties of hy- drated intermediate filaments: Insights from hagfish slime threads. Biophysical Journal 85: 2015-2027. Helfman, G.S., B. B. Collette, and D. E. Facey. 1997. The diversity of fishes. Blackwell Science, London. Lalli, C. M., and T. R. Parsons. 2001. Biological Oceanog- raphy: An Introduction. 2? Edition. Butterworth/Heine- mann, Great Britain. Lesser, M. P., F. H. Mar’tini, and J. B. Heiser. 1996. Ecolo- gy of the hagfish, Myxine glutinosa in the Gulf of Maine. I. Metabolic rates and energetics. Journal of Experimental Marine Biology and Ecology 37: 215-225. Nybakken, J. W. 2001. Marine biology: An ecological ap- proach. 5" edition. Benjamin Cummings, Montreal. Stockton, W. L., and T. E. DeLaca. 1982. Food falls in the deep sea: occurrence, quality and significance. Deep-Sea Research 29: 157-169. Strahan, R. 1963. The behaviour of Myxine and other Myxi- noids. Pages 22-33 in Biology of Myxine. Edited by A. Broda and R. Fange. Universitetsforlaget, Oslo, Norway. Tamburri, M. N., and J. P. Barry. 1999. Adaptations for scavenging by three diverse bathyal species, Eptatretus stoutii, Neptunea amianta and Orchomene obtusus. Deep Sea Research I 46: 2079-2093. Received 26 October 2005 Accepted 16 February 2007 Notes A New Record Size Wolf, Canis lupus, Group for Ontario Liv S. Vors! and Puitie L. WILSON? Trent University, Peterborough, Ontario Canada 'Current address: Department of Biological Sciences, University of Alberta, Edmonton, Alberta. T6G 2E9, Canada; e-mail: vors@ ualberta.ca Current address: c/o Sir Sanford Fleming College, Fish and Wildlife Department, Frost Campus, Albert Street South, P.O. Box 8000, Lindsay, Ontario Canada Vors, Liv S. and Philip L. Wilson. 2006. A New Record Size Wolf (Canis lupus) Group for Ontario. Canadian Field-Naturalist 120(3): 367-369. This report documents a group of 19 Wolves (Canis lupus) in northwestern Ontario. This is the largest group observed since record keeping in the Moose Aerial Inventory commenced in 1995. This large group may be a response to a high Moose (Alces alces) population in the Red Lake area. Key Words: Gray Wolf, Canis lupus, social organization, pack size, Moose, Alces alces, Ontario. The close association between Gray Wolves (Canis lupus) and their ungulate prey greatly influences the ecology of this cursorial predator, which is capable of killing prey much larger than itself (Kolenosky and Standfield 1975). Pack size is believed to be influ- enced by numerous factors. Pack sizes do not neces- sarily differ among Wolf populations whose major prey species are different. For example, average sizes of Wolf packs mainly depredating on Moose (Alces alces) are not necessarily larger than those feeding on White- tailed Deer (Odocoileus virginianus), but Wolf packs tend to be largest where Wolves prey on the largest ungulates (Fuller et al. 2003). This note documents one large group of Wolves observed in Ontario, Canada. Observation A group of 19 wolves was observed on 15 February 2005 between 1300 and 1400 hours on a frozen lake west of the settlement of Red Lake, Ontario, at approx- imately 51°OSN and 94°05W. Red Lake (51°04N, 93°47W) is a settlement located in the boreal forest of northwestern Ontario. This sighting took place during a transect flight to map areas of Woodland Caribou (Rangifer tarandus caribou) habitat activity. The air- craft, a De Havilland Turbo Beaver, was flying at a speed of 90 knots and an altitude of 400 feet when the group was observed. Ontario Ministry of Natural Resources (OMNR) pilot Carl Hansson, Trent Univer- sity graduate student Liv Vors, Sir Sanford Fleming College Fish and Wildlife Technologist graduate Philip Wilson, and Red Lake OMNR employee Claire Que- wezence observed and counted the Wolves. The aircraft made several circles over the animals, which were loosely congregated near the western shore of the lake. The Wolves ran around the ice, evidently disturbed by the aircraft, but made no attempts to flee into the neighbouring forest. No kill was visible. The animals were counted and confirmed by all observers. After about two minutes of observation, the aircraft resumed its transect flight. Discussion We believe this group to be the largest on record for Ontario, and possibly all of Canada east of the Rocky TABLE |. Records of maximum Canadian Wolf pack sizes west of the Rocky mountains, 1969-2005. Location _ Riding Mountain National Park, Manitoba Jasper National Park, Alberta Southeastern Quebec | Northwestern Alberta Algonquin Provincial Park, Ontario Southwestern Quebec Parry Sound Forest District, Ontario _ Pukaskwa National Park, Ontario Size Source 16 Carbyn (1980) 14 Carbyn (1974) 10 Potvin (1988) 10 Bjorge and Gunson (1985) 9 K. Mills, personal communication, 2005 8 Messier (1985) 8 Kolenosky (1972) 6 Forshner (2000) 367 368 Mountains. Wolf pack records are kept in the Moose Aerial Inventory Database 1995—2006*. Before this sighting the largest pack on record was 13, and was sighted in WMU (Wildlife Management Unit) 15A in 1998. Reported average pack sizes in Ontario range from 3.8 (Bergerud et al. 1983) to 5.9 (Pimlott et al. 1969) to 6.0 (Forbes and Theberge 1996), with maxi- mums of 7,9 and 13, respectively. Our observation of 19 wolves is significantly higher than most packs re- ported in Canada (Table 1). Gray Wolves found in the North American land- scape represented in their northern Ontario range pre- dominantly prey upon ungulates and Beaver (Castor canadensis) (Carbyn 1987; Gauthier and Theberge 1987). Large packs likely provide increased hunting efficiency when prey is scarce, or may alternatively be a response to abundant prey (Schmidt and Mech 1997). We suggest the large group we observed is a response to the latter. Since 1975, Moose populations in the Red Lake area have increased from a density of 0.059- 0.209 animals/km? to 0.359-0.509 animals/km? (Mc- Kenney et al. 1998). Current Moose densities in the Red Lake area are lower than McKenney’s estimates. Moose in the Red Lake WMU were estimated to be 0.265 animals/km? in 2004 (unpublished data, Ontario Ministry of Natural Resources, Bracebridge, Ontario). White-tailed Deer are also expanding their range north- ward, providing additional prey (Red Lake OMNR, personal communication). Wolf density is positively correlated with ungulate numbers (Fuller 1989; Mes- sier 1994). Although Moose density has declined since 1998, the ungulate biomass may still be high enough to support a large Wolf population. The relationship between pack size and prey size, however, is imperfectly understood. While food ac- quired per Wolf may be positively correlated with pack size (Nudds 1978), this relationship is not consistent. A review of prey use by Wolves in several geographic areas suggested a negative relationship between pack size and food acquisition (Schmidt and Mech 1997). In addition, prey size and pack size may not be closely linked. Mech (2000) observed a pack of 22-23 wolves in Minnesota, and this group preyed primarily on White-tailed Deer. Consequently, prey size and food acquisition are unlikely to be the only factors explain- ing the large group we observed. An additional caveat is the group we observed may have consisted of sev- eral packs that converged at the time of observation. An additional point of interest was the colour of the wolves. All were dark brown or black, with the excep- tion of one silvery-white individual. Most wolves in eastern Canada are gray, with black and white indi- viduals constituting approximately four and less than one per cent of the population, respectively (Gipson et al. 2002). THE CANADIAN FIELD-NATURALIST Vol. 120 Acknowledgments The aerial surveys were funded by the Ontario Min- istry of Natural Resources, The Wildlands League, The Wildlife Conservation Society, and The National Geo- graphic Society. Many thanks to Brent Patterson for his comments on the manuscript. Documents Cited (marked * in text) Moose Aerial Inventory Database. 1995-2006. Ontario Min- istry of Natural Resources, Bracebridge, Ontario. Literature Cited Bergerud, A. T., W. Wyett, and J. B. Snider. 1983. Role of wolf predation in limiting a moose population. Journal of Wildlife Management 47: 977-988. Bjorge, R. R., and J. R. Gunson. 1985. Evaluation of wolf control to reduce cattle predation in Alberta. Journal of Range Management 38: 483-487. Carbyn, L. N. 1974. Wolf predation and behavioural inter- actions with elk and other ungulates in an area of high prey density. Canadian Wildlife Service, Edmonton, Alberta. 233 pages. Carbyn, L. N. 1980. Ecology and management of wolves in Riding Mountain National Park, Manitoba. Canadian Wild- life Service, Edmonton, Alberta. 184 pages. Carbyn, L. N. 1987. Gray wolf and red wolf. Pages 358- 376 in Wild Furbearer Management and Conservation in North America. Ontario Ministry of Natural Resources. Published by The Ontario Trappers Association under the authority of the Licensing Agreement with the OMNR. Printed in Ontario, Canada. 1150 pages. Forbes, G. J., and J. B. Theberge. 1996. Response by wolves to prey variation in central Ontario. Canadian Journal of Zoology 74: 1511-1520. Forshner, A. 2000. Population dynamics and limitations of wolves (Canis lupus) in the greater Pukaskwa Ecosystem, Ontario. M.Sc. Thesis, University of Alberta, Edmonton. Fuller, T. K. 1989. Population dynamics of wolves in north- central Minnesota. Wildlife Monographs 110, The Wildlife Society, Bethesda, Maryland. 37 pages. Fuller, T. K., L. D. Mech, and J. F. Cochrane. 2003. Wolf population dynamics. Pages 161-191 in Wolves: Behavior, ecology and conservation. Edited by L. D. Mech and L. Boitani. University of Chicago Press, Chicago, Illinois. Gauthier, D. A., and J. B. Theberge. 1987. Wolf predation. Pages 119-127 in Wild Furbearer Management and Con- servation in North America. Published by The Ontario Trappers Association under the authority of the Licensing Agreement with the OMNR. Printed in Ontario, Canada. 1150 pages. Gipson, P.S., E. E. Bangs, T. N. Bailey, D. K. Boyd, H. D. Cluff, D. W. Smith, and M. D. Jiminez. 2002. Color pat- terns among wolves in western North America. Wildlife Society Bulletin 30: 821-830. Kolenosky, G. B. 1972. Wolf predation on wintering deer in east-central Ontario. Journal of Wildlife Management 36: 357-369. Kolenosky, G. B., and R. O. Standfield. 1975. Morpholog- ical and ecological variation among gray wolves (Canis lupus) of Ontario, Canada. Pages 62-72 in The wild canids: Their systematics, behavioral ecology, and evolution. Edited by M. W. Fox. Nostrand Reinhold, New York. 2006 McKenney, D. W., R. S. Rempel, L. A. Vernier, Y. Wang, and A. R. Bisset. 1998. Development and application of a spatially-explicit moose model, Canadian Journal of Zool- ogy 76: 1922-1931. Mech, L. D. 2000. A record large wolf, Canis lupus, pack in Minnesota. Canadian Field-Naturalist 114(3): 504-505. Messier, F. 1985. Social organization, spatial distribution and population density of wolves in relation to moose density. Canadian Journal of Zoology 63: 1068-1072. Messier, F. 1994. Ungulate population models with predation: a case study with the North American moose. Ecology 75: 478-488. Nudds, T. D. 1978. Convergence of group size strategies by mammalian social carnivores. American Naturalist 112: 957-960. NOTES 369 Pimlott, D. H., J. A. Shannon, and G. B. Kolenosky. 1969 The ecology of the timber wolf in Algonquin Provincial Park, Ontario. Research Report (Wildlife) 87. Ontario Department of Lands and Forests, Toronto. 92 pages. Potvin, F. 1988. Wolf movements and population dynamics in Papineau-Labelle reserve, Quebec. Canadian Journal of Zoology 66: 1266-1273. Schmidt, P. A., and L. D. Mech. 1997. Wolf pack size and food acquisition. American Naturalist 150: 513-517. Received 27 September 2005 Accepted 10 May 2007 A Tribute to Nicholas Stephen Novakowski 1925-2004 JOSEPH E. BRYANT 447 Thessaly Circle, Ottawa, Ontario K1H 5W7 Canada Bryant, Joseph E. 2006. A tribute to Nicholas Stephen Novakowski 1925-2004. Canadian Field-Naturalist: 120(3): 370-378. Nicholas Stephen Novakowski was an enthusiastic and very knowledgeable wildlife and fisheries scien- tist. Although his list of published works is modest, he was well known and respected across Canada and in several international fora. He has been credited as “the person who saved the Wood Bison as a discrete sub- species” (W. E. Stevens, personal communication) and was highly engaged in the early years of the sur- veys and recovery of the Whooping Crane (Novakows- ki, 1965b, 1966). Following those early days at Fort Smith, North West Territories, Nick was transferred first to the Edmonton office of the Canadian Wildlife Service and then to CWS headquarters in Ottawa where he was appointed Staff Specialist, Mammalogy, and took on national and international responsibilities. He retired in 1983 but continued his strong support for endangered wildlife and their habitats until his death on 3 July 2004 in Ottawa. He is survived by his wife, Amy, and two sons, Kent, a professor of hydrogeolo- gy and environmental engineering at Queen’s Univer- sity and Erin Nicholas, a professor of geography and environmental studies at Memorial University. Nick was born 28 November 1925 in Mundare, Al- berta (about 70 km east of Edmonton), the second child and eldest son in a family of 10 children. His father owned a Chrysler dealership and garage and from an early age Nick learned about all things mechanical — a fact that would stand him in good stead in later years as a field biologist. He also developed an abiding loy- alty to Chrysler products and, briefly, owned only one automobile that didn’t bear that company’s mark. Find- ing the alternate car highly unreliable, he soon returned to his original choice and did not waver again. In his younger years he was a superb athlete. He and his brothers Archie and Van were well known in central Alberta hockey circles as a high-scoring powerhouse. Nick, who was ambidextrous, could play either wing and was a sharp goal tender (Ken Novakowski, person- al communication). Also in those early years he became an ardent angler and as often as possible visited mem- bers of his extended family who lived in towns north of Edmonton. One of his favourite spots was Siebert Lake. Just a couple of weeks before his death he very happi- ly returned there with his sister, Sharon, and her hus- band to renew those earlier experiences — with consid- erable success. Time had not dulled his expertise. Nick entered the University of Alberta in 1947 and in the following two summers worked on forest fire suppression crews based in Fort Smith, North West Territories. There he became acquainted with Bill Ful- ler, the resident mammalogist for the Canadian Wildlife Service. He graduated with a B.Sc. in Chemistry in 1950 and that summer returned north but this time he was hired by Northern Transportation Company Limit- ed (NTCL) for summer work on the docks at the com- pany’s shipping base near Fort Smith. At that time NTCL gave hiring priority to young men who could enhance its baseball team and Nick “was a very good ball player” (W. A. Fuller, personal communication). Nick returned to the University of Alberta that autumn to begin a Master’s program and went back to Fort Smith in the spring of 1951. That autumn Dr. Fuller was assigned to conduct research of Wolves in Wood Buffalo National Park (WBNP) and adjacent areas of the Mackenzie District and employed Nick as an assis- tant (see Fuller and Novakowski 1955). Nick also helped Fuller with his studies of Beaver and with the “autopsies of lone, crippled, bison we encountered from time to time” (W. A. Fuller, personal communication). It was the start of Nick’s wildlife and fisheries career, and perhaps of equal importance, the start of the romance that led in June 1952 to his marriage to the love of his life, Amy Swartz, a Fort Smith school teacher from Saskatchewan. Also in 1952, with Fuller’s recommendation, Dr. D. S. Rawson accepted Nick for graduate studies in fisheries at the University of Saskatchewan. Over the next few years he worked very happily at his studies and as a fisheries biologist for the Government of Sas- katchewan. Although only one published paper appears to have resulted from that work (Atton and Novakows- ki 1956), he authored or co-authored many in-house reports for the Saskatchewan government dealing with fisheries research and resource management. Amy No- vakowski tells me that those were the happiest years of Nick’s life — helped, no doubt, by the fact that Amy was able to accompany him on many of his extended field studies in northern Saskatchewan. The photograph accompanying this article was taken at that time. After completing his M.Sc., (Novakowski 1955c*) Nick was hired by the Canadian Wildlife Service in 1956 and sent to Fort Smith. His mentor, Dr. Fuller, was transferred that year to Whitehorse, Yukon, and Nick dived immediately into Dr. Fuller’s continuing studies of Beaver populations in the southwestern Northwest Territories, Wolves in Keewatin District (Flook and | 370 2006 A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004 FIGURE 1. Nick Novakowski, fisheries researcher, Lac des Iles. Saskatchewan, 1955. 3/72 Novakowski 1956*; see also Novakowski 2003) and Bison and Whooping Cranes in Wood Buffalo Nation- al Park. In 1957, in the course of an aerial survey of Bison in WBNP, he discovered the presence of an isolated herd of Bison in the Nyarling River area in the northwest corner of the Park (Banfield and Novakowski 1960). Subsequent aerial and ground studies indicated that the herd might be Wood Bison genetically uncontaminated by the Prairie Bison that had been transferred to WBNP in the 1920s. In 1959, Nick was able to obtain speci- mens which were confirmed by the National Museum to be Wood Bison (Banfield and Novakowski 1960). In the following three years he arranged for over 70 live specimens to be trapped and tested for tuberculosis and brucellosis. Of those, 18 that were disease-free were moved in February 1963 to a large holding corral north- west of Fort Smith. Three calves arrived that spring and in early summer all of the animals were again test- ed for disease. All were clear. Nick had previously scouted out appropriate habitat north of Fort Provi- dence, North West Territories (Novakowski, 1959d*), and 18 disease-free Wood Bison, including the new calves, were transported there in August 1963. That area, now known as the Mackenzie Bison Sanctuary, proved to be an excellent site and the small nucleus soon swelled. Subsequently other specimens were taken from the Nyarling River site, transferred to Elk Island National Park, and after a rigorous clean-up protocol, produced another disease-free herd of Wood Bison (Gates et al. 2001). Some of Nick’s less well-known Bison research included “the first use of dental annuli to estimate the age of bison” (Christianson et al. 2005). The same authors also credit him with being the first to measure and to demonstrate the usefulness of other Bison dentition measurements in aging large samples. In 1958, Nick was also in charge of stocking Pine Lake and a few smaller lakes in WBNP with Rainbow Trout. He established and operated a small hatchery for that purpose, putting his Saskatchewan fisheries experi- ence to good purpose (Novakowski 1958c*, 1959c*). In 1962, when anthrax struck the Bison population just outside WBNP, it was the first recognized instance in Canada of anthrax in wildlife and all concerned were faced with a very steep learning curve. Nick worked closely with a team of scientists from the Canadian Wildlife Service, the Health of Animals Branch of the federal Department of Agriculture, and medical per- sonnel to bring it under control (Novakowski et al. 1963; Choquette et al. 1972). Dr. Fuller had engaged the interest of a long-time trapper in the southern portion of WBNP, Solomon Lacaille, to help with Muskrat studies (Fuller 1951). Nick followed through and set up a multi-year study on Mr. Lacaille’s trapline to study many aspects of beaver biology and ecology, leading to his Ph.D. from the Uni- versity of Saskatchewan in 1965 (Novakowski 1965a*, 1967a, 1969a). THE CANADIAN FIELD-NATURALIST Vol. 120 The nesting grounds of the Whooping Crane, long an international mystery, were discovered in 1954 in the northern portion of Wood Buffalo National Park. In the subsequent two years the site was monitored by Dr. W. A. Fuller. Nick took over those studies in 1956 and monitored the small breeding population for sever- al years (Novakowski 1966, 1967d). He was also res- ponsible for capturing an injured juvenile whooper in 1964 and for ensuring its safe conduct to the breeding facility of the United States Fish and Wildlife Service near Washington, D.C. (Novakowski 1965b, 1965e*). That bird, a male named “Canus” for its dual nationali- ty, sired “over 180” (E. Kuyt, personal communication) Whooping Cranes over its long career. It died in Jan- uary 2003. The mounted body was repatriated to Cana- da and is now installed at the Northern Life Museum in Fort Smith, N.W.T. (E. Kuyt, personal communication). Nick’s work on Wood Bison and Whooping Cranes introduced him to the burgeoning field of rare and endangered species. It was a field that would occupy much of his attention for the remainder of his life. The Canadian Wildlife Service moved Nick to a Staff role in Ottawa in 1966. J. Anthony Keith, who was Nick’s immediate superior in the Canadian Wildlife Service for much of his time in Ottawa, reported that “Nick Novakowski was the founding voice in CWS headquarters for wide-scale collaborative action on endangered species, beginning in 1967 when he was Staff Specialist, Mammalogy. While his own earlier work on Whooping Cranes became the trademark fed- eral endangered species project, there was a continuing debate amongst headquarter staff about how far CWS should get involved in endangered species work. Nick had to argue against those who saw it as a diversion of limited resources from urgent wildlife conservation issues. He worked with Ted Mosquin to produce the first set of scientific articles in Canada on endangered vertebrates (The Canadian Field-Naturalist 84(1) 1970), writing the article on mammals.” [See Nova- kowski 1970b]. In 1973, Nick represented the Canadian Wildlife Service at the international negotiations in Washing- ton of what became the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). While John Heppes joined CWS to estab- lish the Canadian management authority for CITES, Nick chaired the scientific authority until he retired in 1983 (Burnett 1999, page 162). Nick Novakowski and John Heppes worked well together. Their first task was to convince provincial gov- ernments that CITES would not interfere with provin- cial authority over wildlife resources. John dealt with the administrators and enforcement personnel while Nick met with the wildlife managers and scientists. “Nick showed an amazing understanding and ability in persuading all concerned to fully cooperate ...” (J. Heppes, personal communication). Canada ratified the Convention in early 1975. Every two years there was 2006 a “Conference of the Parties” at which member nations negotiated changes. “It was at these Conferences that Nick really proved his worth. In his role as Scientific Advisor he was always able to present the Canadian viewpoint in a clear and concise manner. While usually a calm and collected person, Nick could at times be very forceful in getting the Canadian opinion across ... he was respected by many international authorities who often sought his advice on biological status of en- dangered species and his assistance in presenting their positions.” (J. Heppes, personal communication). Eugene Lapointe, the first Secretary General of CITES, remembers Nick’s contributions for their sci- entific rigour and said, “My recollection of Nick is that he played a key role at the CITES meeting held in Gab- arone, Botswana in 1983. The international commu- nity had an opportunity to realize what real and cred- ible science was all about, and that thanks to Nick. I always perceived him as one of the most candid scien- tists. He was highly respected by his peers from the CITES community” (E. Lapointe, personal communi- cation). I recall one meeting in Ottawa where Canadian Non- Government Organizations (NGOs) were presenting their concerns about an upcoming Conference of the Parties. Two delegates were particularly vociferous in their assault on a proposed Canadian position. Nick as usual was thoroughly prepared and met both scientific and emotional arguments with calm, well-reasoned, non-confrontational responses which won the day. Nick was also front and centre in the discussions lead- ing up to the establishment of the Committee on the Status of Endangered Wildlife in Canada (COSEWIC). Ted Mosquin recalled that as he was preparing draft recommendations for an upcoming symposium on Canada’s Threatened Species and Habitats in 1975 Nick suggested that he look into the UK model that had “a unique committee that addressed the matter of the status of endangered species in the UK. [Nick] came over to the CNF office to fill me in on how the UK system worked. The result was a draft recommenda- tion ... which led to almost immediate establishment of COSEWIC by agreement of CWS and provincial agencies via the Federal Provincial Wildlife Confer- ence” (T. Mosquin, personal communication). “Throughout Nick’s work as a senior scientist in headquarters, he maintained a network of colleagues in provincial and territorial governments. Nick’s credibil- ity with these colleagues was vital in their governments’ agreement in 1977 to the initiative that became the Committee on the Status of Endangered Wildlife in Canada (COSEWIC), with Nick elected as its first per- manent secretary. During the first formative years of COSEWIC Nick did outstanding work helping to forge a new way of doing business, helping to bring federal, provincial, territorial and non-government agency peo- ple to agree on publicly-released status reports. Making public the facts about endangered species was what Nick had been calling for since 1967, believing that this A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004 373 would lead to remedial actions supported by the public COSEWIC continued to grow in the spirit of Nick's early aspirations, and recently was established as a for- mal entity under the Species at Risk Act.” (J. A. Keith, personal communication). In addition to Nick’s work as COSEWIC secretary, he maintained his avid interest in species biology. He wrote the original Wood Bison status report and con- tributed to others. He also chaired the Marine Mammal and Terrestrial Mammal subcommittee in 1980 and, when it split, he chaired the Terrestrial Mammal sub- committee from 1981 to 1984. Dr. Novakowski was involved in a much broader scope of activities than this review can encompass but three items may help to round out the story. For a num- ber of years in the 1960s and 1970s he was involved with the search for more humane traps for capturing furbearers. His field experience with trappers proved very useful. He thoroughly understood the trappers’ needs (Novakowski 1958d*) and also the political pres- sure being created to do away with the leg-hold trap. In 1967 he gave a brief but cogent review of the problem to the Federal-Provincial Wildlife Conference (Nova- kowski 1967e) and continued to be involved with the resulting program to develop humane traps until his retirement. Another example of his varied work in- volved blocking the introduction by an Ontario fur rancher of several dozen Raccoon Dogs (Nyctereutes procyanoides) from Finland. When the news arrived on Nick’s desk, he nearly blew a fuse! He was one of the very few people in Canada who had the knowledge to foresee a disastrous outcome if those animals were allowed to stay. He knew from the literature and from his many European colleagues that if the animals ever got loose the species could have devastating ecological effects. And he knew that no matter what precautions were taken, sooner or later some of them would get loose. He sought the assistance of Dr. Eric Broughton, a veterinarian with the CWS, to have Agriculture Cana- da deny the import permit. No dice — there was no basis for denial on environmental grounds. Nevertheless, based on knowledge of the ecology of the species, he convinced Canadian authorities that the animals posed an unacceptable threat to Canadian wildlife and ob- tained authorization to have all of the imports killed, the pelts processed by the Hudson’s Bay Company and compensation made to the owner. But he didn’t leave it there. Nick pursued the problem domestically and with the United States authorities resulting in a formal Canada—USA agreement to bar the future importation or Raccoon Dogs into either Canada or the USA (E. Broughton, personal communication). A third dimen- sion of Nick’s professional work involved teaching a variety of courses, including biology, terrestrial ecol- ogy, and wildlife management at University of Ottawa, the University of Guelph, and Sir Sandford Fleming College in Lindsay, Ontario. After retiring from the Canadian Wildlife Service in 1983, Dr. Novakowski became active as a scientific 374 consultant and engaged in a wide range of projects. In some cases he worked independently. In most he served as a member of a consulting team. In one of his early projects, he worked with Ted Mosquin to con- solidate and review many earlier reports on Caribou and fur-bearers in Pukaskwa National Park (Novakows- ki and Mosquin 1986a*, b*, c*, d*). Dr. Mosquin recently commented: “Nick provided the publications/ reports, guidelines, policies, appendices, survey data, illustrations, etc. as well as some text. I basically organ- ized and wrote the reports based on the material and dis- cussions. He was the expert. We worked well together and I learned a lot! ... We sometimes sat for more than a few hours trying to figure out what needed to be said and how to best say it.” Several of Dr. Novakowski’s post-retirement proj- ects dealt with major Environmental Impact Assess- ments of such projects as the Military Low-Level Fly- ing program in Labrador, uranium mining in northern Saskatchewan, the Rafferty-Alameda dams proposal in southern Saskatchewan, military base proposals in the arctic, and bison management in Wood Buffalo National Park. He became very knowledgeable about environmental impact assessments in Canada and con- tributed behind the scenes on a variety of policy issues. He also contributed to a review paper on the evolution of environmental impact assessments of watershed mod- ification projects (Dirschl et al. 1992). In 1982, a year before Nick retired from the Cana- dian Wildlife Service, Dr. Husain Sadar arrived at the Federal Environmental Assessment Review Office (FEARO). In a letter following Nick’s death, Dr. Sadar wrote that one of his “major responsibilities was to organize teams of highly recognized, accomplished and credible independent experts and specialists to assist and advise [the independent review panels] for analyz- ing and addressing often complex scientific, technical and socio-economic issues ... Nick served as an inde- pendent expert for several panels [dealing with] some of the most difficult, controversial and complex mega project proposals ... the panels benefited immensely from Nick’s unique experiences and expertise in wild- life management and habitat protection. Whether the issue was protection of caribou herds, moose, large car- nivores, furbearers, waterfowl, seabirds, birds of prey, marine birds or deer, Nick’s critical analysis, evalua- tion and resolution of all such issues was simply out- standing ... But the most important quality of Nick’s wonderful character was his ability to interact and work with other professionals in a highly profession- al, civilized and friendly fashion. He truly was a real gentleman.” Dr. Herman Dirschl, with whom Dr. Novakowski collaborated on many contracts, recalled their joint efforts on some of those major projects and comment- ed, inter alia that, “Nick could be very passionate in his views, particularly when political decisions seemed to often favour short term economic gains at the expense THE CANADIAN FIELD-NATURALIST Vol. 120 of the long term environmental health. However, he managed to always stay calm and collected in the face of such adversity. I admired him for this ability and will greatly miss him.” Dr. Dirschl also recalled their work “on an assessment of priority habitat sites throughout Canada to be considered for protection by the Canadi- an Wildlife Service in concordance with the Federal Government’s ‘Green Plan’ goals. Nick took on the res- ponsibility for reviewing the site records for a total of 262 sites in all provinces and territories; designing a classification system and defining latitude/longitude centroids for each site.” Even Canada Post sought out Dr. Novakowski’s ad- vice in planning for a commemorative postage stamp featuring Beaver — the 150" anniversary of the origi- nal 3¢ Beaver stamp (Novakowski 1997*). Nick the scientist was also Nick the practical envi- ronmentalist. Early in their stay in Ottawa he and his wife, Amy, bought 150 acres of forested land in near- by Lanark County. It was covered primarily with sec- ond growth hardwoods but also had a small plantation of Red Pine which Nick carefully nurtured. He became very interested in the “Model Forest” concept and was a very keen member of the “Eastern Ontario Certified Forest Owners.” I leave the closing words to his good friend, car pool partner and long-time professional colleague, Gerry Lee: “On a house visit one day, Nick invited me into his garage to see his latest pride and joy. There sat a beautiful cedar strip canoe, like a collector’s piece of rare furniture, hand built by canoe builder Nick for one of his offspring. Apart from the odd comment previous- ly about “building a boat someday, I had no inkling of his interest or talent in this area. So much for thinking you know someone! To sum up, Nick to me was the consummate good guy; respected professional, devot- ed family man, equally devoted environmentalist, polite, humble and honest. A good friend I'll never forget.” Acknowledgments Nick’s wife, Amy, was very supportive, provided copies of important papers, and authorized the use of the photograph used in this tribute. His brother, Ken, provided details of Nick’s early family life. Several librarians were especially helpful in assembling the lists of Dr. Novakowski’s publications and unpublished reports: Terri Fraser, Canadian Wildlife Service, Edmonton; Jean-Francois Bélanger, Environment Canada Library, Gatineau, Quebec; Carol Schurr, former librarian, and Rob Wallace, Fisheries Ecologist, Saskatchewan Re- source Stewardship Branch, Saskatoon; Staff, Library and Archives Canada, Ottawa, and Keith Van Cleave, Librarian, Northern Prairie Wildlife Research Centre, Jamestown, North Dakota. Many former colleagues of Dr. Novakowski provided assistance: I thank in par- ticular Bill Fuller, Tony Keith and Gerry Lee for their contributions and for their very helpful comments on an earlier draft of this paper. Additional thanks are due 2006 Eric Broughton, Herman Dirschl, Don Flook, Andy Hamilton, John Heppes, Ernie Kuyt, Eugene Lapointe, Alan Loughrey, Ted Mosquin, Doug Pollock, Dick Russell, Husain Sadar, Ward Stevens, John Tener and Don Thomas. I also thank my wife, Mary Bryant, for her constant support and assistance. Literature Cited (exclusive of items listed in Bibliography) Burnett, J. A. 1999. A passion for wildlife: A history of the Canadian Wildlife Service, 1947-1997. The Canadian Field-Naturalist 113(1) 1-214. Christianson, D. A., P. J. P. Gogan, K. M. Podruzny, and E. M. Olexa. 2005. Incisor wear and age in Yellowstone bison. Wildlife Society Bulletin 33(2): 669-676. Fuller, W. A. 1951. Natural history and economic importance of the muskrat in the Athabasca-Peace delta, Wood Buffalo Park. Canadian Wildlife Service, Wildlife Management Bul- letin Series 1, Number 2, 82 pages, charts, photographs. Gates, C. C., R. O. Stephenson, H. W. Reynolds, C. G. vanZyll de Jong, H. Schwantje, M. Hoefs, J. Nishi, N. Cool, J. Chisholm A. James, and B. Koonz. 2001. National Recovery Plan for the Wood Bison (Bison bison athabascae). National Recovery Plan Number 21. Recov- ery of Nationally Endangered Wildlife (RENEW). Ottawa, Ontario. 50 pages. Munro, D. A. 1973. Introductory remarks. Pages 28-29 in Convention on International Trade in Endangered Species of Wild Fauna and Flora. Transactions, 37" Federal-Provin- cial Wildlife Conference, Ottawa. Bibliography This bibliography is divided into published and un- published sections. While many of Nick’s reports were never published, their listing helps to demonstrate the breadth of his professional activities. Most reports pre- pared for the Federal Environmental Assessment and Review Office (FEARO) are not listed, in line with the terms of the contracts under which they were produced. Listings are chronological irrespective of whether Nick was the sole author, lead author or junior author. Publications Fuller, W. A., and N. S. Novakowski. 1955. Wolf control operations, Wood Buffalo National Park, 1951-52, Canadi- an Wildlife Service. Wildlife Management Bulletin, Series 1, Number 11. [2] + 20 pages, photographs. Novakowski, N.S. 1956. Additional records of bats in Sas- katchewan. Canadian Field-Naturalist 70: 142. Atton, F. M., and N. S. Novakowski. 1956. The value of freshwater fish as a mink food. Fur Trade Journal of Cana- da 33 (11): 12-13, 38, 40-42. Banfield, A. W. F. and N. S. Novakowski. 1960. The survival of wood bison (Bison bison athabascae Rhodes) in the Northwest Territories. Natural History Paper Number 8, National Museum of Canada. 6 pages, illustrations, map, tables. Novakowski, N. S., J. H. G. Cousineau, G. B. Kolenosky, G.S. Wilton, and L. P. E. Choquette. 1963. Parasites and diseases of bison in Canada. II. Anthrax epizootic in the Northwest Territories. Pages 233-239 in the Transactions of the 28" North American Wildlife and Natural Resources Conference. Wildlife Management Institute, Washington, D.C. [CWSC 1357]. A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004 375 Novakowski, N.S. 1965a. Cemental deposition as an age cn terion in bison, and the relationship of incisor wear, eye-lens weight, and dressed bison carcass weight to age. Canadian Journal of Zoology 43: 173-178. [CWSC 1426} Novakowski, N.S. 1965b. The day we rescued a whooping crane. Audubon Magazine 67: 230-233. Novakowski, N.S. 1966. Whooping crane population dynam- ics on the nesting grounds, Wood Buffalo National Park, Northwest Territories, Canada. Canadian Wildlife Service, Report Series Number 1. 19 pages, illustrations, tables, maps. Novakowski, N. S. 1967a. The winter bioenergetics of a beaver population in northern latitudes. Canadian Journal of Zoology 45 (6): 1107-1118. Novakowski, N.S. 1967b. Conservation of rare and endan- gered species of mammals in Canada. Pages 73-75 in the Transactions of the 31" Federal-Provincial Wildlife Con- ference, Ottawa. Novakowski, N.S. 1967c. Humane trapping. Transactions of the 31* Federal-Provincial Wildlife Conference, Ottawa. Pages 67-68. Novakowski, N.S. 1967d. Whooping crane. Modern Game Breeding 3 (6): 19-25, 35-41, 43. [A reprint of most of Novakowski 1966] Novakowski, N.S. 1967e. Beaver — Wood Buffalo National Park. Page 48 in Canadian Wildlife Service ‘66. Queen's Printer, Ottawa. Catalogue Number RR66-3766. Novakowski, N.S. 1967f. Bison — Wood Buffalo National Park. Page 48 in Canadian Wildlife Service ‘66. Queen’s Printer, Ottawa. Catalogue Number RR66-3766. Novakowski, N.S. 1968a. The National Committee on Wild- life Land Meeting, May 28-30, 1968. Pages 22-23 in the Transactions of the 32 Federal-Provincial Wildlife Con- ference, Ottawa. Novakowski, N. S. 1968b. The Export-Import Convention. Page 27 in the Transactions of the 32 Federal-Provincial Wildlife Conference, Ottawa. Novakowski, N. S. 1968c. Humane Trapping. Pages 27-29 in the Transactions of the 32 Federal-Provincial Wildlife Conference, Ottawa. Novakowski, N.S. 1969a. The influence of vocalization on the behavior of beaver, Castor canadensis Kuhl. American Midland Naturalist 81: 198-204. Novakowski, N.S. 1969b. Report on the Administrative Com- mittee for Polar Bear Research and Management in Canada. Page 22 in the Transactions of the 33 Federal-Provincial Wildlife Conference, Ottawa. Novakowski, N.S. 1970a Endangered wildlife —- mammals. Pages 6-7 in Endangered Wildlife in Canada. Canadian Wildlife Federation, Ottawa, Ontario. Novakowski, N. S. 1970b. Endangered Canadian mam- mals. Canadian Field-Naturalist 84(1): 17-23. Jeffrey, W. W., C.S. Brown, M. Jurdant, N.S. Novakow- ski, and R. H. Spilsbury. 1970. Foresters and the challenge of integrated resource management. The Forestry Chron- icle, June: 196-199. Choquette, L. P. E., E. Broughton, A. A. Currier, J. G. Cousineau, and N. S. Novakowski. 1972. Parasites and diseases of bison in Canada. II. Anthrax out-breaks in the last decade in northern Canada and control measures. Cana- dian Field-Naturalist 86: 127-132. Novakowski, N. S. 1973a. Natural resources in the Northwest Territories and the Yukon. Pages 91-97 in Natural Resource Development in Canada. Edited by Phillippe Crabbe and Irene Spry. University of Ottawa Press. 376 Novakowski, N.S. 1973b. Report of the Administrative Com- mittee for Caribou Preservation. Page 75 in the Transactions of the 37" Federal-Provincial Wildlife Conference, Ottawa. Novakowski, N.S., and V. E. F. Solman. 1975. Potential of wildlife as a protein source of food. Journal of Animal Science 40(5): 1016-1019. Novakowski, N.S. 1976. Endangered species. Pages 28-31 in the Transactions of the 39" Federal-Provincial Wildlife Conference, Ottawa. Novakowski, N.S. 1977. [an edited version of Dr. Novakow- ski’s statements as a witness on behalf of the Canadian Wildlife Federation before the National Energy Board hear- ings into a Mackenzie Valley Pipeline.] Pages 24A-24E in Wildlife Report. The Canadian Scene. May-June 1977. Canadian Wildlife Federation. Choquette L. P. E., E. Broughton, J. G. Cousineau, and N.S. Novakowski. 1978. Parasites and diseases of bison in Canada. IV. Serologic survey for brucellosis in bison in northern Canada. Journal of Wildlife Diseases 14(3): 329-332. Novakowski, N. S., and Geoff Robins. Editors. 1982. Pro- ceedings, Symposium on Natural Resources Use and Native Rights in Canada. Canadian Society of Environmental Biol- ogists, Ottawa. 105 pages. [EC-E] [EC-O] Novakowski, N. S. 1984. Control of mammals at airports. Pages 183-186 in Proceedings, Wildlife Hazards to Aircraft. Conference and Training Workshop, Edited by M. J. Har- rison, S. A. Gauthreaux and L. A. Abron-Robinson. U.S. Department of Transportation; Federal Aviation Admin- istration; Office of Airport Standards, Charleston, North Carolina. Hill, E. P., and N. S. Novakowski. 1984. Beaver manage- ment and economics in North America. Acta Zoologica Fennica 172: 259-262. Novakowski, N.S. 1985. Beaver management in Canada; a review. Proceedings of the 4" International Theriological Congress, Edmonton, Alberta. [Abstract] Session S08. Dirschl, H. J., N.S. Novakowski, and M. H. Sadar. 1993. Evolution of environmental impact assessment as applied to watershed modification projects in Canada. Environ- mental Management 17(4): 545-556. Novakowski, N.S. 2003. The Wilson River Escapade. Pages 8-10 in Then: Friends sharing memories. Edited and pub- lished by Mary and Joe Bryant, Ottawa. vii + 256 pages, illustrations. ISBN 0-9733372-0-6. Unpublished papers (marked * where cited) [EC-E] = Canadian Wildlife Service Library, Edmonton. [EC-O] = Environment Canada Library, Ottawa (Gatineau, Quebec). [ES-S] = Environment Saskatchewan, Saskatoon. Atton, F. M., and N. S. Novakowski. 1953a. Utilisation of fish on mink ranches in the Buffalo Narrows Region. Fish- eries Technical Report 53-1. 17 pages. [ES-S] Atton, F. M., and N. S. Novakowski. 1953b. The value of freshwater fish as mink food. Fisheries Technical Report 53-5. 10 pages. [ES-S] (See also published version, Atton and Novakowski, 1956.) Atton, F. M., and N.S. Novakowski. 1954a. Analysis of sam- pling the commercial fishery of Lake Athabasca, 1951-1953. Fisheries Technical Report 54-4. 11 pages. [ES-S] Atton, F. M., and N. S. Novakowski. 1954b. Analysis of sampling the commercial fishery of Lake Athabasca, 1954. Fisheries Technical Report 55-3. 7 pages. [ES-S] [EC-E] THE CANADIAN FIELD-NATURALIST Vol. 120 Novakowski, N. S. 1955a. Report on biological and fish- eries survey of Reindeer Lake, 1954. Fisheries Technical Report 55-1. 62 pages. [ES-S] Novakowski, N.S. 1955b. The ecology of Reindeer Lake with special reference to fish. Fisheries Manuscript Report M-4. 62 pages. [ES-S] Novakowski, N.S. 1955c. The ecology of Reindeer Lake with special reference to fish: a thesis submitted to the Faculty of Graduate Studies in partial fulfillment of the require- ments for the degree of Master of Science in the Depart- ment of Biology, University of Saskatchewan. 99 pages, 2 fold-out maps, figures. Atton, F. M., and N. S. Novakowski. 1956a. Biological sur- vey and fisheries management of Waterhen Lake and adjoin- ing lakes. Fisheries Technical Report 56-4. 50 pages. [ES-S] Atton, F. M., and N.S. Novakowski. 1956b. Biological sur- vey and fisheries management of Lac des Iles. Fisheries Technical Report 56-3. 34 pages. [ES-S] Atton, F. M., and N. S. Novakowski. 1956c. Biological survey and fisheries management of Jackfish and Murray Lakes. Fisheries Technical Report 56-2. 66 pages. [ES-S] Atton, F. M., and N. S. Novakowski. 1956d. The value of freshwater fish as mink food. Fisheries Technical Report. 2 pages. [ES-S] (See also published version, Atton and Novakowski 1956.) Novakowski, N. S. 1956a. Analysis of autopsy records of wolves taken in the Eskimo Point area 1956. Canadian Wildlife Service, CWSC 51: 12 pages, 2 tables. [EC-E] [EC-0] Novakowski, N.S. 1956b. Resurvey of beaver habitat in the Fort Simpson, Fort Liard and Wrigley areas, 1956. Cana- dian Wildlife Service, CWSC 753: 4 pages, maps, illustra- tions. [EC-E] [EC-0] Flook, D. R., and N. S. Novakowski. 1956. An experiment in poisoning wolves on the arctic tundra. Part A. CWSC 49: 9 pages, 2 maps. [EC-O] Novakowski, N.S. 1957a. Aerial resurvey of bison in Wood Buffalo National Park and surrounding areas, 1957. Cana- dian Wildlife Service, CWSC 216: 12 pages, maps. [EC-E] [EC-O] Novakowski, N. S. 1957b. Report on tagging, testing and slaughtering of bison in the Lake Claire area Wood Buf- falo National Park, October, November 1957. Canadian Wildlife Service, CWSC 728: 14 pages. [EC-E] [EC-O] Novakowski, N.S. 1958a. Report on the testing, tagging and slaughter of bison in Wood Buffalo National Park, October- November, 1958. Canadian Wildlife Service, CWSC 764: 18 pages, illustrations, charts. [EC-E] [EC-O] Novakowski, N.S. 1958b. Fur resources survey of Wood Buf- falo National Park. Canadian Wildlife Service, CWSC 749: 45 pages, tables. [EC-E] [EC-0] Novakowski, N.S. 1958c. Report on the hatching and plant- ing of rainbow trout Fort Smith, N.W.T., June 10-26, 1958. Canadian Wildlife Service, CWSC 1050: 5 pages. [EC-E] [EC-O] Novakowski, N.S. 1958d. Report on the testing of killer traps as a substitute for leghold traps. Canadian Wildlife Service. CWSC 772: 8 pages, photographs. Novakowski, N.S. 1959a. Aerial resurvey of the higher den- sity bison areas in Wood Buffalo Park and the N.W.T. Cana- dian Wildlife Service, CWSC 883: 5 pages, maps. [EC-E] [EC-O] 2006 Novakowski, N.S. 1959b. Analysis and appraisal of a three year aerial survey of beaver habitat in the southern and cen- tral areas of the Mackenzie District, N.W.T., 1956, 1957, 1958. Canadian Wildlife Service, CWSC 884: 17 pages, tables, maps. [EC-E] [EC-O] Novakowski, N.S. 1959c. Fish hatchery operations at Pine Lake — June 1959. Canadian Wildlife Service, CWSC 1051: 7 pages. [EC-E] [EC-O] Novakowski, N. S. 1959d. Report on the investigation of probable Wood Bison in the Nyarling River area and an investigation of historical Wood Bison range in the Fort Providence area. Canadian Wildlife Service, Fort Smith, CWSC 3493: 12 pages, tables, photographs. [EC-E] [EC-O}] Novakowski, N. S. 1961a. Cemental deposition as an age cri- terion in bison, and the relationship of incisor wear, eye- lens weight and dressed bison carcass weight to age. Cana- dian Wildlife Service, Fort Smith, N.W.T. CWSC 1426: 10 pages [EC-E] [EC-O] See also published version, Nova- kowski 1965a. Novakowski, N.S. 1961b. Estimates of the bison population in Wood Buffalo National Park and the Northwest Territo- ries based on transect and total counts. Canadian Wildlife Service, CWSC 882: 6 pages, maps. [EC-E] [EC-O] Novakowski, N.S. 1961c. Total counts of bison in the higher park density areas of Wood Buffalo Park. Canadian Wildlife Service CWSC 889: 11 pages, illustrations, map. [EC-E] [EC-O] Novakowski, N.S., and L. P. E. Choquette. 1961. Report on the testing and slaughter of bison at Sweetgrass, Wood Buf- falo National Park. Canadian Wildlife Service, CWSC 1102: 10 pages. [EC-E] [EC-O] Novakowski, N.S. 1962. Proposals for a bison management policy. Canadian Wildlife Service: 3 pages. [EC-E] Novakowski, N.S. 1963a. Report on anthrax in bison in the Hook Lake and Grand Detour areas, N.W.T., 1963. Cana- dian Wildlife Service: 4 pages. [EC-E] Novakowski, N. S. 1963b. Report on the transfer of Wood Bison, 1963. Canadian Wildlife Service: 5 pages, maps. [EC-E] Novakowski, N.S. 1963c. Wood bison transfer: completion report. Canadian Wildlife Service: 4 pages. [EC-E] Novakowski, N.S., and L. P. E. Choquette. 1963. Bison test- ing and slaughter at Hay Camp, Wood Buffalo National Park 1963. Canadian Wildlife Service, CWSC 1103: 9 pages. [EC-E] [EC-O] Novakowski, N.S. 1964a. The effect of disease syndromes on the ecology of the bison in Wood Buffalo National Park. Canadian Wildlife Service: 9 pages. [EC-E] Novakowski, N. S. 1964b. Report and recommendations con- cerning the occurrence and control of anthrax in bison in Wood Buffalo National Park and vicinity. Canadian Wild- life Service: 5 pages, maps. [EC-E] Novakowski, N.S. 1964c. Report on the assessment of pro- posed bison range in the Saskatchewan River delta. Cana- dian Wildlife Service: 11 pages, illustrations, table, maps. [EC-E] Novakowski, N. S. 1964d. Slaughter report - Hay Camp, November 1964. Canadian Wildlife Service, CWSC 1104: 9 pages. [EC-E] [EC-O] Novakowski, N.S. 1964e. The use of eye-lens weight, tooth replacement, cementum deposition and dressed weight as aging criteria in the bison. Canadian Wildlife Service: 11 pages, illustrations. [EC-E] Novakowski, N.S. 1965a. Population dynamics of a beaver population in northern latitudes. Ph.D. thesis, University of A TRIBUTE TO NICHOLAS STEPHEN NOVAKOWSKI 1925—2004 377 Saskatchewan, CWSC 1019: x + 154 pages, figures. [EC-E] [EC-0} Novakowski, N.S. 1965b. The effects of disease upon bison, Mackenzie District. Canadian Wildlife Service, CWSC 1696: 2 pages. [EC-E] [EC-O] Novakowski, N.S. 1965c. Possible bison range, Cumberland marshes. Canadian Wildlife Service: 2 pages, maps. [EC-E} Novakowski, N.S. 1965d. Slaughter report, Grand Detour, 1964-65. Canadian Wildlife Service, CWSC 1105: 2 pages [EC-E] [EC-O} Novakowski, N.S. 1965e. Wilderness rescue of an injured whooping crane: parents try and [sic] fend off an invad- ing helicopter in the wild muskegs of northern Canada. Canadian Wildlife Service: 5 pages. [EC-E} Novakowski, N. S., and W. E. Stevens. 1965. Survival of wood bison (Bison bison athabascae Rhodes) in Canada. Canadian Wildlife Service, Edmonton, 5 pages. [EC-E} [Paper delivered to the 45" annual meeting of the American Society of Mammalogists, Winnipeg, 20-26 June, 1965. An abstract is filed under CWSC 1020 in EC-O.] Novakowski, N.S. 1967a. Anticipated ecological effects of possible changes in the water levels of the Peace River- Athabasca River delta as a result of the damming of the Peace River. Canadian Wildlife Service, CWSC 1086: 17 pages, maps. [EC-E] [EC-O] Novakowski, N. S. 1967b. Bighorn sheep study meeting, Edmonton, February 28, 1967. Canadian Wildlife Service: 10 pages. [EC-E] Novakowski, N.S. 1967c. Concepts of multiple-use and inte- grated resource management. Canadian Wildlife Service: 10 pages. [EC-E] Novakowski, N.S. 1967d. Investigation of the Killdeer-Val Marie area in southern Saskatchewan as possible bison range. Canadian Wildlife Service, CWSC 1085: 9 pages, [+2] , maps. [EC-E] [EC-O] Novakowski, N. S. 1967e. Rare and endangered mammals in Canada. Canadian Wildlife Service, CWSC 2991: 141 pages. [EC-E] [EC-O] Novakowski, N. S. 1967f. Wood Buffalo National Park. Canadian Wildlife Service: 8 pages, map. [EC-E] Novakowski, N.S., and L. G. Blight. 1967. Investigation of Athabasca delta to assess feasibility, cost of constructing a rock plug and subsequent effects resulting from restricting the flow of water from the Lake Claire area. Canadian Wildlife Service. CWSC 1357: 5 pages, photographs, chart, map. [EC-E] [EC-O] Novakowski, N.S., and G. More. 1967. Mammalian pest and predator problems in Canada. Canadian Wildlife Service, Ottawa. CWSC 1703: 67 pages. [EC-E] [EC-O] Novakowski, N. S., and G. More. no date. Contributions toward a national pest control policy for Canada — mam- mals. Canadian Wildlife Service, CWSC 3444: 56 pages. [EC-E] [EC-O] Novakowski, N.S. 1970. Fire priority report, Wood Buffalo National Park. Canadian Wildlife Service, CWSC 1305: 6 pages, maps. [EC-E] [EC-O] Novakowski, N.S. 1971a. An overview study of the tourist potential of the Northwest Territories — wildlife. Canadian Wildlife Service, CWSC 1370: 8 pages. [EC-E] [EC-O] Novakowski, N. S. 1971b. Report to the National Energy Board on staffing requirements for an Environmental Divi- sion. CWSC 1369: 5 pages, 10 appendices, chart. [EC-O] Novakowski, N. S. 1975. Evidence concerning impacts of pipelines on rare and endangered species and aquatic and terrestrial animals. Paper presented to the Mackenzie Pipe- 378 line enquiry at the request of the Canadian Arctic Resources Committee. Canadian Wildlife Service, CWSC 2143: 33 pages, appendices. [EC-E] [EC-O] Novakowski, N. S., G. More, and P. Reilly. 1975. Exotic mammals and birds in Canada — a historical review. Cana- dian Wildlife Service, CWSC 1704: 47 pages. [EC-E] [EC-O] Novakowski, N. S., and T. Mosquin. 1986a. Caribou Man- agement Plan; Pukaskwa National Park, Volume I. 53 pages. Natural Resource Conservation, Ontario Region, Environment Canada, Ottawa. Consultant Report to Parks Canada by Mosquin Bio-Information Limited, Box 279 Lanark, Ontario. Novakowski, N. S., and T. Mosquin. 1986b. Caribou Man- agement Plan; Pukaskwa National Park, Volume II (Imple- mentation Strategy). 32 pages. Natural Resource Conser- vation, Ontario Region, Environment Canada, Ottawa. Consultant Report to Parks Canada by Mosquin Bio- Information Limited, Box 279 Lanark, Ontario. Novakowski, N. S., and T. Mosquin. 1986c. Furbearer Management Plan; Pukaskwa National Park, Volume I. 69 pages. Natural Resource Conservation, Ontario Region, Environment Canada, Ottawa. Consultant Report to Parks Canada by Mosquin Bio-Information Limited, Box 279 Lanark, Ontario. Novakowski, N. S., and T. Mosquin. 1986d. Furbearer Man- agement Plan; Pukaskwa National Park, Volume II (Imple- mentation Strategy). 69 pages. Natural Resource Conser- vation, Ontario Region, Environment Canada, Ottawa. THE CANADIAN FIELD-NATURALIST Vol. 120 Consultant Report to Parks Canada by Mosquin Bio- Information Limited, Box 279 Lanark, Ontario. Novakowski, N.S. 1989. Historical origin of National Park Plains Bison. Environment Canada, Canadian Park Service, Natural Resources Branch, Ottawa. 31 pages. Novakowski, N.S. 1990. Northern Diseased Bison Environ- mental Assessment Panel compendium of supplementary submissions, March 1990. Federal Environmental Assess- ment Review Office, Ottawa. 308 pages, maps. [EC-E] Novakowski, N.S., and H. J. Dirschl. 1992. Assessment of priority sites for the establishment of a Network of Con- servation Areas by the Canadian Wildlife Service. Environ- mental Social Advisory Services (ESAS) Inc., Ottawa, Ontario. Dirschl, H., N.S. Novakowski, and L. C. N. Burgess. 1992. An overview of the biophysical environmental impact of existing uranium mining operations in northern Saskat- chewan. vi + 90 pages, appendices. Report prepared for the Saskatchewan Uranium Mine Development Review Panel. Environmental Social Advisory Services (ESAS) Inc., Ottawa, Ontario. Novakowski, N.S. 1997. The beaver in Canada. A textual and visual exposition of the life history of the beaver, Castor canadensis. A report commissioned by the Stamp Products Division of the Canada Post Corporation. ii + 35 pages, illustrations. Received 13 January 2006 Accepted 3 May 2006 Book Reviews ZOOLOGY The Birder’s Companion By Stephen Moss. 2007. Firefly Books Ltd., 66 Leek Cres- cent, Richmond Hill, Ontario L4B 1H1 Canada. 208 pages. U.S. $16.95 This book is an entry in the “everything you ever wanted to know about birds” category. Each of its ten chapters deals with some aspect of birds — where do birds live? How do birds eat? Where do birds go? And, yes, each chapter does consist of a series of ques- tions and answers — over 450 in all — together with side-bars giving what can best be described as “gee- whiz” facts relevant to the chapter’s topic, all in rough- ly 20 pages or less each. Maybe a disclaimer is appropriate at this point — this kind of book turns me off. With that out of the way, Moss has done a creditable job in amassing a formi- dable series of facts and presenting them in a simple, colloquial style. In the introduction the author quotes a friend who suggests the book would appeal to “intel- ligent, inquiring eleven-year-old boys”, and while the author clearly hopes to appeal to a much wider audi- ence, he makes no apology for his approach. In spite of its disarming level of simplicity, the book is very well thought-out, up-to-date, and quite accurate. I imagine I could find an error or two in all this mass of data if I tried hard enough, but the fact remains that I didn’t. He even — glory be — gives the correct origin for the word “jizz” , with no inanities about “general impression of size and shape” usually parroted as the origin nowadays. My main quarrel with the text is the inevitably high level of simplification, which constant- ly left me feeling “yes, but...” Some of the author’s generalizations seemed very broad, and I wondered how he arrived at them; but again the ones I could check seemed within reasonable bounds. The author is British, and this can sometimes be a problem when writing for a North American audience. Birds of the Dominican Republic and Haiti By Steven Latta, Christopher Rimmer, Allan Keith, James Wiley, Herbert Raffaele, Kent McFarland, Eladio Fernan- dez. Princeton Field Guides, Princeton University Press. 2006. 258 pages. U.S. $35. Hispaniola, the second largest island in the Carib- bean, has a wealth of habitats and of birds, making it an emerging destination for birdwatchers. It also has immense development challenges, most acutely felt in Haiti, the poorest country in the western hemisphere. However, in this case the book is very carefully tailored to North America. One minor area where it could be misleading was in references to “buzzards”, the Euro- pean name for Buteo hawks, but also a term colloqui- ally used in parts of North America for vultures. Not only are we not told what buzzards are, but there are no buzzards in the index. One reference, to them being eaten by the European Eagle Owl [also not in the index]; was a particularly poor choice, given that Great Horned Owls will attack Red-tailed Hawks similarly. The weaknesses in the index are by no means con- fined to these two examples. It seems to place most of its emphasis on the material in the side-bars, and the text as a whole is far less well covered. This is particularly unfortunate because for me, at least, this is not a book to read from cover to cover, but is best dipped into from time to time, as some topic of interest arises. For such uses, a thorough index is very impor- tant. There is a one-page bibliography, although again some transatlantic emphasis creeps in, as many of the references are British — it’s hard to see the relevance of the UK breeding bird atlas — but more importantly they may be hard to find on this continent. The line illus- trations are not one of the strengths of the book. My carping aside, this book would appeal to novice birders or anyone with a mild interest in birds. Its appeal to more experienced birders is perhaps more limited, but it’s an easy read, best taken in small doses, and you'll probably find some things you didn’t know. And I seriously think it could be a very good gift for intelligent, inquiring eleven-year-olds. CLIVE E. GOODWIN 1 Queen Street, Suite 405, Cobourg, Ontario K9A 1M8 Canada These development challenges translate into conser- vation challenges, putting Hispaniola’s ecological rich- ness at peril. This book, the latest in several Caribbean guides issued by Princeton, is thus doubly important, both as an aid to identifying the island’s birds, but also as a valuable conservation tool which maps the cur- rent avian diversity of the island and communicates the value and beauty of the island’s birds to local popula- tions. 379 380 The book will be instantly familiar to anyone who has seen any of Princeton’s earlier Caribbean guides, such as A guide to the birds of the West Indies by Raf- faele et al. Indeed, some of the plates have been taken from that earlier guide. The book follows a standard formal, with front pieces speaking to the topography and habitats of the island (from the highest mountains in the Caribbean, at 3098 m, to the lowest lake, at 44 m below sea level), endemic species (31) and subspecies (50), avian conservation, and the island’s ornithological history. This is followed by detailed species accounts with information on similar species, vocalizations, hab- itat preferences, status, nesting, and global range, as well as a “comments” section containing interesting anecdotes such as the place of the bird in local culture. Each species account is accompanied by an up-to-date, detailed map of the bird’s distribution on the island. This information is complemented by 56 colour plates, most of which group up to 10 similar species however, pride of place is given to the island’s endemic species and one endemic family (Palmchat), which merit larg- er portraits. The book closes with descriptions of 14 birding sites and a checklist. The species accounts are accurate and provide suf- ficient information to identify most species. Particular attention is paid to the resident species most likely to be of interest to visitors, thus ensuring that one is indeed able to safely differentiate Long-billed and Narrow- billed Todys, the vireos, and other potentially confus- ing groups. Canadian readers will find the discussion of Bicknell’s Thrush of interest as Hispaniola is the chief wintering ground of this enigmatic breeding spe- cies. In general, the treatment of migrants is better than in many other regional guides; while you might want to lug around a good North American guide for some of the shorebirds and basic plumaged warblers, the level of detail in the plates is impressive and will usu- ally suffice (e.g, if you look closely you can see the differently marked tertials in the illustrations of Long- billed and Short-billed dowitchers). In general the plates have the rather washed out and sombre tones of the earlier Princeton guides, and in some of the por- traits the feathers have a rather messy look, not unlike old museum specimens, but this is a stylistic quibble; on the whole the plates are very good. While the deci- THE CANADIAN FIELD-NATURALIST Vol. 120 sion to illustrate all 306 species which have been re- corded on the island has considerable merit, having three illustrations of a bird which has only occurred once (Swainson’s Hawk) would seem an inappropriate use of always scarce space. The authors largely follow standard taxonomy, but have jumped the gun on American Ornithologists’ Union orthodoxy by accepting recently proposed revisions, such as treating the Hispaniolan Nightjar as distinct from the Cuban Nightjar, both traditionally combined as one species known as Greater Antillean Nightjar (confusingly, the latter name is used in the similar spe- cies account under Least Pauraque, one of relatively few editorial errors in the book). Similarly, the authors have abolished the names of two endemics, Green- tailed Ground-Warbler and White-winged Warbler which have now morphed into Green-tailed Ground- Tanager and Hispaniolan Highland-Tanager, respec- tively. Some other incipient splits and lumps are sug- gested in the text, reflecting the fact that yet more work needs to be done on the island. Likewise, the species accounts indicate that several species have been added to the island’s avifauna in the last decade, suggesting a recent increase in coverage and the possibility that the island’s bird list will grow: the local occurrence of several pelagic species in particular seems to be poor- ly known. Who should buy this book? Someone with a gen- eral interest in Caribbean birds may be better served by A guide to the Birds of the West Indies by Raffaele et al., which covers Hispaniola along with the rest of the islands. But anyone planning to travel to this beautiful island, or with a keen interest in Caribbean ornithology, would be well advised to acquire a copy; the range maps and site guides alone are worth the price if you are planning a self-guided visit (but bet- ter still, go with one of the organizations cited in the book). Finally, and most importantly, there should be a copy in every school and public library on the island: one hopes that Spanish and French versions are in the works. MARK GAWN Mission of Canada, Chemin de |’Ariana 5, Geneva 1202, Switzerland Atlas of Bird Migration — Tracing the Great Journeys of the World’s Birds Edited by J. Elphick. 2007. Firefly Books Ltd., 66 Leek Cres- cent, Richmond Hill, Ontario L4B 1H1 Canada. 176 pages. $35. Firefly Books has come out with another decent book. They’ve claimed the target audience to be, “bird enthusiasts, naturalists and the inquiring reader” and those who “wish to comprehend the perilous voyages that constitute one of the great wonders of the natural world.” The book is written to be consumable by peo- ple in grade school and perhaps almost fully under- stood by those as young as 12 or 14 years of age; it will be an enjoyable, and not too taxing to read for adults who have a passing interest in the topic. Serious birders will want to go elsewhere for more in-depth descrip- tions of the topics. The book starts out with 15 two-to-four page chap- ters on the biology and environmental conditions of migration, including timing, flight styles, genetics and weather. For the most part, these are well-illustrated 2006 synopses of the topics. The subsequent chapters are organized into sections covering major geographic areas (e.g, Eurasia, southern hemisphere, oceans), with two to eighteen chapters per section. Each chapter cov- ers an order or family of birds, using one to four species as examples. There is a map as well as a calendar show- ing migration and breeding times for most of the illus- trated birds. Although the book is worldwide in scope, it is dom- inated by a British point of view; after the typical migra- tion pattern of a species is noted, the accidental occur- rences in Britain (but nowhere else) are described (if such has occurred). The global perspective is worth- while, though only a small fraction of the planet’s migratory birds can be described. There are a few mistakes scattered throughout the book; these include classifying the Great Horned Owl Handbook of the Birds of the World Volume II By Josep del Hoyo, Andrew Elliott and David A. Christie (Editors). 2006. Lynx Edicions, Montseny, 8, 08193 Bel- laterra, Barcelona, Spain. 800 pages. 199 Euros Cloth. Handbook of the Birds of the World. Volume IT cov- ers eight families containing 733 species each with its own account and distribution map. The first of these is the Old World Flycatchers which, with the prejudices of youth, conjure up visions of Little Brown Jobs (LBJs). LBJs are the birders bane; small obscure birds that are difficult to see and identify. My first encounter with an Old World Flycatcher was the Spotted Fly- catcher. This is an LBJ where “spotted’ is an over- statement. They are only faint marks on the breast. This is a Sweeping and somewhat unfair assessment as more than 40 percent of Old World Flycatchers are colour- ful. Indeed, the 22 members of Cyorrus are as colour- ful as North American Bluebirds. In the next family, the Batises are small and largely black-and-white, whereas Wattle-eyes are somewhat more colourful with some orange and yellow. Many of the Monarch-flycatchers are a smart blue, but others are black and brown or black-and-white. However, the members of the small family of Paradise Flycatchers are among the most spectacular of the world’s birds. One of my life’s highlights was seeing the display of the black-and-white morph of the Madagascar Paradise Flycatcher. Similar and equally impressive species live in Africa and India. Kinglets and Firecrests is a family that is split between the old and new worlds. They are woodland sprites that can be surprisingly “cute” des- pite an initial dull impression. Gnatcatchers is a fami- ly of small, grey, long-tailed birds and is the only one in this volume that is confined to the new world. To my mind their counterparts are the Prinias of Africa and Asia. Cisticolas and allies are a bit of a nightmare. Most of them are true LBJs being streaky or dull brown. Knowing their songs and distribution helps. Even the BOOK REVIEWS 38] as an irruptive species, labelling Lake Winnipeg and its neighbours as the Great Lakes and describing the furcula as a pit (it is the wishbone). Many folks will gloss over these, but the more demanding reader will catch them; mistakes are never desirable, but most of these are relatively minor. In addition to covering distance migrants, the book pays some attention to altitudinal migrants. When ap- propriate, the authors have included conservation mes- sages on topics such as introduced species, habitat loss and the implications of commercial fisheries on birds. Overall, this wouldn’t be a book I'd give one of my serious birder friends as a gift, but it would do nicely for the beginner who is just migrating into the hobby. RANDY LAUFF St. Francis Xavier University, Antigonish, Nova Scotia — Old World Flycatchers to Old World Warblers easier to recognise birds like the Common Jery of Madagascar and the long-billed Tailor Bird of Asia can be a problem. They are so active that critical charac- teristics can be hard to see. Whenever you think you have seen all the relevant characteristics, after the bird has flown, you often discover there was something critical you missed! The 270 species of Old World Warblers in the Sylvi- idae (with 14 species confined to the antique island of Madagascar) are mostly LBJs. They are some of the most difficult birds to identify in the field. This night- mare is composed of several genera with hyphenated names like bush-warbler, reed-warbler, leaf-warbler and swamp-warbler. While there are discernable differences between, say, leaf- and swamp-warblers within each group there are many look-alike species which vary only by minuscule differences in eye-stripe or bill length. I spent every lunch hour for two weeks beside a Willow Warbler’s nest so that I could identify this species with confidence. This meant I could better see the subtle difference between Willow Warblers and Chif- fchafs [greener versus browner, etc.]. Unfortunately these birds are variable. The reader need only compare the photo of a pale, greyish Willow Warbler on page 551 with the brownish-yellow one on page 514 to under- stand this difficulty. It gets even worse with birds like the newly split Booted Warbler [Hippolais caligata] and Syke’s Warbler [H. rama] Syke’s Warbler winters all over India whereas H. caligata is found only on the eastern coast between the Ganges and Pondicherry. So the two species overlap in the east and are almost iden- tical. As they do not sing in winter it will take a DNA test to separate them! So how did Handbook of the Birds of the World. Volume II fare in dealing with birds that are not as mag- nificent as albatross or wildly coloured like parrots? It actually does remarkably well. Once again we have a well-organized and researched volume that adds to the growing status of this project. As each new volume is produced it adds to the benchmark status of the exist- ing volumes as a primary source of information on the world’s birds. The format has remained stable for the last several volumes, which makes it easy to use and compare. The photos are top rate despite the small and obscure nature of these species. I know how difficult Conservation of the Black-tailed Prairie Dog: Edited by John Hoogland. 2006. Island Press, Washington, DC. 350 pages paperback U.S. $35. It is amazing how the conservation of an animal as interesting and entertaining as the Black-tailed Prairie Dog can be so controversial. But controversial it has been in the last few decades as environmental inter- ests have woken up to the fact that the campaign of Prairie Dog poisoning and habitat loss has resulted in a range contraction to less than 2% of the historical geographic distribution. To this day many ranchers actively dislike Prairie Dogs for their competition with livestock for rangeland resources. In contrast, some conservation practitioners think that enough other spe- cies rely on the habitat created on Prairie Dog colonies that Prairie Dogs should be considered a keystone species. In my own practice as a biologist at Grass- lands National Park, Saskatchewan, I have frequently been taken aback by how polarized the opinions on Prairie Dogs can be between individuals and groups that often share many similar core values. The book, Conservation of the Black-tailed Prairie Dog: Saving North America’s Western Grasslands is the brain child of Dr. John Hoogland who has re- searched the behaviour of Prairie Dogs at Wind Cave National Park in South Dakota since the mid-1970s. Dr. Hoogland felt that there was a need to summarize the extensive scientific literature on the Black-tailed Prairie Dog in a non-technical format designed to high- light the information’s relevance to conservation. To accomplish this Dr. Hoogland engaged 30 other spe- cialists and challenged them to draw out the lessons for conservation from their areas of specialty. These lessons are explored through 18 chapters on topics such THE CANADIAN FIELD-NATURALIST Vol. 120 it is to get any photographs of such little and active sprites, let alone artistic ones of birds singing or dis- playing. Handbook of the Birds of the World has now reached biblical reference status. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Saving North America’s Western Grasslands as natural history, social behaviour, competition with livestock, keystone species, human attitudes, and how to establish new colonies of Prairie Dogs. The chapter authors are all respected specialists and do an excel- lent job of relating their topic area to the central theme of the book: the conservation of Black-tail Prairie Dogs. The only significant criticism I have is that Dr. Hoog- land repeatedly states in Chapter 2 that Black-tailed Prairie Dogs do not hibernate. This is odd because pub- lished studies have shown that Black-tailed Prairie Dogs use facultative torpor (Lehmer et al. 2001). In fact, in southwestern Saskatchewan, at the northern edge of their distribution, Black-tailed Prairie Dogs spend as much as 95 days a year in winter torpor bouts lasting 7.6 to 13.6 days with minimum core body temperatures ranging from 7.1 to 11.6°C (Gummer 2005). Clearly, this behaviour deserved to be recognized and discussed as to how it might relate to efforts to conserve this spe- cies. Despite this small failing, I fully recommend this book to anyone interested in the conservation of this fascinating social rodent or who wants to become better informed on conservation issues on the Great Plains. Literature Cited Gummer, D. L. 2005. Geographic variation in torpor patterns: The northernmost populations of prairie dogs and kangaroo rats. Ph.D. dissertation, University of Alberta. 210 pages. Lehmer, E. M., B. Van Horne, B. Kulbartz, and G. L. Florant. 2001. Facultative torpor in free-ranging black-tailed prairie dogs (Cynomys ludovicianus). Journal of Mammology, 82(2): 551-557. Pat FARGEY Grasslands National Park, PO Box 150, Val Marie, Saskat- chewan SON 2T0 Canada Migrating Raptors of the World: Their Ecology and Conservation By Keith L. Bildstein. Cornell University Press, Sage House, 512 East State Street, Ithaca, New York 14850 USA. 332 pages. U.S.$35.00 Cloth. Raptors and bird migration both hold great fascina- tion throughout the world, so the combination of the two topics makes for an irresistible book. Bildstein is a foremost world authority. Based at Hawk Mountain, the world’s first site dedicated to the conservation of hawks and the watching of their migration, he is the Sarkis Acopian Director of Conservation Science. Bildstein tells us that “a bad day at Hawk Mountain Sanctuary is better than a good day anywhere else.” His wide knowledge derives in part from his travels to each of the world’s main hawk-watching sites. Fig- ure | displays the five major hawk pathways through- out the world and Figure 7 shows the 12 locations where 100 000 or more raptors pass by. In addition to stud- ies at Hawk Mountain, banding of thousands of raptors has occurred at Cedar Grove, Wisconsin; Cape May, New Jersey; Hawk Ridge on Lake Superior, Minneso- ta; the Goshute Mountains, Nevada; the Manzano Mountains, New Mexico, and Golden Gate Observa- tory, California. Notable Old World banding sites are Chokpak Pass, Kazakhistan, and Elat, Israel. Switching 2006 to birds counted overhead, 851 600 Honey Buzzards flew over Elat in the spring of 1985; there the spring flights are much larger than in fall. Elsewhere large numbers are counted in the fall: an average of 730 O00 at Corpus Christi, Texas; 5 200 000 near Cardel in the state of Veracruz, Mexico; | 950 000 at Kekéldi Indi- genous Reserve, Costa Rica, and 380 000 over the Strait of Gibraltar at the western end of the Mediter- ranean. In Veracruz, sometimes dubbed the “river of raptors,’ 2 677 355 Turkey Vultures were counted in 2003, 2 389 323 Broad-winged Hawks in 2002 and 1 197 850 Swainson’s Hawks in 2003, including 782 653 in one day, 17 October that year. Recent advances in technology have provided a big boost to raptor migration studies. Application of satel- lite radios to 117 Ospreys and 51 Golden Eagles added immeasurably to our understanding of the speed and variability of migration. Nine satellite-tracked Honey Buzzards averaged 270 km/day across the Sahara Desert, with brief stopovers suggesting that most fast- ed during at least this leg of their trip south. Since rap- tors often reduce energy requirements by soaring in thermals, Bildstein has included a map and three dia- grams of the major deflection/updraft corridors. Rap- tors are able to orient and navigate simultaneously (set- ting a direction and then achieving it), using internal magnetic cues, visible landmarks, the sun by day, the stars for the few species that migrate by night. Despite a great deal of research, their navigational prowess still exceeds human comprehension. In at least nine species, including harriers, accipiters and two falcons, the juve- niles precede the adults in southward migration, yet come spring the adults often head north first. In the Osprey and a few other species, juveniles spend an extra year in the tropical wintering habitat before returning north at two years of age. A positive feature is that a number of hawk watch- ing stations, including Hawk Mountain, were found- Land Snails of British Columbia By Robert G. Forsyth. 2004. Royal British Columbia Museum, Victoria, British Columbia, Canada. 188 pages. $35. Paper. The study of snails has certain advantages for a nat- uralist. For one thing, unlike birds or butterflies, they move slowly enough to be identified! However, anyone who has ever tried to identify snails will know that there is a dearth of readily accessible information out there to help with the task. This is a lack that Robert Forsyth has set out to remedy with his guidebook, pro- viding help with the identification of 92 species of land snails and slugs found in British Columbia. This book is similar in layout and format to other recent guidebooks from the Royal British Columbia Museum, a handy 5.5” x 8.5” soft cover book, easy to slip into a backpack or daysack. Each taxon is provided with a “species account” that consists of a description of the animal, its distribution, Book REVIEWS 383 ed as conservation measures. Incensed by the car- nage along the Kittatinny Ridge at Hawk Mountain, Rosalie Edge purchased the property and hired Mau- rice Broun to patrol it and count the raptors. Rachel Carson, in Silent Spring in 1962, used the 25-year Hawk Mountain counts, especially of declining Bald Eagles, to support her arguments concerning the im- pact of organochlorine pesticides, especially DDT, on the reproductive success of birds of prey. Bildstein’s final chapter is perhaps the strongest. It explains the widespread effects of bounties, not com- pletely eliminated in Pennsylvania until 1969. From 1917 to 1952, Alaska territory paid bounties on 128 000 Bald Eagles. Today, the main threats to raptors are habi- tat loss and environmental contaminants, an example being a painkiller, diclofenac, fed to cattle in India, that has led to population declines of over 90 per cent for three species of vulture that eat dead cows. My only criticisms are small ones. Because owls are also raptors, the absence of any mention of their no- madic movements, cyclical irruptions and migrations is somewhat contrary to the title of this book that is restricted to diurnal raptors. Readers of the main text might believe that DDT has long been banned every- where, but in fact it is still being used in tropical coun- tries to combat malaria and yellow fever. Only in the Glossary does Bildstein inform his readers that DDT is “still used by public health organizations to control malarial mosquitoes.” The writing is clear. Each chapter (I have only touched on a few highlights) ends with a synthesis and summary of the main points. This book is a treasure. It belongs in every high school and college library and in the library of every raptor enthusiast. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Canada and its natural history (basically its habitat preferences). Each account also includes information on the etymol- ogy of the animal’s name, some remarks, usually deal- ing with taxonomic issues or species with which it may be confused, and references. The species are arranged in taxonomic order. These accounts form the bulk of the book (128 pages). Most taxa are illustrated by line drawings or black-and-white photographs. For the pla- nispiral snails, the images generally include the upper (apical) surface, lower (umbilical) surface, and apertur- al view. For the conispiral snails, the images generally consist of an apertural view and a distal view. There are also 33 colour images, of which 23 are of slugs. These colour images are generally much crisper and more useful than the black and white photos. Forsyth includes some brief discussion of eleven other taxa that he considers of doubtful occurrence in British Colum- 384 bia or that he judges to have equivocal taxonomic sta- tus. These are often species that were perhaps only recorded from one locality or for which the record was made many decades ago and has not been confirmed or supported by subsequent observations. These species accounts highlight some interesting points about the land molluscs of British Columbia, as they are known at present. Of the species discussed, about one third (24) are slugs. Of these, more than half of them (13) are introduced. Many of these slug species are described as being common in gardens, which per- haps suggests that eagle-eyed gardeners may be respon- sible for drawing attention to them. Forsyth notes that many of these exotic species are significant agricul- tural pests, in contrast to members of the native fauna, which are not. He points out (page 19) that most may have been accidental introductions, brought in on plants or associated soil. Most of the snail species, in contrast, are native (53) with less than a quarter (15) identified as introduced. This striking pattern raises the question as to whether it is a consequence of the method of introduction (that is, on horticultural or agricultural plants) or simply a lower level of investi- gation of the mollusc fauna outside urban and inten- sively farmed areas. Despite its focus on British Columbia, many native species described by Forsyth have wider distributions. Some are found in other areas in the Pacific Northwest, from Oregon to Alaska. Oth- ers are more widespread and some occur east of the continental divide in other areas of western North America. Despite its focus on British Columbia, there- fore, this book has wider applicability in western Can- ada. The remainder of the book offers useful supplemen- tal information. An extensive introduction (19 pages) describes the biology of the animals, including, for the snails, some discussion of shell characteristics, which are important for species identification. There is also a checklist of the species described in the volume, and two dichotomous keys, one for snails and one for slugs, to help narrow down the selection to genus. Ten other keys are interspersed among the species accounts and focus on genera, such as Vallonia, Vertigo, and Arion, in which there are more than a couple of species. All the snails described are dextral coiling species, as are most land snails. Following the species accounts, there is a lengthy (14 pages) reference list, a glossary of terms, and a species index. I obtained this book primarily to help with the iden- tification of mollusc shell remains recovered from fine- fraction analysis of sediment samples from archaeolog- ical sites and other postglacial depositional contexts. THE CANADIAN FIELD-NATURALIST Vol. 120 In these situations, the soft parts of the animals are not preserved and only the shells remain. Hence, identifi- cations rely primarily on the morphology and struc- ture of the shells. So I was particularly interested in the shell descriptions. In this regard, the book has been very helpful although I have noticed some limitations. My main complaint is that the line drawings are not repro- duced well. This is a significant problem. On many drawings, the lines are faint and details of shell sur- face features and ornamentation are extremely diffi- cult to discern. Moreover, Forsyth only provides one shell measure for each taxon. For the planispiral mol- luscs, he provides the width of shells, but not the spire height. For the conispiral molluscs, he provides the height of the shells, but not the width. The complemen- tary measures can be approximated from the drawings, but it would have been useful to have them included in the text. The width/spire height ratio is an easily acquired descriptive character that helps discriminate different taxa. Two additional sets of information would have made this book even more helpful. First, I thought each species could have been illustrated by a “dot map” showing the localities in British Columbia where it has been collected or observed. I imagine that the dots would have been very sparse for many species! Nev- ertheless, such maps could highlight areas that may well repay survey and collecting effort. Second, it would have been useful to have a list of the accession numbers for the specimens that were studied to com- pile this volume. This is important so that in future scholars could re-examine the specimens to confirm and refine the taxonomy or acquire additional meas- urements and images. Because the Royal British Col- umbia Museum is the publisher, I assume that it may be the institution housing these collections. However, this information (that is, the institutional repository and accession numbers) could have easily been in- cluded, for example, in the species checklist, making such detective work unnecessary. Compiling this volume obviously involved a formi- dable amount of painstaking work and study. I heartily commend Forsyth’s commitment to a neglected but rewarding faunal group. This book is a fine introduc- tion to these animals and deserves to be on all west- ern Canadian naturalists’ bookshelves. Anyone who pays attention to the information in this volume will never look at a snail on a rockery or a slug on a cab- bage leaf in quite the same way again! ALWYNNE B. BEAUDOIN Royal Alberta Museum, Edmonton, Alberta TSN 0M6 Canada 2006 Book REVIEWS 385 Songbird Journeys: Four Seasons in the Lives of Migratory Birds By Miyoko Chu. 2006, Walker & Company, 104 Fifth Avenue, New York, New York 10011 USA. 312 pages. U.S. $23. Cloth, This is a welcome, attractive, and highly recom- mended book. Each chapter is preceded by a well- chosen quotation, often an appropriate full-page poem. Miyoko Chu has combined the best of investigative journalism with a passionate account of bird migra- tion that holds the reader’s attention. Four chapters deal with spring migration and two with fall migration. She also must deal with where these migrating birds summer and winter, so five chapters deal with these intervening seasons. Not only does she tell of the lat- est research into bird migration, but she interviews at length some of the leading authorities. Thus, we read fascinating stories about exceptional people who are studying interesting birds. I will mention a few high- lights. Her opening chapter deals with a long-unsolved question: do large numbers of birds make the seem- ingly impossible long-distance flight over the Gulf of Mexico, as George Lowery fervently believed, or was this a preposterous fiction of people’s imaginations, as George Williams posited? Lowery began by stud- ies through a telescope, and admitted that trans-Gulf migration was “fraught with enigmas and strained one’s credulity.” Williams accused Lowery of jumping from minute observations to sweeping conclusions, of extrap- olating from the 12 birds he had seen through the tel- escope to claim a migration of 21 million birds. Sid Gauthreaux, when 17 years old in Louisiana, inveigled his way into one of the first weather radar stations and persuaded the meteorologists to let him look at the fuzzy dots that he thought must be movements of birds. He then persuaded Lowery to accept him as a graduate student, embarking on a master’s thesis and then a doctoral dissertation that proved to doubtful ornithol- ogists everywhere that radar was a superb modality for studying many aspects of migration. In so doing, he proved beyond doubt that Lowery was correct. Frank Moore’s group at Johnsons Bayou, Louisiana, studied weights and condition of migrants as they first reached landfall, after crossing the Gulf. Almost half the birds arrived with their fat reserves totally depleted. Richard Graber, at Urbana, Illinois, was an incred- ibly early pioneer, four full decades ahead of the pack, in placing tiny radio transmitters on small birds. His friend, Bill Cochran, made him a number of tiny, 3 g transmitters, which he glued to the backs of thrushes that weighed about 30 g each. He tuned his receiver to the appropriate frequency and then followed the signals by truck, and once by aeroplane for 400 miles in eight hours. Cochran then took over the study and with special equipment he logged 150 000 miles by truck to follow individual birds for a total of 30 000 miles. He followed one Peregrine Falcon from Wiscon- sin to Tampico, Mexico, a Sharp-shinned Hawk from Cedar Grove, Wisconsin, to Huntsville, Alabama, and a Common Nighthawk from Illinois to South Carolina. Another hero 1s a bird, a Yellow Warbler, nicknamed “Wally,” that carried band 1750-17109, and returned for nine years in a row, to sing near the Long Point Bird Observatory in Ontario. At Hubbard Brook in New Hampshire, where Richard Holmes and his col- leagues have studied birds since 1969, bird numbers have declined by about sixty percent. There, Black- throated Blue Warblers produce more young in years when caterpillars are common and in La Nifia years. When Olin Sewall Pettingill was a boy in Maine, his mother showed him a Bobolink nest. But they faced hazards. About 720 000 Bobolinks were shipped as game from one place in South Carolina in 1912 alone. In Argentina, Pettingill learned in 1978, Bobolinks were now being shot as pests for eating rice crops. It is not surprising that the number of breeding bobolinks has dropped by 75 percent in 25 years. A new branch of science, studying the calls of noc- turnal migrants to recognize individual species, was pioneered particularly by Bill Evans in 1985, who pro- duced a cassette tape of the night calls of the thrush- es. His compilations were joined by those of Michael O’Brien in 1991. Four years later they had “nailed down” the final two species that had eluded them, the night calls of Canada and Wilson’s Warblers. By 2001, they produced the landmark guide that identified the flight calls of 211 species of migrating landbirds in eastern North America. Collaboration with researchers and software programmers at the Cornell Laboratory of Ornithology allowed analysis of different bird species from their individual characteristics. There are now thirty rooftop stations monitoring nocturnal migra- tion with these new methods. Although Chu fails to list the Evans/O’Brien CD in her references, it became available in 2002 and can be ordered through http: oldbird.org. Only in 1995 was Bicknell’s Thrush recognized by the American Ornithologists’ Union as a separate species from the similar Gray-cheeked Thrush. Chris Rimmer studied this thrush, one of the rarest migratory birds in North America, in its breeding habitat among the stunted firs near the top of Mount Mansfield in Vermont. But where did they winter? Rimmer’s crew found them in the Sierra de Bahoruco in the Dominican Republic and, miracle of miracles, in 1995 captured in a mistnet one of the banded birds they had tagged in Vermont the previous summer — and then recaptured the same bird again in Vermont during the next two summers! The reader is also given detailed directions to North American birding migration hotspots, in both spring and fall, and mid-winter hotspots in Yucatan, Belize. Costa Rica, and Panama. She provides instructions for amateurs to help contribute to knowledge, by partici- 386 pating in Nest Record Card programs and Project Feed- erWatch. Chu concludes by calling attention to the many haz- ards facing bird populations, such as loss of habitat, in- creasing parasitism by cowbirds, deaths from striking windows, communication towers, pesticides, carbon dioxide emissions, and acid rain. Twenty-nine species BOTANY Wildflowers of the Rocky Mountains By G. Scotter and H. Flygare. Whitecap Books, 351 Lynn Avenue, North Vancouver, British Columbia V7J 2C4 Canada. 255 pages. $ 29.95 Paper. Plants of Alberta By F. Rover and R. Dickinson. 2007. Lone Pine Publishing. 10145 — 81 Avenue, Edmonton, Alberta T6E 1W9 Cana- da. 527 pages. $ 29.95 Paper. It is most interesting that these two books have been published within such a short space of time. They were of immediate interest to me as I visit family in Alberta and usually manage a trip or two to the Rockies or the prairies. While they cover very different areas they include many of the same species. Furthermore, the authors have taken significantly different approaches. Wildflowers of the Rocky Mountains [Wildflowers] contains 350 species of selected plants in six sections by flower colour [white, yellow and cream, green, pink, red, orange, and brown, purple and blue.] Each page depicts typically two species, all with a description, a photograph and a range map. The authors say they have chosen the most common species (typically the pretty species) likely to be encountered during visits to the Rockies, plus a few rarities. The area covered is the Rocky Mountain zone from northern British Columbia to New Mexico. This is a complete revision and expan- sion of a 1986 guide, that covered 228 species. The authors state that over the years they have learned to use non- technical language. They have been successful in apply- ing this talent and has produced a very readable book. I have a few problems with Wildflowers. I have a dif- ferent view on the choice of colours. For example, I consider Mountain Sorrel to be reddish-brown not pink as the authors contend. Similarly, I think of Flame- coloured Lousewort (Pedicularis flammea) as red, but the authors have placed this plant in the cream section (because the flowers have a cream base). To further add to my confusion this plant is called Oeder’s Lousewort (P. oederi) but P. flammea is given as the synonym. (However a web search of P. oederi showed images of cream-only flowers, while an image search for P. flammae showed cream-based flowers with significant amounts of red on the tips. A search for synonyms of P. oederi did not include P. flammae.) The alternative names are not included in the index. Many of the Eng- lish, and some of the scientific names are not the ones I am used to. As an example, this book’s Four-part THE CANADIAN FIELD-NATURALIST Vol. 120 have declined by more than fifty percent in the last 40 years. Despite these losses, the arrival of songbirds every spring remains a cause for celebration. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N O0J8 Canada Dwarf Gentian or Felwort is called Alpine Gentian in other texts. The range maps are coloured in for the entire state or province regardless of how widespread the plant is. So Bear Grass is shown as occurring in Alberta, where- as it can only be found in the vicinity of Waterton Lakes National Park. This greatly reduces the usefulness of these maps. I thought the approach used by Wildflowers — arrang- ing the species by flower colour — was useful only for beginners. After a couple of years most people know the major families, like vetch, violet, saxifrage and so forth and I thought this is the way people progressed. Recently I was told that women prefer books organ- ized by flower colour and I found my wife agreed. The photographs are very good and will be useful in identifying blooms in the field. The descriptions in Wildflowers are written in prose, often in a poetic style that makes for pleasant reading. There is an excellent introduction on the eco-zones and a wonderfully clear glossary. The second book is Plants of Alberta (Plants) which covers over 1500 native plants that can be found in Alberta. It is arranged by family and includes aquatic plants, grasses, ferns and trees (so not just the pretty flowers). Certainly I prefer the Plants method of arrang- ing by families so I can reference all the orchids, vio- lets, louseworts, etc. together. It too has a description, a photograph and a range map also shown two to a page. There is a short introduction and a decent glossary. Plants’ descriptions are cryptic and scientific. Com- pare “While other saxifrages may dress in flowers of white or yellow, Purple Mountain Saxifrage is garbed with flowers of the richest rose-purple to royal purple” (in Wildflowers) with the terse “Flowers: Purple (occa- sionally pinkish or white.)” (in Plants) The photos in Plants average about twice the size of those in Wildflowers — a considerable benefit. This is achieved by having no margins, giving 20% extra space. Also the headers giving the plant names and family are 60% smaller [but less artistic!]. As the text is shorter the text size is bigger — great advantage for older eyes. The larger script comes at the loss of poetic text to terse notes. So despite the book’s slightly smaller pager size (10%), by the creative use of space the publishers have created a more appealing looking book. 2006 The range maps cover Alberta and show the area within the province where you can see the plant. This is useful because it shows over 35% of Alberta’s species are confined to the southern border with British Colum- bia and the United States. As the maps are small (1 x 2 cm) they require some interpretation. | photographed a Striped Coralroot in Bow Valley Provincial Park, so I verified that the range map showed this plant at this location. It does, but it required careful examination and analysis due to the size. The other distributions are equally valid. There is also a key arranged by leaf structure and flower colour. This uses thumbnail (4") photos of the MISCELLANEAOUS BooK REVIEWS 387 families. So Violas appear in the white, yellow and pur- ple sections. Is this enough to satisfy the ladies? Despite my clear preference for Plants as a book, | would say that both of these books are good guides. The photographs are excellent and the text, while very different in style, gives solid information. Your choice should be dictated by your intentions or location. Al- ternatively, and this I suggest is the best idea, you could buy both as both, are good value for money. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Pilgrimage to Vallombrosa: From Vermont to Italy in the Footsteps of George Perkins Marsh By John Elder. 2006. University of Virginia Press, Charlot- tesville, Virginia, USA. xv + 282 pages. U.S. $29.95 Cloth. This book is a story of pilgrimage, of memory, of hope. It’s a beautiful book, a book of life, of family, of nature and culture. John Elder, a professor of English at Vermont’s Middlebury College, invites the reader on a pilgrimage. It is a story of connections; from the sec- ond and third growth sugar bush hills of Vermont to the centuries-old, culturally prestigious forests of Vallom- brosa in the Tuscany hills; from the sugar bush to the olive grove; from Robert Frost to Dante. Vermont-born George Perkins Marsh was Abra- ham Lincoln’s diplomat to the new Kingdom of Italy from 1861 until his death in 1882. Considered to be America’s first environmentalist, Marsh was the author of the celebrated Man and Nature (1864). Inspired by his study of the rise and fall of Mediterranean cultures and his extensive experience of the region, Marsh’s manifesto focused on environmental decline as central to the collapse of these once powerful cultures. Marsh was the first to describe and document the drastic, long-term impact of deforestation on civilization. His Man and Nature stirred the American imagination as America itself experienced the wave of deforestation flowing from east to west across the American land- scape. A sabbatical year permitted Elder and his wife to set out on a pilgrimage across France and Italy. Elder’s pur- pose was to pursue the footsteps of Marsh who died in Vallombrosa in 1882. A pilgrimage usually has a set route, but novelty and surprise are to be expected. A pilgrimage on foot always becomes a pilgrimage of the heart and of the mind. Elder skillfully invites the read- er on a pilgrimage of ideas, one that examines our place in nature. Elder divides his pilgrimage into three landscapes, each connected to Marsh’s vision of conservation. First is the journey to and within Tuscany, the crucible of Marsh’s thinking in the city of Florence, the surround- ing hill towns and the celebrated forests of Vallombrosa. Second, is the literary landscape where Elder situates Marsh within the lineage of William Wordsworth, Mat- suo Basho and Robert Frost. The final section returns to Vermont, to Marsh’s native Woodstock, the sugar bush of the Elder family, and to the Vermont-wide conser- vation initiatives. Those who set out on a pilgrimage usually have a destination in mind. But that is often the only sure thing about a pilgrimage. It’s what happens in between that counts and that often surprises. Nothing is hard and fast for Elder, nothing black-or-white. He writes skillfully of what he terms the “middle landscape,” that ecotone where wilderness and civilization meet. It is upon that middle ground that Elder chooses to focus, weaving the theme of “stewardship” throughout his work. The term “stewardship” has fallen on hard times in certain intellectual circles. Its anthropocentric conno- tations are too aristocratic, too old-fashioned for some. Not so for Elder, who calls for a mutuality of human history and natural history. He calls for preservation of not only biodiversity but also human values and community, a place where technology and religious faith both have their own special role to play. For Elder, the environmental crisis is ultimately a crisis of culture, a crisis of the human soul, a crisis of the human heart. Stewardship, brought into proper dialogue with con- temporary ecological, feminist, religious and social justice thought, can engender the cultural and moral resources necessary to address the environmental needs of our time. Elder’s attempt to re-frame the discussion is wel- come. Too often has environmental conversation frac- tured into dualistic dichotomies, into opposing camps of nature and culture. We need both voices to forge a social ecology, where ecosystems are not considered in the absence of culture and vice-versa. It’s that mid- dle ground that provides the vital connection between 388 remote regions of wilderness and the places were we live, work and play. We must thank Elder for eschewing sectarian lan- guage. His goal takes much more effort. It seeks a com- mon ground that does not reduce to the lowest common denominator. The built environment, the cultivated envi- ronment, the wilderness environment; they all have a voice that needs to be heard. Elder speaks in the tradi- tion of Aldo Leopold, René Dubos and Wendell Berry, voices that seek a dialogue between culture and the wild. For all three, the steward is the facilitator of such a needed task. The steward lives in that “boundary zone where the wilderness ethic may engage with recent dev- elopments of environmental history, and where the ideal of preservation transcending our narrow utilitarianism may engage with the tradition of stewardship.” (page 218) This is marvellous meditation. It’s no wonder, since Elder’s trade and tackle are words and stories. He weaves a story, a meditation, a contemplation that opens up our mind and heart to new possibilities for our rela- tionship to forests, to those wild and wonderful places. It is a story of people tending the olive groves under the warm Tuscany sun, and of his own family’s stew- arding of the sugar maple groves of their Vermont Where to Watch Birds — World Cities By Paul Milne. 2007. Yale University Press P.O. Box 209040, New Haven, Connecticut 06520-9040 USA. 496 pages. U.S. 28.00 Paper. This type of book is a reviewer’s dream. Before you open it you know the author, Paul Milne, must make choices and these will not be the same as your choic- es. This means you know you will have the opportu- nity to criticize. So how well do his choices match those I would have made? First he has included only one Canadian city, Toronto. It is Canada’s largest city but it is not the capital nor the best for the naturalist. One out of the 60 cities covered in the book seems a little thin for second largest country in the world [even if we rank 36" for population]. Putting my obvious bias aside, I noted that other key cities are missing. Oslo, with its great royal park, and Kiev, a city of wonderful chestnut trees, are two evident examples. Indeed 60% of the cities are in Europe and Asia; a disproportionate number for these two continents over North and South America and Australia. Accepting the author’s choices, how useful is this book? First this book covers more than the city itself. When I visit Victoria, British Columbia, I get up at dawn and walk from my downtown hotel through Bea- con Hill park to the coast. I return before breakfast and then go to my meeting. These two hour jaunts have net- ted me some very interesting birds [e.g. Ancient Mur- relet]. These activities were all within the downtown core of Victoria and did not involve any travel. The author includes in his version of “city” many areas that THE CANADIAN FIELD-NATURALIST Vol. 120 home. Landscape and culture provide the nexus of dis- cussion. Forgetting one or the other dehumanizes us and sets the stage for either pillage or misanthropism. In this era of apocalyptic predictions, environmen- tal and social haemorrhaging, a widening gap between culture and nature, between rich and poor, urban and rural, Elder sets out a path of reconciliation, a path that seeks the common good. It’s a path laced with com- munity, dialogue, a sense of the sacred, good work, jus- tice, stewardship, care and respect. These virtues don’t come easily. A final word to Elder: “I have come to believe that without the stories that integrate the face of nature with the drama of our human lives, society will not have the power to restrain our appetite and respect the larger balance of nature.” (page 67) If you accept this conclu- sion, read this book. It will draw you into a world of memory and elegance. If you don’t agree with Elder, read this book anyway. The full, inclusive fare offered by Elder will call you into a respectful dialogue. JOHN MCCARTHY Ecology Project, Ignatius Jesuit Centre of Guelph, P.O. Box 1238, Guelph, Ontario N1H 6N6 Canada require significant travel — you will need a car. The equivalent of me driving to Goldstream Park, 17 kilo- metres from downtown Victoria, and more. Indeed some sites are over 100 km away (distances are given in the measurement used in the country; e.g., they are in miles in the U.S. and kilometres in Canada.) Accepting the author’s broad boundaries how useful is this book? I would say very useful indeed. I have birded in almost half of the cities mentioned [gener- ally without a car] and have visited many of the sites he describes. He has an introductory section that explains the city in birder’s terms and has a good section on trans- portation. Where there is a good public transportation to suitable birding locations (e.g., London, Moscow) the author provides enough information to get the traveller started. The site descriptions are clear and accurate and I wish I had owned this book years ago. There are loca- tor maps where they are appropriate. There is a list of birds typically seen at each location. This worked well for all places and I verified with my notes that I too had seen a similar list of species. I can quibble on some minor points. For example, under Johannesburg he includes Suikerbosrand [upland species — about 70 km from Johannesburg] but omits the nearby Pilansburg — perhaps the best park to visit [lowland species — about 100+ km from Johannes- burg]. His information on travel omits to say it would be almost suicidal for a tourist to drive in Mumbai. Taxis are cheap and you can get a car and driver, sym- pathetic to your needs, from the Bombay Natural His- 2006 tory Society for reasonable cost. If you take a photo booth shot of yourself plus a glue stick to Austria, you can get a multi-day pass to Vienna’s excellent public transit; a most economical way to travel. All in all this is a great and useful book. My next trips are to Halifax and then Anchorage, but neither of these is included by the author. However, I look for- ward to using it on my next trip to one of the cities in Marshes: The Disappearing Edens By W. Burt. 2007. Yale University Press, P.O. Box 209040 New Haven, Connecticut 06520-9040 USA. 192 pages. U.S. 35.00 Cloth. William Burt has a very deserved reputation. His photographs are amazing. As a naturalist who also takes photos I have some idea of the skill and the patience needed to get a good shot. Burt goes beyond good. Not only are the photos amazing in their quality, but he spe- cialises in birds that are typically difficult to see! Take Black Rail as an example. I spent several hours at night wading in a swamp with an ardent group of seekers before I had my first glimpse of a Black Rail. It ran over my foot. It took another hour to actually get my binoculars [and flashlight] on one of the cute but elu- sive beasts. The idea of a photograph never entered my head. I do have some slides of other rail species I took mostly by good luck. They are nice, but they are not artistic and they all have little flaws. Burt’s photos are technically crisp and clear. They are also artistic; the kind you would frame and hang on the wall. In addition to birds, the author has also included a number of pictures of marsh vegetation and some of the more picturesque flowers. Actually this book is over 50% photographs. Once you have finished drooling over the illustra- tions you could read the text. The author describes his visits to wetlands throughout North America. Start- ing with his home base in Connecticut river marshes he travels to Maryland (Elliot Island), Manitoba (Sewall Lake), Saskatchewan (Crane Lake), Oregon (Malheur) and California (Klamath). He also takes us on a trip Book REVIEWS 389 this book, which will likely be Moscow. | would encour- age Paul Milne to travel more in North and South Amer- ica. Then I can hope he plans a second volume to cover some of the cities he has missed. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada through wetlands in Texas, Louisiana, Florida and Vir- ginia. At each site he describes the value that each place brings and some of the issues it now faces. Burt make an emotional, almost poetic appeal for marshes. He describes his favourite haunts along the east coast of the United States with such verve that I realised, somewhat for the first, that I too had experi- enced the same feelings. I tend to look at life more clin- ically, but Burt is more passionate and has shown that I have similar emotions to him below the surface. I believe you should learn something new every day. I began to learn as soon as | started to read. For the first time I properly understand the issue with Phrag- mites. The ones I see are more likely the aggressive — and therefore dangerous — European plant and not the look-alike native version. Alien invaders are a key threat and Phragmites and loosestrife lead that charge. His description of the work of the photographers Walter Finley and Herman Bohlman in the 1800s is enlightening on the persistence and dedication of these pioneers plus their resounding contribution to conser- vation. This a lovely book and would make a wonderful present for both naturalists and non-naturalists. The beautiful writing style and powerful messages might even convert some folks to be conservationists. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario, K1J 6K5 Canada 390 NEw TITLES Prepared by Roy John + Available for review * Assigned ZOOLOGY Albatrosses, Petrels and Shearwaters of the world. By D. Onley and P. Scofield. 2007. Yale University Press, P.O. Box 209040, New Haven, Connecticutt 06520-9040 [North America] or Christopher Helm Publishers Ltd./A&C Black Publishers Ltd., 38 Soho Square, London W1D 3HB [Europe and the Commonwealth]. 256 pages. U.S. $29.95. Catalogue of Aleocharine Rove Beetles of Canada and Alaska (Coleoptera, Staphylinidae, Aleocharine). By N. Gouix and J. Klimaszewski. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 166 pages. EURO 60 hard- back. Atlas of Bird Migration. Edited by J. Elphick. 2007. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 176 pages. $35. An Enchantment of Birds. By R. Cannings. 2007. Greystone Books, #201 — 2323 Quebec Street, Vancouver, British Colum- bia VST 487 Canada. 211 pages. $29.95. Aves amenazadas de Espana. By J. Varela. [In Spanish]. 2007. Lynx Edicions, Passeig de Gracia, 12, 4rt. 2a, 08007 Barcelona, Spain. 325 pages. U.S. $38.40. The Lapwing. By Michael Shrubb. 2007. T & AD Poyser (A&C Black Publishers Ltd, 38 Soho Square, London W1D 3HB). 320 pages. $100 (approximately). Lista Comentada de las Aves de Guatamala — Annotated Checklist of Birds of Guatamala [bilingual English Span- ish]. By K. Eiserman and C. Avendano. Lynx Edicions, Passeig de Gracia, 12, 4rt. 2a, 08007 Barcelona, Spain. U.S. $15.60 * The Birder’s Companion. By Stephen Moss. 2007. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 239 pages. All the Birds of Brazil — An Identification Guide. Second Edition. By D. De Souza. 2006. Editora Dall/Subbuteo Nat- ural History Books, The Rea, Upton Magna, Shrewsbury SY4 4UR UK. 325 pages. U.S.$60. 500 Butterflies. By K. Preston-Mafham. 2007. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 528 pages. $29.95 Cloth. Ecology and Behaviour of Chickadees and Titmice. Edit- ed by K. Otter. 2007. Oxford University Press, 198 Madison Avenue, New York, New York 10016 USA. 352 pages. U.S. $102. Cloth. *Deep Alberta — Fossil Facts and Dinosaur Digs. By John Acorn. 2007. University of Alberta Press, Ring House 2, Edmonton, Alberta T6G 2E1 Canada. 186 pages. $26.95 Paper. * Ladybugs of Alberta. By J. Acorn. 2007. University of Alberta Press Ring House 2, Edmonton, Alberta T6G 2E1 Canada. $ 29.95 Lost Land of the Dodo. By A. Cheke and J. Hume. 2007. Christopher Helm Publishers Ltd./A&C Black Publishers Ltd., 38 Soho Square, London W1D 3HB. 504 pages. U.S. $84. THE CANADIAN FIELD-NATURALIST Vol. 120 300 Frogs. By C. Mattison. 2007. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 528 pages. $29.95 Cloth. Pocket Guide to the Insects of Britain and Western Europe. By M Chinery. 2007. Christopher Helm Publishers Ltd./ A&C Black Publishers Ltd, 38 Soho Square, London W1D 3HB UK. 320 pages. U.S. $26.20. The Red Kite. By I. Carter. 2007. Arlequin Press, The Rea, Upton Magna, Shrewsbury SY4 4UR U.K. 200 pages. U.S. $44. Cloth. The Lepidoptera of Israel. Volume II: Noctuidae. By V. D. Kravchenko, M. Fibiger, A. Hausmann and G. C. Muel- ler. 2007. Pensoft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. In English, 320 pages. EURO 110. Hardback. Where to Watch Mammals in Britain and Ireland. By R. Moores. 2007. Christopher Helm Publishers Ltd /A&C Black Publishers Ltd, 38 Soho Square, London W1ID 3HB U.K. 304 pages. U.S. $29.60. Paper. Manipulative Monkeys — the Capuchins of Lomas Barbu- dal. By S. Perry and J. Manson. 2007. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 326 pages. U.S. $45. A Field Guide to New Zealand Birds. By B. Parkinson. 2007. New Holland Publishers (UK) Ltd., Garfield House, 86- 88 Edgware Road, London W2 2EA U.K. 136 pages. U'S. $22.75. Know Your New Zealand Birds. By L. Moon. 2007. New Holland Publishers (UK) Ltd., Garfield House, 86-88 Edg- ware Road, London W2 2EA U.K. 126 pages. U.S. $22.75. Collins Field Guides — Birds of the Palearctic: Passerines. By N. Arlott. 2007. Harper Collins Publishers Ltd., 2 Bloor Street East, 20th Floor, Toronto, Ontario M4W 1A8 Canada. 384 pages. U.S $46. Photographic Guide: Birds of Peru. By C. Byers. 2007. New Holland Publishers (UK) Ltd., Garfield House, 86-88 Edgware Road, London W2 2EA U.K. 144 pages. U.S. $15.95. The Sand Wasps — Natural History and Behavior. By H. Evans and K. O’ Neill. 2007. Harvard University Press, 79 Gar- den Street, Cambridge, Massachusetts 02138 USA. 340 pages. + At Sea Distribution and Abundance of Seabirds off Southern California: A 20 Year Comparison. Studies in Avian Biology Number 33. By J. Mason et al. Cooper Ornitho- logical Society, c/o Western Foundation of Vertebrate Zoology, 439 Calle San Pablo, Camarillo, California 933012-8506 USA. 101 pages. U.S. $15. Sharks of the Pacific Northwest. By A. De Maddalena, A. Preti and T. Polansky. 2007. Harbour Publishing, P.O. Box 219, Madeira Park, British Columbia VON 2HO Canada. 160 pages. $21.95 Paper. The Smaller Majority. (Insects) By P. Nasrecki. 2007. Har- vard University Press, 79 Garden Street, Cambridge, Massa- chusetts 02138 USA. 288 pages. U.S. $24.95. 2006 The Birds of Thai-Malay Peninsula — Volume 2. By D. Wells. 2007. Christopher Helm Publishers Ltd./A&C Black Publishers Ltd, 38 Soho Square, London WID 3HB U.K. 704 pages. U.S. $120. Oceanic Wilderness. By Roger Steene. 2007. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 340 pages. $59.95 Cloth. Shells. By P. Starosta and J. Senders. 2007. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 384 pages. $85. Cloth. Whales and Dolphins of the North American Pacific. By G. Cresswell, D. Walker and T. Pusser. 2007. Harbour Pub- lishing, P.O. Box 219, Madeira Park, British Columbia VON 2HO Canada. 216 pages. $21.95 Paper. BOTANY * Plants of Alberta. By F. Rover and R. Dickinson. 2007. Lone Pine Publishing. 10145 — 81 Avenue, Edmonton, Al- berta T6E 1W9 Canada. 527 pages. $ 29.95. Paper. Comparative Plant Ecology. J. Grime, J. Hodgson and R. Hunt. 2007. Springer Science+Business Media Deutschland GmbH, Heidelberger Platz 3, 14197 Berlin, Germany. 752 pages. $168. Cloth. Pocket Guide to the Trees of Britain and Northern Europe. By A Mitchel. 2007. Christopher Helm Publishers Ltd. /A&C Black Publishers Ltd, 38 Soho Square, London W1D 3HB U.K. 288 pages. U.S. $26.20 * Wild Flowers of the Rocky Mountains. By G. Scotter and H. Flygar. 2007. Whitecap Books Ltd, 351 Lynn Avenue, North Vancouver. British Columbia V7J 2C4 Canada. 255 pages. Illustrations $29.95. OTHER Digital Wildlife Photography. By D. Tipling. 2007. Firefly Books Ltd., 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 160 pages. $24.95 paper. Drawing and Painting Plants. By Christina Brodie. 2007. Timber Press, 133 SW 2"™ Avenue, Ste. 450, Portland, Oregon, USA. 144 pages. C$27.95. Paper. Book REVIEWS 39] Where to Watch Birds in Donana. By F. Chiclana and J Garzon. 2006. Lynx Edicions, Passeig de Gracia, 12, 4rt. 2a, 08007 Barcelona, Spain. 160 pages. U.S. $28. * Gilbert White. By Richard Mabey. 2007. University of Vir- ginia Press, P.O. Box 400318, Charlottesville, Virginia 22904- 4318 USA. 230 pages. Hispaniola. Edited by E. Fernandez. 2007. Harvard Univer- sity Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 400 pages. U.S. $60. * Pioneering Conservation in Alaska. By Ken Ross. 2007. University Press of Colorado, 5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado 80303 USA. 540 pages. $34.95. Cloth. This Tender Place — The Story of a Wetland Year. By Lau- rie Lawlor. 2007. The University of Wisconsin Press, 1930 Monroe Street, Third Floor, Madison, Wisconsin 53711-2059 USA. 166 pages. U.S. $19.95. Paper. * Marshes: The Disappearing Eden. By W Burt. 2007. Yale University Press P.O. Box 209040New Haven, Connecticut 06520-9040 USA. 192 pages. U.S. $35. Oregon’s Dry Side — Exploring East of the Cascade Crest. By Alan D. St. John. 2007. Timber Press, 133 SW 2° Avenue, Ste. 450, Portland, Oregon 97204 USA . 324 pages. U.S. $29.95. The Ornitholologist’s Dictionary. By J. Erritzoe, K. Kampp, K. Winker and C. Frith. 2007. Lynx Edicions, Montseny, 8, 08193 Bellaterra, Barcelona, Spain. U.S. $25. Paper. * A Paddlers’ Guide to Quetico and Beyond. By K. Callan. 2007. Firefly Books, 132 Main Street, Erin, Ontario NOB 1TO Canada. 192 pages. $ 24.95. Intelligent Courage: Natural Resource Careers that Make a Difference. By M. Fraideburg. 2007. Krieger Publishing Company, P.O. Box 9542, Melbourne, Florida 32902-9542 USA. 220 pages. U.S. $32.50. YOUNG NATURALISTS * Gracie, the Public Gardens Duck. By Judith Meyrick. 2007. Nimbus Publishing, 3731 Mackintosh Street, Halifax, Nova Scotia B3K SAS Canada. 32 pages. $16.95. News and Comment Marine Turtle Newsletter (113) July 2006. 24 pages: EDITORIAL: New editors for MTN — ARTICLES: Hawksbill and Olive Ridley nest- ing on Masirah Island, Sultanate of Oman: an update (A. F. Rees and S. L. Baker) — Preliminary data from an increasing Olive Ridley population in Sergipe, Brazil (J. C. Castilhos and M. Tiwari) — NOTES: First report of Green and Kemp’s Ridley Turtle nesting on Wassaw, Georgia, USA (K. L. Williams, M. G. Frick, and J. B. Pfaller) — LETTERS — MEETING REPORTS — MST UPDATE — ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICATIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Re- search Group, Centre for Ecology and Conservation, University of Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ United Kingdom; e-mail MTN @sea turtle.org; Fax +44 1392 263700. Subscriptions and donations towards the production of the MTN can be made online at or postal mail to Michael Coyne (online Editor) Marine Turtle Newsletter, 1 Southampton Place, Durham, North Caro- lina 27705 USA (e-mail: mcoyne @seaturtle.org). The Boreal Dip Net/L’Epuisette Boréale January 2006 Newsletter of the Canadian Amphibian and Reptile Conservation Network (CARCNET) 10(1). CON- TENTS: Editor’s Note — Faces and Places/In the Bag — Snake habitat manipulation — A request for infor- mation — CARCNET field trip — Natural Heritage Building of the Canadian Museum of Nature & Parc de la Gatineau, QC — Presentations made at the 2005 annual meeting of CARCNET/RECCAR — Reintro- duction of Leopard Frogs at Bummer’s Flats — North- ern Leopard release marks fifth year — Lost Leopard? — A significant range extension for the Northern Alligator Lizard in British Columbia — Faces and Places/Out of the bag. Membership information can be obtained from Bruce D. Pauli, Canadian Wildlife Service, National Wildlife Research Centre, Carleton University, Raven Road, Ottawa, Ontario K1A 0H3 Canada. Additional information is at website http://www.carcnet.ca. 392 Chairperson: Attendance: Mike Murphy, President 1. Minutes of the Previous Meeting Under Report of the Nominating Committee/Com- mittee Chairs: The Chair of the Education and Publicity Committee should be corrected to read Gillian Marston not John Cameron. It was moved by Gillian Marston seconded by Fenja Brodo that the minutes be accepted. (Motion Carried) 2. Business Arising from the Minutes There was no business arising from the Minutes. '3.Communications Relating to the Annual | Business Meeting There were no communications relating to the Annual ' Business Meeting. 4. Treasurer’s Report Frank Pope reviewed the financial report for the year ending 30 September 2005, noting that the Club’s net ) assets had increased by approximately $86,350. This was due in large part to a bequest from the estate of | Mildred Groh of $62,250. There was an amendment to | the Report. Moved by Frank Pope and seconded by Diane Lepage that the Financial Report be accepted as amended. (Motion Carried) 5. Committee Reports _ Mike Murphy introduced each of the Committee \reports and a representative of the appropriate Commit- tee and asked for questions and comments. He thanked _ the committee chairs and committee members for their /work over the previous year. Moved by David Hobden seconded by Gillian Marston, that the reports be accepted with suggested ‘amendments. | (Motion Carried) at '6. Nomination of the Auditor Moved by Frank Pope, seconded by Bill Cody, that Janet Gehr continue as Auditor for another year. | (Motion Carried) { Minutes of the 127" Annual Business Meeting of The Ottawa Field-Naturalists’ Club 10 January 2006 Place and time: Canadian Museum of Nature, Ottawa, Ontario, 7:30 p.m. Forty-two persons attended the meeting. Attendees spent the first half-hour reviewing the minutes of the previous meeting, the Treasurer’s report and the Report of Council. The meeting was called to order at 7:40 p.m. with some opening remarks from the President. 7. Report of the Nominating Committee President Mike Murphy Vice President Ken Allison Secretary Susan Laurie-Bourque Treasurer Frank Pope Past President Business Manager Gary McNulty Bill Cody Editor, CFN Francis Cook Editor, T&L Karen McLachlan Hamilton Committee Chairs Birds Chris Traynor Computers Dan Millar Conservation Stan Rosenbaum E&P Gillian Marston E&L Fenja Brodo Finance Ann MacKenzie FWG David Hobden Macoun rep. Diane Kitching Membership Henry Steger Publications Ron Bedford ON Rep Eleanor Zurbrigg Members at large Susan Howell Julia Cipriani Justin Peter Chairs not on Council Awards Ernie Brodo Macoun Rob Lee Nominations Fenja Brodo Retiring from the council: Kathy Conlan, Louise Schwartz, Dave Smythe Ken Allison, Julia Cipriani, Susan Howell, Ann MacKenzie, Dan Millar and Justin Peter. Moved by Fenja Brodo, seconded by Frank Pope, that the slate of nominations for the 2006 Council be accepted. New on the council: (Motion Carried) 393 394 8. New Business There was no new business. 9. Presentation of Awards for the 2005 OFNC Photo Contest There was a presentation of the awards to the win- ners of the 2005 OFNC Photo Contest by Suzanne Deschene and Gillian Marston. The grand prize winner received a digital camera and photo printer. Amphibians: Ruth Allison Birds: Lois Naggs Insects and spiders: Christine Hanrahan Mammals and vegetation: Jim Robertson Scenery: Stephen Darbyshire Honorable mention — Birds and vegetation: Grand prize winner: Gwen Williams Dave Sankster THE CANADIAN FIELD-NATURALIST Vol. 120 The Education and Publicity committee was already planning the next photo contest, tentatively titled “The Tale of the Trail”. Details to be published in Trail & Landscape. The presentation of the awards was followed by a slide show of the contest entries. This was viewed with admiration and enjoyment by the members in attendance. 10. Adjournment Moved by Henry Steger/Ron Bedford that the meet- ing be adjourned at 9:35 pm. (Motion Carried) SUSAN LAURIE-BOURQUE Recording Secretary The Ottawa Field-Naturalists’ Club Committee Reports for 2005 Awards Committee At the club’s Annual Soirée, on 30 April 2005 at St. Basil’s Church in Ottawa, awards were once again given to members, and two non-members, who distinguished themselves by ac- complishments in the field of natural history and conservation, or by extraordinary activity within the club. There isn’t al- ways a winner for every potential award, and this year, once again, the Anne Hanes Natural History Award was not given. On the other hand, a new award was approved by Council and awarded for the first time this year: the Mary Stuart Educa- tion Award. This new award has been established for members, non-members or organizations in recognition of outstanding achievements in the field of natural history education in the Ottawa Region. The new Education Award was appropriately named in honour of Mary Stuart, who was so passionately in- terested in establishing a new generation of young naturalists. The winners of the 2004 Awards were: HONORARY MEMBER: Charles D. Bird — a long-time member of the OFNC and one of Canada’s most outstanding naturalists in the fields of cryptogamic botany and entomology. He has been an associate editor of The Canadian Field-Naturalist for over 25 years. MEMBER OF THE YEAR: Marilyn Ward — for her efforts in handling the club phone, helping to make sure lecture and excursion events are suc- cessful, and keeping the Fletcher Wildlife Garden spic and span. GEORGE MCGEE SERVICE AWARD: Martha Camfield — for her many years of service with the Macoun Field Club both at meetings and in the field, and for her botanical work on the Leitrim wetlands. Mary STUART EDUCATION AWARD: Andrea Howard — for her extraordinary, innovative work in teaching natural history at the Eastern Ontario Biodiversity Museum in Kemptville and her “Museum in a Suitcase” school programs. CONSERVATION AWARD — MEMBER: Ghislaine Rozon — for her extraordinary efforts in trying to save the Larose Forest from destruction by developers. CONSERVATION AWARD — NON-MEMBER: Barbara Barr — for attending municipal and regional meet- ings and hearings regularly, mostly representing the Green- space Alliance, gathering information, presenting briefs, and being a source of sage advice to policy makers on conservation issues. The full text of the citations for each of these awards was read at the annual soirée and will be published in vol. 119, no. 4 of The Canadian Field-Naturalist. IRWIN BRODO Chair, Awards Committee Birds Committee The Birds Committee participated with the Club des Orni- thologues de |’Outaouais in both the Christmas Bird Count and the Fall Bird Count. The compilation dinners for both events were very well attended. The committee again orga- nized the Peregrine Falcon Watch at the nest site downtown. Donations to the watch in 2005 totalled $275. While both chicks fledged, one disappeared and its fate remains unknown. Sadly, the adult female, Horizon, was injured and subsequently euthanized at the request of the Ministry of Natural Resources. We continued to provide seasonal bird summaries for Trail & Landscape. Several members have also enhanced our com- mittee’s presence on the OFNC’s web site. The 5" and final year of the Ontario Breeding Bird Atlas has been completed and a report summary has been prepared. The Bird Record Sub-committee met several times during the year to review rare bird records. The committee also provided several leaders for OFNC bird trips. The committee has also added a new bird feeder, provided by the National Capital Commission, near the new picnic shelter at the Mer Bleu. The 2005 seed-a- thon raised over $300 in pledges. We continue to operate the rare bird alert and Ottawa’s bird status line which provides updates and information on both rare birds and regular report summaries. CHRIS TRAYNOR Chair, Birds Committee Computer Management Committee During 2005, the Club purchased two new computers, one for use by the Treasurer’s Assistant; old equipment has been retired. 2006 The Computer Management Committee reviewed how club volunteers manage key information holdings of the club, to ensure that club data are secure and accessible. Key files include the membership database, financial records, Trail & Landscape electronic archives, and Canadian Field-Naturalist files. The committee concluded that current data management practices are adequate. There were no requests to the Computer Management Committee from other committees during the year for com- puter support, as they had expertise available in-house. The club upgraded the capabilities of its web site consid- erably when it moved to a new service provider. These im- provements are allowing the Web Master to continue to improve the content of our popular web site. ELEANOR ZURBRIGG Chair, Computer Management Committee Conservation Committee Alfred Bog Frank Pope continues to chair the Alfred Bog Committee, while planning to hand over to a new person who has pro- visionally accepted this position. Planning Reform In follow-up to our 2004 brief and letters to Ontario Min- isters, additional letters were sent on 19 May 2005, including a joint letter to the Premier and four Ministers, and individual covering letters to each. Frank Pope and Eleanor Zurbrigg presented our position on planning reform at the Ontario Nature/Federation of Ontario Naturalists regional meeting at Alfred College on 15 October. A meeting to discuss the issues with the ON/FON regional Board Member was set for 6 December. South Gloucester (5309 Bank Street) We sent notification by registered letter to both the Minis- try of Natural Resources and R. W. Tomlinson, stating our continuing opposition to the granting of an aggregate extrac- tion licence to which we had previously objected. We pointed out that R. W. Tomlinson’s proposed mitigation measures for damage to significant plant species were dismissed in short work by Brunton Consulting Services in a second Peer Review specially ordered by the City. The issue was presently on hold pending hearings at the Ontario Municipal Board of appeals by the Greenspace Alliance and others. Pesticide Use on Lawns and Gardens Email letters were sent to the Mayor of Ottawa and Coun- cillors, supporting a campaign that is trying to get the pre- sent (ineffective) policy of voluntary reduction in pesticide use replaced by a ban on the cosmetic use of pesticides. City Council rejected a ban, but the Mayor and some Councillors in favour of a ban stated that they would make this an elec- tion issue in 2006. STAN ROSENBAUM Chair, Conservation Committee Education and Publicity Committee Kiosks, DISPLAYS AND OTHER EVENTS The committee enhanced the material available for displays and kiosks by adding several light weight portable displays. Committee members participated with displays at the Ottawa- Carleton Elementary Teachers’ Federation Publishers’ Dis- play, National Wildlife Week’s Wildlife Festival, Environment Week Fair and Eco Fair at Friends of the Farm and the OFNC Soirée. Once again the OFNC participated in the Ottawa Sci- MINUTES OF THE 127™ ANNUAL BUSINESS MEETING 395 ence Fair sponsoring a prize for a project judged by Education and Publicity Committee member Kathy Conlan. The com- mittee also arranged speakers for outside groups. OFNC MEemBeERrSHIP Drivt The portable kiosks were used for approximately 10 out- ings for a successful spring club membership drive DIGITAL LIBRARY The committee has initiated the OFNC’s ‘digital photo library and catalogue’ as a successor to the slide collection that the committee currently manages. OFNC Nature PHOTOGRAPHY CONTEST The committee ran a members’ photography contest to en- courage discovery of nature and enhance the club’s digital library. Over 200 entries were received and catalogued in the digital photo library. BROCHURES The committee continued to distribute the club brochure to area libraries and nature shops. GILLIAN MARSTON Chair, Education and Publicity Committee Executive Committee No report Excursions and Lectures Committee In 2005 this committee arranged 29 events, ten monthly meetings (including the Annual Business Meeting) plus the Soirée. The majority of our trips (twelve) were of general inter- est, two were devoted to plants, two to geology, four to mam- mals and invertebrates and nine were birding trips. The latter included our club’s biannual bus trip to Pt. Pelee which was very well attended, and two winter birding trips in conjunc- tion with the Kingston Field-Naturalists Club. There have been fewer birding trips the last five years because potential leaders were heavily involved with the Breeding Bird Atlas. Something new in 2005 was a winter wildlife tracking ex- cursion, on snowshoes, in the Gatineau. Also a first, club mem- bers were invited to join members of the Dragonfly Society of America, meeting in Armprior, on their scheduled field trips. Unfortunately a woodworking workshop had to be cancelled for lack of interest. Our policy remains to arrange meeting places that are ac- cessible by bus. The signed waiver forms allow us to track the popularity of events and help leaders to recognize parti- cipants. Our monthly meetings cover a great diversity of subjects and most of our speakers are OFNC members. FENJA BRODO Chair, Excursions and Lectures Committee Finance Committee The Finance Committee met four times in 2005. Highlights include: 1. Increase in membership fees for the calendar year 2007 Fees had not been increased for at least four years. and some concerns were noted about gradually declining mem- bership and operating deficits experienced over the previous few years. The committee recommended to Council that fees not be increased for calendar year 2006, but that the member- ship fees increase by at least $5 in 2007. These recommen- dations were approved by OFNC Council. 2. Meeting with the OFNC Auditor The Treasurer, Frank Pope, and the Chair of the Finance Committee met with the OFNC Auditor, Janet Gehr. She advised that the financial health of the club was good com- pared to similar non-profit organizations and that we had ample reserves. She also advised that our computer hardware and software were adequate, as well as the club’s record keep- ing and documentation. It was recommended that future finan- cial statements include a note on potential liabilities related to the backlog of publication of the Canadian Field-Naturalist. The amount of deferred revenue which would cover a part of this financial commitment should also be noted. This recom- mendation was acted upon by the Treasurer. 3. Budget for 2005-06 Fiscal Year A proposed budget for the OFNC and the Canadian Field-Naturalist for the period 1 October 2005 to 30 Septem- ber 2006 was submitted to Council. A deficit of over $18,000 was forecast, and if realized would be a draw on the unrestrict- ed reserve. Council approved this proposed budget. 4. Other Items During the year the committee discussed the impact of changes in the federal government’s Publication Assistance Program on mailing costs for The Canadian Field-Natural- ist and Trail & Landscape, the increase to the audit fee from $1,000 to $2,000, a credit card for club business (not ap- proved), the bequest from the estate of M. Groh, and reviewed the draft financial statements for 2004-05 before finalization. LOUISE SCHWARTZ Chair, Finance Committee Fletcher Wildlife Garden Committee The Garden completed another successful year with over 2000 volunteer hours contributed. We held or participated in a number of events. In February, it was the Great Backyard Bird Count and then an information table at the Pesticides Conference. April brought the Wildlife Festival with a dis- play at Billings Bridge Shopping Centre, joint sponsorship of a seminar at the Canadian Museum of Nature and an event at our Centre for Annual Migratory Bird Day. Our annual plant sale in June raised over $2000. We also participated in a Cen- tral Experimental Farm Open House held in September. Without a grant we were only able to have a part-time employee who worked on the interpretation program. Our preferred employee was unable to start immediately, so the program began later and ran through September. Remodelling of the Backyard Garden continued. Some beds have been reshaped, a new entrance sign added and the patio re-laid. In 2006 most of the beds will contain only regional native flowers. CANADIAN Type Local Other Family 312 (301) 23 (19) Individual 307 (310) 101 (107) Honorary 14 (14) 11 (10) Life 21 (21) 20 (20) Sustaining 10 (11) 2 (4) Total 664 (657) 157 (160) THE CANADIAN FIELD-NATURALIST Vol. 120 Our work on invasive plants continues. There is some suc- cess with buckthorn and garlic mustard, but swallow-wort continues to be a problem. All methods of removing it end up destroying everything else as well. We are also working on burdock, Canada thistle and amur maple which have become too common. Flowering rush and cat-tails are being removed from the pond. The pond itself is showing decreased biodi- versity with very few insect larvae observed in 2005. The pri- mary water source is agricultural run-off so this trend may be difficult to reverse. In the spring we published a new general brochure in En- glish, using a grant from TD Friends of the Environment Foundation and intend to seek another grant to publish the French version. Information sheets on burdock and garlic mustard were also produced. For the Central Experimental Farm Open House a bilingual miniguide was prepared to give an overview of what the Fletcher Wildlife Garden has to offer. This is now available to all visitors. We have renewed the main entrance sign, enhanced the boundary sign system and the direction signs along the Bill Holland Trail. Other improvements include a small shed and gravel on heavily used parts of the trail system. During the winter there was considerable rabbit damage. We must expect this problem to continue until the local fox population recovers. DaviD HOBDEN Chair, Fletcher Wildlife Garden Committee Macoun Club Committee The committee met once during the year, with most of the planning being coordinated by telephone and e-mail. Commit- tee members supervised or gave presentations at 18 indoor meetings (down from 44 last year) and led 14 field trips and one camping trip (roughly the same). The much reduced number of meetings stems from the failure of a Senior (high-school-age) group to form this year, apparently for the first time in the Club’s 57-year history. In part, this follows from weak participation over a period of years in the Intermediate (middle-school) group, which would normally feed into the Seniors. But it also appears to be a social phenomenon of a proliferation of competing activities, and ever more tightly scheduled lives. Participation in the Junior group remains strong, although the same social pressures have become evident in the fami- lies of younger children, too. With no Senior members, the Macoun Field Club’s annual publication (The Little Bear) had to be edited and produced by a committee member for the first time since 1988. The content, however, continues to be dominated by the children’s work. ROBERT E. LEE Chair, Macoun Club Committee FOREIGN USA Other Total 2 (1) 1 (1) BS 8ma(622) 18 (19) 5 (4) 431 (440) 0 (0) 0 (0) 25 (24) 7 (6) 1 (1) 49 (48) 0 (0) 0) (0) 12 S)) 27 (26) 7 (6) (855) (849) 2006 Membership Committee The distribution of memberships for 2005 is shown in the table on previous page, with the comparable numbers for 2004 in parentheses. These statistics do not include the 23 affiliate organizations which receive complimentary copies of Trail & Landscape. During 2005, the club lost a long time member with the death of Eileen Evans, who joined the club in 1974 and served on Council as Corresponding Secretary from 1990 to 1996. Eileen was an important organizer and contributor to the social part of the monthly meetings at the Canadian Museum of Nature and the annual Soirée for many years. DAVE SMYTHE Chair, Membership Committee Publications Committee The Publications Committee met three times in 2005. Only two issues of The Canadian Field-Naturalist were published in 2005: Volume 118, numbers two and three. These MINUTES OF THE 127™ ANNUAL BUSINESS MEETING 397 two issues contained a total of 336 pages; 32 articles; 15 notes; 41 book reviews; 71 new titles; | commemorative trib- ute; 14 pages of News and Comments; and 2 pages of miscel- lany. The journal fell further behind schedule largely because of some problems with the (relatively new) printer. These problems seem to have been solved, and the next four issues were near to being ready for production. The possibility of eventual electronic publication was considered at some length, and is continuing. For the present, the Abstracts of CFN papers will be posted on the OFNC website (to accompany the Tables of Contents which have been appearing there for some time). None of the Manning funds were drawn upon in 2005. Volume 39 of Trail & Landscape was published in four issues containing a total of 200 pages having the usual excel- lent mix of articles. RONALD E. BEDFORD Chair, Publications Committee 398 THE CANADIAN FIELD-NATURALIST Auditor’s Report To The Members of THE OTTAWA FIELD NATURALISTS’ CLUB I have audited the balance sheet of THE OTTAWA FIELD NATURALISTS’ CLUB as at 30 September 2005, the statement of changes in net assets, and the statements of operations. These financial statements are the res- ponsibility of the organization’s management. My res- ponsibility is to express an opinion on these statements based on my audit. Except as explained in the following paragraph, I conducted my audit in accordance with Canadian gen- erally accepted auditing standards. Those standards re- quire that I plan and perform an audit to obtain reason- able assurance whether the financial statements are free of material misstatement. An audit includes exam- ining evidence supporting the amounts and disclosures in the financial statements. An audit also includes as- sessing the accounting principles used and significant estimates made by management, as well as evaluating the overall financial statement presentation. In common with many non-profit organizations, the Ottawa Field-Naturalists’ Club derives some of its revenue from donations and fund raising activities. These revenues are not readily susceptible to complete audit verification, and accordingly, my verification was limited to accounting for the amounts reflected in the records of the organization. In my opinion, except for the effect of the adjust- ments, if any, which I might have determined to be necessary had I been able to satisfy myself concern- ing the completeness of the revenues referred to in the preceding paragraph, these financial statements present fairly, in all material respects, the financial position of the OFNC as at September 30, 2005, and the results of its operations and changes in net assets for the year then ended in accordance with Canadian generally accepted accounting principles. JANET M. GEHR Chartered Accountant North Gower, Ontario 3 January 2006 The Ottawa Field-Naturalists’ Club Balance Sheet September 30, 2005 ASSETS CURRENT Cash (Note 1) Investment certificates (Note 1) Marketable securities (Note 2) Accounts receivable Prepaid expenses LAND — ALFRED BOG 2005 $ 27,819 29 401 0 21,837 1,000 80,057 3,348 MARKETABLE SECURITIES (Note 2) 338,834 LIABILITIES AND FUND BALANCES CURRENT Accounts payable and accrued liabilities Deferred revenue LIFE MEMBERSHIPS NET ASSETS Unrestricted Club reserve Manning principal Manning interest - OFNC - CFN Seedathon Anne Hanes memorial de Kiriline-Lawrence Macoun Baillie Birdathon Alfred Bog $ 422,239 $ 3,500 12,049 15,549 14,079 147,823 100,000 100,000 1,014 20,490 810 870 16,619 1,180 3,805 392,611 $ 422,239 Vol. 120 2004 $ 22,306 29,175 69,310 16,703 1,000 138,494 3,348 191,740 $ 333,582 $ 2,000 11,714 13,714 13,607 62,667 100,000 100,000 2.457 15,805 1,281 870 18,429 1,052 3,700 306,261 $ 333,582 2006 MINUTES OF THE 127™ ANNUAL BUSINESS MEETING The Ottawa Field-Naturalists’ Club Statement of Operations for the Year Ended September 30, 2005 2005 REVENUE Memberships $ 13,635 Trail and Landscape 220 Interest 2,129 GST rebate 720 Other nae OLE 17,326 OPERATING EXPENSES Affiliation fees 652 Computer 1,846 Membership 1,363 Office Assistant 1,000 Telephone 1,733 Insurance WS) Audit 2,500 GST ISS) Other 1,072 12,646 CLuB ACTIVITY EXPENSES Awards 390 Birds 618 Education and Publicity 1,149 Excursions and Lectures (1,805) Macoun Field Club 466 Origins and History of OFNC 3,800 Soiree 228 Trail and Landscape 8,887 Fletcher Wildlife Garden (Note 4) 3,667 Other 0 17,400 Excess EXPENSES OVER REVENUE $ (12,720) 2004 $ 13,969 280 1,835 768 373 17,225 670 1,294 1,168 1 000 1,640 U25 1,000 1,214 952 9,663 150 540 775 (508) 606 0 270 9578 (1,734) 84 9,761 $ (2,199) The Ottawa Field-Naturalists’ Club 399 The Canadian Field-Naturalist — Statement of Operations for the Year Ended September 30, 2005 2005 REVENUE Memberships $ 9,090 Subscriptions 25,135 Reprints 6578 Publication charges 23,270 Interest and exchange 9 824 GST rebate 2,503 Other 1 23t 77,631 EXPENSES Publishing 20,777 Reprints 4,658 Circulation 8532 Editing 2394 Office Assistant 5,000 Honoraria 9 000 GST rebate 2,693 Other 191 53,245 EXcCEss EXPENSES OVER REVENUE $ 24386 2004 $ 9313 23 936 5 574 23,685 8 290 3,889 1,477 76,164 76,823 $ (659) 400 THE CANADIAN FIELD-NATURALIST The Ottawa Field-Naturalists’ Club Notes to the Financial Statements September 30, 2005 1. CASH Chequing Savings Nesbitt Burns Fletcher Wildlife Garden Investment Certificates: 2005 2. MARKETABLE SECURITIES 2005 Investment Certificates: Province of Newfoundland Coupon Province of Ontario Coupon Province of Manitoba Coupon Res CIBC Int BB6 CMHC Global Debs Province of Ontario Bond Province of Newfoundland Bond Government of Canada Coupon Province of New Brunswick Bond Province of New Brunswick Bond 3. CAPITAL ASSETS Equipment at a cost of $16,748 is fully amortized. 4. FLETCHER WILDLIFE GARDEN REVENUE Human Resources and Skills Dev. Canada TD Friends of the Environment Fund Taverner Cup Sales GST Donations EXPENSES Program Backyard Habitats Interpretation centre Administration Publications GST Library Maturity Value $ 29,963 Maturity Value $ 44,782 5237/16 45,740 70,827 52,000 30,000 20,000 30,167 60.000 20,000 Maturity Date 05/01/06 Maturity Date 10/17/11 12/02/12 09/05/13 10/31/14 12/01/06 09/12/07 10/07/08 12/01/09 12/03/15 06/15/10 2005 $ 16,144 6,963 459 4,253 $ 27,819 Yield 1.90% Yield 4.525% 4.591% 4.694% 4.144% 5.250% 6.125% 6.263% 5.605% 3.965% 6.231% 1,479 388 112 “aan $(3,667) Vol. 120 2004 $ 11,899 4,49] 0 5,916 $ 22,306 Book Value $ 29.401 Book Value $ 34,509 11,106 31,909 48 934 53,539 31,187 20,538 23,534 62,510 21,068 $338,834 8460 $(1,734) 2006 MINUTES OF THE 127™ ANNUAL BUSINESS MEETING 40) The Ottawa Field-Naturalists’ Club Statement of Changes in Net Assets for the Year Ended September 30, 2005 (Note 5) Net Beginning Excess Excess Other Other Ending Assets Balance Revenue CFN —_ Expenses OFNC Revenue Expenses _—_ Balance Unrestricted $ 62,667 $ 24,386 $ (12,720) $ 73490a $ $ 147,823 Club reserve 100,000 ~ ~ : 100,000 Manning Principal 100,000 - ~ 100,000 Manning - CFN 15,805 - - 4,685 20,490 Manning - OFNC 2,457 - - 1,172 (2,615 b) 1014 Seedathon 1,28] - - 807 (1,278 c) 810 Anne Hanes Memorial 870 - - ~ ~ 870 de Kiriline-Lawrence 18,429 - - 190 (2,000 d) 16,619 Macoun Baillie Birdathon 1,052 - - 128 - 1,180 Alfred Bog 3,700 - - 105 ~ 3,805 $ 306,261 $ 24,386 $ (12,720) $ 80,577 $ (5,893) $ 392,611 5. STATEMENT OF CHANGES IN NET ASSETS a) Unrestricted other revenue includes a bequest of $62,250 from the Estate of Mildred Groh. b) Manning OFNC expenses: Innis Point Bird Observatory $2,000, Peregrine Falcon Watch $649. c) Seedathon Expenses: birdseed for club feeders, $809. d) Louise de Kiriline-Lawrence Expenses: Contribution to the defence of the Larose Forest at a hearing of the Ontario Municipal Board. 6. PUBLICATION LIABILITY An annual membership fee or subscription entitles the member/subscriber to four issues of The Canadian Field- Naturalist based on a calander year. As the year end of the club is September 30, the Club incurs a liability for publishing the fourth issue of each publication. At this time, however, the publication of The Canadian Field-Naturalist is ranning late. At September 30, 2005, the club owes members/subscribers of 2005 issue number 4 of 2004 and three issues of 2005. Although most of the work preparing these publications is done by volunteers, the club must pay for printing and mailing. Based upon recent costs, it is estimated that the club has a liability of $68,000 for the outstanding issues. This amount will be reduced by page charg- es to the authors in the amount of 40% of the printing costs. The Ottawa Field-Naturalists’ Club Summary of Significant Accounting Policies September 30, 2005 1. Nature of Business The organization is non-profit and incorporated under the laws of Ontario (1884). The organization promotes the appre- ciation, preservation, and conservation of Canada’s natural heritage. It encourages investigation and publishes the results of the research in all fields of natural history and diffuses infor- mation on these fields as widely as possible. It also supports and cooperates with other organizations engaging in pre- serving, maintaining or restoring environments of high quality for living things. 2. Financial Instruments The organization’s financial instruments consist of cash, accounts receivable, marketable securities, and accounts pay- able. Unless otherwise noted, it is the management’s opinion that the organization is not exposed to significant interest, cur- tency, or credit risks arising from these financial instruments. The fair value of these instruments approximate their carry- ing values, unless otherwise noted. 3. Capital Assets Capital assets in excess of $4,000 cost are recorded as assets at cost and amortized on a straight-line basis. These assets have been fully amortized. 4. Statement of Changes in Financial Position A statement of changes in financial position has not been provided as it would not provide additional meaningful infor- mation. 5. Foreign Currency Transactions during the year in U.S. dollars have been con- verted in the accounts to Canadian dollars at the exchange rate effective at the date of the transaction. All monetary assets in U.S. dollars at year end have been converted to Canadian dollars at the rate effective on Sept. 30, 2005. Gains or losses resulting therefrom are included in revenue or expenses. Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit by post to the Editor, in either English or French, three complete manuscripts written in the journal style. Manuscripts may also be submitted (one copy) by e- mail. The research reported should be original. It is recom- mended that authors ask qualified persons to appraise the paper before it is submitted. All authors should have read and approved it. Institutional or contract approval for the publica- tion of the data must have been obtained by the authors. Also authors are expected to have complied with all pertinent leg- islation regarding the study, disturbance, or collection of ani- mals, plants or minerals. The place where voucher specimens have been deposited, and their catalogue numbers, should be given. Latitude and longitude should be included for all indi- vidual localities where collections or observations have been made. Manuscripts should be printed on standard-size paper, dou- blespaced throughout, generous margins to allow for copy marking, and each page numbered. For Articles and Notes provide a bibliographic (citation) strip, an abstract, and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manu- scripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in a separate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “Canadian” or “American” spellings are acceptable in English but should be consistent within one manuscript. 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Cook, Editor RR 3 North Augusta, Ontario KOG IRO Canada 402 TABLE OF CONTENTS (concluded) Volume 120 Number 3 2006 Book Reviews Zoo.oGy: The Birder’s Companion — Birds of the Dominican Republic and Haiti — Atlas of Bird Migration: Tracing the Great Journeys of the World’s Birds — Handbook of the Birds of the World Volume II — Old World Flycatchers to Old World Warblers — Conservation of the Black-tailed Prairie Dog: Saving North America’s Western Grasslands — Migrating Raptors of the Word: Their Ecology and Conservation — Land Snails of British Columbia — Songbird Journeys: Four Seasons in the Lives of Migrating Birds 379 Borany: Wildflowers of the Rocky Mountains — Plants of Alberta 386 MISCELLANEOUS: Pilgrimage to Vallombrosa: From Vermont to Italy in the Footsteps of George Perkins Marsh — Where to Watch Birds: World Cities — Marshes: The Disappearing Edens 387 New TITLES News and Comment Marine Turtle Newsletter 113 — The Boreal Dip Net/L’Epuisette Borale January 2006 392 inutes of the 127" Annual Business Meeting of The Ottawa Field-Naturalists’ Club 393 Advice to Contributors 402 Mailing date of the previous issue 120(2): 1 November 2007 THE CANADIAN FIELD-NATURALIST Volume 120 Number 3 Articles An ethogram developed on captive eastern Coyotes, Canis latrans JONATHAN G. WAY, DEAN-LORENZ M. SZUMMMYLO, and ERIC G. STRAUSS First observations of an Eastern Screech-Owl, Megascops asio, population in an apple-producing region of southern Quebec NGAIO L. RICHARDS, PIERRE MINEAU, DAVID BIRD, PIERRE WERY, JACQUES LARIVEE, and JASON DUFFE Effects of timber harvesting and plantation development on cavity-nestng birds in New Brunswick STEPHEN J. WOODLEY, GREG JOHNSON, BILL FREEDMAN, and DAVID A. KIRK Recent invasion, current status, and invasion pathway of European Common Reed, Phragnites australis subspecies australis, in the southern Ottawa District PAUL M. CATLING and SUSAN CARBYN Pack Size of Wolves, Canis lupus, on Caribou, Rangifer tarandus, winter ranges in west-central Alberta GERALD W. KUZYK, JEFF KNETEMAN, and FIONA K. A. SCHMIEGELOW Bur Buttercup, Ranunculus testiculatus , new to eastern Canada MICHAEL J. OLDHAM, CLIVE E. GOODWIN, and SEAN BLANEY Assessment of effects of an oil pipeline on Caribou, Rangifer tarandus granti, use of riparian habitats in arctic Alaska, 2001-2003 LYNN E. NOEL, MATTHEW W. BUTCHER, MATTHEW A. CRONIN, and BILL STREEVER First records of the Southern Red-backed Vole, Myodes gapperi, in the Yukon Territory THOMAS S. JUNG, ANY RUNCK DAvID W. NAGORSEN, BRIAN G. SLOUGH, and TODD POWELL A conservation evaluation of Smooth Goosefoot, Chenopodium subglabrum (Chenopodiaceae), in Canada DIANA BIZECKI ROBSON Summer movements and impact of individual Striped Skunks, Mephitis mephitis, on duck nests in Saskatchewan SERGE LARIVIERE, LYLE R. WALTON, and FRANCOIS MESSIER Life history phenology and sediment size association of the dragonfly Cordulegaster dorsalis (Odonata: Cordulegasteridae) in an emphemeral habitat in southwestern British Columbia LAURIE B. MARCZAK, JOHN RICHARDSON, and MARIE-CLAIRE CLASSEN An analysis of the vascular flora of Annapolis heathlands S. CARBYN, P. M. CATLING, S. P. VANDER KLOET, and S. BASQUILL Pacific Hagfish. Eppptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and scavenger activity on tethered carrion in subtidal benthic communities off western Vancouver Island SARAH DAVIES, ALI GRIFFITHS, and T. E. REIMCHEN Notes A new record size Wolf, Canis lupus, pack for Ontario Liv S. Vors and PHILIP L. WILSON Tributes A tribute to Nicholas Stephen Novakowski 1925-2004 JOSEPH E. BRYANT ISSN 0008-3550 2006 263 (continued on inside back cover) CAIN IS AY The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 1 Volume 120, Number 4 October-December 2006 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Michaélle Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- ments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Donald M. Britton Anthony J. Erskine Hue N. 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It is available to Libraries at $33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: October- December 2006 (January 2008). ————- > | Cover: Incubating female Northern Hawk Owl Surnia ulula, remained seated on the nest for long periods of time, changing her position infrequently (nest number 1). Photo by Michael Patrikeev. 30 May 2001. See article pages 433-437. The Canadian Field-Naturalist | ™ 04 2008 Volume 120, Number 4 October—Decémber 2906 A Reevaluation of Sexual Dimorphism in the Postcranium of the Chasmosaurine Ceratopsid Chasmosaurus belli (Dinosauria: Ornithischia) JORDAN C. MALLON! and ROBERT B. HoLmMeEs2+ 'Department of Earth Sciences, Carleton University, Ottawa, Ontario K1S 5B6 Canada; e-mail: jmallon@ucalgary.ca Research Division, Palaeobiology, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario KIP 6P4 Canada Current address: Faculty of Veterinary Medicine, University of Calgary, 3330 Hospital Drive, Calgary, Alberta T2N 4N1 Canada; e-mail: jmallon@ucalgary.ca ‘Current address: Department of Biological Sciences, University of Alberta, CW 405 Biological Sciences Building, Edmonton, Alberta T6G 2E9 Canada; e-mail: holmes! @ualberta.ca Mallon, Jordan C., and Robert B. Holmes. 2006. A reevaluation of sexual dimorphism in the postcranium of the chasmosaurine ceratopsid Chasmosaurus belli (Dinosauria: Ornithischia). Canadian Field-Naturalist 120(4): 403-412. The sexual dimorphism attributed to Chasmosaurus belli by Sternberg (1927) is revisited and reevaluated. A reexamination of the two specimens originally considered by Sternberg reveals that they are less complete than first suggested, with only a moderate amount of overlapping material between them. Only a few of the postcranial elements (humeri, sternal plates, and presacral vertebrae) show evidence of dimorphism, the significance of which is either doubtful or equivocal. Instead of repre- senting sexual dimorphs, it is likely that the two specimens belong to separate species, C. belli and C. russelli, as evidenced by their distinct frill morphologies and by their stratigraphic segregation within the Dinosaur Park Formation of Alberta. These findings emphasize the need to remain sceptical about claims advocating sexual dimorphism in the fossil record in the absence of statistical significance or stratigraphic control. Key Words: Chasmosaurus, Ceratopsia, Dinosaur Park Formation, sexual dimorphism, variation, Alberta. Chasmosaurus are modest-sized (~1500 kg) ceratop- sid dinosaurs known from the upper Campanian-aged beds (~75 Ma [million years ago]) of western North America. They are characterized by the possession of a premaxillary flange, recurved supraorbital horncores, and a broad parietosquamosal frill with a straplike pos- terior border (Forster et al. 1993). The type species, Chasmosaurus belli (Figure 1), was discovered in the middle strata of the Dinosaur Park Formation in Alber- ta and was originally named Monoclonius belli (Lambe 1902). Two additional species of Chasmosaurus, C. russelli and C. irvinensis, are currently recognized from elsewhere in the formation (Godfrey and Holmes 1995; Holmes et al. 2001; Ryan and Evans 2005). A further species, C. mariscalensis, has been described from the Aguja Formation in Texas (Lehman 1989), though it was recently given the new genus name Agujaceratops (Lucas et al. 2006) and may prove to be more closely related to Pentaceratops than to Chasmosaurus (Holmes et al. 2001; Diem and Archibald 2005). Charles M. Sternberg (1927) reported on two spec- imens of Chasmosaurus, both identified as C. belli, on display at the National Museum of Canada (now the Figure 1. Life restoration of Chasmosaurus belli, based on Canadian Museum of Nature) in Ottawa (Figure 2). CMN 2245. Illustration by JCM. 403 404 the Canadian Museum of Nature). As restored, the skeletons measured “approximately the same length but the bones of one... are much lighter in construction than those of the other” (Sternberg 1927: 67). According to Sternberg, the total length of the ‘gracile’ skeleton measured 4.93 m, while that of the “robust” skeleton measured 4.95 m. Unfortunately, their supports have since been disassembled and these original measurements cannot be verified. Sternberg attributed the difference in robustness between the two specimens to sexual dimorphism, stating, “it is thought that they represent male and female as it is common, among reptiles, for the female to be larger than the male” (Sternberg 1927: 67). This assumption has gone largely unchallenged in the literature (e.g., Lull 1933; Carpenter and Currie 1990; Dodson 1996), except for an implication by Godfrey and Holmes (1995) that the reported differences between the specimens do not reflect sexual dimorphism because they pertain to two separate species of Chasmosaurus. The recognition of sexual dimorphism in a fossil species must begin with the establishment that the vari- ation observed within that species does represent dimor- phism. With only two reasonably complete skeletons of Chasmosaurus belli at his disposal, Sternberg could not have established this with certainty, and indeed, he neither quantified nor qualified the differences he observed between the specimens. It is also important to note that many of the bones were highly reconstruct- ed with plaster and subsequently painted over so that the reconstructed portions cannot be easily identified. THE CANADIAN FIELD-NATURALIST Vol. 120 FIGURE 2. CMN 2280 (left) and CMN 2245 (right) on display at the National Museum of Canada (circa 1926). (Courtesy of As chronicled by C. H. Sternberg (C. M. Sternberg’s father) regarding the restoration of one of the skulls: “we had colored our plaster to resemble the fossil bone —no small task, by the way, as we had to learn to mix colors as well as do the work of a sculptor — with wax” (Sternberg 1917: 83). As such, the possibility remains that the difference in robustness between the two specimens might in part be attributable to the extensive restoration of certain elements. With these concerns in mind, we set out to identify and describe the variation noted between the skeletons, and con- ducted a reevaluation of Sternberg’s original attribu- tion of sexual dimorphism to C. belli. Materials The two Chasmosaurus belli skeletons (Figure 3) described by Sternberg (1927) reside in the collec- tions of the Canadian Museum of Nature in Aylmer, Quebec. The “gracile” specimen was designated CMN 2245; the “robust,” CMN 2280. CMN 2245 was col- lected from quarry Q037, and CMN 2280 from QO10, of the Dinosaur Park Formation (upper Campanian, ~75 Ma) in Dinosaur Provincial Park, Alberta (origi- nally referred to by Sternberg [1927: 67] as the “Belly River series’). CMN 2245 (Figure 3A) is represented by the pos- terior half of a skull and mandibles (missing only the predentary), complete presacral series, synsacrum com- plete to the sixth sacral vertebra, twenty-four caudal vertebrae, most cervical and thoracic ribs, pectoral gir- 2006 MALLON AND HOLMES: SEXUAL DIMORPHISM IN CHASMOSAURINE CERATOPSID 405 [ ’ . Utter re... = : SETAE Vii" cee FIGURE 3. Material (in grey) attributed to (A) CMN 2245 and (B) CMN 2280. Scale bar = 1 m. Modified from original skeletal drawing by Gregory S. Paul. Used with permission. 406 dle (missing only the right coracoid), both humeri, left ulna, pelvic girdle, both femora, right tibia and fibula, and several carpal and tarsal elements. CMN 2280 (Figure 3B) consists of a complete skull and mandibles, complete presacral series, synsacrum complete to the fourth sacral vertebra, most cervical and thoracic ribs, pectoral girdle (missing only the left coracoid), both humeri, anterior halves of both ilia, both pubes, and the proximal third of the left femur. A return visit to the quarry has yielded more hindlimb material, but it was in poor condition and not collected (D. H. Tanke, personal communication). Overlapping material between the two specimens therefore consists of the posterior region of the skull, presacral and anterior sacral vertebrae, most ribs, most of the pectoral girdle, humeri, anterior ilia, pubes, and left proximal femur. Methods In order to facilitate distinction between the original bone and plaster, the brown paint was removed from many of the bones using acetone and a stiff-bristled brush. In some instances, the removal of the paint was not warranted when it was obvious that the element of interest had been created entirely of plaster (in which case a simple tap with the knuckle would confirm sus- picion). In this way, the original elements common to CMN 2245 and CMN 2280 could be identified. Three hundred and fourteen measurements were then taken from the postcranium of each specimen to the nearest millimetre, primarily according to the standards of Chinnery (2001). For those elements not considered by Chinnery, such as those belonging to the axial skele- ton, one of us (JCM) devised our own measurement parameters. Measurements below 300 mm were gen- erally taken with Mitutoyo SD type dial callipers and measurements above 300 mm were taken either with simple outside callipers and measuring tape or with large (1.04 m) Helios brand Vernier callipers. Circumferences were measured using a tailor’s measuring tape. Angles were measured from photographs using a protractor. All available postcranial elements were measured, but only those bones deemed reliably complete and shared between the two skeletons were considered in this study. Comparative Osteology Description of Overlapping Material Although CMN 2245 and CMN 2280 exhibit mod- est variability in the dimensions of their overlapping elements (Figure 4), much of this appears to be due to the extensive restoration and occasional postmortem deformation of the skeletons. Despite this, the best pre- served elements usually differ in size by only a few percent, which is insufficient to produce visually dis- cernable dimorphism, and is probably within the realm of individual variation that might be expected in tet- rapods (e.g., Rising and Somers 1989; Zaaf and Van Damme 2001; Kelly et al. 2006). The coracoids of THE CANADIAN FIELD-NATURALIST Vol. 120 CMN 2245 and CMN 2280 are noticeably different in shape, although the left and only coracoid preserved in the former specimen has been badly weathered and highly reconstructed as a result, and is therefore of limited use. Only a few bones differ considerably in several dimensions and are described below. Humerus The humerus (Figure 4A) of CMN 2280 is consid- erably more robust than that of CMN 2245 in most dimensions (Table 1), particularly those of the humer- al head, deltopectoral crest, and midshaft circumfer- ence. Besides proportional differences in the humerus, the insertional scar for the /atissimus dorsi muscle on the deltopectoral crest is much less pronounced in CMN 2245. Sternal Plate Although the sternal plate (Figure 4B) is quite sim- ilar in size and shape between the two specimens, the process at the posterior end of the element extends 42-55% further laterally in CMN 2280 than in CMN 2245. Presacral Vertebrae The presacral vertebrae (Figure 4C) of these speci- mens are uniform in most dimensions, with most appar- ent differences being attributable to the addition of plaster. However, in the region of the withers (anteri- or thoracics), the vertebrae of CMN 2280 average 21% taller (Figure SA) and their neural spines 19° more erect (Figure 5B) than in CMN 2245. Similarly, the transverse processes of the cervical and anterior thoracic vertebrae of CMN 2280 average 17° more erect than in CMN 2245 (Figure 5C), although this is difficult to state with confidence given the artificial coalescence of the vertebrae and the incompleteness of many of their spinous processes in CMN 2245. It seems, too, that the transverse widths of the vertebral centra anterior to the sixth thoracic average 10% wider in CMN 2280 (Figure 5D). Discussion Preservation of CMN 2245 and CMN 2280 While CMN 2245 is the most complete skeleton of Chasmosaurus known to date, it is less complete and less well preserved than generally supposed (e.g., Dod- son 1996: 107). Some of the elements touted by Stern- berg (1927) as being complete, such as the femora and caudal vertebrae, are actually only partially represent- ed by nondescript scraps of bone buried in plaster, causing confusion among later authors (e.g., Lull 1933). Many of the presacral vertebrae have also been artifi- cially “fused” together with plaster, obscuring much of the detail. CMN 2280 is by far the better preserved of the two specimens, although it is less complete. Consequently, there is only a moderate amount of mate- rial shared between the two specimens, and additional overlapping material would be desirable to make a con- vincing case for dimorphism in C. belli. 2006 MALLON AND HOLMES: SEXUAL DIMORPHISM IN CHASMOSAURINE CERATOPSID 407 FiGurE 4. Dimorphic overlapping material of (A) CMN 2245 and (B) CMN 2280. i, right and left humeri; ii right and left sternal plates; iii posterior thoracic vertebrae in anterior (left) and left lateral (right) views. Note that, because of the poor preservation of the overlapping vertebrae, the eighth thoracic vertebra is figured for CMN 2245 and the tenth thoracic vertebra is figured for CMN 2280. Scale bar = 10 cm. A Reevaluation of Sexual Dimorphism in Chasmosaurus __ et al. 2005). Chinnery (2001, 2004) attempted to discern belli morphological variation, including sexual dimorphism, Recognizing sexual dimorphism in a fossil species in the appendicular skeleton of the Ceratopsia, but is rarely a straightforward task, especially given small _ could find none. The dimorphism originally ascribed by Sample sizes where subtle but statistically significant Sternberg (1927) to CMN 2245 and CMN 2280 was dimorphic characters are difficult to resolve (Padian _ presumably based on variation in the postcranial skele- 408 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Selected measurements (after Chinnery 2001) for left humeri of CMN 2245 and CMN 2280, showing the large differences between their dimensions. Dimensions in square brackets are estimated. Measurement parameter Width of deltopectoral crest Deltopectoral crest length, from external tuberosity to distal end of deltoid muscle scar Craniolateral view; proximal width Head height Head width Circumference about midshaft* * = parameter not used in Chinnery (2001). ton, although distinct differences in frill morphology are also present (see below). These differences cannot be attributed to either allometry or ontogeny since both specimens are of comparable size. Nor are they like- ly due to geographic variation since both specimens were found less than nine kilometres from one anoth- er. Post-mortem distortion cannot account for the dif- ferences either, as most of the elements are relatively uncrushed. A review of the overlapping material reveals that the most striking differences between CMN 2245 and CMN 2280 lie with the humerus. The disparity in the robustness of the humerus and in the size of the inser- tional scar for the Jatissimus dorsi muscle seems to follow the prediction made by Chapman et al. (1997) that sexual dimorphism in ceratopsians would be ex- pressed in the limb bones. It is also reminiscent of the condition seen in the humerus of many pachypleu- rosaurid sauropterygians (e.g., Sander 1989; Cheng et al. 2004) and of the sauropod Camarasaurus (Ike- jiri 2005). However, if these humeral characters alone separate male from female, then other specimens of Chasmosaurus belli should exhibit the same disparity in humeral robustness as well. A bivariate plot derived from a Principle Components Analysis by Chinnery (2001: 134) revealed that the humerus of CMN 2245 plots as an outlier from the other eleven Chasmosaurus humeri included in the study. As the deviation lies along the y-axis, the differences are likely due to shape rather than size. Thus, while the shape differences between the humeri of CMN 2245 and CMN 2280 are real, the highly asymmetrical sex ratio implied is very unlikely. The inability of Lehman (1990) to identify dimorphism in the humeri from a population of C. mariscalensis, and M. J. Ryan’s (personal communication) failure to find the same in several populations of Centrosaurus apertus, further supports this position. Sternberg (1927) specifically states that “‘[t]here have been no bones of other individuals used in the mounts”, so the differ- ences cannot be ascribed to the incorporation of mate- rial from a smaller individual into the mount. It seems that the humeri of CMN 2245 are simply unusually small and gracile, although the reason for this is unclear. The reasons for the disparity in the dimensions of the Dimensions (mm) Disparity CMN 2245 CMN 2280 (%) 95 72 24 225 Dil, IES 44 76 42 [56] 78 28 55 86 36 216 275 21.5 sternal plates and of the presacral vertebrae are likewise uncertain in the absence of more material. The appar- ently longer and more erect neural spines and transverse processes of CMN 2280 coincide with observations made by Tereshchenko (2001) of protoceratopsids, in which the spinous processes of the vertebrae were reputedly shown to be longer and more erect in males than in females. If this apparent dimorphism is sexual, however, it would imply the “robust” specimen (CMN 2280) was male rather than female (contra Sternberg 1927). This type of “normal” sexual dimorphism, in which the male is more massive than the female, is typ- ical of most reptiles (Fitch 1981; Shine 1989; Brochu 2002). Thus, undoubted differences (albeit fewer than orig- inally implied by Sternberg) between the two speci- mens do exist. The marked difference in the size and shape of the humerus may simply be anomalous, unless we accept the unlikely hypothesis that of the twelve Chasmosaurus humeri measured by Chinnery (2001), eleven are of one sex and only one is of the other. Other differences, such as the specific shape of the sternal plates and the relative size and orientation of the spinous processes of the presacral vertebrae, might represent sexual dimorphism or intraspecific variation. It is also possible that all of the postcranial discrep- ancies noted here are functionally related, reflecting a developmental compensation of the extrinsic muscles of the forelimb in response to the atrophied humeri of CMN 2245. However, in the absence of a statisti- cally significant sample, it is impossible to resolve this question. Whatever the significance of the variation, there seems little choice but to reject the hypothesis of sexual dimorphism. Two-Species Hypothesis It is possible that the few differences documented here within the postcranial skeletons of CMN 2245 and CMN 2280 may simply reflect that these two speci- mens represent separate species. In a review of the systematics of the genus Chasmosaurus (Godfrey and Holmes 1995), two species were diagnosed based on differences in parietosquamosal frill structure. C. belli was defined as possessing a “parietal frill with nearly straight transverse posterior bars, each bearing one large 2006 vertebra (mm) ee A = SBSBSSBSERREFKRKE Vertebra T8 Tg T10 mT T12 —t— CMN 2245 —@- CMN 2280 Vertebra —*— CMN 2245 —®- CMN 2280 MALLON AND HOLMES: SEXUAL DIMORPHISM IN CHASMOSAURINE CERATOPSID 409 70 Ls 8) A A hee é / / e “~ be / —. ya Poe » tn | \™* Oe = 3 : . / low © ; 2 en ae i] S8BSBSFFFFRF REF RFP SES Vertebra —e- CMN 2245 —@- CN 2280 120 - 110 = aN gp ete 3E \ tA s 2 § ee = 2" VA se J \_~< 2s 70 ce a ——- 50 SS8SSBSFFFFFRFERFSSES FiGuRE 5. Graphical depictions of vertebral disparity between CMN 2245 and CMN 2280. A, Differences between maxi- mum heights of vertebrae. The anterior thoracic vertebrae of CMN 2280 are consistently taller than those of CMN 2245. B, Differences between angles of neural spines. In the anterior thoracic region, the neural spines of CMN 2245 are angled further posteriorly than in CMN 2280. C, Differences between angles formed by transverse processes. The transverse processes of the cervical and anterior thoracic vertebrae of CMN 2280 are more erect than those of CMN 2245. D, Differences between transverse widths of centra. The cervical and anterior thoracic centra of CMN 2280 are consistently wider transversely than those of CMN 2245. Abbreviations: C, cervical vertebra; T, tho- racic vertebra. triangular epoccipital on its posterolateral corner; other parietal epoccipitals variable in number and degree of coossification with the parietal, but always much small- er. The lateral bar of the parietal completely encloses the parietal fenestra.” C. russelli was characterized as having a frill in which the posterior margin is “broadly arched on either side of [the] median emargination. Each side bears three low triangular, roughly equal- sized epoccipitals. The lateral ramus of the parietal is reduced and does not completely encircle the fenes- tra in all but one specimen, permitting the squamosal to form a part of its lateral border.’ Under these amended diagnoses, CMN 2245 was retained within the hypo- digm of C. belli, but CMN 2280 was reassigned to C. russelli. This distinction was challenged by Lehman (1998), who argued that the discrete characters used to distinguish C. belli from C. russelli actually represent end-members of a gradational spectrum of frill mor- phologies. In support of his argument, Lehman (1998: figure 9) figured seven skulls attributed to C. belli and C. russelli, illustrating the range of frill morpholo- gies he observed. However, the lateral parietal bars of AMNH 5402 used by Lehman are not illustrated as being complete as they are in the original specimen, which creates the false impression that the frills exhibit graded variation. Correcting for this splits the series into two discrete groupings of frill morphologies (Fig- ure 6), the contents of which agree with the original hypodigms of Godfrey and Holmes (1995). Further support for the distinction between C. belli and C. russelli stems from their stratigraphic segrega- tion within the Dinosaur Park Formation (Figure 7). This observation was made previously by Godfrey and Holmes (1995) and Holmes et al. (2001), and more recent quarry data support this claim, with C. russelli confined to the lower part of the formation and C. belli located much higher in the section (Ryan and Evans 2005). These independent lines of evidence strongly suggest that the variation between CMN 2245 and CMN 2280 is interspecific in nature, negating Stern- berg’s case for sexual dimorphism. Stratigraphic sep- aration has also been noted recently among the suppos- edly sexually dimorphic lambeosaurine hadrosaurids of the Dinosaur Park Formation (Evans et al. 2006), 410 viii THE CANADIAN FIELD-NATURALIST Vol. 120 q FiGureE 6. Parietosquamosal frills of (A) Chasmosaurus belli and (B) C. russelli in dorsal view. i, CMN 491 (holotype of C. belli); ii, YPM 2016; iii, CMN 2245 (paratype of C. belli); iv, AMNH 5402; v, ROM 843; vi, CMN 8803 (paratype of C. russelli); vii, CMN 2280; viii, RTMP 83.25.1; ix, AMNH 5656. Frills not to scale. suggesting a need for more critical consideration of sexual dimorphism in the fossil record. Conclusion After a careful review of CMN 2245 and CMN 2280, it seems that the postcranium of neither specimen is quite as complete or as well preserved as originally described by Sternberg (1927). Consequently, the case for dimorphism (sexual or otherwise) is weaker than previously thought. Most of the variation exhibited by the shared elements is attributable either to poor recon- struction, postmortem distortion, or individual variation. Exceptional cases involve the humeri, sternal plates, and presacral vertebrae. While the disparity in the shape of the humerus is indeed genuine, the same dimorphism is not seen in other specimens of Chasmosaurus, so this feature is likely anomalous rather than dimorphic. The sternal plates of CMN 2245 and CMN 2280 differ in the lengths of their posterolateral processes, and the presacral ver- 2006 tebrae differ in the lengths and orientations of their spin- ous processes. However, analysis of additional Chas- mosaurus material — preferably derived from some yet undiscovered monodominant bonebed (Eberth and Getty 2005) — would be necessary to determine the significance of this variation with regards to sexual dimorphism. The results presented here are therefore only preliminary, pending further investigation of indi- vidual variation in the ceratopsian postcranial skeleton. Finally, previous detailed reviews of the frill mor- phology of either specimen seem to suggest that CMN 2245 and CMN 2280 may, in fact, belong to separate species (C. belli and C. russelli, respectively), as first put forth by Godfrey and Holmes (1995). This view is further supported by the stratigraphic segregation of the two specimens within the Dinosaur Park Formation of Alberta. Whether the postcranial differences iden- tified here represent interspecific distinctions remains to be demonstrated. Therefore, although there do seem to be a few legitimate differences between the two specimens, the weight of the evidence so far accumu- lated does not support sexual dimorphism as Stern- berg (1927) first suggested it, and there remains little choice but to reject this hypothesis. In the future, sex- ual dimorphism should be attributed to fossil species only when statistical significance and stratigraphic control have been demonstrated. Claims made in the absence of such evidence should be regarded with scepticism. Acknowledgments We would like to thank Gilles Danis, Margaret Feuerstack and Kieran Shepherd for offering access to specimens in their care. Philip Currie and Hans Larsson provided valuable resources and discussion. Andrew Farke, Michael Ryan, and two anonymous reviewers provided constructive criticism on earlier versions of this manuscript. This research stems from a B.Sc. Hon- ours thesis written by JCM under the supervision of RH. Institutional Abbreviations AMNH, American Museum of Natural History, New York, New York; CMN, Canadian Museum of Nature, Ottawa, Ontario; RTMP, Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta; YPM, Yale Peabody Museum, New Haven, Connecticut. Documents Cited (marked * in text) Currie, P. J., and E. B. 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Shine, R. 1989. Ecological causes for the evolution of sexu- al dimorphism: a review of the evidence. Quarterly Review of Biology 64: 419-461. Sternberg, C. H. 1917. Hunting Dinosaurs in the Bad Lands of the Red Deer River, Alberta, Canada. Self-published, Lawrence, Kansas. 232 pages. Sternberg, C. M. 1927. Horned dinosaur group in the Na- tional Museum of Canada. Canadian Field-Naturalist 41: 67-73. Tereshchenko, V. S. 2001. Sexual dimorphism in the post- cranial skeleton of protoceratopsids (Neoceratopsia, Pro- toceratopsidae) from Mongolia. Palaeontological Journal 35: 415-425. Zaaf, A., and R. Van Damme. 2001. Limb proportions in climbing and ground-dwelling geckos (Lepidosauria, Gek- konidae): a phylogenetically informed analysis. Zoomor- phology 121: 45-53. Received 1 November 2005 Accepted 5 June 2007 Use of Saltmarsh by Dragonflies (Odonata) in the Baie des Chaleurs Region of Quebec and New Brunswick in Late Summer and Autumn PAUL M. CATLING!, RAYMOND HUTCHINSON?, and PAUL M. BRUNELLE? '170 Sanford Avenue, Ottawa, Ontario K2C OE9 Canada; e-mail: catlinp@agr.ge.ca ? 12 Ch. de la Savane, Appartment 12, Gatineau, Québec J8T 1P7 Canada; e-mail: Raymond.hutchinson@sympatico.ca 44 Hilltop Terrace, Dartmouth, Nova Scotia B2Y 3T1 Canada; e-mail: pmb@ns.sympatico.ca Catling, Paul M., Raymond Hutchinson, and Paul M. Brunelle. 2006. Use of saltmarsh by dragonflies (Odonata) in the Baie des Chaleurs region of Quebec and New Brunswick in late summer and autumn. Canadian Field-Naturalist |20(4): 413-420. During late summer and autumn, in the Baie des Chaleurs region of Quebec, 18 species of adult dragonflies were recorded during one or more visits of at least 2 hours each to 14 saltmarshes. Three species, Aeshna canadensis, Sympetrum danae and S. internum, were present in more than half of the sites. The most abundant species was S. internum with over 100 seen at some locations. Adults of several species, including Aeshna canadensis, A. umbrosa, Enallagma civile, E. hageni, Ischnura verticalis, Lestes disjunctus, Libellula quadrimaculata, Sympetrum danae, S. internum and S. obtrusum, occurred in relatively high frequencies in both Baie des Chaleurs saltmarshes and in those elsewhere in Acadia. Within Baie des Chaleurs observations of emergence and/or presence of larvae, as well as regional abundance, were recorded for Aeshna canadensis, A. umbrosa, Ischnura verticalis, Sympetrum costiferum, S. internum and S. vicinum. Oviposition in saltmarsh pools was recorded for Aeshna canadensis, Enallagma civile, E, hageni, Ischnura verticalis, Lestes congener and Sympetrum danae. The saltmarsh dragon- fly fauna of Baie des Chaleurs is significantly different from that of the rest of Acadia based on frequencies predicted from the latter region. To a large extent this difference is a result of significantly increased use of saltmarsh habitat by adults of six species including Lestes congener, Sympetrum danae, Aeshna canadensis, Sympetrum costiferum, Lestes disjunctus, and Sympetrum internum (in order of decreasing significance) in Baie des Chaleurs in comparison with saltmarshes elsewhere in Acadia. Local amelioration of salty conditions in certain saltmarshes, superimposed on regional amelioration as a result of protection from storms and saltwater dilution in the Baie des Chaleurs estuary, may contribute to an environment where adaptation can occur or where already tolerant species can exist. Dragonflies use saltmarsh habitat on the northeast coast of North America more extensively than is currently documented. Key Words: Odonata, dragonflies, saltmarsh, Baie des Chaleurs, Quebec, New Brunswick. Dragonflies are generally not highlighted as a group of insects that inhabits saltmarshes (e.g. Cheng 1976). Foster and Treherne (1976) outline the difficulty of characterizing marine insects because many species wander in from adjacent habitats. They define marine insects as having a “habitual requirement for some part of the saltmarsh environment at some stage of their life cycle.” They show Odonata species comprising 3% of saltmarsh insect fauna which is dominated by Diptera, Coleoptera and Hemiptera which collectively make up 75% of the species. Corbet (1999, page194) notes the only truly marine dragonfly is Erythrodiplax ber- enice, but that there is also a “heterogenous assemblage of Zygoptera and Anisoptera that occupy brackish waters, usually of relatively low salinity compared with seawater.” Dragonflies have been reported to prey on saltmarsh tabanid flies and mosquitos, both groups of aggressive biting insects that interfere with human activities. As a consequence, questions periodically arise as to the extent of use of saltmarsh by dragonflies. Very little information on this subject is available for the maritime tegion of eastern Canada. Hutchinson and Bélanger (1999, 2004) have reported on adults of Sympetrum danae and Enallagma civile (respectively) using salt- marsh in the Baie des Chaleurs region. These obser- vations suggest that use of saltmarsh by dragonflies may be more extensive than suspected. Preliminary observations in the Baie des Chaleurs region of Quebec and New Brunswick supported this view and sug- gested the use of saltmarsh by widespread species of dragonflies. The objective of the work reported here was to sum- marize information on the late summer and fall use of saltmarsh by dragonflies in this region and to provide a basis for further analysis of evolutionary and eco- logical aspects of occurrence. Methods The study area — Baie des Chaleurs Baie des Chaleurs is bounded on the north by the Gaspé Peninsula and on the south by northern New Brunswick (Figure |). Flowing into it at its far western end is the Restigouche River, famous for fishing of Atlantic Salmon, Salmo salar. The tides are approxi- mately 2 m high in the western basin and seawater is encountered in the estuary where it expands at Camp- bellton, New Brunswick. More information on Baie des 413 414 THE CANADIAN FIELD-NATURALIST Vol. 120 0 100 Kilometers | a eT - : FL id a-0eeer -| & Baie des Chaleurs locations @ Locations outside Baie des Chaleurs 163 61 FicureE |. Region of Acadia showing saltmarshes visited in Baie des Chaleurs (triangles) and elsewhere (dots). Chaleurs is available from Gauthier (2005), but there is apparently little information other than this recent reference (National Wetlands Working Group 1988). Data gathering Summarized below for the Baie des Chaleurs region are our late summer and fall observations of dragon- flies in saltmarshes as well as some from the Atlantic Dragonfly Inventory Project (ADIP 2007°) for the same region. These observations cover the period from 22 July to 1 October over the past decade. The period of observation at any of these sites was at least 1-2 hours on any particular date. A list of observations at each site is followed by an annotated list of species and additional summary comments. In discussing the ob- servations, the term “frequent” indicates a species seen often (e.g. at many sites) whereas “abundant” refers to a relatively large number of individuals at one or more Sites. All of the sites noted were dominated by plant species considered to be obligate halophytes, including two species of Spartina (S. alterniflora and S. patens), and Ruppia maritima was present in pools at most sites. The presence of these and other obligate halo- phytes indicates clearly that the sites were saltmarsh- es subject to periodic inflow of saltwater. Voucher specimens for the records of dragonflies for Baie des Chaleurs and elsewhere in Acadia are at the Canadi- an National Collection at the Agriculture and Agri-food (AAFC) Central Experimental Farm in Ottawa (Cana- dian National Collection CNC), and in the insect col- lections of the Nova Scotia Museum (NSMNH), the New Brunswick Museum (NEM) and the personal collection of P. M. Brunelle. Comparisons To explore differences between dragonfly occur- rence in Baie des Chaleurs and elsewhere on the North Atlantic coast, the number of sites recorded for species of dragonflies in Baie des Chaleurs was com- pared with similar data (ADIP 2007*) for the rest of Acadia (Canadian Maritime Provinces and the state of Maine) outside of Baie des Chaleurs. Only records for the period fromm 22 July until the end of October are included in the comparison to represent late sum- mer and autumn use of saltmarshes, and all records are for the past decade. Number of sites where a species was present was expressed as a percentage of total salt- marsh sites visited. The extent to which the Baie des Chaleurs dragonfly fauna is different was explored us- 2006 CATLING, HUTCHINSON, and BRUNELLE: USE OF SALTMARSH BY DRAGONFLIES 415 none 2 Bcickish pool in CS at St. Omer (site 4) wbere two re of Lay Totee were seen ovipositing in the foreground bed of floating algae and Ruppia maritima. The pool is surrounded by Scirpus spp. , Juncus gerardii and Spartina alterniflora. Eight male Sympetrum danae patrolled the edge of this saltwater during high spring and fall tides. ing a Chi-square test and percentage frequency else- where in Acadia to predict percentage frequency in Baie des Chaleurs. The test also allows the comparison of Chi-square values for individual species, but consider- ing the sample size, the values are most useful to iden- tify species contributing substantially to differences be- tween Baie des Chaleurs and elsewhere, rather than to indicate a specific level of significance. Results Sites and species present For each site, observation dates are given followed by species in alphabetical order and number of indi- viduals seen in brackets. The total for each site and the size and nature of the area surveyed are also given. In cases where we are not the observers, the observers are listed at the end. A total of 18 species of Odonata was encountered in 14 saltmarshes in the Baie des Chaleurs in late summer and autumn. (1) Bathurst (Queen Elizabeth and Youghall streets), New Brunswick (47.6648°N, 65.6328°W), 14 August 2005: Aeshna canadensis (25), Enallagma hageni (40), Libellula quadrimaculata (2), Somatochlora elongata (4), Sympetrum 25 m? pool, which is inundated by costiferum (5), S. internum (25), S. vicinum (4). Total: 7 species. Size: The site included a brackish % acre pool. (2) 4 km E of Pointe-a-la-Croix, Quebec (48.0520°N, 66.6528°W), 15 August 2005: Aeshna canadensis (30), Lestes congener (4), Lestes disjunctus (2), Sympetrum costiferum (1), S. danae (9), S. internum (200), S. obtrusum (2). Total: 7 species. Size: 50 acres traversed, including creeks, pools and a river. (3) Pointe-a-la-Garde, Quebec (48.0773°N, 66.5425°W), 15 August 2005: Aeshna canadensis (50), Enallagma hageni (43). Lestes congener (10), L. disjunctus (5), Sympetrum costiferum (1), S. danae (15), S. internum (150). Total: 7 species. Size: 10 acres traversed with many pools and creeks. (4) Saint-Omer, Quebec (48.1087°N, 66.2514°W), 15 August 2005: Aeshna canadensis (18), Lestes congener (5), L. dis- junctus (1), Enallagma hageni (1), Sympetrum danae (15). Total: 5 species. Size: 3 acres with pools beside creek (Fig- ure 2). (5) Charlo, New Brunswick (48.0182°N, 66.3666°W), 16 August 2005: Aeshna canadensis (1), Lestes congener (3). Sympetrum danae (1), Sympetrum internum (2). Total: 4 spe- cies. Size: 4 acre with one small pool. (6) Nash Creek, New Brunswick (47.9229°N, 66.0840°W), 16 August 2005: Aeshna canadensis (6), Sympetrum danae 416 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Locations of saltmarshes visited in Baie des Chaleurs and elsewhere in Acadia over the past ten years within the period 22 July to 1 October. The order is increasing latitude from left to right and top to bottom. Baie des Chaleurs 47.6648, 65.6328 48.0520, 66.6528 48.0182, 66.3666 47.9229, 66.0840 48.1000, 66.1348 48.0500, 65.4833 47.64, 65.62 47.65, 65.61 Acadia outside Baie des Chaleurs 43.5694, 65.4139 44.15, 68.25 43.5702, 65.3747 44.2518, 68.3398 44.3889, 68.7667 44.4417, 68.8833 44.479, 68.013 44.5556, 67.8722 44.7194, 67.45 44.72, 67.41 45.1699, 66.3135 45.19, 66.32 46.1635, 63.8258 46.2083, 60.6333 46.6532, 64.0748 46.732, 63.9946 48.0773, 66.5425 47.9166, 66.0241 48.1333, 66.3166 44.0667, 64.6417 44.2535, 68.3406 44.45, 67.9 44.605, 67.48 44.8111, 66.9806 45.7321, 64.6765 46.4, 63.86 47.1667, 65.0493 48.1087, 66.2514 47.0657, 65.0750 47.7846, 65.0841 44.1483, 64.5106 44.35, 68.35 44.4587, 64.3215 44.6194, 67.7472 45.107, 61.7266 46.06, 62.95 46.5403, 63.56 47.1721, 65.0372 (1), S. internum (4), S. obtrusum (1). Total: 4 species. Size: 5 acres with pools and creeks. (7) Jacquet River Marsh, New Brunswick (47.9166°N, 66.0241°W), 16 August 2005: Aeshna canadensis (30 adults 1 larva probably this species), Aeshna umbrosa (1 larva prob- ably this species), Lestes congener (50), L. disjunctus (1), Ischnura verticalis (200 adults, 4 larvae), Sympetrum danae (1), S. internum (5). Total: 7 species. Size: traversed 10 acres with pools and creeks. (8) North of Eel River, New Brunswick (48.0657°N, 66.4000°W), 16 August 2005: Aeshna canadensis (22), A. interrupta (3), Enallagma civile (26), Enallagma hageni (1), E. ebrium (1), Ischnura verticalis (100 ), Lestes congener (10 ), L. disjunctus (4 ), L. forcipatus (2), Sympetrum internum (70), S. obtrusum (40). Total: 11 species. Size: at least SO acres traversed with numerous creeks and pools, all brackish, at least periodically. (9) Carleton, Quebec (48.1000°N, 66.1348°W), 29 Septem- ber—1 October 2004: Aeshna umbrosa (5+), Enallagma civile (3), Sympetrum danae (7), S. internum (1), S. vicinum (1); S. costiferum (1); 10 September 1999, Enallagma civile (2), Sympetrum danae (4); 4 September 1995, Sympetrum danae (3), Sympetrum sp. (1 reddish specimen). Total: 6 species. Size: pools within an area of 3 acres. (10) Bonaventure, Quebec (48.0500°N, 65.4833°W), 3 Sep- tember 1995: Aeshna sp. (2), Pantala flavescens (1), Sym- petrum costiferum (1), S. danae (10). Total: 4 species. Size: 3 acres. (11) Nouvelle (saltmarsh of the Nouvelle River Basin), Que- bec (48.1333°N, 66.3166°W), 4 September 1995: Lestes con- gener (1), Sympetrum danae (1), Sympetrum sp. (1, reddish). Total 3 species. (12) Caraquet Bay, New Brunswick (47.7846°N, 65.0841°W) saltmarsh, 12 August 2002: Sympetrum danae (3), Sympetrum internum (1), Sympetrum obtrusum (1). Observer: Dwayne L. Sabine. Total: 3 species. (13) Daly Point Marsh, Bathurst Harbour, New Brunswick (47.64°N, 65.62°W), 12 August 1993: Sympetrum danae (1). Observer: Reginald P. Webster. 21 July 1999, Libellula quadrimaculata (2), Aeshna canadensis (2). Total 3 species. (14) Carron Point Marsh, Bathurst Harbour, New Brunswick (47.65°N, 65.61°W), 22 July 1999: Aeshna canadensis (1), Libellula quadrimaculata, Sympetrum danae (2), Sympetrum obtrusum (1). Total: 4 species. Annotated List of Species (alphabetical order) Aeshna canadensis: at 10 sites and frequent at some, appar- ently uses saltmarsh regularly as an adult hunting on salty pools of the outer marshes which are influenced by tides reg- ularly, as well as inner brackish pools; seen to oviposit in saltmarsh pools. A larva 20 mm long that was probably (but not definitely, due to difficulty of identifying young larvae) referable to this species was retrieved from a periodically brackish pool dominated by Ruppia maritima (site 7). Aeshna interrupta lineata: only one at one site but of interest because this subspecies exists at low frequencies in the Atlantic provinces and is largely confined to the Gulf of St. Lawrence area. The specimen was referred to subspecies /ineata on the basis of uninterrupted lateral stripes. This species is reported to tolerate saline waters (Dunkle 2000). Aeshna umbrosa: only from two sites but five adults seen patrolling saltmarsh ponds; at the other site two young larvae probably referable to this species (21 and 23 mm long) were found in a brackish pool dominated by Ruppia maritima (site 7) — due to their small size, identification is tentative; mature larvae of this species were found in water-filled pock- ets within the storm splash zone of a sea cliff near Port-au- Saumon, Quebec (Hutchinson 1999); and D. Sabine (personal communication) found larvae in a small depression pool in a seaside boulder within the tidal splash zone on the Fundy shore of New Brunswick. Enallagma civile: two sites; twenty-six captured (most re- leased), many paired, were flying over a periodically brackish creek in saltmarsh with mostly Ruppia maritima, also Zan- nichellia palustris, Potamogeton pectinatus, lined by Spartina alterniflora (site 8). Enallagma civile was first reported from this region by Hutchinson and Bélanger (2004), and their re- port was from a saltmarsh habitat at Carleton (site 9), where they observed oviposition behaviour (but did not actually see eggs). Osborn (1906) reported a tolerance of brackish water. Enallagma ebrium: only one at one site, so likely vagrant. Enallagma hageni: At four sites and abundant at two; 40 seen and several ovipositing on Ruppia maritima around a brackish, 4 acre pool with dense beds of Ruppia maritima surrounded by Spartina alterniflora, S. patens, S. pectinata, and Carex 2006 CATLING, HUTCHINSON, and BRUNELLE: USE OF SALTMARSH BY DRAGONFLIES 417 TABLE 2. Comparison of species of dragonflies recorded from Acadia outside of Baie des Chaleurs and Baie des Chaleurs based on number (left) and percentage (right) of total saltmarsh sites visited. Acadia includes the Maritime Provinces (Nova Scotia, New Brunswick and Prince Edward Island) of Canada and the state of Maine. Records from Acadia are from the ADIP database (2007). Records from Baie des Chaleurs are those reported here and are also in ADIP 2007. Only records for the period from 22 July until the end of October are included in the comparison to represent late summer and autumn use of saltmarshes. Chi-square values for each species based on comparison of frequency elsewhere in Acadia to predict frequency in Baie des Chaleurs are included to highlight species most different in their occurrence in these two regions in bold. Names are taken from Catling et al. (2005). Species Lestes congener Hagen, Spotted Spreadwing Lestes disjunctus Sélys, Northern Spreadwing Lestes forcipatus Rambur, Sweetflag Spreadwing Lestes inaequalis Walsh, Elegant Spreadwing Lestes rectangularis Say, Slender Spreadwing Lestes unguiculatus Hagen, Lyre-tipped Spreadwing Enallagma aspersum (Hagen), Azure Bluet Enallagma civile (Hagen), Familiar Bluet Enallagma durum (Hagen), Big Bluet Enallagma ebrium (Hagen), Marsh Bluet Enallagma hageni (Walsh), Hagen’s Bluet Ischnura verticalis (Say), Eastern Forktail Aeshna canadensis Walker, Canada Darner Aeshna constricta Say, Lance-tipped Darner Aeshna eremita Scudder, Lake Darner Aeshna interrupta Walker, Variable Darner Aeshna umbrosa umbrosa Walker 1908, Shadow Darner Anax junius (Drury), Common Green Darner Dorocordulia lepida (Hagen in Sélys), Petite Emerald Somatochlora elongata (Scudder), Ski-tipped Emerald Somatochlora kennedyi Walker, Kennedy’s Emerald Celithemis elisa (Hagen), Calico Pennant Erythrodiplax berenice (Drury), Seaside Dragonlet Leucorrhinia frigida Hagen, Frosted Whiteface Libellula luctuosa Burmeister, Widow Skimmer Libellula pulchella Drury, Twelve-spotted Skimmer Libellula quadrimaculata Linnaeus, Four-spotted Skimmer Pantala flavescens (Fabricius), Wandering Glider Plathemis lydia (Drury), Common Whitetail Sympetrum costiferum (Hagen), Saffron-winged Meadowhawk Sympetrum danae (Sulzer), Black Meadowhawk Sympetrum internum Montgomery, Cherry-faced Meadowhawk 1 Sympetrum obtrusum (Hagen), White-faced Meadowhawk Sympetrum semicinctum (Say), Band-winged Meadowhawk Sympetrum vicinum (Hagen), Autumn Meadowhawk Acadia Chaleurs Chi-square n= 32 n=14 | 3.) Te, 500 352.8872 SG Sau 21.4439 | 3.1 | a] 2.6130 l 3.1 - _ 1.5500 4 12.5 - = 10.0160 2 6.2 - - 4.1333 | 3.1 - 1.5500 ht Z29( 0, Dr NAS 4.1125 ] 3.1 - - 1.5500 2 6.2 | Tal 0.0920 LOW S12 4 28.6 0.2125 6 18.7 2 43 0.9084 hin eels 10 =—71.4 98.2049 3 9.3 - - 6.9759 2 6.3 _ = 4.2223 Sy ISG" ] Tel 3.8409 42-5 2 143 0.1963 1 3.1 - 1.5500 2 6.2 - - 4.1333 3 9.3 Tk 0.3815 1 3.1 = 1.5500 1 3.1 = - 1.5500 8 25.0 - - 22.2420 1 Bel -- - 1.5500 1 Bel - 1.5500 2 6.2 _ - 4.1333 9 28.1 2) yAlee! 1.4495 3 9.3 l Ted 0.3815 2 6.2 - - 4.1333 3 9.3 By SY 55.8876 5) ISHS [2a S57 261.2840 22 SieD 9 64.3 17.4604 OF 28el Ss) SPH 1.8149 Sp els!6 - 13.0139 2) 6.2 — - 4.1333 paleacea, flooded by high tides in fall according to local res- idents (site 1); 43 around pools and creeks with Ruppia mar- itima, Zannichellia palustris, Potamogeton pectinatus and Enteromorpha sp. (site 3). Ischnura verticalis: abundant at each of two locations; more than 200 seen around pools with Potamogeton pectinatus and Ruppia maritima with Spartina pectinata fringe, some ovi- positing, and at the the same location four larvae 17-21 mm long referable to this species were collected from the pools (site 7); more than 100 were seen around spring-fed pools sur- rounded by Spartina pectinata, Spartina alterniflora, Spartina patens and species of Scirpus (site 8). In both cases the pools were fed by freshwater springs but were definitely subject to saltwater during high tides. Lestes congener: at seven sites and at least 82 individuals overall; given the number of sites and the number of individ- uals this species may be a resident of saltmarsh; several pairs seen ovipositing on Juncus balticus and Juncus gerardti around brackish pools with Potamogeton pectinatus and Ruppia mari- tima (site 7). Lestes disjunctus: at five sites; but not as abundant as L. congener. Lestes forcipatus: only one at one location. Libellula quadrimaculata: at three sites: adults patrolled a brackish pool at site 1; said to tolerate saline waters (Dunkle 2000). Pantala flavescens: only one at one location and probably a migrant. Somatochlora elongata: only found at one location, but four seen over pool with Ruppia maritima surrounded by Spartina alterniflora, S. patens, S. pectinata, and Carex paleacea, flood- 418 ed by high tides in fall according to local residents (site 1). This species has also been reported from four other salt- marshes in Acadia (Table 2). Sympetrum danae: at 12 sites, and abundant on the north side of the bay, patrolling saltmarsh pools very much like Ery- throdiplax berenice does in saltmarshes further south on the Atlantic coast. Two females were seen ovipositing in a pool 25 m* with a thick bed of Ruppia maritima surrounded by Scirpus spp., Juncus gerardii and Spartina alterniflora (site 4, Figure 2). The pool was said by local people to be flooded by high tides in spring and fall. Ovipositing behaviour was also observed in saltmarsh ponds at Carleton but eggs were not actually seen. The species appears to be less frequent in the saltmarshes on the south side of the bay. Hutchinson and Bélanger (2004) have previously drawn attention to the uti- lization of saltmarsh by adults of this species and suggested the possibility that larvae may mature in brackish pools. High salinity of spring and fall high tides may be avoided by the resistant egg stage that overwinters. Larval development may be relatively rapid (Corbet 1999) and allow use of saltmarsh pools during periods of minimal tidal impact. Although the larval habitat of this species is widely reported in both Europe and North America as oligotrophic pools and small lakes on bogs and heathlands, it also inhabits very alkaline fens in east- ern and northern Canada. Consequently saltmarsh pools would not be so remarkable a habitat. Interestingly in Ireland, males are said to be non-territorial (Nelson and Thompson 2000’), contrary to some of our observations on saltmarsh pools. In some places within its holarctic range, S. danae is reported to migrate (e.g. Corbet 1999, page 420). Ability to utilize brackish habitats may be beneficial to migrants over water in the mar- itime regions where landfall may have only brackish habitat. Sympetrum internum: at nine sites and abundant at some. One emerging from a pool surrounded by Spartina pectina- ta in upper marsh (site 2) and tenerals in several sites; uses saltmarsh extensively as an adult and the upper marsh at least to some extent as larval habitat. Sympetrum costiferum: at five sites; two emerging from brack- ish 4 acre pool with dense beds of Ruppia maritima surround- ed by Spartina alterniflora, S. patens, S. pectinata and Carex paleacea, flooded by high tides in fall according to local resi- dents (site 1); one emerging from sandy bottom creek with Ruppia maritima and Enteromorpha sp. (site 3). Apparently uses Saltmarsh habitats for development but in low numbers, possibly due to limited amount of sandy larval habitat in most marshes. According to Walker and Corbet (1975), S. costiferum appears to be more tolerant of saline waters than other spe- cies of Sympetrum, but they are not clearly referring to sodium salts. Dunkle (2000) also refers to tolerance of saline waters. Sympetrum obtrusum: at five sites but no strong evidence of use of saltmarsh as larval habitat by this widespread and gen- erally abundant species. Sympetrum vicinum: at two locations but at one of these two were emerging from brackish 4 acre pool with dense beds of Ruppia maritima surrounded by Spartina alterniflora, S. patens and S. pectinata, and Carex paleacea, flooded by high tides in fall according to local residents (site 1). Discussion Comparison with saltmarshes elsewhere in Acadia A total of 35 species has been recorded from saltmarsh- es in Acadia (Table 2). Several occurred in relatively THE CANADIAN FIELD-NATURALIST Vol. 120 high frequencies in both Baie des Chaleurs saltmarshes and saltmarshes elsewhere in Acadia suggesting some degree of salinity tolerance. Included in this category are Aeshna canadensis, A. umbrosa, Enallagma civile, E. hageni, Ischnura verticalis, Lestes disjunctus, Libel- lula quadrimaculata, Sympetrum danae, S. internum and S. obtrusum. With a Chi-square value of 912.707, the saltmarsh dragonfly fauna of Baie des Chaleurs is significantly (well below the 0.005 level) different from that of the rest of Acadia based on frequencies predicted from the latter region. Based on Chi-square values exceed- ing 10.0 (significance below the 0.005 level), Baie des Chaleurs saltmarshes differed from sites elsewhere in having significantly more sites for six generally wide- spread species that occur throughout Acadia (Table 2, Donnelly 2004a, b, c) including Lestes congener, Sym- petrum danae, Aeshna canadensis, Sympetrum costi- ferum, Lestes disjunctus, and Sympetrum internum (in order of decreasing significance). Three species were significantly less frequent in Baie des Chaleurs includ- ing Erythrodiplax berenice, Sympetrum semicinctum, and Lestes rectangularis all of which do not extend as far north as Baie des Chaleurs (Donnelly 2004b, c). Erythrodiplax berenice for example does not occur north of 45°N, whereas Baie des Chaleurs is situated at 48°N. General Observations With 35 species of adult dragonflies recorded in salt- marshes in Acadia and 18 in Baie des Chaleurs, and in both cases, approximately half of them occurring in more than 15% of sites, substantial use by dragonflies is suggested. This is further supported by records of emergence, presence of larvae, ovipositing behaviour and relatively large numbers of individuals. In a number of saltmarsh habitats further south on the coast of New England, the only abundant dragon- fly species that we have observed using the saltmarsh habitat in late summer is the saltmarsh specialist, Ery- throdiplax berenice. It is often present in large numbers (to 30 per acre), the males patrolling pools, the females along the marsh edges but pairs frequent over the marsh in early to mid-morning. Other Odonatists have also noted that in many saltmarshes the only dragonfly present is E. berenice (D. Paulson, personal commu- nication). Other species fly over the marshes includ- ing the migratory Anax junius, Tramea spp., and Pan- tala spp. All of these may hunt over the marsh but seem much more often to hunt and congregate in open shrub- by areas around the marsh edge. We have seen species of Enallagma, Lestes and Sympetrum only occasion- ally in these more southern saltmarshes. There may be a trend toward more extensive use of saltmarsh by dragonflies in a northerly direction. Possibly saltmarsh- es further south are less utilized by widespread drag- onfly species because salt may be more concentrated in pools of upper marsh due to higher evaporation. 2006 Baie des Chaleurs The fact that Baie des Chaleurs saltmarshes dif- fered from sites elsewhere in Acadia in having very significantly more sites for six generally widespread species, including Lestes congener, Sympetrum danae, Aeshna canadensis, Sympetrum costiferum, Lestes dis- junctus, and Sympetrum internum (in order of decreas- ing significance), suggests either that there has been some adaptation to saltmarsh conditions by these wide- ranging species in this region, or that saltmarshes in this region are more readily colonized as a result of unique characteristics, or both. Baie des Chaleurs is home to an endemic and endangered saltmarsh butter- fly, the Maritime Ringlet (Coenonypha tullia nipisiq- uit, McDunnough 1939). Recent studies have sug- gested that this butterfly has evolved physiological adaptation to saline wetland conditions including tol- erance of tidewater submergence for up to 24 hours (Webster 1998; Sei 2004). The endemic but somewhat more widespread Maritime Copper Butterfly (Lycae- na dospassosi McDunnough) is also confined to the saltmarshes of northern New Brunswick and Gaspé (Thomas 1980). Unique charaterisitics of Baie des Chaleurs salt- marshes may be widespread or local, or both. Parts of saltmarsh are prohibitively salty for non-adapted spe- cies only during high tides in the spring and fall, and even then saltwater may be somewhat diluted due to the estuary effect. There are distinctive east and west basins and in the west basin, where most of our obser- vations were made and where the Maritime Ringlet occurs, the tides are higher and salt concentrations lower than in the east basin (Gauthier 2005). It has been suggested that saltmarshes can be divided into two kinds, estuarine such as Baie des Chaleurs, and coastal (National Wetlands Working Group 1988). Al- though the difference between the two types is not perfectly clear, conditions of inundation and salt con- centrations in coastal saltmarsh would be more severe. Saltmarshes have periodic inundation in common, but the timing and duration may vary as well as the salt concentrations. Furthermore, occasional lack of ice development in the bay may result in severe impact of high tide associated with winter storms on some saltmarshes, while others may be protected by barrier beaches (Webster 1994", 1998). Local amelioration of salty conditions in certain saltmarshes, superimposed on regional amelioration, may provide an environ- ment where tolerant species can exist and adaptation can occur. Tolerance of salt Although the places where dragonflies were record- ed in Baie des Chaleurs saltmarshes may not have been consistently high in salt concentration, they were definitely brackish, were part of a tidal salt marsh sub- ject to periodic inundation with salt water and subject to salt to the extent of excluding non-halophytic flora. Species of dragonflies differ in their ability to cope with CATLING, HUTCHINSON, and BRUNELLE: USE OF SALTMARSH BY DRAGONFLIES 419 salt (Cannings and Cannings 1987). Of the 18 species found in Baie des Chaleurs, Aeshna interrupta, Libel- lula quadrimaculata, Sympetrum danae, and S. costi- ferum are reported by Dunkle (2000) to tolerate saline water. However, Walker (1953, 1958), Walker and Cor- bet (1975), Pilon and Lagacé (1998) do not report any of the species listed here as occurring in saltmarsh. Observations in Baie des Chaleurs suggest that drag- onflies occupy saltmarshes to a greater extent than is currently documented. Ecological and evolutionary as- pects, including extent to which local adaptation has occurred, require more study, particularly directed to- ward larval occurrences and comparisons with other habitats in the surrounding landscape. Acknowledgments Brenda Kostiuk and Gilbert Bélanger assisted with field work. Reginald P. Webster and Dwayne L. Sabine contributed information to the Atlantic Dragonfly In- ventory Project, which was used here for comparison with Baie des Chaleurs. Robert Anderson of the Can- adian Museum of Nature provided helpful comments. Documents Cited Atlantic Dragonfly Inventory Program (ADIP). 2007. Data- base maintained by Paul M. Brunelle, Nova Scotia Muse- um, Halifax (searched March 2007). Nelson, B., and R. Thompson. 2000. The natural history of Ireland’s dragonflies. www.habitas.org.uk/dragonflyireland/ 5651d.htm. Webster, R. P. 1994. The life history and ecology of the Mar- itime Ringlet butterfly, Coenonympha inornata nipisiquit McDunnough. Report for the Endangered Species Recoy- ery Fund, World Wildlife Fund Canada and the New Bruns- wick Department of Natural Resources and Energy, Fish and Wildlife Branch. 18 pages Literature Cited Cannings, R. A., and S. G. Cannings. 1987. The Odonata of some saline lakes in British Columbia, Canada: ecolog- ical distribution and zoogeography. Advances in Odona- tology 3: 7-21. Catling, P. M., R. A. Cannings, and P. Brunelle. 2005. An annotated checklist of the Odonata of Canada. Bulletin of American Odonatology 9(1): 1-20. Cheng, L., Editor. 1976. Marine insects. Elsevier Publishing Co., New York. 581 pages. Corbet, P. S. 1999. Dragonflies, behaviour and ecology of the Odonata. Cornell University Press, Ithaca, New York. 829 pages. Donnelly, T. W. 2004a. Distribution of North American Odonata. Part I: Aeshnidae, Petaluridae, Gomphidae, Cor- dulegastridae. Bulletin of American Odonatology 7(4): 61- 90. Donnelly, T. W. 2004b. Distribution of North American Odonata. Part II: Macromiidae, Corduliidae and Libellul- idae. Bulletin of American Odonatology 8(1): 1-32. Donnelly, T. W. 2004c. Distribution of North American Odonata. Part III: Calopterygidae, Lestidae, Coenagrion- idae, Protoneuridae, Platystictidae with data sources and bibliography, parts I-III. Bulletin of American Odonatol- ogy 8(2-3): 33-99. 420 Dunkle, S. 2000. Dragonflies through binoculars. Oxford University Press. 266 pages. Foster, W. A., and J. E. Treherne. 1976. Insects of marine saltmarshes, problems and adaptations. Pages 5-42 in Marine insects. Edited by L. Cheng. Elsevier Publishing Co., New York. Gauthier, B. 2005. L’estuaire de la baie des Chaleurs — Réstigouche. Le Naturaliste canadien 129(2): 51-56. Hutchinson, R. 1999. Aeshna umbrosa larvae found in ex- tremely shallow water. Argia 2(2): 7. Hutchinson, R., and G. Bélanger. 1999. Découverte d’ Enal- lagma civile (Hagen) (Odonata: Coenagrionidae) dans la baie des Chaleurs en Gaspésie (Québec). Fabreries 24(4): 82-84. Hutchinson, R., and G. Bélanger. 2004. Le marais salé comme habitat potentiel pour Sympetrum danae (Sulzer) (Odonata: Libellulidae). Fabreries 29(1): 16. McDunnough, J. 1939. A new Coenonympha race from north- east New Brunswick. Canadian Entomologist 71: 266. National Wetlands Working Group. 1988. Wetlands of Canada. Canadian Wildlife Service, Ecological Land Clas- sification Series 24. 452 pages. Osborn, R. C. 1906. Observations and experiments on drag- onflies in brackish water. American Naturalist 40: 395-399. THE CANADIAN FIELD-NATURALIST Vol. 120 Pilon, J.-G., and Lagacé. 1998. Les odonates du Québec. Entomofaune du Québec. Chicoutimi. 367 pages. Sei, M. 2004. Larval adaptation of the Endangered Maritime Ringlet (Coenonympha tullia nipisiquit McDunnough (Lepidoptera: Nymphalidae) to a saline wetland habitat. Environmental Entomology 33(6): 1535-1540. Thomas, A. W. 1980. New locality records for the salt marsh copper, Epidemia dorcas dospassosi (Lycaenidae). Journal of the Lepidopterists’ Society 34: 315. Walker, E. M. 1953. The Odonata of Canada and Alaska. Volume 1. University Toronto Press, Toronto. 292. Walker, E. M. 1958. The Odonata of Canada and Alaska. Volume 2. University Toronto Press, Toronto. 318 pages. Walker, E. M., and P. S. Corbet. 1975. The Odonata of Cana- da and Alaska. Volume 3. University of Toronto Press. 307 pages. Webster, R. P. 1998. The life history of the maritime ringlet, Coenonympha tullia nipisiquit (Satyridae). Journal of the Lepidopterist’s Society 52(4): 345-355. Received 30 November 2005 Accepted 29 March 2007 First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia G. F. HANKE!?, M. C. E. MCNALL!, and J. RoBERTS* 'Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8W 9W2 Canada 2e-mail ghanke @royalbcmuseum.be.ca; corresponding author ‘Fish and Wildlife — Science and Allocation, Ministry of Environment, Lower Mainland Region, 10470-152 Street, Surrey, British Columbia V3R OY3 Canada Hanke, G. F., M. C. E. McNall, and J. Roberts. 2006. First records of the Yellow Bullhead, Ameiurus natalis, a \oricariid catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia. Canadian Field- Naturalist 120(4): 421-427. In Canada, there are no native catfish west of the continental divide and until recently, the list of extant exotic catfishes in British Columbia only included introduced Black Bullhead (Ameiurus melas) and Brown Bullhead (Ameiurus nebulosus). We report that a single Yellow Bullhead (Ameiurus natalis) was collected from Silvyermere Lake in the Lower Fraser River drainage. This represents the first record of the Yellow Bullhead in western Canada, and its introduction likely was accidental with a shipment of Largemouth Bass (Micropterus salmoides) rather than dispersal from Washington. Warm, eutrophic, weedy habitat in the Fraser Delta provides ample habitat for Yellow Bullheads and other exotic fishes. A Blue-eyed Panaque (Panaque suttonorum), a loricariid catfish found in 1995 in Shawnigan Lake, Vancouver Island, probably represents a single, illegally released aquarium fish, as does a large Silver Pacu (Piaractus cf. P. brachypomus), which was found in Green Lake on Vancouver Island in 2004. Key Words: Yellow Bullhead, Ameiurus natalis, Blue-eyed Panaque, Panaque suttonorum, Silver Pacu, Piaractus sp., exotic introductions, British Columbia. Most populations of native species were established in British Columbia during and after deglaciation, but this slow process of natural colonization and dis- persal pales in comparison to the wave of accidental and intentional species introductions in the province since the arrival of Europeans. Escaped pets, inten- tionally released pets, the illegal and intentional release of exotic game fishes by local fishermen, intentional telease of game fishes by government agencies, con- taminants in poorly sorted shipments of game fish, and northward dispersal of fishes introduced in Washing- ton, now are the principal sources of exotic freshwa- ter fishes in British Columbia. The pathways used to enter the province and the dispersal potential once present, varies with each species. Carl and Guiguet (1957), Carl et al. (1967), Scott and Crossman (1973), Hart (1973), Crossman (1991), Coad (1995), Moyle (2002), and Wydoski and Whitney (2003) detail the exotic fish species (or hybrids) known to have been introduced into British Columbia prior to this report (Table 1). Approximately 7.4% of the exotic fishes in Table 1 are ornamental (including Mos- quitofish [Gambusia affinis] and Fathead Minnows [Pimephales promelas] introduced for mosquito con- trol; Fathead Minnows also may be introduced illegal- ly as live bait, or dispersed into the province via the Peace River drainage [Smith and Lamb 1976]), 24.1% are non-game fishes (e.g., stickleback, Redside Shin- ers [Richardsonius balteatus]), and 68.5% are game/ food fishes and hybrids between game fish species. In many cases, fishes that are native to the province (or their hybrids) are legally stocked in new locations to enhance local fisheries and satisfy anglers (e.g., Wall- eye [Sander vitreus|, Rainbow Trout [Oncorhynchus mykiss], West-slope Cutthroat [O. clarki lewisi], and Pacific salmon [O. gorbuscha, O. nerka, O. keta, O. kisutch, O. tshawytscha}), and in others, game fishes are illegally introduced to new waterbodies by private individuals (e.g., Northern Pike [Esox /ucius], Small- mouth Bass [Micropterus dolomieu]) (McPhail and Carveth 1994). While most fishes released in this pro- vince are game fishes, it is likely that the number of exotic aquarium fishes released in British Columbia will grow with the burgeoning pet trade (see Padilla and Williams 2004). Species Accounts Silver Pacu (Piaractus cf. PR. brachypomus) — Family Characidae (tetras) On 4 June 2004, a large pacu was caught by a Federal Fisheries Officer angling in Green Lake (~49°13'50"N, 124°03'39"W) near Nanaimo (Figure 1). This fish likely is a Silver Pacu (Figure 2), although initially it was thought to be a piranha (Serrasalmus sp./Pygocentrus sp.) and subsequent media reports called the fish a Red-bellied Pacu (a commonly used name in the pet trade). The species identity as used in this report cannot be verified unless the body is located 421 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Fishes introduced British Columbia according to Wydoski and Whitney (2003), Moyle (2002), Coad (1995), Crossman (1991), Hart (1973), Scott and Crossman (1973), Carl et al. (1967), and Carl and Guiguet 1957, and Royal British Columbia Museum (RBCM) collection records; EX = exotic introductions, NA = native populations also in BC, MM = man-made hybrids or hybrids in nature due to stocking of exotics. Family Petromyzontidae Acipenseridae Amiidae Hiodontidae Clupeidae Cyprinidae Catostomidae Ictaluridae Salmonidae Esocidae Gasterosteidae Gadidae Percopsidae Poeciliidae Moronidae Centrarchidae Percidae Cichlidae Species species not mentioned Acipenser transmontanus Amia calva Hiodon alosoides Alosa sapidissima Couesius plumbeus Platygobio gracilis Notropis hudsonius Pimephales promelas Richardsonius balteatus Tinca tinca Cyprinus carpio Carassius auratus Catostomus catostomus Catostomus commersoni Ameiurus melas Ameiurus nebulosus Ictalurus punctatus Oncorhynchus mykiss Oncorhynchus mykiss aguabonita Oncorhynchus clarki lewisi Oncorhynchus clarki lewisi x O. mykiss Oncorhynchus keta Oncorhynchus nerka Oncorhynchus tshawytscha Salvelinus alpinus Salvelinus fontinalis Salvelinus malma Salvelinus fontinalis x S. malma Salvelinus fontinalis x S. confluentus Salvelinus fontinalis x S. namaycush Salvelinus namaycush x S. malma Salvelinus namaycush x S. confluentus Salmo salar Salmo trutta Thymallus arcticus Coregonus clupeaformis Esox lucius Culaea inconstans Gasterosteus aculeatus Lota lota Percopsis omiscomaycus Gambusia affinis Poecilia reticulata (?) Xiphophorus helleri Morone saxatilis Micropterus dolomieu Micropterus salmoides Lepomis macrochirus Lepomis gibbosus Pomoxis nigromaculatus Perca flavescens Sander vitreus Tilapia sp. Common Name White Sturgeon Bowfin Goldeye American Shad Lake Chub Flathead Chub Spottail Shiner Fathead Minnow Redside Shiner Tench Common Carp Goldfish Longnose Sucker White Sucker Black Bullhead Brown Bullhead Channel Catfish Rainbow Trout California Golden Trout West-slope Cutthroat Chum Salmon Sockeye Salmon Chinook Salmon Arctic Charr Brook Trout Dolly Varden Atlantic Salmon Brown Trout Arctic Grayling Lake Whitefish Northern Pike Brook Stickleback Threespine Stickleback Burbot Troutperch Western Mosquitofish Guppy Swordtail Striped Bass Smallmouth Bass Largemouth bass Bluegill Pumpkinseed Black Crappie Yellow Perch Walleye Tilapia Status 2006 and examined in detail. Furthermore, only estimates of size can be taken from images and notes provided by staff of British Columbia, Ministry of Water, Lands, and Air Protection (now Ministry of Environment). The fish measured about 40 cm total length, and 35 cm fork length, and had been feeding on aquatic vegetation based on its gut contents (milfoil was the dominant item in the gut) [M. deLaronde, personal communication, 2005]. Blue-eyed Panaque (Panaque suttonorum) — Family Loricariidae (suck- ermouth armoured catfishes) The Royal British Columbia Museum fish collection contains a single Blue-eyed Panaque (RBCM 996- 190-001) (Figure 3), which was collected in 1995 from a ditch where Royce Road crosses Shawnigan Creek (approximately at 48°40'05", 123°37'32") (Figure 1). The creek leads into the north end of Shawnigan Lake, Vancouver Island. The specimen is 21.8 cm total, and 19.2 cm fork length. This specimen originally was misidentified as Hypostomus plecostomus when it was received in 1996. Yellow Bullhead (Ameiurus natalis) — Family Ictaluridae (North America Freshwater Catfishes) The first Yellow Bullhead found in British Colum- bia (Figure 4) was collected on 21 July 2005, by elec- trofishing the southeast corner of Silvermere Lake (49°10'01"N, 122°24'27"W) (Figures | and 5). The Yellow Bullhead was in stained but clear water about 10 cm deep, and emerged from a clump of submerged, algae-covered terrestrial grass which was approximate- ly 30 cm offshore; the water in the bay was 29.5°C at the surface. The specimen (RBCM 005-079-001) was anaesthetized/euthanized with 2-phenoxy ethanol, prior to fixing in 10% formaldehyde. It is possible that other Yellow Bullheads have been caught in this lake, but were not recognized as different from Black or Brown Bullheads by local anglers. The specimen is 12.9 cm total length and 10.7 cm standard length. The follow- ing features from identification keys in Scott and Cross- man (1973, page 589), Becker (1983, pages 143-145), Moyle (2002, pages 85-86), and Wydoski and Whit- ney (2003, pages 153-154), confirm the identification of the Yellow Bullhead from Silvermere Lake: body lacks spots or blotches; lower jaw does not project be- yond upper jaw; distance between isthmus and lower jaw notch is less than the distance between the lower jaw notch and the tip of the lower jaw (see illustra- tions in Scott and Crossman 1973, page 589); mental barbels yellow-white without melanophores; depressed anal fin overlaps leading edge of caudal fin; and cau- dal fin with round trailing margin. Yellow Bullheads are native to Atlantic and Gulf Slope drainages from New York to Mexico, the St. Lawrence River, the Great Lakes, and the Mississippi River Basin from North Dakota to the Gulf of Mexi- HANKE, MCNALL, AND ROBERTS: BULLHEAD, CATFISH, PACT 423 IN BRITISH COLUMBIA e 4 Shawnigan Lake ae “| 1) Panaque suttonorum 2) Piaractus cf P. brachypomus 3) Ameiurus natalis FiGuRE |. A map showing the general location of the three new records mentioned in this report, relative to southern Vancouver Island and the Fraser River delta; scale bar = 10 km. FIGURE 2. The single Silver Pacu (Piaractus cf. P. brachypo- mus) taken from Green Lake, Vancouver Island, 4 June 2004; surrounding parts of image (stainless-steel table- top) cropped to simplify the illustration. Scale bar = 5 cm. Scom FiGuRE 3. A photograph of the preserved Panaque suttono- rum specimen (RBCM 996-190-001) collected from the Shawnigan Lake area, Vancouver Island; sur- rounding parts of image (dissection tray) cropped to simplify the illustration. Scale bar = 5 cm. co (Scott and Crossman 1973; Lee et al. 1980), and they have been introduced to at least 14 states outside of their native range (Fuller et al. 1999). They also are known from headwaters of the Hudson Bay drainage in Minnesota but have not been found downstream in Manitoba to date (Koel and Peterka 1994; Stewart et al. 2001; Stewart and Watkinson 2004). 424 Discussion Two of the three species documented in this report are not threatening to the ecology of British Columbia, especially since they are tropical and appear to result from the release of single fish rather than groups, which could form breeding populations. However, all three species are unnecessary additions to the provincial ich- thyofauna, and there always is a risk that exotic intro- ductions may carry unknown pathogens which may infect native fishes (e.g., Dove and Ernst 1998; Yama- moto and Tagawa 2000; Font 2003). In British Columbia, a large portion of the north- er waterways are fast-flowing, cool to cold glacier-fed streams and oligotrophic lakes running through mon- tane areas, and as such, are inhospitable to warm water adapted exotic organisms. However, many exotic spe- cies can survive in the southwestern region for a sum- mer or longer, and fortunately, cold meltwater from higher elevation may limit their long-term survival and dispersal. In some regions (e.g., the Okanagan, the Lower Fraser River Valley, and southern Vancou- ver Island), there are many ponds, ditches, reservoirs, and slowly flowing low-gradient streams running through agricultural land that may support a range of exotic species, and in places, may be warm enough in summer to allow tropical species to breed. The mild climate of southwestern British Columbia already is hospitable to a wide range of organisms presently in the pet trade (e.g., fishes such as Lepisosteus osseus, L. platostomus, L. oculatus, Ictalurus punctatus, Clar- ias sp., Tanichthys albonubes, Rhodeus sp., Cyprinella lutrensis, Carassius auratus, Cyprinus carpio [koi], Myxocyprinus asiaticus, Misgurnus anguillicaudatus, Gastromyzon sp., and some hillstream loaches [Homa- lopteridae]) or fishes imported live to Canada for human consumption (e.g., Tilapia sp., Sarotherodon sp., Ore- ochromis sp., Channa sp., Hypophthalmichthys sp., and Monopterus sp.). Given predictions of warming aquatic environments in North America (Mandrak 1989; Meis- ner 1990; Rahel et al. 1996; Eaton and Scheller 1996; Schindler 2001; Stefan et al. 2001; Jackson and Man- drak 2002; Vander Zanden et al. 2004) it is likely many more exotics could become established in the water- ways of this province; those that already are present in Canada likely will expand their range northward where habitat and water flow permits. Characid Fishes The large size of most pacu and related “silver dol- lar” species limits their suitability for home aquaria, and likely is the reason why they are occasionally re- leased in North America. Fuller et al. (1999) discuss a variety of pacu and silver dollars (Colossoma sp., Metynnis sp., Myleus sp., Piaractus sp.), which have been found outside of captivity in many of the United States including Washington. Coad (1995, page 11) noted that a specimen of Colossoma bidens was found in Ontario (although the location was not given), and Renaud and Phelps (2001) mention another pacu THE CANADIAN FIELD-NATURALIST Vol. 120 which was found in the Rideau Canal, Ontario. It is not known if there are any wild/naturalized reproduc- ing populations of pacu and silver dollars in North America (Fuller et al. 1999). Apparently the Silver Pacu found in Green Lake on Vancouver Island was the second pacu to be col- lected in British Columbia; the first was taken from the Lower Mainland a few years earlier (R. Dolighan, personal communication, 2005), but no detailed reports or literature references to this fish exist. It has been suggested that the Silver Pacu from Green Lake was from a large display aquarium in a local restaurant. The other pacu in the display aquarium also may have been released when the restaurant closed down, but attempts to locate other specimens in Green Lake were unsuc- cessful. The intentional release of large, unwanted aquarium pets (including large piranha [probably Py- gocentrus nattereri| which may have been released into Langford Lake on Vancouver Island [T. Duce, per- sonal communication, 2005]) is a continuing threat to British Columbia’s waterways, and a threat which is difficult if not impossible to control. Piranhas such as P. nattereri have been found in the wild as far north as Minnesota, Idaho, and the Columbia River in Wash- ington (Fuller et al. 1999). Fortunately pacu and pira- nha usually are kept in small numbers or singly, and so any released specimens are unlikely to find mates and reproduce, even if they survive in British Colum- bia’s waters. Loricariid Catfishes Some loricariid fishes (“‘plecostomus”) can tolerate cool water aquaria and commonly are kept with gold- fish in home aquaria, as are weatherloaches (Misgur- nus anguillicaudatus), and surprisingly, it is the expen- sive Panaque species, not the inexpensive Hypostomus, which have been found in the wild in Canada. In addi- tion to the single Blue-eyed Panaque from British Columbia, a single specimen of the Royal Panaque or Royal “Pleco” (Panaque nigrolineatus), is known from southern Ontario (Coad 1995, page 11). Both the Royal Panaque and the Blue-eyed Panaque are native to South America and are unlikely to survive long in Canada’s present climate, unless they are released downstream of hotsprings or near warm industrial effluent (the same can be said for most tropical fishes, including pacu). Illegally released tropical aquarium fishes persist in Alberta downstream of hotsprings despite cold win- ters in that province (Nelson and Paetz 1992), and the possibility of tropical fish introductions is a persistent threat to hotsprings in British Columbia. Ictalurid Catfishes Washington state waterways contain several ictalurid species, including: Blue Catfish (/ctalurus furcatus), Channel Catfish (/. punctatus), Tadpole Madtom (Notu- rus gyrinus), Flathead Catfish (Pylodictis olivaris), White Catfish (Ameiurus catus), Yellow Bullhead (A. natalis), Black Bullhead (A. melas), and Brown Bull- 2006 head (A. nebulosus) (Wydoski and Whitney 2003). None of these fishes are native to the state and exhibit varying degrees of success in Pacific slope drainages. Black, Brown, and Yellow Bullhead catfishes now are found all over Washington (Wydoski and Whitney 2003), but since casual observers find the three species difficult to differentiate, it is likely that the true range of each is underestimated. Carl et al. (1967), Scott and Crossman (1973), McPhail and Lindsey (1986), McPhail and Carveth (1994), and Coad (1995) detail the distribution of cat- fishes in British Columbia and until this year, re- searchers thought that only the Black Bullhead and Brown Bullhead existed in this province. As in Wash- ington, the range of Black and Brown bullheads like- ly is underestimated because of the difficulty of iden- tification, and limited survey sampling in appropriate habitat. Yellow Bullheads are hardy, and given their pres- ence in the Columbia River system and the Olympic Peninsula in Washington (Wydoski and Whitney 2003), it was only a matter to time before they appeared in British Columbia. It is possible that Yellow Bullheads were introduced to British Columbia as a contaminant in an unsorted or poorly sorted shipment of Large- mouth Bass. The electrofishing sample from Silver- mere Lake taken on 21 July 2005 contained the single Yellow Bullhead, | Brown Bullhead, 5 Pumpkinseeds (Lepomis gibbosus), 28 Prickly Sculpins (Cottus asper), 30 Black Crappie (Pomoxis nigromaculatus), and 57 Largemouth Bass (Micropterus salmoides). Note that only one species (Cottus asper) in the six caught in Silvermere Lake is native to the Fraser River drainage, and that single native species made up only 23% of the catch. The date of introduction of Brown Bullheads into British Columbia is unknown, but they were stocked in Washington from 1874 onwards, and those on Van- couver Island date back to 1906 (Carl and Guiguet 1957; Carl et al. 1967; Wydoski and Whitney 2003). According to anecdotal evidence, the initial stocking in Beaver Lake on Vancouver Island was accomplished from the window of a passing train, and these few bullheads were from an aquarium displayed in a local restaurant (Carl and Guiguet 1957; Carl et al. 1967). Langford Lake and other lakes were stocked soon after, from the Beaver Lake population. It is possible that Black Bullheads were a contaminant in the early stocks of Brown Bullhead since the two species can be diffi- cult to differentiate, especially when young (Becker 1983; Lever 1996). However, if present, Black Bull- heads failed to reproduce and were extirpated while the Brown Bullhead survived on Vancouver Island. Bullhead catfish have not spread far in British Columbia because the warm, eutrophic conditions they require are found only in lakes and ponds of southern Vancouver Island, the Fraser River Delta, and the southern portions of the Columbia River drainage in HANKE, MCNALL, AND ROBERTS: BULLHEAD, CATFISH, PACU IN BRITISH COLUMBIA 425 FIGURE 4. Photographs of (A) the live Yellow Bullhead (Ameiurus natalis) just after capture, and (B) the same specimen once preserved (RBCM 005-079- 001); ruler in (B) is in centimetres. FiGuRE 5. The southeast bay on Silvermere Lake just north of Highway 7 (A) and detail of shoreline habitat (B) where the Yellow Bullhead was collected. this province. Where they have been introduced, bull- heads are reputed to be effective nest predators, and have been implicated in the elimination of Three- spine Stickleback (Gasterosteus aculeatus) popula- tions in smaller lakes along the coast (McPhail and Lindsey 1986). Male stickleback cannot repel intrud- ing adult bullheads, and in one lake near Mission, a large population of sticklebacks went extinct only two years after bullheads were introduced (McPhail and Lindsey 1986; Wydoski and Whitney 2003). The addi- tion of the Yellow Bullhead (Ameiurus natalis) to 426 British Columbia’s aquatic biota is not to be celebrat- ed, and probably represents one more case of a con- taminant species accidentally released with a transplant of game fish. Acknowledgments Initial information on the pacu from Green lake was provided by Craig Wightman, Fish Biologist, and Randy Dolighan, Ecosystem Biologist, both with the Ministry of Water, Land and Air Protection, Environ- mental Stewardship Division, 2080-A Labieux Road, Nanaimo V9T 6J9. Mike deLaronde, Conservation Officer, Industrial Investigations, 2080-A Labieux Road, Nanaimo V9T 6J9, provided the photograph of the pacu from Green Lake. Thanks to Kelly Sendall for information on the Panaque in RBCM collections and changing the data record to reflect the new and cor- rect species identity. Financial support for field work in 2005 was provided by the Royal British Columbia Museum. Ken Stewart (University of Manitoba, Depart- ment of Zoology), and James Cosgrove (Royal BC Museum, Department of Natural History), and two anonymous individuals reviewed and greatly improved an earlier, rushed draft of this manuscript; their com- ments and suggestions are greatly appreciated. Literature Cited Becker, G. C. 1983. Fishes of Wisconsin. University of Wis- consin Press, Madison, USA. 1052 pages. Carl, G. C., W. A. Clemens, and C. C. Lindsey. 1967. The freshwater fishes of British Columbia. Handbook 5, British Columbia Provincial Museum, Victoria, British Columbia. 192 pages. Carl, G. C., and C. J. Guiguet. 1957. Alien animals in British Columbia [revised by C. J. Guiguet 1972]. Hand- book 14, British Columbia Provincial Museum, Victoria, British Columbia. 103 pages. Coad, B. E. 1995. Encyclopedia of Canadian Fishes. Cana- dian Museum of Nature and Canadian Sport Fishing Pro- ductions, Inc. Ottawa, Ontario. 928 pages. 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. Dove, A. D. M., and I. Ernst. 1998. Concurrent invaders — four exotic species of Monogenea now established exotic freshwater fishes in Australia. International Journal for Parasitology 28(11): 1755-1764. Eaton, J. G., and R. M. Scheller. 1996. Effects of climate warming on fish thermal habitat in streams of the United States. Limnology and Oceanography 41: 1109-1115. Font, W. F. 2003. The global spread of parasites: What do Hawaiian streams tell us? BioScience 53: 1061-1067. Fuller, P. L., L. G. Nico, and J. D. Williams. 1999. Non- indiginous Fishes Introduced into Inland Waters of the United States. American Fisheries Society Special Publi- cation 27, American Fisheries Society, Bethesda, USA. 613 pages. Hart, J. L. 1973. Pacific fishes of Canada. Bulletin 180, Fisheries Research Board of Canada. Ottawa, Ontario. 740 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Jackson, D. A., and N. E. Mandrak. 2002. Changing fish biodiversity: Predicting the loss of cyprinid biodiversity due to global climate change. American Fisheries Society Symposium 32: 89-98. Koel, T. M., and J. J. Peterka. 1994. Distribution and dis- persal of fishes in the Red River of the North basin: a progress report. Pages 159-168 in Proceedings of the North Dakota Water Quality Symposium, North Dakota State University, Water Resources Research Unit, Fargo, U.S.A. Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer. 1980. Atlas of North American Freshwater Fishes. Publication (1980-12), North Carolina Biological Survey, North Carolina State Museum of Natural History. 854 pages. Lever, C. 1996. Naturalized fishes of the World. Academic Press, Inc., London. 408 pages. Mandrak, N. E. 1989. Potential invasion of the Great Lakes by fish species associated with climatic warming. Journal of Great Lakes research 15: 306-316. McPhail, J. D., and R. Carveth. 1994. Field key to the fresh- water fishes of British Columbia. Resources Inventory Committee, Government Publications Centre, Victoria, British Columbia. 239 pages. McPhail, J. D., and C. C. Lindsey. 1986. Pages 615-638 in Zoogeography of the freshwater fishes of Cascadia (the Columbia River system and rivers north to the Stikine). Edited by C. H. Hocutt and E. O. Wiley. The zoogeogra- phy of North American freshwater fishes. John Wiley and Sons, New York, USA. Meisner, J. D. 1990. Effect of climatic warming on the south- erm margins of the native range of brook trout, Salvelinus fontinalis. Canadian Journal of Fisheries and Aquatic Sciences 47: 1065-1070. Moyle, P. B. 2002. Inland fishes of California. University of California Press, Berkeley. USA. 502 pages. Nelson, J. S., and M. J. Paetz. 1992. The fishes of Alberta. University of Alberta Press, Edmonton, Alberta. 437 pages. Padilla, D. K., and S. L. Williams. 2004. Beyond Ballast Water: aquarium and ornamental trades as sources of inva- sive species in aquatic ecosystems. Frontiers in Ecology and the Environment 2(3): 131-138. Rahel, F. J., K. J. Keleher, and J. L. Anderson. 1996. Potential habitat loss and population fragmentation for cold water fish in the North Platte River drainage of the Rocky Mountains: response to climate warming. Limnol- ogy and Oceanography 41: 1116-1123. Renaud, C. B., and A. Phelps. 2001. A Pacu/Piranha in the Rideau Canal. Trail and Landscape 35(2): 86-89. Schindler, D. W. 2001. The cumulative effects of climate warming and other human stresses on Canadian freshwaters in the new millennium. Canadian Journal of Fisheries and Aquatic Sciences 58: 18-29. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada, Bulletin 184. Ottawa, Ontario. 966 pages. Smith, R. J. F., and A. Lamb. 1976. Fathead minnows (Pimephales promelas Rafinesque) in northeastern British Columbia. Canadian Field-Naturalist 90: 188. Stefan, H. G., X. Fang, and J. G. Eaton. 2001. Simulated fish habitat changes in North American lakes in response to projected climate warming. Transactions of the Ameri- can Fisheries Society 130: 459-477. 2006 Stewart, K. W., W. G. Franzin, B. R. McCulloch, and G. F. Hanke. 2001. Selected case histories of fish species invasions into the Nelson River system in Canada. Pages 63-81 in Science and Policy: Interbasin Transfer of Aquat- ic Biota. Edited by J. A. Leitch and M. J. Tenamoc. Insti- tute for Regional Studies, North Dakota State University, Fargo, USA. Stewart, K. W., and D. A. Watkinson. 2004. The freshwater fishes of Manitoba. University of Manitoba Press, Win- nipeg, Manitoba. 276 pages. Vander Zanden, J. M., J. D. Olden, J. H. Thorne, and N. E. Mandrak. 2004. Predicting occurrences and impacts HANKE, MCNALL, AND ROBERTS: BULLHEAD, CATFISH, PACU IN BRITISH COLUMBIA 427 of smallmouth bass introductions in north temperate lakes. Ecological Applications 14(1): 132-148. Wydoski, R. S., and R. R. Whitney. 2003. Inland fishes of Washington. University of Washington Press, Seattle, USA. 320 pages. Yamamoto, M. N., and A. W. Tagawa. 2000. Hawaii's native and exotic freshwater animals. Mutual Publishing, Hon- olulu, Hawaii. 200 pages. Received 14 November 2005 Accepted 10 April 2007 Regional Variation in Amelanchier in the Whitewood area of Southeastern Saskatchewan and the First Saskatchewan Records of Amelanchier sanguinea PauL M. CATLING and G. MITRow Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada; e-mail: catlingp @agr.gc.ca Catling, Paul M., and G. Mitrow. 2006. Regional variation in Amelanchier in the Whitewood area of southeastern Saskatchewan and the first Saskatchewan records of Amelanchier sanguinea. Canadian Field-Naturalist 120(4): 428-432. Based on measurements and evaluation of 111 specimens from 51 localities in the Whitewood area of southeastern Saskat- chewan, most Amelanchier alnifolia from the prairie parkland is represented by short, stoloniferous shrubs, less than 2 m tall with 5 — 100+ stems. In the Moose Mountain area, A. alnifolia is represented by shrubs that are taller, sometimes to 7 m, not stoloniferous and usually having fewer than five stems. Although many of these plants have the inflorescence characteristics of A. alnifolia, some are referable to the eastern A. sanguinea, not previously reported from southeastern Saskatchewan. Such plants appear to be confined to the wooded environment of Moose Mountain. Amelanchier alnifolia is not as clearly differentiated by habit or leaf tip characteristics as the literature suggests. Inflorescence characteristics, habit and actual genetic variation may be associated with biogeographic patterns, especially historically wooded islands within the prairie ecozone. These patterns are an important consideration in classification studies and in protecting and utilizing Amelanchier germplasm. Key Words: Juneberry, Serviceberry, Shadbush, Saskatoon, Amelanchier alnifolia, A. sanguinea, status, biogeophaphic vari- ation, hybrid, agamospermy, Moose Mountain, Saskatchewan. At least three species of Amelanchier are known to occur in the prairie region of Canada. Amelanchier alni- folia (Nutt.) Nutt. is widespread, Amelanchier florida Lindl. is confined to ravines in the Cypress Hills (re- ported by some authors and not by others) and A. san- guinea (Pursh) DC. occurs in southeastern Manitoba (e.g. Boivin 1967; Looman 1979). Reports of A. san- guinea from the northern Great Plains (Stephenson 1973; McGregor et al. 1977) are based on A. humilis Wieg. (McGregor et al. 1986), whereas Boivin’s (1967) report of A. sanguinea in southeastern Manitoba may be based on material of both A. sanguinea and A. hu- milis since he combined the latter with the former and both occur in the general region and in northern Min- nesota (Nielsen 1939; Jones 1946). In general, flowering material of A. sanguinea 1s dis- tinguished from A. alnifolia by having flowering pedi- cels more than 15 mm long instead of less than 10 mm long (Boivin 1967; Cinq-Mars 1971; Gleason and Cronquist 1991) and petals 11-22 mm long instead of 6-10 mm long (e.g., Jones 1946). In addition, A. san- guinea is characterized as a tree with oblong, pointed leaves and A. alnifolia is generally characterized as a stoloniferous shrub with truncate or rounded leaves (e.g., Nielsen 1939; Jones 1946; Fernald 1950). Ame- lanchier humilis is distinguished from A. alnifolia by its acute or obtuse instead of truncate or broadly round- ed leaves (Cinqg-Mars 1971; McGregor et al. 1986). McGregor et al. (1986) have indicated that it is diffi- cult to separate A. alnifolia and A. humilis in the north- ern Great Plains and they have suggested that some colonies “appear as hybrid swarms and the two could easily be considered as variants of one species.” The concept of including A. humilis with A. alnifolia is not new since it was the basis for McKay’s (1973, 1975, 1976) transfer of Nielsen’s A. humilis var. com- pacta to A. alnifolia (A. alnifolia var. compacta (Niel- sen) McKay). During a survey of Amelanchier in the Whitewood and Moose Mountain areas of southeastern Saskatche- wan in early June 2005, it appeared that plants in the prairie-parkland region (ecodistricts 752, 756, 760, Agriculture and Agri-Food Canada 1995) differed from those on the elevated Moose Mountain plateau (eco- district 761) which was forested in presettlement times (Catling and Kostiuk 2006). This rolling plateau, al- most 400 km? in extent with numerous lakes and well- drained wooded slopes with aspen (Populus) and birch (Betula) forest, is 100 — 150 m above the surrounding plains. The objective of the work reported here was to doc- ument the variation and to provide a context for the first report of Amelanchier sanguinea in Saskatchewan. This information is relevant to systematic studies of Amelanchier and is also of interest with regard to the : protection of native Amelanchier germplasm. Although Saskatoon (Amelanchier alnifolia) is one of Canada’s fastest growing native crops, its variation is not well understood and its domestication has yet to take advan- tage of the full range of variability available (Catling and Small 2003). 428 2006 103° 00° CATLING AND MITROW: AMELANCHIER IN SASKATCHEWAN 429 50° 00° { w 49° 30° 102° 00° FiGurE |. Saskatchewan (left) showing location of Whitewood study area (right). Collection localities of Amelanchier used in the study are indicated by dots. The Moose Mountain ecodistrict is shaded. The collection site of A. sanguinea is indicated by “S” and dashed line connected to the dot. Methods . In June 2005, during late flowering, 111 specimens of Amelanchier were collected and measured from 51 sites in the Whitewood — Moose Mountain area of southeastern Saskatchewan (Figure 1). No more than five specimens were collected at each site and sites are defined as at least 0.1 km apart. Specimens were select- ed to represent the extremes of variation present at a site. For each specimen, lengths of lowest pedicel in an inflorescence of average size, average petal length in lowest flower in the inflorescence, average sepal length in lowest flower in the inflorescence, the plant habit (stoloniferous or not, height, number of stems), leaf pubescence (scale 1-4, 1 = glabrous, 4 = densely pubes- cent) and shape of the tip of a larger leaf (scale 1-5 with | = truncate, 5 = pointed). Stoloniferous shrubs were defined as having more than 6 separated stems and being less than 2 m in height. Non-stoloniferous shrubs were defined as having 5 or fewer stems together and exceeding 2 m in height. The data were analysed using Principal Components Analysis (PCA), regression and histograms. PCA re- duced dimensionality of six characters (Table 1), to two allowing for an evaluation of grouping and indi- cation of pattern of variation. Histograms evaluated pedicel length with respect to habitat for different eco- districts. F-ratios based on the comparison of the mean Square variance of the regression model to the mean Square of the residual indicated relationship and sig- nificance of individual characters noted above. Stat- graphics software (www. statgraphics.com) was used to perform all the statistical analyses. Results and Discussion On the PCA plot (Figure 2) that reduces the dimen- sionality of six characters, the first two components account for 32.53% and 25.75% of the sample varia- tion and the third and fourth account for 16.90% and 11.68%, respectively. On the first component, pedicel length, sepal length and height are heavily weighted whereas on the second component, leaf tip and leaf shape are heavily weighted. Leaf pubescence was not heavily weighted on either of the first two components. With a cumulative 58.28% of the sample variation accounted for in the diagram, the plot is marginally useful as an indication of potential groups, and groups are unclear. However, the plants from historically wood- ed Moose Mountain area occupy a specific part of the dispersion suggesting a different pattern of morpho- logical variation from that represented elsewhere in the region. The plants from Moose Mountain tend to have longer pedicels, and they are mostly non-stoloniferous (Fig- ure 3). Variation in leaf tip shape, overall leaf shape, leaf pubescence and the inflorescence characters is extensive. Most plants on Moose Mountain that are presumably A. sanguinea on the basis of height and non-stoloniferous habit have to be treated as A. alni- 430 ‘ | ° ae | 2 ° 20 e 9 © = i o 2 roe) R fo) Go % ° = oo oem o oO oe nN eAlihs coe oe °, = 9 7 = o° od 3 OU = fo} oe) , Cy 0 a fo) o 02a Ce ee er) e aeie I oP i O nics ogo %e ° Ga 2 cee “ 3 2 | eal | al ii Component 1 (32.53%) FIGURE 2. Plot of 111 specimens used in the study on the first two principal components derived from correllation among six morphological characters. The specimens from Moose Mountain are shown with solid dots and these occupy a specific portion of the dispersion sug- gesting distinctive variation within this region. folia on the basis of pedicels less than 15 mm long and petals less than 10 mm long. The ten plants poten- tially referable to A. sanguinea on the basis of pedicels over 15 mm (Figure 3) had leaves with variable tips. For example five shrubs had more or less truncate leaves, four had most leaves broadly rounded and two had many pointed leaves. Leaf shapes also varied. Eight of these are moderately pubescent but two are more or less glabrous. All of these ten had petals over 9 mm but under 12 mm long. The two specimens ac- cepted as A. sanguinea (e.g., Figure 4 left) had pedicels 17 mm, petals 11-12 mm, pointed, moderately pubes- cent, oblong leaves. These two plants were distinctive in blooming later than other plants nearby that pos- sessed fewer A. sanguinea characteristics. The speci- men label data for these are: SASKATCHEWAN: Moose Mountain: White Bear Lake, 49.7550°N, 102.2565°W, 8 June 2005, P. M. Catling 05-27, B. Kostiuk (DAO); Good Bird Point, White Bear Lake, 49.7639°N, 102.2328°W, 6 June 2005, P. M. Catling 05-16, B. Kostiuk (DAO). Amelanchier sanguinea has not been reported previously from Saskatchewan (Harms 2003). Amelanchier alnifolia from the prairie parkland is represented mostly by short, stoloniferous shrubs, less than 2 m tall with 5—100+ stems, but taller, non-stolonif- erous shrubs also occur in poplar groves. The smaller stoloniferous plants often have distinctively short and dense racemes with pedicels little if at all longer than the petals (Figure 4, right). The leaves may be truncate or more or less pointed. The regressions indicated that pedicel length had the most significant relationships with other characteris- tics and was significantly correlated with ecodistrict (Table 1). Sepal length was also significantly correlat- THE CANADIAN FIELD-NATURALIST Vol. 120 = o =a NO © Fa DN © © stoloniferous non-stoloniferous =a O&O Prairie Region Number of Specimens stoloniferous non-stoloniferous Oy os C5 hy ey fy oes Moose Mountain i Mindi ilk anise 6 SU One omen 4G We AD ee! Pedicel length (mm) FIGuRE 3. Histograms showing pedicel lengths and growth habits for specimens of Amelanchier from the prairie region and the Moose Mountain region of southeastern Saskatchewan. ed with other characters, but was not significantly asso- ciated with leaf tip which is regarded as a significant character for distinguishing A. alnifolia by some authors (see above). In overall addition of significance values, pedicel length was three time more significant than sepal length. Futhermore, sepals are more often dam- aged or shrunken (as in fruit) whereas pedicels are more often intact. This represents the first quantitative support for the use of pedicel length in defining taxa. Leaf shape was only highly correlated with leaf tip and was not significantly related to ecodistrict suggesting limited use in defining taxa. These suggestions are based on the operational concept of taxa being defined by a number of correlated morphological characters and by some ecological discreteness. Thus the more recent- ly defined (see above) characters are supported and these are included in the following key which will dif- ferentiate the four species known from the prairie provinces. la. Pedicels short, less than 10 mm long; petals 4-10 mm long; mostly stoloniferous shrubs 2a. Leaves glabrous or sparsely tomentose at flowering, truncate or broadly rounded ... .A. alnifolia 2b. Leaves persistently pubescent at flowering, OIE TORUS go cabcoccccscuapnocoune A. humilis 1b. Pedicels longer, over 15 mm long; petals 11-22 mm long; non-stoloniferous shrubs .......... 3 3a. Leaves glabrous at flowering ............. A. florida 3b. Leaves more or less tomentose atiflowering Wier feared antec A. sanguinea 2006 CATLING AND MITROW: AMELANCHIER IN SASKATCHEWAN 43] ‘Reserva Brasil, Foz do Iguacu 85851-970 Brazil; e-mail: annesophie @reservabrasil.org.br 2Ontario Puma Foundation, Beeton, Ontario LOG 1A0 Canada 3Kouchibouguac National Park of Canada, Kouchibouguac, New Brunswick E4X 2P1 Canada 4 Associate Vice-President, Research, Laurentian University, Sudbury, Ontario P3E 2C6 Canada °Fundy National Park of Canada, Alma, New Brunswick E4H 1B4 Canada Bertrand, A.-S., S. Kenn, D. Gallant, E. Tremblay, L. Vasseur, and R. Wissink. 2006. MtDNA analyses on hair samples confirm Cougar, Puma concolor, presence in southern New Brunswick, eastern Canada. Canadian Field-Naturalist 120(4): 438-442. For the last 40 years, the presence of Cougars (Puma concolor) in eastern Canada has been highly controversial. The purpose of this study was to collect physical evidence of Cougars using a passive detection method. Baited hair-traps combined with © camera-traps were installed in New Brunswick and Nova Scotia, Canada. DNA analyses on two hair samples confirmed that the species was present in southern New Brunswick in 2003. A footprint photographed after an observation of a Cougar by reliable observers was examined by experts and was consistent with a Cougar footprint. Additional data are required to deter- mine the status of Cougars in the northeastern part of its historical range. Key Words: Cougar, Puma concolor, hair-traps, camera-traps, physical evidence, mtDNA analyses, tracks, eastern Canada. Historically, the Cougar (Puma concolor) was dis- tributed all across the American continent, from south- eastern Alaska to southern Argentina and Chile (Park- er 1998). It was the most widely distributed terrestrial mammal in the western hemisphere (Godin 1977). The subspecies referred to as Eastern Cougar (Puma con- color couguar) was known to occur in Nova Scotia [NS], New Brunswick [NB], Quebec [QC], Michi- gan, Tennessee and South Carolina (Goldman 1946). While 32 subspecies were initially listed by Goldman (1946), a recent DNA study by Culver et al. (2000) showed that only six genetically distinguishable sub- species are now believed to occur in the western hemi- sphere, with only one for North America; 1.e., P. c. couguar. Populations in northeastern America drasti- cally declined at the beginning of the last century (Parker 1998). Weaver et al. (1996) argued that con- flicts for resources and land uses may be the main causes of most Cougar extirpation in North America. Previous studies across North America have indicated that road density, urbanization, agriculture and timber harvesting may also limit Cougar expansion (van Dyke et al. 1986; Maehr et al. 2002; Taylor et al. 2002; Dick- son et al. 2005). The status of the Cougar in eastern North America has always been a highly controversial topic (Cumber- land and Dempsey 1994). Due to the secretive habits of this species and the highly fragmented regions where few scattered individuals possibly remain, solid evi- dence (i.e., a dead animal, DNA collected from scat or hair, a clear photograph or indisputable confirmation of authenticated tracks) is difficult to collect, making decision and species-at-risk designation difficult. Prov- incially, Cougars are listed as undetermined in NS, and | endangered in NB. At the federal level, the status of the eastern cougar population was changed from endan- gered to data deficient in 1998 (Scott 1998"). No reli- able estimate of the number of Eastern Cougars has ever been made because authentic, scientifically-based evidence is lacking (Cumberland and Dempsey 1994). However, numerous credible sightings and physical evidences have since then reopened the Eastern Cougar debate. An extensive review carried out by the Com- mittee on the Status of Endangered Wildlife in Canada [COSEWIC] in 1978 leaves little doubt as to the his- torical presence of Cougars in eastern Canada (van Zyll de Jong and van Ingen 1978"). Today, the major concern of provincial wildlife agencies is whether the reported animals are natives or feral escaped or re- leased captives (Scott 1998’). While extensive, logistically demanding fieldwork) is usually required to search for such a wide-ranging ' species, this paper focuses on new data obtained for’ NB through non-invasive detection methods (i.e., bait- ed hair-trap and camera-trap). Material and Methods Study Areas This project was conducted in NB and NS with 12. traps set up in three national parks, 1.e., Fundy Nation-) al Park of Canada [FNP], Kouchibouguac National Park’ of Canada [KNP] in NB, and Cape Breton Highlands National Park of Canada [CBHNP] in NS. Conserva- tion units being the largest wilderness areas available, they are more likely to shelter wide-ranging top pred- ators than urban or agricultural lands. Two additional, 438 | 2006 er eae Quebec DE me ‘ j # ; New Brunswick | “t F “Frederfcfon, X Saint, ohn ’ + p Nova A Scotia BERTRAND ET AL.: MTDNA ANALYSES ON HAIR CONFIRM COUGAR 439 a bs Prince Edward Island S re \ (3 Se Halifax Ap N FiGuRE 1. Baited hair-trap (n = 14) locations (triangles) in New Brunswick and Nova Scotia, eastern Canada. traps were installed in remote areas in the Miramichi region (northeastern NB) because sightings from reli- able observers (biologists and trappers) were fre- quently reported to provincial wildlife agents. All traps were located according to the most recent credi- ble sighting report or physical evidence (Figure 1). Four traps were set up in FNP in forested habitats on the rolling upland plateau. FNP (205.9 km?) is char- acteristic of the Maritime Acadian Highlands region and is part of the Appalachian Mountain range (Wood- ley et al. 1998). Three other traps were installed in KNP (239.2 km?) in northeastern NB. Two traps were in- stalled in coniferous stands, one in a relatively remote area near Black River in KNP’s central region, and the other, less than | km from human habitations along the southeastern boundary of the park. The last KNP trap was installed in a hilly mixed wood stand 3 along the Major Kollock creek. Finally, five traps were installed in CBHNP (948.0 km?), the largest protected wilder- ness area in NS, protecting 20% of northern Cape Bre- ton. Traps were installed based on recent convincing Cougar sightings in deciduous, mixed wood and dead conifer forests. All trap locations had a closed canopy, an important amount of coarse woody debris or a thick leaf litter, and were close to water sources. Non-invasive detection techniques Field work began during the fall of 2003. The pas- Sive detection methods used entail collecting hairs, tracks, and scats in a non-invasive manner. Baited hair- traps (Figure 2), consisting of 2-m-high posts sur- rounded by 3 x 3 m? squared enclosures, were installed in areas where credible sightings had recently been reported (Figure 1). Two barbed wire strands were stretched around the wooden posts delimiting the enclo- sure, at 45 cm and 75 cm from the ground respectively. This was intended to catch hair as the animal enters the enclosure, attracted by the species-specific lure. Drip- ping lure (i-e., Cougar urine obtained from captives, sex unknown) was hung inside the perforated central post and fresh lure was added every month. In 2004, in an attempt to get photographic records of animals entering the enclosure, we equipped each station with a camera-trap unit which consists of either a 35-mm or a digital camera triggered by an infrared motion sensor. Each camera was fastened to a tree near the scent post, at a height of 1.8 m (Figure 2). Any motion up to 7 m away within an angle of 120 degrees activated the camera. Traps were checked year-round every 4-5 weeks. DNA analyses Samples were submitted to a sequence-based analy- sis of 16S mitochondrial DNA (mtDNA: Johnson and O’Brien 1997). DNA was extracted from hair samples using QIAGEN DNeasy Tissue kits. In order to deter- mine whether DNA samples were from felid or other mammal species (i.e., felids, canids, cervids, 4 mustelids, procyonids, and ursids), 300 base pairs of 16S mtDNA were amplified using Polymerase Chain Reaction (PCR). 440, FIGURE 2: Experimental design for Cougar detection, a passive method combining a baited hair-trap (central post and THE CANADIAN FIELD-NATURALIST cae "le §=enclosure Vol. 120 ed hair-tr my So : q Barbed-wired PR LAS = as barbed-wired enclosure) protocol and a camera-trap (upper left corner, strapped on a tree). A volume of 15uL contained 100-500 ng template DNA, |x reaction buffer, 1.5 mmol (millimoles) MgCl, 0.3 mmol each primer, 250 mmol dNTPs and 1 U Tag polymerase. After initial incubation at 94°C for 2 min- utes, 40 cycles of PCR were performed at 94°C for 1 minute, 53°C for 1.5 minutes, 72°C for 1.5 minutes, and 72°C for 10 minutes. Results were then visualized on a 2.0% agarose gel and compared to 20 species from the six families cited above. A second amplification of 600 bp of a specific region of the 16S mtDNA was then conducted according to the protocol detailed by Mills et al. (2000). Restriction enzymes Haelll, Hpall, and Rsal then digested PCR products at 37°C during 12 hours, and results were visualized on 2.5% agarose gel. Since restriction enzymes produce species-specific patterns, it was then possible to distinguish Cougar from other felid species. Results Baited hair traps A total of 207 samples was collected from hair traps (Table 1). MtDNA analyses conducted on hair samples collected in 2003 revealed that two samples collected in FNP were Cougar hair. The first hair sam- ple was found near the main road entering the park TABLE |. Number of hair samples collected per location and per year for 2003-2006. Location Sampling periods 2003 2004 2005 2006 KNP 0 6 15 7 FNP 29 34 49 23 CBHNP 1 3 17 16 MIRAM —* - 6 1 Total 30 43 87 47 *Baited hair-traps and camera-traps were installed in the Miramichi (MIRAM) area during the fall of 2004. (i.e., Kinnie Brook), in a young Red Spruce (Picea rubens) and Balsam Fir (Abies balsamea) stand (6 to 12 m high, canopy closure of 70%). Ground cover mostly consisted of Sphagnum spp. and Bunchberry (Cornus canadensis). The other hair sample was found in the northern area of the park, along an old logging trail (i.e., Big Dam trail) relatively overgrown with Red Spruce, Balsam Fir and birch (Betula spp.). In this case, trees were taller (12-20 m) but canopy closure was | only 40-50%. A 6-m-high understory composed of Balsam Fir, Red Spruce and White Birch (Betula papy- rifera) allowed no ground cover other than birch leaves. 2006 BERTRAND ET AL.: MTDNA ANALYSES ON HAIR CONFIRM COUGAR 44] FIGURE 3. Footprint found west of Blackville (GPS coordinates (NAD 1983): 20N0724671; UTM 5188486), New Brunswick, Canada (22 September 2004). Picture provided by P. Boucher. Box (to left of footprint) diameter: 8.5 cm. Further DNA sequencing revealed that the two hair samples came from two distinct animals, i.e., a North American specimen and a South American specimen. Camera-traps were only installed in 2004, so there is no photographic record of these two individuals. No Cougar hair or picture were collected after the cameras had been installed. Additional Observation Sets of tracks can also be used to identify species (van Dyke and Broke 1987; Cumberland and Dempsey 1994). A footprint (Figure 3) was photographed in 2004 by three reliable observers in Blackville (southwest of Miramichi, New Brunswick) 15 minutes after they had seen the Cougar at 12:25 PM and watched it for several seconds as it was crossing a dirt road 45 m from them. This track was examined by experts and declared not to be of a Coyote (F. Scott and P. G. Crawshaw Jr., personal communication). Discussion Cougar detection program for the Maritimes Hair-traps and camera-traps are alternative and con- venient non-invasive methods that are cost- and time- efficient techniques for animal surveys (Carbone et al. 2001). After four years of permanent cougar monitor- ing in the Maritimes, the amount of field data keeps increasing. As suggested by Cardoza and Langlois (2002), it would be valuable to establish a systematic monitoring program supervised by an expert team for the Eastern Cougar population, especially for the prov- inces where the status of this species is indeterminate. This would allow collection of a series of undisputable physical evidence (Cardoza and Langlois 2002; Maehr et al. 2003). As mentioned by others, the probability of proving the presence of a species which has no es- tablished viable populations in a given area is almost nil (Broke and van Dyke 1985). DNA hair analyses confirmed that two Cougars oc- curred in southern NB in 2003. Additional data (e.g., individual genetic haplotype) are nonetheless crucial to better understand the status of the species in this part of its range. One of the individuals detected in Fundy turned out to be a South American animal, recalling the Chilean specimen shot in Abitibi, Québec, in May 1992 (Jolicoeur et al. 2006). This may indicate that a few scattered escapees from zoos or captives released by private owners remain (Stocek 1995). However, the other positive hair sample caught in the Big Dam Trail in FNP was from an animal of North American origin. This opens up a range of possibilities. In eastern North America, estimates of the number of escaped captives kept increasing during the last 30 years. In Pennsyl- vania for instance, McGinnis (1996) reported only 31 cougars held in captivity in 1979, among which four or five eventually escaped. Then, less than 20 years later, according to J. Seidensticker (cited in Scott 1998°*), there was an increase of 640% in the number of licensed private Cougar owners. This dramatic in- crease in the number of captive individuals probably suggests many more escapees of North and South American origins. Further DNA identifications will en- able researchers to shed more light on this question. The fact that cougars are efficient colonizers (Nero and Wrigley 1977) with large home ranges (Seidenticker et al. 1973) ensures that the debate about the poten- tial presence of animals in eastern Canada continues. Reported cougars may be transients, escapees or mem- bers of a remnant population, but they could act as dis- persers for the (re)establishment of viable populations in eastern Canada (Scott 1998"; Maehr et al. 2002). The 442 determination of these cats’ origin should help define the protection status they deserve. Acknowledgments The authors acknowledge the information provid- ed by F. Scott, M. Mazzolli, P. G. Crawshaw Jr., and J. Dalponte, technical assistance by P.-E. Hébert (KNP), G. Sinclair (FNP), S. Lambert, J. Hudson and L. A. Reeves (CBHNP). Comments by J. Tischendorf, H. McGinnis, J. Bridgland, G. T. Campbell and two anony- mous reviewers significantly increased the quality of this paper. This project was supported by Parks Cana- da — Species at Risk Program Grant to L. V. Documents Cited (marked * in text) Scott, F. W. 1998. Updated COSEWIC status report on Cougar (Puma concolor couguar) [eastern population]. Committee on the Stauts of Endangered Wildlife in Canada (COSEWIC), Ottawa. van Zyll de Jong, G. G., and E. van Ingen. 1978. Status report on eastern cougar Felis concolor couguar in Canada. COSEWIC, Ottawa. Literature Cited Broke, R. H., and F. G. van Dyke. 1985. 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G., J. S. Jenness, and P. Beier. 2005. Influence of vegetation, topography, and roads on cougar movement in southern California. Journal of Wildlife Management 69: 264-276. Godin, A. J. 1977. Wild mammals of New England. John Hopkins University, Baltimore. THE CANADIAN FIELD-NATURALIST Vol. 120 Goldman, E. A. 1946. Classification of the races of the puma. The puma: mysterious American cat. Reprint. Dover Pub- lications, New York. Johnson, W. E., and S. J. O’Brien. 1997. Phylogenetic reconstruction of the felidae using 16S rRNA and NADH- 5 mitochondrial genes. Journal of Molecular Evolution 44: S98-S116. Jolicoeur, H., A. Paquet, and J. Lapointe. 2006. Sur la piste du couguar (Puma concolor) au Québec, 1955-2005 : ana- lyse des rapports d’ observation. Le Naturaliste canadien 130: 49-58. Maehr, D. S., E. D. Land, D. Shindle, O. L. Bass, and T. S. Hoctor. 2002. Florida panther dispersal and conserva- tion. Biological Conservation 196: 187-197. Maehr, D. D., M. G. Kelly, C. Bolgiano, T. Lester, and H. McGinnis. 2003. Eastern cougar recovery is linked to the Florida panther: Cardoza and Langlois revisited. Wildlife Society Bulletin 31: 849-853. McGinnis, H. J. 1996. Reports of pumas in Pensylvania, 1890-1981. Proceedings of the 1‘ Eastern Cougar Confer- ence, 1994. American Ecological Research Institute — AERIE, Fort Collins. Mills, L. S., K. L. Pilgrim, M. K. Schwartz, and K. McKel- vey. 2000. Identifying lynx and other North American felids based on MtDNA analysis. Conservation Genetics 1: 285- 288. Nero R. W., and R. E. Wrigley. 1977. Status and habits of the Cougar in Manitoba. Canadian Field-Naturalist 91: 28-40. Parker, G. R. 1998. The eastern panther. Nimbus Publishing, Halifax, Nova Scotia. Seidensticker, J. C., M. G. Hornocker, W. V. Wiles, and J. P. Messick. 1973. Mountain lion social organization in the Idaho primitive area. Wildlife Monographs number 35. Stocek, R. F. 1995. The Cougar, Felis concolor, in the Mar- itime Provinces. 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Received 19 December 2005 Accepted 27 December 2006 The Leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its North America Distribution and Additional Notes on Species Description PETER HOVINGH 721 Second Avenue, Salt Lake City, Utah 84103 USA; e-mail: phovingh@xmission.com Hovingh, Peter. 2006. The leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its North American distribution and additional notes on species description. Canadian Field-Naturalist 120(4): 443-451. The geographic range of Haemopis lateromaculata Mathers 1963 (Hirudinea: Haemopidae) is extended across North Amer- ica. Its distribution in the coastal region of Alaska and British Columbia suggests a coastal Pleistocene refugia separate from the populations in the lower United States and suggests that H. /ateromaculata and the Eurasian H. sanguisuga Linnaeus 1758 are sister taxa. Support of the identification and geography is based on the anatomical positions of the reproductive organs in H. lateromaculata and H. marmorata Say 1824. The variations within these species are described, noting that no specific variation was confined to a geographical region. Key Words: leech, Haemopis laterormaculata, description, taxonomy, distribution, North America. In an effort to understand the limits of leech (Hi- rudinea) fauna distribution in western North America, a geographical study defined by hydrological basins was undertaken and is still in progress. The report herein describes a taxonomic problem with Haemopis marmorata (Say 1824) in western North America, a species known to be difficult to distinguish by external characteristics from H. grandis (Verrill 1874) (Sawyer 1972) and H. lateromaculata. Four populations of Haemopis were found without teeth and these popu- lations were initially identified as H. grandis (Hovingh 1993). Examination by dissection revealed that the penis sheaths of these populations were H. marmorata and that these specimens had jaws. As a result of this identification problem, the penis sheaths of all mature Haemopis were examined. In examining the reproductive organs of Haemopis, it became apparent that H. /ateromaculata specimens in museum collections had often been misidentified as H. marmorata. A study of Utah (Beck 1954) and Alas- kan (Moore and Meyer 1951) Haemopis vouchers de- posited in the United States National Museum (USNM) contained several lots identified as H. lateromaculata. Three lots of specimens that were collected from British Columbia and Newfoundland, two of which were iden- tified by J. Madill as H. lateromaculata plus one un- identified due to poor conditions are deposited in The Canadian Museum of Nature (CMN) in Ottawa, On- tario, Canada. The two lots at CMN are the first obser- vations of H. lateromaculata outside of the type local- ities of Minnesota and Iowa (Mathers 1963). A specimen from Michigan was utilized in a leech phylogenetic studies (Borda and Siddall 2004). The reproductive organs of Haemopis marmorata and H. lateromaculata specimens collected from west- ern North America examined are presented in this paper. Based on accurate diagnosis of the species, the external characteristics (pigmentation patterns) were determined to distinguish these two species from one another. The results herein describe the geographical distribution of H. lateromaculata, and support some taxonomic and paleogeographic hypotheses concern- ing the family Haemopidae. Methods Specimens of Haemopis were collected during the last 20 years from localities in western North America. The surveys consisted of visual encounters and hand picking from the underside of stones, logs, and aquatic vegetation. The specimens were relaxed with 10% eth- anol, blotted by paper towel to removed the mucous excreted from the relaxation process, fixed with 10% formalin in phosphate buffered saline between two sheets of paper towels overnight, and placed in 70% ethanol for preservation. Identification of all large Haemopis specimens was determined by examining the reproductive organs through a mid-dorsal cut from the annulus with the female gonopore posteriorly approximately 20 annuli. If the penis sheath was not located with this examina- tion, the dorsal cut was extended anteriorly for some 10 annuli. Teeth and jaws were examined by a mid- ventral cut of the lower lip. Posterior sucker width, pig- mentation, and gonopore positions were noted. Small specimens were identified by pigmentation. Identifi- cations followed original descriptions and taxonomic keys (Mathers 1963; Klemm 1985; Sawyer and Shelley 1976). The facilities and specimens of the United States National Museum (USNM), Washington D.C. and the Canadian Museum of Nature (CMN), Ottawa, Ontario were utilized. The western specimens of Haemopidae in these collections were confirmed by dissection if the length was greater than 40 mm, and pigmentation pat- terns and the presence of teeth were examined. The eastern specimens of Haemopis marmorata were select- 443 +44 ed by pigmentation with dissection on random samples. Specimens were also examined from water quality studies in Wyoming (Wyoming Department of Water Quality), now residing in the Albertson College of Idaho Orma J. Smith Museum of Natural History (ALBRCIDA), Boise, Idaho. Taxonomic status There is presently full acceptance of the haemopisine species (Davies 1991; Klemm 1985; Sawyer 1986). The higher classification of leech groupings, and leech- es within the Annelida, is presently contentious, how- ever. Historically, all the haemopisine species were placed in the genus Haemopis including the original description by Mathers (1954, 1963) of H. kingi Math- ers 1954 and H. lateromaculata. Richardson (1969) revised the family Hirudinidae, established the family Haemopidae, and revised the genus classification by introducing the genera Percymoorensis including P. terrestris (Forbes 1890), P. marmorata, P. lateromac- ulata, and P. kingi), Mollibdella grandis, and Bdellaro- gatis plumbeus Moore 1912). He retained Haemopis for the Eurasian H. sanguisuga Linnaeus 1758. Rich- ardson (1971) suggested that Percymoorensis and Haemopis (the Eurasian species) were associated in the subfamily Haemopinae, and that Mollibdella and Bdellarogatis were in the subfamily Mollibdellinae; he also identified a Mexican species Percymoorensis caballeroi Richardson 1971 in that paper. Davies (1991); Davies and Govedich (2001); and Sods (1969) accepted the generic nomenclature of Richardson (1969) but retained the earlier familial clas- sification of Hirudinidae. Sods (1969) expressed reser- vation about Richardson’s classification of Hirudinidae (including Haemopidae) based on his use of type-speci- mens, and that “long standing genera are strongly het- erogeneous in content and cannot any longer be sepa- rated one from the other entirely by single factors, or contained within single divisions in the key”. Klemm (1995), Manoleli et al. (1998), and Sawyer (1986) re- jected the generic nomenclatural changes of Richard- son but accepted the family Haemopidae classification. A new species, Haemopis septagon Sawyer and Shelley 1976, was described from specimens collected from North Carolina in eastern United States. In their description Sawyer and Shelley noted that if the classi- fication of Richardson (1969) was accepted, H. septa- gon warranted placement in a new genus. Davies (1991) placed this species with the genus Percymoorensis. The uncertainty of Richardson’s classification with respect to H. kingi and H. lateromaculata was revealed when Richardson (1969, 1971) stated that these two species most likely belong to a new genus. He did not exam- ine either species but relied on Mathers (1954, 1963) for his paper (Richardson 1969). Borda and Siddall (2004) examined six species of Haemopis for phylo- genetic studies involving anatomical characteristics, 12 S mitochondrial rDNA, 18S and 28S nuclear rDNA, THE CANADIAN FIELD-NATURALIST Vol. 120 and cytochrome c oxidase subunit I gene fragments. Of the 24 anatomical criteria that were utilized, 23 were common to all six species, thus not significant in deter- mining genera or species relations. Borda and Siddall (2004) confirmed the family Haemopidae. I maintain use of the genus Haemopis and the family Haemopi- dae until additional and more conclusive phylogenet- ic studies have been reported for all the hemopisine species, using population samples representing the entire range of these widespread species. Results Distribution of Haemopis lateromaculata in North America The Nearctic distribution of Haemopis lateromac- ulata is shown in Figure | and listed in Table 1. This distribution is based on museum specimens (USNM, ALBRCIDA, and CMN), from my (PH) collections from the geographical studies in western North America and from water quality studies in Wyoming (ALBR- CIDA). Anatomical dissections distinguished H. lat- eromaculata from H. marmorata in western North America when the plain ventrum and plain, cream, or yellow colored lateral margins were prominent, and thus could be used to distinguish these two congeners from one another. Based on this pigmentation distinc- tion, juvenile specimens (< 40 mm) and museum spec- imens in eastern North America H. marmorata, were examined and those with no ventral pigmentation were considered H. lateromaculata. Some specimens with- out pigmentation were dissected and the species deter- mined. The following are the locations of Haemopis lateromaculata (underlined accession numbers refer to museum specimens catalogued as H. marmorata or as Haemopis). Most of these misidentified specimens had been catalogued and accessioned into the museum collections before the work of Mathers (1963) had been completed and published. Alaska: USNM 19230 Yes Bay, 20732 Afognak Is- land, 27255 Yes Bay, 21256 Sanak Island, 43455 Revillagigedo Island, CMNA 1982-0746 Esther Island, and PH collections from Susitna River drainage, Kenai Peninsula, and Kodiak Island. Western Canada: Yukon Territory: PH collection from Pelly River drainage (Yukon River drainage); British Columbia: USNM 1/923] North Fork Moose River (at least two Moose Rivers occur in British Col- umbia), and USNM 38352 “D” [Dee] Lake; CMNA 1985-0289 and CMNA 1990-0056 (identified by J. Madill, but was not found in the collection) Fraser River drainage; and from PH collections from the Fras- er River drainage (CMNA 2006-0036) and Haida Gwaii (Queen Charlotte Islands) (CMNA 2006-0037); Alber- ta: CMNA 1978-0426 Mackenzie River drainage; and from PH collection Peace River drainage (Mackenzie River drainage) (CMNA 2006-0034). Lower western United States: Idaho: from PH col- lection in the Snake River drainage (Columbia River 2006 0 1000 Kilometers HOVINGH: HAEMOPIS LATEROMACULATA IN NORTH AMERICA 2008 Ficure |. The distribution of Haemopis lateromaculata in North America. Locations are shown by (a) squares, museum specimens (NMNH, CMN); (b) circles, western North America survey sites (preliminary results) and identifications from Wyoming water quality studies; and (C) diamonds, Mathers (1963) locations. drainage); Oregon: from PH collection in the Klamath River drainage and the Great Basin; Wyoming: from PH collection in the Snake River drainage (Columbia River drainage) and Water Quality Studies in Missouri River drainage (ALBRCIDA 61050-61052); California: from PH collection in the Pit River drainage (Sacra- mento River drainage); Nevada: from PH collection in the Great Basin; Utah: USNM 38308, 39056, USNM 38624, 38627 (the latter two identified to genus), and from PH collection in the Great Basin and Colorado River drainage; Colorado: from PH collection in the South Platte River drainage (Missouri River drainage); New Mexico: USNM 50/92 Rio Grande drainage (did not dissect) . Eastern North America: Canada: Saskatchewan: USNM 38353; Manitoba: USNM 38347, CMN 1985- 0292, 1985-0301; Ontario: USNM 38360, 38348, CMN 1985-0253, 1985-259, 1985-279, 1987-0670, 1988-0036, 1988-0163; Newfoundland: CMN /982- 0654, 1985-0378 (both identified by J. Madill): lower United States: Michigan: USNM 3833/; Missouri: USNM 2068/; District of Columbia: USNM 30223; Virginia: USNM 42650. 446 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Distribution of Haemopis lateromaculata in North America, from the field collections in western North America and from the Orma J. Smith Museum of Natural History (ALBRICIDA), Canadian Museum of Nature (CMNA), and the United States Museum of Natural History (USNM). (1) In poor condition or not found in the collection. (2) At least two Moose Rivers in British Columbia. (3) Haemopis unidentified. PH, Peter Hovingh collection work in progress. Province or State Alberta Alberta British Columbia British Columbia British Columbia British Columbia British Columbia British Columbia Manitoba Manitoba Manitoba Newfoundland Newfoundland Ontario Ontario Ontario Ontario Ontario Ontario Ontario Ontario Ontario Saskatchewan Yukon Territory Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska Alaska California Colorado District of Columbia Idaho Michigan Missouri Nevada New Mexico Catalogue MNA 1978-0426 MNA 2006-0034 MNA 1985-0289 MNA 1990-0056 MNA 2006-0036 MNA 2006-0037 SNM 19231 SNM 38352 MNA 1985-0301 MNA 1985-0292 SNM 38347 MNA 1982-0654 MNA 1985-0378 MNA 1985-0259 MNA 1985-0262 MNA 1985-0253 M M Aeoeeoee@e G©@ NA 1985-0279 NA 1988-0036 MNA 1988-0163 MNA 1987-0670 USNM 38348 USNM 38360 USNM 38353 CMNA 2006-0035 CMNA1982-0746 CMNA 1985-0297 CMNA 1978-0360 USNM 19230 USNM 21225 USNM 20732 USNM 21256 USNM 43455 PH 1671-1677 PH 1660 PH 1661-1664 PH 1667, 1669 PH 1665, 1666 PH 1668, 1669 PH 2029 PH (0000) PH (0000) USNM 30223 PH (0000) USNM 38331 USNM 20681 USNM (PH 848) USNM 50192 USNM (PH 852) PH (0000) USNM 38308 USNM 38624 USNM 38627 USNM 39056 USNM (PH 497) USNM (PH 498) USNM (PH 499) USNM (PH 489) USNM (PH 491) USNM (PH 487) USNM (PH 488) USNM (PH 486) AOeeee eee Date 1907 Locator Beaver Lake Slave River drainage 150 Mile House Cariboo District Chilcotin River drainage(1) Haida Gwaii North Fork Moose River (2) [Dee] “D” Lake Vita LaSalle River Placenta Bay St Marys Bay Abitibi River Missinaibi River Ottawa River Albany River Thunder Bay Rainy River Kenora Lake Abitibi Lake Nipissing Waskasov Pelly River drainage Esther Island Hinchinbrook Island (3) Chuckagof Island (3) Lake Mc Donald, Yes Bay Lake Mc Donald, Yes Bay Afognak Island Sanak Island Revillagigedo Island Kodiak Island Cook Inlet Cook Inlet Kenai Peninsula Kenai Peninsula Kenai Peninsula Cold Bay, Alaska Peninsula Pit River Great Plains Easter Branch Snake River drainage Mecosta County Sikeston Ruby Marsh Pecos River Catlow Valley Klamath Lake Provo Mapleton Provo Provo River Provo River Provo River Sevier River Sevier River Weber River Weber River Bear River 2006 TABLE |. continued HOVINGH: HAEMOPIS LATEROMACULATA IN NORTH AMERICA 447 Province or State Catalogue Virginia USNM 42650 Wyoming PH (0000) Wyoming ALBRICIDA 61050 Wyoming ALBRICIDA 61051 Wyoming ALBRICIDA 61052 Identification of leech specimens: external characteristics TEETH. All of the Haemopis lateromaculata con- tained teeth (denticles). This characteristic would sep- arate these specimens from H. plumbea and H. grandis. Four populations (Utah, Nevada, Wyoming, Montana) of H. marmorata were without teeth. One population (Utah) was collected in 1941 (USNM 50/95) and again in 1986 suggesting that the loss of denticles may be due to environmental or genetic factors resulting from col- onization after the dessication of Lake Bonneville after 13 000 years ago. POSTERIOR SUCKER. Although the posterior sucker is large (about % the width of the body) in Haemopis lateromaculata and small (less than % the width of the body) in H. marmorata (Klemm 1985), the state of body relaxation was highly variable. This variability made the relative size of the posterior sucker to body width rather arbitrary. PIGMENTATION. The dorsal surface of Haemopis lat- eromaculata was patterned with darker blotches or mottled, or large to small speckles, of frequent to rare numbers. The lateral margins were cream or yellow (shared with H. terrestris) and interrupted with irreg- ular dark intrusions dorsally (a very diagnostic charac- ter), visible from both the dorsal and ventral surface [specimens from 14 sites: the Great Basin (8), Col- orado River drainage (2), Great Plains (1), Columbia River drainage (2), and the Yukon River drainage (1)]; or with the lateral margins the same shade as the ven- trum, without cream or yellow coloring [specimens from 21 sites: the Great Basin (2), Pacific Coast drain- ages (2), Fraser River drainage (1), Mackenzie River drainage (1), coastal islands and their adjacent shores of British Columbia and Alaska (17)]. The external characteristic that identified all of the specimens was the plain ventral surface. Mathers (1963: page 170) noted “with a few indistinct black and yellow blotches” on the ventral surface. I observed one specimen with an occasional speckle on the ventral surface. The pigmentation of H. marmorata in western North America ranged from heavily blotched or mottled to heavily speckled or lightly speckled on both the dor- sum and ventrum (61 sites), with the ventrum having subdued pigmentation of the same pattern. If either H. lateromaculata or H. marmorata were of the light- and/ or dark-colored immaculate phase (Klemm 1985: page 155), pigmentation could not be used to identify these two species. The light-colored immaculate phase was not found in the living state in western North America, but only in preserved specimens in museums. The lack Date Locator 1957 Dismal Swamp 199] Snake River drainage 1994 Johnson County 1997 Crook County 1997 Niobrara County of a dorsal stripe excluded H. terrestris, H. kingi, and H. septagon from further consideration in this study. MAXIMUM LENGTH (MM). Haemopis lateromaculata: Great Basin (90), Colorado River drainage (67), Colum- bia River drainagae (31), Mackenzie River drainage (83), Fraser River drainage (30), Queen Charlotte Is- lands (81), Yukon River drainage (82), Alaska (112), and Great Plains (50). Haemopis marmorata: Great Basin (112), Colorado River drainage (95), Columbia River drainage (121), Pacific Coast drainages (98), Fraser River drainage (98), and Great Plains (94). Length is highly variable, dependent upon both the age of living specimens and their state of relaxation dur- ing fixation. GONOPORE POSITIONS. Haemopis lateromaculata: Male gonopore XI b6 anterior 4; female gonopore XII b6 anterior 4, as determined on two specimens with developed clitellum from Nevada and Yukon Territory. All gonopore positions were separated by 5 annuli and occurred on the anterior 4 of the annulus. Haemopis marmorata: Male gonopore XI b6 anterior 4; female gonopore XII b6 anterior 4, as determined on three specimens with developed clitellum from Nevada and Colorado. Most gonopore positions were separated by 5 annuli and occurred on the anterior 4 of the annulus. Three exceptions: male XI 66 mid-annulus and female XII 56 mid-annulus, and male XI 66 mid-annulus and female XII b5 posterior 4 both from Arizona; and male and female on mid-annulus from British Columbia. Sawyer (1972) noted additional variations of the gono- pores from mid-annulus to the furrow. The male and female gonopore separations excluded H. septagon from further consideration during this study. Identification of leech specimens by reproductive organs Figure 2 illustrates the mean measurements of the reproductive organ position and Table 2 lists the mean + standard deviation. The posterior position of the left and right ejaculatory bulbs, the posterior position of the penis sheath loop and the vagina organs, and the ante- rior position of the prostate organ (anterior end of male atrium) were selected for their defined shape, largely attributed to the muscular nature of the organs (Rich- ardson 1969). The organ positions were determined by vertical positioning of the organ with the annulus, and hence counting the number of annuli to the annu- lus with the female gonopore. There is a possible error of annuli count of +1 annulus due to the positioning in individual specimens. The nomenclature of the organs follows that of Klemm (1985). Left and right positions are determined from the dorsal side. Comparison to other species is based on literature (Klemm 1985; Mann 1954; Richardson 1971). PENIS SHEATH LOOP. Haemopis lateromaculata: The penis sheath loop was located at annulus XIII b5 (XI b6 to XIV a2). The penis sheath was generally U- shaped and distinguished H. lateromaculata from H. marmorata, H. plumbea, H. terrestris, H. septagon, and H. caballeroi. Haemopis marmorata: The penis sheath loop was located at annulus XVII b/ (XVbI to XVIII 5S), a range comparable to Sawyer (1972). The penis sheath was generally J-shaped and on the left side. The ranges did not overlap between H. lateromacula- ta and H. marmorata. The average length of the penis sheath of H. lateromaculata was 21 annuli and of H. marmorata 38 annuli between the male gonopore and the prostate gland. Abnormal variations for the penis sheath of H. lateromaculata include position on the right side (4 specimens from Yukon Territory, Alaska, and Colorado) and highly contorted (1 specimen from Alaska). Abnormal variations for the penis sheath of H. marmorata include: positioned on the right side (11 specimens from Great Basin, Colorado River Basin, and Great Plains), and highly contorted (6 specimens from Great Basin, Great Plains) with one specimen in which the terminal loop turns medially and is twisted to form a topless “8”. PROSTATE GLAND. Haemopis lateromaculata: The anterior end of the male atrium is covered by the pros- tate gland and was located at annulus XI a2 (X a2 to XII 56), positions that are anterior to the female gono- pore and for the most part anterior to the male gono- pore. These locations are associated with the U-shaped penis sheath. Haemopis marmorata: The anterior end of the prostate gland was located at annulus XIV b5 (XIII b/ to XVI 55), defining the J-shaped penis sheath. EJACULATORY BULBS. Haemopis lateromaculata: The posterior end of the left ejaculatory bulb (seminal vesi- cle) was located at annulus XIII b2 (XII b2 to XIV THE CANADIAN FIELD-NATURALIST Vol. 120 a2) whereas the posterior end of the right ejaculatory bulb was at annulus XIII b/ (XII b/ to XIV a2). Al- though the locations show left-right symmetry (Figure 2), large variations were found in which the left bulb was 10 annuli posterior to 7 annuli anterior of the right bulb. The distribution of the positions of the ejaculato- ry bulbs was: right anterior to the left, 13 specimens; right even with the left, 13 specimens; and right pos- terior to the left, 8 specimens). Haemopis marmorata: The posterior end of the left ejaculatory bulb was locat- ed at XIV b2 (XIII b/ to XV b5) whereas the posterior end of the right ejaculatory bulb was at annulus XIV b/ (XII a2 to XV b6). The left bulb ranged from 9 annuli posterior to 5 annuli anterior of the right bulb. The distribution of the ejaculatory bulbs was: right anteri- or to the left, 19 specimens; right even with the left, 16 specimens; and right posterior to the left, 17 spec- imens), a pattern noted by Sawyer (1972). Two speci- mens had only one ejaculatory bulb. VAGINA. Haemopis lateromaculata: The posterior end of the vagina (vaginal bulb) was found at XIV b6 (XIII a2 to XVII a2). The vagina was always posterior to the posterior reach of the penis sheath loop and was generally in the median region. Haemopis marmorata: the posterior end of the vagina was found at XVI b/ (XIV b2 to XVII a2), comparable to the findings of Sawyer (1972). The vagina was always anterior to the posterior reach of the penis sheath loop and posterior to the anterior end of the prostate gland, and generally located in the medial region. Three specimens had the vagina location anterior to the anterior end of the pros- tate gland. The position of the vagina of H. caballeroi had a relative position to the penis sheath loop and the prostate gland (atrium) as in H. marmorata, but the two specimens of H. caballeroi were highly contracted and hence not typical (Richardson 1971). The extensive penis sheath and its relation to the vagina suggests that these two species deserve a separate genus — in TABLE 2. Annuli location of selected reproductive organs in Haemopis marmorata and H. lateromaculata. Annulus X b/ was number | and annuli posterior to this annulus were sequentially numbered. Organ nomenclature was taken from Klemm (1985). Figure 2 illustrates the organs and the points of measurement. H. lateromaculata N=31 Penis sheath loop Mean + standard deviation 18.8 + 2.6 Range 1493 Anterior prostate gland Mean + standard deviation DIED ES Range 2-14 Posterior left ejaculatory bulb Mean + standard deviation 16.7 + 3.0 Range 11-23 Posterior right ejaculatory bulb Mean + standard deviation Sys) se Stl Range 10 — 23 Posterior vagina Mean + standard deviation 24.6 + 4.7 Range 17-38 H. marmorata N=55 35.8 + 4.6 25 —44 23.9 + 3.7 16 —35 2006 36 EAN ine ran er HOVINGH: HAEMOPIS LATEROMACULATA IN NORTH AMERICA 449 FiGurE 2. Measured reproductive organs in Haemopis lateromaculata (A) and H. marmorata (B), viewed from the dorsum. The segments are shown in Roman numerals starting with segment X and extend to segment XVIII. Each segment has five annuli as denotated names shown on the right side of segment X. For the purposes of this paper, the first annulus is numbered 1| and is sequenced posteriorly as noted on the left side with only the first annulus of each seg- ment numbered (see Table 2). Male gonopore on IX b6 and female gonopore on XII 56. Denotations: Elf, posterior reach of the left ejaculatory bulb; Ert, posterior reach of the right ejaculatory bulb; P, posterior reach of the penis sheath loop; V, posterior reach of the vagina; and A, the prostate gland at the anterior end of the male atrium. The vertical lines represent one standard deviation (thick lines) and total range (thin lines) (see Table 2). this case Percymoorensis, if one follows the distinct genera classification in Erpobdellidae of Nephelopsis, Mooreobdella, and Erpobdella, each having morpho- logical distinct male atriums and sperm ducts. Discussion Haemopis lateromaculata is now considered to be widely distributed throughout North America; prior to this study, its known distribution had been limited to two states (Figure 1). The presence of this leech on the Alaskan Peninsula and adjacent islands, the Cook Inlet region, the Alexander Archipelago of southeast Alaska, and Haida Gwaii (Queen Charlotte Islands) of British Columbia suggest that these leeches occupied off-shore coastal refugia during the Pleistocene and were sepa- rated from those populations south of Canada’s cordil- 450 leran and continental glaciers. These northwest refugia were limited by extensive glaciers from Haida Gwaii to Umnak Islands (Coulter et al. 1965; Clague 1989; Kaufman and Manley 2004) with coastal cordilleran glaciers occurring to 12 000 years ago (Peltier 1994; Dyke 2004; Fulton et al. 2004). While the occurrence of small terrestrial refugia have been postulated from Kodiak to the Queen Charlotte Islands (Heusser 1989; Ramsey et al. 2004), present day freshwater habitats (ponds, lakes, slow moving and meandering streams) from which leeches have been collected add a new dimension to the terrestrial refugia. The Pleistocene-Holocene environments were chang- ing due to eustasy (the sea level rising at least 120 m) and glacio-isostatic surface adjustments involving the advance and recession of the glaciers. Hetherington et al. (2003, 2004) mapped this environment for the Haida Gwaii and described a land bridge between the islands and the British Columbia mainland that existed between 11 700 and 11 200 years ago as a result of the presence of a forebulge from the retreating cordilleran glacier. This process, if applied throughout the glaciated region of coastal Alaska, may have allowed island by island transfer of leeches from coastal Pleistocene refugia, thus accounting for the modern day presence of H. lateromaculata populations on the coastal northwest- ern North America mainland (Reimchen and Byun 2005). Upstream movement of H. marmorata has been noted (Herrmann 1970; Richardson 1942; Sawyer 1970), and active movement (to Prince Edward Island) must have occurred by direct means, and not as pas- sive movement by birds or other agents (Richardson 1943). It is assumed herein the cordilleran glaciers and marine habitats prevented any leeches from continen- tal populations colonizing the coastal habitats. Lukin (1976) suggested that Haemopis originated in the Nearctic and penetrated the Palaearctic from the northeast [Siberia] when northern Asia had a warmer climate. He further suggested that H. sanguisuga emerged at that time and is now found across central Eurasia to the Amur Basin and near Vladivostok in east- ern Asia, but not in Kamchatka Peninsula. The center of biodiversity of Haemopis is in eastern North Amer- ica, supporting Lukin’s postulate. Only two species (H. lateromaculata and H. marmorata) moved out of this region to occupy most of North America, and only H. lateromaculata has been identified in Alaska and the coastal islands, suggesting range expansion in an earli- er epoch before the range expansion of H. marmorata. From the Late Cretaceous (80 million years ago) to the middle Pliocene (3 million years ago), North America and eastern Asia were connected between Alaska and eastern Siberia (Repenning and Brouwers 1992; Smith et al. 1994). For much of this time, the climate in this region was comparable to that of the southeastern Unit- ed States today (Bassinger 1991). I propose that H. lateromaculata and H. sanguisuga are sister species. Two anatomical features of H. lat- THE CANADIAN FIELD-NATURALIST Vol. 120 eromaculata and H. sanguisuga support this relation- ship: (1) the vaginal bulb lies posterior to the penis sheath loop (shared with H. grandis and H. kingi) and (2) the prostate gland (anterior end of the male atrium) occurs mostly anterior to the male gonopore (shared with H. kingi and H. caeca) (Mann 1954; Klemm 1985; Manoleli et al. 1998). Note that H. kingi occurs in both groups, a confirmation of Richardson’s (1969, 1971) views that H. kingi and H. lateromaculata are closely related. The distribution of a species is as important as the original description of a species. As noted by Newton (2003: page 90) with respect to birds, “how species are formed is central to understanding their distributions” and “genealogy and distribution are not separate issues: they are simply different sides of the same biological coin”. This paper contributes to the definition of H. lat- eromaculata by the addition of geographical limits of its distribution and its evolutionary implications. Acknowledgments I thank Jacqueline Madill and Jean-Marc Gagnon at the Canadian Museum of Nature, Bill Moser at United States National Museum (Smithsonian Institution), and John Keebaugh at the Orma J. Smith Museum of Nat- ural History for assistance and for providing facilities at these museums which greatly contributed to this study; Robert Wisseman (Aquatic Biology Associates) and Kurt King (Wyoming Department of Environmental Quality) for furnishing leeches for identification from western United States. Early versions of this document were reviewed by Bill Moser and Donald Klemm, and Jacqueline Madill and Mark J. Wetzel reviewed the manuscript for the journal. I thank these reviewers for their suggesting and improvements. Literature Cited Bassinger, J. F. 1991. 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Received 17 January 2006 Accepted 21 April 2007 Predation on Artificial Nests of Northern Bobwhites, Colinus virginianus, by Mammalian Mesopredators: Does the Problem-Individual Paradigm Fit? J. B. JENNiNGs!*, M. L. KENNEDY!, A. E. Houston”, and B. D. CARVER? ‘Ecological Research Center and the Department of Biology, The University of Memphis, Memphis, Tennessee 38152 USA ?Ames Plantation, The University of Tennessee, P. O. Box 389, Grand Junction, Tennessee 38039 USA 3Department of Biology, Freed-Hardeman University, Henderson, Tennessee 38340 USA 4Department of Natural Sciences, Unit 208, Southwest Tennessee Community College, 737 Union Avenue, Memphis, Tennessee 38103 USA Jennings, J. B., M. L. Kennedy, A. E. Houston, and B. D. Carver. 2006. Predation on artificial nests of Northern Bobwhites, Colinus virginianus, by mammalian mesopredators: Does the problem-individual paradigm fit? Canadian Field-Naturalist 120(4): 452-456. Using mark/recapture procedures, predation on artificial nests of Northern Bobwhites (Colinus virginianus) by mammalian mesopredators (Raccoons, Procyon lotor; Virginia Opossums, Didelphis virginiana; and Striped Skunks, Mephitis mephitis) was assessed in relation to the “problem-individual” paradigm. The paradigm, which is untested among mammalian mesopredators, predicts that most predation on a prey species is by a small number of individuals repeatedly involved. By examining number of captures and recaptures on artificial nests during non-nesting and nesting periods in 2000-2003, predation within and among species were gauged. Results varied by species, sampling period, and year; however, predation was by a small percentage of individuals and only within the population of Striped Skunks were individuals (2 of 49) captured on an artificial nest more than once. Raccoons, Virginia Opossums, and Striped Skunks were responsible for 10, 2, and 12% of the loss of all nests avail- able to predators, respectively. Based on low occurrences of individuals repeatedly involved in predation on nests, the problem- individual paradigm was unsubstantiated. Although Raccoons, Virginia Opossums, and Striped Skunks (as individual predators) had only moderate impact on the population of Northern Bobwhites, they had a much greater impact collectively (preying upon 24% of all nests available). To maximize Northern Bobwhite success, the most productive management is probably best directed toward a vertebrate guild that includes mammalian, avian, and reptilian predators, and management strategies that use multiple techniques. Key Words: Raccoons, Procyon lotor, Striped Skunks, Mephitis mephitis, Virginia Opossums, Didelphis virginiana, Northern Bobwhites, Colinus virginianus, artificial nests, problem individuals, predation, Tennessee. Due to several factors (e.g., release of predators and reduced hunting; Young and Ruff 1982; Amarasekare 1993; Mosillo et al. 1999), populations of Raccoons (Procyon lotor), Virginia Opossums (Didelphis virgini- ana), and Striped Skunks (Mephitis mephitis) have in- creased in abundance throughout much of their range. These increases have caused widespread removal to be attempted as one means of controlling populations of these species (United States Department of Agriculture 2004"). Linnell et al. (1999), in discussing carnivore- livestock conflicts, pointed out that individuals or demographic groups within a predator population can show different behavioral traits where only a small pro- portion of individuals is accountable for taking sig- nificant numbers of a prey species. They refer to the phenomena as the “problem-individual” paradigm. Al- though this paradigm appears to fit some larger carni- vores (Rabinowitz 1986; Ross et al. 1997; Sacks et al. 1999; Stahl et al. 2001), the hypothesis rarely has been tested (Linnell et al. 1999). Presently, the existence of individuals or a demographic group within mid-sized mammalian predators that prey upon a disproportionate number of nests of ground-nesting birds is unknown. Different rates of predation, by individuals or species, could allow for targeted removal of individuals rather than widespread removal as a means of controlling nest predation which may have broad conservation and management application as fewer individual predators would need to be removed. Mammalian mesopredators are among the known predators of Northern Bobwhites (Colinus virginianus; Leimgruber et al. 1994; Fenske-Crawford and Nieme 1997), and populations of this game bird have declined throughout most of its range (Brennan 1991). However, causes of mortality for Northern Bobwhites are not well understood (Burger et al. 1995), and rates of predation within and among species of mammalian predators on Northern Bobwhites are uncertain. Additionally, mid- sized mammalian predators have been successfully captured and recaptured in live traps (Johnson 1970; Baldwin et al. 2004) and have been shown to prey upon artificial nests of Northern Bobwhites (Leimgruber et al. 1994; Fenske-Crawford and Niemi 1997); therefore, they make a good model for assessing the “problem- individual” paradigm using capture/recapture proce- dures. ; The purpose of the present study was to examine the “problem-individual” paradigm in light of predation by populations of mammalian mesopredators on artificial nests of Northern Bobwhites. Specifically, the predic- tion was tested that predation on artificial nests of Northern Bobwhites by Raccoons, Virginia Opossums, 452 2006 and Striped Skunks is not widespread within and among species. This work should assist in developing sound conservation and management programs for Northern Bobwhites and control of mammalian meso- predators. Methods This investigation was conducted on the Ames Plan- tation (Ames; 35°4'N, 89°13'W), an agricultural exper- iment station of The University of Tennessee and home to the National Field Trial Championship for bird dogs, located in Hardeman and Fayette counties, Tennessee. Ames was approximately 7500 ha and consisted of a mosaic of habitat types that included agricultural fields, pastures, upland hardwoods, bottomland hardwoods, and pine plantations. On the site, about 2000 ha were managed extensively for Northern Bobwhites. The site provided sufficient bird populations (autumn densities of ca. 2-3 birds/ha; Seckinger 2004) and associated predator suite (Leberg and Kennedy 1988; Baldwin 2003; Baldwin et al. 2004) to allow study of a ground nesting bird. For a more detailed description of Ames see Baldwin et al. (2004). Two study sites (separated by 1.6 km) were sampled. Each was 1.6 km? and was similar in habitat (frag- mented, upland hardwood forest interspersed with early successional fields). Because results obtained at each site were similar, results are combined and are present- ed here. Trapping during the non-nesting season (Octo- ber-December 2000-2002) was conducted using Rac- coon-size live traps (81.3 x 25.4 x 30.5 cm; Tomahawk Live Trap Co., Tomahawk, Wisconsin, or Havahart Live Traps, Woodstream Corporation, Lititz, Pennsylvania), which were baited with cat food and placed in an 8 x 8 configuration with traps spaced at about 230-m inter- vals. Sampling was conducted for 32 nights/season (for a total of 2000 trap nights/season). Once captured, sex and age were determined, and individuals were tagged (National Band and Tag Company, Newport, Kentucky) in both ears. Additionally, to confirm that individuals remained on the study site during the nesting season, 17, 3, and 3 Raccoons; 10, 9, and 5 Virginia Opossums; and 7, 8, and | Striped Skunks were fitted with radio- collars in 2000, 2001, and 2002, respectively. Animals were verified as being on the study site using a three- element hand-held antenna and a portable receiver. Telemetry equipment was from Advanced Telemetry Systems, Inc., Isanti, Minnesota. All individuals were released at the site of capture. Work during this period provided a known number of marked individuals that could be compared to those individuals captured dur- ing the nesting trapping season. Results obtained during the nesting trapping season (June-September 2001-2003) were derived using arti- ficial nests (10 cm wide by 6 cm deep) and baited with two eggs of Northern Bobwhites (Hernandez et al. 2001). The nest itself, constructed from wicker, was commercially purchased. Before placement, nests were JENNINGS ET AL: PREDATION ON ARTIFICIAL NESTS 453 left outdoors in an undisturbed state for 5-7 days to minimize human odors as suggested by Donalty and Henke (2001). Predators can cue on human odors at artificial nests (Whelan et al. 1994), but this can be min- imized by rain or scents (Donalty and Henke 2001). To address this concern, eggs were rinsed with distilled water and handled with rubber gloves when placed on nests (Small and Hunter 1988). By comparing individ- uals of the known population (non-nesting season cap- tures) that also were captured on artificial nests during the nesting season, the percentage of the population actively involved in predation on artificial nests was determined. Artificial nests were placed inside live traps. At each site, 80 traps were established; 50 were in actual use at any one time. Traps were placed either selectively or randomly which resulted in an irregular distribu- tion over each of the 1.6 km? study areas. Because of this placement of traps, artificial nests occurred in a variety of habitat types which can be typical of North- ern Bobwhites (Stribling 1996). Traps that were with- out nests were wired open. All traps were camouflaged with available vegetation. Once a nest was preyed upon, another trap (randomly or selectively placed) in anoth- er location had an artificial nest placed inside. Traps with a preyed-upon nest were closed and no longer used. This provided 50 nests on each of the two sites to be in continuous use. Traps in operation were open for 90 nights/season (for a total of 9000 trap nights/season), and were checked daily. To minimize animals associat- ing traps with nests, 25 pseudo-nests were established on each site. These were made of chicken wire in the shape and size of the actual traps used to contain artifi- cial nests. Pseudo-nests were not baited and were ran- domly placed in the study areas. Results For comparisons between nesting and non-nesting seasons, radiotelemetry procedures confirmed that 9 Raccoons, | Virginia Opossum, and 2 Striped Skunks, which were radiocollared during the non-nesting sea- son (2000), were present during the nesting-trapping season (2001). Three Raccoons collared in the non- nesting sessions (2000) along with 1 Striped Skunk were present on the study area for the nesting-trapping session (2002). There also was one Raccoon and one Striped Skunk collared during the non-nesting-trap- ping session (2002) present on the study area for the nesting-trapping session (2003). Capture data derived from the two sampling periods varied somewhat by species, year, and sampling period (Table 1). Sampling during the non-nesting period re- vealed the presence of all target species, with the great- est number of individuals varying by species each year. Sampling during the nesting period also showed pres- ence of all targeted species during each year but in smaller numbers and with less variation in data across years than observed in the non-nesting period. 454 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE |. Capture data derived in a study of predation by mammalian mesopredators (Raccoons, Virginia Opossums, Striped Skunks) on artificial nests of Northern Bobwhites during two trapping sessions (non-nesting, nesting) on the Ames Plantation in western Tennessee 2001-2003. C = number of individuals captured; R = number of individuals recaptured at least one time during the same trapping session of the same year; %R = percentage of marked individuals recaptured (R/C x 100); M = number of individuals captured > 2 times; %COL = the percentage of radio-collared animals captured (number of collared animals recaptured/total number collared x 100%); T = number of individuals captured during the non-nesting period that were recaptured during the nesting period; %K = percentage of individuals captured during the non-nesting period that were recap- tured during the nesting period (T/C of the Non-nesting period x 100%). Nesting Non-nesting Captured during (October-December) (June—September) both seasons Year Species C R %R M %COL GIR Gar IM Gao al %K 2001 Raccoon 41 24 3) © 18 13) OQ" @ 0 11 4 10 VirginiaOpossum 33 = 13 39 8 10 3 0 O 0 0 0 0 Striped Skunk 12 5 42 4 43 22 Ae’ 5 if 0 1 8 2002 Raccoon 19 8 42 1 0 14 0 O OSS) 0 0 Virginia Opossum 19 9 47 6 0 3 ORO 0 0 0 0 Striped Skunk DAT 2 SO 7 0 11 1 9 0 0 1 4 2003 Raccoon 38) le 45 4 0 lO OO 0 0 3 8 Virginia Opossum 89 24 27 9 20 4 0 O 0 0 1 1 Striped Skunk 46 $18 39 8 0 1610. .O 0 0 0 0 ' This reflects predation by the same individual. The total number of nests available to predators was 433. Raccoons were responsible for 43 nests preyed upon (10% of all nests available to predators). Virginia Opossums were responsible for predation on 10 nests (2%), and Striped Skunks destroyed 52 nests (12% of all nests available). Collectively, these predators preyed upon 24% of all available nests. No significant pattern existed in the age or sex of the predators involved in predation on artificial nests. Discussion Results of this study documented that at most only 10% of a known predator population (Table 1) was actively involved in predation on artificial nests of Northern Bobwhites. Additionally, individual predator species were involved in predation on artificial nests to varying levels (Table 1). Based on these results, the prediction that predation on artificial nests of North- ern Bobwhites by Raccoons, Virginia Opossums, and Striped Skunks is not widespread within and among species, is supported, and the “problem-individual” par- adigm is unsubstantiated. All predators assessed had members captured multiple times during the non-nesting season. These results were similar to those reported in other studies of mesopredators conducted during autumn and winter (non-nesting seasons) in the region (Ladine 1995; Baldwin 2003). Such results demonstrate that individuals can be captured and recaptured in suitable numbers to explore the “problem-individual” paradigm. However, only two Striped Skunks were captured on artificial nests more than once. Given that these indi- viduals represented only 2 of 49 Striped Skunks cap- tured and were only captured 2 and 3 times, respective- ly, in the present study and that Vickery et al. (1992) reported predation by Striped Skunks on nests of birds was not targeted, these results do not corroborate the “problem-individual” paradigm. Additionally, in com- parison with larger mammalian predators, this para- digm has been associated, generally, with much higher levels of predation and by greater numbers of individ- uals repeatedly involved in taking prey (Claar et al. 1986; Stander 1990; Ross et al. 1997; Sacks et al. 1999; Stahl et al. 2001). Scalet et al. (1996) pointed out that predator-prey interactions influence both prey and pred- ator populations. However, based on results of the pres- ent study, these interactions appear to be complex and differ among mammalian taxa. Lack of support for the “problem-individual” para- digm among target predators could be explained by distribution of nests of Northern Bobwhites on the land- scape and the predators involved. Stribling (1996) noted that nests of Northern Bobwhites are located in a number of habitat types. Linnell et al. (1999) suggest- ed that a scattered distribution of prey, such as the dis- tribution of artificial nests in this study, would increase encounter rates by predators without any search be- havior required, and hypothesized that, under these con- ditions, problem individuals are less likely to appear because most individuals have opportunity to take the target prey without developing specialized behaviors. Raccoons, Virginia Opossums, and Striped Skunks are omnivorous and opportunistic predators with strong preferences for seasonally available foods (McManus 1974; Gardner 1982; Godin 1982; Kaufmann 1982; Schwartz and Schwartz 2001). Wiens (1976) discussed population responses to patchy environments and noted the importance of prey density to responses of popula- tions to patchiness. It appears likely that the habitat usually occupied by Northern Bobwhites (forest open- ings, open woods, fallow fields, and edges of cultivat- ed fields; Stribling 1996) provides favorable and abun- dant foods (e.g., insects and other invertebrates, wild fruits, and small mammals; Merritt 1987; Schwartz and Schwartz 2001) for mammalian predators. Given that 2006 number of individuals known alive for each species of predator was greater on the study site during the non- nesting period than the nesting period, as well as over- all recapture rate and number of individuals captured more than 2 times (Table 1), it seems that the presence of eggs of Northern Bobwhites and Northern Bobwhites themselves had no unusual attraction for mesopreda- tors. Neither Raccoons nor Virginia Opossums had a significant affinity for fields, which are managed inten- sively for Northern Bobwhites on Ames, and, because of the lack of individuals recaptured on artificial nests, the problem-individual hypothesis is not supported for these species. However, based on the number of cap- tures in the present study, there was a distinct associ- ation between fields and Striped Skunks, which reflects a preference for this habitat (Nowak 1991). These habi- tats also are good for Northern Bobwhites throughout the year and not just during their breeding season (Stri- bling 1996). These mesopredators apparently used the study site and adjacent areas during the nesting peri- od of Northern Bobwhites for foods other than those associated with Northern Bobwhites. Being opportunistic, mesopredators may prey upon nests of Northern Bobwhites when they encounter them. Because of increased protein intake during formation and laying of eggs (Rosene 1969), eggs and Northern Bobwhites themselves represent foods of high nutri- tional value. However, because Northern Bobwhites on Ames provide only a limited source of food, opti- mal foraging is likely focused on more abundant and easily captured food items. Eggs and birds probably serve as a valuable supplemental food to more abundant food items. For example, Vickery et al. (1992) reported a positive correlation between predation by Striped Skunks on bird eggs and nestlings when that species was foraging for invertebrates. Such foraging patterns could partly explain reports that birds are preyed upon by Raccoons, Virginia Opossums, and Striped Skunks, although their occurrence in diets of these predators is small (McManus 1974; Kaufmann 1982; Godin 1982: Schwartz and Schwartz 2001). In the present investigation, there was only one year where the most abundant species captured during the non-nesting season was responsible for the greater num- bers of artificial nests preyed upon during the nesting- trapping season (Table 1). It appears that Raccoons, Virginia Opossums, and Striped Skunks, as individual species, had only a moderate impact on artificial nests (Table 1). However, as a subset of the community or mesopredator guild—a collection of species that use common resources in similar ways (Root 1967)—they had a much greater impact. In fact, Nicolaus (1987) stated that Raccoons, along with other mammals and birds, belonged to a guild of nest predators. Of artificial nests preyed upon in this study, Raccoons, Virginia Opossums, and Striped Skunks accounted for 10, 2, and 12% of losses, respectively. Collectively, there was a 24% loss of artificial nests attributable to this meso- predator guild. JENNINGS ET AL.: PREDATION ON ARTIFICIAL NESTS 455 Overall, the pattern of predation on artificial nests of Northern Bobwhites is apparently driven by a larg- er vertebrate-predator guild similar to the granivore guild (birds, rodents, and insects) reported in deserts of the southwestern United States (Brown and David- son 1977). This vertebrate-predator guild, including both mammalian and reptilian predators, also has been documented by Staller et al. (2005). Because of the collective predation rates on artificial nests, most pro- ductive management (using multiple techniques; Jimi- nez and Conover 2001) for ground-nesting birds prob- ably is best directed toward a vertebrate guild that includes not only mammalian predators but avian and reptilian predators as well. Because of the dynamic nature of community interactions (Meffe et al. 1997), it is not likely that single actions will resolve complex predator-prey relationships, which reflect the dynamic nature of the ecological world. At present, interaction among mammalian mesopredators is uncertain (Ladine 1997; Kissell and Kennedy 1992) and the impact of large vertebrate-predator guilds on prey species remains unclear. Future investigations are needed to more clear- ly understand these ecological issues. Acknowledgments Thanks are extended to T. L. Best, S. B. Franklin, J. W. Grubaugh, and B. A. Simco who provided a critical review of an earlier draft of the manuscript. R. A. 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Population responses to patchy environ- ments. Annual Review of Ecology and Systematics 7: 81- 120. Young, B. F.,, and R. L. Ruff. 1982. Population dynamics and movements of black bears in east central Alberta. The Journal of Wildlife Management 46: 845-860. Received 3 January 2006 Accepted 17 February 2007 Reactions of Narwhals, Monodon monoceros, to Killer Whale. Orcinus orca, Attacks in the Eastern Canadian Arctic KRISTIN L. LAtpRE!*, MADS PETER HEIDE-JORGENSEN2Z, and JACK R. ORR? 'Polar Science Center, Applied Physics Laboratory, University of Washington, 1013 NE 40" Street, Seattle, Washington 98105 USA *Greenland Institute of Natural Resources, Box 570, DK-3900 Nuuk, Greenland Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Canada Laidre, Kristin, L., Mads Peter Heide-Jgrgensen, and Jack Orr. 2006. Reactions of Narwhals, Monodon monoceros, to Killer Whale, Orcinus orca, attacks in the eastern Canadian Arctic. Canadian Field Naturalist 120(4): 457-465. A Killer Whale attack on Narwhals was observed at Kakiak Point in Admiralty Inlet, Nunavut, Canada, in August 2005. Behavioral responses of both Narwhals and Killer Whales were documented by direct observation. Data collected from Narwhals instrumented with satellite-linked transmitters 5 days prior to the arrival of Killer Whales were used to examine changes in Narwhal movement patterns (e.g., dispersal and clumping) five days before the attack, during the attack, and five days after Killer Whales left the area. A minimum of four Narwhals were killed by 12-15 Killer Whales in a period of 6 hours. Narwhals showed a suite of behavioral changes in the immediate presence of Killer Whales including slow, quiet movements, travel close to the beach (<2 m from shore), use of very shallow water, and formation of tight groups at the surface. These behavioral changes are consistent with Inuit accounts of Killer Whale attacks on Narwhals. During the attack, Narwhals dispersed broadly, the groups were less clumped (standard deviation of inter-whale mean latitudes and longitudes), Narwhal space-use doubled from pre-attack home ranges of 347 km? to 767 km? (kernel 50% probability), and Narwhals shifted their distribution further south of the attack site. After the disappearance of Killer Whales, north-south dispersal of Narwhals contracted and was similar to pre-attack levels, total space use decreased slightly (599 km), yet west-east dispersal remained high. Narwhals were distributed significantly (P < 0.001) more broadly offshore in areas not used before the occurrence of Killer Whales. In general, short-term reactions of Narwhals to Killer Whale presence were obvious; yet normal behavior (as observed from shore) resumed shortly after Killer Whales left the area. Long-term (five day) Narwhal behavioral responses included increased dispersal of Narwhal groups over large offshore areas. This is among the few reports of eyewitness Killer Whale attacks on Narwhals in the high Arctic and is the first time changes in Narwhal behavior have been documented in response to a predation event through the use of satellite telemetry. Key Words: Killer Whale, Orcinus orca, Narwhal, Monodon monoceros, predation, satellite telemetry, Admiralty Inlet, Nunavut, Canada. Killer Whales (Orcinus orca) occur widely in Arc- tic waters, yet their abundance, movements, site fidelity, and specific distribution are poorly known (Forney and Wade 2006). Generally, Killer Whales are sighted during ice-free months and often near areas with high densities of other marine mammals, such as summer- ing grounds for Belugas (Delphinapterus leucas) and Narwhals (Monodon monoceros). In the eastern Cana- dian High Arctic, this includes eastern Lancaster Sound and associated inlets or fjords around Baffin Island (e.g., Eclipse Sound, Admiralty Inlet, and Prince Regent Inlet) (Markham 1874; Reeves and Mitchell 1988). Few eyewitness accounts are available of Killer Whale attacks on Narwhals (Steltner at al. 1984). This is because Killer Whales occur over large, sparsely inhabited Arctic areas and the timing and location of attacks are unpredictable. Thus, the majority of des- criptions, both of Killer Whale attacks themselves and of the response of Narwhals, come from Inuit obser- vations (Degerbgl and Freuchen 1935; Freuchen and Salomonsen 1961) or from scattered opportunistic re- ports (Campbell et al. 1988). Generally these accounts made by locals are limited in that observations extend only as far as what can be observed from shore and generally cover short time scales (hours). Also, Inuit observations of Killer Whale occurrence or Killer Whale attacks may not always be reported. There is little information about the natural mortality of Nar- whals and generally, Killer Whales are assumed not to significantly depress Narwhal populations (Davis et al. 1980). Furthermore, the extent to which Killer Whales affect the behavior of Narwhals (e.g., move- ment patterns) is unknown. In August 2005, a Killer Whale attack on Narwhals was observed from a camp operating a month-long Narwhal satellite tagging operation in Admiralty Inlet, Canada. Several kills of Narwhals by Killer Whales were inferred from vigorous surface and diving activity of Killer Whales within large oiled areas on the sur- face and congregations of seabirds. In combination with eyewitness observations of behavior of Narwhals and Killer Whales, data collected from satellite tags deployed on Narwhals up to five days before Killer Whales arrived in the area were used to examine clus- tering and dispersal of Narwhals prior to and follow- ing the predation events. These are the first quantita- tive movement data obtained from Narwhals subjected to Killer Whale predation. 457 458 THE CANADIAN FIELD-NATURALIST Visual Observations of Attack Killer Whales were reported by Inuit hunters in a camp about 3 km north of Kakiak Point (72°31'N 86°38'W), in Admiralty Inlet, Nunavut, Canada, on 19 August 2005. On 20 August, we witnessed a Killer Whale predation event on Narwhals just offshore of Kakiak Point. Approximately 12-15 Killer Whales arrived in a single group at Kakiak Point at 12 noon local time. At least several hundred Narwhals had been seen daily in the area for at least two weeks preceding the arrival of the Killer Whales and also on the morn- ing of 20 August. On several occasions thousands of Narwhals had been seen moving past Kakiak Point over periods of 2-3 hours. The Killer Whale group consisted of one adult male, 7-10 adult females, and several juveniles of undeter- mined sex. Narwhals were attacked less than 1 km from shore. At least two Narwhals were killed during the first observed attack between 12:00-13:00 local time. After this event, the Killer Whales left the vicin- ity of Kakiak Point for approximately 3 hours and returned at 16:00, when at least two more Narwhals were killed ~0.5 km from the coast. Killer Whales were not seen or reported again in the area until 25 August. Active subsistence hunting for Narwhals was practiced at several hunting camps along the shore north and south of the research camp. The four independent kill events occurred over a 6 hour period in daylight hours. Narwhal kills were in- ferred from large oil/blubber slicks at the surface, con- gregations and diving of Fulmars (Fulmarus glacialis) in surface waters, and focal movements of Killer Whales diving in the center of oiled areas for 15-30 minutes at a time (Figure 1). No body parts or pieces of Nar- whals were directly seen. Narwhal Movements Narwhal satellite tagging operations were ongoing prior to the Killer Whales’ arrival at Kakiak Point. Nar- whals were captured using nets set perpendicular to the shoreline 50-100 m from shore (details described in Dietz et al. 2001) and instrumented with SPOT4 and SPOTS satellite tags made by Wildlife Computers (Redmond, Washington). Tags recorded daily geogra- phic positions based on Doppler shift of tag transmis- sions, received by polar orbiting satellites through Vol. 120 Service Argos (Harris et al. 1990). Transmitters were attached to Narwhals on the dorsal ridge with three 6-10 mm polyethylene pins. The Narwhals were usu- ally released after less than 30 minutes (Table 1). Only data collected from Narwhals instrumented prior to the Killer Whales’ arrival were used in the analysis to control for differences in sample size (num- ber of individuals monitored before and after the pre- dation event). Since the Narwhals were instrumented and released on different occasions, they were assumed to represent different groups or pods. Average daily geographic positions were calculated for each Narwhal based on good quality Argos locations (location quali- ty >=0). Average daily positions for each whale were averaged to create inter-whale means. The standard deviation of daily average inter-whale positions was calculated for each of three time periods: five days before Killer Whales arrived (14-18 August), during the immediate Killer Whale observations or attacks (19-20 August), and five days after the Killer Whale attack (21-25 August) when no Killer Whales were seen in the area. Narwhals were distributed along a north-south gradient along the west coast of Admiralty Inlet; therefore the variation in latitude was interpreted as coastwise dispersion and the variation in longitude was interpreted as inshore-offshore dispersion. Narwhal space-use patterns were quantified in each of the three time periods with a kernel home range polygon (prob- abilistic measure of space use) [Arc View 3.1 Environ- mental Systems Research Institute] based on average daily positions from each whale. Results Pre-attack Narwhal behavior (> 5 days) Narwhals observed passing Kakiak Point before the Killer Whale attack moved in groups of 3-8 whales at a distance of 20-200 m from the beach. Most whales were heading south. Between 14 and 17 August, seven Narwhals were instrumented with satellite tags before the Killer Whales arrived in the vicinity of Kakiak Point (Table 1). These Narwhals moved along the west- ern side of Admiralty Inlet south of Kakiak Point and less than 1-2 km from shore. No tagged Narwhals were located in the center or eastern side of the inlet and the linear home range along the west coast was <100 km. TABLE |. Seven Narwhals instrumented with satellite transmitters prior to Killer Whale occurrence and attack at Kakiak Point, Admiralty Inlet, Canada, 2005. Calf accompanying female on 17 September 2005 was not tagged. ID Tagging date Time of release Sex Body length (cm) 20685 14 September 2005 15:06 F 360 20686 14 September 2005 15:56 M 483 20689 17 September 2005 06:25 F 360 37235 17 September 2005 10:45 F + calf 358 37236 17 September 2005 21:20 F 380 37280 17 September 2005 QeDi, F 380 37282 17 September 2005 21:45 F 364 2006 —e — LAIDRE, HEIDE-JORGENSEN, and Ork: REACTIONS OF NARWHALS 459 ae SF Ficure 1. Oil film on the surface created by Killer Whales feeding on Narwhals in Admiralty Inlet, Canada, August 2005. Large aggregations of Fulmars appeared in the area shortly after each kill. Photo by M. P. Heide-Jorgensen. The standard deviation of average daily positions indicated that dispersal of Narwhal groups varied ap- proximately 19.9 km north-south, and 13 km west-east before Killer Whales arrived (Table 2). The core of the pre-attack kernel home range (50% probability area) was concentrated just offshore of Kakiak Point and distributed along the west coast of the inlet encom- passing 347 km? (Table 2). Immediate Reactions of Narwhals to Killer Whale Attack On 19 August, Killer Whales were reported by Inuit hunters north of the camp, yet no detailed observations of predation events or behavior were collected. On that day, we observed Narwhals passing very close to the shore (within the surf zone approximately 2-3 m from land) and remaining still at the surface in large groups. TABLE 2. Dispersal of Narwhals before, during, and after a Killer Whale attack in August 2005 as measured by the distance between average daily satellite positions obtained from seven Narwhals instrumented at Kakiak Point prior to the arrival of the Killer Whales. The standard deviation (SD) in latitude and longitude is reported and also converted to north-south dis- persal and west-east dispersal with distance units of km. Latitude SD Longitude SD N-S dispersal (km) W-E dispersal (km) Kernel core (50%) km? Before attack (5 days) 0.18 0.12 19.9 13.0 347 During attack (2 days) 0.33 0.29 36.4 32.8 767 After attack (5 days) 0.18 0.25 19.7 27.6 599 460 ies Sees wk cot = ca . kon FIGURE 2. the shore lying still. Photo M. P. Heide-Jgrgensen. On 20 August, Killer Whales were visually observed arriving at Kakiak Point traveling a northbound route along the west side of the inlet. Narwhals were already present in the coastal area (<500 m from the shore) around Kakiak Point and in a small bay just behind the point. When the Killer Whales were within 2-4 km, Narwhals suddenly moved closer to the shore in shal- low water (<2 m). Some Narwhals formed tight groups THE CANADIAN FIELD-NATURALIST Vol. 120 St a Se When Killer Whales arrived in the vicinity of Kakiak Point, Narwhals formed tight groups and remained close to and others moved slowly or lay very still at the surface (Figure 2). One Narwhal stranded on a flat gravel beach (<0.5 m of water) and made violent tail thrash- es for >30 seconds (Figure 3), either as a warning sig- nal or in attempts to remove itself from the beach. Satellite telemetry data indicated instrumented Narwhals clearly responded to the presence of Killer Whales (Figure 4). During the attack, both the north- FiGurE 3. During the Killer Whale attack, Narwhals beached themselves in sandy areas and made tail slaps. Coastline can be seen in lower left. Photo by K. L. Laidre. 2006 LAIDRE, 110°0'0"W 100°0'0"W 70°0'0"N 65°0'0"N 90°0'O"W 80°0'0"W Kakiak Point HEIDE-JORGENSEN, and Orr: REACTIONS OF NARWHALS 70°0'0" J 46] 90°0'O"W 80°0'0"W \ ? 75°0'0"N J it AC Admiralty nll } @ 8/14 - 8/18 [} 8/19 - 8/20 A\ 8/21 - 8/25 FiGuRE 4. Average daily positions received from seven Narwhals instrumented with satellite tags before the Killer Whale attack. Average positions are shown in 3 time periods: the five days prior to Killer Whale arrival (14-18 August), the two days Killer Whales were observed (19-20 August), and the five days after the Killer Whale departure (21-25 south and west-east dispersal (as measured by inter- whale mean latitude and longitude standard deviations) doubled (Table 2). When Killer Whales were in the area, the core kernel home range shifted south by ap- proximately 80 km and doubled in size (767 km/’) (Fig- ure 5, Table 2). Generally, the groups of Narwhals were more widely dispersed in all directions during the attack than before the Killer Whales arrived. Post-Attack Narwhal Behavior (>5 days) Narwhals resumed their normal swimming behay- ior and distance from the coast within an hour after the Killer Whales left the locality. No whales were ob- 462 served closer than 20 m from the beach and tight groups at the surface broke-up and dispersed. The instrumented Narwhals moved offshore and utilized a wider area after the attack (ANOVA on longi- tude, F, ,=17.6, P < 0.001). Standard deviations of the mean inter-whale latitude (19.7 km) were reduced by half after the departure of the Killer Whales and were nearly identical to pre-attack values (19.9 km), demon- strating a contraction of the north-south dispersal (Table 2). The mean inter-whale longitude standard deviation was slightly lower than during the Killer Whale attack (27.6 km), yet did not return to pre-attack value, indicating a general inshore-offshore dispersal from the attack site. Post-attack kernel home range core area (599 km?) was less than that during the attack, yet the core area was located east of the pre-attack core in the center of Admiralty Inlet extending across to the eastern shore- line (Figure 5, Table 2). In general the Narwhal groups maintained a wider offshore dispersal and ranged more widely than before the Killer Whale attack. Discussion Narwhals exhibited clear reactions to the immedi- ate presence of Killer Whales. Their suite of behav- iors included slow, quiet movements, travel very close to the beach in the surf zone, movement into very shal- low water less than 2 m, and formation of tight groups at the surface. All of these observations are consistent with Inuit accounts. Within an hour after the Killer Whale attack and when Killer Whales had left the area, direct observations suggested Narwhals resumed nor- mal behavior (e.g., distances from the coast increased and tight groups dispersed). Satellite telemetry data revealed long-term (multi- day) observations of Narwhal movements before and after the attack, in addition to observations during the attack. Average daily positions indicated that Narwhal groups were more dispersed during and after the Killer Whale attack. Dispersal primarily increased in the off- shore (west-east) direction as measured by longitude standard deviations and home ranges. Cores of area use calculations demonstrated that the range of Nar- whal movements increased two-fold during the Killer Whale attack, shifted south in a direction away from the attack point, and then expanded to offshore areas afterwards (where Narwhals had not been observed previously in 2005). Later in the season, Killer Whales were reported in “fairly large numbers” in Admiralty Inlet and vicinity (especially Adams Sound) until October 2005 (Niore Annie Igqalukjuak, Arctic Bay, personal communica- tion). The exact location and movements of these Killer Whales between August and September are not known, nor is it known whether other attacks on Narwhals oc- curred. Satellite tagging data showed that Narwhals remained in Admiralty Inlet until the third week of October, after which the southbound fall migration was underway. Thus, it appears that even on longer time THE CANADIAN FIELD-NATURALIST Vol. 120 scales (~2 months) Narwhals do not alter their site fideli- ty to the summering grounds (see Heide-Jgrgensen et al. 2003) or depart early in response to the presence of Killer Whales. Killer Whales were also reported in Admiralty Inlet in late August 2004 but no data or specific observa- tions were available. It is unknown if the occurrence observed in 2004 involved the same group of Killer Whales as observed in 2005. On 25 August 2004, one of the authors (J.R.O.) observed a young Bowhead Whale (Balaena mysticetus) killed by Killer Whales at Kakiak Point, and bones were found on the beach less than 1 km away from the attack site the following year in 2005. Furthermore, on the same date as the at- tack reported here, several Killer Whales were observed attacking Narwhals in Repulse Bay, Hudson Strait, Canada (Joani Kringayark, personal communication). Therefore, at least two separate groups of Killer Whales feed on Narwhals on their summering grounds in the eastern Canadian Arctic in August. When Killer Whales attacked Narwhals, there was limited action at the surface with no struggle or splash- ing observed. Killer Whales consumed the Narwhals below the surface of the water, and made multiple dives in the center of oiled areas (presumed kills) for 15-20 minutes. An eyewitness account of a Killer Whale attack on Narwhals in Inglefield Bredning, West Green- land in August 2004 noted Killer Whales jumping on top of Narwhals and holding them under the surface until they drowned (Gretchen Freund, personal com- munication). If the predation level observed at Kakiak Point (4 Narwhals over 6 hours) is representative of the daily predation level of a similar sized pod of Killer Whales in the High Arctic, then consumption of Narwhals dur- ing a two-month stay in Admiralty Inlet would amount to approximately 200-300 Narwhals. A photographic survey estimated 5,556 (CV=0.22) Narwhals in Admi- ralty Inlet in 1984 (Richard et al. 1994). If the photo- graphic survey results are corrected for whales that were diving using a correction factor of 0.38 (0.06) developed by Heide-Jgrgensen (2004), then the abun- dance would have been in the order of 14,621(0.23). A mortality of 200-300 Narwhals from Killer Whale predation would represent <3% of this abundance esti- mate. It is possible that predation constitutes a larger proportion of the annual natural mortality of Narwhals than previously expected; however, it is not known if this natural mortality is fully compensatory. In this case, a continued high level of Killer Whale predation in combination with a population reduction by subsistence harvest could reinforce a decline. Killer Whale preda- tion has been responsible for the decline of sea otters in Alaska (Estes et al. 1998; Doroff et al. 2003) and has been hypothesized as the driver for a decline of other top predators in the North Pacific (Springer et al. 2003). The reason Narwhals occupy deep and narrow fjords during summer is unknown and it has been hypothe- sized that their summer distribution may be related to 463 IN, and ORR: REACTIONS OF NARWHALS :IDE-JORGENSI LAIDRE, HE 2006 ‘JSEOD JSAM AY} UO (JOQUIAS IR}S) JULOg YELYRY wv poasasqo sem yorne seyAA J9[PY oy.L “Asnsny ¢z-isnsny 17) sanyedoap sey JQIPEN Oy) Joye SAPP 2AY ay puke ‘AsNSny OZ-G]1) Parsosqo as9M sapeYyAA JOT[PY Shep OM) oy) ‘Asnsny gi-p]) [PALUE a[PYAA Ja[[Py 0) JoLd skvp aay oY, :spotiod own do1y) JO} Pale[MI[HO Qa soBues owWlOH “(NYA SI %OG) SUOSATOd oSuRs sWOY [OUIOY YIM payewNsea yoRneE aeY AA JO[[OY AY) JaIye pue ‘BuLNp ‘asojoq susoyed asn-dords [RYMIMN ‘¢ AUNOLY = = - _——yayuy Ayesmupy_” ' a Tomy Ayesupy_ yorRye-]sod yore suLing yoRyye-od 464 potential predation by Killer Whales (Kingsley et al. 1994). Narwhals feed very little during the summer season (Laidre and Heide-Jgrgensen 2005a), and sum- mering grounds do not appear to be related to calving needs given calves are born in the spring. The summer refuge from Killer Whales hypothesis lacks conclusive evidence as Narwhal distribution in other seasons is clearly driven by biological needs, such as sea ice for- mation, open water, or access to prey resources (Laidre and Heide-Jgrgensen 2005b) and Narwhals did not seek refuge or depart early in response to the observed at- tack. Little is known about what proportion of different types of Killer Whales occurs in Arctic waters (i.e., mammal-eating vs. fish-eating). Stomach contents of 30 Killer Whales harvested in Disko Bay, West Green- land in February 2003 contained only Lumpsucker fish (Cyclopterus lumpus), despite the fact these Killer Whales were taken in an area with a large abundance of Bowhead Whales, Ringed Seals (Phoca hispida), Narwhals, Belugas, and other potential marine mam- mal prey items (Greenland Institute of Natural Re- sources, unpublished data). Few observations have been collected of Killer Whales during winter in Arctic ice conditions. It is gen- erally assumed that Killer Whales avoid the Arctic pack ice (Heide-Jorgensen 1988) despite the fact many Kil- ler Whales occur and thrive in the dense pack-ice of the Antarctic. It is possible that changes in sea ice (lighter sea ice cover and earlier break-up) will alter (or have already altered) the occurrence of Killer Whales in Arctic waters. These changes may facilitate increased or longer visits by Killer Whales to ice-free Arctic areas. The predicted reduction of annual sea ice, together with a longer open water season, will likely lead to an increase in Killer Whale predation on Narwhals. At the same time, reduced sea ice will also decrease the probability that Narwhals succumb in ice entrapments (also known as Sassats), another important source of natural mortality (Laidre and Heide-Jgrgensen 2005b). It is unclear how the interplay between these two ele- ments of natural mortality will evolve under chang- ing sea ice regimes. Acknowledgments Research was funded by the Greenland Institute of Natural Resources, Fisheries and Oceans Canada, and the Nunavut Wildlife Management Board. The Polar Continental Shelf Project in Resolute supported logis- tics for field work in August 2005. The work was also supported by the Arctic Bay Hunters and Trappers Organization. Narwhal tagging was conducted under permits from Fisheries and Oceans Canada and Animal Care Permit FWISLACC#013. 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Proceed- ings of the National Academy of Sciences 100(21): 12, 223- 12, 228. Steltner, H., S. Steltner, and D. E. Sergeant. 1984. Killer Whales, Orcinus orca, prey on Narwhals, Monodon mono- ceros: An Eyewitness Account. Canadian Field-Naturalist 98: 458-462. Received 9 February 2006 Accepted 18 June 2007 Urine-marking and Ground-scratching by Free-ranging Arctic Wolves, Canis lupus arctos, in Summer L. Davip MeEcu!? 'U. S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 — 37" Street, SE, Jamestown, North Dakota 58401- 7317 USA Mailing address: The Raptor Center, 1920 Fitch Avenue, University of Minnesota, St. Paul, Minnesota 55108 USA Mech, L. David 2006. Urine-marking and ground-scratching by free-ranging Arctic Wolves, Canis lupus arctos, in summer. Canadian Field Naturalist 120(4): 466-470. Urine-marking and ground-scratching were observed in an Arctic Wolf (Canis lupus) pack on Ellesmere Island, Nunavut, Canada, during 16 summers between 1986 and 2005. All previously known urination postures and ground-scratching by breeding males and females were seen, and incidence of marking and scratching was greatest when non-pack wolves were present. Observations of urine-marking of food remains supported the conclusion from a captive Wolf study that such mark- ing signals lack of edible food. Key Words: Arctic Wolf, Canis lupus arctos, ground-scratching, odor, scent-marking, territoriality, urine-marking, Ellesmere Island, Nunavut, Canada. Scent-marking by Wolves (Canis lupus) includes both urine-marking and ground-scratching, usually by dominant breeding males and females (Peters and Mech 1975; Harrington and Asa 2003). Most infor- mation about Wolf urine-marking has been collected during winter by tracking Wolves in the snow (Peters and Mech 1975; Rothman and Mech 1979; Paquet and Fuller 1990; Paquet 1991; Zub et al. 2003) or during other seasons by studying captive Wolves (Harring- ton 1981; Raymer et al. 1984, 1986; Asa et al. 1986, 1990; Mertl-Millhollen et al. 1986; Ryon and Brown 1990; Barja and Miguel 2004). Only scattered men- tions can be found of free-ranging Wolves urine-mark- ing during summer (Clark 1971; Haber 1977; Mech 1991, 1995), and only one study of Wolf ground- scratching in summer has been done (Zub et al. 2003). Detailed information about urine-marking in wild Wolves during summer is lacking because, with a few exceptions (Murie 1944; Clarke 1971; Haber 1977; Mech 1988), it has been difficult to observe Wolves in the wild during summer because of their usual fear and avoidance of humans. I observed free-ranging Wolves in the Canadian High Arctic over a period of 20 summers, where the Wolves are highly tolerant of humans (Parmelee 1964; Miller 1978; Mech 1988). There I obtained detailed data on their urine-marking and ground-scratching behavior. Study Area The study area includes about 2600 km? of the Fosheim Peninsula east, north and west of Eureka on Ellesmere Island (80°N latitude, 86°W longitude), Nunavut, Canada. During summer, daylight is constant. The area includes shoreline, hills, lowlands, creek bot- toms and the area around Blacktop Ridge. Contrary to much of the surrounding region, this area is generally snow- and ice-free in summer, and contains rock, grav- el, bare soil and scattered tundra and northern wetland vegetation. Wolves, Muskoxen (Ovibos moschatus) and Arctic Hares (Lepus arcticus) have long been common in the area (Tener 1954), and Wolves have denned there over decades or possibly centuries (Parmelee 1964; Grace 1976; Mech 1988; Mech and Packard 1990). Aside from intermittent scavenging around a weather station and military base, the main foods of the Wolves I studied were Muskoxen and Arctic Hares (Tener 1954), although seals (Phoca spp.) and Lemmings (Dic- rostonyx groenlandicus) are also taken occasionally. Methods The Wolves I studied live far enough from exploita- tion and persecution by humans that they are relatively unafraid of people (Mech 1988, 1995). During 1986, I habituated a Wolf pack to my presence and reinforced the habituation each summer I observed them. The pack frequented the same area each summer and usually used the same den or nearby dens. The habituation allowed me and an associate to remain with the Wolves daily, to recognize them individually, and to watch them regular- ly from 10 to 200-m away, and often as close as | m (Mech 1988, 1995). This pack had disappeared by 2001, but by 2003 another pack, similarly habituated, was present (Mech 2005). Although I made no attempt to systematically or completely record every urination or scratching made by each Wolf, I did note as many such events as I could, usually those made by breeding males and females, while I was observing general behavior. I identified breeding females by observing them nursing pups or by the presence of obvious teats. Breeding males were identified by their dominance over other males and their | 466 2006 MECH: URINE-MARKING AND GROUND SCRATCHING BY WOLVES 467 TABLE |. Yearly distribution of observed urine marks made by breeding male (RLUs and STUs) and breeding female (FLUs and SQUs) Arctic Wolves, Ellesmere Island, Nunavut, Canada, 1986-1996, including only years in which observations cov ered at least four weekly periods.! Number of Male Weekly Marks Summers Periods Marks Per Period 1986 4 7 1.8 1987 5 7 1.4 1988 7 9 eS 1989 8 5) 0.6 1990 8 11 1.4 199] 8 1] 1.4 1992 6 29 4.8 1993 6 12 2.0 19942 5 3 0.6 1996 6 27 4.5 Female Total Marks Marks Marks Per Period Marks Per Period 0 0 7 1.8 0 0 7 1.4 0 0 9 1.3 5 0.6 10 hes 18 2.3 29 3.6 22, 2.8 33 4.1 14 25 43 V2 13 ZR, 25 4.2 5 1.0 8 1.6 14 PS, 4] 6.8 ' Because of potential biases in times and methods of data collection, fine comparisons are not necessarily valid. 2 1995 covered only two periods. overall initiative and control of pack activities (Schen- kel 1947; Mech 1970, 1999; Peterson et al. 2002). I collected data over periods of 1-8 weeks from June through early August 1986-2005, except 1997, 1999, 2000, and 2001. Most of the longest data col- lection took place 1986-1996 (Table 1). Because study periods, pack behavior, and observation emphasis var- ied over the summers and years (Table 2), biases in the collection of urine-marking and ground-scratching data over both the total duration of the study and over the seasonal extent of the study each summer proba- bly affected the data collected. Thus only the data on ratios of marking postures should be unbiased enough to warrant statistical testing, and I make no attempt to draw detailed conclusions from the untested data. However, these data are still useful to make broader conclusions and to establish several facts that hereto- fore were unknown or poorly documented in free- ranging Wolves during summer. Results I identified three breeding female and three breed- ing male Wolves, plus one pair without pups, that trav- eled, hunted and marked together (Mech 1995, 2005, and unpublished). I recorded males making 121 raised- leg urinations (RLU) 22 June — 8 August and 30 stand- ing urinations (STU) 15 June — 10 August; and females, 47 flexed-leg urinations (FLU) 15 June — 3 August, and 70 squat urinations (SQU) 15 June — 29 July (Table 2). The proportion of male RLUs was significantly high- er than the proportion of female FLUs (x? = 45.01; P < 0.0001; df. = 1). The number of both male and female marks I observed per weekly observation period appeared to decrease over the summer (Table 2). The number of recorded urinations per observation period varied by year from 1.3 to 7.2. (Table 1). In addition to FLUs by breeding females, I also observed a pre-breeding female (bred the next year) FLU twice on 6 August. An associate had also seen this animal do “slight FLUs” three times on 13 July (see Discussion.) I observed double marking (Rothman and Mech 1979) 24 times between 26 June and 3 August. Double marking is marking of a single location by both mem- bers of a mated pair within a few minutes of each other and can be initiated by either member. Of eight pos- sible combinations of double-marking postures (RLUs or STUs initiated by males, FLUs or SQUs initiated by females) I observed five of the combinations, with those involving RLUs and FLUs significantly pre- dominating (Table 3). I observed ground-scratching by both male and fe- male breeding Wolves or by either individual as early as 15 June and as late as 31 July. The pre-breeding female (above) scratched as late as 6 August. Besides the breeders’ and pre-breeder’s ground-scratching, a non-breeding female was observed scratching during two different summers, the same Wolf each year. Of 16 summers when I observed breeding or mated Wolves, I recorded no scratching by them during seven of those years and three or less scratching events during six other summers. Only during 1992, 1993, and 1996 did I see Wolves do much scratching, and both male and female breed- ers did so. During the above years, the breeding pair I studied, the same individuals each year, scratched reg- ularly. Those two individuals had also been the breeders since 1989, but I saw little scratching during 1989- 1991, 1994 or 1995. During 1992, the breeding pair scratched (and urine-marked — Table 2) much more than I had seen them in the past. They scratched (and urine-marked) especially (1) near a Muskox they had killed and at which a non-pack Wolf had also been feeding a few minutes before, (2) near a garbage dump where at least one non-pack Wolf had been feeding regularly and (3) along the easternmost location (ter- 468 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 2. Weekly distribution of observed urine marks made by breeding male (RLUs and STUs) and breeding female (FLUs and SQUs) Arctic Wolves, Ellesmere Island, Nunavut, Canada, 1986-2005.! Male Number Marks Summers Periods Marks Per Period 15-22 June 5 3 0.6 23-30 June 9 39 43 1-7 July 11 31 2.8 8-14 July 13 36 2.8 15-21 July 10 16 1.6 22-28 July 9 20 DD 29 July—4 August 9 4 0.4 5-10 August 6 2 0.3 Female Total Marks Marks Per Period Marks Per Period 9 1.8 12 2.4 18 2.0 Si 6.3 31 2.9 62 5.6 36 2.8 72 5.5) 16 1.6 32 39) 4 0.4 24 De 3 0.3 7 0.7 0 0) 2 0.3 ' Because of potential biases in times and methods of data collection, fine comparisons are not necessarily valid. ritory boundary?) where we saw them that summer. During 1996, the breeding female scratched and urine- marked while trying to force a Muskox away from the den. In another study area, in the Canadian Northwest Territories, I also observed wolves scratching several times at a Red Fox (Vulpes fulva) den (Mech unpub- lished) during August. I observed ground-scratching in various elimina- tion contexts: (1) with no other marking behavior, (2) followed by FLU or RLU, (3) preceded by FLU, RLU or SQU, (4) preceded and followed by FLU, or (5) pre- ceded by defecation, similar to the findings of Peters and Mech (1975) and Mertl-Millhollen et al. (1986). I also observed urine-marking in relation to food remains as Harrington (1981) also did. Breeding males and females used all four kinds of urine postures in marking uneaten food remains or prey odors including the following (1) Arctic Hare intestines; (2) chewed bones; (3) Arctic Hare stomach contents; (4) locations where regurgitated or other food had recently been eaten; (5) old kills; (6) the head of a recent Muskox kill; (7) a putrid Muskox carcass which they sometimes fed on but often passed up; (8) where another Wolf had tried but failed to catch a lemming; and (9) at caches of which they or another Wolf (including a pup) had just consumed the contents. Discussion Captive breeding male and female Wolves urine- mark at all times of the year, with the peak frequency during the winter breeding season, and with all studies except that by Ryon and Brown (1990) observing that males tend to mark more year around than females (Peters and Mech 1975; Mert!-Millhollen et al. 1986; Asa et al. 1990; Barja and Miguel 2004). Information on urine-marking frequencies during summer in captive wolves varies considerably. In one case, where the breeding pairs had pups, both males and females increased their marking rate during sum- mer (Mertl-Millhollen et al. 1986). In another, where “reproductive pairs” were studied, but no pups were born, the males increased their rate of RLU during TABLE 3. Order and type of urination postures in double urine marks! by arctic Wolves observed during summers 1986-2005 on Ellesmere Island, Nunavut, Canada. (Males do raised-leg [RLU] and standing [STU] urinations, and females, flexed-leg [FLU] and squat [SQU] urinations.) Male Initiated RLU-FLU 6 RLU-SQU 1 STU-FLU 2 STU-SQU 0 Female Initiated FLU-RLU 9 FLU-STU 0 SQU-RLU 6 SQU-STU 0) ' Urination sequences involving RLUs or FLUs occurred sig- nificantly more often than expected by chance (x7 = 6.86; P=0.03; d.f. = 1). summer, but the females did not increase their FLU rate (Barja and Miguel 2004). In two others, where no pups were involved, the RLU and FLU rates of domi- nant males and females reached their minimum in summer (Asa et al. 1990; Ryon and Brown 1990). Testosterone tends to decrease to a minimum during summer (Asa et al. 1990), as does testis size (Mech 2006). This relationship suggests that urine-marking, which depends on testosterone (Asa et al. 1990), would also decrease then. However, the variations found in the studies cited above suggest that the social milieu can affect the relationship between testosterone and urine-marking, so that the relationship might not al- ways be so tight, as Asa et al. (1990) also found. Be- cause of possible biases mentioned earlier, my data can add little to this subject, although conceivably the ex- treme variation seen in my data reflect actual behavioral variation similar to that reported in captive Wolves. In that respect, my data on ground-scratching are instruc- tive. This study documents for the first time that wild Wolves urine-mark throughout summer, using all known urine postures, including double-marking, but 2006 that the proportion of RLUs and FLUs to STUs and SQUs is lower then than in captive Wolves during winter (Asa et al. 1990) when RLUs and FLUs tend to predominate (Peters and Mech 1975). The fact that a pre-breeding female was seen FLU- ing (“RLU” in Mech 1995) is of special interest. Based on my observation of this pack the previous three summers, I inferred that she had been born two years earlier. This pre-breeder began dominating the breed- ing female, her apparent mother, about the time the pre-breeder began FLUing. The pre-breeder bred the next year, apparently with her father, and remained bonded to him for the next six years, producing pups during four of them while the mother remained with the pack for two more years without being seen FLU- ing, and then disappeared (Mech 1995 and unpub- lished). This behavior of a young offspring starting to FLU or RLU and challenging its same-sex parent for dominance is similar to that seen in a captive colony (Asa et al. 1990). Previous information about free-ranging Wolves ground-scratching in summer comes only from a single study which concluded that between April and Septem- ber, ground-scratching was low and stable (Zub et al. 2003). I could make no similar seasonal comparison, but it was clear from my multi-summer observations that the degree of ground-scratching varied consider- ably from summer to summer, with no scratching seen some summers and regular scratching during others. In 1992, the year of most scratching, the presence of non- pack Wolves seemed to be the pertinent stimulus. It was the only year I observed the Wolves I had been study- ing chase or attack outsiders on three occasions (Mech 1993). It was also the year when I recorded the high- est amount of urine-marking (Table 1), when the Wolves usually were extra alert and looking around intently when they marked and scratched, and when they marked their easternmost locations, suggesting a territory boundary. My impression during these times was that the Wolves were in a high state of arousal and aggressiveness. Both breeding males and females scratched about equally, contrary to Mertl-Millhollen et al. (1986) who found primarily females scratching. Regarding marking of food remains, my observa- tions confirm those of Harrington (1981) on captive Wolves and support his conclusion that marking of recently emptied food caches (and by extension from my observations, other inedible food remains) signals that a site contains no more edible food despite lin- gering odors. In summary, this study documents that much infor- mation learned about Wolf urine-marking behavior in Captivity during summer applies to free-ranging Wolves; it extends information about ground-scratch- ing; and it places information from captive studies about urine-marking of food remains in its natural context. MECH: URINE-MARKING AND GROUND SCRATCHING BY WOLVES 469 Acknowledgments This research was funded by the U. S. Geological Survey. Polar Continental Shelf Project (PCSP), Atmos- pheric Environment Services of Canada, and USDA North Central Research Station provided logistical sup- port. I also thank numerous field assistants, including L. G. Adams, F. H. Harrington, W. Medwid, J. Sanders, G. Breining, and H. D. Cluff. The following reviewed the manuscript and offered helpful suggestions for its improvement: L. G. Adams, S. M. Barber, and H. D. Cluff, and M. E. Nelson. This is PCSP/EPCP paper 00905. Literature Cited Asa, C.S., U. S. Seal, E. D. Plotka, and L. D. Mech. 1986. 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Canadian Wildlife Service Wildlife Management Bulletin Series 1, Number 9. 40 pages. Zub, K., J. Theuerkauf, W. Jedrzejewski, B. Jedrzejewska, K. Schmidt, and R. Kowalczyk. 2003. Wolf pack terri- tory marking in the Bialowieza Primeval Forest (Poland). Behaviour 140: 635-648. Received 26 September 2005 Accepted 22 December 2006 Notes Double Marking in Arctic Wolves, Canis lupus arctos: Influence of Order on Posture FRED H. HARRINGTON Department of Psychology, Mount Saint Vincent University, Halifax, Nova Scotia B3M 2J6 Canada Harrington, F. H. 2006. Double marking in Arctic Wolves, Canis lupus arctos: Influence of order on posture. Canadian- Field Naturalist 120(4): 471-473. Double marking by Arctic Wolves (Canis lupus arctos) was recorded by Mech (2006) from a pack on Ellesmere Island, Nunavut, Canada, during 16 summers between 1986 and 2005. Using his data on the frequency of occurrence for each of the four postures used by Wolves for urine marking (males — raised leg and stand urinations; females — flexed leg and squat uri- nations), the probabilities of occurrence for each of eight possible double mark sequences were determined and compared with observed frequencies. Females were somewhat but not significantly more likely to initiate double mark sequences. There was no evidence for any bias in the posture used to initiate a double mark sequence, but assertive postures by both males and females nearly always completed the sequence, occurring much more often than expected by chance. Key Words: Arctic Wolves, Canis lupus arctos, scent-marking, double marking, pair bonding, mate guarding, urine-marking, Ellesmere Island, Nunavut, Canada. Double marking, in which a mated pair urine marks a single site in quick succession, is commonly observed in Wolves (Canis lupus) (Harrington and Asa 2003), although evidence from wild Wolves is often inferred from the pattern of urine marks on snow (Peters and Mech 1975; Rothman and Mech 1979). Direct observa- tions, such as those of Mech (2006), permit the identi- fication of the postures used in marking, from which underlying motivation might be inferred. Males may use either raised-leg urinations (RLUs) or standing urinations (STUs) and females either flexed-leg urina- tions (FLUs) or squat urinations (SQUs), with the for- - mer posture for either sex considered to be more reflec- _ tive of aggressive or assertive motivation (reviewed by _ Harrington and Asa 2003). Thus there are eight pos- _ sible forms that a double marking sequence can take. _ Mech (2006) has presented data on urine-marking in _ Wolves, recorded by direct observation from Arctic _ Wolves, Canis lupus actus, during the summer. Among _ those data are 24 cases of double marking (Table 1), _ and the reported finding that the more assertive pos- _ tures (RLUs and FLUs) occur more frequently in dou- ble marking. The purpose of this note is to extend the analysis of those data in order to offer more insight ) into the motivation behind double marking. Methods A dominant adult Wolf may urine-mark using either posture appropriate for its sex. This analysis seeks to determine whether there is a non-random pattern in the _ use of either posture, by either sex, in double marking _ compared to urine-marking in general. Therefore I first _ determined the overall frequency of each urination pos- TABLE |. Urination postures used during double marking by Arctic Wolves during summers 1986-2005 on Ellesmere Island, Nunavut, Canada. Male postures: raised-leg [RLU] and standing [STU] urinations; female postures: flexed-leg [FLU] and squat [SQU] urinations. (Data from Mech 2006). Male Initiated Double Marks RLU—-FLU 6 RLU-SQU l STU-FLU 2 STU-SQU 0 Female Initiated Double Marks FLU-RLU 9 FLU-STU 0) SQU-RLU 6 SQU-STU 0 ture that occurred exclusive of double marking (Table 2) to yield two base rates, one for the probability of occurrence for each of the postures overall and anoth- er for the probability of the occurrence of each form within a sex. From these probabilities, one can calcu- late expected frequencies for each form of double mark, under the assumption that the postures used in double marking simply represent random combinations of the postures utilized otherwise. This is done by multiply- ing the value from row | (the random probability the posture will occur overall and thus initiate a sequence) with the value from row 2 (the random probability the other sex will use one of its two alternative postures to complete the double mark). Thus, for example, the 47] 472 THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 2. Calculation of the probability of occurrence for each of the urination postures used by Arctic Wolves on Ellesmere Island, Nunavut, Canada during summers 1986-2005. Urination posture RLU STU Total marks 121 30 Minus double marks -22 -2 Non-double mark urinations 99 28 (1) Probability of posture (overall) 0.45 0.13 (2) Probability of posture (within sex) 0.78 0.22 FLU SQU Total marks 47 70 268 -17 -7 -48 30 63 220 0.14 0.28 1.00 0.32 0.68 1.00/sex TABLE 3. Expected and observed frequencies of occurrence for each of the eight different forms of Wolf double marking, for Arctic Wolves on Ellesmere Island, Nunavut, Canada during summers 1986-2005. Expected frequencies are based on the combined probability of occurrence for each posture in the sequence (from data in Table 2), under the assumption of independence of occurrence for each posture type and based on a sample size of 24. Double urination mark sequence Male initiated sequences RLU-FLU RLU-SQU STU-FLU STU-SQU Expected frequency 3.5) eS 1.0 Dell Observed frequency 6 1 2 0 Probability of occurrence 0.144 0.306 0.042 0.088 Female initiated sequences FLU-RLU FLU-STU SQU-RLU SQU-STU Expected frequency 2.6 0.7 2 NES) Observed frequency 9 0 6 0 Probability of occurrence 0.109 0.031 0.218 0.062 probability for an RLU/SQU double mark to occur at random is 0.45°0.68=0.306, whereas a double mark in the reverse order (SQU/ RLU) is expected less often (0.28°0.78=0.218), primarily because males are more likely to use dominant postures overall than are females (Table 3). Results Three questions can be addressed using the proba- bilities generated in Table 3. First, is one sex more likely to initiate a double marking sequence? Al- though females initiated 63% of the double marks observed, compared to the expected frequency of 42%, this increase was not greater than expected by chance (x? = 2.10, df = 1, ns). Second, is the first mark more likely to be from an assertive posture than otherwise? Again, although twice as many double marks were initiated using the more assertive RLU and FLU pos- tures, this was expected overall, given the high rate of assertive posturing (59%) among dominant Wolves (x? = 0.322, df = 1, ns). Finally, is the second mark which completes the double mark, more likely to be from an assertive posture? Here, the data indicate a high- ly significant deviation from expectations (x7 = 13.90, df = 1, P < 0.001). All but one of the second marks involved either RLU or FLU postures. The one excep- tion was an RLU by the male followed by a SQU by the female. Males increased their use of RLUs from a base rate of 78% in other situations to 100% when com- pleting a double mark, while females increased their use of FLU postures even more dramatically, from 32% overall to 89% in double marks. Discussion Several captive studies have found that dominant female Wolves are usually the initiators of double marks (93% of double marks, n=23, Rothman and Mech 1979; 64%, n=36, Mertl-Millhollen et al. 1986). In a sample of over 700 double marks by free-ranging Coyotes (Canis latrans), Gese and Ruff (1997) found that double marking was initiated by females 75% of the time. Unfortunately, none of the previous studies tested observed initiation rates against expected rates, based on the overall frequency of marking by indi- viduals of each sex, as I have done here. However, the females observed by Mech (2006) did initiate double marking more often than expected, at a rate (63%) comparable to previous studies, so the lack of statistical significance for this result may be an artifact of small sample size. Thus it seems likely that double marks are more likely to begin as urinations by dominant females. Mertl-Millhollen et al. (1986) found. that 94% of double marks in their captive pack were initiated by RLUs (they combined RLUs and FLUs in their analyses). Unfortunately, although they indicate that RLUs comprised the majority of the marks they re- corded, they do not present their data in a manner to allow testing of expected versus observed rates for the postures. In the present study, although the initial posture used in a double mark was most often an 2006 assertive one (RLU or FLU), this is to be expected, given the relatively high rate of these postures in dom- inant individuals. This analysis suggests that the ini- tial mark of a double mark occurs as an independent event, influenced by a combination of internal factors (arousal, hormones, urine load, etc.) and features of the environment (smells of old marks, food odors, con- spicuous visual objects, etc.), but not with the delib- erate intent to initiate a double mark sequence. For example, when Wolves are traveling, the lead Wolf is typically the initiator of double marks (Mech 1999); as it encounters sites which elicit its marking, the op- portunity for double marking is then created for the trailing Wolf. The second mark, however, is highly influenced by the presence of the first, and either its freshness, indi- vidual identity, chemical composition, or its mere pres- ence at a specific location induces the second individ- ual to use the more assertive RLU or FLU posture. Mertl-Millhollen et al. (1986) likewise found that all but one second mark was either an RLU or FLU; the one exception was a SQU (they combined SQUs and STUs) that followed another SQU. Thus the second mark of a double mark likely represents an increased level of arousal on the part of the marker. Whether that arousal represents generalized excitement, increased sexual motivation or increased aggressive (protective) motivation is not possible to determine here. A vari- ety of hypotheses have been advanced to account for double marking: pair bonding, reproductive synchro- nization, mate guarding, and territorial defense being among the most common (Rothman and Mech 1979; Mertl-Millhollen et al. 1986; Gese and Ruff 1997). The tight relationship between the frequency of dou- ble marking and successful reproduction in Wolves (Rothman and Mech 1979: Asa et al. 1986; Mertl-Mill- hollen et al. 1986; Peterson et al. 2002) suggests that mating related functions, including mate guarding by the male, are most likely. NOTES 473 Acknowledgments This note grew out of a manuscript review. | would like to thank L. D. Mech and F. R. Cook for the invi- tation to prepare this paper and for their willingness to expedite its consideration. I also thank my stu- dents for their patience in receiving their final grades while I worked on the manuscript. Literature Cited Asa, C. S., U. S. Seal, E. D. Plotka, and L. D. Mech. 1986. Effect of anosmia on reproduction in male and female Wolves (Canis lupus). Behavioral and Neural Biology 46: 272-284. Gese, E. M., and R. L. Ruff. 1997. Scent-marking by Coy- otes, Canis latrans: the influence of social and ecological factors. Animal Behaviour 54: | 155-1166. Harrington, F. H., and C. S. Asa. 2003. Wolf communica- tion. Pages 66-103 in Wolves: Behavior, ecology, and con- servation. Edited by L. D. Mech and L. Boitani. University of Chicago Press, Chicago, U.S.A. 428 pages. Mech, L. D. 1999. Alpha status, dominance, and division of labor in Wolf packs. Canadian Journal of Zoology 77: 1196- 1203. Mech, L. D. 2006. Urine-marking and ground-scratching by free-ranging Arctic Wolves, Canis lupus arctos, in sum- mer. Canadian Field-Naturalist 120: 475-479. Mertl-Millhollen, A. S., P. A. Goodmann, and E. Kling- hammer. 1986. Wolf scent marking with raised-leg urina- tion. Zoo Biology 5: 7-20. Peters, R., and L. D. Mech. 1975. Scent-marking in Wolves: A field study. American Scientist 63: 628-637. Peterson, R. O., A. Jacobs, T. D. Drummer, and L. D. Mech. 2002. Leadership behavior in relation to domi- nance and reproductive status in gray Wolves. Canadian Journal of Zoology 80: 1405-1412. Rothman, R. J., and L. D. Mech. 1979. Scent-marking in lone Wolves and newly formed pairs. Animal Behavior 27: 750-760. Received 24 December 2006 Accepted 24 December 2006 474 THE CANADIAN FIELD-NATURALIST Vol. 120 Use of “Micro”-Corridors by Eastern Coyotes, Canis latrans, in a Heavily Urbanized Area: Implications for Ecosystem Management JONATHAN G. Way! and Davip L. EATOUGH? ‘Biology Department, Boston College, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA. e-mail: jw9802 @ yahoo.com. Present address: 64 Cranberry Ridge Road, Marstons Mills, Massachusetts 02648 USA. *Science Department, Revere High School, 101 School Street, Revere, Massachusetts 02151 USA Way, Jonathan G., and David L. Eatough. 2006. Use of “micro”-corridors by eastern Coyotes, Canis latrans, in a heavily urbanized area: implications for ecosystem management. Canadian Field-Naturalist 120(4): 474-476. We document the use of very narrow, linear corridors (termed “micro-corridors’’) that facilitated movements by both a transient and a resident group of eastern Coyotes (Canis latrans) in a heavily urbanized area in north Boston, Massachusetts. Two corridors are discussed: one, a railroad line through downtown Boston; and two, a hole in a cemetery fence giving access to two separated cemeteries in a region of intense human development. Coyotes can be good subjects to illustrate the use of fragmented landscapes because they are common and thus are abundant enough to study yet are wary and avoid novel things and generally avoid people. Key Words: Eastern Coyote, Canis latrans, corridor, fragmentation, Massachusetts, micro-corridor, urbanization. The use of corridors is becoming increasingly rec- ognized as an important conservation tool in frag- mented landscapes (Beier 1995; Gilbert et al. 1998; Meffe and Carroll 1994). Corridors are strips of habi- tat connecting otherwise isolated habitat patches and have been promoted as important features of reserve design that allow movement among high(er) quality habitats (Meffe and Carroll 1994). Connecting land- scapes rather than maintaining a large unfragmented core ecosystem is becoming the only alternative in many urbanized areas, especially where unregulated sprawl is occurring (e.g., Beier 1995). Yet, there is a paucity of data on animal use of corridors and how effective they are in connecting landscapes (Gilbert et al. 1998; Meffe and Carroll 1994). Because of the natural history of predators (e.g., large territories, long- distance movements), a knowledge of carnivore biol- ogy can predict minimum areas where ecosystems can function relatively naturally (Beier 1993; Gittleman et al. 2001; Mech and Boitani 2003; Meffe and Carroll 1994; Way et al. 2002a). However, the effectiveness (positive or negative) of corridors could greatly affect the size needed to maintain predators in certain sized refuges. Coyotes are common in North America (Parker 1995), yet are elusive and difficult to capture (Way et al. 2002b); they are known to avoid novel objects and structures that are dangerous to them, such as box traps (Way et al. 2002b) or foothold traps (Conner et al. 1998; Sacks et al. 1999). Because Coyotes are ubiqui- tous (Parker 1995), neophobic and wary (Sequin et al. 2003), and have relatively large home ranges and move- ment rates (Way et al. 2004), they are a good species to use in the determination of corridor use, especially in urban areas. In other words, although they are common and frequently use human-dominated areas (Grinder and Krausman 2001; Way et al. 2004), they are natural- ly shy and avoid people by being nocturnal and travel- ing quickly in and out of human-dominated areas. For example, Way et al. (2004) found that Coyotes travel extensive distances on linear pathways such as pow- erlines, railroad tracks and golf courses in urbanized areas of Cape Cod, Massachusetts. Findings of Coyote use of corridors in highly urbanized and fragmented areas can potentially be used by managers to help pro- tect more rare species and/or better design reserves for more common species. In this note, we describe the use of very narrow, linear (also called line corri- dors by Meffe and Carroll 1994) “micro”-corridors. Study Area and Methods Coyotes were captured for an ecological study on the north edge of Boston (42.43°N, 71.06°W), in east- ern Massachusetts, in the bordering cities of Everett (4345.0 people/km7), Malden (4290.5 people/km7), and Revere (3089.0 people/km’) (U.S. Census Bureau, 2000 estimates). Coyotes were captured by box trap (Way et al. 2002a) or by ground-based netlauncher (one coyote; Coda Enterprises, Mesa, Arizona) and outfitted with radio-collars (Mod-400, Telonics, Mesa, Arizona, USA and M1900, Advanced Telemetry Sys- tems, Isanti, Minnesota, USA). Tracking protocols were described by Way et al. (2002a) and Way et al. (2004). Portable receivers (Custom Electronics, Urbana, Ilh- nois, USA) and hand-held 3-element Yagi antennas were used to radio-track Coyotes both on foot and from a vehicle. Because of the highly developed landscape with many roads we mostly restricted our activities to automobiles as Coyotes did not react to them as much as to people (e.g., by running away; J. Way, unpub- lished data); occasionally we approached radio-col- lared Coyotes as close as possible on foot without disturbing them. We used binoculars and video-cam- eras when observing Coyotes, and city street lights, nightscopes, and occasionally headlights when fol- lowing Coyotes at night with a vehicle (Way et al. 2002a; Way et al. 2004). 2006 Observations On 13 April 2004 we box-trap captured a dispers- ing 13.6 kg female Coyote (“Fog” — ID #BN0402) in a wooded section of Revere. Based on her behavior, she seemed to have been dispersing from the north to the south when she reached our study area. Her move- ments the week after capture were mostly to the south, including traveling >1 km through densely populated neighborhoods and streets. She localized in four dif- ferent areas for 2-3 days each before moving to a new location. On 26 April 2004 we located her in a fenced- in abandoned field of ca. 4 ha behind a shopping mall at the north edge of the city limits of Boston. She reached that location by either swimming a 200 m river or traveling along railroad tracks over that same river. Based on data taken earlier that night we sus- pect that she went on the railroad tracks. For 3 days she remained in the fenced-in area and ate many Nor- way Rats (Rattus norvegicus) and Cottontail Rabbits (Sylvilagus floridanus) (J. Way, unpublished data). On the night of 29 April she traveled 1.8 km south- west on the railroad line (which was bordered by industrial buildings on both sides), going under Inter- state 93 and the Zakim Bridge, and entered an aban- doned railroad yard in Boston where she spent the following day in a sparsely vegetated 200 x 50 m area. At 02:00 h on 30 April she followed the railroad tracks west through the Cambridge part of Boston and found the first wooded area available 4.9 km from her pre- vious day’s location. She was inactive through 23:37 h on 30 April 2004 and could not be located until 19 November 2004 when she was found alive 100.5 km south of her capture location (and 88.9 km south of her last location in Cambridge) in the town of Dartmouth, Massachusetts, near the Rhode Island border. To make that voyage Fog had to have crossed (over or under) six major interstate highways, including route 93 des- cribed above. From 17 May 2004 to 3 April 2005 we monitored a pack of Coyotes (“The Cemetery Pack’’) in the border- ing towns of Everett, Malden, and Revere. Four Coy- otes were captured and radio-collared in this pack: one, “Maeve” (#BN0404), a 14.5 kg lactating female, captured 17 May 2004, was the breeding female; two, “Jet” (#BN0403), a 15.9 kg breeding male, captured by netlauncher on 29 June 2004, was Maeve’s mate; three, “Jem” (#BN0406), a 10.0 kg 4.5-month-old pup, was captured on 26 August 2004; and four, “Cour” (#BN0405), a 12.3 kg 5-month-old pup, was captured on 15 September 2004. The pack consisted of two to three adults (i.e., one additional uncollared coyote was occasionally sighted in the pack’s territory — its status Was never determined but it was probably a helper Coyote [Way et al. 2002a] to Jet and Maeve) and four pups (two of which were not collared). The group went from six to seven members to four individuals by mid-winter 2004-2005 when it was presumed that some of the pack members (two of the pups [includ- NOTES 475 ing Jem, last successfully located on 11 December 2004] and probably the uncollared adult) dispersed. The group resided almost exclusively in a green area (including some thicker woods) surrounding four large connected cemeteries. The entire area was about 2.5 km? (J. Way, unpublished data) and aside from inactive railroad tracks (i.e., no trains used them) the pack’s territory was surrounded by high-density hous- ing units and/or commercial spaces (malls) on all sides. Two roads transected their range, including a straight east-west road (Fuller Street) in the central part of their territory that connected the two cemeteries that they most frequently used. Most of the cemeteries were surrounded by fencing consisting of vertical metal bars spaced 10 cm apart. The narrowness of this space prevented the Coyotes from crossing through the fence at random locations. However, a small corridor connected two cemeteries where a 34 cm opening occurred at a height of 55 cm in the south cemetery because of a missing metal post and two bent poles, one on each side. After crossing Fuller Street diagonally for about 15 m the Coyotes could then go through an open 305 cm x 198 cm gate (this door was never shut) followed by six steps that lead down into the north cemetery. Besides that cross- ing, there were two roads (one open to cars, the other gated) approximately 50 m east of the main corridor that the Coyotes could use (by diagonally crossing Fuller Street) to access either cemetery. There was one other opening (< 20 cm) in the south cemetery fence about 400 m west of the main corridor that led into a residential neighborhood (after crossing Fuller Street) bordering the west part of the north cemetery. Cour was the only coyote to use the road crossing cor- ridor regularly and Maeve was the only one observed to use the neighborhood crossing corridor. We radio-tracked the Coyotes six to seven days per week on average and, when tracking them at night (street lights illuminating the area), usually made sight- ings (range: one to ten sightings per night) from our parked, turned off vehicle of one to five Coyotes togeth- er crossing the main corridor (exceptions were during a heavy snowstorm [ca. 60 cm] when they did not use the south cemetery at all). They often also crossed the road in daylight during the early morning (06:00- 08:00). During their first 6 months (i.e., April to Octo- ber) sightings included some of the pups going under the fence (17 cm from fence to dirt) about | m west of the opening in the fence leading to the south ceme- tery, often during the same crossing (i.e., they were together) as other Coyotes went through the opening in the fence. This section of Fuller Street was straight. thus the Coyotes could see in both locations and we often watched them, especially the adults, standing on a hill in the south cemetery watching traffic and cross- ing when there were not any cars going by. Except for two instances when the Coyotes were almost hit by cars (the cars had to brake) all crossings were successful. 476 Monitoring of this group ended after the non-dispers- ing members (Maeve, Jet, Cour and one uncollared) were illegally poisoned. Discussion These data indicate that very small areas, if posi- tioned in the right place, can be very important for Coyotes. These micro-corridors gave access to city habitats with few to no trees (along the railroad tracks) and also connected fragmented areas (cemeteries with fences) in already very urban landscapes. While not ideal habitat for Coyotes, these corridors were cer- tainly better than nothing, and echoing the statement of Beier (1995: 235) when discussing Cougar (Puma concolor) dispersal, “any connection between two isolated patches is better than no connection.” Man- agers can use these data for =3 purposes: one, to pro- vide better habitat connectivity in already fragment- ed landscapes (e.g., opening sections of fence where wildlife is likely to cross); two, proactively establish- ing these types of corridors (ideally larger than des- cribed in this paper) where development is planned; and three, more regional planning where important habitat exists and maintaining linkages between sep- arated core habitats. The importance of sub-marginal habitats can not be overstated to facilitate animal movement. For exam- ple, Beier (1995) noted that Cougars can use corri- dors 100 m wide if the distance is < 800 m and 400 m wide if 1-7 km, yet adult Cougars have enormous home ranges in the hundreds of km? (Beier 1993, 1995). Likewise, species previously thought to inhabit only wilderness such as Wolves (Canis lupus) can often live at least at the edge of human-dominated areas if not persecuted (Mech and Boitani 2003) and no doubt corridors facilitate travel in and out of these landscapes. While preserving larger areas (e.g., Beier 1995) is preferred, this is not always possible in the real world and it is increasingly being discovered that animals are adaptable and can often respond to human-induced changes (e.g., Mech and Boitani 2003). Therefore, it is imperative to at least link what is remaining of our wild lands and this can start at the very specific micro- corridor landscape scale. For example, bridges that exist because of waterways (common in many cities) could also promote wildlife movement if cover and some space are provided. THE CANADIAN FIELD-NATURALIST Vol. 120 Acknowledgments We thank the Way and Cifuni families, N. Lima, S. Cifuni, E. Strauss, Environmental Studies Program, and Urban Ecology Institute at Boston College, B. Binder and Saugus Animal Hospital, and Revere High School. Literature Cited Beier, P. 1993. Determining minimum habitat areas and habi- tat corridors for cougars. Conservation Biology 7: 94-108. Beier, P. 1995. Dispersal of juvenile cougars in fragmented habitat. Journal of Wildlife Management 59: 228-237. Conner, M. M., M. M. Jaeger, T. J. Weller, and D. R. McCullough. 1998. Effect of coyote removal on sheep depredation in northern California. Journal of Wildlife Management 62: 690-699. Gilbert, F., A. Gonzalez, and I. Evans-Freke. 1998. Corri- dors maintain species richness in the fragmented land- scapes of a microecosystem. Proceedings of the Royal Society of London 265: 577-582. Gittleman, J. L., S. M. Funk, D. Macdonald, and R. K. Wayne. Editors. 2001. Carnivore conservation. Cambridge University Press, New York. 675 pages. Grinder, M. I., and P. R. Krausman. 2001. Home range, habi- tat use, and nocturnal activity of coyotes in an urban envi- ronment. Journal of Wildlife Management 65: 887-898. Mech, L. D., and L. Boitani. Editors. 2003. Wolves: behav- ior, ecology, and conservation. University of Chicago Press, Chicago, Illinois. 448 pages. Meffe, G. K., and C. R. Carroll. 1994. Principles of con- servation biology. Sinauer Associates, Sunderland, Mas- sachusetts. 601 pages. Parker, G. R. 1995. Eastern coyote: the story of its success. Nimbus Publishing, Halifax, Nova Scotia. 254 pages. Sacks, B. N., K. M. Blejwas, and M. M. Jaeger. 1999. Rel- ative vulnerability of coyotes to removal methods on a northern California ranch. Journal of Wildlife Management 63: 939-949. Sequin, E. S., M. M. Jaeger, P. F. Brussard, and R. H. Barrett. 2003. Wariness of coyotes to camera traps rela- tive to social status and territorial boundaries. Canadian Journal of Zoology 81: 2015-2025. Way, J. G., I. M. Ortega, and P. J. Auger. 2002a. Eastern coyote home range, territoriality and sociality on urbanized Cape Cod, Massachusetts. Northeast Wildlife 57: 1-18. Way, J. G., I. M. Ortega, P. J. Auger, and E. G. Strauss. 2002b. Box-trapping eastern coyotes in southeastern Mas- sachusetts. Wildlife Society Bulletin 30: 695-702. Way, J. G., I. M. Ortega, and E. G. Strauss. 2004. Move- ment and activity patterns of eastern coyotes in a coastal, suburban environment. Northeastern Naturalist 11: 237- 254. Received 4 October 2005 Accepted 14 March 2007 2006 NOTES 477 Bald Eagle, Haliaeetus leucocephalus, Preying on Maritime Garter Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova Scotia STORRS L. OLSON Division of Birds, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 USA Olson, Storrs L. 2006. Bald Eagle, Haliaeetus leucocephalus, preying on Maritime Garter Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova Scotia. Canadian Field-Naturalist 120(4): 477. At 1053 hours on 23 July 2006 a Bald Eagle, Haliaeetus leucocephalus, was observed in flight transporting in its talons a live Maritime Garter Snake, Thamnophis sirtalis pallidulus, at South Harbour (46°52'01.7"N, 60°26'45.8" W), Cape Breton Island, Victoria County, Nova Scotia. Key Words: Bald Eagle, Haliaeetus leucocephalus, predator, Maritime Garter Snake, Thamnophis sirtalis pallidulus, South Harbour, Cape Breton Island, Nova Scotia. The Bald Eagle, Haliaeetus leucocephalus, is a large North American raptor that usually frequents aquatic environments where it feeds predominantly on cap- tured or scavenged fish. Eagles may be seen daily in South Harbour, at the northern end of Cape Breton, Nova Scotia. This area appears to be occupied by a resident pair based on an observation of two adults seen standing very close together on a mud flat at the mouth of the harbour on 16 July 2006. South Harbour is a Shallow, brackish, nearly landlocked estuary with an artificially maintained channel through the sandbar across its mouth providing exchange of water with the Atlantic Ocean. On 23 July 2006 at 10:53 h, the screams of eagles drew my attention to two adult birds wheeling and skir- mishing in the air over the southern shore of the har- bour (46°52'01.7"N, 60°26'45.8"W). When two birds eventually flew directly overhead I saw that the lead bird was carrying a snake in its talons. The grayish- brown coloration and very attenuated, pointed tail writhing in characteristic ophidian fashion precluded any possibility of the prey being an American Eel (Anguilla rostrata). Two other witnesses (Johanna R. Humphrey, Katherine Allen) also confirmed that the prey, which was very much alive, was a snake. The individual carried by the eagle appeared to be some- what in excess of 600 mm in total length. The only snake of this size in Nova Scotia would be a Maritime Garter Snake, Thamnophis sirtalis pallidulus, females of which range from 412 to 916.9 mm, whereas males do not exceed 602 mm (Gilhen 1984). Because most of the area surrounding South Harbour, especially the southern shore, where the observation was made, is very heavily vegetated, the eagle most likely captured the snake while the latter was swimming across the surface of the water. Maritime Garter Snakes are “excel- lent swimmers” and have been seen on “freshwater lakes 100 metres or more from shore” (Gilhen 1984: 132). The Bald Eagle feeds almost exclusively on verte- brates, paticularly on fish, followed by birds, and then mammals (Stalmaster 1987; Buehler 2000). Although Stalmaster (1987: 93) emphasized the eagle’s “vast diversity of prey’, he considered that snakes “are an un- usual dietary item” (pages 95-96) and cited only the report of McEwan and Hirth (1980) of the remains of a single Black Swamp Snake, Seminatrix pygaea, in a nest in central Florida. Bald Eagles are notorious scay- engers, and large numbers of snakes, including Semi- natrix, are victims of vehicular traffic in central Florida (Smith and Dodd 2003), so there is a strong possibility that that food item was obtained as road kill. I have found no other references to Bald Eagles feeding on snakes, although they probably do so opportunistical- ly wherever snakes and eagles occur together. Literature Cited Buehler, D. A. 2000. Bald Eagle Haliaeetus leucocephalus. In The Birds of North America (506: 1-40). Edited by A. Poole and F. Gill, The Birds of North America, Inc., Philadelphia, Pennsylvania. Gilhen, J. 1984. Amphibians and reptiles of Nova Scotia. Nova Scotia Museum, Halifax. McEwan, L. C., and D. H. Hirth. 1980. Food habits of the Bald Eagle in north-central Florida. Condor 82: 229-231. Smith, L. L., and C. K. Dodd. 2003. Wildlife mortality on U.S. Highway 441 across Paynes Prairie, Alachua County, Florida. Florida Scientist 66: 128-140. Stalmaster, M. V. 1987. The Bald Eagle. Universe Books, New York. Received 8 August 2006 Accepted 16 February 2007 478 THE CANADIAN FIELD-NATURALIST Vol. 120 Rat Poison Kills a Pack of Eastern Coyotes, Canis latrans, in an Urban Area JONATHAN G. Way!, STEPHEN M. CiFuN?, Davip L. EatouGH?, and Eric G. STRAUSS * ‘Environmental Studies Program and Lynch School of Education, Boston College, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA; e-mail: jw9802@yahoo.com Present address: Eastern Coyote Research, 64 Cranberry Ridge Road, Marston Mills, Massachusetts 02648 USA 232 Maple Street, Revere, Massachusetts 02151 USA 3Science Department, Revere High School, 101 School Street, Revere, Massachusetts 02151 USA 4Urban Ecology Institute, Environmental Studies Program, and Biology Department, Boston College, Higgins Hall, Chestnut Hill, Massachusetts 02467 USA Way, Jonathan G., Stephen M. Cifuni, David L. Eatough, and Eric G. Strauss. 2006. Rat poison kills a pack of eastern Coyotes, Canis latrans, in an urban area. Canadian Field-Naturalist 120(4): 478-480. We document the death of a pack of Eastern Coyotes (Canis latrans) from high levels of brodifacoum, a second generation poison that is the active ingredient in some forms of rat poison (e.g., d-Con®). The Coyotes died within a week of each other during late March/early April 2005. This incident indicates the vulnerability of wild animals to commercial over-the- counter rodenticides. Key Words: Canis latrans var., Eastern Coyote, anticoagulant, brodifacoum, poison, rat poison, Massachusetts. Coyotes (Canis latrans) live successfully in a vari- ety of habitats ranging from rural to urbanized areas (Gese et al. 1996; Harrison et al. 1991; Patterson and Messier 2001; Riley et al. 2003; Way et al. 2001, 2004). However, in almost all of these settings (except nation- al parks; Gese et al. 1996) people constitute the major source of mortality for Coyotes usually via trapping, shooting and automobile strikes (Grinder and Kraus- man 2001; Parker 1995). Poison was historically used to kill predators but was banned in 1973 in the United States (Mech 2000; Mech and Boitani 2003). Mech (1970) noted that “poison is no doubt the most effec- tive and efficient method of controlling or exterminat- ing Wolves (Canis lupus).’ However, the use of poisons is controversial because of their relative non-selectivity and reputation for inhumaneness (Cluff and Murray 1995). Most poisons, such as strychnine, cyanide, and sodium fluoroacetate (compound 1080), are not readily obtainable today. Because these poisons are illegal, many canid populations have greatly increased in the past 30 years (Mech and Boitani 2003; Parker 1995). Anticoagulants are present in urban areas (for rat con- trol), and Riley et al. (2003) found them to be a signifi- cant cause of death for Coyotes in southern Califor- nia. This paper details the poisoning deaths of a family group of Coyotes in urban north Boston, Massachu- setts, most likely deliberately poisoned by someone . Study Area and Methods Coyotes were captured for an ecological study on the north edge of Boston (42.43°N, 71.06°W), in east- ern Massachusetts, in the bordering cities of Everett (4345.0 people/km7), Malden (4290.5 people/km?), and Revere (3089.0 people/km*) (U.S. Census Bureau, 2000 estimates). Box traps were used to capture Coy- otes (Way et al. 2002a) except for one (“Jet”) cap- tured via a ground-based netlauncher (Coda Enterpris- es, Inc., Mesa, Arizona). Four Coyotes were radio-col- lared in this pack: “Maeve” (#BN0404), a 14.5 kg lactating female, captured 17 May 2004, was the breed- ing female; “Jet” (#BN0403), a 15.9 kg breeding male, captured 29 June 2004, was Maeve’s mate; “Jem” (#BN0406), a 10.0 kg 4.5 month-old pup, was captured on 26 August 2004; and “Cour” (BN0405), a 12.3 kg 5 month-old pup, was captured on 15 September. The behavior of Maeve and Jet (i.e., frequently being locat- ed with each other and with the pups, including obser- vations of them feeding the pups) indicated that they were the parents of these pups (see Way et al. 2001). The Coyote pack consisted of two to three adults (1.e., one additional uncollared Coyote occasionally sight- ed in the pack’s territory — its status was never deter- mined but it was probably a helper Coyote [see Way et al. 2002b] to Jet and Maeve) and four pups (two of which were not collared). The group was named the Cemetery Pack as it resided almost exclusively at one green area (including some thicker adjacent woods) surrounding four large connected cemeteries. The entire area was about 2.5 km? and on the north side, aside from unused railroad tracks the pack’s territory was surrounded by high-density housing units and/or com- mercial spaces (malls) on all sides. Results and Discussion The group went from six or seven members in the fall of 2004 to four individuals by mid-winter 2004- 2005 when it was presumed that some of the pack members (two of the pups [including Jem] and proba- bly the uncollared adult) dispersed. Jem was last suc- cessfully located on 11 December 2004. Snow track- ing and sighting data indicate that only one uncollared Coyote (a light yellowish-brown animal) remained along with the breeding pair and Cour. Similarly, Way et al. (2002b) found three to four individuals to be a 2006 typical winter pack size of Coyotes in eastern Massa- chusetts. The winter of 2004-2005 was harsh with much snow yet the Coyotes remained in their small territory. Res- idents often reported to us that they fed the Coyotes and people noticed some of the animals were collared (J. Way, unpublished data), indicating that the group obtained food from people living in and/or around the cemeteries. Aside from their abnormally small home range (see Way et al. 2002b), they behaved much like Coyotes studied in other locations, including the avoid- ance of people by being nocturnal (Way et al. 2004), crossing streets quite often, denning in wooded or rel- atively undisturbed (including under a gravestone) regions of their territory (Way et al. 2001), and acting territorial (Way et al. 2002b), including observations of Jet scent marking on the railroad tracks at the north edge of his pack’s home range. There was no sign of the Coyotes’ ill-health until just prior to them dying. On 27 March 2005 Maeve was found dead in the middle of a cemetery in the central part of the pack’s territory. She was an emaciated 12.7 kg despite ap- pearing normal when sighted on 24 March. A gross necropsy revealed significant internal bleeding and no fetuses, indicating that she was not pregnant. A labo- ratory (Idexx Veterinary Services, www.vetconnect. com) diagnosis indicated erosive acute gastritis, severe necrotizing hemorrhagic endometritis with retained placental decidual tissue, and subacute suppurative endocarditis and myocarditis. On 31 March 2005 Jet was found dead, 100 m from where Maeve died, and was also emaciated (14.3 kg). Radio-telemetry data indicates date of death was 30 March. He was observed moving normally 2-3 days before his death. Because of massive internal bleeding and similar gross necropsy results as Maeve, we only tested for poisoning (specifically for common chemi- cals found in household rat poisons) on Jet. Brodifa- coum was detected in the liver at 0.733 parts per mil- lion (ppm), and the laboratory (Idexx) indicated that the results supported a diagnosis of brodifacoum poi- soning. On 3 April 2005 Cour was found dead in a shallow (< 1 m deep) canal that he frequently (i-e., daily) crossed prior to his death. He appeared healthy and weighed 17.3 kg, heavier than both of his parents. His relatively robust physique support observations from residents indicated that Cour (with a red ear tag) was the radio- collared Coyote most commonly seen eating food left by people. He was previously observed up-close and, besides limping on his right hind leg, appeared healthy on | April 2005. Not having obtained the results from Jet or Maeve at the time, we had a full necropsy per- formed at Tufts University (Grafton, Massachusetts). Internal bleeding (subcutaneous hemorrhage) was noted and based on autopsy findings and toxicological analy- sis of the liver (brodifacoum = 0.542 ppm), Cour died from an anticoagulant rodenticide. Because of the way NOTES 479 Jet and Cour died, and the similar necropsy findings from Maeve (i.e., massive internal bleeding), we con- clude that Maeve also died from brodifacoum poi- soning. Because all three of the Coyotes’ behavior seemed normal prior to their death and that they all died close together (< 1 week), it appears that someone purpose- fully poisoned them at high concentrations rather than the coyotes having eaten enough poisoned prey to have died (i.e., from bioaccumulation; Riley et al. 2003). Most likely Maeve and Jet were poisoned around the same time, and then Cour was given a later dose(s) judging by a sighting of him traveling alone after Maeve and Jet were documented as dead. However, Cour’s healthier condition might have allowed him to survive longer than his parents. We extensively searched for the source of the poisons (especially near where the coyotes died and where they spent the majority of their time when they were alive), including informal- ly talking to numerous people, but we never managed to locate any substantial leads. Throughout summer 2005 only a couple of sightings were made by residents and cemetery staff indicating that either the uncollared Coyote survived or, more likely, a new Coyote (i.e., formerly not part of this pack) dispersed into this location. Regardless of those individual sightings, this pack was decimated in a short period of time via rodenticides. The public should be better informed of the dangers that common household poisons present for wildlife, especially in urbanized areas, and the potential health threat to humans and pets. For example, about two months later, pet dogs (Canis familiaris) died or were injured from rat poison in neighboring towns (S. Cifu- ni, personal communication). The slow, painful death of an animal internally bleeding is not pleasant and should not be acceptable in our society. These animals were diagnosed because they were part of an ecological study; due to the expense of testing and difficulty of finding non-radio-collared animals that die in the woods, it would be difficult to estimate how many non- target (i.e., not rats or mice) animals die from antico- agulants. We recommend that these poisons be strictly controlled either through making the over the counter sale of them illegal to all but licensed, professional exterminators and/or through required public education campaigns explaining the dangers of these poisons. Acknowledgments This study would not be possible without the sup- port of the Way and Cifuni families, N. Lima, the Saugus Animal Hospital and Dr. Bob Binder, Fiore’s Meat Market and The Meat Market (in Winthrop), D- L. Szumylo, the Environmental Studies Program and Urban Ecology Institute at Boston College. Numer- ous private individuals donated to have the necrop- sies performed and numerous residents in the area of the poisonings helped out in myriad ways. Care and 480 capture of free-ranging Coyotes was approved by Boston College’s Institutional Animal Care and Use Committee Protocol Number 01-02 (renewed in 2005) and by the Massachusetts Division of Fisheries and Wildlife permit #003.04LP. Literature Cited Cluff, H. D., and D. L. Murray. 1995. Review of wolf con- trol methods in North America. Pages 491-504 in Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S. H. Fritts, and D. R. Seip. Canadian Cir- cumpolar Institute, Occasional Publication Number 35, Edmonton, Alberta, Canada. Gese, E. M., R. L. Ruff, and R. L. Crabtree. 1996. Foraging ecology of coyotes (Canis latrans): the influence of extrin- sic factors and a dominance hierarchy. Canadian Journal of Zoology 74: 769-783. Grinder, M., and P. R. Krausman. 2001. Morbidity-mor- tality factors and survival of an urban coyote population in Arizona. Journal of Wildlife Diseases 37: 312-317. Harrison, D. J., J. A. Harrison, and M. O’Donoghue. 1991. Predispersal movements of coyote pups in eastern Maine. Journal of Mammalogy 72: 756-763. Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. 1995, Reprint. University of Minneso- ta Press, Minneapolis, Minnesota, 384 pages. Mech, L. D. Editor. 2000. The wolves of Minnesota: howl in the heartland. Voyageur Press, Stillwater, Minnesota. 128 pages. THE CANADIAN FIELD-NATURALIST Vol. 120 Mech, L. D., and L. Boitani. Editors. 2003. Wolves: behav- ior, ecology, and conservation. University of Chicago Press, Chicago, Illinois. 448 pages. Parker, G. R. 1995. Eastern coyote: the story of its success. Nimbus Publishing Halifax, Nova Scotia, Canada, 254 pages. Patterson, B. R., and F. Messier. 2001. Social organization and space use of coyotes in eastern Canada relative to prey distribution and abundance. Journal of Mammalogy 82: 463-477. Riley, S. P. D., R. M. Sauvajot, T. K. Fuller, E. C. York, D. A. Kamradt, C. Bromley, and R. K. Wayne. 2003. Effects of urbanization and habitat fragmentation on bob- cats and coyotes in southern California. Conservation Biol- ogy 17: 566-576. Way, J. G., P. J. Auger, I. M. Ortega, and E. G. Strauss. 2001. Eastern coyote denning behavior in an anthropogenic environment. Northeast Wildlife 56: 18-30. Way, J. G., I. M. Ortega, P. J. Auger, and E. G. Strauss. 2002a. Box-trapping eastern coyotes in southeastern Massachusetts. Wildlife Society Bulletin 30: 695-702. Way, J. G., I. M. Ortega, and P. J. Auger. 2002b. Eastern coyote home range, territoriality and sociality on urbanized Cape Cod, Massachusetts. Northeast Wildlife 57: 1-18. Way, J. G., I. M. Ortega, and E. G. Strauss. 2004. Movement and activity patterns of eastern coyotes in a coastal, subur- ban environment. Northeastern Naturalist 11: 237-254. Received 4 October 2005 Accepted 7 April 2007 A New Record of Deepwater Sculpin, Myoxocephalus thompsonii, in Northeastern Alberta M. STEINHILBER! and D. A. NEELY 'Royal Alberta Museum, 12845 102nd Avenue, Edmonton, Alberta TSN 0M6 "Department of Biology, St. Louis University, St. Louis, Missouri 63103 USA Corresponding author: e-mail: dave.neely @ gmail.com. Present address: Department of Ichthyology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103 USA Steinhilber, M., and D. A. Neely. 2006. A new record of Deepwater Sculpin, Myoxocephalus thompsonii, in northeastern Alberta. Canadian Field-Naturalist 120(4): 480-482. We present the first documented records of Deepwater Sculpin, Myoxocephalus thompsonii, from northern Alberta, and the sec- ond record for the province. Three specimens of Deepwater Sculpin were taken in gill nets set at 17 to 20 m depth in Colin Lake, Alberta, on 15 September 2001. Colin Lake, located in the Canadian Shield region of northeastern Alberta about 125 km north- east of Fort Chipewyan, drains into Lake Athabasca via the Colin River. The only other known Alberta population of Deepwater Sculpin inhabits Upper Waterton Lake in the southwestern corner of the province. This record is approximately 300 km SSE of the nearest verified record in the Northwest Territories and 400 km NW of the nearest verified record in Saskatchewan. Key Words: Deepwater Sculpin, Myoxocephalus thompsonii, distribution, Colin Lake, Alberta Three individuals of Deepwater Sculpin, Myoxoce- phalus thompsonii, were collected in two separate gill net sets in Colin Lake, Alberta (59°34'N, 110°08'W) on 15 September 2001. One set was in 17 m of water, and fished for a period of 15 hours. The other was set at 18-20 m depth for 14 hours. Each net set consisted of one 60 m multi-mesh net with six 10 m panels rang- ing from 10 to 25 mm bar mesh and one 60 m net with six 10 m panels of 10 to 50 mm bar mesh. All nets were 1.8 m in depth. The two nets were fished in series on the bottom of the lake. Both collecting sites were locat- ed on the gently sloping periphery of the two deep basins in the lake (Figure 1). The maximum depth of the lake is approximately 25 m. The composition of the substrate at the sampling sites was not determined. Species taken syntopically with the sculpins included Lake Whitefish (Coregonus clupeaformis), Cisco (Co- regonus artedi), Burbot (Lota lota), Northern Pike 2006 Colin Lake 1 2%, #3 kilometers Colin River NN (drainage to Lake Athabasca) * FiGuRE |. Bathymetric map of Colin Lake, with Deepwater Sculpin capture localities indicated. Depths given in metres. (Esox lucius), Lake Trout (Salvelinus namaycush), and Longnose Sucker (Catostomus catostomus). Lake Whitefish, Cisco, and Longnose Suckers were the most common species. All three specimens of Deepwater Sculpin are gravid females, ranging from 62-74 mm SL (Standard Length), with four prominent, large, and straight preopercular spines, a large gap between the dorsal fins, and the gill covers free at the isthmus. Meristic counts of these specimens are consistent with data presented by McAI- lister (1961) for adjacent populations in Saskatchewan and the Northwest Territories. Dorsal fin spines were (number of specimens in parentheses) 7(2) or 8(1); dorsal fin rays 14(1), 16(1), or 17(1); anal fin rays 14(1), 15(1), or 17(1); pelvic rays 3(6); pectoral rays 16(2) or 17(4). Two of the three had an incomplete lateral line (ending under the last dorsal fin ray) with 32 pores; in the remaining specimen the lateral line continued posteriad to the caudal peduncle and had 36 pores. These specimens represent the first record of Deep- water Sculpin from northern Alberta, and only the sec- ond record for the province (Roberts 1988). The only other known population in Alberta occupies Upper Waterton Lake (49°02'N, 113°54'W) on the Alberta/ Montana border (McAllister and Ward 1972) (Figure 2). Nelson and Paetz (1992) suggested the likely pres- ence of Deepwater Sculpin in northeastern Alberta, given the proximity of populations in adjacent Saskat- chewan and the Northwest Territories. The closest pop- ulations supported by voucher specimens are Great Slave Lake, Northwest Territories (~300 km NNW) and Wollaston (400 km SE) and Reindeer Lakes (520 km SE), Saskatchewan (COSEWIC 2006*). Records from Lake Athabasca in northwestern Saskatchewan (~100 km ESE, COSEWIC 2006*) are not supported by voucher specimens. The specimens from Colin Lake fill this gap, and further suggest that additional popula- tions of this enigmatic cottid remain undiscovered in the northern portions of its range. New populations con- NOTES 451 Re ho 100"W FIGURE 2. Distribution of Deepwater Sculpin in western Canada (data from Scott and Crossman 1973, and Murray et al. 2003). Colin Lake records indicated by open circle. tinue to be discovered as deepwater habitats are system- atically sampled (i.e., Murray et al. 2003; COSEWIC 2006*). Although typically encountered at depths greater than 44 m (Scott and Crossman 1973), this species is occasionally found at much shallower depths, espe- cially as larvae and juveniles (COSWEIC 2006 ); Raw- son (1951) collected specimens in | m of water in Great Slave Lake, while McPhail and Lindsey (1970) docu- mented specimens from 4-6 m in Great Bear Lake. Alex Peden (in McPhail and Lindsey 1970) suggested that this fish is probably concentrated in deep water during the summer and moves inshore in the fall. The discoy- ery of Deepwater Sculpin in Colin Lake is thus some- what unusual in that the lake is not particularly deep, with a maximum depth of only 25 m. Further studies to determine seasonal distribution patterns within the lake and any unique biological characteristics of this population would be valuable. It is worth noting that Colin Lake is connected via the Colin River to the Saskatchewan side of Lake Athabasca (approximate- ly 20 km to the SE) where depths can reach 124 m. It is likely that trapping efforts in deepwater habitats of Lake Athabasca will provide additional records. How- ever, there are currently no data to support the use of the Colin River as a post-glacial movement corridor between Colin Lake and Lake Athabasca. The only populations of Deepwater Sculpin current- ly given protective status in Canada are the Great Lakes — Western St. Lawrence populations, which were list- ed as Threatened by the Committee on the Status of Endangered Wildlife in Canada [COSEWIC] (Parker 1988) but have recently been downgraded to Special Concern (COSEWIC 2006 ), and are on schedule 2 of 482 the Canadian Species At Risk Act (SARA). Western populations (which include the new population dis- cussed herein) were designated Not at Risk in 2006 (COSEWIC 2006’). The gill nets used to capture these Deepwater Scul- pins were deployed as part of a Shortjaw Cisco (Core- gonus zenithicus) survey in the Canadian Shield region of Alberta (Steinhilber 2004"). Our recent experience suggests that deployment of cyalume light stick-bait- ed wire minnow traps in deepwater habitats is more effective at sampling Deepwater Sculpin than are small- mesh gill nets. The effectiveness of this technique has not been previously discussed in the literature. Long- term surveys of Lake Saganaga on the Ontario/Min- nesota border have primarily used small-mesh gill nets (Etnier and Skelton 2003). Between 1986 and 2003, only one specimen of M. thompsonii was taken. Use of light stick-baited cylindrical wire minnow traps during summer 2004 per our request resulted in cap- ture of 10 specimens (D. A. Etnier, personal commu- nication). Similarly, during 2003 we provided Fish- eries and Oceans Canada personnel on Lake Nipigon with light sticks and traps that were set simultaneously with their gill nets, with similar results (A. Van Ogtrop, personal communication). Other researchers working on Deepwater Sculpins have had similar experiences (T. Sheldon, personal communication; COSEWIC 2006"). We thus strongly recommend the use of this technique over small-mesh gill nets for assessing pres- ence/absence of Deepwater Sculpin. Acknowledgments We thank B. Meagher for field assistance. Collect- ing was conducted under permit 401 FR issued to M. Steinhilber. D. Neely was supported by a grant from the British Columbia Ministry of Fisheries. Materials Examined All specimens are deposited in the holdings of the Royal Alberta Museum, Edmonton. PMA L01.33.319, Myoxocephalus thompsonii, n=1 (74.0 mm SL), Colin Lake, 59°33'12"N, 110°9'30" W. 15 September 2001. Collectors: M. Steinhilber, B. Meagher. PMA LO1. 33.311, M. thompsonii, n=2 (62.7-73.0 mm SL), Colin Lake, 59°33'28"N, 110°5'8" W. 15 September 2001. Collectors: M. Steinhilber, B. Meagher. THE CANADIAN FIELD-NATURALIST Vol. 120 Documents Cited (marked * in text) COSEWIC. 2006. COSEWIC assessment and update status report on the deepwater sculpin Myoxocephalus thomp- sonii (Western and Great Lakes-Western St. Lawrence pop- ulations) in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. 39 pages. (www.sararegistry. gc.ca/status/status_e.cfm). Steinhilber, M. 2004. Shortjaw cisco species at risk assess- ment 2001. Alberta Sustainable Resource Development, Fish and Wildlife Division, Alberta Species at Risk Report Number 82. Edmonton, Alberta. 21 pages. Literature Cited Etnier, D. A., and C. Skelton. 2003. Analysis of three Cisco forms (Coregonus, Salmonidae) from Lake Saganaga and adjacent lakes near the Minnesota/Ontario border. Copeia 2003: 739-749. McAllister, D. E. 1961. The origin and status of the deepwa- ter sculpin, Myoxocephalus thompsonii, a Nearctic glacial relict. National Museum of Canada Bulletin 172: 44-65. McAllister, D. E., and J. C. Ward. 1972. The deepwater sculpin, Myoxocephalus quadricornis thompsoni, new to Alberta, Canada. Journal of the Fisheries Research Board of Canada 29: 344-345. McPhail, J. D., and C. C. Lindsey. 1970. Freshwater fish of northwestern Canada and Alaska. Bulletin 173, Fisheries Research Board of Canada, Ottawa, Ontario. 381 pages. Murray, L., M. H. Papst, and J. D. Reist. 2003. First record of the Deepwater Sculpin, Myoxocephalus thomp- sonii, from George Lake in Whiteshell Provincial Park, Manitoba. Canadian Field-Naturalist 117: 642-645. Nelson, J. S., and M. J. Paetz. 1992. The fishes of Alberta. 2™4 edition. The University. of Alberta Press, Edmonton. 437 pages. Parker, B. J. 1988. Status of the deepwater sculpin, Myoxo- cephalus thompsoni, in Canada. Canadian Field-Naturalist 102: 126-131. Rawson, D. S. 1951. Studies of the fish of Great Slave Lake. Journal of the Fisheries Research Board of Canada 8: 207-240. Roberts, W. E. 1988. The sculpins of Alberta. Alberta Nat- uralist 18: 121-127; 153. Addendum 19:35. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Bulletin 184, Fisheries Research Board of Cana- da, Ottawa, Ontario. 966 pages. Received 29 December 2005 Accepted 27 March 2007 Book Reviews ZOOLOGY The Clements Checklist of Birds of the World 6th Edition By James F. Clements. 2007. Cornell University Press, Sage House, 512 East State Street, Ithaca, New York 14850 USA. 864 pages. U.S.$59.95. Cloth. If you travel outside of North America [or any other home territory] you will have to face the question. Is this bird species one I have seen before? There are so many look alike birds that are different species in some areas and the same species in others. You can wonder about crows, oystercatchers, stilts and the plethora of wagtails. Or is this bird with the same or a similar name to a look-alike home species, the same or a differ- ent bird. For example are Willow Grouse, Red Grouse and Willow Ptarmigan the same species? It can be real- ly confusing. Enter Dr. James Franklin Clements [1927-2005], an orphan who served in the U.S. merchant marine and navy before going to college. In the Korean war he was in the air force, before finally settling to get his Ph.D. in 1975. His thesis became the first edition of a check list of the world’s birds. Because of ever-evolving tax- onomy we are now at the 6th edition. Clements has produced a list, simply a list, of all the known bird species and subspecies. Producing such a simple list is a very complex task indeed. It must have felt like trying to resolve the unresolvable on many occasions. So here we have this simple list and can ask how useful is it? Theoretically, I should be able to check out a bird that I am unsure about. Looking it up in Clements should allow me to clarify the bird’s status and decide whether it can be added to my life list or not. How well does this work? The first problem is you must use the same English name as Clements. Nowhere is there a list of alternative names. If you look for Melba Finch in the index you will be disappointed. You will need to use either Green-winged Pytilia or look for the genus Pytilia. The North American bias is shown by Circus cyaneus being called Northern Harrier as op- posed to Hen Harrier [the older name]. Now we come to the sticky issue of splits and lumps. The North American Magpie is split from its European counterpart, as is Hooded and Carrion Crow. Clements lists the Somali Ostrich as a subspecies, Struthio cam- elus molybdophanes, whereas most recent literature gives this distinctive blue-necked bird separate species status [Struthio molybdophanes}. Similarly the bright- ly coloured barn swallow found only in the Nile delta is not split from its more common cousin. Again this is listed by some as a separate species, Egyptian Swal- low, Hirundo savignii. The vegae Herring Gull is sep- arated as East Siberian Gull. This is open to question and recent work suggests more research is needed. Certainly I have found it difficult to see a difference in the field. Is it a little darker than a Herring or was I being deluded by poor light? Each entry is accompanied by a description of the species range. These ranges are useful when compar- ing species. For example, my list for Mongolia includes Pink-footed Goose, Anser fabalis. The range for Pink- footed Goose in Clements does not include Mongolia, whereas the Bean Goose, correctly Anser fabalis, is present. Clearly this is an error in the Mongolian list, probably due to translation problems, that was quick- ly resolved. However, the ranges given for other species are less useful. Clements can be forgiven for minor lapses such as the range of the Northern Wheatear or Crested Auk- let — given as Siberia. The birds I saw on the eastern- most coast of Russia this year were technically in the Russian Far East — not Siberia. Siberia does not extend to the east coast. This is minor compared to Houboura and Macqueens Bustards. Clements splits these two birds and gives Macqueens a range of the Nile to .... Pakistan. The Handbook of the Birds of the World, Volume 3, cited as the major reference, does not split this species but lists it as a subspecies, Chlamydotis undulata macqueenii, and correctly gives the range as NW Kazakhstan to .... Mongolia and Northern China. Similarly Clements does not mention the range exten- sion of Cardinal and House Finch into eastern Canada. I have checked to see if some of the recent discover- ies are included in this check list. Scarlet-banded Bar- bet and Long-whiskered Owlet are, but the two most recent discoveries, Large-billed Warbler [the 1867 refer- ence is given but not the recent March 2007 capture of two birds] and Gorgeted Puffleg, [May 2007] are not. Having one coherent checklist is extremely valu- able for the travelling birder. It still does not solve all the issues with a changing taxonomy, but at least you can use it as a solid foundation for your records. You will still have to research current taxonomy and recent discoveries and decide which birds you will accept as a full species. Incidentally, I make it a point never to directly mark books. I use post-it notes or equivalent. I will make an exception for this one book. Not only is it set up to be 483 A84 ticked, but it is a good idea to add information on species as new data become available. That is until version seven is printed. THE CANADIAN FIELD-NATURALIST Vol. 120 Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Birds of Europe, Russia, China and Japan: Passerines, Tyrant Flycatchers to Buntings By Norman Arlott. 2007. Princeton University Press, 41 Wil- liam Street, Princeton, New Jersey 08540-5237 USA. 240 pages. U.S.$29.95. Paper. My wife and I recently decided to visit northern China and Mongolia [the Silk Road]. This left me with a dilemma; which bird book or books would I need to take to get adequate coverage. I did not want to pay $100 or more for a paperback copy of A Field Guide to the Birds of China (John MacKinnon and Karen Phillips, Oxford University Press, New York). My Birds of Russia (Flint et al.) gives fair coverage but the illustrations are not as precise as I would like them, particularly for warblers, etc. Birds of China (De Schauensee) only illustrates some of the species and gives descriptions of range only and you need a thor- ough knowledge of China’s geography to follow these descriptions. Birds of India (Grimmett et al.) is a much better guide with a lot of species covered [as winter visitors], but the coverage is far to the south so the range maps are useless. Then Princeton announced Birds of Europe, Russia, China and Japan and I thought it would be just right for my purposes. While the book covers only the passer- ines (from Tyrant Flycatchers to Buntings), it is exactly these groups for which I needed a better, more up-to- date book. The non-passerines, like ducks and gulls, tend to be more distinctive and are suitably covered in previous books. Technically Princeton calls this book an illustrated checklist. Each species has an illustration or two with very brief facing-page text. At the back of the book are range maps. While the text presents little about each species but key features, it is perfect for the traveller. It measures only 5" x 7" x 4" [12.5 x 17.5 x 2 cm] and weighs 14 oz [100 g or around half a normal field guide]. The range maps are 2" x %" [5 x 2 cm] and cover from Britain to Kamchatka, from Svalbard to North- ern India. As there are no country boundaries [these would make the little map too cluttered] I found it helpful to create a mask with a piece of clear plastic sheet. I traced the outline of the map in blue, then, using Google Earth as a guide [Google Earth has a similar orientation to these maps] I shaded in transparent yel- low the region I plan to visit. I can now place the mask over each map and quickly see if I will be in any bird’s range. This was somewhat satisfactory. I compared my list created with the mask to actual trip lists and I was in error by around 15 percent. Although the book’s title suggests it covers Europe, Russia, China and Japan, the maps go much further. For example it includes the Nile Valley Sunbird of the Middle East and the Purple Sunbird of India, as well as the geographically restricted Sinai Rosefinch and Sub-Sahara’s Blandford’s Lark. I am not sure of the author’s rationale for including birds like the Rosy- patched Shrike, a purely African species [but I am hap- py he did so]. The critical illustrations are those of difficult to iden- tify birds such as warblers and some finches. Arlott has been illustrating bird books for many years [my first Arlott book is over 40 years old]. His skill has grown dramatically and his most recent work is top quality. My copy has all the illustrated species look- ing a little dark. [Technically the hue is correct but the illustrations are over saturated and this is part of the printer’s quality control. North American readers will see this when looking at the Cedar Waxwing and Varied Thrush illustrations. Both are way more intense than the real birds I have seen this year.] In detail though the artwork is excellent. When I looked at Arc- tic Warbler and try to separate it from it congener leaf warblers or the Beautiful Rosefinch from its relatives then the relevant field marks are apparent. North Amer- icans will have no difficulty identifying wood warblers using this guide, even with the intense colours. [I was surprised to find over 40 “North American” species, such as Wood Warblers and Icterids, have been seen in this region as vagrants. ] This book solved my problem and will be ideal for other travelers to this large region. I look forward to a similar book on the non-passerines. I hope Princeton does well with their checklist series [I believe they have a dozen titles] as they are so much more convenient when luggage weight is critical. Literature Cited Flint V. E., R. Boehme, Y. Kostin, and A. Kuznetsov. A Field Guide to Birds of Russia and Adjacent Territories. Princeton University Press, New Jersey. De Schauensee, Rodolphe Meyer. The Birds of China. Smithsonian Institution Press, Washington. Grimmett, Richard, Carole Inskipp, and Tom Inskipp. Birds of India, Pakistan, Nepal, Bangladesh, Bhutan, Sri Lanka and the Maldives. Princeton University Press, New Jersey. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada 2006 Deep Alberta. Fossil Facts and Dinosaur Digs By John Acorn. 2007. The Royal Tyrrell Museum and The University of Alberta Press, Edmonton, Alberta Canada. XII + 186 pages. $26.95. Early on the recognition of our country’s rich re- sources was often a reactive response, rather than proac- tive endeavours. It was in response to the U.S. activities, primarily those of Barnum Brown (1873-1963) of the American Museum of Natural History, New York, that the Canadian government in early the 20" century began their vertebrate paleontological activities in the Canadian west. First under the Geological Survey of Canada, followed by the National Museum of Canada (now, Canadian Museum of Nature, Ottawa). These activities were central to what has been called “The Great Canadian Dinosaur Rush” (see David Spalding’s Into the Dinosaurs’ Graveyard, Doubleday Canada, Toronto, 1999). Despite these efforts, and the bountiful treasures col- lected and studied, it wasn’t until the late 1960s that the Province of Alberta, host of the majority of these activities, was able to begin to take control of its own paleontological resources. Provincialism — restriction of outside exploitation — emerged. This occurred with the materialization of the University of Alberta, Edmon- ton, establishing its own paleontological program of education and research, supported by the Vertebrate Paleontology In Alberta conference in 1963 (Univer- sity of Alberta, 1965) in addition to the establishment of the Provincial Museum of Alberta (Edmonton), which opened in 1967. What followed was a slow eruption of discovery, research, tourism and marketing savvy. John Acorn’s Deep Alberta skims the surface of all that is sexy in paleontology in Alberta. And as most will see, there is a lot to witness. Acorn’s guide, based on his CKUA radio series by the same name, is not quite similar to the other popular guides to Alberta’s paleontological wealth, The Land Before Us, The Making of Ancient Alberta by the Royal Tyrrell Museum of Palaeontology (Lone Pine Press, Edmonton, 1994) is more of a complement to its own gallery (somewhat similar to the Royal Saskatchewan Museum’s Earth Sciences gallery guide, Geological History of Saskatchewan by John E. Storer (Govern- ment of Saskatchewan, Regina, 1989)). Nor is it obvi- ously as in-depth as Dinosaur Provincial Park (edited by P. J. Currie, Indiana University Press, Blooming- ton, 2005), a symposium volume summarizing all the Book REVIEWS 485 major taxonomic groups found within the park. Deep Alberta is designed similar to a field-guide, with one- page descriptions and colorful illustrations of species, or non-taxonomic themes. Alphabetical in its listings of themes, creatures, and places, each item listed is given usually a page of text, accompanied by a full page illustration, often a photograph of an exhibit at the Royal Tyrrell Muse- um of Palaeontology. These range from mosasaurs, those snake/lizard derivatives from the marine environ- ment of the Cretaceous Period, the Chasmosaurus, the horned-dinosaur familiar to the fields of Dinosaur Provincial Park, to the singularly rare Atrociraptor, the “savage robber” skull found near Drumheller — only one partial skull of this is known. Of place, from the badlands near Drumheller to the Milk River regions are few. The thematic questions in- clude “How do you Know Where to Dig?”, an often made public query. Yet, surprisingly, a hot topic like extinction is not addressed. Of people, the likes of Barnum Brown and the Sternberg family (three gen- erations of which have worked in Alberta) are briefly brought to the fore. Appended is a list of most of the Ph.D.’s who are currently at play in the paleontological field of Alberta. Those who have contributed research but lack graduate documentation, or who have provided discoveries without institutional support, are not rec- ognized in this summation. Since the discovery by Joseph B. Tyrrell (1858- 1957) over a century ago of an Albertosaurus jaw, later culminating with erection of a provincial museum spe- cifically dedicated to paleontology (The Royal Tyrrell Museum of Palaeontology) and the designation of Dinosaur Provincial Park as a World Heritage Site, the Province of Alberta has ample reason to show off its fossil resources. Acorn’s Deep Alberta is a primer of a primer, a very first introduction to this wealth. Acorn has much to choose from in his summation of pale- ontology of Alberta and will likely be another useful marketing tool to maintain the visibility of this science in the minds of the public. The treatment of these re- sources by other provincial jurisdictions should take note. Alberta is not the sole place for paleontology. There is a lot more to see. TIM T. TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada Gibson’s Guide to Bird Watching & Conservation By Merritt Gibson. 2007. Nimbus Publishing Limited, Hali- fax, Nova Scotia, Canada. 214 pages. $16.46. This small bird watching guide to the maritime prov- inces is a tribute to the birds of Nova Scotia, Prince Edward Island and New Brunswick. It is also a trib- ute to the ornithologists and bird-watchers of the area, the unsung heroes who for the past several decades have been the backbone of bird conservation and study to preserve habitat, protect endangered species and pro- 486 mote birding. The reader is introduced to a variety of birds and a variety of birders following the colourful pages of anecdotes, life histories and illustrated birds. The book does not try to describe all of the birds which regularly visit the maritime provinces of Cana- da. It is not a field guide to all birds. It is a book for bird-watchers and conservationists and introduces the reader to several birds, each of the sea, the coast, the shore, freshwater, the forest, agricultural lands, and finally towns and gardens, in that order. The book is divided into the seven sections named above with an introduction to each section and ending comment intro- ducing birders amateur and professional studying with- in each of the sections and involved in the steward- ship of habitat. Three to seven birds are described in some detail in each section. The description includes anecdotal stories of the birds, their habitats, origins of their common name, behaviour, nesting and egg-laying behaviours and chick hatching and rearing behaviour. Whether one is ready to brave the wind and waves to spot birds, or maybe just sit in an armchair by the win- dow, the text leads you to appreciate the diversity of birds present in the Maritime Provinces and the pos- sibilities of discovering or re-acquaintance with species which have always been present and probably have been visible many times. Conservation is one important focus of the book and the end of each chapter gives some detail on the work of amateurs and professionals who research, monitor, THE CANADIAN FIELD-NATURALIST Vol. 120 and care for the different populations of each bird spe- cies mentioned. Throughout Nova Scotia, New Bruns- wick, Prince Edward Island, and Newfoundland, many different bird populations are in decline, especially but not exclusively, species mentioned in the book. Differ- ent programs are in place to monitor and conserve lands inhabited by these birds, and programs are being devel- oped to make the public aware of the trends in bird pop- ulations. This book is one of those awareness projects. I read the book quickly and then went back over the chapters more slowly to savour the stories of birds I know and the birds mentioned which I would like to see one day. It is an exciting book for a part-time birder and one which taught me not just about the birds but even more about the state of the shoreline, the wet- lands, the forests and the agricultural fields. To the vis- itor, the land changes little, but to the resident, there are problems and potential solutions. The solution is often awareness and information. I applaud the re- searchers for their work and Merritt Gibson for bring- ing it together. The book also serves as a delightful picture book illustrated by Twilia Robar-DeCoste with drawings of birds, birds in their natural settings, land- scapes and seascapes to show perspective and the beau- ty involved in the study and enjoyment of birding on Canada’s East Coast. JIM O’ NEILL 28718 Five Mile Road, Livonia, Michigan 48154 USA Ladybugs of Alberta: Finding the Spots and Connecting the Dots By John Acorn. 2007. University of Alberta Press, Edmon- ton, Alberta, Canada. 169 pages. $29.95. This is an important book about insects for natural- ists. It has field credibility. Acorn reviewed and under- stands the writings of the experts on lady beetle identi- fication and ecology in North America. Acorn studied the specimens that collectors in Alberta deposited in collections. But most importantly, he was in the field in Alberta chasing and watching and photographing lady beetles. The result is a guide, written by an expert, which will be valuable before someone starts chasing lady beetles and will continue to provide insights as a person’s expertise grows. And it is accessible; virtu- ally anyone will be able to use it. What can you do with this book? First and foremost it is possible to identify the lady beetles (a.k.a. lady- bugs, ladybird beetles, Coccinellidae) that live in Al- berta. The combination of excellent colour drawings, photographs, plus key features highlighted in text make it possible. In addition the range maps, both range in Alberta and much of North America, notes on food and habitat preferences give some guidance, to reduce the probability of an embarrassing misidentification. The difficult identifications are not glossed over. The neces- sity of dissecting genitalia to distinguish some Hippo- damia (e.g., glacialis and quinquesignata) and the sig- nificance of chromosomal differences between Chilo- corus stigma and C. hexacyclus are covered clearly, with humour and insight. Most authors would have taken the easy route and restricted themselves to the big showy lady beetles with which most naturalists are familiar. However Acorn tackled the “lesser ladybugs”, a daunting task to make what is known about Scymnus and Hyperaspis and Brumoides widely available, including both features that make species level identifications possible and some natural history. Acorn deserves credit for perse- vering and including the lesser species, especially con- sidering the extra work, almost double the number of species, and that they are not as charismatic to gener- al naturalists and that little is known about them. He engenders interest in these little beetles. In addition to identification and species accounts, there are substantial sections on ecology, behaviour, and history of coccinellid study in Alberta. The hot topic of the influence of introduced species on native species is attacked head on. It starts with a perceptive critique of the functionally useless Nature Canada at- tempt to coordinate a citizen science project on lady beetles in the 1990s. His says that a prejudgment, intro- duced species are bad, led to many of the problems. In my view, the other key problem was that there was little or no quality control that compromised the value 2006 of the data. What was needed was Acorn’s book, or listening to field entomologists, during the planning process. The discussion continues with an assessment of the Evans’ hypothesis that native species have con- stricted their range to ancestral habitats, that is have become less common in extensively human altered habitats (farmland and suburbia) after the arrival of introduced species such as the seven-spotted lady bee- tle. Refreshingly, he is able to present his own data to support his views. His views about ecological change, conservation and invasion biology are clear, logical and worthy of discussion. The style deserves special mention. It is highly cred- ible, yet at the same time there is a breath of whimsy and fireside chattiness. A fine example is the descrip- tion of taste-testing. You will not look at a brightly coloured lady beetle the same after you read this. What bothered me during my initial readings? The first was that I wanted more detail, more data. But, this is a book for generalists and not the appropriate place for the level of detail I want. Second were those cou- plets. Each species account includes a quirky, rhyming couplet. I did not understand many at first, but they did grow on me. Perhaps they are like beer, an acquired taste. Third were the common names. The Entomolog- ical Society of Canada is working to provide consis- tent, standard common names for insects, just like we Book REVIEWS 487 have for birds. However, many were ignored, for exam- ple Mulsantina picta here is the Painted Ladybug, not pine lady beetle. However, some of the newly coined names hit the nail on the head and deserve to be main- tained. My favourite was the Once-squashed Lady- bug, a cryptic species in the same genus as the Twice- stabbed Ladybug. Yes, it needs to be squashed on a microscope slide to examine its chromosomes in order to be identified. Brilliant. These are minor complaints. The identification sections are worth the whole price of the book. The discussion of introduced species is worth the whole price of the book. The description of how to taste a ladybug to assess its palatability is worth the whole price of the book. Our understanding of foods, habitat use and phenol- ogy (seasonal activity) is not as well developed in North America as it is in Great Britain and Europe. People armed with this guide, will have the tools to be at the forefront in remedying this situation. Compilations often function as a catalyst for a quantum leap in inter- est and new findings by curious naturalists. I predict this book will provide another great example in west- ern Canada. DaviID MCCORQUODALE Department of Biology, Cape Breton University, Sydney, Nova Scotia B1P 6L2 Canada A New World of Animals: Early Modern Europeans on the Creatures of Iberian America By Miguel de Asua and Roger French. 2005. Ashgate Pub- lishing Ltd., Aldershot, England. 235 pages. $89.95 How would you describe a beaver to someone who has never seen a beaver? Is it simply a 20 kg rat with thick fur and with something that has the texture and consistency of an elephant’s ear sticking out of its back- side? That person to whom you are giving the descrip- tion might be forgiven for disbelieving you. What about a whole forest of new animals including muskrats, otters, woodchucks, skunks and any of the larger for- est-dwelling species? In North America we have an abundance of animals which were unknown in the Old World before European exploration. Combine these animals with their South American counterparts and add birds, reptiles, amphibians, and insects known only to the New World and you have the basis for the book given to us from Miguel de Asua. For the history buff and the technophile who wants to track down the classic literature of the conquest of the Americas, this book is a new door to obscure liter- ature of natural history. Miguel de Asua is an Argentin- ian professor at the Universidad Nacional de San Mar- tin and Roger French was English, from the department of History and Philosophy of Science at Cambridge University, UK. Their work focuses on the animals found by early explorers of the Americas, particularly South America though reference is made to animals occurring in North America as well. The reader is introduced to many of the early writers describing the newly discovered lands, and emphasis is placed on the animals, birds, reptiles, amphibians and insects which the authors noted at the time. From the earliest explorers, Columbus and his fol- lowers, the discovery of the New World has been a dis- covery of the beauty and ferocity of many new and exciting creatures. Columbus returned with birds, skins and many stories trying to describe the richness of nature which he found and impress his audiences with tales of strange and exciting beasts. Written accounts which survived from his time sought to identify the animals, birds and reptiles using the experience of known animals and the descriptions are interesting associations with European and Asian animal forms put together to construct forms which would be with- in the experience of the audience. So bison looked like camels or cows, armadillos looked like striped foxes and manatees looked like large puppies. For two hundred years the animals of the New World were exotic, unique and objects of wonder in Europe. Even as the years rolled on with more and more litera- ture being accumulated from explorers, naturalists and then colonists with animals imported into Europe as exotic pets and zoo specimens, the earliest descrip- tions survived and were repeated. De Asua takes us on a literature search, naming writers, quoting their writ- 488 ings and showing the parallels as information, disin- formation and wild speculation was accumulated, re- peated and refuted. This is a new study of scientific literature because of its origins and perspectives. First of all it is centred on South America with only some brief mentions of North American mammals. Secondly it is restricted to the writers first in Spanish then in central Europe, all of whom wrote in their own language for audiences in their local areas. Only toward the end of the literature covered in the book are we introduced to English sources though the names are obscure here as well. A good conclusion is added where the ideas of the sailors, soldiers, functionaries, friars and adventurers each had their day and their utility in telling a part of the natural Penguins of the World By Wayne Lynch. 2007. Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 175 pages. U.S.$24.95. Paper. If you take one of the most photogenic birds in the world, living in the most pristine habitat and send forth a remarkable photographer, can you fail? No! The hard part about penguins is getting to where they live. Then anyone can point- and-shoot and get a fine photo. The appearance of penguins is so charming that failure is not an option. A quick thumb through the photos in this book will show how Lynch can take such a good photo opportunity and make the result mag- nificent. But let us look at the text first. In a smooth flowing narrative the author takes us through the lives of the world’s 17 penguin species. He covers the origins and ancestry of this long-lived family. Most intriguing is Lynch’s description of a five foot seven inch prehistoric bird. Now that would have been a sight! The author ex- plains why penguins are shaped the way they are and the uniqueness of their adaptation to sea life. He des- cribes their habitat. You may be surprised to realise that most species live in the temperate [not polar] region. With humour, he explains their sex lives and the advan- tages of their marriage and divorce systems. He follows their lives from egg to adulthood. In all of this he make these birds sound human. Or is it that we humans are like penguins — after all they have a much longer lineage? From sex to family life, from feeding to predators the author weaves a delightful tale of the delightful creatures. In addition you will get to know some of the penguins, sometimes less-than-delightful, neighbours. Only a photographer as good as Lynch could make a THE CANADIAN FIELD-NATURALIST Vol. 120 history story. The opinions and wisdom of the native peoples of the Americas are not given much consider- ation because of course they did not write to Europe and the conquerors formed their own opinions based on the utility of the natural wildlife to their own needs, not the inherent worth of the ages of native wisdom. This is neither an easy book to read nor a simple his- torical storybook. It is a history of intellectual thought and discovery. There are many gems of thought in- cluded but the reader needs to be attentive to the the- sis of the writer not simply to the many repetitions of the details. Jim O’ NEILL 28718 Five Mile Road, Livonia, Michigan 48154 USA pair of Sheathbills look cuddly. He also adds a chap- ter on the northern counterpart of penguins, the alcids. An appendix lists all the penguin species, with a range map and photo for each one. This is not a field guide as there is no species description. There is, how- ever, a small photo of an adult. The range maps and distribution notes are clear and useful. Lynch notes the population size and the species status. These range from a scant 1000 or more pairs of Galapagos Pen- guins to 10 million Macaronis. The photographs are wonderful. Many of them are so balanced in format as to look posed. This speaks of the infinite patience and possibly some good luck that Lynch had on his trips. One notable point is the sun- shine. In my trips to polar regions I rarely see the sun, yet many of his photos bask in glorious light — more patience I think. There is a photo of a braying penguin showing the backward-facing spikes on his tongue in remarkable detail. There are numerous portraits of chicks, with and without parents, that are irresistibly charming. I have seen 10 out of the 17 living species. To see six of the remaining species I must visit New Zealand and the islands that lie to its south, a difficult and very ex- pensive area to reach. But the lure for a penguinophile is undeniably very strong. However, if you have never seen wild penguins and are not likely to do so, then this book is the next best thing. Buy it and enjoy it, for only the smell is missing. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada 2006 BOOK REVIEWS 489 Raptors of Western North America and Raptors of Eastern North America By Brian K. Wheeler. Princeton University Press, Prince- ton, 41 William Street, Princeton, New Jersey 08540 USA. Hardcover. 544 pages and 430 pages, respectively. U.S.$29.95. The hardcover editions of these two volumes were reviewed in Canadian Field-Naturalist 118(1): 142-143 by Houston and John. These books have now been is- sued as paperbacks, reducing their cost [U.S. $49.50 Cloth] and weight. There are no updates or additions. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6KS Canada The Sand Wasps: Natural History and Behavior By Howard E. Evans and Kevin M. O’ Neill. 2007. Harvard University Press. Cambridge, Massachusetts, USA. 340 pages. U.S.$49.95 Cloth. As a baby-boomer “bugster’, two writers had a tre- mendous effect on my early enthusiasm for insects: Edwin Way Teale and Howard Ensign Evans. Sadly, both have passed on, but their legacies live on, and it was with pleasure that I agreed to review a book on the sand wasps (the subfamily Bembicinae of what is now the family Crabronidae, formerly included in the Sphecidae) coauthored by Evans, and created from his notes, ably revised and updated by his former student and colleague Kevin O’ Neill. As one might expect, this is indeed a book for spe- cialists. However, it is also clearly written, and a model for studies of how comparative natural history and be- haviour can be informed by phylogeny and brought together in a synthetic fashion. If you have read Wasp Farm, Evans’ 1963 classic of popular science writing, you understand what this is all about. If not, the first two paragraphs of the introduction to Sand Wasps provide a concise, if not downright brilliant, justifica- tion for this sort of work. Later, on page 5 in the section on “Sand Wasp Natural History/Sand Wasp Science” the authors provide a very fine defense of natural his- tory in general. The overall purpose of the book is to update Evans’ previous works on the sand wasps, and it’s clear that to appreciate all of the details of the pres- ent volume, one really should have read the works that preceded it. But having skipped that step, I still found the book a pleasant and interesting read. The book begins by putting the sand wasps in con- text, both with respect to their biology and their clas- sification. After that, chapter-length summaries are provided of what is known about the wasps, on a tribe- by-tribe basis. I found myself getting a bit bogged down in details during Chapters 2-7, but a student of sand wasp biology would find them riveting, I’m sure. My only criticism of this part of the book, which lays out the evidence to be synthesized in the remaining chapters, is that no summary is provided of the clas- sification used, to which one could refer back while working through the tribal accounts. I also found my- self wishing for a gallery of illustrations, showing the various genera of sand wasps side by side. For general readers from eastern North America, the most useful portion of these chapters will probably be the account of cicada killer wasp biology (genus Sphecius), in Chapter 3. The synthetic portion of the book begins in Chapter 8, “Comparative Ethology of Sand Wasps.” Patterns begin to emerge, but so does an overall impression of great diversity, tremendous complexity, and many un- answered questions. The theme of phylogenetic recon- struction enters strongly at about page 260, at which point I became quite nostalgic, since this was always my favourite aspect of Evans-school wasp studies in the past. I found myself wishing that I were reading an updated account of the evolution of all solitary wasp behaviour, not just that of one subfamily, but the focus here is clearly on the sand wasps alone. My interest was strongly piqued again, however, by the section on sand wasp conservation. The preservation and promo- tion of open, erosional habitats is becoming increasing- ly important to those of us who study organisms that require such places (yes, I’m thinking here of tiger beetles), and it is encouraging that sand wasp biology adds further impetus to this cause. As a work of entomology, this book is superb. I doubt many people outside the hymenopterists’ guild will read it cover to cover, but I do think that various portions of the text have important implications for evolutionary biology, entomology, natural history, and conservation biology. And as a final offering from one of the great naturalists of the last century, it is a wel- come book indeed. JOHN ACORN Department of Renewable Resources, University of Alberta, Edmonton, Alberta TST 5L7 Canada 490) Tracks By D.C. Jackson. 2006. University Press of Mississippi, 3825 Ridgewood Road, Jackson, Mississippi USA. 279 pages. U.S.$25. Paper. Tracks is a collection of 23 short essays, each rang- ing from 2 to 26 pages. Each essay is built around a personal story of hunting, fishing, trapping or ram- blings, and Jackson uses these narratives to provoke thought and contemplation regarding our interactions with and place within nature. While I think his goal is very worthwhile and of great value to anyone interested in understanding our relationship with the environment, I found that he was only sometimes successful. Often the stories are simply entertaining anecdotes, and some essays miss the mark entirely. This is not to detract from the compilation however, as a range of success is to be expected in such a compilation of wide-ranging indi- vidual stories. When successful, the author is very in- sightful and provides much food for thought. Given this objective however, the musings and philosophies are somewhat confined and repetitive in that all are focused around stories of exploitative nature (hunting, fishing, trapping) whereas some of a non- exploitative nature (camping, hiking, canoeing, drifting, etc.) would be welcome. Such a broader approach would illustrate the connection between a contemplative man and nature in a way that does not require the taking of life. As is, | am left with the vague unease that to “con- nect” with nature it is necessary to kill something. Dr. Jackson is a professor at Mississippi State University and, given that, I would have appreciated some stories and deep reflections arising from his time in the field on his research. It would show the natural environment giv- ing rise to musings in some other aspect of his life be- sides the exploitative areas of fishing/hunting/trapping. THE CANADIAN FIELD-NATURALIST Vol. 120 Tracks is an interesting book, though I would not recommend it as a must-read. Sometimes the “philos- ophy” or musings felt forced, but at other times they were very appropriate, flowing from the personal anec- dote. The first chapter “Wilderness Before Dawn”, really captured my attention and made me hope that I was in for great insight and original perspective. This hope, while not entirely fulfilled, was not entirely de- feated either. These essays are clearly intended for a broad audience of readers and in many ways refreshed exquisitely enjoyable past memories for me. I grew up on stories from the magazines Outdoor Life and Field and Stream, and the stories here are a cross between that type of article and a primer on environmental ethics. As a bridge between outdoor adventure stories and a contemplative approach to nature it is very successful. In addition, while I may not have gleaned profound new insight from the writings, I did learn, and think I have a better appreciation, and respect for, the Deep South of the United States over that when I began the book. All-in-all Tracks is a pleasant read but I would not recommend it as delivering particular insight or a message that we have not heard previously (with the exception of “Wilderness Before Dawn’). For those interested in the rural culture of the Deep South (at least several decades ago), it would be a very useful book. It is the type of book; I think, which is best read in front of a warm fireplace on a cold winter night. SEAN MITCHELL St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada Why Don’t Woodpeckers Get Headaches? And Other Bird Questions You Know You Want To Ask By Mike O’ Connor. 2007. Beacon Press, Fitzhenry & White- side Limited, 25 Beacon Street Boston, Massachusetts 02108 USA. 209 pages. U.S.$14.95. The book Why Don’t Woodpeckers Get Headaches? is a collection of letters submitted to The Cape Codder in the “Ask the Bird folks” column, and answered by Mike O’Connor who is a bird watcher, the owner of a birding store, and a born comedian. The letters ask questions of interest to any backyard birder, ranging from how to attract Bluebirds, to buying the bird bath that’s right for you (and your birds) and how to get rid of a squirrel infestation. The book is divided into eight chapters, each related to a particular area of backyard birding. The first sec- tion deals with bringing birds into your yard. O’Connor gives several important tips to attract various birds to your yard, such as having a proper nest box, leaving food that is suited to the birds you hope to attract, hav- ing suitable habitat, and most importantly, if you want these birds to stay in your yard, offering birdhouses with low rent, an example of the author’s humourous approach. Other chapters of the book deal with food issues such as: Peanut Butter is it safe? What birds can you attract with oranges? Should you replace your rotten seeds? Another chapter discusses providing some basic com- forts for your avian friends: selecting a quality bird house is obviously a key issue, and is talked about extensively. It is safe to conclude from the various entries that, in housing birds, simplicity is key, and while the four story summer villa with the small pick- et fence and lawn included may look great at the store, your birds would probably prefer the cheaper wooden box. Also, providing a heated birdbath is quite impor- tant; contrary to popular belief, birds don’t enjoy a chilly swim in the middle of December any more than we do. Even though this book mostly deals with attracting birds, and encouraging their visits, there is an entire section devoted to bird-related problems. People write in to ask: Why are some woodpeckers more attracted 2006 to metal chimney caps than the peanut feeder? Why do birds insist on chirping in a new day at 4:30 in the morning, right outside their bedroom window? And of course, the timeless question; How to stop their new bird feeder from becoming an all-you-can-eat diner for the cunning gray squirrel? O’Connor provides excel- lent and thorough advice for his readers, even if he hasn’t found a sure-fire solution to the squirrel problem. Why Don’t Woodpeckers Get Headaches is a fasci- nating collection of people’s backyard bird triumphs and tragedies. O’Connor is a talented writer, whose quick wit and edgy humour had this reader crying with laughter. Not only does he answer readers’ queries with accurate and detailed opinions, he also provides a brief BOTANY Alaska Trees and Shrubs. Second Edition By L. Viereck, and E. L. Little (with contributions by D. F. Murray and E. L. Little, Jr.). 2007. Snowy Owl Books, University of Alaska Press, Fairbanks, Alaska. 265 pages. US$24.95. Paper. Measured in Cranberries, Alaska must be among the wealthiest states in the world. Although this Alaskan hardcopy publication was printed in China, it was pro- duced with major U.S. governmental support. A long- awaited second edition, its underlying data exist else- where in digital geo-referenced format online. For some of the most abundant Alaskan species (willows, over 37 Salix species occur in Alaska) the authors present text that simplifies relevant details and identi- fications. These complex details are all part of Alaska’s wild reality and a management that is mainly oriented on commercial goals. The lack of a GAP project in Alas- ka (otherwise found throughout all states within the U.S.; see http://gapanalysis.nbii.gov/portal/server.pt) leaves it a large and harmful heritage; e.g., lack of rel- evant biodiversity management information and digital administrative culture. At least the digital data from this book are available in an older GIS format (ArcGIS 9.0 + Metadata) at the Bonanza Creek LTER website (http://www.lter.uaf.edu) raising hopes of great things to come in the future. This matters a lot, because as the book outlines: “An increasing number of people look to Alaska for wilderness that is no longer present in the more developed areas of the world”. It therefore represents a global natural resource leadership issue and role model on how we deal and publish on Alas- ka’s biodiversity and wilderness management compo- nents that are so crucial to business and human welfare of the global village. Together with contributions by D. F. Murray and G. W. Argus, the authors summarize in this book (first edition published in 1972) their life-time achievement presenting up-to-date information on over 14 tree and 115 shrub species. The book offers four identification Book REVIEWS 49] background look into the bird's appearance and behav- iour. The illustrations by Catherine E. Clark are well done and the birds are easily identifiable. This book is aimed at readers of all ages, and all birding-skill levels, from novice to expert. It is well written and gives a new and often amusing perspective to the world of back yard birding. Even people who are not interested in birds will have a great time reading this book and I would recommend it to anyone. So why don’t woodpeckers get headaches? Well, if you want to know, buy the book! JESSICA GAWN c/o Mission of Canada, 5 Ave de |’Ariana, Geneva 1202, Switzerland keys (Key to Alaska Trees and Tall Shrubs Based Main- ly on Leaves, Winter Key to Deciduous Trees and Tall Shrubs of Alaska, Key to Genera of Alaska Shrubs, Winter Key to Alaska Shrubs). For each of the 132 spe- cies, distribution maps, a descriptive text (usually coy- ering leaves, twigs, bark, wood, flowers, capsules, hy- brids, habitat, distribution, uses and notes), paintings and photos (44 color plates overall) are presented. | enjoyed the numerous drawings (mostly taken from Forest Service publications). The plant distribution maps present traditional-style, expert-drawn “polygon monsters” with mathematically exact shapes and mar- gins but are unlikely to represent biological reality or relevant landscape gradients. Some of the general spe- cies text information one might find in other botanical key references. However, readers interested in Alaska- nia will appreciate the many species details provided. For instance “Alaska Athabascans eat the fruits raw or cooked in moose fat” (for Silverberry), or “Alaska Indians used the wood for totem poles, dugout canoes, and houses, and made mats, baskets, and ropes from the stringy bark” (for Western Red Cedar). The reader of this book will appreciate the incredi- ble diversity (and abundance) of willows, birches, and berries (e.g., Blueberry, Bearberry, Cranberry, Huckle- berry, Snowberry) in Alaska. But environmental prob- lems, widely cited in geo-botanical publications, such as climate change, endemic species loss, road impacts, overcutting, nitrogen input and general human foot- print found throughout Alaska are sparsely treated). Oil development, or pipeline impacts, key features in Alaska’s landscape and recent history, and discussed almost worldwide with high-profile books devoted to this issue (see for instance National Research Coun- cil of the National Academies 2003), are hardly even mentioned. The authors maintain a dubious and ecologically non-sensitive view that “...the distribution of trees and shrubs have changed little” since the 1970s. However, 492 at least the changes for Greene Mountain Ash and Creeping Juniper are described. The concepts of (Land- scape) Ecology, Biogeography, Fragmentation, Bio- diversity, and related loss of plant DNA are virtually omitted in this book. With great regret one has to see once more that the impact of the Intensive Manage- ment Law (signed 1994 by Governor Hickel; maximiz- ing moose numbers through non-science based pred- ator control, and thus severely affecting browsing on shrubs and vegetation Alaska-wide) remains virtually unmentioned. But the impact of ANILCA (Alaska Na- tional Interests Land Conservation Act) in 1980, and the number one problem of the Alaskan National Parks, invasive species, is mentioned (over 100 invasive spe- cies are referred to, but only 9 are described in this book; for a better description the reader is advised to visit websites such as http://www.ucsusa.org/invasive_ species/state-invasion-portfolios.html and http://www. uaf.edu/ces/ipm/plants/index.html). Many of these spe- cies were introduced with governmental help, even by agencies that now have to spend huge amounts of their budget to deal with these earlier sins. The book section on Transplanted Native Species I found very informative (e.g., for Sitka Spruce: “During World War II, the U.S. military had an active program of plant- ing trees near bases in the Aleutian Islands.”). Other environmental topics mentioned are facts like “Much Alaska cedar is exported to Japan in log form...”, or for Sitka Spruce: “Low-grade lumber is made into packing boxes for the Alaska salmon industry”. Keep in mind that the Tongass and Chugach Forest is the largest temperate rainforest in the world, with an ex- tensive network of forest roads reaching more than 10 times around the world! Fortunately, the authors leave no doubt that more forest exploitation will occur in Alaska. Such an outlook makes one shiver for Alas- ka’s future well-being; plants included. But Cranberry juice apart, “Alaska is a land of con- trasts in climate, physical geography, and vegetation” providing a great country for botanists with a thirst to explore. Besides many endemics, it harbours the Quak- ing (Trembling) Aspen (most widely distributed tree in North America) as well as the largest cottonwood known until 1965. And so, the book is full of fascinat- ing Alaskan landscape details such as nunataks, ice fields, permafrost, and some fire history. The promi- nent botanist Robert Marshall and his historic seedling experiments are also mentioned. But relevant Russ- ian, Chukotkan, Kamchatkan and Bering Sea botany are not further dealt with, nor the links to the Yukon, British Columbia or the circumpolar Arctic. It appears a botanical religion that a vegetation clas- sification always has to be carried out in such botani- cal books and landscapes. Therefore, this publication THE CANADIAN FIELD-NATURALIST Vol. 120 offers 8 main vegetation types and their leading species for Alaska (published by the U.S. Forest Service 1971; but relevant quantitative analysis details are not pro- vided, and one has to trust the experts blindly once more). The underlying botanical species taxonomy of this book is still in flux, as can be seen in the large num- ber of hybrids, or in the alder, birch and willow species complexes. Such dynamic and complex taxa are not easy to deal with by using classic governmental hard- copy black-and-white publication philosophies (the authors followed for instance the Flora of North Amer- ica, Furlow 1997 and Argus 2004). It is here where readers need more leadership from the experts. Per- haps presenting links with Genbank (http://www.ncbi. nim.nih.gov/Genbank/GenbankOverview.html) and ITIS (http://www.itis.gov/) would start this concept? The general audience will appreciate the two Alaskan botanical maps and the four page glossary of botanical terms. A seven page index of Common and Scientific Names, and the six pages of literature are of further interest. The authors state the contributions from several agencies and funding institutions, including the U.S. Forest Service and the Museum of the North Herbari- um Database. Unfortunately, botany is often equated with timber, and, as elsewhere, a governmental forest service focus and its cultural bias can be found in this book. For instance, from an Arctic tundra perspective, I am disappointed not to find any relevant references from S. Walker in this book; the reader is advised to see online for CAVM http://www.geobotany.uaf.edu/ cavm. But despite the forestry involvement, the authors still make a nice case that “...the forests of Alaska provide more than timber to the people of Alaska, and the rest of the United States”. Well spoken. This book well describes what Alaska currently has to offer: wide terrain, many new botanical findings that are waiting to be put in order (a la: “we sort ’em post- mortem’’), and much relevant ecology and sustainable management waiting to be done. Curious naturalists might also want to compare the state of the art in Alas- ka with the Flora of adjacent British Columbia (http:// www.eflora.bc.ca). One assumes that the future will likely see digital online species pages, such as already started in Wikipedia and coming forward in GAP and GBIF, with a Biology Google (“Boogle”, or Poogle for Plants) just being around the corner. This books makes for a great step towards this achievement for Alaska, its cranberries and beyond. FALK HUETTMANN Biology and Wildlife Department, Institute of Arctic Biology, University of Alaska, Fairbanks Alaska 99775-7000 USA 2006 ENVIRONMENT BOOK REVIEWS 493 The Atlas of Climate Change: Mapping the World’s Greatest Challenge By Kirstin Dow and Thomas E. Downing. 2006. University of California Press, Berkeley, California, USA. 112 pages. US$23.95 Paper. An atlas suggests maps, and maps there are in abun- dance in this volume, though it also contains many graphs, charts, diagrams, and informative explanatory text. Although short, this book is packed with useful information and well repays careful study. The atlas is divided into seven sections, each focusing on a dif- ferent aspect of the climate change story and each pre- ceded by a one-page introduction. For the first five sec- tions, topics are illustrated primarily by mapped data, usually at a global scale, though occasionally focusing on a specific region, such as the Antarctic. The maps are usually supplemented by a few paragraphs provid- ing some additional background or explanation for the displayed data. The atlas is sprinkled with quotes, though these are mostly undated, which rather dimin- ishes their urgency or relevance to the issues. Part 1, Signs of Change, highlights some warning signs, such as the European heat wave of 2003, and then covers three aspects of climate change that set the context for the rest of the book. These three aspects comprise glacier retreat, weather-related disasters, and changes in the polar regions, areas that have been iden- tified as having particular sensitivity to climate change. These topics are likely to be familiar to Canadian read- ers through various recent media stories. Part 2, Forc- ing Change, focuses more directly on the climate sys- tem, covering four topics, mostly through charts and diagrams. Two aspects of the climate system are high- lighted, explanations for the greenhouse effect and how climate operates in terms of heat transfer across the globe. This section also provides a long-term perspec- tive on climate change, showing the record of carbon dioxide fluctuations for the last 400 000 years derived from ice-core records. Finally, there are some scenar- ios, based on various model projections, showing pro- jected carbon dioxide increases and related tempera- ture increases. The following three sections form the bulk of the atlas. With six topics, Part 3, Driving Climate Change, examines greenhouse gas production in more detail, highlighting several critical aspects of modern indus- trial activity and linkages to fossil fuels, especially transportation and agriculture. Covering seven topics, Part 4, Expected Consequences, paints a grim picture, showing, amongst other consequences, the impacts of rising sea levels, water shortages, and threats to human health. Africa stands out as the region likely to be most severely affected in the last category. One of the most interesting maps in this section deals with cultural loss- es, including historical and archaeological sites. These kinds of impacts are not often highlighted in discus- sions of climate change. On the other hand, biodiver- sity losses and biogeographic changes are given very little attention. Since alterations in plant and animal distributions are some of the more obvious and visible signs of climate change, I found this rather surprising. For Canadian readers, these sections may induce both complacency and embarrassment. Complacency be- cause the maps show that on an absolute basis, Canada does not contribute as much to the problems as other more populous industrialized countries. Embarrassment because on a per capita basis we consume so much compared to so many countries of the Third World. Such reflections form a natural transition to the last sections of the book. Having provided a gloomy outlook in the previous section, Part 5, Responding to Change, shows what we can do about it, or how we might be able to modify the outcomes at a macro-scale. Eight topics are presented, most of which are focused on institutional or adminis- trative responses, such as carbon trading or develop- ment of alternate renewable energy sources. Finally, Part 6, Committing to Solutions, shifts the focus from the global to the local, with some charts showing how individual actions might affect the trajectory of change. For those who want the “nitty-gritty” details, Part 7, Climate Change Data, tabulates various measures of economic activity and consumption on a country-by- country basis. These data underpin the maps and graph- ics in the rest of the book. Generally the book concentrates on the more obvi- ous impacts of climate change and the ones that are often the focus of media attention, such as potential limits on water and food supplies. As such, most read- ers are likely to have at least some passing familiarity with the issues. This book therefore serves as a use- ful source or companion book for those seeking more background on current issues. More subtle effects of climate change, such as water quality degradation, or cascading effects, such as wildfire frequency changes or aquifer depletion, are not shown, perhaps because these are less amenable to measurement or estimation and mapping. Moreover, the emphasis throughout is on impacts to the terrestrial realm; other than sea-level changes, very little attention is paid to marine sys- tems. Again, this is perhaps surprising given the link- ages that have been found between, for example, the ENSO (EI Nino-Southern Oscillation) with rainfall and droughts. Cartographically, the book is attractive, clear, and well-designed. It is apparent that a great deal of thought and care has been paid to the layout, presentation, and overall design. In my view, the design is one of the book’s greatest strengths. Many people are “visual learners” and this book will definitely appeal to anyone that prefers graphics to text. Most of the global presen- 494 tations use the same map projection, with Europe and Africa in the centre and the Americas and Australasia at the extreme left and right. For North American readers, this projection can seem strange, since we are more used to seeing the Americas displayed cen- trally with other continents on the periphery. Much of the base information is quite complicated but the maps show good use of colour and symbols to con- vey information and summarize complex ideas. One of the more interesting maps shows greenhouse gas emissions, with countries’ size distorted according the amounts emitted. The effect is quite startling, with the dominance of the industrialized regions (USA, Europe, CIS, and East Asia) standing out. Perhaps more telling is the minuscule contribution from Africa, which shows up as a small polygon, despite its large land area and population. Even with the references and sources, I did find my- self wondering exactly how some of the figures under- lying these maps are derived, especially because so many of the data are presented at a regional level, where regions may include several countries, and are “binned” into a few categories to simplify cartographic presenta- tion. Uncertainties or error estimates are not provided for the data. Inevitably, there must be some fairly broad assumptions and generalizations underlying the figures. To take one example, what kinds of measurements could be available to allow an estimate of methane emissions on a per capita basis? Are the same or com- parable measurements, that is, the same data quality, really available for all areas of the world? How are data integrated even within one country, given that there THE CANADIAN FIELD-NATURALIST Vol. 120 are different ecoregions and uneven population distri- bution? How much of the disparity shown is real and how much attributable to varying levels of data avail- able for different areas of the world? In fact, how far are the numbers behind the maps realistic estimates based on reliable instrumentation or simply informed “guesstimates”? Notwithstanding the beguiling pres- entation, I did find myself asking many questions about the methodology behind the mapping. Perhaps the greatest drawback with this book is that, because it deals with current and rapidly changing is- sues, the information presented will quickly become dated. The authors note that no new material was added after spring 2006, but they also indicate that they in- tend to update the volume, especially after the next Intergovernmental Panel on Climate Change (IPCC) report, expected in 2007 (page 13). It will be interest- ing to see how future editions compare to this one and whether detectable changes in some of the indicators show up on that time scale. One could argue that such information is better presented on a website, where it could be updated as needed. However, having the information gathered together in book form makes it easier to consult and compare the different maps. The book’s greatest advantage is its accessibility, with its clear presentation and distillation of an enormous amount of complex information into a readily under- standable format. ALWYNNE B. BEAUDOIN Royal Alberta Museum, Edmonton, Alberta T5N OM6 Canada An Inconvenient Truth: The Planetary Emergency of Global Warming and What We Can Do About It By Al Gore. 2006. Rodale, New York, USA. 328 pages. US$28.95 Paper. I freely admit that I picked up this book already pre- disposed to dislike it. After all, it’s supposedly a book about climate science by a politician, so how good could that possibly be? I am relieved to report that this volume pleasantly surprised me; the book is consider- ably more readable and worthwhile than I expected it to be. Surprisingly, it also projects, an air of intelligence and careful consideration. Yes, the book is a call to action and a strong statement of a particular point of view, but the hype and hyperbole are at a lower volume than I had feared. More importantly, the book is largely based on solid science and the presentation of that sci- ence is done in a remarkably straightforward fashion. In his introduction, Gore says that the book was developed from a slide show and, judging by the lay- out and design, that presentation mode has carried over and set the book’s style. It is arranged as a continuous narrative, with clear graphs and graphics, some on fold- out pages, colourful eye-catching photographs, short sentences and paragraphs, and large type. Graphics are sourced, though complete citations are not provided. Text and explanatory captions are generally written in plain conversational language. The book surveys the evidence for global warming and outlines the main forcing factors, highlighting the importance of green- house gases and the role of human activities in the in- creasing rate of, especially, carbon dioxide content in the atmosphere. Gore drives home the message that human activities, in particular industrial activities, are responsible for much of the recent increase in greenhouse gas concen- trations in the atmosphere and therefore the increasing rate of global warming. He sets these trends in context by showing, for example, recent rates compared to the carbon dioxide trend inferred from Antarctic records extending back to 600 000 years before present. Other long-term records, such as tree-rings and Greenland ice-cores, are also used to provide perspective. Several times, the point is made that it is not simply the absolute amount of increase that is important but the increasing rate of increase. Taking it as given that global warming will continue, even if actions are taken now to reduce rates of carbon dioxide emissions, Gore then exam- 2006 ines what the implications are likely to be during the next few decades. The book presents many different types of evidence for global warming and covers a lot of territory. It draws on data and images from across the world. Gore amasses a formidable array of information to support his argument. Some of the topics he features include: droughts and shrinking lakes (including an intriguing sequence of images showing the reduction in Lake Chad), reduction in Arctic ice, shrinking of large ice sheets (notably on Greenland and Antarctica), effects on temperatures and seasonality, impact of rising sea- levels (including the inundation of islands), damage to coral reefs, effects of climate on human health and diseases rates, frequency and occurrence of extreme weather events (including, not surprisingly, hurricanes), and climate impacts on biodiversity and distribution of plants and animals. The evidence for global warming includes some striking “then and now” photographs of glacier forefields showing recent ice retreat. Two issues that also stand out are the effects of exceptionally high summer temperatures, as seen in recent years in Europe especially, and increased storminess, especially on low- lying islands and coastal areas. The evidence and impli- cations are shown from a global and not just a U.S. perspective. This worldwide coverage underlines the urgency of the issues presented, since some of the im- pacts are not just hypothetical but are being felt by people across the world now. This leads Gore into a consideration of the roots of the problem, in particular the main sources of green- house gas emissions. Some striking graphics visually emphasize the large contribution to greenhouse gas emissions made by the U.S. Gore points out some of the global inequities in consumption that have led to this situation and militate against any remedial action. He identifies energy production as the main culprit and highlights the need to move to alternate energy sources, such as wind power. Here, the text gets more overtly political with Gore taking aim at the Bush administra- tion for obfuscating the issue and refusing to act, for example, by not ratifying the Kyoto Protocol. Interspersed within the text are eight short essays, most of which recount anecdotes and incidents from Gore’s life. These stand out from the rest of the narra- tive by being printed in smaller font on tan-coloured pages. The essays highlight some of the reasons for his passionate concern about global warming. What they show is that this is not a recent preoccupation but something that has engaged his attention for many years, growing out of life-long interest in environmen- tal issues. Clearly, he wants to let readers know that he is not simply “jumping on a bandwagon” and taking BOOK REVIEWS 495 up a trendy issue out of opportunism, but is following an interest that has underpinned his political career. Here, I did feel that I was reading a political manifesto at times. There’s a lot of name-dropping and there are rather too many “warm and fuzzy” pictures of Gore and his family for my taste. Nevertheless, these essays bring the issue from the abstract to the personal, which does sharpen the sense that these concerns matter to individuals. By being set off from the rest of the text, it is clear that the message in these essays differs from the rest of the book. The final section of the book, offset by being print- ed on light-green paper, is the call to action, outlining some things that individuals can do to make a differ- ence. Most of these actions will already be familiar to most readers: buy a hybrid car, bag your groceries in re-usable shopping bags, replace incandescent light- bulbs with compact fluorescent bulbs, and so on. Gen- erally, these actions are an appeal to a sense of guilt in affluent people living in western countries, where, Gore maintains, most of the problem originates any- way. Some of the suggestions run counter to many driving forces in the economy. When the TV screen is filled with ads encouraging us to be tourists and travel to sunny beachside resorts, how realistic is it to tell peo- ple to reduce air-travel? Interspersed among these ex- hortations are ten common misconceptions about glob- al warming and Gore’s recommended responses. These misconceptions include the ones you might expect, such as the assertion that “it’s all just natural variation”, though there was one I hadn’t heard before, attributing global warming to after-effects of the Tunguska event. In such a brief statement of the global warming issue, Gore obviously does not get into any topic in detail. There’s clearly a lot more that could be said about all lines of evidence he presents but, unfortu- nately, there is no reading list for readers to follow up on any topics raised in the book. Nevertheless, Gore does assemble a persuasive argument that global warm- ing is a problem and we need to do something about it immediately. However, this is very much a book that is “preaching to the converted”. I can’t see many peo- ple who are not already convinced there is a problem reading it. On the other hand, Gore is a well-known figure so perhaps some people may take a look at this book because of name-recognition. As a politician, Gore has to overcome a good deal of cynicism in order to reach people. This is a pity because this book is an articulate and thought-provoking presentation of a global phenomenon that affects us all. ALWYNNE B. BEAUDOIN Royal Alberta Museum, Edmonton, Alberta TSN OM6 Canada 496 MISCELLANEOUS THE CANADIAN FIELD-NATURALIST Vol. 120 Gilbert White: A Biography of the Author of The Natural History of Selborne By Richard Mabey. University of Virginia Press, Char- lottesville. 2006. 239 pages. U.S.$16.50 Paper. In his introduction, Mabey admits that, at his first reading of Gilbert White’s The Natural History and Antiquities of Selborne, he could barely cope “with its rambling disorder.” Only on visiting Selborne in per- son did Mabey change his view decisively. I marvel, as Mabey did, that White “could produce something so wholly original and appealing out of such unpromis- ing ingredients.” In this Whitbread Prize-winning biography, the pain- staking research of Mabey — whom the Times has called ‘Britain’s foremost nature writer’ — results in a remark- ably detailed account of the life of Gilbert White. Born in the tiny village of Selborne, White obtained his uni- versity education in theology at Oriel College, Oxford, beginning in 1739. Here he met John Mulso, in the year behind him. Although “constitutionally lazy and hypochondriacal,’ Mulso’s life-long correspondence to and from White became a major source for Mabey. Perhaps it was equally providential that White was not particularly successful in his first three appointments to curacies at a distance from his home. When he returned to Selborne he began, somewhat inauspiciously, to com- mit his nature observations to paper in 1756. White was patient and inquisitive. His early descrip- tion in 1761 of the life of crickets was both “vivid and sensuous.” In 1767 he began writing to the naturalist and author, Thomas Pennant; two years later he began A Paddler’s Guide to Quetico and Beyond By Kevin Callan. 2007. The Boston Mills Press, Erin, Ontario Canada. 192 pages. U.S.$24.95 Paper. It’s the dead of winter on a Friday evening when friends lay out the maps on the floor in front of a burn- ing fire. Over wine, possible routes are discussed and commitments are made for a canoe trip six months hence into one of the most intact, accessible and largest protected wilderness areas in North America: Quetico Provincial Park. This is a book that will lead you beyond your usual canoe routes; in fact, it may inspire you to plan a trip celebrating Quetico’s 100" anniversary in 2009! Kevin Callan is no stranger to writing books on wilderness canoeing with seven “‘Paddler’s Guides” for Ontario canoe routes to his name. Kevin reminds us that there is a great story to be told after every canoe trip we make, as a result of spending quality time with a group of family or friends on a journey through the natural world of wind, water, wildlife and plants, com- plete with portages, bannock and fireside stories. Each of the sixteen canoe routes described in this book is a well-written informative story, weaving important prac- his original observations that helped prove that birds migrate. In 1774, White’s lively letters to Sir Daines Barrington included one about house martins, which Barrington read to a meeting of the Royal Society. The result was publication of these “models of lucidity and insight” in Britain’s earliest scientific journal, Philo- sophical Transactions. Finally, late in 1788, at age 68, White had the sat- isfaction of seeing his book in print. He died in June 1793, a month short of his 73" birthday, modestly un- aware of the prominence his book would achieve in the centuries to come. Second and third editions did not appear until 1802 and 1813, respectively, but there have been more than two hundred subsequent editions, one of the most published books in the English language. I appreciated Mabey’s delightful biography all the more because I, too, have visited Selborne. By learn- ing more about Gilbert White, we can better appreci- ate White’s impact on all subsequent nature writing. I congratulate the University of Virginia Press for mak- ing this inexpensive reprint, first published in hard cover in 1986, readily available to North Americans. I heartily recommend it to anyone with the slightest interest in English literature or the history of natural history. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada tical information such as the take-out and put-in spots for portages, special sites to see, and danger spots — into historical accounts of those who were here long before our forays using Kevlar canoes and Gortex. The stories also convey the hardships and joys shared by the group traveling with Kevin; the “real stuff” that makes canoe trips memorable. The book does not overwhelm the reader with un- necessary detail on the gear to bring, what kind of canoe paddle is better than another, etc., rightly assuming someone heading to Quetico is not a novice. At the same time, simple, straightforward descriptions of canoe routes make them immensely doable for the aver- age paddler. Exceptional pictures and tidy maps are also included, with references for the “proper” maps a canoeist must purchase in order to undertake these trips Sixteen canoe routes are described in Quetico and Beyond; eleven in Quetico Park and five within neigh- boring areas of northern Ontario. Most routes require about one week to complete, although some are as short as 2-4 days and others as long as 28 days. Sum- maries provided for each of the canoe routes includes 2006 number of days required to complete the route, the number of portages, the longest portage, route difficul- ty, access points, and the maps needed. Scanning the summaries provides a quick method to narrow down which routes look the most interesting within the time frame and abilities of the group. Quetico is a book that provides very readable pleas- ant conversation that, in a personal way, brings canoe- BooK REVIEWS 497 ing to life through its past and more recent adven- tures. The many reminders of the subtle rewards of canoeing that are conveyed provide a real enticement to begin planning your next trip. BRENT TEGLER North-South Environmental Inc., 35 Crawford Crescent, Unit BUS, Campbellville, Ontario LOP 1B0O Canada More than Kin and Less than Kind: The Evolution of Family Conflict By Douglas W. Mock. 2004. Belknap Press of Harvard Uni- versity Press, Cambridge, Massachusetts, USA. 267 pages. U.S.$27.95. If you wish to understand how complicated and how variable reproductive behaviours can be, this is the book! Doug Mock offers a wide vista of nature in all its infinite variability and complexity. To explain bird behaviour, Mock invokes human examples, as diverse as his three older brothers, chess, Wall Street, sports teams, and vote counting. Mock offers creative chapter headings and begins each chapter with a brilliant quotation. He simplifies difficult-to-understand concepts in an instructive and often entertaining manner. As he says, this book is a “mixture of theory and data.” He is especially inter- ested in brood size and in siblicide. I will offer a few highlights, some of which mention Canadian research. Mock attempts to explain both the “hows” and “whys” of concepts that include Darwinian fitness, evolutionary game theory, inclusive fitness theory, evo- lutionarily stable strategy, parent-offspring conflict, parental manipulation, optimal clutch size, replacement offspring, and the insurance egg hypothesis. Reproduction is costly. When Collared Flycatcher broods were enlarged artificially, there were fewer breeding adults the following season, and when two eggs were removed, females laid larger clutches the following year. Seychelles Warblers produced 77% male chicks in poor habitat and 88% female chicks in good habitat. Lifetime monogamy is uncommon among birds but there are notable exceptions. A sample of 919 pairs of Bewick’s Swans showed not a single divorce, but if a mate died the survivor would re-pair. From a sample of over 6 000 banded birds, 99.6% of Barnacle Goose pairs were socially monogamous. Mock discusses Hamilton’s rule, promulgated in 1964, whereby “two full siblings should be the evo- lutionary equivalent of one Self’ — or eight cousins — based on the amount of shared genetic material. Altru- istic behaviour, such as nest helpers, relates to the mix of relatedness. What happens in nature often seems counter-intu- itive if not outright bizarre. Biologists have difficulty understanding, much less explaining, such events. What conceivable evolutionary advantage could result from a surplus egg or surplus young? Why do some species of eagle and pelican regularly lay two eggs, yet raise only one young? Why does a parent bird passively watch one of its nestlings kill another, even when sur- plus food is within reach? Why does a hawk nestling sometimes eat its sibling after killing it, but on other occasions does not? At Delta Marsh, Manitoba, Spencer Sealy video- taped nests with a single cowbird and a single warbler; the cowbird out-hustled, out-begged and out-stretched its warbler nestmate. In British Columbia, Bruce Lyon studied American Coots, which lose some chicks to starvation in nearly half the broods. Adult coots make a point of getting more food to the youngest, but when Lyon clipped the bright ornamental plumage of the tiny chicks, the clipped birds survived less often. The late Roger Evans of the University of Manitoba did experiments to study the role of the extra or “insur- ance” egg in the American White Pelican. He and his student Kevin Cash carefully marked the first-hatching or A egg, and the second-hatching or B egg. Twenty percent of the B eggs hatched. The parent pelican would brood its young, but as soon as the parent stood up, the older young attacked its smaller sibling. Wahlberg’s Eagle, which ranges across central Africa, lays a single egg; not more than three percent of pairs lay a second egg and then the larger chick regularly kills the smaller one. The Verraux’s (Black) Eagle lays two eggs, but an observed first chick began assaulting the second chick soon after it hatched, giving 1 569 blows with its beak to kill its sibling, even though food was plentiful in the nest. The Crested Penguin lays two eggs, but the first egg is up to 40 percent smaller; four days later the sec- ond, larger egg is laid and is given the optimal brood patch position beneath the mother, and is usually the sole survivor. Mock does not restrict himself to birds. He writes about plants, insects and fish, using the firefly as an example of signalling. He tells how, in a pig litter, ante- rior teats produce more milk than posterior teats, hence the colloquialism, “sucking hind teat.” As Marlene Zuk says on the dust cover, this book about family conflict “is a model of how behavioral ecology can and should be done ... Just don’t take it to family reunions.” C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Canada 498 To See Every Bird on Earth By Dan Koeppel. 2005. Hudson Street Press, New York, New York, USA. 278 pages. $24.95 U.S.$35. Dan Koeppel tells the story of his father, Dr. Richard Koeppel, a chronically unhappy medical doctor with two overwhelming obsessions: to see as many species of birds as possible, world-wide, and to read, cover to cover, every novel that was short-listed for or won the Booker Literary Prize. Each chapter begins engagingly with a postage stamp that features a bird from the collection of Chris Gibbins who has amassed stamps depicting 2950 species (a record list!), and a paragraph from Richard Koeppel’s notebooks telling of a numbered addition to his life-list. One learns about the methodology of listers, par- ticularly the rules laid down by the American Birding Association (pages 195-197). Dan explains, in layper- son’s terms, “lumping” and “splitting” of species. Despite the ever-escalating CPB (cost per bird), Dan fully realizes that the listing process is “intensely arcane, fascinating, and absurd.” This is a quick and easy read, but in no way uplift- ing. Dan’s brutally frank, very personal psychoanaly- sis of his father and mother is somewhat excessive and distracting. Dr. Richard Koeppel, it seems, entered med- icine to please his parents, and sadly failed to find a fulfilling niche in medicine. His marriage failed early. He became a morose, rather pathetic man who found solace in excessive use of marijuana, alcohol and nico- tine. Most of his career was in emergency medicine, where roughly half his time could be spent birding. When his bad habits caught up with him in the form of larynx cancer, Richard Koeppel’s personal list was at 7080 species. I most enjoyed the behind-the-scenes accounts of Jim Clements, who compiled the first one-volume list of birds of the world, and has amassed a personal life list of 7200; Victor Emmanuel, who began the first field trips designed to add maximum numbers of birds to each customer’s life-list; and Bret Whitney, who lacks formal postgraduate education in ornithology but has succeeded the late Ted Parker as a finder and des- criber of new species in South America. One also meets Richard’s competitors. Within our region, North and Central America, Dan Canterbury Wildfire in the Wilderness By Chris Czajkowski. 2006. Harbour Publishing, Madeira Park, British Columbia, Canada. 221 pages. $19.95 Paper. A woman who lives a solitary life in the wilderness of the British Columbian mountains, dozens of kilo- metres from her nearest neighbours, has not isolated herself from her community. Her life is a heart-warm- ing Set of tales explaining the connection she has to the community of mountain dwellers, outfitters, and bush pilots in her immediate neighbourhood as well as THE CANADIAN FIELD-NATURALIST Vol. 120 has seen a record 1731 species. Peter Kaestner, who traveled widely in the diplomatic service, was the first person to have seen a representative of all 159 avian families. Others in competition with Richard Koeppel for the world list have been Harvey Gilston of Britain, who, without ability to identify the bird himself, would check off a bird when a guide called out its name (he reached nearly 8000); Michael Lambarth and Sandra Fisher of England (Michael quit when his beloved part- ner died); Joel Abramson, a medical doctor who hired top birders to lead endurance-test expeditions; Stuart Keith, founder of the American Birding Association, who was recognized by the Guinness Book of World Records when his list reached 4300; Roger Tory Peter- son, who reached the “half-way total” of 4800 in the 1980s; Bill Rapp, nearing 7000; Jim Plyler, a retired oil executive at 7200; Peter Winter and John Danzen- backer, military men, at 7800 and about 7750, respec- tively; Peter Kaestner, with 7958; Tom Gullick, leader of those still alive, with 8114; Phoebe Snetsinger, still tops, who reached nearly 8500 before she died in a car accident in Madagascar in 1999. What are the downsides to this book? The title is a bit misleading, since Dan Koeppel knows full well that no human will ever see every bird (of about 9600 species) on earth. An index to birder’s names would have been helpful. Worse, Richard Koeppel, the protag- onist of this story, is a sad-and pathetic person, lacking enthusiasm for his profession, with complicated mar- ital and family situations. He is a perfectionist but only on rare occasions does he show much feeling, even for birds. But Richard has made amends with his son, Dan, and has recently turned his attention to but- terflies. All readers of this review, especially the writer, are sinners. We claim to be environmentalists, yet many or perhaps most of us drive gas-guzzling vehicles to pur- sue our sometimes obsessive hobby. At least we fall short of the obscenely high travel expenses of certain of the Big Listers. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N O0J8 Canada friends, publishers, fans of her books and international tourists in the wider community. The reader is intro- duced to her dogs, her history of publications, her inge- nuity in building and maintaining a wilderness site and even the state of her bank account while following her stories of maintaining solitude in the mountains and contacts with the outside world. A hiker, camper and outdoorsman would certainly relate to Chris Czajkowski’s stories and her descriptions 2006 which make the mountains a home. The well-being of the environment becomes close to the heart of any dweller of the wilderness and the threat of nature tak- ing a turn for the worst can send dwellers of the back- woods to worry over their personal charge, the nature and landscape around them. Sustained inclement weather, freak storms, too much snow in a season, frost at the wrong time, drought in the wrong season or pro- longed drought in any season, are all causes for worry but a natural disaster like a wildfire is feared above all. Though the title tells us only of one dramatic event in the history of her lake, cabins and near environment, the reader is drawn into her whole lifestyle of pro- tecting, nurturing and living within the environment which she has come to call home. As the story unfolds, Czajkowski leads us into her world, travelling long distances, keeping up with mod- ern technology, camping and hiking with her tourist helpers. She is home in her particular isolated part of the world a long way from her nearest neighbours but connected with them by regular radio conversations, listening to radio broadcasts and occasionally eaves- dropping on her neighbours’ conversations, and finally email as that technology became available. At the same time, she is host to the world by operating a small guest house for people willing to travel the distance to spend a few days with her, or international wanderers who pay by passing a few weeks working and helping her with projects. Finally the reader is led to the hot dry summer of 2004 and the devastating fire, the Lonesome Lake Fire which was well reported to the Canadian public as one of many fires in a hot dry summer, but to people of YOUNG NATURALISTS Tale of a Great White Fish By Maggie de Vries. Greystone Books, Douglas & McIntyre Publishing Group, Vancouver, British Columbia, Canada. 34 pages. $12.95. The book Tale of a Great White Fish follows the story of a giant white sturgeon. Through its long life, the sturgeon overcomes many obstacles in its battle to survive and reproduce. The story deals with the inter- action between sturgeons and people, along the way providing the reader with a wealth of information and insight into the natural history of the sturgeon. The story starts in 1828, when the sturgeon, named “Big Fish,” was a small and defenceless egg hatched in the Fraser River. She matures into a larva and then into a fry. As a fry she must eat thousands of other small water creatures to grow. Fry are also especially vulnerable to predators, and most of the young fry never reach adulthood. The fry that survive mature into adults before returning to their spawning ground to lay their own eggs. There are many dangers that the sturgeon must over- come. For most of their long existence sturgeon have BooK REVIEWS 499 that remote region, a disaster which shaped their envi- ronment for years to come. The reader is invited to travel with Czajkowski through the fear and uncertain- ty of watching gathering smoke and distressing news from various watchers in neighbouring valleys. We are inspired to evacuate with her to safer areas, but with- out any assurance that home or lifestyle could be pre- served. Czajkowski leads us along her tale with inter- esting anecdotes and humorous stories about her dogs, friends, and people who come to visit her. She is an engaging story-teller and keeps the reader’s attention for hours or just a few minutes at a time as the book is filled with short stories, portions of her diary and longer accounts of specific incidents. The story builds in an entertaining way through her history of building the cabins, receiving visitors, includ- ing facts about making one’s life in the wilderness at the same time maintaining connections firmly planted in the outside world. The fire is the high point of the book and gets the reader to the excitement of living her experience of the fear, doubt, loneliness and response of care for her own animals and the natural dwellers of her wilderness world. The reader enters into the story with her and can live the excitement she feels. In our sheltered world, our decisions are not always the same as the author’s but she puts her concern and care for her environment in front of the reader and shows us a different world to enjoy for a little while then look forward, like her globe-trotting visitors, to the next visit we might have with her. Jim O° NEILL 28718 Five Mile Road, Livonia, Michigan 48154 USA been around since the time of the dinosaurs, the threats have been natural: predators, natural calamities and disease. Today, human fishermen and the caviar indus- try are one of the greatest perils. Big Fish was hooked by a fisherman, but fortunately she was freed by some young boys and was able to escape. During the Hells Gate Slide in 1913 many migrating sturgeon were killed. Big Fish survived, but she was forced to find a new place to spawn. Later, Big Fish had to find a new home because the lake she was living in was drained. Later still, an unknown disease swept through the Fras- er River and many sturgeon died and were washed up on the shores of the lake. When the story closes, almost a hundred years later, Big Fish is still surviving, but continues to face many challenges. Scientists study sturgeons to help them survive. They keep track of their weight and size, and try to stop over- fishing. There are now laws which prevent the fishing of these magnificent animals. The book is written as a story and at the end there is a useful list of sturgeon facts, a labelled diagram of a 500 White Sturgeon, a glossary, and a letter from Rick Hansen, a famous Canadian para-athlete. Rick Hansen is the chairman of the Fraser River Sturgeon Conser- vation Society. The drawings by Renné Benoit are done in water colours and they provide a magnificent glimpse into the beautiful and dangerous world of the great White Sturgeon. The book is very informative. In it, the reader will learn a great deal about the White Sturgeon’s envi- ronment and behavior and appearance. The Sturgeon Facts and a glossary at the book of the book provide NEw TITLES Prepared by Roy John + Available for review * Assigned ZOOLOGY Bird — The Definitive Visual Guide. Edited by Peter Frances et al. 2007. Dorling Kindersley, DK Enquiries, Tourmaline Editions Inc., 662 King Street West, Suite 304, Toronto, Ontario, MSV 1M7. 512 pages. GBP 25.99 Cloth. The Black Bear Almanac. By David Smith. 2007. Globe Pequot Press, 246 Goose Lane, P.O. Box 480, Guilford, Con- necticut 06437 USA. 288 pages. U.S.$29.95. * The Clements Checklist of Birds of the World (6" edi- tion). By James F. Clements. 2007. Cornell University Press, Sage House, 512 East State Street, Ithaca, New York 14850 USA. 864 pages. U.S.$59.95 Cloth. The Biology of African Savannahs. By Bryan Shorrocks. 2007. Oxford University Press, New York, New York. 268 pages. GBP 27.50 Paper. Arctic Flight - Adventures Amongst Northern Birds. 2007. James McCallum. Langford Press, 10 New Road, Langtoft Peterborough, UK. 180 pages. GBP 37.99 Cloth. Endangered. By George C. McGavin. 2007. Cassell Ilustrat- ed, 2-4 Heron Quays, London E14 4JP. 192 pages. U.S.$35. * Birds of Europe, Russia, China, and Japan: Passerines: Tyrant Flycatchers to Buntings. By Norman Arlott. 2007. Princeton University Press, 41 William Street, Princeton, New Jersey 08540 USA. 240 pages. U.S.$29.95 Paper. Feathers: Identification for Bird Conservation. By Marian Cieslak and Bolestaw Dul. 2007. Natura Publishing House, ul. Olbrachta 118c/20, 01-373 Warszawa, Poland. 320 pages. GBP 25.62. Forensic Entomology: An Introduction. By Dorothy Gen- nard. 2007. John Wiley & Sons Canada Ltd., 6045 Freemont Boulevard, Mississauga, Ontario LSR 4J3 Canada. 244 pages. U.S.$55 Cloth. Homalopsid Snakes, Evolution in the Mud. By J. Murphy. 2007. Krieger Publishing, P.O. Box 9542, Melbourne, Florida 32902-9542 USA. 260 pages. U.S.$68.50 Cloth. The Inner Bird — Anatomy and Evolution. By G. Kaiser. 2007. The University of Washington Press, P.O. Box 50096, Seattle, Washington 98145-5096 USA. 464 pages. U.S.$85 Cloth. Insects of Britain and Western Europe. By Michael Chinery. 2007. A & C Black Publishers Ltd. (United Kingdom), 37 Soho Square, London, W1D 3QZ. 320 pages. GBP 14.99 Paper. THE CANADIAN FIELD-NATURALIST Vol. 120 a very useful summary, and is well presented. The book is aimed at a child’s reading level. Older readers would find this book less interesting; howey- er, it provides a basic insight into the world of a great white fish and is nicely illustrated. This book is rec- ommended for any child with an interest in the out- doors and in nature. JESSICA GAWN c/o Mission of Canada, 5 Avenue de |’ Arianna, Geneva 1202, Switzerland Insect Conservation Biology. Edited by A. J. A. Stewart, T. R. New and O. T. Lewis. 2007. Oxford University Press, 198 Madison Avenue, New York, New York 10016 USA. 480 pages. U.S.$190 Cloth. Australia’s Mammal Extinctions. By Chris Johnson. 2007. Cambridge University Press, 40 West 20" Street, New York, New York 10011-4221 USA. 278 pages. GBP 34.99 Paper. Owls of the United States and Canada. By W. Lynch. The University of Washington Press, P.O. Box 50096, Seattle, Washington 98145-5096 USA. 256 pages. U.S.$44.95 Cloth. Spotted Owls. By J. Hobbs. 2007. Greystone Books, Suite 201, Quebec Street, Vancouver, British Columbia V5T 4S7 Canada. 144 pages. $39.95 Cloth. Birds of Pakistan. By Richard Grimmett and Tim Inskipp. 2007. Helm Field Guides. 256 pages. GBP 24.99 Paper. Birds of Peru. By Daniel F. Lane, John P. O’ Neill, Theodore A. Parker II, Thomas S. Schulenberg, and Douglas F. Stotz. 2007. Helm Field Guides. 656 pages. GBP 29.99 Paper. Raptors: A Field Guide to Survey and Monitoring. By J. Hardey, H. Crick, C. Wernham, H. Riley, B. Etheridge and D. Thompson. Stationery Office (TSO), 71 Lothian Road, Edin- burgh EH3 9A2 UK. Scotland. 300 pages. email: esupport@ tso.co.uk *Reptiles and Amphibians of Canada. By Chris Fisher and Ronald Brooks. 2007. Lone Pine Publishing, 10145 — 81 Avenue, Edmonton, Alberta T6E 1W9 Canada. 208 pages. $18.95. * Rodent Societies. By J. Wolff and P. Sherman (Eds). 2007. University of Chicago Press, 1427 East 60" Street, Chicago, Illinois 60637 USA. 610 pages. U.S.$49 Paper, $125 Cloth. The New Encyclopaedia of Snakes. By Christopher Matti- son. 2007. Cassell Illustrated, 2-4 Heron Quays, London E14 4JP. 272 pages. GBP 30 Cloth. Native Trees of the Southeast — An Identification Guide. By L. Katherine Kirkman, Claud L. Brown, and Donald J., Leopold. 2007. Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, Oregon 97204 USA. 372 pages. $45.95. Waterbirds Around the World. By Gerard Boere, Colin Galbraith and David Stroud [Editors]. 2007. TSO (The Sta- tionery Office), 71 Lothian Road, Edinburgh EH3 9A2 UK. Scotland. 940 pages. GBP 50. 2006 The Last Wild Wolves. By I. McAllister. 2007. Greystone Books, Suite 201, Quebec Street, Vancouver, British Colum- bia VST 487 Canada. 192 pages. $45 Cloth. + Whales and Seals — Biology and Ecology. By P. H. Fontaine. 2007. Schiffer Publishing Ltd., Lower Valley Road, Atglen, Pennsylvania. 448 pages. U.S.$34.95. * Why Don’t Woodpeckers Get Headaches? By M. O'Connor. 2007. Fitzhenry and Whiteside Limited, 195 All- state Parkway, Markham, Ontario L3R 4T8 Canada. 209 pages. $14.95. ENVIRONMENT The Sacred Balance. By D. Suzuki. 2007. Greystone Books, Suite 201 Quebec Street, Vancouver, British Columbia VST 487 Canada. 320 pages. $22.95 Cloth. Scaling Biodiversity. By David Storch, Pablo Marquet and James Brown. 2007. Cambridge University Press, The Edin- burgh Building, Shaftesbury Road, Cambridge, CB2 8RU. GBP 38. Climate Change. Edited by J. DiMento and P. Doughman. 2007. The MIT Press, 55 Hayward Street, Cambridge, Mas- sachusetts 02142 USA. 232 pages. U.S.$19.95 Paper. What We Know About Climate Change. By K. Emmanuel. 2007. The MIT Press, 55 Hayward Street, Cambridge, Mas- sachusetts 02142 USA. 96 pages. U.S.$14.95 Cloth. BooK REVIEWS 501 Saving Planet Earth. By Tony Juniper. 2007. BBC Books, Random House, 1745 Broadway, 3rd Floor, New York, New York 10019 USA. 256 pages. $39.95 Cloth. + Foraging — Behavior And Ecology. By B. Stephens, J. Brown and R. Ydenberg. 2007. University Chicago Press, 1427 E. 60" Street, Chicago, Illinois USA. 608 pages. U.S. Cloth. The Great Lakes — The Natural History of a Changing Region. By W. Grady. 2007. Greystone Books, Suite 201, Quebec Street, Vancouver, British Columbia VST 487 Cana- da. 320 pages. $49.95 Cloth. MISCELLANEOUS * Beneath My Feet: the Memoirs of George Mercer Daw- son. By Phil Jenkins. 2007. McClelland & Stewart Ltd., 75 Sherbourne Street, 5th Floor, Toronto, Ontario MSA 2P9 Canada. 368 pages. $34.99 Cloth. Modelling for Field Biologists and Other Interesting Peo- ple. 2007. By Hanna Kokko. Cambridge University Press, 32 Avenue of the Americas, New York, New York. 230 pages. GBP 27.99. The World Without Us. By Alan Weisman. 2007. St. Martin’s Press, 175 Sth Avenue New York, New York 10010 USA. 324 pages. U.S.$24.95 Cloth. News and Comment Marine Turtle Newsletter (114) October 2006. 32 pages: EDITORIALS: Editorial (H. H. Godfrey and L. M. Campbell) — Revision of the Kemp’s Ridley Recovery Plan (C. W. Caillouet, Jr.) — Sea turtle subpopulations and the IUCN Red List: A complementary role for conservation genetics (E. Naro-Maciel and A. Formia) — Commentary: What is a subpopulation? (N. Mrosovsky) — ARTICLES: Transatlantic migration of juvenile Loggerhead Turtles from the Strait of Gibraltar (D. Cejudo, N. Varo-Cruz, A. Liria, J. J. Castillo, J. J. Bellido, and L. F. Lopez- Jurado) — On the occurrence of Columbus Crabs (Planes minutus) from Loggerhead Turtles in Florida, USA (M. G. Frick, K. L. Williams, M. Bressette, D. A. Singewald, and Richard M. Herren) — Hawksbill Turtle tracking as part of initial sea turtle research and conservation at Groote (S. D. Whiting, S. Hartley, S. Lalara, D. White, T. Bara, C. Maminyamunja, and L. Wurramarrba) — Lower nesting success of Flatback Turtles caused by disorientation (A. U. Koch and M. L. Guinea) — IUCN-MTSG Quarterly report — MEET- ING REPORTS — LETTER — BOOK REVIEWS — | ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — — RECENT PUBLICATIONS. The Marine Turtle Newsletter is now edited by Lisa | M. Campbell, Nicholas School of Environment and Earth Sciences, Duke University, 135 Duke Marine Lab Road, Beaufort, North Carolina 28516 USA; and Mat- | thew H. Godfrey, North Carolina Wildlife Resources Commission, 1507 Ann Street, Beaufort, North Caro- lina 28516 USA. The Managing Editor is Michael | Coyne, A321 LSRC, Box 90328, Nicholas School of the Environment and Earth Sciences, Duke University, | Durham, North Carolina 27708-0328 USA. Fax +44 1392 263700. Subscriptions and donations towards the production of the MTN can be made online at www. seaturtle.org/mtn or postal mail to Michael Coyne (Man- aging Editor) Marine Turtle Newsletter, | Southampton Place, Durham, North Carolina 27705 USA (e-mail: mcoyne @seaturtle.org). Canadian Association of Herpetologists/Association Canadienne des Herpetologistes Bulletin Fall 2006 14(1) 28 pages. CONTENTS: Instructions for authors — EDITORIAL: On the return of CAH/ACH — A mes- sage from the editor — MEETINGS: CARCNet 2006 Meeting Announcement (Pat Gregory) — JMIH 2006 (Leslie Lowcock) — CAH/ACH Membership and Executive meetings (Jacqueline Litzgus) — FEATURE ARTICLE: On the origins and dispersal of Neotropical rattlesnakes in South America (Adrian Quijada-Mas- carenas and Wolfgang Wuster) — FIELD NOTES: Interesting nesting behaviour of Wood Turtles (Glyp- temys insculpta) (William Greaves) — NEW FACUL- TY IN HERPETOLOGY AT CANADIAN UNIVERSITIES: Gabriel Blouin-Demers, University of Ottawa; David Lesbarreres, Laurentian University; Jacqueline Litzgus, Laurentian University — THESIS ABSTRACTS IN CANADIAN UNIVERSITIES: Karine Beriault, M.Sc. Wildlife Afield 3 (1 and Supplement to 1) Volume 3, Number 1, contains FROM THE EDITOR: Coming together — On the covers — FEATURE ARTI- CLE: Supplement to Wildlife Afield: Roads and wildlife — NOTES: American Avocets breeding at Bechers Prairie near Riskke Creek, British Columbia in 2005 and 2006 (Phil Ranson) — Manx Shearwater in British Columbia: comments on a pioneering seabird (Michael Force, Ken Morgan, and Jukka Jantunen) — A note- worthy record of the Pigmy Short-horned Lizard (Glenn R. Ryder, R. Wayne Campbell, and Lawrence Powell) 2005; Joseph Crowley M.Sc. 2006; Amanda Duffus M.Sc. 2006; Raymond Saumure Ph.D. 2004; Sara Wick | M.Sc. 2004 — RECENT PUBLICATIONS IN CANADIAN HERPETOLOGY: Citations 2005-06 — ANNOUNCE- MENTS: Former CAH/ACH President Michael Rankin remembered (Francis Cook) — MEMBERSHIP FORM. Editor Dr. Jacqueline D. Litzgus, Laurentian Uni- versity, Sudbury, Ontario P3E 2C6; e-mail: jlitzgus@ laurentian.ca; phone 705-675-1151 extension 2314. Membership in CAH/ACH is $20.00 regular member, $10.00 student member payable to the Canadian Asso- ciation of Herpetologists. Mail to Dr. Patrick Gregory, President and Treasurer of CAH/ACH, Department! of Biology, University of Victoria, British Columbia : V8W 2Y2. — Cactus traps California Myotis in British Columbia ‘ (Christian W. Gronau) — Whip-poor-will Caprimulgus\\ vociferus: a new species for British Columbia, 1968-': 2006 (Adrian Dorst) — Status of the Yellow-breasted: Chat in the Creston Valley, British Columbia, 1968- 2006 (R. Wayne Campbell and Edward McMackin) — Unusual mortality of a male Rufous Hummingbird in Burton, British Columbia (Arthur Schoeddert) — Common Raven steals and caches eggs of the Double- crested Cormorant in the Creston Valley, British Col- 502 2006 umbia (Linda M. Van Damme) — Impact of Cooper’s Hawk predation on a nesting colony of Brewer’s Black- birds on Vancouver Island, British Columbia (R. Wayne Campbell and Barbara Begg) — Golden Eagle attacks and kills yearling Mountain Goat (Barbara Zettergreen) — Skeletal remains of the Northern Flicker with a gross bill deformity discovered in a popular tree cavity (Sherry L. Lidstone) — WILDLIFE DATA CENTRE: — Featured species — Common Nighthawk (R. Wayne Campbell, Martin K. McNicholl, R. Mark Brigham, and Janet Ng) — Report of the Wildlife Data Centre: 1 January 2006 to 30 June 2006 (Michael I. Preston) — BRITISH COLUMBIA ROUND-UP: Tips for the field — Field notes for caring people — Errata — News of friends — Life membership — a personal commitment to wildlife — From the Archives — Final Flight — Announcements, Publications of Interest — Volume 3(1), (and Supplement to 3(1), contains: FROM THE EDITOR: At a crossroads — On the covers — ROADS & WILDLIFE: Collisions with wildlife: An overview of major wildlife vehicle collision data collection sys- tems in British Columbia and recommendations for the future (S. Gayle Hesse) — A comparison of roads and rivers as barriers to animal movements: implica- tions for roads in protected areas (Joanna Preston) — High incidence of vehicle-induced ow] mortality in the lower mainland and central Fraser Valley, British Col- umbia (Michael I. Preston and Gerry A. Powers) — Errata The Canadian Field-Naturalist 120(2) Table of contents outside back cover: NEWS AND COMMENT 503 A study on the incidence of amphibian road mortality between Ucluelet and Tofino, British Columbia (Barbara Beasley) — Mitigation efforts to reduce mammal! mor- tality on roadways in Kootenay National Park, British Columbia (Michael I. Preston, Larry Halverson and Gayle Hesse) — Using wildlife vehicle collision data, expert opinion and GPS technology to more accurately predict and mitigate vehicular collisions with wildlife in northern British Columbia (Roy V. Rea, Eric K. Rapaport, Dexter P. Hodder, Michael V. Hurley and Nicole A. Klassen) — Road watch BC: A program for collecting, centralizing, and synthesizing information from vehicle-induced wildlife mortalities in British Columbia (R. Wayne Campbell and Michael I. Pres- ton) — Estimating the probability of potential vehi- cle collision from birds cross roads in interior British Columbia (Michael I. Preston and Joanna Preston) — USEFUL RESOURCES: Books — Symposia, Pro- ceedings, Handbooks, and Reports — Websites — Up- coming meetings — Book review: Road ecology — Road watch BC — Wildlife collision prevention: Hints for the highway. Details on membership can be obtained from the Biodiversity Centre for Wildlife Studies, PO Box 6218, Station C, Victoria, British Columbia V9P 5L5 Canada. Tel/Fax: 250-477-0465; e-mail: editor@ wild lifebc.org; web site: www.wildlifebc.org. Conservation evaluation of Dwarf Wolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada GEORGE W. DOUGLAS, JENIFER L. PENNY, and KSENIA BARTON “Wolly-heads” should read Woolly-heads. Article page 235: First record of a River Otter, Lontra canadensis, captured on the northern coast of Alaska SHAWN P. HASKELL In abstract and citation “Lutra“ should be Lontra. Errata The Canadian Field-Naturalist 120(3) Table of contents outside back cover: Recent invasion, current status, and invasion pathway of European Common Reed, Phragnites australis subspecies australis, in the southern Ottawa District PAUL M. CATLING and SUSAN CARBYN “Phragnites* should read Phragmites. Pacific Hagfish. Eppptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and scavenger activity on tethered carrion in subtidal benthic communities off western Vancouver Island SARAH DAVIES, ALI GRIFFITHS, and T. E. REIMCHEN “Eppptatretus* should read Eptatretus The Ottawa Field-Naturalists’ Club Awards for 2005 IRWIN M. BRobo, CHRISTINE HANRAHAN, BEVERLY MCBRIDE, and ELEANOR ZURBRIGG At the Club’s Annual Soirée, held on 29 April 2006, at St. Basil’s Church in Ottawa, awards were once again given to members, and two non-members, who distin- guished themselves by accomplishments in the field of natural history and conservation, or by extraordinary activity within the Club. For the first time in several years, we had a recipient of the Anne Hanes Natural History Award, and the Mary Stuart Education Award was given for the second time. The following citations for those who received an award were read to the mem- bers and guests assembled for the event. Gillian Marston and Suzanne Deschenes — Members of the Year The OFNC’s Member of the Year award recognizes the member(s) judged to have contributed the most to the club during the previous year. Education and Pub- licity Committee members Gillian Marston and Suz- anne Deschenes have made stellar accomplishments over the past year in developing educational and pub- licity materials for OFNC, and leading a successful membership drive through publicity efforts. They have also brought their considerable computer skills to mod- ernize the Club’s photo library and displays. Gillian and Suzanne are the driving force behind the OFNC’s new photography contest which began in 2005 and is set for a second round in 2006. Meant to encour- age people’s appreciation of nature through photogra- phy, and to boost the club’s library of digital photo- graphs on natural themes, the first contest reaped 275 images, all of which have the potential to be used in displays or slideshows. The contest resulted in a profes- sional, stunning slide show for the 2006 Annual Busi- ness Meeting. All these new images are now catalogued in a digital library, also developed and managed by Suzanne and Gillian. This library is the successor to the club’s long-standing and well used slide collection. Suzanne and Gillian together bring a special synergy of creativity and management skills. They further ap- plied their talents in 2005, taking the club’s new image collection to the public by designing and constructing several portable exhibits to highlight Ottawa nature, the club’s activities, and to attract new members. Members and non-members alike will recall seeing volunteers ee, ee eT z ia | staffing these exhibits at the entrance to popular nature _ walk areas such as the Britannia Conservation Area. Y They also appeared on site for various club excursions \_ and conferences where they successfully attracted a) good number of new members and caught the eye of '} i educators. Gillian and Suzanne and the rest of the com- © mittee continue to develop more portable exhibits and to refine the design. For example, they put together a — set of slide shows on different themes that can be shown) — at exhibits using a table-top projector or laptop com- puter. many special contributions during 2005. Designating you as members of the year is our way of saying “thank you”! Christine Hanrahan — George McGee Service Award Most successful organizations have a handful of members that devote themselves, body and soul, to the workings of the group, serving on committees, on Council, and in any way that they are needed to make the Club succeed. Christine Hanrahan is one of these people. The Ottawa Field-Naturalists’ Club has bene- fited from her efforts and devotion for over 25 years, and we recognize these efforts with the George McGee Service Award. This award honours members who have volunteered their time to the betterment of the Club over several years. The award commemorates George McGee, who for more than three decades actively devoted much of his spare time to teaching people about birds and natural history through his talks and numerous outings. Christine has been a member of the OFNC since the late 1970s. She has been an active member of many Club committees, including the Fletcher Wildlife Gar- den Committee, Conservation Committee, Bird Com-) mittee, and Awards Committee. On top of this, she has: recently coordinated the local group working on they Gillian and Suzanne, the club is grateful for your | | | { Ontario Breeding Bird Atlas, which is now drawing to’ | a close. Co-ordinating over a hundred volunteers for 5 years (plus an additional start-up trial year, plus the) | final data compilation/coordination in 2006) is no small feat. Not only was there all the coordination between’ Atlas headquarters and the region’s 24 volunteers.: which included distributing information packages.’ forms, CDs and other material, but also there were the) annual meetings to coordinate, the annual reports te! write, the articles for Trail & Landscape, and hours anc hours spent on-line writing e-mail messages and check- ing the data of the central database. And this is the second time Christine has done this! She was the leac 504 i q >: r 2005 coordinator in region 24 for the first Atlas project 20 years ago. After that she coordinated the Loggerhead Shrike survey in this area. But our members probably know Christine best through her work on the Fletcher Wildlife Garden Com- mittee over many years. She has diligently kept long- term and short-term lists of many of the FWG occu- pants and visitors (e.g., birds and butterflies). This in- cludes preparing the current sightings board and the checklists. She has published many articles in Trail & Landscape about the activities and happenings at FWG. She has prepared posters, many pamphlets, checklists, nesting reports, etc., if not solely, then as the main driv- ing force. She has organized many important activities at the Centre to draw in and educate visitors. Every year she has prepared the FWG display and organized the hosting (including herself) at the National Capital Region Wildlife Festival. She has also prepared displays at other events such as the Health Canada-sponsored Environment Day. Christine has represented the OFNC and the FWG on the National Capital Wildlife Festival Committee for many years and puts a huge amount of time into organizing workshops and conservation awards. The workshops she has organized have been on topics of great interest to the OFNC, such as Invasive Alien Spe- cies, conservation (Protecting Wild Places) and bird conservation. Another of Christine’s favourite activities is the OFNC Conservation Committee. She has participated in many committee activities and battles including, most recently, the Larose Forest issue. Here she worked on developing checklists of the flora and fauna with vari- ous specialists. She also worked for hours and hours with Ghislaine Rozon reviewing documents, preparing strategy, briefs, writing letters, attending all kinds of meetings in an effort to save the forest from develop- CLUB AWARDS 505 ment. Her work on conserving Petrie Island also had a major impact. Working with The Friends of Petrie Is- land, she compiled lists, wrote articles for Trail & Landscape, organized field trips, as well as attending endless meetings, reviewing documents, and preparing OFNC responses. There have been many other conser- vation issues around Ottawa that have received Chris- tine’s attention and hard work, including: Moffat Farm, Montfort Woods, Nortel Woods, Leitrum wetlands, 5309 Bank Street (Tomlinson quarry) and Ferguson Forest. The list seems endless. And who among us has not read, enjoyed and learned from Christine’s many articles in Trail & Landscape. She has published on birds, plants, habitats and a wide range of conservation issues. Her short article on snags and old logs is a well known, authoritative and influ- ential document. It has been reproduced and widely distributed within the Ontario Ministry of Natural Re- sources and has been used and cited by people even in other parts of the world. Her article on hedgerows and their importance to wildlife is another gem and has become an important resource for ecologists. Since the OFNC has been on the Internet, Christine has been an active contributor to our website with informative and interesting material on a variety of natural history subjects. Because Christine has also served as a knowledge- able member of the Awards Committee for a number of years, recognizing all her achievements and work with an appropriate prize “slipped through the cracks”. Not until her nomination came from outside the com- mittee by a number of members all wondering how she could have been overlooked so long could we make this presentation. The Club is therefore pleased to present Christine Hanrahan with this long overdue and well- deserved Service Award. Friends of the Carp River — Conservation Award (Non-Member) The OFNC Conservation Award, non-member, is given to a group or individual who has made an out- standing contribution in the cause of natural history conservation in the Ottawa Valley, with particular em- phasis on activities within the Ottawa District. The Friends of the Carp River have worked for many years to protect and restore the ecosystem of the Carp River, and this dedication makes them very worthy recipients of this award. The Friends of the Carp River grew out of the West Carleton Township’s Environmental Advisory Commit- tee which began an extensive study of the Carp River in 1993. The organization was formed in 1997 as a citi- zen’s group whose goal was to help restore and improve the health of the Carp River. The river has its headwa- ters just south of the Corel Centre and flows 42 km to _ the Ottawa River at Fitzroy Harbour, and is “the only river that flows entirely within the boundaries of the _ City of Ottawa”. Over the years the river has suffered from shoreline erosion, siltation, flooding, low oxygen levels and subsequent loss of important wildlife habitat. Working with various groups, including landowners, businesses, government agencies and recreational users, the Friends are developing some substantial, achievable goals to continue rehabilitating the “ribbon of life”, their wonderfully descriptive term for the Carp River. Studies show that the Carp River has suffered overall degradation, including silting problems due in part to agricultural tilling practices and channelization. Shore- line erosion further contributes to siltation, which puts at risk the 40 species of fish inhabiting the river. Tribu- taries of the Carp River have severely degraded water quality due largely to agricultural fertilizers and ani- mal wastes. In spring 2006, the threat of development in the floodplain could bring more problems such as future flooding as well as damaging restoration projects. The Friends of the Carp River, an entirely volunteer- run organization, is dedicated to reversing the decline 506 of the river and the Friends have already made tremen- dous improvements. The Friends have planted over 20 000 trees and shrubs since 1997 with the help and cooperation of local landowners living along the river and many volunteers, and this will have a substantial positive impact on controlling erosion and siltation. With assistance from provincial foresters, site plans for tree planting have been created. The Friends have worked with the City of Ottawa on its Watershed study, and gathered support from the Ottawa Stewardship Council and the Rural Clean Water Program. Their work was recognized with a Trillium Foundation grant of $50 000 to help them achieve their goal of rejuvenating the Carp River. One of their objec- tives for this grant was to commission a study of the river, which they did, and the Carp River Remediation Project report was prepared by consultants in April 2003. The Friends of the Carp River have produced a pam- phlet which details ways and means for landowners to achieve best management practices. The Friends are available to help steer farmers and others to specific programs, including funding, to enable them to imple- ment improvements that benefit both landowners and the river. The Friends have been ingenious, creative and tire- less in coming up with ways to garner support for the river. Believing that an “image change” for the river was a necessary part of creating a positive public per- ception, they held a year-long photography contest that encouraged people to submit photos of the Carp River, and also persuaded them to look at the river as a thing Iola Price — Conservation Award (Member) The OFNC Conservation Award is given in recogni- tion of a club member who has made an outstanding contribution towards protecting our natural environ- ment. Iola Price is a very worthy recipient of this award in recognition of her effective membership in the City of Ottawa’s Forest and Greenspace Advisory Committee (OFGAC) to promote conservation of trees and forests. As the current Chair of the Ottawa Forests and Greenspace Advisory Committee, and co-chair before that, Iola provides an articulate and credible voice on issues related to conservation of the trees, forests and greenspace of our Nation’s capital. The Ottawa Forests and Greenspace Advisory Com- mittee was established in the fall of 2001 to advise Ottawa City Council and provide a forum for citizens on issues related to trees and forests. Iola’s accom- plishments include reviewing OFGAC position papers before transmission to appropriate authorities, prepa- ration of numerous written briefs and making power- ful submissions to city committees. For example, Iola recently exhorted City Council at a budget planning meeting to “think of trees as green infrastructure, as essential to a well-ordered city as sewers, lighting and roads”. THE CANADIAN FIELD-NATURALIST Vol. 120 of beauty . Winning entries were used to create a Mil- lennium calendar, Flowing into the next Millennium: A year in the life of the Carp River, which was sold at cost and sold out in three months. The photos were also used to create sets of notecards, again sold at cost. A benefit concert with Terry Tufts and Kathryn Briggs was held on Earth Day, 2001 to help raise funds for continued work on the river. A CD of a song, The Mighty Carp by Dan Mayo, inspired by the river and dedicated to the Friends was released in 2002. The Friends explain that “shorelines are breeding grounds, nurseries, food sources, shelter, and hiding places for many species. Ninety per cent of all living things in a lake or river are found along the shoreline. Shorelines are effective natural filters and buffer zones, acting as the ‘kidneys’ for the watershed. Shorelines help to improve the quality and quantity of ground water. Polluted run-off is trapped and absorbed by the vegetation, thus protecting water quality. The complex interplay of plants, animals, land, and water combine to make the shorelines the single most important part of the river ecosystem.” The Friends have been remarkably successful in drawing the community into active involvement with the health of the river as evidenced not only by the many volunteers who turn out for tree-planting activities but by the tremendous support they have received from local businesses and granting agencies. For these reasons and more, the OFNC is very pleased to give the Friends of the Carp River this year’s Conservation Award for non- members. ; On many occasions, Iola has made a powerful and persuasive intervention during the City’s planning pro- cess in support of the importance of greenspace and protection of sites of urban natural ecological signifi- cance. Her recent interventions regarding the Nepean Creek Corridor as well as her efforts for improving the criteria utilized for determining appropriate urban density development levels underscores this dedica- tion towards the maintenance and protection of trees and greenspace in the National Capital Region (NCR). . Iola has provided recommendations to strengthen 1 ecological provisions in urban design studies such as $ Riverside South Community Design and Leitrim Com- - munity Design. She spearheaded opposition to devel- - opment projects such as the highly controversial Carp ) Ridge housing project and the Phase 2 Leitrim Wetlands development, which, if approved, would extend hous- ing into the wetland itself. Iola is recognized as a very effective and valuable “networker”, regularly liaising with other tree and greenspace-oriented community groups operating in the © NCR. This includes our Club through the Conserva- tion Committee and Fletcher Wildlife Garden. Members © of these committees wholeheartedly appreciate that Iola | | ! i { : | ' | | 2005 is very quick and efficient in replying to queries and in exploring issues and ideas, which helps so much when it allows one to continue one’s work without delay. CLUB AWARDS 507 Through this award, OFNC members recognize the outstanding efforts towards protecting our natural envi- ronment by one of our members — lola Price. Macoun Field Club — Anne Hanes Natural History Award The Anne Hanes Natural History Award, named in honour and memory of one of the Club’s most dedi- cated amateur naturalists, is made in recognition of a member who, through independent study or investiga- tion, has made a worthwhile contribution to our knowI- edge, understanding and appreciation of the natural his- tory of the Ottawa Valley. The award is designed to es- pecially recognize work that is done by amateur natu- ralists. The OFNC feels that there can be no more worthy recipient than the Macoun Field Club in recognition of the remarkable studies the members of the MFC have made of the Macoun Study Area in Bells Corners over a 35 year period, documenting rare plants and animals, tracking changes to plant and animal populations as well as the environment, producing and up-dating vegetation maps of the area, and much more. The MFC, spon- sored jointly by the OFNC and Canadian Museum of Nature, has been a haven for youngsters interested in nature since 1948, and accepts members from grades 4 through 12. The idea of having a special area of natural land des- ignated as a “study area” for members of the Macoun Field Club came from the senior (high school) mem- bers, not from the adult leadership. The first study area in the Bells Corners area was actually where the Wild Bird Centre is now, but because the OMNR was cutting out the dead and dying elms at the time, they suggest- ed an area across the road in a less disturbed site west of Moodie Drive. Within two years, the members had drawn a vegetation map of most of the 800 acres mak- ing up the site. This valuable base map was based on aerial photographs supplemented by field trips to deter- mine what was there. There have been several versions made to keep pace with changes: 1972, 1986, 1992, and 2000. A colour version was created for the Club’s web site (www.magma.ca/~rel/mfc/studyarea/msa.html). In this way, the MFC members traced the changes that occurred in a significant part of the Ottawa Greenbelt brought on by urbanization, beaver activity, tree dis- eases and people. The MFC members, on their bi-week- ly field trips over many years, mostly led by Rob Lee, created lists of trees (54 species), wildflowers, birds, mammals (29 species), reptiles and amphibians, and lichens (215 species). In the process, they discovered many rarities and have documented the disappearance of certain species. Their studies continue on every- thing from “the sand grains in the soil” to “Leather- wood Treehoppers.” Because each member is encouraged to “adopt a tree” and study its growth and health, its inhabitants in the crown, on the trunk and at its base, the MFC has amassed an amazing amount of information about sea- sonal fluctuations in a suburban forest system. Much of this information has been organized on the MFC’s amazing website designed and maintained by Rob Lee, but the information itself was gathered by the members. It is improbable that any urban greenspace in Canada has such a well-documented history of its flora and fauna. Observations on the Macoun Study Area have been regularly published by the members in their annual magazine, The Little Bear, and, because of the Internet, is now readily available to the general public as well. Most important, the experience that the members have gained in doing careful field research has encouraged many past members to embark on a career in conser- vation biology, botany, zoology, park management or teaching. The OFNC feels that the Macoun Field Club mem- bers have done a remarkable job, under the leadership of Rob Lee, and well deserves the Anne Hanes Natural History Award. Michael Léveillé — Mary Stuart Education Award The Mary Stuart Education Award was established to recognize members, non-members or organizations for their outstanding achievements in the field of nat- ural history education in the Ottawa Region. Michael Léveillé, a science teacher at the Educarium, an Ottawa private school, is without doubt a natural history edu- cator par excellence, and a most worthy recipient of this Award. Michael has taught at the Educarium since 1996. His passion for nature is infectious and inspirational and is observed most keenly by his students. Michael brings an impressive background to his role as science _ teacher. He founded the Ottawa Paleontology Society in 1991 to advance an understanding of fossils. He has long been associated with the Canadian Museum of Nature where he created a number of popular programs, most notably the “Collector's Corner’, a museum dis- covery centre. At the museum he worked with people of all ages, instilling an awareness of, and appreciation for, the natural world. However, there is no doubt that Michael excels at working with young people, encour- aging them to be attuned to the rhythms of the natural world around them. Michael is also an accomplished artist with a degree in Visual Arts from the University of Ottawa. His work has been exhibited at local universities, museums, as well as at the Educarium. Michael was the artist for the children’s storybook series “Fergi the Frog”. 508 Under Michael’s knowledgeable direction, in 2004 his students designed and implemented the “Macoun Marsh Project” at Beechwood Cemetery, to demon- strate how a thriving and diverse ecosystem can exist inside a large urban centre. This became one of their most ambitious and high profile projects, and is ongo- ing. Because the marsh was unnamed when they began working on it, they held a “Marsh Naming Contest” and encouraged neighbours and the community at large to join in. Overwhelming support led to naming it after John Macoun (1831-1920) who is buried at Beech- wood. The contest, and the work done by the students, led to increased public awareness of the site and even- tually resulted in the Directors of Beechwood Cemetery making a firm commitment to protect and maintain it. The Macoun Marsh project encourages students to study the marsh over different seasons. They are taught to record scientific observations in journals, identify and study the different species found, and learn the fun- damentals of ecology. They also design, construct, install and maintain bird feeders at the site. Under Michael’s THE CANADIAN FIELD-NATURALIST Vol. 120 careful instruction, the Educarium students have doc- umented nearly 900 species at the marsh. The project has grown beyond their wildest expec- tations. Funding to continue with studies and enhance- ments has been obtained from a variety of sources in- cluding the Wetland Habitat Fund and the City of Ottawa. In conjunction with the Beechwood Cemetery Foundation, plans are underway for habitat enhance- ment, a boardwalk, and an outdoor classroom in spring 2006. The students continue to keep nature journals and will present this work at various venues around the city over the coming year. Michael invited the neigh- bouring Jean Vanier Catholic School to join his students on the project, and in May 2006 both schools sent rep- resentatives to Géteborg, Sweden, for the Volvo Adven- ture International contest for natural history projects. Michael’s inspired teaching guarantees that his stu- dents will continue to be engaged by the world of nature, and so it is for this most important work, that OFNC is pleased to present Michael Léveillé with the Mary Stuart Education Award for 2005. \ Index to Volume 120 Compiled by Leslie Cody Abies amabilis, 172 balsamea, 100,236,238,299,433,440 lasiocarpa, 100,331 Accipiter striatus, 46 Acer spp., 226 circinatum, 11,171,348 glabrum var. douglasii, 171 macrophyllum, 11,180 rubrum, 299,354 saccharum, 299 Achillea millefolium, 354 Achnatherum hymenoides, 336 Acipenser fulvescens, 71 transmontanus, 422 Acipenser fulvescens, Movements in Rainy Lake, Minnesota and Ontario, Lake Sturgeon, 71 Acorn, J., Reviews by, 245,489 Adams, W.E. Jr., L.W. Kallemeyn, and D.W. Willis. Lake Sturgeon, Acipenser fulvescens, Movements in Rainy Lake, Minnesota and Ontario, 71 Adiantum capillus-veneris, 138 Aegolius acadicus, 294,303 Aeshna sp., 416 canadensis, 415 constricta, 417 eremita, 417 interrupta, 416 interrupta lineata, 416 interrupta lineata ssp. lineata, 416 umbrosa, 416 umbrosa umbrosa, 417 Agalinis neoscotica, 357 Agriotes collaris, 19 stabilis, 19 Agropyron cristatum, 337 Agrostis exerata, 214 tenuis, 354 Agujaceratops, 403 Aira caryophyllea, 320 praecox, 154,320 Aix sponsa, 236,303 Alaska, First Record of a River Otter, Lontra canadensis, Cap- tured on the Northeastern Coast of, 235 Alaska, Physical Characteristics, Hematology, and Serum Chemistry of Free-ranging Gray Wolves, Canis lupus, in Southcentral, 205 Alaska, 2001-2003, Assessment of Effects of an Oil Pipeline on Caribou, Rangifer tarandus granti, Use of Riparian Habitats in Arctic, 323 Alberta, A New Record of Deepwater Sculpin, Myoxocephalus thompsonii, in Northeastern, 480 Alberta Boreal Forests, Winter Habitat Use by American Marten, Martes americana, in Western, 100 Alberta, Pack Size of Wolves, Canis lupus, on Caribou, Ran- gifer tarandus, Winter Ranges in Westcentral, 313 Alces alces, 205,313,367 alces americana, 5| Alder, 236,240 Red, 11,348 Speckled, 22 Allium amplectens, 160 cernuum, 154 Alnus sp., 236,238,240 incana, 22 rubra, 348 rubus, \\ Alopecurus carolinianus, 164 Alopex lagopus, 36 Alosa sapidissima, 422 Ambystoma gracile, 90 macrodactylum, 90 Ameiurus catus, 424 melas, 422 natalis, 421 nebulosus, 422 Ameiurus natalis, a Loricariid Catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia, First Records of the Yellow Bull- head, 421 Amelanchier, 428 alnifolia, 176,428 alnifolia var. compacta, 428 florida, 428 humilis, 428 humilis var. compacta, 428 laevis, 354 laevis x lucida, 354 lucida, 354 sanguinea, 428 Amelanchier in the Whitewood area of Southeastern Saskat- chewan and the First Saskatchewan Records of Ame- lanchier sanguinea, Regional Variation in, 428 Amelanchier sanguinea, Regional Variation in Amelanchier in the Whitewood area of Southeastern Saskatchewan and the First Saskatchewan Records of, 428 Amia calva, 422 Ammodytes americanus, 96 Ammospiza leconteii, 22 Ammospiza leconteii, from Northeastern Ontario, with Some Notes on Nesting Behaviour, New Nesting Records of the Le Conte’s Sparrow, 22 Anaphalis margaritacea, 354 Anas clypeata, 5 platyrhynchos, 6 Anax junius, 417 Anderson, C.S., D.B. Meikle, A.B. Cady, and R.L. Schaefer. Annual Variation in Habitat Use by White-footed Mice, Peromyscus leucopus: The Effects of Forest Patch Size, Edge and Surrounding Vegetation Type. 192 Andrena, 19 509 510 Anguilla rostrata, 238,477 Anguille d’ Amérique, 238 Antennaria flagellaris, 183 howellii ssp. neodioica, 354 Antennaria flagellaris, in Canada, Conservation Evaluation of Stoloniferous Pussytoes, 183 Anthophora, 180 Anthoxanthum odoratum, 148,154,160 Antilocapra americana, 210 Apera interrupta, 320 Apis mellifera, 294 Apocynum androsaemifolium vat. incanum, 354 Apodemus sylvaticus, 195 Apple-Producing Region of Southern Quebec, First Obser- vations of an Eastern Screech-Owl, Megascops asio, Population in an, 289 Aralia hispida, 354 Arbutus menziesii, 148,180 Arbutus, 148,180 Arctostaphylos uva-ursi, 172,336,354 Arenaria capillaris ssp. americana, 184 Aronia sp., 353 Aronia, 360 Artemisia cana ssp. cana, 336 tridentata, 164,184 Ascophyllum nodosum, 236 Asio otus, 295 Aspen, 314,428 Trembling, 22,100,342 Astragalus miser, 176 Athous acanthus, 19 Auditor’s Report, 398 Aulnes, 238 Aythya sp., 6 affinis, 5 americana, 109 collaris, 6 marila, 109 valisneria, 109 Baccha elongata, 17 Badger, 209 American, 31,342 European, 52 Badiou, P.H.J. and L.G. Goldsborough. Northern Range Expan- sion and Invasion by the Common Carp, Cyprinus car- pio, of the Churchill River System in Manitoba, 83 Balaena mysticetus, 462 Balaenoptera acutorostrata, 95 borealis, 97 musculus, 95 physalus, 95 Balanus improvisus, 106 Baldwin, R.A., A.E. Houston, M.L. Kennedy, and P.S. Liu. Predicting Raccoon, Procyon lotor, Occurrence Through the Use of Microhabitat Variables, 225 Balkwill, D., B.W. Coad, I. Galvez, and J. Gilhen. First Record of the Great Barracuda, Sphyraena barracuda from Canada, 241 Ballard, W.B., 205 Balsamorhiza sagittata, 176 Balsamroot, Arrow-leaved, 176 Barley, 343 Barracuda, Great, 241 Barracuda, Sphyraena barracuda from Canada, First Record of the Great, 241 THE CANADIAN FIELD-NATURALIST Vol. 120 Barton, K., 163,183 Bartonia virginica, 357 Basquill, S., 351 Bass, Largemouth, 422 Smallmouth, 421 Striped, 422 Bat, Big Brown, 39 Little Brown, 39,51 Long-legged, 39 Northern Long-eared, 39,51,332 Silver-haired, 39 Western Long-eared, 39 Bat, Myotis septentrionalis, in the Yukon Territory, First Records of the Northern Long-eared, 39 Bdellarogatis, 444 plumbeus, 444 Bear, American Black, 51 Black, 57,68,110,209,314 Brown, 57 Grizzly, 314 Bear, Ursus americana, Retrieval of an Elk, Cervus elaphus, Carcass from a Small Lake in Riding Mountain Na- tional Park, Manitoba, Probable Black, 110 Bear, Ursus americanus L., Long-range Homing by an Adult Female Black, 57 Bearberry, Common, 336 Beaudoin, A.B., Reviews by, 383,493,494 Beaver, 51,62,205,331,368 Bee, 15 Honey, 294 Beetle, 296 Click, 15 Ground, 16 Beluga, 457 Bertrand, A.-S., S. Kenn, D. Gallant, E. Tremblay, L. Vasseur, and R. Wissink. MtDNA Analyses on Hair Samples Confirm Cougar, Puma concolor, Presence in South- ern New Brunswick, Eastern Canada, 438 Betula spp., 428,440 alleghaniensis, 299 cordifolia, 299 papyrifera, 22,50,238,299,354,433,440 populifolia, 354 Birch, 428,440 Mountain, 299 Paper, 433 White, 22,50,299,440 Yellow, 299 Bird, D.M., 289 Bison bison athabascae, 68 Bison, Wood, 68 Bitter-cress, Little Western, 180 Bizecki Robson, D. A Conservation Evaluation of Smooth Goosefoot, Chenopodium subglabrum (Chenopodi- aceae), in Canada, 335 Blackberry, 359 Blaney, S., 319 Blarina brevicauda, 294,436 Blastocerms dichotomus, 64 Blinn, B.M., V. Violette, and A.W. Diamond. Osprey, Pandion haliaetus, Depredates Common Eider, Somateria mol- lissima, Duckling, 236 Blue-eyed Mary, Large-flowered, 180 Blue-Eyed-Grass, Coastal-Plain, 359 Blueberry, 360 Alaskan, 172 2007 Bluegill, 422 Bluegrass, Annual, 320 Bulbous, 320 Kentucky, 160 Bluejoint, Canada, 22 Bluet, Azure, 417 Big, 417 Familiar, 417 Hagen’s, 417 Marsh, 417 Blush, Sea, 154,158 Bobcat, 230 Bobwhite, Northern, 452 Bobwhites, Colinus virginianus, by Mammalian Mesopreda- tors: Does the Problem-Individual Paradigm Fit?, Predation on Artificial Nests of Northern, 452 Bombus, 180 perplexus, 19 vagans, 19 Bombycilla cedrorum, 294 Boreal Dip Net/L’Epuisette Boréale January 2006, The, 392 Bouleau blanc, 238 Bowfin, 422 Brachyopa, 20 notata, 17 Brachypalpus oarus, 17 Branta canadensis, 109 British Columbia, A Test of Interspecific Effects of Introduced Eastern Grey Squirrels, Sciurus carolinensis, on Dou- glas’s Squirrels, Tamiasciurus douglasii, in Vancou- ver, 10 British Columbia, Early Ontogenetic Diet in Gray Wolves, Canis lupus, of Coastal, 61 British Columbia, First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque sut- tonorum, and a Silver Pacu, Piaractus cf. P. brachy- pomus, in, 421 British Columbia, Life History Phenology and Sediment Size Association of the Dragonfly Cordulegaster dorsalis (Odonata: Cordulegastridae) in an Ephemeral Habitat in Southwestern, 347 Brome, 148,154 Barren, 160,180 Bromus, 148,154 sterilis, 160,180 tectorum, 176 Broom, Scotch, 148,155,160 Brunelle, P.M., 413 Bryan, H.M., C.T. Darimont, T.E. Reimchen, and P.C. Paquet. Early Ontogenetic Diet in Gray Wolves, Canis lupus, of Coastal British Columbia, 61 Bryant, J.E. A Tribute to Nicholas Stephen Novakowski 1925-2004, 370 Bubo virginianus, 290 Bucephala albeola, 6 clangula, 5,109 Buckwheat, 343 Cushion, 184 Parsnip-flowered, 184 Budworm, Spruce, 50,299 Bufflehead, 6 Bufo americanus, 240 boreas, 87 Bullhead, Black, 422 Brown, 422 INDEX TO VOLUME 120 S11 Yellow, 421 Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque sut- tonorum, and a Silver Pacu, Piaractus cf. P. brachy- pomus, in British Columbia, First Records of the Yellow, 42] Bulrush, American Three-square, 213 Bunchberry, 440 Bunker-Popma, K. Scoter, Melanitta spp., Migrations Inter- rupted by Confederation Bridge: An Update, 232 Bur-reed, Narrowleaf, 240 Burbot, 422,480 Burchsted, F., 106 Burchsted, J.C.A. and F. Burchsted. Lady Crabs, Ovalipes ocellatus, in the Gulf of Maine, 106 Butcher, M.K., 323 Butler, M.J., W.B. Ballard, and H.A. Whitlaw. Physical Char- acteristics, Hematology, and Serum Chemistry of Free-ranging Gray Wolves, Canis lupus, in Southcen- tral Alaska, 205 Buttercup, Bur, 319 Buttercup, Ranunculus testiculatus, New to Eastern Canada, Bur, 319 Butterfly, Maritime Copper, 419 Cabbage, Skunk, 172 Cady, A.B., 192 Calamagrostis sp., 354 canadensis, 22 Calamovilfa longifolia, 336 Calidris spp., 5 Caligus, 199 clemensi, 199 Camarasaurus, 408 Camas, Common, 148 Camassia quamash, 148 Canada, A Conservation Evaluation of Smooth Goosefoot, Chenopodium subglabrum (Chenopodiaceae), in, 335 Canada, Bur Buttercup, Ranunculus testiculatus, New to East- ern, 319 Canada, Conservation Evaluation of Pacific Rhododendron, Rhododendron macrophyllum, in, 169 Canada, Conservation Evaluation of Slender Collomia, Col- lomia tenella, in, 175 Canada, Conservation Evaluation of Stoloniferous Pussytoes, Antennaria flagellaris, in, 183 Canada, Conservation Evaluation of the Bog Bird’s-foot Tre- foil, Lotus pinnatus, in, 157 Canada, Conservation Evaluation of the Pacific Population of Dwarf Woolly-heads, Psilocarphus brevissimus vat. brevissimus, in, 163 Canada, Conservation Evaluation of the Prairie Lupine, Lupinus lepidus var. lepidus, in, 147 Canada, Conservation Evaluation of the Seaside Birds-foot Trefoil, Lotus formosissimus, in, 153 Canada, Conservation Evaluation of the Small-flowered Tonel- la, Tonella tenella, in, 179 Canada, Extension of Coyote, Canis latrans, Breeding Range in the Northwest Territories, 67 Canada, First Record of the Great Barracuda, Sphyraena barracuda trom, 241 Canada, MtDNA Analyses on Hair Samples Confirm Cougar, Puma concolor, Presence in Southern New Brunswick, 438 Canada’s Maritime Provinces, Recent Declines of House Spar- rows, Passer domesticus, in, 43 2 Canadian Arctic, Reactions of Narwhals, Monodon monoc- eros, to Killer Whale, Orcinus orca, Attacks in the Eastern, 457 Canadian Association of Herpetologists/Association Cana- dienne des Herpetologistes Bulletin, 502 Canadian Field-Naturalist, Advice for Contributors to The, 134,262,402,534 Canadian Field-Naturalist 119(3), Errata The, 133 Canadian Field-Naturalist 120(1), Errata The, 259 Canadian Field-Naturalist 120(2), Errata The, 503 Canadian Field-Naturalist 120(3), Errata The, 503 Cancer magister, 364 productus, 363 Canis sp., 263 familiaris, 263479 latrans, 27,31,40,50,67,110,209,263,314,342,472,474, 478 lupus, 27,57,61,67,111,205,263,313,367,466,47 1,476, 478 lupus arctos, 466,471 lupus beothucus, 50 lycaon, 27 rufus, 27 Canis latrans, An Ethogram Developed on Captive Eastern Coyotes, 263 Canis latrans, Breeding Range in the Northwest Territories, Canada, Extension of Coyote, 67 Canis latrans, Conditions for Sexual Interactions Between Wild Grey Wolves, Canis lupus, and Coyotes, 27 Canis latrans, in a Heavily Urbanized Area: Implications for Ecosystem Management, Use of “Micro’-Corridors by Eastern Coyotes, 474 Canis latrans, in an Urban Area, Rat Poison Kills a Pack of Eastern Coyotes, 478 Canis lupus, and Coyotes, Canis latrans, Conditions for Sexual Interactions Between Wild Grey Wolves, 27 Canis lupus arctos: Influence of Order on Posture, Double Marking in Arctic Wolves, 471 Canis lupus arctos, in Summer, Urine-marking and Ground-scratching by Free-ranging Arctic Wolves, 466 Canis lupus, Group for Ontario, A New Record Size Wolf, 367 Canis lupus, in Southcentral Alaska, Physical Characteristics, Hematology, and Serum Chemistry of Free-ranging Gray Wolves, 205 Canis lupus, of Coastal British Columbia, Early Ontogenetic Diet in Gray Wolves, 61 Canis lupus, on Caribou, Rangifer tarandus, Winter Ranges in Westcentral Alberta, Pack Size of Wolves, 313 Canola, 343 Canvasback, 109 Carassius auratus, 422 Carbyn, L., Review by, 249 Carbyn, S., 307 Carbyn, S., P.M. Catling, S.P. Vander Kloet, and S. Basquill. An Analysis of the Vascular Flora of Annapolis Heath- lands, Nova Scotia, 351 Cardamine oligosperma, 180 Cardinal, Northern, 294 Cardinalis cardinalis, 294 Carduellis tristis, 294 Carex atlantica, 357 cumulata, 354 deflexa var. deflexa, 354 emonsii, 354 THE CANADIAN FIELD-NATURALIST Vol. 120 feta, 160 lucorum, 360 lyngbyei, 213 nigra, 354 oligosperma, 22 paleacea, 416 praegracilis, 310 scoparia, 354 siccata, 360 stipata, 22 tonsa, 358 tonsa vat. rugosperma, 354 tonsa var. tonsa, 354 tumulicola, 160 Caribou, 313,323 Barren-ground, 205 Woodland, 50,313,367 Caribou, Rangifer tarandus granti, Use of Riparian Habitats in Arctic Alaska, 2001—2003, Assessment of Effects of an Oil Pipeline on, 323 Caribou, Rangifer tarandus, Winter Ranges in Westcentral Alberta, Pack Size of Wolves, Canis lupus, on, 313 Carp, Common, 83,422 Carp, Cyprinus carpio, of the Churchill River System in Mani- toba, Northern Range Expansion and Invasion by the Common, 83 Carver, D.B., 452 Carya spp., 226 Castor canadensis, 62,205,331,368 canadensis caecator, 51 Cat, Domestic, 46 Cat’s-ear, Hairy, 154 Caterpillar, Forest Tent, 433 Catfish, Black, 425 Blue, 424 Brown, 425 Channel, 422 Flathead, 424 Loricariid, 421 White, 424 Yellow, 425 Catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia, First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid, 421 Catling, P., Review by, 249 Catling, P.M., 351 Catling, P.M. and G. Mitrow. Regional Variation in Amelanchi- er in the Whitewood area of Southeastern Saskatche- wan and the First Saskatchewan Records of Amelanchi- er sanguinea, 428 Catling, P.M. and S. Carbyn. Recent Invasion, Current Status and Invasion Pathway of European Common Reed, Phragmites australis subspecies australis, in the Southern Ottawa District, 307 Catling, P.M., R. Hutchinson, and P.M. Brunelle. Use of Salt- marsh by Dragonflies (Odonata) in the Baie des Cha- leurs Region of Quebec and New Brunswick in Late Summer and Autumn, 413 Catostomus catostomus, 422,481 commersoni, 422 Cedar, White, 300 Celithemis elisa, 417 Centaurea nigra, 354 Centrosaurus apertus, 408 Cerastium pumilum, 320 2007 semidecandrum, 320 Ceratocephalus testiculatus, 319 Ceratodon purpureus, 358 Certhia americana, 303 Cervus elaphus, 110,188,314 Cervus elaphus, Calf Observed Near Bancroft, Ontario, Late- born Elk, 188 Cervus elaphus, Carcass from a Small Lake in Riding Moun- tain National Park, Manitoba, Probable Black Bear, Ursus americana, Retrieval of an Elk, 110 Chalcosyrphus, 20 nemorum, \7 Chamaecyparis nootkatensis, 172 Chamaedaphne calyculata, 23 Channa sp., 424 Char [Charr], Arctic, 199,235,422 Chasmosaurus, 403 belli, 403 humeri, 408 irvinensis, 403 mariscalensis, 403 russelli, 403 Chasmosaurus belli (Dinosauria: Ornithischia), A Reevalu- ation of Sexual Dimorphism in the Postcranium of the Chasmosaurine Ceratopsid, 403 Cheatgrass, 176 Cheilosia rita, 17 tristis, 17 Chen caerulescens caerulescens, 213 Chen caerulescens caerulescens, Exceeding the Carrying Capacity of the Fraser River Delta’s Brackish Marsh- es?, Are Lesser Snow Geese, 213 Chenopodium album, 337 bonus-henericus, 340 fremontii, 340 pratericola, 336 quinoa, 340 subglabrum, 335 Chenopodium subglabrum (Chenopodiaceae), in Canada, A Conservation Evaluation of Smooth Goosefoot, 335 Cherry, 360 Pin, 50 Chickadee, Black-capped, 294,302 Boreal, 302 Chickweed, Curtis’ Mouse-ear, 320 Lesser, 320 Small Mouse-ear, 320 Chimpanzee, 264 Chipmunk, 264,295 Eastern, 51 Chitonotus pugetensis, 364 Choristoneura fumiferana, 50,299 Chub, Flathead, 422 Lake, 422 Cifuni, S.M., 478 Circus cyaneus, 109 Cisco, 480 Shortjaw, 482 Cladina rangiferina, 358 stellaris, 358 Cladonia spp, 171 Cladonia, 171 Clarias sp., 424 Clarke, J.B., Review by, 247 Classen, M.-C., 347 Claytonia perfoliata, 180 INDEX TO VOLUME 120 513 Cleavers, 180 Clethrionomys sp., 291 gapperi, 50,331 Clethrionomys gapperi, in Newfoundland: Implications for the Endangered Newfoundland Marten, Martes amer- icana atrata, First Record of the Southern Red-Backed Vole, 5O Clintonia borealis, 357 Cluff, H.D. Extension of Coyote, Canis latrans, Breeding Range in the Northwest Territories, Canada, 67 Clupea harengus, 96 Coad, B.W., 241 Coad, B.W. and A.M. Murray, Review by, 116 Coad, B.W., Review by, 114 Cody Receives Yukon Biodiversity Awareness Award, Bill, 258 Coenonypha tullia nipisiquit, 419 Colaptes auratus, 294,303 Coleoptera, 296 Colinus virginianus, 452 Colinus virginianus, by Mammalian Mesopredators: Does the Problem-Individual Paradigm Fit?, Predation on Artificial Nests of Northern Bobwhites, 452 Collinsia, 180 grandiflora, 180 Collomia linearis, 176 tenella, 175 Collomia, Narrow-leaved, 176 Slender, 175 Collomia, Collomia tenella, in Canada, Conservation Evalu- ation of Slender, 175 Collomia tenella, in Canada, Conservation Evaluation of Slen- der Collomia, 175 Colorado, Multiple Scale Den Site Selection by Swift Foxes, Vulpes velox, in Southeastern, 31 Colossoma sp., 424 bidens, 424 Columba spp., 57 livia, 294 Comptonia peregrina, 354 Coot, 2 Cordulegaster bidentata, 350 boltonii, 347 dorsalis, 347 heros, 350 maculata, 347 Cordulegaster dorsalis (Odonata: Cordulegastridae) in an Ephemeral Habitat in Southwestern British Columbia, Life History Phenology and Sediment Size Association of the Dragonfly, 347 Coregonus artedi, 480 clupeaformis, 422,480 zenithicus, 482 Corema, 357 conradii, 351 Corn, 226,294 Cornus canadensis, 354,440 Corvus brachyrhyncos, 109 corax, 109,110 Corylus cornuta, 170 Cotton, 226 Cottontail, Nuttall’s, 165 Cottonwood, 226 Cottus asper, 425 Couesius plumbeus, 422 Cougar, 40,314.438.476 Eastern, 438 514 Cougar, Puma concolor, Presence in Southern New Bruns- wick, Eastern Canada, MtDNA Analyses on Hair Samples Confirm, 438 Cow-wheat, 170 Cowbird, Brown-headed, 294 Coyote, 27,31,40,67,110,209,314,342,472 Eastern, 50,263,474,478 Coyote, Canis latrans, Breeding Range in the Northwest Territories, Canada, Extension of, 67 Coyotes Canis latrans, An Ethogram Developed on Captive Eastern, 263 Coyotes, Canis latrans, Conditions for Sexual Interactions Between Wild Grey Wolves, Canis lupus, and, 27 Coyotes, Canis latrans, in a Heavily Urbanized Area: Impli- cations for Ecosystem Management, Use of “Micro”- Corridors by Eastern, 474 Coyotes, Canis latrans, in an Urban Area, Rat Poison Kills a Pack of Eastern, 478 Crab, Dungeness, 364 Lady, 106 Redrock, 363 Crabs, Ovalipes ocellatus, in the Gulf of Maine, Lady, 106 Crane, Sandhill, 68 Crappie, Black, 422 Crataegus cf. chrysocarpa, 354 Creeper, Brown, 302 Crepis tectorum, 354 Crins, W.J., 15 Crocuta croenta, 52 Cronin, M.A., 323 Crow, American, 109 Ctenicera cruciatus, 20 hieroglyphica, 19 insidiosa, 19 resplendens, 19 spinosa, 19 triundulata, 18 vulnerata, 18 Cudweed, Lowland, 164 Culaea inconstans, 422 Curran, W.J., 50 Cutthroat, West-slope, 421 Cyanocitta cristata, 294 Cyclopterus lumpus, 464 Cygnus buccinator, 220 Cynomys ludovicianus, 31 Cynosurus echinatus, 148 Cyprinella lutrensis, 424 Cyprinus carpio, 83,422 Cyprinus carpio, of the Churchill River System in Manitoba, Northern Range Expansion and Invasion by the Com- mon Carp, 83 Cypripedium acaule, 354 Cytisus scoparius, 148,155,160 Dactylis glomerata, 154,160 Daisy, Cut-leaved, 184 Salish, 137 Dalopius spp., 19 cognatus, 19 fuscipes, 19 Dandelion, Common, 320 Danthonia spicata, 354 unispicata, 164,184 Darimont, C.T., 61 Darner, Canada, 417 THE CANADIAN FIELD-NATURALIST Vol. 120 Common Green, 417 Lake, 417 Lance-tipped, 417 Shadow, 417 Variable, 417 Dasysyrphus pauxillus, 17 Davies, S., A. Griffiths, and T.E. Reimchen. Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and Scavenger Activity on Tethered Carrion in Subtidal Benthic Communities off Western Vancouver Island, 363 Deer, 68 Black-tailed, 63 Marsh, 64 Mule, 40,314 Pampas, 64 Sitka Black-tailed, 61 White-tailed, 29,40,64,68,209,3 13,367 Delphinapterus leucas, 457 Delphinium menziesii, 154 Delphinus delphis, 97 Demarchi, M.W. Are Lesser Snow Geese, Chen caerulescens caerulescens, Exceeding the Carrying Capacity of the Fraser River Delta’s Brackish Marshes?, 213 Deschampsia danthonioides, 164 flexuosa, 354 Desert-parsley, Swale, 184 Desroches, J.-F. Extension de |’ aire de distribution de 1’ Umbre de vase, Umbra limi, dans le nord-est du Québec, 238 Desroches, J.-F., I. Picard, and J.E. Maunder. The Mink Frog, Rana septentrionalis, in Southeastern Labrador, 239 Desrosiers, J., 169 Dewey, T.A., 39 Diamond, A.W., 236 Dianthus armeria, 354 Dichanthelium (sub Panicum) depauperatum vat. psilophyl- lum f. cryptostachys, 360 acuminatum vat. fasciculatum, 354 depauperatum, 354 linearifolium, 354 sabulorum var. thinium, 360 Dicranum sp., 180 scoparium, 358 Dicrostonyx groenlandicus, 466 Didelphis virginiana, 229,452 Diervella lonicera, 354 Distichlis spicata, 214 Doellingeria umbellata var. umbellata, 354 Dog, 479 African Wild, 52 Domestic, 263 Dogtail, Hedgehog, 148 Dollar, Silver, 424 Dolphin, Atlantic White-sided, 95 Common Bottlenose, 97 Dusky, 97 Short-beaked Common, 97 White-beaked, 95 White-sided, 95 Donaldson, M.R., Review by, 121 Donovan, M. Conservation Evaluation of the Bog Bird’s-foot Trefoil, Lotus pinnatus, in Canada, 157 Dorocordulia lepida, 417 Douglas, G.W. and J. Desrosiers. Conservation Evaluation of Pacific Rhododendron, Rhododendron macrophyllum, in Canada, 169 2007 Douglas, G.W. and J.L. Penny. Conservation Evaluation of Slender Collomia, Collomia tenella, in Canada, 175 Douglas, G.W. and J.L. Penny. Conservation Evaluation of the Small-flowered Tonella, Tonella tenella, in Canada, 179 Douglas, G.W. and M. Ryan. Conservation Evaluation of the Prairie Lupine, Lupinus lepidus var. lepidus, in Cana- da, 147 Douglas, G.W. and M. Ryan. Conservation Evaluation of the Seaside Birds-foot Trefoil, Lotus formosissimus, in Canada, 153 Douglas, G.W., J.L. Penny, and K. Barton. Conservation Eval- uation of Stoloniferous Pussytoes, Antennaria fla- gellaris, in Canada, 183 Douglas, G.W., J.L. Penny, and K. Barton. Conservation Eval- uation of the Pacific Population of Dwarf Woolly- heads, Psilocarphus brevissimus var. brevissimus, in Canada, 163 Douglas, H., 15 Douglas 1938 — 2005, A Tribute to George Wayne, 135 Douglas-fir, 148,158,164,169,176,184,348 Coastal, 180 Dove, Mourning, 294 Rock, 294 Dragonflies (Odonata) in the Baie des Chaleurs Region of Quebec and New Brunswick in Late Summer and Autumn, Use of Saltmarsh by, 413 Dragonfly, 347 Dragonfly Cordulegaster dorsalis (Odonata: Cordulegastri- dae) in an Ephemeral Habitat in Southwestern British Columbia, Life History Phenology and Sediment Size Association of the, 347 Dragonlet, Seaside, 417 Drosera intermedia, 354 Dryocopus pileatus, 299 Duck, 2 Common Eider, 236 Ring-necked, 6 Wood, 236,303 Duck Nests in Saskatchewan, Summer Movements and Impact of Individual Striped Skunks, Mephitis mephitis, on, 342 Ducks, Ring-billed Gull, Larus delawerensis, Food Piracy on Diving, 109 Duffe, J., 289 Eagle, Bald, 109,237,477 Eagle, Haliaeetus leucocephalus, Preying on Maritime Garter Snake, Thamnophis sirtalis pallidulus, on Cape Bre- ton Island, Nova Scotia, Bald, 477 Eatough, D.L., 474,478 Eel, American, 477 Editor’s Report for Volume 119, 260 Eider, Common, 233 Eider, Somateria mollissima, Duckling, Osprey, Pandion hali- aetus, Depredates Common, 236 Elk, 110,188,314 Elk, Cervus elaphus, Calf Observed Near Bancroft, Ontario, Late-born, 188 Elk, Cervus elaphus, Carcass from a Small Lake in Riding Mountain National Park, Manitoba, Probable Black Bear, Ursus americana, Retrieval of an, 110 Elymus canadensis, 336 lanceolatus ssp. lanceolatus, 336 repens, 355 trachycaulus, 184 INDEX TO VOLUME 120 515 Emerald, Kennedy's, 417 Petite, 417 Ski-tipped, 417 Emphoropsis, 180 Enallagma aspersum, 417 civile, 413 durum, 417 ebrium, 416 hageni, 415 Enteromorpha sp., 417 Epigaea repens, 355 Epilobium angustifolium, \7\ densiflorum, 160 Epinettes, 238 Epistrophe nitidicollis, \7 Eptatretus stoutii, 363 Eptatretus stoutii, Spotted Ratfish, Hydrolagus colliei, and Scavenger Activity on Tethered Carrion in Subtidal Benthic Communities off Western Vancouver Island, Pacific Hagfish, 363 Eptesicus fuscus, 39 Equus cabalus, 314 Ericameria nauseosus var. speciosa, 184 Erigeron, 137 compositus var. glabratus, 184 leibergii, 137 salishii, 137 strigosus, 355 Eriogonum heracleoides var. angustifolium, 184 ovalifolium var. nivale, 184 Eristalis dimidiatus, 17 Erophila verna, 320 Erpobdella, 449 Erskine, A.J. Recent Declines of House Sparrows, Passer domesticus, in Canada’s Maritime Provinces, 43 Erythrodiplax berenice, 413 Esox lucius, 421,481 Eubalaena glacialis, 95 Eupeodes americanus, 17 perplexus, 17 pomus, 17 Euphausiid, 96 Euphorbia esula var. esula, 337 Fairybells, Hooker’s, 171 Falco columbarius, 46 sparverius, 290,303 Falsebox, 169 Fargey, P., Review by, 382 Feathermoss, 169 Felis catus, 46 concolor, 314 Fern, Bracken, 172 Maidenhair, 138 Fescue, 226 Festuca spp., 226 filiformis, 355 trachyphylla, 355 Fir, Amabilis, 172 Balsam, 50,100,236,299,433,440 Douglas, 11 Subalpine, 100 Fireweed, 171 Fisher, 62,101,229 Flax, 343 Fleabane, 137 516 Leiberg’s, 137 Flicker, Northern, 294,302 Forktail, Eastern, 417 Fox, Arctic, 36 Red, 36,51,68,263,342,468 San Joaquin Kit, 52,210 Santa Cruz Island, 210 Swift, 31,52 Foxes, Vulpes velox, in Southeastern Colorado, Multiple Scale Den Site Selection by Swift, 31 Foxglove, Middleton False, 357 Fragaria virginiana, 355 Freedman, B., 298 Frog, Boreal Chorus, 87 Columbia Spotted, 87 Green, 294 Mink, 239 Western Spotted, 87 Wood, 87 Frog, Rana septentrionalis, in Southeastern Labrador, The Mink, 239 Frostweed, Long-Branch, 359 Fucus vesiculosus, 236 Fulmar, 458 Fulmarus glacialis, 458 Gale, Sweet, 172,240 Galium aparine, 180 Gallant, D., 438 Galvez, I., 241 Gambusia affinis, 421 Gasterosteus aculeatus, 240,422 Gastromyzon sp., 424 Gaultheria ovatifolia, 171 procumbens, 355 shallon, 11,172 Gawn, J., Reviews by, 490,499 Gawn, M., Reviews by, 119,379 Gaylussacia baccata, 355 dumosa, 360 Geese, Chen caerulescens caerulescens, Exceeding the Car- rying Capacity of the Fraser River Delta’s Brackish Marshes?, Are Lesser Snow, 213 Gese, E.M., 31 Gilhen, J., 241 Glider, Wandering, 417 Globicephala melas, 97 Glycine max, 226 Gnaphalium palustre, 164 Goat, Mountain, 314 Golden-aster, Hairy, 336 Goldeneye, Common, 5,109 Goldenrod, 226 Goldeye, 422 Goldfinch, American, 294 Goldfish, 422 Goldsborough, L.G., 83 Gomphid, 350 Good King Henry, 340 Goodwin, C.E., 319 Goodwin, C.E., Review by, 379 Goose spp., 5 Canada, 109 Lesser Snow, 213 White, 214 Goosefoot, Fremont’s, 340 THE CANADIAN FIELD-NATURALIST Vol. 120 Narrow-leaved, 336 Smooth, 335 Goosefoot, Chenopodium subglabrum (Chenopodiaceae), in Canada, A Conservation Evaluation of Smooth, 335 Gossypium spp., 226 Gowans, S., 93 Grackle, Common, 294 Grass, Orchard, 154,160 Sand, 336 Switch, 226 Grayling, Arctic, 422 Grebe, 2 Horned, 109 Red-necked, 109 Greenling, Kelp, 363 Griffiths, A., 363 Grouseberry, 172 Grus canadensis, 68 Gull, 2 Bonaparte’s, 5 Ring-billed, 109 Gull, Larus delawerensis, Food Piracy on Diving Ducks, Ring-billed, 109 Gulo gulo, 111 Gum, Sweet, 226 Guppy, 422 Haemopis, 443 caeca, 450 grandis, 443 kingi, 444 lateromaculata, 443 marmorata, 443 plumbea, 447 sanguisuga, 444 septagon, 444 terrestris, 447 Haemopis lateromaculata (Hirudinea: Haemopidae): Its North America Distribution and Additional Notes on Species Description, The Leech, 443 Hagfish, Pacific, 363 Hagfish, Eptatretus stoutii, Spotted Ratfish, Hydrolagus col- liei, and Scavenger Activity on Tethered Carrion in Subtidal Benthic Communities off Western Vancou- ver Island, Pacific, 363 Hairgrass, Annual, 164 Early, 154,320 Silvery, 320 Haliaeetus leucocephalus, 109,237,477 Haliaeetus leucocephalus, Preying on Maritime Garter Snake, Thamnophis sirtalis pallidulus, on Cape Breton Is- land, Nova Scotia, Bald Eagle, 477 Hall, M.N., 57 Hamr, J., 57 Hanke, G.F., M.C.E. McNall, and J. Roberts. First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque suttonorum, and -a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia, 421 Hare, Arctic, 51,466 Snowshoe, 51,68,205 Harrier, Northern, 109 Harrington, FH. Double Marking in Arctic Wolves, Canis lupus arctos: Influence of Order on Posture, 471 Haskell, S.P. First Record of a River Otter, Lontra canadensis, Captured on the Northeastern Coast of Alaska, 235 2007 Hawk, Sharp-shinned, 46 Hawkweed, White, 170 Hazelnut, Beaked, 170 Hearn, B.J., J.T. Neville, W.J. Curran, and D.P. Snow. First Record of the Southern Red-Backed Vole, Clethri- onomys gapperi, in Newfoundland: Implications for the Endangered Newfoundland Marten, Martes amer- icana atrata, 50 Hedge-parsley, Upright, 180 Heide-Jgrgensen, M.P., 457 Helianthemum canadense, 355 Helianthus petiolaris ssp. petiolaris, 336 Helophilus fasciatus, \7 Hemlock, Coastal Western, 172 Eastern, 290 Western, 11,170,348 Herring, 96 Heterotheca villosa var. villosa, 336 Hexagrammos decagrammus, 363 Hickory, 226 Hieracium albiflorum, 170 pilosella, 355 piloselloides, 355 Hiodon alosoides, 422 Holcus lanatus, 154 Holmes, R.B., 403 Holodiscus discolor, 158,171 Horse, Wild, 314 Houston, A.E., 225,452 Houston, C.S., Reviews by, 124,382,385,496,497,498 Hoverfly, 15 Hovingh, P. The Leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its North America Distribution and Additional Notes on Species Description, 443 Huckleberry, 348,360 Red, 11,172 Hudsonia ericoides, 355 Huettmann, F., Reviews by, 125,254,491 Hutchinson, R., 413 Hwang, Y.T. and S. Lariviére. A Test of Interspecific Effects of Introduced Eastern Grey Squirrels, Sciurus caro- linensis, on Douglas’s Squirrels, Tamiasciurus dou- glasii, in Vancouver, British Columbia, 10 Hydrolagus colliei, 363 Hydrolagus colliei, and Scavenger Activity on Tethered Car- rion in Subtidal Benthic Communities off Western Vancouver Island, Pacific Hagfish, Eptatretus stoutii, Spotted Ratfish, 363 Hyena, Spotted, 52 Hylaeus elliptica, 19 Hylocomium splendens, 171 Hymenoptera, 295 Hypericum perforatum, 355 Hypochaeris radicata, 154 Hypophthalmichthys sp., 424 Hypostomus, 424 Ictalurus furcatus, 424 punctatus, 422 Ischnura verticalis, 416 Tsoetes nuttallii, 160 Jackal, 263 Jay, Blue, 294 INDEX TO VOLUME 120 517 Jennings, J.B., M.L. Kennedy, A.E. Houston, and B.D. Carv- er. Predation on Artificial Nests of Northern Bob- whites, Colinus virginianus, by Mammalian Meso- predators: Does the Problem-Individual Paradigm Fit?, 452 John, R., Reviews by, 113,114,119,123,244,251,255,38 1 386, 388,389,483,484,488 489 Johnson, G., 298 Junco hyemalis, 294 Junco, Dark-eyed, 294 Juncus balticus, 417 effusus, 355 gerardii, 417 Juneberry, 360 Jung, T.S., A.M. Runck, D.W. Nagorsen, B.G. Slough, and T. Powell. First Records of the Southern Red-backed Vole, Myodes gapperi, in the Yukon, 331 Jung, T.S., B.G. Slough, D.W. Nagorsen, T.A. Dewey, and T. Powell. First Records of the Northern Long-eared Bat, Myotis septentrionalis, in the Yukon Territory, 39 Juniper, 32 Creeping, 336 Juniperus communis var. depressa, 355 horizontalis, 336 monosperma, 32 Kallemeyn, L.W., 71 Kalmia, 240 angustifolia, 355 Kenn, S., 438 Kennedy, A., 233 Kennedy, M.L., 225,452 Kestrel, 302 American, 290,303 Kindbergia oregana, 180 Kinnikinnick, 172 Kirk, D.A., 298 Kitchen, A.M., E.M. Gese, and S.G. Lupis. Multiple Scale Den Site Selection by Swift Foxes, Vulpes velox, in Southeastern Colorado, 31 Kneteman, J., 313 Knotweed, Close-flowered, 164 Common, 164 Kogia breviceps, 97 Koi, 424 Kudzu, 226 Kuzyk, G.W., J. Kneteman, and F.K.A. Schmiegelow. Pack Size of Wolves, Canis lupus, on Caribou, Rangifer tarandus, Winter Ranges in Westcentral Alberta, 313 l’Umbre de vase, 238 l’Umbre de vase, Umbra limi, dans le nord-est du Québec, Extension de l’aire de distribution de, 238 Labrador, The Mink Frog, Rana septentrionalis, in South- eastern, 239 Lactuca canadensis var. longifolia, 355 Lagenorhynchus acutus, 95 albirostris, 95 obscurus, 97 Laidre, K.L., M.P. Heide-Jorgensen, and J.R. Orr. Reactions of Narwhals, Monodon monoceros, to Killer Whale, Orcinus orca, Attacks in the Eastern Canadian Arctic, 457 518 Lamb’s-quarters, 337 Lambdina fiscellaria fiscellaria, 50 Landriault, L.J., M.N. Hall, J. Hamr, and F.F. Mallory. Long- range Homing by an Adult Female Black Bear, Ursus americanus, 57 Larch, American, 433 Larivée, J., 289 Larivieére, S., 10 Lariviére, S., L.R. Walton, and F. Messier. Summer Move- ments and Impact of Individual Striped Skunks, Mephitis mephitis, on Duck Nests in Saskatchewan, 342 Larix laricina, 314,355,433 Larkspur, Menzies, 154 Larus delawerensis, 109 Philadelphia, 5 Larus delawerensis, Food Piracy on Diving Ducks, Ring- billed Gull, 109 Lasioglossum admirandus, 19 divergens, 19 laevissimus, 19 nymphaearum, 19 quebecensis, 19 rohweri, 19 versans, 19 zephyrus, 19 Lasionycteris noctivagans, 39 Latissimus dorsi, 406 Lauff, R., Reviews by, 118,121,243,380 Lawlor, J.L., 93 Leatherleaf, 23 Lechea intermedia, 355 intermedia var. intermedia, 361 intermedia vat. juniperina, 361 Ledum groenlandicum, 172 Leech, 443 Leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its North America Distribution and Additional Notes on Species Description, The, 443 Lejota aerea, 17 cyanea, 17 Lemming, 466 Lentil, 343 Leontodon autumnalis, 355 Lepeophtheirus salmonis, 199 Lepeophtheirus salmonis Infestation Levels on Juvenile Wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, Salmon to Arrival of Parasitized Wild Adult Pink Salmon, Response of the Sea Louse, 199 Lepidosetta bilineata, 364 Lepisosteus oculatus, 424 osseus, 423 platostomus, 424 Lepomis gibbosus, 422 macrochirus, 422 Lepus americanus, 68,205 americanus struthopus, 51 arcticus, 466 arcticus bangsii, 51 Lestes congener, 415 disjunctus, 415 forcipatus, 416 inaequalis, 417 rectangularis, 417 unguiculatus, 417 Leucanthemum vulgare, 355 THE CANADIAN FIELD-NATURALIST Vol. 120 Leucorrhinia frigida, 417 Libellula luctuosa, 417 pulchella, 417 quadrimaculata, 415 Lichen, 172 Limonius aeger, 19 confusus, 19 Linaria canadensis, 355 genistifolia ssp. dalmatica, 176 Lindell, J., Review by, 252 Linnaea borealis, 172 Lion, 52 Liquidambar styraciflua, 226 Liriodendron tulipifera, 226 Listera cordata, 170 Liu, P.S., 225 Lobelia dortmanna, 240 Lobelia, Water, 240 Lomatium ambiguum, 184 Lontra canadensis, 235 canadensis degener, 51 Lontra canadensis, Captured on the Northeastern Coast of Alaska, First Record of a River Otter, 235 Loon, 2 Looper, Hemlock, 50 Loosestrife, Purple, 307 Lota lota, 422,480 Lotus corniculatus, 157 formosissimus, 153,157 pinnatus, 153,157 Lotus formosissimus, in Canada, Conservation Evaluation of the Seaside Birds-foot Trefoil, 153 Lotus pinnatus, in Canada, Conservation Evaluation of the Bog Bird’s-foot Trefoil, 157 Louse, Sea, 199 , Louse Lepeophtheirus salmonis Infestation Levels on Juvenile Wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, Salmon to Arrival of Parasitized Wild Adult Pink Salmon, Response of the Sea, 199 Lumpsucker, 464 Lupine, Prairie, 147 Silky, 176 Lupine, Lupinus lepidus var. lepidus, in Canada, Conservation Evaluation of the Prairie, 147 Lupinus lepidus, 147 lepidus var. lepidus, 147 lepidus var. lobbii, 147 lyallii, 147 minimus, 147 sericeus, 176 Lupinus lepidus var. lepidus, in Canada, Conservation Eval- uation of the Prairie Lupine, 147 Lupis, S.G., 31 Lutra canadensis, 62 Luzula multiflora, 355 Lycaena dospassosi, 419 Lycaon pictus, 52 Lycopodium dendroideum, 355 obscurum, 357 tristachyum, 355 Lygodesmia juncea, 336 Lynx lynx susolanus, 51 pardinus, 229 rufus, 230 Lynx, Canada, 51 Iberian, 229 2007 Lysichitum americanum, 172 Lythrum salicaria, 307 MacKinnon, C. and A. Kennedy. An Observation of the Spring 2006 Migration of Black Scoter Melanitta nigra, in Northumberland Strait, Interrupted by the Confeder- ation Bridge, New Brunswick — Prince Edward Island, 233 Madtom, Tadpole, 424 Mahonia nervosa, 171 Maianthemum canadense, 355 Maine, Lady Crabs, Ovalipes ocellatus, in the Gulf of. 106 Malacosoma disstria, 433 Mallard, 6 Mallon, J.C. and R.B. Holmes. A Reevaluation of Sexual Dimorphism in the Postcranium of the Chasmosaurine Ceratopsid Chasmosaurus belli (Dinosauria: Ornithis- chia), 403 Mallory, F.F., 57 Malus pumila, 355 Manitoba, Northern Range Expansion and Invasion by the Common Carp, Cyprinus carpio, of the Churchill River System in, 83 Manitoba, Probable Black Bear, Ursus americana, Retrieval of an Elk, Cervus elaphus, Carcass from a Small Lake in Riding Mountain National Park, 110 Maple, 226 Big Leaf, 11,180 Douglas, 171 Red, 299 Sugar, 299 Vine, 11,171,348 Marczak, L.B., J.S. Richardson, and M.-C. Classen. Life History Phenology and Sediment Size Association of the Dragonfly Cordulegaster dorsalis (Odonata: Cor- dulegastridae) in an Ephemeral Habitat in South- western British Columbia, 347 Marine Turtle Newsletter, 132,258,392,502 Marten, 62 American, 100 Newfoundland, 50 Pine, 52 Marten, Martes americana atrata, First Record of the South- ern Red-Backed Vole, Clethrionomys gapperi, in Newfoundland: Implications for the Endangered New- foundland, 50 Marten, Martes americana, in Western Alberta Boreal Forests, Winter Habitat Use by American, 100 Martes americana, 62,100 americana atrata, 50 martes, 52 pennanti, 62,101,229 Martes americana atrata, First Record of the Southern Red-Backed Vole, Clethrionomys gapperi, in New- foundland: Implications for the Endangered New- foundland Marten, 50 Martes americana, in Western Alberta Boreal Forests, Win- ter Habitat Use by American Marten, 100 Mary, Blue-eyed, 180 Maunder, J.E., 239 McCarthy, J., Review by, 387 McCorquodale, D., Review by, 486 MeNall, M.C.E., 421 Meadow-foxtail, Carolina, 164 Meadowhawk, Autumn, 417 Band-winged, 417 INDEX TO VOLUME 120 519 Black, 417 Cherry-faced, 417 Saffron-winged, 417 White-faced, 417 Mech, L.D. Urine-marking and Ground-scratching by Free- ranging Arctic Wolves, Canis lupus arctos, in Sum- mer, 466 Meganyctiphanes norvegica, 96 Megaptera novaeangliae, 95 Megascops asio, 289 Megascops asio, Population in an Apple-Producing Region of Southern Quebec, First Observations of an East- ern Screech-Owl, 289 Meikle, D.B., 192 Melampyrum lineare, 357 lineare var. lineare, 170 Melanitta spp., 232 nigra, 233 Melanitta nigra, in Northumberland Strait, Interrupted by the Confederation Bridge, New Brunswick — Prince Ed- ward Island, An Observation of the Spring 2006 Migration of Black Scoter, 233 Melanitta spp., Migrations Interrupted by Confederation Bridge: An Update, Scoter, 232 Melanostoma, 19 mellinum, 17 Melanotus castanipes, 19 Melanpyrum lineare, 355 Meles meles, 52 Melic, Harford’s, 180 Melica harfordii, 180 Melilotus spp., 337 Meliscaeva cinctellam 17 Mennell, R.L., 87 Mephitis mephitis, 229,342,452 Mephitis mephitis, on Duck Nests in Saskatchewan, Summer Movements and Impact of Individual Striped Skunks, 342 Merganser, Red-breasted, 109 Mergus serrator, 109 Merlin, 46 Messier, F., 342 Metynnis sp., 424 Mice, Peromyscus leucopus: The Effects of Forest Patch Size, Edge and Surrounding Vegetation Type, Annual Variation in Habitat Use by White-footed, 192 Micropterus dolomieu, 421 salmoides, 422 Microtus sp., 52,291 pennsylvanicus, 264,294,332,436 pennsylvanicus terraenovae, 50 Milk-vetch, Timber, 176 Mimulus guttatus, 158 Mineau, P., 289 Miner’s-lettuce, 180 Mink, 51,62 Minnesota and Ontario, Lake Sturgeon, Acipenser fulvescens, Movements in Rainy Lake, 71 Minnow, Fathead, 421 Misgurnus anguillicaudatus, 424 Mitchell, S., Review by, 490 Mitrow, G., 428 Mollibdella, 444 grandis, 444 Mollugo verticillata, 355 Molothrus ater, 294 520 Monkey-flower, Yellow, 158 Monoclonius belli, 403 Monodon monoceros, 457 Monodon monoceros, to Killer Whale, Orcinus orca, Attacks in the Eastern Canadian Arctic, Reactions of Nar- whals, 457 Monopterus sp., 424 Monotropa hypopithys, 355 Montia parvifolia, 158 Montia, Small-leaved, 158 Mooreobdella, 449 Moose, 51,205,313,367 Morone saxatilis, 422 Morton, A. and R. Williams. Response of the Sea Louse Lepeophtheirus salmonis Infestation Levels on Juve- nile Wild Pink, Oncorhynchus gorbuscha, and Chum, O. keta, Salmon to Arrival of Parasitized Wild Adult Pink Salmon, 199 Mosquitofish, 421 Western, 422 Moss, Heron’s-bill, 180 Lanky, 171 Oregon Beaked, 180 Step, 171 Mouse, Deer, 50,57,332 House, 51 Meadow Jumping, 332 White-footed, 192 Mousetail, Tiny, 164 Mus musculus domesticus, 51 Muskox, 466 Muskrat, 51 Mustela ermina, 62 erminea richardsonii, 51 vison, 51,62 Myleus sp., 424 Myodes californicus, 332 gapperi, 331 glareolus, 332 rufocanus, 332 rutilus, 331 Myodes gapperi, in the Yukon, First Records of the Southern Red-backed Vole, 331 Myosotis stricta, 320 Myosurus minimus, 164 Myotis evotis, 39 keenii, 40 lucifugus, 39 lucifugus lucifugus, 51 septentrionalis, 39,51,332 volans, 39 Myotis septentrionalis, in the Yukon Territory, First Records of the Northern Long-eared Bat, 39 Myoxocephalus thompsonii, 480 Myoxocephalus thompsonii, in Northeastern Alberta, A New Record of Deepwater Sculpin, 480 Myrica gale, 172,240 pensylvanica, 355 Myxocyprinus asiaticus, 424 Nagorsen, D.W., 39,331 Narwhal, 457 Narwhals, Monodon monoceros, to Killer Whale, Orcinus orca, Attacks in the Eastern Canadian Arctic, Reac- tions of, 457 Neely, D.A., 480 THE CANADIAN FIELD-NATURALIST Vol. 120 Neoascia distincta, 17 Neohypdonus tumescens, 19 Nephelopsis, 449 Neuhold, J., 188 Neville, J.T., 50 New Brunswick, Eastern Canada, MtDNA Analyses on Hair Samples Confirm Cougar, Puma concolor, Presence in Southern, 438 New Brunswick, Effects of Timber Harvesting and Plantation Development on Cavity-nesting Birds in, 298 New Brunswick in Late Summer and Autumn, Use of Salt- marsh by Dragonflies (Odonata) in the Baie des Chaleurs Region of Quebec and, 413 New Brunswick — Prince Edward Island, An Observation of the Spring 2006 Migration of Black Scoter Melanitta nigra, in Northumberland Strait, Interrupted by the Confederation Bridge, 233 Newfoundland: Implications for the Endangered Newfound- land Marten, Martes americana atrata, First Record of the Southern Red-Backed Vole, Clethrionomys gapperi, in, 50 Newfoundland Marten, Martes americana atrata, First Record of the Southern Red-Backed Vole, Clethrionomys gapperi, in Newfoundland: Implications for the En- dangered, 50 Newt, Roughskin, 90 Nine-bark, 158 Noel, L.E., M.K. Butcher, M.A. Cronin, and B. Streever. Assessment of Effects of an Oil Pipeline on Caribou, Rangifer tarandus granti, Use of Riparian Habitats in Arctic Alaska, 2001-2003, 323 Nol, E., H. Douglas, and W.J. Crins. Responses of Syrphids, Elaterids and Bees to Single-tree Selection Harvesting in Algonquin Provincial Park, Ontario, 15 North America Distribution and Additional Notes on Species Description, The Leech Haemopis lateromaculata (Hirudinea: Haemopidae): Its, 443 Northwest Territories, Canada, Extension of Coyote, Canis latrans, Breeding Range in the, 67 Notropis hudsonius, 422 Noturus gyrinus, 424 Nova Scotia, An Analysis of the Vascular Flora of Annapolis Heathlands, 351 Nova Scotia, Bald Eagle, Haliaeetus leucocephalus, Prey- ing on Maritime Garter Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island, 477 Nova Scotia, Temporal Variability of Cetaceans near Halifax, 93 Novakowski 1925-2004, A Tribute to Nicholas Stephen, 370 Nuphar variegata, 240 Nuthatch, 290 Red-breasted, 302 White-breasted, 302 O’Neill, J., Reviews by, 120,485,487,498 Oak, 226 Garry, 150,154,157,180 Oat, 343 Oatgrass, One-spike, 164,184 Oceanspray, 171 Odocoileus sp., 68 heminous, 40,314 hemionus sitkensis, 61 virginianus, 29,40,64,68,209,3 13,367 Oldham, M.J., C.E. Goodwin, and S. Blaney. Bur Buttercup, Ranunculus testiculatus, New to Eastern Canada, 319 2007 Olson, S.L. Bald Eagle, Haliaeetus leucocephalus, Preying on Maritime Garter Snake, Thamnophis sirtalis pal- lidulus, on Cape Breton Island, Nova Scotia, 477 Omble de fontaine, 238 Oncorhynchus clarki lewisi, 421 gorbuscha, 199,421 keta, 199,421 kisutch, 421 mykiss, 90,421 mykiss aguabonita, 422 nerka, 421 tshawytscha, 421 Oncorhynchus gorbuscha, and Chum, O. keta, Salmon to Arrival of Parasitized Wild Adult Pink Salmon, Res- ponse of the Sea Louse Lepeophtheirus salmonis In- festation Levels on Juvenile Wild Pink, 199 Oncorhynchus keta, Salmon to Arrival of Parasitized Wild Adult Pink Salmon, Response of the Sea Louse Lep- eophtheirus salmonis Infestation Levels on Juvenile Wild Pink, Oncorhynchus gorbuscha, and Chum, 199 Ondatra zibethicus obscurus, 51 Onion, Nodding, 154 Slim-leaf, 160 Ontario, A New Record Size Wolf, Canis lupus, Group for, 367 Ontario, in 2001, Observations on the Nesting of the Northern Hawk Owl, Surnia ulula, near Timmins and Iroquois Falls, Northeastern, 433 Ontario, Lake Sturgeon, Acipenser fulvescens, Movements in Rainy Lake, Minnesota and, 71 Ontario, Late-born Elk, Cervus elaphus, Calf Observed Near Bancroft, 188 Ontario Natural Heritage Information Centre Science and Information Newsletter 11(1) Winter 2006, 132 Ontario, Responses of Syrphids, Elaterids and Bees to Sin- gle-tree Selection Harvesting in Algonquin Provin- cial Park, 15 Ontario, with Some Notes on Nesting Behaviour, New Nest- ing Records of the Le Conte’s Sparrow, Ammospiza leconteii, from Northeastern, 22 Onychogomphus uncatus, 350 Opossum, Virginia, 229,452 Orcinus orca, 457 Orcinus orca, Attacks in the Eastern Canadian Arctic, Reac- tions of Narwhals, Monodon monoceros, to Killer Whale, 457 Oreamnos americanus, 314 Oregon-grape, Dull, 171 Oreochromis sp., 424 Orr, J.R., 457 Orthasterias koehleri, 364 Orthocarpus barbatus, 137 Oryzopsis asperifolia, 358 Osmia, 180 atriventris, 19 Osprey, 236 Osprey, Pandion haliaetus, Depredates Common Eider, Soma- teria mollissima, Duckling, 236 Ottawa Field-Naturalists’ Club Awards for 2005, The, 504 Ottawa Field-Naturalists’ Club 10 January 2006, Minutes of the 127th Annual Business Meeting of The, 393 Otter, 51 River, 62,235 Otter, Lontra canadensis, Captured on the Northeastern Coast of Alaska, First Record of a River, 235 INDEX TO VOLUME 120 52] Ovalipes ocellatus, \06 Ovalipes ocellatus, in the Gulf of Maine, Lady Crabs, 106 Ovibos moschatus, 466 Ovis canadensis, 314 dalli, 205 Owl, Barred, 290,302, Great Horned, 290 Long-eared, 295 Northern Hawk, 433 Northern Saw-whet, 294,303 Saw-whet, 302 Owl, Surnia ulula, near Timmins and Iroquois Falls, North- eastern Ontario, in 2001, Observations on the Nesting of the Northern Hawk, 433 Owl-clover, Grand Coulee, 137 Ozotocerus bezarticus, 64 Pacu, Red-bellied, 421 Silver, 421 Pacu, Piaractus cf. P. brachypomus, in British Columbia, First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque suttonorum, and a Silver, 42] Pan troglodytes, 264 Panaque sp., 424 nigrolineatus, 424 suttonorum, 421 Panaque, Blue-eyed, 423 Royal, 424 Panaque suttonorum, and a Silver Pacu, Piaractus cf. P. brachypomus, in British Columbia, First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid Catfish, 421 Pandalus platyceros, 364 Pandion haliaetus, 236 Pandion haliaetus, Depredates Common Eider, Somateria mollissima, Duckling, Osprey, 236 Panicum virgatum, 226 Pantala spp., 418 flavescens, 416 Panthera leo, 52 Paquet, P.C., 61 Parasyrphus sp., 17 semiinterruptus, 17 Passer domesticus, 43,264 Passer domesticus, in Canada’s Maritime Provinces, Recent Declines of House Sparrows, 43 Patrikeev, M. New Nesting Records of the Le Conte’s Spar- row, Ammospiza leconteii, from Northeastern Ontario, with Some Notes on Nesting Behaviour, 22 Patrikeev, M. Observations on the Nesting of the Northern Hawk Owl, Surnia ulula, near Timmins and Iroquois Falls, Northeastern Ontario, in 2001, 433 Paxistima myrsinites, 169 Pea, 343 Pennant, Calico, 417 Penny, G., Reviews by, 117,246 Penny, J.L., 163,175,179,183 Penny, J.L. A Tribute to George Wayne Douglas 1938 — 2005, 135 Pentaceratops, 403 Perca flavescens, 422 Perch, Yellow, 422 Percopsis omiscomaycus, 422 Percymoorensis, 444 327) THE CANADIAN FIELD-NATURALIST caballeroi, 444 kingi, 444 lateromaculata, 444 marmorata, 444 terrestris, 444 Peromyscus sp., 294 leucopus, 192 maniculatus, 50,57,332 Peromyscus leucopus: The Effects of Forest Patch Size, Edge and Surrounding Vegetation Type, Annual Variation in Habitat Use by White-footed Mice, 192 Phacelia linearis, 176 Phacelia, Thread-leaved, 176 Phenacomys ungava, 332 Phoca spp., 466 hispida, 464 vitulina, 62 Phocoena phocoena, 95 Phoebe, Eastern, 294 Phragmites australis, 307 australis subspecies americanus, 307 australis subspecies australis, 307 Phragmites australis subspecies australis, in the Southern Ottawa District, Recent Invasion, Current Status and Invasion Pathway of European Common Reed, 307 Physalis alkekengi, 355 Physocarpus capitatus, 158 Piaractus sp., 424 brachypomus, 421 Piaractus brachypomus, in British Columbia, First Records of the Yellow Bullhead, Ameiurus natalis, a Loricari- id Catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf., 421 Piaractus cf. P. brachypomus, in British Columbia, First Records of the Yellow Bullhead, Ameiurus natalis, a Loricariid Catfish, Panaque suttonorum, and a Silver Pacu, 421 Picard, I., 239 Picea sp., 20,110,236,238,431 abies, 299 engelmannii, 100 glauca, 100,299,314,331,355 mariana, 22,100,240,299,3 14,33 1,355,433 rubens, 299,355,440 sitchensis, \1 Picoides arcticus, 303 pubescens, 303 villosus, 294,303 Pigeon, 57 Pike, Northern, 421,480 Pimephales promelas, 421 Pine, 290 Jack, 299,433 Loblolly, 226 Lodgepole, 100,148,170,314 New Jersey, 351 Pinyon, 32 Ponderosa, 164,176,184 Red, 351 Scots, 320 Western White, 172 White, 20 Pinus sp., 290 banksiana, 299,355,433 contorta, 100,170,314 contorta var. contorta, 148 Vol. 120 edulis, 32 monticola, 172 ponderosa, 164,176,184 resinosa, 351 strobus, 20,355 sylvestris, 298,320,355 taeda, 226 Pipiza femoralis, 17 Piranha, 421 Plagiobothrys scouleri, 158,164 Plantago lanceolata, 154 Plantain, Ribwort, 154 Plathemis lydia, 417 Platycheirus, 19 confusus, 17 obscurus, 17 Platygobio gracilis, 422 Pleco, Royal, 424 Plectritis congesta, 154,158 Pleurozium schreberi, 169,358 Poa annua, 320 bulbosa, 320 compressa, 356 pratensis, 160,356 Podiceps auritus, 109 grisegena, 109 Poecile atricapillus, 294,303 hudsonicas, 303 Poecilia reticulata, 422 Polygala sanguinea, 357 Polygonella articulata, 360 Polygonum sp., 356 aviculare, 164 polygaloides ssp. confertiflorum, 164 Polytrichum commune, 358 Juniperinum, 358 Pomoxis nigromaculatus, 422 Pondlily, Yellow, 240 Popcornflower, Scouler’s, 158,164 Poplar, 433 Balsam, 22 Tulip, 226 Populus, 428,433 balsamifera, 22 deltoides, 226 grandidentata, 356 tremuloides, 22,100,314,343,356 Porpoise, Harbour, 95 Potamogeton pectinatus, 416 Potentilla argentea, 356 canadensis, 356 pacifica, 214 recta, 356 simplex, 356 Poulin, R., Reviews by, 128,253 Powell, T., 339,331 Prairie Dog, 36 Black-tailed, 31 Primrose, Dense-spike Evening, 160 Prince Edward Island, An Observation of the Spring 2006 Migration of Black Scoter Melanitta nigra, in North- umberland Strait, Interrupted by the Confederation Bridge, New Brunswick —, 233 Procyon lotor, 57,225,290,342,452 Procyon lotor, Occurrence Through the Use of Microhabitat Variables, Predicting Raccoon, 225 2007 Pronghorn, 210 Prosartes hookeri var. oregana, \7\ Proulx, G. Winter Habitat Use by American Marten, Martes americana, in Western Alberta Boreal Forests, 100 Prunus pensylvanica, 50,356 serotina, 356 susquehanae, 360 virginiana, 356 Pseudacris maculata, 88 septentrionalis, 90 triseriata maculata, 88 triseriata septentrionalis, 88 Pseudoroegneria spicata, 176 Pseudotsuga menziesii, 11,148,158,164,169,176,180,184,348 Psilocarphus brevissimus var. brevissimus, 163 brevissimus var. multiflorus, 163 elatior, 163 tenellus var. tenellus, 163 Psilocarphus brevissimus vat. brevissimus, in Canada, Con- servation Evaluation of the Pacific Population of Dwarf Woolly-heads, 163 Psoralea, Lance-leaved, 336 Psoralidium lanceolatum, 336 Pteridium acquilinum var. latiusculum, 357 aquilinum, 172 aquilinum var. latiusculum, 356 Pueraria lobata, 226 Puma concolor, 40,438,476 concolor couguar, 438 Puma concolor, Presence in Southern New Brunswick, East- ern Canada, MtDNA Analyses on Hair Samples Con- firm Cougar, 438 Pumpkinseed, 422 Pussytoes, Stoloniferous, 183 Pussytoes, Antennaria flagellaris, in Canada, Conservation Evaluation of Stoloniferous, 183 Pycnopodia helianthoides, 364 Pygocentrus sp., 421 nattereri, 424 Pylodictis olivaris, 424 Quebec and New Brunswick in Late Summer and Autumn, Use of Saltmarsh by Dragonflies (Odonata) in the Baie des Chaleurs Region of, 413 Québec, Extension de l’aire de distribution de 17? Umbre de vase, Umbra limi, dans le nord-est du, 238 Quebec, First Observations of an Eastern Screech-Owl, Megascops asio, Population in an Apple-Producing Region of Southern, 289 Quebec, Herpetological Survey 2006 Atlas of Amphibians and Reptiles of, 258 Quercus spp., 226 garryana, 150,154,157 rubra, 356 Quillwort, Nuttall’s, 160 Quinoa, 340 Quiscalus quiscula, 294 Rabbit, Cottontail, 475 Rabbit-brush, Common, 184 Raccoon, 57,225,290,342,452 Raccoon, Procyon lotor, Occurrence Through the Use of Microhabitat Variables, Predicting, 225 Rana clamitans, 294 luteiventris, 88 INDEX TO VOLUME 120 WwW Nm w septentrionalis, 239 sylvatica, 90 Rana septentrionalis, in Southeastern Labrador, The Mink Frog, 239 Rangifer tarandus, 313 tarandus caribou, 313,367 tarandus granti, 205,323 tarandus terraenovae, 5\ Rangifer tarandus granti, Use of Riparian Habitats in Arctic Alaska, 2001-2003, Assessment of Effects of an Oil Pipeline on Caribou, 323 Rangifer tarandus, Winter Ranges in Westcentral Alberta, Pack Size of Wolves, Canis lupus, on Caribou, 313 Ranunculus falcatus, 319 testiculatus, 319 Ranunculus testiculatus, New to Eastern Canada, Bur Butter- cup, 319 Raphanus raphinistrum, 356 Rat, 478 Brown, 264 Norway, 51,264,475 Rat Poison Kills a Pack of Eastern Coyotes, Canis latrans, in an Urban Area, 478 Ratfish, Spotted, 363 Ratfish, Hydrolagus colliei, and Scavenger Activity on Teth- ered Carrion in Subtidal Benthic Communities off Western Vancouver Island, Pacific Hagfish, Eptatretus stoutii, Spotted, 363 Rattus norvegicus, 264,475 norvegicus norvegicus, 51 Raven, Common, 109,110 Redcedar, Western, 11,170,348 Redhead, 109 Reed, Common, 307 European Common, 307 Reed, Phragmites australis subspecies australis, in the South- ern Ottawa District, Recent Invasion, Current Status and Invasion Pathway of European Common, 307 Reimchen, T.E., 61,363 Rhingia nasica, 17 Rhizobium, 154,159 Rhodeus sp., 424 Rhododendron albiflorum, 169 californicum, 169 canadense, 356 lapponicum, 169 macrophyllum, 169 Rhododendron, Pacific, 169 White-flowered, 169 Rhododendron macrophyllum, in Canada, Conservation Eval- uation of Pacific Rhododendron, 169 Rhododendron, Rhododendron macrophyllum, in Canada, Conservation Evaluation of Pacific, 169 Rhytidiadelphus loreus, 171 Rice-grass, Indian, 336 Richards, N.L., P. Mineau, D.M. Bird, P. Wery, J. Larivée, and J. Duffe. First Observations of an Eastern Screech- Owl, Megascops asio, Population in an Apple-Pro- ducing Region of Southern Quebec, 289 Richardson, J.S., 347 Richardsonius balteatus, 421 Ringlet, Maritime, +19 Roberts, J., 421 Robin, American, 294 Rocksole, 364 524 Ronconi, R.A. Predicting Bird Oiling Events at Oil Sands Tailings Ponds and Assessing the Importance of Alter- nate Waterbodies for Waterfowl: a Preliminary Assess- ment, | Ronquil, Northern, 364 Ronquilus jordani, 364 Rosa carolina, 353 nutkana, 158 virginiana, 353 Rosatte, R.C. and J. Neuhold. Late-born Elk, Cervus elaphus, Calf Observed Near Bancroft, Ontario, 188 Rose, Nootka, 158 Rosebay, Lapland, 169 Rubus sp., 236 allegheniensis, 356 arenicola, 353 hispidus, 353 idaeus vat. strigosus, 18 particeps, 359 parviflorus var. parviflorus, 171 spectabilis, 11,348 vermontanus, 356 Rudbeckia hirta, 356 Rumex acetosella, 356 Runck, A.M., 331 Ruppia maritima, 414 Russian-thistle, 337 Ryan, M., 147,153 Rye, 343 Canada Wild, 336 Sagebrush, Big, 164,184 Hoary, 336 Salal, 11,172 Salamander, Long-toed, 90 Northwestern, 90 Salix sp., 158,314 exigua ssp. interior, 336 humilis, 356 Salmo gairdneri, 199 salar, 199,413,422 trutta, 199,422 trutta trutta, 199 Salmon, Atlantic, 199,413,422 Chinook, 422 Chum, 199,422 Pacific, 421 Sockeye, 422 Wild Pink, 199 Salmon, Response of the Sea Louse Lepeophtheirus salmonis Infestation Levels on Juvenile Wild Pink, Oncorhyn- chus gorbuscha, and Chum, O. keta, Salmon to Arrival of Parasitized Wild Adult Pink, 199 Salmonberry, 11,348 Salmonella, 46 Salsola tragus, 337 Salvelinus alpinus, 199,235,422 confluentus, 422 fontinalis, 238,422 malma, 422 namaycush, 422,481 Sander vitreus, 421 Sandlance, 96 Sandwort, Thread-leaved, 184 Sapin baumier, 238 THE CANADIAN FIELD-NATURALIST Vol. 120 Sapsucker, Yellow-bellied, 302 Sarotherodon sp., 424 Saskatchewan and the First Saskatchewan Records of Ame- lanchier sanguinea, Regional Variation in Amelanchi- er in the Whitewood area of Southeastern, 428 Saskatchewan Records of Amelanchier sanguinea, Regional Variation in Amelanchier in the Whitewood area of Southeastern Saskatchewan and the First, 428 Saskatchewan, Summer Movements and Impact of Individual Striped Skunks, Mephitis mephitis, on Duck Nests in, 342 Saskatoon, 176 Sayornis phoebe, 294 Scaup, 6 Greater, 109 Lesser, 5 Schaefer, R.L., 192 Schmiegelow, F.K.A., 313 Scirpus sp., 214,295,417 americanus, 213 maritimus, 214 paludosus, 215 Sciurus carolinensis, 10,264 vulgaris, 10 Sciurus carolinensis, on Douglas’s Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia, A Test of Interspecific Effects of Introduced Eastern Grey Squir- rels, 10 Scorpion-grass, Blue, 320 Scoter, 232 Black, 233 Scoter Melanitta nigra, in Northumberland Strait, Interrupt- ed by the Confederation Bridge, New Brunswick — Prince Edward Island, An Observation of the Spring 2006 Migration of Black, 233 Scoter, Melanitta spp., Migrations Interrupted by Confedera- tion Bridge: An Update, 232 Screech-Owl, Eastern, 289 Screech-Owl, Megascops asio, Population in an Apple-Pro- ducing Region of Southern Quebec, First Observa- tions of an Eastern, 289 Sculpin, Deepwater, 480 Prickly, 425 Roughback, 364 Sculpin, Myoxocephalus thompsonii, in Northeastern Alberta, A New Record of Deepwater, 480 Seal, 466 Harbour, 62 Ringed, 464 Seburn, D., Review by, 248 Sedge, Awl-fruited, 22 Few-seeded, 22 Foothill, 160 Green-sheathed, 160 Lyngbei, 213 Sedum acre, 356 spathifolium, 154 Selatosomus pulcher, 19 Seminatrix pygaea, 477 Sericomyia chrysotoxoides, 17 Serrasalmus sp., 421 Shad, American, 422 Sheep, Bighorn, 314 Dall, 205 Shepherdia canadensis, 172 2007 Shiner, Redside, 421 Spottail, 422 Shoveler, Northern, 5 Shrew, Dusky, 332 Masked, 50,332 Northern Short-tailed, 436 Pygmy, 332 Short-tailed, 294 Shrimp, Spot, 364 Silene antirhina, 356 Simard, P., J.L. Lawlor, and S. Gowans. Temporal Variability of Cetaceans near Halifax, Nova Scotia, 93 Sisyrinchium arenicola, 359 fuscatum, 357 Sitta sp., 290 canadensis, 303 carolinensis, 303 Skeleton-weed, 336 Skimmer, Four-spotted, 417 Twelve-spotted, 417 Widow, 417 Skunk, Santa Cruz Island Spotted, 210 Striped, 229,342,452 Skunks, Mephitis mephitis, on Duck Nests in Saskatchewan, Summer Movements and Impact of Individual Striped, 342 Slough, B.G., 39,331 Slough, B.G. and R.L. Mennell. Diversity and Range of Amphibians of the Yukon Territory, 87 Snake, Black Swamp, 477 Maritime Garter, 477 Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova Scotia, Bald Eagle, Haliaeetus leucocephalus, Preying on Maritime Garter, 477 Snow, D.P., 50 Solidago spp., 226 bicolor, 356 canadensis, 356 graminifolia, 356 Juncea, 356 nemoralis, 356 puberula, 356 Somateria mollissima, 233,236 Somateria mollissima, Duckling, Osprey, Pandion haliaetus, Depredates Common Eider, 236 Somatochlora elongata, 415 kennedyi, 417 Soopolallie, 172 Sorex sp., 294 cinereus, 50,332 cinereus acadicus, 51 hoyi, 332 monticolus, 332 Soybean, 226 Sparganium angustifolium, 240 Sparrow, House, 43,264 Le Conte’s, 22 Sparrow, Ammospiza leconteii, from Northeastern Ontario, with Some Notes on Nesting Behaviour, New Nest- ing Records of the Le Conte’s, 22 Sparrows, Passer domesticus, in Canada’s Maritime Provinces, Recent Declines of House, 43 Spartina, 414 alterniflora, 414 patens, 414 INDEX TO VOLUME 120 pectinata, 356,416 Speedwell, American, 158 Spring, 320 Spermophilus spp., 31 Sphaerophoria novaeangliae, \7 Sphagnum sp., 172,440 Sphegina brachygaster, \7 campanulata, \7 flavomaculata, \7 keeniana, \7 Sphyraena barracuda, 24} borealis, 241 guachancho, 241 Sphyraena barracuda from Canada, First Record of the Great Barracuda, 24] Sphyrapicus varius, 303 Spilogale gracilis amphiala, 210 Spiraea alba, 356 Spiranthes lacera, 356 Spray, Ocean, 158 Spreadwing, Elegant, 417 Lyre-tipped, 417 Northern, 417 Slender, 417 Spotted, 417 Sweetflag, 417 Spruce, 20,110,236,431 Black, 22,100,240,299,314,331,433 Englemann, 100 Norway, 299 Red, 299,440 Sitka, 11 White, 100,299,314,331 Spruce-fir, 331 Spurge, Leafy, 337 Squirrel, 295 Douglas’s, 10 Eastern Grey, 10 European Red, 10 Gray, 264 Ground, 31 Red, 10,51,57 Squirrels, Sciurus carolinensis, on Douglas’s Squirrels, Tami- asciurus douglasii, in Vancouver, British Columbia, A Test of Interspecific Effects of Introduced Eastern Grey, 10 Squirrels, Tamiasciurus douglasii, in Vancouver, British Columbia, A Test of Interspecific Effects of Intro- duced Eastern Grey Squirrels, Sciurus carolinensis, on Douglas’s, 10 Star, Long-rayed, 364 Sunflower, 364 Starling, 264 European, 109,290 Steinhilber, M. and D.A. Neely. A New Record of Deepwater Sculpin, Myoxocephalus thompsonii, in Northeastern Alberta, 480 Stellaria graminea, 356 pallida, 320 Stickleback, 421 Brook, 422 Threespine, 240,422 Stonecrop, Lance-leaved, 154 Strauss, E.G., 263,478 Strawberry, 360 526 Streever, B., 323 Strix varia, 290,303 Sturgeon, Lake, 71 White, 422 Sturgeon, Acipenser fulvescens, Movements in Rainy Lake, Minnesota and Ontario, Lake, 71 Sturnus vulgaris, 109,264,290 Stylasterias forriei, 364 Sucker, Longnose, 422,481 White, 422 Sunflower, Prairie, 336 Surnia ulula, 433 Surnia ulula, near Timmins and Iroquois Falls, Northeastern Ontario, in 2001, Observations on the Nesting of the Northern Hawk Owl, 433 Swallow, Tree, 302 Swan, Trumpeter, 220 Sweet-clover, 337 Swordtail, 422 Sylvilagus spp., 165 floridanus, 475 nuttallii, 165 Sympetrum sp., 416 costiferum, 415 danae, 413 internum, 415 obtrusum, 415 semicinctum, 417 vicinum, 415 Symphyotrichum laterifloum, 356 Synhalonia, 180 Syrphus rectus, 17 Szumylo, D.-L.M., 263 Tachycineta bicolor, 303 Tamarack, 314 Tamias striatus, 264,295 striatus lysteri, 51 Tamiasciurius sp., 295 douglasii, 10 hudsonicus, 10,57 hudsonicus ungavensis, 51 Tamiasciurus douglasii, in Vancouver, British Columbia, A Test of Interspecific Effects of Introduced Eastern Grey Squirrels, Sciurus carolinensis, on Douglas’s Squirrels, 10 Tanichthys albonubes, 424 Taraxacum officinale, 320,358 Taricha granulosa, 90 Taxidea taxus, 31,209,342 Tea-berry, Western, 171 Tea, Labrador, 172 Tegler, B., 496 Tegler, B., Review by, 252 Temnostoma balyras, 17 Tench, 422 Thamnophis sirtalis pallidulus, 477 Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova Scotia, Bald Eagle, Haliaeetus leucocephalus, Prey- ing on Maritime Garter Snake, 477 Thiel, R.P. Conditions for Sexual Interactions Between Wild Grey Wolves, Canis lupus, and Coyotes, Canis latrans, 27 Thimbleberry, 171 Thrasher, Brown, 294 THE CANADIAN FIELD-NATURALIST Vol. 120 Thuja occidentalis, 300 plicata, 11,170,348 Thymallus arcticus, 422 Tilapia sp., 422 Tilapia, 422 Tinca tinca, 422 Toad, American, 240 Western, 87 Toadflax, Dalmation, 176 Tokaryk, T., Reviews by, 127,485 Tonella tenella, 137,179 Tonella, Small-flowered, 137,179 Tonella tenella, in Canada, Conservation Evaluation of the Small-flowered Tonella, 179 Tonella, Tonella tenella, in Canada, Conservation Evaluation of the Small-flowered, 179 Torilis japonica, 180 Toxomerus geminatus, 17 Toxostoma rufum, 294 Tramea spp., 418 Trefoil, Bird’s-foot, 157 Bog Bird’s-foot, 153,157 Seaside Bird’s-foot, 153,157 Trefoil, Lotus formosissimus, in Canada, Conservation Eval- uation of the Seaside Birds-foot, 153 Trefoil, Lotus pinnatus, in Canada, Conservation Evaluation of the Bog Bird’s-foot, 157 Tremblay, E., 438 Trientalis borealis, 356 Trifolium arvense, 357 procumbens, 357 Triglochin maritimum, 214 Tringa flavipes, 5 Triteleia hyacinthina, 158 Triteleia, White, 158 Troglodytes troglodytes, 303 Trout, Brook, 422 Brown, 422 California Golden, 422 Lake, 481 Rainbow, 90,199,421 Sea, 199 Troutperch, 422 Tsuga canadensis, 290 heterophylla, 11,170,348 Turdus migratorius, 294 Tursiops truncatus, 97 Twayblade, Heart-leaved, 170 Twinflower, 172 Typha latifolia, 214 Umbra limi, 238 Umbra limi, dans le nord-est du Québec, Extension de |’ aire de distribution de 1’ Umbre de vase, 238 Urocyon littoralis, 210 Urosalpinx cinera, 106 Ursus americanus, 57,68,110,209,314 americanus hamiltoni, 51 arctos, 57,314 Ursus americanus, Retrieval of an Elk, Cervus elaphus, Car- cass from a Small Lake in Riding Mountain National Park, Manitoba, Probable Black Bear, 110 Ursus americanus L., Long-range Homing by an Adult Female Black Bear, 57 2007 Vaccinium alaskense, 172 angustifolium, 357 angustifolium X corymbosum, 357 myrtilloides, 357 ovatum, 348 parvifolium, 11,172 scoparium, 172 Vander Kloet, S.P., 351 Varden, Dolly, 422 Vasseur, L., 438 Velvet-grass, Common, 154 Vernalgrass, Sweet, 148,154,160 Veronica beccabunga ssp. americana, \58 officinalis, 357 verna, 320 Vetch, Common, 180 Viburnum nudum var. cassinoides, 357 Vicia sp., 157 cracca, 357 sativa, 180 tetrasperma, 357 Viola sagittata var. ovata f. glabrata, 360 Sagittata var. ovata, 357 Violette, V., 236 Vole, Eastern Heather, 332 Meadow, 50,264,294,332,436 Northern Red-backed, 331 Southern Red-backed, 50,331 Vole, Clethrionomys gapperi, in Newfoundland: Implications for the Endangered Newfoundland Marten, Martes americana atrata, First Record of the Southern Red- Backed, 50 Vole, Myodes gapperi, in the Yukon, First Records of the Southern Red-backed, 331 Volucella bombylans, 17 Vors, L.S. and P.L. Wilson. A New Record Size Wolf, Canis lupus, Group for Ontario, 367 Vulpes fulva, 468 macrotis, 52 macrotis mutica, 210 velox, 31 vulpes, 36,68,263,342 vulpes deletrix, 51 Vulpes velox, in Southeastern Colorado, Multiple Scale Den Site Selection by Swift Foxes, 31 Walley, W.J. Probable Black Bear, Ursus americanus, Retrieval of an Elk, Cervus elaphus, Carcass from a Small Lake in Riding Mountain National Park, Manitoba, 110 Walley, W.J., Ring-billed Gull, Larus delawerensis, Food Piracy on Diving Ducks, 109 Walleye, 421 Walton, L.R., 342 Wasp, 295 Waxwing, Cedar, 294 Way, J.G. and D.L. Eatough. Use of “Micro’-Corridors by Eastern Coyotes, Canis latrans, in a Heavily Urban- ized Area: Implications for Ecosystem Management, 474 Way, J.G., D.-L.M. Szumylo, and E.G. Strauss. An Ethogram Developed on Captive Eastern Coyotes Canis latrans, 263 Way, J.G., S.M. Cifuni, D.L. Eatough, and E.G. Strauss. Rat Poison Kills a Pack of Eastern Coyotes, Canis latrans, in an Urban Area, 478 INDEX TO VOLUME 120 527 Weasel, Short-tailed, 51,62 Weatherloach, 424 Wery, P., 289 Whale, Blue, 95 Bowhead, 462 Fin, 95 Humpback, 95 Killer, 457 Long-finned Pilot, 97 Minke, 95 North Atlantic Right, 95 Pilot, 97 Pygmy Sperm, 97 Sei, 97 Whale, Orcinus orca, Attacks in the Eastern Canadian Arctic, Reactions of Narwhals, Monodon monoceros, to Kil- ler, 457 Wheat, 343 Wheat-grass, Bluebunch, 176 Crested, 337 Northern, 336 Slender, 184 Whiteface, Frosted, 417 Whitefish, Lake, 422,480 Whitetail, Common, 417 Whitlaw, H.A., 205 Whitlow-grass, Spring, 320 Wildlife Afield, 132,502 Williams, R., 199 Willis, D.W., 71 Willow, 158,314 Sandbar, 336 Wiison, P.L., 367 Windgrass, Interrupted, 320 Wissink, R., 438 Wolf, 27,57,67,263,313,367,466,47 1,476,478 Arctic, 466,471 Eastern, 27 Eastern Timber, 51 Gray [Grey], 27,61,111,205,367 Newfoundland, 50 Red, 27 Timber, 27 Wolf, Canis lupus, Group for Ontario, A New Record Size, 367 Wolverine, 111 Wolves, Canis lupus arctos: Influence of Order on Posture, Double Marking in Arctic, 471 Wolves, Canis lupus, and Coyotes, Canis latrans, Conditions for Sexual Interactions Between Wild Grey, 27 Wolves, Canis lupus arctos, in Summer, Urine-marking and Ground-scratching by Free-ranging, 466 Wolves, Canis lupus, in Southcentral Alaska, Physical Char- acteristics, Hematology, and Serum Chemistry of Free- ranging Gray, 205 Wolves, Canis lupus, of Coastal British Columbia, Early Onto- genetic Diet in Gray, 61 Wolves, Canis lupus, on Caribou, Rangifer tarandus, Winter Ranges in Westcentral Alberta, Pack Size of, 313 Woodley, S.J., G. Johnson, B. Freedman, and D.A. Kirk. Effects of Timber Harvesting and Plantation Devel- opment on Cavity-nesting Birds in New Brunswick, 298 Woodpecker, Black-backed, 302 Downy, 302 528 Hairy, 294,302 Pileated, 299,302 Woolly-heads, Dwarf, 163 Woolly-heads, Psilocarphus brevissimus var. brevissimus, in Canada, Conservation Evaluation of the Pacific Popu- lation of Dwarf, 163 Wren, Winter, 302 Xiphophorus helleri, 422 Xylota, 20 confusa, 17 quadrimaculata, 17 Index to Book Reviews Botany Burger, W.C. Flowers How They Changed the World, 249 Carder, A. Giant Trees of Western America and the World, 120 Rover, F. and R. Dickinson. Plants of Alberta, 386 Scotter, G. and H. Flygare. Wildflowers of the Rocky Moun- tains, 386 Viereck, L. and E.L. Little. Alaska Trees and Shrubs. Second Edition, 491 Environment Dow, K. and T.E. Downing. The Atlas of Climate Change: Mapping the World’s Greatest Challenge, 493 Fish and Wildlife Historical Society. Fish, Fur and Feathers: Fish and Wildlife Conservation in Alberta, 1905-2005, 124 Gore, A. An Inconvenient Truth: The Planetary Emergency of Global Warming and What We Can Do About It, 494 Kruse, G.H., V.F. Gallucci, D.E. Hay, R.I. Perry, R.M. Peterman, T.C. Shirley, P.D. Spencer, B. Wilson, and, D. Woodby. Fisheries Assessment and Management in Data-Limited Situations, 121 McGavin, G.C. Endangered: Wildlife on the Brink of Extinc- tion, 252 Mundy, P. The Gulf of Alaska: Biology and Oceanography, 125 Redcliff, D. Lapland A Natural Historym 123 Saab, V.A. and H.D.W. Powell. Fire and Avian Ecology in North America Studies in Avian Biology Number 30, 121 Warren, A. and T. Allan. Guide to Deserts, 251 Miscellaneous Burt, W. Marshes: The Disappearing Edens, 389 Callan, K. A Paddler’s Guide to Quetico and Beyond, 496 Czajkowski, C. Wildfire in the Wilderness, 498 Elder, J. Pilgrimage to Vallombrosa: From Vermont to Italy in the Footsteps of George Perkins Marsh, 387 Hudson, R. A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s Drive of Vancouver, 128 Hudson, R. A Field Guide to Gold, Gemstones & Mineral Sites of British Columbia, Sites Within a Day’s Drive of Vancouver, 253 Koeppel, D. To See Every Bird on Earth, 498 Mabey, R. Gilbert White: A Biography of the Author of The Natural History of Selborne, 496 Mayor, A. Fossil Legends of the First Americans, 127 Milne, P. Where to Watch Birds World Cities, 388 Mock. D.W. More than Kin and Less than Kind: The Evolu- tion of Family Conflict, 497 THE CANADIAN FIELD-NATURALIST Vol. 120 Yellow-cedar, 172 Yellowlegs, Lesser, 5 Yukon, First Records of the Southern Red-backed Vole, Myo- des gapperi, in the, 331 Yukon Territory, Diversity and Range of Amphibians of the, 87 Yukon Territory, First Records of the Northern Long-eared Bat, Myotis septentrionalis, in the, 39 Zannichellia palustris, 416 Zapus hudsonius, 332 Zea mays, 226 Zenaida macroura, 294 Quammen, D. The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His Theory of Evolution, 252 Shtilmark, F. History of the Russian Zapovedniks 1985-1995, 254 Van der Flier-Keller, E. A Field Guide to the Identification of Pebbles, 255 Zoology Acorn, J. Deep Alberta. Fossil Facts and Dinosaur Digs, 485 Acorn, J. Ladybugs of Alberta: Finding the Spots and Con- necting the Dots, 486 Adler, P., D. Currie, and D.M. Wood. The Black Flies (Simu- liidae) of North America, 245 Arlott, N. Birds of Europe, Russia, China and Japan: Passer- ines, Tyrant Flycatchers to Buntings, 484 Beadle, D. and J. Rising. Tanagers, Cardinals, and Finches of the United States and Canada, 119 Bildstein, K.L. Migrating Raptors of the World: Their Ecology and Conservation, 382 Bonin, F., B. Devaux and A. Dupre. Turtles of the World, 248 Chu, M. Songbird Journeys: Four Seasons in the Lives of Migratory Birds, 385 Clements, J.F. The Clements Checklist of Birds of the World 6th Edition, 483 de Asua, M. and R. French. A New World of Animals: Early Modern Europeans on the Creatures of Iberian America, 487 del Hoyo, J., A. Elliott and D.A. Christie. Handbook of the Birds of the World Volume IT Old World Flycatchers to Old World Warbler, 381 Eisner, T., M. Eisner, M. Siegler. Secret Weapons Defenses of Insects, Spiders, Scorpions, and Other Many-Legged Creatures, 118 Elbroch, M. Animal Skulls, A Guide to North American Spe- cies, 243 Elphick, J. Atlas of Bird Migration: Tracing the Great Jour- neys of the World’s Birds, 380 Evans, H.E. and K.M. O’Neill. The Sand Wasps: Natural History and Behavior, 489 Firouz, E. The Complete Fauna of Iran, 114 Forsyth, R.G. Land Snails of British Columbia, 383 Gibson, M. Gibson’s Guide to Bird Watching & Conservation, 485 Hoogland, J. Conservation of the Black-tailed Prairie Dog: Saving North America’s Western Grasslands, 382 Jackson, D.C. Tracks, 490 Latta, S., C. Rimmer, A. Keith, J. Wiley, H. Raffaele, K. McFarland, E. Fernandez. Birds of the Dominican Republic and Haiti, 379 2007 Lynch, W. Penguins of the World, 488 Marshall, S.A. Insects: Their Natural History and Diversity: With a photographic guide to insects of eastern North America, 117 Marshall, S.A. Insects: Their Natural History and Diversity: With a Photographic Guide to Insects of Eastern North America, 246 Maybank, B. The Birding Sites of Nova Scotia, 114 Morrison, M.L. The Northern Goshawk: A technical Assess- ment of its Status, Ecology and management, 247 Moss, S. The Birder’s Companion, 379 Nagorsen, D.W. Rodents and Lagomorphs of British Col- umbia, 118 Nelson, J.S. Fishes of the World, 116 O’Connor, M. Why Don’t Woodpeckers Get Headaches? And Other Bird Questions You Know You Want To Ask, 490 INDEX TO VOLUME 120 529 Panov, E.N. Wheatears of Palaearctic: Ecology, Behaviour and Evolution of the Genus Oenanthe, 119 Proctor, N. and P. Lynch. A Field Guide to North Adantic Wildlife, 244 Sale, R. A Complete Guide to Arctic Wildlife, 244 Stevenson, T. and J. Fanshawe. The Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, Burundi, 113 Theberge, J.B. and M.T. Theberge. The Wolves of Algonquin Park: A 12-year Ecological Study, 249 Wheeler, B.K. Raptors of Western North America and Raptors of Eastern North America, 489 Young Naturalists de Vries, M. Tale of a Great White Fish, 499 530 THE CANADIAN FIELD-NATURALIST Vol. 120 Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit by post to the Editor, in either English or French, three complete manuscripts written in the journal style. Manuscripts may also be submitted (one copy) by e- mail. The research reported should be original. It is recom- mended that authors ask qualified persons to appraise the paper before it is submitted. All authors should have read and approved it. Institutional or contract approval for the publica- tion of the data must have been obtained by the authors. Also authors are expected to have complied with all pertinent leg- islation regarding the study, disturbance, or collection of ani- mals, plants or minerals. The place where voucher specimens have been deposited, and their catalogue numbers, should be given. Latitude and longitude should be included for all indi- vidual localities where collections or observations have been made. Manuscripts should be printed on standard-size paper, dou- blespaced throughout, generous margins to allow for copy marking, and each page numbered. For Articles and Notes provide a bibliographic (citation) strip, an abstract, and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. 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The Oxford English Dictionary, Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Illustrations Photographs should have a glossy finish and show sharp contrasts. Electronic versions should be high resolution. Pre- pare line drawings with India ink on good quality paper and letter (don’t type) descriptive matter. Photographic reproduc- tion of line drawings should be no larger than a standard page. Write author’s name, title of paper, and figure number on the lower left corner or on the back of each illustration. Reviewing Policy Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor (who reviews it or asks another qualified person to do so), and at least one other reviewer, who is a specialist in the field, cho- sen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—some- times extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by pay- ing $90 for each page, plus $30 for each illustration (any size up to a full page), and up to $90 per page for tables (depend- ing on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Editor. Limited joumal funds are available to help offset publi- cation charges to authors without grants or institutional sup- port. Requests for financial assistance should be made to the Editor when the manuscript is submitted. Reprints An order form for the purchase of reprints or pdf will ac- company the galley proofs sent to the authors. Invoices for publication costs will be sent when the submission is pub- lished. FRANCIS R. Cook, Editor RR 3 North Augusta, Ontario KOG IRO Canada TABLE OF CONTENTS (concluded) Volume 120 Number 4 300k Reviews ZOOLOGY: The Clements Checklist of Birds of the World 6th Edition — Birds of Europe, Russia, China and Japan: Passerines. Tyrant Flycatchers to Buntings — Deep Alberta: Fossil Facts and Dinosaur Digs — Gibson’s Guide to Bird Watching & Conservation — Ladybugs of Alberta: Finding the Spots and Connecting the Dots — A New World of Animals: Early Modern Europeans on the Creatures of Iberian America — Penguins of the World — Raptors of Western North America and Raptors of Eastern North | America — The Sand Wasps: Natural History and Behavior — Tracks — Why Don’t Woodpeckers Get Headaches? And Other Bird Questions You Want to Ask B30TANY: Alaska Trees and Shrubs Second edition INVIRONMENT: The Atlas of Climate Change: Mapping the World’s Greatest Challenge — An Inconvenient Truth: The Planetary Emergency of Global Warming and What We Can Do About It ISCELLANEOUS: Gilbert White: A Biography of the Author of The Natural History of Selborne — A Paddler’s Guide to Quetico and Beyond — More than Kin and Less than Kind: Evolution of Family Conflict | — To See Every Bird on Earth — Wildfire in the Wilderness ‘OUNG NATURALISTS: Tale of a Great White Fish arine Turtle Newsletter (114) — Canadian Association of Herpetologists/Association Canadienne des Herpetologists Bulletin Fall 2006 — Wildlife Afield 3 (1 + Supplement to 1) — Errata The Canadian-Field Naturalist 120(2) and 120(3) | ttawa Field-Naturalists’ Club Annual Awards for 2005 ‘dex to The Canadian Field-Naturalist Volume 120 Compiled by LEsLte Copy | dvice to Contributors i {ailing date of the previous issue 120(3): 30 November 2007 2006 483 49] 493 497 499 500 502 530 THE CANADIAN FIELD-NATURALIST Volume 120 Number 4 Articles A reevaluation of sexual dimorphism in the postcranium of the chasmosaurine ceratopsid Chasmosaurus belli (Dinosauria: Ornithischia) | JORDAN C. MALLON and ROBERT B. HOLMES Use of saltmarsh by dragonflies (Odonata) in the Baie des Chaleurs region of Quebec and New Brunswick in late summer and autumn PAUL M. CATLING, RAYMOND HUTCHINSON, and PAUL M. BRUNELLE First records of the Yellow Bullhead, Ameiurus natalis, a loricariid catfish, Panaque suttonorum, and a Silver Pacu, Piaractus cf P. brachypomus. in British Columbia G. F. HANKE, M. C. E. MCNALL, and J. ROBERTS Regional variation in Amelanchier in the Whitewood area of southeastern Saskatchewan and the first Saskatchewan records of Amelanchier sanguinea PAUL CATLING and G. MITROW Observations on the nesting of Northern Hawk Owl, Surnia ulula, near Timmins and Iroquois Falls, northeastern Ontario, in 2001 MICHAEL PATRIKEEV MtDNA analyses on hair samples confirm Cougar, Puma concolor, presence in southern New Brunswick, eastern Canada A.-S. BERTRAND, S. KENN, D. GALLANT, E. TREMBLAY, L. VASSEUR, and R. WISSINK The leech Haemopis lateromaculata (Hirudinea; Haemopidae): Its North American distribution and additional notes on species description PETER HOVINGH Predation on artificial nests of Northern Bobwhites, Colinus virginianus, by mammalian mesopredators: Does the problem-individual paradigm fit? J. B. JENNINGS, M. L. KENNEDY, A. E. HOUSTON, and B. D. CARVER Reactions of Narwhals, Monodon monocerus, to Killer Whale, Orcinus orca, attacks in eastern Canadian Arctic KRISTIN L. LAIDRE, MADS PETER HEIDE-JORGENSEN, and JACK R. ORR Urine-marking and ground-scratching by free-ranging Arctic Wolves, Canis lupus arctos, in summer L. DAvID MECH Notes Double marking in Arctic Wolves, Canis lupus arctos: Influence of order on posture FRED H. HARRINGTON Use of “micro”-corridors by Eastern Coyotes, Canis latrans, in a heavily urbanized area: Implications for ecosystem management JONATHAN G. Way and Davib L. EATOUGH Bald Eagle, Haliaeetus leucocephalus, preying on a Maritime Garter Snake, Thamnophis sirtalis pallidulus, on Cape Breton Island, Nova Scotia STORRS L. OLSON Rat poison kills a pack of eastern Coyotes, Canis latrans, in an urban area JONATHAN G. Way, STEPHEN M. CIFUNI, DAviD L. EATOUGH, and Eric G. STRAUSS A new record of Deepwater Sculpin, Myoxocephalus thompsonii, in northeastern Alberta M. STEINHILBER and D. A. NEELY (continued on inside back cover) ISSN 0008-3550 403 413 421 428 433 438 443 AT8h 480 2006 LAN SB a4 The CANADIAN FIELD-NATURALIST | Published by THE OTTAWA FIELD-NATURALISTS” CLUB, Ottawa, Canada | peated, PAT ' ; 1 | 1 | H Volume 121, Number 1 January—March 2007 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Michaille Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- ments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Charles D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson 2007 Council President: Mike Murphy Annie Belair Diane Kitching Dan Millar Vice-President: Ken Allison Ronald E. Bedford Karen McLachlan Hamilton Stanley Rosenbaum Recording Secretary: Susan Laurie-Bourque Fenja Brodo David Hobden Henry Steger Treasurer: Frank Pope Julia Cipriani Diane Lepage Chris Traynor Past President: Gary McNulty William J. Cody Ann Mackenzie Eleanor Zurbrigg Francis R. Cook Gillian Marston To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2; or e-mail: ofnc @ofnc.ca. For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward our mailing costs. PAP Registration Number 9477. Canada Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KO G IRO; (613) 269-3211; e-mail: cfn@ofnc.ca Copy Editor: Elizabeth Morton Honorary Business Manager: William J. Cody Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5; e-mail: r.john@rogers.com Associate Editors: Robert R. Anderson Charles D. Bird Paul M. Catling Anthony J. Erskine Donald F. McAlpine Warren B. Ballard Robert R. Campbell Brian W. Coad David Nagorsen William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail or e-mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. Subscriptions and Membership Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: January—March 2007 (December 2007). Cover: “When Blackfeet and Sioux Meet” 1908 oil painting by Charles M. Russell (1864-1926). Courtesy of Sid Richardson Museum where the original is housed. Born in St. Louis, Missouri, Charles M. Russell went west to Montana as a boy of 16 with a dream of becoming a real cowboy. Beginning in 1882, Charlie worked on various cattle drives wrangling horses where he quickly established a reputation as a likeable cowboy who loved to draw and paint. During his 11 years on the range, he witnessed the changing of the West as the days of free grass and unfenced range were ending. Charlie quit the cowboy way of life in 1893 and married Nancy Cooper three years later. With Nancy’s keen business sense and belief in her husband’s talent, Russell’s paintings began to be known nationally. Charlie Russell felt deeply the passing of the West, the most evident theme of his art. The old ways of Indian and cowboy life had disappeared. His works reflected the public’s demand for authentic pictures of the West, yet the soul of his art was romance. (Jan Scott, Director, Sid Richardson Museum, 309 Main Street, Fort Worth, Texas 76102 USA; e-mail: jan @sidricharsonmuseum.org) See article by Charles M. Kay “Were native people keystone predators: A continuous-time analysis of wildlife observations made by Lewis and Clarke in 1804-1806” pages 1-16. MCz LIBRARY JUN 19 2008 HARVARD UNIVERSITY THE CANADIAN FIELD-NATURALIST Volume 121 2007 Volume 123 The Ottawa Field-Naturalists’ Club Transactions Promoting the study and conservation of northern biodiversity since 1880 THE OTTAWA FIELD-NATURALISTS’* CLUB OTTAWA CANADA The Canadian Field-Naturalist January—March 2007 Volume 121, Number | Were Native People Keystone Predators? A Continuous-Time Analysis of Wildlife Observations Made by Lewis and Clark in 1804-1806 CHARLES E. KAY Department of Political Science, Utah State University, Logan, Utah 84322-0725 USA Kay, Charles E. 2007. Were native people keystone predators? A continuous-time analysis of wildlife observations made by Lewis and Clark in 1804-1806. Canadian Field-Naturalist 121(1): 1-16. It has long been claimed that native people were conservationists who had little or no impact on wildlife populations. More recently, though, it has been suggested that native people were keystone predators, who lacked any effective conservation strategies and instead routinely overexploited large mammal populations. To test these hypotheses, I performed a continuous- time analysis of wildlife observations made by Lewis and Clark because their journals are often cited as an example of how western North America teemed with wildlife before that area was despoiled by advancing European civilization. This included Bison, Elk, Mule Deer, Whitetailed Deer, Blacktailed Deer, Moose, Pronghorn Antelope, Bighorn Sheep, Grizzly Bears, Black Bears, and Grey Wolves. I also recorded all occasions on which Lewis and Clark met native peoples. Those data show a strong inverse relationship between native people and wildlife. The only places Lewis and Clark reported an abundance of game were in aboriginal buffer zones between tribes at war, but even there, wildlife populations were predator, not food-limited. Bison, Grizzly Bears, Bighorn Sheep, Mule Deer, and Grey Wolves were seldom seen except in aboriginal buffer zones. Moose were most susceptible to aboriginal hunting followed by Bison and then Elk, while Whitetailed Deer had amore effective escape strategy. If it had not been for aboriginal buffer zones, Lewis and Clark would have found little wildlife anywhere in the West. Moreover, prior to the 1780 smallpox and other earlier epidemics that decimated native populations in advance of European contact, there were more aboriginal people and even less wildlife. The patterns observed by Lewis and Clark are consistent with optimal foraging theory and other evolutionary ecology predictions. Key Words: Native hunting, aboriginal buffer zones, Lewis and Clark, keystone predation, reference conditions, Elk, Bison, Grizzly Bears, Mule Deer, Whitetailed Deer, western North America. It has long been postulated that native people were conservationists who had little or no impact on wildlife populations (e.g.; Speck 1913, 1939a, 1939b). Studies of modern hunter-gatherers, however, have found little evidence that native people purposefully employ con- servation strategies (Alvard 1993, 1994, 1995, 1998a, 1998b; Hill and Hurtado 1996), while archaeological data suggest that prehistoric people routinely overex- ploited large-mammal populations (Broughton 1994a, 1994b, 1997; Jones and Hilderbrant 1995: Janetski 1997; Butler 2000; Chatters 2004). Elsewhere, I have proposed that native people were keystone predators, who once structured entire ecosystems (Kay 1994, 1995, 1997a, 1997b, 1998, 2002). To test these competing hypotheses, I performed a continuous-time analysis of wildlife observations made by Lewis and Clark on their expedition across North America in 1804-1806 because their journals are often cited as an example of how the West teemed with wild- life before that area was despoiled by advancing Euro- pean civilization (Botkin 1995, 2004; Patten 1998: 70; Wilkinson and Rauber 2002; Nie 2003: 1). Lewis and Clark were the first Europeans to traverse what eventually became the western United States, and many of the native peoples they met had never before en- countered Europeans. In addition, historians universal- ly agree that Lewis and Clark’s journals are not only among the earliest, but also the most detailed and ac- curate, especially regarding natural history observations (Burroughs 1961; Ronda 1984; Botkin 1995, 2004). Thus, the descriptions left by Lewis and Clark are thought by many to represent the “pristine” state of western ecosystems (Craighead 1998: 597; Patten 1998: 70; Wilkinson and Rauber 2002; Botkin 2004). Botkin (1995: 1), for instance, described Lewis and Clark’s journey as “the greatest wilderness trip ever recorded.” Methods and Study Area Recently, Martin and Szuter (1999a, 1999b, 2002, 2004), Lyman and Wolverton (2002), and Laliberte and Ripple (2003) presented contrasting interpretations of western ecosystems based on Lewis and Clark’s jour- nals, but those analyses are flawed, in part, because they either did not separate ungulates by species or they did DD THE CANADIAN FIELD-NATURALIST not correlate wildlife sightings with the abundance of native people on each day of the entire trip. Instead, I developed three measures to quantify the wildlife ob- servations recorded by Lewis and Clark in their orig- inal journals, which have recently been re-edited and republished (Moulton 1986, 1987a, 1987b, 1988, 1990, 1991, 1993 — hereafter cited only by volume and page). First, game seen. If Lewis and Clark reported old sign of a species, that was assigned a value of one, fresh sign a two, and if they actually saw the animal, a three. This included Bison (Bison bison), Elk (Cervus elaphus), Whitetailed Deer (Odocoileus virginianus), Mule Deer (O. hemionus hemionus), Blacktailed Deer (O. h. co- lumbianus), Moose (Alces alces), Pronghorn Antelope (Antilocapra americana), Bighorn Sheep (Ovis can- adensis), Grizzly Bears (Ursus arctos), Black Bears (U. americanus), and Grey Wolves (Canis lupus). This was done each day for the entire 863 days of the expe- dition. Second, game killed. On each day, Lewis and Clark recorded the exact number of animals that were killed to provision their party. In three instances, though, Lewis and Clark reported that “some” Whitetailed Deer (day 78), Elk (day 365), or Bison (day 413) were killed. In these cases, “some” was recorded as three animals killed. In 12 instances, Lewis and Clark re- ported that “several” Whitetailed Deer (days 46, 365, 367, 373, 408, and 811), Bison (days 354, 406, 408, and 413), Mule Deer (day 404), or Blacktailed Deer (day 602) were killed. In those cases, “several” was recorded as seven animals killed. Similar to game seen, the number of animals killed was recorded for all spe- cies on all days. Third, herd size. If Lewis and Clark reported sight- ing large numbers of a particular animal, a value of ten was assigned to that species on that day. A value of ten was also assigned if Lewis and Clark reported killing 10 or more of one species on a single day. I then added game seen, game killed, and herd size values for all species on each day to obtain a daily measure of wild- life abundance. Again, this was done for all 863 days of the expedition. I also developed a similar convention to quantify the relative abundance of native people that Lewis and Clark encountered each day of their journey. If Lewis and Clark observed old sign, that was assigned a value of one, fresh sign a two, and if Lewis and Clark actu- ally saw native people, a three. If Lewis and Clark met more than ten native people on a given day that was as- signed a value of ten. On most days Lewis and Clark traveled together but on a few occasions they took sep- arate routes, most notably on the return trip. In those cases, Lewis’ observations were recorded separately from Clark’s. These conventions produced nearly 40 000 numerical data entries. To facilitate analysis, Lewis and Clark’s route was divided into 55 trip segments (Table 1), for which mean daily abundances of wildlife and mean daily abundances of native people were cal- Vol. 121 culated. It should be noted that Lewis and Clark gen- erally sent their best hunters ahead of the main party so that game would more readily be encountered. Lewis and Clark left St. Louis, Missouri on 14 May 1804 and proceeded, via watercraft, up the Missouri River through present-day Missouri, Kansas, Nebraska, Iowa, South Dakota, and into North Dakota where they built Fort Mandan in close proximity to the Mandan and Hidatsa villages. Lewis and Clark over-wintered at Fort Mandan, and then ascended the Missouri River into present-day Montana during the spring of 1805. After leaving their larger boats and portaging the Great Falls, Lewis and Clark continued up the Missouri to Three Forks before ascending the Jefferson and Beaver- head Rivers, on whose upper reaches they met the Sho- shone. After obtaining horses from the Shoshone, Lewis and Clark cached their canoes where Clark Canyon Reservoir is now situated and traveled over the Con- tinental Divide into Idaho and down the Lemhi and Salmon Rivers. From there, Lewis and Clark ascend- ed the North Fork of the Salmon and crossed Lost Trail Pass, re-entering Montana. Next, Lewis and Clark traveled down the Bitterroot Valley to Lolo Creek, which they traced to its source. Lewis and Clark then followed the high ridges north of Idaho’s Lochsa River and eventually descended to the lower Lochsa, where the explorers met the Nez Perce. At this point, Lewis and Clark left their horses and proceeded via canoe down the Clearwater, Snake, and Columbia Rivers through present-day Oregon and Washington state. Finally, Lewis and Clark built Fort Clatsop and overwintered on the south bank of the Columbia near the Pacific Ocean. During the spring of 1806, Lewis and Clark retraced their route, with minor variations, until the expedition reached present-day Lolo, Montana, where the party divided. Lewis ascended the Blackfoot River, crossed the Continental Divide, and proceeded to the Great Falls on the Missouri River, where the party split a sec- ond time. Lewis left most of his men to repair the boats cached in 1805, while he and three companions trav- eled by horseback to Cutbank Creek, where they met the Blackfeet. After the only fatal encounter with native people on the entire trip, Lewis retreated to the Mis- souri, where he rejoined the rest of his men and togeth- er they floated down that river until reunited with Clark below the Yellowstone in present-day North Dakota. Clark, on the other hand, left Lolo, Montana, and as- cended the Bitterroot River to Chief Joseph Pass, where he entered the Big Hole. From there, Clark crossed to the Beaverhead and refloated the canoes cached in 1805. Clark’s party then proceeded by land and water to Three Forks, where the group split a second time. Clark sent some of his men and the canoes down the Missouri to meet Lewis at Great Falls, while he trav- eled overland via Bozeman Pass to the Yellowstone. At this point, Clark fashioned canoes and floated down the Yellowstone and Missouri Rivers until reunited with 2007 KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? TABLE |. Trip segments and itinerary of Lewis and Clark 1804-1806. Trip segment CSmeAIDUNLWNH— Segment length (days) 30 o>) DNOMMANWAO HW Description St. Louis to Grand River, Missouri Grand River to Council Bluffs, Nebraska Council Bluffs to Big Sioux River, Nebraska Big Sioux River to above Vermillion River, Nebraska Above Vermillion River Above Vermillion River to Bad River, South Dakota Bad River to below the Cannonball River, North Dakota Cannonball River to Mandan Villages, North Dakota Mandan Villages, North Dakota Fort Mandan, North Dakota Fort Mandan to Little Missouri River, North Dakota Little Missouri River to Yellowstone River, North Dakota Yellowstone River to Milk River, Montana Milk River to Musselshell River, Montana Musselshell River to Marias River, Montana Marias River to Great Falls, Montana Great Falls portage Great Falls to Three Forks, Montana Three Forks to Big Hole River, Montana Big Hole River to meeting Shoshone in Idaho — Lewis’ account Present Clark Canyon Reservoir across divide to Lemhi River, Idaho — Lewis’ account Above Three Forks to present Clark Canyon Reservoir, Montana — Clark’s account Clark Canyon Reservoir to Lemhi River to Salmon River and return to Lemhi — Clark’s account Lemhi River, Idaho, to present Lolo, Montana Lolo, Montana, over Lolo Trail to lower Lochsa River, Idaho Lower Lochsa River — canoe camp Canoe Camp to the Dalles, Washington Cascade Mountains and portage, Washington Cascade Mountains to the mouth of Columbia River on the north bank, Washington Crossed to south bank of the Columbia River, Oregon Ft. Clatsop, Oregon Ft. Clatsop to the Cascade Mountains, Washington Cascade Mountains to the Dalles, Washington The Dalles to Camp Chopunnish, Idaho Camp Chopunnish, Idaho Attempted crossing of mountains Crossed mountains on Lolo Trail Present Lolo, Montana — here Lewis and Clark separated Lewis — up Blackfoot River and across Continental Divide, Montana Lewis — Divide to Great Falls (split party) Lewis to Cutbank Creek, Montana Lewis — Upper Cutbank Creek, met Piegan, fled back to Missouri River Lewis — Marias River down Missouri to Yellowstone River, North Dakota Lewis — Down Missouri until reunited with Clark Clark — Lolo, Montana, up Bitterroot into Big Hole Valley then to present Clark Canyon Reservoir and down to Three Forks where the party again split Clark — Three Forks to Bozeman Pass and on to the Yellowstone River, Montana Clark — Down Yellowstone River to Missouri River, North Dakota Clark — Down Missouri River until reunited with Lewis, North Dakota Little Missouri River to Mandan Villages to Ft. Mandan, North Dakota Heart River to Cannonball River, North Dakota Aricara villages, South Dakota Moreau River to White River, South Dakota White River to Vermillion River, Nebraska Big Sioux River to Council Bluffs, Nebraska Platte River to Grand River, Missouri Grand River to St. Louis, Missouri Dates 5/14-6/12/1804 6/13-8/1/1804 8/2-8/21/1804 8/22-8/25/1804 8/26-9/2/1804 9/3-9/24/1804 9/25-10/16/1804 10/17-10/23/1804 10/24-11/1/1804 1 1/2/1804-4/6/1805 4/7-4/12/1805 4/13-4/25/1805 4/26-5/7/1805 5/8-5//19/1805 5/20-6/2/1805 6/3-6/15/1805 6/16-7/14/1805 7/15-7/24/1805 7/25-8/6/1805 8/7-8/12/1805 8/13-8/26/1805 8/1-8/16/1805 8/17-8/28/1805 8/29-9/10/1805 9/11-9/19/1805 9/20-10/6/1805 10/7-10/24/1805 10/25-1 1/2/1805 11/3-11/25/1805 11/26-12/6/1805 12/7-3/22/1806 3/23-4/6/1806 4/7-4/18/1806 4/19-5/13/1806 5/14-6/9/1806 6/10-6/24/1806 6/25-6/29/1806 6/30-7/2/1806 7/3-7/7/1806 7/8-7/21/1806 7/22-1/2711806 7/28-8/7/1806 8/8-8/11/1806 7/3-7/12/1806 7/13-7/15/1806 7/16-8/2/1806 8/3-8/11/1806 8/12-8/17/1806 8/18-8/20/1806 8/2 1-8/22/1806 8/23-8/29/1806 8/30-9/3/1806 9/4-9/8/1806 9/9-9/17/1806 9/18-9/23/1806 4 THE CANADIAN FIELD-NATURALIST Lewis. Lewis and Clark then descended to St. Louis (2: 64; 3: 6; 4: 6; 5: 6, 110, 176; 6: 6, 80; 7: 6; 8: 8-9, 49, 84). Results Lewis and Clark’s observations show an inverse rela- tionship between wildlife and native people (Figure 1). Wildlife was abundant only where native people were absent, and if it had not been for the presence of abo- riginal buffer zones between tribes at war (Hickerson 1965; Steffian 1991; Martin and Szuter 1999a, 1999b, 2002, 2004; Farr 2001; Laliberte and Ripple 2003), there would have been little wildlife anywhere in the West. Yankton Sioux buffer zone As Lewis and Clark ascended the Missouri River, they met the Omahas and Ottes on day 97 and the Yankton Sioux on day 108 (Figure 2). These two groups were at war (2: 488), and wildlife was abundant only in the buffer zone between the tribes. Bison, in particu- lar, were found only in the center of the buffer zone. Sioux-Mandan buffer zone Lewis and Clark met the Teton Sioux on day 135, the Arikaras on day 148, and the Mandan-Hidatsa on day 164. Wildlife was not abundant in the area between the Teton Sioux and the Arikaras, but was abundant between the Arikaras and the Mandan-Hidatsa (Figure 3). This was because the Teton Sioux and Arikaras were allied against the Mandan-Hidatsa (3: 156, 161, 195- 196, 207, 226, 233-234, 243-244, 251, 272-273, 295- 297, 304-305; Porsche and Loendorf 1987; Bouchet- Bert 1999). That is, peace had a negative impact on wildlife populations while war had a beneficial effect, similar to conditions Hickerson (1965) reported in the upper Mississippi Valley (Farr 2001). Missouri- Yellowstone buffer zone In 1804-1806 all of Montana between the Missouri and Yellowstone Rivers was a six-sided buffer zone between warring tribes (4: 21-22, 67, 108-109, 159- 160, 216, 222, 354, 379, 401, 426, 437; 5: 8-9, 45, 68- 71, 77-80, 85, 87-91, 96-97, 102-106, 123-124, 178, 197, 259, 318; 7: 242, 250; 8: 88, 93-94, 104, 113, 123, 143, 182, 195, 278, 321, 323). The north was con- trolled by the Blackfeet Confederation, which consist- ed of five tribes (Ewers 1958), while on the west were the Flathead, Salish, Kootenay, and their allies. The Shoshone occupied the southwest (Trenholm and Car- ley 1964), the Crow the south-central, and the Sioux, Cheyenne, and their allies the southeast. To the east were the Mandan, Hidatsa, and their allies (Ahler et al. 1991). Within this large buffer zone (Martin and Szuter 1999a, 2002, 2004; Farr 2001), wildlife was rel- atively more abundant (Figures 4-7) because the war- ring factions did not hunt along the Yellowstone and Missouri as frequently as they did more secure envi- ronments closer to each tribe’s core area. As noted by Lewis and Clark, tribes did venture into the buffer zone, Vol. 121 MEAN DAILY ABUNDANCE OF ALL WILDLIFE 0 2 4 8 10 MEAN DAILY ABUNDANCE OF NATIVE PEOPLE Ficure |. Relationship between the abundance of native peo- ple and the abundance of wildlife as observed by Lewis and Clark in 1804-1806. Plotted are the mean daily abundance of all wildlife species and the mean daily abundance of native people by trip segments — seg- ments 1, 2, 54, and 55 were excluded because those areas were near European settlements. Line fitted us- ing a smoothing spline with cross validation (Math- soft 1997: 158-167). X and Y axes are offset. Note that there are no data points in the upper right as might be expected if cultural beliefs fostered conservation. Clearly, it made little difference what native people believed, or said they believed. Instead, aboriginal hunting followed predictions derived from optimal for- aging theory and other evolutionary ecology models. but only in force due to fear of attack. So the Missouri- Yellowstone buffer zone was not unhunted (4: 232), in- stead the area was just hunted less frequently (Farr 2001), which apparently was sufficient to permit great- er numbers of wildlife. Deer Lewis and Clark killed more Whitetailed Deer than all other large mammals combined. By comparison, Mule Deer were rare and were found only in tribal boundary zones, while blacktails were restricted to the Cascade Mountains west to the Pacific (6: 328, 331, 403-404). Even along the lower Columbia, though, Lewis and Clark encountered more whitetails than blacktails. This was because whitetails had a more ef- fective escape strategy than the other deer (Geist 1998; Whittaker and Lindzey 2001; Lingle 2002; Robinson et al. 2002) and thus were less affected by native hunt- ing. Even where native people were abundant, some whitetails were usually able to survive (Figure 8a) be- cause, when discovered, whitetails quickly fled into riparian thickets from which they could not easily be dislodged (5: 87; 6: 403). Lewis and Clark noted that Mule Deer and Elk when chased fled into the open (4: 136-137; 6: 403), making those species easier to hunt. Elk Lewis and Clark reported that Elk were easier to kill than Whitetailed Deer (6: 85, 242), which 1s reflected in the fact that native hunters had a greater impact on 2007 KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? 5 35 Yankton Sioux buffer zone All wildlife 30 + All wikslifen, 25 Bon! & 20 £ 2 ES 5 i E Se eiane OMes 5 Yankton Sioux e I\ / B fay / > 104 4 \ : a Bison / \ ; Y | / \ i B \ ison 5 4 yy) / \ x Y Lat | \ / A) AGEN / \ / VY) 7 N \ / Z a Ts ete 5 - a 4 o4 ig ph tel 101 102 103 104 105 111 112 113 Day FiGuRE 2. Yankton Sioux buffer zone along the Missouri River as reported by Lewis and Clark in 1804. Wildlife was abun- dant only in the zone between warring tribes. 80 All oe Sioux - Mandan buffer zone 70 All wildlife = 60 | wf °o a a As cs c oo g 3 40 - 3 8 é 30 4 =] 2 sS > ® 20-4 4 Qa Bison Teton ; wi A Sioux yee Mandan - Hidatsa Newwe ¥ ' 10 | jeer) MAGA TT PAA ABAR y WHAT V AHH HH HLF HHH HH ‘ . onffhtuvolsog oe EEEELSE CHU YE 130 135 140 145 165 170 are FiGureE 3. Sioux-Mandan buffer zone along the Missouri River as reported by Lewis and Clark in 1804. The Teton Sioux were allied with the Arikaras against the Mandan-Hidatsa and wildlife was abundant only in the zone between war- ring factions. There was little wildlife and no Bison in the area between the allied tribes. 6 THE CANADIAN FIELD-NATURALIST Vol. 121 40 5 The abundance of wildlife in the Missouri River buffer zone as reported by Lewis and Clark in 1805. “6 Direction of travel was from east to west. Ga All Pua 25 | 20 4 154 Mean daily abundance of wildlife and native people 9 11 12 13 14 16 17 18 19 20 21 Trip Segment FIGURE 4. The mean daily abundance of wildlife and native people along the Missouri River buffer zone as reported by Lewis and Clark in 1805. On their trip across this section of Montana, Lewis and Clark did not see a single native person from the time they left the Mandan (trip segments 9 and 11) until they met the Shoshone along the Montana- Idaho border (trip segment 21). Bison were observed only where native people were absent. 40 5 The abundance of wildlife in the Missouri River buffer zone as reported by Lewis in 1806. Direction of travel was west to east. Mean daily abundance of wildlife and native people 34 35 36 39 40 42 43 48 Trip segment FIGURE 5. The mean daily abundance of wildlife and native people along the Missouri River buffer zone as reported by Lewis in 1806. Trip segment 34 included tribes in central Washington, while Lewis and Clark spent trip segment 35 with the Nez Perce. Lewis and Clark then crossed the Bitterroot Mountains (trip segment 36) and separated at Lolo, | Montana. By trip segment 40, Clark was back at Great Falls on the Missouri River, which he descended to the Man- dan villages (trip segment 48). As on the upstream journey (Figure 4), Clark did not see a single native personon this section of the Missouri and wildlife was abundant only where native people were absent. Bison, in particular, occurred only in the center of the buffer zone. 2007 KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? 7 40 The abundance of wildlife in the Yellowstone River buffer zone as reported by Clark in 1806 Direction of travel was from west to east 35 s x Nn o . Mean daily abundance of wildlife and native people 8 g GY Trip Segment FIGuRE 6. The mean daily abundance of wildlife and native people along the Yellowstone River buffer zone as reported by Clark in 1806. Trip segments 34, 35, and 36 are the same as those in Figure 5. After Lewis and Clark separated, Clark returned to Three Forks by trip segment 44 and was on the Yellowstone River by trip segment 46. Although the Crow stole all of Clark’s horses on the Yellowstone, he did not actually see a single native person on his return trip across Montana until he neared the Mandan villages (trip segment 48). The only place wildlife was abundant was along the Yellowstone River and Bison were only seen in the center of that buffer zone. 35 Lewis -- Marias River buffer zone 30 = N nN oa o uo Daily abundance of wildlife and native people r= Ficure 7. Marias River buffer zone. After returning to Great Falls, Lewis ascended the Marias River and Cutbank Creek in what is now the Blackfeet Indian Reservation. As Lewis traveled from the Missouri, wildlife became less and less abundant, while Bison disappeared. Wildlife was reported on days 803 and 804 only because Lewis sent his hunters downstream 30-40 km to kill Whitetailed Deer for food. On day 804 Lewis met seven Blackfeet who told him their village was less than one-half day’s travel. Lewis camped with this small group of Blackfeet and next morning (day 805a) awoke to find the Blackfeet attempting to steal his guns and horses. An altercation followed and at least one Blackfoot was killed, the only native person killed by Lewis and Clark on their entire journey. Fearing retaliation and annihilation, Lewis fled back to the Missouri, and by his own account, traveled more than 160 km by nightfall (day 805b). Thus within one day’s hard travel, Lewis went from an area with no game and native people to an area with abundant game and no natives. Bison were found only where native people were absent (8: 112-140). 8 THE CANADIAN FIELD-NATURALIST the abundance of Elk (Figure 8b) than they did deer (Figure 8a). Lewis and Clark did kill a number of Elk at Fort Clatsop, but only because they purposefully built the fort where Elk were relatively more common and native people infrequent (6: 92-93, 95-96, 105, 108, 112). That is, Lewis and Clark constructed Fort Clat- sop in an intervillage buffer zone to take advantage of the more abundant Elk. Nevertheless, Lewis and Clark observed that most of the Elk they killed during the winter of 1805-1806 had old arrow wounds (6: 208, 210), indicative of intense native hunting. “Many of the Elk we have killed since we have been here, have been wounded with these arrows, the short piece with the barb remaining in the animal and grown up in the flesh” (6: 208). Lewis and Clark also described how native people used pit traps to kill Elk. “Then pits are emp- loyed in taking the Elk, and of course are large and deep, some of them a cube of 12 or 14 feet. These are usually placed by the side of a large fallen tree, which as well as the pit [lie] across the [trails] frequented by the Elk. [The] pits are disguised with the slender boughs of trees and moss; the unwary Elk in passing the tree precipitates himself into the pit which is sufficiently deep to prevent his escape” (6: 208). Thus, even in thick coastal forests, Elk were intensely hunted by native people. Pronghorn Antelope Native hunting had an even greater impact on the abundance of Pronghorn Antelope (Figure 8c). Despite their great speed, pronghorns were relatively easy for native people to kill (3: 176; Frison 1991). Bison Native hunting controlled the distribution and num- ber of Bison on the northern Great Plains (Figure 8d). The only places Lewis and Clark saw Bison, and espe- cially in large numbers, were in the center of aborigi- nal buffer zones between warring tribes. This is similar to what West (1995) documented on the central Great Plains — if it had not been for warring tribes and buffer zones, there would have been few Bison anywhere in North America (Kay 2002). Bighorn Sheep Native hunting had an even greater effect on Bighorn Sheep (Figure 8e). Lewis and Clark reported an abun- dance of bighorns only in the center of buffer zones far removed from native people. Grizzly Bears Native hunters also controlled the distribution and abundance of Grizzly Bears (Figure 8f). This is simi- lar to what Birkedal (1993) reported in Alaska. Aside from a few grizzlies killed in the rugged Idaho moun- tains, Lewis and Clark only observed grizzlies in abo- riginal buffer zones. Black Bears Based on Lewis and Clark’s observations and kill rates, Black Bears were less common than grizzlies, even in forested areas. Vol. 121 Moose Despite spending substantial amounts of time in what is currently prime Moose habitat, Lewis and Clark recorded Moose only once (6: 313; 7: 326; 8: 95) and that was in the center of the buffer zone between the Blackfeet and the Flathead-Salish. As explained else- where, native hunting controlled the distribution and abundance of Moose throughout western North Ameri- ca (Kay 1997b). Contrary to what is generally believed, Moose are more abundant in western North America today (Stevens 1971; Pierce and Peek 1984) than they were in Lewis and Clark’s time, or any other point in the past (Kay 1997b). Grey Wolves Lewis and Clark observed a direct relationship be- tween the abundance of game and the abundance of wolves. Wolves were common only where game was relatively abundant (4: 85). Thus, wolves were largely restricted to the same aboriginal buffer zones as were Bison, Elk, and other ungulates. Dogs and horses I also recorded the number of dogs (Canis familaris) Lewis and Clark purchased when game was in short supply, and the number of horses (Equus caballus) the explorers killed for food. Lewis and Clark killed nine horses and bought (ate) 210 dogs, primarily in the Columbia Basin, where native people were particu- larly abundant and wildlife was virtually non-existent (7: 49, 92). Lewis and Clark also bought large quan- tities of other foodstuffs from various native peoples, especially corn from the Mandan-Hidatsa and salmon from tribes throughout the Columbia Basin. Discussion Optimal-foraging theory According to optimal-foraging theory, high-ranked diet items are more susceptible to overexploitation than lower-ranked items (Smith 1983; Stephens and Krebs 1986; Smith and Winterhalder 1992; Butler 2000). The- oretical considerations and studies of modern hunter- gatherers both indicate that large mammals are the high- est-ranked diet items, and that, in general, the larger the animal, the higher its rank (Smith and Winterhalder 1992; Hill and Hurtado 1996). Moreover, if risk to the hunter or travel distances are great, only the highest- ranked diet items should be pursued (Smith and Win- terhalder 1992). Thus, optimal-foraging theory would predict that when native people entered aboriginal buf- fer zones, they should have concentrated their hunt- ing on the larger species, such as Bison and Elk, caus- ing those species to decline accordingly. This would also imply that native people lacked any effective con- servation strategy regarding these prey items. This pat- tern was, in fact, observed by Lewis and Clark as they left various native peoples and entered buffer zones, first Whitetailed Deer increased, then Elk and then Bison. Conversely, as Lewis and Clark exited a buffer zone, Bison disappeared first, followed by Elk, while 2007 = no = co MEAN DAILY ABUNDANCE OF DEER r.) 2 4 6 8 MEAN DAILY ABUNDANCE OF NATIVE PEOPLE MEAN DAILY ABUNDANCE OF PRONGHORN ANTELOPE MEAN DAILY ABUNDANCE OF BIGHORN SHEEP 0 2 4 6 8 10 MEAN DAILY ABUNDANCE OF NATIVE PEOPLE KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? Y b “x 10 ad | w | 5 | w 8 Z < | z 6 5 rs) < 4 < ra) are ; = — — 0 6 coud 0 2 4 6 8 10 MEAN DAILY ABUNDANCE OF NATIVE PEOPLE 11 d MEAN DAILY ABUNDANCE OF BISON 2 “ 6 i) 10 MEAN DAILY ABUNDANCE OF NATIVE PEOPLE wo 4 _ nN =k —t MEAN DAILY ABUNDANCE OF GRIZZLY BEARS ° 0 2 a 6 8 10 MEAN DAILY ABUNDANCE OF NATIVE PEOPLE Ficure 8. Relationship between the abundance of native people and the abundance of various wildlife species as observed by Lewis and Clark in 1804-1806. Plotted are the mean daily abundance of species and the mean daily abundance of native people by trip segments — segments 1, 2, 54, and 55 were excluded because those areas were near European settlements. Lines fitted using a smoothing spline with cross validation (Mathsoft 1997: 158-167). X and Y axes are offset. (a) Whitetailed Deer, (b) Elk, (c) Pronghorn Antelope, (d) Bison, (e) Bighorn Sheep, and (f) Grizzly Bears. some Whitetailed Deer were usually able to escape native hunters (Figure 9). Furthermore, Lewis and Clark noted that native hunters preferred to kill female ungu- lates (3: 61, 270) due to that sex’s higher fat content. Now, killing females runs counter to any conservation strategy (Kay 1994, 1997b, 1998; Kay and Simmons 2002). Alvard (1998b, 2002) recently reviewed the condi- tions under which evolution by natural selection might favor resource conservation by humans. In short, con- servation will be favored by evolution only if the resource is economical to defend. For instance, if 1000 kcal are spent defending a resource, but less than 1000 keal are derived from that resource, evolution will not favor conservation. For a variety of reasons, including competition from carnivore predators, large mammals were seldom, if ever, economical to defend (Kay 1994, 1998, 2002). Instead the logical, rational thing to do was to kill-out the large mammals as quickly as possible and then move on to other resources, 10 THE CANADIAN FIELD-NATURALIST which is exactly what aboriginal people did (Kay 1998, 2002). Counter-intuitively, once that was accomplished, native populations actually increased because people were forced to consume lower-ranked, but more abun- dant diet items (Hawkes 1991, 1992, 1993). There is also an evolved discount rate that acts to negate a wide range of possible conservation practices (Rogers 1991, 1994). Predator-limited Even within buffer zones, though, wildlife was not as abundant as one might think, because the animals were predator, not food, limited (Kay 1998, 2002). Food-limited ungulates invariably destroy berry-pro- ducing shrubs and woody riparian vegetation due to repeated browsing, and once willows (Salix spp.), cot- tonwoods (Populus spp.), and aspen (Populus tremu- loides) decline, so do associated species like Beaver (Castor canadensis) (4: 189-190), which are dependent upon those plants for food (Kay 1998 and references therein; Nietvelt 2001). Lewis and Clark, however, re- ported that riparian thickets were common in buffer zones, as were Beaver and berry-producing shrubs (e.g.; 4: 70, 145-146, 189-190, 247, 278, 332, 374, 391- 392, 399, 414, 419, 428, 435, 451; 5: 14, 42, 46, 59). In addition, Lewis and Clark noted that Whitetailed Deer often had twin fawns or triplets, and that even lactat- ing deer were fat (4: 165), which would not have been physiologically possible if ungulate populations had been food-limited. Thus, carnivore predation and occa- sional hunting by native people (4: 232) kept buffer zone ungulate populations well below what the habi- tat could otherwise support (White et al. 1998; Kay 2002). Estimate of pre-Columbian wildlife populations A number of investigators have cited Lewis and Clark’s descriptions of abundant wildlife without real- izing that those accounts only apply to the center of buffer zones (Craighead 1998; Wilkinson and Rauber 2002). Botkin (1995: 49-86; 2004: 141-147), for in- stance, used Lewis and Clark’s observations of griz- zlies along the Missouri and Yellowstone Rivers to estimate the number of bears in the western United States prior to European contact, and arrived at a fig- ure of 56 000, which others increased to 100 000 for the entire continent (e.g.; Flores 1998: 61). Although Botkin (1995: 165-169) acknowledged that native peo- ple could be important ecological factors, he failed to realize that native hunting controlled the distribution and numbers of grizzlies throughout North America (Fig- ure 8f; Birkedal 1993). During pre-Columbian times, there may have been no more than 4000-5000 grizzlies in all of North America because the bears were simply large packages of fat meat that native hunters killed at will (Hallowell 1926: 31-37; Birkedal 1993). Similar- ly, there never were 60 million Bison on the Great Plains (Seton 1929; Roe 1951), as is commonly believed (Shaw 1995; Geist 1996; Kay 2002). Vol. 121 Canadian buffer zones Aboriginal buffer zones also occurred throughout western Canada. Palliser, for instance, reported that “As a general rule the more dangerous the country [due to Indian attack] the greater the probability of finding [an] abundance of game, showing in more ways than one the truth of the old sportsman’s adage the more danger the more sport. This part of the country is so evidently the line of direction [demarcation] between the three hostile tribes, that none of them dare venture into it for hunting, except when driven to desperation by hunger, they endeavor to snatch their game from between the jaws of Scylla and Charybdis. Much there- fore as I enjoyed [this] locality for a hunting camp, see- ing buffalo on all sides, elk feeding in the distance, and fresh deer tracks in every direction... Boucharville [Palliser’s companion] did not relish it at all, and began already to calculate how soon we were to go away” (Palliser 1969: 266-267). “The abundance of game here [and not anywhere else] is accounted for by its being the neutral ground of the Crees, Assineboines, and Blackfeet; none of these tribes are in the habit of re- sorting to its neighborhood except in war parties.... We are now in the heart of the buffalo country. This region may be called a buffalo preserve, being the battle-ground between the Crees and the Blackfeet...” (Spry 1968: 146). Like Lewis and Clark, Palliser observed grizzlies primarily in aboriginal buffer zones. Alexander Henry the Younger, describing conditions on the Red River, also noted that grizzlies were found almost exclusively in aboriginal buffer zones — “they [grizzlies] are — very numerous, and seldom are molest- ed by the hunters [Indians], it being the Frontier of the Sieux [sic] and their enemy where none can hunt in safety. Here they [the bears] breed and multiply in the greatest security” (Gough 1988: 72). While Henry Hind (1971: 28-29) reported the presence of five aboriginal buffer zones on the northern Great Plains — four in Canada and one in the United States. “The following are celebrated “‘war-paths,’ where hunting is generally disallowed although game from that circumstance is usually more abundant. 1. “The War-path River’ and war road of the Lac la Pluie, Ojibways and the Sioux, from Rainy River to Red Lake River, thence across the prairies in the Valley of Red Lake River to Miniwahen or Devil’s Lake, in Dakotah [sic] Territory. 2. ‘War- path River, from the southwest corners of Lake of the Woods to Roseau River, thence to the prairies west of Red River — same tribes. 3, “War-path River’, from Lake Winnipeg to the Little Sashatchewan [sic], thence to the prairies south of Manitobah [sic] Lake — the old war-path of the Swampy Crees, the Assinaiboines and Sioux, also of the Swampy Crees and the Lake Win- nipeg Ojibways. 4. The ‘war-road’ near the Elbow of the South Branch of the Sashatchewan [River], on the flanks of the Grand Coteau, of the Blackfeet and Plain Crees. 5. The ‘war-road’ of the Sioux, Blackfeet and 2007 10 Species buffer zone effects Mean daily abundance of wildlife and native people 17 18 KAY: WERE NATIVE PEOPLE KEYSTONE PREDATORS? 1] Native 19 20 21 Trip segment FiGurE 9. The effect of native hunting on Bison, Elk, and Whitetailed Deer. As Lewis and Clark ascended the Missouri River from Great Falls (trip segment 17) and finally met the Shoshone (trip segment 21), wildlife became less and less abundant. First, Bison disappeared, and then Elk, until only a few Whitetailed Deer remained. This is the pattern that would be expected if native hunters foraged optimally without regard to conservation. Crows, in the Valley of the Yellowstone [River]’. In addition, both Alexander Mackenzie and Samuel Black noted the presence of an aboriginal buffer zone along the lower Peace River in northern British Columbia (Rich 1955: 112; Lamb 1970: 279, 288-289, 404-407). Similarly, Alexander Mackenzie reported a major In- dian-Inuit buffer zone in the far north on his 1789 trip to the Arctic Ocean (Lamb 1970: 181-233) as did Frank- lin (1969) in 1821-1822. As in the United States, most observations of wild- life in western Canada recorded by early European explorers, including virtually all sightings of Grizzly Bears, occurred in aboriginal buffer zones. Heretofore these buffer zone observations have been interpreted to mean that all of western Canada once teemed with game, which is simply not true. Instead, many areas of western Canada were almost devoid of wildlife due to intense native hunting (Kay et al. 2000). Prey behavior Lewis and Clark also reported a direct relationship between prey behavior and native hunting. In the cen- ter of buffer zones, where native people hunted only infrequently, game was relatively tame and could eas- ily be approached (e.g., 4: 67, 108). Elsewhere, how- ever, game was exceedingly wary. “The country about the mouth of this river [Little Missouri] had been recently hunted by the Minetares, and the little game which they had not killed and frightened away, was so extreemly [sic] shy that ... [our] hunters could not get in shoot [range] of them” (4: 26). “The Borders of the river [Missouri] has so much hunted by those Indi- ans ... [that] the game is scerce [sic] and veery [sic] wild” (4: 39). This also applied to Grizzly Bears and other animals. “[The bears] appear more shy here [near the Shoshone] than on the Missouri below the moun- tains” (4: 426). “These anamals [sic] [beaver] in conse- quence of not being hunted [in a buffer zone] are ex- treemly gentle, where they are hunted [though] they [the beaver] never leave their lodges in the day” (4: 100). Similarly, in 1819 Long observed that Bison fled in panic at the mere scent of humans. “The wind hap- pening to blow fresh from the south, the scent of our party was borne directly [to the Bison], and we could distinctly note every step of [our scent’s] progress through a distance of eight or ten miles, by the conster- nation and terror it excited among the buffaloes. The moment the tainted gale infected their atmosphere, [the Bison] ran with as much violence as if pursued by a party of mounted hunters” (Thwaites 1905: 255-256) —not unexpectedly, these observations were made in an aboriginal buffer zone along the Platte River in eastern Colorado (West 1995). This is identical to what Dia- mond (1984) reported in New Guinea, where even low- intensity aboriginal hunting completely altered the be- 2 THE CANADIAN FIELD-NATURALIST havior of prey species, as well as significantly reducing prey number (Kay 2002). Habitat Over the years, I have retraced most of Lewis and Clark’s route across North Dakota, Montana, Idaho, Washington, and Oregon, and there are no habitat fea- tures that could explain the distribution and abundances of the various species observed by the explorers. Lewis and Clark, for instance, did not find any “buffalo” in the large, treeless valleys of southwest Montana. This they attributed to the fact that Bison had been driven- out and/or killed-out by Shoshone hunters, not habitat characteristics (8: 182). At another point in their jour- ney, Lewis and Clark commented on how they could see no difference between the country west of the mountains and the plains along the Missouri, except that wildlife was common only on the latter. “I see very little difference between the apparent face of the country here [eastern Washington and western Idaho] and that of the plains of the Missouri only that these [the Columbia Basin grasslands] are not enlivened by the vast herds of buffaloe [sic] Elk [etc] which orna- ment the other” (7: 196). Bighorn Sheep are certainly restricted to areas with precipitous escape terrain, but Lewis and Clark found bighorns common only in the center of aboriginal buffer zones. Other suitable habi- tat was unoccupied because those areas were more frequently used by native people. Moreover, there is no evidence to support the asser- tion by Mack and Thompson (1982) or Lyman and Wolverton (2002) that Bison and other ungulates were rare in the Columbia Basin because those grasslands were nutritionally deficient. First as Daubenmire (1985) and others have noted, the Columbia Basin supported an open range livestock industry for nearly 40 years during the late 1800s (Oliphant 1968; Galbraith and Anderson 1971). Since cattle are less efficient herbi- vores than Bison, there is no physiological reason Co- lumbia Basin grasslands could have supported large numbers of free-ranging cattle and not Bison (Van Vuren 1987; Urness 1989). In addition, a modern Elk herd on Columbia grasslands not only grew at near the maximum rate of increase for that species, but pro- duced huge record-book antlers, as well — all indica- tive of excellent nutritional conditions (McCorquodale et al. 1988, 1989; McCorquodale 1991, 1993). Free- ranging Bison on other intermountain ranges have also shown high rates of increase (Van Vuren and Bray 1986; Keiter 1997; Bjornlie and Garrott 2001). There is also no evidence to support the notion that histori- cally Columbia Basin Bison populations were kept at low levels by severe winter weather, as proposed by Daubenmire (1985). After all, Bison thrive in Yellow- stone National Park (Keiter 1997; Bjornlie and Garrott 2001) and Wood Buffalo National Park (Carbyn et al. 1998) where winter climates are much more severe than in the Columbia Basin (Urness 1989). Instead, Bison and other ungulates were rare or absent from Vol. 121 most of the Columbia Basin because large runs of sal- mon, and other alternative resources, supported high numbers of native people (Hunn and French 1981), who took their preferred ungulate prey to low levels or local extinction (Kay 1994, 1998, 2002; Chatters 2004). Native populations and European diseases It has long been known that native people in the Americas had no immunological resistance to Euro- pean diseases, but only recently has it been learned that those diseases had a significant impact on native peo- ple prior to direct European contact (Dobyns 1983), or how this, in turn, caused abnormal increases in wildlife populations (Neumann 1985; Preston 1996, 1997, 2002; Kay 1998, 2002; Kay and Simmons 2002). European diseases, for instance, preceded Lewis and Clark. The smallpox epidemic of 1780 was especially devastating (Boyd 1985; Trimble 1985), and its aftermath was noted by Lewis and Clark (2: 478-482; 3: 285, 295, 311-312; 6: 81-82, 285, 308). In 1804-1806, Lewis and Clark found four Mandan villages along the mid- dle Missouri but observed that there had been 12 prior to the 1780 epidemic. Similarly, Arikaras villages were reduced from 32 to 2 (Ahler et al. 1991: 57). Thus, if Lewis and Clark had journeyed west in 1775 instead of 1804-1806, they would have met more native people and correspondingly there would have been even less wildlife (Geist 1998: 4-5; Kay 1998, 2002). Further- more, European diseases may have decimated native populations throughout western North America as early as 1550-1600 (Ramenofsky 1987; Campbell 1990; Kornfeld 1994: 198; Preston 1996, 1997, 2002), which suggests that pre-Columbian wildlife populations were likely much lower than even what Lewis and Clark experienced. Butler (2000), who studied resource de- pression in the Columbia Basin, reported that high- ranked diet items, such as ungulates, increased only after epidemic diseases decimated native populations ca. 1550. Similarly, Chatters (2004: 72-73) reported that Bison numbers increased only when native pop- ulations declined. Conclusions Contrary to prevailing paradigms (Lyman and Wol- verton 2002; Moore 2002; Wilkinson and Rauber 2002), native people controlled the distribution, abundance, and behavior of wildlife, and large mammals were com- mon only in boundary or buffer zones between warring tribes (Martin and Szuter 1999a, 1999b, 2002, 2004; Farr 2001). It is also clear that Lewis and Clark recog- nized this phenomenon, for Clark (8: 328) “observed that in the country between the [Indian] nations which are at war with each other the greatest numbers of wild animals are to be found.” This pattern can only be ex- plained if native hunters pursued an optimal-foraging strategy and did not employ any effective conservation measures (Alvard 1998b, 2002). Only twice did Lewis and Clark report high wildlife values and encounter 2007 large numbers of native people on the same day. In both cases, native hunters were killing as many animals as possible (3: 176, 253-255). Moreover, Lewis and Clark were only able to complete their journey because of the food, horses, and above all else, knowledge that they received from native people. There were no un- named streams, there were no unnamed mountains, and there was no wilderness (Kay and Simmons 2002). As noted by Lewis and Clark, the West was even more densely populated prior to the smallpox pandemic that decimated native people in 1780. These data have important implications for anthro- pology and archaeology, as well as other disciplines. Most anthropological subsistence models, for instance, incorporate the view that native people harvested un- gulates at or near sustained yield levels, yet these and other data do not support that assumption (Kay and Simmons 2002). Similarly, cultural or religious beliefs are often invoked to explain how aboriginal peoples interacted with their environment (Krech 1999), yet irrespective of what the 40 or so native groups encoun- tered by Lewis and Clark believed, or said they be- lieved, the ecological patterns were identical, at least regarding large mammals (see Figure 1). This is simi- lar to what Jerozolimski and Peres (2003) reported for modern subsistence hunters in South America, where neither ethnicity nor culture slowed the depletion of game stocks. Finally, these data support the hypothe- sis that native people were keystone predators, who once structured entire ecosystems (Kay 1998, 2002; Kay and Simmons 2002); i.e., ecologists need to aban- don the myth of once abundant wildlife and instead recognize that unhunted ungulate populations are out- side the range of historical variability. 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The way to the West: Essays on the central plains. University of New Mexico Press, Albuquerque, New Mexico, USA. White, C. A., C. E. Olmsted, and C. E. Kay. 1998. Aspen, elk, and fire in the Rocky Mountain national parks of North America. Wildlife Society Bulletin 26: 449-462. Whittaker, D. G., and F. G. Lindzey. 2001. Population char- acteristics of sympatric mule and white-tailed deer on Rocky Mountain Arsenal, Colorado. Journal of Wildlife Management 65: 946-952. Wilkinson, T., and P. Rauber. 2002. Lewis and Clark’s America: The Corps of Discovery left us a blueprint for a wild West. Sierra 87: 43-46. Received 4 July 2005 Accepted 27 June 2007 The Influence of Air Pollution on Corticolous Lichens near the Strathcona Industrial Area, Alberta M. EvsinGer!, E. BURRELL', N. DEBRUYN!, K. TANASICHUK!, and K. TIMONEY- 'Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E1 Canada * Treeline Ecological Research, 21551 Township Road 520, Sherwood Park, Alberta T8E 1E3 Canada; email: ktimoney @ compusmart.ab.ca; corresponding author Elsinger, M., E. Burrell, N. deBruyn, K. Tanasichuk, and K. Timoney. 2007. The influence of air pollution on corticolous lichens near the Strathcona Industrial Area, Alberta. Canadian Field-Naturalist 121(1): 17-23. Lichens that grow on the bark of mature trees were studied at 35 sites along an air pollution gradient east of Edmonton, Alberta. Data on species composition, richness, and cover were recorded in October 1999 in a matrix of sites that extends from a known source of pollutants (the Strathcona Industrial Area) east across Strathcona County. Air pollution is affecting the corticolous lichen community. Lichen species richness and total cover increased with distance from the pollution source. Species richness in areas distant from pollution was roughly twice that in areas near the Strathcona Industrial Area. Xantho- ria fallax and Phaeophyscia orbicularis were the most pollution tolerant lichens. Xanthoria hasseana, Ochrolechia arborea, Physcia adscendens, Parmelia sulcata, and Melanelia albertana were rare or absent near the pollution source and common in more distant areas. Most of the 15 species assessed were sensitive to air quality to some degree. Some lichens near the refineries and in Sherwood Park showed abnormal coloration and poor thallus integrity indicative of stress. We dis- cuss implications for human health. Key Words: corticolous lichens, air pollution, health, refineries, vehicle emissions, Alberta. Lichens are well-known indicators of air quality. While most lichen biomonitoring studies have been conducted in Europe and the United States, there is a growing body of such studies in Canada. In western Canada, 49 lichen species have been used to study gen- eral air pollution, while 39, 23, and 18 species have been used to monitor sulphur dioxide, photochemicals, and fluoride, respectively (Thormann 2006). Lichens have proven useful to characterize the concentrations and deposition patterns of air pollutants over large areas, particularly for metals, sulphur, and nitrogen compounds (Enns 2001*). The fungal and algal partners of the lichen symbiosis live in a delicate balance. Any pollutant that affects the well-being of either partner affects the lichen as a whole (Brodo et al. 2001). Sensitivity to air pollution is relat- ed to several features of lichens. They obtain nutri- ents only from air and precipitation; they accumulate - compounds in their thalli and have inefficient mecha- nisms of excluding or eliminating wastes, they have no | protective epidermis or cuticle, and they are quick to show symptoms of stress (Case 1981*; USGS 2005*). The concentration and constituents of air pollution vary in time and space. In Alberta, there are several important constituents and sources of air pollution (Alberta Environment 1996*; Alberta Health and Well- ness 2003). Sulphur dioxide (SO) is released from na- tural gas processing plants, oil sands facilities, power plants, gas flaring, oil refineries, pulp and paper mills, and fertilizer plants. In Alberta, 42% of SO, is emitted from natural gas processing plants (Myrick and Hunt 1998*). Hydrogen sulphide is released from fugitive emissions (leakages) at petroleum refineries, tank farms for unrefined petroleum products, natural gas and petro- chemical plants, oil sands plants, sewage treatment plants, kraft pulp and paper plants, and animal feedlots. Natural sources include sulphur hot springs and some wetlands and lakes. Nitrous oxides are produced from high temperature combustion of natural gas, coal, oil, and gasoline. Nitro- gen dioxide (NO,) is a pungent toxin. Ozone near ground level comes from reaction of nitrous oxides and volatile organic compounds in the presence of sunlight and from transport of upper atmosphere ozone down to ground level. Ozone levels are generally lower in urban and polluted areas as ozone is destroyed by nitric oxide (NO). Hydrocarbons come from a variety of in- dustrial and natural sources. The most abundant hydro- carbon in the Earth’s atmosphere is methane, a potent greenhouse gas. Reactive hydrocarbons such as alkenes, alkynes, and aromatics are toxic to humans and other animals and vegetation. They react with nitrous oxides in the presence of sunlight to form ozone. Particulates come from a variety of sources (typi- cally combustion) and can act as carriers for biologi- cally active metals and combustion products. Particu- late monitoring is based on the mass of particles filtered per m° of air; PM,, and PM, . refer to particles <10 um and <2.5 um respectively. In Alberta, the ratio of PM, .: PM, in recent years has ranged from 0.52 to 0.59 (Pol- lution Watch 2006*). Alberta is the largest emitter of air pollutants in Canada, with a 2005 release of 1.339 billion kg, 28.4% of the national total, an increase of 62% over its 2002 pollutant release (Pollution Watch 2006*). Among the criteria air contaminants that affect human and ecosys- 17 18 THE CANADIAN FIELD-NATURALIST tem health, Alberta released 412.8 million kg of sulphur dioxide and 403.9 million kg of oxides of nitrogen in 2005 (Pollution Watch 2006*). Lichens can be harmed by a variety of air pollutants, including sulphur dioxide, sulfuric and nitric acids, ozone, hydrocarbons, fluorides, and metals such as copper, lead, and zinc (Brodo et al. 2001). Sulphur dioxide impacts on lichens have been noted in Europe since the early 19" century (Simonson 1996*) and can cause decreased species richness, diversity and vitality (Case 1981*). Pollution-tolerant lichens may exhibit increased cover in polluted areas because of decreased competition with more sensitive species, such as Mel- anelia albertana (Muir and McCune 1988; Forest Health Highlights 1994*). Comparative study of the lichens in an urban or industrial area with those dis- tant from pollution can provide a good indicator of air quality. Our objective was to determine if there are spatially detectable effects on the lichen communities of mature tree bark within and downwind of pollution sources. We focused on the corticolous lichen species that grow on Balsam Poplar (Populus balsamifera) and Trembling Aspen (Populus tremuloides) trunks. Study Area The study area and the Strathcona Industrial Area (“Refinery Row”, Figure 1) are located east of Edmon- ton, Alberta. Refinery Row contains a large group of refineries and other industrial plants and is downwind of Edmonton. Sherwood Park, Fort Saskatchewan, and rural Strathcona County are downwind of Refinery Row. Major sources of air pollution from Refinery Row are industrial plants and vehicle emissions (Alberta Environment 1997*). Methods Study sites spanned an area roughly 28 km east-west and 14 km north-south downwind (east) of the primary pollution source in the Strathcona Industrial Area. Study sites were located in naturally forested areas both public and private. Two crews of two people each sam- pled a total 35 sites in mid-October 1999. Only mature trees with diameters >20 cm at breast height were sam- pled. Balsam Poplar was the preferred tree as its rough bark afforded abundant microsites for lichens. Aspen were also sampled. Prior to sampling, we learned to recognize a subset of the lichen flora. We focused on 15 taxa readily identifiable in the field. At each site, we chose the four trees with the high- est lichen cover. These trees had rough or fissured bark and were usually the largest and oldest trees present. Our data indicate average maximum cover rather than average cover. For each tree, we noted the tree species and diameter at breast height and determined the per- cent cover of each lichen species growing on the bark in the zone 0.5-1.5 m above the ground. Locations were recorded with a GPS unit. Lichen names follow Brodo Vol. 121 et al. (2001). For each site, we determined mean percent cover for each species and all combined species and the total number of species. Lichen cover and location data were analysed statistically. While air pollutants origi- nate from the entire industrial area, and from Edmon- ton in general, we defined a single pollution source point to quantify distance from source. That point was defined as the junction of Baseline Road and 34" Street on the west side of Refinery Row. Distances from each site to the defined pollution source were determined in a geographic information system. Results Of the 15 taxa of corticolous lichens assessed, six tended to be common and dominant and nine uncom- mon and non-dominant (Tables 1, 2). Lichen species richness and cover increased with distance from the pollution source (Figure 2). Relationships with dis- tance were strongest for species richness and for non- dominant species cover (Table 3). Both richness and cover appeared to rise exponentially with distance. Species richness and total cover increased gradually from west to east over the first 15-20 km of the pollu- tion gradient, then increased more rapidly (Figure 3). Minimum richness and cover were found in the north- western sites. Richness and cover continued to rise over the gradient indicating that air quality influenced the lichen community across the entire area. Some lichens near the refineries and in Sherwood Park showed ab- normal coloration (e.g., pale colors, or pinkish hues) and poor thallus integrity (partly disintegrated thalli) indicative of stress. Of the six dominant species, all but one species (Phaeophyscia orbicularis) increased in cover with dis- TABLE 1. Corticolous lichen species observed on mature Balsam Poplar and Trembling Aspen bark in the study.* Dominant Species Melanelia albertana Ochrolechia arborea Phaeophyscia orbicularis Physcia adscendens Xanthoria fallax Xanthoria hasseana Non-dominant Species Arthonia patellulata Bryoria spp. Evernia mesomorpha Flavopunctelia flaventior Hypogymnia physodes Parmelia sulcata Parmeliopsis ambigua Physcia aipolia Usnea spp. * non-dominant species occurred at <5% cover at most sites 2007 ELSINGER, BURRELL, DEBRUYN, TANASICHUK, and TIMONEY: AIR POLLUTION 19 Ficure 1. (a) Location of the 35 study sites. The area marked with an X, at the corner of Baseline Road and 34" Street (Edmonton), was the reference point for measurement of distances from the pollution sources. (b) Part of the Strathcona Industrial Area as seen from the corner of Baseline Road and Broadmoor Boulevard (west side of Sherwood Park. 9 June 2006). 20 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 2. Cover (%) of six common, dominant lichen taxa, non-dominant species cover, total cover, species richness, distance from pollution source, and location (UTM, Alberta 10TM projection) for the 35 study sites. e S aS} 3 a a eS Ss s aS 5 aS FOES SUS US) WUE Sis 5 SUS eS © 8 2 8 S 2 SiS) oS <8 © 3 5 a =3 O8 CaS) as > | 1.53 1.28 9.50 10.50 9.25 2 1.53 WS) 6.25 27.50 15.50 3 1.25 So} 6.25 30.75 13.00 4 9.50. 6.25 5.50 18.75 9.25 5) 1.03 0.33 10.50 DBS) 27.00 6 1.03 1.05 17.00 9.50. 8.50 iT 4.03 5198 Te 9.50 13.50 8 0.08 SM Si20 25 8.25 9 0.28 18.25 15.00 DAS 10 0.55 12250 20:50 14.25 11 0.30 35) 9.53 ITS 34.50 12 0.28 3.53} 4.78 17.00 11.25 13 0.55 5.28 18.28 10.50 14 0.10 12.00 11.28 18.00 15 0.10 13.75 13.25 9.50 16 0.05 9.00 18.75 15.00 17 0.08 6.25 15.03 17.50 18 0.08 15.25 0.28 12.00 19 30.00 0.08 4.75 20 0.03 16.25 0.08 9.00 21 0.30 3.50 26.25 0.03 14.50 2D) 0.03 0.30 23.75 1.55 16.25 23 0.08 38.00 4.00 eS 24 10.00 18.78 1.03 25 0.55 14.25 0.28 20.00 26 0.05 14.50 155 15.00 27 0.03 MBAS 6.75 3.53 28 0.05 9.50 0.55 1.05 29 0.55 14.28 1.03 3.50 30 10.50 0.08 16.25 31 1.03 16.25 0.08 S25) 29) 3.03 26.25 6.30 4.78 33 0.53 35.00 155 DN DS 34 31.25 3.00 26.52 35 0.05 10.75 6.25 3.25 tance from the pollution source (Figure 4). Xanthoria fallax and Phaeophyscia orbicularis were the most hardy lichens, present in virtually all sample sites. Xan- thoria hasseana and Ochrolechia arborea increased appreciably in cover with distance from the refineries. Physcia adscendens, Parmelia sulcata, and Melanelia albertana were rare or absent near the pollution source and common in more distant areas. Similarly, Flavop- unctelia flaventior, Hypogymnia physodes, Usnea sp., Evernia mesomorpha, and Arthonia patellulata were rare to absent near the refineries. Most species were sensitive to air quality to some degree. Discussion Lichens are good bioindicators of air quality in the region. There were clear trends of decreasing cover and richness with increasing proximity to Refinery Row. He Ee > = S bo Ss 2G oe 2 es = BS SS een = 23) Ss S = S33 Sh 6 Be B ‘ e 1, reduced levels of ARA and high EPA/ARA ratios decrease the inci- dence of pseudoalbinism, incomplete eye migration and hooked dorsal fin (Ness et al. 1995; McEvoy et al. 1998a; Neess and Lie 1998; Shields et al. 1999; Hamre et al. 2002). Other nutritional differences between arte- mia and copepods may contribute to pigmentation, such as the greater proportion of polar lipids (McEvoy et al. 1998b), vitamin A (or its precursors, carotenoid com- pounds) and iodine (Hamre et al. 2005) in copepods. Less attention has been paid to the development of ambicoloration. It can be voluntarily produced in the Japanese Flounder, Paralichthys olivaceus, by immers- ing larvae in water containing retinoic acid (Haga et al. 2005). Ambicoloration is an undesirable trait in hatcheries that is not normally induced voluntarily, but rather occurs spontaneously at incidence rates that depend on diet. Most diets that reduce the incidence of CHABOT AND MILLER: AMBICOLOURED ATLANTIC HALIBUT 37 pseudoalbinism also increase the incidence of hyper melanosis of the blind side (for a review, see Haga et al 2004). For instance, an excess of vitamin D, (Hasegawa et al. 1998; Haga et al. 2004) or A (Miwa and Yamano 1999) can increase the rate of ambicoloration in the Japanese Flounder. In Atlantic Halibut, copepod-based diets sometimes resulted in ambicoloration for up to 30-40% of the fish (Shields et al. 1999; Hamre et al. 2002; Hamre et al. 2005), but in other studies it did not (Ness et al. 1995; Ness and Lie 1998). This is likely the result of variability in nutritional properties of the copepods used, as the quantities and ratios of unsatu- rated fatty acids, vitamins and other nutrients changed between batches or size-classes of copepods within and between studies (e.g., Ness et al. 1995; McEvoy et al. 1998a; Shields et al. 1999; Hamre et al. 2002). Diet- induced ambicoloration is conceivable in wild halibut as well: hatching time, egg condition and environmen- tal conditions all influence when larvae begin to meta- morphose and the availability and quality of prey items. Laboratory studies have shown that the critical period during which diet influences the development of post- metamorphic pigmentation is short, making it possible for wild fish to have a suboptimal diet near the begin- ning of metamorphosis and become ambicoloured. Malpigmentation can also occur in post-metamor- phic flatfish. Many wild fingerlings of Speckled Floun- der (Paralichthys woolmani) that were normally pig- mented when captured developed white spots on their ocular side within a few days. Most of them returned to a normal pigmentation within one year, except for a few that retained some pigmentation on their blind side (Benetti 1997; Venizelos and Benetti 1999). Light reaching the blind side where it is not in contact with the bottom, and lack of sediment in which fish can bury themselves can also result in some pigmentation on the blind side of flatfish (Cunningham 1893; Ottesen and Strand 1996). However these are examples of staining, not ambicoloration. True ambicoloration appears to be in place soon after metamorphosis. It is difficult to assess whether factors leading to malpigmentation in hatchery-reared halibut would do so in the wild, or if these factors are even relevant in the wild. Halibut larvae likely encounter appropriate prey items in the wild. However, considering the high incidence of malpigmentation when diet is inappro- priate for only a few days in hatcheries, and the fact that non-diet related causes of malpigmentation in cap- tive Atlantic Halibut (light, sediment type, stocking density, etc.) are even less likely to be problematic in the wild, diet is the most likely of the known causes of malpigmentation to occur in the wild. Other possi- ble causes, such as a genetic causes, remain possible, however. True albinism (i.e., presence of red eyes) is inheritable in the Japanese Flounder (Shikano 2005), but diet appears much more important than genetic fac- tors in the development of pseudo-albinism and ambi- colouration (Seikai and Matsumoto 1994). 38 THE CANADIAN FIELD-NATURALIST Few wild ambicoloured Atlantic Halibut have been reported, but reporting rate is unknown. Reporting malpigmentation should be encouraged in scientific surveys for this species as well as others to assess incidence and mortality rate of ambicoloured fish and improve our understanding of the causes of ambi- colouration in Atlantic Halibut, and of taxonomically related incidence rates in flatfish (e.g., Diaz de Astar- loa et al. 2006). The specimen presented in this study has been pre- served and is part of the Maurice-Lamontagne Insti- tute Collection (accession number 11633). Acknowledgments We thank G. Hogan and V. Puvanendran for their helpful comments on an earlier version of the manu- script. Literature Cited Benetti, D. D. 1997. Spawning and larval husbandry of floun- der (Paralichthys woolmani) and pacific yellowtail (Seriola mazatlana), new candidate species for aquaculture. Aqua- culture 155: 307-318. Bolker, J. A., and C. R. Hill. 2000. Pigmentation development in hatchery-reared flatfishes. Journal of Fish Biology 56: 1029-1052. Brewster, B. 1987. 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Cybium, Revue Internationale d’Ichtyologie 30: 73-76. Gartner, J. V., Jr. 1986. Observations on anomalous condi- tions in some flatfishes (Pisces: Pleuronectiformes), with a new record of partial albinism. Environmental Biology of Fishes 17: 141-152. Gudger, E. W., and F. E. Firth. 1935. An almost totally ambi- colorate halibut, Hippoglossus hippoglossus, with partially rotated eye and hooked dorsal fin—The only recorded spec- imen. American Museum Novitates 811: 1-7. Gudger, E. W., and F. E. Firth. 1936. Three partially ambi- colorate four-spotted flounders, Paralichthys oblongus, two each with a hooked dorsal fin and a partially rotated eye. American Museum Novitates 885: 1-9. Gudger, E. W., and F. E. Firth. 1937. Two reversed partially ambicolorate halibuts: Hippoglossus hippoglossus. Amer- ican Museum Novitates 925: 1-10. Vol. 121 Haga, Y., T. Takeuchi, Y. Murayama, K. Ohta, and T. Fuku- naga. 2004. 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Relationship between fat soluble vitamins in diet and the occurrence of colour abnormality on the blind side of juvenile Japanese flounder. Suisanzoshoku 46: 279-286. McEvoy, L. A., A. Estevez, J. G. Bell, R. J. Shields, B. Gara, and J. R. Sargent. 1998a. Influence of dietary levels of eicosapentaenoic and arachidonic acids on the pigmenta- tion success of turbot (Scophthalmus maximus L.) and hal- ibut (Hippoglossus hippoglossus L.). Bulletin of the Aqua- culture Association of Canada 98: 17-20. McEvoy, L. A., T. Neess, J. G. Bell, and @. Lie. 1998b. Lipid and fatty acid composition of normal and malpig- mented Atlantic halibut (Hippoglossus hippoglossus) fed enriched Artemia: a comparison with fry fed wild cope- pods. Aquaculture 163: 235-248. Miwa, S., and K. Yamano. 1999. Retinoic acid stimulates development of adult type chromatophores in the flounder. Journal of Experimental Zoology 284: 317-324. Neess, T., M. Germain-Henry, and K. E. Naas. 1995. First feeding of Atlantic halibut (Hippoglossus hippoglossus) using different combinations of Artemia and wild zoo- plankton. Aquaculture 130: 235-250. Neess, T., and @. Lie. 1998. A sensitive period during first feeding for the determination of pigmentation pattern in Atlantic halibut, Hippoglossus hippoglossus L., juveniles: the role of diet. Aquaculture Research 29: 925-934. Norman, J. R. 1934. A systematic monograph of the flatfish- es (Heterosomata) Volume I Psettodidae, Bothidae, Pleu- ronectidae. British Museum, London. viii + 459 pages. Ottesen, O. H., and H. K. Strand. 1996. Growth, develop- ment, and skin abnormalities of halibut (Hippoglossus hip- poglossus L.) juveniles kept on different bottom substrates. Aquaculture 146: 17-25. Pittman, K., A. Jelmert, T. Nzess, T. Harboe, and K. Watanabe. 1998. Plasticity of viable postmetamorphic forms of farmed Atlantic halibut, Hippoglossus hippoglos- sus L. Aquaculture Research 29: 949-954. Sele, @., J. S. Solbakken, K. Watanabe, K. Hamre, D. Power, and K. Pittman. 2004. Staging of Atlantic halibut (Hippoglossus hippoglossus L.) from first feeding through metamorphosis, including cranial ossification independ- ent of eye migration. Aquaculture 239: 445-465. Scott, W. B., and M. G. Scott. 1988. Atlantic fishes of Cana- da. Canadian Bulletin of Fisheries and Aquatic Sciences 219. xxx + 731 pages. j 2007 CHABOT AND MILLER: AMBICOLOURED ATLANTIC HALIBUT 39 Seikai, T., and J. Matsumoto. 1994. Mechanism of pseudoal- binism in flatfish: an association between pigment cell and skin differentiation, Journal of the World Aquaculture Soci- ety 25: 78-85. Shields, R. J., J. G. Bell, F. S. Luizi, B. Gara, N. R. Bro- mage, and J. R. Sargent. 1999. Natural copepods are superior to enriched artemia nauplii as feed for halibut lar- vae (Hippoglossus hippoglossus) in terms of survival, pig- mentation and retinal morphology: relation to dietary essen- tial fatty acids. Journal of Nutrition 129: 1186-1194. Shikano, T. 2005. Marker-based estimation of heritability for body color variation in Japanese flounder Paralichthys olivaceus. Aquaculture 249; 95-105, Venizelos, A., and D. D. Benetti. 1999. Pigment abnormali- ties in flatfish. Aquaculture 176: 181-188. Wheeler, A. C. 1969. The fishes of the British Isles and North-West Europe. Macmillan, London, xvii + 613 pages. Received 3 August 2006 Accepted 22 February 2007 Micro-Propagation of White-top Aster, Sericocarpus rigidus, a Threatened Species from the Garry Oak Ecosystem in British Columbia BRENDA FREY, CHAIM KEMPLER, and DAvID L. EHRET! Pacific Agri-Food Research Centre, Agriculture and Agri-Food Canada, PO Box 1000, Agassiz, British Columbia VOM 1A0 Canada ‘Author to whom correspondence should be addressed; e-mail: ehretd @agr.gc.ca Frey, Brenda, Chaim Kempler, and David L. Ehret. 2007. Micro-propagation of White-top Aster, Sericocarpus rigidus, a threatened species from the Garry Oak Ecosystem in British Columbia. Canadian Field-Naturalist 121(1): 40-45. A study was undertaken to examine the feasibility of using micro-propagation techniques to regenerate White-top Aster (Seric- ocarpus rigidus), a threatened species native to the Garry Oak ecosystem in coastal British Columbia. Shoot multiplication from explants was compared in three different basal media formulations. The effects of varying [AA (indole-3-acetic acid) and kinetin on shoot production were also investigated using one of the three basal media. Shoot production was most suc- cessful on modified MS (Murashige and Skoog) media containing various concentrations of BAP (benzylaminopurine) and GA, (gibberellic acid), followed by media containing IAA and kinetin at concentrations of 2 mg l'!. Root formation occurred readily on modified MS media with IAA and reduced macronutrient and micronutrient concentrations. The study showed that S. rigidus can be successfully propagated using in vitro shoot multiplication. Key Words: Sericocarpus rigidus, Aster curtus, White-top Aster, micro-propagation, Garry Oak ecosystem, British Columbia. The Garry Oak ecosystem has a geographic distri- bution that spans the west coast of North America, from southwestern British Columbia to southern Califor- nia. In recent years, the survival of this ecosystem in Canada has been threatened by fire suppression and by residential, industrial and agricultural develop- ment. These activities have resulted in loss of species habitat, habitat degradation and the spread of inva- sive exotic species (Garry Oak Ecosystem Recovery Team 2002"). The Garry Oak ecosystem contains more plant species than any other terrestrial ecosystem in coastal British Columbia, and many plants that live in the ecosystem do not occur elsewhere in Canada. It is now estimated that less than 5% of all Garry Oak sites remain in their natural state (Garry Oak Ecosystem Recovery Team 2002’). The White-top Aster, Sericocarpus rigidus (formerly Aster curtus), is an herbaceous perennial species of Asteraceae native to low elevation grasslands in the Pacific Northwest in the United States and the associ- ated Garry Oak ecosystem on Vancouver Island, British Columbia (Clampitt 1987). Due to the serious loss of habitat, the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) designated White-top Aster as a threatened species in 1996, and the species is now protected under the Species At Risk Act (SARA) (Environment Canada 2006*). Sericocarpus rigidus pro- duces underground rhizomes, and as a result, occurs in patches which can vary tremendously in size. Shoots emerge from rhizomes in April, flower in July and Au- gust, then die back in the autumn. Seedlings are rarely, if ever, observed, making rhizomes the major form of propagation (Douglas and Illingworth 1997). Even though S. rigidus produces an abundance of seeds, via- bility is low and often uneven. In pollination studies, Clampitt (1987) found that excluding pollinators by bagging ramets significantly reduced the percentage of viable seeds (measured with tetrazolium chloride) from 20% in the unbagged treatment to 5% in the bagged. Bigger (1999) found that exposed and hand- pollinated flowers generally had the same number of viable seed (as tested with tetrazolium chloride), but differences in seed viability between hand-pollinated and exposed plants varied greatly among sites, possi- bly due to pollinator limitations in some patches. Patch size was also found to influence the degree of preda- tion by insects. Overall, seed viability varied from 0 to 83%. In a field comparison of open pollinated, between patch pollinated, within patch pollinated and pollina- tion excluded treatments, Giblin (1997) found that open and between patch pollination treatments produced sig- nificantly more filled seed (39% and 35%, respectively) than within patch and pollination exclusion treatments (18% and 9%, respectively). In terms of conditions re- quired for germination, Clampitt (1987) showed that stratification and exposure to light increased germina- tion. Ehret et al. (2004) have shown that the seeds re- quired an 8-12 week period of stratification for germi- nation and seedlings are slow to establish. Even so, Giblin and Hamilton (1999) suggest that the reproduc- tive biology of the species does not contribute to its rarity. Sericocarpus rigidus was one of three species “at risk” plants studied at the Pacific Agri-Food Research Centre (PARC) with the objective of developing prop- agation methods that would allow for the generation of plant material that could be reintroduced into pro- tected Garry Oak ecosystems, parks and ecological | AO 2007 reserves or made available to native plant nurseries. Because of the difficulties associated with seed germi- nation, micro-propagation was studied with the objec- tive of developing a method that could rapidly and reli- ably produce plantlets. Materials and Methods Plant Material Seeds were collected from two patches of S. rigidus growing at an elevation of 220 m on Mt. Tzuhalem (48°47'N, 123°38'W) on Vancouver Island in Septem- ber 2001 and from another patch at the same location in September 2002. The collection sites were 100 to 300 m apart. Given the close proximity, all plants may have been derived from the same clone. Collected seed count varied from 50 to 200 per plant. Seeds from 2001 were dried and stored at room temperature in the dark for one year. In September 2002, all seeds were placed in the dark in a cooler at 4°C. In October 2002, 15 to 30 seeds from each patch were placed on damp filter paper in petri dishes and incubated in a growth cham- ber maintained at 20°C and 14/10 h light/dark. After three weeks, only four seeds had germinated and all were collected from the same patch in 2001. These were potted in 10 cm pots containing Sunshine Mix 4, (Sun Gro Horticulture Inc., Vancouver, British Colum- bia) and were over-wintered in a greenhouse. In May 2003 the seedlings were hardened off in a cold frame for four weeks before being transplanted into a silt loam soil in irrigated raised beds at PARC. After one season of growth and flowering, seeds were collected from those plants in October 2003, and after air drying, were stored in a cooler at 4°C for 3 months. Surface sterilization of 10 seeds was conducted by suspending the seeds in a 5% solution of PPM (Plant Preservative Mixture) (Plant Cell Technology Inc., Washington D.C., USA) and deionized water (v/v) in a beaker on a shaker for 24 h. Seeds were aseptically transferred to a petri dish containing a 1.5% solution of sterile Bacto agar in deionized water (w/v). The petri dish was sealed with parafilm and placed in a plastic bag in a refriger- ator at 3°C in the dark. After nine weeks at 3°C, one seed germinated. This was the only seed that germinated during the next five months. The seedling was removed from the petri dish under sterile conditions and placed in a 350 ml jar con- taining half-strength MS (Murashige and Skoog 1962) media containing 1.0 mg L' TAA and 0.1 mg L" kinetin. The jar was placed in a growth room maintained at 22—25°C with a 16/8 h light/dark cycle using fluores- cent light with a PPFD (photosynthetic photon flux density) of ~50 uwE m? s"!. After five weeks, the fully developed, healthy seedling was transferred to a mod- ified MS medium containing half the concentration of ammonium nitrate (825 mgeL), 2.0 mgeL IAA and 2.0 mg |! kinetin. After four weeks on this medium, shoot and roots were excised and cultured on fresh media. Cultures were maintained by bimonthly trans- FREY, KEMPLER, AND EHRET: MICRO-PROPAGATION OF WHITE-TOP ASTER 4] fer to fresh media until enough shoots were generated to conduct shoot multiplication and rooting experi- ments. The excised root tissue failed to grow after being removed from the shoot and was discarded. Shoot Induction A replicated shoot multiplication experiment was initiated to determine the effects of various concen- trations of IAA plus kinetin or BAP plus GA, on the growth of explant shoots produced on modified MS media. The experimental design was a randomized complete block design consisting of seven media treat- ments with six replicate jars, each containing five ex- plants. The experiment included three modified MS basal medium (BM) formulations with each formula- tion containing a different concentration of macronu- trients, micronutrients and organic additives (Table 1). BM1 was a formulation used in earlier micropropa- gation media development using the cultivated violet species, Viola praemorsa (Ehret et al. 2004). BM2 and BM3 were modified MS media used routinely in the PARC strawberry and raspberry breeding programs for shoot proliferation. The ammonium nitrate concentra- tion of BM1 was half that of the other media because higher rates of ammonium nitrate have been found to be toxic to cultures of some plant species because the uptake of ammonium ions can acidify the medium and increase the frequency of vitrification (Slater et al. 2003). BM1 was the basal media used in media treat- ments | through 5 and it was supplemented with five combinations of IAA and kinetin as follows: 0.5 mg*L IAA + 1.0 mgeL" kinetin (M1), 0.5 mg*eL! IAA + 2.0 mg*L"! kinetin (M2), 2.0 mg*L! IAA + 1.0 mg*L'! kinetin (M3), 2.0 mg*L IAA + 2.0 mg*L" kinetin (M4) and 3.0 mgeL"! IAA +2.0 mg I! kinetin (M5). M4 was the initial shoot multiplication media used to generate plant material for the experiment. BM2 was the basal media used in treatment 6 (M6) and it was supplement- ed with 2.5 mg*L' BAP and 0.01 mg*L"! GA,. BM3 was the media used in treatment 7 (M7) and it was supplemented with 1.0 mg*L' BAP and 0.01 mgeL'! GA,,. The ferrous sulphate and Fe-EDTA supplied in BMI and BM? was replaced by Sequestrene 138 (Beck- er Underwood Inc., Ames, Iowa, USA) in BM3. The pH of all media treatments was adjusted to 5.6— 5.7 and the media were then dispensed into 350 mL jars which were covered with cellophane and autoclaved. Explants which had a mean weight of 0.164 g + 0.008 (SE), were aseptically transferred into treatment jars which were incubated in a growth room maintained at 22-25°C with a 16/8 h light/dark cycle using fluores- cent light with a PPFD of ~50 uE m” s' for four weeks. After four weeks, the number of explants that produced shoots and roots, the number of shoots produced per ex- plant, and shoot and explant fresh weights were recorded. Root Induction A separate rooting experiment was conducted using BM4 (Table 1) which is used in the PARC berry breed- 42 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Four modified Murashige and Skoog (MS) basal media (BM) formulations used for S. rigidus culture. Rates are expressed in mgel! with the exception of Plant Preservation Mixture, which is expressed in ml I". Induction Media BM1 Macronutrients Ammonium nitrate 825 Calcium chloride (anhydrous) 332 Magnesium sulphate (MgSO,-7H,O) 370 Potassium phosphate 170 Potassium nitrate 1900 Micronutrients Boric acid 6.2 Cobalt chloride 0.025 Cupric sulphate 0.025 Ferrous sulphate 27.8 Manganese sulphate DS, Potassium iodine 0.83 Sodium molybdate 0.25 Zinc sulphate 8.6 Na,EDTA SHk2 Sequestrene 138 - Sodium phosphate (NaH,PO,-H,O) - Organic Additives Sucrose 20 myo-inositol 1000 Nicotinic acid 0.5 Pyridoxine-HCL 0.5 Thiamine 0.1 Glycine 20 Adenine — Plant Preservation Mixture 2 Gerlite guar gum 2 ing program for root development. Fifteen shoot sam- ples initiated on M4 were placed on the rooting medium containing 1.0 mgeL"! IAA. After four weeks, rooted plants were transferred to Sunshine Mix 4 potting mix and moved to a greenhouse where they were acclima- tized. Statistical Analysis Data were analysed using the general linear model procedure in SAS (SAS Institute, Cary, North Carolina, USA), for one-way analysis of variance (ANOVA). All independent factors were fixed. Where ANOVA results were significant, treatment means were separated by Duncan’s multiple range tests. Data expressed in per- centages were transformed by arcsin transformations before statistical analysis. Results and Discussion In general, explants grew rapidly in culture, with extensive shoot multiplication (Figure 1 a, b). After four weeks of culture, explant survival ranged from 83.3 to 100 percent with no significant survival differ- ences among media treatments (Table 2). The percent- age of surviving explants that produced shoots ranged from 61 to 100%. Just over 60% of M1 and M3 ex- plants produced shoots compared to 90-100% of ex- Shoot Root BM2 BM3 BM4 1650 1237 464 332 B52) 332 370 370 370 170 170 64 1900 1425 534 6.2 6.2 3.1 0.025 0.025 0.013 0.025 0.025 0.013 27.8 = A DOS 3} Lil 0.83 0.83 0.42 0.25 0.25 0.13 8.6 8.6 4.3 BD) — = = 200 100 170 170 85 30 30 30 100. 100 100 0.8 0.8 0.4 80 80 — D 2; 2 2 2 D plants cultured on M4—M7. In addition, explants cul- tured on M1, M2 or M3 produced roots while being cultured on shoot multiplication media (data not pre- sented). M1 and M3 which had the lowest concentra- tions of kinetin produced the greatest number of roots, 13 and 20 % respectively. These results suggest that aster media should contain kinetin at minimal rates of 2.0 mgeL to avoid root initiation during the shoot multiplication stage. This supports previous recom- mendations that kinetin be added to a medium for cell division, shoot multiplication and axillary bud pro- liferation while IAA should be added to promote cell enlargement, root initiation and adventitious bud for- mation (Kyte and Klein 1999). Mean explant plus shoot weights ranged from 0.241 to 0.855 grams. No statistically significant differences were observed among BMI treatments. M6 produced a significantly greater biomass than all other media, followed by M7 which had a significantly greater bio- mass than M1 to M3 (Table 2, Figure 1c). Explants and shoots cultured on M6 appeared slightly chlorot- ic compared to those cultured on M7 which were the darkest green of all treatments (data not presented). Explants cultured on M2 and M3 were the least vigor- ous; explant and shoot leaves appeared mottled and there 2007 FREY, KEMPLER, AND EHRET: MICRO-PROPAGATION OF WHITE-TOP ASTER 43 Treatment 7 bis) a — 2 — — cw — he Ficure 1. A. Seriocarpus rigidus explant on culture media after two weeks in culture. B. Explants and developed shoots on media after 4 weeks. C. Representative sam- ples of explants and shoots cultured on seven media treatments after 4 weeks of culture. Treatment labels 1 through 7 represent treatments M1 through M7, respectively. D. Four-month-old rooted aster plants in the greenhouse. ly fewer shoots than M4. Treatments M6 and M7, both of which contained BAP and GA, and which were on BM2 and BM3, respectively, produced a greater num- ber of shoots per explant than most BM1 treatments containing combinations of [AA and kinetin. The mean weight of individual shoots cultured on the different media ranged from 0.037 to 0.097 grams per shoot. Few consistent differences were observed were several dead leaves at the base of the explants. among the five BMI treatments. Compared to those The number of shoots per surviving explant ranged _ treatments, M6 (on BM2 with BAP and GA,) pro- from 1.72 to 6.19 shoots per explant (Table 2). Within duced shoots with the greatest weight. Despite some the BMI treatments, M2 and M3 produced significant- differences among BMI treatments, the results show 44 THE CANADIAN FIELD-NATURALIST Vol. 121 that the ratio and the concentration of IAA and kinetin in BM] did not have a consistent effect on overall shoot multiplication. Shoots were small and difficult to handle after four weeks of culture. Extending the culture time from four to five weeks would likely have increased shoots to a size that would have made them easier to transfer and a more desirable size for rooting. Root initiation was 100% on the BM4 medium (data not shown). Roots appeared on the shoots after 7 days of culture, and after 4 weeks a healthy root system had fully developed. The rooted plants were easily acclima- tized to a greenhouse environment and showed vigor- ous growth (Figure 1d). Other methods to induce root development were not attempted since this technique was so highly successful. Micro-propagation methods for commercially im- portant species of Asteraceae such as sunflower (Hel- ianthus annuus) employ a variety of techniques to re- generate plants (Alibert et al. 1994). There are also methods to micro-propagate other Asteraceae species of commercial value, such as feverfew (Tanacetum parthenium) (Simmons 1998), marigold (TJagetes erec- ta) (Vanegas et al. 2002), Echinacea pallida (Koroch et al. 2003) and safflower (Carthamus tinctorius) (Orli- kowska and Dyer 1993). These techniques have in- creasingly been used on medicinally important Aster- aceae species such as goldenrod (Solidago virgaurea, S. canadensis, S. gigantea and S. graminifolia) (Kelem- ba and Thiem 2004) and those which are rare, such as Saussurea obvallata (Joshi and Dhar 2003). In Canada, micro-propagation of the Gulf of St. Lawrence Aster, a rare species in Prince Edward Island, has been shown to have potential in conservation efforts (Stewart and Lacroix 2001). Meristem culture has been extensively used for the clonal propagation of horticultural plants. Since the constituent cells of meristems are genetically stable, plants regenerated by in vitro culture of shoot apices without the callus-mediated process of organogenesis should result in the recovery of genetically identical progeny. Because the culture was started from a single seed and mass regenerated in tissue culture, genetic var- iability is limited. Sericocarpus rigidus propagates clonally in the wild by underground rhizomes, so in vitro mass clonal propagation could be a good strategy for preservation and reintroduction of this threatened species. Our study is the first to show that the threatened Asteraceae species, S. rigidus, can also be successfully propagated using micro-propagation techniques. Only one germinated seed was available for use in the study. However, by its very nature, micro-propagation works at the level of single seeds or explants, so our situa- tion is not atypical. The genetic variability of the Mt. Tzuhalem population of S. rigidus is not known, but since the plant occurs in clonal patches, it is unlikely that other seeds of S. rigidus would respond different- ly to micro-propagation. The best results for shoot mul- Shoot weight (g/shoot) 0.056 + .009 bed 0.056 + .007 bed 0.040 + .005 cd 0.037 + .004 d 0.070 + .013 be 0.097 + .009 a 0.079 + .005 ab Number of shoots per surviving explant 2.68 + 0.29 cd 1.94+0.13 d 1.72+0.14d 4.13 + 0.50 be 2.97 + 0.58 cd 6.19+0.96 a 4.63 + 0.47 ab oh oK 0.542 + 0.04 b 0.855 + 0.13 a 2K Explant plus shoot weight (g) 0.262 + 0.02 c 0.278 + 0.02 c 0.241 + 0.04 c 0.439 + 0.07 be 0.379 + 0.08 be Surviving explants producing shoots (%) 63.3 +6.1b 81.7 + 6.4 ab 61.1+17.1b 90.0 + 6.8 a 90.0+4.4a 96.7+3.3a 100.0+0a 2K Explant survival (%) 100 86 83.3 100 93.0 100 96.7 NS (mgeL!) IAA + kinetin 0.5 + 1.0 0.5 + 2.0 2.0 + 1.0 regulators 2.0 + 2.0 Growth BAP + GA, 2.5+0.01 1.04+ 0.01 3.0 + 2.0 Basal medium BM] BM2 BM3 Means (+ SE) within columns followed by the same letter are not significantly different. NS: Non-significant or *significant at P <0.05 and **0.01, respectively. TABLE 2. Growth and survival of Sericocarpus rigidus explants on basal medium amended with growth regulators after four weeks in culture. Treatment Significance M1 M2 M3 M4 M5 M6 M7 2007 tiplication were obtained with combinations of BAP and GA,, a regime which has not often been used in other Asteraceae studies. Further work is required to determine the optimum basal medium formulation to use with the most successful combinations of BAP plus GA, in order to generate the greatest number of vigorous S. rigidus shoots. Additionally, research with vegetative tissue as a source of explants, rather than seeds, could be conducted in order to avoid the time and difficulties associated with seed germination. Howev- er, from a conservation point of view, harvesting veg- etative tissue, particularly rhizomes, from a threatened or endangered species may be less desirable than har- vesting seeds because of damage to the plant and dis- turbance of the site. The costs of micro-propagation were not evaluated against more conventional propagation techniques. Tis- sue culture is cost effective and is used commercially in the propagation of many plants. Although it requires more knowledge and extra capital to start a micro- propagation operation, the rate of propagation is much faster than in conventional propagation. Given that hundreds of propagules of an endangered or threatened species may be generated using micro-propagation, the effort seems reasonable. Acknowledgments The authors thank Georgia Kliever and Frederic Bounaix for technical assistance, and Matt Fairbarns for collecting seed and providing advice. This study was partially funded by the Interdepartmental Recoy- ery Fund of Environment Canada. Documents Cited (marked * in text) Environment Canada 2006. Species at Risk, White-top Aster. http://www.speciesatrisk.gc.ca/search/speciesDetails_e.c fm?SpeciesID=237. Garry Oak Ecosystem Recovery Team. 2002. Recovery Strategy for Garry Oak and Associated Ecosystems and their Associated Species at Risk in Canada 2001 — 2006, 1 page. Literature Cited Alibert, G., C. Aslane-Chanabé, and M. Burrus. 1994. Sun- flower tissue and cell cultures and their use in biotechnol- ogy. Plant Physiology and Biochemistry 32: 31-44. Bigger, D. S. 1999. Consequences of patch size and isolation for a rare plant: pollen limitation and seed predation. Nat- ural Areas Journal 19: 239-244. FREY, KEMPLER, AND EHRET: MICRO-PROPAGATION OF WHITE-TOP ASTER 45 Clampitt, C. A. 1987. Reproductive biology of Aster curtus (Asteraceae), a Pacific Northwest endemic. American Jour nal of Botany 74: 941-946, Douglas, G. W., and J. M. Illingworth. 1997. Status of the white-top aster, Aster curtus (Asteraceae), in Canada. Cana dian Field-Naturalist 111: 622-627. Ehret, D., B. Frey, and T. Forge. 2004. Recovery of En- dangered Species Inhabiting Garry Oak Ecosystems, PARC Technical Report (172), Agriculture and Agri-Food Canada, 12 pages. Giblin, D. E. 1997. The relationships of reproductive biology and disturbance to the rarity of Aster curtus (Cronq.), a Pacific Northwest endemic. Master of Science dissertation, University of Washington. Giblin, D. E., and C. W. Hamilton. 1999. The relationship of reproductive biology to the rarity of endemic Aster curtus (Asteraceae). Canadian Journal of Botany 77: 140-149. Joshi, M., and U. Dhar. 2003. In vitro propagation of Saus- surea obvallata (DC.) Edgew.— an endangered ethnoreli- gious medicinal herb of Himalaya. Plant Cell Reports 21: 933-939. Kalemba, D., and B. Thiem. 2004. Constituents of the essen- tial oils of four micropropagated Solidago species. Flavour and Fragrance Journal 19: 40-43. Koroch, A. R., J. Kapteyn, H. R. Juliani, and J. E. Simon. 2003. In vitro regeneration of Echinacea pallida from leaf explants. In Vitro Cellular and Developmental Biology — Plant 39: 415-418. Kyte, L., and J. Kleyn. 1999. Plants from Test Tubes, an Introduction to Micropropagation. Timber Press Inc., Port- land, Oregon. Murashige, T., and F. Skoog. 1962. A revised medium for rapid growth and bioassays with tobacco tissue cultures. Physiologium Plantarum 15: 473-479. Orlikowska, T. K., and W. E. Dyer. 1993. In vitro regener- ation and multiplication of safflower (Carthamus tincto- rius L.). Plant Science 93: 151-157. Simmons, C. B. 1998. Pharmacognosy and micropropagation of feverfew (Janacetum parthenium (L) Schultz Bip). Master of Science dissertation, University of Guelph. Slater, A., N. W. Scott, and M. R. Fowler. 2003. Plant bio- technology: The genetic manipulation of plants. Oxford University Press, Oxford. Stewart, S. E., and C. R. Lacroix. 2001. Germination poten- tial, updated population surveys and floral, seed and seedling morphology of Symphyotrichum laurentianum, the Gulf of St. Lawrence Aster, in the Prince Edward Island National Park. Canadian Field-Naturalist 115: 287-295. Vanegas, P. E., A. Cruz—Hernandez, M. E. Valverde, and O. Paredes—L6pez. 2002. Plant regeneration via organo- genesis in marigold. Plant Cell Tissue and Organ Culture 69: 279-283. Received 5 September 2006 Accepted 26 March 2007 Pattern and Potential Causes of White-faced Ibis, Plegadis chihi, Establishment in the Northern Prairie and Parkland Region of North America Jct A. SHAFFER!, GREGORY A. KNUTSEN?, RON E. MArtTIN?, and JoEL S. BRICE* 'U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 37" Street SE, Jamestown, North Dakota 58401 USA; e-mail: jshaffer@usgs.gov °U.S. Fish and Wildlife Service, Long Lake National Wildlife Refuge Complex, 12000 353" Street SE, Moffit, North Dakota 58560 USA 316900 125" Street SE, Sawyer, North Dakota 58781 USA ‘Delta Waterfowl Foundation, 1305 East Central Avenue, Bismarck, North Dakota 58501 USA Shaffer, Jill A., Gregory A. Knutsen, Ron E. Martin, and Joel S. Brice. 2007. Pattern and potential causes of White-faced Ibis, Plegadis chihi, establishment in the northern prairie and parkland region of North America. Canadian Field- Naturalist 121(1): 46-57. The Northern Prairie and Parkland Waterbird Conservation Plan calls for renewed attention to determining the current status of waterbird populations, their distributions, and conservation needs. It highlights the need for baseline information on the White-faced Ibis (Plegadis chihi). In response, we examined the historical and current distribution of the ibis in North Dakota and summarized first sightings and nest records for the provinces and other states composing the northern prairie and parkland region. The establishment of breeding colonies of White-faced Ibis here may be due to climate and precipitation patterns, invasion and spread of Narrowleaf Cattail (Typha angustifolia), changes in agricultural practices, habitat loss and range expansion in the southern and western portions of the species’ range, and increases in ibis populations in the Intermountain West. We placed special emphasis on North Dakota, a state for which there is scant published information concerning the current status of this species. In recent decades, the ibis has become a regular breeding-season resident in North Dakota and in other areas of the northern prairie and parkland region. From 1882 to 2002, there were 145 reports of one or more White- faced Ibis in North Dakota, including 93 reports during the breeding season (15 May to 31 August), 49 during the non- breeding season (1 September to 14 May), and three for which the season of occurrence was not reported. Prior to the 1960s, there were only three records of the species in North Dakota. Observations of White-faced Ibises in North Dakota increased dramatically between the 1960s and the early 21st century, and the species has been observed nearly annually since 1971. The first White-faced Ibis nesting activity in the state was recorded in 1978, and to date, there have been 21 known records of nesting activity in the state. The species nested in large (>300 ha) semipermanent or permanent wetlands within mixed- species colonies ranging in areal extent from small (0.1 ha) to fairly large (27 ha), and colonies were located in patches of emergent vegetation dominated by cattails (Typha) and bulrushes (Scirpus). We classify the White-faced Ibis as a fairly com- mon migrant and a locally uncommon breeder east of the Missouri River and a casual migrant west of the Missouri River. Key Words: White-faced Ibis, Plegadis chihi, colonial-nesting waterbird, distribution, status, northern prairie and parkland region, North Dakota. The White-faced Ibis (Plegadis chihi; hereafter ibis) is a wading bird that nests in wetlands with emergent vegetation and forages in shallow wetlands, flooded hay meadows, or agricultural fields (Ryder and Manry 1994). Northernmost populations undertake regular north-south migrations, whereas populations in Texas and Louisiana are mainly year-round residents (Ryder and Manry 1994). Extralimital wanderings seem to be more prevalent before the nesting season than afterward and may represent individuals seeking new nesting areas when traditional sites are dry or have been destroyed by humans (Ryder 1967; Ryder and Manry 1994). Dur- ing the twentieth century, numerous sightings of the species east and north of its historical range were report- ed (Sabo 1992; Jorgenson and Dinsmore 2005). The recently completed North American Waterbird Conservation Plan (Kushlan et al. 2002) and its regional offshoot, the Northern Prairie and Parkland Waterbird Conservation Plan (NPPWCP) (Beyersbergen et al. 2004), called for renewed attention to determining the current status of waterbird populations, their distribu- tions, and conservation needs. The NPPWCP em- phasized that information on the breeding distribution of ibis in the northern prairie and parkland region (por- tions of Alberta, Saskatchewan, Manitoba, Montana, North Dakota, South Dakota, Minnesota, Iowa, and Nebraska) is needed to serve as a baseline for future monitoring needs (Beyersbergen et al. 2004). In re- sponse to these information needs, we examined the historical and current distribution of the ibis in North Dakota and summarized first sightings and nesting records for the provinces and states composing the northern prairie and parkland region. We placed spe- cial emphasis on North Dakota, a state for which there is scant published information concerning the current status of this species. Lokemoen (1979) and Schmidt (1980a) summarized the first occurrences and nest records of White-faced 46 2007 Ibis in North Dakota. Sabo (1992) described the status, range, first occurrences, and first nesting observations for some Great Plains states and Canadian provinces. Goossen et al. (1995) provided a detailed list of obser- vations and breeding records for the species through- out Canada. Jorgenson and Dinsmore (2005) provided an update of breeding records throughout the Great Plains. In this paper, we summarize the reported occur- rences of White-faced Ibis in North Dakota up to 2002, and we report nest records through 2003, beyond which time the U.S. Fish and Wildlife Service (USFWS) no longer quantified colony and nest numbers. We discuss factors influencing the species’ expansion into the state and provide data on the species’ breeding habitat. We also explain how the pattern of occurrence of the spe- cies in North Dakota compares to the general pattern throughout the rest of the northern prairie and park- land region. Methods Records of ibis occurrences in North Dakota were summarized from published reports, including those in American Birds, Audubon Field Notes, National Audu- bon Society Field Notes, and North American Birds, and unpublished reports filed with the North Dakota Birding Society (REM, unpublished data). We included data from the North American Breeding Bird Survey (BBS; 1966-2002; Sauer et al. 2005*), the National Audubon Society’s Christmas Bird Count (NAS 2007*), and banding records from the U.S. Geological Survey Bird Banding Laboratory (USGS 2007*). We defined a record as a museum specimen or a reported occurrence of an individual, group of individ- uals, or a colony. Records were summarized by date (year, month, and day if known) and location (to coun- ty level at a minimum). Nest records were defined as the presence of nests, eggs, dependent young, or adults exhibiting breeding behavior (e.g., carrying nesting material). Observations from 15 May to 31 August were considered breeding-season records, and obser- vations from 1 September to 14 May were considered nonbreeding-season records, based on authors’ obser- vations of ibis behavior and phenological information as described by Ryder and Manry (1994). Occurrences of ibises at the same location over a range of days to months were counted as a single observation for pur- poses of annual summaries. To summarize data for the breeding and nonbreeding seasons, reports of the spe- cies at the same location within both seasons were counted as separate records, unless information indi- cated that individuals had nested at the location and were still present during the nonbreeding season. We summarized information on use of wetlands for nesting from one published nest record and from three studies that measured habitat attributes of nesting ibis in North Dakota. Schmidt (1980a) provided a brief description of the habitat attributes of a wetland that supported several flightless young in 1978 and of one SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 47 nest in southern North Dakota in 1979. During June and July 2000, Brice (2003) documented a multi-species colony that included ibis nests at Lake Alice National Wildlife Refuge (NWR; 48°19'54"N, 99°06'38"W), western Ramsey County, in northeastern North Dako- ta. Colony size was defined as the area occupied by nests of all colonially nesting species within the colony boundary, not just by ibis nests. Within the colony boundary, number of ibis nests, dominant vegetation, and presence of other bird species nesting within the colony were recorded. Two transects were placed lengthwise through the colony. The percent of water surface that was not covered or obstructed by stand- ing or horizontal residual vegetation (1.e., percent open water), residual stem density, and water depth were measured within 86 1-m? sampling frames placed at 10-m intervals along the two transects (Table 1). Percent open water within the colony was recorded by visual estimation. Water depth was recorded using an approx- imately 3-cm diameter closet rod marked in |-dm inter- vals. In 2001, this same colony was monitored for the presence of breeding ibises by USFWS personnel (Phelps and Meeks 2001*). Percentages of colony area composed of residual emergent vegetation and open water were visually estimated. During June and July 2001 and 2002, the USFWS (2001*, 2002*) document- ed ibis nesting activity on USFWS lands in Burleigh, Emmons, and Kidder counties in south-central North Dakota. The number of nests, the number of colonies, the dominant vegetation within colonies, and the pres- ence of other nesting bird species were recorded on one NWR and one waterfowl! production area (WPA). The ratio of live-to-residual emergent vegetation within colonies was visually estimated. In 2001, water depth at nests was measured using a meter stick. During June 2003, Knutsen (2004*) conducted a colonial-nesting waterbird inventory on 183 wetlands located on NWRs and WPAs in the Long Lake Wetland Management District (Knutsen 2004*). Wetlands were divided into three strata (high, moderate, and low) related to their likelihood of supporting a waterbird colony, based on habitat conditions in 2003, as well as the size and per- manence of the wetlands (sensu Stewart and Kantrud 1971). Number of nests, number of colonies, dominant vegetation, and the presence of other colonial bird species nesting within colonies were recorded. The ratio of live-to-residual emergent vegetation within colonies and the percent of the colony’s area that was composed of residual emergent vegetation and open water were visually estimated. Water depth was not measured in 2003. Vegetation density was classified into one of three categories: dense (heavy, thick stands of emer- gent vegetation with few visible openings), moderate (contiguous stands of emergent vegetation with numer- ous small openings), or sparse (large “canoe-sized™ openings within the emergent vegetation of open-water floating mat colonies). THE CANADIAN FIELD-NATURALIST Vol. 121 Source Schmidt (1980a) Phelps and Meeks (2001*) Knutsen (2004*) Schmidt (1980a) Brice (2003) USFWS (2002*) Knutsen (2004*) USFWS (2001*) Water Other Yes 57-77 Yes Yes Yes Yes Yes depth species (cm) present 80 80 Ratio of live-to-dead vegetation 1:1 at 1 colony, more live than 1:1 at the smaller colony, more live than dead at larger colony 9:1 dead at 3 colonies % % open emergent water vegetation 20-50 30-50 73 Dominant vegetation Scirpus sp. Typha spp., Schoenoplectus sp. Typha spp. Typha sp. Typha spp. Typha spp. Phragmites australis, Typha spp., Schoenoplectus sp. Phragmites australis, 15 Typha spp., Schoenoplectus sp. ibis nests or pairs Number of 2-4 nests in 1 colony 0.1-1.2 23 nests in 4 colonies 31 pairs in 3 colonies 11 nests in 2 colonies 7 nests in | colony 8 nests in | colony 1 nest 8 nests Colony area (ha)! 10-27 2.8, 6.0 6.5 1] Wetland size (ha) 2 380 4913 4913 345 345 1978 2001 1979 2000 2001 2002 2003 Year 'Colony area refers to the area occupied. by nests of all colonially nesting species within the colony, not just to ibis nests. The wetland size for the 1978 ibis observation includes the entire wetland basin size for Long Lake. The wetland size in 2001 and 2003 reflects the size of an impoundment that did Long Lake NWR, Burleigh Co not exist in 1978. Kraft Slough, Sargent Co. Lake Alice NWR, Ramsey Co. location TABLE |. White-faced Ibis colonies in North Dakota for which wetland and colony sizes, vegetation characteristics, and water depths were recorded. Colony (Dewald Slough), Kidder Co. Kleppe Lang WPA Results Ibis in North Dakota The only reported observation of ibis in North Dako- ta in the nineteenth century was of a specimen that was collected in 1882 and deposited in the University of Michigan’s Museum of Zoology (Wood 1923). Wood (1923) also reported the first two occurrences in the twentieth century, including single individuals in Morton County in 1912 and in Slope County in May 1920. The fourth report of an ibis occurred over 40 years later, on 5 June 1961, in Stutsman County (fide REM). This observation marked the beginning of the species’ documented range expansion into North Dakota (Figure 1). Observations of ibis increased mark- edly between the 1960s and 1990s. Between 1961 and 2002, the species has been observed nearly annually, occurring in all but nine of the 42 years. From 1882 to 2002, there were 145 reports of one or more White-faced Ibis in North Dakota, including 93 reports during the breeding season, 49 during the nonbreeding season, and three for which the season of occurrence was not reported. Of the 107 occurrences of the species during the twentieth century, 71 occurred during the breeding season, 34 during the nonbreeding season, and the season of occurrence for two early records was unknown (Figure 1). During the twenti- eth century, no occurrences were reported from Janu- ary through March, and two occurrences each were reported in November and December (Figure 2). The earliest observation within a year was 14 April, which occurred in 2002 at Long Lake NWR (46°43'59"N, 100°05'43"W) in Burleigh County (P. C. Van Ningen, U.S. Fish and Wildlife Service, Moffit, North Dakota, personal communication). The latest observation with- in a year was 17 December 1999 in Sargent County (D. Kinzler, Oakes, North Dakota, personal commu- nication). The largest group of birds observed during the breeding season was about 400 individuals on 11 August 2001 in Sargent County; this group was un- doubtedly a post-breeding flock (Martin 2002). The largest group of birds observed during the nonbreeding season was over 50 individuals in September 2000 in Kidder County by H. F. Duebbert (Martin 2001). From 1912 to 2002, ibises have been sighted in 28 (53%) of the 53 counties in North Dakota (Figure 3). Observations first occurred west of the Missouri River. The two observations in the early part of the twentieth century both occurred in the western part of the state (Wood 1923). Of the eight occurrences in the 1960s, all were reported from southern counties, but only one occurred west of the Missouri River, in Slope County (G. B. Berkey, personal communication). Most obser- vations after the 1960s were east and north of the Missouri River. The first occurrences in the northern half of the state occurred in 1973 at Upper Souris NWR (48°30'11"N, 101°31'50"W), Renville County, by I. O. Rostad (Houston 1974) and in 1974 at J. Clark Salyer NWR (48°40'52"N, 100°47'23"W), McHenry County, 2007 70 ——— () Unknown Season Nonbreeding Season @ Breeding Season Number of observations 1901- 1910 1911- 1920 1921- 1930 1931- 1940 194]- 1950 SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 49 1951- 1960 1961- 1970 1971- 1980 1981- 1990 1991- 2000 FicureE 1. Records of White-faced Ibis occurrences in North Dakota during the twentieth century, by decade. by R. C. Fields (Houston and Houston 1974). The first sightings in five counties (Bottineau, Foster, Mercer, Pierce, and Ward) occurred recently, within 2001-2002. To date, there are 21 records of nesting activity in North Dakota (Table 2). The first nest was reported by R. A. Schmidt and R. N. Randall in 1978 (Serr 1978: Schmidt 1980a). From 1978 to 2003, nests have been reported in ten of the 26 years, with the majority occur- ring between 2000 and 2003. Nests or nesting activity have been reported in each of the four recent years from 2000 to 2003. The number of nests discovered within an individual wetland varied from one to 31. All 21 nesting observations were recorded in June, July, or August. Observations in June indicated that nest- ing had been initiated in May. By August, young or fledglings were observed. Nesting ibises have been observed in eight of the 53 counties (Table 2, Figure 3). Most nesting activity was recorded in Burleigh and Kidder counties (Table 2). Of the ten known wetlands that have contained nesting ibises, six are federally owned; two involve a mixture of federal, state, and/or private ownerships; and two are privately owned. The ibis has been recorded on a Breeding Bird Sur- vey route in North Dakota in only one year since the survey's inception in 1966. On 10 June 1999, three individual ibis were reported by H. C. Talkington (Mandan, North Dakota, personal communication) on his BBS route south of Tappen in Kidder County. According to banding records, three flightless young were banded in late July 1979, in western Sargent Coun- ty (K. Klimkiewicz, U.S. Geological Survey, Laurel, Maryland, personal communication). No ibises have been recorded in North Dakota on a Christmas Bird Count, which have been conducted in the state since the early 1900s. In North Dakota, ibises have been found breeding only in large (>300 ha) semipermanent or permanent wetlands. Colonies, all of which were mixed-species colonies, were located in patches of emergent vegeta- tion dominated by cattails and bulrushes and ranged in areal extent from 0.1 to 27 ha (Table 1). Ibises appeared to prefer a fairly equal proportion of open water and emergent vegetation within a wetland basin, with half or more of the vegetation composed of live rather than residual stems. Water depths varied from 60 to 80 cm. Schmidt (1980a) reported that one ibis nest was built just above the water level in residual cattails inter- spersed with open water and was located 600 m from shore and 20 m from the central open water area of the wetland basin. Flightless ibises were located in emer- gent vegetation about 700 m from shore and 35 m from open water. Brice (2003) reported that the mean densi- ty of residual cattail stems within an ibis colony was 44 stems/m?. Knutsen (2004*) found that two of 40 waterbird colonies contained ibis nests. Both of these ibis colonies were located on wetlands categorized as having a high probability of supporting a waterbird colony. One of the colonies was located on an im- poundment of Long Lake that contained ibises in 2001, and the other was located on a WPA that contained 50 THE CANADIAN FIELD-NATURALIST Vol. 121 | nN oo Sp i) tN i) tO Ne i) & o Number of observations a ie Jan Feb Mar 18 + 14 + 12 + 10 + 3h 6 4 2 0 r T T T r 53 T r Apr May Jun Jul Aug Sep Oct Nov Dec Month FIGURE 2. Records of White-faced Ibis occurrences in North Dakota during the twentieth century, by month. An observation) was counted once, regardless of whether the observation was of one or of multiple individuals. ibises in 2002 (Table 1). Vegetation density within the colony in the impoundment was categorized as mod- erate, whereas vegetation density within the colony in the WPA was categorized as dense. Several other waterbird species occurred at colonies with ibises (Table 1). These species included Ruddy Duck (Oxyura Jamaicensis), Horned Grebe (Podiceps auritus), Eared Grebe (Podiceps nigricollis), Western Grebe (Aech- mophorus occidentalis), Clark’s Grebe (Aechmophorus clarkii), Snowy Egret (Egretta thula), Cattle Egret (Bubulcus ibis), Black-crowned Night-Heron (Nycti- corax nycticorax), American Coot (Fulica americana), Franklin’s Gull (Larus pipixcan), Forster’s Tern (Sterna forsteri), and Black Tern (Chlidonias niger) (Schmidt 1980a; Phelps and Meeks 2001*; USFWS 2001*, 2002*; Brice 2003; Knutsen 2004*). Ibis expansion in the northern prairie and parkland region There appears to be a consistent pattern throughout the northern prairie and parkland region concerning the chronological progression from the first occurrences to the establishment of breeding colonies (Table 3). From our examination of regional patterns of ibis dis- tribution, we determined that the species rarely visited the northern prairie and parkland region in the late nineteenth and early twentieth centuries and did not develop breeding populations until the late twentieth century. North Dakota, Iowa, and Minnesota reported ibis occurrences as early as the late nineteenth century (Peabody 1896; Lokemoen 1979; Schmidt 1980a; Jans- sen 1987; Kent and Dinsmore 1996), and Nebraska and Alberta reported occurrences during the early twenti- eth century (Kondla et al. 1973; Goossen et al. 1995; Mollhoff 2001). Montana, South Dakota, Manitoba, and Saskatchewan did not report ibis occurrences until the mid-to-late twentieth century (Skaar 1969; Serr 1975; DuBois 1989; Goossen et al. 1995; Tallman et al. 2002). The first nests in the region were reported in’ 1894 in Minnesota (Peabody 1896) and in 1916 in Nebraska (Mollhoff 2001), but other states and prov- inces did not report nesting ibis until the mid-to-late twentieth century. Two nest records have been report- ed for Saskatchewan, both of which occurred in 2000 (B. Hepworth, Ducks Unlimited Canada, Regina, Sas- katchewan, personal communication; P. Taylor, Cana- dian Wildlife Service, Environment Canada, Saska- toon, Saskatchewan, personal communication). Bazin and Artuso (2006) documented the first nest record for Manitoba in 2005. Jorgenson and Dinsmore (2005) summarized recent nesting records for Alberta and! some of the states in the northern prairie and parkland region. An additional nest record in Alberta was report- ed by Flockhart (2001). Discussion The number of occurrences of ibises in North Dako- ta reached the point that the North Dakota Birding Society stopped recording new nonbreeding observa- tions after about 2002 (REM, unpublished data). Nest- ing ibises are becoming increasingly common, with multiple reports of nests every year from 2000 to 2003 (Table 2). Spring arrival in North Dakota typically begins in mid-to-late April, and birds begin departing in September (Figure 2). Few birds remain in the state in November and December, and no records exist for January through March. Nesting occurs from mid-May | to late August (Table 2). The ibis appears to nest colo- | nially with other waterbird species as has been report- 2007 ii ae Hettinger SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 5] FiGURE 3. Records of White-faced Ibis occurrences in North Dakota during the twentieth century and 2001-2002, by county. Asterisks (*) denote counties in which nesting activity has been observed during the twentieth century and 2001-2003. ed by other researchers in the northern prairie and park- land region (Peabody 1896; Dinsmore and Dinsmore 1986; Mollhoff 2001). Based on our findings, we be- lieve that the status of ibis in North Dakota is that of a fairly common migrant and a locally uncommon breed- er east of the Missouri River and a casual migrant west of the Missouri River. Of the ten known wetlands that have contained nest- ing ibis, most are under federal management. These wetlands are protected from drainage and tillage; there- fore, thick stands of cattails and other tall wetland veg- etation are able to become established in some years (Table 1). These wetlands also may experience higher levels of human visitation than privately owned wet- lands. More human visits, either by federal employees conducting biological surveys or by bird watchers and nature enthusiasts visiting the NWRs, WPAs, and other federal properties, could explain the greater number of reports of nesting activity on federally managed properties. Because ibises were present as vagrants in some parts of the northern prairie and parkland region in the late nineteenth century, several authors have speculated that the presence of breeding colonies in the twentieth cen- tury was not so much an expansion of the species’ range as it was the reclamation of portions of the species’ for- mer range (Ryder and Manry 1994; Dinsmore et al. 1984). Goossen et al. (1995) advanced several explana- tions for the expansion of breeding ibises into Canada, which included population growth; loss of habitat to draining, flooding, or drought in other portions of the Species’ breeding range; dispersal; and an increase in Suitable habitat due to the advent of managed or re- stored wetlands and irrigation. The virtual absence of sightings from the late nineteenth century through the early twentieth century in the northern prairie and park- land region (Table 3) coincided with the withdrawal of ibises from inland areas in Texas, suggesting to Jor- genson and Dinsmore (2005) that sightings in northern regions in the late nineteenth century and first half of the twentieth century were linked to fluctuations in the Gulf Coast populations. They examined the historical status of the species from Alberta to Texas and sug- gested that the current range expansion appears to be unprecedented. They contended that the idea that ibis- es occupied the northern Great Plains in the nineteenth century was based on only three nesting records that occurred at a time when extralimital breeding of the species was being documented elsewhere in the conti- nent. They suggested that the development of breeding populations of ibises was linked to population fluctu- ations in the species’ core breeding areas of Texas, Louisiana, and, more likely in modern times, to fluctu- ations occurring in the Intermountain West, the region of North America lying between the Rocky Moun- tains to the east and the Cascades and Sierra Nevada to the west. Drawing upon knowledge of habitat, climate, and socioeconomic conditions in the northern Great Plains, using North Dakota as an example, we suggest fur- ther explanations for the pattern of ibis observations. 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Alng Z ZOOT Atne aIv| 1007 aung ZZ 1007 oung 1007 sung ooo Aine QOOT eunr 9661 Arne 066] Isnsny T] — A[n¢ Cz 0661 Aine 81 S861 Aine I 8861 Alne 1 B86 Arne ZRH] uns | 6L6| Isnsny Z| — une ¢ 8161 AINE OF-8I (s)aleqd COOT YBNosy) RIOYLE YON UT SIQ] PIovJ-a}TY MA JO SpsO99I SUNSON 7 AAV 2007 and Dinsmore’s (2005) suggestion that ibis occurrences in the northern Great Plains during the late nineteenth century and early twentieth century may have been due to extralimital wandering as was occurring in other parts of the U.S. at the same time, combined with the species’ predilection for post-breeding dispersal north of its nesting grounds (Ryder 1967; Ryder and Manry 1994). No ibises were reported in North Dakota between 1920 and 1961 (Table 3). Plausible explanations include a scarcity of suitable nesting habitat and a low level of human engagement in nature-based activities. Potential suitable wetland habitat for ibises deteriorated with the onset of drought that characterized the 1930s, the Dust Bowl years (Robinson 1966). Given the duration and intensity of that drought, it is highly likely that suitable wetland habitat was largely nonexistent in the 1930s. The widespread drought of the 1930s, combined with economic collapse during the Great Depression, resulted in a human exodus from North Dakota (Rob- inson 1966). The human population in North Dakota reached the highest levels to date, according to the 1930 census, and experienced a 9% decline between 1930 and 1950 (USCB 2005), with much of the decline occur- ring in rural areas. The human population continues to fluctuate and recovered somewhat from the 1950s, but never has reached the levels experienced prior to the 1930s. During the 1940s, fewer people lived in North Dakota than in the 1920s or 1930s (Kazeck 1956), and some people that remained were called to war or were engaged in war-related activities. During the war, am- munition was rationed and hunting activities were sub- sequently curtailed. Overall, then, there likely were fewer people afield and thus there were fewer chances of people encountering ibises during this time. Also during the 1940s, moisture conditions, and thus potential ibis habitat, began to improve, with the high- est annual precipitation at Bismarck between 1900 and 2000 occurring in 1941 with 79.1 cm (NCDC 2004*). Concomitant with improved moisture condi- tions was the invasion into North Dakota of an intro- duced species of European cattail, the Narrowleaf Cat- tail (Typha angustifolia), which was first collected in North Dakota in 1942 at Long Lake NWR (Kantrud 1992). Prior to that time, there were few cattail-dom- inated wetlands in North Dakota, and the few stands that did exist were in boggy areas around freshwater lakes (Kantrud 1992). Narrowleaf Cattail was restrict- ed largely to the southeastern portion of North Dakota in the early 1960s and spread to many wetlands in central North Dakota by the 1970s (Kantrud 1992). However, the invasive species hybridized with the native Broadleaf Cattail (Typha latifolia), and the resulting hybrid (7ypha xglauca) spread rapidly throughout the prairie pothole region of North Dakota during the 1950s, becoming the most abundant large hydrophyte in the state (Kantrud 1992). Wetlands dominated by the hybrid cattail potentially provided prime breeding habi- tat for ibises. Also, despite the decrease in the number TABLE 3. Pattern for the northern prairie and parkland region, USA and Canada, that exhibits the lapse between time of first White-faced Ibis observations, time of consistent (>5 observations in a given decade) observations, and year of first nest record. Consistent sightings Early sightings Mid-to-late 1900s Mid-to-late Late Early 1900s Early Late Sources First nest 1900s 1800s 1900s 1912 1800s 1882 189] Lokemoen 1979; Schmidt 1980a; this paper SHAFFER ET AL.; WHITE-FACED IBIS IN NORTHERN PRAIRIE Dinsmore and Dinsmore 1986; Kent and Dinsmore 1996 2 AND PARKI _ / Peabody 1896; Janssen 1987 Mollhotf 2001 Skaar 1969; DuBois 1989 + Tallman et al. 200 Kondla et al. 1973; Goossen et al. 1995 SDOU 1991; 1960s North Dakota lowd Min 1980s = 1960s 1910 1894-1895 nesota 983-1984 1960s 1970s 1970s AND 006 Goossen et al. 1995; personal communications: B. Hepworth’; R. Luterbach?: 5 O05 O00 5 Serr 1975; Goossen et al. 1995; Bazin and Artuso > 1970s 1916 = h Dakota = Sou 194] Manitoba Saskatchewan P. Taylor’. *P. Taylor, Canadian *R. Luterbach, Regina, Saskatchewan, personal communication 'B. Hepworth, Ducks Unlimited Canada, Regina, Saskatchewan, personal communication. Wildlife Service, Saskatoon, Saskatchewan, personal communication. 53 54 THE CANADIAN FIELD-NATURALIST of temporarily flooded wetlands due to drainage, the number of permanent wetlands in the form of basin wetlands and riverine impoundments increased in the state (Lokemoen 1979). Kantrud (1992) suggested that prairie wildfires and grazing by large ungulates, two major ecological forces during presettlement times, were responsible for the restricted distribution of Broadleaf Cattail prior to the invasion of Narrowleaf Cattail. Many semipermanent wetlands in eastern North Dakota, eastern South Dako- ta, and western Minnesota that were grazed and that were dominated by semi-open stands of Hardstem Bul- rush (Schoenoplectus acutus) presently are not grazed and are dominated by dense stands of cattail. Gary Krapu (U.S. Geological Survey, Jamestown, North Da- kota, personal communication) recalls that, prior to the mid-1950s in North Dakota, most wetlands expe- rienced grazing pressure by livestock or were hayed. In the late 1950s and 1960s, however, many farmers began to focus exclusively on small-grain production, phasing out livestock production. Livestock were ben- eficial in stopping the growth of dense stands of cat- tails by eating young, green cattail shoots. With the loss of grazing pressure, Narrowleaf Cattail more easily became established in wetlands. The decline in farm operations that raised livestock was hastened by the Soil Bank Program in the 1950s, which provided farmers with annual payments under 5-to-10 year contracts to idle their cropland by planting to perennial cover. This program indirectly also encouraged the idling of wet- lands as farmers reduced the number of livestock that they produced. During the 1950s and 1960s in southern portions of the species’ breeding range, ibis breeding habitat was being lost due to human encroachment or drought. Breeding colonies of ibises disappeared or decreased in size due to habitat loss or pesticide contamination in Oregon, California, and Nevada (Ryder 1967; Ryder and Manry 1994). This decline coincided with increased sightings in Minnesota, North Dakota, South Dakota, and Montana (Table 3). It is plausible that this highly nomadic species sought and found better breeding con- ditions in the northern prairie and parkland region than in its more traditional breeding range. Banding studies during the 1960s found that shoot- ing was the major cause of reported deaths for this species, with many bands recovered from Mexico (DuBois 1989). The species was a gamebird until the early twentieth century, when international treaties con- ferred protection from hunting or harassment. Loke- moen (1979) postulated that ibis population increases might have been due in part to federal protection. Al- though federal protection of migratory birds began in 1918 with the signing of the Migratory Bird Treaty (Legal Information Institute 2005*), resources were not available for widespread federal enforcement in North Dakota until the 1940s (G. Krapu, personal com- munication). Vol. 121 After the late 1960s, pesticide contamination and continued habitat loss occurred in southern states, coin- ciding with more frequent ibis sightings and first nest records in several northern states. King et al. (1980) monitored eggshell thickness, pollutant residues, and the population status of the species in Texas from 1969 through 1976. They found that nesting populations de- clined by 42%, associated with low reproductive suc- cess caused by DDE-induced shell thinning and diel- drin mortality. Taylor et al. (1989) reported that the ibis population doubled in the late 1970s in Utah and also increased in Oregon. They attributed these increas- es to flooding in the Great Basin during the early 1980s, recovery from reduced productivity caused by DDT in the 1970s, and/or natural fluctuations in ibis popu- lations. Other Cicontiformes species also increased in oc- currences and expanded their distribution during the same time period in North Dakota and other areas of the northern prairie and parkland region (Jones and Malcolm 1978; Lokemoen 1979; Schmidt 1979a,b, 1980b; Lambeth 1980). The first Cattle Egret in North Dakota was reported in 1971 (Lokemoen 1979), and the first nesting colony was discovered in 1976 among nests of Black-crowned Night-Herons and Little Blue Herons (Egretta caerulea) (Jones and Malcolm 1978). More Cattle Egret colonies were discovered in the en- suing years (Schmidt 1980b). It is possible that the presence of the colonies of Cattle Egret and other colo- nial-nesting birds provided the stimulus that prompt- ed ibis to establish colonies. More recently, the increase in White-faced Ibis ob- servations in North Dakota has coincided with the wet conditions experienced in the northern prairie and park- land region from 1993 to the present (NCDC 2004*). The current wet cycle may be the wettest period dur- ing the past 130 years (Winter and Rosenberry 1998). These wet conditions may be contributing to suitable habitat conditions for the ibis. Johnson (1994) docu- mented large-scale pioneering expansion of nesting dis- tributions of several species of western North American birds over the last three decades. Although Johnson (1994) did invoke climatic warming as a possible expla- nation, he believed that it was neither the sole expla- nation nor even the primary cause. Rather, he hypothe- sized that many of the species were responding to a decades-long increase in summer rainfall in regions beyond their former ranges, based on regional climatic trends. Summer moisture, perhaps coupled with a high- er mean temperature, encouraged avian range expan- sions. Detailed correlations of local climatic data with temperature and moisture requirements of individual species would be necessary to establish causality (John- son 1994), We believe that the combination of adequate rain- fall and invasion of Narrowleaf Cattail in the 1940s in North Dakota, the rapid establishment of Narrowleaf Cattail and its hybridization with the Broadleaf Cat- 2007 tail during the ensuing decades, the general expansion of other cicontiform species, the conversion from ranch- ing to small-grain production, wetland loss and poor ibis reproduction in the southern portions of the ibis breeding range, growing ibis populations in the Inter- mountain West region, wet conditions suitable for colo- nial waterbird establishment, and climatic changes have contributed to the growth of ibis populations in North Dakota. We suspect that the above factors also may have played a role in population changes elsewhere in the northern prairie and parkland region. Ibises are considered rare, localized breeders in the northern prairie and parkland region (Beyersbergen et al. 2004) and, more specifically, are considered locally uncom- mon in North Dakota (this paper), rare to locally un- common in South Dakota (Sabo 1992), and a rare sum- mer resident in Canada (Goossen et al. 1995). The future status of ibis within the northern prairie and parkland region likely will depend upon precipi- tation patterns. With continuing years of adequate rain- fall, wetlands that harbored colonies should continue to provide suitable habitat. There are no indications that the spread of monotypic stands of cattail within wet- lands will abate, which should only aid in the spread of ibis into new breeding areas. Already, breeding sites in North Dakota that once contained only a few birds and nests now contain larger groups of nesting birds. With an onset of a prolonged drought, the quality of nest- ing habitat could decline in some wetlands, but water levels in other wetlands that had been inundated to the degree that they no longer supported emergent veg- etation would recede and again allow the growth of deep-marsh vegetation that could harbor new nesting colonies. Even in a prolonged drought, colonies could persist at managed wetlands, such as those maintained within certain NWRs. From these colonies, ibis could disperse when precipitation levels again rise at previ- ously dry wetlands. Acknowledgments We thank the many individuals who have con- tributed records of White-faced Ibis in North Dakota. We thank Richard Crawford, Todd Grant, Dave Grif- fiths, Craig Hultburg, Lawrence Igl, Will Meeks, and Mike Norton for providing detailed information on unpublished sightings. Will Meeks of the U.S. Fish and Wildlife Service procured funding and equipment for Joel Brice’s graduate field research, and Richard Crawford served as his graduate advisor. Kathy Klim- kiewicz of the Bird Banding Laboratory provided infor- mation on ibis banding records. Special thanks to Gary Krapu for sharing his knowledge and experiences of North Dakota birdlife and to him and Lawrence Igl for prompting discussions on factors relating to ibis pop- ulation increases. Thank you to Rachel Bush for con- ducting library searches. Early drafts of the manuscript were reviewed by Steve Dinsmore, Lawrence Ig], Dou- glas Johnson, Gary Krapu, and Terry Shaffer. SHAFFER ET AL.: WHITE-FACED IBIS IN NORTHERN PRAIRIE AND PARKLAND 55 Documents Cited (marked * in text) Knutsen, G. A. 2004. Colonial nesting waterbird distribution and abundance within Long Lake National Wildlife Refuge Complex. U.S. Fish and Wildlife Service, Moffit, North Dakota. Legal Information Institute. 2005. Cornel! University Law School Legal Information Institute home page: U.S. Code Title 16. http://www.law.cornell.edu/uscode/html/uscode 16/usc_sup_01_16_10_7_20_IU.html. NAS (National Audubon Society). 2007. National Audubon Society Christmas Bird Count home page. http://audubon2. org/birds/cbe/hr/table.html. NCDC (National Climatic Data center). 2004. National Climatic Data Center home page: climate division, precipi- tation, temperature, and drought data. http://www.ncde. noaa.gov/onlineprod/drought/xmegr.html. Phelps, W. A., and W. Meeks. 2001. Waterbird nesting colonies at Lake Alice National Wildlife Refuge. Unpub- lished report. U.S. Fish and Wildlife Service, Devils Lake Wetland Management District, Devils Lake, North Dakota. Sauer, J. R., J. E. Hines, and J. Fallon. 2005. The North American Breeding Bird Survey, Results and Analysis 1966 — 2004. Version 2005.2. USGS Patuxent Wildlife Re- search Center, Laurel, Maryland. http://www.mbr-pwre. usgs.gov/bbs/bbs.html. USFWS (U.S. Fish and Wildlife Service). 2001. Long Lake National Wildlife Refuge and Long Lake Wetland Manage- ment District annual narrative report, Calendar Year 2001. Moffit, North Dakota. USFWS (U.S. Fish and Wildlife Service). 2002. Long Lake National Wildlife Refuge and Long Lake Wetland Man- agement District annual narrative report, Calendar Year 2002. Moffit, North Dakota. USGS (U.S. Geological Survey). 2007. USGS Patuxent Wild- life Research Center, Laurel, Maryland. Bird Banding Lab- oratory home page. http://www.pwrc.usgs.gov/bbl. Literature Cited Bazin, R., and C. Artuso. 2006. First documented breeding record of White-faced Ibis in Manitoba. Blue Jay 64: 64-68. Berkey, G. 1988. The nesting season: northern Great Plains. American Birds 42: 1305-1307. Beyersbergen, G. W., N. D. Niemuth, and M. R. Norton. 2004. Northern prairie and parkland waterbird conserva- tion plan. Prairie Pothole Joint Venture, Denver, Colorado. Brice, J. S. 2003. Distribution, abundance and habitat use of overwater colonial nesting waterbirds at Lake Alice Na- tional Wildlife Refuge. M.S. thesis. University of North Dakota, Grand Forks, North Dakota. Dinsmore, J. J., T. H. Kent, D. Koenig, P. C. Petersen, and D. M. Roosa. 1984. Iowa birds. Iowa State University Press, Ames, Iowa. Dinsmore, S., and J. J. Dinsmore. 1986. White-faced Ibis nesting in Dickinson County. lowa Bird Life 56: 120-121. DuBois, K. 1989. Arising, alighting ibis. Montana Outdoors 20(6): 30-33. Flockhart, D. T. Tyler. 2001. Rare or unusual bird sightings for Beaverhill Lake, AB, 1996-2000. Blue Jay 59: 33-40. 56 THE CANADIAN FIELD-NATURALIST Goossen, J. P., D. M. Ealey, H. Judge, and D. C. Duncan. 1995. Distribution and breeding status of the White-faced Ibis, Plegadis chihi, in Canada. Canadian Field-Naturalist 109: 391-401. Houston, C. S. 1974. Fall migration: northern Great Plains. American Birds 28: 67-70. Houston, C. S., and M. I. Houston. 1974. Nesting season: northern Great Plains. American Birds 28: 916-918. Janssen, R. B. 1987. Birds in Minnesota. University of Min- nesota Press, Minneapolis, Minnesota. Johnson, N. K. 1994. Pioneering and natural expansion of breeding distributions in western North American birds. Studies in Avian Biology 15: 27-44. Jones, L. A., and J. M. Malcolm. 1978. First nesting record of Cattle Egret and Little Blue Heron in North Dakota. Prairie Naturalist 10: 122. Jorgenson, J. G., and S. J. Dinsmore. 2005. An assessment of the status of White-faced Ibis (Plegadis chihi) in the Great Plains. North American Birds 59: 376-381. Kantrud, H. A. 1992. History of cattails on the prairies: wild- life impacts. Pages 9-12 in Proceedings of the Cattail Man- agement Symposium. Edited by G. M. Linz. North Dakota State University, Fargo, North Dakota. Kazeck, M. E. 1956. North Dakota, a human and economic geography. North Dakota Institute for Regional Studies, North Dakota Agricultural College, Fargo, North Dakota. Kent, T. H., and J. J. Dinsmore. 1996. Birds in Iowa. Pub- lished by the authors, Iowa City and Ames, Iowa. King, K. A., D. L. Meeker, and D. M. Swineford. 1980. White-faced Ibis populations and pollutants in Texas, 1969- 1976. Southwestern Naturalist 2: 225-240. Kondla, N. G., H. W. Pinel, C. A. Wallis, and C. R. Wersh- ler. 1973. Avifauna of the Drumheller area, Alberta. Cana- dian Field-Naturalist 87: 377-393. Kushlan, J. A., M. J. Steinkamp, K. C. Parsons, J. Capp, M. Acosta Cruz, M. Coulter, I. Davidson, L. Dickson, N. Edelson, R. Elliot, R. M. Erwin, S. Hatch, S. Kress, R. Milko, S. Miller, K. Mills, R. Paul, R. Phillips, J. E. Saliva, B. Sydeman, J. Trapp, J. Wheeler, and K. Wohl. 2002. Waterbird conservation for the Americas: the North American waterbird conservation plan, Version 1. Water- bird Conservation for the Americas, Washington, D.C. Lambeth, S. 1980. Green Heron nesting in North Dakota. Prairie Naturalist 12: 109. Lokemoen, J. T. 1979. The status of herons, egrets, and ibises in North Dakota. Prairie Naturalist 11: 97-110. Martin, R. 2001. Fall migration: northern Great Plains. North American Birds 55: 68-69. Martin, R. 2002. Fall migration: northern Great Plains. North American Birds 56: 66-67. Martin, R. 2004. Fall migration: northern Great Plains. North American Birds 58: 95-97. Mollhoff, W. J. 2001. Nebraska breeding bird atlas. Nebras- ka Ornithologists’ Union Occasional Papers Number 7. Nebraska Technical Series Number 20. Nebraska Game and Parks Commission, Lincoln, Nebraska. Vol. 121 Peabody, P. B. 1896. White-faced Ibis breeding in Minnesota. Auk 13: 79. Robinson, E. B. 1966. History of North Dakota. University of Nebraska Press, Lincoln, Nebraska. Ryder, R. A. 1967. Distribution, migration and mortality of the White-faced Ibis (Plegadis chihi) in North America. Bird Banding 38: 257-277. Ryder, R. R., and D. E. Manry. 1994. White-faced Ibis. In The birds of North America, No. 130. Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadel- phia and the American Ornithologists’ Union, Washing- ton, D.C. Sabo, T. 1992. Plegadis ibis — a change in status. Birder’s Journal 1: 241-256. chmidt, R. 1979a. Burleigh County, second nesting loca- tion of Cattle Egrets in North Dakota. Prairie Naturalist le 10): Schmidt, R. 1979b. First nesting record of a Louisiana Heron in North Dakota. Prairie Naturalist 11: 93-95. Schmidt, R. A. 1980a. First breeding records of the White- faced Ibis in North Dakota. Prairie Naturalist 12: 21-23. Schmidt, R. A. 1980b. Third nesting of Cattle Egret in North Dakota. Prairie Naturalist 12: 19-20. Serr, E. M. 1975. Breeding season — northern Great Plains region. American Birds 29: 867-870. Serr, E. M. 1978. Breeding season — northern Great Plains region. American Birds 32: 1176.Serr, E. M. 1979. Breeding season — northern Great Plains region. Ameri- can Birds 33: 874-875. Skaar, P. D. 1969. Birds of the Bozeman latilong. Published by the author. Bozeman, Montana. South Dakota Ornithologists’ Union. 1991. The birds of South Dakota. SDOU (South Dakota Ornithologists’ Union), Aberdeen, South Dakota. Stewart, R. E., and H. A. Kantrud. 1971. Classification of natural ponds and lakes in the glaciated prairie region. U.S. Fish and Wildlife Service, Resource Publication 92. Bureau of Sport Fisheries, Washington, D.C. Svingen, D., and R. E. Martin. 2003. First report of the North Dakota Rare Bird Committee. Prairie Naturalist 35: 257-272. Tallman, D. A., D. L. Swanson, and J. S. Palmer. 2002. Birds of South Dakota. Midstates/Quality Quick Print, Aberdeen, South Dakota. Taylor, D. M., C. H. Trost, and B. Jamison. 1989. The biol- ogy of the White-faced Ibis in Idaho. Western Birds 20: 125-133. USCB (U.S. Census Bureau). 2005. Decennial census. Wash- ington, DC. Winter, T. C., and D. O. Rosenberry. 1998. Hydrology of prairie pothole wetlands during drought and deluge: a 17- year study of the Cottonwood Lake wetland complex in North Dakota in the perspective of longer term measured and proxy hydrological records. Climate Change 40: 189- 209. Wood, N. A. 1923. A preliminary survey of the bird life of North Dakota. University of Michigan Museum of Zool- ogy Miscellaneous Publications, Number 10. University of Michigan Press, Ann Arbor, Michigan. Received 12 October 2006 Accepted 7 May 2007 Recent Distribution Records of the Little Brown Bat, Myotis lucifugus, in Manitoba and Northwestern Ontario JouN E. Dupois!? and KIMBERLY M. MONSON? 'Natural History Division, The Manitoba Museum, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2 Canada *Present address: Wildlife and Ecosystem Protection Branch, Manitoba Conservation, Box 24, 200 Saulteaux Crescent, Winnipeg, Manitoba R3J 3W3 3Geography Department, University of Winnipeg, 515 Portage Avenue, Winnipeg, Manitoba R3B 2E9 Canada ‘Dubois, John E., and Kimberly M. Monson. 2007. Recent distribution records of the Litthe Brown Bat, Myotis lucifueus, in Manitoba and northwestern Ontario. Canadian Field-Naturalist 121(1): 57-61. ‘Until recently, the distribution of the Little Brown Bat (Myotis lucifugus) in Manitoba and northwestern Ontario was poorly ‘documented. Since 1988, we have been banding and recapturing little browns throughout Manitoba and adjacent Lake of the Woods region in Ontario. All known hibernacula in the study area are recorded here for the first time, along with time of emer- gence. Connections between some hibernacula and summer nurseries are verified by band returns, ranging from 37 to 540 km. We began our study of the Little Brown Bat (Myotis lucifugus) in 1988. The objectives of the study were: (1) to map the distribution of Little Brown Bats in ‘Manitoba; (2) to find as many hibernacula and summer nursery roosts as possible; (3) to learn where bats found in hibernacula spend the summer; and (4) where bats found in summer roosts spend the winter. An overriding objective has been to encourage awareness and conser- vation of bats among the public and responsible agen- cies in Manitoba. All work was done under Manitoba Conservation “Wildlife Scientific Research”, “Ecologi- cal Reserves Research”, and Ontario Ministry of Natural ‘Resources (Kenora District) permits issued to Dubois. _ Little previous research had been done on the six “species of bats known to occur in Manitoba (Banfield 1974). Robert Barclay worked on some species and made incidental observations of occurrences and migra- tion dates of little browns at the University of Manitoba | ar Marsh Field Station from 1981 to 1983 (Barclay 1984). Barclay (1993) stressed “Even for species that are well studied elsewhere, such as the Little Brown ‘Bat, information about the biology of prairie popula- ‘tions is scarce. More than anything else, it is this lack ‘of information that hinders conservation efforts for /prairie bats.” This study is meant to aid conservation | efforts by filling gaps in our knowledge. A. L. Rand (1948) reported sightings of the Little Brown Bat in the Flin Flon district. J. Dewey Soper (1961) reported the first specimens taken at Max Lake, ‘in Turtle Mountain Provincial Park, 20 August 1949. Little browns were reported from the Delta area (Tam- -sitt 1962). In 1978 the late Walter Cook of Grand Rapids showed provincial biologist Bill Koonz a cou- ' ple of caves north of the town that over-wintered bats _(Koonz, Manitoba Conservation, personal communica- | tion). Since 1988 systematic searching for caves with the Speleological Society of Manitoba (SSM) has pro- ‘Key Words: Little Brown Bat, Myotis lucifugus, distribution, Manitoba, northwestern Ontario. duced more information on the Litthke Brown Bat in Manitoba. Soliciting the public and various agencies to report caves, summer bat roosts and sightings has also been very productive. Methods We banded Little Brown Bats to monitor their distri- bution and seasonal movements. Plastic coloured bands were used in the first year of banding, 1988. Each cave or location was identified by a different colour. For more precise tracking of individual bats and their movements, numbered aluminium bands (National Band & Tag Co., Kentucky), size 2 were used in 1989 and thereafter. Hibernacula were located with the aid of the consid- erable efforts of the members of the Speleological Soci- ety of Manitoba and other volunteers. Starting from reported hibernacula sites, the authors and other mem- bers of the SSM used a combination of surficial geology, topographic maps, and air photos to identify likely areas for methodical ground searches. Karst features such as caves, sinkholes and trenches were detected on air pho- tos in areas where limestone, dolomite and gypsum bed- rock are near the surface. These areas were then visited and searched in a grid pattern, using map and compass, and latterly, hand-held global positioning system (GPS) units. Information provided by home and cottage owners, Manitoba Department of Conservation, Ontario Min- istry of Natural Resources, and other agencies led to the banding of bats in residential and cottage summer nursery colonies throughout the province, spilling over into adjacent northwestern Ontario around the north end of Lake of the Woods. Greatest frequency of banding occurred in caves in the Grand Rapids and Gypsumville areas, and in one large cave (St. George Cave) north of Hodgson. [See map, Figure 1] The karst features, where the caves are found in gypsum and dolomite formations of these areas, are SV/ 58 THE CANADIAN FIELD-NATURALIST described in detail in the field reports of the Manitoba Mines Branch for 1988, 1989 and 1990 (e.g., Sweet et al. 1988) and in McRitchie and Monson (2000). We used standard techniques to capture and band the bats (Kunz and Kurta 1988). We captured them in caves by plucking the torpid bats carefully by hand Vol. 121 from the cave wall, and placing them in a cloth bag. We live-trapped Little Brown Bats with a harp trap (Tut- tle 1974). Once captured, the bats were examined to determine sex, reproductive status, age, injuries (both healed and fresh), and, on a cursory basis, external par- asites present. According to custom (Kunz and Kurta TABLE |. Hibernacula banding locations in Manitoba and northwestern Ontario for Little Brown Bats. Location Cook’s Cave [GR] Dale’s Cave [GR] Firecamp Cave [GR] Fold Cavern [Gyp] Iguana Crypt [GR] Longcrawl Cave [Gyp] Microwave Cave [GR] Squeaky Cave [GR] St. George Cave [Hod] Stormcloud Cave [Gyp] Richard Lake, Ont. [LW] Abyss Cave [GR] Okaw Cave [GR] [GR] = Grand Rapids area; [Gyp] = Gypsumville area; [Hod] = Hodgson area; [Wh] = Whiteshell area; [LW] = Lake of the Woods TABLE 2. Summer nursery colony banding locations in Manitoba and northwestern Ontario for Little Brown Bats. Location Brereton Lake, Manitoba Caliper Lake Provincial Park, Ontario Deleau, Manitoba French Portage Narrows, Ontario Grand Rapids, Manitoba Jackfish Lake, Manitoba Keewatin, Ontario Melita, Manitoba Moosehorn, Manitoba Pebble Beach, Manitoba Rush Bay, Lake of the Woods, Ontario Rushing River Provincial Park, Ontario Schist Lake, Manitoba Sherridon, Manitoba Wasagaming, Manitoba Wekusko Falls, Manitoba Latitude Longitude 53°26'10"N 99°32'10" W 53°39'32"N 99°26'19" W B29 25NN 99°20'50" W 51°51'02"N 98°29'30"W 53°26'14"N 99°30'58" W 51°47'10"N 98°37'00"W 53°34'05"N 99°26'25"W 53°37'42"N 99°20'3 1" W 51°36'07"N 97°24'52"W 51°50'28"N 98°29'40" W 49°50'54"N 94°41'30"W 53°42'55"N 99°25'40"W 53°51'53"N 100°07'12"W Latitude Longitude 49°54'16"N 95°32'40"W 49°03'42"N 93°54'46"W 49°34'5S2"N 100°34'3 1" W. 49°26'58"N 94°40'14"W 53°12'30"N 99°18'00" W 50°18'05"N 100°02'10"W 49°45'33"N 94°33'20"W 49°16'05"N 100°59'45"W SS W29EN 98°25'22"W 50°59'34"N 98°26'53"W 49°39'55"N 94°53'28"W 49°41'13"N 94°13'44"W 54°39'45"N 101°48'19"W 55°07'26"N 101°05'12"W 50°39'30"N 99°58'07" W 54°47'22"N 99°58'21"W TABLE 3. Additional Little Brown Bat locations in Manitoba and northwestern Ontario verified by the authors. Location Souris, Manitoba Reston, Manitoba Minnedosa, Manitoba Falcon Lake, Manitoba Deception Lake, Ontario Setting Lake, Manitoba Woodridge, Manitoba vicinity of Oak Lake, Manitoba Kerr’s Lake, Manitoba Pointe du Bois, Manitoba Otter Falls, Manitoba Taiga Biological Station, Wallace Lake, Manitoba Winnipeg, Manitoba Latitude Longitude 49°37'02"N 100°15'37"W 49°33'27"N 101°05'36"W 50°14'43"N 99°50'34" W 49°41'11"N 95°14'54"W 49°44'27"N 94°50'33" W 55°04'11"N 98°29'46"W 49°17'02"N 96°08'52" W 49°40'17"N 100°37'34"W 50°30'00"N 99°40'57" W 50°18'03"N 95°33'05"W 50°08'48"N 95°48'38"W 51°02'40"N 95°20'40""'W 49°53'04"N 97°08'47"W DUBOIS AND MONSON: RECORDS OF LITTLE BROWN Bat 59 wr. FA . a f Dale’s Cave ‘owave Fire Camp Cave Lake » Winnipeg Dauphin : Brandon’ \_~ Winnipeg > LY Turtle Mountain \ Prov. PArk MINNESOTA aa NORTH DAKOTA 98° \ 96° Vine ees Ficure 1. Map of Manitoba showing the portion of the Boreal Ecozone and major settlements. Circles are settlements, triangles are other localities (see text). 60 THE CANADIAN FIELD-NATURALIST 1988), we banded males on the right forearm and fe- males on the left. Those bats found to be previously banded were duly recorded and injuries and/or para- sites noted. Time in capture and handling was kept to a minimum to avoid stressing the animals. Banding locations are summarized in Tables | and 2, below. Many of the Manitoba caves containing bats were mapped, the annual maximum and minimum tempera- tures recorded (average +5°C) and any other vertebrate or invertebrate inhabitants noted, by us, other SSM members, and by staff of the Manitoba Museum and the Manitoba Geological Survey. These details and more on the caves and surrounding landscapes are thor- oughly described in McRitchie and Monson (2000). Results To date we have banded more than 9000 Little Brown Bats. J. Dewey Soper (1961) speculated that Little Brown Bats “probably extend almost to the Hud- sonian Zone’, now called the Boreal Shield ecozone (Ecological Stratification Working Group 1995), while van Zyll de Jong (1985) stated: “It is found from the Atlantic to the Pacific coast and occurs to the limits of the boreal forest.” In addition to those locations where banding has taken place (Tables 1 and 2), we have verified the pres- ence of this species throughout southern Manitoba and at least to the latitude of Taiga Biological Station, Uni- versity of Manitoba, Wallace Lake (51°01'50"N), east of Lake Winnipeg (Table 3). On the west side of the province, Little Brown Bats occur to the latitude of Sherridon (55°07'26"N), and in north-central parts of the province to Setting Lake (55°04'11"N) (Table 3) — locations slightly north and just south of Herb Lake (reported in van Zyll de Jong 1985, page 70). Similar to many species across Canada, a combina- tion of degree-days and night-length likely constrains their range in the north — see discussions in Holroyd et al. (1994), Nagorsen and Brigham (1993), Humphries et al. (2002). A maternity roost of Little Brown Bats was reported recently at 58°06'36"N on the Alaskan coast (Parker and Cook 1996), as was one at Squanga Lake, Yukon at 60°28'40"N (Jung and Slough 2005) . Sherridon (55°07'26"N) is the site of the northern- most nursery colony we have found in Manitoba to date, while Okaw Cave (53°51'53"N) is the northern- most hibernaculum. Individual Little Brown Bats have been reported during summers from as far north as Thompson (55°44'36"N). Little browns have been re- ported to us as showing up at summer nursery colonies in central and southern Manitoba as early as the last week in April, and more often, the first week of May. However, we routinely find large numbers of a given hibernaculum still in torpor when entering the caves in the Grand Rapids area on the Victoria Day long week- end in May (normally the third weekend of the month, range 17-28 May). Band returns from this study have shown little browns are very loyal to both their summer and winter Vol. 121 roosts, with very low incidences of transference (>1%). Recoveries from subsequent, different roosts ranged from 5 km to 533 km distant from point of original banding. Seasonal migration distance from hibernac- ula to summer nursery colony for adult females ranged from 37 km to 540 km. Details of inter-roost and sea- sonal movements will be published separately. We started banding Little Brown Bats around Lake of the Woods (mainly in the Kenora area) in 1993 to learn whether those populations use Manitoba caves to hibernate. Band returns demonstrate that at least some of those females forming nursery colonies in that region definitely use central Manitoba caves to over-winter, as well as one nearby abandoned mine. It has been spec- ulated for some time that abandoned mines in the Keno- ra area have served as hibernacula (Nagorsen 1980). In July 1995, we were directed to an abandoned mine at Richard Lake, 37 km east of Kenora and, upon re- turning in late September, found it to contain at least 250 Little Brown Bats, none previously banded. In the Richard Lake hibernaculum in subsequent years we recaptured banded bats from nursery colonies on Lake of the Woods (Keewatin, French Portage Narrows, Rushing River Provincial Park), as well as bats previ- ously banded in Manitoba hibernacula as far as 533 km away (St. George, Iguana Crypt and Microwave caves). Conservation This study has verified that at least 11 caves in Mani- toba and one abandoned mine in northwestern Ontario serve as hibernacula for Little Brown Bats (Table 1). Bats of many species are at their most vulnerable at roost sites, particularly hibernacula (Brigham 1993). The largest hibernaculum discovered to date in Man- itoba, by several orders of magnitude, is St. George Cave, 200 km north of Winnipeg. More than 20 000 lit- tle browns spend each winter in that cave. This critical cave and a small area around it was declared an ecolog- ical reserve by the Province of Manitoba on 15 March 1997, after several years of effort by Willard Anderson, the authors, and other members of the Speleological Society of Manitoba. This is the most stringent legal protection a natural feature can get in this province. The Province of Manitoba, on 15 January 1996 official- ly declared six species of bats to be “a wild animal” under The Wildlife Act (by the Designation of Wild Animals Regulation MR 3/96), as a first step in their protection. As of December 1997, Manitoba Depart- ment of Natural Resources (now Conservation) has issued guidelines to direct all activities occurring on Crown land around the known over-wintering caves (Asmundson and Larche 1996*). Acknowledgments We thank our employers, the Manitoba Museum and the University of Winnipeg, for their support during the majority of the time period of this study. Additional funding was granted by the Manitoba Museum Foun- dation Fund. We were also substantially assisted by the 2007 staff of Manitoba Department of Conservation (Natural Resource Officers, Operations Branch, particularly of the Grand Rapids and Gypsumville offices, as well as Parks and Natural Areas Branch), and Manitoba Geological Survey, Department of Industry, Trade and Mines. Thanks to Tolko Industries Ltd. and the hos- pitable people of Grand Rapids and Gypsumville. Many family members and friends have worked with us for many hours. We would like to particularly thank Dale Brown and David Wright. Other volunteer banders and helpers over the years include Al Bisset, Melanie Dubois, Michelle Bell, Liza McClintock, Marie Voth, Lori Bilecki, Christy McDonald, Marg Rubin, Gloria Goulet, Cecile Foster, Janis Klapecki, Dennis Peristy, Kim Ottenbreit, Victoria Shemeliuk, Mike Claussen, Justin Harrison, Donna Kurt, Rick Nash, Paul Buisson, Robert Senkiw, Jocelyn Hildebrand, Marianne Setliff, Chris Caslake, Bob Gill, Darren Keam, Amy Barker, Lidi Kuiper, Lyndon Kivi, Kevin Campbell, Christine Abraham, John Fitzmaurice, Lisa Lagerge and Trapper John Christie. We thank Jim Duncan and Robert Nero for reviewing an earlier draft of this paper. Documents Cited (marked * in text) Asmundson, J. D., and R. A. Larche. 1996. Bat Hibernacula Management Guidelines. (unpublished manuscript) Wild- life Branch, Manitoba Department of Natural Resources. 31 pages. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. National Museum of Natural Sciences. University of Toronto Press. 438 pages. Barclay, R. M. 1984. Observations on the migration, ecology and behaviour of bats at Delta Marsh, Manitoba. Canadian Field-Naturalist 98: 331-336. Barclay, R. M. 1993. The biology of prairie bats in Proceed- ings of the Third Prairie Conservation and Endangered Species Workshop. Edited by G. L. Holroyd, H. L. Dick- son, M. Regnier, and H. C. Smith. Provincial Museum of Alberta, Natural History Occasional Paper Number 19. Brigham, R. M. 1993. The implications of roost sites for the conservation of Bats in Proceedings of the Third Prairie Conservation and Endangered Species Workshop. Edited by G. L. Holroyd, H. L. Dickson, M. Regnier, and H. C. Smith. Provincial Museum of Alberta, Natural History Occasional Paper Number 19. Ecological Stratification Working Group. 1995. A National Ecological Framework For Canada. Agriculture and Agri- Food Canada, Research Branch, Centre for Land and Bio- DUBOIS AND MONSON: RECORDS OF LITTLE BROWN BAT 6) logical Resources Research and Environment Canada State of the Environment Directorate, Ecozone Analysis Branch, Ottawa/Hull. Holroyd, S. L., R. M. R. Barclay, L. M. Merk, and R. M. Brigham. 1994. A survey of the bat fauna of the dry interi- or of British Columbia. Wildlife Working Report Number WR-63, Wildlife Branch, Ministry of Environment, Lands & Parks, B.C. Humphries, M. M., D. W. Thomas, and J. R. Speakman. 2002. Climate-mediated energetic constraints on the dis- tribution of hibernating mammals. Nature 418: 313-316 Jung, T. S., and B. G. Slough. 2005. Mortality of Little Brown Bats, Myotis lucifugus, in a rodent trap in the boreal forest Canadian Field-Naturalist 119: 589-590. Kunz, T. H., and A. Kurta. 1988. Capture methods and holding devices. Pages 1-28 in Ecological and behavioral methods for the study of bats. Edited by T. H. Kunz. Smith- sonian Institution Press, Washington, D.C. 533 pages. McRitchie, D. W., and K. M. Monson. 2000. Caves and Karst of Manitoba’s Interlake Region. Speleological Soci- ety of Manitoba, Second Edition. Winnipeg. 181 pages. Nagorsen, D. W. 1980. Records of hibernating Big Brown Bats (Eptesicus fuscus) and Little Brown Bats (Myotis lucifugus) in northwestern Ontario. Canadian Field-Nat- uralist 94: 83-85. Nagorsen, D. W., and R. M. Brigham. 1993. The Bats of British Columbia. Volume 1: The Mammals of British Columbia. Royal British Columbia Museum Handbook. UBC Press, Vancouver. Parker, D. I., and J. A. Cook. 1996. Keen’s Long-eared Bat, Myotis keenii, confirmed in southeast Alaska. Canadian Field-Naturalist 10: 611-614. Rand, A. L. 1948. Mr. W. H. Bryenton’s notes on Manitoba mammals of the Herb Lake-Flin Flon area. Canadian Field- Naturalist 62: 140-150. Soper, J. D. 1961. The mammals of Manitoba. Canadian Field-Naturalist 75: 171-219. Sweet G., P. Voitovice, and W. D. McRitchie. 1988. Karst Investigations in Palaeozoic Carbonates of the Grand Rapids Uplands and Southern Interlake. Geological Paper GP88-1, Province of Manitoba, Mines Branch. Tamsitt, J. R. 1962. Mammals of the Delta Marsh region of Lake Manitoba, Canada. Canadian Field-Naturalist 76: 71- 76. Tuttle, M. D. 1974. An improved trap for bats. Journal of Mammalogy 55: 475-477. van Zyll de Jong, C. G. 1985. Handbook of Canadian mam- mals. Volume 2, Bats. National Museum of Natural Sci- ences, Ottawa, Ontario. Received 12 August 2003 Accepted 2 October 2007 Immobilization of Elk, Cervus elaphus, with Telezol and Xylazine and | Reversal with Tolazine or Yohimbine RICK ROSATTE Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, DNA Building, 2140 East Bank Drive, Peterborough, Ontario K9J 7B8 Canada; e-mail: rick.rosatte @ ontario.ca. Rosatte, Rick. 2007. Immobilization of Elk (Cervus elaphus) with telezol and xylazine and reversal with tolazine or yohim- bine. Canadian Field-Naturalist 121(1): 62-66. During January 2003 and March 2005, a Telazol®/xylazine mixture was used to immobilize 4 free-ranging Elk in Ontario, Canada. A dosage of 3.3-3.6 mg/kg of Telazol® and 1.7-2.0 mg/kg of xylazine proved to be effective for the rapid immobi- lization of Elk. Induction time for those dosages was as short as 3-4 minutes. The advantage of using Telazol® is that only small volumes (3-4 ml) are needed to immobilize Elk-sized animals. In addition, Tolazine® and yohimbine both proved to be effective antagonists for xylazine with recovery times of 8 to 15 minutes when administered at dosages of 3.3-3.6 and 0.08-0.14 mg/kg, respectively. The use of oxygen proved to be effective for treatment of hyoxemia in Elk immobilized with a Telazol®/xylazine mixture. The immobilization procedures and the drug and antagonist dosage information will be useful to researchers planning to capture free-ranging Elk for activities such as radio-collaring and blood sampling. Key Words: Cervus elaphus, Elk, antagonist, immobilization, reversal agent, Telazol®, Tolazine, Yohimbine, Ontario. During 1998 — 2001, 443 Elk (Cervus elaphus), also known as American Elk, or Wapiti, were transported from Elk Island National Park (EINP), Alberta, and released in four areas of Ontario, Canada (Bellhouse and Rosatte 2005; Rosatte et al. 2007). One of the release sites, near Bancroft, Ontario (approximately 44°5'N, 77°30'W), received 120 Elk during 2000 and 2001. All Elk were aged and weighed at EINP during processing, which included ear-tagging and fitting the Elk with telemetry collars (VHF and GPS) (Rosatte et al. 2002). The weights of Elk at EINP were used as guidelines to estimate the weight of Elk in Ontario for drug dosage calculations. As ten of the Elk were fitted with GPS collars during processing at EINP, some of the animals had to be immobilized to retrieve the collars and download the data. On 16 January 2003 (-20°C), an 8 year old, free- ranging adult cow Elk (ear tag 341), previously fitted with a GPS collar (148.188 Mhz) at EINP, Alberta, was immobilized with Telazol® (tiletamine hydrochloride and zolazepam hydrochloride) (Fort Dodge Animal Health, Fort Dodge, Iowa) and AnaSed (xylazine hydrochloride) (Vet-A-Mix, Shenandoah, Iowa) near Bancroft, Ontario, for the purpose of collar removal. The drug dosage (target dosage was about 3 mg/kg Telazol® and approximately 1.5 mg/kg of xylazine) was calculated using an estimated weight for the Elk of 236 kg — her weight during processing at EINP in January 2001. The immobilizing drug mixture was pre- pared by injecting the desired volume and weight of xylazine into a bottle containing lyophilized Telazol®. The Telazol®/xylazine mixture was then drawn off with a syringe and injected into the immobilization dart. The dart was then plugged with vasoline to prevent the leakage of drugs from the dart. A Model 193 Pneu-Dart 50 caliber rifle and 4x scope (Pneu-Dart Inc., Williams- port, Pennsylvania) with a yellow CCI 22 caliber brass power charge (Omaha Industries, Lewiston, Idaho) | and a diffuser setting of 4, was used to project a 50 cal- iber, 4 cc aluminum type C dart with a 3 cm long barbed needle (Pneu-Dart Inc., Williamsport, Pennsylvania), that contained the Telazol®/xylazine mixture, into the hind leg musculature of the Elk. The distance of the shot was about 35 m. As soon as the animal was in a lateral recumbent position and there was no evidence ~ of leg or eye reflex, she was moved into a sternal re- cumbent position and the head was elevated to mini- mize the chance of bloat and regurgitation of rumen contents. Ophthalmic ointment was applied to the eyes of the immobilized Elk to prevent drying. The respiration rate of the cow Elk was about 18/min at 30 min post-darting. At 36 min post-darting, the res- piration rate increased to 54/min and breathing was very shallow. At this point, bloat was evident and the gum/ tongue area was blue in color (indicative of low oxygen content in the blood). The animal was admin- istered oxygen orally as a treatment for hypoxemia, as well as rolled side to side in an effort to relieve the bloat. The head was elevated and lowered as well to stimulate belching. At 50 min post darting, respiration | rate decreased to 42/min and the tongue/gum area returned to a pinkish color following the administra- - tion of oxygen. Hibitane antibacterial veterinary oint- - ment (1% chlorhexidine acetate) (Ayerst, Guelph, On- tario) was applied to the dart wound. After the animal was processed an intramuscular injection of Tolazine (tolazoline hydrochloride, Lloyd Laboratories, Shenan- doah, Iowa) was administered in the right leg muscu- lature to speed recovery. The concentration and dosage of drugs as well as the induction, down and recovery times are shown in Table 1. After the animal was ina standing position, she orally expelled gas for several © 62 2007 ROSATTE: IMMOBILIZATION OF ELK 63 seconds and the bloat decreased dramatically. The Elk was monitored for the next few days and no adverse co effects were noted. She was still in good health when 3 5 we w.8 observed near Bancroft, Ontario in April 2005. Z Bs gZal4AR== On 19 February 2004 (-6°C), a 10.5 year old cow £ Bagalezza Elk (ear tag 360) was immobilized near Bancroft, On- es weak tario, to remove a GPS collar (149.560 Mhz). The _ = weight of the animal at EINP during January 2001 was i 234 kg. The animal was darted with a mixture of Tela- i ee ime zol”/xylazine as described above using a Pneu-Dart 5 z E atl ee a eee rifle, with a yellow 22 caliber power charge and a dif- = zg “Ss fuser setting of 3, and a 4 cc dart. The distance of the 5 shot was about 25 m with the dart hitting the right upper z hind leg musculature. As the drug took effect the Elk 2 ee. a assumed a recumbent position beginning with the hind 3 5 E = $RHRF end first (Figure 1). Some bloat was evident so oxygen A AT} was administered for treatment of hypoxemia and the z animal was rolled from side to side and the head elevat- @ a ed to expel gases (Figure 2). Respiration rate increased g PIN Son ae ae Ps from about 14/min to 70/min when bloating occurred. 2 | = =| ae i > The gums and tongue remained pink colored through- S =) E = out the handling period. When processing was com- fe S s plete, Tolazine was injected intramuscularly to ® Cie ye 3 reverse the effects of xylazine and speed recovery = b Z eee = 5 from the drug. Induction, immobilization and recoy- 5 25a g/9NSS 3 = ery times are noted in Table 1. || 2 = S On 11 January 2005 (-17°C ambient temperature), = = = = a yearling bull Elk, born in Ontario, suspected of being A oe & S = fc infected with meningeal worm (Parelaphostrongylus 2 2 = fBalooa aes tenuis), was immobilized with an intramuscular 5 22 El sor (Begs injection of Telazol® and xylazine hydrochloride for the e er ERES% purpose of collecting a blood sample. The dosage of z f Z7se drug was calculated using mean weights of yearling 3 —q |eo wioe 8 oe bull Elk at EINP. As the Elk had no fear of humans, = pag =l| spose = 2 oe a 25 possibly due to the effects of P. tenuis, the drugs were = Salss S S of Slt administered intramuscularly in the right upper hind 3 Flare = 2885 limb area via a 5 cc sterile syringe (Burron Medical Z Oba 3 5 Products, Bethlehem, Pennsylvania, USA) and 22 g = eS ete oN 255 (4 cm) needle (Terumo, Belgium) attached to a jab = 26 SRTeonyss =a) = stick made from a piece of copper tubing. When pro- > 2S Sa SSaalsus 225 cessing was complete, which included radio-collaring 8 AaX S| aaaalo 2 ass (148.412 Mhz) and ear-tagging (#139) (as reported in 3 a5 252 McIntosh et al. 2007), the Elk was injected intramus- S Ham ig gpergelll eS 2 SUS ew cularly in the hind limb musculature with Yobine (yo- 2 Sora aa cess = Mess ES himbine hydrochloride) (Lloyd Laboratories, Shenan- S SeSE/SSssii<2e eee doah, Iowa, USA) as a reversal agent for xylazine. Drug 5 ES re PaO fa Ee eo dosages, induction, immobilization and recovery times : =o = = 2 = 5 are presented in Table 1. While immobilized, respira- aa r eSs5ce3 tion rate for the yearling bull Elk was 14-16/min, there i Helunooll SE Ese = was no evidence of bloat, and the gums and tongue 8 2 S|SASS sR wee a's remained pink in color. E s SE Ex On 30 March 2005 (+5°C ambient temperature), a a v. Sgetecs 2% year old bull Elk (born in Ontario) was immobilized 22 Sel RR SS| PS sss gz with Telazol® and xylazine for the purpose of de-antler- 2S Sosy te Ba otees ing as the animal had broken the pedicle in the fall of oo S BY oe Be 2004. The drugs were administered as described above alee = = E i: > 3 using a Pneu-Dart rifle, with a green 22 caliber power = z oss S <= Sseacs charge, a diffuser setting of 5, and a 4cc dart. The dis- agi 2 SSeS 2 53 2 ys tance of the shot was about 30 m with the dart hitting a 3a Se She a ss 64 tne a be THE CANADIAN FIELD-NATURALIST Vol. 121 Ficure |. A GPS collared adult cow Elk (ear-tag 360) exhibiting hind limb paralysis 2 minutes after being darted with a mixture of Telazol® and Xylazine. (Photo by R. Rosatte). the right upper hind leg musculature. Respiration rate was stable at about 12/min during processing (de- antlered and ear-tagged — #141) and the gums and tongue remained pink colored with no evidence of hypoxemia or bloat. A 3 cc intramuscular injection of Dystosel (Vitamin E — 136 IU/mL and selenium — 3 mg/mL) (Pfizer Canada Inc, London, Ontario) was administered as a preventative measure against cap- ture myopathy. When processing was complete, Yobine was injected intramuscularly to speed recovery from the effects of xylazine. Induction, immobilization and recovery times are shown in Table 1. Discussion A dosage of 3.3-3.6 mg/kg (body weight) of Tela- zol® and 1.7-2.0 mg/kg of xylazine proved to be an effective drug combination for the rapid immobiliza- tion of four free-ranging Elk in Ontario. Induction time or the time from darting until the animal was prone for those dosages was 3-4 minutes. This calcu- lation did not include the Elk for which induction time was noted as 10 minutes, as that animal was not immediately located in the forest. Similarly, Mills- paugh et al. (1995) reported a mean induction time of 4.6 minutes when Elk were immobilized with 2.5 mg/kg Telazol® and 0.3 mg/kg xylazine. The advantage of using a Telazol®/xylazine mixture over a drug combi- nation such as ketamine hydrochloride (at a commer- cially available concentration 100 mg/mL)/xylazine is that a much lower volume of Telazol is required to at- tain a state of immobilization. For example, only a 4 mL (800 mg of Telazol® reconstituted in 3.5 mL of xyla- zine) volume of drug was required to immobilize a 225 kg cow Elk. About 15 mL of a ketamine hydrochlo- ride/xylazine hydrochloride (100 mg/mL ketamine) mixture was required to immobilize a bull Elk in On- tario during 2003. The advantage of a smaller volume of Telazol®/xylazine mixture is that a much smaller dart can be used (4 cc compared to a 10 or 15 cc) to administer the drugs, which means less weight and less damage to the animal when the dart is administered via an immobilization gun. A smaller dart (4 cc) has a better trajectory and is more accurate at longer dis- tances (> 65 m) than a larger/heavier dart (10-15 cc). In addition, as the induction time for ketamine in Elk is generally longer than that for Telazol®, animals tend to move farther before being completely immobilized (Golightly and Hofstra 1989; Miller et al. 2004), which could result in problems locating the animal in heavi- ly forested areas. 2007 ROSATTE: IMMOBILIZATION OF ELK 65 FIGURE 2. Oxygen being administered to the immobilized cow Elk (ear tag 360) for treatment of hypoxemia (Photo by R Rosatte). One drawback with using a Telazol®/xylazine mix- ture for the immobilization of Elk is the extended re- covery time which could lead to decreased survival. In one study, an Elk immobilized with a Telazol®/xylazine mixture was not administered an antagonist and was immobilized for 5 hours (Golightly and Hofstra 1989). As well, there is no known antagonist for the tiletamine component of Telazol® (Miller et al. 2004). Tolazine and yohimbine both proved to be effective antagonists for xylazine with recovery times of 8 to 15 minutes when administered at dosages of 3.3-3.6 and 0.08- 0.14 mg/kg (body weight), respectively. However, the key to rapid recovery was administering the antagonist 45-60 minutes (or longer) post -immobilization. This allowed sufficient time for Telazol® to be metabolized and diminish side-effects after injection of the rever- sal agent. Animals may stagger for some time due to the effects of Telazol® if the antagonist is adminis- tered too soon after immobilization as the antagonist only neutralizes the effects of xylazine. Millspaugh et al. (1995) reported a mean recovery time of 14 min- utes when yohimbine was administered intravenous- ly to reverse xylazine administered to Elk at a dose of 0.3 mg/kg (in a mixture with Telazol® at a dose of 2.5 mg/kg). However, a recovery time in that study of 125 min was reported when yohimbine was given intra- muscularly. Hypoxemia or deficient oxygenation of the blood may occur during the immobilization of wild rumi- nants. This can lead to organ failure and capture myopa- thy (Read et al. 2001). Bloating occurred in two of the Elk immobilized in this study despite their being in a sternal recumbent position. As well, respiration rate was high in those two Elk {normal respiration rate for elk is about 13 breaths/min (Hudson and Haigh 2002)}. Bloat is a result of gas production in the rumen, which may result in pressure being applied to the diaphragm (due to the weight of the abdominal viscera on the diaphragm) with resultant respiratory/circulatory prob- lems and oxygen depletion in the blood. Bloating in this study may have been aggravated due to the ani- mals foraging at feeding stations (with alfalfa) prior to darting. Oxygen quickly relieved the symptoms of hypoxemia (blue tongue, lips), and the suggested rate of oxygen administrated nasally for preventing hypox- emia was 10 L/min for a period of about 5 minutes (Read et al. 2001). The four immobilized Elk were later (1 month to 2 years) located by radio-telemetry and had recovered fully, with no observable side effects from immobiliza- 66 THE CANADIAN FIELD-NATURALIST tion or handling. Capture procedures, drugs and antag- onist data outlined above will be useful to researchers planning to capture free-ranging Elk. For “off-label” use of the above drugs on wild animals such as Elk, an “Emergency Drug Release Authorization” must be approved by the Veterinary Drugs Directorate, Health Canada, and animals immobilized with agents such as Telazol® are not to be used as food items. Acknowledgments The Ontario Elk Restoration program is supported by the Provincial Elk Technical Team and the Ontario Ministry of Natural Resources (OMNR), Wildlife Sec- tion, and the Wildlife Research and Development Sec- tion (WRDS). Thanks to Tom Simpson and Mike Scafie, OMNR, Bancroft District; John O’ Donnell, Barry Wannamaker, Ryan Cavanaugh, Jason Langis, Bancroft Elk Program; Jim Young, Provincial Elk Pro- gram Coordinator; and Terese McIntosh and Arthur Dupuis, Trent University, for assisting with the immo- bilization and processing of the elk. Special thanks to Joe Neuhold for all of his assistance and dedication to the Elk Program. Thanks to Norm Cool and Rob Kay, Parks Canada, EINP, Alberta, and Bruce Chisholm, Canadian Food Inspection Agency, Edmonton, Alberta, for weighing and aging the elk during processing at EINP. J. C. Davies and Mike Allan, OMNR, WRDS, re- viewed the manuscript and provided helpful comments. Literature Cited Bellhouse T., and R. Rosatte. 2005. Assessment of the poten- tial for negative interaction between re-introduced elk (Cer- vus elaphus) and resident white-tailed deer (Odocoileus virginianus) in their wintering areas in Ontario, Canada. Mammalia 69 (1): 35-56. Vol. 121 Golightly R., and T. Hofstra. 1989. Immobilization of elk with a ketamine-xylazine mix and rapid reversal with yo- himbine hydrochloride. Wildlife Society Bulletin 17: 53- 58. Hudson R., and J. Haigh. 2002. Physical and physiological adaptations. Pages 199-257 in North American elk, ecology and management. Edited by D. Toweill and J. W. Thomas. Smithsonian Institution Press, Washington, D.C. McIntosh, T., R. Rosatte, D. Campbell, K. Welch, D. Fornier, M. Spinato, and O. Ogunremi. 2007. Evidence of Parelaphostrongylus tenuis infections in free-ranging elk (Cervus elaphus) in southern Ontario. Canadian Veteri- nary Journal 48: 1146-1154. Miller B., L. Muller, T. Doherty, D. Osborn, K. Miller, and R. Warren. 2004. Effectiveness of antagonists for tileta- mine-zolazepam/xylazine immobilization in female white- tailed deer. Journal of Wildlife Diseases 40: 533-537. Millspaugh J., G. Brundige, J. Jenks, C. Tyner, and D. Hustead. 1995. Immobilization of Rocky Mountain elk with Telazol and xylazine hydrochloride, with antagonism by yohimbine hydrochloride. Journal of Wildlife Diseases 31: 259-262. Read M., N Caulkett, A. Symington, and T. Shury. 2001. Treatment of hypoxemia during xylazine-tiletamine- zolazepam immobilization of wapiti. Canadian Veterinary Journal 42: 861-864. Rosatte R., J. Hamr, B. Ranta, J. Young, and N. Cool. 2002. Elk restoration in Ontario, Canada: infectious dis- ease management strategy, 1998-2001. Annals of the New York Academy of Sciences 969: 358-363. Rosatte, R., J. Hamr, J. Young, I. Filion, and H. Smith. 2007. The restoration of Elk (Cervus elaphus) in Ontario, Canada: 1998-2005. Restoration Ecology 15: 34-43. Received 19 January 2006 Accepted 11 January 2008 First Confirmed Breeding of the Marbled Godwit, Limosa fedoa, in Québec FRANCOIS MorNEAU!, BENOIT GAGNON,” and SIDNEY WHISKEYCHAN® '63 rue Champagne, Saint-Basile-le-Grand, Québec J3N 1C2 Canada Corresponding author. Hydro-Québec, Direction Environnement, 855 Sainte-Catherine Est, 9° étage, Montréal, Québec H2L 4P5 Canada; e-mail: gagnon.benoit@hydro.qc.ca > P.O. Box 167, Waskaganish, Québec JOM 1RO Canada Morneau, Francois, Benoit Gagnon, and Sydney Whiskeychan. 2007. First confirmed breeding of the Marbled Godwit, Limosa fedoa, in Québec. Canadian Field-Naturalist 121(1): 67—70 The objectives of this study were to confirm breeding of the Marbled Godwit (Limosa fedoa) in Rupert Bay, on the Québec coast of James Bay, and to assess population abundance. Nest search was conducted 17—20 June 2003, using several tech- niques. Two nests were found in the high marsh of Cabbage Willows Bay: these are the first confirmed breeding records of this species in Québec. The Rupert Bay region probably holds a breeding population of no more than a few dozen pairs. Cette étude visait a confirmer la reproduction de la Barge marbrée (Limosa fedoa) dans la baie de Rupert située sur la céte est de la Baie James (Québec) et 4 estimer l’abondance de la population. Une recherche de nids a été réalisée entre le 17 et le 20 juin 2003 a l’aide de différentes techniques. Deux nids ont été découverts dans le haut marais de la baie Cabbage Willows. Ils confirment pour la premiére fois le statut nicheur de cette espéce au Québec. La région de la baie de Rupert abrite probablement une population nicheuse d’au plus quelques douzaines de couples. Key Words: Marbled Godwit, Limosa fedoa, nest search, breeding, abundance, James Bay, Rupert Bay, Cabbage Willows Bay, Québec. The Marbled Godwit (Limosa fedoa) breeds only in North America, in three disjunct areas: the grassland of the northern United States and southern Canada, the Alaska Peninsula, and James Bay (Gratto-Trevor 2000). The James Bay area probably holds only 1000-2000 birds (Gratto-Trevor 2000), but the species breeding has been confirmed only in Ontario (Morrison et al. 1976; Peck and James 1983; Morrison 1987). In Québec, the species was first recorded on the James Bay coast by Spreadborough (in Macoun and Macoun 1909), who stated that the species bred on both coasts in 1904, although no direct evidence for breeding was reported. The Marbled Godwit was not recorded again on the east coast of James Bay before 1990. Since then it has been observed during the breed- ing season in and near Rupert Bay (Létourneau and Morrier 1996). Breeding behaviours were noted there in 1990 and 1991, but nesting was not confirmed. In 2003, as part of the Eastmain-1-A and Rupert diversion hydroelectric project environmental studies, we sought to clarify the breeding status of the Marbled Godwit in Rupert Bay, on the Québec side of James Bay. The objectives of this study were to confirm breed- ing of the species in Québec and to assess population abundance. Study Area and Methods Field work was carried out in the coastal wetlands of Rupert Bay, Québec, Canada (Figure 1). These wet- lands are particularly well developed owing to the sandy deposits and flat landscape. They are especially extensive in Cabbage Willows Bay, which opens into Rupert Bay, and Boatswain Bay, which opens directly into James Bay, but are narrower in Hall Cove and else- where. They consist, in sequence from upland to open water, of swamps, fens, high marshes and low marshes (FORAMEC 2004*). The high marsh grounds are somewhat spongy but not covered by water. Low marshes are covered twice daily by the tide but the other wetlands are flooded only by storm tides and equinoc- tial tides. Nest searches were conducted on 17—20 June 2003, using several techniques. The rope-drag method (Klett et al. 1986) was used on 0.3—1.4 km transects in parts of the high coastal marsh of Cabbage Willows Bay and Hall Cove. Nests were also searched for in the same areas by watching birds flying from the low marsh to potential nesting areas or vice versa. In addition, play- backs of taped breeding calls were used regularly on the high marsh and once in the low marsh. Finally, to increase ground coverage, we conducted a helicopter survey in Cabbage Willows Bay, in Hall Cove, over some Tamarack (Larix laricina) fens on Ministikawatin Peninsula, and on Jacob Island, north of Rupert Bay. We searched along 50-m parallel transects, at speeds that varied from 20 km/h to hover, and at altitudes varying from 3 to 9 m. Breeding chronology was estimated assuming a mean egg-laying interval of 2 days and an incubation period of 25 days (Gratto-Trevor 2000). Abundance was estimated with counts of birds performing aerial display and nests. 67 68 THE CANADIAN FIELD-NATURALIST Vol. 121 —78° 45 * ] Boatswain Bay Migratory Bird La bs ‘Bay lleTent'> £ ft 4 Sanctuary i Di. 0 Island Cabbage Willows Bay Biodiversity Reserve == —_ onli Migratory Bird Sanctuary Marbled Godwit nest 6675_hq_146_f1_080115.fh9 ag cd ’ Proposed Boatswain Bay Biodiversity Reserve Rupert Waskagani Ficure |. Location of Marbled Godwit study area and nests. Results The Marbled Godwit nest search was conducted on slightly over 20 km of line transects using the rope- drag method. It covered about 70 ha. Two Marbled Godwit nests, 3.6 km apart, were dis- covered in the high coastal marsh of Cabbage Willows Bay in 2003 (Figure 1). The first nest (51°32.61'N, 79°16.13'W), containing four eggs (Figure 2) and one adult incubating, was found on 17 June using the rope-drag technique. The second nest (51°30.77'N, 79°15.13'W), discovered on 20 June following obser- vation of courtship behaviour, contained only one egg, but two adults were within 30 m of the nest. The first nest was revisited again on 19 June and still contained four eggs. Nests were 5 cm in depth and 20 cm in diameter at the rim. They were mostly, if not completely, made of dry stems of Sweet Grass (Hierochloe odorata). Both nests were found in the largest clumps of that species in the vicinity. The first nest was partly covered by 2007 MORNEAU, GAGNON, and WHISKEYCHAN: MARBLED GOpDWIT 69 FIGURE 2. First Marbled Godwit nest discovered in Rupert Bay, Québec. green stems. Throughout the high coastal marsh, grass cover, about 10—15 cm tall, was relatively sparse, with occasional clumps up to 15—25 cm. In 2003, laying and potential incubation in Cab- bage Willows Bay occurred at least between 11 June and 19 July. No Godwit responded to several playback trials in the high coastal marsh. However, two individuals answered spontaneously to the single trial on the low coastal marsh, by flying and calling around the inves- tigators. Over the same biotope, four different birds were seen engaging in aerial courtship behaviour dur- ing nest searches. Besides the birds associated with the nests, only one Marbled Godwit was seen in the high marsh: it was flying from the low marsh and landed in the high marsh, on a potential nesting area. Despite a long search, it was not seen again. Partial aerial coverage of the Cabbage Willows Bay low coastal marsh enabled us to identify four adults. Another bird was spotted as it flew toward the low marsh during the aerial survey; it seemed to be com- ing from the second nest area. No birds were seen in Hall Cove or in Ministikawatin Peninsula or Jacob Island during aerial surveys. Discussion The two Marbled Godwit nests that we found in Cabbage Willows Bay represent the first definitive breeding evidences of this species in Québec. Some previous observations had led us to believe the species was nesting there. On 17 June 1991, two copulating birds were observed in Cabbage Willows Bay. The same year, three Marbled Godwits were seen on 21 July in the same area, including two agitated birds giving alarm calls, indicating the probable presence of young (Létourneau and Morrier 1996). Finally, a similar ob- servation was recorded on 27 July 2002, again in Cab- bage Willows Bay, north of the Novide River (Hydro- Québec, unpublished data). Breeding habitat in Cabbage Willows Bay is simi- lar to that described elsewhere. In North Dakota, the Marbled Godwit shows a strong preference during breeding season for short, sparse to moderately dense cover along wetland shorelines and shorter vegetation in upland habitats (Ryan et al. 1984). In James Bay, Ontario, one nest was found in a raised, grassy ridge between two ponds, in an open coastal marsh (Peck and James 1983). Estimated breeding chronology suggested that nest search was carried out in the egg-laying and early in- cubation period. This is consistent with the observed copulation (17 June 1991) and other previous records suggesting breeding (see above). The flight displays observed in 2003 also support the breeding chronolo- gy outline. Display behaviour is most frequent early in the breeding season and is commonly performed by 70 THE CANADIAN FIELD-NATURALIST unpaired or nest-scraping males until incubation duties are initiated (Nowicki 1973). In Ontario, two breeding records from James Bay indicate that egg-laying was initiated approximately between 24 May and 31 May (Morrison et al. 1976; Peck and James 1983). In 2003, the Marbled Godwit breeding population included at least four pairs in Cabbage Willows Bay. In 2002, the species was also recorded in Boatswain Bay, Jacob Island and Tent Island, between 23 July and 1 September (Hydro-Québec, unpublished data). However, considering the late dates and that no breed- ing behaviour was observed, it is likely that these birds were migrants rather than breeders. Although no Mar- bled Godwit was recorded during the aerial survey inland of Ministikawatin Peninsula, the species was located there in June 1990 and 1991, in three areas, in- cluding a bird on a shallow lake fen (Létourneau and Morrier 1996). In Ontario, the Marbled Godwit is also found up to 20 km inland (Gratto-Trevor 2000). Thus, the species probably breeds inland of Ministikawatin Peninsula. Its abundance there could exceed that of Cabbage Willows Bay because of the vast expanse of Tamarack fens. On 15 June 1991, 80 Marbled Godwits were counted in Cabbage Willows Bay (Létourneau and Morrier 1996). Their breeding status is question- able. As noted earlier, that date probably coincides with egg-laying and early incubation. Either these birds were on the point of dispersal for local breeding or were migrants on the way to northern James Bay breeding grounds in Ontario. However, if Marbled Godwits fly directly to breeding areas in spring from wintering sites on southeastern U.S. coasts, as the re- verse is presumed to occur in the fall (Morrison et al. 1976), then those birds may have been local or regional breeders. Therefore, the Rupert Bay region probably holds a breeding population of between four pairs and a few dozens pairs. Acknowledgments This study was funded by Hydro-Québec. We are grateful to Francoise Lebrun and Gilles Lefebvre, from Hydro-Québec, for production of the map. We thank Jeffrey Salt, from the Waskaganish Cree community, for his contribution to this study. We are grateful to helicopter pilot Laurie Gagnon for her field assistance. Jean Doucet and Mary Antico provided helpful com- Vol. 121 ments on an earlier draft of the manuscript. A. J. Ersk- ine and an anonymous reviewer provided useful com- ments to improve the manuscript. Document Cited (marked * in text) FORAMEC Inc. 2004. Centrale de l’Eastmain-1-A et déri- vation Rupert. Etude de la végétation et des espéces floris- tiques et fauniques menacées ou vulnérables. Préparé pour la Société d’ énergie de la baie James. Québec. Literature Cited Gratto-Trevor, C. L. 2000. Marbled Godwit (Limosa fedoa). The Birds of North America, No. 492. Edited by A. Poole et F. Gill. The Birds of North America, Inc., Philadelphia. Klett, A. T., H. F. Duebbert, C. A. Faanes, and K. F. Hig- gins. 1986. Techniques for studying nest success of ducks in upland habitats in the prairie pothole region. U.S. Fish and Wildlife Service, Resource Publication (158). Wash- ington, D.C. Létourneau, V., and A. Morrier. 1996. Marbled Godwit. Pages 1138-1139 in The breeding birds of Québec: atlas of the breeding birds of Southern Québec. Edited by J. Gauthier and Y. Aubry. Association québécoise des groupes d’ornithologues, Montréal, Québec. Province of Québec Society for the Protection of Birds, Canadian Wildlife Service, Environment Canada, Québec Region, Montréal. Xvill + 1295 pages. Macoun, J., and J. Macoun. 1909. Catalogue of Canadian birds, 24 edition. Geological Survey of Canada, Depart- ment of Mines, Ottawa. Bulletin (973). 761 pages. Morrison, R. I. G. 1987. Marbled Godwit. Page 528 in Atlas of the breeding birds of Ontario. Edited by M. D. Cadman, P. F. Eagles, and F. M. Helleiner. University of Waterloo Press, Waterloo, Ontario. Morrison, R. I. G., T. H. Manning, and J. A. Hagar. 1976. Breeding of the Marbled Godwit, Limosa fedoa, in James Bay. Canadian Field-Naturalist 90: 487-490. Nowicki, T. 1973. A behavioral study of the Marbled Godwit in North Dakota. MLS. thesis, Central Michigan University, Mount Pleasant, Michigan. Peck, G. K., and R. D. James. 1983. Breeding birds of On- tario: nidiology and distribution, vol. 1: Non-passerines. Life Sciences Miscellaneous Publications, Royal Ontario Museum, Toronto. xii + 321 pages. Ryan, M. R., R. B. Renken, and J. J. Dinsmore. 1984. Marbled Godwit habitat selection in the northern prairie region. Journal of Wildlife Management 48: 1206-1218. Received 23 January 2006 Accepted 18 January 2008 Swift Fox, Vulpes velox, Den Use Patterns in Northwestern Texas BRADY K. MCGEE, WARREN B. BALLARD, and Kerry L. NICHOLSON Department of Natural Resources Management, Texas Tech University, Box 42125, Lubbock, Texas 79409 USA McGee, Brady K., Warren B. Ballard, and Kerry L. Nicholson. 2007. Swift Fox, Vulpes velox, den use patterns in northwestern Texas. Canadian Field-Naturalist 121(1): 71-75. Predator avoidance may be a reason why Swift Foxes (Vulpes velox) are one of the most burrow-dependent canids in North America. Typically Swift Foxes have multiple dens, which they frequently move among. As part of a larger study to reduce Coyote (Canis latrans) related mortalities on Swift Foxes, we installed artificial escape dens in areas occupied by Swift Foxes on Rita Blanca National Grassland, Dallam County, Texas. For this paper, our objective was to determine the effects of artificial escape dens on Swift Fox den use patterns. From January 2002 to August 2004 we captured, radio-collared, and monitored 55 Swift Foxes. We documented annual number of dens used, rate of den use (fidelity), distance between dens, den area, and den sharing. We compared treated (artificial dens installed) and untreated (no artificial dens) areas but found no differences in annual number of dens (P = 0.64; ¥ = 8), rate of den use (P = 0.96; ¥ = 35%), mean distance between dens (P = 0.99; X = 2,311 m), den area (P = 0.55; X = 5.72 km’), or den sharing (P = 0.46; ¥ = 42% of time). We did not observe an effect of artificial escape dens on Swift Fox den use patterns probably because artificial escape dens were designed for . | | | if | 1 \ temporary escape cover rather than diurnal den use. Key Words: Swift fox, Vulpes velox, den range, den sharing, den use, fidelity, Texas. Swift Foxes (Vulpes velox) were once abundant throughout the short and mid-grass prairies of North America but have rapidly declined with expansion of human settlement (Egoscue 1979). Studies have shown that the Swift Fox primarily inhabits areas of native rangeland (Allardyce and Sovada 2003). Much of the historical Swift Fox range has been fragmented into patches of native rangeland, Conservation Reserve Program, and agricultural fields (Allardyce and Sova- da 2003). Habitat loss has been one of the reasons Swift Foxes were temporarily a candidate for endan- gered species listing with the United States Fish and Wildlife Service (USFWS) from 1992 to 2001 (USFWS 1995; USFWS 2001; Allardyce and Sovada 2003). Predation has been another limiting factor on Swift Fox populations (Kamler et al. 2003a). Many studies have shown that Coyotes (Canis latrans) are the pri- mary source of Swift Fox mortality, with annual sur- vival rates ranging from 43 to 53% (Sovada et al. 1998; Kitchen et al. 1999; Matlack et al. 2000; Anderson et al. 2003; Allardyce and Sovada 2003). Because Coy- otes rarely consume Swift Foxes they kill (Sovada et al. 1998; Kitchen et al. 1999; Allardyce and Sovada 2003), Coyote predation seems to be the result of both interference and exploitative competition (Kamler et al. 2003b). Predator avoidance may be one reason why Swift Foxes are one of the most burrow-dependent canids in North America (Moehrenschlager et al. 2004). Dens _ are an integral part of Swift Fox ecology. Swift Foxes use dens year-round not only for protection from pred- { ; ators, but also for reproduction, resting, and avoidance of extreme climatic conditions (Egoscue 1979). In sparsely vegetated habitats occupied by Swift Foxes, dens may constitute crucial escape cover. Arjo et al. (2003) found the number of dens used by Kit Foxes (Vulpes macrotis) was positively correlated with Coy- ote numbers. In addition, White et al. (1994) suggest- ed that Kit Foxes established a number of dens (220) to facilitate escape. It has also been suggested that Swift Foxes move frequently among different den sites (Kilgore 1969; Hines and Case 1991), but little detail on den use pat- terns has been reported. Understanding Swift Fox den behavior may be an important factor in sustaining viable populations throughout their range. Because Swift Foxes use dens year-round (Egoscue 1979), we installed artificial escape dens as part of a larger study to determine if lack of den sites limited Swift Fox populations in northwest Texas, USA (McGee 2005). The objective of this paper was to doc- ument the effects of artificial escape dens on Swift Fox den use patterns. We determined annual number of dens used, rate of den use (fidelity), distances between dens, den area, and den sharing. We predicted that with more dens available in treated areas Swift Foxes would use more dens, have less fidelity for certain dens, have greater distances between dens, and be less likely to share a den. We made comparisons between treated (artificial escape dens installed) and untreated (no arti- ficial dens installed) areas. Study Area We collected data from a contiguous 100-km? area on the Rita Blanca National Grassland (NG) in Dallam County, Texas, approximately 43 km northwest of Dalhart, Texas (Figure 1). The NG consisted of native rangelands with short-grass prairie dominated by Blue Grama (Bouteloua gracilis), Side-oats Grama (Boute- loua curtipendula), Burrograss (Haplopappus tenui- wal YD THE CANADIAN FIELD-NATURALIST sectus), and Buffalograss (Buchl6e dactyoides) that were moderately to intensively grazed by cattle (Bos taurus; Kamler et al. 2003a,b; Nicholson 2004; McGee 2005). Methods We captured, handled, and radio-collared Swift Fox using methods described by McGee (2005). We tracked Swift Fox to their diurnal resting sites (dens) using a hand-held antenna 1—2 times per week. We recorded each den location with a Garmin global positioning system receiver (Garmin International Inc., Olathe, Kansas, USA). We only used Swift Fox that were monitored =8 months of the year in data analysis. We calculated annual estimates from September to August of each year for all analyses. This was to allow Swift Foxes an adjustment period after installation of artifi- cial dens and to perform two full years of data analysis. During April 2002, we placed 108 artificial escape dens in three spatially separated areas (Figure 1). We considered Swift Foxes belonging to a treated area if their home ranges overlapped an artificial escape den area by =50%. We considered untreated Swift Foxes as those whose home ranges did not overlap artificial escape den areas (McGee 2005). No Swift Foxes in un- treated groups were ever located within an artificial escape den treated area. Also, we considered foxes to belong to the same family group if they used the same area and dens concurrently (Kitchen et al. 1999; Kam- ler et al. 2003a,b). Escape dens consisted of corrugated plastic sewer pipes 4.04 m long, 20.32 cm diameter with 20.32 cm holes cut in the middle to allow foxes to modify and expand subterranean dens ($6.41/m U.S.; Amarillo Plumbing Supply, Inc., Amarillo, Texas, USA). The diameter size of our artificial escape dens was based on previous studies that reported a mean den opening height of 20.0 cm for Swift Fox dens (Cutter 1958; Hill- man and Sharps 1978; Pruss 1999; Jackson and Choate 2000). Coyote dens were reported to be 30-37 cm in diameter (Bekoff 1977; Althoff 1980; Bekoff 1982; Harrison and Gilbert 1985). We assumed that artifi- cial escape den entrances, being the same diameter as natural Swift Fox dens, were too narrow for Coyotes. A John Deere 260 skid loader (Deere and Company World Headquarters, Moline, Illinois, USA) was used to install and cover the sewer pipe with only the two open ends exposed. Escape cover was randomly ori- ented and spaced approximately 322 m apart in a 2.59 km? grid pattern for a density of 36/2.59 km? (McGee 2005). We calculated rate of den use (fidelity) by dividing number of dens by the number of times the fox was located in dens (x 100%). Lower values represented higher den fidelity. We calculated distances between dens using Bearing and Distance Extension for ArcView 3.2 (Environmental Systems Research Insti- tute, Redlands, California, USA). We used den loca- Vol. 121 FiGurE 1. Map of the 100-km? study area located on the Rita Blanca National Grassland in northwest Dallam County, Texas, USA. One hundred and eight artifi- cial escape dens (black dots) where installed in three separate grid locations. tions to estimate annual den areas for Swift Foxes us- ing 100% minimum convex polygon (MCP) method (Mohr 1947) as calculated by Home Range extension (Rodgers and Carr 1998) for ArcView 3.2 (Environ- mental Systems Research Institute, Redlands, Cali- fornia, USA). We calculated den sharing as percent- age of time a radio-collared Swift Fox was found in a den with another radio-collared fox. We found no statistical differences among years so we pooled data to increase power. We used 1-way ANOVAs in SPSS 12.0 (SPSS 2003) to determine differences between treated and untreated areas for annual number of dens used, mean distances between dens, and mean den areas. We compared average rate of den use and sharing between treated and untreated areas using Yates’ corrected chi-square tests (Zar 1999). Differences were deemed significant when P <0.05. Results From January 2002 to August 2004, we captured and radio-collared 55 Swift Foxes (31 males, 24 fe- males). We documented a total of 104 Swift Fox dens a 2007 McGEE, BALLARD, and NICHOLSON: Swift FOX DEN Usi 73 TABLE |. Average annual number of dens and den area of Swift Foxes on Rita Blanca National Grassland (NG) in northwest Texas, 2002—2004. Annual number of dens Study area n % +t SE range treated 8 7.88 + 0.48 6-10 untreated 4 7.50 + 0.50 7-9 during our study including four separate occasions when we radio-tracked and observed Swift Foxes within arti- ficial escape dens during the day. Due to the high turnover rate in our Swift Fox population, only 12 Swift Foxes (n = 8 treated, n = 4 untreated) were monitored for 28 months a year from September 2002 to August 2004. There was no difference (P = 0.64) in annual num- ber of dens used (mean + SE) by Swift Foxes between treated (7.88 + 0.48, n = 8) and untreated (7.50 + 0.50, n = 4) areas (Table |). Average rate of den use (fideli- ty) was similar (Yates’ 7 = 0.003, P = 0.96) between treated (35.9%) and untreated (35.4%) areas. Mean dis- tance between dens (+ SE) was not different (F < 0.001, P=0.99, 1-B = 0.05) between treated (2308 + 442 m, n = 8) and untreated (2317 + 654 m, n = 4) areas. There was no difference in den area (F = 0.38, P = 0.55, 1 — 8 = 0.09) between treated (6.34 + 1.86 km’, n = 8) and untreated areas (3.81 + 1.91 km?, n = 4). Den shar- ing only occurred between mated pairs. Average rate of den sharing was similar (Yates’ x? = 0.56, P = 0.46) between treated (44.9%) and untreated (39.0%) areas. Discussion We have shown that average number of dens used by Swift Fox each year was eight in northwest Texas (Table 1). Schauster et al. (2002) documented num- ber of Swift Fox dens used was 2-8 for breeding and gestation season, 5-10 for pup-rearing season, and 3-8 for the dispersal season. Similar multiple den use has been documented with other small canids that share a dependence on dens, such as Artic Fox (Alopex lago- pus; Eberhardt et al. 1983) and Kit Fox (Tannerfeldt et al. 2003; Moehrenschlager et al. 2004). Koopman et al. (1998) reported an average of 11.8 dens per year for Kit Foxes in California. Arjo et al. (2003) sug- gested to an increase in the number of dens used by Kit Foxes in western Utah could have been related to the increase in Coyote presence. We found that Swift Foxes had relatively high fidelity to particular dens for both treated and un- treated areas. Koopman et al. (1998) found that Kit Foxes exhibited a strong affinity for particular dens because Kit Foxes were located in their most frequent- ly used den 32% of the time. Artic Foxes also preferred certain dens while others were used infrequently (Eber- hardt et al. 1983). We suspect that den fidelity may be the result of den quality. Distinctions have been made between good and bad dens for Artic Foxes (Tanner- Den range (km*) ¥+SE range 6.34 + 1.86 0.52-13.21 4.50 + 1.91 0.81-9.20 feldt et al. 2003). On the other hand, Swift Foxes fre- quently switch between dens (Egoscue 1979; Hines and Case 1991), suggesting poor den quality. One of the reasons for frequent changes between dens has been attributed to the large numbers of fleas found in Swift Foxes’ dens (Kilgore 1969). Other factors like human disturbance, leaking, shifting towards food re- sources, and predator avoidance may play a role in Swift Foxes den switching and fidelity (Kilgore 1969; Tannerfeldt et al. 2003). In addition, we documented a mean distance be- tween dens of 2311 m (range 729-3998 m). Similarly, Moehrenschlager (2000) found that Swift Foxes in Canada moved to dens up to 1900 m away. In contrast, Cutter (1958) noted that Swift Fox dens in overgrazed pastures of northern Texas were concentrated but did not state the distance between dens. Cutter (1958) noted that up to six dens were frequently observed within 65 hectares of pastureland. Greater distances between dens would allow Swift Foxes access to more resources within their environment if dens were used to escape predators. It has been suggested that carnivore home-range size can be affected by habitat composition and food distribution (Macdonald 1983). White and Ralls (1993) observed that larger Kit Fox home ranges were asso- ciated with low prey availability. We suspect that these effects can be applied to den area of Swift Foxes as well. Our results indicated a mean den area of 5.72 km”. It is possible that Swift Foxes with larger den areas may have had greater access to food resources while avoid- ing predators. Ables (1969) recognized that Red Fox (Vulpes vulpes) home range size was affected by food abundance and availability. In addition, Hines and Case (1991) speculated that carrion availability and prey distribution probably affected Swift Fox home range size and shape. Likewise, Olson and Lindzey (2002) suggested that intensive hunting by Swift Foxes near natal dens may have reduced prey availability and con- sequently forced adults to expand the areas in which they hunted. Thus, larger den areas may be the result of fewer resources (Hines and Case 1991). Although additional adult foxes have been observed with mated pairs at Swift Fox (Egoscue 1979: Covell 1992; Kitchen et al. 1999; Lemons et al. 2003), Arctic Fox (Eberhardt et al. 1983), and Kit Fox (Ralls et al. 2001; Tannerfeldt et al. 2003) den sites, we did not document this occurrence. Of the Swift Foxes that we monitored, only mated pairs were ever found occupy- 74 THE CANADIAN FIELD-NATURALIST ing the same den. Adult radio-collared Swift Foxes in our study denned with their mate 39-44% of the time. Similarly, Koopman et al. (1998) found that mated adult Kit Foxes denned together about 45% of the time. Also, Ralls and White (2003) found that Kit Fox pair members shared the same den 51% of the time. It is possible that we underestimated the rate of den shar- ing. We believe that on occasion a radio-collared fox denned with adult foxes that were not radio-collared. In conclusion, we were able to describe Swift Fox den use patterns in northwest Texas even though we did not observe an effect of artificial dens. One possi- ble reason for not observing an effect may be the low sample sizes. Even though we captured 55 Swift Foxes during our study, residents were not abundant as data analysis was only performed on eight Swift Foxes in treated and four Swift Foxes in untreated areas that were monitored =8 months per year. Also, no observed effect of artificial dens may be due to the fact that arti- ficial dens were designed for temporary escape from predators, specifically Coyotes, while Swift Foxes were away from their natural dens. As part of the larger study mentioned before (McGee 2005), we found higher annual Swift Fox survival (P = 0.07) in artificial den treated areas (0.81) than in untreated areas (0.52) on the same study site. Higher survival in treated areas suggests that Swift Foxes were using artificial dens for escape during their normal nocturnal activities. Diur- nal use of artificial dens was limited. We only tracked Swift Foxes to artificial escape dens on four occasions during the day. Therefore, alternative artificial den designs need further study to find a more suitable diur- nal artificial den for Swift Foxes. Acknowledgments Funding was provided by the National Fish and Wildlife Foundation and Texas Tech University. We thank the United States Forest Service personnel for letting us conduct research on the Rita Blanca Nation- al Grasslands and helping install artificial escape dens. We thank F. Pronger for allowing us to use his ranch as a study site. We also thank E. and B. Hampton for providing us with a place to stay while conducting field research. Research protocols were approved by the Animal Care and Use Committee at Texas Tech University. This is Texas Tech University, College of Agricultural Sciences and Natural Resources techni- cal publication T-9-1091. Literature Cited Ables, E. D. 1969. Home-range studies of Red Foxes (Vulpes vulpes). Journal of Mammalogy 50: 108—120. Allardyce, D., and M. A. Sovada. 2003. 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Journal of Wildlife Management 57: 861-867. White, P. J., K. Ralls, and R. A. Garrott. 1994. Coyote- Kit Fox interactions as revealed by telemetry. Canadian Journal of Zoology 72: 1831-1836. Zar, J. H. 1999. Biostatistical analysis, 4" edition. Pren- tice-Hall, Inc., Upper Saddle River, New Jersey, USA. Received 3 February 2006 Accepted 17 January 2008 Flowering During January in Antigonish County, Nova Scotia* DAVID J. GARBARY and BARRY R. TAYLOR Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia, B2G 2W5 Canada; e-mail: dgarbary @ gmail.com *Contribution from the Herbarium of St. Francis Xavier University Garbary, David J., and Barry R. Taylor. 2007. Flowering during January in Antigonish County, Nova Scotia. Canadian Field- Naturalist 121(1): 76-80. Over 85 records of 23 species of blooming, herbaceous angiosperms were made at 19 sites in Antigonish County between 7 and 21 January 2006, when daytime temperatures reached 15°C. These observations followed an unusually warm fall and early winter. All species were observed on waste ground or in fields and garden plots, except for Epigaea repens L. which was part of ground vegetation in a sparsely wooded site. The primary families represented were Asteraceae (six species), Brassicaceae (six species) and Carophyllaceae (four species). The most commonly observed plants were Taraxacum officinale (L.) Weber (11 sites), Capsella bursa-pastoris (L.) Medik. (nine sites), Thlaspi arvense L. (three sites) and Stellaria media (L.) Vill. (four sites). Many plants and inflorescences were conspicuously frost-damaged, and flowers were rarely fully open. In several species, e.g., TZ: arvense and Cerastium vulgatum L., many individual plants looked normal and there was no evidence of frost damage. These observations are the latest flowering records for Nova Scotia. Key Words: Asteraceae, Brassicaceae, Caryophyllaceae, winter flowering, phenology, Antigonish County, Nova Scotia. In Nova Scotia most species of flowering plants have ceased to flower by early November (Zinck 1998). Taylor and Garbary (2003) carried out an ex- tensive survey of flowering during an unusually warm late fall in Antigonish County and found 92 species in bloom during November-December. These were the latest flowering records of all but one of these species, and extended the known flowering periods for plants in Nova Scotia by an average of 45 days. Of particular note were 15 species that were in bloom on 14-16 De- cember 2001. A detailed evaluation of the phenology of the entire late fall flora was again undertaken in 2005 after anoth- er warm fall in which abundant flowering was still oc- curring in early November. This study will be described elsewhere in the context of regional climate change (Garbary et al.). Here, we provide records of 23 species of herbaceous plants observed in flower during an ex- traordinary warm period in January 2006. Our obser- vations complement those on spring phenology by Vasseur et al. (2001). They compared flowering times in 12 species in the late 1990s with dates a century earlier and showed significant changes in only two species, of which only Syringa vulgaris L. had earlier flowering. Materials and Methods Collection sites Nineteen sites in Antigonish Town and County, northern Nova Scotia, were explored between 7 and 21 January 2006 in three collection periods. Adjacent sites were a minimum of 200 m apart, and the maxi- mum distance between two sites was 23 km. Ten sites were visited on 7 January, 16 sites during the period 11-16 January, and seven sites on 21 January. Five sites were visited three times, and three other sites were visited twice. Most of these sites had been previously visited by Taylor and Garbary (2003). Sites were typi- cally highly disturbed areas of waste ground or gravel (Figure 1), or agricultural fields or garden plots. Each site was surveyed for 15-45 minutes. Notes were taken on the flowering status and plant condition of each species in bloom. Representative plants were collected; plants with cryptic flowers were dissected and exam- ined under a stereomicroscope to ensure that flowers were intact. Voucher specimens for most species were prepared and deposited at St. Francis Xavier Univer- sity. Nomenclature follows Zinck (1998). Weather Weather data were taken from the nearest active Environment Canada meteorological station at Tra- cadie, approximately 25 km from Antigonish (http:// climate.weatheroffice.ec.gc.ca/climateData/canada _e.html). The first collecting period was on 7 January when daytime temperatures reached 4.3°C. This fol- lowed several days with daily maximum temperatures above 0°C and daily minimal temperatures no lower than -2.5°C. Each of the succeeding four nights had frost, although daytime maxima were always above freezing. The second collecting period, between 12 to 16 January, had minimum and maximum tempera- tures of 0.4°C and 15.9°C, respectively. Temperatures between 15 to 20 January were cooler and frosts oc- curred most nights (minimum -6.8°C) although mean daily temperatures were below freezing on only one day (-4.6°C). The final collecting day was 21 January when temperature ranged from 2.0 to 8.4°C. This was followed by at least ten days when temperatures dropped below freezing (-0.3 to -9.2°C) and condi- tions were more typical of the season. 76 2007 r GARBARY AND TAYLOR: FLOWERING DURING JANUARY Ficure 1. Sample habitat. Waste ground on campus of St. Francis Xavier University adjacent to Trans-Canada Highway, 7 January 2006. Four species were found in flower at this site. Results ASTERACEAE Taraxacum officinale (L.) Weber was the most wide- ly distributed species, occurring at ten sites. One to four plants were observed at each site during all three collecting periods and blossoms occurred at two sites on 21 January. Each plant had one or two ragged inflorescences with short pedicels. Leaves had some frost damage and were often flattened against the substratum, although they had good colour on some individuals. Achillea millefolium L. occurred as a single plant in a farmer’s field and as six plants on waste ground (Figure 2). Plants had green leaves, but the flowering shoots were often reclining, with several to dozens of white florets. The flowering shoots observed 14 Jan- uary were wilted and most of the flowers had turned brown. Two species of Matricaria were found: M. mariti- ma L. and M. matricarioides (Less.) Porter. Plants of M. maritima were found at two sites. A single, well- developed inflorescence was found on 7 January as well as several partially intact flower heads. By 15 Jan- uary remaining ray florets were mostly lost; however, the disc florets on many heads remained bright yellow. The plants were often partly wilted but the leaves had healthy colour. Up to three plants of M. matricari- oides were found at each of three sites, and it was observed during the first two collecting periods. In- florescences typically had a mixture of dull yellow and brownish florets. The plants appeared no less viable than others observed in early December. Leontodon autumnalis L. occurred at two sites on 7 January where single individuals were found with only one or two wilted inflorescences on each plant. Basal rosettes had good colour but were slightly wilt- ed. Examination of both sites one week later revealed no flowering individuals. Two plants of Senecio vulgaris L. occurred at one site on waste ground; a single individual was found at a second site. Florets on most inflorescences had turned brown but several had bright yellow florets. Plants were a maximum of 10 cm in height and flow- ering was observed only in the first two observation periods. 78 THE CANADIAN FIELD-NATURALIST ayy i@ FIGURE 2. Achillea millefolium photographed in situ on 7 January 2006 at site shown in Figure 1. Flowering shoot is about 30 cm tall. BRASSICACEAE Capsella bursa-pastoris (L.) Medik. was among the most common plants flowering in January. It oc- curred at nine sites with one to three plants per site. Flowering individuals were up to 30 cm tall and car- ried a few flowers at the apex of each plant. Leaves were typically green and healthy, although larger plants had generally become decumbent. During the second week, plants were mostly very small (5-10 cm high) with only a few florets. On 21 January flowers were found at two sites. Thlaspi arvense L. was among the most robustly flowering species observed. The plants were erect, dark green and with prominent white flowers. 7: arvense in flower was as common in January as in any previ- ous collecting period, and flowers were found at one site on 21 January. Raphanus raphanistrum L. occurred at two sites where there were typically many plants. The stems and leaves of these plants were badly wilted and reclin- ing; however a few bright yellow florets and numer- ous clusters of buds remained intact at many apices on many plants. In the second week only a single, badly damaged plant was found at one site. Cardamine pensylvanica Muhl. ex Willd. was the only species clearly protected by the heat shadow of a large building. Dozens of small, prostrate plants each with one to several flowers occurred in a flowerbed along the side of a large brick building. No decline in flowering was noted in the second week and flowers persisted to 21 January. Erysimum cheiranthoides L. was found at two sites and only in the second week. Four large plants were found along a bare concrete foundation for a house. The surrounding lot supported numerous plants of this species but all had gone to seed. A second site had a few small individuals up to 5 cm tall. Both sites were covered with snow on 7 January and so were not examined until the second week. Vol. 121 A single plant of Lepidium campestre (L.) R. Br. was found 12 January on waste ground. The plant was about 20 cm tall and had two clusters of flowers with the most vigorous close to the ground and im- mersed in a cluster of leaves. CARYOPHYLLACEAE Stellaria media (L.) Vill. was found at two or three sites in all collecting periods (four sites overall), and was among the most prominent of the plants in flower. At two locations it was found as scattered individual plants; however, at two other sites it formed large ex- panses of continuous ground cover with numerous inflorescences (Figure 3). FiGure 3. Stellaria media photographed in situ on 7 January 2006 showing part of a large colony about 1 m in diameter. Flowering individuals of Cerastium vulgatum L. were few and occurred at only three sites, although many vegetative plants or plants with remains of inflo- rescences were present at most locations. The repro- ductive individuals appeared healthy and showed no signs of frost damage. They grew on loose gravel where there was no competition from surrounding vegeta- tion. Flowering persisted at one site to 21 January. A few flowers were present on several plants of Spergula arvensis L. from a farmer’s field in the first two weeks. The plants showed considerable frost dam- age and dissection showed that very few bore intact stamens and pistils. At a second site where vegetative parts of the plants were less damaged, no intact flow- ers occurred. An extensive patch of Spergularia rubra (L.) J. & C. Presl. occurred on waste ground. The prostrate plants were 5-10 cm diameter and showed all reproductive stages from buds to fruits. The site was only exam- ined during the second week of observation. OTHER FAMILIES Euphorbia helioscopa L. (Euphorbiaceae) was pres- ent at only one of the sites. There were hundreds of individuals. Large plants (20-40 cm tall) were con- 2007 spicuously frost damaged, with the wilted stems caus- ing the upper parts of the plants to topple over. The terminal portions of these plants looked healthy, and subsequent dissection of inflorescences showed intact flowers. Small plants (5-15 cm tall) showed less col- lapse of the stems and also had numerous intact flow- ers. Flowering persisted until 21 January. Individual plants of Fumaria officinalis L. (Fumari- aceae) were found on each of the first two visits to one site. The plants had two small inflorescences in which a few flowers still retained their characteristic two- toned colour. Inflorescences on other plants at this site had been severely damaged and turned brown. On most individuals the finely dissected leaves and deli- cate stems showed no conspicuous wilting. At another site dozens of plants grew in soil mounds, but there were no inflorescences. Viola tricolor L. (Violaceae) occurred at two sites. Both were in flower beds and one was adjacent to a building. Plants at both sites had leaves and stems in excellent condition and several to dozens of fully open flowers with blossoms persisting until 21 January. There were numerous plants of Veronica persica Poir. (Scrophulariaceae) at one site where they grew as prostrate clumps or straggling plants over the bare soil. The bilobed fruits were conspicuous, and a single intact but only partially open flower was found on sin- gle individuals returned to the laboratory at each of the three visits. Vegetative plants of Polygonum arenastrum Jord. ex Boreau (Polygonaceae) were common at most sites. Although most plants had conspicuous frost damage, a few plants had two to three white-tipped buds and open flowers that persisted until week two. Epigaea repens L. (Ericaceae) was the only wood- land plant found in the study. An extensive population occurred at one site (14 January) among other ground vegetation beneath an open, mixed forest growing on sandy soil. Although buds on many plants appeared swollen, only a single plant had open blossoms. Discussion That any plants should be flowering in January in northern Nova Scotia, much less 23 species, is remark- able. This late phenology may reflect the ability of plants to have late flowering strains (e.g., Thlaspi ar- vense, Warwick et al. 2002), or possibly reflect a west- ern European origin where fall and early winter tem- peratures would be much milder. Zinck (1998) does not include December in the flowering periods of any of these species; November is included in the flower- ing periods only for Capsella bursa-pastoris, Matri- caria matricarioides, Senecio vulgaris, Stellaria media and Viola tricolor. The region falls within plant har- diness zone 5B of Agriculture and Agri-Food Canada (http://sis.agr.gc.ca/cansis/nsdb/climate/hardiness/intro. html), in which harsh winters kill non-hardy species. January is normally associated with severe winter weather; the 30-year daily average temperature at Col- GARBARY AND TAYLOR: FLOWERING DURING JANUARY 79 legeville (the nearest meteorological station for which climate normals are available) is -6.6°C, and the aver- age minimum is —11.6°C. Winters here are among the coldest in the province (Davis and Browne 1997). About 50 cm of snow falls on average in January, and a heavy snowpack is common. Snowpack depths in a wooded area of the St. Francis Xavier University cam- pus were 40-50 cm in mid-January every year but one in 1999-2005 (B. Taylor, unpublished data). In the single low-snow year, the weather was very cold and the ground was frozen. The circumstances that led to plants blossoming in mid-January were more complex than an extended peri- od of warm weather. While fall and early winter of 2005-2006 were unusually mild, there had been reg- ular nights of frost since November and two major snowfalls (5 and 15 December) prior to the collection period. The sites were visited over the course of three weekends, and in the intervening periods there were frosts on most night at Tracadie. These plant species appear to be very resistant to short-term freezing, as long as warm temperatures return quickly. In addition, the most severe weather may have occurred when the plants were covered with 20 cm of snow from the 15 December snowstorm. The snow would have insulat- ed the plants from the deep frosts, but it was subse- quently washed away by heavy rain. None of the plants in bloom was restricted to the first round of collecting; indeed, five species were found in bloom only in the second round and ten species had blossoms on 21 January. The two sites supporting Erysimum cheiranthoides were snow-covered on 7 Jan- uary and the single specimen of Lepidium campestre found in week two may have been missed on the ear- lier visit to the site. Only one site visited in this study could be considered highly protected and influenced by radiant heat from a building, and Cardamine pen- sylvanica was the only species unique to this site. The remaining sites were exposed to ambient air tempera- tures and precipitation. Nevertheless, even though plants in bloom were found at 17 sites in mid-January, this was not a con- spicuous phenomenon; surveys of additional sites re- vealed no plants in blossom. The sites with plants in flower comprised highly specialized habitats mostly of limited extent: land cleared for development, margins of abandoned or active agricultural fields, garden plots and soil mounds. Even within these sites, most plants of many species reported here had entirely succumbed to the rigours of winter. The plant species in bloom were not a cross-section of the provincial flora. Most of the species reported here are opportunistic, weedy species that thrive in highly disturbed habitats, including agricultural set- tings (e.g., Raphanus raphanistrum L., Warwick and Francis 2005). Indeed many of our sites had less than 50% cover of vegetation. In addition, all species ob- served in flower except Epigaea repens, Cardamine pensylvanica and Cerastium vulgatum are non-native 80 THE CANADIAN FIELD-NATURALIST members of the flora (Zinck 1998). Most of these species are weeds that were introduced from Europe with grain. It would appear that the ability to tolerate harsh physical conditions (in contrast with competi- tive ability) extends to tolerance of harsh weather as well (Taylor and Garbary 2003). Flowering in January in a cold climate would ap- pear to be futile, and therefore likely of little evolution- ary value. Excepting Epigaea repens, all of the species reported here are annuals or short-lived perennials (Zinck 1998). As early colonizers of disturbed ground, they would be adapted to growing and reproducing quickly in advance of competition from later species. It seems likely that these species are adapted to grow, flower and set seed as long as the weather permits (e.g., Senecio vulgaris, Robinson et al. 2003). This phenol- ogy contrasts with that of longer-lived perennials in which above-ground plant parts senesce in late autumn in anticipation of winter. The ruderal species observed here evidently lack this adaptation, presumably be- cause either cold temperatures (annuals) or competi- tion (perennials) normally curtails reproduction and kills the plant. Consequently, when those events are delayed, the plants have no physiological mechanism to end flowering, even when seed production may be impossible. An important exception to the above is Pomquet Beach Provincial Park, an undisturbed, coastal site where Epigaea repens was part of a dense community of ground vegetation beneath a diffuse tree layer. Unlike the remaining fall-flowering species, Epigaea repens normally flowers in April and May (Zinck 1998), when it forms a conspicuous part of the spring-flowering flora. The plants in flower in January may be consid- ered either a second, late-fall flowering, or an excep- tionally early spring flowering. This phenomenon con- trasts with the situation of the remaining species, in which normal late-fall flowering merely persisted for a long time. It is doubtful that new flowers were initi- ated among these species. Epigaea repens was observed in flower in the An- napolis Valley, about 250 km southwest of our study area, in early November 2005 (N. Nickerson, Agricul- ture and Agri-Food Canada, Kentville, personal com- munication). Hence our flowering material from Antigonish County probably should be considered a further extension of a late fall return to flowering. The same probably applies to Fumaria officinalis, for which the latest flowering month given by Zinck (1998) is August. It would be of interest to determine details of plant phenology for E. repens and other species at the extreme southern tip of the province, where Janu- ary mean temperate is only just below freezing (plant hardiness zone 6B; Davis and Browne 1997). Vol. 121 Changes in plant phenology are indicators of climate change and have been used as a proxy for tempera- ture (Badeck et al. 2005; Menzel 2002; Sherry et al. 2007; Walther 2004). Consequently, the January flow- ering records reported here might be considered as evidence of regional climate warming. Thus, even if regional climate warming has not been demonstrated, the changes that we observed would be a natural con- sequence of such a warming trend. Acknowledgments We thank Robert Garbary and Jonathan Ferrier who assisted with field work and Nancy Nickerson for in- formation on flowering of Epigaea repens. This work was supported by grants from the Natural Sciences and Engineering Research Council of Canada to both authors. Literature Cited Badeck, F.-W., A. Bondeau, K. Boéttcher, D. Doktor, W. Lucht, J. Schaber, and S. Sitch. 2005. Responses of spring phenology to climate change. New Phytologist 162: 295-309. Davis, D. S., and S. Browne (Editors). 1997. The natural history of Nova Scotia, Volume 1. Topics and habitats. Nimbus Publishing and Nova Scotia Provincial Museum, Halifax, Nova Scotia. 518 pages. Menzel, A. 2002. Phenology: its importance to the global change community. Climate Change 54: 379-385. Robinson, D. E., J. T. O’Donovan, M. P. Sharma, D. J. Doohan, and R. Figueroa. 2003. The biology of Canadian weeds. 123. Senecio vulgaris L. Canadian Journal of Plant Science 83: 629-644. Sherry, R. A., X. H. Zhou, S. L. Gu, J. A. Arnone, D. S. Schimel, P. S. Verburg, L. L. Wallace, and Y. Q. Luo. 2007. Divergence of reproduction phenology under climate warming. Proceedings of the National Academy of Sci- ences of the United States of America 104: 198-202. Taylor, B. R., and D. J. Garbary. 2003. Late-flowering plants from northern Nova Scotia. Rhodora 105: 118-125. Vasseur, L., R. L. Guscott, and P. J. Mudie. 2001. Moni- toring of spring flower phenology in Nova Scotia: com- parison over the last century. Northeastern Naturalist 8: 393-402. Walther, G. R. 2004. Plants in a warmer world. Perspectives in Plant Ecology Evolution and Systematics 6: 169-185. Warwick, S. I., and A. Francis. 2005. The biology of Cana- dian weeds. 132. Raphanus raphanistrum L. Canadian Journal of Plant Science 85: 709-733. Warwick, S. I., A. Francis, and D. J. Susko. 2002. The biology of Canadian weeds. 9. Thlaspi arvense L. (up- dated). Canadian Journal of Plant Science 82: 803-823. Zinck, M., Editor. 1998. Roland’s flora of Nova Scotia, 3 edition. Nimbus Publishing Ltd. and Nova Scotia Provincial Museum, Halifax, Nova Scotia. 1297 pages. Received 24 January 2006 Accepted 15 January 2008 The Occurrence of Muskoxen, Ovibos moschatus in Labrador | Tony E. Cuusss! and J. BRAZIL* 'Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and Labrador AOP 1SO Canada; e-mail: techubbs @cablelab.net *Endangered Species and Biodiversity Section, Wildlife Division, Department of Environment and Conservation, Govern- ment of Newfoundland and Labrador, P.O. Box 2007, Corner Brook, Newfoundland and Labrador A2H 7S! Canada Chubbs, Tony E., and J. Brazil. 2007. The occurrence of Muskoxen, Ovibos moschatus in Labrador. Canadian Field-Natu- ralist 121(1): 81-84. Muskoxen were first observed in Labrador in 1988 and have since been recorded at several locations in two separate river valleys in northern Labrador. The occurrence of Muskoxen in Labrador is attributed to the dispersal of captive releases from / Québec in 1973 and 1983. Here we document observation records and examine the future management of the species in The Muskox (Ovibos moschatus) is a relic from ‘the Pleistocene and is native to northern Canada, ‘northwestern Greenland, and the northern coast of Alaska (Banfield 1974; Reynolds 1988). The elimi- “nation of the species from Alaska and the reduction ~of Muskox numbers in Canada by the early 1900s re- ‘sulted in efforts to introduce and restore populations ‘in several arctic areas (Klein 1988; Reynolds 1988). At present, Muskox populations are experiencing a period of success, with high numbers throughout most ‘of their range (Groves 1997). Muskox apparently did ‘not colonize the Québec-Labrador peninsula after the i last glaciation as no remains have been found at archae- _ dlogical sites (Banfield 1975). ' In 1967, 15 Muskoxen (12 females, 3 males) were ‘captured near Eureka, Ellesmere Island (Centre D’E- vudes Nordique 1981*), and brought to the Uming- maqautik farm, near Kuujjuag in northern Québec _ Figure 1) for Inuit to establish a Muskox farm, cre- Ate employment, and build up a stock to introduce the Bpecies into northern Québec (Le Hénaff and Créte 1989; Nault et al. 1993). A project of raising the ani- “nals to collect their quviut (Muskox wool) was unsuc- >essful. Subsequently, from 1973 to 1983, 54 animals vere released at three different locations in northern Québec (Le Hénaff and Créte 1989). The first release sites were located to the north of Kuujjuaq and to the Benuivest of Tasiujaq (Le Hénaff and Créte 1989). In +983, Muskoxen were released east of Kuujjuag near he outlet of the Whale River. The Umingmagqautik arm was closed in 1983 and the remaining animals vere given to the St-Félicien and the Orsainville zoos -n Québec (Jean and Rivard 2005*). __ The introductions in Québec proved successful. In _ 983 a total of 148 Muskoxen were counted and by 986, with a finite rate of increase of 1.25, the popu- _ ation had increased to 290 animals. By 1986, indi- viduals and mixed groups of Muskoxen had dispersed ‘nall cardinal directions up to 650 km from the origi- Labrador and speculate on its possible effect on this northern ecosystem. / Key Words: Muskox, Ovibos moschatus, range, distribution, management, Labrador, Québec. nal release sites throughout the province of Québec (Le Hénaff and Créte 1989). In 2003, the Muskox population around Kuujjuaq and Tasiujaq was estimated to be 1400 animals, includ- ing calves (Jean and Rivard 2005*). In June 2005, aer- ial surveys on Diana Island, near Quaqtaq, counted a total of 112 adults and 25 calves. This new population apparently originated from the Tasiujaq area (Jean and Rivard 2005*). Study Area Over the past two decades, Muskoxen have been ob- served on several occasions in the Low Arctic Alpine- Torngat ecoregion in northern Labrador. This ecore- gion has short, cool summers and long, cold winters with mean average daily temperatures ranging from -19 to -22°C in February and 7 to 9°C in July. Aver- age annual precipitation is 500 to 700 mm with a total average snowfall of 3 m (Meades 1990*). The vegeta- tion is tundra (Alpine heath) with sedge meadows domi- nating plateaus and Mountain Alder (Alnus crispa) and willow (Salix spp.) thickets dominating on the lower slopes and river valley slopes. Groves of White Birch (Betula papyrifera) and Balsam Poplar (Populus bal- samifera) occur sporadically in some river valleys. Graminoids occur on the valley bottoms but are sparse to absent on the upper slopes and at high elevations. Dwarf Birch (B. glandulosa), Labrador Tea (Ledum groenlandicum) and Black Crowberry (Empetrum nigrum) dominate drier mountain ridges. Conifer trees are absent from this region (Meades 1990*). Major features of the terrain include braided east-flowing rivers, foothills, elevated plateaus, rugged coastal cliffs and inland mountains. Methods We recorded Muskox sightings during remote aeri- al field surveys in northern Labrador. We queried res- idents from remote communities throughout northern 81 82 THE CANADIAN FIELD-NATURALIST Labrador and helicopter and fixed-wing aircraft pilots for sightings of Muskoxen. We found only one record of Muskoxen in other areas of Labrador apart from our own observations. We classified individual Muskox to age and sex from oblique aerial photographs. When possible, animals were classified as adult male or fe- male, based on body size, coat color and size and shape of horns. Observations Five sightings of Muskoxen have been recorded in Labrador over the past two decades in three different river valleys, approximately 340 km apart. Four sight- ings were made from a helicopter in different years (1988, 1995, 2000, 2005) and included three sight- ings of a single Muskox, likely different animals, and one sighting of a group of three. The fifth sighting (2006 — 2008) was reported by residents of Nain while travelling by snowmobile to Voisey’s Bay. Site 1. Komaktorvik River Valley The first sighting of Muskoxen in Labrador was recorded on 24 July 1988 at approximately 59°14'N, 63°56'W along the Komaktorvik River valley, which flows into Seven Islands Bay (Figure 1). Three indi- viduals were observed in a large patch of willows (Salix spp.) in the river valley. All individuals had horns with a slight separation in the butte and were all approxi- mately the same size indicating that these animals were likely adult males, although definite sex and age remain uncertain (J. Brazil, personal observation). On 28 July 1995, one Muskox was again observed in the Komaktorvik River valley (59°14.65'N, 63°55.51'W) very near the original 1988 sighting. From a photo- graph, this animal, browsing in a large thicket of wil- lows (Salix spp.) along the north slope of the river val- ley, appeared to be an adult male. On 30 July 2000, during aerial surveys for Harle- quin Ducks (Histrionicus histrionicus) and Peregrine Falcons (Falco peregrinus), we observed and pho- tographed a lone Muskox in Seven Islands Bay at (59°14.16'N, 63°57.66'W) 3 to 4 km west of the previ- ous sightings in 1988 and 1995. This animal was large, had well formed horns, and light coloration on the back and saddle and was identified as a sub-adult male (Ole- sen and Thing 1989). The sightings along the Komaktorvik River valley occurred approximately 250 km northeast of the re- lease site at Whale River providing a straight-line rate of dispersion of 50 km/year. The most probable dis- persion route was along the Koroc River valley, which traverses the Labrador Peninsula. Site 2. Hebron Fiord On 9 June 2004, a single Muskox was observed near the outlet of Hebron Fiord (58°03.82'N, 63°12.71’ W) (Figure 1, Site 2). This animal was feeding in a thick- et of alders (Alnus spp.) and Willows (Salix spp.) at the foot of a small plateau on the north side of Hebron Brook where it flows into Hebron Fiord. A photo- Vol. 121 graph identified this animal as an adult male > 4 years old (D. Jean, personal communication). The straight- line dispersal from the Whale River release site was a distance of 220 km providing a rate of dispersion of approximately 11 km/year. The most probable route of dispersion was south along the George River val- ley and then the Tasirlaq River to Hebron Fiord. Site 3. Voisey’s Bay In April 2006, the fifth Muskox reported in Lab- rador was observed on a small unnamed island south of Tabor Island in Voisey’s Bay at approximately 54°20'N, 61°43'W (Figure 1, Site 3). This animal was readily identifiable as its right horn tip was broken at the curl. This 4-5 year old male has remained on the island and is in its second winter at this location. The Straight-line dispersal from the Whale River release site is a distance of 400 km providing a rate of dis- persion of approximately 17 km/year. The most prob- able route of dispersion was south along the George River valley and then along either the Fraser or Kogaluk river to Voisey’s Bay. Discussion Muskoxen have successfully dispersed and colo- nized many remote regions of northern Québec. The dispersal of Muskoxen from Québec into Labrador may have been influenced by unoccupied habitat and lack of dispersal barriers. No mixed-aged groups (the reproductive segment of the population) have been observed to date in Labrador. It is thus likely that the separate observations at Kom- aktorvik River valley, Hebron Fiord and Voisey’s Bay are of different animals that immigrated to Labrador through separate overland routes and at different times. Based on the dispersion of Muskox south along the George River in Québec through 1986 (Le Hénaff and Créte 1989), it is possible that other individuals or groups have traveled eastward into Labrador as far south as Nain along the numerous river valleys ex- tending inland. Although Muskoxen and Caribou (Rangifer taran- dus) are occasionally found together on the same veg- etation type, caribou usually move through an area rapidly, feeding on willows and flowering forbs where- as Muskoxen are more sedentary and forage on sedges and grasses (Parker 1978). Abundance and distribu- tion of Muskoxen on many of the islands of the High Arctic appear to be related to the abundance and dis- tribution of sedge-producing meadows (Parker et al. 1975; Parker and Ross 1976). Initial colonization by Muskox may be slow and sporadic consisting mainly of adult males (Le Hénaff and Créte 1989; Reynolds 1988). This may explain why reproductive, mixed-aged groups have do not appear to have been observed yet in Labrador. As Muskoxen had populated many remote regions of Québec in all cardinal directions north of 54°30'N latitude by 1986 (Le Hénaff and Créte 1989) and populations continue 2007 CHUBBS and BRAZIL: MUSKOXEN IN LABRADOR 83 Labrador Legend Sightings Communities Rivers FiGuRE 1. Location of Muskox (Ovibos moschatus) sightings in northern Labrador. The two release sites were near Tasiujagq and Kujjuaq in Québec. to grow (lambda = 1.25) near their maximum finite rate of increase (1.30), breeding groups may have be- come established in Labrador but have gone undetect- ed. Emigration of mixed-sexed groups out of regions first occupied may result in shifts in population dis- tribution and range expansion (Reynolds 1988). The dispersal and colonization of Muskoxen to adjacent ranges can be slow. However, with the present high rate of population increase (Jean et al. 2004*) in northern Québec, we believe that emigration may soon result in a shift in an expansion of the distribution and range of Muskoxen into Labrador. Reynolds (1988) identi- fied three stages in the expansion of Muskox into new range: (1) slow growth following release (2) the irrup- tive phase and (3) decline and stabilization. It is likely that in the next decade during the decline and stabi- lization of Muskox numbers in Québec that the dis- persal of mixed groups into Labrador will occur and a breeding population be established. In Alaska, where Muskoxen were extirpated and later reintroduced, range expansion occurred only over 20% of the former Muskox range and populations grew to fewer than 1000 individuals over nearly 30 years (Reynolds 1988). Management Implications Hunting of Muskox under a limited permit system is conducted on Nunivak and Nelson islands and in the Arctic National Wildlife Refuge of Alaska and in Nunavut, Northwest Territories, and Québec in Can- ada. Muskoxen are considered a unique and valuable trophy by the outfitting industry and Muskox meat is highly valued as an exported delicacy. This survivor of the ice ages is an important attraction for tourists, photographers, researchers and students of wildlife in areas where the species is endemic or introduced. The establishment of a Muskox population in Labra- dor may provide an alternative to Caribou as a source of protein for northern communities as well as a new resource for developing the regional economy through sport hunting similar to northern Québec and Alaska (Le Hénaff and Créte 1989; Jean et al. 2004*; Reynolds 1988). It appears that habitat for Muskoxen is widely dis- tributed and largely unchanged in northern Labrador. With public support and proper management, Muskox- en may eventually become a more visible and famil- iar wildlife species in Labrador. We recommend that systematic aerial surveys be conducted to document the current and future status and distribution of Muskoxen in Labrador. Acknowledgments Thanks are extended to D. Jean, Ministére des Res- sources naturelles et de la Faune, who provided liter- ature on Muskoxen introduction and releases in Québec 84 THE CANADIAN FIELD-NATURALIST as well as assistance in sexing and aging Muskoxen from photographs. M. Créte, F. Phillips, G. Parker, the late N. P. P. Simon and an anonymous reviewer pro- vided comments on earlier drafts of the manuscript. L. Elson drafted the figure. The Department of Environ- ment and Conservation, Government of Newfoundland and Labrador, provided funding for this publication. Documents Cited (marked * in text) Centre D’études nordiques. 1981. Rapport d’évaluation du projet d’élevage de boeuf musqué, Umingmagautik, Nouveau-Québec. (1967-1980). Centre d’ Etudes nordiques et Faculté des sciences de l’agriculture et de l’alimentation, Université Laval. Présenté au ministére de I’ Agriculture, des Pécheries et de |’ Alimentation du Québec, Québec. 168 pages. Jean, D., and S. Rivard. 2005. Inventaire de la population de boeeufs musqués (Ovibos muschatus) sur Vile Diana, prés de Quagtaq. Ministére des Ressources naturelles et de la Faune — Secteur Faune Québec, Direction de l’aménage- ment de la faune du Nord-du-Québec. 8 pages. Jean, D., S. Rivard, and M. Bélanger. 2004. Inventaire et structure de population du boeuf musqué (Ovibos mos- chatus) au sud-ouest de la baie d’Ungava, aott 2003. Ministére des Ressources naturelles, de la Faune et des Parcs — Secteur Faune Québec, Chibougamau. 22 pages. Meades, S. J. 1990. Natural regions of Newfoundland and Labrador. Protected Areas Association, St. John’s, New- foundland and Labrador. 103 pages. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. University of Toronto Press, Toronto. Vol. 121 Banfield, A. W. F. 1975. Les mammiferes du Canada. Musées nationaux and University of Toronto Press. Toronto. Groves, P. 1997. Intraspecific variation in mitochondrial DND of Muskoxen, based on control-region sequences. Canadian Journal of Zoology 75: 568-575. Klein, D. R. 1988. The establishment of Muskox populations by translocation. Pages 298-317 in Translocation of wild animals. Edited by L. Nielsen and R. D. Brown. Wisconsin Humane Society, Milwaukee, Wisconsin, USA. Le Heénaff, D., and M. Créte. 1989. Introduction of muskoxen in northern Québec: the demographic explosion of a colo- nizing herbivore. Canadian Journal of Zoology 67: 1102- 1105. Nault, R., C. Mathieu, and M. Créte. 1993. Vegetation bio- mass and habitat selection by a newly introduced population of muskoxen in northern Québec. Rangifer 13: 71-77. Olesen, C. R., and H. Thing. 1989. Guide to field classifi- cation by sex and age of the Muskox. Canadian Journal of Zoology 67: 1116-1119. Parker G. R. 1978. The diets of muskoxen and Peary caribou on some islands in the Canadian High Arctic. Canadian Wildlife Service Occasional Paper (35). 21 pages. Parker G. R., and R. K. Ross. 1976. Summer habitat use by muskoxen (Ovibos moschatus) and Peary caribou (Rangi- fer tarandus pearyi) in the Canadian High Arctic. Polar- forschung 1976(1): 12-25. Parker, G. R., D. C. Thomas, E. Broughton, and D. R. Gray. 1975. Crashes of muskoxen and Peary caribou pop- ulations in 1973-74 on the Parry Islands, Arctic Canada. Canadian Wildlife Service Progress Note (56). 15 pages. Reynolds, P. 1988. Dynamics and range expansion of a re- established Muskox population. Journal of Wildlife Man- agement 62: 734-744. Received 15 March 2006 Accepted 8 March 2008 The Spindle Tree, Euonymus europaea L. (Celastraceae): A Newly Naturalized Shrub in Nova Scotia Davip J. GARBARY! and Amy E. DEVEAU 'Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2WS5 Canada; e-mail: dgarbary @ gmail.com Garbary, David J., and Amy E. Deveau. 2007. The Spindle Tree, Euonymus europaea L. (Celastraceae); a newly naturalized shrub in Nova Scotia. Canadian Field-Naturalist 121(1): 85-88. Over 100 plants, including over 750 individual shoots, of Euonymus europaea L. are reported from the Town of Antigonish in northern Nova Scotia. Plants were found in five discrete areas separated by 200-1000 m. A few individuals at one site may represent human planting; however, over 90 % of plants occur on waste ground or in dense shrubbery where natural colonization has occurred. Many individuals were observed in fruit in the fall of 2005, and over 95% of the plants had well developed buds and flowers in May and early June 2006. Euonymus europaea is considered a newly naturalized species for the flora of Nova Scotia. Key Words: Spindle Tree, Euonymus europaea, introductions, naturalized, Nova Scotia. Euonymus (Celastraceae) is a cosmopolitan genus of over 200 species of vines, deciduous and evergreen shrubs, and small trees (Gleason and Cronquist 1991). Many species are ornamentals because of their col- ourful fruits and fall foliage and their unusual winged stems. At least six species are recorded from Canada, of which one is regarded as doubtful (Scoggan 1978; Catling 1997; Hinds 2000). Only E. occidentalis Nutt. ex Torr. (Western Wahoo), E. atropurpurea Jacq. (East- em Wahoo) and E. obovata Nutt. (Running Strawberry- Bush) are native in Canada, with E. occidentalis res- tricted to the west coast. Euonymus europaea L., E. fortunei (Turcz.) Hand.-Mazz., E. alata (Thun.) Sie- bold, and E. nana Bieb. are escapes from cultivation in eastern Canada, although they have restricted distri- butions in New Brunswick (Catling 1997; Hinds 2000). Other species occur in the northeastern United States (i.e., E. alata and E. americana L., Gleason and Cronquist 1991). No species is included in the flora of Nova Scotia (Zinck 1998). Euonymus europaea was first noted in Antigonish as a single, non-cultivated plant in 2001 in Antigonish County, Nova Scotia (DJG unpublished). In a sub- sequent survey during fall and early winter of 2005- 2006 (Garbary and Taylor 2007), fruiting plants of E. europaea were noted from the previously recognized site as well as additional sites in Antigonish. The spe- cies likely represents an escape from cultivation. Here We report E. europaea as a newly naturalized plant in Nova Scotia. Materials and Methods During the course of plant surveys in late fall and early winter of 2001 and 2005-06 (Taylor and Garbary 2003; Garbary and Taylor 2007), fruiting specimens of Euonymus europaea were noted. In May 2006, when flower buds were well developed but unopened, a sur- vey of all previously identified locations was under- taken. Further surveys were conducted to determine the size of the overall population in Antigonish and the extent to which the species was reproducing. The four primary sites are designated as University, Clay- more, Hospital, and Brierly Brook (see Table 1). Two additional sites with only 1-2 primary plants were found but are not included in Table 1. Sites were surveyed with hand-held GPS units (Gar- min 12, Olathe, Kansas, USA). The largest stem in any cluster was considered the primary plant, and adjacent shoots were assumed to be derived from the same base, or progeny from seed. The height of the largest plant at each site was measured with a meter stick, and the numbers of subsidiary shoots and nearby juveniles were counted. These data were gathered to demonstrate that plants were successfully reproducing in the area. For each primary plant, the slope, exposure to full sun- light, proximity to water (ditches, ponds or saturated soil) and reproductive condition were recorded. The number of flower buds on each peduncle was counted on the population on St. Francis Xavier University. Voucher specimens are deposited in the herbarium of St. Francis Xavier University (STFX) with duplicates deposited in herbariums of ACAD (Acadia University) and NSPM (Nova Scotia Provincial Museum). Results With its opposite leaves on short petioles, the longi- tudinal whitish markings on many young stems that often gives them a four-angled outline, and the bril- liant scarlet and orange fruits, Euonymus europaea 1s a distinctive species during all seasons (Figure 1). Most primary shoots (> 95%) and many adjacent smaller plants had abundant clusters of flower buds in mid- to late May 2006. The bud scales of branch buds and developing inflorescences had red tips and a ciliate 85 86 THE CANADIAN FIELD-NATURALIST Vol. 121 FIGURE 1. Euonymus europaea from Antigonish. A) Branch with leaves, developing inflorescences and bud scale (arrow). Scale = 2 cm. B) Inflorescence with central bud partially open. Scale = 1 cm. C) Fully open flower. Scale = 5 mm. D) Mature fruit after dehiscence showing single seed. Scale = 1 cm. E) Longitudinal view of stem showing leaf scar and longitudinal ridge on bark (arrow). Scale = 2 cm. F) Transverse section of stem showing squarish outline and position of longitudinal ridges. Scale = 5 mm. margin. Flowers did not open until late May when a single bud in each peduncle opened. The flowers are not conspicuous, and plants are cryptic against a back- ground of the larger and showier species of Prunus, Crataegus and Amelanchier. The flowers are ca. 1 cm diameter, and have diagnostic greenish-white petals. The flowering peduncles had from 1-9 buds (5.0 + 1.8), with most inflorescences having odd numbers of buds. Over 100 primary plants over 50 cm high and over 650 secondary shoots and adjacent progeny were found at six locations in Antigonish (Table 1). The primary axes were 1.5 + 0.8 m (mean + s.d.) tall, and many plants were over 3 m (Figure 2) with a trunk diameter of up to 6 cm. There was wide variation (0 — >50) in number of secondary shoots and progeny associated with each primary plant, although about 30% had 2007 Frequency o HE Din EASES laiae Bian ee os | spr es sree ht ae Shy YO SI NPI oR <= EN oe =e! | Set eatin VA cA Plant height (m) FIGURE 2. Histogram showing sizes of 112 primary stems of Euonymus europaea from Antigonish. only 1-2 additional axes (Figure 3). The largest single clump was at the Brierly Brook site. It occupied about 4 x 3 m and had about 50 separate stems that formed a dense thicket. Plants were typically found in the open where they experienced full sun, or at the edges of dense shrub thickets where they were either crowded or overtopped by the surrounding plants. Most plants grew in moist locations. Plants in the open tended to occur on bogg ground, and plants were common close to ditches, ponds or small pools or on the flood plain of a stream. Some plants were present on steep slopes on ground with obvious seeps. Only a few plants occurred on drier soils; these were in grass on mowed areas and may have been planted. GARBARY and DEVEAU: SPINDLE TREE IN NOVA SCOTIA 87 50 40 30 20 10 y Sy rt | Number of shoots FIGURE 3. Histogram showing number of secondary shoots and possible progeny associated with each primary plant. Discussion All of the Euonymus plants we observed were E. europaea. The plants are scattered over several square kilometres in at least five separate sites. The abundance of small plants within 3 m of larger individuals and widely scattered small plants (1.e., less than | m) show that successful seed germination is occurring. These observations are consistent with the definition of na- turalized of Pysek et al. (2004); i.e., that successful reproduction has been occurring for at least ten years in the vicinity of the parent plant. The site where most of the plants were found is an open field where EF. europaea is mixed with many other shrubs. These shrubs are mostly widely scattered and TABLE |. Populations of Euonymus europaea in Antigonish. Additional sites with only one or two primary plants not included in the table. Hospital Latitude and longitude 45°37.743'N 61°59.033'W Number of primary plants 12 Secondary shoots and juveniles 61 Number with flower buds 11/12 Habitats Steep slope with extensive bed of lupins; mowed and unmowed grassy area Shade Full sun Claymore Brierly Brook 45°37.056'N 45°37.394'N 61°58.877'W 61°59.179'W 4 5 11 95 4/4 5/5 Shrub thicket Border of beside hayfield at small stream margin of thicket adjoining stream Partial to heavy shade Partial to high shade University 45°37.074'N 61°59.182'W 90 683 87/90 Open areas with herbaceous plants and shrubs: on flat ground to steep slopes Full sun but some heavily shaded 88 THE CANADIAN FIELD-NATURALIST have an overall cover of less that 15%. The occasional individuals that we observed in dense shrubbery had few secondary shoots. Thus, once vegetation closes in, E. europaea may not be able to establish or spread. Species with fleshy fruits have low ability to recruit under a canopy of E. europaea relative to nine other tall shrub species (Kollmann and Grubb 1999). This has not yet become important in Antigonish, since we have observed only one continuous canopy of E. europaea. Some Euonymus species, including E. alata, have become invasive in several northeastern and mid- western states (e.g., http://tncweeds.ucdavis.edu/alert/ alrteuon.html) following escape from ornamental plantings. FE. alata is considered a threat in woodland areas, fields and coastal scrubland, where it can out- compete native species. There is no record of E. europaea as a problematic invasive species, although it has become widely naturalized in central Canada. Montgomery (1957) refers to sites where it has be- come “very common” in woods. The abundance of mature and immature individuals in Antigonish sug- gests that this is possible for E. europaea in Nova Scotia. The extensive field survey carried out by DJG in the fall and early winter of 2005-2006 identified four of the sites reported here. Additional surveys in 2006- 2007 did not reveal further populations. While there are no specimens based on wild collections of E. europaea in other local herbaria (1.e., NSPM, ACAD, NSAC [Nova Scotia Agriculture Canada]), other urban areas may be expected to host wild plants (e.g., Wolf- ville, Nova Scotia — Sam Vander Kloet, personal com- munication). To satisfy the Pysek et al. (2004) defini- tion of invasive, many offspring must be produced at a distance from the parent individuals. The limited dis- persal of E. europaeus in Antigonish has not yet satis- fied this criterion. Acknowledgments We thank Greg Power, Kelly Clement and Meghan Hines for assistance with fieldwork, Randy Lauff who Vol. 121 pointed out the location of an additional Euonymus site, and Barry Taylor for comments on the manuscript. Curators of ACAD, NSPM and NSAC provided in- formation and access to collections. This research was supported by research grants from the Natural Sci- ences and Engineering Research Council of Canada to DJG. Literature Cited Catling, P. M. 1997. The problem of invading alien trees and shrubs: some observations in Ontario and a Canadian checklist. Canadian Field-Naturalist 111: 338-342. Garbary, D. J., and B. R. Taylor. 2007. Plant flowering dur- ing January in Antigonish County, Nova Scotia. Canadian Field-Naturalist 121(1): 76-80. Gleason, H. A., and A. Cronquist. 1991. Manual of vascular plants of Northeastern United States and adjacent Canada, 2™4 edition. New York Botanical Garden, Bronx, New York. Hinds, H. R. 2000. Flora of New Brunswick. University of New Brunswick, Fredericton, New Brunswick. 695 pages. Kollmann, J., and P. J. Grubb. 1999. Recruitment of fleshy- fruited species under different shrub species: control by under-canopy environment. Ecological Research 14: 9-21. Montgomery, F. H. 1957. The introduced plants of Ontario growing outside of cultivation. Transactions of the Royal Canadian Institute 32: 3-35. Pysek, P., D. M. Richardson, M. Rejmanek, G. L. Webster, M. Williamson, and J. Kirschner. 2004. Alien plants in checklists and floras: towards better communication be- tween taxonomists and ecologists. Taxon 53: 131-143. Scoggan, H. J. 1978. The Flora of Canada Part 3 — Dicotyle- donae (Sauruaceae to Violaceae). National Museum of Natural Sciences, National Museum of Canada, Ottawa. 568 pages Taylor, B. R., and D. J. Garbary. 2003. Late-flowering plants from northern Nova Scotia. Rhodora 105: 118-135. Zinck, M. Editor. 1998. Roland’s Flora of Nova Scotia, 3" edition Nimbus Publishing Ltd. and Nova Scotia Museum, Halifax, Nova Scotia, Canada. 1297 pages. Received 16 June 2006 Accepted 7 March 2008 Notes Serviceberry, Amelanchier intermedia, Escaped from Cultivation in Eastern Ontario PAUL M. CATLING and GISELE MITROW Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Saunders Building, Central Experimental Farm, Ottawa, Ontario KIA 0C6 Canada; e-mail: catlingp @agr.ge.ca Catling, Paul M., and Gisele Mitrow. 2007. Serviceberry, Amelanchier intermedia, escaped from cultivation in eastern Ontario Canadian Field-Naturalist 121(1): 89-91. Although not generally recognized as a cultivated plant, Amelanchier intermedia is widely cultivated in eastern Ontario under the incorrect name, A. canadensis. Wild plants referable to A. intermedia in old fields south of Ottawa are likely to be escaped from cultivation. Amelanchier intermedia likely originated as a hybrid involving A. canadensis and A. laevis. A key to the Canadian species of Amelanchier with glabrous ovaries is provided. Key Words: Saskatoon, Juneberry, Serviceberry, Shadbush, Amelanchier intermedia, status, cultivation, hybrid, Ontario. Fernald (1950) and Kartesz and Meachum (1999) reported Amelanchier intermedia Spach from New- foundland to Minnesota and south to North Carolina. It was not included in Michigan by Voss (1985), but Cing-Mars (1971) showed a relatively extensive distri- bution in southern Quebec. Although within the gen- eral range outlined by Fernald (1950) and frequent in adjacent Quebec (Cingq-Mars 1971), A. intermedia was ig reported in Ontario by Soper and Heimburger (1982) or McKay (1973). Although it was listed for Ontario by Newmaster et al. (1998*), the listing was without ranking indicating a lack of information on its ‘status. Over several years of examining populations of Amelanchier in Ontario we have found plants that cor- ‘respond to the description of A. intermedia at only two locations 1 in old field habitats south of Ottawa. Plants similar to these wild plants are widely cultivated but are never called A. intermedia. ' Most eastern Ontario nurseries offering shrubs for sale offer Serviceberry (also called Juneberry, Shad- bush and Saskatoon, Amelanchier spp.). The plants that are most frequently available are either multi-stemmed ‘shrubs labelled as A. canadensis or they are named cultivars referable to A. xgrandiflora Rehder, a hybrid of A. arborea (Michx. f.) Fern. and A. laevis Wieg. The plants referred to as Amelanchier canadensis (L.) ‘Medic. are particularly popular with companies pro- ‘viding landscaping for office buildings, whereas the larger small trees are increasingly popular for use as ‘Street trees under utility wires (Gerhold 1999). “> Identification of wild and similar cultivated ' plants __ With petals 9-13 mm long and reddish-tinged leaves prominently acute at the tip, the plants growing in old os fields and woodland edges south of Ottawa (cited be- low) and similar plants labelled as A. canadensis in 10 local nurseries as well as many plants in Ottawa gar- dens could not be placed with A. canadensis, which has petals 3-10 mm long and green young leaves with leaf tips more or less rounded (Cruise 1964; Cing-Mars 1971). They are not A. /aevis because the leaves are tomentose at flowering instead of glabrous. Nor are they A. arborea because these are multi-branched shrubs with reddish-tinged leaves and flowers with a rela- tively broad hypanthium 4.5—5.5 mm across instead of single-trunked trees with green leaves and flowers with a relatively narrow hypanthium 2.5—3 mm across. They also differ from A. arborea in having sepals that are less reflexed and lanceolate with a concave outer edge instead of strongly reflexed at the base and oblong- lanceolate with the outer edge convex. In addition the berries of these shrubs are pleasant-tasting rather than insipid as in A. arborea. These plants do correspond to A. intermedia Spach of Fernald (1950) and Cing-Mars (1971). While it is clear that A. intermedia occurs both as a cultivated plant and outside of cultivation in eastern Ontario, it is not clear whether or not the wild plants are escapes from cultivation. It seems most likely that they are escaped since one of the parents, A. canaden- sis, does not occur in the local area and the similar widely cultivated plants have become increasingly pop- ular over the past few decades and are currently widely available in nurseries and widely used by landscaping companies. A key to the group with glabrous ovaries is provided below. Specimens referable to A. intermedia growing out- side of cultivation in Ontario include: OTTAwA-CAR- LETON: open, sandy ground 3 km N of Herbert’s Cor- 89 90 THE CANADIAN FIELD-NATURALIST ners, Osgoode Twp., 45.2359°N, 75.5733°W, 17 May 2002, P. M. Catling s. n. (DAO); UNITED COUNTIES OF STORMONT, DUNDAS AND GLENGARRY: open old field, 2 km WNW of Ormond and about 5 km N of Winches- ter at 45.1552°N, 75.4173°W, 21 May 2005, P. M. Catling 2005-41, 2005-42 (DAO). Cultivated speci- mens referable to A. intermedia collected as part of this study in Ottawa area gardens are also preserved in DAO (see DAO 798819, 801264, 801265, 801272, 801275, 801280, 801328, 801329, 801330, 801332) and vouch- ers obtained from nurseries are also in the collection (see DAO 800923, 800924, 800925, 801262, 801267, 801269, 801270, 801271, 801333, 801335). Taxonomic history and possible hybrid origin of A. intermedia Jones (1946) included the name intermedia with his relatively narrowly defined A. canadensis showing a largely eastern coastal plain distribution. Wiegand (1920) used the name to represent a species related to A. canadensis which he defined narrowly. Later Fer- nald (1950) expanded the concept to include plants clearly intermediate between A. canadensis and A. lae- vis and this concept was maintained in the classic work of Cing-Mars (1971). Amelanchier intermedia may have arisen as a hybrid of A. canadensis and A. laevis as noted by various authors (e.g., Gleason and Cronquist 1991). Plants referable to A. intermedia have leaves that are only moderately pubescent and losing their pubescence at flowering. This and their purplish tinge suggests that one parent is a species with leaves purple-green and glabrous and unfolded at flowering. The only species with a glabrous ovary fitting this description is A. laevis. The relatively short petal lengths and also the widely open hypanthium of the putative hybrids suggest that the other parent may have short petals, barely reflexed sepals and a saucer-shaped hypanthium. This is true of A. canadensis but not A. arborea which has relatively long petals, prominently reflexed sepals and a rather narrow, campanulate hypanthium. Furthermore a cross of A. arborea and A. laevis would be expected to pro- duce a more tree-like hybrid, one form of which, A. xgrandiflora Rehd. is well known. Other species of Amelanchier have pubescent ovary summits and/or rounded and more coarsely toothed leaves and are thus excluded from consideration as putative parents. Ame- lanchier canadensis x laevis hybrids have been report- ed to be common in some areas, whereas hybrids of A. arborea and A. canadensis are less common (Cruise 1964). The only material that can be confused with Ame- lanchier interior that occurs in the lower Ottawa valley are depauperate specimens of A. arborea which differ in their early flowering with unexpanded and very hairy leaves and apparent hybrids of A. /aevis and A.spicata Vol. 121 which differ in their broad spatulate petals and partially hairy ovaries. The typical habitat of these latter plants is dried and disturbed heath bogs where A. spicata also occurs. Such plants may have been the basis for reports of A. intermedia from edges of bogs in Minnesota (Nielsen 1939). Alternatively these reports may have been based on fruiting material of A. interior of the northwestern Great Lakes region which had lost some of the pubescence on the ovary (as found to be the case for reports of A. intermedia from northwestern Ontario). The following key will readily separate flowering A. intermedia from related Canadian species. Key to Canadian flowering Amelanchier with glabrous ovaries: la. Leaves glabrous and purple at flowering; petals 10-20 mm long; lowest pedicels 25-S 0 MMVON Sees eto Aesth eee A. laevis 1b. Leaves tardily pubescent and green or purple-tinged at flowering; petals 3-15 mm long; lowest pedicels 8-35 mm long 2a. Petals 3-10 mm long; leaves FOUNG=(ippPed eee eee A. canadensis 2b. Petals 9-15 mm long; leaves pointed at tip ......... B) 3a. Leaves purple-tinged, more or less expanded and becoming glabrous beneath during flowering; SHEUDS 4c SHES as eT Ee, se A. intermedia 3b. Leaves green, small and folded at flowering, pubescent beneath after flowering; trees ...A. arborea Acknowledgments Brenda Kostiuk assisted with field studies. W. J. Cody provided useful comments on the manuscript. Document Cited (marked * in text) Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. McMurray, and M. J. Oldham. 1998. Ontario plant list. Ontario Forest Research Institute, Ontario Ministry of Natural Resources (Sault Ste. Marie, Ontario), Forest Research Information Paper 123. www.uoguelph.ca/foibis. Literature Cited Cing-Mars, L. 1971. Le genre Amélanchier au Québec. Le Naturaliste canadien 98: 329-346. Cruise, J. E. 1964. Studies of natural hybrids in Amelanchier. Canadian Journal of Botany 42: 651-663. Fernald, M. L. 1950. Gray’s manual of Botany. American Book Company, New York. 1632 pages. Gleason, H. A., and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2"4 edition New York Botanical Garden, Bronx, New York. 993 pages. Gerhold, H. D. 1999. Serviceberry cultivars tested as street trees: initial results. Journal of Arboriculture 25(4): 189- 192. Jones, G. N. 1946. American species of Amelanchier. Illinois Biological Monographs 20(2): 1-126. Kartesz, J. T., and C. A. Meachum. 1999. Synthesis of the North American flora. Version 1.0. Biota of North America Program, University of North Carolina, Chapel Hill. 2007 CHUBBS and PHILLIPS: A TRIBUT! rO NEAL PHILIP PERRY SIMON, 197 42006 FiGurE 1. Flowering inflorescence of Amelanchier intermedia showing relatively long petals and pedicels and leaves pubescent below and more or less expanded at flowering time. Photo of a cultivated plant in Ottawa by P. M. Catling. McKay, S. M. 1973. A biosystematic study of the genus Amelanchier in Ontario. M. Sc. thesis, University of Toron- to. 241 pages. Nielsen, E. L. 1939. A taxonomic study of the genus Ame- lanchier in Minnesota. American Midland Naturalist 22: 160-205. Soper, J. H., and M. L. Heimburger. 1982. Shrubs of Ontario. Royal Ontario Museum Publications in Life Sciences. 495 pages. Voss, E. G. 1985. Michigan flora, part I. Dicots (Saururaceae — Cornaceae). Cranbrook Institute of Science Bulletin 59. 724 pages. Wiegand, K. M. 1920. Additional notes on Amelanchier. Rhodora 22: 146-151. Received 15 October 2005 Accepted 23 February 2007 9] 92 THE CANADIAN FIELD-NATURALIST Vol. 121 A Freshwater Hydrobiid, cf. the Squat Duskysnail, Lyogyrus granum (Mollusca), Widespread in the Hampton Marsh, New Brunswick DONALD F. MCALPINE!, RACHEL H. MAILLET!, ANDREW J. ALBERT!, LAURA M. CROSSMAN!, REBECCA R. SmitH!, and ANDRE L. MARTEL? 'New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada 2Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada McAlpine, Donald F., Rachel H. Maillet, Andrew J. Albert, Laura M. Crossman, Rebecca R. Smith, and André Martel. 2007. A freshwater hydrobiid, cf. the Squat Duskysnail, Lyogyrus granum (Mollusca), widespread in the Hampton Marsh, New Brunswick. Canadian Field-Naturalist 121(1): 92-94. In Canada, the Squat Duskysnail, Lyogyrus granum, is an apparently rare freshwater hydrobiid. We document putative L. granum to be widespread throughout the Hampton Marsh, New Brunswick, with densities sometimes exceeding 150 snails/m?* on substrates ranging from vegetated organic detritus over mud, to sparsely-vegetated cobble/sand lakeshore. That our identi- fication of L. granum from Hampton Marsh remains tentative, in spite of large samples, emphasizes the need for detailed taxonomic study of any putative L. granum populations in Canada. Such study will be necessary before the true conservation status can be determined for this and any related taxa, as yet unrecognized in Canadian material. Key Words: Squat Duskysnail, Lyogyrus granum, Hydrobiidae, conservation status, New Brunswick. Of the two species of freshwater hydrobiid snails recorded in Maritime Canada (Clarke 1981), the Ordi- nary Spire Snail, Amnicola limosa, and the Squat Duskysnail, Lyogyrus granum, the latter is very poorly known. In part, this is due to difficulties in identifica- tion; Smith (1991) notes that in Massachusetts L. granum and L. pupoidea will rarely be confidently distinguished from each other and Jokinen (1992) has reported sexual dimorphism within the genus that is poorly understood. In Canada, L. granum has been reported from only two locations in south-central New Brunswick (Jem- seg and Baker Brook) and 11 sites in Nova Scotia (COSEWIC 2003*). COSEWIC (2003*) suggested that L. granum is rare in Canada and that the only known Canadian populations in New Brunswick and Nova Scotia should be considered nationally significant. Nonetheless, a paucity of information on this species has led to a designation of Data Deficient by the Com- mittee on the Status of Endangered Wildlife in Canada (COSEWIC). Here we document a L. granum-like hydrobiid to be abundant and widespread in the Hamp- ton Marsh, New Brunswick. Considered one of the largest and most diverse fresh- water marshes in the Saint John River floodplain, the 2000 hectare Hampton Marsh (45°33'N, 65°50'W) con- sists of a series of eutrophic oxbows, tributaries and backwaters of the Kennebecasis and Hammond Rivers. The marsh has repeatedly been identified as a site of ecological importance within the province (Stocek 1984*: Dionne et al. 1988; Austin-Smith 1994*) and is currently designated as a wetland of significance by the Wetlands and Coastal Habitat Program of the New Brunswick Department of Natural Resources and Ener- gy. Although about 40% of the land adjacent to the marsh remains forested (Austin-Smith 1994*), shore- line development, sedimentation, eutrophication, and pesticide introduction from upstream agricultural areas are issues of concern. The marsh is located largely with- in the municipal boundaries of Hampton and Quispam- sis, two of the fastest growing urban areas in New Brunswick, and the Local Service District of Nauwige- wauk (Austin-Smith 1994*). Putative L. granum were discovered in the Hampton Marsh during the course of ongoing mollusc surveys on the lower Saint John River and tributaries (Sabine et al. 2004; McAlpine et al. 2005). Data reported here were collected from 20 June to 22 August 2005 from 25 survey sites scattered across the marsh. Survey sites had been selected based on study of aerial photos, topographic and nautical maps, and field inspection, to represent a variety of habitat types (i.e., lakeshore, river channel, backwaters, oxbows) within the marsh complex. Molluscs were collected using a combina- tion of techniques that included SCUBA, snorkeling, dip netting, and sieving bottom sediments; voucher material of all mollusc species reported here has been placed in the New Brunswick Museum mollusc col- lection. To provide abundance estimates for putative L. granum in contrasting habitats within the marsh, 15 0.25-m? random quadrats were also sampled in ~ | m of water in August at three of the sites. Only live snails were tabulated in quadrats following the com- plete removal, sieving, and hand-sorting of the top 5 cm of sediment along with all aquatic vegetation. Hampton Marsh specimens of Lyogyrus were identified largely on the basis of the structure of the verge in fresh mate- rial as described and illustrated in Smith (1991) and Jokinen (1992), but reference was also made to shell and opercular morphology as noted by these authors, and by Clarke (1981). As well, comparison was made with Atlantic Canadian material in the collection of the Canadian Museum of Nature. Although L. pupoidea has not been recorded in Canada, our extensive series of specimens agree with Smith (1991), in that inter- mediates and extremes showing external morpholog- 2007 NOTES 93 TABLE |. Densities of putative Lyogyrus granum in contrasting habitats in the Hampton Marsh, New Brunswick GPS N 45° 30.025' W 65° 52.50' N 45° 31.358' W 65° 52.867' N 45° 31.856! W 65° 51.297’ Habitat rocky lakeshore, clearwater Ceratophyllum marsh murky backwater channel Mean + sd(/m*) 96.8 + 52.6 76.0 + 63.9 17.6 + 12.5 Range (/m*) Associated molluscs 48-184 Ai, Al, Cd, Ec, Gd, Ha, Lr, Pc, Ph, Sc, Se, Vt, 4-160 Ai, Al, Ec, Gd, Ha, Pa, Ph, Se, Ss, Vs, Vt 8-20 Al, Cd, Ec, Fp, Gd, Pe, Ph, Se, Ss. Vs, Vt ' Ai=Anodonta implicata, Al\=Amnicola limosa, Cd=Campeloma decisum, Ec=Elliptio complanata, Fp=Ferrissia parallela, Gd=Gyraulus deflectus, Ha=Helisoma anceps, Pa=Planorbella campanulata, Lr=Lampsilis radiata, Pe=Pyganodon cataracta, Ph=Physa heterostropha, Ss=Sphaeriidae sp., Sc=Stagnicola catascopium, Se=Stagnicola elodes, Vs=Valvata sincera, Vt=Valvata tricarinata ical features of L. pupoidea are present. COSEWIC (2003*) notes the possibility of L. pupoidea, as well as closely related L. browni, overlapping the Canadian range for L. granum. Clearly there is a need for detailed taxonomic study of any putative L. granum populations in Canada before the true distribution and conservation status of this and related species in the country can be determined. Putative L. granum are widespread throughout the Hampton Marsh, although they were noticeably ab- sent from sites in the river channel, where currents of 0.1-0.3 m/sec were recorded, and in the lowest por- tion of the marsh where there is periodic intrusion of brackish water. Discounting two sites where brackish water clearly intrudes on occasion, we recorded L. granum at 17 of the 22 survey sites remaining. The limited habitat data available for Lyogyrus indicates that this species is generally found on organic debris and vegetation in standing water (Clarke 1981; Smith 1987*: Jokinen 1983). Currents at sites where we re- corded Lyogyrus did not exceed 0.1 m/sec. COSEWIC (2003*) reports L. granum densities of up to 400/m”; densities in the Hampton Marsh were lower but some- times exceeded 150/m? (Table 1). We found the highest densities along an open, sparsely-vegetated cobble/sand lake shore on rocks and on the living shells of the unionids Elliptio complanata, Anodonta implicata and Lampsilis radiata. Lyogyrus granum was also abundant in clear water on submersed vegetation where Cerato- phyllum demersum dominated (lakeshore vs clear water; Mann-Whitney U = 15, P > 0.05) but signifi- cantly less abundant in a murky vegetation-clogged backwater channel characterized by Vallisneria amer- icana and Potamogeton perfoliatus than on the rocky lake shore (lake shore vs backwater; Mann-Whitney U=25, P < 0.05) (Table 1). Both of the more heavily vegetated sites were characterized by deep bottom layers of organic detritus over mud. COSEWIC (2003*) suggests that the few sites from which the species has been recorded in Canada indi- cate either that L. granum is a habitat specialist, that human impacts are reducing the availability of suit- able habitat, that L. granum is a truly rare species, or some combination of these factors. That the rarity of this species may only be apparent, due to a lack of col- lecting, is not mentioned. Jokinen (1983, 1992) found this snail to be uncommon in Connecticut, and both Smith (1987*) and Jokinen (1992) found the species to be rare in Massachusetts and New York, respectively. While L. granum may indeed prove to be naturally rare, this species is small and easily overlooked and there have been few detailed surveys of the freshwater gas- tropods of Atlantic Canada. That L. granum has been recorded from 11 widely distributed sites across Nova Scotia also suggests that further field investigation may show this species, in association with related taxa as yet unrecognized in Canada, to be more prevalent in the region than current records indicate. Acknowledgments Financial support for this project was provided through grants to DFM from the Community-Univer- sity Research Alliance program of the Social Sciences and Humanities Research Council of Canada, The Sala- mander Foundation, the New Brunswick Wildlife Trust Fund, and the Province of New Brunswick Summer Mentorship Program for Female Students. Laurie Mills and James Wilson of the Hampton Area Environmen- tal Group assisted with field arrangements while Archie and Connie Downey were unfailingly gracious in pro- viding access to the marsh through their property and providing use of a wharf and other amenities; Gart Bishop and Geoffry McBriarty assisted with the field sampling of molluscs. Documents Cited (marked * in text) Austin-Smith, P. 1994. Hampton-Kennebecasis Marsh Com- plex — Status Report. Town of Hampton, New Brunswick, 99 pages. COSEWIC. 2003. Assessment and update status report on the Squat Duskysnail, Lyogyrus granum, in Canada. Com- mittee on the Status of Endangered Wildlife in Canada. Ottawa. [unpublished report]. vi + 27 pages. Smith, D. G. 1987. Keys to the freshwater invertebrates of Massachusetts. Number 2. Mollusca Mesogastropoda (oper- culate snails). Massachusetts Department of Environmental Quality Engineering, Division of Water Pollution Control (Westborough). 34 pages. 94 THE CANADIAN FIELD-NATURALIST Stocek, R. 1984. Environmentally significant areas in the Saint John Planning Region. Environmental Services Branch, Environment New Brunswick, Fredericton, New Brunswick. 116 pages. Literature Cited Clarke, A. H. 1981. The freshwater molluscs of Canada. National Museum of Natural Sciences, National Museums of Canada, Ottawa. 446 pages. Dionne, L. A., D. M. Farmer, and M. C. Young. (Compilers). 1988. Critical natural areas in New Brunswick. New Hori- zons Critical Natural Areas Committee, Fredericton, New Brunswick. 313 pages. Jokinen, E. H. 1983. The freshwater snails of Connecticut. State Geological and Natural History Survey of Connecti- cut. Department of Environmental Protection Bulletin 109: 1-83. Vol. 121 Jokinen, E. H. 1992. The freshwater snails (Mollusca: Gas- tropoda) of New York State. New York State Museum Bul- letin 482. 112 pages. McAlpine, D. F., D. L. Bateman, and C. A. Davis. 2005. Spurwinkia salsa (Pilsbry 1905) (Gastropoda: Hydrobi- idae) in the Kennebecasis River estuary, New Brunswick: A brackish water snail new to Canada. Journal of Conchol- ogy 38: 602-604. Sabine, D. L., S. Makepeace, and D. F. McAlpine. 2004. The Yellow lampmussel (Lampsilis cariosa) in New Bruns- wick: A population of significant conservation value. North- eastern Naturalist 11: 407-420. Smith, D. G. 1991. Keys to the freshwater macroinverte- brates of Massachusetts. Department of Zoology, Univer- sity of Massachusetts, Amherst, Massachusetts. 236 pages. Received 27 February 2006 Accepted 5 November 2007 Northern Range Extension of the Pygmy Shrew, Sorex hoyi, in the Yukon THOoMaAs S. JuNG!, TRoy D. PRETZLAW2, and Davip W. NAGORSEN? 'Fish and Wildlife Branch, Yukon Department of Environment, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada; e-mail: thomas.jung @ gov.yk.ca "Department of Natural Resource Sciences, McGill University, 21111 Lakeshore Blvd., Montreal, Quebec H9X 3V9 Canada 3Mammalia Biological Consulting, 4268 Mechosin Road, Victoria, British Columbia V9C 3Z4 Canada Jung, Thomas S., Troy D. Pretzlaw, and David W. Nagorsen. 2007. Northern range extension of the Pygmy Shrew, Sorex hoyi, in the Yukon. Canadian Field-Naturalist 121(1): 94-95. A Pygmy Shrew, Sorex hoyi, was captured in a pitfall trap on the Blackstone River (65°04.6'N, 138°10.8'W) in central Yukon. This represents a northern range extension of about 110 km for S. hoyi in the Yukon. Key Words: Distribution, Pygmy Shrew, Sorex hoyi, Yukon. Shrews, Sorex, have been undersampled in north- western North America because they are often not the focus of surveys (Jarrell 1986; Nagorsen 1996) and they are not readily captured in traps designed for small rodents (Nagorsen 1996). As such, our knowledge of the distributions of shrews in northwestern North Amer- ica (i.e., Alaska, Yukon, northern British Columbia, western Northwest Territories) is limited. In recent years, however, directed surveys of shrews in this vast region have resulted in discovery of a new species, the Alaskan Tiny Shrew, Sorex yukonicus (Dokuchaev 1997), and range extensions for a number of species (e.g., American Water Shrew, S. palustris, Jarrell 1986; Cook et al. 1997; Tundra Shrew, S. tundrensis, Nagors- en and Jones 1981; Alaskan Tiny Shrew, S. yukonicus, and Pygmy Shrew, S. hoyi, Peirce and Peirce 2000). Nevertheless, much work is needed to better understand the distributions of shrews in northwestern North Amer- ica. Herein, we describe a northern range extension for S. hoyi in the Yukon. During an extensive survey of the small mammal fauna along the Dempster Highway in central Yukon, a gmy Shrew was captured on 12 July 2005, 2 km west of the Blackstone River (65°04.6'N, 138°10.8'W), 129 km northeast of Dawson City. The shrew was cap- tured in an unbaited pitfall trap installed flush with the substrate. Pitfall traps have been found to be particu- larly effective for Pygmy Shrews (Prince 1941). No other shrews were captured at this site. The specimen was identified using dental characteristics and keys in Nagorsen (2002) and van Zyll de Jong (1983). The specimen (Field ID: DEMP-0015) is held by the Yukon Department of Environment, but will be deposited at the Museum of Southwestern Biology. Our record represents a northern range extension of approximately 110 km for S. hoyi in the Yukon. Youngman (1975) mapped the hypothetical range as being as far north as the Old Crow Flats (approximate- ly 68°N). Prior to our capture, however, the northern- most specimen records of S. hoyi in the Yukon were from 22 km east of Dawson City (Youngman 1975; van Zyll de Jong 1983) and the abandoned village of Forty- mile (64°25'N, 140°32'W), 67 km northwest of Dawson City (B. G. Slough and T. S. Jung, unpublished data). This Yukon range extension of S. hoyi is not surpris- ing as there are more than 125 specimens held at the University of Alaska Museum of the North that were collected north of 65°N in Alaska, with the most northerly taken near the Dalton Highway at the Diet- rich River (67°37'N, 149°46.8'W; UAM 23050). In 2007 addition, van Zyll de Jong (1983) reported specimens from Chick Lake and Fort Franklin, two locations north of 65°N in the Northwest Territories. It is likely that S. hoyi ranges further north in the Yukon; this is a species of the boreal forest and it likely extends north to the treeline. Further observations are needed to doc- ument the range of S. hoyi and other soricids in the Yukon and elsewhere in northwestern North America (Cook et al. 1997). Acknowledgments We thank Kevin Lake, Elise Bolduc, Kyle Russell, Kieran O’Donovan, Shannon Barker, Lea Randall, Brian Bell, Sara Nielsen, Michael Smith, Becky Cad- sand and Matthias Clyde for able assistance in the field or lab. Funding for this work was provided by McGill University, the Yukon Department of Environment, and NatureServe Yukon. We thank Murray Humphries, Julie Frisch and Gordon McRae for logistical support while on the Dempster Highway. Literature Cited Cook, J. A., C. J. Conroy, and J. D. Herriges, Jr. 1997. Northern record of the water shrew, Sorex palustris, in Alaska. Canadian Field-Naturalist 111: 638-639. Dokuchaey, N. E. 1997. A new species of shrew (Soricidae, Insectivora) from Alaska. Journal of Mammalogy 78: 811- 817. NOTES 95 Jarrell, G. H. 1986. A northern record of the water shrew, Sorex palustris, from the Klondike River, Yukon Territory Canadian Field-Naturalist 100: 39) Nagorsen, D. W. 1996. The mammals of British Columbia Volume 2: Opossums, shrews and moles of British Colum- bia. Royal BC Museum Handbook. UBC Press, Vancouver 169 pages. Nagorsen, D. W. 2002. An identification guide to the small mammals of British Columbia. Ministry of Sustainable Resource Management, Ministry of Water, Land, and Air Protection, Biodiversity Branch, and Royal BC Museum 153 pages. Nagorsen, D. W., and D. M. Jones. 1981. First records of the tundra shrew (Sorex tundrensis) in British Columbia. Canadian Field-Naturalist 95: 93-94. Peirce, K. N., and J. M. Peirce. 2000. Range extensions for the Alaska tiny shrew and pygmy shrew in southwestern Alaska. Northwestern Naturalist 81: 67-68. Prince, L. P. 1941. Water traps capture the Pygmy Shrew (Microsorex hoyi) in abundance. Canadian Field-Naturalist Se: van Zyll de Jong, C. G. 1983. Handbook of Canadian mam- mals. 1: Marsupials and insectivores. National Museums of Canada, Ottawa. 210 pages. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museums of Canada, Ottawa, Ontario. Publica- tions in Zoology 10: 1-192. Received 25 January 2006 Accepted 15 January 2008 A Tribute to Neal Philip Perry Simon 1973-2006 Tony E. Cuupss! and FRANK R. PHILLIPS? 'Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and Labrador AOP 1S0 Canada; e-mail: techubbs @cablelab.net *Department of Natural Resources, Government of Newfoundland and Labrador, Box 175, Station C, Happy Valley-Goose Bay, Newfoundland and Labrador, Newfoundland AOP 1CO Canada Chubbs, Tony E., and Frank R. Phillips. 2006. A tribute to Neal Philip Perry Simon, 1973-2006. Canadian Field-Naturalist 121(1): 96-99. Neal Philip Perry Simon, our friend, hunting com- panion, and colleague, was born in Labrador City, Newfoundland and Labrador, on Sunday 30 December 1973, to Dave and Doreen Simon. He passed away sud- denly and unexpectedly in a tragic accident on 23 Sep- tember 2006, at the age of 32 near Happy Valley — Goose Bay where he resided since 1988. His parents, Dave and Doreen Simon, sister Nicole and fiancée, Leanne Elson, survive him. Neal had a mutual adora- tion for his grandparents, Carrie and Gordon Temple- man, who will miss him dearly. Neal’s early education was in Catholic schools: Notre Dame Academy (primary and elementary) and Labra- dor City Collegiate (high school) both of which were in Labrador City and have since closed. Neal, along with his grade school pal Colin Carroll, developed a keen interest in biology at an early age, trying his hand at taxidermy in grade seven. Francis Schwab, Neal’s first year university professor, was instrumental in introduc- ing him to scientific research, especially the effects of post-fire succession on songbird abundance. His post- secondary education began at Memorial University of Newfoundland in St. John’s, Newfoundland and Lab- rador, where in 1995 he obtained a B.Sc.(Hons) with a major in ecology and evolution and a minor in statis- tics. It was there, under the instruction of Eric Baggs and the late Dr. Gary Cowen, he was instilled with an appreciation of the importance of small mammals in understanding forest structure dynamics. In 1996, Neal began his Master of Science in For- estry at the University of New Brunswick under the supervision of Dr. A. W. Diamond. As part of the Atlantic Cooperative Wildlife Ecology Research Net- work (ACWERN), Neal regularly contributed presen- tations at the annual workshop from 1998 to 2003. In 2000, Neal continued under the supervision of Dr. Diamond to pursue his Ph.D. on the effects of site productivity and heterogeneity on bird habitat quality and species richness. His research ranged from mod- elling species distributions to studying the effects of landscape features on habitat use by passerines, with a focus on their demography. Much of Neal’s work had an overlying conservation theme linked quite strongly to ecological theory. Neal completed his Ph.D. in 2005 and was about to embark on his post-doctoral research in the spring of 2007. Neal taught in various capacities at Memorial Uni- versity of Newfoundland, the University of New Bruns- wick and St. Mary’s University. From 1996 to 1998, Neal worked as a contract biologist for the College of | ' the North Atlantic in Labrador City and Happy Valley — — Goose Bay where he also offered his services as a university tutor in various disciplines, including chem- — istry and biology. Neal was always there to help out whenever or wherever he could and would always with- out hesitation say “Yes, just provide me the materials.” Students found him friendly and informative and they usually had few problems grasping concepts from him. He will be remembered for his willingness to help, his ability to teach a difficult content area and his friendly — personality. From November 1998, Neal was employed by the Provincial Government of Newfoundland and Labra- dor, Department of Natural Resources, occupying the position of Regional Ecologist for the Labrador Region. Neal’s work concentrated on experimental design, data collection, and analysis of research projects relating to forest management plans in Labrador. Neal’s research interests included effects of forest management and changing forest structures and specifically how these changes may affect the spatial and temporal distribu- tion of threatened Woodland Caribou (Rangifer taran- dus caribou) in central Labrador. Neal’s most recent interests involved sustainable forest management in) collaboration with the Government of Newfoundland: and Labrador, Department of Natural Resources, and) the Université du Québec 4 Montréal. Neal was about to embark on his post-doctorate research using inno- vative simulation models to ensure the sustainable man- agement of Labrador’s forests while his fiancée and co-; worker Leanne would be completing her Master of) Science in a supporting area of research. Neal was one¢ of those rare ecologists who not only recognize thes power of quantitative methods, but also actually under- stand how to appropriately apply these methods. Hav- ing just barely embarked upon his research career, Neal, as attested by his already growing list of publi- cations, was destined for greatness in the world of research and statistics. Since the summer of 2000, Neal had been an active participant in the North American Breeding Bird Sur- | vey (BBS) program and ran the Newfoundland anc: 96 2007 CHUBBS and PHILLIPS: A TRIBUTI rO NEAL PHILIP PERRY SIMON, 1973-2006 g 75 Ficure |. Neal P. P. Simon holding his first seal darted off Mulligan Point, 2 May 2003. (Photo courtesy Sarah Townley). Labrador, Happy Valley — Goose Bay survey route for Environment Canada. Neal was also a committed par- ticipant in the Christmas Bird Count (CBC). Neal was a dedicated member of both the local Squash League and Goose Bay Judo Club. He exem- plified the philosophy of Judo in all aspects of his life, striving to maximize his physical, intellectual and per- sonal character by doing his best; and being his best, Neal truly reflected this teaching through his physical strength and fitness, his academic accomplishments and desire to learn. As a competitor, Neal was intense and spirited while always maintaining his youthful grin and sense of humour. Neal’s enthusiasm for research in ecology and his jubilant attitude radiated from him, rapidly instilling in his colleagues a source of continual positive motivation. With an acute and intense sense of humour, Neal added a unique and welcome aspect to his interaction with both friends and fellow researchers. As an editor and reviewer of prospective manuscripts, his dedication and support were immeasurable and will be greatly missed. In 1999, Neal was an original member of the Lab- rador Wolverine Working Group, and became one of the founding members of the Labrador Woodland Cari- bou Recovery Team in 2001. Neal was a member of The Society for Conservation Biology, the Wildlife Society, the Atlantic Region of the Canadian Climate Impacts and Adaptation Research Network, the Atlan- tic Society of Fish and Wildlife Biologists, Atlantic Co-operative Wildlife Ecology Research Network 98 THE CANADIAN FIELD-NATURALIST (ACWERN), Newfoundland and Labrador Biodiver- sity working group, Forest Management Planning Team Member for Labrador Districts 19, 20, 21 and 22 and a member of the Institute for Environmental Monitoring and Research — Osprey Technical Com- mittee. Neal was about to join the Groupe de recherche en écologie forestiére interuniversitaire at 1’ Université du Québec 4 Montréal to complete his post-doctoral research developing habitat and timber harvesting- conservation trade-off models for Woodland Caribou. Neal’s interests were varied and included, but were not limited to, modelling species movements, predict- ing species distributions and understanding the effects of forest management on species occurrence and demo- graphy. He spent most summers in Labrador, along with his students and co-workers; in the field conducting songbird point count samples and vegetation plot counts. We came to know Neal and build a strong friendship when he moved to Happy Valley — Goose Bay in 1998 and shortly thereafer began work as the Ecosystem Ecologist for Labrador. Neal was friendly, energetic and an avid waterfowl and big game hunter. We anxiously awaited mid spring each year when we would travel by snowmobile over the bay ice to Portage Island, Mul- ligan Point. Here we would participate in the eons old traditional Inuit hunting practice of waiting patiently at breathing holes for Ringed Seals (Pusa hispida). Evenings would be filled with laughter as we swapped humorous exaggerated stories of past good times. Such wonderful times were set against the backdrop of a blazing driftwood fire and the assurance of deep slum- ber later in our traditional Labrador canvas tents heat- ed with the warmth of woodstoves. Neal often swapped related stories of how he had enjoyed the outdoors with his best hunting buddy, his father Dave, in West- ern Labrador. It was more important to Neal to enjoy the outdoors than to be successful on a hunt, though more often than not he was. As he so often mentioned to his hunting partner Colin Carroll, “If we don’t get anything today, it doesn’t matter. Just being here doing this is making my day.” His friends will best remember Neal as a shining, though comedic intellectual, with a love of life and a passion for the outdoors. Through discussions with some of Neal’s colleagues and friends in the prepara- tion of this tribute, we have found that Neal existed in several parallel universes, having the same mannerisms, characteristics and sharp wit despite the varied per- sonalities and context of his friends. He will be remem- bered for his comic additions to his work and conver- sation slipping easily between extreme seriousness and laughter. Acknowledgments We thank Neal’s parents (Dave and Doreen), his sister Nicole, fiancée Leanne, and others for personal information. Colin Carroll, Neal’s schoolmate and col- league provided information on Neal’s early schooling in Labrador City. We thank Tony Diamond for provid- Vol. 121 ing a compilation of Neal’s academic achievements. Matt Betts and Joe Nocera provided insight on Neal’s life that could only be shared between close colleagues and best friends. Bob Simms shared personal interac- tions with Neal through their judo club and Bernice Tracy provided details of Neal’s association with the College of the North Atlantic. Finally, we would like to thank all of Neal’s friends, family, colleagues and acquaintances that contributed to the Neal Simon Memorial Trust to fund an academic scholarship in Neal’s honour and memory. Neal appreciated the out- doors and enjoyed hunting and fishing and spending time in the Labrador wilds with Leanne and his dog, Boss. He will be greatly missed by his family, his many friends and his academic colleagues. Bibliography of Neal P. P. Simon Compiled by Tony E. CHUBBS Peer Reviewed Publications: Elson, L. T., N. P. P. Simon, and D. Kneeshaw. /n press. Regeneration differences between fire and clearcut logging in southeastern Labrador: a multiple spatial scale analysis. Canadian Journal of Forest Research: 000-000. Elson, L. T., and N. P. P. Simon. /n press. Plant abundances following clearcutting and stripcutting in central Labrador. Northern Journal of Applied Forestry, 00: 000-000. Elson, L. T., F. E. Schwab, and N. P. P. Simon. Jn press Winter food habits of willow ptarmigan (Lagopus lagopus) as a mechanism to explain winter sexual segregation. North- eastern Naturalist 00: 000-000. Roberts, B. A., N. P. P. Simon, and K. W. Deering. Jn press. The forests and woodlands of Labrador, Canada: Ecology, distribution, and future management. Ecological Research 00: 000-000. Schwab, F. E., N. P. P. Simon, S. Stryde, and G. Forbes. /n press. Effect of post-fire snag removal on breeding birds of western Labrador. Journal of Wildlife Management 00: 000-000. Schwab, F. E., N. P. P. Simon, and A. R. E. Sinclair. 2006. Breeding birds related to vegetation structure in southeast British Columbia. Journal of Wildlife Management 70: 189-197. Schwab, F. E., N. P. P. Simon, and S. Nash. 2005. Sex and age segregation of wintering willow ptarmigan in Labrador. Northeastern Naturalist 13: 113-118. Simon, N. P. P., and F. E. Schwab. 2005. Plant community structure following wildfire in the subarctic forests of Lab- rador. Northern Journal of Applied Forestry 22: 229-235. Newbury, T. L., and N. P. P. Simon. 2005. The effects of clearcutting on snowshoe hare (Lepus americanus) rela- tive abundance in central Labrador. Forest Ecology and Management 210: 131-142. Betts, M., N. P. P. Simon, and J. J. Nocera. 2005. Point count summary statistics differentially predict reproductive activ- ity in bird-habitat relationship studies. Journal of Ornithol- ogy 146: 151-159. Simon, N. P. P., and F. E. Schwab. 2005. The response of conifer and broad-leaved trees and shrubs to wildfire and clearcut logging in the boreal forests of central Labrador. Northern Journal of Applied Forestry 22: 35-41. Otto, R. D., N. P. P. Simon, S. Couturier, and I. Schmelzer. 2003. Evaluation of satellite collar sample size require- ments for mitigation of low-level military jet disturbance 2007 of the George River caribou herd. Rangifer special issue number 14; 297-302. Simon, N. P. P., A. W. Diamond, and F. E. Schwab. 2003. Do northern forest bird communities show more ecological plasticity than southern forest bird communities in eastern Canada? Ecoscience 10: 298-296. Simon, N. P. P., F. E. Schwab, and R. D. Otto. 2002. Song- bird abundance in clear-cut and burned stands: a compari- son of natural disturbance and forest management. Cana- dian Journal of Forest Research 32: 1343-1350. Simon, N. P. P., C. B. Stratton, G. J. Forbes, and F. E. Schwab. 2002. Similarity of small mammal abundance in post-fire and clearcut forests. Forest Ecology and Manage- ment 165: 163-172. Schwab, F. E., N. P. P. Simon, and C. G. Carroll. 2001. Breeding songbird abundance in the subarctic forests of western Labrador. Ecoscience 8: 1-7. Schwab, F. E., F. G. Pitoello, and N. P. P. Simon. 2001. Relative palatability of green manure crops and carrots to white-tailed deer. Wildlife Society Bulletin 29: 317-321. LeCoure, M. I., F. E. Schwab, N. P. P. Simon, and A. W. Diamond. 2000. Effects of post-fire salvage logging on breeding birds in western Labrador. Northeast Wildlife 55: 39-46. Simon, N. P. P., F. E. Schwab, and A. W. Diamond. 2000. Patterns of bird abundance in relation to logging in west- ern Labrador. Canadian Journal of Forest Research 30: 257-263. Simon, N. P. P., F. E. Schwab, M. I. LeCoure, F. R. Phillips, and P. G. Trimper. 1999. Effects of trapper access on mar- ten population in central Labrador. Northeast Wildlife 54: 73-76. Simon, N. P. P., F. E. Schwab, M. I. LeCoure, and F. R. Phillips. 1999. Fall and winter diet of Martens, Martes americana, in central Labrador related to small mammal densities. Canadian Field-Naturalist 113: 678-680. Simon, N. P. P., F. E., Schwab, E. M. Baggs, and G. I. McT. Cowan. 1998. Distribution of small mammals among suc- cessional and mature forest types in western Labrador. Canadian Field-Naturalist 112: 441-445. Peer Reviewed Manuscripts in Preparation: Newbury, T. L., N. P. P. Simon. Moose (Alces alces) browse use in central Labrador Submitted to Canadian Field-Nat- uralist. Simon, N. P. P., and A. W. Diamond. Songbird habitat quality across a timber productivity gradient within an unfragment- ed northern boreal forest: local and landscape effects. Sub- mitted to Landscape Ecology. Simon, N. P. P., and A. W. Diamond. Evaluation of environ- mental factors influencing vegetation structure in mature Picea mariana forests using constrained ordination and constrained classification. Submitted to Canadian Journal of Forest Research. Simon, N. P. P., F. E. Schwab, and J. K. Colbert. Vegetation response to summer and winter logging of Black Spruce Picea mariana forests in central Labrador. Submitted to Northern Journal of Applied Forestry. Theses: Simon, N. P. P. 1996. The effects of secondary succession on vegetation structure and its effects on small mammal bio- diversity in western Labrador. B.Sc. Honours thesis. Memorial University of Newfoundland, St. John’s, New- foundland and Labrador. CHUBBS and PHILLIPS: A TRIBUTE TO NEAL PHILIP PERRY SIMON, 1973-2006 YY Simon, N. P. P. 1998. Patterns of bird abundance in relation to logging in western Labrador. M.Sc. University of New Brunswick, Fredericton, New Brunswick. 83 pages Simon, N. P. P. 2006. The effects of site productivity and heterogeneity on bird habitat quality and species richness Ph.D. thesis. University of New Brunswick, Fredericton, New Brunswick. 184 pages. Reports: Schmelzer, L., J. Brazil, T. Chubbs, S. French, S. B. Hearn, R. Jeffery, L. LeDrew, H. Martin, A. McNeill, R. Otto, F. Phillips, G. Mitchell, G. Pittman, N. Simon, and G. Yetman. 2004. Recovery Strategy for three Woodland cari- bou herds (Rangifer tarandus caribou; Boreal population) in Labrador, Canada. Department of Environment and Con- servation, Government of Newfoundland and Labrador, Corner Brook. 51 pages. Simon, N. P. P. 2003. A summary of some effects of full tree logging — literature review. Internal report — Department of Forest Resources and Agrifoods, Wildlife Division. Simon, N. P. P., E. Baggs, and G. I. McT. Cowan. 1998. the effect of forest fire on small mammals in western Lab- rador. Internal report — Department of Forest Resources and Agrifoods, Wildlife Division. 23 pages. Simon, N. P. P., F. E. Schwab, and M. I. LeCoure. 1998. Labrador marten demographics as determined by winter diet and trapper access. Interim report submitted to the Newfoundland and Labrador Department of Forest Re- sources and Agrifoods, Wildlife Division. Simon, N. P. P., F. E. Schwab, and A. W. Diamond. 1998. Patterns of bird abundance in relation to logging in western Labrador. Interim report submitted to: Wildlife Habitat Canada, Newfoundland and Labrador Department of For- est Resources and Agrifoods, Wildlife Division. Simon, N. P. P., F. E. Schwab, and A. W. Diamond. 1997. The effects of logging on breeding birds in western Lab- rador. Interim report submitted to: Wildlife Habitat Cana- da, Newfoundland and Labrador Department of Natural Resources and Wildlife Division. Workshop and Conference Presentations: Simon, N. P. P. 2003. Evaluating the effects of landscape change on abundance, productivity, and survival of forest birds (presented with M. Betts). ACWERN, Wolfville, Nova Scotia. Simon, N. P. P. 2003. Natural disturbance and forest man- agement. Workshop on Natural Disturbance Management. Sustainable Forest Management Network, Corner Brook, Newfoundland and Labrador. Simon, N. P. P. 2002. The relationship between forest pro- ductivity and songbird habitat quality. ACWERN, Rocky Harbour, Newfoundland and Labrador. Simon, N. P. P. 2000. Songbird abundance and fecundity in relation to forest structure and productivity. ACWERN, St. Andrews, New Brunswick. Simon, N. P. P. 1998. Patterns of bird abundance in relation to logging in western Labrador/Bird plasticity in relation to latitude. Institute of Environmental Monitoring and Re- search Seminar Series — Happy Valley — Goose Bay, New- foundland and Labrador. Simon, N. P. P. 1997. The effects of logging on birds in west- erm Labrador. ASWFB/ACWERN conference — Alma. New Brunswick. Simon, N. P. P. 1996. The effects of logging on birds in westem Labrador. ACWERN, Bon Portage Island, Nova Scotia. Received 14 February 2007 Book Reviews ZOOLOGY Amphibians and Reptiles of British Columbia By Brent M. Matsuda, David M. Green, and Patrick T. Gre- gory. 2006. Handbook, Royal British Columbia Museum, Victoria, British Columbia. 266 pages. $17.33. The British Columbia Provincial Museum (now the Royal British Columbia Museum) was second only to the Royal Ontario Museum to issue guidebooks to their provincial amphibian and reptile faunas for the public (by E. B. S. Logier in 1937 and 1939 for Ontario, and Clifford Carl in 1943 and 1944 for British Columbia). The Carl guides (Handbooks 2 and 3) subsequently went through reprintings with some revisionary materi- al added in 1951 and 1968. In 1984 completely rewrit- ten guides were published on reptiles (Handbook 44 by Gregory) and amphibians (Handbook 45 by Green), both coauthored with R. Wayne Campbell. Now a new handbook (unnumbered) is a combined revised and partially rewritten text from the earlier Green and Gregory texts with updated distribution maps and the addition of a new coauthor. The previ- ous authors are long-established, still active research herpetologists with extensive publications, Green pri- marily on frogs (in the broad sense including, as well as typical “frogs”, toads, treefrogs, spadefoots, tailed frogs) and Gregory primarily on snakes but also includ- ing lizards, turtles and some amphibians. Dr. Gregory has spent his entire post-graduate career at the Univer- sity of Victoria; Dr. Green was born British Columbian but his post-graduate career has been at the Redpath Museum at McGill in Quebec. Both have extensive field experience in British Columbia. The new addition, Matsuda, is characterised in the tongue-in-cheek auth- ors’ profiles (page 258) as an “overachieving ecto- therm” (although this is surely not a unique charac- terization within this triumvirate) with a UBC M.Sc. who is now settled in British Columbia as an envi- ronmental consultant. This handbook, like its predecessors, follows a stan- dard guide layout. Preface, introduction (covering a wide range of aspects of amphibians and reptiles in general), checklist and conservation status, individual species accounts (for 39 species of which 4 are defi- nitely and 2 possibly introduced) and a further section which includes additional introduced species which apparently have not persisted or whose occurrence is only speculative, an appendix which gives contact in- formation for regional authorities, glossary of terms, bibliography (divided into additional reading and ref- erences), acknowledgements (about authors with illus- trator credits) and an index. The black-and-white text drawings are mostly repeats from the earlier guides except for newly included wall lizard and all of the amphibians. A few of the latter are superior to those of the earlier amphibian guide but many are not. The new salamander drawings are particularly disappoint- ing with the number of costal grooves shown rarely agreeing with the counts given in the text. However, all are sufficient for identification. New are 60 small colour photographs inserted between pages 186 and — 187. Additions since the last guide new to British Colum- | bia and Canada are the secretive salamander Plethodon _ idahoensis and the introduced European wall lizard Podacris muralis. Also new is the conclusion from molecular evidence by T. R. Jackson in 1998 that the Aneides salamander on Vancouver Island is an import with bark from California, A. vagrans, and not the geographically closer A. ferreus to which it had long been assigned. Other nomenclature updates from recent revisionary studies are Dicamptodon tenebrosus, Asca- phus montanus, Spea intermontanus, Actinemys mar- morata, Pseudacris maculata, Rana luteiventris, Pitu- ophis catenifer, and Crotalus oreganus. One accepted elsewhere that was not made is Pseudacris for Hyla regilla, and the more controversial generic changes Lithobates for Rana sylvatica and Rana pipiens (but not for R. aurora, R. pretiosa, and R. luteiventris) and Anaxyrus for Bufo boreas, even though one of the authors (David Green) is a coauthor on the paper detailing the latter changes (American Museum of Natural History Bulletin 297). Many incidental facts liven the text (such as the observation that frogs always close their eyes when) they leap (page 14)), that frogs can “hear” ground- borne vibrations by transmissions through their front! legs (pages 21-22) and that females of many turtles: species can store sperm from a single male for up to four years (page 32). The marine Cadborosaurus and the Okanagan Lakee “Ogopogo” are only mentioned under folklore (pagee 49) with the (arguable by some) statement “Sporadic¢ sightings of these beasts continue, but evidence of! their existence is lacking.” Few slips were missed in proofing such as the fail- ure to italicise Hyla versicolor and H. chrysoscelis on page 29, and the only British Columbia specimens ol the horned lizard Phrynosoma given as “two neal Osoyos in 1898” on page 182 but as “two records from Osoyos area in 1910” on the facing page 183 (the firs is correct). 100 2007 This guide brings British Columbia herpetology effectively into the 21" century as to be expected from the distinguished authors. It is a must for any west coast naturalist’s bookshelf and/or field jacket with an out- sized pocket. BooK REVIEWS 10] FRANCIS R. Cook Canadian Museum of Nature, Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada Blue Grouse: Their Biology and Natural History By Fred C. Zwickel, and James F. Bendell. 2004. National Research Council of Canada, Ottawa, Ontario. 284 pages. $69.95 Paper. This monograph is the culmination of two life- times devoted to studying Blue Grouse population biology and behaviour, both in the field and in aviaries, predominantly in coastal British Columbia. One of the advantages of a lengthy monograph is that the authors were able to synthesize much unpublished data and “gray” literature in addition to published references, making it the most comprehensive reference on this species group that is currently available. As such, it is not a book for the general reader of natural history (the colour and black-and-white photographs scattered throughout are quite small), but a scientific reference full of tables and graphs and lots of detail (there are 12 pages each of references and results of statistical tests). Unfortunately, only two years after Zwickel and Ben- dell’s monograph was published, the American Ornith- ologists’ Union (Banks et al. 2006) re-split Blue Grouse into two taxa, Dendragapus obscurus in the Pacific Coast Range and the Sierra Nevada, and D. fuliginosus in the Rocky Mountains. These two taxa had origi- nally been considered two species (Brooks 1929), but were conspecific for most of the 20" century. The re- split resulted from the recent DNA-based work of Barrowclough et al. (2004), who also found that the New Mexico populations of Dusky Grouse were per- plexingly different, but not enough to consider them separate species. Populations in the northern parts of the range (i.e., north of the Chilcotin in central B.C., through to Alaska) were not included in the Barrow- clough et al. study, but the separation into coastal and inland species is assumed to hold true. Unfortunately, the AOU split makes the monograph more difficult to use, but in their favour, Zwickel and Bendell use the scientific names, often with full trinomial reference Birds of Peru By Thomas S. Schulenberg, Douglas F. Stotz, Daniel F. Lane, John P. O'Neill, and Theodore A. Parker III. 2007 Prince- ton University Press. 656 pages. U.S. $49.50 Cloth. In 2001 I was delighted to find Birds of Peru, by J. Clements and N. Shany. It was a good modern field guide and proved its worth in Peru. However, there were a few times when I needed to borrow a more com- prehensive text to be certain of my identification. Now to the eight accepted races at the time of publication, throughout, as well as referring to “coastal” and “interior” subspecies analogous to the new split. I recommend that the serious reader really study the chapter on taxonomy and distribution with Barrow- clough’s paper (available on the internet) in hand. As a serious naturalist, | found the chapters on histor- ical review, physical environment, integument (plumage especially), behaviour, habitat use and movement, pop- ulation parameters, predators and disease the most interesting. The seven chapters on form and function — integument, morphology, reproduction, growth and development, food and nutrition, energetics and genet- ics — will be of interest mostly to serious students of Blue Grouse. Although Zwickel and Bendell refer to studies from other parts of Blue Grouse range, they admit that the strong focus on the coastal species (D. fuliginosus) is unfortunate because there are a number of character- istics, such as vocalizations, that are clearly different between the two species. The authors hint at a future publication that will explore the population ecology of Blue Grouse, and it is to be hoped that they are able to overcome this geographical disparity. Literature Cited Banks, R. C., C. Cicero, J. L. Dunn, A. W. Kratter, P. C. Ras- mussen, J. V. Remsen, J. D. Rising, and D. F. Stotz. 2006. Forty-seventh supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 123(3): 926-936. Barrowclough, G. F., J. G. Groth, L. A. Mertz, and R. J. Gutierrez. 2004. Phylogeographic structure, gene flow and species status in Blue Grouse (Dendragapus obscurus). Molecular Ecology 13: 1911-1922. Brooks, A. 1929. On Dendragapus obscurus obscurus. Auk 46: 111- 113. CyYNDI M. SMITH Box 5, Waterton Park, Alberta TOK 2MO Canada we have a new guide by Schulenberg et al. and the obvious question is whether it is an improvement. The first difference is that the new guide has range maps. This means I do not have to struggle with phras- es such as “on the east bank of the Rio Utacamba at the south end of Cordillera de Colon.” [Marvelous Spatule- tail] Now I can look at the map and have an immediate understanding. The range maps are conveniently set 102 next to the text in what is normally the margin, making great use of space. This alone is a great improvement. Peru has about 1800 species or one fifth of the world’s birds. Most sources cite this vague number. The few that are more precise give numbers that vary by up to 100 species. It is not surprising that a state of flux exists in such a vast and complex area. This book covers 1792 species which is probably as good as you do under such fluctuating circumstances. Try- ing to reconcile the list of species covered by both books or given in other sources is very difficult. It is a bit like trying to compile a list of “honest” politi- cians. The new book drops some questionable records like Plumbeous Ibis [one old, suspect, badly labeled specimen] but adds Bogota Rail [based on a current valid sighting.] Some birds are treated by Shulenburg as sub-species; Andean Ibis is listed as a sub-species of Black-faced Ibis whereas other sources give it full species status. There are complications due to name changes. For example, there are four toucans listed in both books. However, the names do not correlate well. This includes the scientific binomials, normally a very stable source of information. The old book has Toco Toucan Ramphastos toco, Black-mandibled Toucan Ramphastos ambigus Yellow-ridged Toucan Ramphas- tos culminates, and Cuvier’s Toucan Ramphastos cu- vieri. The new book has Toco Toucan Ramphastos toco, Black-mandibled Toucan Ramphastos ambigus plus White-throated Toucan Ramphastos tucanus, and Channel-billed Toucan Ramphastos vitellinus. The last two are new names for Cuvier’s and Yellow-ridged Toucans. [Speciation in the whole toucan complex is confused.] I also noted that Andean Flamingo and James’s [Puna] Flamingo have been updated to Phoeni- coparrus andinus and Phoenicoparrus jamesi, respec- tively, but the Guanay and Red-legged Cormorants The Freshwater Fishes of British Columbia By J. D. McPhail, illustrated by D. L. McPhail, foreword by Joseph S. Nelson. 2007. University of Alberta Press, Edmonton. lxxiv + 620 pages. $90.00. This book is the latest treatment covering the fresh- water fishes of British Columbia. Earlier books ap- peared in 1948 (132 pages) and 1967 (192 pages). They were published by the British Columbia Provin- cial Museum (now Royal British Columbia Museum) while the latest book is published, strangely, in Alberta. The present volume is 694 pages long with a larger format and is indicative of both the growth in knowl- edge and the need for a comprehensive work on this provincial fauna. The book is composed of introductory sections coy- ering purpose, use of keys, names of fishes, layout of the book, origins of the fauna, present distributions, and conservation. The sections on glacial events and THE CANADIAN FIELD-NATURALIST Vol. 121 have not been changed to Leucocarbo bougainvillii and Leucocarbo gaimardi. So those with a penchant for lists may need to do some research. That being said, this book is more than adequate as a field guide for the resident and visiting birder alike. The color plates are opposite the descriptions and distribution maps. There are 13 illustrators, so there is some difference in style. The most disparate are Lau- rence McQueen and John O’ Neill. O’ Neill’s work has the precise, almost photographic rendering we see in most modern field guides. McQueen’s paintings are more flowing and look like paintings. My junior school art teacher would have said they were more “painterly.” Daniel Lane’s work is somewhat in between these two styles. Which style you like is a matter of taste. I have looked at many of the illustrations for accuracy and can find no problems. In fact, I particularly like the tou- canets by O’ Neill and the jaegers by McQueen. There are 4000 color illustrations at about six to a plate, mean- ing the illustrations are quite large [again the publish- er has used the margins]. In addition to the species accounts there are well- written sections on Habitats, Molts-and-Plumages, and Conservation. In particular the terminology used for habitat in the species accounts is clearly documented. The text in the species accounts covers the key iden- tification features, status, distribution, and song. This book will please birders and biologists living in or visiting Peru. It is also useful beyond the borders of Peru into adjacent areas of South America. It is a lit- tle on the heavy side for dragging through a hot jungle, but what else can you expect with such an enormous bird list. Roy JOHN 2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada current distributions are a very useful synthesis as is the discussion on what a species is and why the vari- ous forms of sticklebacks and others have not been named taxonomically. The bulk of the text is the descriptions of species. Family accounts give general information on the in- cluded species and the identification keys are found there. The species accounts are preceded by a pictori- al key to families that works for the more distinctive body shapes but would have benefited from some anno- tations for those fish that are more similar in shape to enable the naive reader to correctly identify the family. It is always a debate as to whether keys should be with the family or should form a separate section. The for- mer works when reading through the book but the latter is much easier for the actual identification process in a laboratory or the field. The keys are well illustrated al- 2007 though both illustrations and text are too small. Partic- ularly useful are keys to fry, parr, and adult salmons and trouts as these life history stages differ in appearance. Each species account has a scientific name (but no date with the author), English common name, a black- and-white illustration, sections on distinguishing fea- tures, taxonomy, sexual dimorphism, hybridization, dis- tribution with a spot map, life history, reproduction, age, growth and maturity, food habits, habitat and con- servation comments. A total of 81 species are covered, including 10 exot- ics. The main families of the 17 known from British Columbia are salmons, trouts and whitefishes with 23 species, minnows (20), and sculpins (8). Minor er- rors are inevitable in any work; e.g., Lindsey is mis- spelled in the dedication, Qadri as Quadri in the Ref- erences, but some can be confusing; e.g., on page 498 Pomoxis nigromaculatus is said to have 5 anal spines in the sunfish identification key but most fish have 6- 7 spines; the stickleback key separates two species by non-overlapping dorsal fin spine counts (which do, however, overlap occasionally) where in one species (Pungitius) the spines are staggered left to right, an easily seen and unique character (mentioned in Dis- tinguishing Characters). The book ends with an extensive Bibliography, an Appendix I (a checklist of the fishes, with no other appendices), a Glossary and an index to scientific and common names. The Glossary explains a variety of terms, even that the plural of dentary is dentaries which is fairly standard English, but gives axillae without explaining it is the plural of axilla (a knowledge of Latin is ever more dead). One obscure word is snye, defined as a dead-end side channel. This is listed in dictionaries as an Ontario word and can also mean a channel joining two rivers. Cost is increasingly a factor in production of books and this may explain the absence of any colour plates. Owls of the World By J. Duncan. 2003. Key Porter Books, Toronto, Ontario. 319 pages. Hardcover, $60. This beautiful book actually fulfills the two roles that many natural history books aspire to but usually fall short of ... being both a detailed scientific reference and easy to read. The first two-thirds of the book cover the biology and life history of owls, stories by people who have uncovered these amazing details and the methods they used, the substantial role that owls have in mythology around the world, and threats to their sur- vival (but also the passion and hope that people hold out for owls). The U.S. edition, published by Firefly Books, carries the apt subtitle “their lives, behaviour and survival”. The last third of the book is a detailed look at 205 species of owls: physical description, habitat, natural history, general distribution and their global conserva- Book REVIEWS 103 In my mind this argues for including a CD/DVD with the book to carry photographs of habitats and coloured depictions of fishes. The text can also be put on the disk, making the book searchable. I still prefer a book in the hand to a computer screen but suspect this does not apply to a younger generation. Even the CD in a pock- et at the back of the book is a transitional stage and “books” should now appear on the internet, thereby saving trees, allowing a plethora of colour plates and other illustrations, enabling the author to receive and incorporate feedback and new data, and facilitating correction of errors of fact and of omission. A “book” on a website can be built up incrementally although some will inevitably fail to reach a completed form. Each stage allows for criticism and correction, from the initial checklist of species, the identification keys, distribution maps (allowing zoom-in for finer details than part of a book page allows), and finally text descriptions. For those worried about vanishing websites, EM pulses and cyber trashing, a printout of say 20 copies of the digital “book” can be deposited in various libraries. This trend can be seen in some scientific works such as the “Annotated Checklists of Fishes” produced by the California Academy of Sciences which are on-line at that institution but have copies deposited in a minimum of six natural history institutions world- wide, admittedly as CDs. This is the definitive book on this fish fauna and an essential tome for any student of the aquatic environ- ment in British Columbia. Future versions will require multiple authors to match and improve on the experi- ence and knowledge exhibited here by Don McPhail. BRIAN W. CoAD Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada tion status. Even a quick look highlights how difficult it is to study some of these species, as many do not have photographs. Duncan brings not only his own wealth of knowl- edge and field experience (particularly with Great Gray Owls) to this task, but also those of a “whoo’s whoo” in the world of owls, such as Robert W. Nero, Irina Menyushina, C. Stuart Houston, and Aki Higuchi. Duncan’s use of anecdotes helps to de-mystify sci- ence and raises the readability level, such as when he relates being dared to taste a freshly regurgitated owl pellet in front of a crowd. He said it tasted “sweet,” but his incredulous darer (R. W. Nero) bit into it and quickly spat it out, declaring that it tasted “bitter.” They were both right ... the pellet is coated with a pH-neutral film of mucous that protects the owl’s throat and mouth from the low-pH acid-soaked mass of fur and bones, which Nero had bitten into! 104 As with many species, owl populations are often in trouble around the world for the usual reasons: habitat destruction, direct (vehicles, shooting, trapping) and indirect (hitting barbed wire fences) mortality, and in- vasive alien species such as West Nile virus. But Dun- can argues that an awareness and understanding of how people have perceived owls in the past may help support efforts to conserve them today. The one small section of the book that many people will struggle to read describes the DNA sequencing and classification of owls. It would have been more understandable if Duncan had linked this highly tech- nical description, especially the Maximum Likelihood Tree showing ow] relationships, to some of the species accounts, where it is obvious from a glance at range maps for many of the Scops Owls in the Malaysian THE CANADIAN FIELD-NATURALIST Vols 121 archipelago that these species probably had a com- mon ancestor, and allopatric speciation occurred due to island isolation. The photos in this book are luscious, with rich colour. Today, I randomly opened it and there was a full-page picture of Verraux’s Eagle Owl — the best photo that I’ve seen of this owl that adequately shows its pinkish- purple eyelids. I recall driving along a sandy track in Chobe National Park in Botswana when one of my companions excitedly sputtered “purple eyelids, pur- ple eyelids,” as she pointed at the bird that had caught our attention in the guidebook! Excuse me while I read up on the Tucuman Pygmy Owl before heading off to Argentina .... Cynpbi M. SMITH Box 5, Waterton Park, Alberta TOK 2MO Canada Ecology, Conservation, and Status of Reptiles in Canada Edited by C. N. L. Seburn, and C. A. Bishop. 2007. Herpeto- logical Conservation volume 2. Society for the Study of Am- phibians and Reptiles, Salt Lake City, USA. x + 246 pages. illus., hardbound. Available from the Publications secretary of SSAR, ssar@herplit.com/U.S. $40 plus shipping. This book is an edited compilation of peer-reviewed contributions on several aspects of conservation and more specifically, on the ecology of the reptiles that occur in Canada. It contains chapters by many (figu- rative) heavyweights of Canadian herpetology, who bring combined centuries of experience to the proj- ect. The book is a very good application of accumu- lated biological knowledge to the question of conser- vation. It is also a good example of how a professional community can collaborate in a common cause. It begins with a short introduction, then a histori- cal review of work done on reptiles in Canada. Next are four chapters summarizing the life history of lizards, snakes, freshwater turtles, and marine turtles. Contributions on the applications of genetics to con- servation, health and disease in reptile populations, and traffic mortality follow. The future of Canadian reptiles and a strategy for conservation are discussed. The book ends with an illustrated checklist of species, with the official conservation status of each. There is also a comprehensive summary of literature, both his- toric and recent, with few omissions. The book succeeds in its aim of providing a sum- mary of the biology and conservation status of Cana- dian reptiles. The chapters are of a consistently high calibre — a testimony to the knowledge of the authors and the thoroughness of the editors. The book is use- ful on more than one level — as a summary of reptile ecology in Canada and as a conservation guide; the extensive literature is also a valuable resource. Several themes are common to the contributions. Reptiles living in Canada face two problems — the cli- mate and humans. Because climate affects life history, reptiles are concentrated in southern Canada, where it’s warmer. Unfortunately for the reptiles, humans are concentrated in the same areas. Human activities that result in habitat modification are usually in direct conflict with the best interests of reptiles. The book makes the point that although the Cana- dian climate may not be ideal for reptiles, they have adapted to life here. A reader soon realizes that many, if not all, declines in reptile numbers are due to human action. For example, road mortality is a common theme in several chapters. Human alterations to the landscape can have direct effects such as habitat destruction, and also secondary effects such as increases in reptile pred- ators like cats and raccoons. Several authors make the point that declines in reptile, especially turtle, popula- tions can occur slowly, drawing little public attention. Although I was familiar with much of the informa- tion presented, there were some surprises, one being the high number of marine turtles off the Atlantic and Pacific coasts. This is but one example of how much is still not known about our reptile fauna. Quibbles are few and minor. Colour variations and aberrant scutellation in turtles should not be included among diseases and parasites; the authors themselves admit that these “..are not detrimental...” (page 149). What is the prognosis for reptile conservation? Al- though knowledge of the need for conservation is high among the public, and although the number of conser- vation programs is high, the future still looks bleak, because of the juxtaposition of human and reptile pop- ulations. The point is made repeatedly that while atlases and monitoring programs are good, they are ultimate- ly ineffective without concrete action; “...ecological knowledge alone will not save species when the real problems facing them are social and political.” (page 47). Ross D. MACCULLOCH Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6 Canada; e- mail: rossm@rom.on.ca 2007 BOOK REVIEWS 105 Rodent Societies: An Ecological & Evolutionary Perspective Edited by J. O. Wolff and P. W. Sherman. 2007. The University of Chicago Press, Chicago, 610 pages. U.S. $49. Paper, $125 Cloth. This opus on rodent behaviour brings together 61 re- searchers in a single edited volume. There are 42 chap- ters grouped into various topics including systematics, sexual behaviour, life histories, behavioural develop- ment, social behaviour, antipredator behaviour, com- parative socioecology, conservation and disease. The main objectives are to present the latest research on the behaviour of rodents and to place it in the context of ecology and evolution. The resultant compilation of papers is firmly based on comparative biology, which goes beyond merely describing phenomena and ex- plains it in terms of the larger ecosystem and within a phylogenetic framework. There is an emphasis on hypothesis testing and explicit experimentation that results in robust and scientifically-sound studies. This approach fostered by the editors gives more than just summaries of particular topics but highlights the on- going nature of our understanding of behaviour in not only rodents but also as related to other organisms. At over 2200 species, Rodentia is the most speciose order of mammals and accounts for approximately 40% of the diversity. In terms of breadth of coverage, almost half of the chapters summarize behavioural data across all major lineages of rodents or within a higher-level classification. In addition, many different groups or spe- cies of rodents are directly studied as case examples in this compendium, with taxa examined ranging from squirrels to beavers to rats to capybara. After an introductory chapter on rodents as model systems, the evolutionary background is set with a paper discussing phylogenetics and biogeography. This is followed by several chapters on sexual behaviour dealing with mating and reproductive strategies. Life histories of rodents are covered by topics on dimor- phism, sex ratios, stress, dispersal and philopatry, gene dynamics, and self-regulation. There is discussion of behavioural development such as neural regulation, ontogeny, learning, and kin recognition. Social behay- iour is examined with papers on parental care, ecology of sociality, scent marking, non-parental infanticide, monogamy, and pacifism. There is a small section with three chapters on antipredator behaviour including Seashells: Jewels from the Ocean By Budd Titlow. 2007. Voyageur Press, 729 Prospect Avenue, P.O. Box 1, Osceola, Wisconsin 54020. 112 pages. US $20. Cloth. Sea shells have fascinated people since the dawn of time. Intrinsically beautiful, they have inspired artists and architects, been used to fashion tools, and have even served as money. They are also the prize of col- lectors the world over. All of these themes feature in alarm communication. Comparative socioecology has several papers on social organization and structure of a number of different species or higher-level groups. The book ends with a section on conservation and disease followed by a chapter summarizing conclusions and future directions on the research of rodent behaviour. One weakness in an otherwise comprehensive anthol- ogy was the lack of molecular study on the genetic basis for much of the evolution of behaviour. In particular, DNA sequence data nowadays seems to be churning out by the genome. Was this an editorial oversight, or is there just nobody looking into this field of research for rodents? If so, this highlights an area that needs attention, especially since there are several candidate model-system species in rodents to choose from. Like- wise, I found it odd that only four of the chapters have figures of phylogenies, considering that an evolutionary perspective is supposed to be one of the major underly- ing themes of the volume. Perhaps this is an indication that there is still an obvious gap that requires bridg- ing between micro- and macroevolution. Another observation is that over 80% of the authors are based at institutions in the Americas. Has this biased the presentation of the state of knowledge of rodent behaviour? A more subjective criticism is the com- bined literature cited section for the whole book. For edited volumes, I prefer references listed separately at the end of each chapter, especially when topics are as varied as this monograph. However, there are some advantages such as the elimination of repeated cita- tions for general publications on rodents and a single compiled source useful for searching purposes. None- theless, these minor detractions by no means diminish from the scientific merits of the book. This book is definitely aimed at the specialist re- searching the behaviour of rodents, but will also be of interest to biologists working on other organisms and studying similar social systems. At over 600 pages, the price is reasonable for purchase by a university library, or would be a valuable addition to any ecolo- gist’s bookshelf because it is packed with the latest information on ethology. BURTON K. LIM Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6 Canada Budd Titlow’s handsome book Seashells: Jewels from the Ocean. This relatively slender volume, copiously illustrated with photographs by the author or from stock footage, provides a good introduction to the world of mollusks and the shells which adorn so many of them. Succes- sive chapters deal with the physiology of mollusks, their classification, their habitats, their cultural and econom- 106 ic importance to humans, and the threats that mollusks face. The latter include marine and land-based sources of pollution and over-exploitation of commercially valuable species. There is also a primer on shell collecting, and also short notes on good collecting sites in North America. At its heart, this is a book aimed at shell collectors, be they casual beachcombers or fanatical conchologists. The former will learn much, the latter will no doubt admire the stunning photographs. There is a distinct American bias, both in the selection of photographs and in the shell collecting sites profiled; indeed at times the book reads as an ode to Sanibel Island, Florida, the Mecca of American shell collectors. The book oscillates between straightforward expla- nations in layman’s terms of the science underpinning malacology, and a rather folksy, and at times highly personal, appreciation of the cultural aspects of sea shells and shelling. While occasionally rather affected, on the whole it is highly readable. Many of the pho- tographs selected sacrifice ecological verisimilitude for art; for example, there are several very beautiful assemblages that would be highly improbable in nature. And, belaying the book’s American bias, it would be a fortunate day indeed where one found a Spider Conch (from the tropical Indo-Pacific) on a North American beach. For a field naturalist, the book is faintly disap- pointing in that it tends not to identify in specific terms THE CANADIAN FIELD-NATURALIST Vol. 121 most of the shells portrayed. Shelling, unlike birding, is One pastime where getting a grip on the Latin ter- minology and the taxonomy is essential; for this the reader would be well advised to turn to any one of the several excellent works cited in the short but useful reference section. By the same token, a few sketches illustrating such things as mollusk morphology would have been helpful. There are a few errors in the text; for example the swans that winter at Chesapeake Bay are Tundra Swan, not Trumpeter Swan, but these errors do not detract from the overall value of the book. One somewhat dis- appointing aspect of the book, and one in counterpoint to the otherwise conservationist approach, is a section dealing with the collection of live specimens. While pale in comparison to other threats, collectors can pose a considerable threat to vulnerable species and it would have been preferable if the author had concentrated the reader’s attention on the collection of non-living material. That said, the author does emphasize the need to respect local regulations. In conclusion, this book is not a scientific reference; however, it does provide a useful primer for anyone keen to explore the world of shelling, and a handsome addition to any collection of “coffee table” books. MARK GAWN Mission of Canada, 5 Avenue de |’ Ariana, Geneva 1202 Switzerland Turtles: An Extraordinary Natural History 245 Million Years in the Making By Carl J. Franklin. 2007. Voyageur Press, 729 Prospect Avenue, PO Box 1, Osceola, Wisconsin 54020. 160 pages. U.S. $35. Turtles provides a brief overview of the turtles of the world along with abundant colour photographs of many of the species. The book is divided into two parts. Part 1, the life and lifestyle of turtles, consists of three chapters. The first chapter covers the taxonomy and internal biology of turtles in roughly 10 pages. Chap- ter two, the ecology of turtles, covers topics such as thermoregulation, feeding, reproduction, and preda- tors. The third chapter, covering the origin and fossil history of turtles, is only 3 pages long. Part 2, the diver- sity of modern chelonians, consists of two chapters. One chapter covers the families of Pleurodira, or side- necked turtles, and the other discusses the families of Cryptodira, or hidden-necked turtles. For each family account there is a map illustrating the global distribu- tion of the family, then text discussing shared traits among members of the family and then information on many of the species within the family. Roughly two-thirds of the books is allocated to Part 2, the family accounts. I found this an unsatisfactory balance. Part 1, which should provide a coherent pic- ture of the various different turtle life history strategies, is just too short and superficial. Likewise, the family accounts remain superficial because they do not pro- vide systematic species accounts. The information sel- ected for individual species should highlight the unique qualities of that species, but all too often the species profiles rely on just some basic numbers (size, number of eggs laid), distribution (which countries it occurs in) and some general habitat info. The text also has a number of glaring mistakes. In a text box on Linnaean taxonomy of turtles, the author states that turtles belong to the class Anapsida (page 14). In Linnaean terms, Anapsida is the subclass, while Reptilia (or Chelonia, for splitters) would be the class. The author claims that both McCord’s Box Tur- tle (Cuora mccordi) and Zhou’s Box Turtle C. zhoui) are believed to be extinct (pages 39-40), when really he should state they are likely extinct in the wild, as both species still occur in captivity. The author makes use of the most recent phylogenetic work on the genus Clemmys, by transfering the Wood Turtle (Glyptemys insculpta) to its new genus, but still leaves the Bog Tur- tle (G. muhlenbergii) in the genus Clemmys. Other careless errors include stating that there is more than one species within the genus Actinemys (page 98), and claiming that there are four species of Painted Turtle (Chrysemys picta; page 101), when really there are four subspecies. The Indian Flapshell Turtle (Lissemys 2007 punctata) receives two different and somewhat con- tradictory species accounts, one brief account under its correct subfamily (Cyclanorbinae; page 134), and a longer account under an incorrect subfamily (Tri- onychinae; page 139). Despite these errors, there are some good things about this book. Part | does pro- vide a very brief and readable introduction to the world Whales and Seals Biology and Ecology By Pierre-Henry Fontaine. 2007. Schiffer Publishing, 4880 Lower Valley Road, Atglen, Pennsylvania 19310 USA. 448 pages. U.S. $35. Paper. During my career as a marine mammal research scientist I was reluctantly dragged in on several occa- sions to necropsy and dispose of large dead mammal carcasses found along the beaches of Canada’s east or west coast. Almost always these mammoth moun- tains of mammalian flesh were in an advanced stage of putrefaction. Upon opening the body cavity one would confront what has been termed a “gaseous pud- ding” and it was challenging to properly identify even the most conspicuous organs. It is with this in mind that I say that I am impressed with the work in hand, which has yielded for the author a large amount of useful, detailed information, gathered with fortitude and persistence, from a source that many would have been wont to avoid. The illustrations, particularly the excellent photo- graphs of the organs and musculature, will serve as a valuable resource for field biologists learning their way around marine mammal carcasses. The author’s expert- ise in marine mammals comes primarily from his exten- sive experience in performing necropsies of beach cast cetaceans and to a lesser extent pinnipeds. This book uses this as the point of departure relating the specific adaptations, the senses, nutrition, and locomotion, to the anatomical structures which he has studied and illustrated during his years of studies. The remainder of the book attempts to give a gener- al account of whales and seals, particularly from the North Atlantic, where the author has worked. The chap- ter on strandings is devoted to the question of why live cetaceans beach themselves. As the author points out, almost every imaginable reason has been offered from geomagnetic anomalies to suicide. The chapter leaves one hanging as do most articles on this subject. One very important recent work on this subject is the mono- graph Marine Mammals Ashore (J. R. Geraci and V. Loundsbury. 1993. Texas A&M Sea Grant Publica- tion). This not mentioned by the author and would be a valuable source of additional information. The chapter on mounting marine mammal skele- tons will be found especially useful by museums and institutions wishing to do such displays. Little detailed BoOoK REVIEWS 107 of turtles. The photographs are generally quite good. Ultimately, however, this book could have been great- ly improved by a thorough review from an expert on turtles. DAVID SEBURN Seburn Ecological Services, 2710 Clarenda Street, Ottawa, Ontario K2B 785 Canada information exists on this subject and the author has learned valuable lessons in avoiding errors in skeletal reconstruction from his numerous and careful field dissections. The remaining chapters, while not based on the au- thor’s direct experience, are well researched and illus- trated. The long association of people with whales and seals is presented in an interesting manner. Some spe- cific examples of marine mammal hunting in the St. Lawrence are little known. The technique on Walrus hunting in the Magdalene Islands, which led to the early extinction of the southern-most Atlantic Walrus stock, is one such example. The chapter dealing with the complicated subject of marine mammal paleontology gives a good intro- duction to a discipline which is both difficult to assimi- late and where the information is not well organized in any single comprehensive source. This will serve as a good point of departure for students who want to learn more about the origins of marine mammals. The last chapters, the fact sheets on Cetaceans and Pinnipeds, might be considered useful additions to this general work on marine mammals only because they include the species found in the North Atlantic, which is where most of the author’s work has been done. One wonders why the author has included the Northern Elephant Seal (only found in the Pacific) and why the Pacific Walrus (Odobenus divergens) is mentioned and shown in one of the illustrations. The Arctic Ringed Seal, which is found as far south as the Labrador coast (Lake Melville) is omitted. A number of up-to-date field guides exist, which give a comprehensive account of the seals and whales in the North Atlantic and would serve their readers more effectively. I recommend this book to the non-specialist who has an interest in whales in particular. It will also be a useful source of information for field biologists and naturalists who might deal with beach cast remains of marine mammals. It succeeds in what the author set out to accomplish by sharing his considerable know]- edge of a subject, in which he has thoroughly immersed himself. THOMAS G. SMITH EMC Eco Marine Corp, 5694 Camp Comfort Road, Beaulac- Garthby, Quebec GOY 1B0 Canada 108 ENVIRONMENT China Ecosystems Sun Honglie, Editor-in-chief: 2005. The Science Press of China. 1823 pages. 350 Yuan RMB Cloth. The publication of such a magnum opus can be re- garded as a landmark achievement accomplished by this group of Chinese ecologists. The book summed up the main results of the long-term research on the ecosystem structure and function at more than 80 eco- logical research stations, extensively distributed over China, of the Chinese Academy of Science. The re- search comprehensively covered the various fields such as ecosystem classification, ecosystem structure and dynamics, ecosystem energy flow, water and nutrient cycling in ecosystems, ecosystem management and sustainable use and so on. China is a vast territory, richly endowed by nature and abounding in diverse types of ecosystems. Tradi- tional Chinese integrative philosophy, an abundance of resources and the fast increasing economic power confer on China satisfactory conditions and capacity for ecosystem research. However, in China, the initiation of ecosystem research was rather late compared with many other countries. At the beginning stage in the 1950s, ecosystem research in China was very rare and scattered. This was nearly stopped during the ten-year period of the Cultural Revolution. China began to par- ticipate in the large international ecosystem research plans, such as IBP [International Biological Program], MAB [Man and Biosphere Program], IGBP [Interna- tional Geosphere-Biosphere Program], GTOS [Glob- al Terrestrial Observation System], EHA [Ecosystem Health Assessment], PAGE [Pilot Analysis of Global Ecosystem] [Pesticide Assessment Guidelines?] and MA [Millennium Ecosystem Assessment] from the late 1970s. Driven by these international programs, Chinese ecosystem research developed quickly, espe- cially in recent years. Long-term observational studies play an irreplace- able role in understanding and assessing the ecosys- tem structure, function and their health status. In this course, the ecological research stations made great contributions. The Chinese ecological stations, which are distributed throughout the typical ecological zones of China, have accumulated long-term continuous and dynamic data, not only on the changes of the same ecosystem types under different driving forces, and on the integrated features of different types of ecosystems at the regional scale, but also on the long-term ecologi- cal experiments and management practices conducted in these stations. Most of Chinese ecosystems were generally healthy in the past; however, in recent years, at least some of them have undergone serious degradation, and some are even close to irreversible damage. Not long ago, China Environmental Protection Administration an- THE CANADIAN FIELD-NATURALIST Vol. 121 nounced the first Report on the Ecological Protection in China, which admits that the trend of environmental deterioration in China has not been effectively curbed yet, and more than 60% of lands are becoming envi- ronmentally fragile. Obviously, to achieve sustainable development, China must continually improve its dete- riorating environment, maintain and continuously up- grade the capacity of its ecosystem service, which needs a sound understanding of the effective management of various types of ecosystems in China. The results of the long-term research on the Chinese ecosystems were achieved with a background of large temporal and spatial environmental changes. There- fore, the data accumulated by the Chinese ecological research stations (data used in this book), include the information not only on the progressive but also de- grading succession of Chinese ecosystems, and the summary of some restoration practices. These results are particularly unique and valuable, and could be used as references for some other regions and countries. The objective of the Millennium Ecosystem Assess- ment of the UN, which involves a large number of ecologists from different regions and countries, is to systematically and comprehensively reveal the trends of changes of various ecosystems, and accordingly pro- pose effective countermeasures for the future. China ecosystems is certainly helpful for such an assessment and its publication is timely. The book is composed of eight parts. The first part is General review, including Chapter one: the history, evolution and perspective of the ecosystem research; Chapter two: the natural conditions for the formation of the Chinese ecosystems; Chapter three: the evolu- tion of the Chinese ecosystems; Chapter four: the im- pacts of human activity on the ecosystems; Chapter five: the principles and the systems of the Chinese eco- system classification. The second part is Forest ecosystems, including Chapter one: introduction; Chapter two: cold-temper- ate zone northern conifer forest ecosystem; Chapter three: middle-temperate zone conifer-broad-leaved for- est ecosystem; Chapter four: warm-temperate zone deciduous forest ecosystem; Chapter five: subtropical zone evergreen and deciduous broad-leaved mixed for- est ecosystem; Chapter six: subtropical zone evergreen broad-leaved mixed forest ecosystem; Chapter seven: subtropical zone western mountainous region ever- green hard-and-broad-leaved forest ecosystem; Chap- ter eight: subtropical zone alpine conifer forest and mountainous region conifer-broad-leaved mixed for- est ecosystem; Chapter nine: tropical zone seasonal rain forest ecosystem; Chapter ten: tropical zone rain forest ecosystem; Chapter eleven: sustainable devel- opment of China forest ecosystems. 2007 The third part is Grassland ecosystems, including Chapter one: the formation and classification of the grassland ecosystems; Chapter two; temperate zone grassland ecosystems; Chapter three: high and cold region grassland ecosystems; Chapter four: warm- temperate zone grassland ecosystems; Chapter five: tropical zone grassland ecosystems. The fourth part is China’s water ecosystems, in- cluding Chapter one: introduction; Chapter two: fresh water ecosystems; Chapter three: lake ecosystems; Chapter four: the succession and primary productivity of the lake ecosystems; Chapter five: ocean ecosystems. The fifth part is Farmland ecosystems, including Chapter one: introduction; Chapter two: northeastern Song-nen Plain farmland ecosystems; Chapter three: lower field of Liao River Plain farmland ecosystems; Chapter four: north China Taihang Piedmont Plain farmland ecosystems; Chapter five: Huang-huai Plain farmland ecosystems; Chapter six: Tai Lake Plain farm- land ecosystems; Chapter seven: South China hilly region farmland ecosystems; Chapter eight: Dongting Lake Plain farmland ecosystems; Chapter nine: Central Sichuan hilly region farmland ecosystems; Chapter ten: Loess Plateau farmland ecosystems; Chapter eleven: oasis farmland ecosystems in desert regions. The sixth part is nutrient cycling in terrestrial eco- systems, including Chapter one: a review on the re- search progress of biogeochemical cycling; Chapter two: cycling of carbon and main nutrient elements in farmland ecosystems; Chapter three: cycling of main New Zealand — A Natural History By Tui De Roy and Mark Jones. 2006. Firefly Books Ltd., Richmond Hill, Ontario. 160 pages. $40. Hardcover. The authors, experienced professional photogra- phers Tui De Roy and Mark Jones, moved to New Zealand about 16 years prior to the publication of the book, and, during this time, they explored many of New Zealand’s distinct and often rugged and remote natural regions. The result of these explorations and hard work is this beautifully illustrated coffee table book containing more than 450 magnificent photo- graphs which offer a wonderful introduction to many of the unique species and habitats of New Zealand. In keeping with one of the main areas of expertise of the authors — nature photography, the book is largely made up of beautiful pictures. Early in the book, there is a helpful map, which includes many of the locations and national parks referred to later on. The book also contains a reasonably detailed and useful index. As a biologist, I appreciated the inclusion — in an appendix just before the index — of a list of scien- tific and common names for many of the species named in the previous pages. The text sections are fairly brief overall. I counted 39 pages covered mainly by text (not including the acknowledgments and index), out of the total 160 pages Book REVIEWS 109 nutrient elements in forest ecosystem; Chapter four: cycling of main nutrient elements in grassland ecosys- tems. The seventh part is water cycling in terrestrial eco- systems, including Chapter one: introduction; Chapter two: water cycling in farmland ecosystems; Chapter three: water cycling in forest ecosystems; Chapter four: water cycling in grassland ecosystems. The eighth part is the theories and methods for the ecosystem network research, including Chapter one: the purposes and tasks for the ecosystem network re- search; Chapter two: the establishment and develop- ment of the ecosystem network; Chapter three: the methods for the ecosystem network research; Chapter four: the scales for the ecosystem network research. The book was well written with few errors. Abun- dant illustrations are helpful for readers to easily under- stand the research results. The book is suitable for the professionals who engage in ecology or environmental science, or other persons who are interested in these fields. Li Dezut', Qin AIL’, AND QIN HonG? 'Lab of Urbanization and Ecological Restoration of Shang- hai; National Field Observation and Research Station in Tiantong Forest Ecosystem of Zhejiang; Department of Environmental Science, East China Normal! University, 3663, Zhongshan Rd (N). Shanghai, China 200062) *Jilin Forestry Staff School, Jilin, China Zhejiang University, Hangzhou, China of the book. Thus, if one is planning a visit to New Zealand, reading the book prior to the departure may be easier than putting such a weighty, hardcover tome in the luggage. Although brief, the text is not without merit. The authors have done their best to include information on a wide variety of animals and plants, as well as a few details about New Zealand’s geological past and present. There are some interesting, although usually very short, stories about several species, including fist- sized giant carnivorous land snails — some of them endemic (restricted) to a single hill — that hunt large earthworms at night (some of the local earthworms can reach | meter in length) or the inquisitive kea, the very active mountain parrots of New Zealand, which are apparently capable of sometimes killing sheep and dismantling cars. There is a strong conservation theme throughout the book, and the main purpose of the text is clearly to highlight the plight of the many endan- gered species unique to New Zealand, and to discuss some of the conservation programs undertaken to save these rare species. This is a worthy goal, of course, and the authors’ commitment to conservation is evident throughout. However, as a scientist, I would have liked to see a more balanced and in-depth discussion of certain envi- 110 ronmental and conservation issues. The authors seem to take an uncritical approach towards some controver- sial control programs undertaken by the local Depart- ment of Conservation against species introduced to New Zealand by people, over the centuries. It is some- what unsettling, for example, to read about the “Her- culean aerial poisoning operation” — “the largest rodent control program ever undertaken anywhere in the world”, on the same page where the authors refer to the “humane integrity” associated with the conserva- tion initiatives such massive poisoning operations are a part of. It should perhaps have been mentioned, for example, that such massive poisoning operations can occasionally backfire, and affect rare native species as well. For instance, poisons used to eradicate mice also killed the North Island saddleback, a rare New Zealand native forest bird (Davidson and Armstrong 2002). Also, the magnitude of the effects of introduced herbi- vores on New Zealand’s native plants can sometimes be difficult to determine (Veblen and Stewart 1982). In addition, it is worth noting that some species, such as the swallows and silvereyes mentioned by De Roy and Jones, expanded their ranges naturally into New Zealand in recent times. When it comes to the many interactions among introduced species and their new environments, the story is often considerably more complex than this book suggests. Thus, the repeated demonization of introduced species found in the text is often overly simplistic. The authors often mention the negative impact of introduced wild mammals such as rats or Australian brush-tailed possums in New Zealand, but, curiously, seem to largely ignore the huge impact of New Zea- land’s domestic animals, particularly the country’s enormous sheep population. According to Ponting (1993), there are more than 70 million sheep and 8 million cattle in New Zealand, and many of the habi- tats of this country have been changed drastically and MISCELLANEOUS THE CANADIAN FIELD-NATURALIST Vol. 121 irreversibly as a result. Also largely ignored in the book is a discussion of the early, pre-European, human- caused species extinctions. New Zealand’s fauna used to be dominated by very large (the largest ones reached 3 meters in height) flightless birds called moa. These birds became extinct mainly due to overhunting by the Maori — the country’s first human colonizers (Ellis 2004). The disappearance of the moas had important effects on some of the native plant species these giant birds were browsing on and had coevolved with. Thus, although conservation is, of course, a very important goal, it is also essential to recognize that nature is dynamic, and that the initial, pre-human colonization web of life found in New Zealand long ago can no longer be re-created due to the many irreversible changes that have taken place since then. A much more thorough and detailed discussion of New Zealand’s prehistoric life and ecology can be found in the book by Worthy and Holdaway (2002). Despite the shortcomings, the current work by De Roy and Jones represents a nice introduction to the unique natural world of this fascinating and remote country. Literature Cited Davidson, R. S., and D. P. Armstrong. 2002. Estimating impacts of poison operations on non-target species using mark-recapture analysis and simula- tion modelling: an example with saddlebacks. Biological Conservation 105: 375-381. Ellis, R. 2004. No turning back: The life and death of animal species. Harper Collins Publishers, New York. 428 pages. Ponting, C. 1993. A green history of the world: The environment and the collapse of great civilizations. Penguin Books, New York. 430 pages. Veblen, T. T., and G. H. Stewart. 1982. The effects of introduced wild animals on New Zealand forests. Annals of the Association of American Geographers 72: 372-397. Worthy, T. H., and R. N. Holdaway. 2002. The lost world of the moa: Prehistoric life of New Zealand. Indiana University Press, Bloomington and Indianapolis. 575 pages. RADU CORNEL GUIASU Environmental and Health Studies Programme, Glendon College, York University, 2275 Bayview Avenue, Toronto, Ontario M4N 3M6 Canada Beneath My Feet: The Memoirs of George Mercer Dawson By Phil Jenkins. McClelland & Stewart, Toronto. 2007 Hard- cover. 350 pages. 18 photographs, 6 sketches. $34.99. Phil Jenkins set out to write the biography of George M. Dawson, the son of Sir John William Dawson, the first principal and then first president of McGill Uni- versity. Jenkins has instead taken nearly 5000 of George Dawson’s letters, most of them previously unpublished, from the McGill University archives, and has wisely allowed Dawson to write his own “autobiography.” Jenkins has created an introduction in Dawson’s style, has abridged an article published by Dawson in Harp- er’s Magazine, and has written over 5% of the text (we don’t know which segments) to fill in gaps not avail- able in Dawson’s own words. The result is a personal, frank, well-written, entertaining life of George M. Daw- son, CMG, LLD. A designer, “Mr. Richardson” (unaccountably, Jenk- ins does not provide his first name) has produced a charming book cover that fits perfectly with what would have been expected in the 1890s. George M. Dawson was crippled in late childhood by spinal tuberculosis which left him a hunchback and a good foot shorter than he should have been. Nev- ertheless as an adult he struggled manfully through arduous hiking, canoeing and horseback riding that would have deterred many men of normal stature. He obtained superb training at the Royal School of Mines in London, England, where he was taught by Professor 2007 Thomas Henry Huxley, a leading proponent of Dar- win’s theory of evolution. Young George was fav- ourably inclined to Huxley’s views and had spirited discussions on the subject with his father. George M. Dawson was a geologist, author, teacher, civil servant, anthropologist, palaeontologist, surveyor, mapper, photographer, artist and poet. A number of his photographs, sketches, and poems are interspersed throughout this book. Although he was named geolo- gist and naturalist to the North American Boundary Commission in 1873-74, his skills in natural history were weak (Jenkins fails to share this fact). In north- ern Montana, just south of the 49" parallel, he observed the last great herds of Bison and came across skele- tons resulting from the slaughter of 21 plains Indians. Dawson named a number of geographic features in the Queen Charlotte Islands and the interior of British Columbia that are still in use today. He suffered the pain of unrequited love and remained a bachelor until his premature death from an acute chest infection at age 51. I agree with Jenkins (page 2), that Dawson was “a writer ... of endearing wit, evocative description, and illuminating fact.” He described an Orkneyman in the employ of the Hudson’s Bay Company as “an amiable Hudson Bay Fossil.” He used the term “liquid excis- able articles” to describe spirituous liquors. This book has not achieved its full potential. I spot- ted the following misspellings and factual errors, pre- sumably made by Dawson and not by Jenkins: alumni (when alumnus was indicated), latitude (when longi- tude was meant), astrological (astronomical), penin- sular (peninsula), tit bits (tid bits), 100 meridian (110" meridian), prairie chickens (Sharp-tailed Grouse), Dr Helckin (Dr Helmcken), F.R.C. (E.R.S.), silver load (silver lode), and Bearing Sea (Bering Sea). There is The Wolf Totem By Jiang Rong. 2004. Changjiang Literary Art Press. Pages: 478; $18. Yuan RMB. [Editors note: The Penguin Group, under the British Pearson Education Company, bought the copyright and will publish an English version in 2007. This researched- based novel has topped China’s bestseller list for 16 straight months, selling about | million copies. ] The Chinese press has not published a significant book in many years. However, in April of 2004, the Changjiang Literary Art Press of China published the novel-style book The Wolf Totem, which was written by an author bearing the pseudonym Jiang Rong. The response from the media and the public was rapid and it became a best seller. The book is composed of various relatively inde- pendent stories. The narration of each story is full of craftsmanship, firing one’s imagination and stirring one’s emotion. The novel-style book is a marvellous, BOOK REVIEWS 11] little doubt that a University press would have en- couraged Jenkins to insert the correct spellings in square brackets, as well as to include an index, explanatory footnotes (e.g., to add the initials and explain the im- portance of Dr. W. F. Tolmie, surgeon to the Hudson's Bay Company), and a bibliography. The latter would have included Life and Letters of George Mercer Daw- son, 1849-1901, by Lois Winslow-Spragge, Dawson's niece, and would have listed the available photocopies of Dawson’s detailed field notebooks which Jenkins seems to have ignored. Similarly, we are not told that Sapper Duckworth was Dawson’s collector and taxi- dermist in 1873-74, that white cranes were Whoop- ing Cranes, that the blue jay was probably a Steller’s Jay, that the hummingbird nest was likely that of a Rufous Hummingbird, and that the name of the des- tructive but now extinct Rocky Mountain Locust has since changed from Caloptenus spretus to Melanoplus spretus. A longer introductory essay or a postscript could have allowed a more complete assessment of Dawson’s great achievements which Dawson modestly did not mention. We learn, for example, that Dawson received an honorary doctorate from the University of Toronto, but those from Princeton, Queen’s and McGill, and other high honors, are not mentioned. In spite of the shortcomings mentioned, Dawson's skills make this book a treasure, a lasting contribution to Canadian history. I recommend it highly to the gen- eral public. It deserves a place in every university and high school library. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Canada unprecedented way to present many stories of the enigmatic animal of the steppe, the wolf. The stories are not only charming in their sense of humanity, but also colourful and valuable in their understanding of the ecological role played by the Wolf. After reading the book, readers will be enlightened, moved and even shocked by the vivid ecological ethics conveyed in the book. The publication of the book can be regarded as an important event for both literature and ecologists. From the viewpoint of ecology, a steppe may be re- garded as a special type of ecosystem seemingly sim- ple in structure and function, but essentially diverse, complicated and unpredictable. From the perspective of humanity, a boundless steppe may be mysterious, thought inspiring and soul stirring, sometimes touching and sometimes plaintive. In the cultural evolutionary history of the world, the acted steppe as an endless well- spring of artistic creation and a boundless field for sci- 112 entific exploration. The anecdotes and the legends, the music and the paintings related to the steppe are un- countable; the study cases concerning the steppe eco- system are also voluminous. Turning aside from the fascinating visions and thrill- ing scenes the author describes in each story, and turn- ing aside from the totem of wolf at the level of politics and humanities the author elucidates in his narration, the book is of important ecological values. The descrip- tions of the behaviours of the Wolf populations in team- work, reconnaissance, ambuscade, and surprise attack, and the unusual wisdom exhibited in making use of weather and topography, as well as the discussions on the relationships among the populations of Wolf and human, sheep and grass can be regarded as very vivid and enlightening material. However, it is a pity that in an otherwise valuable book that the author does not give the basic scientific and quantitative relationships among the component populations of the ecosystem. This may be beyond the scope of this style of book. All these mysterious stories took place in the steppes of northern China during a special period more than nearly 40 years ago when the author and his classmates as urban school graduates laboured in the countryside in response to the political call of Chairman Mao. At that time, the vast area of steppe of northern China had plenty of water and lush grass, creating a beauti- ful landscape. But afterward, due to overgrazing and farming, the steppe ecosystem degenerated rapidly. China is the second largest country in the world in terms of the steppe area. However, due to the increase in large scale agriculture in the half century from 1949, 5% of the natural steppe area was turned into farmland [This constituted one fifth of the total farm- land of China]. This has resulted in more than a quarter of Chinese land being threatened by desertification. More than 90% of the steppe has been degenerated and such degeneration is expanding at the rate of 2 million km?/yr. Since steppe restoration is becoming harder and harder in China, all the stories about the steppe in the past are becoming almost unrepeatable. This further implicates the extreme rarity and the important value of the material in this book. Ecosystems with multiple-trophic levels are com- plicated and maintaining the stability is critical. In a typical steppe ecosystem with three trophic levels: carnivore, herbivore and grass, the carnivore usually plays a top-down effect on the community structure, and by this means grass tends to be protected to some extent. The theory of trophic cascade and HSS hypoth- esis is useful in explaining, “Why the terrestrial world tended to be green.’ McLaren and Peterson (1994) used tree ring data to show how a top-down effect of Wolf populations on Isle Royale reduced Moose abun- dance resulting in an increase in the growth increment of forage trees. The Wolf Totem shows that in the north- em China about one century ago, harmonious rela- tionships among the component populations of the THE CANADIAN FIELD-NATURALIST Vol. 121 ecosystem did exist, and the relative stability of the pasture was naturally maintained. This supports the rationality of the theories as mentioned above. How- ever, considering the present situations of the rapidly degenerated steppe ecosystem in northern China, the above-mentioned theories encounter limitations, because they do not deal with how and why “the ter- restrial world turned from green to yellow or black” due to over-use and human disturbance. If a distur- bance dimension was added to the theories and each dimension were given a threshold, the theories would become more general, compatible and extendable. The Mongolians, the native people in the northern China, fully understood the natural laws of mutual restriction and dynamic balance among the popula- tion of wolf, sheep and grass in the steppe ecosystem in their long history dealing with nature, and knew how to efficiently live within these laws. They never killed all of the Wolves, never bred the sheep exces- sively and never over-grazed the pasture. The accu- mulation of such ecological wisdom imitating nature, acclimating nature and making use of nature, moulded the Mongolian nationality. On the contrary, by disobey- ing the laws of nature, they have unavoidably learnt a painful lesson about the degeneration of the steppe. When human or social-economic dimensions are added into a natural ecosystem, it would become a social-economic-natural-complex-ecosystem (SENCE). The present steppe ecosystem in northern China is such a SENCE. Keeping such a SENCE developing in a harmonious way is a new challenge to China with ever-increasing population. The classical Chinese philosophy, such as The Doctrine of the Mean, emphasizes human’s active roles such as participation, promotion, coordination and cultivation rather than domination, so as to reach harmony between human and nature. Combining the classical Chinese philoso- phy with the modern ecological theories and using the actual experiences and lessons for references, steppe- like SENCE would be managed more effectively. The Wolf Totem with its stories of a rural youth living among Inner Mongolian nomads and wolves tells of this grassland environment and nomadic culture. Reading The Wolf Totem will bring the reader a better understanding of the ecology of the Chinese-Mongo- lian steppe. Literature Cited McLaren, B. E., and R. O. Peterson. 1994. Wolves, moose and tree rings on Isle Royale. Science 266: 1555-1558. Li Dezut', QIN AILP, AND QIN HONG? 'Lab of Urbanization and Ecological Restoration of Shang- hai; National Field Observation and Research Station in Tiantong Forest Ecosystem of Zhejiang; Department of Environmental Science, East China Normal University, 3663, Zhongshan Rd (N). Shanghai, China 200062) “Jilin Forestry Staff School, Jilin, China Zhejiang University, Hangzhou, China 2007 YOUNG NATURALISTS Gracie. The Public Gardens Duck By Judith Meyrick. Illustrations by Richard Rudnicki, 2007. Nimbus Publishing Limited. Halifax, Nova Scotia B3K SMB Canada. 32 pages. $16.95. Cloth. This is a delightful children’s book, told from the perspective of Gracie the duck. The story is about the duck’s quest for food crumbs in a public park near the CBC building in Halifax. Gracie spends most of her time, quite successfully, getting the attention of differ- ent people she sees in the park so that they will share their food with her. However, one day, people are not giving food to her. “Mr. Black Suit’, the person who usually feeds her, does not proffer up any of his muffin; nor does a pair of school girls, who actually apologize to Gracie. A young girl almost shares a piece of her peanut butter sandwich, until the girl’s mother reminds her of the sign near the park entrance. What sign? What is going on? Gracie asks herself, puzzled about this change in her daily feeding regimen. NEw TITLES Prepared by Roy John + Available for review * Assigned ZOOLOGY Book REVIEWS 113 Finally, hungry Gracie reads some of the words on the sign and discovers that park policy has changed and feeding the ducks is no longer allowed. Gracie takes action, but even this doesn’t work. The hungry duck is then forced to try the insects in the grass and then some plants in the pond, and finds that these are very satisfying. Pleased with herself, she now realizes that there is plenty of food in the park without rely- ing on people for crumbs such as Mr. Black Suit — who comments to Gracie that she has adapted nicely. The story is very compelling and suitable for ages 3 to 8 (at least) —- my 12-year-old niece picked up the book and enjoyed it as much as her younger sisters did. The illustrations are dynamic and it is almost as if the people and ducks in this story are moving as you read it. Truly a delight — with a good lesson learned along the way. ELLEN WOODLEY Liana Environmental Consulting Ltd., RR 4, Fergus, Ontario N1M 2W5 Canada [Editor’s note: The following publications are available from China Scientific Book Services, Jian Nei Da Jie Post Office, 100001-88, Beijing, China or e-mail order: order-csbs@94book.com.cn, Online order: http://www.-hceis. The publication dates and costs vary and not all have English text. Fauna Sinica Invertebrata Volume 1: Crustacea Freshwater Cladocera Fauna Sinica Invertebrata Volume 2: Arthropoda Crustacea Freshwater Copepoda Fauna Sinica Invertebrata Volume 3: Platyhelminthes: Trematoda: Digenea (1) Fauna Sinica Invertebrata Volume 4: Cephalopode Fauna Sinica Invertebrata Volume 5: Hirudinea Fauna Sinica Invertebrata Volume 6: Holothuroidea Fauna Sinica Invertebrata Volume 7: Gastropoda: Meso- gastropoda: Cypraeacea Fauna Sinica Invertebrata Volume 8: Arachnida: Ara- neae:Thomisidae and Philodromidae Fauna Sinica Invertebrata Volume 9: Polychaeta: Phyllo- docimorpha Fauna Sinica Invertebrata Volume 10: Arachnida: Araneae: Araneida Fauna Sinica Invertebrata Volume 11: Gastropoda: Opis- thobranchia: Cephalaspidea Fauna Sinica Invertebrata Volume 12: Bivalvia: Mytiloida Fauna Sinica Invertebrata Volume 13: Arachnida Araneae Theridiidae Fauna Sinica Invertebrata Volume |4: Sacodina: Acan- tharia and Spumellaria Fauna Sinica Invertebrata Volume 15: Myxosporea Fauna Sinica Invertebrata Volume 16: Anthozoa: Acti- niaria, Ceriantharia and Zoanthidea Fauna Sinica Invertebrata Volume 17: Crustacea: Decapoda: Parathelphusidae and Pota midae Fauna Sinica Invertebrata Volume 18: Protura Fauna Sinica Invertebrata Volume 19: Gastropoda: Pul- monata: Stylommatophora: Clausiliidae) Fauna Sinica Invertebrata Volume 20: Bivalvia: Proto- branchia Anomalodesmata Fauna Sinica Invertebrata Volume 21: Crustacea: Mysidacea Fauna Sinica Invertebrata Volume 22: Monogenea Fauna Sinica Invertebrata Volume 23: Anthozoa: Sclerac- tinia: Hermatypic coral 114 Fauna Sinica Invertebrata Volume 24: Bivalvia Veneridae Fauna Sinica Invertebrata Volume 25: Nematoda: Rhab- ditida: Strongylata Fauna Sinica Invertebrata Volume 26: Phylum Granulore- ticulosa Class Foraminiferea Agglutinated Fauna Sinica Invertebrata Volume 27: Hydrozoa and Scy- phomedusae Fauna Sinica Invertebrata Volume 28: Crustacea: Amphi- poda:Hyperiidea Fauna Sinica Invertebrata Volume 29: Gastropoda: Archaeogas-tropoda: Trochacea Fauna Sinica Invertebrata Volume 30: Crustacea: Brachy- ura: Marine Primitive Crabs Fauna Sinica Invertebrata Volume 31 : Bivalvia: Pteriina Fauna Sinica Invertebrata Volume 32: Polycystinea: Nasel- laria: Phaeodarea: Phaeodaria Fauna Sinica Invertebrata Volume 33: Annelida Polychaeta II Nereidida Fauna Sinica Invertebrata Volume 34: Mollusca Gastropoda Tonnace Fauna Sinica Invertebrata Volume 35: Arachnida: Araneae: Tetragnathidae Fauna Sinica Invertebrata Volume 36: Crustace Decapoda Atyidae Fauna Sinica Invertebrata Volume 37: Mollusca Gastropoda Fauna Sinica Invertebrata Volume 38: Chaetognatha: Sagittoidea Fauna Sinica Invertebrata Volume 39: Arachnida Araneae Gnaphosidae Fauna Sinica Invertebrata Volume 40: Echinodermata Ophiuroidea Fauna Sinica Invertebrata Volume 41: Crustacea Amphipoda Gammaridea (1) Fauna Sinica Invertebrata Volume 44: Crustacea Decapoda Palaemonoidea Fauna Sinica Invertebrata Volume 46: Sipuncula Echiura Seashells of China Illustrated Animals of China-Mollusca (Volume 3) Economic Mollusca of China Bivalve Mollusca of China Seas Pearl Science Biology of the Economic Species of Cephalopods in the World Oceans Fossil Bivalves of Yunnan Marine Molluscs and Their Collection Rotifera of China Latin-Chinese Names of Invertebrate Animal THE CANADIAN FIELD-NATURALIST Vol. 121 Invertebrates of Wuling Mountains Area, Southwestern China Vermiculture & vermiprotein Polychaetous Annelids Commonly Seen From the Chinese Waters The Parasitic Helminth Fauna of Shaanxi A List of Parasites for Livestock and Poultry in China Nematology Advances and Perspective (2 volume set) 1: Nematode Morphology, Physiology and Ecology 2: Nematode Management and Utilization Parasitic Nematodes from Equus spp. The Studies on the cestodes in China Monogeneans of Chinese Marine Fishes Parasites and Parasitic Disease of Fishes Fauna of Zhejiang-Trematoda Offshore Radiolaria in China Pathogenic Protozoa in Mariculture Foraminifera and Ostracoda in Bottom Sediments of the East China Sea The Echinoderms of Southern China Crabs of China Seas Illustrated Animals of China-Class Crustacea (Volume 1) Illustrated Animals of China-Class Crustacea (Volume 2) Illustrated Animals of China-Class Crustacea (Volume 3) Fauna of Zhejiang-Crustacea Pictorial Keys To Soil Animals Of China Atlas of shrimps in Taiwan straight Penaeoid Shrimps of the South China Sea Economic Shrimp and Crab of the East China Sea Pelagic Siphonophora in China Seas The Sichuan Farmland Spiders Spiders in Qinghai-Tibet Plateau of China The fauna of Hebei, China-Araneae Economic Fauna of China: Terrestrial Mollusks Economic Fauna of China: Freshwater Mollusks Economic Fauna of China: Marine Mollusks The Ferocious Summer — Adelie Penguins and the Warm- ing of Antarctica. By M. Hooper. 2008. Greystone Books, #201 — 2323 Quebec Street, Vancouver, British Columbia VST 4S7. 336 pages. $29.95 Cloth. * The Ecology and Behavior of Amphibians. By K. Wells. 2007. University of Chicago Press, 1427 E. 60" Street, Chicago, Illinois 60637.1400 pages. US$75. Cloth. 2007 Threatened Amphibians of the World. Edited by S. N. Stuart, M. Hoffmann, J. S. Chanson, N. A. Cox, R. J. Ber- ridge, P. Ramani and B. E. Young. 2008. Lynx Edicions, Montseny, 8, 08193 Bellaterra, Barcelona, Spain. 850 pages. US$24. The Atlas of Breeding Birds of Alberta: A Second Look. The Federation of Alberta Naturalists fan@fanweb.ca * Birds in Love. By Jean Levelle. 2007. Voyageur Press, 729 Prospect Avenue, PO Box 1, Osceola, Wisconsin 54020. 160 pages. $25. Birder’s Conservation Handbook — 100 North American Birds at Risk. By J. Wells. 2007. Princeton University Press, 41 William Street, Princeton, New Jersey, USA, 08540-5237. 464 pages. US$35. Paper. 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Speciation in Birds. By Trevor Price. 2007. Roberts & Company Publishers, 4950 S. Yosemite Street, F2 #197, Greenwood Village, Colorado 80111. U.S. $71 Paper * Handbook of the Birds of the World. Volume 12: Pica- thartes to Tits and Chickadees. Edited by Josep Del Hoyo, Andrew Elliot and David A Christie. 2007. Lynx Edicions, Montseny, 8, 08193 Bellaterra, Barcelona, Spain. 798 pages US$294, Euros 205,00 Cloth. + What Bugged the Dinosaurs. By G. and R. Poinar. 2008. Princeton University Press 41 William Street, Princeton, New Jersey, USA 08540-5237. 312 pages. US$29.95 Cloth. * The Return of Caribou to Ungava. By A. T. Bergerud, Stuart N. Luttich and Lodewijk Camps. 2007. McGill- Queen’s University Press, 3430 McTavish Street, Montreal, Québec. Cloth. * The Freshwater Fishes of British Columbia. By J. McPhail. 2007. University of Alberta Press, Ring House 2 , University of Alberta, Edmonton, Alberta T6G 2E1 Canada. 696 pages. $90. Cloth. How and Why Species Multiply — The Radiation of Dar- win’s Finches. P. and R. Grant. 2008. Princeton University Book REVIEWS 115 Press 41 William Street, Princeton, New Jersey USA 08540- 5237. 272 pages. US$35. Cloth. Handbook of European Freshwater Fishes. By Maurice Kottelat and J. Freyhof. 2007. Steven Simpson Books, 5 Hardingham Road, Norwich, Norfolk NR9 4LX, United Kingdom. US$122. Cloth. The Natural History and Management of the Gopher Tortoise. By R. and P Ashton. Krieger Publishing, P.O. Box 9542, Melbourne, Florida USA 32902-9542. 288 pages. US$60. Cloth. The Wild Mammals of Wisconsin. By Charles A. Long. Pensoft Publishers, Sofia-Moscow, Pensoft Series Faunistica 68. In English. 524 pages. EURO 45 (Paper), EURO 78 (Cloth). + Ecology and Management of the North American Moose, Second Edition. By A. Franzmann and C. Schwartz. 2007. University Press of Colorado, 5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado 80303. 776 pages. US$75. Cloth. + Operation Orca. By D. Francis and G. Hewlett. 2007. Harbour Publishing, Box 219, Madeira Park, British Columbia VON 2HO. 280 pages. $34.95. Cloth. Owls of the United States and Canada: A Complete Guide to Their Biology and Behavior. Wayne Lynch. 2007. The Johns Hopkins University Press, 2715 North Charles Street, Baltimore, Maryland 21218-4363 USA. 264 pages. US$39.95 Cloth. A Field Guide to Seashells and Shellfish of the Pacific Northwest. By R. Harbo. 2008. Harbour Publishing, Box 219, Madeira Park, British Columbia, VON 2HO. Laminated pamphlet, $7.95. *Biology of the Snapping Turtle (Chelydra serpentina). Edited by A.C. Steyermark, M. S. Finkler, and R. J. Brooks. 2008. The Johns Hopkins University Press, 2715 North Charles Street, Baltimore, Maryland 21218-4363. 240 pages. US$75. Cloth. Choosing Wildness — My Life Among the Ospreys. By C. Arbour. 2008. Greystone Books, #201 — 2323 Quebec Street, Vancouver, British Columbia, VST 487. 256 pages. $24.95. Cloth. Behavioral Ecology of Insect Parasitoids. Edited by E. Wajnberg, C Bernstein and Jacques van Alphen. 2007. Blackwell Publishing Inc., Commerce Place, 350 Main Street, Malden, Massachusetts 02148, USA. 464 pages. $89.99 Cloth. * Seashells — Jewels from the Ocean. By B. Titlow. 2007. Voyageur Press. Galtier Plaza, Suite 200, 3800 Jackson Street, St. Paul, Minnesota 55101. 112 pages. US$20. Cloth. * The Smaller Majority. By P. Naskrecki. 2007. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138, USA. 288 pages. US$24.95. Paper. The Snakes of Europe. By Guido Kreiner. 2007. Chimaira. Publishing, Groenesteinlaan 22, 9722 BX Groningen. The Netherlands [distributed in the US by Eco Publishing] GBP 44. 116 Orangutans. By J. Payne and C. Prudente. 2008. The MIT Press, Five Cambridge Center, 4th Floor, Cambridge, Massachusetts 02142-1493. 160 pages. $29.95 Cloth. * Turtles. By C. Franklin. 2007. Voyageur Press, 729 Prospect Avenue, PO Box 1, Osceola, Wisconsin 54020. 160 pages. US$45. BOTANY The Curious World of Carnivorous Plants. By Wilhelm Barthlott, Stefan Porembski, Riidiger Seine, and Inge Theisen [Translated by Michael Ashdown]. Timber Press, 133 SW 2™4 Avenue, Suite 450 Portland, Oregon 97204 USA. 224 pages. US$39.95 Cloth. Figs, Dates, Laurel, and Myrrh — Plants of the Bible and the Quran. By Lytton John Musselman. Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, Oregon 97204, USA 336 pages. US$24.95. Cloth. *Grasses of Colorado. By R. Shaw. 2008. University Press of Colorado. 5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado 80303. 662 pages. US$75. Cloth. Bringing Nature Home — How Native Plants Sustain Wildlife in Our Gardens. By Douglas W. Tallamy. 2007. Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, Oregon 97204 USA. 288 pages. US$27.95. Cloth. Wild Plants of Eastern Canada. By M. Walker. 2008. Nim- bus Publishing Ltd., P.O. Box 9166, Halifax, Nova Scotia B3K 5MB8. 240 pages. $29.95 Paper. OTHER + The Explorer’s Guide to Algonquin Park. By M. Runtz. 2008. The Boston Mills Press, 132 Main Street, Erin, Ontario NOB 1TO. 222 pages. $29.95 Paper. Antarctica. By Jonathan Scott and Angela Scott. 2007. Har- per Collins Publishers Ltd. 2 Bloor Street East, 20" Floor, Toronto, Ontario M4W 1A8. 256 pages. $49.00. Cloth. * Arc of the Medicine Line: Mapping the World’s Longest Undefended Border across the Western Plains. 2007. Duncan and McIntyre, # 201 — 2323 Quebec Street, Vancouver, British Columbia VST 4S7. 384 pages. $36.95 Cloth. * Arctic Hell Ship. By W. Barr. 2007. University of Alberta Press, Ring House 2, Edmonton, Alberta T6G 2E1. 318 pages. $34.95 Paper. The Big Earth Book. By James Bruges. 2007. Alistair Sawday Publishing, The Old Farmyard Yanley Lane, Long Ashton, Bristol BS41 9LR U.K. £19.99. Cloth. Birds and People. Nigel J. Collar, Adrian J Long, Patricio Robles Gil and Jamie Rojo. CEMEX-Agrupaci6én Sierra Madre-BirdLife International, Mexico. 360 pages. GBP $29.99. Cloth. Go Wild with your Camera — DVD. 2007. Chris Gomer- sall, Peter Cairns and Mark Hamblin. http://www.gowildtv. com/index.htm. GBP 19.95. THE CANADIAN FIELD-NATURALIST Vol. 121 The Ornithologist’s Dictionary. By Johannes Erritzoe, Kaj Kampp, Kevin Winker and Clifford B. Frith. 2007. Lynx Edicions, Montseny, 8, 08193 Bellaterra, Barcelona, Spain. 290 pages. 19 Euros Paper. Earth. By Iain Stewart and John Lynch. 2007. The Random House Group Limited, Random House, 20 Vauxhall Bridge Road, London SW1V 2SA. US$31 Cloth. Culturing Wilderness in Jasper National Park — Studies in Two Centuries of Human History in the Upper Atha- basca River Watershed. By I. S. MacLaren, Michael Payne, Peter J. Murphy, PearlAnn Reichwein, Lisa McDermott, C. J. Taylor, Gabrielle Zezulka-Mailloux, Zac Robinson, and Eric Higgs. 2007. University of Alberta Press, Ring House 2, Ed- monton, Alberta T6G 2E1. 400 pages. $45. Paper. Natural Sable Island. By Z. Lucas. 2008. Nimbus Publishing Ltd., P.O. Box 9166, Halifax, Nova Scotia B3K 5M8. 128 pages. $18.95. Paper. + Nikkei Fishermen of the BC Coast. By M. Fukawa. Harbour Publishing, PO Box 219, Madeira Park, British Columbia VON 2HO. 208 pages. $ 39.95. Cloth. Ornithology, Evolution, and Philosphy: The Life and Science of Ernst Mayr 1904-2005. By Jiirgen Haffer. 2007. Springer Medizin Verlag, Heidelberger Platz 3, 14197 Berlin, Germany. 473 pages. US$119 Cloth. Reefs Revealed. By Alex Mustard. 2007. Constable & Robinson Ltd., 3 The Lanchester, 162 Fulham Palace Road, London W6 9ER U.K. 192 pages. GBP 30. Cloth. Beneath the Surface. By David Miller. 2007. Langford Press 10 New Road, Langtoft, Peterborough PE6 9LE UK. £37.99. Cloth. Encyclopedia of Tidepools and Rocky Shores. Edited by Mark W. Denny and Steven D. Gaines. 2007. University of California Press, 120 Berkeley Way, Berkeley, California 94704-1012. GBP 56. Cloth. * Hunters at the Margin: Native People and Wildlife Con- servation in the Northwest Territories. By John Sandlos. UBC Press, 2029 West Mall, Vancouver British Columbia V6T 1Z2 . 360 pages. $85 Cloth, 39.95 Paper. Paddling and Hiking the Georgian Bay Coast. By Kas Stone. 2008. The Boston Mills Press, 132 Main Street, Erin, Ontario, NOB 1TO. 256 pages. $29.95 Paper. Science and Conservation of Vernal Pools. By A. Calhoun and P. Maynadier. CRC press LLC Headquarters 2000 NW Corporate Boulevard, Boca Raton, Florida 33431, USA. 363 pages. US$99.95. Cloth. How to be Wild. By Simon Barnes. 2007. Short Books, 3A Exmouth House, Pine Street, Exmouth Market, London, ECIR OJH. 282 pages. GBP 14.99 Cloth. 2007 BooK REVIEWS 117 News and Comment Marine Turtle Newsletter (115) January 2007. 32 pages: EpiroriALs: Guest Edi- torial (P C. H. Pritchard) — Beach Driving Manage- ment Cape Hatteras and Cape Lookout National Seashores, USA (L. R. Nester and N. B. Frazer) — ARTICLES: Interactions Between Marine Mammals and Turtles (D. Fertl and G. L. Fulling) — Identifica- tion of Bacterial Isolates from Unhatched Loggerhead Sea Turtle Eggs in Georgia, USA (K. S. Cravem. J. Awong-Taylor, L. Griffiths, C. Bass, and M. Musca- rella — Nores: Captive-raised Loggerhead Turtle Found Nesting Eight Years After Release (A. de Padua Almeida, C. Baptistotte, A. S’A dos Santos, and T. Z. Serafinai) — Tarballs and Early Life Stages of Sea Turtles in Paraiba, Brazil (R. G. Santos and E. F. Mariano) — Rapid Survey of Marine Turtles in Agalega, Western Indian Ocean (O. Griffiths and V. Tatayah) — Kemp’s Ridley Shell Damage (W. N. Witzell) — Live Loggerhead Observed in Newfoundland, Canada in Late Autumn (W. Ledwell) — IUCN-MTSG Quar- TERLY REPORT — MEETING REPORTS — OBITUARY: Boyd Nathaniel Lyon 1969-2006 (Peter C. H. Pritchard) — ANNOUNCEMENTS — News & LEGAL BRIEFS — RECENT PUBLICATIONS. The Marine Turtle Newsletter is edited by Lisa M. Campbell, Nicholas School of Environment and Earth Sciences, Duke University, 135 Duke Marine Lab Road, Beaufort, North Carolina 28516 USA; and Matthew H. Godfrey, North Carolina Wildlife Re- sources Commission, 1507 Ann Street, Beaufort, North Carolina 28516 USA. Subscriptions and donations towards the production of the MTN can be made online at or postal mail to Michael Coyne (Managing Editor) Marine Turtle Newsletter, 1 Southampton Place, Durham, North Carolina 27705 USA; e-mail: mcoyne @seaturtle.org. 118 THE CANADIAN FIELD-NATURALIST Vol. 121 Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit by post to the Editor, in either English or French, three complete manuscripts written in the journal style. Manuscripts may also be submitted (one copy) by e- mail. The research reported should be original. It is recom- mended that authors ask qualified persons to appraise the paper before it is submitted. All authors should have read and approved it. Institutional or contract approval for the publica- tion of the data must have been obtained by the authors. Also authors are expected to have complied with all pertinent leg- islation regarding the study, disturbance, or collection of ani- mals, plants or minerals. The place where voucher specimens have been deposited, and their catalogue numbers, should be given. Latitude and longitude should be included for all indi- vidual localities where collections or observations have been made. Manuscripts should be printed on standard-size paper, dou- blespaced throughout, generous margins to allow for copy marking, and each page numbered. For Articles and Notes provide a bibliographic (citation) strip, an abstract, and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manu- scripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in a separate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “Canadian” or “American” spellings are acceptable in English but should be consistent within one manuscript. The Oxford English Dictionary, Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Illustrations Photographs should have a glossy finish and show sharp contrasts. Electronic versions should be high resolution. Pre- pare line drawings with India ink on good quality paper and letter (don’t type) descriptive matter. Photographic reproduc- tion of line drawings should be no larger than a standard page. Write author’s name, title of paper, and figure number on the lower left corner or on the back of each illustration. Reviewing Policy Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor (who reviews it or asks another qualified person to do so), and at least one other reviewer, who is a specialist in the field, cho- sen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—some- times extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by pay- ing $90 for each page, plus $30 for each illustration (any size up to a full page), and up to $90 per page for tables (depend- ing on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Editor. Limited joumal funds are available to help offset publi- cation charges to authors without grants or institutional sup- port. Requests for financial assistance should be made to the Editor when the manuscript is submitted. Reprints An order form for the purchase of reprints or pdf will ac- company the galley proofs sent to the authors. Invoices for publication costs will be sent when the submission is pub- lished. FRANCIS R. Cook, Editor RR 3 North Augusta, Ontario KOG IRO Canada TABLE OF CONTENTS (concluded) Volume 121 Number 1 Tributes A tribute to Neal Philip Simon 1973-2006 TONY E. CHUBBS and FRANK R. PHILLIPS Book Reviews Zoo.oGcy: Amphibians and Reptiles of British Columbia — Blue Grouse: Their Biology and Natural History — Birds of Peru — The Freshwater Fishes of British Columbia — Owls of the World — Ecology, Conservation, and Status of Reptiles in Canada — Rodent Societies: An Ecological & Evolutionary Perspective — Seashells; Jewels from the Ocean — Turtles: An Extraordinary Natural History 245 Million Years in the Making — Whales and Seals Biology and Ecology ENVIRONMENT: China Ecosystems — New Zealand — A Natural History MISCELLANEOUS: Beneath My Feet: The Memoirs of George Mercer Dawson — The Wolf Totem YOUNG NATURALISTS Gracie, The Public Gardens Duck New TITLES News and Comment ‘Marine Turtle Newsletter (115) Advice to Contributors Mailing date of the previous issue 120(4): 12 February 2008 2007 100 108 110 113 113 117 118 THE CANADIAN FIELD-NATURALIST Volume 121 Number 1 2007 Articles Were native people keystone predators? A continuous-time analysis of wildlife observations made by Lewis and Clark in 1804-1806 CHARLES E. KAy 1 The influence of air pollution on corticolous lichens near Strathcona industrial area, Alberta M. ELSINGER, E. BURRELL, N. DEBRUYN, K. TANASICHUK, and K. TIMONEY Ay The hepatic flora and floristic affinity of hepatics around Takakia Lake, Queen Charlotte Islands, British Columbia WON SHIC HONG 24 Comparison of scales, pectoral fin rays and opercles for age estimation of Ontario Redhorse, Moxostoma species S. M. REID 29 A rare case of completely ambicoloured Atlantic Halibut, Hippoglossus hippoglossus, from the lower St. Lawrence estuary, Quebec DENIS CHABOT and ROBERT MILLER 35 Micro-propagation of White-top Aster, Seriocarpus rigidus, a threatened species from the Garry Oak ecosystem in British Columbia BRENDA FREY, CHAIM KEMPLER, and DAviD L. EHRET 40 Pattern and potential causes of White-faced Ibis, Plegadis chihi, establishment in the northern prairie and parkland region of North America JILL A. SHAFFER, GREGORY A. KNUTSEN, RON E. MartIN, and JOEL S. BRICE 46 Recent distribution of the Little Brown Bat, Myotis lucifugus, in Manitoba and northwestern Ontario JOHN E. DUBOIS and KIMBERLY M. MONSON 57 Immobilization of Elk, Cervus elaphus, with Telezol and Xylazine and reversal with Tolazine or Yohimbine RICK ROSATTE 62 First confirmed breeding of the Marbled Godwit, Limosa fedoa. in Quebec FRANCOIS MORNEAU, BENOIT GAGNON, and SIDNEY WHISKEYCHAN 67 Swift Fox, Vulpes velox, den use patterns in northwestern Texas BRADY K. MCGEE, WARREN K. BALLARD, and KERRY L. NICHOLSON 71 Flowering during January in Antigonish County, Nova Scotia DaviD J. GARBARY and BARRY R. TAYLOR 76 The occurrence of Muskoxen, Ovibos moshatus, in Labrador Tony E. CHUBBS, and J. BRAZIL 81 The Spindle Tree, Euonymus europaea L. (Celastraceae): A newly naturalized shrub in Nova Scotia Davib J. GARBARY and Amy E. DEVEAU 85 Notes Serviceberry, Amelanchier intermedia, escaped from cultivation in eastern Ontario PAuL M. CATLING 89 A freshwater hydrobiid, cf. the Squat Duskysnail, Lyogyrus granum (Mollusca), widespread and abundant in the Hampton Marsh, New Brunswick | DONALD F. MCALPINE, RACHEL H. MAILLET, ANDREW J. ALBERT, LAURA M. CROSSMAN, REBECCA R. SMITH, and ANDRE MARTEL 92 Northern range extension of the Pygmy Shrew, Sorex hoyi, in the Yukon | THOMAS S. JUNG, TRoy D. PRETZLAW, and Davip W. NAGORSEN 94 (continued on inside back cover) ISSN 0008-3550 CAN 27 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 121, Number 2 April-June 2007 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Michaille Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage: to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields | as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- ments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Charles D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson 2007 Council President: Mike Murphy Annie Belair Diane Kitching Dan Millar Vice-President: Ken Allison Ronald E. Bedford Karen McLachlan Hamilton Stanley Rosenbaum Recording Secretary: Susan Laurie-Bourque Fenja Brodo David Hobden Henry Steger Treasurer: Frank Pope Julia Cipriani Diane Lepage Chris Traynor Past President: Gary McNulty William J. Cody Ann Mackenzie Eleanor Zurbrigg Francis R. Cook Gillian Marston To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2; or e-mail: ofnc @ofne.ca. For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward our mailing costs. PAP Registration Number 9477. Canada Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KO G IRO; (613) 269-3211; e-mail: cfn@ofne.ca Copy Editor: Elizabeth Morton Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5; e-mail: r.john@rogers.com Associate Editors: Robert R. Anderson Charles D. Bird Paul M. Catling Anthony J. Erskine Donald F. McAlpine Warren B. Ballard Robert R. Campbell Brian W. Coad David Nagorsen William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail or e-mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. Subscriptions and Membership Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 pe year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) ant $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) mus! add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Loca Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries a $33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be maile: to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publication Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: April-June 2007 (December 2007). Cover: Four Maritime Garter Snakes, Thamnophis sirtalis pallidulus, observed in courtship behavior on the branches C’ Rhododendron shrub at 325 Prince Albert Road, Dartmouth, Nova Scotia, on 5 September 2006. See note by Gilhen an Strum pages 210-211. WIZ LIBRARY DEC 04 2008 HARVARD PHY ER SEY April-June 2007 The Canadian Field-Naturalist Volume 121, Number 2 Diversity and Distribution of the Terrestrial Mammals of the Yukon Territory: A Review BRIAN G. SLOUGH! and THoMas S. JUNG? '35 Cronkhite Road, Whitehorse, Yukon Territory Y! A 589 Canada; e-mail: slough@northwestel.net Yukon Department of Environment, Box 2703, Whitehorse, Yukon Territory Y 1A 2C6 Canada; e-mail: thomas.jung @ goy.yk.ca Slough, Brian G., and Thomas S. Jung. 2007. Diversity and distribution of the terrestrial mammals of the Yukon Territory: A review. Canadian Field-Naturalist 121(2): 119-127. The diversity and distribution of the terrestrial mammals of the Yukon has not been systematically reviewed since 1975, a time when the occurrences of many species were not well known. Since then, single species and community studies, biological col- lections and expert observations have increased our knowledge of the land mammals of the Yukon. Taxonomic studies have resulted in some additional species. We provide an update on the diversity and distribution of recent land mammals of the Yukon, including previously unreported range extensions, and include a revised checklist. Research in adjoining jurisdictions has contributed more hypothetical species for the Yukon. The mammalian fauna of the Yukon is in a relatively dynamic state, and human-induced changes, particularly climate warming, will undoubtedly influence mammalian diversity and distributions in the coming decades. Key Words: biological diversity, Beringia, checklist, distribution, Mammalia, Yukon. Philip Youngman (1975) provided the most com- prehensive survey of the recent mammals of the Yukon. That information was based on a survey of observa- tions and biological collections from the late 1800s to mid-1970s, and is now over 30 years old. A recent increase in interest in mammals, particularly non-har- vested species, rare species, and species at risk, has resulted in new information and corrections to previ- ously reported distributions. The evolution of the ter- ritorial wildlife management agency, the establishment of a conservation data centre, several university-based research projects (most notably near Kluane Lake; see Krebs et al. 2000), documentation of local and tradi- tional knowledge, and a number of resident and visit- ing naturalists have all contributed much new infor- mation. Changes in taxonomy and nomenclature have also occurred since Youngman (1975), with the former particularly advanced with the use of phylogeographic analyses to differentiate taxa. DNA analyses have also helped detect the presence of cryptic species (e.g., Myotis, Myodes, and Peromyscus spp.) that are difficult to identify to the species level in the field. Herein, we provide an update of the terrestrial mammals of the Yukon Territory; an interesting zoogeographic area of Canada with both Beringian and post-glacial immi- grant species from southern refugia. Methods We reviewed information from published and un- published work, collections, and sightings made since Youngman (1975), primarily chronicling the period from the mid-1970s to the present. We identified and report on changes in species known to be present and update their distributions since Youngman (1975) and others (e.g., Banfield 1974; van Zyll de Jong 1983, 1985). Species without noteworthy changes are not dis- cussed in our review; however, all documented species occurring in the Yukon are provided in a checklist (Table 1). Scientific names and English common names follow Wilson and Reeder (2005) except Caribou which replaces Reindeer. In addition, we have also extrapo- lated ranges of species found in neighboring jurisdic- tions to compile a list of hypothetical species. We did not consider the marine mammals of the Beaufort Sea. Order Soricomorhpa (Insectivores) Seven species of soricomorphs, representing one family (Soricidae), are recorded from the Yukon (van Zyll de Jong 1983; Table 1). Few surveys specifically for shrews have been undertaken in the Yukon and because they are most efficiently captured in pitfall traps (Prince 1941; MacLeod and Lethiecq 1963), they are likely undersampled using standard methods for small rodents (i.e., live-traps or snap-traps). My) 120 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Checklist of the terrestrial mammals of Yukon. Scientific and English common names follow Wilson and Reeder (2005). Order / Family Scientific Name English Common Name SORICOMORPHA Soricidae Sorex arcticus Arctic Shrew Sorex cinereus Cinereus Shrew Sorex hoyi American Pygmy Shrew Sorex monticolus Dusky Shrew Sorex palustris American Water Shrew Sorex tundrensis Tundra Shrew Sorex ugyunak Barren Ground Shrew CHIROPTERA Vespertilionidae Eptesicus fuscus Big Brown Bat Myotis lucifugus Little Brown Myotis Myotis septentrionalis Northern Myotis LAGOMORPHA Ochotonidae Ochotona collaris Collared Pika Leporidae Lepus americanus Snowshoe Hare RODENTIA Sciuridae Glaucomys sabrinus Northern Flying Squirrel Marmota caligata Hoary Marmot Marmota monax Woodchuck Spermophilus parryii Arctic Ground Squirrel Tamias minimus Least Chipmunk Tamiasciurus hudsonicus Red Squirrel Castoridae Castor canadensis American Beaver Cricetidae Dicrostonyx groenlandicus Nearctic Collared Lemming Dicrostonyx nunatakensis Ogilvie Mountains Collared Lemming Lemmus trimucronatus Nearctic Brown Lemming Microtus longicaudus Long-tailed Vole Microtus miurus Singing Vole Microtus oeconomus Root Vole Microtus pennsylvanicus Meadow Vole Microtus xanthognathus Taiga Vole Myodes gapperi Southern Red-backed Vole Myodes rutilus Northern Red-backed Vole Neotoma cinerea Bushy-tailed Woodrat Ondatra zibethicus Common Muskrat Peromyscus keeni Northwestern Deermmouse Peromyscus maniculatus North American Deermouse Phenacomys ungava Eastern Heather Vole Synaptomys borealis Northern Bog Lemming Dipodidae Zapus hudsonius Meadow Jumping Mouse Zapus princeps Western Jumping Mouse Erethizontidae Erethizon dorsatum North American Porcupine CARNIVORA Canidae Canis latrans Coyote Canis lupus Wolf Vulpes lagopus Arctic Fox Vulpes vulpes Red Fox Ursidae Ursus americanus American Black Bear Ursus arctos Grizzly Bear Ursus maritimus Polar Bear Mustelidae Gulo gulo Wolverine , Lontra canadensis North American River Otter Martes americana American Marten Martes pennanti Fisher Mustela erminea Ermine Mustela nivalis Least Weasel Neovison vison American Mink Felidae Lynx canadensis Canadian Lynx Puma concolor Cougar 2007 TABLE |. (continued) Order / Family Scientific Name ARTIODACTYLA Cervidae Alces americanus Cervus elaphus Odocoileus hemionus Odocoileus virginianus Rangifer tarandus Bison bison Oreamnos americanus Ovibus moschatus Ovis dalli Bovidae Stewart et al. (2003) reported a specimen of Pygmy Shrew, Sorex hoyi, from the Ogilvie Mountains which would be a northern range extension; however the co- ordinates given for the specimen align with Dawson City (64°03'N, 139°25'W). Subsequently, a specimen was retained in 2005 from the Blackstone River in the Ogilvie Mountains (65°05'N, 138°11'W; Jung et al. 2007), representing a northern range extension of about 110 km. The range of the American Water Shrew, S. palus- tris, has been extended northward since Youngman (1975) and van Zyll de Jong (1983) and it is now known to as far north as the Tombstone Range of the southern Ogilvie Mountains (Jarrell 1986; Cook et al. 1997). It has been observed near the tree line on Hamilton Creek, 30 km southeast of the Tombstone Campground (64°19'N, 137°38'W; J. Schou, person- al communication). The Tundra Shrew, S. tundrensis, is known from the northern Yukon south to Fortymile (64°26'N, 140°32'W Youngman 1975), where three additional specimens were obtained in 2003 (B. Slough and T. Jung, unpub- lished data). In the late 1970s five Tundra Shrews were trapped in alpine and subalpine habitats on the Haines Road in British Columbia within 20 km of the southwest Yukon, and about 500 km south of its pre- viously known limits (Nagorsen and Jones 1981). It was thought to be limited to Beringia, a region unglaciat- ed during the Wisconsin glaciation; however, the British Columbia locations are 500 km from the lim- its of Beringia and confound the expected distribution pattern (Nagorsen and Jones 1981). It may yet be found in the intervening area in the Yukon. The Arctic Shrew, S. arcticus, remains known only from a single skull collected in 1965 (Youngman 1975), despite substantial subsequent survey effort for shrews (e.g., T. Jung and colleagues, unpublished data; J. Cook and colleagues, unpublished data). Further col- lections are required to assess its distribution in the Yukon. Order Chiroptera (Bats) Three species of bats, representing one family (Ves- pertilionidae), are reported from the Yukon (Table 1). SLOUGH AND JUNG: TERRESTRIAL MAMMALS OI rHE YUKON 12] English Common Name Moose Elk Mule Deer White-tailed Deer Caribou American Bison Mountain Goat Muskox Dall’s Sheep Youngman (1975) reported only the Litthe Brown Myotis, Myotis lucifugus, from the Yukon, where it is widespread and common south of 64°N. Since Young- man (1975), the Northern Myotis, M. septentrionalis, has been confirmed (Jung et al. 2006a) and another species, either the Big Brown Bat (Eptesicus fuscus) or Silver-haired Bat (Lasionycteris noctivagans), has been recorded via acoustic sampling (B. Slough, un- published data). Until recently, few mist-net or echolo- cation monitoring surveys for bats had been under- taken in the Yukon, and it is likely that the Order remains undersampled. Based on surveys in adjacent jurisdictions, there may be other species of bats pres- ent in the Yukon that are not yet documented. The Northern Myotis, M. septentrionalis, was first found in the Yukon in 2004 when three individuals were captured at the La Biche River (60°08'N, 124°04'W; Jung et al. 2006a). Five additional specimens were cap- tured in 2007 near Watson Lake (60°07'N, 128°49'W; Lausen et al. 2008). This species has also been cap- tured at several locations in the Liard watershed of northern British Columbia (Wilkinson et al. 1995*; Bradbury et al. 1997*; Crampton et al. 1997*; Vonhof et al. 1997*; Vonhof and Wilkinson 1999*) and in the southern Northwest Territories as far west as Nahanni National Park (van Zyll de Jong 1985; Lausen 2006*). Youngman (1975) had listed the Big Brown Bat, E. fuscus, as a hypothetical species for the Yukon based on a record from interior Alaska (Reeder 1965). In September 1999, B. Slough (unpublished data) made a recording of a bat echolocation call sequence with an AnaBat II bat detector (Titley Electronics, Ballina, New South Wales, Australia) at Morris Lake (60°25'N, 131°38'W) that was most likely E. fuscus (C. Corben, personal communication; Keinath 2004*). This is pos- sibly the first record of the species from the Yukon, although it may be difficult to distinguish the echolo- cation calls of E. fuscus from those of Silver-haired Bats, as recorded with an AnaBat detector (Betts 1998). There is a single record from central Alaska (Reeder 1965) and unsubstantiated reports exist for southeast Alaska (MacDonald and Cook 1996). E. fuscus had been recorded on bat detectors in the Liard watershed in northern British Columbia (Wilkinson et al. 1995*: 122 Bradbury et al. 1997*; Crampton et al. 1997*) and southwestern Northwest Territories (Lausen 2006*). Order Lagomorpha (Pikas and Hares) As noted by Youngman (1975), two species of lago- morphs, representing two families (Ochotonidae and Leporidae) are recorded from the Yukon (Table 1). The southeastern limit for the Collared Pika, Ocho- tona collaris, shown by Youngman (1975) was the Logan Mountains. It has since been sighted approxi- mately 200 km east-southeast in the Kotaneelee (2004; 60°27'N, 124°12'W) and La Biche (1998; 60°14'N, 124°32'W) ranges (B. Bennett, personal communica- tion). Order Rodentia (Rodents) There are 26 known species of rodents, represent- ing five families (Sciuridae, 6 species; Castoridae, | species; Cricetidae, 16 species; Dipodidae, 2 species; Erithizontidae, 1 species), recorded from the Yukon (Table 1). The Northern Flying Squirrel, Glaucomys sabrinus, occurs throughout the forests of the southern Yukon (Youngman 1975). It has been observed near the tree- line on Hamilton Creek in the Ogilvie Mountains (64°20'N, 137°39'W; J. Schou, personal communica- tion), confirming the northern range limit projected by Youngman (1975). The Hoary Marmot, Marmota caligata, has been confirmed in the La Biche Range in the extreme south- east Yukon (60°14'N, 124°32'W; B. Bennett, personal communication). The Woodchuck, Marmota monax, apparently has a broad range through the southern Yukon; however, sightings and specimens indicate a geographic sepa- ration of at least 370 km between the southern (Ross River; 61°59'N, 132°27'W) and central (Klondike; 63°39'N, 138°40'W) populations which Youngman (1975) shows as contiguous. The Least Chipmunk, Neotamias minimus, was re- ported by Youngman (1975) at Bonnet Plume Lake. It has since been sighted approximately 100 km north- west on the Bonnet Plume River (seven locations from Bonnet Plume Lake to Fairchild Creek (64°58'N, 133°46'W; B. Slough, unpublished data) and approx- imately 80 km northwest on the Wind River (65°03'N, 134°54'W; S. Gilbert, personal communication). The species appears well distributed in the Wernecke Moun- tains. The distribution of the Ogilvie Mountains Lemming, Dicrostonyx nunatakensis, is poorly understood, since it is known from only 13 specimens collected at Angel- comb Peak in the Ogilvie Mountains (64°36'N, 138°16'W; Youngman 1964, 1967, 1975), two of which were recorded recently (T. Jung and B. Slough, un- published data). Krebs and Wingate (1976, 1985) captured Nearctic Brown Lemmings, Lemmus trimucronatus, near the Alaska Highway south of Kluane Lake and from the THE CANADIAN FIELD-NATURALIST Vol. 121 Chilkat Pass, near the Haines Road, in British Colum- bia, extending its known range about 170 km southwest of that reported by Youngman (1975). The Singing Vole, Microtus miurus, is known at present only from the British Mountains, the southern Ogilvie Mountains, and from the Kluane region of the southwestern Yukon (Youngman 1975; Krebs and Win- gate 1976, 1985). Krebs and Wingate (1976, 1985) trapped extensively in the Kluane region and found the southern limit of distribution to be the Slims River area at the south end of Kluane Lake (61°N, 138°31'W). Nagorsen (2002) reported that it has not been found south to the British Columbia border as suggested by Banfield (1974) and Youngman (1975). Jung et al. (2006b) reported the Southern Red- backed Vole, Myodes gapperi, from the Yukon at the La Biche River (60°08'N, 124°04'W). Youngman (1975) did not report M. gapperi from the Yukon; however, he considered M. gapperi and the Northern Red-backed Vole (M. rutilus) to be conspecific. Based on the dis- tribution of M. gapperi in adjacent northeastern British Columbia and southwestern Northwest Territories (Banfield 1974; Nagorsen 2005) we suspect that M. gapperi may be more widespread in the Liard River watershed. Youngman (1975) gave the northwestern limit of the Bushy-tailed Woodrat, Neotoma cinerea, as Klu- ane Lake. It has since been been observed on Ogilvie Island, Yukon River, near Youngman’s projected north- ern limit (63°34'N, 139°44'W; B. Bennett, personal communication). In 2003, a nest of this species was observed at Bear Creek (64°02'N, 139°15'W; B. Slough, unpublished data), representing a 160 km northward range extension. Muskrat, Ondatra zibethicus, inhabit wetlands across the territory (Youngman 1975). The northern- most observations were made in the Arctic wetlands of the Mackenzie Delta in the Yukon (68°46'N, 136°32'W; B. Slough, unpublished data). The Northwestern Deermouse, Peromyscus keeni, had been confirmed through DNA analysis as being present in the Coast Mountains and adjacent areas of the southwestern Yukon (Wike 1998; Lucid and Cook 2004; M. Lucid, personal communication) within the range of the former P. maniculatus algidus. In July 2005, a North American Deermouse, P. man- iculatus, was captured in the Ogilvie Mountains (64°30'N, 138°13'W; T. Jung et al. unpublished data); representing a 70 km northward range extension, but within the range projected by Youngman (1975). Interestingly, Lucid and Cook (2007) provide DNA evidence of an undescribed species of Peromyscus in southwestern Yukon. Wike (1998), working in central Yukon, also reported the possibility of an undescribed species of Peromyscus in the Yukon. Banfield (1974) and Youngman (1975) show the Northern Bog Lemming, Synaptomys borealis, ranging throughout the Yukon north to the Porcupine River drainage. Smits et al. (1989) reported S. borealis teeth 2007 and bone fragments in the scat of an Arctic Fox, Vulpes lagopus, collected in 1985, on Herschel Island, extend- ing the range of S. borealis about 150 km north to the Beaufort Sea (69°35'N, 139°05S'W). The Western Jumping Mouse, Zapus princeps, is known from only two specimens collected in 1944 (Rand 1945; Youngman 1975) and another collected in 1980 (S. Beare, unpublished data); all three speci- mens are from the South Canol Road, near the Rose and Nisutlin rivers. Order Carnivora (Carnivores) Sixteen species of carnivores, representing four fam- ilies (Canidae, 4 species; Ursidae, 3 species; Mustel- idae, 7 species; Felidae, 2 species), are recorded from the Yukon (Table 1). Coyotes, Canis latrans, expanded northward from 55°N to southern Alaska between 1829 and 1907 (Banfield 1974). The earliest reported sighting for the area is from “near Whitehorse, on the Alsek River Alaska, in February 1907” (J. A. Allen 1908, report- ed in Hall 1981). Cairnes (1909) reports the presence of Coyotes in the southern Yukon in 1908. The range has been represented as the entire Yukon (Banfield 1974; Youngman 1975); however, trapper and hunter returns and observations suggest that Coyotes are absent or scarce north of 65°N (Yukon Department of Environment, unpublished data). Extraordinary obser- vations, such as a sighting at Old Crow (67°N, 135°W, C. P. Charlie reported in Youngman 1975) should be considered as extra-limital and not evidence of range expansion. The Arctic Fox, occurs on the Yukon Coastal Plain and Herschel Island, roughly northward of the 150 m ASL contour (Smits and Slough 1993). There is no evidence of denning or other range use further inland, therefore extraordinary specimens from the Old Crow Flats and the Porcupine River (reported in Youngman 1975) should be considered as extra-limital and not evidence of range expansion. Similarly, extraordinary observations of Polar Bear, Ursus maritimus, near Old Crow (reported in Youngman 1975) should be consid- ered as extra-limital and not evidence of range expan- sion. The Yukon range of Polar Bears is also restricted to the North Coastal Plain and Herschel Island. An analysis of fur harvest data from registered trap- lines to 2006 shows that the Fisher, Martes pennanti, is largely confined to the Liard River watershed in the southeast Yukon, ranging west to the Teslin River (60°28'N, 133°18'W) as projected by Youngman (1975). Youngman’s northern limit was approximately 61°N; however, the species ranges 80 km north to 61°40'N (upper Hyland River and Finlayson Lake; B. Slough and T. Jung, unpublished data). Several observations have been made west of the Teslin River, and Fishers have been trapped north of Whitehorse (Klusha Creek; 61°13'N, 135°58'W) and near Haines Junction (60°51'N, 137°36'W). We believe that the species is expanding its range northwards and westwards. SLOUGH AND JUNG: TERRESTRIAL MAMMALS OF THE YUKON 123 The Least Weasel, Mustela nivalis, occurs through- out the Yukon (Banfield 1974) and, contrary to Young man (1975), it is present in the Liard watershed. In- dividuals have been captured by trappers in the Liard watershed (B. Slough, unpublished data; Yukon Depart- ment of Environment, unpublished data). American Mink, Mustela vison, occurs throughout the Yukon, as far north as the Old Crow Flats (Young- man 1975) where it has been captured by trappers (Yukon Department of Environment, unpublished data) but does not appear to be a regular inhabitant north of the treeline (68°25'N; Bee and Hall 1956; Banfield 1974) as suggested by Youngman (1975). Similarly, the North American River Otter, Lontra canadensis, occurs throughout the Yukon, as far north as the British and Richardson mountains (Youngman 1975), but does not likely inhabit the Arctic Coastal Plain (north of the 150 m ASL contour, Bee and Hall 1956; Banfield 1974) as suggested by Youngman (1975). We suggest that an observation from Herschel Island (Wildlife Management Advisory Council (North Slope) and the Aklavik Hunters and Trappers Commit- tee 2003*) was an extra-limital movement. A similar occurrence was reported for the Beaufort Sea of north- eastern Alaska by Haskell (2006). Additionally, we suggest that the Arctic range for Canadian Lynx, Lynx canadensis, shown by Youngman (1975) is extra-limital, and occurs only following years during a population crash. The range of Cougar, Puma concolor, in the Yukon remains somewhat obscure, given the species’ propen- sity for long-distance extra-limital movements. Spo- radic sightings have occurred throughout the southern half of the territory. Although most sightings are unsub- stantiated, they occasionally occur in concurrent geo- graphic clusters and are often associated with the occurrence of Mule Deer, Odocoileus hemionus, (Yukon Department of Environment, unpublished data), pro- viding some credibility to the sighting data. The first Yukon specimen was found dead at Watson Lake (60°04'N, 128°43'W) in November 2000 (Jung and Merchant 2005). Successful breeding by P. concolor in the Yukon has not been documented. Order Artiodactyla (Even-toed Ungulates) Ungulates have long been the management priority in the Yukon, and consequently the best known Order. Nine species of ungulates, representing two families (Cervidae, 5 species; Bovidae, 4 species), are recorded from the Yukon (Table 1). Since Youngman’s (1975) review, White-tailed Deer, Odocoileus virginianus, have been recorded in the Yukon (Hoefs 2001). Since Youngman (1975), introduced populations of Elk, Cervus elaphus, north and west of Whitehorse have grown substantially and now may number over 200 animals, although no reliable estimates are avail- able. In addition to introduced populations, Elk have been reported regularly since 1977 in southeastern Yukon, primarily in the Hyland, Beaver, Coal, La- 124 Biche, and Liard river valleys (Hoefs et al. unpub- lished data*). The most recent reported location in southeast Yukon is from October 2004 at Beaver River, where the resident trapper has seen Elk with some reg- ularity over the past several years (60°'N, 124°25'W; D. MacDonald, personal communication). Whether these animals in the southeast represent populations expanding their range northward from British Colum- bia (Shackleton 1999), or are extra-limital movements, is unknown. Mule Deer, Odocoileus hemionus, appear to have extended their range in the Yukon since the review by Youngman (1975). They are now found throughout the Yukon, south of the Ogilvie Mountains (64°05'W) and including westward to Alaska (Hoefs 2001). Hoefs (2001) reported a 1998 observation north of the Ogilvie Mountains at Chapman Lake (64°50'N, 138°25'W). We regard this observation, however, as an extra-limital movement and not evidence of Mule Deer being dis- tributed north of the Ogilvie Mountains. Reliable observations of White-tailed Deer, Odo- coileus virginianus, some including photographs, have been made in the Yukon (Hoefs 2001). The first obser- vation was reported in 1975 near Tagish Lake (60°10'N, 134°20'W, Hoefs 2001). Mayo (63°05'N, 135°53'W) appears to be the most northern location that White- tailed Deer have been observed in the Yukon (M. O’ Donoghue, personal communication). Hoefs (2001) reported an antlered skull found near Ross River (62°N, 132°25'W) as further evidence of White-tailed Deer in the Yukon. American Bison (Bison bison) (Wood Bison sub- species, B. b. athabascae) have been reintroduced to the Yukon since the review by Youngman (1975). An attempt in 1951 to establish a small herd of Plains Bison, B. b. bison, in the Yukon, by releasing five animals from Alaska, had failed (Youngman 1975). Three Wood Bison herds may now be found in the Yukon; all are reintroduced. The Aishihik Herd ranges in southwestern Yukon, near Aishihik Lake (61°36'N, 137°30'W). Two other, smaller herds — the Nahanni and Nordquist herds — mainly range in the Northwest Territories and British Columbia; Shackleton 1999), respectively, but a portion of their annual range is in southeastern Yukon. The Nahanni Herd ranges into the LaBiche River valley (60°07'N, 124°03'W); the Nordquist Herd ranges along the Alaska Highway, north of Liard Hot Springs in British Columbia, and into the Yukon at Contact Creek (60°01'N, 127°37'W). Both of the southeastern herds were also reintroduced to their native range in the 1990s (Gates et al. 2000). Muskox, Ovibos moschatus, were reintroduced to Barter Island (70°08'N, 143°38'W) and the adjacent mainland in northern Alaska in 1969 and 1970. Muskox were sighted almost yearly in the Yukon since the rein- troduction but they did not appear to establish a year- round breeding population at the time of Youngman (1975). The first mixed sex groups including calves were reported in the Yukon in 1986 (D. Cooley, per- THE CANADIAN FIELD-NATURALIST Vol. 121 sonal communication). They currently range as far east as the Blow River (ca. 137°W; D. Cooley, personal communication). Summer movements to the Old Crow Flats (ca. 68°N, 140°W) have been somewhat com- mon since 1995. A new resident breeding population appears to have formed in the Richardson Mountains (ca. 68°N, 137°W) that remains all year in the moun- tains (D. Cooley, personal communication). Hypothetical Species The following species are found in neighbouring Alaska, British Columbia or the Northwest Territories, and their ranges may extend into the Yukon (Table 2). Some taxa, such as shrews and bats, are undersampled, as are some of the more remote geographic areas. The Alaskan Tiny Shrew, Sorex yukonicus, likely occurs in the Yukon (Table 2), but has yet to be col- lected outside Alaska. This species has been recently described from seven specimens from Alaska in 1993 (Dokuchaev 1997). Subsequently, other specimens (ca. 30) have been obtained from Alaska as a result of directed pitfall trapping efforts (Peirce and Peirce 2000; MacDonald and Cook 2001*; Cook and Mac- Donald 2003*), indicating that the species is wide- spread, but uncommon, in Alaska. Specimens were obtained from Yukon-Charley Rivers National Pre- serve and Wrangell-St. Elias National Park and Pre- serve only 50 km and 12 km, respectively, from the western Yukon border (MacDonald and Cook 2001*:; Cook and MacDonald 2003*). Directed sampling effort in 2003 near the abandoned village of Fortymile (64°25'N, 140°32'W) failed to secure a specimen (B. Slough and J. Jung, unpublished data). Youngman (1975) noted the Long-legged Bat, Myotis volans, as a hypothetical species for the Yukon, based on a record near Atlin, British Columbia, re- ported by Swarth (1936). The species was recently captured in the southwestern Northwest Territories (Lausen 2006*). In addition to M. volans, we add the Western Long-eared Bat, M. evotis, and Silver-haired Bat, Lasionycteris noctivagans, to the list of hypotheti- cal bat species in the Yukon. All three of these hypo- thetical species have recently been captured within 150 km of the Yukon border in northeastern British Columbia (see: Wilkinson et al. 1995*; Bradbury et al. 1997*; Crampton et al. 1997*; Vonhof et al. 1997*; Vonhof and Wilkinson 1999*; Lausen 2006*) and occur in southeast Alaska (Parker et al. 1997). These captures, however, have not been substantiated with a specimen, echolocation recording or a genetic sample (D. Nagors- en, personal communication). Acoustic records also exist for the Hoary Bat, Lasiurus cinereus, and the Eastern Red Bat, L. borealis, from the southwestern Northwest Territories (Lausen 2006*), suggestive that these species may also occur in southeastern Yukon. Parker and Cook (1996) reported Keen’s Myotis, Myotis keenii, from southeastern Alaska (north to ca. 58°N), and it may yet be reported from in or near Klu- ane National Park Reserve in southwestern Yukon. 2007 SLOUGH AND JUNG: TERRESTRIAL MAMMALS OF THE YUKON 125 TABLE 2. Hypothetical terrestrial mammals that may occur in the Yukon. Order / Family Scientific Name SORICOMORPHA Soricidae Sorex yukonicus CHIROPTERA Vespertilionidae Lasionycteris noctivagans Lasiurus borealis Lasiurus cinereus Myotis evotis Myotis keenti Myotis volans LAGOMORPHA Leporidae Lepus arcticus Lepus othus RODENTIA Sciuridae Marmota broweri Muridae Mus musculus CARNIVORA Memphitidae Mephitis mephitis In addition to the resident Snowshoe Hare, Lepus americanus, Youngman noted the possibility of Arc- tic Hare, L. arcticus, or Alaskan Hare, L. othus, being found in northern Yukon. There is an unsubstantiated sight record of L. arcticus from the eastern Richard- son Mountains of the Yukon in the 1970s (P. Frank- ish, personal communication). The Alaska Marmot, Marmota broweri, may occur in the Yukon, but it has not yet been recorded from there. M. broweri is found as far east in Alaska as the Brooks Range in the Arctic National Wildlife Refuge (Bee and Hall 1956); it may also range in the British Mountains in Ivvavik National Park, Yukon. The Wild- life Management Advisory Council (North Slope) and the Aklavik Hunters and Trappers Committee (2003*) reports “two people said they remembered seeing (mar- mots) in 1960s and 1970s in the Babbage and Firth River areas”. The House Mouse, Mus musculus, occurred in the vicinities of Dawson and Whitehorse in the 1970s (Youngman 1975); however, no further specimens have been obtained. This species may be periodically intro- duced, but does not appear capable of sustaining com- mensal populations in the Yukon. Banfield (1974) shows the Striped Skunk, Mephitis mephitis, entering the Yukon in the extreme southeast (La Biche River region) and coming very close in the Teslin area. It has not been documented from the Yukon with the nearest record being two skins taken at the junction of the Fort Nelson and Liard rivers in British Columbia (Rand 1944; Hatler et al. 2008), about 52 km SE of the Yukon. In the Northwest Terri- tories, it is present at Fort Smith, and is expected at Fort Liard (S. Carriére, personal communication), the latter being only 30 km from the Yukon border. English Common Name Alaskan Tiny Shrew Silver-haired Bat Eastern Red Bat Hoary Bat Long-eared Myotis Keen's Myotis Long-legged Myotis Arctic Hare Alaskan Hare Alaska Marmot House Mouse Striped Skunk Conclusions Knowledge of the diversity and distribution of the terrestrial mammals of the Yukon has improved steadi- ly since the last review was published in 1975 (Young- man 1975). Excluding Humans, there are now 63 spe- cies of terrestrial mammals that are believed to occur in the Yukon, representing 40 genera, 15 families and 6 orders (Table 1). There are an additional 11 hypo- thetical species (Table 2). All species, except Mus mus- culus, are native. Only one species, Dicrostonyx nuna- takensis, is endemic. Populations of some species native to the Yukon have been introduced (Cervus elaphus) or reintroduced (Martes americana, Oreamnos amer- icanus, Bison bison). The mammalian fauna of the Yukon is in a relatively dynamic state. Several species have colonized the Yukon in the past 125 years (e.g., Canis latrans, Odocoileus hemionus, and Puma concolor). The history of mam- mal colonization and evolution in Yukon and neigh- bouring Alaska have been discussed by Youngman (1975) and MacDonald and Cook (1996). Approxi- mately one-third of the recent terrestrial mammals of the Yukon originated in Beringia; the remainder are southern immigrants (Youngman 1975). Many of the southern immigrants have limited distributions in the Yukon today. For example, the newly recorded bat species are likely southern immigrants, and Chiropter- ans may still be colonizing northwestern North Amer- ica (Jung et al. 2006a). Myodes gapperi and Peromyscus keeni are similarly southern immigrants. Further cryp- tic species may be present in the Yukon, for example, in the genus Peromyscus (Lucid and Cook 2007) and Myotis (Jung et al. 2006a). Human-induced changes, particularly climate change, will undoubtedly affect mammalian diversity and distributions, particularly in 126 northern regions (e.g., Humphries et al. 2002, 2004). Ranges of some species in the Yukon will likely grow with anthropogenic habitat changes and climate warm- ing (e.g., Mule Deer), while others may contract (e.g., Caribou). Detailed mapping of distribution in the Yukon can assist in monitoring any change in distributional patterns. Appropriate sampling techniques for species typi- cally undersampled, such as echolocation monitoring and mist-netting for bats and pitfall trapping of shrews, has rapidly increased our knowledge of these secre- tive and cryptic species in the Yukon. Sampling in remote areas not well surveyed for mammals will also likely yield interesting records. Further mammal sur- veys, particularly for the smaller species (<1 kg) and in geographic areas with inadequate sampling, will con- tinue to improve our understanding of Yukon mam- mals. Such knowledge is imperative for the conserva- tion of biodiversity and predicting the impact of climate change and landscape change on the diversity, distri- bution and conservation status of Yukon’s mammals. Acknowledgments We thank the numerous individuals and agencies that contributed species-specific information, includ- ing collections and observations, or funding, that sub- stantially improved our knowledge of the diversity, distribution and status of terrestrial mammals of the Yukon. Financial support was provided by the Yukon Department of Environment and the Northern Re- search Institute, Yukon College. David Nagorsen and an anonymous referee reviewed an earlier version of the manuscript. Documents Cited [marked * in text] Bradbury, S. M., S. Morris, and S. McNally. 1997. Bat survey of the Liard watershed in British Columbia: the lower Liard River and Highway 77 area. Wildlife Branch, British Columbia Ministry of Environment, Land and Parks, Victoria. 29 pages. Cook, J. A., and S. O. MacDonald. 2003. Mammal inventory of Alaska’s National Parks and Preserves: Wrangell-St. Elias National Park and Preserve. Annual Report 2001- 2002. Idaho State University. 32 pages. 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A new subspecies of varying lem- ming, Dicrostonyx torquatus (Pallas), from Yukon Terri- tory (Mammalia, Rodentia). Proceedings of the Biologi- cal Society of Washington 80: 31-34. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museums of Canada, Ottawa, Ontario. Publica- tions in Zoology 10: 1-192. Received 21 March 2006 Accepted 9 May 2008 State Records and Habitat of Clam Shrimp, Caenestheriella gynecia (Crustacea: Conchostraca), in New York and New Jersey ROBERT E. ScHMipT! and ERIK KIVIAT Hudsonia Limited, P.O. Box 5000, Annandale, New York 12504 USA 'Present address and correspondence to: Bard College at Simon’s Rock, 84 Alford Road, Great Barrington, Massachusetts 01230 USA; e-mail: schmidt @simons-rock.edu Schmidt, Robert E., and Erik Kiviat. 2007. State records and habitat of Clam Shrimp, Caenestheriella gynecia (Crustacea: Conchostraca), in New York and New Jersey. Canadian Field-Naturalist 121(2): 128-132. We report new records of the Clam Shrimp, Caenestheriella gynecia Mattox 1950, from three localities in the Hudson Valley of New York and one locality in northeastern New Jersey. All of our specimens were collected in puddles on dirt roads. The New Jersey specimens have meristics that are well within the range of Caenestheriella Daday 1915; however, the eastern New York specimens resemble those reported from Massachusetts with meristics that are closer to Cyzicus Audouin 1837. We hypothesize that C. gynecia was established as a parthenogenetic species due to an unlikely dispersal event in the west- ern part of its range and has migrated eastward since the last glacial maximum. Dispersal of this species may occur by wind, in the gut of birds, or stuck to animal feet or fur, or to vehicles. We recommend that C. gynecia be treated as rare and vulner- able to extinction throughout its range unless demonstrated otherwise. Key Words: Clam Shrimp, Caenestheriella gynecia, temporary pond, biogeography, dirt road, New York, New Jersey. Clam Shrimps are small freshwater bivalvular arthro- pods in the order Conchostraca. Clam Shrimps inhabit isolated ephemeral pools, temporary ponds, and some- times lakes; the taxon is found nearly worldwide (Wil- liams 1987). Caenestheriella gynecia Mattox 1950 is a parthenogenetic species with no records of males, although males can be uncommon in related species and may be discovered (see Eder et al. 2000). Until Smith and Gola (2001) reported two populations of this species from western Massachusetts, the known range extended east from Ohio (Smith 2001) to northeastern Pennsylvania. Here we report four localities of C. gynecia in the eastern United States, representing range extensions to New York and New Jersey. Methods Specimens were captured from rain puddles and pre- served in 50% isopropanol. Preserved specimens were measured with digital calipers (greatest length in mm). Spines on the telson and second antennal segments were counted at 320x with a stereomicroscope. Results Here we document four collections of Caenestheriel- la gynecia, representing the first records of this species from New Jersey and New York. New JERSEY: rain puddles on unimproved Paterson Lateral Gas Pipeline road between Empire Avenue and the New Jersey Turnpike, Borough of Carlstadt and Township of South Hack- ensack, Bergen County, 74°02'26"W longitude, 40°49'37"N latitude, Hackensack Meadowlands. [Weehawken, New Jersey— New York. 1967, U.S. Geological Survey (USGS) 7.5 minute topographic map sheet], E. Kiviat, 2 September 2001. 6 spec- imens (Kiviat and MacDonald 2004). New York: rain puddle on abandoned farm road, 3.5 km south of Rhinebeck, Dutchess County, 73°55’15”W longitude, 41°53’ 19”N latitude, Town of Rhinebeck, [Kingston East, New York. 1963, photorevised 1980 USGS 7.5 minute topograph- ic map sheet], S. Nyman, 27 May 1994. 1 specimen, 7.6 mm, dried (Site C-3 in Kiviat et al. 1994*). NEw York: rain puddles on dirt road, ca. 1.4 km NNW of East Park, Dutchess County, 73°55'20"W longitude, 41°48'19"N latitude, Town of Hyde Park, [Hyde Park, New York. 1963, USGS 7.5 minute topographic map sheet] E. Kiviat & J. Bridges, 26 June 2002. 5 specimens. NEw YorRK: rain puddles on ATV trail, ca. 3.4 km NNE of Saugerties, Greene County, 73°55'58"W longitude, 42°06'33"N latitude, Bristol Beach State Park [Saugerties, New York. 1963, USGS 7.5 minute topographic map sheet], E. Kiviat, 25 July 2007. 1 specimen. Clam Shrimp habitats (Figure 1) at the New York and New Jersey localities were all rain puddles on dirt roads. Clam Shrimp were observed swimming slowly on the underside of the surface film in New Jersey, and swimming in the water column and along the bottom in New York. The puddles were ca. 1.5-30 m? and ca. 5-15 cm deep at the dates of collection. The rain puddles essentially lacked vascular plants. We have visited the puddles at the New Jersey locality several times at dif- ferent seasons 2002-2007; some of the puddles main- tained their depths even during dry periods, but did dry during the summer 2005 drought. The New Jersey pud- dles were consistently very turbid and Clam Shrimp were not visible below the surface; the Hyde Park pools were clear enough to see Clam Shrimp swimming along the bottom. The roads at the New Jersey and Saugerties, New York localities are regularly used by all-terrain 128 2007 ts Ee, ans - Sas SCHMIDT AND KIVIAT: CLAM SHRIMP IN NEW YORK AND NEW JERSEY 129 FiGurE |. Habitat of Caenestheriella gynecia in New Jersey. Photograph by Erik Kiviat. vehicles (including “‘four-wheelers” and motorcycles), and road vehicles (pickup trucks and sport-utility vehi- cles) were observed using the road at the Hyde Park locality. The puddles at the Bristol Beach State Park site were created and maintained by all-terrain vehicle use. We believe the puddle habitats at these four sites were created and maintained by wheeled vehicles; the puddles at the Rhinebeck locality appeared less recent- ly disturbed and may have been created by farm vehi- cles. The dirt road at the Hyde Park locality was bor- dered by hardwood forest and woodland pools. The dirt road at the Rhinebeck locality was fringed by tall shrubs and small trees in the midst of a wet meadow and old field. In New Jersey, the dirt road was fringed by Com- mon Reed (Phragmites australis), tall shrubs, and small trees, and was raised ca. 1.5 m above the level of extensive, formerly tidal, marsh now dominated by common reed. The ATV trail at the Bristol Beach site was bordered by second-growth hardwood forest. The Hyde Park and Rhinebeck roads appear to com- prise in situ soil materials which are mapped as Nas- sau-Cardigan complex (Lithic and Typic Dystrochrepts) and Sun silt loam on glacial till, and Hudson-Vergennes soil (Glossaquic Hapludalf) on glaciolacustrine silty clay, respectively (Faber 2002*). The Saugerties site has soils mapped as Hudson silt loam, a Glassaquic Hapludalf and Madalin silty clay loam, a Mollic Ochra- qualf according to Tormes (1979*). The pH of these soils varies from strongly or very strongly acidic at the surface to moderately akaline in the substratum (Faber 2002*). Wetlands adjoining the New Jersey site were mapped as Sulfhemists and Sul- faquents (Goodman 1995*); however, the dirt road ap- pears to have been constructed from imported materi- al with a light reddish color that may have originated from nearby red Triassic shale and sandstone (Schu- berth 1968). We have not found Clam Shrimp in wet- lands adjoining the dirt road habitats. Smith and Gola (2001) discussed generic placement of this species and showed that the meristics on their Massachusetts specimens were not typical of Caenes- theriella but were intermediate or closer to species of Cyzicus Andouin 1837. The most significant difference in the genera is the shape of the male rostrum which cannot be determined in our specimens. There are dif- ferences in the number of spines on the telson, but the counts on the Massachusetts specimens overlapped the genera and Smith and Gola (2001) questioned whether the two genera were distinct. The size and meristics of our New Jersey specimens (Table 1) fall clearly within Caenestheriella Daday 1915 (all specimens contained eggs) as did the single specimen from Saugerties (west of the Hudson River). Our New York specimens from east of the Hudson River, however, are similar to those in western Massachusetts. It is possible that the eastern New York and Massachusetts populations are Cyzicus. Discussion Caenestheriella gynecia inhabits temporary fresh- water pools and was considered a “warm water” species 130 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Size and meristics of Caenestheriella gynecia (Clam Shrimp) specimens from New Jersey and New York. State Number Mean Number of Number of of specimens (range) size, mm telson spines segments, 2nd antennae New Jersey 6 4.75 (4.2-5.2) 18-22 15-19 New York 5 9.10 (8.6-10.2) 27-32 15-20 by Weeks and Marcus (1997). This is typical habitat for a suite of clam shrimp and relatives (Anostraca, Noto- straca) and for certain distantly related species of Cla- docera and Ostracoda (Williams 1987). The diversity of aquatic animals in a given pool is low, however, probably due to the habitat’s inherent instability and simplicity. Caenestheriella gynecia appears to be dif- ferent from its close and distant relatives in that it is known mostly from rain puddles on dirt roads; the type locality (Mattox 1950), a second location in Ohio (Em- berton 1980), one of the populations in Massachusetts (Smith and Gola 2001), and this paper. Although the puddles in which we found C. gynecia are more-or- less permanent, their small size (and high turbidity in the case of the New Jersey pools) may exclude some aquatic predators much as does impermanence. Wiggins et al. (1980) and Williams (1985) character- ized the resident biota of ephemeral aquatic habitats as having high fecundity, short life spans, and rapid mat- uration. Additionally Clam Shrimp produce “resting eggs” (Hann 1996*) which can tolerate heat, cold, and dessication and may require freezing or drying to hatch (Mattox and Velardo 1950; Belk and Belk 1975). Hann (1996*) observed that the small scale distribu- tion of Caenestheriella setosa was patchy, with some puddles inhabited while nearby puddles were empty. This suggests that, despite the Clam Shrimp’s invest- ment in large numbers of resting eggs, egg survival is low, even short-distance dispersal is difficult, or local extinction (in a pool) is frequent. Boileau and Taylor (1994) showed that related organisms had poor disper- sal over even very short distances and calculated that there were 0.6-4.5 dispersers per generation in small pools in South Carolina. We presume that dispersal of Clam Shrimp is primarily in the resting egg stage and we can envision several modes of dispersal of these eggs. Eggs of C. gynecia from desiccated puddles could be blown along with soil particles from one puddle or locality to another. Animals passing through puddles, especially species that spend time in muddy habitats, could pick up eggs in mud adhering to their feet, shells, or fur. Potential vectors of dispersal include Raccoons (Procyon lotor, see Maguire 1963), Mallards (Anas platyrhynchos), Snapping Turtles (Chelydra serpenti- na), and humans. More recently, human traffic in the form of wheeled or tracked vehicles could act as a dispersal agent. Although our few specimens of C. gynecia do little to establish a pattern that would resolve the taxonomic question posed by Smith and Gola (2001), there is a geographic pattern that deserves discussion. All known localities for this species are within the maximum ex- tent of the Wisconsinan glaciation (Figure 2) except one location in central Ohio (Emberton 1980). It is, of course, possible that not all populations of this species have been documented (especially since we are here reporting several) and that there may be populations to the south of the known distribution. This is impor- tant because robust biogeographic hypotheses do require accurate knowledge of an animal’s distribu- tion. Although this species is rare, there are a number of researchers actively working with Clam Shrimp in the midwestern and southern United States and there have been no other reports of this species. This distribution suggests the following hypotheses. Caenestheriella gynecia is derived from a sexually re- producing species, probably C. setosa, whose range is closest to C. gynecia. Dispersal into the glaciated area occurred in the Ohio Valley and was an unlikely event given the dispersal mechanisms of the species. Unlikely dispersal would favor establishment of a parthenoge- netic individual over at least two sexually reproducing ones. This scenario suggests that C. gynecia is relative- ly young, less than 12 500 years old (the approximate age of the last glacial retreat). Dispersal eastward from the Ohio Valley would be equally unlikely except that seasonally muddy dirt roads were created soon after European colonization of the northeastern states. Caenestheriella gynecia could have spread as eggs in mud splattered on wagons or horses’ hoofs and its distribution would follow traffic flow. Re- cently, many of these colonial roads have been paved, leaving C. gynecia to persist in the remaining dirt road habitats, including farm roads, logging roads, and serv- ice roads for powerlines and pipelines. The above scenario could be corroborated by phy- logenetic and genetic analyses. The closest living rel- ative of C. gynecia should be geographically very close to C. gynecia but we are unaware of any phylogenetic analyses of this genus. Additionally, the genetic vari- ation in C. gynecia should be minimal because it is parthenogenetic, has been derived from a very small founder population, and is geologically young. Caenestheriella gynecia or its ancestors could have evolved in association with American Bison (Bison bison), living in “buffalo wallows” and dispersing in mud caked on Bison fur or hoofs. Butler (1995) noted buffalo wallows somewhat larger than the puddles where we collected C. gynecia and alluded to observa- tions of mud stuck to fur. Dispersal of resting stages in the guts of birds is also possible for C. gynecia. Proctor et al. (1967) found that Artemia eggs and the eggs and adults of Ostracoda could remain viable after passing through the gut of Killdeer (Charadrius vociferus). 2007 SCHMIDT AND KIVIAT: CLAM SHRIMP IN NEW YORK AND NEW JERSEY 13] Lae FIGURE 2. Map of the eastern United States showing collection localities of Caenestheriella gynecia in relation to the south- ern extent of the most recent glaciation. Long distance (1400 km) dispersal of Caenestheriel- la belfragei was presumed due to eggs being carried on or in waterfowl (Donald 1989). Proctor (1954) showed that viable eggs of some phyllopods could pass through the gut of ducks. If C. gynecia had been car- ried to the glaciated areas by waterfowl, its close rel- atives could be anywhere along the waterfowl flyways. Two of the New York and the New Jersey Clam Shrimp localities are privately owned. The Hyde Park road is part of a residential subdivision and reportedly has been excavated for installation of a water pipeline. The New Jersey road is subject to gas pipeline main- tenance activities and adjoins The Richard P. Kane Natural Area, possibly subject to habitat restoration. Because field surveys have not been conducted to determine the distribution of, and degree of threat to, C. gynecia populations in the northeastern states, we recommend this species be treated as rare and vulner- able to extinction unless demonstrated otherwise. Only 11 localities (Figure 2) have been reported for C. gyne- cia, including our localities, although a recent Ohio survey listed just one site (Weeks and Marcus 1997). If this Clam Shrimp were reviewed by state Natural Heritage Programs in New York and New Jersey it would be ranked as G2, S1 assuming all the reported sites are extant. We recommend that dirt roads one year or more old with long-lasting rain puddles > 1 m* and >5 cm deep be checked for Clam Shrimp during June- September as part of land management or environmen- tal review processes. Clam Shrimp may sometimes be observed swimming in the pools and may be captured by sweeping a fine-mesh dipnet through the pools. Research, conservation, management, and restoration of biodiversity are most often done in wildlands and natural areas. Yet urban, industrial, and other altered landscapes are expanding rapidly in North America. Certain rare or declining native species survive or even thrive in human-altered landscapes. Examples of rare species occupying human-disturbed soils are: Knies- kern’s Beaked Rush (Rhynchospora knieskernii) in a tire rut and on a waste dump in the New Jersey Pine Barrens (Schuyler 1999) and larvae of the Regal Fritil- lary (Speyeria idalia) overwintering in soils disturbed by tanks in Pennsylvania (Rosenzweig 2003). These ecological distributions can be the result of human activities creating a surrogate habitat type or physi- cal-chemical environment not otherwise available, an abundance of food, reduced predation or competition, or persistence during landscape change (see, e.g., Adams 1994). It can be said that “Biodiversity is where you find it.” Therefore conservation biologists and environ- mental planners need to pay more attention to organ- isms such as Caenestheriella gynecia and the artificial or altered habitats that support them. Altered or arti- ficial habitats of types known to support rare organ- isms should be surveyed for those species routinely, much as natural and semi-natural habitats are surveyed for rare species. Those sites discovered to support a rare 132 species, such as the Clam Shrimp sites reported here, should be considered for conservation management. In the case of Caenestheriella gynecia, management may need to include a degree of controlled motor vehicle activity or its equivalent to maintain the puddle habitat and potentially disperse Clam Shrimp among puddles. We recommend that this species be considered rare and vulnerable to extinction throughout its range unless demonstrated otherwise. Acknowledgments Joseph T. Bridges and Stephen Nyman collaborated in the field. Kathleen Schmidt drew Figure 2. Clam Shrimp were collected and the manuscript prepared dur- ing studies funded by the Geraldine R. Dodge Foun- dation, H2O Fund (Highlands to Ocean Fund), Geof- frey C. Hughes Foundation, Mary Jean and Frank P. Smeal Foundation, Hackensack Meadowlands Part- nership, Matthew D. Rudikoff Associates, and U.S. Environmental Protection Agency. This paper has not been reviewed by the funders; opinions herein are sole- ly those of the authors. This is Bard College Field Sta- tion — Hudsonia Contribution 87. Documents Cited [marked * in text] Faber, M. 2002. Soil survey of Dutchess County, New York. U.S. Department of Agriculture, Natural Resources Con- servation Service. 356 pages + folded maps in case. Goodman, S. D. 1995. Soil survey of Bergen County, New Jersey. U.S. Department of Agriculture, Soil Conservation Service. 142 pages + folded maps in case. Hann, B. J. 1996. Ecology of aquatic invertebrates in tempo- rary habitats: Caenestheriella setosa (Conchostraca, Crus- tacea) in Delta Marsh, MB, Canada. University Field Sta- tion (Delta Marsh) Annual Report 31: 43-46. Kiviat, E., P. M. Groffman, G. Stevens, S. Nyman, and G. C. Hanson. 1994. 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Received 30 January 2006 Accepted 1 May 2008 Colonization of Non-Traditional Range in Dispersing Elk, Cervus elaphus nelsoni, Populations FRED VAN DYKE Montana Department of Fish, Wildlife and Parks, Red Lodge, Montana 59068, USA Present address: Department of Biology, Wheaton College, Wheaton, Illinois 60187 USA; e-mail: fred.g. vandyke @ wheaton.edu Van Dyke, Fred. 2007. Colonization of non-traditional range in dispersing Elk, Cervus elaphus nelsoni, populations. Canadian Field-Naturalist 121(2): 133-141. As ungulate populations increase in density on traditional range, resulting increases in intraspecific competition can encour- age dispersal of some individuals to new areas. Such areas, although lower in density of conspecifics, might present unfamil- iar arrays and types of habitat that could require altered patterns of home range and habitat use by dispersers. However, the specific adaptations employed by dispersers in such circumstances are not well documented or understood. I investigated three cases of range expansion by Elk (Cervus elaphus nelsoni) populations experiencing population growth on traditional ranges in south-central Montana, USA. Each source population produced a group that dispersed to non-traditional areas. Compared to source populations, dispersing groups increased average size of home ranges, changed patterns of use in core areas of home ranges, and used habitats differently than Elk on traditional range. Dispersing groups demonstrated fidelity to new ranges equal to that of source populations, but their seasonal tenure on non-traditional range was strongly linked to envi- ronmental conditions, especially rainfall. Dispersal of groups increased the overall range of the population and its range of habitat use. In growing populations of Elk, managers should determine if dispersing groups exist and whether they should be protected to establish new populations in marginal areas or be reduced to limit potential Elk-landowner conflicts. Key Words: Elk, Cervus elaphus, dispersal, habitat use, population, Montana. On traditional ranges, animals typically concentrate use in the most productive patches of habitat (Charnov 1976; Parker and Stuart 1976). However, intraspecific population pressure, often associated with population growth, can cause a species to increase its range of habitat use in less predictable ways (Rosenzweig 1991). For non-territorial species, the tendency to increase range of habitat use with increasing population densi- ty has been formalized as the Ideal Free Distribution (IFD) Theorem. The Ideal Free Distribution Theorem asserts that individuals choose habitats at densities asso- ciated with equal levels of fitness (Fretwell and Lucas 1970). Thus, IFD predicts that when populations occu- py new ranges at low densities, individuals will occu- py only optimal habitat. As density increases, so does interference with conspecifics, causing individual fit- ness to decline. Individuals leave optimal habitat when fitness drops below what they could achieve in a sub- optimal habitat at a lower population density. Among ungulates, the behavior of Elk (Cervus ela- phus nelsoni) in populations experiencing long-term population growth corresponds to predictions of IFD. Dispersive behavior associated with population growth and an increasing range of habitat use might contribute to the initial stages of colonization and the establishment of a permanent population in a previously unoccupied area (Safriel and Ritte 1983). The colonization of vacant habitat without decreased fitness of colonists is an alter- native to increased intraspecific competition on tradi- tional range. Although the IFD Theorem makes den- sity-dependent dispersal theoretically plausible, such dispersal has been difficult to demonstrate in ungulates (Clutton-Brock et al. 1985; Boyce 1989), and determi- nation of the mechanisms of colonization by individu- als dispersing from established ungulate populations, or their relation to population growth, has been prob- lematic. In three increasing populations in south-central Mon- tana, USA, Elk demonstrated changes in habitat use and home-range characteristics, increasing spatial separa- tion of adjacent populations, and fissioning of individ- ual populations into multiple populations (Van Dyke and Klein 1996; Van Dyke et al. 1998), all of which can lead to an increasing range of habitat use. The last response, population fissioning, was initiated by the formation of a dispersing group that made seasonal movements to new range, often containing a distribu- tion of vegetation communities different from the tradi- tional range of the established population. In investigat- ing the role of such groups in expanding range and habitat use of established populations, I sought to deter- mine what changes dispersing Elk make in their use of home range and habitats compared to their popula- tions of origin that might enable them to persist on new ranges that the established population did not use. Study Area Three populations of Elk in south-central Montana (USA) were investigated, locally known as the Line Creek (LC), Picket Pin (PP), and Silver Run (SR) populations (Van Dyke and Klein 1996: Van Dyke et al. 1998). 133 134 THE CANADIAN FIELD-NATURALIST These populations summer in the Absaroka-Bear- tooth Wilderness Area and winter in lower elevation foothills and prairies on the Custer National Forest and adjoining private land (Figure 1). Climate in this area was characterized by long, cold winters and short, cool summers during the study peri- od. Mean January and July temperatures at the Mys- tic Lake, Montana, reporting station (elevation 2339 m) near the approximate center of the study area were -4.1 and 17.1°C, respectively. Average annual rainfall and snowfall reported at the same location were 64.7 and 497.9 cm, respectively (NOAA 1992). Forests were dominated, from lowest to highest ele- vations, by Limberpine (Pinus flexilis), Lodgepole Pine (P. contorta), Douglas Fir (Pseudotsuga menziesit), Engelmann Spruce (Picea engelmannii), and White- barked Pine (Pinus albicaulis). Non-forested commu- nities at lower elevations were dominated by Bunch- grass (Pseudoroegneria spicata)-forb and sagebrush (Artemisia spp.) associations. Alpine areas were dom- inated by sedges (Carex spp.), cottongrass (Eriopho- rum spp.), Alpine Bluegrass (Poa alpina), and vari- ous alpine forbs. Detailed descriptions of these plant communities are provided in Van Dyke et al. (1991) and Van Dyke et al. (1994). Methods Sampling Procedures Thirty-five female Elk were captured and radio- collared in January-March 1988 (19), December and February 1989 (11), March 1990 (1), and March 1991 (4). All but three were captured by darting from a helicopter (Nielson and Shaw 1967) with Carfentanil (dosage 0.0082 mg/kg). Two Elk were captured in bait- ed traps and one Elk by a net-gun fired from a helicop- ter. All were fitted with telemetry collars (AVM Instru- ment Co., Telonics, Inc.). Procedures associated with locating individual animals have been described (Van Dyke and Klein 1996; Van Dyke et al. 1998). Detection of Dispersing Groups I documented movements of small groups (<40 in- dividuals) away from source populations (120-160 indi- viduals) through relocations of =1 radio-collared Elk in each group. Such groups used different ranges and remained isolated from individuals in the source pop- ulations for 6-9 months each year. One group separat- ed from each source population. These were the Trout Creek (TC) group from the PP population, the Hell- roaring (HR) group from the SR population, and the Wolf Creek (WC) group from the LC population. The relationship of dispersing groups to source pop- ulations, as well as the boundaries of seasonal ranges of the dispersing groups, was unknown prior to the study. As a result, individuals in dispersing groups were unintentionally collared when they were near source populations on winter range. Although only a small number of Elk were radio-collared in each group, the percentages of radio-collared Elk in such groups were Vol. 121 = those collared in source populations (5-12% vs. 4- 6%, respectively). Given the sociability of Elk, previ- ous studies have been successful in tracking large groups even when only one radio-collared animal was present (e.g., Craighead et al. 1973), and that also was the case in this study. Locations of radio-collared individuals were considered representative of the entire group because: (1) dispersing groups contained few (7-40) individuals; (2) dispersing groups remained con- stant in size during occupancy on non-traditional range, indicating that groups possessed a high degree of cohe- siveness; and (3) radio-collared individuals always were present in the group when they were located. Estimation and Analysis of Home Range Characteristics Home range of each population or group was defined as the 95% contour minimum convex polygon (MCP) area and associated activity centers of source popula- tions and dispersing groups as computed by HOME RANGE, a home range analysis program (Ackerman et al. 1990). To estimate seasonal home ranges, sea- sons were defined as winter-spring (1 December — 31 May) and summer-autumn (1 June — 30 November). Seasons were combined in this manner because source populations had substantial overlap between winter and spring ranges and between summer and autumn ranges (Van Dyke and Klein 1996; Van Dyke et al. 1998). Because MCPs are sensitive to sample size, I eval- uated only populations with = 21 locations/season for calculation of seasonal range and =45 total locations for calculation of annual range. All samples were equal to or larger (seasonal x = 97, range 21-438; annual x = 189, range 45-787) than these minimums. Previous tests of the relation between numbers of loca- tions and home area estimates indicate that such sam- ple sizes estimated = 81% of population home ranges for ungulates in this area (Van Dyke et al. 1995; Van Dyke et al. 1998). Core areas within home ranges (Kaufman 1962) were identified by comparing actual range use distributions generated from harmonic mean estimates of home range (Dixon and Chapman 1980) to hypothetical uniform use distributions (Samuel et al. 1985). I also determined the percent contribution of core areas to total home areas and core area per- cent contribution to total use, and then determined the ratio of use contribution to area contribution as an index of the concentration of use in core areas. Evaluation of Range Use Home areas of dispersing groups and source popu- lations were compared in four ways. First, as a first approximation of potential movement differences in landscape pattern and scale, I compared distances Elk in dispersing groups moved from winter to spring or summer ranges with the same movements in their associated traditional populations using linear, straight- line distances between seasonal geometric activity centers in each group as determined by pooled loca- tions from 1988-1992. I compared distances associ- 2007 30 km STONE RNG Ry x ~ GALLATIN NATIONAL FOREST VAN DYKE: COLONIZATION OF NON-TRADITIONAL RANGE BY ELK Ny NORTH ABS Sd Sebi PICKET PIN Nair POPULATION x ewarer ae SILVER RUN POPULATION LINE CREEK POPULATION te SHOSHONE NATIONAL FOREST Ficure 1. Study area and approximate ranges of the Line Creek, Picket Pin, and Silver Run populations of Elk in south-central Montana, USA, 1988-1991. ated with such movements to the radius of the shared winter home area as an index of relative movement scale. Second, I compared home range size between source populations and dispersing groups using 2- sample t-tests. Third, I evaluated Euclidean distances between points in associated dispersing group and source populations through a multiple range permuta- tion procedures test, which evaluates whether distri- butions of distances in different groups come from a common probability distribution (Mielke et al. 1976). Fourth, I compared use-area ratios of core areas be- tween dispersing groups and associated source popu- lations to determine if differences existed in degree of concentration of core area use using a 2-sample rtest. The ratio of core to non-core use is important because core areas would be likely to include the most opti- mal habitat within a home range. Therefore I treated the concentration of use in core areas as an index of relative differences between optimal and suboptimal habitats within a home range. Use of Vegetation Communities and Production of Forage The types of vegetation communities present on all seasonal ranges of all populations varied, but were simplified for analysis to categories of alpine, grass- forb (meadows), Douglas Fir, Limberpine, Lodgepole Pine, sagebrush, Spruce (subalpine forest), and White- bark Pine. Together, these vegetation communities cov- ered the range of elevations used by all populations in all seasons as well as broadly covering the array of vegetation communities encountered and used by EIk. Comparisons of use of vegetation communities by Elk in source populations, based on locations of animals, to availability of such communities visually identified at 568 randomly selected points within home areas of such populations were evaluated using the Design I version of Manly et al. (2002) resource selection function (RSF) model. This version of the RSF model, which uses independently sampled counts of available and used resources, assumes that the availability of vegetation communities does not change over the course of the study period, that vegetation communities were cor- rectly identified, and that Elk had free and equal access to all vegetation communities within their designated ranges, assumptions that were met in this study. The model’s null hypothesis that Elk select vegetation com- munities in proportion to their occurrence was tested by comparing use to availability via a Chi-square good- ness of fit test (Manly et al. 2002). The application of this procedure to similar tests for random selection of 136 THE CANADIAN FIELD-NATURALIST each vegetation community was adjusted using Bon- ferroni corrections to prevent error accumulation. I used Manly’s standardized selection ratio to represent relative strength of selection for a given vegetation community. Vegetation communities with 0 availability in a given range were omitted from analysis. Locations of Elk during 1988-1991 were used to determine core foraging areas of dispersing groups on summer ranges. Using radio telemetry locations and visual observation, I established five 404-m? (20.1 m x 20.1 m) macroplots selected within core foraging areas determined in each dispersing group and in the core foraging area of the PP population in 1991. Ten macroplots already had been established in core foraging areas of LC and SR populations (five in each area) as part of another study (Van Dyke et al. 1994). I compared volumes of forbs and graminoids produced from each core area associated with dispers- ing groups to core areas of respective source popula- tions through 2-sample t-tests. Sampling methods have been described (Van Dyke et al. 1991, 1994). Shrubs were not included in the analysis because shrub cov- erage was low on most plots and summer diets of Elk in all populations averaged <10% shrubs (Van Dyke et al. 1994). Although species-specific forage selec- tion was evident (Van Dyke et al. 1994), all available graminoids and forbs were consumed. Thus, estimates of forb and graminoid volume were considered to rep- resent volume of edible browse. Results Seasonal dispersal and home range characteristics All dispersing groups moved greater absolute and relative distances from winter to spring or summer ranges than traditional populations, and, in two of three cases, traveled in entirely different directions and followed different elevational gradients (Table 1). Rela- tive to the size of winter home areas, traditional pop- ulations migrated distances 2.3-7.5x the radius of their winter home areas to reach spring and summer ranges, but dispersing groups moved 5.6-11.0x the radius of their winter home areas to reach new, non-traditional ranges in the same seasons. Elk in dispersing groups used annual or seasonal ranges differently than Elk in associated source pop- ulations (Table 2). Differences also existed in home range sizes of dispersing groups and source populations (Figure 2). The annual and seasonal home ranges of Elk in the TC and WC groups were larger than home ranges of Elk in associated source populations. In con- trast, seasonal home ranges of Elk in the HR group were similar to those estimated for individuals in the associated source population. Differences between source and dispersing group populations also were evi- dent in use of core areas (Figure 3). Hellroaring Elk made greater use of smaller areas (higher use/area ratios) in summer and autumn than did Elk in their source (SR) population. A similar trend was evident in the more concentrated use of winter and spring Vol. 121 core areas by the WC group. In contrast, the TC group demonstrated more dispersed range use than its source (PP) population, and had no identifiable core area in winter-spring (use/area ratio = 1). Patterns of habitat and elevation use in source populations: Comparisons of source populations and dispersing groups No population showed random proportional selec- tion of vegetation communities (P < 0.001, all cases), but selection patterns differed in different populations, and, in two of three cases, patterns of selection in source populations differed from selection patterns in asso- ciated dispersing groups. The SR source population and HR dispersing group were the only source popu- lation-dispersing group pair in which selection of vege- tation communities did not differ. Both used alpine areas less frequently than expected and used Lodge- pole Pine communities more than expected (Table 3). The Line Creek source population and its associated WC dispersing group differed in selection patterns in four of seven vegetation communities (Table 4), and the PP source population and its TC dispersing group in four of eight (Table 5), although patterns were dif- ferent in each case. Wolf Creek Elk used sagebrush communities most commonly in the spring, followed by an intensive shift to alpine communities in the sum- mer, resulting in nearly equal rates of use in these com- munities over the combined period. However, WC Elk did not use any one vegetation community differently than expected, although their overall pattern of use was different from availability. In contrast, LC Elk made more equitable use of all available vegetation types, although they were most commonly located in stands of Limberpine (33%). Line Creek Elk used grass-forb and Limberpine communities more than expected, and sagebrush and alpine communities less than expected. Actual proportional use in grassland- forb communities was identical in the two populations. Observed differences in significance in this case were attributable to differences in the number of observa- tions in the two groups. The majority of PP Elk locations were in grass-forb vegetation (56%), and this population used this vegeta- tion type and Limberpine at greater than expected fre- quencies. Picket Pin Elk used Lodgepole Pine less than expected. In contrast, TC Elk were most commonly located in stands of Douglas Fir (42%). They used this vegetation community more than expected, but used spruce-dominated vegetation less than expected. Core foraging areas of dispersing groups and source populations differed in volumes of forbs and gram- inoids produced (Table 6). Trout Creek core areas had higher volumes of graminoids and forbs compared to PP core areas, but differences in other comparisons displayed no consistent pattern. Two dispersing groups, TC and WC, showed dif- ferent patterns in seasonal elevational use compared to their source populations (Figure 4). Trout Creek 2007 VAN DYKE: COLONIZATION OF NON-TRADITIONAL RANGE BY ELK 137 TABLE 1. Scale considerations in distance, direction, and elevational differences in pairs of traditional source populations and dispersing groups of Elk in south-central Montana, U.S.A., 1988-1991. In each pair, the traditional source population is listed first. Migration distance (km) reflects distance from shared winter range activity center to traditional source popula- tion or dispersing group spring or summer range. Home range radii estimated as square root of home range area ( km*) divided by st. Migration Migration Elevation Population home Population distance direction change range radius — winter Line Creek 4.9 W Higher 2.12 Wolf Creek 11.8 NE Lower Picket Pin 11.6 W Higher 1.54 Trout Creek 17.0 NE Same Silver Run 11.8 SW Higher 1.70 Hellroaring 14.6 SW Higher TABLE 2. Results of Multiple Range Permutation Procedures analysis comparing range utilization between dispersing groups and associated source populations in south-central Montana 1988-1991. P < 0.05 interpreted as evidence of significantly different distributions between dispersing group and associated source population. n = number of Elk locations used in analysis (source population and dispersing group, respectively). Seasons selected represent periods of movement to new ranges by dispersing groups. Population Season n Standardized test statistic 12 Picket Pin-Trout Creek Annual 348/71 -102.89 <0.0001 Silver Run-Hellroaring Summer-Autumn 291/58 - 46.24 <0.0001 Line Creek-Wolf Creek Spring-Summer 263/73 - 10.99 <0.0001 2 Se ae ees ee ee Seasonal Home-range Size Core Area Concentration 300 54 @ Trout Creek O Picket Pin @ Trout Creek els O Picket Pin v Heliroaning v_ Hellroaring v Silver Run Vv Silver Run 44 @ Wolf Creek Lf @ Wolf Creek 0 Line Creek Y= 200 0 Line Creek e =? ==+ ‘6 e co = > s a o T ~- | 2 & 34 | < ® ro) on = = y | a T = We it 100 i> ml) 5 1 2 4 j I a Mg I] ! a & 5 | : a = | : A | 8 | Winter/Spring ‘Summer/Autumn Annual | Season 0 = Winter/Spring Summer/Autumn Annual FiGuRE 2. Seasonal and annual home area sizes of source populations and associated dispersing groups of Elk in south-central Montana, USA, 1988-1991. Bars indicate 95% CI. Elk used lower elevations in summer (f,,, = 3.62, P < 0.0001), consistent with their tendency to move to traditional, high elevation summer range later than PP Elk. Similarly, WC Elk used lower elevations in summer (t,,, = 2.03, P = 0.05) and autumn (,,, = 2.46, P = (0.02), reflecting their tendency to spend at least part of these periods off the Line Creek Plateau and away from the main LC population. Hellroaring Elk used elevations similar to SR Elk in all seasons, had non-overlapping ranges in summer and autumn, and Season FiGuRE 3. Core area concentration indices (% use of core area/% contribution of core area to home area) of source populations and associated dispersing groups of Elk in south-central Montana, USA, 1988-1991. Bars indicate 95% Cl. persisted on their new ranges throughout both seasons in all years. In contrast, TC and WC Elk always aban- doned the non-traditional ranges at some point during the summer. Over time, it became apparent that dispersing groups abandoned non-traditional ranges more quickly in drier years. I examined the strength of this relationship at the conclusion of the study through a logistic regres- sion analysis relating June and July rainfall (estimated 138 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 3. Proportional availability and use of vegetation communities on summer-autumn range by the Silver Run Elk popu- lation and an associated population (Hellroaring) colonizing non-traditional range in disjunct areas in south-central Mon- tana, USA, 1988-1991. P values (Bonferroni corrected) equal the probability that proportional use equals availability. Vegetation Proportion Proportion used Standardized selection ratio P community available Silver Run Hellroaring SilverRun Hellroaring Silver Run Hellroaring Alpine 0.70 0.48 0.48 0.08 0.07 <0.001 0.02 Lodgepole Pine 0.04 0.22 0.29 0.60 0.70 <0.001 <0.001 Sagebrush 0.08 0.14 0.12 0.21 0.16 0.27 1.00 Whitebark Pine 0.18 0.16 0.11 0.11 0.07 1.00 1.00 TABLE 4. Proportional availability and use of vegetation communities on spring-summer range by the Line Creek Elk popu- lation and an associated population (Wolf Creek) colonizing non-traditional range in disjunct areas in south-central Mon- tana, USA, 1988-1991. P values (Bonferroni corrected) equal the probability that proportional use equals availability. Vegetation Proportion Proportion used Standardized selection ratio P community available Line Creek Wolf Creek Line Creek Wolf Creek Line Creek Wolf Creek Alpine 0.33 0.17 0.39 0.05 0.16 <0.001 1.00 Douglas Fir 0.08 0.14 0.08 0.15 0.13 0.11 1.00 Grass-forb 0.02 0.06 0.06 0.31 0.44 0.001 0.17 Limberpine 0.10 0.33 0.07 0.30 0.09 <0.001 1.00 Sagebrush 0.30 0.18 0.40 0.05 0.18 <0.001 0.60 Spruce 0.08 0.09 0.00 0.04 0.00 1.00 0.17 Whitebark Pine 0.09 0.04 0.00 0.10 0.00 0.21 0.13 TABLE 5. Proportional availability and use of vegetation communities on annual range by the Picket Pin Elk population and an associated population (Trout Creek) colonizing non-traditional range in disjunct areas in south-central Montana, USA, 1988-1991. P values (Bonferroni corrected) equal the probability that proportional use equals availability. Vegetation Proportion Proportion used Standardized selection ratio JB community available Picket Pin Trout Creek Picket Pin Trout Creek Picket Pin Trout Creek Alpine 0.07 0.02 0.12 0.04 0.25 0.57 1.00 Douglas Fir 0.15 0.13 0.42 0.11 0.39 1.00 <0.001 Grass-forb 0.25 0.56 0.33 0.28 0.18 <0.001 1.00 Limberpine 0.04 0.10 0.01 0.35 0.05 0.01 1.00 Lodgepole Pine 0.26 0.05 0.09 0.02 0.05 <0.001 0.06 Sagebrush 0.05 0.02 0.03 0.06 0.08 1.00 1.00 Spruce 0.16 0.11 0.00 0.09 0.00 1.00 0.03 Whitebark Pine 0.02 0.01 0.00 0.05 0.00 1.00 1.00 TABLE 6. Volume (m*/ha) of forbs and graminoids in core foraging areas of dispersing population groups and associated source populations in south-central Montana 1988-1991. n (number of sampled macroplots) = 5 except for Line Creek (n = 4) and Silver Run (n = 3). Means of plant volume in areas used by dispersing groups and areas used by source populations compared by t-test, same categories. Volume (m?/ha) Area Forbs Graminoids m3/ha SE m?/ha SE Trout Creek SIS 62 1218 230 Picket Pin 228 32 307 2D Hellroaring 3184 11 326° 84 Silver Run 215 15 1002 376 Wolf Creek 698 18 B24 65 Line Creek 415 108 210 i * Different from source population. P < 0.0001. 2007 3000 Seasonal Elevation Use 2800 —@®- Trout Creek ff Oo Picket Pin / —¥— Hellroaring | ff o v 2600 9 SilverRun | . —f— Wolf Creek | / © LineCreek | yi Elevation (m) ~ s ~ Sy | ~ Ny aa Pe | / ° 2000 ~ 7 oY ae a Ol pad en OZ 1800 ie Winter Spring Summer Autumn Season FIGURE 4. Seasonal elevational use (m) of source populations and associated dispersing groups of Elk in south-cen- tral Montana, USA, 1988-1991. from the Red Lodge, Montana recording station for WC Elk and the Nye, Montana recording station for TC Elk, NOAA 1992) to the total estimated number of days in June, July, and August on new range. Vari- ation in combined June and July rainfall explained two-thirds (77 = 0.67, P < 0.0001) of the variation in days on non-traditional summer range in the TC and WC groups (Figure 5). Discussion Dispersal and density of populations Historical expansion and contraction of ranges of ungulate species has been influenced by many fac- tors, both natural and anthropogenic (Laliberte and Ripple 2004), but range expansions at local levels are often achieved by dispersal, which in turn may be mediated by density (Andersen et al. 2004). An eco- logical correlate of a population’s capacity to colonize new range is its ability to yield a large colonizing group (Safriel and Ritte 1983), something which can be best achieved during periods of population growth. In this area, annual aerial counts indicated that all source populations experienced population growth on tradi- tional range in the decade prior to this study. From 1979 to 1988, the PP population increased from 44 to 190 individuals (355%), the SR population from 72 to 139 individuals (93%), and the LC population from 85 to 127 individuals (49%) (Van Dyke et al. 1998), and all populations continued to increase through 1991. These increases are small relative to fluctuations in historically large populations of Elk like the nearby Northern Yellowstone Population which has ranged from 10000 to 20 000 individuals in recent years (Committee on Ungulate Management in Yellowstone National Park US National Research Council 2002). Nevertheless, such increases were proportionately large relative to past population size and occurred on very small annual ranges (84 — 166 km?) (Van Dyke VAN DYKE: COLONIZATION OF NON-TRADITIONAL RANGE BY ELK 139 60 ° 7 e . 50 P | | e 40 , 2 y = 36 44L n(x) - 17 707 f = 06724 30 / Sa 20 e 10 + r =" —— —r 2 a 6 ) Rainfall (cm) FiGuRE 5. Relation between June and July rainfall (combined) and estimated number of days on non-traditional sum- mer range in the Trout Creek and Wolf Creek dispers- ing groups in south-central Montana, USA, 1988-1991. et al. 1998) compared to those of the Northern Yel- lowstone Population, whose annual range encompass- es thousands of km? (Committee on Ungulate Man- agement in Yellowstone National Park U.S. National Research Council 2002). Thus, it is reasonable to believe that the numerical increases observed in these herds on these restricted ranges would increase den- sities and competition for space and other resources, contributing to the formation of dispersing groups, although the dispersive behavior seen in these small populations might be very different from that observed in larger populations. Size of home range and use and productivity of vegetation communities Variability in use and selection patterns of vegeta- tion communities in different populations is, in part, evidence of the behavioral plasticity of Elk. As these populations experienced relatively large proportional increases, the movement and fidelity of colonists to new ranges appeared similar to observed seasonal move- ments of White-tailed Deer (Odocoileus virginianus) in areas fragmented by agriculture (Nixon et al. 1991). However, whereas deer occupied traditional vegetation communities in the new areas to which they dispersed, Elk in these dispersing groups demonstrated their plas- ticity in habitat selection by changing their patterns of use of vegetation communities as they moved to new ranges. It is not only the proportion of a vegetation commu- nity in the landscape that determines its use by Elk, but also its juxtaposition and interspersion with other such communities (Porter and Church 1987), a dimen- sion of availability that was beyond the scope of this study to measure. Thus, use of vegetation communities by specific populations would be a poor guide for landscape scale habitat management. In these popu- lations Elk adjusted use of vegetation communities according to local and seasonal availabilities. 140 Use of marginal habitats is associated with increased rates of range expansion in some ungulate species (Andersen et al. 2004). The larger home ranges of TC and WC groups using non-traditional lower elevation ranges in summer might indicate the same trend. Fur- ther, larger home ranges in these groups also suggest- ed that forage on non-traditional ranges might have been more dispersed. However, comparisons of forage volume in core foraging areas of group and source pop- ulations did not support the premise that productivity was lower in ranges of dispersing groups. On the TC range, forage volume was higher in both graminoids and forbs. However, foraging areas for TC Elk were concentrated in riparian areas and probably could not support higher densities or more extended use by Elk in most years. Nutritional quality of vegetation is often better at higher elevations (Johnston et al. 1968; Boyce 1989), a trend confirmed on these ranges in measure- ments of protein and other nutrients in low vs. high- elevation plants (Van Dyke et al. 1991; Van Dyke et al. 1994). However, McCorquodale (1991) found that, in sagebrush habitats used by Elk in central Washing- ton similar to those used by the WC group, low inter- community variability in forage production, lack of a nonforage overstory, and the relative abundance of foraging areas all mitigated against lower primary production. The low-elevation, mostly treeless habitat of the WC group appeared to provide the greatest potential for heat stress of Elk during summer months because of its absence of thermal cover. However, Peek et al. (1982) have argued that thermal cover is not a requirement for Elk. Elk historically occurred on the Great Plains and in other areas that lacked thermal cover. Loss of Elk in these areas was due to human influence, not lack of adaptability to the habitat (Laliberte and Rip- ple 2004). In recent years Elk have recolonized areas in treeless, sagebrush-steppe habitats in central Wash- ington (Rickard et al. 1977; McCorquodale et al. 1986; McCorquodale 1991) similar to those used by the WC Elk. Both source populations and dispersing groups demonstrated fidelity to their selected home ranges from year to year. Such fidelity suggested that dis- persing groups had already established some degree of tradition in their use of new ranges and were not dispersing randomly. However, early abandonment of non-traditional range at low elevations under drier conditions suggested that such new ranges might have been sub-optimal environments for Elk compared to traditional range. Dispersing groups responded to peri- ods of environmental stress by rejoining source pop- ulations on traditional areas. By doing so, individuals in dispersing groups might have benefited from the source population’s collective knowledge of available resources on traditional range and its successful, long- term strategies of range use (Edge et al. 1985). THE CANADIAN FIELD-NATURALIST Vol. 121 Management Implications Elk in dispersing groups demonstrated an ability to locate and adapt to non-traditional range through var- ious combinations of altered habitat use, changes in patterns of seasonal elevational migration and adjust- ments in home range size and use. Given this reper- toire of adaptive strategies, managers should assume that areas adjacent to existing Elk populations but presently without Elk could be colonized, and could be of importance to future growth of such populations and expansion of their range. For example, in restora- tions of Elk in Kentucky (USA), translocated Elk demonstrated fidelity to a variety of habitat types, in- cluding some not previously encountered on their orig- inal ranges (Larkin et al. 2004). Such considerations are of importance to managers as efforts to restore Elk populations in several U. S. states (Didier and Porter 1999; McClafferty and Parkhurst 2001; Larkin et al. 2004) are being considered or have commenced. As noted previously, dispersing groups from these smaller populations might not exhibit the same pat- terns or behaviors associated with dispersal as has been observed in historically large herds. Thus, it would be advisable to consider and evaluate dispersive behav- ior associated with populations of different sizes before making broad generalizations about patterns of dis- persive behavior that could apply to populations of all sizes. However, even with this caveat in hand, it will be true that expanding ranges and growing populations of Elk, regardless of size, might create added opportu- nity to view and hunt Elk, but such expansion could contribute to increasing conflict between Elk and other land uses such as agriculture and must be examined in a regional context. Where colonization of new range by Elk enhances the persistence of marginal popula- tions and provides desired recreational opportunities for hunting and viewing Elk, managers should identify, provide access to, and, if possible, acquire adjacent range to which individuals might disperse, especially within the broader context of regional strategies that follow the principle of retaining large contiguous or connected areas that contain critical habitats (Dale et al. 2000). In these cases, dispersing groups should be protected from disturbance until they have established fidelity to new ranges (Larkin et al. 2004). Where dispersion to adjacent, but non-traditional ranges and habitats is likely to create or exacerbate land use con- flicts between Elk and humans, managers should con- sider lowering densities of Elk on traditional range to reduce the likelihood of such dispersal. Acknowledgments R. J. Mackie and E. H. Merrill offered comments on earlier versions of this manuscript. S. H. Yoo assisted in preparation of text, tables, and figures. C. E. Eustace, Montana Department of Fish, Wildlife and Parks, pro- vided logistical support for fieldwork. 2007 Literature Cited Ackerman, B. B., F. A. Leban, M. D. Samuel, and E. O. Garton. 1990, User’s manual for program HOME- RANGE. Second edition, Technical Report 15, Forestry, Wildlife, and Range Experiment Station, University of Idaho, Moscow, Idaho, USA. Andersen, R., I. Herfindel, B. E. Szther, J. D. C. Linnell, J. Oddén, and O. Liberg. 2004. When range expansion is faster in marginal habitats. Oikos 107: 210-214. Boyce, M. S. 1989. 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Proceedings of the first biennial scientific con- ference on the Greater Yellowstone Ecosystem. Edited by D. L. Despain. NPS/NRYELL/NTTR-93/XX. Van Dyke, F. G., B. L. Probert, and G. M. Van Beek. 1995. Moose home range fidelity and core area characteristics in south-central Montana. Alces 31: 93-104. Van Dyke, F. G. W. C. Klein, and S. T. Stewart. 1998. Long- The Rumsfeld Paradigm: Knowns and Unknowns in Characterizing Habitats Used by the Endangered Sharp-tailed Snake, Contia tenuis, in Southwestern British Columbia SHANNON F. WILKINSON!*, PATRICK T. GREGORY! >, CHRISTIAN ENGELSTOFT?, and KARI J. NELSON? 'Department of Biology, University of Victoria, P.O. Box 3020 STN CSC, Victoria, British Columbia V8W 3N5 Canada *Alula Biological Consulting, 1967 Nicholas Road, Saanichton, British Columbia V8M 1X8 Canada Ecosystems Branch, British Columbia Ministry of Environment, P.O. Box 9338 STN PROV GOVT, Victoria, British Columbia V8T 5J9 Canada 4Present Address: 824 Leslie Drive, Victoria, British Columbia V8X 2Y4 Canada Author for correspondence: viper @uvic.ca Wilkinson, Shannon F., Patrick T. Gregory, Christian Engelstoft, and Kari J. Nelson. 2007. The Rumsfeld paradigm: knowns and unknowns in characterizing habitats used by the endangered Sharp-tailed Snake, Contia tenuis, in southwestern British Columbia. Canadian Field-Naturalist 121(2): 142-149. The Sharp-tailed Snake, Contia tenuis, has a small and highly fragmented range in British Columbia, where it is considered | _ endangered. Known sites are few in number and generally small in spatial extent; numbers of snakes apparently are correspond- ingly low. Furthermore, most known sites for the species are on private lands in areas that are fairly heavily developed or being developed. Thus, the species is under serious threat of habitat alteration or loss. Although land stewardship has been a valuable conservation tool in this case, we also need to identify the key habitat requirements of Sharp-tailed Snakes to identify potential new sites, modify former or potential ones, or even create new ones. In this study, we compared sites known to harbour Sharp- tailed Snakes with those that seemed subjectively similar and therefore potentially suitable. We also compared these known and | potential sites with randomly chosen nearby locations. Variability of most measured features was high, both within and among site/location categories. Nonetheless, we found significant differences between known and potential sites and between those locations and random ones. Overall, locations known to be used by snakes had a more southerly aspect, more rock cover, shal- lower soil and litter, and less shrub cover than other sites. This study was constrained by the small number of known sites for Sharp-tailed Snakes in southwestern British Columbia, making our conclusions suggestive rather than definitive. Future work should incorporate additional variables. It also might be useful to undertake comparative habitat studies elsewhere in the range of the Sharp-tailed Snake where it is more common. Key Words: Sharp-tailed Snake, Contia tenuis, habitat, British Columbia. On 12 February 2002, U.S. Secretary of Defense Donald Rumsfeld made the following widely cited observation: “As we know, there are known knowns; there are things we know we know. We also know there are known unknowns; that is to say, we know there are some things we do not know. But there are also unknown unknowns, the ones we don’t know we don’t know. And if one looks throughout the history of our country and other free countries, it is the latter category that tend to be the difficult ones.” Although Rumsfeld was not discussing science, his circumlocutional comments nonetheless may well be apt in science (Shermer 2005). In conservation biology, the unknowns, whether known or unknown, usually exceed the known knowns and we often have to make educated guesses in, for example, trying to determine where there might be additional suitable habitats for a threatened or endangered species. To manage and conserve any species, it is crucial to understand its habitat requirements. The fitness of any species should be greatest in a habitat for which it is best suited and that it presumably prefers (Garshelis 2000). Determining critical habitats is an important component of the background knowledge that allows researchers and managers to develop strategies to either recover or maintain an endangered or threatened species and to choose areas in which to explore such strategies. The Sharp-tailed Snake (Contia tenius) is a small secretive snake, from 20 to 45 cm in total length when adult (Leonard and Ovaska 1998), that ranges from southern California to southern British Columbia (Cook 1960). Although the southern part of its range, to north- ern Oregon, is more-or-less continuous, the distribution of this species is highly fragmented in Washington and British Columbia (Leonard and Ovaska 1998). In ~ British Columbia, Sharp-tailed Snakes are known only | from a small number of scattered locations on southern Vancouver Island and some of the adjacent Gulf Is-- lands. A previous report from Chase in the interior of | British Columbia is of doubtful origin (Matsuda et al. 2006). Because of the low numbers of snakes found in) Canada and few known sites, the snake was consid- ered an endangered species by COSEWIC in 1999 (COSEWIC 2002*). Although formal study of Sharp- tailed Snakes has been limited, recent studies done in British Columbia and adjacent Washington cover as- pects of their ecology such as defensive behaviour | (Ovaska and Engelstoft 1999; Leonard and Stebbins 1999), aggregative behaviour (Leonard et al. 1996), 142 2007 movements (Engelstoft et al. 1999), habitat (Leonard and Leonard 1998), and cover use (Engelstoft and Ovaska 2000). Most of the areas where the Sharp-tailed Snake is found in British Columbia have fairly dense human populations and highly disturbed habitat. Consequent- ly, many of the sites at which snakes are found are on private land. Although the Sharp-tailed Snake seems somewhat tolerant of anthropogenic habitat distur- bance, habitat loss is still considered the primary threat to its persistence in Canada. Therefore, management and stewardship by landowners are important factors in conservation programs for this species. The aim of this study was to determine the key char- acteristics that define habitats used by the Sharp-tailed Snake, in order to guide future habitat management. We did this by comparing sites known to be used by Sharp-tailed Snakes with sites that were subjectively similar but not known to be used (potential sites), and with locations chosen at random with respect to both known and potential site types. The first comparison objectively tested whether potential sites actually are similar to those sites known to be used by these snakes and the second tested whether Sharp-tailed Snakes use a subset of those habitats that are available. Initial observations suggested that south-facing rocky slopes and forest clearings appeared to provide the most suitable habitat for this species (see also Leonard and Leonard 1998), but no quantitative assessment of habi- tat characteristics has previously been done. Thus, we measured a wide range of possibly influential habitat variables at each sample location and compared them statistically among locations. Although our comparisons among sites and loca- tions were objective, selection of potential sites obvi- ously was biased by our general knowledge of snake ecology and our previous experience with Sharp-tailed Snakes. This mainly reflected an aim of the recovery program for this species to find as many occupied sites as possible. Study Area In the past few years, federal lands in the Capital Regional District (including Gulf Islands National Park, Department of National Defence lands, and Coast Guard properties) have been assessed and rated for suitability as habitat for the Sharp-tailed Snake (Engel- stoft and Ovaska 1998*). These sites are in various locations on North and South Pender Islands, Salt- spring Island and southern Vancouver Island. In addi- tion, artificial cover objects (ACOs) have been installed at all sites at which Sharp-tailed Snakes are known to occur, including private lands, and at sites rated as hay- ing high potential as habitat for these snakes. ACOs provide easily sampled, attractive cover and, combined with naturally occurring cover objects, are an effective means of determining presence of snakes (Engelstoft and Ovaska 2000). Sharp-tailed Snakes are secretive WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKE 143 Potential 2004 Known and potential 2004 New since 2004 FiGurE |. Map of study area with sites known to be occupied by Sharp-tailed Snakes up to 2004 and since, and potential sites identified in 2004. Squares labeled “known and potential 2004” indicate known and potential sites that are too close together to represent separately on this map. For reference, latitude and lon- gitude for Victoria are 48°25'N and 123°19'W, respec- tively, and for Ganges 48°51'N and 123°31'W, respec- tively. Modified from a map obtained from the Natural Areas Atlas on the Capital Regional District (2008*) website. and rarely seen in the open, and ACOs provide the main means by which they are discovered in the field. Poten- tial sites in this study were identified by CE based on his extensive field experience with Sharp-tailed Snakes. The sites where ACOs have been placed can be classified into two general types. Unrestricted sites are those with relatively large extents of contiguous habi- tat under the same ownership surrounding the location of the ACO. These areas typically have experienced relatively little disturbance in the last century (e.g.. Department of National Defence lands or Canada Parks lands). Restricted sites are those on relatively 144 small, privately owned properties surrounded by private lands in relatively developed areas. These locations have generally experienced a greater degree of habitat modification (buildings, roads, gardens, etc.). Methods Fieldwork for this study was undertaken in summer and autumn of 2004. Our general sampling scheme was straightforward. The 31 sites were divided into two site types, those known to harbour Sharp-tailed Snakes (known knowns) and those that appeared to have the potential to harbour them (known unknowns). Hereafter, we call these known and potential sites, respectively (Figure 1). At each of these sites, in addi- tion to the known or potential assigned location, we also chose a corresponding random location (unknown unknowns) for sampling. Random locations were 50 m from an assigned location, but in a randomly chosen direction. Because 50 m was well within the known movement capabilities of Contia (Engelstoft et al. 1999), yet far enough away for the environment to vary significantly, this distance provided a test of non-ran- dom use of the local habitat. However, because most known sites are restricted, we were able to obtain a random location for only seven of the 16 known sites (compared to 14/15 for potential sites). Our sampling protocol was thus dictated to a large degree by access to private property, and by the amount of contiguous habitat under the same ownership surrounding the site. The total number of assigned and random locations at all sites combined was 52. At each of these locations, we took samples along three randomly chosen 10-m- long transects. The first transect at each assigned location was placed in a random direction from a randomly chosen ACO, so that the O0-m mark was at the centre of the ACO. The other two transects were placed at two dif- ferent randomly chosen locations (and in random directions) within a 25-m radius of the initial ACO. We measured canopy cover, aspect, and slope of loca- tion at the 0-m mark of the first transect; that is, these variables were measured once per location. All other variables were measured along each of the three tran- sects (i.e., three replicates for each location). This same protocol was repeated for random locations, with a point on the ground standing in for an ACO. We ensured that transects around an assigned location and its associated random location did not overlap or abut each other. We measured the following variables at each of the 52 locations: 1. Percentage canopy cover — Canopy cover was measured once for each location at the starting point of the first transect, directly overhead, and then at a 45° angle in each of the cardinal directions (south, north, east and west — surrounding canopy cover). The tree species making up the canopy cover also were recorded. . Aspect — The aspect of each location was measured in degrees using a compass at the starting point of the first i) THE CANADIAN FIELD-NATURALIST Vol. 121 transect. A measurement of 180° indicates a southerly aspect. 3. Slope — The general slope of the location was recorded in degrees as well as the slope for each separate transect. Both were measured using a compass equipped with a clinometer. 4. Substrate — At the l1-m mark along each transect, we determined the depth of both soil and litter layer by measuring the distance a thick wire penetrated into the ground. The composition of the litter (needles, leaves, etc.) also was recorded. 5. Rock coverage — Rock coverage was determined at each transect by measuring the total distance intercepted by rocks along the length of the transect. The rocks were rated by size as follows: loose gravel (pebbles <7.5 cm), cobbles (7.5-19.9 cm), rocks (20-50 cm), boulders (>50 cm), and bedrock. 6. Coarse woody debris — This was determined by measuring the total distance intercepted by coarse woody debris along the transect. Each piece of coarse woody debris was identified to species (if possible), and assigned a decay class (1-5) as per Table 5.13 in Meidinger (1998*). 7. Vegetation — Shrub cover and ground cover were determined by measuring the total distance intercepted by vegetation along the transect. For logistical reasons, coverage by each of lichens, bryophytes, grass and forbs was expressed as the total distance intercepted by these plants between the 2-m and 4-m marks of each transect. Statistical Analyses We used SAS 8.0 to analyze the data, except for aspect, which we also analyzed using ORIANA soft- ware. We compared locations and site types using both univariate and multivariate statistics. Although one level of the factor “location” was a randomly selected partner to a known or potential assigned location for Sharp-tailed Snakes, all factors (site, site type, location) were treated as fixed factors because all sites initially were chosen deliberately as particular site types and locations. In cases of non-orthogonal data, we used Type III sums of squares for F-tests. As this was essen- tially an exploratory study, we used a liberal criterion for tests of significance, highlighting all results for which P < 0.10. Most variables showed no clear pattern of hetero- geneity among categories of sites, so we combined them in various ways, which also reduced the number of variables to manageable levels, as follows: total shrub coverage (all shrubs — e.g., broom, Oregon grape, salal); tall, non-shrub coverage (ferns and asters); thick ground cover (e.g., ivy, periwinkle, trailing blackberry); light ground cover (lichens, bryophytes, grasses, forbs); rock cover (sum of all rock-size classes); coarse woody debris (sum of all CWD decay classes); and surround- ing canopy cover (average of directional values around sample location). All other variables (e.g. overhead canopy cover, soil depth, slope) were left unaltered. Because assigned and random locations were paired at unrestricted sites, we compared them via a 2-way factorial analysis of variance (ANOVA), with the three transects treated as replicates within each combination 2007 of site and location. We did this analysis separately for known and potential site types, as they were in differ- ent places. In each case, we had to drop restricted sites from the analysis because they lacked a random loca- tion. In all cases in which the interaction between fac- tors was non-significant, we dropped it from the analy- sis to test main factors only. In cases in which the interaction was significant, compromising interpreta- tion of main effects, we then did a comparison of loca- tions for each site separately. To compare known and potential site types, we used a nested ANOVA (sites nested within site type), ana- lyzing assigned and random locations separately, with transects serving as replicates within locations. F-tests were adjusted to those appropriate for nested ANOVA (Sokal and Rohlf 1981). Because aspect is an angular variable, with the low- est possible value (0°) identical to the highest possible (360°), we treated it differently from the other variables. We categorized aspect at each location as one of four quadrants representing the four cardinal directions and then compared frequency of directions among site type/location categories (known/assigned, known/ran- dom, potential/assigned, potential/random) by contin- gency-table analysis. We also used circular (angular) Statistics, as described by Zar (1984), to compare mean aspect between pairs of site type/location categories. In addition to aspect, a few other variables (e.g. can- opy cover) were recorded only once for each location, rather than separately for each transect. We therefore reduced the data set to average values for each loca- tion and then analyzed those new variables using two- way ANOVA (site x type x location, sites serving as replicates within each combination of factor levels). Again, we dropped the interaction term when it was non-significant, but did a separate comparison of locations at each site type when the interaction was significant. We also tested for correlations among these variables across all sites/locations. Studies of habitat relationships are necessarily mul- tivariate (Reinert 1993) and typically involve methods such as Principal Components Analysis (PCA) and Discriminant Function Analysis (DFA). In comparing categories of habitats, such as we have in this study, two approaches are possible (Quinn and Keough 2002). One is the multivariate analysis of variance (MANO- VA) approach embodied by Discriminant Function Analysis (DFA). A second is to do a Principal Com- ponents Analysis (PCA) of all groups combined and then compare the derived components among groups using univariate ANOVA. Although the two approach- es do not produce identical results, they generally result in broadly similar patterns of group differences (Quinn and Keough 2002). Advantages of the PCA approach include ease of analysis of multiple derived compo- nents and straightforward post hoc comparisons of particular groups (Quinn and Keough 2002), so we used it here. WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKI 145 To avoid pseudo-replication, especially for variables for which only one measurement was made per loca- tion (see above), we used the reduced data set (i.e., means of the three transects treated as data) for mul- tivariate analysis. Using the same variables as in the ANOVAs above, we subjected the data to PCA, ex- amined each of the first three principal components for influence of the original variables, and then treat- ed each of the principal components as variables in separate two-way ANOVAs (as above). We also used the first principal component (PC 1) as the independ- ent variable in a logistic regression comparing sites/ locations known to be used by Contia ys. all other sites. Results Two-way ANOVAs using individual transect data as values yielded few differences attributable to loca- tion, although sites differed significantly for several variables. In comparisons of assigned vs. random loca- tions for known site types, we found an effect only of rock cover, which was higher at assigned than at ran- dom locations (F, ,, = 3.42, P = 0.07). In cases of variables for which there was a significant interaction between site and location (extent of litter, slope of transect, coarse woody debris), differences between locations were significant at some sites, but not in any consistent direction. For potential site types, assigned and random loca- tions differed only in thick ground cover (F, ,.. = 38.8. P = 0.053), which was higher in assigned locations. Sites again differed significantly for several variables and the interaction between site and location was sig- nificant for four variables (extent of litter, rock cover, light ground cover, shrub cover). Patterns from inter- actions again generally were indistinct, but for the three sites that differed significantly in rock cover, the value was higher at assigned than random locations; for three sites differing significantly in shrub cover, the value was lower at assigned locations. Nested ANOVAs comparing assigned locations be- tween known and potential site types showed only that known site types had lower soil depth (F, ,. = 3.20. P = 0.08) and less coarse woody debris (F, ,. = 7.45. P = (0.01) than potential site types. Again, there were significant differences among sites for some variables. Similar comparison of random locations between known and potential site types revealed only that rock cover was higher at known site types (F, ,, = 3.15. P = 0.09), with sites again varying significantly for some variables. Two-way ANOVAs, using mean values (per 3 tran- sects) as data (and all site type x location combinations, n = 52), revealed more differences (Table 1). Overall (i.e., assigned and random locations considered togeth- er), known site types had shallower soil (F, ,, = 2.92. P = 0.09), shallower litter (F, ,, = 2.95, P = 0.09), 1.49 and higher rock cover (F, 4 = 4.67. P = 0.04) than 146 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Statistical summary of main habitat variables by site type (known vs. potential) and location (assigned vs. ran- dom). Only variables that were significantly different in at least one analysis (see text) are shown. Individual data are means for 3 transects at each site type/location combination; shown in the table are mean (top row) and + standard deviation, mini- mum value — maximum value (bottom row) of these transect means. Site type Known Potential Location Assigned Random Assigned Random (n= 16) (n=7) (n= 15) (n= 14) Rock Cover (m) Dell 2.1 1.6 0.9 2.45, 0 — 7.46 2.24, 0.02 — 6.03 1.78, 0.02 — 6.40 1.04, 0.01 — 3.11 Coarse Woody Debris (m) 0.1 0.6 0.4 0.5 0.18, 0 —0.71 1.37, 0 — 3.70 0.43, 0- 1.71 0.72, 0 — 2.07 Thick Ground Cover (m) 0.6 0 0.3 0.03 0.96, 0 — 3.50 0,0-0 0.56, 0 — 1.61 0.05, 0 — 0.14 Shrub Cover (m) 1.8 2.3 1.8 3.5 1.83.0-—5.29 1.42, 0.41 — 4.00 1.85, 0- 6.95 3.04, 0 — 9.95 Soil Depth (cm) 6.6 8.4 9.0 9.4 2.44, 3.5 -— 10.9 3.58, 3.6 — 13.2 4.85, 4.3 — 22.2 4.30, 2.7 — 17.0 Litter Depth (cm) Dal De) 2.7 Bs 1.26, 0- 4.3 1.46, 1.0—5.3 1.14, 0.7-4.2 2.23, 0.7 — 8.7 Overhead Canopy (%) 28.8 17.1 eS 28.2 21.17, 0 — 60.0 22.15, 0 — 50.0 14.25, 0 — 50.0 21.6, 0 — 50.0 Surrounding Canopy (%) 25.4 21.1 19.1 Spl 1239825) 52-5 16.51, 0 — 42.5 14.74, 0 - 46.3 18.66, 0 — 52.5 Transect Slope (°) 13.0 15.4 11.2 Holl 7.76, 0 — 28.33 15.39, 0 — 45.33 9.56, 0 — 27.67 7.24, 0.67 — 27.33 potential site types, on average. Assigned locations had F, ,. = 3.02, P = 0.1; known random vs. potential ran- more thick ground cover (F, ,, = 4.15, P = 0.047), dom —F, ,, = 3.55, P = 0.07, although this last result less shrub cover (F, ,, = 3.59, P = 0.06), less overhead canopy cover (potential site types only, F, ,, = 9.55, P =0.005) , less surrounding canopy cover (potential site types only, F,,, = 6.62, P = 0.02), and steeper transect slopes (potential site types only, F, ,, = 5.90, P =0.02) than random locations, on average. As expected, many of the variables above were sig- nificantly correlated over all site types/locations (n = 52 in all cases). These included mean soil depth vs. mean litter depth (r = 0.40, P = 0.004), mean rock cover vs. mean litter depth (r = -0.29, P = 0.04), mean soil depth vs. mean rock cover (r = -0.38, P = 0.005) and overhead canopy cover vs. surrounding canopy cover (r = 0.58, P < 0.0001). Direction (aspect) differed significantly among site type/location categories (Fisher’s exact test, P = 0.004). Thirteen known/assigned locations (81%) faced south and the rest (3) all faced west, whereas only six (40%) of potential/assigned locations faced south and another six faced east. Potential/random locations were partic- ularly deviant from known/assigned, with eight (57%) facing either north or east. These results were consis- tent with those from analysis via circular statistics (mean vectors: known assigned — 203.99°; known ran- dom — 215.15°; potential assigned — 155.73°; potential random — 125.86°). Significant differences between site type/location categories were as follows: known assigned vs. potential assigned — F, ,. = 4.27, P = 0.05; known assigned vs. potential random — F, ,, = 7.21, P = 0.01; known random vs. potential assigned — may not be reliable because of a uniform distribution of points. We ran PCA using various combinations of vari- ables, dropping those that had low weights on PC 1. Results were broadly similar in all cases, but they were most clearly defined when we used mean rock cover, mean soil depth, mean surrounding canopy cover, and mean shrub cover as variables in the analysis. In that case, PC 1 accounted for 44% of the total variance and PC 2 and PC 3 23% and 20%, respectively. This high cumulative percentage variance for the first three prin- cipal components suggests that they summarized the data efficiently (McGarigal et al. 2000). PC 1 was pos- itively weighted by soil depth, surrounding canopy cover, and shrub cover, but negatively weighted by rock cover (Table 2). Two-way ANOVAs on the PCs showed that PC 1 differed overall between both site types (F, 4. = 4.19, P = 0.046) and locations (F, ,, = 4.84, P = 0.03). Known site types had lower average values of PC 1 than potential site types and assigned locations had lower values than random, consistent with the two-way ANOVAs performed above on the individual variables and with the signs of coefficients in PC 1. Random locations had higher values of PC 2 than assigned loca- tions, but only for potential site types (F, ,, = 3.86, P = 0.06). Neither site types nor locations differed with respect to PC 3. Despite statistically interesting differences in PC 1 between site types and locations, differences overall 2007 WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKE 147 TABLE 2. Summary of weights (coefficients) of four variables on first three principal components in analysis of habitat vari ables potentially influencing occurrence of Sharp-tailed Snakes. PC | accounts for 44% of variance and PC 2 and PC 3 23% and 20%, respectively. Variable PC | Coefficients Soil Depth 0.497 Surrounding Canopy Cover 0.440 Shrub Cover ().489 Rock Cover -0.566 Site — type Known Potential tS 5 2 S 1 £ Suio Penal: 3° = -2 —3 A R A R Location FIGURE 2. Box-and-whisker plots of values of principal component | by site type (known vs. potential) and location (actual, A, vs. random, R). Horizontal line in each box is median. Upper and lower boundaries of box (hinges) indicate quartiles (25" and 75" per- centiles). Whiskers from box reach to the most ex- treme value that does not exceed 1.5 times the dis- tance between the quartiles; n = 52. among the four site type/location combinations were not large (Figure 2). In fact, differences among these categories were graded rather than discrete. We there- fore divided site type/location combinations into two categories, those known to be occupied by Sharp-tailed Snakes and those not. The resulting binomial variable was then regressed (logistic regression) against PC 1. The logistic model was a satisfactory fit to the data (Homer and Lemeshow test, x *, = 4.46, P = 0.81) and the relationship was significantly negative (Wald’s xX z = 4.25, P = 0.04; Figure 3). We also compared PC | of known/assigned locations successively, via logis- tic regression, to known/random, potential/ assigned and potential/random locations. The trend also was negative in each case, but significant (and strongly so) only in the last one. Thus, sites known to be occu- pied by Sharp-tailed Snakes differed from those not known to be occupied. Discussion The study of how and why animals occur in partic- ular habitats has a long history in ecology and has been the subject of numerous reviews and syntheses (e.g. PC 2 Coefficients PC 3 Coefficients -0.59] 0.346 0.549 0.670 0.490 -(). 562 0.331 0.339 P (Presence of Contia) ee aT =) 0 3 Principal Component 1 FIGURE 3. Logistic regression of presence(1)/absence(0) of Sharp-tailed Snakes vs. principal component | for all site type/location combinations. Absence implies “not recorded” rather than confirmed absence. Verti- cal lines are data points, solid line is predicted regression, and dashed lines are upper and lower 95% confidence limits on regression; n = 52. Morse 1980; Gray and Craig 1991; Reinert 1993; Garshelis 2000). Habitat use by a species is a function not only of habitat selection based on resource require- ments and physical factors such as temperature, but of other factors such as population density and the pres- ence of competitors and predators (Huston 2002: Mor- ris 2003). Thus, the habitats used by animals may not be those that are optimal for them in terms of fitness. In fact, it is even possible that some occupied habi- tats have negative implications for fitness (Van Horne 1983), for example if those habitats are sinks in a source-sink system (Pulliam 1988). Conversely, opti- mal habitats that could be used might not be occu- pied simply because the species’ ability to disperse there is limited (Huston 2002). In short, the analysis and interpretation of habitat-use studies is fraught with difficulties (Garshelis 2000). In that light, this study 148 is but a small first step towards understanding the habitat relationships of the Sharp-tailed Snake. It is further limited by its reliance on presence/absence (really presence/not recorded) data rather than data on abundance, or better, demographic performance (Pulliam 1988; Garshelis 2000). In terms of Garshelis’ (2000) classification of methods, our approach is a variant of the site-attribute design, but with the added inclusion of predetermined potential sites for the species’ occurrence. Animals typically do not use their environments at random, but non-randomly, selecting some sites over others because of particular habitat and micro-habitat features. Thus, the mere demonstration that places used by a particular species are different from those not used is unremarkable by itself. For the conservationist or manager, however, determining which habitat charac- teristics are important is fundamental to preservation, restoration, or even creation of habitats. What are the key features that describe suitable habi- tats for the Sharp-tailed Snake? The data collected in this study were highly variable and can only hint at differences between site types and locations that appear to be important. More detailed, longer-term work with larger samples would allow us to reach stronger con- clusions, but this study was limited by the small num- ber of sites at which the Sharp-tailed Snake is known to occur in southwestern British Columbia. It also was limited by the subjectivity with which potential sites were identified. For example, south-facing slopes are often favoured by snakes and this is reflected in the distribution of known and potential sites in this study. However, a Sharp-tailed Snake recently was found on a north-facing slope on Pender Island (D. Spalding, personal communication), suggesting that the species may occur in a wider range of conditions than previously thought. Currently, a more randomized approach to sampling sites for Sharp-tailed Snakes is under way to reduce this kind of bias, but we still have some way to go before we can convincingly move known and unknown unknowns to the fully known side of the ledger. Aside from the necessary constraints of small num- bers of known sites for Sharp-tailed Snakes, this study also might have been limited by failure to measure key environmental variables. For example, as is the case for other ectotherms, the ecology of Sharp-tailed Snakes is no doubt strongly influenced by temperature, which will affect the kinds of habitats and microhabi- tats they use at different times (Huey 1991; Reinert 1993; Blouin-Demers and Weatherhead 2001). Simi- larly, given the potential for desiccation in this small species, as well as a requirement for incubation sites for eggs, moisture also is likely to be an important physical factor. Therefore, cracks and openings for underground access and suitable sites for egg-laying or hibernation are potentially critical habitat features for these snakes. However, measuring subterranean access would be very difficult and we know nothing THE CANADIAN FIELD-NATURALIST Vol. 121 about the kinds of sites favoured by Sharp-tailed Snakes for egg deposition or for spending the winter. To deal adequately with temperature and moisture, we would need to measure them across all seasons and in multiple microhabitats per location simultane- ously. Despite these deficiencies, we found fairly clear in- dications that sites used by Sharp-tailed Snakes tend to be south-facing and to have relatively shallow soil and litter, relatively high rock cover, and relatively low shrub cover. Differences from randomly chosen sites emphasize this species’ non-random use of habitats, but differences between locations actually used by snakes and those deemed to be potential habitat underline the subjectivity of visual habitat assessment. That said, this subjective approach based on field experience has proven effective. Although none of the potential sites used in this study have since yielded evidence of Sharp-tailed Snakes, two of the known sites were initially identified by CE as potential sites and anoth- er site designated as potential by CE after this study was completed has since turned out to be occupied by Sharp-tailed Snakes. Even so, quantitative measures for objective assessment of potential sites also would be valuable. Future work could focus on variables found to be important in this study and on others that we were unable to measure, such as those mentioned above as well as shrub height, tree density, soil com- position, drainage, and type of bedrock. Given the rarity of the Sharp-tailed Snake in British Columbia and the obvious difficulties in studying rare species in general, perhaps an alternative (or sup- plementary) approach would be to undertake compar- ative studies of this species in more southerly parts of its range where it is more widespread and more abun- dant. Extrapolating from a species’ ecology in one part of its range to another distant one requires caution as there may be significant adaptive differences in differ- ent areas. For example, Sharp-tailed Snakes in Cali- fornia occupy a wide diversity of habitats (D. Wake, personal communication), most of which do not occur in British Columbia. Nonetheless, this approach might reveal fundamental species’ characteristics that can be used to delineate more clearly the critical habitats required for conservation of the Sharp-tailed Snake in British Columbia. Acknowledgments We thank Laura Matthias and Michelle Masselink for showing us sites known to be occupied by Sharp- tailed Snakes and Andrea Pilski for assisting in the field. The Sharp-tailed Recovery Team offered useful help and advice and both David Wake and David Spalding provided us with important personal obser- vations. We also thank various landowners for allow- ing access to their land: Pender Island — Don and Theresa Williams, Sylvia O’Reilly, Robert Dill, Jon Crawford, Pat and Ian Haugh; Saltspring Island — Frances Hill and Dave Kerman, Sandra and Vince 2007 Knight, Leslie Northey, Mark and Brenda Haughley. The British Columbia government provided staff time. The views presented are those of the authors, not their respective agencies. The Endangered Species Recovery Fund (World Wildlife Fund, Canada) and the Habitat Conservation Trust Fund (BC) provided financial support. Documents Cited (marked * in text) Capital Regional District. 2008. Natural areas atlas. http:// www.crd.be.ca/maps/natural/atlas.htm (Accessed May, 2008). COSEWIC. 2002. Canadian Species at Risk. Committee on the Status of Endangered Wildlife in Canada. http:// www.cosewic.gc.ca/eng/sct5/index_e.cfm (Accessed Jan- uary, 2005). Engelstoft, C., and K. Ovaska. 1998. Sharp-tailed snake study on the Gulf Islands and southern Vancouver Island, March — November 1997. Final report. Ministry of Envi- ronment, Lands and Parks, Nanaimo, British Columbia. Meidinger, D. 1998. Field Manual for Describing Terrestri- al Ecosystems. B.C. Ministry of Environment, Lands and Parks and B.C. Ministry of Forests. http://www.for.gov. be.ca/hfd/pubs/Docs/Lmh/Lmh25.htm (Accessed August 2004). Literature Cited Blouin-Demers, G., and P. J. Weatherhead. 2001. Thermal ecology of black rat snakes (Elaphe obsoleta) in a ther- mally challenging environment. Ecology 82: 3025-3043. Cook, S. F. 1960. On the occurrence and life history of Con- tia tenius. Herpetologica 16: 163-173. Engelstoft, C., and K. Ovaska. 2000. Artificial cover-objects as a method for sampling snakes (Contia tenius and Thamnophis spp.) in British Columbia. Northwestern Naturalist 81: 35-43. Engelstoft, C., K. Ovaska, and N. Honkanen. 1999. The harmonic direction finder: a new method for tracking move- ments of small snakes. Herpetological Review 30: 84-87. Garshelis, D. L. 2000. Delusions in habitat evaluation: meas- uring use, selection, and importance. Pages 111-164 in Research techniques in animal ecology. Edited by L. Boi- tani and T. K. Fuller. Columbia University Press, New York. 442 pages. Gray, R. D., and J. L. Craig. 1991. Theory really matters: hidden assumptions in the concept of “habitat require- ments’. Acta XX Congressus Internationalis Ornithologici: 2553-2560. Huey, R. B. 1991. Physiological consequences of habitat selection. American Naturalist, Supplement 137: S91- SIS: Huston, M. A. 2002. Introductory essay: critical issues for improving predictions. Pages 7-21 in Predicting species WILKINSON, GREGORY, ENGELSTOFT, and NELSON: SHARP-TAILED SNAKE 149 occurrences — issues of accuracy and scale. Edited by J.M Scott, P. J. Heglund, M. L. Morrison, J. B. Haufler, M. G Raphael, W. A. Wall and F. B. Samson. Island Press, Washington.. Leonard, W. P., D. Darda, and K. R. McAllister. 1996 Aggregations of sharptail snakes (Contia tenuis) on the east slope of the Cascade Range in Washington state Northwestern Naturalist 77: 47-49. Leonard, W. P., and M. A. Leonard. 1998. Occurrence of the sharptail snake (Contia tenuis) at Trout Lake, Klickitat County, Washington. Northwestern Naturalist 79: 75-76. Leonard, W. P., and K. Ovaska. 1998. Contia Girard. Cat- alogue of American Amphibians and Reptiles 677.1- 677.7. Leonard, W. P., and R. C. Stebbins. 1999. Observations of antipredator tactics of the sharp-tailed snake (Contia tenuis). Northwestern Naturalist 80: 74-77. Matsuda, B. M, D. G. Green, and P. T. Gregory. 2006. Amphibians and reptiles of British Columbia. Handbook, Royal British Columbia Museum, Victoria, British Colum- bia. 266 pages. McGarigal, K., S. Cushman, and S. Stafford. 2000. Mult- variate statistics for wildlife and ecology research. Springer, New York. 283 pages. Morris, D. 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Density as a misleading indicator of habitat quality. Journal of Wildlife Management 47: 893- 901. Zar, J. H. 1984. Biostatistical Analysis, 2™ ed. Prentice-Hall. Englewood Cliffs, NJ. 718 pages. Received 11 April 2006 Accepted 20 May 2008 Do Repugnant Scents Increase Survival of Ground Nests? A Test with Artificial and Natural Duck Nests VANESSA B. HARRIMAN! 4, JusTIN A. Pitt?, and SERGE LARIVIERE? 'Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan S7N SE2 Canada "Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada 3Cree Hunters and Trappers Income Security Board, Quebec, Quebec G1 V 4K5 Canada 4Corresponding author: vanessa.harriman@ec.gc.ca Harriman, Vanessa B., Justin A. Pitt, and Serge Lariviere. 2007. Do repugnant scents increase survival of ground nests? A test with artificial and natural duck nests. Canadian Field-Naturalist 121(2): 150-154. Ground-nesting birds typically experience high predation rates on their nests, often by mammalian predators. As such, researchers and wildlife managers have employed numerous techniques to mitigate nest predation. We investigated the use of scents as repel- lents to deter predators from both artificial and natural ground nests. Survival rates of artificial nests did not differ among six groups of substances (Wald y?,,_ ;= 4.53, P < 0.48); however the chronology of predation among groups differed. A commercial Coyote urine based deterrent (DEER-D-TER™), human hair, and Worcestershire sauce were depredated faster than the control (F, ; = 40.3, P < 0.001). Nest survival of natural nests differed among those groups tested (Wald ren — SSP 01005); the eight mothball treatment decreased survival (Wald x? ,,_ , = 11.5, P < 0.005), which indicated that novel smells may attract predators or result in duck nest abandonment when coupled with natural duck scent. Chronologies of predation events among treatment groups were not different for natural nests (F,, = 1.9, P = 0.22). These findings indicate an interaction between novel scents and predator olfactory cues. Key Words: Coyote, Canis latrans, urine, human hair, mothballs, napthaldehyde, nest predation, olfactory cues, waterfowl, Manitoba. In the prairies of North America, predation on nests of ground-nesting ducks is considered the most impor- tant factor limiting waterfowl populations (Chafloun et al. 2002). As such, wildlife managers have devel- oped and employed various techniques in an attempt to mitigate nest predation with varying degrees of success. Typically, a predation event by a mammalian predator such as the Striped Skunk (Mephitis mephitis), Red Fox (Vulpes vulpes), or Raccoon (Procyon lotor) is oppor- tunistic (Lariviere and Messier 1997). These predators are nocturnal and locate nests using audition (when the hen flushes) and olfaction (Lariviére and Messier 1997). Although substances that may attract predators to nests have been investigated (Whelan et al. 1994; Clark and Wobeser 1997), scents that may repel pred- ators from nests have yet to be tested. Chemical compounds have been used successfully to repel animals, including mammals. Some animals have been effectively repelled by means of conditioned taste aversion (e.g., Hoover and Conover 1998) and through the use of noxious scents such as toxic com- pounds or odor from predatory animals (Sullivan et al. 1985; Rosell 2001). We investigated the effectiveness of various scents to increase survival of duck nests in the Canadian prairies. Specifically, we tested whether applying DEER-D- TER™ (a commercial product containing Coyote (Canis latrans) urine), human hair, Worcestershire sauce, and mothball treatments could increase survival of eggs in artificial nests and the capability of DEER- D-TER and mothball treatments to increase the sur- vival of eggs in natural duck nests. We also tested for differences in speed of predation events to determine whether specific scents may temporarily deter preda- tors from a nest. Finally, we compared survival of eggs in artificial and natural nests. Study Area This study was conducted southeast of Minnedosa, Manitoba (50°14'54"N, 99°50'30"W), in an area sup- porting numerous small (<3 ha) seasonal to permanent wetlands (ca. 14% of study site) and characterized by intensive cultivation (63%) of small grains (wheat, bar- ley, and oats) and oil crops (canola and flax). Inter- mixed among agricultural fields are small aspen bluffs composed of poplar species (largely Quaking Aspen (Populus tremuloides) and Bur Oak (Quercus macro- carpa). The predator community is diverse and includes Raccoons, Striped Skunks, Coyotes (Canis latrans), Red Foxes, American Badgers (Taxidea taxus), and Ameri- can Mink (Neovison vison). Predation of bird nests in the area is largely attributed to mammals (Pasitschni- ak-Arts and Messier 1995) but avian predation also occurs, primarily by birds such as the Common Raven (Corvus corax), American Crow (Corvus brachyrhyn- chos), and Black-billed Magpie (Pica pica). Methods The effectiveness of five groups of substances was compared to a control using artificial nests. The repel- lent substances were chosen based on unsubstantiated reports of their efficacy against mammals (human hair 150 2007 HARRIMAN, PITT, AND LARIVIERE: DO REPUGNANT SCENTS REPEL PREDATORS? 15] 10 ie re 08 S 06 $ 06 g § 3 a - D 8 0.4 ZS 04 0.2 02 0.0 ue r T T T 7 0.0 cots? ow 5554 ee ow? worn wor Control Coyote urine 8 Mothballs quo" Treatment Treatment FiGurE |. Daily nest survival (all eggs remaining in nest) of artificial duck nests treated with scents in Manitoba, 2002. Control 1.2 Koyrcae Coyote urine ———*-—— Human hair — —-y—-- Worcestershire sauce 1.0 — -#— 4 Mothbalis 8 Mothballs 08 2 © 06 se) oa’ ee << S "OR asl © ~ Oo---0 no ~S 2 0.4 ~- Vy o Zz 0.2 0.0 a0 SS ree fe 0 5 10 15 20 25 30 Exposure days FIGURE 2. Percent of surviving artificial duck nests (all eggs remaining in nest) treated with scents at X exposure day in Manitoba, 2002. and Worcestershire sauce) or from their commercial claims (DEER-D-TER™ and mothballs). Nests in the control group were each treated with 0.1 L of marsh water, which was poured along the outside edge of the nest. A packet of DEER-D-TER™ was placed on the outside edge of each nest of the second group. DEER- D-TER™ consisted of a green plastic packet contain- ing a mixture of sand and Coyote urine and was adver- tised to repel deer (Odocoileus spp.) and skunks. According to the manufacturer, this product was effec- tive in the field for ca. 90 days. Group three received one cup of human hair which was collected from three different salons and was combined to create a homoge- neous mixture. Group four received 0.15 L of Worces- tershire sauce applied evenly to each egg and was re- applied on every subsequent visit. The fifth group FIGuRE 3. Daily nest survival (all eggs remaining in nest) of natural duck nests treated with scents in Manitoba, 2002. 125 —®— _ Control ° Coyote urine | 104 lms 8 Mothbalis | 084 2 g 064 oS oO 2 044 o z 024 re) 02000 [5 es 0.0 1 ee hea L er ee. = 0 5 10 15 20 25 30 Exposure days FIGURE 4. Percent of surviving natural duck nests (all eggs remaining in nest) treated with scents at X exposure day in Manitoba, 2002. received four mothballs double wrapped in cheesecloth to prevent accidental ingestion by ducks or predators. The sixth group received eight mothballs, to determine if a greater amount yielded a different or stronger res- ponse, with two cheesecloth-wrapped groups of four mothballs placed on opposite sides of the outside edge of the nest. Three groups of substances were applied to natural nests: marsh water, DEER-D-TER™, and eight mothballs. Methodology and application of treatments for natural nests followed the same protocol as those of artificial nests. Data were collected from 12 June to 7 July 2002. Artificial nests were created by depositing six medium- sized chicken eggs in a depression which was then cov- ered with grass. Nests were marked by a thin welding rod at a distance of | m. To avoid biases associated with 2 habitat, all artificial nests were deployed in rights-of- way. Each nest was randomly placed within a 0.8 km stretch along four different, but intersecting, second- ary unpaved grid roads. Artificial nests were located 300-1000 m apart on alternating sides of the road. Nests were placed in the densest vegetation as close as possi- ble to the randomly selected location to reduce possi- ble effects of concealment on survival. Artificial nests were visited every five days for 25 days and were ap- proached from a different angle each visit to avoid the creation of a pathway to the nest. Rubber boots and latex gloves were worn at all times in hopes of con- cealing human scent. Each nest was randomly assigned one test substance until an equal number of nests per treatment was obtained. Natural nests were located via the drag chain method. Only upland areas of dense nesting cover were searched and nests were marked with stakes placed 10 m away in arandom direction. Nests were located on four sep- arate quarter sections within 1.6-8.0 kilometers of each other. Each quarter section consisted of dense nesting cover and was bordered by either a crop or fallow land and a secondary gravel road. Treatments of natural nests were assigned systemat- ically because we were unaware of how many nests we would eventually locate and we sought to have a sim- ilar number of nests per treatment. If, on return to nests treated with mothballs, any of the mothballs had fallen out of the cheesecloth, the entire packet was replaced with a new one. Hens often incorporated the wrapped mothballs into their clutch and occasionally moved the packet(s) directly (< 0.5 m) outside of the nest bowl. In these cases, the wrapped mothballs were left where the hen placed them. The first re-visit to the nest oc- curred between five and eight days after initial loca- tion and treatment application. Subsequent visits were made every seven days for up to a total of five visits depending on the fate of the nest. Natural nests were excluded from the analyses if they were abandoned but not depredated. However, we were unable to determine whether a nest had been abandoned and subsequently depredated. All nests were deemed depredated (failed) after at least one of the eggs was removed from or destroyed in the nest. Mayfield logistic regression (Hazler 2004) was used to determine whether the nest survival of each treatment group differed from the nest survival of con- trol nests within each nest type (artificial or natural). Because nests were not visited daily the exact date of failure was unknown. The number of days a nest was active (“exposure days”) was calculated differently depending on the fate of the nest. Exposure days of nests that hatched (or survived to the last day of the study in the case of artificial nests) was the number of days that nest was under observation. Exposure days of failed nests was the number of days the nest was known to be active plus the midpoint between the last THE CANADIAN FIELD-NATURALIST Vol. 121 check and the last known active day. Nests that fail early are less likely to be detected and the Mayfield logistic regression reduces this bias by incorporating exposure days. Also, because this is a logistic-regres- sion analysis, covariates can be incorporated, allowing for easy analysis of experimental data (Hazler 2004). All artificial nests were deployed on the same day and nest dragging was completed within three days which minimized the potential impact of seasonality on the data. An analysis of variance with repeated measures was used with respect to nest survival as a function of exposure days to determine whether there was a differ- ence in the chronology of predation events between treatments as time progressed. All post-hoc tests were conducted using an all-pair-wise comparison Tukey test. Survival of artificial and natural nests was com- pared using Mayfield logistic regression. Results Survival of eggs in artificial nests was higher than that of natural nests (Wald y? ,,_ , = 43.4, P < 0.001). Artificial nest daily survival was 0.54 (SE + 0.003) for all substances combined, while daily nest survival for natural nests was 0.04 (SE + 0.01) for all substances combined. We deployed 120 artificial nests, with each treatment group containing 20 nests. Due to flooding, seven arti- ficial nests were lost at various phases of the experi- ment and were excluded from further analyses. We found no significant difference in artificial nest survival rates among treatment groups (Wald 7 Aes =e P = 0.48; Figure 1). For all treatments combined, there was a positive correlation between survival of artificial nests and exposure days, with survivorship of nests increasing with time (F, ,,= 6.77, P < 0.001) and the first five exposure days being the most fatal (Figure 2). Addi- tionally, the chronology of predation differed among groups (F, , = 40.3, P < 0.001). Artificial nests con- taining both four and eight mothball treatments were depredated similarly to control nests (q = 1.63, p = 6, P > 0.05;.q = 1.40, p = 6, P > 0.05). However, nests treated with DEER-D-TER™ (q = 9.14, p= 6, P < 0.05), human hair (q = 4.84, p = 6, P < 0.05), and Worcester- shire sauce (q = 13.9, p= 6, P < 0.05) were depredated more quickly than control nests. In total, 70 natural duck nests were located, with 24 receiving the control, 23 treated with DEER-D-TER™, and 23 treated with eight mothballs. Natural nest sur- vival of the treatment groups differed from that of the control (Wald x? ,,_5= 11.82, P < 0.005; Figure 3) with survivorship of nests treated with eight moth- balls being significantly lower (Wald x? ,,_ , = 11.47, P < 0.005). Nest survival did not vary as a function of exposure days for all natural duck nests combined (F, , = 2.14, P =().17) despite a notably rapid decrease in nest sur- 2007 vival during the first five exposure days (Figure 4). Nest survival did not vary as a function of exposure days among treatments for natural duck nests (F, , = 1.99, P= 22): Discussion We assessed the effect of various scent treatments on the survivorship of artificial and natural duck nests in the prairie pothole region of Canada. The application of scents did not increase nest survival of artificial or natural nests. Addition of mothballs decreased survival of natural nests. These results not only indicate that DEER-D-TER™, human hair, Worcestershire sauce, and mothballs were ineffective at repelling predators, but that certain scents may actually attract predators and/or increase duck nest abandonment rates. The chronology of predation differed among treat- ments for artificial nests but not for natural nests. Con- trary to expectations, artificial nests treated with DEER- D-TER™, human hair, and Worcestershire sauce were depredated more quickly than control nests. Again these results may indicate a neophilic predator response. In particular, human-associated scents (i.e., hair and food items) may elicit predator attraction depending on the study site and the animal’s experience with people. Al- though this study took place in a rural location, it is plausible that local predators may have positive asso- ciations with human dwellings and food, especially opportunistic species such as Striped Skunks and Rac- coons (Pitt et al. 2008). Raccoons in particular often rely on anthropogenic foods at the northern edge of their range distribution (Lariviére 2004; Pitt et al. 2008). Additionally, DEER-D-TER™ may have also attracted predators to nests, especially con-specifics. Mothballs and hair regularly went undisturbed at de- predated nest sites whereas urine packets were often chewed on and/or ripped open. The result of predators exhibiting neophilic tenden- cies may have important implications for waterfowl due to the propensity for which they are studied. The concealment of human scent should be an important consideration for researchers. Neophilic predators may increase their search radius when a particular scent cue is present (Whelan et al. 1994), possibly rendering human-visited nests more susceptible to predation. Our study indicates that the survival rates of artificial and natural duck nests in the prairie pothole region are different and this may indicate that these nests are viewed differently from the perspective of a predator. Artificial nests realized higher survival rates compared to natural nests, which has been previously document- ed (Guyn and Clark 1997). The disproportionate dif- ference in survival between artificial and natural nests could also in part be attributed to the presence of scents associated with natural nests, as well as the presence of a hen and the ability to cue in on the nest after a flush event (Lariviere and Messier 1997). Alternatively, the HARRIMAN, PITT, AND LARIVIERE: DO REPUGNANT SCENTS REPEL PREDATORS? 153 observed difference may have been mediated by uncon- trolled factors affecting natural nest survival, such as nest abandonment, hen species and behavior, nest con- cealment, and variation in nest spacing. Specifically, the eight mothball treatment may have provoked aban- donment by nesting female ducks as some ejected mothballs from nests. Conditioned taste aversion has been effective in deterring predators from nests (Nicolaus 1986; Con- over 1990). Napthaldehyde (the noxious substance in mothballs) has been found to be effective in causing adverse responses by Coyotes when ingested (Hoover and Conover 1998). After repeated exposure, Coyotes avoided eggs injected with napthaldehyde. Although highly successful, it is expensive and time consum- ing to set up nests with injected chemicals. For future studies, it may be equally effective and less time con- suming to combine the use of conditioned taste aver- sion with scents. Our results indicate that applied by themselves, these scents do not increase survival of either artificial or natural duck nests. Acknowledgments Delta Waterfowl Foundation provided funding and assistance for this study. Ducks Unlimited Canada kind- ly granted permission to search nesting cover. D. Coul- ton, H. Remenda, J. Estrella provided assistance in the field. Two reviews contributed to the clarity of the man- uscript. This research was approved and conducted under Manitoba Wildlife Animal Care Committee #2002-10. SL was supported through a National Science and Engineering Research Council discovery grant. Literature Cited Chafioun, A. D., F. R. Thompson, and M. J. Ratnaswamy. 2002. Nest predators and fragmentation: a review and meta-analysis. Conservation Biology 16: 306-318. Clark, R. G., and B. K. Wobeser. 1997. Making sense of scents: effects of odour on survival of simulated duck nests. Journal of Avian Biology 28: 31-37. Conover, M. R. 1990. Reducing mammalian predation on eggs by using a conditioned taste aversion to deceive pred- ators. Journal of Wildlife Management 54: 360-365. Guyn, K. L., and R. G. Clark. 1997. Cover characteristics and success of natural and artificial duck nests. Journal of Field Ornithology 68: 33-41. Hazler, K. R. 2004. Mayfield logistic regression: a practical approach for analysis of nest survival. Auk 121: 707-716. Hoover, S. E., and M. R. Conover. 1998. Effectiveness of volatile irritants at reducing consumption of eggs by cap- tive coyotes. Journal of Wildlife Management 62: 399-405. Larivieére, S. 2004. Range expansion of raccoons in the Cana- dian prairies. Wildlife Society Bulletin 32: 955-963. Lariviere, S., and F. Messier. 1997. Characteristics of water- fowl nest depredation by the striped skunk: can predators be identified from nest remains? American Midland Nat- uralist 137: 393-396. Nicolaus, L. K. 1986. Conditioned aversions in guild of egg predators: implications for aposematism and prey defence mimicry. American Midland Naturalist 117: 405-419. 154 Pasitschniak-Arts, M., and F. Messier. 1995. Risk of preda- tion on waterfowl nests in the Canadian Prairies: effects of habitat edges and agricultural practices. Oikos 73: 347-355. Pitt, J. A., S. Lariviere, and F. Messier. 2008. Survival and body condition of raccoons at the edge of their range. Journal of Wildlife Management 72: 389-395. Rosell, F. 2001. Effectiveness of predator odours as gray squir- rel repellents. Canadian Journal of Zoology 79: 1719-1723. THE CANADIAN FIELD-NATURALIST Vol. 121 Sullivan, T. P., L. O. Nordstrom, and D. S. Sullivan. 1985. Use of predator odors as repellents to reduce feeding damage by herbivores II. Black-tailed deer (Odocoileus hemionus columbianus). Journal of Chemical Ecology 11: 921-935. Whelan, C. J., M. L. Dilger, D. Robson, N. Hallyn, and S. Dilger. 1994. Effects of olfactory cues on artificial-nest experiments. Auk 111: 945-952. Received 8 May 2006 Accepted 11 April 2008 Translocation and Recovery Efforts for the Telkwa Caribou, Rangifer tarandus caribou, Herd in Westcentral British Columbia, 1997-2005 ASTRID VIK STRONEN!?, PAUL PAQUET!, STEPHEN HERRERO!, SEAN SHARPE®, and NIGEL WATERS* 'Faculty of Environmental Design, University of Calgary, 2500 University Drive NW, Calgary, Alberta T2N 1N4 Canada *Present Address: Department of Biology, University of New Brunswick, Fredericton, New Brunswick E3B 6C2 Canada: Mailing Address: 6533 rue de Normanville, Montreal, Quebec H2S 2B8 Canada; e-mail: astrid.stronen@unb.ca ‘Ministry of Environment, #325, 1011 Fourth Avenue, Prince George, British Columbia V2L 3H9 Canada ‘Department of Geography, George Mason University, 4400 University Drive MS 1E2, Fairfax, Virginia 22030 USA Stronen, Astrid Vik, Paul Paquet, Stephen Herrero, Sean Sharpe, and Nigel Waters. 2007. Translocation and recovery efforts for the Telkwa Caribou, Rangifer tarandus caribou, herd in westcentral British Columbia, 1997-2005. Canadian Field- Naturalist 121(2): 155-163. During 1997-1999, 32 Woodland Caribou (Rangifer tarandus caribou) were translocated from the Sustut Herd to the Telkwa Mountains in westcentral British Columbia to augment recovery of the Telkwa Caribou Herd. The animals were fitted with radiocollars and located during 1997-2000 to determine selection of habitat features and terrain variables. Six Caribou calves were also collared to determine causes and timing of calf mortality during summer 1999. Defining available habitat for newly translocated animals is often arbitrary and subjective, and we based the analyses on ranks for habitat use and availability as this is less sensitive to the inclusion or exclusion of a questionable resource. This method represents some loss of information but provides indications of the relative importance of various habitat types without classifying any as avoided. High elevation habitat (> 1700 masl) on moderate slopes (16 — 45°) received the highest ranks, as did “warm” (136 — 315°) aspects and forests > 250 years old. Three calves died shortly after birth. One calf appeared to have been killed by predation, likely by a Golden Eagle (Aquila chrysaetos), and one calf was abandoned by the cow. Cause of death for the third calf is unknown. To assess habitat use associated with calving we compared summer locations with data obtained throughout the rest of 1999 for eight cows with calves and eight without calves. We found significant difference in use of elevation during calving time, when cows with calves remained at high elevations and barren cows generally descended to lower elevation habitat. Surveys conducted in 2005, five years after the completion of the initial study, produced a count of approximately 90 Caribou. This suggests that in the short term, the translocation was successful in re-establishing a self-sustaining Caribou population in the Telkwa - Mountains. Key Words: Woodland Caribou, Rangifer tarandus caribou, calf mortality, calving habitat, habitat selection, rank analysis, wildlife management, British Columbia. Woodland Caribou (Rangifer tarandus caribou), pop- ulations have generally experienced decline and range reduction over the last century in British Columbia (Heard and Vagt 1998), across Canada (COSEWIC 2000*) and throughout the circumpolar region (Mal- lory and Hillis 1996). Causes of decline include a com- bination of habitat loss, habitat fragmentation, increased | human access, hunting and predation (Bloomfield 1980; Edmonds and Bloomfield 1984; Heard and Vagt 1998), _ poaching (Edmonds 1988), and possible interactions _ between disturbance and predation (Mallory and Hillis 1996). Woodland Caribou in the Southern Mountain _ National Ecological Area of British Columbia (includ- | ing the Telkwa Caribou Herd) were designated as | threatened in 2000 (COSEWIC 2000*). Caribou were once widely distributed throughout the mountainous areas surrounding Telkwa and Smithers, British Col- umbia, and might have formed part of a larger herd _ whose range extended south into Tweedsmuir Provin- cial Park (TCHRP 1998*). Few data are available on _ the Telkwa Caribou Herd before 1997; however, sur- yveys indicated fluctuations in herd size from less than 100 in 1949 to a high of 271 in 1965. A hunting clo- sure was implemented for the herd in 1973 but only 68 animals were counted in 1984 (van Drimmelen 1986*) and monitoring flights in the 1990s resulted in counts of less than 20 Caribou (TCHRP 1998*). During 1997-1999, 32 Caribou were captured from the Sustut Herd about 150 km north of Smithers and translocated to the Telkwa Mountains to help the recovery of the Telkwa Caribou Herd. Translocation is now a common tool in the recovery of populations at risk (Griffith et al. 1989) and the need for documenting habitat use and availability is critical in efforts to preserve endangered species and manage small populations (Manly et al. 1993). We studied habitat use and calf survival of the Telkwa Caribou Herd from 1997 to 2000 to determine preliminary habi- tat selection of translocated animals and assess timing and causes of calf mortality. With small populations following a translocation, there are usually limited data available to investigate habitat selection, and therefore little or no information for land managers. Although defining available habitat for newly translocated ani- mals is often arbitrary and subjective, a preliminary evaluation of habitat selection can provide some infor- 155 156 mation on priority areas for protection when assess- ing other land uses such as forestry and recreation. Translocation success can be evaluated in three phases: (1) initial occupation and reproduction in the new eco- system; (2) presence of a short-term self-sustaining population, and (3) ultimate population recovery (Compton et al. 1995). We outline findings on habitat selection and calf mortality between 1997 and 2000 and report for phases | and 2 of the Telkwa Caribou Herd Recovery Project. We also discuss challenges involved in providing habitat management priorities based on preliminary data from a small reintroduced population. Study Area The study area comprises approximately 25 000 km? in westcentral British Columbia, with elevations rang- ing from 300-3000 meters above sea level (Figure 1). It is situated between 53.40 and 55.20 degrees of lati- tude north and 125.10 to 128.30 degrees of longitude west. Glaciers, rugged peaks, high alpine plateaus, and deep glaciated U-shaped valleys with extensive wetlands characterize the alpine areas. Major tree spe- cies in the study area include Subalpine Fir (Abies lasiocarpa), Engelmann Spruce (Picea engelmannii), and Hybrid White Spruce (Picea glauca x engelman- nii). The climate has both coastal and inland influ- ences, with cool summers and moderately cold winters. Precipitation averages 500 mm per year, of which about half falls as snow. The Telkwa Mountains have a population of approximately 250 Mountain Goats (Oreamnus americanus), and lower elevations have populations of Moose (Alces alces), Mule Deer (Odo- coileus hemionus hemionus), and some Elk (Cervus elaphus). Large predators within the range of the Telk- wa Caribou Herd include: Grizzly Bears (Ursus arc- tos), Black Bears (Ursus americanus), Wolverine (Gulo gulo), Wolves (Canis lupus), Coyotes (Canis latrans), and Golden Eagles (Aquila chryseatos). To account for the difficulty in defining available resources for the herd and thus delineating our study area, where most animals have had < 2 years of experience, we selected an area of available habitat based on three factors: (1) locations of collared caribou 1997-1998 (G. Schultze, British Columbia Ministry of Environ- ment, personal communication); (2) Traditional Eco- logical Knowledge of the Telkwa Caribou Herd (B. Holland, Wet’suwet’en First Nation, personal commu- nication in Stronen 2000) and (3) availability of forest cover data from the British Columbia Ministry of Forests (Inventory Branch, Victoria, scale 1: 20 000). Methods We located radiocollared animals throughout the year from fixed wing aircraft (Cessna 206, 185, or 172) using standard telemetry equipment. Tracking intervals varied from every four days to every three weeks, de- pending on funding, season, and weather conditions. THE CANADIAN FIELD-NATURALIST Vol. 121 We determined geographic location using yoke-mount- ed, handheld, or panel-mounted Global Positioning System (GPS) units and noted habitat characteristics, activity, group size, and presence of young. The accu- racy of relocation data was tested and averaged + 300 meters. Most locations were determined during morn- ing or mid-day. For the purpose and scale of this study, the data were assumed representative of Caribou habi- tat use for day and night. Six calves were captured shortly after birth and equipped with radiocollars dur- ing the summer of 1999. Caribou calves were captured as close to within 24 hours of birth as possible using a Bell 206 Jet Ranger and a Long Ranger helicopter and a capture crew of three people. Calves were sexed, weighed, and fitted with VHF radiocollars (Lotek En- gineering, Newmarket, Ontario) with motion sensitive mortality sensors. We estimated age by assessing the presence/condition of the umbilical cord, condition of hooves and pelage, posture, and general appearance (Haugen and Speake 1958; Miller et al. 1988), and calculated weight by subtracting 0.6 kg for each day of estimated age (Adams et al. 1995). Handling time was approximately five minutes from capture to release. Calves were monitored daily, weather permitting, from date of capture until the end of August. During Sep- tember and October, we monitored calves weekly and after October they were monitored as a part of regular telemetry flights for the herd. We investigated mortal- ity signals via helicopter the day of discovery. Data for elevation (meters above mean sea level), slope (%), and aspect (in azimuth measured from north) were derived from a Digital Elevation Model using Arc/INFO v. 7 software. Tiles from the Digital Eleva- tion Model were merged to form a seamless grid cov- ering the study area. Floating point grids were gener- ated for slope and aspect and we created a variable combining elevation and slope. Data on habitat type (leading tree species or non-productive timber area), crown closure (%), and tree age (using leading tree spe- cies) were derived from Forest Cover Inventory Poly- gons (British Columbia Ministry of Forests, Inventory Branch, Victoria, scale 1: 20 000). Values for terrain and habitat variables were summarized (km?) for the entire study area to determine percent availability of the various classes of each variable. Habitat availability was analyzed by the British Columbia Ministry of En- vironment’s regional GIS data analyst, who also pro- duced cartographic output. We used data collected between November 1997 and April 2000 to analyse habitat use and determined terrain and forest cover values for all telemetry locations. To assess habitat sel- ection, we followed sampling design 2 of Manly et al. (1993), where individual animals were identified and the use of resources was measured for each individual, but availability was measured at the population level. To assess habitat use associated with calving, we used locations for eight cows with calves (n = 123 loca- tions) and eight without calves (n = 128 locations) 2007 200 0 STRONEN, PAQUET, HERRERO, SHARPE, and WATERS: TELKWA CARIBOL 157 Qe rie, rte anal — ervey (ter peereil eae “ } are | GB an ve at may Corte) 200 Kilometers FiGuRE |. The study area in British Columbia, 1997 — 2000. collected during the summer of 1999. We compared summer data (n = 150 locations) with locations ob- tained during the rest of the year (n = 145 locations) to account for differences not attributed to calving. Locations were divided into two groups: (1) calving (25 May — 6 July) (2) all-year (1 January — 24 May, and 7 July — 31 December) We analyzed habitat selection using Resource Sel- ection for Windows (RSW), a 32-bit Windows program for analyzing preference (Leban 1999). Using Johnson’s (1980) method, which orders all variable components by rank for usage and availability, we measured habitat preference for each animal as the difference between the rank of usage and the rank of availability. We aver- aged these differences across animals to obtain a mean value for each habitat component and then compared the values to determine a ranking from least preferred habitat component to most preferred. We tested for habitat selection using alpha = 0.05. We compared habitat use of reproductive cows and cows without calves using analysis of variance (repeated measures, alpha = 0.05) within each group and within each sea- son using SPSS 10.0 (SPSS Inc. 1999). The herd con- sisted primarily of translocated Caribou during the study and we considered the radio-collared animals to be a random sample of the Telkwa Herd. Results High elevation habitat on moderate slopes (> 1700 meters above sea level, 16 — 45%) received the highest ranks (Table 1). Caribou also made extensive use of the steep rugged mountains in the Telkwa range (>1700 m, > 45%). Aspects 136 — 315° received the highest rank (Table 2). The rank values for aspect ap- pear to contradict the information on use versus avail- ability. However, data on use report only overall percen- tage of use, and taking behavior of individual animals into account; the 136 — 315° aspects receive the highest rank. Less than 4 of locations (405 of 1424) were in forested habitats (Table 3), and of these locations, forests > 250 years old received the highest rank. For analysis within groups, we found a significant difference in mean elevation between calving time and the rest of the year for cows without calves, but not for cows with calves. For analysis within season, we found a significant difference in mean elevation for calving time but not for the rest of the year. Although sample sizes were smaller for calving time, both groups had a narrower range in elevation during calving and individual differences in elevation use were also high- er during calving than the remainder of the year (Fig- ures 2 — 5). Figure boxes represent the interquartile range containing 50% of the values, with a line show- ing the median value. Error bars represent the highest and lowest values, excluding outliers (extreme values). Most cows gave birth in alpine locations, in exposed areas free of snow. Of the 16 cows sampled, only cow #151 made extensive use of elevations below 1400 meters throughout the year (Figure 4). Calving time for Telkwa Caribou ranged from 26 May until 10 June, and six Caribou calves were col- lared between 3 June and 10 June 1999. Five were males born to transplanted Caribou cows and one was a female calf born to the only resident collared cow. Average birth weight was 8.4 kg (range 6.9 — 9.4 kg). Three of the six calves died shortly after birth. One calf likely died from abandonment, whereas cause of 158 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Use, availability and rank of classes for elevation and slope. Telkwa Mountains, British Columbia, Canada 1997-2000. N = 1424 Elevation (m) Slope (%) Class Locations <1100 0-15 1 116 <1100 16-45 2 38 <1100 >45 3 4 1100 — 1700 0-15 4 166 1100 — 1700 16-45 5) 303 1100 — 1700 >45 6 168 >1700 0-15 7 144 >1700 16-45 8 313 >1700 >45 9 172 % use % available Rank 8.15 38.43 9 2.67 20.75 8 0.28 6.40. 7 11.66 17.98 6 21.28 9.71 4 11.80 3.00 5 10.11 2.18 3 21.98 1.18 1 12.08 0.36 2 TABLE 2. Use, availability and rank of classes for aspect. Telkwa Mountains, British Columbia, Canada 1997—2000. N = 136 — 315 1 627 14162 Aspect (azimuth) Class Locations 316 — 135 D) 789 % use % available Rank 44.03 47.24 1 55.41 48.23 2 4 Areas of zero aspect (usually where relocation error positioned animals in water) are excluded. TABLE 3. Use, availability and rank of classes for forest age. Telkwa Mountains, British Columbia, Canada 1997-2000. N = 405° 8.47 4 Forest age (years) Class Locations 1-40 I 18 41 — 100 D 84 101 — 140 3 105 141 — 250 4 161 >250 5 Sy) > Including only locations below tree line (1700 m). death for the second calf appeared to be avian preda- tion, most likely Golden Eagle. Cause of death for the third calf was unknown. The calf had remains of what appeared to be milk curds in the stomach and small amounts of vegetation. However, membranes inside the hide appeared dry and indicated dehydration. No signs of predation or scavenging were evident. We found adult tracks, likely made by the mother, circling the carcass. During the first year after the transplant (1998), we observed five calves in the herd. The following year, we saw one yearling in the herd, as well as 12 new calves and one cow with an extended udder, indicat- ing she recently had given birth and lost her calf. We observed eight calves during the 1999 October calf count. Recent surveys of the herd found 16 cows with calves in October 2003 and 23 the following year. In October 2005 ten cows with calves were counted. How- ever, the decrease in cow/calf numbers from 2004 to 2005 may reflect fewer functioning radio collars in the herd during 2005, thus increasing the probability that some Caribou groups went undetected (G. Schultze, personal communication). % use % available Rank 1.26 5.90 17.82 3 7.37 14.85 Dp, 11.31 24.60 5 2.53 7.08 1 Discussion The Telkwa Caribou Herd appears to select high elevation habitat with moderate slopes, which corre- sponds with earlier observations (van Drimmelen 1986*; Bill Holland, personal communication in Stro- nen 2000). The herd uses gentle to moderate slopes throughout much of the year, which is common for Woodland Caribou (Edmonds and Bloomfield 1984; Seip 1990*; Terry et al. 1996; Wood and Terry 1999). However, use of steep slopes is also reported (Hatler 1987*; Terry et al. 1996). Woodland Caribou have been found using southern/warm aspects (Edmonds and Bloomfield 1984; Warren et al. 1996), northern/cool aspects (Warren et al. 1996) or to show no apparent selection (Seip 1990*). Use also varies between years and seasons (Terry et al. 1996; Wood and Terry 1999). When in forested habitat the Telkwa Caribou Herd appears to select old forests, which corresponds with other studies (Cichowski 1993; Terry et al. 1996; Wood and Terry 1999; Apps and Kinley 2000*). Woodland Caribou in British Columbia have been classified into two ecotypes (Stevenson and Hatler 1985). The north- : ! { : ’ 2007 z oO m > TT) i TT) Pay w = ce lu a Ke 1s Zz 7 001 O71 272 CARIBOU COWS STRONEN, PAQUET, HERRERO, SHARPE, and WATERS: TELKWA CaArisot 159 610 FiGuRE 2. Barren cows’ use of elevation all-year, 1999. Telkwa Mountains, British Columbia. ‘ 4s DERIVED ELEVATION Ke 1¢ 1 10 004 o71 272 CARIBOU COWS i. . i" On Os Oss =) 7 = = s 283 291 177 610 801 FiGure 3. Barren cows’ use of elevation at calving time, 1999. Telkwa Mountains, British Columbia. er ecotype (including the Telkwa Caribou Herd) depends primarily on terrestrial lichens during winter and the mountain ecotype depends more on arboreal lichens. Arboreal lichens are most abundant in old for- est (Apps and Kinley 2000*), often on trees > 120 years old (Stevenson and Hatler 1985; Wood and Terry 1999). We found the Telkwa Caribou Herd in forested habitat only one-third of the time. However, divisions between arboreal- and terrestrial lichen-dependent groups are not absolute. The temporal impacts of icing, deep snow, and other environmental conditions may at times pre- vent Caribou from obtaining terrestrial lichens, making areas of arboreal lichens provisionally more impor- tant (van Drimmelen 1986*; Edmonds 1988: Cichow- ski 1993). Forests 141—250 years old received the high- est use but the lowest rank. This is the most abundant forest age class (Table 3). Whereas the small number of Caribou during this study did not use this age class 160 in proportion to availability, the low rank does not equal avoidance or lack of importance. Future monitoring could also help clarify whether certain areas within this forest age class are locally important. The difference in elevation use between barren and reproductive cows during calving season might be due to reproductive status. There are, however, wide dif- ferences within each group. As four of eight barren cows remained at high elevations (which presumably meant forgoing nutritional opportunities, at least in spring/early summer), this could be an antipredator strategy employed regardless of reproductive status. Based on this study, possible antipredator tactics for protecting a calf can not be distinguished from those that cows seem to use for their own protection. Ed- monds and Smith (1991) reported that cow/calf pairs localised movement to relatively small geographic areas in early June (1-15) and did not increase the size of their ranges until after July. The Telkwa Caribou displayed similar behaviour, as cow/calf pairs in the Telkwa Herd remained relatively stationary until the middle of July. Subsequently, some pairs made long distance movements (one pair moved 50 km over a few days). The calving time for Telkwa Caribou cor- responded to that reported by Edmonds (1988) for westcentral Alberta. As only one birth from a resident Caribou was recorded (10 June), it was not possible to examine any differences between transplant and resi- dent animals. Some cows in the Telkwa Caribou Herd appeared more sensitive to fixed-wing aircraft during the first couple of weeks post-calving than during the rest of the year, but we noted a wide range in behaviour. Cichowski (1993) reported that most adult females were found alone on ridge tops during calving. Two Telkwa cows calved on the top of a rocky ridge and were joined by two other cows with calves in the mid- dle of June. The ridge had few access points but bio- physical information suggests low potential for lichen production. All four calves were found alive in Octo- ber. The use of ridges may be a trade-off between pred- ator avoidance and the availability of abundant high protein food (Bergerud and Miller 1984). Cichowski (1993) found that most calving sites were within 10 km of previous calving sites and Seip (1990*) reported many Caribou use the same calving locations each year. We could not analyse fidelity to calving areas within the Telkwa Herd and it would take several years before this could be assessed. Caribou calving areas could change from year to year as an antipredator strategy (Bergerud and Miller 1984), and Valkenburg et al. (1988) warned against making management decisions based on Caribou calving areas located during narrow time intervals. However, previously known calving areas identified by the Traditional Ecological Knowl- edge of the Wet’suwet’en First Nation (B. Holland, in Stronen 2000) should be considered when making management decisions, as this information is based THE CANADIAN FIELD-NATURALIST Vol. 121 on years of observation and would indicate fidelity to specific calving sites. Small sample size precluded analysis of calf mortal- ity for the Telkwa herd but suggested that avian preda- tion was a factor. Recruitment data from November 1998 to April 2000 seemed to indicate that low pro- duction of calves was a factor of equal or higher im- portance than the loss of calves. If the low recruitment of calves was due to calf predation, a higher frequen- cy of Caribou cows with extended udders but without calves would be expected. Females without calves commonly shed their antlers in March and April and exhibit fresh antler growth during the calving period, whereas pregnant females shed their antlers near calv- ing in May and June (Bergerud 1980). During our sur- veys at calving time, many cows exhibited fresh antler growth, indicating that they were not expecting a calf that year. The Caribou transplanted from the Sustut Cari- bou Herd would be expected to show pregnancy rates comparable to those of other populations when they arrived, and reported pregnancy rates usually range from 80% to 90% (Bergerud 1980; Seip 1990*; Ed- monds and Smith 1991). It is possible that stress relat- ed to capture, immobilization, and transplant may have caused abortions or fetal absorptions, or that a large number of the cows were by chance not pregnant. In addition, there may not have been enough bulls to achieve normal pregnancy rates for the Telkwa herd during the first years. At least one resident mature bull was present in the herd during 1998 but due to difficul- ties associated with movement and collaring of adult bulls only younger males were translocated to the Tel- kwa Mountains. Human activities such as recreation and resource extraction may also influence Caribou habitat use. Van Drimmelen (1986*) expressed trepidation over snow- mobiling on the winter ranges of the Telkwa Herd and this has been a concern for over 25 years (Bustard 1977*). The presence of dogs may also cause Caribou to avoid otherwise useful habitat. This is particularly important during calving time, when cows may be extra sensitive to disturbance (Bergerud and Miller 1984). Need for information related to species translo- cations and/or small remnant populations will increase in the future. Where habitat availability is poorly esti- mated it may be particularly important not to label habitats and resources, which may be important in the long term, as avoided. We chose Johnson’s (1980) method because it is relatively insensitive to decisions on what constitutes available habitat for the Telkwa Caribou Herd. If habitat components are used in pro- portion to availability, the rank ordering of use and availability will be the same. However, the converse is not necessarily true; the use and availability propor- tions could be different across habitat components, while still having the same rank, thus resulting in high Type 2 errors (where a difference in proportional selection is not detected) (Alldredge and Ratti 1986, : : | 2007 DERIVED ELEVATION co o 3 CARIBOU COWS STRONEN, PAQUET, HERRERO, SHARPE, and WATERS: TELKWA CARIBOI 16] FIGURE 4. Reproductive cows’ use of elevation all-year, 1999. Telkwa Mountains, British Columbia. 2200 2000 1800 1600 1400 1200 1000 800 DERIVED ELEVATION 024 O32 034 CARIBOU Cows 151 251 770 256 $70 FiGuRE 5. Reproductive cows’ use of elevation during calving time, 1999. Telkwa Mountains, British Columbia. 1992). A bias may also occur against small habitat patches when the size of error area for locations [here + 300 m] is independent of habitat patch size (Rettie and McLoughlin 1999). Many wetlands in our study area were too small to be captured in Forest Cover data and Houwers (1996*) reported that distribution of ar- boreal lichens in the Telkwa Mountains was patchy. These two habitat units, too small to be noted by our analysis, may be vital for long-term survival of the Telkwa Caribou Herd. The size of habitat required by Caribou to avoid predation may be significantly greater than the amounts required to obtain sufficient forage (Stevenson et al. 1994*: Seip and Cichowski 1996), and requirements for predation avoidance could be 10 times larger (Bergerud 1980). It is theorized that the Telkwa Mountains could support approximately 162 300 caribou (van Drimmelen 1986*). Given sufficient habitat and calf survival the herd could recover to historic numbers. At present, the translocation appears to have been successful in establishing a self-sustaining Caribou herd in the Telkwa Mountains. Acknowledgments The Habitat Conservation Trust Fund, Forest Ren- ewal BC, the British Columbia Ministry of Environ- ment, and a Graduate Research Scholarship from the Faculty of Environmental Design, University of Cal- gary, provided funding for this study. Staff at the Bri- tish Columbia Ministry of Environment in Smithers assisted with fieldwork, and G. Schultze provided survey data from 2000 to 2005. We thank J. Warren and other staff at the GlIS-section for help with all GIS-related work. Northern Lights Air piloted fixed- wing telemetry flights, and Canadian Helicopters pilot- ed flights to capture caribou calves and investigate mortalities. An anonymous reviewer provided helpful comments on the manuscript. Documents Cited (marked * in text) Apps, C., and T. Kinley. 2000. Multi-scale habitat associa- tions of Mountain Caribou in the Southern Purcell Moun- tains, British Columbia. East Kimberley Environmental Society, Kimberley British Columbia and Crestbrook Forest Industries Ltd., Cranbrook, British Columbia. Bustard, D. 1977. Caribou observations — Hunter’s Basin — Camel Humps Area. Fish & Wildlife Branch memorandum, Ministry of Environment, Lands and Parks, 14 March, LOTT: COSEWIC. (Committee on the Status of Endangered Wildlife In Canada) 2000. Updated Status Report on the Woodland Caribou (caribou des bois) Rangifer tarandus dawsoni and Rangifer tarandus caribou in Canada. Avail- able at http://www.sararegistry.gc.ca/virtual_sara/files/cose wic/sr_woodland_caribou_e.pdf Hatler, D. F. 1987. Perspectives on inventories of caribou in British Columbia. Spatsizi Association for Biological Research Report (4). Wildlife Report (R-14) July 1987. Houwers, C. 1996. Lichen Abundance Inventory of the Telkwa Range. British Columbia Ministry of Environ- ment, Lands and Parks, Smithers. Seip, D. R. 1990. Ecology of woodland caribou in Wells Gray Provincial Park. Wildlife Bulletin B-68, March 1990. Ministry of Environment, Wildlife Branch, Victoria, British Columbia and Ministry of Parks, Kamloops, British Col- umbia. Stevenson, S. K., H. M. Armleder, M. J. Jull, D. G. King, E. L. Terry, G. S. Watts, B. N. McLellan and K. N. Child. 1994. Mountain Caribou in Managed Forests: Pre- liminary Recommendations for Managers. Ministry of Forests, Research Branch, Victoria, British Columbia. TCHRP (Telkwa Caribou Herd Recovery Plan). 1998. Telkwa Caribou Herd Recovery Team, British Columbia Ministry of Environment, Lands and Parks, Skeena Region, British Columbia, Canada. van Drimmelen, B. 1986. Telkwa Mountains Caribou Tele- metry Project, Progress Report # 2. Ministry of Environ- ment, Smithers, British Columbia THE CANADIAN FIELD-NATURALIST Vol. 121 Literature Cited Adams, L. G., F. J. Singer, and B. W. 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Habitat use and movements of two ecotypes of translocated Caribou in Idaho and British Columbia. Con- servation Biology 10: 547-553. Wood, M. D., and E. L. Terry. 1999 Seasonal movements and habitat selection by woodland caribou in the Omineca Mountains, North-Central British Columbia. Phase |: The Chase and Wolverine Herds (1991-1994). Peace/Williston Fish and Wildlife Compensation Program Report (201). Received 17 May 2006 Accepted 10 March 2008 An Inventory of the Aquatic and Subaquatic Plants in SASK Water Canals in Central Saskatchewan, Canada, Before and After the Application of the Herbicide Magnacide J. HUGo CoTa-SANCHEZ! and KIRSTEN REMARCHUK Department of Biology and Herbarium of the University of Saskatchewan, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada ' Corresponding author: hugo.cota@usask.ca Cota-Sanchez, J. Hugo, and Kirsten Remarchuk. 2007. An inventory of the aquatic and subaquatic plants in SASK Water canals in central Saskatchewan, Canada, before and after the application of the herbicide Magnacide. Canadian Field-Naturalist 121(2): 164-167. This study focuses on the floristic composition of aquatic and semi-aquatic plants in the SASK Water canal system and their potential effect on irrigation systems. A checklist, evaluation, and synthesis of the species identified in this survey before and after the application of the herbicide Magnacide are provided, in addition to a brief discussion of the environmental effects of Magnacide. Thirty-three species in 26 genera within 20 plant families were identified. Two unidentified green algae were also collected. Common aquatics (i.e., green algae, Potamogeton spp., Alisma gramineum, A. plantago-aquatica, Ceratophyllum demersum, and Myriophyllum sibiricum) combined with debris from terrestrial plants were the primary contributors to block- age of irrigation drains. In general, the concentration of Magnacide used in this study had a minor effect on aquatic plant diversity, but effectively reduced plant density. However, the long-term effects of pesticides on the surrounding aquatic and terrestrial environments of the SASK Water irrigation system are unknown. Key Words: aquatic plant inventory, environmental effect, SASK Water canals, Magnacide, acrolein, Saskatchewan. Plants are vital to the function of aquatic ecosystems for their role in providing food, oxygen, and habitat for other organisms. However, floating and submerged vegetation along with debris from terrestrial plants may be problematic in different types of water bodies. For example, in shallow lakes, profuse plant growth creates dense mats that prevent the regular movement of watercraft. In addition, aquatic plants, including algae, can significantly reduce the aesthetic value of water bodies. Furthermore, rapid growth of vegetation and deposition of debris in irrigation canals can result in a decline in water flow rates and a subsequent in- ability to supply water to crops. Hence, the removal of selected aquatic species via physical or chemical meth- ods may be required to improve ecosystem function, aesthetic value, and availability for human consump- tion. Aquatic plants can be placed into four broad cate- gories: (1) algae, (2) floating-leaved, (3) submerged, and (4) emergent plants (Shelton and Murphy 1989). Of the four categories listed above, algae are of the greatest concern in canal systems because of the likelihood of interfering with water flow. Algae grow profusely in slow moving, shallow water. Floating-leaved and sub- merged plants are rooted, with the foliage at or below the water surface. Flowing water or disturbance, such as wind, can uproot plants leading to subsequent ob- struction of water flow. Finally, emergent species typ- ically grow in shallow, fluctuating water, such as drain- age ditches, canals, rivers, periphery of water bodies, and ponds. Plants in this category are of minor concern in canal systems and in some cases may prevent non- aquatic plant debris from entering the system. Aquatic species impede water flow by clogging gates, intake screens, valves, and pumps in irrigation and drainage channels (Lancar and Krake 2002). As a consequence, localized flooding may result under high water conditions. In addition, Holm et al. (1969) indi- cate that abundant vegetation in canals can lead to an excessive loss of water through evapotranspiration. pondweed (Potamogeton L.), Milfoil (Myriophyllum sibiricum), Canada Waterweed (Elodea canadensis), wild celery (Vallisneria L.), and American Eel-grass (Zostera marina L.) are a few of the problematic coarse or large species in shallow aquatic environments (Lan- car and Krake 2002). Of particular interest to this study are aquatic taxa that interfere with canal irrigation systems in central Saskatchewan. Accelerated growth of plants in SASK- Water canals restricts the amount of water available for irrigation. Given the clogging effect of aquatic plant growth on water flow, agricultural canals were treated with Magnacide, a volatile algaecide and aquatic her- bicide, in which the active ingredient is acrolein (Score- card 2006*). Acrolein contains two functional groups, a reactive double bond and an aldehyde group (Nor- done et al. 1996) and is, in turn, toxic to some organ- isms (Albarifio et al. 2007). However, it is widely used in agricultural canals and water bodies as an herbicide to control aquatic weeds (Burland et al. 1984; San- godoyin and Smith 1996). The use of high concentra- tions of acrolein could pose a significant risk to aquat- | ic environments (Nordone et al. 1996); therefore, the controlled application and use of Magnacide and other 164 2007 herbicides is highly recommended. Previous studies have shown that toxic residues deposited in soils and water bodies have detrimental effects in the life cycles of local flora and fauna (Sangodoyin and Smith 1996). Thus, even though several sources, e.g., Sangodoyin and Smith (1996); Nordone et al. (1996); Albarino et al. (2007), claim that the transient use of acrolein in agricultural waters at minimal amounts has no negative impact in natural aquatic environments, rigid control must be enforced to prevent ecological damage. In this study, we investigated the floristic composi- tion of aquatic and semi-aquatic plants in the SASK- Water canal system and their potential effect on ir- rigation systems. Our study is a contribution to the knowledge of Saskatchewan aquatic and sub-aquatic flora and provides a preliminary assessment and syn- thesis of the species identified prior to and after the application of Magnacide. The potential effect of Magnacide on aquatic flora is also discussed. Methods Compared to most wetland inventories, this survey covers a limited geographic area and a restricted range of wetland flora. The site selection and sampling strat- egy were designed to survey all major areas of the SASK Water canal system to be evaluated in terms of aquatic vegetation within the Saskatoon Southeast Water Supply Project Location Plan (Brightwater-Blackstrap- Zelma Reservoir areas; 51°25' to 52°55'N, 106°15' to 107°00'W). This system irrigates an estimated area of 20 120 acres. The survey sites include locations of Magnacide application by SASKWater personnel at intake screens. Acrolein was directly applied to water in strategic sites to control submersed, floating and emergent aquatic weeds. The herbicide was applied at a concentration of 2.5-5.0 ppm/30 minutes every two weeks from June through August 2005 in an attempt to eradicate aquatic species with excessive growth rates (J. Mander, SASK Water, personal communication). Surveys were conducted before and after the chemi- cal treatment to determine the effect of Magnacide on the taxonomic richness and occurrence of aquatic plants in the canal system. Plant collections were con- ducted in 25-50 m areas before and after the point of application. Plants were pressed and identified in the herbarium of the University of Saskatchewan (SASK) using several local and regional references (Hotchkiss 1972: Larson 1993: Johnson et al. 1995; Moss 1996: Vance et al. 1984; Royer and Dickinson 1999). The plants collected were prepared as voucher specimens to serve as a permanent reference of this study in the SASK collection. Results and Discussion The survey indicates that the taxonomic composi- tion of the SASK Water canals includes a total of 33 species in 26 genera and 20 plant families (Table 1). In addition, two different, unidentified, algae were col- lected (Table 1). Our survey also indicates that no plant CotaA-SANCHEZ and REMARCHUK: AQUATIC AND SUBAQUATIC PLANTS 165 species were eradicated from the sites as a result of Magnacide application; that is, the same species were identified before and after the treatment. However, the plant density was noticeably lower after the treat- ment with Magnacide. Various submerged species in- cluding Potamogeton richardsonii, P. pectinatus, and the unknown green algae were present in most of the sites surveyed. Other aquatics, e.g., Elodea canadensis and Lemna trisulca, were less frequent and account- ed for a smaller proportion of plant biomass in the canal system. Emergent species in the Cyperaceae, Equisetaceae, and Asteraceae do not appear to play a role in impeding water flow, but floating debris from non-aquatic taxa, such as Melilotus albus and Hordeum jubatum, contributes to the clogging of intake screens throughout the canal system. Based on this survey, it is not possible to determine a single species that adversely affects water flow by clogging intake screens in the SASK Water canal sys- tems. A combination of common aquatic plants ( Pota- mogeton spp., Alisma gramineum, A. plantago-aquat- ica, Ceratophyllum demersum, and Myriophyllum sibiricum), which are frequently dislodged by flowing water or wind, are the primary contributors to blockage of drains and intake screens. The presence of large, coarse water pondweeds such as Potamogeton spp. and other filamentous species in SASK Water canals is un- desirable because their accumulation affects the reg- ular water flow. A similar situation has been reported in the Alberta irrigation systems (Burland and Catling 1986). In addition, green algae are common and are presumably the greatest problem in clogging the in- take screen systems because of their filamentous nature and massive growth. Furthermore, some plants or frag- ments from non-aquatic plants, particularly Melilotus albus and Hordeum jubatum, are involved in reducing water flow because the species tend to concentrate near the uptake areas, especially in the vicinities of mowed ditches along the canals. Since this survey is mainly concerned with the flo- ristic composition of the SASK Water canal system and the probable effect of some species on irrigation canals, only general comments can be made regarding the chemical treatment and subsequent effect in the aquat- ic and subaquatic plants. In general, the use of Magna- cide at the concentration previously specified appears to have a minor effect on aquatic plants because spe- cies diversity before and after the application remained similar. This suggests that the amount of chemical applied is too low to notably reduce the growth and development of aquatic plants and possibly other non- target organisms. Acrolein is a contact herbicide; there- fore, the root system is not damaged and plants quickly resume growth. This leads to the frequent re-applica- tion of acrolein (J. Mander, SASK Water, personal com- munication). Other studies, [e.g., Bowmer and Sainty (1977)], have shown that acrolein is effective in con- trolling aquatic plant growth, but no concentration rates are given. On the other hand, Sangodoyin and Smith 166 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 1. Taxonomic list of the inventory of plants in the SASK Water canal system in south central Saskatchewan. The arrangement is alphabetical by family, then by genus and species. Family Species Characeae Unknown green algae Chara L. Alismataceae Alisma gramineum Le}. Alisma plantago-aquatica L. Apiaceae/Umbelliferae Sium suave Walter Asteraceae/Compositae Symphiotrichum lanceolatum (Willd.) G.L. Nesom subsp. hesperium (A. Gray) G.L. Nesom Ceratophyllaceae Ceratophyllum demersum L. Cyperaceae Carex rostrata Stokes Eleocharis acicularis (L.) Roem. & Schult. Eleocharis palustris (L.) Roem. & Schult. Schoenoplectus acutus (Muhl. ex Bigelow) A. Love & D. Love Schoenoplectus tabernaemontani (C.C. Gmel.) Palla Scirpus microcarpus J. & K. Presl. Equisetaceae Equisetum palustre L. Fabaceae/Leguminosae Melilotus albus Medik. Halogaridaceae Myriophyllum sibiricum Kom. Hydrocharitaceae Elodea canadensis Michx. Lamiaceae/Labiatae Lycopus asper Greene Lemnaceae Lemna trisulca L. Onagraceae Epilobium ciliatum Raf. Oenothera biennis L. Poaceae/Graminae Glyceria grandis S. Watson Hordeum jubatum L. Phalaris arundinacea L. Potamogetonaceae Potamogeton filiformis Pers. Potamogeton pectinatus L. Potamogeton richardsonii (Benn.) Rydb. Potamogeton vaginatus Turcz. Potamogeton zosteriformis Fern. Ranunculaceae Ranunculus aquatilis L. Salicaceae Salix exigua Nutt. Sparganiaceae Sparganium sp. L. Typhaceae Typha latifolia L. Zannichelliaceae Zannichellia palustris L. Common Name Narrow-leaved Pondweed Broad-leaved Water Plantain Water Parsnip Western Willow Aster Hornwort Beaked Sedge Needle-Spike Rush Common or Pale Spike-Rush Great Bulrush Small-fruited Bulrush Marsh Horsetail White Sweet Clover Water Milfoil Canada Waterweed Water Horehound Duckweed Northern Willow Herb Yellow-Evening Primrose Manna Grass Wild Barley Reed Canary Grass Leafy Pondweed Sago Pondweed Clasping Leaf Pondweed Large-sheath Pondweed Flat-stemmed Pondweed Thread-leaved Buttercup Narrow-leaved Willow Bur-Reed Common Cattail Horned Pondweed (1996) caution that low concentrations of pesticides and agricultural chemical on surface waters have poten- tial effects on fisheries and aquatic environments, but conclude that Canadian water sources in general exhib- it concentrations of pesticides below the guideline lim- its, which suggests the proper use and handling of pes- ticides and herbicides in Canada. Nonetheless, more detailed research is required to draw conclusions on water quality issues. Considering the losses and adverse effects to farm- ing activities caused by obstructed canal systems, the control of aquatic plants is necessary in water bodies, particularly in areas with intake screens, so that irri- gation systems can be most efficiently utilized. In addi- tion to the chemical methods currently used for the eradication of aquatic plants, new research to imple- ment a biological control program (involving perhaps an herbivorous species) of aquatic plants in irrigation canals should be conducted. Since numerous non- aquatic species clog irrigation systems, in our view, the most effective, environmentally friendly, and in- expensive strategy for maintaining levels of water flow for irrigation is to periodically manually clean intake screens and surrounding areas. Nonetheless, cautious and planned applications of chemicals such as acrolein (Bowmer and Sainty 1977), Endothall (Sisneros et al. 1998), and Hydrothall (Corbus 1982) may be effective to control excessive growth levels of species such as Potamogeton pectinatus and filamentous algae species. Since several native, relatively poorly represented species are characteristic of these ecosystems, consis- tent monitoring is recommended to preserve wild nat- ural resources. Furthermore, the long-term effects of pesticides on the surrounding aquatic and terrestrial environments of the SASK Water irrigation system are unknown. The current challenge is to assess the use of herbicides, in particular the long-term effect and impact of Magnacide residues, in the water quality, environ- ment, and diversity of the wetland flora and fauna in Saskatchewan and other Canadian ecosystems. 2007 CoTa-SANCHEZ and REMARCHUK: Acknowledgements We thank SASK Water personnel, in particular K. Berkers for help with surveys, J. Mander for comments and for showing the canal system, and two anonymous reviewers for their comments on the manuscript. We also thank the personnel of the Herbarium of the Uni- versity of Saskatchewan (SASK) for their assistance. Documents Cited (marked * in the text) Scorecard: The Pollution Information Site. http://www. scorecard.org/chemical-profiles/summary.tcl?edf_sub stance_id=107-02-8. Site visited 22 May 2006. Literature Cited Albarino, R., A. Venturino, C. M. Montagna, and A. M. Pechen D’Angelo. 2007. Environmental effect assess- ment of Mangnacide® H herbicide at Rio Colorado irriga- tion channels (Argentina). Tier 4: In situ survey on benth- ic invertebrates. Environmental Toxicology and Chemistry 26: 183-189. Bowmer, K. H., and G. R. Sainty. 1977. Management of aquatic plants with acrolein. Journal of Aquatic Plant Man- agement 15: 40-46. Burland, R., and P. M. Catling. 1986. Vascular aquatic weeds in Alberta. Pages 1-4 in Canadian Chapter Aquatic Plant Management Society 1. Calgary, Alberta. Burland, G. R., M. D. O’Shea, D. Stix, and R. Veger. 1984. Investigation into the behaviour of Magnacide H in Alberta irrigation canals. Alberta Environment Pesticide Chemicals Branch, Edmonton, Alberta. Corbus, F. G. 1982. Aquatic weed control with Endothall in a Salt River Project Canal. Journal of Aquatic Plant Man- agement 20: 1-3. Holm, L. G., L. Weldon, and R. D. Blackburn. 1969. Aquat- ic weeds. Science 166: 699-709. Hotchkiss, N. 1972. Common Marsh, Underwater and Float- ing-leaved Plants of the United States and Canada. Dover Publications Inc. 124 pages. AQUATIC AND SUBAQUATIC PLANTS 167 Johnson, D., L. Kershaw, A. MacKinnon, and J. Pojar. 1995. Plants of the Boreal Western Forest and Aspen Park land. Lone Pine Publishing, Edmonton, Alberta. 392 pages Lancar L., and K. Krake (reviewers). 2002. Aquatic weeds and their management. International Commission on Irri- gation and Drainage, USA. 71 pages. Larson, G. E. 1993. Aquatic and Wetland Vascular Plants of the Northern Great Plains. General Technical Report RM-238. U.S. Dept. Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Col- lins, CO. 681 pages. Moss, E. H. 1996. Flora of Alberta, 2” Edition. Revised by J. C. Packer, University of Toronto Press, Toronto, Ontario 687 pages. Nordone, A. J., R. Matherly, B. Bonnivier, R. Doane, H. Caravello, S. Paakonen, W. Winchester, and R. A. Par- ent. 1996. Effect of Magnacide” H herbicide residuals on water quality within wildlife refuges on the Klamath Basin, CA. Bulletin of Environmental Contamination and Toxi- cology 56: 964-970. Royer, F., and D. Dickinson. 1999. Weeds of Canada and Northern United States. Lone Pine Publishing, Edmonton, Alberta. 434 pages. Sangodoyin, A. Y., and D. W. Smith. 1996. Review of infor- mation on pesticide residues in the Canadian environment. The Environmentalist 16: 187-196. Shelton, J. L., and T. R. Murphy. 1989. Aquatic Weed Man- agement: Control Methods. Southern Regional Aquacul- ture Center, Publication 360. 2 pages. Sisneros, D., M. Lichtwardt, and T. Greene. 1998. Low-dose metering of endothall for aquatic plant control in flowing water. Journal of Aquatic Plant Management 36: 69-72. Vance, F. R., J. R. Jowsey, and J. S. MacLean. 1984. Wildflowers Across the Prairies. Western Producer Prairie Books, Saskatoon, Saskatchewan. 214 pages. Received 10 July 2006 Accepted 23 May 2008 Life History of the Marine Isopod Cyathura polita in the Saint John River Estuary, New Brunswick: a Species at the Northern Extent of its Range SARAH C. MERCER!, GLENYS D. G1BSON2, and MICHAEL J. DADSWELL? Department of Biology, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada 'Sarah_mercer9 @hotmail.com * glenys.gibson @acadiau.ca >Corresponding author. Email: mdadswell @bwr.eastlink.ca Phone: (902) 585-1161. Fax: (902) 585-1059 Mercer, Sarah C., Glenys D. Gibson, and Michael J. Dadswell. 2007. Life history of the Marine Isopod Cyathura polita in the Saint John River estuary, New Brunswick: A species at the northern extent of its range. Canadian Field-Naturalist 121(2): 168- WIGS The marine isopod, Cyathura polita, inhabits estuaries on the east coast of North America from the Gulf of Mexico to the Bay of Fundy, Canada. We studied C. polita in the Saint John estuary to test for potential differences in life history that might occur because of the northern location of the population. In the Saint John, based on our interpretation from a six-month sampling program (May-October), the population exhibits a three-year life cycle, one year longer than more southern populations, and stretching over four summers. Our study supported the occurrence of protogynic hermaphroditism. After two summers as juveniles, individuals matured as females during their third summer, then displayed sex reversal by becoming males that fall, and finally reproducing as males in their fourth summer of life before death. Mean length of C. polita from the Saint John was greater than individuals from more southern populations (females, 13.8 + 2.14 mm; males, 16.3 + 2.41 mm). Annual brood release occurred in late July-early August. Mean fecundity of females was 53.2 + 18.9 embryos per brood, which was greater than found in southern populations. Cyathura polita is rare in Canada and is known only from the Saint John and along the northern shore of the Bay of Fundy to the border of the United States. Key Words: Isopod, Cythura politica, growth, reproductive cycle, fecundity, protogynic hermaphroditism, status, St. John River, New Brunswick. Our knowledge of life history evolution depends on understanding intraspecific variation in life history traits, such as those correlated with latitude. Cyathura polita (Stimpson, 1855) is a benthic isopod that occurs in estuaries along the Atlantic coast of North America from the Gulf of Mexico to the Bay of Fundy, Canada (Burbanck et al. 1979), and is an appropriate model for such a study. Latitudinal variations in life history traits have been demonstrated for C. polita populations from Florida to Cape Cod, and include adult size and life span (Burbanck and Burbanck 1979). Latitudinal dif- ferences in life history traits have been described in the common cyathuran of the eastern Atlantic, Cyathura carinata, among populations from Portugal to Swe- den (Bamber 1985; Olafsson and Persson 1986; Mar- ques et al. 1994; Cruz et al. 2003). In the present study, we investigated life history characteristics of C. polita for its northernmost known population in the Saint John River estuary, New Brunswick. Cyathurans are interstitial isopods that live in bur- rows and feed on detritus and algae in the sediment (Burbanck 1959). They are eurytopic animals, tolerat- ing wide fluctuations in conditions such as temperature and salinity, an adaptation suitable to estuaries. They are also an important indicator organism for healthy estuarine ecotones because they are usually present only in unpolluted waters (Burbanck and Burbanck 1979). Cyathurans in general do not swim as adults nor do they have a pelagic larval stage, resulting in popula- tions that are relatively immobile, sessile and disjunct. According to Burbanck (1959, 1962a, b), C. polita inhabits estuaries where fresh and salt water mix over stable sediment containing gravel, silt and detritus. They are obligatory aquatic animals and cannot with- stand long periods of dryness (Burbanck and Bur- banck 1979). Often C. polita are located in transition- al zones where cattails (Typha) and salt marsh grasses (Spartina) meet along the shoreline (Burbanck 1962b). They are an important food for ducks and fishes (Bur- banck and Burbanck 1979; Dadswell 1979). The life cycle of C. polita varies with latitude (Bur- banck and Burbanck 1979). In southern populations, . the life cycle duration is generally one year. In Florida, young are released from May to October, and grow over the winter until they mature and produce young } the following spring and summer. Most southern cya- - thurans die at the end of their second summer (Kruczyn- - ski and Subrahmanyam 1978). In more northern loca- tions, such as Cape Cod, the young are released in early | summer (late June or early July), grow rapidly in the® fall, and produce young the following year. In these populations some cyathurans die at the end of their second summer while others may survive a second win- ter and reproduce the following year, often exhibiting protogynic hermaphroditism (Burbanck and Burbanck ID), 168 2007 Cyathuran females carry their embryos in a ventral marsupium made of overlapping pairs of oostegites (Stromberg 1972). The embryos develop in the mar- supium until they hatch from a vitelline membrane and are released into the environment as juveniles. This method of reproduction permits detailed study of fec- undity and development in cyathurans, making the species an excellent model for life history studies. Cyathura polita has been studied extensively in the United States, particularly in the southern states (Bur- banck and Burbanck 1979). The Saint John River estu- ary and along the north Bay of Fundy shore to the United States border are so far the only known occur- rences of C. polita in Canada, and these populations are located at the northern extent of its range (Burbanck et al. 1979, Atlantic Reference Center, St. Andrews, New Brunswick). We investigated the life history of C. polita in the Saint John River estuary, and compared these data to published observations to explore the hypothesis that northern cyathurans would exhibit differences in life history relative to those at southern latitudes. We suspected that, with probable slower growth in this population, body size of adults would be greater because of delayed maturity and the larger females would carry more embryos. Also, since this is one of the few Canadian populations of an organ- ism which has poor powers of dispersal caused by its non-swimming life style and a brooding reproductive method, it may be important to characterize its life history for future reference under the Species at Risk Act. Finally, the Saint John River estuary is a unique habitat in Canada containing elements of Virginian, coastal marine ecosystems in the summer-warm shal- lows (Bousfield 1962; Burbanck et al. 1979; Dadswell 1979), boreal, marine species in its deep, fjord-like basins (Trites 1960; Carter and Dadswell 1983) along with the wide range of freshwater, diadromous, and marine species typically found in Atlantic Canada (Rogers 1936; Bousfield 1962; Gorham 1965; Gillis 1974). Study Area and Methods Samples of Cyathura polita were collected from the subtidal shore of the lower Saint John River estuary from four stations: Bell Point, Westfield Beach, Sum- merville and Millidgeville (Figure 1). This region is the junction between the Bay of Fundy and the Saint John River and both estuary outflow and tidal inflow are inhibited by a sill at the Reversing Falls. As a result, there are large fluctuations in salinity because of fresh- water discharge, and inside the estuary tidal range is reduced to 0.5 m and summer surface temperatures are higher than in the Bay of Fundy (Metcalfe et al. 1976). The Saint John River has a salt wedge estuary com- bined with deep, fjord-lake basins, and surface salini- ties at the seaward end fluctuate throughout the year, falling to 0% during spring freshet and increasing to 20% during low freshwater flow (Metcalfe et al. 1976). MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA 169 Samples were collected approximately bi-weekly, usually at low tide, from 29 May to 27 August 2004, with an additional collection on 10 October 2004 Samples of C. polita were collected randomly by re- moving the surface substrate from subtidal sites in water depths of 20 to 150 cm. Substrate was collected with a shovel, and sieved through | mm or 0.25 mm mesh. The smaller mesh size was used to capture juve- niles too small to be retained by the larger mesh. Ap- proximately 30 adults and 30 juveniles, if possible, were collected from each location during a sampling effort. Individuals were fixed on the day collected in 10% formalin buffered in sea water and transferred to 70% isopropyl! alcohol within three days. On each col- lection date at each station, the water temperature at 0.5 m depth was measured using a mercury thermome- ter, and a IL sample of surface water was collected for salinity measurements later in the laboratory. Sedi- ment samples from each station (1L) were collected on 17 July 2004. Salinity was measured using an YSI 30 Salinity meter, after calibration with a mercury thermometer. Sediment samples were analyzed for proportions of gravel, sand, and silt, using a sieve series where gravel was <38.]1 mm, sand <2.0 mm, and silt <63 um (Miall 2000). Within a few days of capture, the total length of all individuals was measured to the nearest mm from the base of the antennae to the tip of the telson. Measur- ing the total body length was preferred to estimating size from prosomal length (Bamber 1985), for com- parison with other literature on C. polita. Males were distinguishable by the presence of an appendix mas- culina on the second pleopod; females could be accu- rately distinguished only when with oostegites forming a brood pouch or marsupium (Figure 2). Individuals without these sexual dimorphisms were considered to be juveniles. All gravid females with intact marsupia were ana- lyzed for number and size of embryos. Embryos were removed from marsupia and classified into four stages of development based on electron microscopy (Mer- cer et al., in press): Class 1 embryos were round and contained within two membranes, the chorion and vitelline membranes; Class 2 embryos were elongated and showed differentiation of the three body regions (head, thorax, abdomen), but lacked differentiation of the appendages; Class 3 embryos were still housed within the vitelline membrane, but had considerable differentiation of appendages and head: and Class 4 embryos were hatched from the vitelline membrane and had well-differentiated appendages. Embryos were measured with a compound microscope to the nearest 17 um. Adults and embryos were prepared for scanning electron microscopy (SEM) with primary fixation in 2.5% glutaraldehyde (1 hr), followed by a post-fixation in 1% osmium tetraoxide (4 hrs, both fixes in 0.1M 170 ES Sa, SSS Ei os =redericton OROMOCTO Des Sl. JOIN: ISIMElR ESTUARY (@) 5 10 15 kilometers THE CANADIAN FIELD-NATURALIST Vol. 121 ZN Zi A GAGE TOWN Ef \\\ Washademoak A\ ee ake ) t, € N Q SIRE / EVANDALE 9 yf Belleisle I5°. , d 207 OAK POINT, , 4 Bay ? So cw oe PUBLIC LANDING cope o. COIS) ; BP—w Va Kennebecasis e WEF BEM eos ; Grano ¢ GAS BAY |, NEW V | BRUNSWICK PE! St. John REVERSING U FALLS Ficure |. Map of the Saint John River estuary, with the four study locations indicated. BP = Bell Point; WF = Westfield; SV = Summerville; MV = Millidgeville. cacodylate buffer in seawater). Tissues were dehydrat- ed in an ascending ethanol series, critical point dried using a Bio-Rad E3000 critical point drier with liq- uid CO, and sputter coated with a Technics Hummer II sputter coater. Samples were viewed with a JEOL 5600LV Scanning Electron Microscope at 10KV. Statistics were estimated with MINITAB 14, Mic- rosoft Office Excel 2003, and SPSS 12. Graphs were plotted in Excel and SigmaPlot 9.0. Images were manipulated for size in Corel PhotoPaint 9 and labelled with CorelDraw 9. Results Water temperature, salinity and substrate were com- pared among all stations. For most of the study period water temperature ranged between 15 and 20°C (mean 19°C), with the highest temperature of 28°C on 17 July 2004 (Mercer 2005). Water temperatures at the four stations were similar on each collection date (one-way ANOVA, P = 0.993). Surface salinity ranged from 0.2 ppt in spring to a maximum of 6.6 ppt dur- ing mid-summer (Mercer 2005). However, mean salin- ity differed among the four stations (one-way ANOVA, 2007 MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA 17] FIGURE 2. Scanning electron micrograph of a female Cyathura polita with marsupium, collected from the Saint John River during the summer of 2004. Symbols are: g = gnathopod, m = marsupium, pll= |“ pleopod. 4.5 5 4.0 | 3.9 201 2.9 5 2.0 1.5 1.0 05 0.0 (3) Sex ratio (males:females) (37) 29 May 4 Jun 10 Jun 18 Jun (41) 1 Jul Date (61) (7) ~~ (13) | x =~ | 17 Jul 314 Jul 17 Aug 27 Aug 10 Oct FiGuRE 3. Male to female sex ratio of Cyathura polita during the study period in the Saint John River estuary. Data were pooled from four stations, and sample size (7) is given in brackets above each column. On 10 October, most adult isopods collected were male, resulting in a different scale, thus the sex ratio is given above the bar. P < 0.001). Mean salinities at Bell Point (0.69 + 0.31 ppt) and Westfield (1.5 + 0.86 ppt) were similar (Fisher’s 95% CI of -0.35, 0.78), while Millidgeville (4.7 + 1.3 ppt) and Summerville (4.9 + 1.5 ppt) were comparable (Fisher’s 95% CI of -0.94, 0.19). Sediment composition varied among the four study sites but all generally consisted of easily drained sub- strates (no anoxia; Mercer 2005). Bell Point had the highest gravel content of the four (84.5%), with little sand (15.1%) and hardly any silt (0.4%). Westfield had the highest percentages of sand (66.4%) and silt (7.3%) and the least amount of gravel (26.3%). Mil- lidgeville contained more gravel (55.9%) than sand (43.5%) and very little silt (0.6%). Summerville was similar to Millidgeville with abundant gravel (47.1%) and sand (51.5%) and little silt (1.4%). The sex ratio of C. polita varied during the study period, from a male:female ratio of 2:1 in May, declin- ing to 0.5:1 in July, and increasing to 21:0 in October (Figure 3). In late spring, numbers of males exceeded females and large juveniles (probably maturing fe- males) were present (Figure 4). From early June to the end of August, there were more females than males. Small, free-living juveniles, 2 to 3 mm in length, were first observed on 31 July 2004 numbers of presum- ably recently released from female marsupia. By the end of August, few distinguishable adults were col- lected at any station (Figure 4), but there were many 172 large juveniles (probably adults without secondary sexual characteristics). In October, despite the rarity of males during late August, there was an abundance of relatively large males (Figure 3). Over the entire study, C. polita in the Saint John River estuary were an average of 10.1 + 4.02 mm long, although the mean length varied among stations (Table 1). The mean length of all females collected was 13.8 + 2.14 mm (n = 163) while males averaged 16.3 + 2.41 mm (n = 68). Males were similar in length at all stations, as were females (Table 1; one- way ANOVA). Mean length of individuals collected from Bell Point and Millidgeville over the study peri- od were similar, but isopods from both stations were significantly longer than those from Westfield and Summerville largely because the size of juveniles varied among stations (Table 1; one-way ANOVA). For three stations, mean juvenile length averaged 10.82 + 2.50 mm while juvenile cyathurans from Summerville were smaller (8.60 + 2.49 mm; Table 1). Most C. polita collected at Summerville throughout the study period were juveniles. This fact was high- lighted when comparing the size of juveniles at Sum- merville (8.60 + 2.49 mm in length) to the size of all individuals collected at Summerville throughout the study (8.71 + 2.92 mm). Seasonal changes in body length were estimated by comparing lengths of C. polita from each collec- tion date, after sorting individuals by sex and repro- ductive stage (Figure 4). Since the physical charac- teristics were relatively similar among all stations growth data were pooled to assess life history. Females (with oostegites) were observed from 29 May to 27 August. Mature males were observed throughout the study, with peak abundance in late spring and early fall. Newly-released juveniles were first observed on 31 July and were abundant into the fall. Cohort growth was estimated by comparing the mean length of each group of cyathurans at each col- lection date over the entire study period (Figure 5). Growth was estimated by extrapolating the mean length of each cohort in October back to the mean length of the next cohort in May. These estimates suggest that cyathurans from Saint John have a three-year life cycle, which spans four summer seasons. Young juveniles (Age 0) released from the marsupia at the end of July had a mean body length of 2.7 + 0.25 mm and aver- aged 7.5 mm by October. Young juveniles grew ap- proximately 0.07 mm/day, the fastest of all groups, where growth was represented by: y = 0.068x -11.9 (R* = 0.99). The second cohort of juveniles (Age 1) had a mean length of 8.8 + 3.4 mm in May and grew approximately 0.02mm/day were growth was repre- sented by: y = 0.018x + 6.44 (R?= 0.22), resulting in a mean length in October of 12.4 + 1.90 mm. Mature females (Age 2) were 11.0 mm mean length in May, and grew slowly at a rate of 0.01 mm/day where growth was represented by: y = 0.009x + 11.3 (R* = 0.05) THE CANADIAN FIELD-NATURALIST Vol. 121 15 July 17 = Ao) & =| S 15 August 17 — 10 ro) B 5 2 ® 0 ial 20 4 15 August 27 10 n=139 5 0 20 15 October 10 10 n=78 5 O A 0 5 10 15 20 25 Body length (mm) FIGURE 4. Size distribution of Cyathura polita in the Saint John River estuary for selected sampling dates during the summer of 2004, pooled from the four stations and grouped by sex and reproductive stage. New juveniles, horizontal bars; juveniles, cross-hatching: females, white; gravid female, stippled; males, black. until they reached a mean length of 12.3 + 1.53 mm by the end of August. Females grew little during the summer; however, it was the reproductive period, sug- gesting resources were partitioned into reproduction, not growth. The size distribution of male cyathurans 2007 MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA 173 TABLE |. Body lengths of Cyathura polita in the Saint John River estuary collected over the entire study period (29 May to 10 October 2004). Differences analyzed with a one-way ANOVA and a Scheffé post-hoc comparison among sites Location N Mean length + SD (mm) Minimum (mm) Maximum (mm) All isopods : nitghrdrs ar: Bell Point 247 11.2 + 3.06 ) He) 19.0 Westfield 231 9.2 + 5.00 2.0 21.0 Summerville 297 8.7 + 2.90 2 Mos 19.0 Millidgeville 272 11.5 + 4.20 2.5 24.0 one-way ANOVA df = 3, 1046, f = 35.90, p<0.000 (sites BP=>MV>WF=SV) Males Bell Point 26 15.1 + 1.77 12.0 18.0 Westfield 11 17.3 + 2.57 14.0 21.0 Summerville 3 16.3 + 2.51 14.0 19.0 Millidgeville 28 16.9 + 2.54 13.0 24.0 one-way ANOVA df = 3, 67, f = 3.633 ns Females Bell Point 50 13.4 + 1.87 9.0 19.0 Westfield 34 14.2 + 2.38 8.0 20.0 Summerville 16 12.7 + 2.79 9.0 18.0 Millidgeville 63 14.1+ 1.9] 9.0 17.0 one-way ANOVA df = 3, 162, f = 2.75 ns Juveniles Bell Point 136 10.6 + 2.01 6.0 15.0 Westfield 96 11.4 + 2.69 6.0 18.0 Summerville 262 8.6 + 2.49 4.0 16.0 Millidgeville 149 LOM 2515 7.0 17.0 one-way ANOVA df = 3, 642, f = 43.75, P < 0.000 (sites BP = MV = WF>SYV) 25.0 20.0 @ Males 15.0 O Females = Juveniles 10.0 O New Juveniles 320 Mean body length (mm) 0.0 4 May 23 Jun 12 Aug Oat 20 Nov Date FicurE 5. Mean length of Cyathura polita from the Saint John River estuary pooled from all stations at each sampling interval during the summer of 2004 and grouped by sex and reproductive stage. Vertical bars are standard deviations. 174 was very different from that of juveniles and females. Males (Age 3) were largest at the end of May (mean length 18.5 + 3.54 mm), but smaller in October (mean length 15.1 + 1.95 mm). A bi-modal size distribution during the growing season suggests the die-off of larg- er and older males in early summer, and the addition of new, smaller males in the fall as females underwent protogynic sex change or some large juveniles matured as males. Other studies noted latitudinal variation in body length of C. polita, with individuals from more northern populations being generally larger than in southern populations. However, environmental con- ditions, such as the type of substrate or depth of water, may also strongly affect body size (Burbanck and Burbanck 1979). To assess whether cyathurans from Saint John followed this latitudinal trend, their mean body lengths were compared to values reported in the literature (Table 2). Females pooled from all stations had a mean length of 13.8 + 2.14 mm (maxi- mum: 20.0 mm), and males had a mean length of 16.3 + 2.41 mm (maximum: 24.0 mm). These lengths were similar to cyathurans populations studied north of North Carolina, although none collected at Saint John were as large as the maximum length (27 mm) of the individuals collected from Chewonki Creek, Maine (Burbanck 1959). The individuals collected in the Saint John estuary differed markedly from south- ern populations such as Georgia and Florida: the largest male collected at St. Mark’s, Florida, was only 1.8 mm larger than the smallest male collected in the Saint John (Kruczynski and Subrahmanyam 1978). Broods were examined from 72 females, pooled from all four stations. Fecundity ranged from 16 to 98 embryos per brood, with a mean of 53.2 + 18.9 embryos (Table 3). Abundant gravid females with embryos were first observed on | July. All of these broods contained early embryos (Class 1), with a mean length of 0.45 + 0.06 mm. Two weeks later, more than half of the broods contained elongated Class 2 and 3 embryos, while the remainder contained Class | embryos. By the end of July, 50% of broods contained Class 4 young that had hatched out of their vitelline membranes but were still developing within the marsupia. By mid-August, only six gravid females were found: two had very large Class 1 broods and two had late-stage Class 4 broods. Within the marsupium, embryos grew from a mean length of 0.49 + 0.08 mm as Class 1 embryos to a mean length of 2.07 + 0.16 mm when they were ready to be released as juveniles. Class 2 and 3 embryos had a mean length of 1.05 + 0.15 mm and 1.12 + 0.19 mm, respectively (Table 3). There was no correlation between female length and fecundity (R*= 0.01). There was also no effect of embryos/brood on embryo size (R? = 0.01) and little mortality within broods during development (Mercer 2005). Within-brood survivorship was 96.4% with only a slight decrease observed in mean embryos/brood THE CANADIAN FIELD-NATURALIST Vol. 121 when compared between Class | (57.9 + 18.77 embry- os/brood) and Class 4 broods (55.8 + 19.64 embryos/ brood). Discussion Cyathura polita from the Saint John River estuary exhibits several modifications in life history associat- ed with its northern distribution. In particular, this population exhibits a longer life span, a shorter and later reproductive season, and produces larger adults with higher fecundity, in comparison to southern populations. Additionally, our data suggest that pro- togynic hermaphroditism occurs, as found in some other populations (Burbanck and Burbanck 1974). In the Saint John River estuary, C. polita exhibits a three-year life cycle. Juveniles are released during July and August (Age 0) and, after little growth dur- ing the winter, remain immature the following sum- mer (Age 1). Over the course of their second winter, juveniles develop into mature females (Age 2), which reproduce the following summer. Then, based on our length and sex ratio data, females either die or become males (Age 3) during the late summer-fall, and repro- duce as males during their final summer of life. The result is a three-year life cycle, which spans four sum- mers. In contrast, C. polita from southern populations have shorter life cycles. For example, in St. Mark’s, Florida, C. polita are mature one year after hatching and reproduce throughout their second summer, sug- gesting a life cycle of less than two years (Kruczynski and Subrahmanyam 1978), while on southern Cape Cod, C. polita reproduce in their second, and some- times third, summer, suggesting a two-year life cycle (Burbanck 1962a; Burbanck and Burbanck 1979). Latitudinal changes in the duration and timing of reproduction are also evident. Cyathura polita repro- duce from April to August in St. Mark’s, Florida (Kruczynski and Subrahmanyam 1978), from June to July at Cape Cod, Massachusetts (Burbanck 1962a) and July and August in the Saint John estuary (pre- sent study). Although the breeding season is short and relatively simultaneous in the Saint John popula- tion, females were found with broods at different stages of embryonic development throughout the brooding period, especially in mid- to late-July. The longer life cycle and delayed reproduction in northern cyathurans probably occur because of a shorter growing season. The population in the Saint John lives in water that averages 19.0°C during sum- mer, but shallow water temperatures decline to near O°C from December until April (Metcalfe et al. 1976). The result is limited opportunity for growth and reproduction in warmer waters, resulting in delayed maturation. In contrast, cyathurans from Florida live at temperatures ranging between 13.8° and 33°C, and above 20°C for ten months of the year (Kruczynski and Subrahmanyam 1978). Previous studies suggest that rapid growth in cyathurans occurs only during periods 2007 MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA TABLE 2. Summary of body lengths for C. polita reported in the literature, as means + S.D. or range, depending on the source, Location of study Saint John, New Brunswick SO 16.3+0,29 9 13.8+0.17 Marsh River, Maine GO 14-21 9 14-19 Marshfield, Massachusetts 13.3 + 0.33 Sapelo Island, Georgia 9.3 + 0.64 St. Mark’s, Florida GO 7.3 - 13.8 9 8.3 - 13.0 Body length (mm) Author(s) present study Haefner et al. 1969 Frankenberg 1962 Frankenberg 1962 Krucsynski and Subrahmanyam 1978 TABLE 3. Fecundity and mean embryo length of Cyathura polita in the Saint John River estuary. Embryos were grouped by class of development where: Class | embryos were round and contained in two membranes, the chorion and vitelline, Class 2 embryos were elongated and differentiated into three body regions (head, thorax and abdomen), Class 3 embryos were within vitelline membrane but had differentiated appendages, Class 4 embryos were hatched from the vitelline membrane (Mercer et al. in press). Date Class of Number of collected development gravid females 1 July 15 17 July 13 2 14 3 7) 31 July 1 3 2, l 3 7! 4 11 17 August I 2 2 1 3 | 4 2 of higher water temperature, and that growth ceases when waters cool (Olafsson and Persson 1986). The two-year life cycle described for Cape Cod may be more similar to the southern populations, perhaps because the southern side of Cape Cod, where the study was conducted, is more Virginian in nature than Boreal (Gosner 1971). Cyathura carinata, common along the coast of Europe, exhibits a similar cline in life history traits. Studies of C. carinata in Portugal have suggested rel- atively short life cycles of 19 (Cruz et al. 2003) or 18 to 22 months (Marques et al. 1994). However, three- year life cycles are reported for C. carinata populations in more northern locations such as Poland (Jazdzewski 1969), Sweden (Olafsson and Persson 1986) and Eng- land (Bamber 1985). Olafsson and Persson (1986) sug- gest that C. carinata populations in Sweden require two years before juveniles are able to mature and repro- duce, in contrast to southern populations, which are able to reproduce after one year. Bamber (1985) sug- gests that a longer life cycle is necessary in northern populations as the isopods cannot grow during the win- ter; whereas southern populations are able to grow continuously and attain mature size sooner. Fecundity Total number Embryo length (mm) Mean + SD of embryos Mean + SD 52.8 + 22.48 956 0.45 + 0.06 59.4 + 11.08 658 0.52 + 0.09 47.0 + 18.88 630 1.05 + 0.15 38.3 + 14.21 207 1.10+0.11 52.0 + 22.61 181 0.52 + 0.01 23 1.03 + 0.11 50.5 + 15.34 34] 1.14+0.24 51.3 + 24.18 540 2.06 + 0.16 90.0 + 11.31 180 0.59 + 0.07 4 1.07 + 0.15 37) 1.13 + 0.07 SS Ee ee 67 2.15 +0.18 Cyathura polita in the Saint John River estuary ap- pears to exhibit protogynic hermaphroditism. Sequen- tial hermaphroditism is rare in marine peracaridans, although relatively common in terrestrial and parasitic species (Johnson et al. 2001). Brooks et al. (1994) report that of the approximately 60 species of crus- taceans known to be sequential hermaphrodites, only a few are protogynous, including three species of Cya- thura, three other isopods and five species of tanaids. Evidence for protogyny from the present study includes sex ratio and adult size. Males were abundant in the population in May and again in October, but rare dur- ing summer when the majority of adults were females. Also, males in the fall were similar in length to spent females at the end of the summer (approximately 15 and 12 mm in length, respectively), while most large juveniles were smaller (approximately 10 mm in length). Males continue to grow during the fall and reach a length of approximately 18 mm in the spring of their final year. These observations support a model of protogynic hermaphroditism, whereby mature fe- males become reproductive in early summer of their second year (i.e., third summer) and release juveniles during late July and August. After the release of young, 176 females molt, change sex, and function as mature males in their fourth spring and summer, before death. The hypothesis that C. polita in the Saint John River are protogynic hermaphrodites is consistent with evi- dence of sex change reported from other populations under both laboratory (Burbanck and Burbanck 1974) and field conditions (Kruczynski and Subrahmanyam 1978). While it is possible that males migrate to other areas of the estuary after reproducing and return to shallow waters in the fall, previous work suggests that mostly immature cyathurans occur in the deeper waters of this estuary, while reproductive adults are in shal- lower water (Gillis 1974; Burbanck et al. 1979). Bur- banck and Burbanck (1974, 1979) have reported that some primary males can develop directly from juve- niles, which may account for some of the variation in sex ratios and body lengths observed here. As hypothesized by Bamber (1985), it would be advantageous for male cyathurans to be larger than females to reduce their chances of mortality. Males are able to mate with more than one female, but need to leave their burrows in order to mate, thus they are at a greater risk of predation. Females do not need to expose themselves to that risk, and thus may mature at a smaller size. Warner (1988) additionally suggest- ed that it would be advantageous to become male at a larger size if there is competition among males for females. With a few exceptions, the development of embryos collected at Saint John was synchronized within a brood, which agrees with observations on cyathurans by Stromberg (1972) and Bamber (1985). Our obser- vations suggest that the brooding period for females in the Saint John River estuary lasts from 30 to 45 days. Class 1 embryos were first observed in marsupia on | July. The first hatched embryos (Class 4) were observed on 31 July, which corresponds with the first appear- ance of new juveniles. While comparable observa- tions are not available for more southern populations of this species, a 30 to 45 day brooding period may be typical for this genus. Olafsson and Persson (1986) found that a C. carinata population on the southern coast of Sweden had a three to four week brooding period, while others studying C. carinata in Portugal determined embryonic development to also last 30 to 45 days (Cruz et al. 2003). Brood size is not widely reported for C. polita, how- ever broods collected from Saint John were much larg- er (53.2 + 18.9 embryos/brood) than broods from Flori- da (mean: 14 embryos/ brood, range 2-32; Kruczynski and Subrahmanyam 1978). A similar south-north cline is known in the European C. carinata. Bamber (1985) found that C. carinata females in England carry a mean of 29.8 embryos/brood (range 14-45), while C. carinata females in Sweden averaged 42 + 12 embryos/ brood (Olafsson and Persson 1986). The large brood size in Saint John C. polita is likely a function of increased female size because of greater age (3 vs. 2 THE CANADIAN FIELD-NATURALIST Vol. 121 years). Larger females would likely have larger marsu- pia and therefore may be able to carry more embryos. Cyathura polita at the northernmost extent of its range exhibits several modifications in life history compared to more southern North American popula- tions. These modifications are similar to those observed among European C. carinata populations. Our results support the existence of a latitudinal cline in life his- tory traits in Cyathura, with implications for life span, size and fecundity. Plasticity in reproduction, coupled with protogynic hermaphroditism, makes C. polita an excellent model for studies of the effects of the envi- ronment on life history traits. Cyathura polita is an important indicator of estu- arine health (Burbanck and Burbanck 1979) and since the Saint John River estuary has one of the few popu- lations in Canada there are compelling reasons to understand its life history and population size. We sug- gest a more comprehensive knowledge of the biology of this isopod in the Saint John could assist both with pollution monitoring and with further the distinction of the Saint John estuary as a distinct ecosystem in Canada. Future study of the Saint John cyathurans should be undertaken to verify the 3-year life cycle with sampling throughout the year and to define its estuarine distribution and abundance. Cyathura poli- ta occurs as far inland in the Saint John estuary as Washademoak Lake (Figure 1; Burbanck et al. 1979) and it is a common prey of the Shortnose Sturgeon (Acipenser brevirostrum) another southern species that is endemic to this estuary in Canada (Dadswell O79): Acknowledgments This project was supported by NSERC (Natural Sciences and Engineering Research Council of Canada) through a grant to G. Gibson. The authors are grateful to Acadia University for use of equipment; D. Mac- Donald and G. Cheeseman of the Acadia Centre for Microstructural Analysis; T. Avery for reviews and corrections, Y. Zhang for statistics assistance (Acadia University Statistical Consulting Centre), and R. Mer- cer and C. Mercer for assistance with field work. Literature Cited Bamber, R. N. 1985. The autecology of Cyathura carinata (Crustacea: Isopoda) in a cooling water discharge lagoon. Journal of the Marine Biological Association U.K. 65: 181-194. Bousfield, E.L. 1962. Studies on littoral marine arthropods from the Bay of Fundy region. National Museum of Can- ada Bulletin 183: 42-62. Brooks, H.J., T. A. Rawlings, and R. W. Davies. 1994. Pro- togynous sex change in the intertidal isopod Gnorimos- phaeroma oregonense (Crustacea: Isopoda). Biological Bulletin 187: 99-111. Burbanck, W. D. 1959. The distribution of the estuarine iso- pod, Cyathura sp., along the eastern coast of the United States. Ecology 40: 507-511. 2007 Burbanck, W. D. 1962a. An ecological study of the dis- tribution of the isopod Cyathura polita (Stimpson) from brackish waters of Cape Cod, Massachusetts, American Midland Naturalist 67: 449-476. Burbanck, W. D. 1962b. Further observations on the biotope of the estuarine isopod, Cyathura polita. Ecology 43: 719-722. Burbanck, M. P., and W. D. Burbanck. 1974. Sex reversal of female Cyathura polita (Stimpson, 1855) (Isopoda, Anthuridae). Crustaceana 26: 110-112. Burbanck, W.D, and M. P. Burbanck. 1979. Cyathura (Arthropoda: Crustacea: Isopoda: Anthuridae). Pages 293- 323 in Pollution Ecology of Estuarine Invertebrates. Edited by C. W. Hart, Jr. and S. L. H. Fuller. Academic Press, New York. Burbanck, M. P., W. D. Burbanck, M. J. Dadswell, and G. F. Gillis. 1979. Occurrence and biology of Cyathura polita (Stimpson) (Isopoda, Anthuridae) in Canada. Crustaceana 37: 31-38. Carter, J. C. H., and M. J. Dadswell. 1983. Seasonal and spatial distribution of planktonic crustacea in the lower Saint John River, a multi-basin estuary in New Brunswick, Canada. Estuaries 6: 142-153. Cruz, S., J. C. Marques, S. Gamito, and I. Martins. 2003. Autecology of the isopod, Cyathura carinata (Kroyer, 1847) in the Rio Formosa (Algarve, Portugal). Crustaceana 76: 781-802. Dadswell, M. J. 1979. Biology and population characteristics of the shortnose sturgeon Acipenser brevirostrum LeSueur, 1818 (Osteichthyes: Acipenseridae), in the Saint John River estuary, New Brunswick, Canada. Canadian Journal of Zoology 58: 2186-2210. Frankenberg, D. 1962. A comparison of the physiology and ecology of the estuarine isopod Cyathura polita (Stimpson) in Massachusetts and Georgia. Unpublished Ph.D. thesis, Emory University, Georgia. Gillis, G. F. 1974. An ecological survey in the Saint John estuary. Huntsman Marine Laboratory, St. John River Basin Report 15: 1-64. Gorham, S. W. 1965. Notes on fishes from the Browns Flat area, Kings County, New Brunswick. Canadian Field- Naturalist 79: 137-142. Gosner, K. L. 1971. Guide to identification of marine and estuarine invertebrates. Wiley-Interscience, New York Haefner, J., P. A. M. Mazurkiewicz, and W. D. Burbanck. 1969. Range extension of the North American estuarine isopod crustacean, Cyathura polita (Stimpson, 1855). Crustaceana, 17: 314-317. MERCER, GIBSON, and DADSWELL: LIFE HISTORY OF CYATHURA POLITA 177 Jazdzewski, K. 1969. Biology of two hermaphroditic Crusta- cea, Cyathura carinata (Kroyer) (lsopoda) and Hetero tanais oerstedi (Kroyer) (Tanaidacea) in waters of the Polish Baltic Sea. Zoologica Poloniae 19: 5-25 Johnson, W. S., M. Stevens, and L. Watling. 200). Re production and development of marine peracaridans Advances in Marine Biology 39: 105-260 Kruczynski, W. L., and C. B. Subrahmanyam. 1978 Distribution and breeding cycle of Cyathura polita (\so- poda: Anthuridae) in a Juncus roemerianus marsh of northern Florida. Estuaries |: 93-100. Marques, J. C., I. Martins, C. Teles-Ferreira, and S. Cruz. 1994, Populations dynamics, life history, and production of Cyathura carinata (Kroyer) (Isopoda: Anthuridae) in the Mondego Estuary, Portugal. Journal of Crustacean Biology 14: 258-272. Mercer, S. C. 2005. Life history strategy and morphogenesis of Cyathura polita (Stimpson, 1855). in the Saint John River, New Brunswick, Canada. Unpublished Honours thesis, Acadia University, Wolfville, Nova Scotia. Mercer, S. C., M. J. Dadswell, and G. D. Gibson. /n press. Morphogenesis of embryos and manca in the direct dev- eloping estuarine isopod, Cyathura polita (Anthuridae). Journal of Invertebrate Biology. Metcalfe, C. D., M. J. Dadswell, G. F. Gillis, and M. L. H. Thomas. 1976. Physical, chemical, and biological para- meters of the Saint John River Estuary, New Brunswick, Canada. Fisheries and Marine Service, Research and Dev- elopment Technical Report 686. Miall, A. D. 2000. Principles of sedimentary basin analysis. Springer-Verlag, New York. Olafsson, E. B., and L.-E. Persson. 1986. Distribution, life cycle and demography in a brackish water population of the isopod Cyathura carinata (Kroyer) (Crustacea). Estu- arine, Coastal and Shelf Science 23: 673-687. Stromberg, J-O. 1972. Cyathura polita (Crustacea, Isopoda), some embryological notes. Bulletin of Marine Science 22: 463-482. Rogers, H. M. 1936. The estuary of the Saint John River. Its physiology, ecology, and fisheries. Unpublished MA thesis, University of Toronto, Toronto, Ontario. Trites, R. W. 1960. An oceanographic and biological recon- naissance of Kennebecasis Bay and the Saint John estuary. Journal of the Fisheries Research Board of Canada 17: 377-408. Warner, R. R. 1988. Sex change and the size-advantage model. Trends in Ecology and Evolution 3: 133-136. Received 8 August 2006 Accepted 30 May 2008 Post-Emergence Movements and Overwintering of Snapping Turtle, Chelydra serpentina, Hatchlings in New York and New Hampshire GorRDON R. Uttscu!”, Matt DRAub?, and BARRY WICKLOW* ‘Corresponding author: Courtesy Professor, Department of Zoology, University of Florida, Gainesville, Florida 32611 USA *Mailing address: 4324 NW 36th St., Cape Coral, Florida 33993 USA; e-mail: gultsch@bama.ua.edu *Department of Biology, C. W. Post — Long Island University, 720 Northern Boulevard, Brookville, New York 11547 USA *Department of Biology, St. Anselm College, 100 Saint Anselm Drive, Manchester, New Hampshire 03102 USA Ultsch, Gordon R., Matt Draud, and Barry Wicklow. 2007. Post-emergence movements and overwintering of Snapping Turtle, Chelydra serpentina, hatchlings in New York and New Hampshire. Canadian Field-Naturalist 121(2): 178-181. Hatchling Common Snapping Turtles (Chelydra serpentina) were captured within, or as they emerged from, their nest cavities in Long Island, New York, and in southeastern New Hampshire. They were fitted with radiotransmitters and released at their nest sites. Their movements were monitored for as long as possible, which for some included tracking them to their overwintering sites and relocating them the following spring. On Long Island, all hatchlings initially moved to water. Later movements were both aquatic and terrestrial, and those that could be located while overwintering had left the water and hibernated in spring seeps, where they were recovered alive the following April. In New Hampshire, hatchlings moved directly to nearby aquatic habitats after emergence, where they spent the winter submerged in shallow water in root masses near banks. Key Words: Common Snapping Turtle, Chelydra serpentina, hatchling turtles, hibernation in turtles, New York, New Hampshire. In northern climates, the behavior of hatchling aquatic turtles, once they have pipped and left the eggshell, is variable both among and within species. In some species the majority of hatchlings remain in the nest until the following spring, although some fall emergence may occur. These include Painted Turtles (Chrysemys picta — Zwiefel 1989; Lindeman 1991; Pappas et al. 2000), Common Map Turtles (Grap- temys geographica — Pappas et al. 2000; Baker et al. 2003; Nagle et al. 2004), and Diamondback Terra- pins (Malaclemys terrapin; Baker et al. 2006). For the two former species, it is assumed that if the hatchlings leave the nest, they overwinter aquatically, while for the latter species fall emergence coupled with aquatic or terrestrial overwintering has been reported (Draud et al. 2004), as well as overwintering in the nest cou- pled with fall emergence (Baker et al. 2006). Other species have been consistently reported to be over- whelmingly fall emergers, including Snapping Turtles (Chelydra serpentina — Obbard and Brooks 1981; Car- roll and Ultsch 2007), Spiny Softshell Turtles (Apalone spinifera — Christiansen and Gallaway 1984; Costanzo et al. 1995), Musk Turtles (Sternotherus odoratus — Cagle 1942; Mitchell 1988), Wood Turtles (Glyptemys insculpta — Harding 1991; Carroll and Ultsch 2007), Spotted Turtles (Clemmys guttata — Ernst 1976; Car- roll and Ultsch 2007), and Blanding’s Turtles (Emy- doidea blandingii — Standing et al. 1997, 1999; Pap- pas et al. 2000). Hatchlings of most of these species are assumed to overwinter aquatically, even though some may remain on land for extended periods before entering the water; e.g., Wood Turtles (Tuttle and Carroll 2005; Castellano, Behler, and Ultsch, unpub- lished data) and Blanding’s Turtles (Standing et al. 1997, 1999). However, there are some reports of sus- pected terrestrial overwintering in the hatchlings of some of these species, based largely on the capture of one or a few individuals in spring in microhabitats that are suggestive of non-aquatic overwintering (Ernst 1966, 1976; Standing et al. 1997; Pappas et al. 2000, Parren and Rice 2004). While such behavior does not | appear common, its degree of occurrence is uncertain. There are no reports of following individual hatch- lings from the time of emergence from the nest until the resumption of activity the next spring. The typical approach of a study of overwintering behavior, a num- ber of which have been done with adult turtles (re- viewed by Ultsch 2006), has been the use of radiotele- metry, permitting tracking of the turtles throughout the hibernation period, including entrance and exit. Recently, transmitters small enough to attach to hatch- lings have become available, which allows tracking of the animals from the nest to their wintering grounds, although battery life is not yet long enough to track the turtles until the following spring. However, once the turtles go into hibernation, they move little, which 1 — potentially allows recovery during the early spring: before movement is resumed. Here we report on the post-emergence and over- - wintering behavior of Snapping Turtle (Chelydra ser- - pentina) hatchlings in Long Island, New York, and in| southeastern New Hampshire. In both localities the hatchlings moved to water rather directly, but their eventual overwintering sites differed considerably at’ the two locations. 178 2007 Methods On Long Island, 15 Snapping Turtle hatchlings were collected in the nest on 11 September 2004 from two nest sites (seven from one and eight from the other) in the vicinity of Cold Spring Harbor, Nassau County. The nests were located along an earthen dam separat- ing a lake from the stream formed by the dam over- flow. They were brought to the laboratory, fitted with 0.9 g BD-2 transmitters (Holohil Systems Ltd., Carp, Ontario, Canada) attached to the carapace with cyano- acrylate glue (Krazy Glue”), and released into their respective nest cavities on 13 September 2004. All hatchlings left the nest cavity within 10 min. Telemetry commenced on 14 September, using a Communication Specialists model R1000 receiver. The expected life of the transmitters was 30 days, and some were replaced on day 30. Whenever possible, a visual location was paired with the telemetric location, which was record- ed with a hand-held Magellan GPS 315. Hatchlings were located on nine days from 14-30 September, 10 days from 2-28 October, and on 5 and 12 November. Five hatchlings were lost during the study. Hatchlings that entered final hibernation sites were marked by surveyor flags for recovery, which occurred on 5 April 2005. In New Hampshire, hatchlings were collected from both riverine and palustrine habitats. Six hatchlings were collected from a nest site 20 m from a third-order stream (riverine site) and 18 hatchlings were collected from two adjacent nests 3 m from a first-order stream (palustrine site) within a wetland complex. On 17 Sep- tember 2005 six hatchlings from the riverine site were brought to the laboratory and fitted with 0.5 g A2415 transmitters (Advanced Telemetry Systems [ATS], Inc., Isanti, Minnesota) attached to the carapace with epoxy resin. They were released at the nest site on 19 Sep- tember 2005. Hatchlings were located using an ATS FM-100 receiver and their GPS coordinates recorded using a Trimble Geoexplorer 3. At the palustrine site, nine hatchlings emerging from a nest (22 September 2005) were fitted on site with Holohil transmitters (see above) attached to the carapace using epoxy resin. These hatchlings were then returned to the nest. After a flash flood event during the first week of October 2004, 12 hatchlings were lost. At that time transmitters were attached on site to eight additional hatchlings captured at the site in shallow water. Hibernation sites were marked using surveyor flags. These sites were searched in March 2006. Results On Long Island, 10 hatchlings were followed. The transmitter became detached from three after their movement to water. Two others were found dead. One moved to the water and was found dead on a beach after 26 days. One exhibited considerable movement activity (34-260 m between location periods), includ- ing some terrestrial movements, and by 5 October had ULTSCH, DRAUD, and WICKLOW: OVERWINTERING SNAPPING TURTLE HATCHLINGS 179 moved to a seep, buried in leaves near the surface; it was found dead there on 3 April at a depth of 5 cm. One moved into and out of water and then back into the water, and was located on 12 November dug into sand under a tree root in about 8 cm of water. This animal could not be located the following April. Four hatchlings were tracked to their hibernation quarters and located the following spring. One moved immediately to water, then was found on 2 October under a log 8 m from water. By 10 October it had moved uphill to a spring seep, and two days later had dug to about 20 cm depth and did not move through 12 November. It was found alive and responsive at the same site on the following 3 April in 8 cm of sat- urated mud. A second moved directly to water and then emerged onto land on 30 September about 1.5 m from the water. By 2 October it had moved to a site about 20 m from the water, where it buried under leaves and remained until 8 October. By 10 October it had moved 15 m and was at the surface under cat briar. On 12 October it had moved to a second spring seep, where it buried, moved, and reburied over a 6-day period to depths of 10-13 cm. By 21 October it had relocated in the seep and buried to about 15 cm and had not moved by the last observation on 12 Novem- ber. It was found alive and alert near the surface at the same location on 3 April. A third moved to water, and moved aquatically until 30 September, when it moved into the same seep area as the second, where it moved intermittently, being found buried at depths of 8-13 cm until 18 October, when it settled into the wet organic detritus along the side of a fallen tree, and had not moved by 12 November. It overwintered there and was found alive and well at the surface of the mud at the same site on 3 April. A fourth moved to and about in shallow water for five days. By the sev- enth day it had left the water and buried into moist sphagnum moss about 0.25 m from shore for 3-5 days without moving. It then moved back to water for 2-3 days and then back onto land on 30 September, where it buried for three weeks about 8 cm into the same seep used by the second and third hatchlings. It then moved downward to about 15 cm by 21 October and did not move through 12 November. It was recoy- ered alive at this site on 3 April. In New Hampshire, at the riverine site, 5 of 6 hatch- lings moved first up a gentle slope and then down a steep slope to the river 6 m below. Three of these entered the river within 3 h of release, one after 24 h, and one after 36 h. The sixth moved 8 m on land, down a slope, and burrowed under leaves and then entered the water the next day, all within 24 h of release. The hatchlings remained within 20 m of their entrance points into the river, remaining in shallow (2-3 cm) water, and often buried in sand or detritus with only the head exposed. After extreme flooding caused by heavy rains (25-30 cm in 3 days) in early October, the hatchlings were lost. Three detached transmitters were 180 found, suggesting significant buffeting of the hatch- lings. At the palustrine New Hampshire site, 15 hatchlings (five with and 10 without transmitters) traveled 3-5 m to enter a nearby wetland stream within 4 h of release. Of the four additional hatchlings with transmitters, one burrowed back into the sand of the nest, where it remained for 2 days until dug out by a predator. Two more entered the water within 24 h, and the fourth trav- eled 5 m, remained under wetland vegetation for one day, burrowed into sand one day, then traveled 20 m to shallow water (8 cm) at the edge of the stream. Six hatchlings were lost after a flood during the first week of October. On 11 October 2005, eight additional hatchlings were located along the stream within 5 m of the nest. Each was fitted with a transmitter on site and then returned to the stream. Of these, one moved 15 m downstream into the flooded wetland within 24 h, two moved 12 and 17 m downstream into the wet- land by 16 October 2005. A fourth moved 15 m into the wetland by 2 November 2005. Four hatchlings re- mained tucked into roots of sedges and silky dogwood in 2-3 cm water along the edge of the wetland stream, 3-5 m from the nest. A fifth had remained in this location since 22 September 2004 (visually located on 2 November 2005). Three hatchlings were relocated in March 2006, and they were all at the locations at which they were last observed in November 2005. One was located on 8 March, 6 cm below the surface and 2.5 m from the nest site, under sedge roots at the edge of the stream in 1°C water. A second was found on 8 March, 6 cm below the surface in 1°C water under leaves at the base of a submerged log in the wetland 15 m from its nest. A third was found on 10 March, 7 cm below the surface in 2°C water under silky dog- wood roots at the edge of the stream 2.5 m from its nest. The three hatchlings increased in mass by 12.9 to 19.5% from autumn release to spring capture, but we cannot partition mass gain between possible autumn feeding and potential winter water uptake. Discussion Although the transmitters were small, they still were up to ~10% of the body mass of the hatchlings, and potentially could snag on vegetation during locomo- tion. While we cannot discount effects on behavior, since the hatchlings within a given group behaved sim- ilarly, we do not think that transmitter attachment had a major effect on behavior relative to that of an unfet- tered hatchling. On Long Island, the four hatchlings that were recov- ered from their hibernacula in April all overwintered in spring seeps away from standing water, in contrast to the behavior of the New Hampshire hatchlings, which all moved to shallow water and remained there to overwinter. The four Long Island hatchlings also initially moved to open water from their release in the nest cavity (two from each nest), but they then followed THE CANADIAN FIELD-NATURALIST Vol. 121 different paths, sometimes including both aquatic and terrestrial movements, before eventually settling into the spring seeps for overwintering, as did the fifth, although we could not find it in April. In the sense that Long Island hibernacula were not under standing water, the hibernacula were terrestrial and differed from those expected by assuming that hatchlings move to aquatic lentic or lotic habitats and overwinter there, as was the case with the New Hampshire hatchlings. However, the Long Island hatchlings were surround- ed by a constantly saturated medium that appeared to have an underground flow, so desiccation was not a threat. We do not know the oxygen content of the interstitial water, but assume that it was appreciable or the turtles would have died if they had remained submerged throughout the winter. Reese et al. (2004) found that Snapping Turtle hatchlings will die in 30 days if submerged in anoxic water at 3°C, compared to survival of adults for at least 125 days under simi- lar conditions (Reese at al. 2002). The limited survival of anoxic hatchlings is attributable to their reduced whole-body buffering capacity, and therefore reduced tolerance of the metabolic acidosis that accompanies the accumulation of lactic acid during anaerobiosis. The reduced buffering capacity is due to the reduced relative amount of bone, especially in the shell, of the hatchlings compared to adults, and is a general char- acteristic of hatchling turtles (Reese et al. 2004). This inability to tolerate anoxia would preclude the turtles from hibernating for extended periods in anoxic con- ditions, such as mud under standing water. Thus the seep water that surrounded the hatchlings must have had a significant amount of dissolved oxygen, perhaps maintained by a constant percolation of oxygenated spring water that exceeded the biochemical oxygen demand of the detritus at the low wintertime temper- atures. Alternatively, it is possible that the hatchlings may have occasionally air-breathed, since the soil sur- face of the seeps did not freeze during the winter. Air- breathing would require that the animals move to the surface to breathe (on the assumption that the water saturation prevents any underground air pockets from forming); since turtles are not completely immobilized by cold, air-breathing cannot be ruled out. Why the hatchlings moved out of open water to hibernate in the seeps, or how they oriented toward them, is uncertain. Thus the overwintering habitats of hatchling Snap- ping Turtles are varied, ranging from the expected shallow-water retreats typical of the New Hampshire hatchlings, to the use of wet but not aquatic sites (e.g., seeps), even though aquatic sites are nearby and easily accessible, as seen with the Long Island hatchlings. The use of seeps as hibernacula for turtles, especially young ones, may be more common than has been ap- preciated. V. Lamoureaux (personal communication), while studying the use of seeps as hibernacula for Green Frogs (Rana clamitans) in New York (Lam- oureux and Madison 1999), found one adult and three 2007 juvenile (9.5 — 11 cm) Snapping Turtles leaving a seep area in late April, where they apparently overwintered, and moving toward a nearby pond. Dunson (1986) radiotracked four snapping turtles in a tidal marsh area in Virginia, and they all moved to seep/spring areas on higher ground, which remained unfrozen throughout the winter, suggesting that the use of seeps as hibernac- ula may not be limited to immature turtles, although almost all reports of hibernacula of adult Snapping Turtles are of those covered by standing or flowing water (Ultsch 2006). Acknowledgments We thank Sara Zimnavoda for help with radiotrack- ing in Long Island and students of the Aquatic Ecology course at St. Anselm College for similar aid in New Hampshire. This study was supported by a National Science Foundation grant to GRU (IBN 0076592). Literature Cited Baker, P. J., J. P. Costanzo, J. B. Iverson, and R. E. Lee, Jr. 2003. Adaptations to terrestrial overwintering of hatchling northern map turtles, Graptemys geographica. Journal of Comparative Physiology B 173: 643-651. Baker, P. J., J. P. Costanzo, R. Herlands, R. C. Woods, and R. E. Lee, Jr. 2006. Inoculative freezing promotes winter survival in hatchling diamondback terrapin, Mala- clemys terrapin. Canadian Journal of Zoology 84: 116-124. Cagle, F. R. 1942. Herpetological fauna of Jackson and Union Counties, Illinois. American Midland Naturalist 28: 164- 200. Carroll, D. M., and G. R. Ultsch. 2007. Emergence season and survival in the nest of hatchling turtles in southcentral New Hampshire. Northeastern Naturalist 14: 307-310. Christiansen, J. L., and B. J. Gallaway. 1984. 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Lactate accumulation, glycogen depletion, and shell composition of hatchling turtles during simulated hibernation. Journal of Experimental Biology 207: 2889-2895. Standing, K. L., T. B. Herman, D. D. Hurlburt, and I. P. Morrison. 1997. Postemergence behaviour of neonates in a northern peripheral population of Blanding’s turtle. Emydoidea blandingii, in Nova Scotia. Canadian Journal of Zoology 75: 1387-1395. Standing, K. L., T. B. Herman, and I. P. Morrison. 1999. Nesting ecology of Blanding’s turtle (Emydoidea blan- dingii) in Nova Scotia, the northeastern limit of the species” range. Canadian Journal of Zoology 77: 1609-1614. Tuttle, S. E., and D. M. Carroll. 2005. Movements and be- havior of hatchling wood turtles (G/yptemys insculpta). Northeastern Naturalist 12: 331-248. Ultsch, G. R. 2006. The ecology of overwintering among tur- tles: Where turtles overwinter and its consequences. Bio- logical Reviews 81: 339-367. Zwiefel, R. G. 1989. Long-term ecological studies on a pop- ulation of painted turtles, Chrysemys picta, on Long Island, New York. American Museum Novitates 2952: 1-55. Received 27 October 2006 Accepted 29 January 2008 Notes on Succession in Old Fields in Southeastern Ontario: the Herbs A. CROWDER!, R. HARMSEN!, and S. E. BLATT? 'Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6 Canada 26 Wallace Place, Wolfville, Nova Scotia B4P 2R6 Canada 3Corresponding author Crowder, A., R. Harmsen, and S. E. Blatt. 2007. Notes on succession in old fields in southeastern Ontario: the herbs. Canadian Field-Naturalist 121(2): 182-190. Vegetation in abandoned hayfields was monitored during 1976-1998. An earlier successional stage followed ploughing. Changes in tree, shrub and vine populations have been reported earlier and showed expected increases in species richness and cover. Highest species richness of herbs occurred three years after ploughing. Non-woody species richness trended irregularly downward, while non-woody cover was variable, peaking in 1987. Within the herbaceous community, year-to- year changes in cover and frequency of species in the following selected groups are reported here: 18 grasses including sown and adventive species; 13 legumes including two sown species; 14 macroforbs of the Compositae, including a goldenrod, Solidago canadensis, which dominated parts of the fields; a rosette weed, Taraxacum officinalis; sedges, horsetails and some other minor components. Grasses and goldenrods were grazed, sometimes intensively and repeatedly, by insects; grasses were impacted by skipper larvae (Thymelicus lineola), and goldenrods by beetle larvae (Trirhabda spp.). Effects of repeated outbreaks on host plant cover are shown for two plots (100 m2) matching the scale of outbreaks. Key Words: old field herbs, phytophagous insects, plant succession, goldenrods, grasses, legumes, Ontario. Two abandoned hayfields in southeastern Ontario were selected as sites in which to investigate possible effects of insect herbivory on early stages of secondary succession. The location, geology, history of land use, and soils have been described earlier (Crowder and Harmsen 1998). Their nutrient status was discussed by Epp and Aarssen (1988). Estimates of increasing cover and abundance of trees, shrubs and vines (51 species) during 1976-1998 were shown to follow classic succes- sional models, although with unusual pioneer species. After two decades the structure of the vegetation had become heterogeneous, with groves of small trees, shrub thickets, patches dominated by macroforbs and patches still dominated by grasses. Cover varied with fluctuations of weather from year to year, while soil differences influenced cover and species richness. Most successional studies on old fields use synoptic records from fields of different ages, a method which obscures minor fluctuations and differences between fields. In this paper, year-to-year observations are used to follow temporal changes in the herbaceous com- munity and to relate its cover and species richness to those of the woody layers. Herbaceous species found in 27 experimental plots are listed to allow comparisons with other areas; lichens and bryophytes are omitted. Differences in soil texture (Crowder and Harmsen 1998) influenced cover and richness. Within the herbaceous community, details of tem- poral change in selected groups of plants and some specific interactions are reported here. Grasses were the main constituents of the hayfields, including both agricultural and adventives species. Legumes also in- cluded sown and invasive species. Invasive macroforbs, 182 mainly Solidago and Aster species, dominated parts of the fields; population changes of 15 species are reported. Taraxacum officinalis, Dandelion, was selected to rep- resent rosette plants. Sedges, horsetails and a few other minor components are described. During the decades when vegetation was monitored, arthropods were also sampled (McBrien et al. 1983). Two insect populations had repetitive outbreaks which caused significant decreases of plant cover, and are therefore the only arthropods reported here. Larvae of © the hesperiid European Skipper (Thymelicus lineola) grazed on grasses, particularly on Timothy, Phleum | pratense (Henderson 1986). Larvae of beetles of the genus Trirhabda Le Conte (Coleoptera, Chrysomeli- | dae), mostly Trirhabda virgata, grazed primarily on leaves of Solidago canadensis, causing intense defo- liation. The strong effect of one or more outbreaks of beetles on plant vigour was replicated experimentally (McBrien and Harmsen 1987). These defoliations can cause changes in successional pathways at the grass- macroforb stage by affecting the relative dominance * of grass or goldenrods. Blatt et al. (2001) have used | these data to develop a successional model including : herbivore effects, showing possible outcomes for the ° herbs. Methods A hayfield abandoned in 1970 and one mown for hay until 1975 were selected at Queen’s University | Biology Station at Opinicon, Frontenac County, On- tario (44°03'N, 76°0'W). Nine permanent plots, each | 100 m?, were demarcated in 1975 in the older field, which was called S field, for Solidago. Eighteen similar \ fi 2007 plots were placed in the younger field, and nine of these were ploughed once; these ploughed plots were called P and the undisturbed plots G, for grass. All plant species present in the 27 plots were listed in June and September of sampling years: 1976-1988, 1995, 1998. Cover was estimated using five randomly placed quadrats within each plot; cover values used in this paper were from September (McBrien et al. 1983; Henderson 1986). Each quadrat was 50 cm x 50 cm in area. Values were compared using Scheffe’s mean separation test (Zar 1984; Blatt et al. 2003). Percentage frequency, the number of quadrats in which a species was recorded at a particular time (Mellinger 1972; Eriksson and Jakobssen 1998), was used as a measure of relative abundance (Maycock and Guzikowa 1984). Nomenclature follows Newmaster et al. (1998); voucher specimens were lodged at the Fowler Herba- rium, Queen’s University. Lichens, mosses and liver- worts have been omitted from this paper. Results and Discussion Species richness of woody species and of herbs per 100 m? plot, through 1976-1998, is shown in Figure 1. Note gaps in sampling after 1988. Herb richness trended downward with high annual variance while woody spe- cies richness increased over time (see Crowder and Harmsen 1998). The range of species richness was 7-65 in ploughed (P) plots, 6-42 in grass-dominated (G) plots, and 6-33 in Solidago (S) plots. Species richness of herbs and of woody plants in a grass plot (G1) and a ploughed plot (P2) are shown in Figures 2 and 3. Because soil texture was found to be correlated with species richness, the selected plots were matched for texture; both are sandy, with relatively low richness (Crowder and Harmsen 1998). Note the difference in scale between Figures 2B and 3B. Species richness peaked in P2 three years after ploughing when several weedy species were pre- sent (cf. Armesto and Pickett 1985). Herbaceous taxa observed in 27 plots during 1976- 1998 are listed in Table 1, in alphabetical order, with English names added (Newmaster et al. 1998). The closest comparable study site is a 50-year-old field, at Erindale (Toronto area), described by Maycock and Guzikowa (1984), in which 93% of the species listed at Opinicon were found. Other old fields in Ontario have been described by Lamoureux (1970), Reader (1990) and Reader and Buck (1991). Fewer of the Opinicon species (37%) were found in abandoned hayfields in New York State described by Mellinger (1972), and only 20% in old fields in Quebec (Dan- sereau and Gille 1949). The proportion of introduced species at Opinicon was 37%, which is higher than the average for the Kingston region (Beschel et al. 1970). They included both crop plants and weeds. During 1976-1998 the cover value of woody plants for all plots combined increased significantly (Crowder and Harmsen 1998): herbaceous cover, however, did CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 183 not decrease significantly over time (Figure 1). Mean percent cover of non-woody species in 1998 was not significantly different from that in 1988, 1986, 1984, 1978, 1977 and 1976. Years with high cover were 1979, 1980, 1981, 1985, or 1987 (Figure 1). These values suggest that herbaceous cover was related to year-to- year changes in weather or herbivory rather than to a steady increase in shading. No correlations of cover values with available climatic data were found, al- though the years with low cover had partial droughts that caused soil cracking. Herb cover in selected plots is shown in Figures 2A and 3A, with relatively low variance between years. Plot G1 consistently had higher values than P2. The main components of herb cover are discussed in the following section. The hayfields had been sown with Phleum pratense (Timothy) and Poa pratensis (Ken- tucky Blue Grass), which maintained their frequency after the cessation of mowing (see Table 2). Mellinger (1972) also reported their high frequency in old hay- fields in New York. In the Opinicon fields the two spe- cies are associated randomly (Epp and Aarssen 1988). Table 2 lists annual percentage frequency of nine grass species. The most frequent invasive grasses have been Elymus repens, Poa compressa and Calamagrostis canadensis. Other less common invasive grasses, not shown on Table 2, were Bromus inermis ssp. pum- pellianus, Echinochloa crusgalli, Elymus trachycaulon ssp. trachycaulon, Muhlenbergia mexicana, Panicum acuminatum, Poa capillare, Poa annua, Setaria pumila and Setaria viridis. Annual cover of all grass species in all plots varied from a mean of 35% in 1998 to a mean of 87% in 1987. Cover in the G (grass) plots ex- ceeded 100% in some years, because layers of two or more species developed. Differences between the three sets of plots were not consistent. Cover of grasses did not show synchronous trends within sets of plots. Changes in grass cover were not correlated with soil texture; the two plots shown in Figures 2 and 3 were matched for soil texture but the undisturbed plot G1 in Figure 2 had high grass cover and the ploughed plot in Figure 3 had not. A temporal pattern in the grass-dominated plot showed successive peaks three to five years apart followed by declines in cover values. This pattern was typical of grass- dominated plots. The disturbed ploughed plot shown in Figure 3 had two less marked peaks and declines. Herbivory was demonstrated by Henderson (1986) to reduce grass cover significantly; he counted larvae of a hesperiid European Skipper (Thymelicus lineola) and found that peaks of larval numbers coincided with low cover of Elymus repens, Phleum pratense and Poa pratensis, and low concentrations of skipper larvae with high grass cover. The grass species differed in their capacity to recover after an insect outbreak, with Phleum pratense being most negatively affected. Out- break behaviour by the European Skipper has been documented by Pengelly (1961) elsewhere in Ontario. 184 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Alphabetic list of herbs recorded within 27 plots during 1975-1998 at two old fields at Opinicon, with English names. Nomenclature follows Newmaster et al. (1998). Acalypha virginica. Copperleaf Achillea millefolium ssp. millefolium. Yarrow Agrimonia gryposepala. Hooked Agrimony Agrostis gigantea. Redtop Ambrosia artemisiifolia. Common Ragweed Anaphalis margaritacea. Pearly Everlasting Anemone canadensis. Canada Anemone Antennaria neglecta. Field Pussytoes Antennaria parlinii. Plantain-leaved Pussytoes Arctium minus ssp. minus. Common Burdock Artemisia absinthium. Absinth Asclepias syriaca. Common Milkweed Aster cordifolius. Heart-leaved Aster Aster ericoides. Heath Aster Aster lanceolatus. Panicled Aster (includes Aster simplex) Aster lateriflorus. Calico Aster Aster umbellatus. Flat-topped White Aster Aster urophyllus. Arrow-leaved Aster Barbarea vulgaris. Wintercress Bromus inermis ssp. pumpellianus. Smooth Brome Calamagrostis canadensis. Canada Blue Joint Capsella bursa-pastoris. Shepherd’s Purse Carex aurea. Sedge Carex communis. Common Sedge Carex debilis. Sedge Carex foenea. Sedge Carex gracillima. Filiform Sedge Carex granularis. Granular Sedge Carex laxiflora. Slender Sedge Carex normalis. Normal Sedge Carex scoparia. Sedge Carex vulpinoidea. Fox Sedge Cerastium fontanum ssp. triviale. Common Mouse-ear Chickweed Chenopodium simplex. Maple-leaved Goosefoot Chrysanthemum leucanthemum. Ox-eye Daisy Cichorium intybus. Chicory Cirsium arvense. Canada Thistle Clinopodium vulgare. Wild Basil Convolvulus arvensis. Field Bindweed Conyza canadensis. Canada Fleabane Dactylis glomerata. Orchard Grass Danthonia spicata. Poverty Oat Grass Daucus carota. Wild Carrot Dianthus armeria. Deptford Pink Echinochloa crusgalli. Echinochloa Echium vulgare. Blueweed Elymus repens. Couchgrass Elymus trachycaulus ssp. trachycaulus. Slender Wheat Grass Epilobium hirsutum. Hairy Willow Herb Equisetum arvense. Field Horsetail Equisetum hyemale. Scouring Rush Erigeron annuus. Daisy Fleabane Erigeron philadelphicus ssp. philadelphicus. Common Fleabane Erigeron strigosus. Lesser Fleabane Eupatorium maculatum. Joe Pye Weed Eupatorium perfoliatum. Boneset Eupatorium rugosum. White Snakeroot Euthamia graminifolia. Grass-leaved Goldenrod Fragaria vesca ssp. americana. Wood Strawberry Fragaria virginiana. Wild Strawberry Galinsoga quadriradiata. Quickweed Galium obtusum. Wild Madder Galium trifidum ssp. trifidum. Small Bedstraw Gentiana andrewsii. Bottle Gentian Geranium bicknellii. Bicknell’s Crane’s Bill Geum aleppicum. Yellow Avens Geum canadense. White Avens Hieracium aurantiacum. Orange Hawkweed Hieracium piloselloides. King Devil Hypericum perforatum. Common St. John’s Wort Juncus dudleyi. Dudley’s Rush Lactuca canadensis. Wild Lettuce Lepidium campestre. Field Peppergrass Lithospermum officinale. Gromwell Lobelia inflata. Indian Tobacco Lycopus americanus. Cut-leaved Water Horehound Lycopus uniflorus. Water Horehound Lysimachia terrestris. Yellow Loosestrife Medicago lupulina. Black Medick Medicago sativa ssp. sativa. Alfalfa Medicago sativa ssp. falcata. Medick Melilotus alba. White Sweet Clover Melilotus officinalis. Yellow Sweet Clover Mentha arvensis ssp. borealis. Wild Mint Muhlenbergia mexicana. Dropseed Oenothera perennis. Sundrops Onoclea sensibilis. Sensitive Fern Osmunda regalis. Royal Fern Oxalis stricta. Yellow Wood Sorrel Panicum acuminatum. Woolly Panic Grass Panicum capillare. Witch Grass Panicum depauperatum. Starved Panic Grass Phleum pratense. Timothy Physalis heterophylla. Clammy Ground Cherry Plantago lanceolata. Narrow-leaved Plantain Plantago major. Common Plantain Plantago rugelii. Rugel’s Plantain Poa annua. Annual Blue Grass Poa compressa. Canada Blue Grass Poa pratensis ssp. pratensis. Kentucky Blue Grass Polygonum convolvulus. Black Bindweed Polygonum persicaria. Lady’s Thumb Polygonum sagittatum Arrowleaved Tear-Thumb Potentilla argentea. Silvery Cinquefoil Potentilla canadensis. Old Field Cinquefoil Potentilla inclinata. Downy Cinquefoil Potentilla norvegica. Rough Cinquefoil Potentilla recta. Erect Cinquefoil Prunella vulgaris ssp. vulgaris. Selfheal Ranunculus abortivus. Kidney-leaf Buttercup Ranunculus acris. Field Buttercup Rudbeckia hirta. Black-eyed Susan Rumex acetosella ssp. acetosella. Sheep Sorrel Rumex crispus. Curly Dock Sanicula marilandica. Black Snakeroot Setaria pumila. Yellow Foxtail Setaria. viridis. Green Foxtail Silene vulgaris. Bladder Campion Sisymbrium officinale. Hedge-Mustard Sisyrinchium angustifolium. Blue-eyed Grass Solidago caesia. Blue-stemmed Goldenrod 2007 TABLE |. (continued) Solidago canadensis. Canada Goldenrod Solidago gigantea. Yall Goldenrod Solidago juncea. Early Goldenrod Solidago nemoralis ssp. nemoralis. Gray Goldenrod Solidago rugosa. Rough-stemmed Goldenrod Sonchus asper ssp. asper. Spiny Sow-thistle Spiranthes romanzoffiana. Hooded Ladies’ Tresses Stellaria borealis ssp. borealis. Northern Stitchwort Stellaria graminea. Stitchwort Stellaria media, Chickweed Taraxacum officinale. Dandelion Teucrium canadense ssp. canadense. Germander Thlaspi arvense. Field Pennycress Larval outbreaks are considered to affect the succes- sional pathway in the Opinicon fields in a model developed from our data by Blatt et al. (2001). When insect herbivory on grasses in old fields was studied by Brown et al. (1988), Elymus repens was the grass species most affected by Homoptera/Auchenorrhycha; these authors considered that both texture and struc- ture of the successional vegetation were affected by the insects. The initial seed mix contained Medicago sativa ssp. sativa (Alfalfa) and possibly Trifolium pratense (Red Clover) which have persisted in the fields. Medicago sativa frequency was 3-5% in 1976-1980, rose to 12% in 1983 and fell again to 2% in the 1990s. Trifolium pratense frequency was 35% in 1976, fell to less than 8% during 1980-1995 and rose again to 22% in 1998. Trifolium aureum, Trifolium campestre and Trifolium hybridum were recorded occasionally. Vetches (Vicia sativa ssp. nigra, Vicia cracca, and Vicia tetrasperma) became more frequent in later years with a maximal frequency of 69% for Vicia cracca in 1987. Possibly these straggling plants benefited from the cessation of mowing. Other legumes recorded were Melilotus alba, Melilotus officinalis and Medicago lupulinus, randomly distributed except for Medicago lupulina, which was most frequent in 1976-1979 in the ploughed plots. Macroforb areas of the fields have been and are dominated by Solidago canadensis (Canada Golden- rod), which we observed to spread by both seeds and rhizomes. Its importance in old field succession in North America has been widely reported (e.g., Dale et al. 1966; Goldberg and Gross 1988). Percentage fre- quency of Solidago canadensis in all plots during 1976-1998 is shown as mean values in Table 2, start- ing with minimal values in 1976 and reaching a maxi- mum in 1987-1988. Frequencies of Solidago nemoralis (Old Field Goldenrod), Solidago caesia (Blue-stemmed Goldenrod), and Euthamia graminifolia (Grass-leaved Goldenrod), also given in Table 2, were higher in the 1980s than in the 1970s or 1990s. When the three sets of plots were compared, Solidago canadensis and Soli- CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 185 Tragopogon dubius. Goatsbeard Tragopogon pratensis ssp. pratensis. Showy Goatsbeard Trifolium aureum. Yellow Clover Trifolium campestre. Smaller Hop Clover Trifolium hybridum. Alsike Clover Trifolium pratense. Red Clover Verbascum thapsus. Common Mullein Verbena hastata. Blue Vervain Veronica serpyllifolia ssp. serpyllifolia. Thyme-leaved Speedwell Vicia cracca. Cow Vetch Vicia sativa ssp. nigra. Common Vetch Vicia tetrasperma. Slender Vetch Viola sp. Violet dago caesia were found to be most frequent in the S$ plots (during 1976-1998 the mean frequency of Solli- dago canadensis in S plots was 47%, in the P plots 37% and in the G plots 26%). Solidago nemoralis and Solidago juncea were generally more frequent in sandy dry plots. This distribution may be explained by dif- ferent soil moisture tolerances among the congeners (Werner and Platt 1976). Cover values for Solidago canadensis ranged from 1 to 100%. Patterns were not synchronous between plots. Two examples are shown. Figure 3 shows Plot P2, with three irregular peaks of goldenrod cover, and Figure 2 shows Plot G1 with only three years of spor- adic goldenrod cover. The plot with three peaks and lows (Figure 3) had a periodicity also observed in other plots, while others had one, two, or three-and-a-half episodes. Some sandy plots similar to that shown in Figure 2 failed to develop dense Solidago cover, and did not have a series of highs and lows. We have found that the cover of Solidago canadensis was negatively and significantly correlated with larval density of three species of Trirhabda beetles in the field (McBrien et al. 1983; Blatt et al. 2001). Trirhabda vir- gata was most abundant. Field experiments showed that effects on biomass and cover similar to those caused by larval grazing could be produced by manual defoliation (McBrien and Harmsen 1987). Within a field, beetle outbreaks were not synchro- nous between plots. Outbreaks in the two fields also occurred at different times. In the P/G field, years with several plot peaks of Solidago cover were 1980, 1981, and 1985, and years with most lows were 1976, 1985 and 1998. That both peaks and lows occurred in 1985 shows the small scale of the beetle outbreaks, some- times confined to a plot 100 m?. Outbreaks on a much larger scale have been reported, for example, on a scale of several hectares (Werner et al. 1980), and over two hundred insect species have been reported on Solidago canadensis. Macroforbs growing to the same size as some gold- enrods included the asters Aster umbellatus (Flat- topped White Aster) and Aster novae-angliae (New 186 THE CANADIAN FIELD-NATURALIST Vol. 121 Mean non-woody species nchness 30 25 finan By pie a a \ es v4 \ ve 20 15 10 AD abca be ia i defg cto bedef'” A” > fb 5 ao & es “st S&S Ss PSH SF SF PS & SS S Year [___] Mean percent cover of non-woody species, all plots ber A...... Mean percent cover of woody species, all plots Ficure |. Comparison of woody and non-woody vegetation. Mean species richness of herbaceous plants in 27 plots (species number/100 m7) for the years 1976-1988, 1995 and 1998 is shown by dots. Mean percent cover of non-woody species in all plots in September is shown in the histogram; means which do not share the same letter are significantly different. Mean percent cover of woody species in all plots is indicated by a dotted line with triangles. England Aster). During the years of observation the percentage frequency of these two species increased (Table 2). Three smaller species, Aster lanceolatus (in- cluded in Aster simplex in Table 2), Aster lateriflorus and Aster ontarionis, have been lumped together be- cause they were generally intermingled in straggling clumps which also increased in frequency. Aster cordifolius (Heart-leaved Aster) and Aster urophyllus (Arrow-leaved Aster), both of which are shade-tolerant (Semple et al. 1996), had low frequency. Five of the aster species found in the Opinicon fields were re- corded by Maycock and Guzikowa (1984). When all aster species are lumped together, mean annual cover ranged from | to 45%. Comparing dif- 2007 CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 187 ferent plots, patterns of peaks and lows like those ob- a bh ns ty my fe a served in Solidago canadensis occurred in some plots, = MANM=OOVVSTOMOVONATO but with a longer periodicity ranging up to ten years. ats Since these fluctuations in cover were not synchronous ES o> ee ce a, oe a, i a between plots or fields, it is possible that they were due to herbivory, like the changes in Solidago and grasses. Two species of horsetail, Equisetum arvense and Equisetum hyemale, were present in the P/G field throughout 1976-1998 with a combined mean per- 1988 4 0 6 ] 5 5 4 0 % ie, alt ES bo os a centage frequency of 68%. Both species were most Bre SS EN ot eovwrer N'| frequent in’ satidy plots and) are locally. common in disturbed habitats such as sandpits (Crowder et al. % mons_s = > ~ 1997). Cover did not exceed 29% for the two species ee OVS VN VIN O ONO Om ST combined in any yearnand was,eenerally low: ss Mean frequency of Jaraxacum officinale ranged oe etd tase er ve es os from 13% in 1981 to 37% in 1988. Dandelions were mE en VV VO VONNANANS™ HAOTAOSH resent in all three sets of plots every year, with highest frequency in the S and lowest in the G plots. The x rosette growth form of Taraxacum officinale is similar FIFA ROR CVV OMS +n CoH ammAc]| to that of many genera in the plots, such as Hieraciwn spp. and Antennaria parlinii, and probably facilitates eS early spring growth. Rloann = oA _ ~ Sedges, Carex spp., were observed in all years and AS WOarIANNOCSOVOOCOTAONODOO VANMOMNTNS : > m in all sets of plots, but were not analysed because of Q difficulty in distinguishing non-flowering stages. Carex DPT oon voted Vornono|| granularis, Carex laxiflora and Carex normalis were relatively abundant. Mean annual frequency of all spe- a Bil, es S cies was 39% and most plants were found in plots with Nemo ms ootast =n vyenee*e™2 high moisture content (Crowder and Harmsen 1998). Mean cover was less than 2% until 1979, rose to 7% SSBmooVVOFate+Voemnac]| in 1987, and then declined again. Species which increased in both frequency and Lis Seal bs a cover during 1976-1998 included Asclepias syriaca SEAS SUG SO OO OS SOO ANd Sil!’ (Milkweed) and Ranunculus acns (Field Buttercup). Species which declined in both frequency and cover Sea Of MAO VOkmm unl aecluded (Rumex acetosella ssp. acetosella (Sheep’s Sorrel), Ambrosia artemisiifolia (Ragweed), Oxalis stricta (Yellow Wood Sorrel), and Prunella vulgaris Bae ealtosg Veotoauaacoaaw sal ssp. lanceolata (Self-Heal). Rianex acetosella and Ambrosia artemisiifolia, which are weedy species, were mostly in the ploughed plots, whereas Prunella 1981 ay 52 1979 1980 1978 1977 TABLE 2. Mean percentage frequency of selected herbs in all plots during 1976-1988, 1995 and 1998 (n = 162) S Recien a = a vulgaris was initially concentrated in grass plots. A ~ PPrepoomomomona = Vom ~~2 1 minor component with low cover value but high fre- quency (40-50%) was Potentilla recta (Erect Cin- 8 quefoil). s When grass cover was lowered by grazing of = Thymelicus lineola, feeding preferentially on Phleum S pratense, cover of Solidago and Aster spp. in both aS fields increased rapidly; they then maintained their Be dominance at levels equal to that of Poa pratensis, 2 S§ preventing the recovery of Phlewm pratense (Hen- S§s s a derson 1986; Blatt et al. 2001; Blatt et al. 2003). In SSs s See 8 = aS addition to interactions between grasses and golden- 2 S as SSSEs . Ses $< _ || rods, other layers were affected. For example, when 28 S oes S Ss Sos ue SS Sse S|} the cover of Solidago canadensis was decreased by . SS LSS = S < : ge SSS SPSS the feeding of beetle larvae a chamaephyte. Fragaria Sis Sg E Sas Bes YS &s £3 S|] virginiana (Wild Strawberry). showed significantly a ls8 = SSEE S ss = = = = =S 3 $2 $]] increased cover (McBrien et al. 1983). This finding FIRQAHQTOCAACHAAANHAHAAXTIICK 188 THE CANADIAN FIELD-NATURALIST Vol. 121 30 A. Non-woody species 25 20 15 / ~ | a ~ a 7 ° ie : Non-woody species nchness oOo WA a B. Woody species 50 C. Grass species 3 2 D. Solidago species oO Oo Mean percent (+ SE) cover or species richness i=) WD SS OO A S > , & “te, 9p N FIGURE 2. Conditions in grass plot G1 during 1976-1988, 1995 and 1998. 2A shows mean percent cover of non-woody species, and non-woody species richness (species number/100 m7). 2B shows cover and species richness of woody plants. 2C shows mean cover of all grass species. 2D shows Solidago species cover, which was only present in 1986, 1988 and 1998. 2007 CROWDER, HARMSEN, and BLATT: SUCCESSION IN OLD FIELDS 189 A. Non-woody species 35 a abe 30 Yani Non-woody species nchness 23 ~ ae. 7 e ~ B. Woody species Woody species nchness 70 — C- Grass species Mean percent (+ SE) cover or species richness 60 — D. Solidago species wo dA oe OD DOD GD & SP OOM NL N ro So Dd FP SF Or OTe ane een ase Year FIGuRE 3. Conditions in ploughed plot P2 during 1976-1988, 1995 and 1998. 3A shows non-woody species richness (species/ 100 m2) and mean annual percent cover. 3B shows woody species richness and cover. 3C shows cover of all grass species with peaks and lows. 3D shows cover of Solidago species with peaks and lows. 190 has been confirmed elsewhere by Carson and Pickett (1990) who attributed it to decreased shading by the macroforb. Shade-tolerant herbs in the plots included Sanicula marilandica (Black Snakeroot), Teucrium canadense ssp. canadense (Germander), Lactuca canadensis (Wild Lettuce), Solidago caesia and Aster cordifolius. The increase in woody plants documented by Crowder and Harmsen (1998) could have been expected to create conditions favouring increases of such shade-tolerant species. They did not, however, increase in frequency or cover over time. Presumably shaded microniches under plants such as goldenrods were present in early years and were later replaced by similar conditions under shrubs or trees. Acknowledgments Partial funding was received from the Ontario Ministry of Natural Resources; we particularly thank R. Wagner, S. Strobl and P. Neave. Preliminary analysis of field results was done by C. Vardy, and we thank her and the many others who helped with field surveys. Our thanks are due to J. Pringle and P. Catling who identified some of the composites and sedges. We are grateful for the continued support of Queen’s University Biology Station. Literature Cited Armesto, J. J., and S. T. A. Pickett. 1985. Experiments on disturbance in old-field plant communities: impact on species richness and abundance. Ecology 66: 230-240. Beschel, R. E., A. E. Garwood, R. Hainault, I. D. Mac- donald, S. P. van der Kloet, and C. H. Zavitz. 1970. List of the vascular plants of the Kingston region. Fowler Herb- arium, Queen’s University, Kingston, Ontario. Blatt, S. E., J. A. Janmaat, and R. Harmsen. 2001. Mod- elling succession to include a herbivore effect. Ecological Modelling 139: 123-136. Blatt, S. E., J. A. Janmaat, and R. Harmsen. 2003. Quan- tifying secondary succession: a method for all sites? Com- munity Ecology 4: 141-156. Brown, V. K., A. C. Gange, and C. W. D. Gibson. 1988. Insect herbivory and vegetational structure. Pages 263-280 in Plant form and vegetation structure. Edited by M. J. A. Werger, P. J. M. van der Aart, H. J. During, and J. T. A. Verhoeven. SPB Academic Publishing, The Hague. Carson, W. P., and S. T. A. Pickett. 1990. Role of resources and disturbance in an old-field plant community. Ecology 71: 226-238. Crowder, A., and R. Harmsen. 1998. Notes on forest suc- cession in old fields in southeastern Ontario: the woody species. Canadian Field-Naturalist 112: 410-418. Crowder A., K. E. J. Topping, and J. C. Topping. 1997. Plants of the Kingston Region: 1996. Fowler Herbarium, Queen’s University, Kingston, Ontario. Dale, H. M., P. J. Harrison, and G. W. Thomson. 1966. Weeds as indicators of physical site characteristics in aban- doned pasture. Canadian Journal of Botany 44: 1319-1327. Dansereau, P., and A. Gille. 1949. Ecologie des principaux types de paturage des environs de Granby. Bulletin du service de biogéographie. Université de Montréal, Québec. THE CANADIAN FIELD-NATURALIST Vol. 121 Epp, G. A., and Aarssen, L. W. 1988. Attributes of competi- tive ability in herbaceous plants. Pages 71-76 in Plant form and vegetation structure. Edited by M. J. A. Werger, P. J. M. van der Aart, H. T. During, and J. T. A. Verhoeven. SPB Academic Publishing, The Hague. Eriksson, O., and A. Jakobssen. 1998. Abundance, distri- bution and life histories of grassland plants: a comparative study of eighty-one species. Journal of Ecology 86: 922- 933. Goldberg, D. E., and K. L. Gross. 1988. Disturbance regimes of midsuccessional old fields. Ecology 69: 1677-1688. Henderson, J. B. 1986. The European Skipper (Thymelicus lineola): population ecology and herbivory in an old-field community. M.Sc. thesis, Queen’s University, Kingston, Ontario. Lamoureux, W. J. 1970. Ecological succession. Garden Bulletin 24. Royal Botanic Gardens, Hamilton, Ontario. Maycock, P. F., and M. Guzikowa. 1984. Flora and vege- tation of an old field community at Erindale, southern Ontario. Canadian Journal of Botany 62: 2193-2207. McBrien, H., R. Harmsen, and A. Crowder. 1983. A case of insect grazing affecting plant succession. Ecology 65: 1035-1039. McBrien, H., and R. Harmsen. 1987. Growth response of goldenrod, Solidago canadensis (Asteraceae) to periodic defoliation. Canadian Journal of Botany 56: 1035-1039. Mellinger, M. 1972. Dynamics of plant succession on aban- doned hay fields in central New York State. Ph.D. thesis, University of Syracuse, Syracuse, New York. Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. McMurray, and M. J. Oldham. 1998. Ontario Plant List. Ontario Ministry of Natural Resources, Ontario Forest Research Institute, Sault Ste. Marie, Ontario. Information Paper 123. Pengelly, D. H. 1961. Thymelicus lineola (Ochs) (Lepidoptera: Hesperiidae) a pest of hay and pasture grasses in southern Ontario. Proceedings of the Entomological Society of Ontario 91: 102-105. Reader, R. J. 1990. Relationship between seedling emergence and species frequency on a gradient of ground cover density in an abandoned pasture. Canadian Journal of Botany 69: 1397-1401. Reader, R. J., and J. Buck. 1991. Control of seedling density on disturbed ground; role of seedling establishment for some mid-successional old-field species. Canadian Journal of Botany 69: 773-777. Semple, J. C., S. B. Heard, and C. S. Xiang. 1996. The Asters of Ontario (Compositae: Asteraceae). University of Waterloo Biology Series 38: 1-94. Waterloo, Ontario. Werner, P. A., I. K. Bradbury, and R. S. Gross. 1980. The biology of Canadian weeds. 45. Solidago canadensis. Can- adian Journal of Plant Science 60: 1393-1409. Werner, P. A., and W. J. Platt. 1976. Ecological relationships of co-occurring goldenrods. (Solidago: Compositae). Amer- ican Naturalist 110: 959-971. Zar, J. 1984. Biostatistical analysis. Prentice Hall, Englewood Cliffs, New Jersey. Received 16 October 2006 Accepted 28 April 2008 Growth of White Spruce, Picea glauca, Seedlings in Relation to Microenvironmental Conditions in a Forest-Prairie Ecotone of Southwestern Manitoba SOPHAN CHHIN! and G. GEOFF WANG? ' Corresponding author: Department of Forestry, Michigan State University, East Lansing, Michigan 48824-1222 USA * Department of Forestry and Natural Resources, Clemson University, 261 Lehotsky Hall, Clemson, South Carolina 29634-0317 USA; e-mail: gwang @clemson.edu | Chhin, Sophan, and G. Geoff Wang. 2007. Growth of White Spruce, Picea glauca, seedlings in relation to microenvironmental! conditions in a forest-prairie ecotone of southwestern Manitoba. Canadian Field-Naturalist 121(2): 191-200. The influence of microenvironmental conditions on the growth performance (1.e., diameter and height growth) of White Spruce [Picea glauca (Moench) Voss] seedlings was examined within three contrasting habitats: White Spruce tree islands, open prairies and Trembling Aspen (Populus tremuloides Michx.) groves. These habitats exist within a disjunct occurrence of White Spruce at its southern limit of distribution in three mixed-grass prairie preserves in the Spruce Woods Provincial Park within the forest-prairie ecotone of southwestern Manitoba. Microenvironmental measurements (i.e., light, temperature, relative humidity, soil moisture) were obtained on 10 sites in each of the three habitats and growth characteristics of 60 White Spruce seedlings were measured in each of three habitats. Higher light and soil temperature conditions occurred within the open prairie. In contrast, lower light and soil temperature conditions occurred under the tree canopy of aspen groves and the north- ern aspect of spruce islands, which moderated the effect of the dry regional climate. Height growth did not vary significantly among the three habitats. The greater diameter growth and decreased slenderness of White Spruce seedlings in the open ver- sus the shaded habitats appears to be a result of increased photosynthesis at higher light intensity and may also represent a morphological adaptation to withstand the effect of increased wind exposure. The increased slenderness of White Spruce in the shaded habitats appears to be a morphological adaptation of increasing carbon allocation towards height growth and thus maximizing effective competition for light. Key Words: White Spruce, Picea glauca, prairie-forest boundary, aspen parkland, growth, microenvironment, seed-seedling conflicts, Manitoba. In the prairie provinces of Canada, the southern limit of conifer species occurs in the aspen parkland. Vegetation in the aspen parkland is dominated by Trembling Aspen (Populus tremuloides Michx.) which exists as continuous forest or as groves intermixed with prairie (Bird 1961). The aspen parkland is a tran- sitional vegetation zone (ecotone) between prairie to the south and boreal forest to the north (Bird 1961; Zoltai 1975). The prevailing regional climate of the aspen parkland and mixed-grass prairie in southwestern _ Manitoba is characterized by low precipitation, high summer temperatures, and dry winds, any of which may be limiting factors to plant distribution and growth - (Coupland 1950). For instance, previous studies which examined the radial growth-climate association of White | Spruce [Picea glauca (Moench) Voss] in the aspen parkland region demonstrated that growth was restrict- ed by moisture deficiency exacerbated by tempera- ture-induced drought stress (Chhin et al. 2004). Tree invasion into prairies under the dry climate of _ the aspen parkland must be assisted by favorable micro- environments or “safe sites” (Harper 1977; De Steven » 1991a, 1991b; Kellman and Kading 1992; Chhin and - Wang 2002). In many prairie environments, shrubs are _ known to facilitate tree seedling establishment (Call- away 1995; Chambers 2001) by suppressing grass com- petition (Bird 1961; Werner and Harbeck 1982) and by providing protective conditions from abiotic forces such as fire (Coupland 1950; Bird 1961). The presence of previously established trees in open prairie not only reduces evaporative water loss from plants (Kellman and Kading 1992) but also increases soil moisture and nutrients (Wilson and Kleb 1996; Li and Wilson 1998: Wilson 1998). Furthermore, woody canopies modify the understory microclimate, including shading, inter- ception of precipitation, litter fall (Vetaas 1992), reduced wind speed (Archibold et al. 1996), lower soil temper- ature, and higher humidity (Carlson and Groot 1997). Within the aspen parkland of southwestern Manitoba, White Spruce typically establishes under a Trembling Aspen canopy and will eventually replace aspen to form White Spruce forests in the absence of disturbances (Bird 1961). Abundant White Spruce seedlings and saplings are commonly observed in many Trembling Aspen stands, indicating a favorable environment for regeneration. However, White Spruce can occasional- ly invade open prairies within previously established Creeping Juniper (Juniperus horizontalis Moench) patches (Bird 1961; Chhin and Wang 2002). Creeping Juniper is believed to reduce mortality of White Spruce seedlings from prairie ground fires (Bird 1961). The progressive recruitment and establishment of White 19] Spruce in the open prairie results in the development of White Spruce tree islands which are surrounded by mixed-grass prairie. The islands have an asym- metric spatial structure since White Spruce regenera- tion is concentrated on the northern aspect of the tree islands. Chhin and Wang (2002) determined that the best environment for White Spruce regeneration with- in mixed-grass prairie was under the influence of estab- lished White Spruce trees (i.e., on the north vs. south aspect and between 4 to 12 m from the spruce island centre, and close association with Creeping Juniper). Other studies have also observed growth of White Spruce in the aspen parkland region on favourable microenvironments such as the north side of trees, north side of river valleys, and other north-facing slopes and microsites (Zoltai 1975; Hogg 1994; Hogg and Schwartz 1997). Together, these previous studies suggested that the suitability for White Spruce estab- lishment decreased from aspen groves (1.e., under a Trembling Aspen canopy), to White Spruce tree islands (i.e., under the influence of established White Spruce trees) and to Creeping Juniper patches in open prairies. However, environmental conditions associated with these habitats have not been documented and how White Spruce growth differs among these habitats has not been quantified. Ecotones are expected to be sensitive to climatic change, and the environment of the aspen parkland is believed to characterize the future boreal forest under a warmer climate (Hogg 1994). Moderated microen- vironmental conditions may potentially serve as micro- scale refugia or “safe sites” (Harper 1977) for tree seedlings in the context of global warming during the 21* century (IPCC 2007) within a climatically sensi- tive ecotonal region of the aspen parkland. The first objective of this study was to examine the microenvi- ronmental conditions of five habitats within the aspen parkland of the Spruce Woods Provincial Park (SWPP) in southwestern Manitoba, Canada. These habitats include one habitat in Trembling Aspen groves (AG) [interior (AG-I)], two habitats in White Spruce tree islands (SI) [northern (SI-N) vs. southern (SI-S) aspect], and adjacent to the islands, two habitats in open prairies (OP) [ground vegetation dominated by grass (OP-G) vs. juniper (OP-J)]. The second objective of the study was to compare White Spruce seedling growth (i.e., height and diameter) among three habitats (OP-J, SI- N, and AG-I; seedlings were not sampled in OP-G and SI-S since it was extremely rare to find them in these habitats) and relate growth patterns to the microenvi- ronmental patterns. Methods Study area The study was conducted in the SWPP which is located in southwestern Manitoba (49°40'N, 99°15'W) (Figure 1). The nearest meteorological station with a long-term climatic record is the Brandon Agriculture THE CANADIAN FIELD-NATURALIST Vol. 121 Station (49°52'N, 99°59'W), which is located about 88.5 km northwest of the SWPP. The area experienced, for the reference period of 1971-2000, an average annu- al temperature of 2.4°C (Environment Canada 2002). Average annual precipitation amounts to 474.0 mm, with 78.3% as rainfall and the remainder as snowfall. Precipitation peaks in June (75.7 mm), and tempera- ture is highest in July, reaching a daily mean of 18.9°C (Environment Canada 2002). Vegetation in the park is characterized as the aspen-oak grove of the boreal forest region (Rowe 1972). A sandhill, mixed-grass prairie community unique to the aspen parkland is located in the SWPP and consists of Trembling Aspen intermixed with a disjunct population of White Spruce at its southern limit of distribution (Schykulski and Moore 1997). The SWPP resides over an extensive belt of deltaic sands (6500 km”) created 12 000 years B.P. when a predecessor of the Assiniboine River flowed into glacial Lake Agassiz. The SWPP is located on this delta, and this area has been designated as the Assiniboine Delta Natural Region (ADNR). Within the ADNR, the SWPP contains some of the best exam- ples of the few remaining remnants of native mixed- grass prairie, since most of the native prairie in the ADNKR has been lost to agricultural cultivation. The mixed-grass prairie preserves of the SWPP are pro- tected under the mandate of the Prairie Management Plan of Manitoba Conservation in order to preserve its unique and rich biodiversity of plants, insects, and wildlife (Schykulski and Moore 1997). Natural and anthropogenic disturbances have con- tributed to the landscape mosaic of the aspen park- land (Bird 1961). In pre-European settlement times, disturbances included fire which occurred naturally or started by Native people in order to maintain the prairie environment for the Bison (Bison bison L.). Grazing by herbivores such as Bison and Elk (Cervus canadensis Erxl.) prevented the encroachment of the forest onto the prairie. European settlement contributed to the decline of the Bison population due to hunting, livestock grazing, intensive agriculture, and the policy of fire suppression (Bird 1961). The reduction of graz- ing and fire pressure has resulted in the encroachment of the forest onto the prairie. A policy of fire suppres- sion continues within the SWPP, with intermittent prescribed burning permitted within the mixed-grass prairie preserves (Schykulski and Moore 1997). Site selection The study was located in three mixed-grass prairie preserves north of the Assiniboine river: Picnic, Aspen Bluff, and Jackfish prairie (Figure 1). Ten White Spruce islands (SI) with adjacent open prairie (OP), and ten aspen groves (AG) were selected using aerial photo- graphs and field reconnaissance. Four sites of each habitat were chosen in Picnic prairie and Aspen Bluff prairie, whereas two sites of each habitat were selected in Jackfish prairie. Although the occurrence of habitat types could be confounded with environmental con- 2007 99°14'W Spruce Woods : Provincial Park | Epinette mag Prairie Sites —— Park Boundary — Roads Township Line 1:75000 500 0 500 1000 Meters Le ae CHHIN AND WANG: GROWTH OF WHITE SPRUCE SEEDLINGS 193 99°06'W MANITOBA / L« of’ ¢ . Lo SYS z < = WwW x O | 0.05). Light intensity in the aspen grove interior (AG-I, 18.3%) was significantly less than any of the other habitats (all P < 0.05). Soil temperature differed among some of the habi- tats (Table |). Soil temperature within spruce islands was lower in the northern (SI-N, 13.5°C) versus south- ern (SI-S, 16.4°C) aspect (P < 0.05). The spruce island habitats (SI-N, SI-S) in turn were not significantly different from soil temperature in the open prairie grass habitat (OP-G, 15.7°C) (all P > 0.05). Both the SI-S and the OP-G habitats had significantly greater soil temperature than the open prairie juniper habitat (OP-J, 13.4°C) (all P < 0.05). Soil temperature of the aspen grove habitat (AG-I, 10.2°C) was lower than any of the other habitats (all P < 0.05). While surface and air temperature did not differ significantly among the habitats they followed similar relative patterns among the habitats as described for soil temperature. Although relative air humidity was greatest in aspen groves (AG-I, 36.7%), lowest in the open prairie (OP- G, 30.4%; OP-J, 31.8%), and intermediate in spruce islands (SI-N, 32.7%; SI-S, 33.7%) there were no significant differences in relative air humidity among these habitats (all P > 0.05) (Table 1). Surface soil moisture in the northern island aspect (SI-N, 7.1%) was not significantly different from that in the southern aspect (SI-S, 4.7%) (P > 0.05) (Table 1). Soil moisture in the southern island aspect in turn was significantly less than in either the open prairie grass (OP-G, 10.9%) or aspen grove (AG-I, 8.9%) (all P < 0.05). Surface soil moisture of the open prairie juniper (OP-J, 8.8%) habitat was not significantly dif- ferent from the other habitats (all P > 0.05). Average soil moisture of all profile depths in the SI-S (1.8%) habitat was less than that in the AG-I (8.0%) habitat (all P < 0.05) (Table 1). Seedling growth White Spruce seedlings in the open prairie and spruce islands established exclusively amongst Creep- ing Juniper. Seedlings in aspen groves established ex- clusively on forest floor litter (i.e., Trembling Aspen leaves). General seedling attributes are shown in Table 2. Mean annual height growth of seedlings did not dif- fer significantly among the three habitats (all P > 0.05) (Figure 2). Mean annual diameter growth differed significantly amongst the three habitats (all P < 0.05) (Figure 3). Open prairie seedlings had the greatest diam- eter followed by seedlings from spruce islands and then aspen groves. The slenderness coefficient also dif- fered significantly amongst the three habitats (all P <0.05) (Figure 4). Aspen grove seedlings were most slender, followed by seedlings from spruce islands and then open prairies. Discussion Less exposure to solar radiation was expected under the canopy of aspen groves and spruce islands because of shading. Light conditions under aspen canopy 196 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE |. Microenvironmental conditions of Trembling Aspen groves [interior (AG-I) habitat], and White Spruce tree islands {north (SI-N) and south (SI-S) aspect habitat] with adjacent open prairies [grass (OP-G) and juniper (OP-J) habitat]. The lower and upper limits of a 95% confidence interval of the mean of ten replicates per habitat are shown in parentheses. For each microenvironmental variable, habitats with different letters are significantly different (P < 0.05). Except for the tem- perature data, comparisons were made based on log-transformed data. Microenvironmental variables with no letters for each habitat were not significantly different (P > 0.05). Habitat Variable! OP-G OP-J SLS SIN AG-I Light (% PAR) 100.0 c 100.0 c 79.1 ¢ 54.5 b 18.3 a Temp. (°C) (100.0, 100.0) (100.0, 100.0) (62.6, 100.0) (44.1, 67.2) (15.9, 21.1) Soil lSeade 13.4 be 16.4e 13.5 cd 10.2 a (14.1, 17.3) (11.8, 14.9) (IS.7/, 7eib)) (12.1, 14.9) (9.7, 10.7) Surface 31.3 30.8 30.1 29.1 28.0 (28.1, 34.5) (27.9, 33.8) (27.3, 32.9) (27.4, 30.7) (25.6, 30.3) Air 31.7 31.6 29.3 30.5 29.1 (28.7, 34.6) (28.7, 34.4) (26.8, 31.9) (28.9, 32.1) (26.7, 31.6) RH (%) 30.4 31.8 33)57/ 32.7 36.7 (25.0, 37.1) (26.0, 38.8) (26.7, 42.6) (28.0, 38.1) (30.5, 44.1) SM (% vol) Surface 10.9b 8.8 ab 47a 7.1 ab 8.9b (8.7, 13.6) (6.5, 11.9) (2.6, 8.5) Gell, 97) (7.4, 10.8) Profile 5.3 ab 2.4 ab 1.8a 2.0 ab 8.0 b (2.0, 12.0) (0.4, 7.2) (0.4, 4.5) (0.7, 4.4) (6.2, 10.1) 'PAR, photosynthetically active radiation; Temp., temperature; RH, relative air humidity; SM, soil moisture TABLE 2. Attributes of White Spruce seedlings from White Spruce tree islands, open prairies, and Trembling Aspen groves. Standard error of the mean in parentheses. Attribute Open Prairie Spruce Island Aspen Grove Sample size (7) 60 60 60 Height (cm) 50.4 (4.9) 49.8 (4.6) 69.6 (6.7) Diameter (cm) 1.66 (0.21) 1.24 (0.13) 1.16 (0.12) Age (year) 11.4 (0.6) 11.9 (0.7) 15.1 (0.8) (18.3%) occurred at levels similar to other studies of aspen stands (Archibold et al. 1996: 23%; Carlson and Groot 1997: 18%). The dominant overstory White Spruce trees cast shade onto the understory, with more shade on the northern versus southern aspect. The reduced soil temperature of the northern versus south- ern island aspect as well as the lowest soil temperature in aspen groves indicated that areas with increased shade were also cooler. This was expected since tem- perature is generally a function of solar radiation (Archibold et al. 1996; Morecroft et al. 1998). Decreased soil temperature may lead to decreased evapotranspirative demands in the aforementioned shaded areas. In our study, soil temperature in aspen groves was 3.2-5.5°C less than in the open prairie. Other studies have also reported similar levels of reduced soil temperature in aspen (Archibold et al. 1996: 5°C) and other deciduous stands (Morecroft et al. 1998; 2.5°C) compared to adjacent open conditions. Reduced solar radiation has the potential to moderate the microclimate in forest/grassland ecosystems and thus provide a less hostile environment for tree regen- eration and growth (Zoltai 1975; Hogg 1994; Bres- hears et al. 1997; Hogg and Schwartz 1997). While light and soil temperature varied significant- ly among different habitats, surface and air tempera- ture as well as relative air humidity did not. Morecroft et al. (1998) also found that a deciduous forest canopy had a greater effect on reducing soil temperature than air temperature. Although air temperature and humid- ity did not differ significantly among the habitats, the lower level of air temperature in aspen groves (2.5- 2.6°C lower) versus the open prairie was comparable to that observed in other aspen (Archibold et al. 1996: 1.5°C lower; Carlson and Groot 1997: 0.7°C) and deciduous stands (Morecroft et al. 1998: 0.6-0.9°C). Relative humidity was 4.9-6.3% higher in aspen groves compared to the open prairie. This is comparable to a study in the aspen parkland of Saskatchewan by Archi- bold et al. (1996). They observed that average relative humidity in the summer was generally 4% higher in- side than outside an aspen grove. Higher air humidity 2007 i a) Open Prairie (4.1 + 0.2) o 3 © = 4 ® ic 1 0 3 + 5 Mean annual height growth (cm y-") b) Spruce Island (3.7 + 0.1) Frequency OF — GN) 16> =| 3 4 5 Mean annual height growth (cm yr-") c) Aspen Grove (4.2 + 0.2) - NYO wo Ff Frequency 3 4 5 Mean annual height growth (cm yr-") FIGURE 2. Frequency histogram of height/age ratio (i.e., mean annual height growth) of White Spruce seedlings from open prairies (a), White Spruce tree islands (b), and Trembling Aspen groves (c). Mean + | standard error of the mean of 10 replicates per habitat shown in parentheses. Mean annual height growth did not differ significantly among habitats (P > 0.05). is usually linked to the effects of reduced wind speed and reduced air temperature (Archibold et al. 1996) Some significant differences in soil moisture were observed among the five habitats. Better moisture re- tention due to increased shading may explain signifi- cantly higher surface and profile soil moisture in aspen groves when compared to SI-S. Due to different degrees of shading, both soil surface and profile moisture were also slightly higher on the northern versus southern aspect of spruce islands. These results support the CHHIN AND WANG: GROWTH OF WHITE SPRUCE SEEDLINGS 197 a) Open Prairie (0.126 + 0.010 C) 4 o 3 Cc 52 oO ® iy i 0 0.05 0.10 0.15 0.20 0.25 Mean annual diameter growth (cm y-") A b) Spruce Island (0.091 + 0.003 B) o 3 S iD ion oO ic 1 0 0.05 0.10 0.15 0.20 0.25 Mean annual diameter growth (cm y-") 4 c) Aspen Grove (0.069 + 0.004 A) an) © ae oO Liter 0 0.05 0.10 0.15 0.20 0.25 Mean annual diameter growth (cm yr-") FIGURE 3. Frequency histogram of RCD/age ratio (i.e., mean annual radial growth) of White Spruce seedlings from open prairies (a), White Spruce tree islands (b), and Trembling Aspen groves (c). Mean + | standard error of the mean of 10 replicates per habitat shown in parentheses, and habitats with different capital letters are significantly different (P < 0.05). hypothesis of soil moisture retention which asserts that soil moisture is expected to be greater in areas of increased shading (Wilson and Kleb 1996; Li and Wilson 1998; Wilson 1998). The significantly higher soil surface moisture found in OP-G, and the lack of significant difference between open prairies and spruce islands or aspen groves in soil profile moisture are, however, difficult to explain. A possible explanation is that tree or shrub canopies, especially those of conifers, intercept rainfall and thus limit the amount of through- 198 fall (Vetaas 1992). Although water may also enter soil via stemflow, it has been shown that such water is more likely to infiltrate to deep soil layers to be used by tree species (Parker 1983; Vetaas 1992; Breshears et al. IGS). Although open prairies represent the worst habitat for White Spruce establishment (Chhin and Wang 2002), they support the best seedling growth. These results support the hypothesis of seed-seedling con- flicts which asserts that the habitat requirements for optimal seedling establishment are discordant with that for seedling growth (Schupp 1995). White Spruce did not show any significant differences in height growth amongst the three habitats. In contrast, White Spruce significantly had the greatest diameter growth within the open prairie followed by tree islands and then aspen groves. These results suggest that diameter growth was more responsive to the environmental factors associ- ated with each habitat than height growth. These results are consistent with that of Groot (1999) who reported that diameter growth of young, planted White Spruce was primarily influenced by light availability, whereas height growth exhibited complex relationships with multiple environmental factors. The larger diameter growth of White Spruce seedlings in the open prairie was not expected given the harsher abiotic microenvironment (i.e., heat stress) of the open prairie compared to the moderated microclimatic con- ditions within the spruce islands and aspen groves. There are a number of factors which may have con- tributed to the better diameter growth of seedlings in the open prairie and the lower than expected diameter growth within spruce islands and aspen groves. These factors include the effect of substrate growth condi- tions, light availability, and morphological adaptations to wind. First, surveying White Spruce seedlings in the open prairie was difficult given the rarity of finding open prairie spruce. Furthermore, those White Spruce individuals that were found grew exclusively in asso- ciation with Creeping Juniper. Therefore, the better than expected growth in the open prairie may be due to the moderating influence of a Creeping Juniper sub- strate, and Creeping Juniper is likely providing White Spruce seedlings protective cover from prairie ground fires (Bird 1961). However, seedlings on the northern aspect of spruce islands also established exclusively on juniper. If substrate was a dominant factor influencing diameter growth it is unclear why seedlings from the open prairie and spruce islands would have different diameter growth rates given the same growth substrate. Corresponding to the diameter growth reduction from OP-J to SI-N to AG-I, the degree of shade in- creased. The reduced supply of solar radiation could have outweighed the benefits of a moderated micro- climate. In other words, while shade is an important moderating microclimatic factor that is conducive to successful White Spruce establishment (Chhin and Wang 2002), shade becomes a limiting factor to sub- sequent seedling growth. Although ecophysiological THE CANADIAN FIELD-NATURALIST Vol. 121 a) Open Prairie (38.3 + 1.9 A) Frequency Oo - NY WwW FA 20), 650) 4055 450)) (GOV e70mco Slendemess coefficient b) Spruce Island (45.9 + 1.3 B) Frequency Oo =] NY WwW A 20° 230) 240 350) 260 e700 Slendemess coefficient c) Aspen Grove (64.2 + 2.3 C) w WwW F#& Frequency 20K. E50) ee AOe 50 a0 eae Oe oO. Slendemess coefficient FiGuRE 4. Frequency histogram of height/RCD ratio (ie., slenderness coefficient) of White Spruce seedlings from White Spruce tree islands (a), open prairies (b), and Trembling Aspen groves (c). Mean + | stan- dard error of the mean of 10 replicates per habitat shown in parentheses, and habitats with different capital letters are significantly different (P < 0.05). studies indicate that White Spruce seedlings general- ly attain maximal height growth at 40% full sunlight (Lieffers and Stadt 1994; Lieffers et al. 1996), seedling diameter growth is greatest at full light intensity (Logan 1969). This may explain the reduced diameter growth of seedlings from the northern aspect of spruce islands which was characterized by light levels of 54.5% of full sunlight. Light levels below 8% generally results in mortality of White Spruce (Lieffers and Stadt 1994). While light conditions under the aspen canopy (18.3%) were greater than this minimal light level, they are 2007 well below that required for maximal diameter growth, Man and Lieffers (1997) also confirmed that light intensity under an aspen canopy was not sufficient to reach the saturation point although light levels were above the compensation point for White Spruce pho- tosynthesis. Another likely factor which may explain the dis- crepancy in diameter growth between the three habi- tats may be morphological adaptations to the effect of wind. Although wind speed was not examined here, it has been shown in another study in Saskatchewan that wind speed in an aspen grove was reduced to 7% of that in adjacent prairie (Archibold et al. 1996). Forest openings are generally characterized by greater wind speeds than forest interiors (Carlson and Groot 1997). Wind causes trees to sway and this can lead to morphological adaptations to resist the effect of wind via increased diameter growth particularly at the base of trees (Kozlowski et al. 1991; Telewski 1995). The effect of shade and wind is further supported by White Spruce being most slender in aspen groves than in tree islands and the open prairie. These results confirmed our morphological observations in the field. White Spruce seedlings from aspen groves were gen- erally tall and thin, while seedlings from the open prairie were short and thick. White Spruce seedlings from spruce islands were intermediate along this mor- phological continuum. The morphology of White Spruce under aspen canopy is thus suited for concen- trating growth in height to counter the effects of over- topping vegetation (Lieffers and Stadt 1994; Lieffers et al. 1996). The morphology of White Spruce in the open prairie is suited for a more robust stature to with- stand the effect of winds during summer (Archibold et al. 1996) and wind and snow abrasion during winter. Wind and snow abrasion have been shown to influence the growth form of open grown spruce at its northern limit of distribution at the tree-line in Churchill, Man- itoba (Scott et al. 1993). The possible discordance in habitat requirements of different developmental stages of White Spruce seed- lings may potentially influence successional dynam- ics in a forest-prairie ecotone in the Spruce Woods Provincial Park. For instance, projecting the initial slow degree of White Spruce seedling establishment in the open prairie onto all developmental stages of White Spruce may underestimate the rate of succes- sional expansion of forest onto prairie, since subse- quent seedling diameter growth in the open prairie appears to benefit from high light availability. Nonethe- less, further research is required to separate the rela- tive effects of high light availability and increased wind exposure on increased diameter growth of White Spruce seedlings in the open prairie. Projecting the initial establishment success of White Spruce under a Trem- bling Aspen canopy onto all developmental stages of White Spruce may overestimate the rate of succession- al replacement of Trembling Aspen by White Spruce, CHHIN AND WANG: GROWTH OF WHITE SPRUCE SEEDLINGS 199 since shade appears to outweigh the benefits of a mod- erated microclimate for diameter growth of White Spruce seedlings under a Trembling Aspen canopy In conclusion, higher light and soil temperature con ditions occurred within the open versus shaded habi- tats. The greater diameter growth and decreased slen- derness of White Spruce seedlings in the open versus shaded habitats appears to be a result of increased photosynthesis at higher light intensity and may also represent a morphological adaptation to withstand the effect of increased wind exposure. The increased slenderness of White Spruce in the shaded habitats appears to be a morphological adaptation of increas- ing carbon allocation towards height growth and thus maximizing effective competition for light. While shade is an important moderating microclimatic factor that is conducive to successful White Spruce establishment (Chhin and Wang 2002), our study has shown that at a later developmental stage, shade is a contributing factor towards limiting diameter growth. Our results support the hypothesis that there are conflicting habi- tat requirements for trees at different developmental stages (Schupp 1995). Thus, in the context of future global warming (IPCC 2007) in a climatically sensitive ecotonal region of the aspen parkland, although mod- erated microclimates may act as micro-scale refugia or “safe sites” (Harper 1977) for tree seedling estab- lishment, this may be at the expense of reduced growth at a later developmental stage. Models of forest growth in response to climate change should therefore take into account the role of conflicting habitat requirements for trees at different developmental stages. Acknowledgments This study was funded through a Natural Sciences and Engineering Research Council of Canada (NSERC) Postgraduate Scholarship to S. Chhin, and a research grant from Global Forest to G. G. Wang. We thank Drs. R. Staniforth and J. Tardif for their contributions to the initial research proposal. Thanks also go to K. Ryan, R. Klos, and M. Kreiner for their assistance in the field data collection; and K. Schykulski and H. Hernandez of Manitoba Conservation for their logis- tical support. Literature Cited Archibold, O.W., E. A. Ripley, and D. L. Bretell. 1996. Comparison of the microclimates of a small aspen grove and adjacent prairie in Saskatchewan. American Midland Naturalist 136: 248-261. Bird, R. D. 1961. Ecology of the aspen parkland of western Canada in relation to land use. Canada Department of Agri- culture, Research Branch, Publication 1066, Ottawa. On- tario. 155 pages. Breshears, D. D., P. M. Rich, F. J. Barnes, and K. Camp- bell. 1997. Overstory-imposed heterogeneity in solar radi- ation and soil moisture in a semiarid woodland. Ecologi- cal Applications 7: 1201-1215. Callaway, R. M. 1995. Positive interactions among plants. Botanical Review 61: 306-349. 200 Carlson, D. W., and A. Groot. 1997. Microclimate of clear- cut, forest interior, and small openings in Trembling Aspen forest. Agricultural and Forest Meteorology 87: 313-329. Chambers, J. C. 2001. 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American Midland Naturalist 108: 124-132. Wilkinson, L. 1990. SYSTAT: the system for statistics. SYSTAT Inc., Evanston, Illinois. Wilson, S. D. 1998. Competition between grasses and woody plants. Pages 231-254 in Population biology of grasses. Edited by G. P. Cheplick. Cambridge University Press, Cambridge, United Kingdom. Wilson, S. D., and H. R. Kleb. 1996. The influence of prairie and forest vegetation on soil moisture and available nitro- gen. American Midland Naturalist 136: 222-231. Yamaguchi, D. K. 1991. A simple method for cross-dating increment cores from living trees. Canadian Journal of Forest Research 21: 414-416. Zoltai, S. C. 1975. Southern limit of coniferous trees on the Canadian prairies. Information Report NOR-X-128, Cana- dian Forestry Service, Edmonton, Alberta. 12 pages. Received 2 January 2007 Accepted 9 May 2008 | Natural Recolonization of Cultivated Land by Native Prairie Plants and its Enhancement by Removal of Scots Pine, Pinus sylvestris PAUL M. CATLING! and BARRY KING? ' Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1 A 0C6 Canada; e-mail: catlingp @agr.ge.ca 2 217 King St. West, Apartment 175, Cobourg, Ontario K9A 2N2 Canada Catling, Paul M., and Barry King. 2007. Natural recolonization of cultivated land by native prairie plants and its enhancement by removal of Scots Pine, Pinus sylvestris. Canadian Field-Naturalist 121(2): 201-205. A combination of native and introduced plants colonized abandoned cultivated land with adjacent relict prairie and savanna in the Rice Lake region of southern Ontario. After 71 years, the native colonizers included 86 species found in regional prairie relicts, but much of the area was also colonized by introduced Scots Pine, Pinus sylvestris, which apparently spread from an adjacent planting. The pines formed expanding patches of dense growth that excluded other species. Removal of the invasive Scots Pine from a 200 m? plot within the abandoned land led to colonization 17 years later by 36 native species characteristic of the prairie, savanna and sand barrens of the region. Included in this group of native prairie colonizers were keystone species such as Andropogon gerardii, Carex siccata, Ceanothus americanus, Comptonia peregrina and Quercus velutina. Areas where patches of Scots Pine had been allowed to continue growing were either devoid of vegetation or had a sparse understory of introduced species and Poison Ivy (Rhus radicans). These observations support the concepts of (1) protecting islands of native dry ground flora which can serve as sources for recolonization and (2) protection of old field and particularly recently cultivated land adjacent to protected native grassland to allow natural restoration. The fact that keystone native species were able to colonize the area from which the introduced Scots Pines had been removed suggests that the pines are aggressive competitors that occupy space to the exclusion of the native species. Scots Pine is thus a driver of ecological change in degraded ecosystems. Management including removal of Scots Pine to support native plant biodiversity is strongly supported. Key Words: Scots Pine, Pinus sylvestris, prairie, sand barren, savanna, recolonization, driver, Andropogon gerardii, Carex siccata, Ceanothus americanus, Comptonia peregrina, Quercus velutina, Rice Lake, Ontario. Although information on restoration of prairie veg- etation is abundant (e.g., Schramm 1992; Packard and Mutel 1997), information on natural recolonization of prairie flora following human impacts (e.g., Inouye et al. 1987; Rabinowitz and Rapp 1985) is relatively scarce. Despite the lack of information it is often con- _ sidered a good idea to preserve degraded habitats that have islands of pristine habitat or have adjacent pris- _ tine habitats that can serve as sources for natural re- colonization. It appears in some cases that invasive alien plants can limit the success of natural recoloniza- tion in these circumstances but a lack of information is apparent in this area as well. While some have expressed concern that invasive plant species are one of the lead- ing direct causes of biodiversity loss (e.g., Catling 2005), others have questioned the extent to which in- troduced species threaten biodiversity (e.g., Davis 2003) and several recent articles have recommended changes in language and attitude in dealing with inva- Sive species problems based to some extent on the need for more accurate information (Gobster 2005; Larson 2005). Recently it has been suggested that invasive plants are the passengers rather than the drivers of eco- logical change in degraded ecosystems (MacDougal and Turkington 2005). As “drivers” the invasives would compete with native species and may also change the environment to reduce the capability of native species to survive. As “passengers” they are not the limiting factor for native species but just along for the ride. MacDougal and Turkington (2005) suggested that if interactive processes were limiting, then removal of in- vasive plants should result in a direct increase in native species richness (the “driver” model). On the other hand, if invasive plants are not the limiting factor for native plants, then removal of the invasive plant(s) should have little effect (the “passenger” model). Although common sense suggests that (1) natural recolonization could occur from pristine nuclei and (2) that invasive alien plants may be either drivers or passengers, the cost of managing natural areas requires stronger evidence. A simple field experiment conduct- ed in the Rice Lake region of eastern Ontario provides information relating to these assumptions. It involves invasive alien Scots Pine (Pinus sylvestris) and a poten- tially diverse savanna, sand barren and prairie flora. In situations where a native sand barren, prairie or savanna has been replaced by a dense stand of Scots Pine with a completely barren and dark understory (e.g., Catling and Carbyn 2005), it is compelling to think of this invasive as a “driver of ecological change” (Did- ham et al. 2005). Indeed, Scots Pine is listed as the fifth most significant invasive alien plant of natural habitats in Canada (Catling and Mitrow 2005). However, the present question is whether or not Scots Pine is a driv- er of ecological change in a degraded ecosystem where the native prairie vegetation has been reduced by other 201 202 factors. If following perturbation the native prairie species respond by colonizing an area of previously cultivated ground and they also respond by colonizing a portion of that same ground from which invading Scots Pine is removed, then native prairie plants do naturally recolonize and the introduced pine is a driver of ecological change that is clearly in the way since it is preventing the natural restoration of native flora fol- lowing the cessation of degrading processes. Methods Natural Recolonization The degraded ecosystem in this experiment origi- nated from an area of sandy soil that was completely utilized for the cultivation of cereal crops and espe- cially potatoes until 1935 when it was abandoned. It is located at 44.1268°N, 77.9839°W in Lot 4, Conces- sion 7, originally Haldimand Twp. but now Alnwick/ Haldimand township in Northumberland County, On- tario. The eradication of native flora is confirmed by BK whose family occupied the land, and further by aerial photographs taken in 1929. Cultivation was es- sentially continuous following settlement in the 1850s, but prior to this the region had been a rolling prairie dominated by prairie grasses with localized Black Oak (Querus relutina) Red Pine (Pinus resinosa) savanna and sand barren flora (Catling et al. 1992). By 2006 a combination of native species and alien species, in- cluding alien Scots Pine, had colonized the previously cultivated field. It had become a semi-open forest with 50-year-old pines, including the Scots Pine, along with the native White Pine (Pinus strobus) and both Red (Querus rubra) and Black oaks, but throughout the forest were openings of various sizes with a combina- tion of native prairie species (see Table 1) and inva- sive alien species, including Hypericum perforatum, Poa compressa, Hieracium pilloselloides, Rumex ace- tosella, Potentilla norvegica, and especially Pinus syl- vestris. Native species that have colonized the aban- doned potato field are believed to have spread from the adjacent Russ Creek pioneer cemetery and from the adjacent Barr property preserve, which was never cultivated. The early pioneer graveyard dates from 1849 (earliest marker, although it had a schoolhouse beside it in 1845 and was likely used much earlier) with few burials after 1900. The prairie plant commu- nity in pioneer graveyards in prairie regions was usu- ally not cultivated or grazed unlike much of the sur- rounding landscape because it was sacred ground. Since it was only controlled occasionally with a scythe, the prairie vegetation survived and this seems to be the case with the Russ Creek Cemetery, which is a nucle- us Of rare and restricted native prairie species. In 2006 an open (treeless) portion of the recolonized field that was 0.8 hectares in extent was selected as a sample plot to document recolonization of native species. All native plant species present in this sample plot were recorded during spring, summer and fall visits in 2006. THE CANADIAN FIELD-NATURALIST Vol. 121 Effect of removing Scots Pine The invading Scots Pines appeared to be preventing the re-colonization of native species in the old field since they grew in dense stands that contained no other species of vascular plants. In 1991 a dense stand of pines was removed from an area of 200 m? adjacent to the sample plot to allow colonization of native species. Beneath this stand, and beneath others within 100 m, the only plants at the time of tree removal were occa- sional depauperate specimens of Poison Ivy (Rhus rad- icans) which does well in the decomposing leaf litter of Scots Pine needles. In 2006 the area from which the trees were removed was surveyed and all native plant species present were recorded during spring, summer and fall visits. Native flora was also recorded from adjacent stands of Scots Pine (where the trees had not been removed). Scientific and common names used in the text gen- erally follow Kartesz and Meacham (1999). Repre- sentative specimens are at the AAFC (Agriculture and Agri-Food Canada) vascular plant herbarium in Ottawa (acronym — DAO). Results and Discussion Natural Colonization In 2006, 71 years after the old potato field was aban- doned as cultivated land, a total of 86 species charac- teristic of natural prairie, savanna and sand barrens in the region was recorded in the 0.8 hectare sample plot of an open (treeless) area (Table 1). It is clear that na- tive species can recolonize seriously degraded habitat in this region but it is evidently a slow process since the occurrence of native species was still very patchy after 71 years despite a relatively uniform substrate and elevation throughout the area. The return of prairie flora to the previously cultivated field is to be expect- ed on the basis of observations elsewhere where dis- persal and colonization from adjacent protected sites have proven to be the major factors in old field succes- sion (e.g., Inouye et al. 1987). The results of this study support the concept of protecting islands of native dry ground flora which can serve as sources for recolo- nization. The results also suggest that even in the north- eastern-most prairie and savanna habitats, protection of previously cultivated land adjacent to protected native grassland is likely to result in an old field suc- cession that includes many native prairie species. Fur- thermore, some studies have shown that the native prairie species increase with time (over 60 years) and with increasing soil nitrogen at the expense of both the introduced plants and native non-prairie plants (Inouye et al. 1987: Figure 8). Thus protection of aban- doned farmlands adjacent to protected sites is likely to result in a degree of natural restoration that can be supported through plantings and management. Effect of Removing Scots Pine It is likely that the colonization by native species in the old potato field would have been more extensive | 2007 CATLING and KING: RECOLONIZATION OF CULTIVATED LAND 203 TABLE |, Native species of prairie, savanna and sand barren habitats recorded in 2006 that naturally colonized a 0.8 hectare plot of previously cultivated land abandoned in 1935, The native species recorded in the 200 m* subplot in 2006, Le., 15 years after removal of invading Scots Pines, are indicated with an asterisk (*). Amelanchier arborea (Michx.) Fern. var. arborea, DOWNY SERVICE-BERRY * Amelanchier laevis Wieg., ALLEGHENY SERVICE-BERRY Amelanchier stolonifera Wieg., RUNNING SERVICE-BERRY * Andropogon gerardii Vitman, BIG BLUESTEM Anemone cylindrica Gray, LONG-HEAD THIMBLEWEED Antennaria howellii ssp. petaloidea (Fern.) Bayer, SMALL PUSSYTOES * Antennaria parlinii ssp. fallax (Greene) Bayer & Stebbins, PARLIN’S PUSSYTOES * Apocynum androsaemifolium L., SPREADING DOGBANE Aquilegia canadensis L., RED COLUMBINE Arabis divaricarpa A. Nels., ROCKCRESS Arabis hirsuta var. pycnocarpa (M. Hopkins) Rollins, HAIRY ROCKCRESS Arabis holboellii var. retrofracta (Graham) Rydb., HOLBOELL’S ROCKCRESS Arctostaphylos uva-ursi (L.) Spreng., RED BEARBERRY Artemisia campestris ssp. borealis var. scouleriana (Hook.) Crong., PACIFIC WORMWOOD Asclepias exaltata L., POKE MILKWEED * Asclepias syriaca L., COMMON MILKWEED * Asclepias tuberosa ssp. tuberosa, BUTTERFLY MILKWEED * Calystegia spithamaea ssp. spithamaea, LOW FALSE BINDWEED * Carex backii Boott, BACK’S SEDGE Carex gracillima Schwein., GRACEFUL SEDGE * Carex molesta Mackenzie ex Bright, TROUBLESOME SEDGE * Carex muehlenbergii var. muehlenbergii, MUHLENBERG’S SEDGE Carex pensylvanica Lam., PENNSYLVANIA SEDGE Carex richardsonii R. Br., RICHARDSON’S SEDGE * Carex siccata Dewey, DRY-SPIKE SEDGE * Carex tonsa var. rugosperma (Mackenzie) Crins * Ceanothus americanus L., NEW JERSEY-TEA Comandra umbellata ssp. umbellata, BASTARD TOADFLAX * Comptonia peregrina (L.) Coult., SWEET-FERN Cornus racemosa Lam., GRAY DOGwWooD Cyperus houghtonii Torr., HOUGHTON’S FLAT SEDGE Cyperus lupulinus (Spreng.) Marcks ssp. lupulinus, GREAT PLAINS FLAT SEDGE * Danthonia spicata (L.) Beauv. ex Roemer & J.A. Schultes, POVERTY WILD Oat GRASS * Dichanthelium sabulorum (Lam.) Gould & C.A. Clark var: thinium (A.S. Hitche. & Chase) Gould and C.A Clark (D. columbianum Scribner), HEMLOCK ROSETTE GRASS Dichanthelium depauperatum (Muhl.) Gould, STARVED ROSETTE GRASS Dichanthelium linearifolium (Scribn. ex Nash) Gould, SLIM LEAF ROSETTE GRASS Dichanthelium perlongum (Nash) Freckmann, LONG SLIM LEAF ROSETTE GRASS Elymus trachycaulus ssp. subsecundus (Link) A.& D. Léve (includes unilaterale), SLENDER WILD RYE * Fragaria virginiana Duchesne ssp. virginiana, VIRGINIA STRAWBERRY Galium boreale L., NORTHERN BEDSTRAW Galium circaezans Michx. var. circaezans, LICORICE BEDSTRAW Helianthus divaricatus L., WOODLAND SUNFLOWER Lechea intermedia var. intermedia, ROUND FRUIT PINWEED Liatris cylindracea Michx., ONTARIO GAYFEATHER Lilium philadelphicum L. var. philadelphicum, Woop LILy * Maianthemum canadense Desf., FALSE LILY-OF-THE-VALLEY * Maianthemum stellatum (L.) Link (Smilacina stellata), STARRY FALSE SOLOMON’S SEAL Melampyrum lineare Dest. var. lineare, AMERICAN COW WHEAT Monarda fistulosa L. ssp. fistulosa var. fistulosa, OSWEGO-TEA * Physalis heterophylla Nees, CLAMMY GROUND CHERRY Penstemon hirsutus (L.) Willd., HAIRY BEARDTONGUE Pinus strobus L., EASTERN WHITE PINE * Poa pratensis L. ssp. pratensis, KENTUCKY BLUE GRASS * Polygala polygama Walt., RACEMED MILKWORT Polygonatum biflorum (Walt.) Ell. var. commutatum (J.A. and J.H. Schultes) Morong, KING SOLOMON’ S-SEAL * Potentilla arguta Pursh ssp. arguta, TALL CINQUEFOIL Potentilla simplex Michx., OLDFIELD CINQUEFOIL Prunus pumila L. var. susquehanae (hort. ex Willd.) Jaeger, SUSQUEHANNA SAND CHERRY * Prunus serotina Ehrh., BLACK CHERRY Prunus virginiana L. var. virginiana, CHOKE CHERRY 204 TABLE |. (cont.) THE CANADIAN FIELD-NATURALIST Vol. 121 Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underwood ex Heller, NORTHERN BRACKEN FERN Quercus alba L., NORTHERN WHITE OAK Quercus rubra L., NORTHERN RED OAK * Quercus velutina Lam., BLACK OAK Ranunculus rhomboideus Goldie, LABRADOR BUTTERCUP * Rhus typhina L., SLIM LEAF ROSETTE GRASS * Rosa acicularis Lindl. ssp. acicularis, PRICKLY ROSE * Rosa blanda Ait. var. blanda, SMOOTH ROSE * Rubus idaeus L. ssp. idaeus, COMMON RED RASPBERRY Rudbeckia hirta L. var. hirta, BLACK-EYED SUSAN Salix humilis Marsh var. humilis, PRAIRIE WILLOW Schizachyrium scoparium (Michx.) Nash var. scoparium, LITTLE FALSE BLUESTEM Shepherdia canadensis (L.) Nutt., RUSSET BUFFALO-BERRY Solidago canadensis L. var. canadensis, CANADA GOLDENROD Solidago gigantea Ait., LATE GOLDENROD * Solidago juncea Ait., EARLY GOLDENROD * Solidago nemoralis Ait. var. nemoralis, GRAY GOLDENROD Sorghastrum nutans (L.) Nash, YELLOW INDIAN GRASS * Symphyotrichum ericoides (L.) Nesom vat. ericoides, WHITE HEATH AMERICAN-ASTER * Symphyotrichum novae-angliae (L.) Nesom, NEW ENGLAND ASTER * Symphyotrichum oolentangiense (Riddell) Nesom var. oolentangiense, SKY-BLUE ASTER * Toxicodendron rydbergii (Small ex Rydb.) Greene, WESTERN POISON Ivy Viola adunca Sm. var. adunca, HOOK-SPUR VIOLET * Viola sagittata Ait. var. ovata (Nutt.) Torr. & Gray, ARROW-LEAF VIOLET Vitis riparia Michx., RIVER-BANK GRAPE and less patchy if the Scots Pines had not invaded. Scattered dense stands of Scots Pine were without other species in 1991 and in 2006. However, within the recol- onized area of the old field where the Scots Pines were removed in 1991, 36 native species of prairie, savanna and sand barren affinity were found in 2006, and had thus colonized this space within 15 years (Table 1). The absence of native species under the dense Scots Pines that remained suggested that this colonization would not have occurred if the pines had not been removed. Although introduced species had also colonized this area from which the Scots Pines were removed, includ- ing particularly Poa compressa, Hieracium pilosel- loides and young Scots Pines, the native species were at least 50% of the cover. Therefore, the pines occupy space to the exclusion of the native species and restrict recolonization of disturbed sites. Since only more pines and Poison Ivy and raspberries Rubus spp. occur in the older (more than 50 years old) Scots Pine stands in the region, there is very good evidence that the Scots Pines interfere with the recolonization of native prairie species following cessation of degrading processes and apparently also change the environment to reduce the capability of native species to survive. In adjacent stands of Scots Pines the understory was bare or cov- ered only by a scant growth of Poison Ivy, raspberries, and introduced plants. The native species that respond- ed to the removal of the pines included the character- istic and keystone native species of prairie, savanna and sand barren such as Andropogon gerardii, Carex siccata, Ceanothus americanus, Comptonia peregrina and Quercus velutina. The fact that these native species were able to colonize the area from which the pines were removed, but did not where the pines remained, suggests that the pines are aggressive competitors that occupy space to the exclusion of the native species and change conditions by competing for nutrients, reduc- ing light and changing soil characteristics with leaf lit- ter accumulation, so that most of the native species of prairie, savanna and sand barren cannot exist. Scots Pine in this situation and evidently in many other situa- tions (e.g., Catling and Carbyn 2005) is thus a driver of ecological change. Management including removal of Scots Pine to support native plant biodiversity is strong- ly supported. Since the trees are relatively slow grow- ing, slow to mature, reproduce only by seed, are con- spicuous at all times, and easily eliminated by pulling and cutting and/or burning, control of Scots Pine is much less problematic than the control of some other invasive plant species. Conclusion These observations suggest that natural recoloniza- tion by native prairie species can occur, but that it is hindered by invasive alien Scots Pine, which drives ecological change in a degraded prairie ecosystem. Acknowledgments Ms. B. Kostiuk provided extensive assistance in the field survey. Earlier surveys in the area were con- ducted by D. White and E. Heuvel. 2007 Literature Cited Catling, P. M. 2005. Effects of invasive alien plants on birds: some examples from North America. Biodiversity 6; 30-39. Catling, P. M., and S. Carbyn. 2005. Invasive Scots Pine, Pinus sylvestris, replacing Corema, Corema conradii, heathland in the Annapolis valley, Nova Scotia, Canadian Field-Naturalist 119: 237-244. Catling, P. M., V. R. Catling, and S. M. McKay-Kuja. 1992. The extent and floristic composition of the Rice Lake Plains based on historical records. Canadian Field- Naturalist 106: 73-86. Catling, P. M., and G. Mitrow. 2005. A prioritized list of the invasive alien plants of natural habitats in Canada. Canadian Botanical Association Bulletin 38: 55-57. Didham, R. K., J. M. Tylianakis, M. A. Hutchison, R. M. Ewers, and N. J. Gemmell. 2005. Are invasive species the drivers of ecological change? Trends in Ecology and Evolution 20: 470-474. Davis, M. A. 2003. Biotic globalization: does competition from introduced species threaten biodiversity? Bioscience 53: 481-489. Gobster, P. H. 2005. Invasive species as ecological threat: is restoration an alternative to fear-based resource manage- ment? Ecological Restoration 23: 261-270. Inouye R. S., N. J. Huntly, D. Tilman, J. R. Teste, M. Still- well, and K. Zinnel.1987. Old-field succession on a Min- nesota sand plain. Ecology 68: 12-26 CATLING and KING: RECOLONIZATION OF CULTIVATED LAND 205 Kartesz, J. T., and C. A. Meachum. 1999. Synthesis of the North American Flora, version 1.0. North Carolina Botan- ical Garden, Chapel Hill, North Carolina. Larson, B. M. H. 2005. The war of the roses: demilitarizing invasion biology. Frontiers in Ecology and Environment 3: 495-500. MacDougal, A. S., and R. Turkington. 2005. Are invasive species the drivers or passengers of change in degraded ecosystems? Ecology 86: 42-55. Packard, S., and C. F. Mutel. Editors. 1997. The tallgrass restoration handbook: for prairies, savannas, and wood- lands. Society for Ecological Restoration and Island Press, Washington, D.C. xxxil + 463 pages. Rabinowitz, D., and J. K. Rapp. 1985. Colonization and establishment of Missouri prairie plants on artificial soil disturbances. III. Species abundance distributions, survi- vorship, and rarity. American Journal of Botany 72: 1635- 1640. Schramm, P. 1992. Prairie restoration: a 25-year perspec- tive on establishment and management. Pages 169-177 in Proceedings of the Twelth North American Prairie Con- ference. Edited by D. D. Smith and C. A. Jacobs. Univer- sity of Northern Iowa, Cedar Falls. Received 19 September 2006 Accepted 22 May 2008 Notes A High Elevation Record of the Star-nosed Mole (Condylura cristata) in Northeastern Vermont RYAN W. Norris and C. WILLIAM KILPATRICK Department of Biology, University of Vermont, Burlington, Vermont 05405 USA; e-mail: rnorris@zoo.uvm.edu and wkilpatr@ zoo.uvm.edu Norris, Ryan W., and C. William Kilpatrick. 2007. A high elevation record of the Star-nosed Mole (Condylura cristata) in northeastern Vermont. Canadian Field-Naturalist 121(2): 206-207. A single male Star-nosed Mole, Condylura cristata, was captured in a rock pile at the top of East Mountain (elevation 1042 m; 3240 ft) in northeastern Vermont. Although known from high elevations (up to 1676 m; 5500 ft) in southern U.S. states, this species was not known to occur above 573 m (1880 ft) in the northern part of its range. This record is also important in confirming that C. cristata has some climbing ability when travelling above ground. Key Words: Star-nosed Mole, Condylura cristata, elevation, talus, Vermont. The Star-nosed Mole, Condylura cristata, has the largest and most northerly geographic range of all North American moles (Peterson 1966; Hall 1981). The species ranges from southwest Manitoba to Labrador and Nova Scotia, down the eastern United States to South Carolina, and to the Great Lakes region (Petersen and Yates 1980). The species is known to be associated with moist soils near marshes and streams and is an excellent swimmer (Hamilton 1931; Petersen and Yates 1980). Habitat preference for the Star-nosed Mole is usually characterized as low wet ground near bodies of water, swamps, wet meadows, wet spots in fields or low-lying woods, and dry soils within mixed hardwoods near water (Godin 1977; DeGraaf and Yamasaki 2001). Beane (1995), Laerm et al. (1997), and McCay et al. (1999) have suggested that C. cristata is quite com- mon in high elevations in states at the southern edge of its distribution such as North Carolina and South Carolina. Elevational records reported for southern U.S. states include a record at 760 m (2494 ft) in South Carolina (Laerm et al. 1997) and several individuals from the Smoky Mountain National Park at elevations ranging from 488 to 1676 m (1600-5500 ft; Linzey and Linzey 1971). Literature records from Canada and northern U.S. states rarely report elevation, but all avail- able records are lower. The highest elevation reported at northern latitudes that we have been able to find is Saunders’s (1988) suggestion that C. cristata ranges up to 573 m (1880 ft) in elevation in the Adirondacks of New York. Study Area and Methods The summit of East Mountain in northeastern Ver- mont was modified in 1955 with the construction of the 206 North Concord Air Force Station. The ridgeline was levelled creating an area of approximately 17 acres on the summit for this radar station. The rubble from this modification was pushed to the edges, levelling the surface and creating substantial areas of talus. The station was deactivated in 1963; the buildings have slowly deteriorated, and have been partially disman- tled. The native fauna and flora have since recolo- nized the modified area. Dense regenerative spruce- fir forest now covers much of the summit and small mammals associated with talus habitat, such as the Yellow-nosed Vole (Microtus chrotorrhinus), have been found in this human-made talus habitat. During a small mammal survey, 30 Sherman live traps, baited with rolled oats, were set in and adjacent | to one of several human-made talus piles at the top of East Mountain (elevation 1042 m or 3420 ft), East | Haven township, Essex County, Vermont, 44°39.5'N 71°46.1'W over three nights (23-25 August 2004). Traps were checked daily and rebaited as needed. The talus pile was approximately 20 m in diameter and contained bare, flattish rocks ranging from ~ 0.5 m to 3 m in diameter. It was located on a gentle slope and had small pools of standing water. No larger streams or bodies of water were observed in the vicinity. Traps ; in the talus were placed on rocks, not soil, but a small | quantity of humus had collected in some areas. Results A single male C. cristata was discovered dead in a Sherman trap placed among rocks approximately 5 m from the edge of the talus pile on the first night of trapping (23 August). This night saw moderate rain with nighttime low temperatures at or approaching’ O°C. The measurements of this specimen are: total 2007 length 174 mm, tail length 80 mm, and hind foot length 27 mm. Skin, skull, and tissue are housed in the Zadock Thompson Natural History Museum at the University of Vermont (catalog number 5273). Other species cap- tured over the course of these three nights of trapping were Short-tailed Shrew, Blarina brevicauda (1); Smokey Shrew, Sorex fumeus (1), Southern Red- backed Vole, Myodes gapperi (5), and Deer Mouse, Peromyscus maniculatus (8). Discussion The Star-nosed Mole, Condylura cristata, is clearly a more northern-adapted species than other moles in eastern North America. The relatively high number of reports of this species in mountainous areas in south- ern states is not surprising considering its distribution as far north as Labrador. Our capture of the species at a high elevation site in Vermont was unexpected, par- ticularly considering that it was taken in a talus pile and not associated with soil or long-term standing water. The single specimen reported here is evidence that the Star-nosed Mole also occurs at high elevations (at least 1042 m; 3420 ft) even in the northeastern USA. This capture also suggests that they can live, or travel through, rocky areas away from soil. Peterson (1966) suggested that Star-nosed Moles spend more time above ground than other mole species and, as a result, are occasionally found dead on the surface or captured on the surface with snap or live traps. The capture of the mole amongst rocks and not on bare soil suggests that it must have been maneuvering within this rock pile. Hickman (1982) reported observ- ing a vertical climb of 60 cm by a Star-nosed Mole in the corner of a concrete column, and Schmidt (1931) reported a Star-nosed Mole climbing two feet out of a swimming tank. Our capture provides further indica- tion that C. cristata is capable of some degree of climb- ing, a feature not often attributed to fossorially adapted mammals. Hickman (1982) hypothesized that the climbing ability of C. cristata may serve to aid in mo- bility across rocky areas. Our capture is also consis- tent with this idea. NOTES 207 Acknowledgments The authors thank Charles A. Woods for important discussion leading to the preparation of this manu- script. This specimen was collected in association with a small mammal survey funded by the Kingdom Com- mons Group. Literature Cited Beane, J. C. 1995. New distributional record for the star- nosed mole, Condylura cristata (Insectivora: Talpidae), in North Carolina, with comments on its occurrence in the Piedmont Region. Brimleyana 22: 77-86. DeGraaf, R. M., and M. Yamasaki. 2001. New England Wildlife: Habitat, Natural History, and Distribution. Uni- versity Press of New England, Hanover, New Hampshire 482 pages. Godin, A. J. 1977. Wild Mammals of New England. Johns Hopkins University Press, Baltimore, Maryland. 304 pages. Hall, E. R. 1981. The mammals of North America. Volume 1. John Wiley and Sons, New York. 600 pages. Hamilton, W. J., Jr. 1931. Habits of the star-nosed mole, Condylura cristata. Journal of Mammalogy 12: 345-355. Hickman, G. C. 1982. Climbing ability of the star-nosed mole, Condylura cristata (Talpidae). Saugetierkundliche Mitteilungen 30: 296-297. Laerm, J., G., Livingston, C. Spencer, and B. Stuart. 1997. Condylura cristata (Insectivora: Talpidae) in the Blue Ridge Province of western South Carolina. Brimleyana 24: 46-49. Linzey, A. V., and D. W. Linzey. 1971. Mammals of Great Smoky Mountains National Park. University of Tennessee Press, Knoxville. 114 pages. McCay, T. S., M. J. Komoroski, and W. M. Ford. 1999. Use of an upland pine forest by the star-nosed mole, Condylura cristata. The Journal of the Elisha Mitchell Scientific Soci- ety 115: 316-318. Petersen, K. E., and T. L. Yates. 1980. Condylura cristata. Mammalian Species 129: 1-4. Peterson, R. L. 1966. The mammals of eastern Canada. Ox- ford University Press, Toronto. 465 pages. Saunders, D. A. 1988. Adirondack Mammals. State Univer- sity of New York. 216 pages. Schmidt, F. K. W. 1931. Mammals of Western Clarke County, Wisconsin. Journal of Mammalogy 12: 99-117. Received 22 February 2006 Accepted 8 February 2008 208 THE CANADIAN FIELD-NATURALIST Vol. 121 Northern Long-eared Bat, Myotis septentrionalis (Chiroptera: Vespertilionidae), on Prince Edward Island: First Records of Occurrence and Over-Wintering JENNIFER A. BRowN!, DONALD F. MCALPINE“, and ROSEMARY CURLEY? University of Guelph, Department of Animal and Poultry Science, Guelph, Ontario N1G 2W1 Canada ’New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada (author to whom correspondence should be addressed; e-mail: donald.mcalpine @nbm-mnb.ca) Forests, Fish and Wildlife Division, Prince Edward Island Department of Environment, Energy, and Forestry, P.O. Box 2000, Charlottetown, Prince Edward Island C1A 7N8 Canada Brown, Jennifer. A., Donald F. McAlpine, and Rosemary Curley. 2007. Northern Long-eared Bat, Myotis septentrionalis (Chiroptera: Vespertilionidae), on Prince Edward Island: first records of occurrence and over-wintering. Canadian Field-Naturalist 121(2): 208-209. First occurrence of the Northern Long-eared Bat, Myotis septentrionalis, on Prince Edward Island is reported. A mixed- species hibernaculum of M. septentrionalis and the Little Brown Bat, M. lucifugus, is also described from southeastern Prince Edward Island. This is the first record of bats over-wintering in the province and the first time either species has been reported making use of a building as a hibernaculum. Key Words: Little Brown Bat, Myotis lucifugus, Northern Long-eared Bat, Myotis septentrionalis, Prince Edward Island, distribution, hibernaculum. The bats of Prince Edward Island are poorly known. Although seven species have been recorded from the adjacent mainland of New Brunswick and Nova Sco- tia (van Zyll de Jong 1985), only the Little Brown Bat (Myotis lucifugus) and Hoary Bat (Lasiurus cinereus) are known from the Island (van Zyll de Jong 1985; McApine et al. 2002). Cameron (1958) reported a sight- ing which he believed to be a Red Bat (Lasiurus bore- alis) near coastal Fortune Bridge, Kings County, Prince Edward Island (46°21'23"N 62°29'57"W) on 7 June 1954, but this species remains unconfirmed on the Is- land. Neither have any bat species been documented to hibernate on Prince Edward Island. Here we docu- ment the first occurrence of the Northern Long-eared Bat (Myotis septentrionalis) from the province and report a mixed-species hibernaculum of M. lucifugus and M. septentrionalis from southeastern Prince Ed- ward Island. On 24 September 1988, two bats retrieved by a house cat (Felis domesticus) were collected in Breadalbane, Queens County, Prince Edward Island (46°21'23"N 63°29'57"W) by JAB and subsequently identified as M. septentrionalis. These specimens, deposited in the Canadian Museum of Nature (CMN), include a male (CMN 52612) and a female (CMN 52613). Measurements for both bats (M (male), TL (total length) 85 mm, T (tail) 37 mm, HF (hind foot) 10 mm; F (female), TL 90 mm, T 36 mm, HF 10 mm) fall within the range for the species presented by van Zyll de Jong (1985). On or about 10 January 1989 a commercial pest con- trol officer removed about 800 bats, both Little Brown and Northern Long-eared bats, from the basement of a home at Culloden, Queens County, Prince Edward Island (46°0'58"N 62°42'8"W; reported in the media as Murray River, the larger nearby community). All bats were destined to be incinerated. However, DFM was able to secure 781 of these bats alive and an unsuc- cessful attempt was made to maintain them in a state of torpor over the winter with the intention of spring release. Various media reports suggest that the hiber- naculum contained 1500 bats (Anonymous 1989*; Cody 1989 a*, b*; White 1989*). However, communi- cation with the pest control officer (personal commu- nication to DFM) confirmed that all hibernating bats observed were removed. Information presented here is based on an examination of all 781 bats, apparently the contents of virtually the entire hibernaculum. We visited the hibernaculum site on 29 November 1990 and found only two torpid female M. septentrionalis, clinging to wooden floor joists. A voucher series of both species from the site have been deposited in the mammal collection of the New Brunswick Museum. The hibernation site was the unheated basement of an occupied home, built about 1980. The house had been recently re-occupied following a 3-5 year vacancy (resident personal communication to DFM). The base- ment had an earthen floor and island stone and mortar foundation walls with inside dimensions about 11 x 8 m and a ceiling height of 1-2 m. Entry to the basement was via a trap door on the first floor. Air temperature in the hibernaculum recorded on the night of 29 No- vember 1990 was 11.3°C. This hibernation tempera- ture, while above the mode of 6°C for vespertilionids, falls within the range of -4° to 13°C and 0.6°C to 13.9°C reported for Little Brown and Northern Long- eared bats; respectively (Webb et al. 1996). The hiber- naculum was dominated by the Little Brown Bat (82.6%); with smaller numbers of the Northern Long- eared (17.4%). In both species sex ratios were slightly ] 4 at ees 2007 female biased (62.1% and 57.1% in M. lucifugus and M. ‘septentrionalis, respectively). We also assessed reproductive status by dissection of all Northern Long-eared Bats following the criteria and terminol- ogy of Sluiter and Bouman (1951), Sluiter (1954, 1961), and Racey (1974), with the following results: males; 5.9% sexually immature, 94.1% sexually mature; females; 17.5% nulliparous, 14.2% non- parous, 68.3% parous. The Northern Long-eared Bat is widespread across Atlantic Canada (van Zyll de Jong 1985). Although less abundant, apparently, in the region than the Little Brown Bat, the presence of the Northern Long-eared Bat on Prince Edward Island is not unexpected. Cullo- den and Breadalbane are 67 km straight-line distance apart, suggesting M. septentrionalis may be widespread in the province. Conversely, the discovery of large num- bers of hibernating bats on the Island was unantici- pated. The province is underlain largely by Permian sandstones, siltstone and glystone (van de Poll 1983) and we are aware of no natural caves or abandoned mines on the Island that might provide suitable hiber- nacula. The closest natural caves serving as bat hiber- nacula occur in adjacent New Brunswick and Nova Scotia (McAlpine 1983; Mosely 2007). Fenton and Barclay (1980) were unable to locate any records of Little Brown Bats hibernating in buildings. Likewise, the Northern Long-eared has previously only been recorded hibernating in caves and mines (van Zyll de Jong 1985). Both these species of Myotis may be more flexible in their hibernation requirements than the lit- erature suggests, being able to rapidly exploit suit- able artificial hibernacula where present. Acknowledgments We are grateful to the late C. G. van Zyll de Jong, both for his encouragement and for confirming the identity of the two bats collected at Breadalbane. We also thank the resident of Culloden who so graciously allowed us access to her home, and Earl Peterson, Mari- time Varmint and Rodent Control, for agreeing to make the bats removed from this home available to DFM. Documents Cited [marked* in text] Anonymous. 1989. Going batty (photo with caption). The Telegraph Journal, Saint John, New Brunswick, 14 January 1989, Page 1. NOTES 209 Cody, J. 1989a. P.E.1. bats likely to become “ex-bats”: people curious about their fate. The Guardian, Charlottetown, Prince Edward Island, 19 January 1989, Page 6 Cody, J. 1989b. Bat removal poses question of animal's right to live. The Evening Patriot, Charlottetown, Prince Edward Island, 18 January 1989, Page 3 White, D. 1989. Bats take up residence in Culloden basement. The Times-Transcript, Moncton, New Brunswick Literature Cited Cameron, A. W. 1958. Mammals of the islands in the Gulf of St. Lawrence. Bulletin 154, National Museum of Canada, Ottawa. Fenton, M. B., and M. R. Barclay. 1980. Myotis lucifugus. Mammalian Species 142: 1-8. McAlpine, D. F. 1983. Status and conservation of solution caves in New Brunswick. New Brunswick Museum. Pub- lications in Natural Science (1). 28 pages. McAlpine, D. F, F. Muldoon, and A. I. Wandeler. 2002. First record of the Hoary Bat, Lasiurus cinereus (Chi- roptera: Vespertilionidae), from Prince Edward Island. Canadian Field-Naturalist 116: 124-125. Mosely, M. 2007. Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick, Canada. International Journal of Speleology 36: 1-21. Racey, P. A. 1974. Ageing and assessment of reproductive status of Pipistrellus bats, Pipistrellus pipistrellus. Jour- nal of Zoology 173: 264-271. Sluiter, J. W. 1954. Sexual maturity in bats of the genus Myotis I. Females of M. mystacinus and supplementary data on female M. myotis and M. emarginatus. Koninklijke Nederlandsche Academie van Wetenschappen 57: 696-700. Sluiter, J. W. 1961. Sexual maturity in males of the bat Myoris myotis. Koninklijke Nederlandsche Academie van Weten- schappen 64: 243-249. Sluiter, J. W., and M. Bouman. 1951. Sexual maturity in bats of the genus Myotis I. Size and histology of the repro- ductive organs during hibernation in connection with age and wear of the teeth in female Myotis myotis and Myotis emarginatus. Koninklijke Nederlandsche Academie van Wetenschappen 54: 595-603. van de Poll, H. W. 1983. Geology of Prince Edward Island. Province of Prince Edward Island, Department of Energy and Forestry. Report $3-1. van Zyll de Jong, C. G. 1985. Handbook of Canadian Mam- mals 2: Bats. National Museum of Natural Sciences, Ottawa. Webb, P. I, J. R. Speakman, and P. A. Racey. 1996. How hot is a hibernaculum? A review of the temperature at which bats hibernate. Canadian Journal of Zoology 74: 761-765. Received 9 May 2007 Accepted 14 January 2008 210 THE CANADIAN FIELD-NATURALIST Vol. 121 Arboreal Late Summer Courtship Behaviour of Maritime Garter Snake, Thamnophis sirtalis pallidulus, in Dartmouth, Nova Scotia, Canada JOHN GILHEN! and ROBERT STRUM2 ' Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada; e-mail: gilhenja @gov.ns.ca 2 325 Prince Albert Road, Dartmouth, Halifax County, Nova Scotia B2Y 1N5 Canada Gilhen, John, and Robert Strum. 2007. Arboreal late summer courtship behaviour of Maritime Garter Snake, Thamnophis sirtalis pallidulus, in Dartmouth, Nova Scotia, Canada. Canadian Field-Naturalist 121(2): 210-211. We report, for the first time, Maritime Garter Snake, Thamnophis sirtalis pallidulus, arboreal, late summer, courtship behaviour in the wild observed 4-5 September 2006. Key Words: Maritime Garter Snake, Thamnophis sirtalis pallidulus, arboreal, late summer, courtship behaviour, Dartmouth, Nova Scotia, Canada. On the morning of 5 September 2006 Robert Strum reported to John Gilhen that a ball of snakes had gathered on a Rhododendron shrub, Rhododendron sp.., in his yard at 325 Prince Albert Road, Dartmouth, Nova Scotia. In the afternoon of the previous day, 4 Sep- tember 2006, Robert Strum could see, from the kitchen window, that a number of brown snakes had gathered inside a gazebo style bird feeder on the Rhododendron shrub and he was successful in moving them out. The following morning, 5 September 2006, he noticed the snakes had gathered, again, on a branch near the bird feeder, about two meters above the ground (Cover). Upon investigation it was determined that there were four (three males and one female) Maritime Garter Snakes, Thamnophis sirtalis pallidulus, involved in courtship behaviour: male number 1, the largest male; males 2 and 3 about equal total length; and 1 female (the largest of the four snakes). The weather at 1200 hours on 5 September 2006 was cloudy and overcast with ambient air temperature of 20°C and temperature near the ground was also 20°C. At 1215 hours the males were observed moving alongside and over the back of the female. At the same time the body of male number | was pulsating rhyth- mically. This male rubbed his chin along the sides of the female. At this time the tail and about one half of the female’s posterior trunk (about 20 cm) drooped downward from one of the supporting branches, with the tails of all three males wrapped around her tail. While the female’s posterior body and tail were dang- ling down, the most active male (male number 1) ap- peared to maintain ventral apposition to copulate with her (Figure 1). All four snakes remained on the Rhododendron shrub from the morning to early afternoon but dispersed at some time late in the afternoon, as they were not seen there again that evening or in the following days. It is not known if copulation occurred over the two-day period these snakes occupied the Rhododendron shrub. One of the two subordinate males was collected as a voucher specimen (NSM55388). Courtship behaviour of the Maritime Garter Snake in captivity has been previously observed by John Gilhen in late August (unpublished). On two separate occasions (August 2001 and August 2004), days after individual females had given birth, two males were observed following alongside the female and trying to attain cloacal apposition. Although other adult females were present in the enclosure, the males were only in- terested in courting with a recently spent female. These observations of captives and the present observation in the wild suggests that the Maritime Garter Snake mates in late summer, sometime after females give birth, but prior to hibernation. Galbraith (2001) reports similar arboreal late sum- mer courtship by Eastern Garter Snakes, Thamnophis sirtalis sirtalis. Cook (1984) mentions snakes in the genus Thamnophis most frequently copulate in spring but have occasionally been noted to copulate in late fall, immediately prior to entering hibernacula. Arboreal spring breeding activity of the Red-sided Garter Snake, Thamnophis sirtalis parietalis, was observed on 8 May 1972 near Inwood, Manitoba (Gregory 1975). Rossman et al. (1996) and Ernst and Ernst (2003) agree mating of Thamnophis sirtalis usually occurs in spring after emergence from hibernation but may also take place in autumn. Rossman et al. (1996) also states that, until recently, most of the studies of mating be- havior and the factors that regulate the timing of re- production in T. sirtalis have focused primarily if not exclusively on spring activities. Previous accounts of the species in Nova Scotia (Barnes et al. 2006; Gilhen 1984, 2000), have not noted fall mating. The urbanization of both Halifax and Dartmouth cities over the past two decades has been extensive leaving many isolated pockets of amphibians and rep- tiles. The Maritime Garter Snakes at the observation site in Dartmouth continue to survive in small numbers in a back yard landscape, surrounded by apartment buildings on the south side, a major city road on the north side and other homes on the east and west sides. | Galbraith’s (2001) observation of the Eastern Garter | | 2007 NOTES 2] Ficure |. An adult male Maritime Garter Snake, Thamnophis sirtalis pallidulus, seeking ventral apposition with adult female in courtship behavior on the branches of a Rhododendron shrub at 325 Prince Albert Road, Dartmouth, Nova Scotia, on 5 September 2006. Snake also took place on private land in a cottage community on the coast sand dune area of the eastert shore of Lake Huron in Ontario In Nova Scotia, the Maritime Garter Snake is most common along the shores of lakes, ponds and stream but has readily adapted to a host of urban habitat Robert Strum has commonly observed Maritime Garter Snakes in his yard and has noted they move down along the base of the swimming pool and prob ably hibernate under the pool. Acknowledgment The authors appreciate the advice offered by Andrew Hebda, Curator of Zoology, Nova Scotia Museum of Natural History, during the preparation of the manu- script. Literature Cited Barnes, S., C. M. Dubesky, and T. B. Herman. 2006 Ecology and morphology of Thamnophis sirtalis pallidulus (Maritime Garter Snake) on Georges Island, Nova Scotia Northeastern Naturalist 13(1): 73-82. Cook, F. R. 1984. Introduction to Canadian amphibians and reptiles. National Museum of Natural Sciences, Ottawa, Ontario. 200 pages. Ernst, C. H., and E. M. Ernst. 2003. Snakes of the United States and Canada. Smithsonian Books, Washington and London. 668 pages. Galbraith, D. A. 2001. Arboreal courtship behaviour by Eastern Garter Snakes, Thamnophis sirtalis sirtalis, in Sep- tember in Bruce County, Ontario. Canadian Field-Naturalist 115: 347-348. Gilhen, J. 1984. Amphibians and reptiles of Nova Scotia Nova Scotia Museum. Halifax. 162 pages. Gilhen, J. 2000. Amphibians and reptiles of Nova Scotia Species recorded in the accession books of Harry Piers from 1899 to 1939. Manuscript Report. Nova Scotia Mu- seum of Natural History. Gregory, P. T. 1975. Arboreal mating behaviour in the Red- sided Garter Snake. Canadian Field-Naturalist 89: 461-462. Rossman, D. A., N. B. Ford, and R. A. Seigel. 1996. The garter snakes: Evolution and ecology. University of Okla- homa Press, Norman, Oklahoma. Received 2 November 2006 Accepted 11 March 2008 PWD THE CANADIAN FIELD-NATURALIST Vol. 121 Human-assisted Movements of Raccoons, Procyon lotor, and Opossums, Didelphis virginiana, between the United States and Canada Rick RosaTTE!, DENNIS DONOVAN!, MIKE ALLAN!, LAURA Bruce!, and CHRIS Davies! ‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, 2140 East Bank Dr., Peterborough, Ontario K9J 7B8, Canada *Corresponding author (e-mail: rick.rosatte @ ontario.ca) Tel. (705) 755-2280, Fax (705) 755-1559 Rosatte, Rick, Dennis Donovan, Mike Allan, Laura Bruce, and Chris Davies. 2007. Human-assisted Movements of Raccoons, Procyon lotor, and Opossums, Didelphis virginiana, between the United States and Canada. Canadian Field-Naturalist 121(2): 212-213. Movements of Raccoons (Procyon lotor) (x = 479 km) and Opossums (Didelphis virginiana) (xX = 688 km) by means of transport trailers and a train from the USA and Quebec into Ontario were significantly greater than Raccoon movements (x = 15 km) from Ontario to New York State determined by mark-recapture. Human-assisted movements of wildlife could have significant impacts with respect to cross-border movements of diseases such as rabies. Proactive communication programs, especially at International border crossing areas, should be encouraged to decrease the occurrence of the unintentional movement of wildlife and associated diseases. Key Words: Didelphis virginiana, Opossum, Procyon lotor, Raccoon, movement, disease, rabies, Ontario. The movement of wildlife and domestic animals by humans has resulted in the national, international, and sometimes global movement of diseases such as rabies, West Nile virus, bovine tuberculosis, parvovirus, chron- ic wasting disease (CWD), and bovine spongiform en- cephalopathy (BSE) (Jenkins and Winkler 1987; Niel- son and Brown 1988; Woodford and Rossiter 1993; Williams and Barker 2001). The international move- ment of Raccoons (Procyon lotor) from the United States to Ontario, Canada, has been documented previ- ously (Rosatte et al. 1997, 2001); however, those pub- lications did not document the distances moved or geographic areas involved. During January 1997 and December 2005, a total of 14 Raccoons were documented as having moved across the St. Lawrence and Niagara Rivers from Ontario, Canada, to New York State. Those Raccoons were ear- tagged and released in Ontario as part of a wildlife rabies control program during 1997-2005 (Rosatte et al. 1997, 2001). They were later recaptured or found dead in the vicinity of Buffalo or Ogdensburg, New York (Figure 1). Exact locations were known for 13 of those. Mean movement of the 13 Raccoons was 15 km (SD=9.0) (range = 3-30 km). The sex and age were known for 11 of the 13 Raccoons. More adult males (9) than adult females (2) moved from Ontario to New York (Chi Square = 8.9; P<0.0028). Mean movement (18 km, SD=8.4) (range = 6-30 km) of adult male Raccoons was greater than mean movement of adult females (7.5 km, SD=6.4) (range=3-12 km) (t=1.65; e—(at)) In addition, during 1998-2005, a total of 14 Rac- coons and two Opossums (Didelphis virginiana) were found in transport trailers and in a box car on a train in the greater Toronto area. The point of departure of those vehicles included Ohio, Ilinois, Tennessee, New York, Michigan, Quebec, and Ontario (Figure 1). In most cases the trailer or container was locked and sealed at the point of departure and was not opened until it arrived in Ontario. Mean movement of Raccoons by vehicle from the USA and Quebec (n=11) to Ontario, Canada, was 479 km (SD=250) (range=250-890 km). This was significantly greater (t=6.9, P<0.00002) than movements of Raccoons noted above that did not in- volve transport via vehicles. The two Opossums moved 1000 km from Tennessee and 376 km from Michigan, respectively. In Ontario, rabid Raccoons have been documented as having moved 0.7-4.1 km (Rosatte et al. 2005, 2006). Median movements of non-rabid Raccoons in Ontario were generally less than 1 km, with movements rarely exceeding 50 km in distance (Rosatte 2000; Rosatte et al. 2006). However, those movements were assumed not to have been due to human involvement (intentional or otherwise) (i.e. by vehicles). The exceptional move- ments for Raccoons, as well as Opossums, documented in this paper, indicate that those species have the poten- tial to move infectious diseases such as rabies signifi- cant distances. In view of this, people should be vigi- lant for wildlife such as Raccoons riding on vehicles. Transport trailer drivers (as well as operators of other vehicles such as ferries, trains, boats) should be encour- aged to check their vehicles and trailers for wildlife prior to departure to minimize the unintentional move- ment of animals. This will assist in decreasing the movement of wildlife diseases. Signage at internation- al border crossing areas advising drivers to check their vehicles for wildlife should also be posted. These tac- tics were used during the 1990s in Ontario in an attempt to prevent the movement of Raccoon rabies from the United States to Canada (Rosatte et al. 1997). Acknowledgments Special thanks to Brad Gates, AAA Wildlife Con- trol, Scarborough, Ontario; Scarborough Animal Con- trol; Mississauga Animal Control staff; Dan Lawrence, | | ) | | | | q 2007 Animal Control, Markham; Brampton Animal Control; Art King, Canadian Food Inspection Agency, Ft. Erie, Ontario; Rick Stevens and Ted Smith New York State Department of Environmental Conservation; James Lucas, Lockport, New York; Laura Bigler, Cornell Uni- versity; Mark Carrera, United States Department of Agriculture, Potsdam, New York; and Jim Masters, Niagara County Health Department, New York. The manuscript was reviewed by Mark Gibson, Tore Buchanan, and Kirk Sobey, Ontario Ministry of Nat- ural Resources, Peterborough, Ontario. Literature Cited Jenkins, S., and W. Winkler. 1987. Descriptive epidemiol- ogy from an epizootic of raccoon rabies in the Middle Atlantic States, 1982-1983. American Journal of Epidemi- olgy 126: 429-437. Nielson, L., and R. Brown. 1988. Translocation of wild ani- mals. Wisconsin Humane Society and the Caesar Kleberg Wildlife Research Institute publishers. 333 pages. Rosatte, R. 2000. Management of raccoons (Procyon lotor) in Ontario, Canada: Do human intervention and disease have significant impact on raccoon populations? Mam- malia 64 : 369-390. Rosatte, R., C. MacInnes, R. Taylor Williams, and O. Williams. 1997. A proactive prevention strategy for rac- coon rabies in Ontario, Canada. Wildlife Society Bulletin 25: 110-116. Rosatte, R., D. Donovan, M. Allan, L. Howes, A. Silver, K. Bennett, C. MacInnes, C. Davies, A. Wandeler, and B. Radford. 2001. Emergency response to raccoon rabies introduction into Ontario. Journal of Wildlife Diseases 37: 265-279. Rosatte, R. M. Allan, R. Warren, P. Neave, T. Babin, L. Buchanan, D. Donovan, K. Sobey, C. Davies, F. Mul- doon, and A. Wandeler. 2005. Movements of two rabid Raccoons, Procyon lotor, in eastern Ontario, Canada. Canadian Field-Naturalist 119: 453-454. Rosatte, R., K. Sobey, D. Donovan, L. Bruce, M. Allan, A. Silver, K. Bennett, M. Gibson, H. Simpson, C. Davies, A. Wandeler, and F. Muldoon. 2006. Behavior, move- ments and demographics of rabid raccoons in Ontario, Canada: management implications. Journal of Wildlife Dis- eases 42: 589-605. Williams, E. S., and I. K. Barker. 2001. Infectious diseases of wild mammals. Iowa State Press. 558 pages. NOTES 213 FIGURE |. Map depicting Raccoon movement to and from Ontario, Canada. Direction of arrows represents direc- tion of Raccoon movements. Arrows depicting move- ments of > 30 km represent Raccoon movement via transport trailers. One arrow in Michigan represents Raccoon movement on a train. Arrows depicting move- ments of < 30 km probably represent natural Rac- coon movements (i.e. not human assisted). Locations of arrows are approximate. Woodford, M., and P. Rossiter. 1993. Disease risks associ- ated with wildlife translocation projects. Revue Scientific et Technique de L’ Office International des Epizootics. 12: 115-135. Received 10 April 2006 Accepted 21 May 2008 214 THE CANADIAN FIELD-NATURALIST Vol. 121 Correlation Between Age Estimates for Elk, Cervus Elaphus, Using Tooth Wear/Eruption Patterns and Counts of Annuli in Tooth Cementum Rick RosAtTE!“*, ANDREW SILVER!, MARK GIBSON!, BRUCE CHISHOLM, and NorRM CooL? ‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, DNA Building, 2140 East Bank Drive, Peterborough, Ontario K9J 7B8 Canada *Canadian Food Inspection Agency, Edmonton, Alberta, T6H 5T6 Canada 3Parks Canada, Elk Island National Park, Fort Saskatchewan, Alberta T8L 2N7 Canada 4Corresponding author e-mail: rick.rosatte @ ontario.ca Rosatte, Rick, Andrew Silver, Mark Gibson, Bruce Chisholm, and Norm Cool. 2007. Correlation between age estimates for Elk, Cervus elaphus, using tooth wear/eruption patterns and annuli in tooth cementum. Canadian Field-Naturalist 121(2): 214-215. Two different techniques, (1) tooth wear, tooth eruption patterns and other attributes and (2) counting annuli and seasonal growth zones in the cementum of incisor teeth, were used to estimate the age of Elk, Cervus elaphus, that were relocated from Alberta to Ontario, Canada. Age estimates for Elk ranged from 2 to 20 years, and a significant relationship was found between the ages acquired by the two methods. Critical to acquiring accurate age estimates for Elk were staff with extensive skill with respect to the aging technique utilized. Key Words: Elk, Cervus elaphus, age estimates, annuli, cementum, tooth wear. During 1998-2001, 443 Elk (Cervus elaphus) were acquired from Elk Island National Park (EINP), Al- berta, Canada, and released in four different areas of Ontario, Canada (Rosatte et al. 2002; Rosatte et al. 2007). Elk were aged according to tooth wear, eruption patterns (deciduous and permanent teeth), colouring and staining of teeth, as well as by comparison to a reference sample of known-age Elk jaws, during pro- cessing at EINP (Hudson et al. 2002) (Figure 1). By about two years of age the upper pair of permanent in- cisiform canines erupt (the ivory) and by about 3 years of age, the 4 pairs of permanent incisors have erupted. The weight of the animal was also considered when estimating the age of Elk at EINP; e.g., the weight of 7-month-old calves, 1¥2-year-old, and = 22-year-old Elk was approximately 100-120 kg, 130-170 kg, and > 180 kg (+ 10 kg), respectively. The weights of the Elk were checked following age estimation to ensure that the age estimate fell into the above age/weight categories. During 1999-2004, incisor teeth were collected op- portunistically from Elk that died in Ontario post- release. These teeth were aged via counting annuli and seasonal growth zones in tooth cementum (Figure 2). The annuli or rings in the cementum represent yearly growth. By counting the number of annuli in the cementum an estimate of the animals age can be det- ermined (Johnston et al. 1999). The age estimates were compared with those obtained during processing at EINP after the tooth wear age estimates were extrap- olated to coincide with the time of death of the Elk in Ficure 1. Photo of an Elk (Cervus elaphus) mandible showing incisor, premolar, and molar teeth. Outer edge of tooth FiGurE 2. Longitudinal section (decalcified, Harris modified hematoxylin stain using bright field illumination) of an incisor tooth (100x magnification) from an Elk (Cervus elaphus) depicting annual growth lines (annuli) in the cementum. Estimated age is 20 years. sue and soaked in 70% ethanol for 5 minutes to prevent Ontario. Staff involved in aging had many years of ex- perience with the aging technique. Incisor teeth were extracted (via tooth extractors) from a sample (7 = 10) of Elk that died in Ontario post-release. Two yearling bulls, for which the age was known, served as controls. Teeth were cleaned of tis- the transmission of infectious pathogens. They were then preserved in 10% formalin or frozen at -20°C. Teeth were sectioned using an Isomet double-bladed saw (Buehler Isomet, Techmet, Scarborough, Ontario) according to Johnston et al. (1999). Each section (60- 100u) was mounted on a microscope slide with Per- 2007 mount and covered with a cover slip. Two microscopic techniques were used to determine an age estimate for each incisor. To distinguish juveniles from adults, the undecalcified, unstained sections were viewed under a compound light microscope (SOx and 100x magni- fication) using polarized light to view seasonal grow zones. In the majority of sections (especially older ani- mals), the annuli were not distinct and not easily dis- cerned under polarized light. These were decalcified (according to Johnston et al. 1999), stained (Harris modified hematoxylin), and placed in 70% ethanol for 2-3 minutes. Slides were then air-dried, covered with Permount and a cover slip, and examined under the microscope (unpolarized light) to count the number of annuli to obtain an estimate of age (Johnston et al. 1999) (Figure 2). Age estimates for Elk acquired by tooth wear (and other indicators as noted above) and annuli or seasonal growth zone counts in incisor teeth, ranged between 2 and 20 years of age (Figure 3). The first week of June was used as the birth date for Elk (Hudson et al. 2002). A simple linear correlation and a f statistic (Zar 1974) were used (Statistica 6.0 software, StatSoft, Tulsa, Oklahoma, USA) to measure the relationship and level of significance between tooth wear and counting annuli for age determination in Elk. The correlation was found to be significant (P < 0.0001; t = 20.1; r = 0.9903; 7 = 0.98). There was a positive slope of 1.09 between the ages acquired by the two methods (Y = 1.0918 x = 1.261) (Figure 3). This study examined the relationship between two methods used to estimate the ages of wild Elk. Tooth wear estimates of age may be inexact as tooth wear could be affected by diet, soil type and substrate (Hud- son et al. 2002). However, as the Elk at EINP were in an enclosed (fenced) situation, these variables were fairly constant as their geographical location remained un- changed among years. Therefore, we believe the age estimates acquired by tooth wear (and other attributes that were examined) were fairly accurate. Some authors (Keiss 1969; Hamlin et al. 2000; Hudson et al. 2002) suggest that staining sections of teeth and counting an- nuli provides the most reliable estimate of age for Elk. Furthermore, Keiss (1969) and Hamlin et al. (2000) only found about a 50% agreement between tooth erup- tion/wear and cementum annuli counts and concluded that wear/eruption estimates are unreliable for aging Elk. On the contrary, in our study, although the sam- ple size was small, age estimates for Elk obtained by tooth wear correlated well with those acquired by counting annuli in stained and unstained sections of incisor teeth. We believe that the different result in this study was due to staff expertise with both aging tech- niques and that this is critical to obtaining accurate age estimates for Elk. Acknowledgments The Ontario Elk restoration and research program is supported by the Ontario Ministry of Natural Resources NOTES 215 ge i © 20 4 y = 1.0918x - 1.261 =e 7 = 0.9807 a3 c c oO a c 7 a o io) < 0 5 10 15 20 Age using tooth wear (years) FIGURE 3. Relationship between age of Elk (Cervus elaphus) determined by tooth wear and annual growth lines (annuli) in tooth cementum (n= 10). (OMNR), Wildlife Research and Development Section, Dr. J. Chris Davies, manager, and the OMNR, Wildlife Section, Deb Stetson, manager. Special acknowledg- ment goes to the staff of the OMNR, Rabies Research and Development Unit, Peterborough, Ontario, to Trent University graduate students, Peterborough, Ontario, to the Canadian Food Inspection Agency, Edmonton, Alberta, and to Parks Canada, EINP, Alberta (especial- ly Rob Kaye), who assisted with the acquisition of teeth for aging and/or the processing of elk at EINP. The manuscript was reviewed by Dr. J. Chris Davies, B. Stevenson, and P. Bachmann, OMNR. Literature Cited Hamlin, K., D. Pac, C. Sime, R. DeSimone, and G. Dusek. 2000. Evaluating the accuracy of ages obtained by two methods for Montana ungulates. Journal of Wildlife Man- agement 64: 441-449. Hudson, R., J. Haigh, and A. Bubenik. 2002. Physical and physiological adaptations. Pages 199-257 in North Amer- ican elk: ecology and management. Edited by D. Toweill and J. W. Thomas. Smithsonian Institution Press, Washing- ton, D.C. Johnston, D. H., D. G. Joachim, P. Bachmann, K. Y. Kar- dong, R. E. A. Stewart, L. M. Dix, M. A. Strickland, and I. D. Watt. 1999. Aging furbearers using tooth struc- ture and biomarkers. Pages 228—243 in Wild Furbearer Management and Conservation in North America. Edited by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. [CD Edition] Ontario Fur Managers Federation, Sault Ste. Marie, Ontario. Keiss, R. 1969. Comparison of eruption-wear patterns and cementum annuli as age criteria in elk. Journal of Wildlife Management 33: 175-180. Rosatte, R., J. Hamr, B. Ranta, J. Young, and N. Cool. 2002. Elk restoration in Ontario, Canada: Infectious dis- ease Management strategy, 1998-2001. Annals of the New York Academy of Sciences 969: 358-363. Rosatte, R., J. Hamr, J. Young, I. Filion, and H. Smith. 2007. The restoration of Elk (Cervus elaphus) in Ontario, Canada: 1998-2005. Restoration Ecology 15: 34-43. Zar, J. 1974. Biostatistical analysis. Prentice Hall Inc., Engelwood Cliffs, New Jersey, 620 pages. Received 6 March 2006 Accepted 14 March 2008 216 THE CANADIAN FIELD-NATURALIST Vol. 121 Predator-Prey Interaction Between an American Robin, Turdus migratorius, and a Five-lined Skink, Eumeces fasciatus E. NATASHA VANDERHOFE!: 2 'Department of Biology, University of Louisville, Louisville, Kentucky 40292 USA *Current address: Department of Biology, Francis Marion University, Florence, South Carolina 29501 USA; e-mail: evanderhoff@ fmarion.edu Vanderhoff, E. Natasha. 2007. Predator-prey Interaction between an American Robin, Turdus migratorius, and a Five-lined Skink, Eumeces fasciatus. Canadian Field-Naturalist 121(2): 216-218. I observed a predator-prey interaction between a juvenile American Robin (Turdus migratorius L.) and a juvenile Five-lined Skink (Eumeces fasciatus L.). Although Robins are considered omnivorous, there are no previous reports of a robin eating lizards although they have been recorded as occasionally taking snakes. I discuss the age of the individuals involved as it relates to prey capture and escape. Key Words: American Robin, Turdus migratorius, Five-lined Skink, Eumeces fasciatus, predation. The American Robin (Turdus migratorius Linnaeus) and Five-lined Skink (Eumeces fasciatus Linnaeus) are commonly found in deciduous forests, as well as rural and urban areas and parks, in eastern North America. However, although they inhabit the same areas, there have been no accounts of predator-prey interactions between these two species. The American Robin, the most common thrush species in North America, is a versatile forager, feeding largely on invertebrates dur- ing the spring and summer and shifting to a diet with more fruits in the fall and winter (Sallabanks and James 1999; Wheelwright 1986). Robins also occasionally consume vertebrate prey, including snakes (Davis 1969; Netting 1969; Richmond 1975; Erickson 1978), shrews (Powers 1973), and even fish (Bayer 1980; Kimball 1944). However, there are no published records of Robins pursuing or consuming any species of lizard (Sallabanks and James 1999). The Five-lined Skink is a small lizard associated with forest openings throughout the eastern United States and southern Ontario (Fitch 1954; Behler and King 1997). Juveniles and adults differ in their appear- ance; adults develop a solid brownish color, whereas juveniles have five light-colored stripes on a black body and a bright blue tail (Behler and King 1997). Juveniles can lose their blue tail (autotomy) during interactions with predators. This, along with their striped pattern, is thought to confuse predators and allow the skink to escape (Fitch 1954; Clark and Hall 1970; Vitt and Cooper 1986; Goodman 2006). Numerous mammalian and reptilian predators of the Five-lined Skink have been documented (Fitch 1954). Although several birds, including mimids, icterids, corvids, and accipiters, prey upon lizards (Cooper and Vitt 1985), the American Robin has not been known to prey upon lizards. Addi- tionally, there have only been a few confirmed avian predators of the Five-lined Skink, including the Amer- ican Kestrel (Falco sparverius) (Heintzelman 1964), Broad-winged Hawk (Buteo platypterus) (Fitch 1974), Cooper’s Hawk (Accipiter cooperii) (Toland 1985), Greater Roadrunner (Geococcyx californianus) (Brown 1963) and Red-shouldered Hawk (Buteo lineatus) (Fitch et al. 1946). I observed a predator-prey interaction between an American Robin and a Five-lined Skink on 24 August 2006 at Joe Creason Park in Jefferson County, Kentucky (38°12'N 85°42'W). The interaction took place in a field at the edge of a forested area and I used 10 x 43 binoculars to aid my observations. The initial observa- tions of two juvenile robins (J1 and J2), and the juve- nile skink were made from a distance of over 20 m. Subsequent observations of J2 were made from about 10 m. At 18:18 I observed J1 with a skink in its beak. At 18:19 J2 approached J1, at which point the skink escaped by jumping out of J1’s beak. The robins pur- sued the skink for approximately 30 cm; J2 captured the skink and carried it approximately 5 m away while J1 flew off. I moved closer and observed the robin han- dle and consume the skink over the next six minutes. The robin, holding the skink in its beak, beat the skink on the ground multiple times and then took several bites. The robin repeated this four times, stopping periodi- cally to stand alert and scan the area. The robin also took one large bite and swallowed a large portion of the skink. After the robin flew off, I examined the spot and found only the skink’s entrails. Closer examination of the site where the skink was initially captured revealed the skink’s blue tail. Although I was unable to measure the skink, based on the tail remnant and entrails the skink was approximately 90 mm. This is the first account of a predator-prey interac- tion between the American Robin and the Five-lined Skink. The Five-lined Skink spends most of its time under cover such as woody debris and thus may large- ly avoid predation from avian foragers (Fitch 1954). However, the ranges and habitat of both species over- lap, and it is likely that skinks may occasionally be pursued and successfully captured by robins. At least one other thrush species is known to consume skinks; Turdus caleanops of Japan has been recorded to prey upon juvenile Eumeces okadae (Hasegawa 1990). It is interesting that the interaction I observed took place 2007 between juveniles of both species. Previous studies have shown that juvenile robins are less skilled when foraging for invertebrates and fruits (Vanderhoff and Eason 2007). Therefore, one might not expect that a juvenile robin would attempt to capture a skink. How- ever, juveniles are often more likely than adults to pur- sue and consume novel food items (Wunderle 1991). Juvenile Five-Lined Skinks may also be less skilled at escaping predators than are adults. I did not witness the initial capture of the skink by JI so | am unsure of any anti-predator behaviors that the skink may have used at that time. However, many species of Eumeces flicker their vividly colored tails to distract predators (Cooper 1998), and studies have shown that juveniles that display this behavior are better able to escape from snakes (Cooper and Vitt 1985). Five-lined Skink populations may be more intermit- tent and less common in Ontario than in Kentucky where this interaction was observed. In recent years Five-lined Skink populations have been in decline in Canada, and under the Species at Risk Act the Five- lined Skink is listed as a species of special concern (Quirt et al. 2006). Human removal of woody debris has eliminated an important microhabitat for the skink, and the decline in appropriate microhabitat is thought to be one of the main causes for the skink’s decline in some areas of Canada (Hecnar and M’Clos- key 1998). Woody debris is important for nest sites, but also acts as an important refuge from predators (Hec- nar 1994), and removal of these refuges may increase the Five-lined Skinks’ encounter rate with avian pred- ators, like the American Robin, as well as increase the skinks’ likelihood of being captured. Birds and lizards are commonly studied organisms, but documentation of predator-prey interactions be- tween these two groups is rare (Blomberg and Shine 2000), although avian predation is thought to be the selective pressure responsible for the vivid blue tail of juvenile skinks (Cooper and Vitt 1985), little informa- tion exists on interactions between birds and skinks. Both quantitative and qualitative information on pred- ator-prey interactions between birds and lizards is need- ed to add to knowledge of both groups. Acknowledgments I thank Catherine Byers, Chadwick Hanna, Lina Rifai, my advisor Perri Eason, and two reviewers for helpful comments on the manuscript. I also thank Louisville Metro Parks for allowing me to conduct my research at Joe Creason Park. Literature Cited Bayer, R. D. 1980. Novel use of an unusual food: American robin eating parts of fish. Journal of Field Ornithology 51: 74-75. Behler, J. L., and F. W. King. 1997. National Audubon Society Field Guide to North American Reptiles and Amphibians. Alfred F. Knopf, New York. 743 pages. NOTES 217 Blomberg, S. P., and R. Shine. 2000. Size-based predation by kookaburras (Dacelo novaeguineae) on lizards (Eulam prus tympanum: Scincidae): What determines prey vul nerability? Behavioral Ecology and Sociobiology 48 484-489. Brown, L. N. 1963. Status of the Roadrunner in Missouri The Condor 65: 242-243. Clark, D. R., and R. J. Hall. 1970. Function of the blue tail- coloration of the Five-lined Skink (Eumeces fasciatus) Herpetologica 26: 271-274. Cooper, W. E. 1998. Reactive and anticipatory display to de- flect predatory attack to an autotomous lizard tail. Cana- dian Journal of Zoology 76: 1507-1510. Cooper, W. E., and L. J. Vitt. 1985. Blue tail and autotomy Enhancement of predation avoidance in juvenile skinks. Zeitschrift fiir Tierpsychologie 70: 265-276. Davis, W. F. 1969. Robin kills snake. The Wilson Bulletin 81: 470-471. Erickson, D. B. 1978. Robin feeding upon snake. The Mur- relet 59: 26. Fitch H. 1954. Life history and ecology of the five-lined skink, Eumeces fasciatus. Kansas University Museum of Natural History Publications 8(1): 1-156. University of Kansas Publications, Lawrence. Fitch H. 1974. Observations on the food and nesting of the Broad-winged Hawk (Buteo platypterus) in northeastern Kansas. The Condor 76: 331-333. Fitch, H., F. Swenson, and D. F. Tillotson. 1946. Behavior and food habits of the Red-tailed Hawk. The Condor 48: 205-237. Goodman, R. M. 2006. Effects of tail loss on growth and sprint speed of juvenile Ewmeces fasciatus (Scincidae). Journal of Herpetology 40: 99-102. Hasegawa, M. 1990. The thrush 7iurdus celaenops as an avian predator of juvenile Eumeces okadae on Mayake-Jima, Izu Islands. Japanese Journal of Herpetology 13: 65-69. Hecnar, S. J. 1994. Nest distribution, site selection, and brood- ing in the five-lined skink (Eumeces fasciatus). Canadian Journal of Zoology 72: 1510-1516. Hecnar, S. J., and R. T. M’Closkey. 1998. Effects of distur- bance on five-lined skink, Eumeces fasciatus, abundance and distribution. Biological Conservation 85: 213-222. Heintzelman, D. S. 1964. Spring and summer Sparrow Hawk food habits. The Wilson Bulletin 76: 323-330. Kimball, J. W. 1944. A fishy bird story. The Auk 61: 646- 647. Netting, M. G. 1969. Does the robin eat DeKay’s snake? The Wilson Bulletin 81: 471. Powers, L. R. 1973. Record of a robin feeding shrews to its nestlings. The Condor 75: 248. Quirt, K. C., G. Blouin-Demers, B. J. Howes, and S. C. Lougheed. 2006. Microhabitat selection of five-lined skinks in northern peripheral populations. Journal of Herpetology 40: 335-342. Richmond, M. L. 1975. American robin feeds garter snake to its nestlings. The Wilson Bulletin 87: 552. Sallabanks, R., and F. C. James. 1999. American Robin (Turdus migratorius). Birds of North America (462). Ameri- can Ornithologists’ Union, Philadelphia, Pennsylvania. 27 pages. Toland, B. 1985. Food habits and hunting success of Cooper’s Hawk in Missouri. Journal of Field Ornithology 56: 419-422. 218 Vanderhoff, E. N., and P. K. Eason. 2007. Disparity be- tween adult and juvenile American Robins Turdus migra- torius foraging for ground invertebrates and cherry fruits. Ethology 113: 1212-1218. Vitt, L. J., and W. E. Cooper. 1986. Tail loss, tail color, and predator escape in Eumeces (Lacertilla: Scincidae): Age- specific differences in costs and benefits. Canadian Journal of Zoology 64: 583-592. THE CANADIAN FIELD-NATURALIST Vol. 121 Wheelwright, N. T. 1986. The diet of American robins: An analysis of U.S. Biological Survey Records. The Auk 103: 710-725. Wunderle, J. 1991. Age-specific foraging proficiency in birds. Pages 273-324 in Current Ornithology. Edited by Dennis Power. Plenum Press, New York. Received 15 December 2006 Accepted 20 March 2008 Book Reviews Book Review Editor’s Note: From this issue forward we will use the current currency codes. Thus Canadian dollars are CAD, U.S. dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars are AUD, and so on. You will find these are the codes now used by financial institutions and internet currency converters. I will include an updated note for the next few issues as a reminder. ZOOLOGY Atlas of Breeding Birds of Ontario 2001 — 2005 By M. D. Cadman, D. A. Sutherland, G. G. Beck, D. Lepage, A. R. Couturier. 2008. Published by Bird Studies Canada, Environment Canada, Ontario Field Ornithologists, Ontario Ministry of Natural Resources, Ontario Nature. xxii + 706 pages. 96.00 CAD Cloth. There’s nothing like a good, hefty book to make reading in bed difficult; there’s also nothing like a good, hefty book that’s full of all the information it’s supposed to have. This second atlas of Ontario birds is both. Like all recent atlases, this book has enough coffee-table appeal to be of interest to non-birders, and could in fact, entice them into the fold. This is not at the expense of the data, since the book typically pres- ents the data thoroughly and well. I am very happy that Ontario (as most jurisdictions) chose to publish a book, and not a CD-ROM, as did Oregon...people will always be able to read books...CDs may already be on their way out. Given that this is a second atlas for Ontario, the opportunity to compare results with the first atlas was there, and was used. The reader can see instantaneous- ly whether 10 x 10 km squares (in southern Ontario) or 100 x 100 km blocks (in northern Ontario) have had a change in breeding status. This segues me to describe the regrettable, but unfortunately practical practice of giving less detail to the north than the south; being a native born-and-raised northern Ontario boy...this has always bothered me, but as stated, I see the practicality. What I don’t understand, though, is the situation where the distribution of a bird like the Palm Warbler is shown as six squares on a large, and large scale south- er Ontario map, even though the distribution of 99% of the population is shown on a smaller, and smaller scale, northern Ontario map. At least northern Ontario is covered, unlike the otherwise-fantastic Quebec atlas, which didn’t attempt to describe birds in the northern two-thirds of that province. This atlas has several important chapters of pream- ble, though, like the Alberta atlas, these are essential- ly limited to what is necessary to best understand the Species accounts. Other jurisdictions have provided a greater diversity of introductory chapters to create an even more useful atlas — these include chapters on the local history of birding and ornithology, aboriginals and their relationships with birds, conservation, and others. Although not necessary, chapters like these would have enhanced the Ontario atlas. Somewhere in the fleet of introductory chapters or appendices, one would have expected to come across a very basic piece of information — the number of squares in Ontario. There are tables in the introducto- ry chapters and appendices which tell you the num- ber of squares in which a species was found...obvi- ously, without knowing the total number of squares, the former information is less valuable than it could be. As with the Alberta atlas, each species account is given one double-page spread. This is convenient for the reader and allows for enough text, graphs and maps in most cases. It must have been an editorial decision to do this — surely some species could have done with three pages, without getting verbose. However, a two- page allotment does give a decent treatment to all. The accounts are divvied into appropriate subheadings in Distribution and Population Status, Breeding Biolo- gy and Abundance. A multitude of authors wrote these species accounts, and even so, I didn’t find a single one lacking. All were well, to me extremely well, written — congratulations go to the authors, reviewers and editors of the species accounts. By using the multi-author approach, the workload is spread out and the project takes advantage of the areas in which these folks spe- cialize. This atlas incorporated point counts into its method- ology as an important way to quantify many species. Very intuitive maps show the distribution of different population densities. By doing this, the third atlas will be better able to quantify population changes on the whole, as well as shifts in population centres and bor- ders within Ontario. In contrast, there are many other surveys from which information could have been incorporated into the atlas; not all species are effectively sampled using point counts. For example, the Ontario Nocturnal Ow! Survey data could have been summarized for each of Ontario’s owls; this would not have been a daunting task. The Breeding Bird Survey data, along with the 19 220 data from other sources, were incorporated into the Oregon BBA, to the overall benefit of atlas users. There are some technical points to the data presen- tation that I did not find appealing or useful, though I will only mention the bigger ones here. For each spe- cies, there is a histogram which illustrates the probabil- ity that a person spending 20 h birding will encounter a species in a square from one of the five biogeograph- ic regions of Ontario (and another bar illustrating the data for all of Ontario). On the y-axis are the labels for the five regions and the whole of Ontario; the x- axis shows the probability. There are two bars for each region, one for the first atlas, one for the second. Hav- ing both bars does give the reader a good idea of the change in abundance of the species. However, there are two issues with these histograms. The first is that the exact value of the probability is put at the end of each of the twelve bars...isn’t that what the x-axis is for? For those very few people who need to know an exact value (e.g., a 59.9% chance of finding a House Finch in Lake Simcoe-Rideau) instead of the ballpark x-axis value of 60%, those data can be retrieved from the atlas project. For the rest of us, the data labels (made popular by many software packages) are simply so much clutter. Secondly, regardless of the data, the x-axis is always calibrated in 20% increments, from 0-100%. That means, for species like the Ruddy Duck and Wilson’s Phalarope, where eight of the twelve bars are at 1% or less, the reader barely sees the bars...why not scale ~ the axis from 0-10% to show the data more effectively? This becomes ridiculous with birds like the Worm-eat- ing Warbler and Northern Wheatear, which have some of the six categories blank, and all of the others with non-existent bars labelled at 0.0%. Each of the species accounts features one photo- graph of the bird and sometimes a habitat and nest shot as well. The quality was from good to great... there were a few shots that I would have replaced, but nothing serious here. To increase the visual appeal, I would have included more habitat shots; both The Birds of British Columbia and Birds of the Yukon Ter- ritory had more of this, and I think that added quite a bit to those books [Neither of those books are atlases per se, but both do show distribution and breeding records]. A few photographers contributed many of The Return of Caribou to Ungava A. T. Bergerud, Stuart N. Luttich, and Lodewiih Camps. 2007. McGill-Queen’s University Press, Montreal, Quebec. 2007. 586 pages. 49.95 CAD Cloth. This is the most comprehensive book on Caribou ecology and predator-prey relationships that has ap- peared in many years, perhaps ever. Not only is the research seminal, but the authors systematically dis- mantle paradigms that have been in vogue for years. According to the authors, Caribou biologists have THE CANADIAN FIELD-NATURALIST Vol. 121 the pictures, though overall there was a good diversity of photographers; that many peoples’ works get fea- tured is always good to see in a volunteer effort...so this is definitely a plus in my mind. Finally, the cover photo. A Prairie Warbler? Really? Why? This bird was recorded in only 45 squares in Ontario — that’s less than 0.5% of the total. If I were to ask you to name the best-known bird of the Yukon, what would it be? And yes, it’s on the cover of their book. The widely-dispersed Red-tailed Hawk was a fine choice for the cover of the first atlas of Maritime birds. Surely something much more widespread and charismatic like their provincial bird, or one known to almost anyone who feeds birds in Ontario, like the Dark-eyed Junco, would have been more appropriate. The decision to have the very local Prairie Warbler as the coverbird just boggles my mind. Overall my impression is that this is quite a good book that could have so easily become a great book. Ontario atlassers should be very satisfied with their second atlas — it largely presents the efforts of their long hours well; users of this book will be faced with a lot of information that is well-organized, and pleas- ing to read. Literature Cited Adamus, P. R., K. Larsen, G. Gillson, and C. R. Miller. 2001. Oregon Breeding Bird Atlas. Oregon Field Ornithologists, Eugene. CD ROM. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. Kaiser, M. C. E. McNall and G. E. J. Smith. The Birds of British Columbia Volume 3. University of British Columbia Press, Vancouver. 693 pages. Erskine, A. J. 1992. Atlas of Breeding Birds of the Maritime Provinces. Nimbus Publishing Company and Nova Scotia Museum, Halifax, Nova Scotia 270 pages. Gauthier, J., and Y. Aubry. 1996. The Breeding Birds of Québec: Atlas of the Breeding Birds of Southern Québec. The Province of Québec Society for the Protection of Birds and the Canadian Wildlife Service. Montréal. 1302 pages. Hess, G. K., R. L. West, M. V. Barnhill and L. M. Fleming. Birds of Delaware. University of Pittsburgh Press. Pittsburgh. 635 pages. Sinclair, P. H., W. A. Nixon, C. D. Eckert, and N. L. Hughes. 2003. Birds of the Yukon Territory. UBC Press, Toronto. 595 pages. The Federation of Alberta Naturalists. 2007. The Atlas of Breeding Birds of Alberta: A Second Look. FAN, Altona. 626 pages. RANDY F. LAUFF St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada wasted the last 50 years measuring lichens on winter ranges, when they should have been documenting plant production on summer ranges. Wolves, along with human hunters, both limit and regulate caribou popu- lations, not habitat. Food on the summer range only regulates at high densities and only after the range has been overgrazed. Wolves are driving Woodland and Mountain Caribou to extinction. Caribou populations where Wolves are absent maintain densities 100 times 2007 greater than predated herds. The reason arctic Caribou migrate to barren ground calving areas is to avoid Wolves tied to den sites at treeline. Even so, if it were not for periodic rabies epidemics, migratory Caribou populations would be severely limited by Wolf pre- dation. Volcanic eruptions half a world away trigger population declines in arctic Caribou at high densi- ties. And this is just for starters. The book chronicles the history of the George River Caribou in Labrador and Quebec from near extinc- tion during the early 1900s to an estimated 600 000 animals before the herd declined. The authors explore the various hypotheses that have been proposed to explain these fluctuations and present data set after data set to separate between competing explanations. In addition, the authors discuss virtually every other Caribou population that has been studied in North America, Scandinavia, and beyond, including the dif- ference between migratory and sedentary herds, which is key to understanding this species’ ecology. To the south of Ungava are small non-migratory populations of Woodland Caribou that are being driven to extinction by Wolf predation. But in reality, Moose and White-tail Deer are to blame. Historically, these areas sustained low-density, widely-spaced Caribou that in and of themselves could support few or no Wolves. Moose and whitetails were absent. But since the early 1900’s, Moose and whitetails have extended their range providing alternative prey for Wolves, where none existed before. The Wolves then drive the more vulnerable Caribou ever downward. That is to say, the addition of alternative prey did not buffer predation on Caribou, but instead increased predation pressure, contrary to what many people would expect. But that is not the most intriguing part. Why were Moose and whitetails absent historically and prehistorically? The authors contend that logging changed coniferous forests to secondary deciduous species favored by Moose and whitetails. In this I believe they erred because fire history data indicate there was always a strong deciduous component in those forests. Besides, Moose and Whitetails can sur- vive on a winter diet of Balsam Fir, as they do on Isle Royale and Anticosti Island. Instead, I believe that native hunters once kept eastern Moose populations in check, as I know native hunters did in western North America, where there are more Moose today than at any time in the last 12 000 years (see Alces 33:141- 164). Historically and prehistorically, native hunters extirpated Moose over large areas because, like the Wolves discussed above, humans had a multitude of alternative prey, including vegetal resources and fish unavailable to carnivores. As aboriginal hunting pres- sure declines, prey populations increase. In fact, the Book REVIEWS 221 authors note that the influenza epidemic of 1918 dec- imated native populations on Ungava, which in turn allowed Caribou to increase. I certainly commend the authors for presenting data on aboriginal peoples since the time Ungava was first inhabited and for describing how human hunting im- pacts Caribou. Most other studies of ungulate ecology begin with the premise that native people are irrele- vant because everything was a “wilderness” untouched by the hand of man prior to the arrival of Europeans: e.g., see The Kruger Experience. As | have explained elsewhere, however, this is a fatal error. The authors did not make that mistake, but I would suggest they need to look deeper into human evolutionary ecology. Take the seemingly random movements of Caribou, a subject covered at length in this book. Unfortunately, the authors neglected to consult Bin- ford’s data on Inuit Caribou hunters — see Numamint Ethnoarchaeology. One of the questions Binford asked was how do caribou hunters select a direction to hunt when they have no prior knowledge of where the Cari- bou are? The Inuit base their decisions on what we in the West would call mysticism. By careful observa- tion, however, Binford determined that Inuit pre-hunt behavior was simply a random number generator. That is to say, in these cases, the Inuit hunted randomly, which makes perfect ecological sense, odd though it may seem. If the Caribou moved in a predictable pattern, they would be easy prey for aboriginal hunters, as the au- thors note when the Ungava herd is forced by topog- raphy to cross the George River at Indian House Lake. If the hunters hunted in a predictable pattern, the Cari- bou would quickly learn to avoid the hunters, and the people would starve. The solution to the Caribou’s problem is to move as randomly as possible, while the solution to the hunter’s predicament is to hunt random- ly. This co-evolution occurred over thousands of years and probably is the only evolutionary stable strategy available to both Caribou and humans and then only because the Caribou’s range was vast and diverse. The authors note that even when Ungava Caribou num- bered only 15 000 animals, spread over an immense area, aboriginal hunting alone kept the herd from in- creasing. Using dog sleds, native hunters would follow caribou tracks for days, until the animals were killed or the trail lost. The Return of Caribou to Ungava should be read by everyone with even a passing interest in northern ecology, caribou management, or predator-prey rela- tionships. It should also be read by historians, anthro- pologists, and archaeologists. CHARLES E. KAy Utah State University, Logan, Utah 84322-0725 USA Dpape A Guide to the Mammals of China Edited by A. Smith and Y. Xie. 2008. Princeton University Press 41, William Street, Princeton, New Jersey USA 08540-5237. 576 pages. 60.00 USD Cloth. China, like many other areas outside North America and Europe, has had good field guides to the birds for some years. Finally it now has a decent guide to the mammals. Unlike the mammal guides for Africa and South America, this new Chinese book includes a more thorough coverage of the small stuff, such as rodents, pikas, shrews and the like. It was produced with contributions from nine people from the United States, the United Kingdom, Italy, and China. It is refreshing to see such a cooperative effort by a multi-national team in a region where politics often clash. The book covers all of China, including Taiwan and Xizang [Tibet]. The artwork is by Federico Gemma of Italy and is top quality. I am particularly impressed by the way he has occasionally moved away from the “classic” guide poses. For example, he portrays Tibetan Gazelle leap- ing through the air. This accentuates the gracefulness of this lovely species. At first it may appear that Gemma has an easy job because so little is known of Chinese mammals in the west. This is not true. We are familiar with the cute poster mammals — the Giant and Red pandas and the Tiger. Many animals are familiar zoo and park species — Muntjac [Red], Sika and Sambar Deer. Others are common species in more accessible places — Ermine, Caribou, Moose, Red Fox. Using these as a guide, the reader will soon see that the illus- trations are accurate. If you want to examine the accu- racy further, then you can verify details like the facial features of the Greater Horshoe Bat. However, not all of the species are covered by this fine artwork. I esti- mate that a little over 67% of the 556 species are depict- ed [although some of the non-illustrated species are in other, readily available, books]. The book has a great introductory section covering geography, the history of mammalogy in China, and conservation. This includes some useful maps and pho- tographs. The text consists of the typical sections on characteristics, distribution, natural history, additional comments, and status. I was pleased to find this infor- mation is current and honest. Chinese literature meant for the West often glosses over key problems. This is not the case here, and the status of the Tiger, for example, is clearly stated. The reasons for its decline are given frankly. As this book covers 81 species more than I have on my existing Chinese mammal list, I tried com- paring the two. I gave up after about 150 species because there were considerable differences. Many additional species, primarily from the fringe areas of Xinjiang, Xizang and Yunnan, were included and many others were omitted. I realised many of the missing ones had been assigned a new genus name making direct comparison more difficult. I did wonder why THE CANADIAN FIELD-NATURALIST Vol. 121 the Eastern Black Crested Gibbon (Nomascus nasutus) was not mentioned except in a list of subspecies. This is generally regarded as a full species and is consid- ered the most endangered primate. There are a few [in the low tens] on Huinan [plus a similar number in Viet- nam]. Other species like the Manchurian Zokor /Myo- spalax epsilanus] and the Chinese Scrub Vole [Micro- tus irene] are not even mentioned, yet are listed in Duff and Lawson’s Mammals of the World and other ref- erences. In addition, there are some changed common names — Onager is called a Kulan, for example. Also only five coastal species of marine whales out of over two dozen pelagic species are included. [However an Ap- pendix lists an additional 29 cetaceans]. Yet all five possible seals recorded, mostly as vagrants, on China’s coast are in the book. Another problem is character- ized by the distribution of the Northern Birch Mouse, Sicista betulina, given by some authorities as being in China, but others as being absent [Including this book]. I think that all of these differences reveal the status of the taxonomy of Chinese mammals and the need to in- crease research. I will now use this book as the author- itative list, but will likely still consider fringe species. The issue I have with the text is in the “characteris- tics” sections. The descriptions will be difficult to use in the field as there is very little comparison with like species. The descriptions are generic enough to over- lap and the illustrations are similar. For example, Jer- boas are very difficult to separate. I know how hard it is to see these small fast creatures [the Great Jerboa goes over 40 km/hr!] in the field. There is a key to each genus, but this is more useful for museum specimens. [In fact there were a few places where I felt strong museum overtones as opposed to field experience. However, the book is not called a field guide.] In addition to re-assigning genera, the authors have split the Eurasian Moose or Elk from the North Amer- ican Moose. In China there are two widely separated populations: one in Xinjiang [north west] and one in Heilongjiang [north east]. They have made this split based on the larger size, redder fur and bigger dewlap of the “American” Moose. No other reference makes this difference. The range maps plot actual museum and other rec- ords rather than shaded estimations of current distribu- tion. This is more precise and factual, but may not rep- resent today’s status and may be more of a historical record. This will mean the reader will have to interpret the data carefully so as not to get a wrong impression. I did spot a couple of errors. Savi’s Pipistrelle’s range is given as Xinjiang ... to Northern Europe. It actually is found in Greece, Italy and eastern Spain, not what I consider northern Europe. The index indi- cates that the Tibetan Antelope is on Plate 60 when it is depicted on Plate 57. I also question the range map of the Spotted Seal, shown with many specimens going 2007 far inland. | would be surprised if this pack-ice loving beast would go up fresh water rivers. Birds of the Saskatoon Area Edited by Anna L. Leighton, Jim Hay, C. Stuart Houston, J. Frank Roy, and Stan Shadick. 2002. Nature Saskatchewan Special Publication Number 23, Regina and Saskatoon Nature Society, Saskatoon. 345 pages. 40.00 CAD [30.00 CAD to Saskatoon Nature Society members] Paper. During my second encounter with the late J. Bernard Gollop in Regina some time between 1968 and 1970, the late Margaret Belcher and her sister, Mary Hous- ton, mentioned that his planned book on the birds of the Saskatoon area would probably be the next of the esteemed Saskatchewan Natural History Society spe- cial publications to appear. Unfortunately, Bernie did not live to see it in print, but 77 authors and six edi- tors finally brought his 1965-1988 efforts to fruition. The long gestation period produced an outstanding addition to Saskatchewan’s ornithological literature. This largest book in the series to date is also the first to be co-published [by the Saskatoon Nature Society] and is the fifth in the Manley Callin series on birds of specific areas of the province. It is jammed with infor- mation, with even the covers and fold-outs from the covers containing information [an area map, a Landsat image of the study area and a few tables]. Introductory material covers the first 68 pages of text, including pub- lication blurbs, tables of contents, drawings, maps and photographs, a dedication to Bernie Gollop, a preface, a foreword by Farley Mowat, a poem, acknowledg- ments, chapters on birding aids, histories of birding in the area and of the project, a detailed chapter on habi- tats and other ecological influences on the local avi- fauna, a detailed chapter on “hot spots” [essentially a bird-finding guide], a list of “unpublished” sources [in- cluding published seasonal compilations] consulted, a list of abbreviations, a topography and a glossary. These are well illustrated with figures, maps and drawings. The bulk of the text consists of 340 species accounts in taxonomic order except for one [Prairie Warbler] that was added while the book was in late stages of production. Eight appendices, a list of references cited and indices to English and French bird names close the book. The species accounts are preceded by a two-page introduction, explaining their contents and definitions used for seasonal occurrence, status in the area, abun- dance, tables of arrival and departure dates and other general information. Species accounts range in length from two paragraphs to two and a quarter pages, de- pending on regularity of occurrence in the area, amount of knowledge of the species’ life history features in the Book REVIEWS 493 Notwithstanding my comments, this is a much- needed guide and it will be indispensable for natural- ists and mammalogists for years to come. Roy JOHN 2193 Emard Crescent, Ottawa, Ontario. K1J 6K5 Canada area, numbers banded in the area, and other factors, such as change in status, locally, provincially, and/or generally. If migratory, earliest and median spring ar- rival and latest and median fall departure dates are included. If banded, numbers of the species banded in the area are listed, as are numbers (if any) recovered in- side and/or outside the Saskatoon bird area. If known to nest, data are included on whatever details of nest- ing biology, chronology, habitat and/or success are available. Remarks on population changes and their believed causes are also included if appropriate. Draw- ings accompany several species accounts. Current En- glish, French and scientific names and one or more four-letter species code(s) are indicated for each spe- cies. Former English names are also indicated for sev- eral. Nineteen nesting species are listed as hosts of Brown-headed Cowbird documented in the Saskatoon area in both the cowbird and the host species accounts. Several miscellaneous observations of biological, his- torical or population trend interest are also mentioned, often in a “side bar” or box. The book ends with a series of eight appendices (on annual Christmas, May Day and fall counts, local Breeding Bird Surveys, numbers of each of 79 species banded in the area, numbers of birds banded by each bander who has banded in the area, nest-box dimen- sions and requirements for 12 species and a list of 12 theses based on birds studied within the area), a list of references cited and indices in English and French to the first page of each species account. This volume continues the trend of improving on the already high quality of earlier contributions in this series, using as many sources of published and unpub- lished sources as possible. Although some variability in style is inevitable in a book with so many authors and editors, the quality of writing and accuracy of information are high throughout. I noted a few minor grammatical deviations, but few errors or omissions of substance. The statement (page 284) that Dickcissels “continue to breed” in southern Manitoba implies reg- ular occurrence there. While they occasionally nest there, rarely in fairly large numbers, their occurrence is irregular, often with several years between sightings (Davis et al. 2003). Minor errors include the omission of the hyphen from screech-owl (page 68), reference to Townsend’s Solitaires as “thrush-sized™ (page 232). implying that they are not thrushes and a few instances of species names in lower case. The vocalization of Baltimore Oriole referred as a “call” (page 291) ap- 1) tO TS pears to be the song. Cross-referencing between species accounts is quite thorough, but the list of 19 known local cowbird host species (page 290) omits Hermit Thrush (page 235) and Common Yellowthroat records (page 259). Also, a record of House Sparrows nesting in Bank Swallow holes is mentioned in the swallow account, but not in that of the sparrow and a case of hybridization between Eastern and Mountain bluebirds is mentioned only in the Eastern species account. Odd omissions from the glossary are the terms “albinism” and “leucism,’ especially since the less common “mel- anism” is included. An odd feature, though not an error, is the use of U.S. spelling for behaviour, centered, colour, favoured, harbour, moulting, neighbourhood and variants of these. The difference in slope of the bill between Tundra and Trumpeter swans, a feature much easier to see than the visual differences noted, would have been a useful addition to the identification tips in the swan accounts (pages 89 and 90). The head- er for Wilson’s Snipe indicates that two nesting records have been documented in the area, whereas three are mentioned in the text. The record of a robin feeding a young cowbird (page 237), while interesting, is not definitive evidence of successful parasitism by cow- birds on robins, as non-parental birds sometimes feed offspring of birds of other species (for example, Salt 1997). Most references cited are included in the ref- erence list, but one by Allen (1952) is cited as both Allan [incorrectly] and Allen on page 138, one by Dunn et al. (2000) cited on page 189 is not listed un- less 2002 is intended, the title is missing from a paper by Greenberg and Droege, the pages of the 1996 owl paper by C. S. Houston should be 125-133 [not 25- 132] and the second author of the book by Robbins et al. should be B. Bruun, not B. Brown. Although such regional works serve primarily to document occurrence, dynamics and changes in local THE CANADIAN FIELD-NATURALIST Vol. 121 bird populations, they also serve to document various life history observations that the author(s) may other- wise never find time to write up. Examples of such tid-bits in this book include fondness of Snow Geese for peas, television tower casualties of Green-winged Teal, albino and/or “partial albino” Ruddy Duck, mag- pies, Mountain Bluebird, Yellow-rumped Warbler and Yellow-headed Blackbird, a pair of Red-necked Grebes incubating golf balls, predation by American Crows on Eared Grebe eggs, cormorants perching on power lines, a dry land nest of an American Bittern, Willet courtship flight, magpies chasing and catching bats and eating berries, a Bank Swallow colony in a hole in a wheat field, House Sparrows nesting in Bank Swal- low holes, robins nesting on antlers and large numbers of starlings flycatching. Another measure of success in such regional works is the degree to which they stimulate publication to fill in gaps and document changes. Judging by the num- ber of notes and papers on bird observations and re- search in the Saskatoon area that have appeared in ornithological and natural history journals since “Birds of the Saskatoon Area” appeared in print, this volume has succeeded in that respect as well. It is an impor- tant contribution towards a forthcoming two-volume account of the birds of Saskatchewan. Literature Cited Davis, S. K., G. Walz, and P. Taylor. 2003. Dickcissel Dickcissel d’amerique Spiza americana (Gmelin). Pages 372-373 in The birds of Manitoba. Edited by P. Taylor. Manitoba Naturalists Society, Winnipeg. Salt, J. R. 1997. MacGillivray’s and Black-throated Gray warblers compete to feed fledgling cowbird. British Columbia Birds 7: 15. MARTIN Mc NICHOLL Apartment 105, 8752 Centaurus Circle, Burnaby, British Columbia V3J 7E7 Canada Book Review Editor’s note: In a brainstorming session between the Editor, Francis Cook and myself, on future directions, we agreed I would investigate suitable websites and, if appropriate, include their reviews. I proposed we use Brian Coad’s site as a test case, hence the following review. If others know of similar suitable sites [Cornel University, USDA and USGS come to mind] please e-mail me [r.john@ rogers.com] with your suggestions. Brian Coad’s Ichthyology Site http://www.briancoad.com/main.asp?page=fishdictionary.htm By Brian Coad. 2008. Unlike books, websites are not static. This is the case with Coad’s site, which is defined as a work in progress. What it currently contains is sections on Projects, Fishes of Canada: Annotated Checklist, Dictionary of Ichthyology, Freshwater Fishes of Iran, Fishes of Canada’s National Capital Region and Fresh- water Fishes of Iran. The first section lists the author’s eight current proj- ects, with a few sentences of explanation. The checklist of Canadian Fish is a straightforward list of scientific and common names in both languages, plus occasion- ally a few comments. The dictionary, co-authored by the late Don E. McAllister, defines terms specifically used in the study of fish and includes some generally used common words. The Freshwater Fishes of Iran has individual species accounts arranged by genera. These start with an intro- duction to the genus. Each detailed account has sec- tions on the Common name in English and Farsi with the phonetic pronunciation, Systematics, Key charac- ters, Morphology, Sexual dimorphism, Colour Size, Distribution, Zoogeography, Habitat, Age and growth, Food, Reproduction, Parasites and predators, Econom- ic importance, Conservation, Further work and Sources. 2007 These accounts are illustrated [black-and-white line drawings] and give the reader a very exhaustive under- standing of the individual species. Fishes of Canada’s National Capital Region [NCR] has similar coverage to the above. Also included are photographs of out-of-water fish where these are avail- able and range maps within the National Capital Region. I decided to try out the site to identify small fish I saw in Mud Lake in Ottawa. After comparing descriptions and other information I decided it might be a Central Mudminnow. To be certain I would need the fish in hand. I might raise eyebrows, or worse, if I started netting fish in a public park. The Freshwater Fishes of Iraq contains only a checklist and bibliography at this date. Some of the advantage that this web site [and all others] have is that material [such as photos] can be included as required, poor quality photos can be re- placed if better become available, multiple photo can shows all features (not the case in a book where cost and space are limitations), text and other errors can be corrected, feedback from readers is easily incorporated, new information can be added as science progresses, in some cases differing view points can be presented at length, saving on costs of paper and distribution BOTANY BOOK REVIEWS 995 over the book format. The site can be accessed from anywhere in the world [with Internet access] without needing to carry around hard copy, it is copied by stu- dents, researchers and naturalist for their own projects This site is a wonderful source of information in a subject area that is often difficult to access. It is very easy to navigate and loads quickly [but it is not as user-friendly as a book]. | am not one to load up my favourites folder unless it is a site I will use frequently. I suggest this is one of those sites, especially if you live in the Ottawa-Gatineau area (National Capital Region). I also need to trust the information supplied. The generic question remains in that sites like this have not been reviewed as a book would be: Readers on the internet will need to verify the quality of a website before trusting the data. As Brian Coad is a respected research scientist (at the Canadian Museum of Nature) with extensive field experience in the Ottawa District and Iran, this is not an issue, despite its being a person- al, rather than an institutional site. At least you should take a look at it as | am sure you will learn as I did. Roy JOHN 2193 Emard Cresent, Ottawa, Ontario K1J 6K5 Canada Exploitation and Utilization of Chinese Spice Plant Resources By Zhang Weiming and Xiao Zhengchun et al. 2007. South- east University Press. 722 pages. Price 186.00 CNY. Spice plants are in a special category in the plant kingdom, and have been popularly used for various purposes in different countries or regions of the world. The history of the use of spice has even exceeded re- corded history of human beings. Early humans might have used spice plants to preserve meat, fish or other food, reduce the annoying odors of various foods, or increase a special flavor when cooking. Gradually, these began to expand the range of use of spice plants. Spice plants are closely related to the daily lives of human beings. Systematic research and summary of them is necessary, however, as this has not been well done before, at least in China. Now, this situation has been changed by the publication of the book Exploita- tion and Utilization of Chinese Spice Plant Resources by Zhang Weiming and Xiao Zhengchun et al. The authors have been engaged in research work on spice plant resources for a long time. They have conducted several research projects relating to Chinese spice plants in the last few decades, and obtained valuable and abundant data in this field. Meanwhile, they also collected other abundant information on spice plant resources at home and abroad. Based on these data and information, they produced this massive book. There are a number of climate zones in China. Dif- ferent environmental conditions allow for the growth of an abundance of plant species, including spice plants. Except for a few tropical species, most of the spice plants grow in China. China was one of the earliest countries to use spices. There are some records on the aromatic and spicy plants in the Book of Songs, the Book of Mountains and Seas, and other book that are more than 2000 years old. Over a long time, due to the various usage of different types of spice plants in cook- ing, the people living in various areas of China formed a variety of eating habits, reflected in the so-called eight genres of cuisine of Chinese dishes. Hence, we might say that one of the most important reasons that Chinese food (or any other regional food) enjoys a world-wide reputation is its unique use of spices in cooking. The indigeneous spice plants in China are abundant. Furthermore, over the years, China introduced many foreign-origin species. The book not only is a system- atic and comprehensive monograph on the spice plant resources of China, but also has its value as a refer- ence for other related countries. The book is divided into two parts. The first part is a general introduction to spice plants, including the history of spice plants, the general situation of Chinese spice plant resources, the main ingredients and the application of spice plants, the processing technology and the development of spice plants, and the strategy of the development of China spice plant resources. The second part is the special issues, including a total of 134 kinds of commonly used spices, and more than 200 spice plant species. For each spice plant species, the ecological environ- ment, history of use, chemical ingredients, compre- hensive methods of use, exploitation of related prod- ucts, standardization and cultivation techniques, and sO on were comprehensively introduced. The typically introduced spice plant species in this part include Capsicum frutescens, Allium sativum, Aglaia adorata, Allium cepa, Allium fistulosum, Allium porrum, Allium sativum, Artemisia vulgaris, Illicium verum, Foenicu- lum vulgare, Levisticum officinale, Mentha haploca- MISCELLANEOUS Arctic Hell Ship By W. Barr. 2007. University of Alberta Press, Ring House 2, Edmonton, Alberta T6G 2E1. 318 pages, 34.95 CAD Cloth. More of a human history than a natural history book, this tale covers part of one of the great sagas of Canada’s north. The Franklin Expedition left Britain with high hopes and the best resources available — well almost. It vanished in 1845 and spawned at least two dozen searches to solve the mystery of the disap- pearance. These efforts in turn have led to a flood of books and scholarly articles. So do we need another treatise? The answer is yes in this case. Barr’s contribution covers the 12" search by Captain R. Collinson aboard the HMS Enterprise with Robert John Le Mesurier M’Clure, as second in command, on the HMS Investigator. Barr gives us an extremely detailed description of life on one of Her Majesty’s ships in the 1800s. So often the other books concentrate on the leaders and their accomplishments, so we learn little of how the crew fares. Barr remedies this by telling us about many of the individual sailors. The “story” is very simple. This was a voyage that pro- duced limited results and suffered a lot of bickering between the officers. As a result, the ship’s, logs and letters contain irritating daily details that show what life was really like. Collinson was a good seaman and surveyor, but he was Clearly not a leader. He picked on very minor transgressions and turned them into major discipli- nary actions. He sowed unnecessary seeds of discord, interspersed by bouts of humanity. In hindsight, his biggest failure was that he did not do enough. He was charged with rescuing Franklin’s men and given the resources to do it, and yet he spent a lot of time sailing back and forth, unable to make a clear decision. His sledging trips were too little and too late. The other ships in his command (which got “separated” early in the expedition) went farther and did more. When THE CANADIAN FIELD-NATURALIST Vol. 121 lyx, Pimpinella anisum, Sesamum indicum, Zanthoxy- lum bungeanum, Zanthoxylum simulans, and Zingiber officinale, among others. The book was well written with few errors, in a style of writing that can be easily understood by non- scientists. As a comprehensive book, it is suitable for anyone who is engaged in the various areas of research and application of spice plants. Li DEZHI Lab of Urbanization and Ecological Restoration of Shanghai; National Field Observation and Research Station in Tiantong Forest Ecosystem of Zhejiang; Department of Environmental Science, East China Normal University, 3663, Zhongshan Road (N). Shanghai, China 200062 Collinson did make progress he found notes from either M’Clure or Dr. John Rae showing that they had searched the area earlier. In the tough conditions on board ship during an Arctic winter this debilitating caution and indecisiveness coupled with petty deci- sions on discipline must have made an unhappy ship full of exasperated people. Collinson returned to Britain, having failed to solve the Franklin issue, demanding courts-martial for his officers. The lords of the Admiralty wisely declined to take action. Public exposure of such silly shenanigans would do the service no good. The most notorious British naval captain of this era is Bligh. In spite of the fact that his reputation owes more to the movies than reality, Bligh was a superb navigator, excellent seaman and a relatively humane captain. He personally intervened in the trials of the captured mutineers, got some set free and obtained reduced sentences for others. He died as a Vice-Admi- ral. Collinson’s dithering and his stubborn need for absolute discipline would have made him a better can- didate for Hollywood’s spite, yet even this would be unfair. How would we all fare under such harsh con- ditions? Collinson did make Vice-Admiral, after serving at Trinity House, the establishment that maintained the aids to navigation, where he rose to deputy master. He remained at Trinity House until retirement, clearly a better bureaucrat than expedition leader. Perhaps the error lies with the lords of the Admiralty. They fell into that awful trap. They promoted a very technically com- petent person to a supervisor, a move that does not al- ways work. In particular, voyages to the Arctic, even today, need competent leaders. Collinson did not come close to Nansen, Shackleton, Amundsen, or even Scott in his ability to deal with people. While Barr has produced a minutely detailed doc- ument, using all available evidence, rather than a dis- 2007 tillation of the key events, it is those details that give the book its appeal. I found I wanted to read just one more paragraph each time before I put the book down. I was surprised when I realized I was over half way through the book and it seemed I had just started. As | stated initially this is really a history text, but it covers a key era in the story of Canada. The only important Kighteenth-century Naturalists of Hudson Bay By Stuart Houston, Tim Ball and Mary Houston. 2003. McGill-Queen’s University Press, 3430 McTavish Street, Montreal, Quebec, H3A 1X9. xxiv + 333 pages 49.95 CAD Cloth. In 1959, C. Stuart Houston and Maurice G. Street began to introduce prairie naturalists to the pioneering contributions of employees and other associates of the Hudson’s Bay Company and to expand our under- standing of the natural history of Canada’s Arctic and prairie regions (Houston and Street 1959) in the first of a long series of articles, books, notes and papers by Houston and numerous colleagues on biographies of specific contributors and the history of various aspects of these contributions. In this volume, Stuart and his chief collaborator (Mary I. Houston) bring many of the details of their earlier publications together, expand on the European collaborators of these naturalists and compare the Hudson’s Bay contributions with approx- imately contemporary efforts in South Carolina. They also collaborate with Timothy E. Ball in outlining the contributions of H.B.C. officials in documenting cli- matological details at various sites and their changes over a long time period and with Arok Wolvengrey and Jean Okimasis in documenting Cree names for numerous natural history species in the lands of the vast territory formerly controlled by H.B.C. The book opens with a series of introductory sec- tions, including a list of McGill-Queen’s “Native and Northern Series” (of which this is the 34"), a dedica- tion to William B. Ewart (whose efforts led to the mov- ing of the extensive Hudson’s Bay Company archives from England to Winnipeg), lists of contents, illustra- tions and tables, eight colour plates of birds originally published in two of four 1743-1751 volumes by George Edwards, acknowledgments, a list of supplementary documents available through the internet, a list of ab- breviations and a foreword by the Keeper of the Hud- son’s Bay Archives, Judith Hudson Beattie. The main text of the book (pages 1-142) consists of an unnum- bered introductory chapter, followed by 12 numbered chapters. Seven appendices occupy almost as many pages (pages 143-251). About 40 pages of footnotes, 25 pages of references and a 14-page index close the book. The main text starts with a short outline of the his- tory of Hudson’s Bay Company activities in North America and the role of HBC officials in documening natural history features of much of the prairie prov- Book REVIEWS 227 note of natural history concerns the surgeon, Edward Adams, for whom the Yellow-billed Loon was named Gavia adamsii. It is well worth the read if you love the Arctic, enjoy history or are a Franklin buff Roy JOHN 2193 Emard Crescent Ottawa, Ontario K1J 6K5 Canada inces, Ontario, Quebec, Nunavut and adjacent areas in collaboration with native North Americans and Euro- pean scientists. The first numbered chapter consists of brief biographies of seven European scientists who received and/or examined animal specimens collected by HBC personnel, described and often named them, and wrote the first accounts of their behaviour and other natural history features. The role of these speci- mens in developing Linnaeus’s system of classification of plants and animals is outlined. Eight chapters fol- low on the contributions and efforts by eight specific HBC officials (Peter Fidler, Andrew Graham, Samuel Hearne, Thomas Hutchins, James Isham, Alexander Light, Humphrey Marten and Moses Norton). The tenth numbered chapter summarizes these contributions, lists Hudson Bay area bird species mentioned in ten early manuscripts, provides several background details and notes several corrections and omissions in earlier pub- lications. The eleventh chapter, based primarily on four years of archival research by Ball, outlines weather, climate and climatic changes in the area, as document- ed through records of 30 or more years at 30 HBC posts. The final chapter outlines early contributions to North American natural history by early settlers of Charles Town, South Carolina, when that area was still a British colony. Brief biographies of two contrib- utors of meteorological data and two natural history collectors are included, as well as a list of species named as new in Mark Catesby’s 1729-1747 two-vol- ume treatise on the birds of South Carolina (58 from “South Carolina,” seven from other parts of eastern U.S. states, seven from the Caribbean, and one from “Quebec.” Seven appendices cover about 100 pages. The longest (about 50 pages) by the Houstons, Jean Oki- masis and Arok Wolvengrey lists words used in the eighteenth century in four Cree dialects for various taxa that live in the territory controlled by the Hudson’s Bay Company. Another lists departure and arrival dates and the captains of ships sailing between York Factory and Europe from 1716 to 1892. Smiles will cross the faces of Saskatchewan natural- ists, long aware that two of their most prominent mem- bers (Margaret Belcher and Mary Houston) were sis- ters, when they read that one of two ships captained by a Belcher (twice) was “Mary.” A two-page appendix by Deirdre Simmons outlines the history of the Hud- son’s Bay Company Archives. Others concern ten man- 228 uscripts that have variously been attributed to Andrew Graham and Thomas Hutchins, and plagiarism of Gra- ham by Hutchins. The Houstons provide an interesting account of their research in the archives, in London and at a symposium in the Orkney Islands that brought them to the conclusion that these manuscripts resulted from collaboration. Other appendices concern the ten- year cycles of various northern birds and mammals, the effects of nineteenth century trade on Trumpeter Swans, and the naming of Canada Goose prior to the official existence of Canada. I assumed that the latter was a combined reprint of two earlier publications in Blue Jay (Houston 1994, 1995), but some of it presents additional findings of historical research that was stim- ulated by the earlier notes. Naturalists interested in historical aspects of nature study will find plenty of interest in this volume. It is written well and researched thoroughly. I found no sub- stantial errors, although a few references cited were either not included in the literature lists or the publica- tion dates and/or author list in the text differed slightly from those in the literature list. One reference (Ross 1834) is listed in a footnote (page 256), but not in the references. Three references cited are not included in the literature list (Gmelin [1788 on page 140], McIn- tyre and Houston [1999] on page 269) and Rousseau [1969] on page 279, although enough information on the latter is included to enable a reader to track it down). The 1983 book by Peck and James should be listed as Volume 1. Footnote 39 on page 255 on William Wales should refer to Chapter 11, not 12. References to Appendix E and Figure 3.13 on page 52 apply to Appendix F and Figure 3.14; respectively. THE CANADIAN FIELD-NATURALIST Vol. 121 The abbreviated forms of the authors’ names from the more usual versions that they use in publications (Tim- othy E. Ball, C. Stuart Houston and Mary I. Houston) provides potential confusion for bibliographers who may think that other, presumably related, authors are involved, especially as their usual publication versions are used in some appendices. As noted by Meyer (2004), some of the appendices would be more appro- priate within the main text. Although the literature list is substantial, readers new to the subject who wished to follow up with further reading would have benefited from inclusion of more of Houston’s previ- ous biographies, book chapters, notes and papers on some of the subjects covered. Similarly, although bio- graphical references are included for many people, none are included for a few (for example, William Rowan) for whom several are available. None of these criticisms detract substantially from the overall high quality of this excellent contribution to the history of nature study in North America. Literature Cited Houston, C. S. 1994. How did the Canada Goose get its name before there was a Canada? Blue Jay 52: 141-143. Houston, C. S. 1995. Earlier use of the name Canada Goose. Blue Jay 53: 116-117. Houston, C. S. and M. G. Street. 1959. The birds of the Saskatchewan River Carlton to Cumberland. Saskatchewan Natural History Soci- ety Special Publication Number 2, Regina. Meyer, D. 2004. [Review of] Eighteenth-century naturalists of Hudson Bay. Blue Jay 62: 120-122. - MARTIN K. Mc NICHOLL Apartment 105, 8752 Centaurus Circle, Burnaby, British Columbia V3J 7E7 Canada Hunters at the Margin: Native People and Wildlife Conservation in the Northwest Territories By John Sandlos. UBC Press, 2029 West Mall, Vancouver, British Columbia V6T 1Z2. 360 pages, 85.00 CAD Cloth, 39.95 CAD Paper Professor Sandlos has written a very well-researched text on three major conservation issues. He examines the plight of the Bison, Muskox and Caribou in Cana- da’s north. The author has examined the questions raised about hunting of these three key species and the development and enforcement of laws relating to them. He has done this with a very sympathetic atti- tude towards Canada’s original inhabitants. He com- pares the lifestyle and requirements of aboriginals to the efforts of Ottawa’s bureaucrats to control hunting and to conserve these mammals. These two philoso- phies are rarely compatible, and this leads to conflict. I had a hard time reading this book. While I under- stand the author’s sympathy with Canada’s aboriginal people I found his constant innuendo very galling. When discussing the native viewpoint he writes with- out emphasis. When writing about the non-native peo- ple’s actions he frequently uses parentheses, causing the reader to doubt the adjective’s veracity. When defending the native people Sandlos ignores much per- tinent information. He proposes the natives are not the wanton killers portrayed by white policemen, wardens and travelers, but long before the arrival of the white man the natives hunted Mammoths, Giant Sloths and others to extinction. This was also before climate change pushed these creatures over the edge [as in Europe]. The latest book on Caribou (Bergerud et al. 2007) notes that after an influenza epidemic decimated native populations, Caribou increased [see review this issue]. In my own experience, two years ago at Cam- bridge Bay, I was informed the hunting ban on Muskox had been lifted a few months earlier and the town’s residents had shot the entire local herd. I had to travel out of town for an hour by truck and an hour and a half on foot to see distant Muskox on this island. Before the arrival of whites, there were few natives in the north; it was too hostile a climate. Living was easier on the plains with its abundant Bison. In fact the author provides evidence for this concept when he quotes a convicted poacher, Joseph Wakwan, as pre- ferring to hunt the larger Bison over the legal Moose. 2007 The natives moved north after they found they could ‘make money for guns and supplies by trapping Beaver. The demand for Beaver hats was purely European. I do not know if the author has been involved in creating laws, but I suspect not. This a long and com- plex process where nobody wins everything they want. At the conclusion you achieve compromise regulations that all can accept, more or less. To constantly criti- cize the Ottawa law-making bureaucrats, who were try- ing their best with the information and attitudes of the day, is a futile process. Trying to judge past events with today’s understanding is fun, but useless. National committees have learned long ago not to react too quickly to the latest reports and studies. It takes time and careful study to be certain any new data are valid. There have been many reports that claimed some- thing was bad, only to be followed by another that said it was good. We do not want our laws flip-flopping with every new announcement. If we applied Sandlos’s logic that white man’s laws do not match aboriginal situations and therefore real- ly do not apply, then I could ignore the laws of Eng- land as I am Welsh, a true English aboriginal. [The Welsh inhabited all of Britain before they were con- quered by the Angles and Saxons. These in turn were reduced to serfdom by the Norman French.] I, like all those in England, have to obey today’s laws. Such is history. The Canadian native people actually fared rea- sonably well for a conquered community. Compare their fate to that of the Inca after the arrival of the Span- ish or the original inhabitants of the Amazon after the Portuguese took over. The arrival of the white man brought white-man’s laws, as well as welfare, modern The Ornithologist’s Dictionary By Johannes Erritzoe, Kaj Kampp, Kevin Winker and Clifford B. Frith. 2007. Lynx Edicions, Montseny, 8, 08193 Bella- terra, Barcelona, Spain. 290 pages, 19 EUR Paper. This is a great little book that will be handy for all who study birds. I can easily slip out of my depth when reading a book or article because I do not understand some terms. Typically I stick a mark in the page and later go to my computer for help. Google searches are normally wonderful ways to fill in your knowledge gaps. But this also disrupts the continuity of your read- ing and lowers the value you can pull from the text. Having a resource that can sit by your side is a big advantage. So how accurate and comprehensive is the text? First, let me say the authors use the English spellings [colour etc.] This does not mean a European bias as there are plenty of references to purely North American terms [National Audubon, AOU etc.] and the U.S. spellings are included [spishing vs pishing!]. I looked up several definitions of words which are frequently misused and could only find one significant error. Parameter is not “any variable” but a constant in an BooK REVIEWS 299 medicine and current technologies. Whether these are good or bad is a moot question. When | wrote my last report on people of the north, the data sets I accessed on diet used by northern residents showed that a high proportion of junk food had replaced food gathered from the wild. In my view, this represents a poor choice; on the other hand, I do not recall any native village starving or freezing to death [as happened in the past, before the arrival of the white man]. We need to remember we all live in today’s conditions, which includes a plethora of laws which we must all obey [Whether we agree with them or not. Ask me about CITES. A well-meaning idea, whose concept I support. In practice it is often counterproductive to conservation and frequently punishes those who are promoting conservation. | Do not let my criticisms put you off reading this book. There is a lot of useful and thought provoking material embedded in the text. The questions and issues surrounding the native people and wildlife in the north are not easy to deal with and | doubt there will ever be “an” answer. This book makes a significant contribution to the continuing research and debate required to make rational decisions. I just wish it was written in a more balanced and critical style. Literature Cited Bergerud, A. T., Stuart N. Luttich, and Lodewiih Camps. 2007. The Return of Caribou to Ungava. McGill-Queen’s University Press, Montreal, Quebec. Roy JOHN 2193 Emard Crescent, Ottawa, Ontario K1J 6K5 equation that varies in other equations of the same gen- eral form [the classic example is the force of gravity — always constant in Ottawa but different yet con- stant in Banff.] This is an incredibly common error, even among scientists who should know better. I have a few other less significant comments, such as the needless use of interrelationship [relationship is fine, particularly in a dictionary] and I would not say the use of “mirrors” for the white tip of a gull’s wing is un- common as it is used in most of my books. I thought, too, that the authors could have included newer terms like sahel [note the lower case s] — A region having characteristics of a savanna or a steppe and bordering on a desert — as well as the classic definition of Sahel — a transition zone between the Sahara and the trop- ical forests to the south. Each letter of the alphabet starts with a neat black- and-white drawing of a bird. These are not identified, but I could easily determine the species in all but X and Y. Also I could not find the identity of the artist. Several pages in my original copy were greyed. As this is totally unlike the publisher, Lynx Edicions, who 230 normally produce beautiful, crystal clear copies, I con- tacted them. It appears this was an isolated incident; however, you should check the particular copy you intend to buy. Lynx sent me another, perfect copy. This will be of greatest use to non-professional avid birders who lack formal training. You can quickly resolve the occurrence of melanin, the nature of re- miges, the difference between granivorous and grami- nivorous, mimesis and mimicry and other sources of Pioneering Women in Plant Pathology Edited by Jean Beagle Ristaino. 2008. The American Phyto- pathological Society, 3340 Pilot Knob Road, St. Paul, Minnesota, U.S.A. 339 pages. 89.00 USD Cloth. As seen by one woman, an M.Sc. graduate in plant pathology from the mid-1970s, it would appear that women scientists in this field are rapidly proving their abilities to shine with accomplishment in what was once an all male profession; moving from being seen as pretty women hobbyists picking wild flowers to seri- ous taxonomists of vascular and non-vascular plants, many of which are plant pathogens. Agriculture has itself moved from its primitive beginnings of slash and burn to a precise science of food production in the 21* century. As the possibility that a woman may progress from a mere technician and housekeeper of the laboratory to the more stellar role of research scientist has im- proved, their research results have proved their abili- ties. The first woman plant pathologist, hired by the United States Department of Agriculture, was Effie A. Southworth, in 1887. Her most significant contri- bution was the description of the pathogen Colleto- trichum gossypii, the cause of anthrocnose on cotton, and the recommendation of measures of control of this disease. In 1895, Flora W. Patterson became the first woman mycologist at United States Department of Agriculture. She is remembered for the develop- ment of the U.S. National Fungus Collections, which are still of importance to mycologists and plant pathol- ogists today. She also was very involved with the in- spection of imported materials for invasive fungal pathogens, and intercepted the dangerous potato wart disease before it was imported into the United States for the first time. Another woman of importance was Edna Marie Buhrer, who brought forward the impor- tance of nematodes in plant diseases; up till 1920 the role of nematodes in crop production was considered unimportant. This led to various treatments of soils with Birdwatcher: The Life of Roger Tory Peterson By Elizabeth J. Rosenthal. 2008. The Lyons Press, Guilford, Connecticut. 437 pages. 34.95 CAD. Roger Tory Peterson’s parents could not afford to send him to college and he had no formal training in science. He moved to New York City, where he paint- THE CANADIAN FIELD-NATURALIST Vol. 121 confusion. This book would have rapidly ended my last bird-term argument — the meaning of leucism — if I had had it handy. It may not be as valuable to pro- fessionals, but I think they will likely find it useful too. Buy this one for yourself! Roy JOHN 2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada nematicides which results in improved crop produc- tion. In England also the importance of women in this field was recognized at Rothamsted Research Center and among those employed there and making a valu- able contribution was Mary Gwynne. She started her career in 1917, worked mainly on diseases of cereal crops, and in 1960 was awarded the Order of the British Empire for her contribution. Margaret Newton, at Mac- Donald College, McGill University, was a Canadian woman who made a considerable contribution to the knowledge of stem rusts of wheat, so important to the agricultural economy of Canada. At this period of time in the early twentieth century, women from European countries were also making their contributions. It is recommended that this book be on the shelves of every plant pathology department library, not only to show the valuable contributions made by these sci- entists, but also to show the courage and dedication of these pioneering women in the face of the prejudices of those times. This book also highlights the impor- tance of this field of science to the agriculture of the American continent and the safety of the world’s food trade. We have here a book about pioneering women plant pathologists who have overcome prejudice to make considerable contributions to the economy of North America. Pioneering Women Plant Pathologists is edited by Jean Beagle Ristaino; and the careers of 26 women scientists are outlined by 37 authors from the United Kingdom, United States of America, Canada, Portu- gal, Italy and the Netherlands. It is most interesting to read, and gives one a very good indication of what these plant pathologists have contributed to this field. ELIZABETH PARNIS 12-1063 Valewood Trail, Victoria, British Columbia V8X 5G5 Canada ed designs on furniture in the mornings to earn enough to attend art classes in the afternoon. He joined the Bronx County Bird Club, following the new concept of identifying birds in the field without the need to shoot any specimens for confirmation. The club’s pres- 2007 ident, Ludlow Griscom, was an early advocate of this method and RTP soon rivaled him in auditory and visual skills. RTP’s first book, A Field Guide to the Birds, pub- lished 27 April 1934 at the height of the depression, made bird identification easy for the first time. His wide-ranging combination of skills and knowledge made possible the succeeding Peterson field guides, all published by Houghton Mifflin in Boston. Each used the “Peterson system of field identification,’ with an arrow pointing to important field marks for each species. Millions of copies were sold. His first bird field guide went through four more revisions, the last posthumously; it was 85% complete when he died a month before his 88" birthday. Through his multifac- eted achievements: books, articles, photographs, movies and paintings, Peterson became the best-known and arguably the best-loved naturalist in the world. RTP’s first teaching experiences were at a boy’s camp in Maine, and in writing pamphlets for members of Junior Audubon Societies. He remained a teacher all his life. His dedicated second wife, Barbara, man- aged his home and business affairs with great distinc- tion, freeing him for extensive world travel. Liz Rosenthal’s informative book is based largely on 147 interviews with 117 people, including many of the “who’s who” of the birding world. Their words and memories provide inside stories about their interactions with Roger. The inevitable result of this methodology is a few minor omissions. As one example, Rosenthal’s book omits one of RTP’s early pupils at the Rivers School near Boston: Eliot Richardson, who much later became U.S. Attorney-General and successfully nom- inated RTP as the teacher who had influenced him most, resulting in the nationwide “Teacher of the Year” Award. Nor we do we learn that RTP’s final total of honorary doctorate degrees reached 23. Conservation themes are interwoven throughout this book. These began with Peterson’s long involvement with the National Audubon Society, as a director, sec- retary, columnist in their magazine, and presenter of some of their most popular screen tours. He helped found the World Wildlife Fund, helped save fragile areas such as the fabulous Coto Donana in Spain, Mid- Book REVIEWS 23] way Atoll in the Pacific and Aldabra Atoll in the Indi- an Ocean, and helped Kenya's Lake Nakuru become a national park, where a million flamingos form “the world’s greatest ornithological spectacle.” He pho- tographed all 17 of the world’s species of penguin. We learn of Peterson’s close friendships with two other world-famous ornithologists, James Fisher and Sir Peter Scott. As evidence of his stature among pro- fessional ornithologists, Peterson served as president of the Wilson Ornithological Society and as Vice-Presi- dent of the American Ornithologists’ Union (AOU). An icon, he received the William Brewster Award from the AOU in 1944 at the early age of 36; Presidential Medal of Honor, presented by Jimmy Carter; Audubon Medal from the National Audubon Society; Gold Medal of the World Wildlife Fund, presented by Prince Bernhard of the Netherlands; Order of the Golden Ark from the Netherlands; Linnaeus Gold Medal from the Swedish Academy of Sciences, and the Smithson Medal from the Smithsonian Institution. He attended the dedication of the Charles Darwin Research Station in the Galapagos. RTP was happiest when serving as naturalist-lecturer aboard Lars-Eric Lindblad’s Explor- er on cruises to the Antarctic, Galapagos, Amazon River and Aldabra Atoll. I detected two errors. Iceland is NOT north of the Arctic Circle (page 133). The Peregrine nest on the Sun Life building was in Montreal, not Toronto (page 229). I also regret that, in a book about people, the peo- ple index is incomplete, omitting some names entirely, and incompletely listing mentions of others. Elizabeth J. Rosenthal, a graduate in both journalism and law, has previously written His Song: the Musical Journey of Elton John. Her Peterson life deals with a man of very different but not lesser talents, who was unusually helpful to beginner and expert alike, and whose personality shines through in the interview process. The potential audience includes the millions of people who began nature study with a Peterson field guide. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Canada Jee New TITLES Prepared by Roy John + Available for review * Assigned ZOOLOGY Audubon: Early Drawings. By S. Edwards [Annotater]. 2008. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 272 pages. 125.00 USD Cloth. * Backyard Birdwatch. By S. Kress. 2008. Dorling Kin- dersley/Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 72 pages. 11.95 CAD Paper. * Beautiful Minds: The Parallel Lives of Great Apes and Dolphins. By M. Bearzi and C. Stanford. 2008. Harvard University Press, 79 Garden Street, Cambridge, Massachu- setts 02138 USA. 322 pages. 24.95 USD Cloth. Butterfly. By T. Marent. 2008. Dorling Kindersley/Tour- maline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 280 pages. 35.00 USD Paper. Life in old Blood. By Sir D. Attenborough. 2008. Princeton University Press, 41 William Street, Princeton, New Jersey, 08540-5237 USA. 288 pages. 29.95 USD Cloth. * Ecology. By M. Cain, W. Bowman and S. Hacker. 2008. Sinauer Associates Inc., Box 407, Sunderland, Massachusetts 01375-0407 USA. 621 pages. 107.95 USD. Egg & Nest. By R. Purcell, L. Hall and R. Corado. 2008. Harvard University Press, 79 Garden Street Cambridge, Massachusetts 02138 USA. 224 pages. 39.95 USD Cloth. Evolution. Edited by M. Ruse and J. Travis. 2009. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. * The Ferocious Summer. By M. Hooper. 2008. Greystone Books, #201-2323 Quebec Street, Vancouver, British Col- umbia V5T 4S7 Canada. 336 pages. 29.95 USD. + Fossil Ecosystems of North America. By J. Nudds and P. Selden. 2008. University of Chicago Press, 5801 Ellis Avenue, Chicago, Illinois 60637 USA. 288 pages. 39.00 USD. * A Guide to the Mammals of China. Edited by A. Smith and Y. Xie. 2008. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 576 pages. 60.00 USD. * A Guide to the Mammals of Southeast Asia. By C. Francis. 2008. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 392 pages. 55.00 USD. Seashore Life in the North Atlantic. By J. D. Duane. 2008. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 240 pages. 19.95 USD. BOTANY * The Macrolichens of New England. By James W. Hinds and Patricia L. Hinds. 2008. The New York Botanical Garden Press, 200th Street & Kazimiroff Boulevard, Bronx, New York 10458 USA. 600 pages. 65 USD Cloth. THE CANADIAN FIELD-NATURALIST Vol. 121 MISCELLANEOUS Culturing Wilderness in Jasper National Park. By LS. MacLaren. 2008. University of Alberta Press, Ring House 2, Edmonton, Alberta T6G 2E1 Canada. 312 pages. 39.95 CAD Paper. Nahanni Journals — R.M. Patterson’s 1927-1929 Journals. Edited by R. Davis. 2008. University of Alberta Press, Ring House 2, Edmonton, Alberta T6G 2E1 Canada. 316 pages. 29.95 CAD Paper. Ocean. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario M5V 1M7 Canada. 512 pages. 27.95 USD Paper. *Birdwatcher: The Life of Roger Tory Peterson. By Elizabeth J. Rosenthal. 2008. The Lyons Press/Globe Pequot Press, Guilford, Connecticutt USA. 437 pages. Smithsonian Rock and Gem. 2008. By Dorling Kindersley, Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario M5V 1M7 Canada. 360 pages. 22.95 USD Paper. Shelter at the Shore — The Boathouses of Muskoka. By J. De Visser and J. Ross. 2008. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario M5V 1M7 Canada. 72 pages. 11.95 USD Paper. YOUNG NATURALISTS Dangerous Dinosaurs Q & A. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 64 pages. 14.99 USD Paper. Encyclopedia of Dinosaurs & Prehistoric Life. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 376 pages. 22.99 USD Paper. My Terrific Dinosaur Book. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario M5V 1M7 Canada. 14 pages. 14.99 USD Cloth. Earth Matters. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, On- tario MSV 1M7 Canada. 256 pages. 27.99 USD Cloth. Eyewitness Climate Change. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 72 pages. 18.99 USD Cloth. Sharpedia. 2008. By Dorling Kindersley. Tourmaline Editions Inc., 662 King Street West, Suite 504, Toronto, Ontario MSV 1M7 Canada. 128 pages. 14.99 USD Cloth. News and Comment Bill Cody Receives Honorary Doctorate of Science from McMaster University, June 5, 2007 William James (Bill) Cody was born in Hamilton, Ontario in 1922. He grew up in Hamilton during the Depression. During summers, he worked on his uncle Bruce’s fruit farm near St. David’s in the Niagara Peninsula. He always enjoyed the outdoors, but it was with the help of McMaster University’s Professor Lulu Gaiser that his love for plants grew. Bill re- ceived his bachelor’s degree from McMaster University in 1946, That same year, Bill joined the Canada Department of Agriculture at the Central Experimental Farm, in Ottawa, as Junior Agricultural Assistant. He was subsequently promoted to Senior Agricultural Assistant, Technical Officer and then Research Assistant. His enthusiasm for the outdoors led him to travel extensively during the summer months across nine provinces and two territories. During these excursions, he collected over 35,000 plant specimens. He has published over 343 articles, including books, scientific papers, and popular articles. He is Canada’s leading expert on the flora of the Yukon and is an Honorary Research Associate with Agriculture and Agri-Food Canada. Bill was appointed Curator of the Vascular Plant Herbarium in 1959. He held this position for 28 years. Under his super- vision it became the largest vascular plant herbarium in Can- ada with more than one million specimens, including the thou- sands collected by him personally. The herbarium provides a basis for research and identification of economically impor- tant plants in Canada. Officially retiring in 1987, after 41 years of service, Bill continues to go to the office. His expertise in identifying spec- imens and his knowledge is sought after by the younger gen- eration of botanists. He remains an internationally recognized Marine Turtle Newsletter (116) April 2007. 36 pages: EDITORIALS: Proceeding to the future (M. S. Coyne) — The oldest place where there is always something new (J. G. Frazier) — ARTICLES: An up- date on marine turtles in Cote d’Ivoire, West Aftrica (J. G. Penate, M. Karamoko, S. Bamba, and G. Djadji) — An update on sea turtle conservation activities in the Republic of Congo (G. Bal, N. Brehert, and H. Vanleeuwe) — Conflicts and social dilemmas associated with the incidental capture of marine turtles by artisanal fishers in Benin (J. S. Dossa, B. A. Somsom and G. A. Mensah) — Marine turtle mortali- ty in southern Gabon and northern Congo (R. Parnell, B. Verhage, S. L. Deem, H. Van Leeuwe, T. Nishihara, C. Mou- koula, and A. Gibudi) — Artificial lights as a significant cause of morbidity of Leatherbacks in Pongara National Park, Gabon (S. L. Deem. F. Boussamba, A. Z. Nguema, G-P Sounguet, S. Bourgeois, J. Cianciolo, and A. Formia) — Observations of marine turtles in relation to seismic air- gun sond off Angola (C. R. Weir) — Fibropapillomatosis confirmed in Chelonia mydas in the Gulf of Guinea, West Africa (A. Formia, S. Deem, A. Billes, S. Ngouessono, R. Parnell, T. Collins, G-P Sounguet, A. Gibundi, A. Villarubia, authority in Canada and continues to contribute to the devel- opment of the collection. He joined the Ottawa Field-Naturalist’s Club when he arrived in Ottawa in 1946. The position of Business Manager of a scientific journal, The Canadian Field-Naturalist was his for over 50 years. His duties included managing finances, monitoring stock, billing for reprints and public relations. Bill was married for 47 years to Lois and together they raised five children: David, Margaret, Leslie, Douglas and Gordon. Having sung in a barber shop quartet and church choir, his joy of music was very evident. Bill’s baritone voice could be heard resonating throughout the halls of his office and apartment building. He often had a pun or quip on the tip of his tongue. In recent years, Bill had the opportunity to return to the Yukon four times. Each time, he took one of his children to help him collect and log specimens in order to determine range extensions and also to let them experience a place that had been a large part of his life. Bill is very proud of his family heritage. His father, William MacPherson Cody, was the first appointed anesthetist to the Hamilton General Hospital. Also, his second cousin, 3 times removed, was William Frederick Cody, known to most as “Buffalo Bill” Cody, a famous hero of the American Wild West. Among the tributes awarded to Bill are the following: the Lawson Medal, most prestigious award of the Canadian Bo- tanical Association (1997); the Distinguished Technical Comm- unication Award of the Society for Technical Communica- tions (1997); induction into the McMaster University Alumni Gallery (2002); The Queen’s Golden Jubilee Medal (2002): and the Yukon Biodiversity Awareness Award (2006). G. H. Balazs, and T. R. Spraker) — Presence of sea turtles in traditional pharmacopoeia and beliefs of West Africa (J. Fretey, G. H. Segniagbeto, and M’M. Soumah) — Discovery of a Gabonese Leatherback in South Africa (J. Fretey, A. Billes, B. Baxter, and Caterine Hughes) — Second TAMAR- tagged Hawksbill recaptured in Corisco Bay, West Africa (A. Grossman, C. Bellini, A. Fallabrino, A. Formia, J. M. Mba, J. N. Mba, and C. Obama) — IUCN-MTSG QUARTERLY UPDATE — MEETING REPORTS — NEWS & LEGAL BRIEFS — RECENT PUBLICATIONS — ACKNOWLEDGEMENTS The Marine Turtle Newsletter is edited by Lisa M. Camp- bell, Nicholas School of Environment and Earth Sciences, Duke University, 135 Duke Marine Lab Road, Beaufort, North Carolina 28516 USA; and Matthew H. Godfrey, North Carolina Wildlife Resources Commission, 1507 Ann Street, Beaufort, North Carolina 28516 USA. Subscriptions and donations towards the production of the MTN can be made online at or postal mail to Michael Coyne (Managing Editor), Marine Turtle Newsletter, 1 Southampton Place, Durham, North Carolina 27705 USA: e-mail: mcoyne @seaturtle.org. 234 THE CANADIAN FIELD-NATURALIST Vol. 121 Atlas of amphibians and reptiles of Quebec, 2007 inventory Instructions for participants in the 20" herpetological survey for the Québec Atlas of amphibians and reptiles, coordinated by the St. Lawrence Natural History Society in collaboration with the Ministere des Ressources naturelles de la Faune du Québec. The data from this annual inventory pro- gram are incorporated into the Centre de donnees sur le pat- rimone natural du Québec database; to date this database con- tains more than 60 000 date entries on Québec species. This provincial inventory is done every year with the important co- operation of volunteers. Contributors are encouraged to send their e-mail address and request for instruction on trasmitting data, Observation cards in paper format are available on re- quest. Mathieu Oulette, Research Officer, Research and Conser- vation, Ecomuseum — St. Lawrence Valley Natural History Society, 21 125 chemin Saint-Marie, Saint-Anne-de-Bellevue, Quebec H9X 3Y7; phone 514-457-9449; fax 514-457-0769; e-mail: info @herpetofaune.org. The Boreal Dip Net/L’epousette boreale 11(1) Spring 2007 Newsletter of the Canadian Amphibian and Reptile Con- servation Network/Réseau Canadien de Conservation des Amphibians et Reptiles Contents: Editor’s Note (Sara Ashpole) — Notes from the chairperson (David Galbraith) — Presentaions at CARC- NET/RECCAR 2006 Annual Meeting, October, Victoria, British Columbia: Student Presentation Awards: Best Platform in 2006 Julie A. Lee-Yaw — Best Poster in 2006 Isabelle Deguise — 2006 Silver Salamander Award Mike Rankin (posthumously) — 2006 Silver Salamander Award: Fish & Wildlife Compensation Program: Jean Wai Jang — 2006 Silver Salamander Award: Gord McAdams and the West Kootenay Ecosociety: They stuck their necks out to save the turtles — 2006 Scholarship for Amphibian and Reptile Re- search: Gregory Bulte — Keynote Speakers with Extended Abstracts : Captive Breeding of Amphibians - Conservation or Cosmetic Surgery? (Richard A. Griffiths); The Way of a Gecko on a Rock — Why the Animal’s Real Environment Matters — How the Student Presentations are Judged — Presentations Made (titles and authors of 50 presentations) — Book notice: Amphibians and Reptiles of British Columbia — Towards a Cooperative Strategy for Dealing with Disease Issues in Amphibian and Reptile Conservation — Blanding’s Turtle Management Symposium and Roads and Ecopassages Forum — Announcement: CARRCNET/RECCAR Annual General Meeting 21 to 24 September 2007 Kingston, Ontario — CARCNET/RECCAR Board of Directors. Canadian Association of Herpetologists/Association Canadienne des Herpetologists Bulletin 14(2) Spring 2007 Contents: Instructions for Authors — EDITORIAL NOTES: Membership and Association News — MEETINGS: CARCNet 11" Annual Meeting 27 to 29 October in Victoria, British Columbia; Joint Meeting of Ichthylogists and Herpetologists in St. Louis, Missouri, 11 to 16 July 2007 — FEATURE ARTICLE: Accuracy and precision of site occupancy estimates of anuran acoustic surveys when incorporating detection prob- abilities (Shane de Solla) — FIELD NOTES: Use of digital images obtained from volunteers to increase sample size and accuracy of morphometric ratios (Wayne Weller) — BOOK REVIEWS: Herpetological Conservation and Biology [an online journal] (assessed by Mare J. Mazerolle, Jim Bogart, Kate Jackson) — Diseases of Amphibians and Reptiles (re- viewed by Dale A. Smith) — THESIS ABSTRACTS IN CAN- ADIAN HERPETOLOGY: Sara M. Delannoy MSc 2006 Univer- sity of Calgary, Alberta (Supervisor Anthony P. Russell) Subdigital setae of the Tokay Gecko: Variation in form and im- plication for adhesion; Patrick S. Druckenmiller PhD 2006 University of Calgary, Alberta (Supervisor Anthony P. Russell) Early Cretaceous Plesiosaurs (Sauroppterygia: Plesiosaura) from northern Alberta: Palaeoenvironmental and systematic implications; Virgil C. Hawkes MSc 2007 University of Vic- toria (Supervisor Patrick Gregory) Riparian management and ampibians: Does buffer width matter?; Geoffrey Hughes Honours BSc 2007 Laurentian University, Sudbury, Ontario (Supervisor Jacqueline Litzgus) Nest site selection by Wood Turtles (Glyptemys insculpta) in a thermally limited environ- ment; Heather A. Jamniezky PhD 2006 University of Calgary, Alberta (Supervisor Anthony P. Russell) Turtle cranial arterial circulation: Intergrative analysis of a systematically influen- tial character complex; Sheila Smith BSc 2006 Laurentian University, Sudbury, Ontario (Supervisor Jacqueline Litzgus) Geographic variation of sexual size dimorphism in Painted Turtles, Chrysemys picta; Eric D. Snively PhD 2006 Univer- sity of Calgary, Alberta (Supervisor Anthony P. Russell) Neck musculoskeletal function in the Tyrannosauridae (Theropoda, Coelurosauria): Implications for feeding dynamics — THESES IN HERPETOLOGY AT THE UNIVERSITY OF VICTORIA (Patrick Gregory) a list by student and title for 33 theses (1974-2005): 9 BSc, 18 MSc, 6 PhD — RECENT PUBLICATIONS IN CANADIAN HERPETOLOGY: 17: 2006 (15), 2007(1), in press (1) — ANNOUNCEMENTS New Fiction Book about Sea Turtles: Sand Turtles. Editor’s Report for Volume 120 (2006) Mailing dates for issues in volume 120(1) 15 July TABLE 2. Number of articles and notes published in The Can 2007, (2) 1 November 2007; (3) 30 November 2007; (4) @4/an Field-Naturalist Volume 120 (2006) by major field of 31 December 2007. study. A summary of circulation to members and sub- Subject. ~~~~~-~Arficiles”~—~—sONotes”~—=~=S*«*STowal scribers (institutional and individual) totals for 2006 is given in Table 1. The number of articles and notes in ne ae = : = volume 121 is summarized in Table 2 by topic; totals for eae: : ; d ; : Amphibians + reptiles | I 2 Book Reviews and New Titles are given in Table 3, and 5, 5 4 7 the distribution of content by page totals per issue in Iycects 3 0 3 Table 4. ' : Other invertebrates | | 2 Council continued to authorize 40% of membership Plants 13 0 13 dues for publication of the journal and all of sub- Paleontology | 0 scriptions (both individual and institutional). Council — Tributes 2 0 2 also has allocated 80% of the annual interest from the ea , Totals 56 16 72 Manning Fund and other capital funds to The Canadian Field-Naturalist. Use of the Manning fund portion was broadened on an issue-by-issue basis to offset the pub- lication costs where authors lacked grant or institutional funds to support publication. tents and abstracts are now posted on the Ottawa Field- The journal was printed at Gilmore Printers, Ottawa, Naturalists’ Club website by webmaster Sandra Gar- and thanks are due business representative Tom Smith; land, and PDFs may be ordered by authors as well as customer representative Chuck Graham for overseeing _ traditional print reprints. production, and Wendy Cotie for typesetting and cor- Manuscripts (excluding book reviews, notices, and rections. Leslie Cody prepared the Index for volume reports) submitted to The Canadian Field-Naturalist 120; Elizabeth Morton proofed the galleys. When Bill _ totalled 67 in 2006, down by 8 from the total for 2005. Cody, Businesss Manager for 60 years, was unable to The following reviewed for papers submitted in 2006 carry on, most of his duties were shouldered by Trea- (with number of manuscripts reviewed in parentheses surer Frank Pope and Assistant Treasurer Jim Ward _ if more than one): with invoice preparation assumed by the editor. Roy Associate Editors: R. Anderson, Canadian Museum of John arranged and edited book reviews and prepared the Nature, Ottawa, Ontario; C. D. Bird, Erskine, Alberta (8); R. New Titles listings. Special thanks are due to Jenifer i woe Pa aCe ace elie pe ae aed eee aie iste, a ae and 2 Canadian Museum of Nature, Ottawa, Ontario (6); A. UE Er- ‘ ; u : : : skine, Sackville, New Brunswick (10); D. F. McAlpine, New ing for 120(1) the eight papers on conservation evalu- Brunswick Museum, Saint John, New Brunswick (4); D. W. ation of some British Columbia plants originated by the — Nagorsen, Mammalia Biological Consulting, Victoria, British late George Douglas and also for preparing the tribute — Columbia (8); W. O. Pruitt, Jr., University of Manitoba, Win- and updated bibliography for Douglas. The journal con- _nipeg, Manitoba (6). TABLE 1. The 2006 circulation of The Canadian Field-Naturalist (2005 in parentheses). Prepared by Robin Murphy, bmr group, Ottawa. Totals are copies mailed. As some members opt not to receive the journal, membership total here is only those receiving the journal. For total club membership see the Annual Report of the membership committee to the 2007 Annual Business Meeting. Canada USA Other Totals Memberships Family & individual 556 (624) 24 (26) 5 (6) 585 (656) Subscriptions Individuals 131 (130) 57 (63) 5 (4) 193 (197) Institutions 158 (156) 232 (229) 32); 64) 422 (419) Ags 845 (910) 313 (318) 42 (44) 1200 (1272) Note: 20 countries (outside Canada and United States) are included under “Other” (number of copies in parentheses): Australia, Austria, Belgium, Brazil, Denmark, United Kingdom (9: including | to Scotland), Finland (2), France (2), Germany (2), Iceland, Ireland, Japan, Netherlands (2), New Zealand, Norway (6), Poland, Russia, South Africa, Spain (2), ) Switzerland, Taiwan (2). 235 236 TABLE 3. Number of reviews and new titles published in Book Review section of The Canadian Field-Naturalist Volume 120 (2006) by topic. Reviews New Titles Zoology 37 126 Botany 4 1S) Environment 9 58 Miscellaneous 10 4 Young Naturalists 1 5 Totals 61 208 Other Reviewers: P. Achuff, Waterton Lakes, Alberta; W. B. Ballard, Texas Tech University, Lubbock, Texas (4); J. R. Bider, Baie d’Urfe, Quebec (2); G. Blouin-Demers, University of Ottawa, Ontario; E. L. Bousfield, Victoria, British Columbia (2); K. Conlan, Canadian Museum of Nature, Ottawa (2); D. Boyd, Helena, Montana; R. J. Brooks, University of Guelph, Ontario; S. Carriere, Environment and Natural Resources, Yellowknife, Northwest Territories; R. W. Campbell, Bio- diversity Centre for Wildlife Studies, Victoria, British Colum- bia; J. A. Cook, Museum of Southwestern Biology, University of New Mexico, Albuquerque; S. Cooke, Carleton University, Ottawa, Ontario (2); W. J. Crins, Ontario Ministry of Natural Resources, Peterborough, Ontario; D. B. Donald, Environment Canada, Regina, Saskatchewan; J. Dubois, Manitoba Natural Resources, Winnipeg, Manitoba; J. Ferron, Universite du Quebec a Rimouski, Quebec; D. R. Galbraith, Royal Botanical Gardens, Hamilton, Ontario; A. J. Gaston, Environment Can- ada, Gatineau, Quebec; D. Giblin, Burke Museum, University of Washington, Seattle, Washington; J. Gilhen, Nova Scotia Museum of Natural History, Halifax, Nova Scotia (2); P. Goossen, Environment Canada, Edmonton, Alberta; P. T. Gregory, University of Victoria, British Columbia; F. H. Har- rington, Mt. St. Vincent University, Halifax, Nova Scotia; S. J. Hecnar, Lakehead University, Thunder Bay, Ontario; E. Hendrycks, Canadian Museum of Nature, Ottawa, Ontario; P. Hicklin, Environment Canada, Sackville, New Brunswick; L. V. Hills, Cochrane, Alberta; S. Hooker, University of St. Andrews, Fife, United Kingdom; B. D. Johnson, Herbarium, Northern Forest Research Centre, Canadian Forest Service, Edmonton, Alberta (2); C. S. Houston, Saskatoon, Saskat- chewan; J. Kamler, Polish Academy of Sciences, Bialowieza, Poland; C. R. Lacroix, University of Prince Edward Island, Charlottetown; J. Lanteigne, Travaux publics et services, gouvernementaux Canada, Gatineau, Quebec; G. Lee, Ottawa, Ontario; R. MacCulloch, Royal Ontario Museum, Toronto, Ontario; J. Madill, Canadian Museum of Nature, Ottawa, Ontario; F. F. Mallory, Laurentian University, Sudbury, Ontario (2); M. Mallory, Environment Canada, Iqaluit, Nunavut; André Martel, Canadian Museum of Nature, Ottawa, Ontario; L. D. Mech, U.S. Geological Survey, The Raptor Center, University of Minnesota, St. Paul (3); J. Metcalfe-Smith, Aquatic Ecosystem Protection Branch, National Water Re- search Institute, Burlington, Ontario; A. Moehrenschlager, Calgary Zoological Society, Alberta; E. L. Mills, Dalhousie University, Halifax, Nova Scotia; G. Morrison, Environment Canada, Gatineau. Quebec; G. Parker, Environment Canada, Sackville, New Brunswick; E. Pip, University of Winnipeg, Manitoba; G. Pohle, Huntsman Marine Science Centre, St. Andrews, New Brunswick; W. Preston, Winnipeg, Manitoba; G. Proulx, Alpha Research & Management Ltd., Sherwood THE CANADIAN FIELD-NATURALIST Vol. 120 TABLE 4. Number of pages per section published in The Cana- dian Field-Naturalist Volume 120 (2006) by issue. (1) (2) (3) (4) Total Articles 105 85 104 68 362 Notes 7 11 3 12 33 Tributes 0 12 9 0 21 Book Reviews 19 15 13 19 66 CEN/OFNC Reports” °0 2 9 5 16 News and Comment 2 2 1 D) 7 Index 0 0 0 21 21 Advice to Contributors | 1 1 1 4 Totals ey ARS IX) 128 530 * Total pages for book review section include both reviews and new titles listings. ™ Includes CEN Editor’s report in (2), OFNC Annual Business Meeting (3) and OFNC Awards (4). Park, Alberta (2); R. Reeves, Okapi Wildlife Associates, Hudson, Quebec (2); T. E. Reimchen, University of British Columbia, Victoria, British Columbia; G. J. Robertson, Envi- ronment Canada, Mount Pearl, Newfoundland; L. Rogers, . Ely, Minnesota; R. Rosatte, Ministry of Natural Resources, Trent University, Peterborough, Ontario; A. Russell, University of Calgary, Calgary, Alberta; F. Scott, Acadia University, Wolfville, Nova Scotia; F. W. Schueler, Bishops Mills Natural History Center, Oxford Mills, Ontario; J. W. Sheard, University of Saskatchewan, Saskatoon; Brian Slough, Whitehorse, Yukon Territory; K. W. Stewart, University of Manitoba, Winnipeg, Manitoba (3); P. Weatherhead, Queen’s Biological Station, Elgin, Ontario; M. J. Wetzell, Illinois, Natural History Survey, Champaign, Illinois; N. Yan, York University, North York, Ontario. I am also indebted once again to the President of the Ottawa Field-Naturalists’ Club, Gary McNulty, and the Club Council for continuing support of the journal: Chairman Ron Bedford and the Publications Com- mittee of the OFNC for editorial encouragement and support, the Finance Committee for input and encour- agement to speed things up, the Canadian Museum of ' Nature for access to its library and the facilities at the « Natural Heritage Building, 1740 Pink Road, Aylmer, | Quebec, and to Joyce for everything else. I must add my special thanks due William J. (Bill) Cody who served the club as council member for more « than 60 years, vitually all ot this time as Business! Manager of The Canadian Field-Naturalist. Those of) us exposed to Bill’s unfailing good humour, determi-' nation, efficient focus on the job on hand, and some-: times his inevitable serenades, have lasting memories’ of the time shared. Bill served with, and kept each on) track, five editors (one twice) — Harold Senn to 1955), Bob Hamilton (1956-1961), myself (1962-1966), Ted Mosquin (1967-1972), Lorraine Smith (1972-1981, and me again (1981 — present). Efficient management and financial stability through good times and bad_ were due in large measure to Bill’s initiative which assured, in some periods, the journals very survival. 2007 A decade ago, to mark 50 years of service, the club paid tribute to Bill’s contribution to that point with a reception at the Central Experimental Farm, Depart- ment of Agriculture, Ottawa, where Bill had spent his whole career, including “retirement”, as a research botanist. Stephen Darbyshire for Awards Committee stated in part (1998. Canadian Field-Naturalist 1 12(4): 719-720): “Such long-term service and dedication is unprece- dented in the history of the Club and is not likely to ever be repeated. The titles of office do not reveal the extent to which Bill has laboured for the Club. One cannot completely list the jobs that he has taken on (even Bill cannot remember them all), many of which are among the most distasteful and time consuming (imagine the years of dealing with government forms for postage rates, charitable status, etc.). Executive officers have 7 EpIror’s REPORT 237 come and gone, Canadian Field-Naturalist editors have come and gone, Council is re-elected every year, but Bill Cody continues as a major constituent of the glue that keeps the Club together and keeps it, and its scien- tific publication, running smoothly.” Bill continued to perform these duties for close to another decade before failing health forced him pass them on to others. The full truth in the earlier words of tribute has become increasingly realized by those of us Bill left to share his multitude of tasks. Most of the burden has fallen on Frank Pope, already contri- buting tremendous service as Treasurer of the Ottawa Field-Naturalist’s Club, and Jim Ward as Treasurer's Assistant. They have made the transition work. FRANCIS R. COOK Editor 238 THE CANADIAN FIELD-NATURALIST Vol. 120 Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit by post to the Editor, in either English or French, three complete manuscripts written in the journal style. Manuscripts may also be submitted (one copy) by e- mail. The research reported should be original. It 1s recom- mended that authors ask qualified persons to appraise the paper before it is submitted. All authors should have read and approved it. Institutional or contract approval for the publica- tion of the data must have been obtained by the authors. Also authors are expected to have complied with all pertinent leg- islation regarding the study, disturbance, or collection of ani- mals, plants or minerals. The place where voucher specimens have been deposited, and their catalogue numbers, should be given. Latitude and longitude should be included for all indi- vidual localities where collections or observations have been made. Manuscripts should be printed on standard-size paper, dou- blespaced throughout, generous margins to allow for copy marking, and each page numbered. For Articles and Notes provide a bibliographic (citation) strip, an abstract, and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manu- scripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in a separate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “Canadian” or “American” spellings are acceptable in English but should be consistent within one manuscript. The Oxford English Dictionary, Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Illustrations Photographs should have a glossy finish and show sharp contrasts. Electronic versions should be high resolution. Prepare line drawings with India ink on good quality paper and letter (don’t type) descriptive matter. Photographic reproduction of line drawings should be no larger than a standard page. Write author’s name, title of paper, and fig- ure number on the lower left corner or on the back of each illustration. Reviewing Policy Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor (who reviews it or asks another qualified person to do so), and at least one other reviewer, who is a specialist in the field, cho- sen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—some- times extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by pay- ing $90 for each page, plus $30 for each illustration (any size up to a full page), and up to $90 per page for tables (depend- ing on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Editor. Limited joumal funds are available to help offset publi- cation charges to authors without grants or institutional sup- port. Requests for financial assistance should be made to the Editor when the manuscript is submitted. Reprints An order form for the purchase of reprints or pdf will ac- company the galley proofs sent to the authors. Invoices for publication costs will be sent when the submission is pub- lished. FRANCIS R. CooK, Editor RR 3 North Augusta, Ontario KOG IRO Canada TABLE OF CONTENTS (concluded) Volume 121 Number 2 2007 Notes (continued) Human-assisted movements of Raccoons, Procyon lotor, and Opossums, Didelphis virginiana, between the United States and Canada Rick C. ROSATTE, DENNIS DONOVAN, MIKE ALLAN, LAURA BRUCE, and CHRIS DAVIES PAV Correlation between age estimates for Elk, Cervus elaphus, using tooth wear/eruption patterns and counts of annuli in tooth cementum RICK ROSATTE, ANDREW SILVER, MARK GIBSON, BRUCE CHISHOLM, and NORM CooL 214 Predator-prey interaction between and American Robin, Turdus migratorius, and a Five-lined Skink, Eumeces fasciatus E. NATASHA V ANDERHOFF 216 Book Reviews Zoo.ocy: Atlas of Breeding Birds of Ontario 2001-2005 — The Return of Caribou to Ungava — A Guide to the Mammals of China — Birds of the Saskatoon Area — Brian Coad’s Ichthyology Site 219 Borany: Exploration and Utilization of Chinese Spice Plant Resources pipe) MISCELLANEOUS: Arctic Hell Ship — Eighteenth-century Naturalists of Hudson Bay — Hunters at the Margin: Native People and Wildlife Conservation in the Northwest Territories — The Ornithologist’s Dictionary — Pioneering Women in Plant Pathology — Birdwatcher: The Life of Roger Tory Peterson 226 New TITLES DBD News and Comment Bill Cody receives Honorary Doctorate of Science from McMaster University, June 2007 — Marine Turtle Newsletter 116 April 2007 — Atlas of amphibians and reptiles of Quebec, 2007 inventory — The Boreal Dip Net/L'epuosette boreale 11(1) Spring 2007 — Canadian Association of Herpetologists/Association Canadien des Herpetologists Bulletin 14(2) Spring 2007 233 Editor’s Report for Volume 120 (2006) FRANCIS R. COOK 235 Advice to Contributors 238 Mailing date of the previous issue 121(1): 6 June 2008 THE CANADIAN FIELD-NATURALIST Volume 121 Number 2 Articles Diversity and distribution of the terrestrial mammals of the Yukon Territory: A review BRIAN G. SLOUGH and THOMAS S. JUNG State records and habitat of Clam Shrimp, Caenestheriella gynecia (Crustacea: Conchostraca), in New York and New Jersey ROBERT E. SCHMIDT and ERIK KIVIAT Colonization of non-traditional range in dispersing Elk, Cervus elaphus nelsoni, populations FRED VAN DYKE The Rumsfeld Paradigm: Knowns and Unknowns in characterizing habitats used by the endangered Sharp-tailed Snake, Contia tenuis, in southwestern British Columbia SHANNON F. WILKINSON, PATRICK T. GREGORY, CHRISTIAN ENGELSTOFT, and KARL J. NELSON Do repugnant scents increase survival of ground nests? VANESSA B. HARRIMAN, JUSTIN A. PITT, and SERGE LARIVIERE Translocation and recovery efforts for the Telkwa Caribou, Rangifer tarandus caribou, herd in westcentral British Columbia, 1997-2005 ASTRID VIK STRONEN, PAUL PAQUET, STEPHEN HERRERO, SEAN SHARPE, and NIGEL WATERS An inventory of aquatic and subaquatic plants in SASK Water Canals in Central Saskatchewan, Canada J. HuGo Cota-SANCHEZ and KIRSTEN REMARCHUK Life history of the marine isopod Cyathura polita in the Saint John River Estuary, New Brunswick: a species at the northern extent of its range SARAH C. MERCER, GLENYS D. GIBSON, and MICHAEL J. DADSWELL Post-emergence movements and overwintering of Snapping Turtles, Chelydra serpentina, hatchlings in New York and New Hampshire GORDON R. ULTScH, MATT DRAuD, and BARRY WICKLOW Notes on succession in old fields in southeastern Ontario: Il The herbs A. CROWDER, R. HARMSEN, and S. E. BLATT Growth of White Spruce, Picea glauca, seedlings in relation to microenvironmental conditions in a forest-prairie ecotone of southwestern Manitoba SOPHAN CHHIN and G. GEOFF WANG Natural recolonization of cultivated land by native prairie plants and its enhancement by removal of Scots Pine, Pinus sylvestris PAUL CATLING and BARRY KING Notes A high elevation record of the Star-nosed Mole, Concylura cristata, in northeastern Vermont RYAN W. Norris and C. WILLIAM KILPATRICK Northern Long-eared Bat, Myotis septentrionalis, (Chiroptera: Vespertilionidae) on Price Edward Island: First records of occurrence and over-wintering JENNIFER A. BROWN, DONALD F. MCALPINE, and ROSEMARY CURLEY Arboreal, late summer, courtship of Maritime Garter Snakes, Thamnophis sirtalis pallidulus, in Dartmouth, Nova Scotia JOHN GILHEN and ROBERT STRUM 210 i MUCUS N (continued on inside back cover) ier erhegmwccrand” SEN ISSN 0008-3550 FSC www.fscorg Cert no. SGS-COC-003284 © 1996 Forest Stewardship Council 2007 1] 12 133 14 15 206 The CANADIAN FIELD-NATURALIST Canada ASTS’ CLUB, Ottawa, Published by THE OTTAWA FIELD-NATURAI July-September 2007 Volume 121, Number 3 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Michaille Jean Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to) encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields| as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-) ments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson 2007 Council President: Mike Murphy Annie Belair Diane Kitching Dan Millar Vice-President: Ken Allison Ronald E. Bedford |= Karen McLachlan Hamilton Stanley Rosenbaum Recording Secretary: Susan Laurie-Bourque Fenja Brodo David Hobden Henry Steger Treasurer: Frank Pope Julia Cipriani Diane Lepage Chris Traynor Past President: Gary McNulty William J. Cody Ann Mackenzie Eleanor Zurbrigg Francis R. Cook Gillian Marston To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069. Westgate P.O. Ottawa, Canada K1Z 1A2 For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed 1 ir this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. | We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) towarc our mailing costs. PAP Registration Number 9477. Canada Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca Copy Editor: Elizabeth Morton Honorary Business Manager: William J. Cody Business Manager: Frank Pope, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5; e-mail: r.john @rogers.com Associate Editors: | Robert R. Anderson Robert R. Campbell Anthony J. Erskine Donald F. McAlpine Warren B. Ballard Paul M. Catling David Nagorsen William O. Pruitt, Jr. Charles D. Bird Brian W. Coad Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal mail or e-mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. Subscriptions and Membership Subscription rates for individuals are $33 per calendar year. Libraries and other institutions may subscribe at the rate of $50 pe; year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $33 (individual) $35 (family) $50 (sustaining) ani $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) mus add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Locz Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries < $33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be maile to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publication Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: July-October 2007 (December 2008). Cover: A Red-winged Blackbird nest with three blackbird eggs and five Brown-headed Cowbird eggs. The nest was found in Conservation Reserve Program grassland in Day County, South Dakota. Photographed on 16 June 2005 by Lawrence I Igl. See article on cowbird parasitism by Ig] and Johnson, pages 239-255. ] | Volume 121, Number 3 April June 2007 Brown-headed Cowbird, Molothrus ater, Parasitism and Abundance in the Northern Great Plains LAWRENCE D. IGL! and DouGLAS H. JOHNSON? 'U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 37" Street SE, Jamestown, North Dakota 5840] USA; e-mail: larry_ig] @usgs.gov 2 U.S. Geological Survey, Northern Prairie Wildlife Research Center, 204 Hodson Hall, 1980 Folwell Avenue, University of Minnesota, St. Paul, Minnesota 55108 USA Igl, Lawrence D., and Douglas H. Johnson. 2007. Brown-headed Cowbird, Molothrus ater, parasitism and abundance in the northern Great Plains. Canadian Field-Naturalist 121(3): 239-255. The Brown-headed Cowbird (Molothrus ater) reaches its highest abundance in the northern Great Plains, but much of our understanding of cowbird ecology and host-parasite interactions comes from areas outside of this region. We examine cow- bird brood parasitism and densities during two studies of breeding birds in the northern Great Plains during 1990-2006. We found 2649 active nests of 75 species, including 746 nonpasserine nests and 1902 passerine nests. Overall, <1% of non- passerine nests and 25% of passerine nests were parasitized by Brown-headed Cowbirds. Although the overall frequency of cowbird parasitism in passerine nests in these two studies is considered moderate, the frequency of multiple parasitism among parasitized nests was heavy (nearly 50%). The mean number of cowbird eggs per parasitized passerine nest was 1.9 + 1.2 (SD; range = 1-8 cowbird eggs). The parasitism rates were 9.5% for passerines that typically nest in habitats characterized by woody vegetation, 16.4% for grassland-nesting passerines, 4.7% for passerines known to consistently eject cowbird eggs, and 28.2% for passerines that usually accept cowbird eggs. The Red-winged Blackbird (Agelaius phoeniceus) was the most commonly parasitized species (43.1% parasitism, 49.6% multiple parasitism, 71.2% of all cases of parasitism). Passerine nests found within areas of higher female cowbird abundance experienced higher frequencies of cowbird parasitism than those found in areas of lower female cowbird abundance. Densities of female cowbirds were positively related to densities and rich- ness of other birds in the breeding bird community. Key Words: Brown-headed Cowbird, Molothrus ater. brood parasitism, grasslands, multiple parasitism, spatial variation, temporal variation, northern Great Plains. The Brown-headed Cowbird (Molothrus ater; here- after cowbird) is an obligate brood parasite that lays its eggs in nests of a variety of host species. Cowbird dis- tribution, abundance, and brood parasitism have long been major topics in avian ecology in North America (Nice 1937; McGeen 1972; May and Robinson 1985; Ortega 1998; Herkert et al. 2003). Cowbird abundance and rates of brood parasitism vary geographically (Robinson et al. 1995; Ortega 1998; Rothstein and Robinson 1998). The cowbird reaches its highest abun- dance in the northern Great Plains (Sauer et al. 2005*), yet much of our understanding of cowbird parasitism and the dynamics of host and parasite populations comes from areas outside of this region. Nonetheless, some studies have reported high rates of cowbird par- asitism and multiple parasitism in the northern Great Plains (e.g., Linz and Bolin 1982; Romig and Craw- ford 1995; Davis and Sealy 2000; Koford et al. 2000). Robinson and Smith (2000), however, suggested that some of these oft-cited studies may not be representa- tive of the northern Great Plains. There are few pub- lished examples of community-wide studies of cow- bird parasitism in the northern Great Plains (e.g., Gran- fors et al. 2001; Woolfenden et al. 2004). Rather, most reports of cowbird parasitism from this region focus on a single species (e.g., Linz and Bolin 1982), a sub- set of species (e.g., Winter et al. 2004), or species that have evolved defenses against cowbird parasitism (Sealy 1996). At the continental level, cowbird abundance dec- lines with distance from the center of the species’ breeding range in the northern Great Plains (Thomp- son et al. 2000). The frequencies of cowbird parasitism appear to follow a similar biogeographical pattern (e.g. Smith and Myers-Smith 1998), especially for grass- land birds (Jensen and Cully 2005a, 2005b), although this relationship may not be generalized across habitats (Robinson and Smith 2000) and may not be indicative of parasitism frequencies at the local level (Chace et al. 2005). Nonetheless, Chace et al. (2005) acknowl- 239 240 edged that, at the continental scale, cowbird abundance appears to be a reasonable predictor of cowbird para- sitism levels. Because of the difficulty in obtaining data on both cowbird abundance and brood parasitism across the continent (or even over a large region), there are few examples of studies that have evaluated this rela- tionship using cowbird and nest data collected concur- rently at the same study sites over a large region (e.g., Robinson et al. 2000; Jensen and Cully 2005a, 2005b). Only a few evaluations of this biogeographical rela- tionship have included data from study sites in the northern Great Plains (e.g., Smith and Myers-Smith 1998; Herkert et al. 2003). At local and regional scales, geographic differences in cowbird distribution and abundance also may result from differences in avian communities (e.g., Farmer 1999; Thompson et al. 2000). That is, given that cow- birds are host generalists, measures of avian abundance or richness might serve as proximate cues for cowbirds in determining where to settle (McGeen 1972; Farmer 1999). Chace et al. (2005) listed four nested levels of an avian community that cowbirds could use as cues for settlement: (1) all breeding birds in the avian com- munity, (2) all passerine species, (3) all host species, and (4) a single host species. Very few studies have evaluated this relationship at all four levels (Farmer 1999). Using BBS (Breeding Bird Survey) data, Hahn and O’Connor (2002) concluded that cowbird distri- bution was not simply the result of shared habitat preferences with their hosts, but rather host abun- dance was an important predictor of cowbird occur- rence in recently colonized regions of the United States (1.e., eastern and western states) and to a lesser extent in the cowbird’s ancestral range (i.e., central Great Plains). Except for Hahn and O’Connor’s (2002) evaluation using BBS data, the influences of avian abundance and richness on cowbird distribution and abundance have not been evaluated using regional data from the northern Great Plains (reviewed in Chace et al. 2005). In this paper, we report cowbird densities and par- asitism frequencies during two breeding-bird studies in the northern Great Plains: a statewide study in North Dakota (1992 and 1993: Igl and Johnson 1997; Igl et al. 1999) and a Conservation Reserve Program (CRP) grassland study in nine counties of four states (1990-2006: Johnson and Schwartz 1993a, 1993b; Igl and Johnson 1995, 1999; Johnson and Igl 1995, 2001). We use data from nests found incidentally during both studies to examine weekly and annual variation in parasitism rates in the northern Great Plains, and nest data from the CRP grassland study to examine region- al variation in parasitism rates. We use data from the long-term CRP grassland study to evaluate the rela- tionships between female cowbird densities and cow- bird parasitism and between female cowbird densities and avian densities and richness in different regions of the northern Great Plains. CRP grasslands are a suit- THE CANADIAN FIELD-NATURALIST Vol. 121 able habitat to evaluate these relationships because this habitat has been shown to be a dominant landscape predictor of cowbird distribution in the United States (Hahn and O’Connor 2002). Study Areas North Dakota statewide study The study area for the North Dakota statewide study was described in detail by Stewart and Kantrud (1972) and Ig] and Johnson (1997) and is only briefly described here. North Dakota was divided into eight major strata based on biogeographical, physiographi- cal, and ecological characteristics. From these eight strata, 130 quarter-sections (about 64.75 ha each) were selected randomly (Figure 1). The number of sample units allocated to each stratum was proportional to the area of the stratum. Landowners denied access to one quarter-section in 1992 and a different quarter-section ims 19935 CRP grassland study The CRP study is an ongoing investigation (1990 to present) that is examining breeding-bird communi- ties annually in nearly 350 CRP grassland fields in nine counties of four states (Figure 2), including Fal- lon and Sheridan counties in Montana; Hettinger, Kidder, and Eddy counties in North Dakota; Butte, McPherson, and Day counties in South Dakota; and Grant County in Minnesota (Johnson and Schwartz 1993a, 1993b). Field size varied from <1.0 to 111.7 ha. In the northern Great Plains, most CRP land was planted to a mixture of native and/or non-native grass- es and legumes (Johnson and Schwartz 1993a). Woody vegetation has encroached into some idle CRP fields, and some CRP fields contain wetlands or were partially inundated by water during a recent wet period (Ig] 2001; Ig] and Johnson, unpublished data). Methods Breeding bird and cowbird surveys In both studies, we conducted total area counts of breeding birds using the strip-transect procedure em- ployed by Stewart and Kantrud (1972: also see Ig] and Johnson 1997). During the North Dakota statewide study, bird surveys were conducted by two observers on foot, and each observer surveyed cowbirds and other breeding birds on a rectangular half (805 * 402 m;: 32.37 ha) of a quarter-section by following a standard- ized survey route. This route was 100 m inside of, and parallel to, the boundary of the rectangle. The rectan- gular halves were usually surveyed simultaneously, and an interval of 400 m was maintained between observers. Deviations of up to 100 m from the route were often necessary to adequately survey all habitats. Bird species were identified by sight or sound. Counts during precipitation and strong winds (> 24 km/h) were avoided. Surveys of open country birds were con- ducted between 0.5 h after sunrise and 15:00 CST. Although some surveys occurred outside the time of 2007 FiGurRE |. Distribution of 130 quarter-sections (64.75 ha each) in North Dakota in which bird surveys were conducted during the breeding seasons in 1992 and 1993, most active bird vocalizations (i.e., early morning or late evening), Stewart and Kantrud (1972) concluded that singing and other activities of open-country birds were not appreciably affected by time of day (also see Vickery 1995*). Quarter-sections containing exten- sive woodland habitats were usually covered on rela- tively calm (<8 km/h), sunny days between 0.5 h after sunrise and 10:00 CST. These limitations were neces- sary because song frequencies and other activities of most woodland birds are reduced on cloudy days, in moderate or high winds, and at mid-day. Counts of breeding birds were based primarily on the number of indicated breeding pairs on territories or home ranges during peak breeding periods. All sample units were surveyed for early-nesting species between 24 April and 7 June, for mid-nesting species between 14 May and 10 July, and for late-nesting species between 22 May and 21 July. When a survey was conducted during an overlapping portion of the peak breeding periods, counts of early-, mid-, and late- nesting species coincided. Thus, quarter-sections that were visited between 22 May and 7 June were only surveyed once, and those that were surveyed before 22 May were surveyed again after 7 June so as to in- clude species from all three breeding periods. For most species, nearly all indicated pairs were observed as segregated pairs or as territorial males. For the sexually dimorphic Brown-headed Cowbird, we separately tallied the males and females seen per sample unit; herein, we report only female cowbird densities. We did not consider certain birds observed during the census- es to be breeding and excluded them from our results. These included late-migrating flocks and other birds passing overhead in high, direct flight. By counting birds only during their peak breeding periods, we max- imized the potential for recording breeding birds and, at the same time, minimized the likelihood of con- founding breeding birds with migrants. In the CRP study, total area counts of breeding birds were conducted by using a minor modification of the strip-transect procedures used by Stewart and Kan- IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS 24) Sheridan e NDISaya Ne es {Ys eA J Hetiinger u Kidder a McPherson zB Day SD FIGURE 2. Counties in which Conservation Reserve Program grassland fields were surveyed for breeding birds in North Dakota, South Dakota, Minnesota, and Mon- tana, 1990-2006. trud (1972) and Ig] and Johnson (1997). Small fields usually were surveyed by a single observer; large fields typically were surveyed by two observers on foot, each simultaneously covering about one-half of the field. The number and configuration of transects were con- sistent among years but varied depending upon the field size and shape to achieve total coverage of a field. We surveyed each CRP grassland field once each year between 21 May and 9 July in 1990-2006. Nests and cowbird parasitism Prior to entering the field in each year, the observers were trained in bird, nest, and egg identification. Nests were found serendipitously during breeding bird sur- veys and related research activities. Most nests were found by unintentionally flushing an adult host (or rarely a female cowbird) from the nest or nest vicinity. The remainder of the nests were found by observers as they were simply walking through a habitat. When a nest was located, we inspected it and recorded the species (based on the identification of the flushing adult, the nest, or the nest contents) and the number of eggs or nestlings of the host and of cowbirds. If the nest species could not be identified at the time of the initial observation, the nest location was temporarily marked and then revisited and identified at the end of the survey by an experienced observer. We classified a nest as parasitized if it contained at least one cow- bird egg or nestling, regardless of the stage of the nest- ing cycle. Cowbird and host nestlings were identified based on size, gape and flange colors, and down or contour feather characteristics and development. We were unable to return to nests to monitor their out- come or to document additional brood parasitism. We rarely located or checked the contents of nests of up- per-canopy nesting species. Most of the nests that we located were found in the egg-laying stage or the in- cubation stage (see below); cowbirds lay most of their eggs before or during the hosts’ egg-laying stage (Johns- gard 1997). Although we found many nests in the nest-building stage (i.e., before the onset of egg lay- ing), we summarize here only active nests, which we defined as nests attended by adults that contained at least one host or cowbird egg and/or nestling. We also 242 found a few abandoned nests that contained cowbird eggs, but these are not reported because cowbirds are known to parasitize inactive nests (Freeman et al. 1990). The presence of cowbird fledglings that were fed by adult hosts also was noted during the surveys of breed- ing birds, although an observation of a potential host feeding a cowbird fledgling does not establish conclu- sively the true foster-parentage of the fledgling cow- bird (sensu Klein and Rosenberg 1986). We use the frequency (%) of nests parasitized as an index to the levels of parasitism. We define the frequen- cy (%) of cowbird parasitism as the total number of parasitized nests (*100) divided by the total number of active nests found for a given species, group of species (e.g., grassland birds), county, or time period (week or year). We define the frequency (%) of multiple par- asitism as the total number of nests containing more than one cowbird egg (*100) divided by the total num- ber of parasitized nests found for a given species, group of species, county, or time period. The average (+ SD) number of cowbird eggs per parasitized nest (1.e., par- asitism intensity) was calculated by summing the num- ber of cowbird eggs found in all parasitized nests in both studies and dividing by the total number of par- asitized nests. Although cowbirds may avoid parasitiz- ing nests of ejector species (Sealy and Bazin 1995), we include nests of all passerine species, because ejector species are sometimes parasitized (e.g., Scott 1977; Peer et al. 2000), because parasitism of ejector species may vary geographically (e.g., Haas and Haas 1998) or with time of sympatry with cowbirds (Briskie et al. 1992), and because it is difficult to estimate how often ejector species are parasitized (e.g., Scott 1977). To evaluate changes in cowbird parasitism within the breeding season, we divided each month into four, roughly weekly, periods (7, 8, 7[8] and 8 days, respec- tively) (sensu Orians et al. 1989). For discussion pur- poses, we grouped rates of parasitism and multiple par- asitism into five categories: low (<10%), low-to- moderate (10-20%), moderate (20-40%), moderate- to-heavy (40-50%), and heavy (>50%) (sensu Ortega 1998: 184). Scientific names of potential host species are included in Table 2. To determine whether cowbirds avoid laying eggs in previously parasitized nests (Mayfield 1965a; Ori- ans et al. 1989), we evaluated the distribution pattern of cowbird eggs (i.e., random or non-random) using a zero-truncated Poisson distribution model for only para- sitized nests (Lindsey 1997). A random distribution of cowbirds eggs suggests that female cowbirds did not discriminate among parasitized nests and that the pro- portion of nests with 1, 2, 3, 4, ..., i cowbird eggs ap- proximated a Poisson distribution (Orians et al. 1989). A non-random distribution of cowbird eggs indicated that the random distribution model was rejected (P <0.01). For both studies, we calculated overall parasitism frequencies, mean densities of female cowbirds, mean avian densities, and mean avian richness. For the CRP THE CANADIAN FIELD-NATURALIST Vol. 121 grassland study, we calculated mean avian density and richness among five groups (all of which excluded cow- birds): (1) all avian species, (2) all passerine species, (3) all passerines known to have raised cowbird young (Ortega 1998; Davis and Sealy 2000), (4) all passer- ines excluding ejectors (Peer and Sealy 2004), and (5) a single, preferred host (i.e., Red-winged Black- bird; see below). We were interested in the regional avian influences that affect cowbird distribution and abundance rather than short-term influences, and thus we averaged across the 17 years within a county and did not analyze the survey data separately for individ- ual years. For the CRP grassland study, we used female cowbird densities both as an explanatory variable in relation to rates of cowbird parasitism and as a res- ponse variable in relation to avian community variables (i.e., density and richness). Linear regression analyses (PROC REG; SAS Institute, Inc. 2004) were performed (1) to determine the relationship between overall cow- bird parasitism and female cowbird densities, and (2) to examine the relationship between female cowbird densities and the five levels of the avian community. Results In the North Dakota study, we observed 160 breed- ing bird species, including 78 nonpasserine species and 82 passerine species (Igl and Johnson 1997). Given that grasslands, croplands, and wetlands are the three most common habitats in North Dakota (Ig] and Johnson 1997), most of the common breeding bird species in North Dakota are associated with such open habitats. The five most abundant species in the North Dakota study, in decreasing order of abundance, were the Horned Lark, Chestnut-collared Longspur, Red-winged Blackbird, Western Meadowlark, and Lark Bunting (scientific names given in Table 2). Average breeding bird densities in a quarter-section, excluding cow- birds, were 126 breeding pairs per 100 ha in 1992 and 143.9 breeding pairs per 100 ha in 1993 (Table 1). The average numbers of species observed in a quar- ter-section (64.75 ha) were 18.5 species in 1992 and 21.6 species in 1993. Average cowbird densities were 3.0 female cowbirds per 100 ha in 1992 and 4.3 female cowbirds per 100 ha in 1993. Female Brown- headed Cowbirds were found in 66.7 and 76.7% of the quarter-sections in 1992 and 1993, respectively. In the CRP grassland study, we observed 143 breed- ing bird species, including 78 nonpasserine species and 65 passerine species. The five most abundant species, in decreasing order of abundance, were Savannah Spar- row, Grasshopper Sparrow, Red-winged Blackbird, Clay-colored Sparrow, and Western Meadowlark. Av- erage breeding-bird densities within a county (aver- aged across years and excluding cowbirds) ranged from 104.1 to 208.1 breeding pairs per 100 ha, and the average number of species observed within a county ranged from 26.8 to 52.7 species (Table 1). Average cowbird densities within a county ranged from 0.6 to 8.0 female cowbirds per 100 ha, and the average annual i 2007 IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS 243 TABLE |. Summary of Brown-headed Cowbird densities (females/100 ha), cowbird occurrence, breeding bird densities (all species excluding cowbirds; breeding pairs/100 ha), and number of breeding bird species per quarter-section in two studies in the northern Great Plains, 1990-2006, Female cowbird Year or Study Location density Mean SE North Dakota statewide study 1992 3.0 0.3 1993 AS» (0/5 CRP grassland study (1990-2006) Butte County, South Dakota 0.6 0.1 Day County, South Dakota 8.0 0.7 Eddy County, North Dakota 6.3 0.4 Fallon County, Montana 0:9" 10:2 Grant County, Minnesota 3h 0.4 Hettinger County, North Dakota 7) 0.5 Kidder County, North Dakota 519, 1033 McPherson County, South Dakota fell 0.5 Sheridan County, Montana 2.8 9 o 30. , frequency of female cowbird occurrence ranged from 12.4 to 54.1%. Average cowbird densities and frequen- cies were lowest in the counties on the eastern and western edges of our study area (Table 1, Figure 2). This pattern mirrors cowbird distribution maps from the North American BBS (Price et al. 1995; Sauer et al. 2005*), which show cowbird abundance declining both east and west of central North Dakota and South Dakota. We found 351 active nests of 51 species (18 non- passerine and 33 passerine species) during the North Dakota statewide study and 2298 active nests of 62 species (23 nonpasserine and 39 passerine species) during the CRP grassland study, for a total of 2649 nests of 75 species (Table 2). Most of the nests of nonpasserine species were found during the egg- laying or incubation stages; 89.9% of the 746 nests of nonpasserine species contained only eggs, and 10.1% contained only nestlings or both eggs and nestlings. Only one nonpasserine nest was parasitized by a cowbird (Table 2). An Upland Sandpiper nest with four sandpiper eggs and one cowbird egg was found in a CRP grassland field in Sheridan County, Montana, on 28 June 1993. Hereafter, we summarize data only for passerine nests. Most of the active nests of passerines were found during the egg-laying or incubation stages; 85.8% of the 1902 passerine nests contained only eggs, and 14.2% of the nests contained only nestlings or both eggs and nestlings. Overall, the parasitism rate of passerine nests was moderate; 476 of the 1902 (25%) passerine nests that we found during the two studies were parasitized by cowbirds (Table 2). Twenty-four of the 45 passerine species were found parasitized (Table 2). Among species with 25 or more nests, parasitism levels were low for Western Kingbird (0%), Eastern Kingbird (1.7%), Brown Thrasher (3.8%), Savannah Sparrow (9.8%), Chestnut-collared Longspur Female Breeding Number cowbird bird of breeding bird occurrence density Species Mean % SE Mean SE Mean SE 0.67 - 126.0 7.8 18.5 0.8 0.77 ~ 143.9 7,3 216 09 0.14 0.02 105.8 5.9 28.5 1.4 0.54 0.03 208.1 14.5 48.8 2.7 0.49 0.03 188.7 15.1 45.5 2.5 0.12 0.02 104.1 9.| 26.8 1.2 0.25 0.02 181.8 13.7 39.6 2.2 0.52 0.03 160.0 9.3 35/74 1.3 0.50 0.01 160.2. 14.1 Sse LS 0.54 0.02 164.5 9.8 43.8 2.4 0.29 0.05 122.3) 30.4 1.1 (7.7%), and Common Grackle (1.1%); low-to-moder- ate for Horned Lark (15.6%), Clay-colored Sparrow (11.7%), Lark Bunting (19.4%), Grasshopper Sparrow (19.7%), Bobolink (19.2%), and Western Meadowlark (16.1%); moderate for Brewer's Blackbird (33.3%); and moderate-to-heavy for Red-winged Blackbird (43.1%). The Red-winged Blackbird was the most commonly parasitized species in the two studies (71.2% of all cases of parasitism), and we classified it as a preferred host in subsequent analyses. Parasitized passerine nests contained from zero to six host eggs and/or nestlings and from one to eight cowbird eggs and/or nestlings (Table 3). Of the 476 parasitized passerine nests, 50.6% contained one cow- bird egg, 28.6% contained two, 12.0% contained three, and 8.8% contained four or more cowbird eggs. The maximum number of eggs or nestlings in a parasitized nest (host and cowbird combined) was nine. The aver- age number of cowbird eggs per parasitized nest was 1.9 + 1.2 (Tables 2 and 3). Multiple parasitism occurred at 12.4% of all passerine nests and at 49.4% of all par- asitized passerine nests. Parasitized nests of 18 passer- ine species contained multiple cowbird eggs and/or nestlings (Table 2). Among parasitized species with 25 or more nests, multiple parasitism rates were low for Eastern Kingbird (0%), Brown Thrasher (0%), and Common Grackle (0%); moderate-to-heavy for Clay- colored Sparrow (43.5%), Lark Bunting (38.9%), Grass- hopper Sparrow (44.4%), Chestnut-collared Long- spur (50.0%), and Red-winged Blackbird (49.6%): and heavy for Horned Lark (80.0%), Savannah Spar- row (62.5%), Bobolink (53.3%), Western Meadowlark (100%), and Brewer’s Blackbird (58.3%). Among these species, the average number of cowbird eggs per par- asitized nest was highest for the Western Meadowlark (3.4 + 1.6); all nine parasitized nests of the Western Meadowlark contained two or more cowbird eggs. 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(panuyuo)) ‘7 ATAV], 246 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 3. Contents of parasitized passerine nests found incidentally during breeding bird surveys in two studies in the northern Great Plains, 1990—2006. Number of host eggs or nestlings per nest J 2 3 0) 37 2) 8 J 40 23 9 2 50 4| 14 3 58 37 18 4 45 13 zy 5 10 | 5 6 | Total number of nests 24\ 136 57 Number of Brown-headed Cowbird eggs or nestlings per nest Total number of nests 1 1 77 83 114 4 5 6 7 8 5 zy 2 | SW BON WwW NN We 22 10 8 ] 1 476 TABLE 4. Yearly variation of Brown-headed Cowbird parasitism in passerine nests (all species combined) found incidentally during breeding bird surveys in two studies in the northern Great Plains, 1990-2006. Frequency Total number of cowbird Year of nests parasitism (%) 1990, 57 29.8 199] 52 15.4 1992! 18] 12.7 1993' 90 20.0 1994 4] ID 1995 32 18.8 1996 20 0) 1997 42 16.7 199% 96 24.0 1999 96 25.0 2000 75 25.3 2001 129 Sbeyei 2002 184 34,8 2003 189 24.9 2004 234 29.9 2005 231 23,8 2006 153 26.8 Mean + SE UMA ete Uhre PLUG) a0) Pel Frequency of multiple parasitism (%) Distribution of cowbird eggs in parasitized nests> Parasitism intensity (mean + SD) 5B) 1.6+0.9 random 50.0 1.8+0.9 random PJP M phe \\ 5) random 55.6 Dd w= MN L6) random 62.5 1.9+0.8 random Bos lgs) (05) random 0 ~ - 42.9 16+0.8 random 60.9 2.0 + 1.4 random 62.5 24+1.6 random 57.9 EO FENRZ random 34.8 1.5+£1.0 random 50.0 f yse ell random 46.8 LS + 1.2 random Sif 1.8+0.8 random 36.4 17+ 1.4 non-random 56.1 19+ 1.2 random 46.9 + 3.8 "Mean number of cowbird eggs or nestlings per parasitized nest. "Random distribution of cowbird eggs in parasitized nests was evaluated using a zero-truncated Poisson distribution model (Lindsey 1997), Random distribution of cowbirds eggs indicated that female cowbirds did not discriminate among parasitized hests and that the proportion of nests with 1, 2, 3,4, .../ cowbird eggs approximated a Poisson distribution, Non-random distibution indicated that the random distribution model was rejected (P< 0.01). "Data for the North Dakota statewide study and the CRP study were combined for 1992 and 1993, Not including nests of the Red-winged Blackbird, our sample of passerine nests from the two studies in cluded mostly nests of grassland-nesting (e.g, West ern Meadowlark) and edge-nesting (¢.¢., Yellow War bler) species and very few nests of cavity-nesting (c.2., House Wren) or woodland-interior (¢.g., Ovenbird) species (Table 2), We found 482 nests of 15 passerine species that typically nest in grasslands, Overall, the parasitism rate Of grassland bird nests was low-to moderate (16.4%), We found 602 nests of 23 passerine species that typically nest in habitats characterized by woody vegetation (Le,, woodland, shrubland, open habitat with scattered trees or shrubs, open or semi open deciduous woodland, and edge), The parasitism rate of bird species associated with woody vegetation was low (9.5%). Of the 45 passerine species for which nests were found, nine species are known to eject cowbird eggs (Table 2; Peer and Sealy 2004), Of the 257 nests of these nine ejector species that we found, t2 (4.7%) contained cowbird eggs. The overall parasitism rate tor nests of all non-ejector (.c,, acceptor) passerine species combined was 28.2% (1 = 1645), In addition, Yellow Warblers often bury the contents of parasitized nests under a new nest floor (Clark and Robertson 198t; Burgham and Picman 1989), Eight of 24 (44.4%) Yel 2007 IGL and JOHNSON; COWBIRDS IN THE NORTHERN GREAT PLAINS 247 TABLE 5. Weekly variation of Brown-headed Cowbird parasitism in passerine nests (all species combined) found incidentally during breeding bird surveys in two studies in the northern Great Plains, 1990-2006 Frequency Frequency Parasitism Distribution of Weekly Total number of cowbird of multiple intensity cowbird eggs in period of nests parasitism (%) parasitism (%) (mean + SD)' parasitized nests’ April IV 5 0 0 May I 0 0 0 May II 6 0 0 May III 1] 9.1 100.0 4.0 + 0.0 May IV 9] 11.0 60.0 2.6+2.0 random June | 274 19.7 50.0 J Vee Wis} random June II 4020 S53 Sill See non-random June Il 421 27.8 53.0 19+ 1.1 random June [V 454 20.9 41.1 1.6+0.9 non-random July I 162 21.0 50.0 2.0 + 1.4 random July U 39 23.1 33.3 1.4+0.7 random July 1 37 16.2 50.0 2,2 E16 random Mean + SE 158.5 +51.9 20.7 + 2.8° 54.3 + 6.2° - - “To evaluate changes in cowbird parasitism within the breeding season, each month was divided into four, roughly weekly, periods (week I = 7 days, week II = 8 days, week III = 7[8] days, and week IV = 8 days) (sensu Orians et al. 1989). See Table 4 for explanation. °Mean includes third week in May to third week in July (i.e., weeks with cowbird parasitism). low Warbler nests each contained a single cowbird egg. During the CRP study, buried (but visible) cow- bird eggs were observed in two Yellow Warbler nests (one in Kidder County, North Dakota, on 27 June 2003 and one in Eddy County, North Dakota, on 25 June 2005) and one Clay-colored Sparrow nest (McPher- son County, South Dakota, on 13 June 2006). Parasitized nests of passerines were found in 16 of the 17 years (Table 4). No parasitized nests were found in 1996, which also is the year that we found the fewest number of active passerine nests. Excluding 1996, an- nual parasitism rates ranged from 12.7 to 35.7% and averaged 22.6% (Table 4). The average annual rate of multiple parasitism was 46.9%. The zero-truncated Poisson distribution model for parasitized nests was rejected in one (i.e., 2005) of the 16 years in which par- asitized nests were found. In both studies combined, passerine nests were found in 11 of 12 weekly periods from the last week of April through the third week of July (Table 5). Eleven passerine nests were found in two of the three weeks before the first cowbird egg was found (i.e., before the third week in May), although cowbirds were recorded in each of those three weeks (Igl and Johnson, unpub- lished data). In the North Dakota statewide study, we found active nests between 27 April and 20 July, par- asitized nests between 20 May and 18 July, and mul- tiply parasitized nests between 20 May and 18 July. In the CRP grassland study, we found active nests be- tween 22 May and 9 July, parasitized nests between 29 May and 9 July, and multiply parasitized nests be- tween 29 May and 9 July. During both studies, non- parasitized and parasitized nests were found largely between late May and early July, which corresponds with the peak breeding periods of most passerine species in the northern Great Plains, including the Brown-headed Cowbird (Stewart 1975; Ig] and Johnson 1997; Tallman et al. 2002). After the third week in May, weekly parasitism rates ranged from 9.1 to 37.3% and averaged 20.7%; the highest parasitism rates were found in the second and third weeks of June (Table 5). The average weekly rate of multiple parasitism was 54.3%. The zero-truncated Poisson distribution model for parasitized nests was rejected in two (i.e., second and fourth week of June) of eight weekly periods. In the CRP grassland study, parasitized and multi- ply parasitized nests of passerines were found in all nine counties (Table 6). Cowbird parasitism of passerine nests was low in Fallon County in southeastern Mon- tana (5.8%) and Grant County in western Minnesota (9.3%). Parasitism rates were low-to-moderate in Butte County in western South Dakota (12.8%) and Sheridan County in northeastern Montana (11.6%), and moder- ate in Hettinger (29.9%), Kidder (25.3%), and Eddy (23.8%) counties in North Dakota and Day County in northeastern South Dakota (28.9%). Heavy parasitism was found in McPherson County in north-central South Dakota (51.5%). The frequency of multiple parasitism was moderate in Butte (21.4%) and Day counties in South Dakota (39.8%) and Fallon County, Montana (25%), and moderate-to-heavy to heavy in the other six counties (range: 42.9-58.7%). The zero-truncated Poisson distribution model for parasitized nests was rejected in one (i.e., Day County, South Dakota) of the nine counties (Table 6). Nests within the counties of highest female cowbird abundance experienced high- er frequencies of cowbird parasitism (R* = 0.64, F = 12.65, P = 0.0093) than those found in counties with the lowest female cowbird abundance (Figure 3). The quadratic term for cowbird densities was non- significant (P = 0.62) and was not included in the final model. 248 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 6. Regional variation in Brown-headed Cowbird parasitism in passerine nests (all species combined) found incidentally during breeding bird surveys in Conservation Reserve Program grassland fields in nine counties in the northern Great Plains, 1990-2006. Frequency of cowbird Total number County of nests parasitism (%) Butte, South Dakota 109 12.8 Day, South Dakota BD 28.9 Eddy, North Dakota We 23.8 Fallon, Montana 69 5.8 Grant, Minnesota 151 9.3 Hettinger, North Dakota 211 29.9 Kidder, North Dakota 332 25.3 McPherson, South Dakota 233) 51.5 Sheridan, Montana 69 11.6 “See Table 4 for explanation. In the CRP grassland study, cowbird densities, total breeding bird densities, and total breeding bird rich- ness were lowest in the southwestern counties (Butte County, South Dakota, and Fallon County, Montana) and highest in counties near the center of the cowbird’s breeding bird range. There was a positive relationship between female cowbird densities and all measures of avian abundance and richness. That is, female cowbird densities increased with increasing density and rich- ness at all levels of the avian community (Table 7). Breeding bird density for all bird species (R* = 0.72) was a better predictor of cowbird abundance than the other four levels of avian densities (Table 7, Figure 4a). Breeding bird richness for all bird species (R* = 0.69) was a better predictor of cowbird abundance than the other three levels of avian richness (Table 7, Figure 4b). There was only a weak suggestion that cowbirds use, as a settlement cue, the number of passerines species known to have raised cowbird young (P = 0.115). During breeding bird surveys in both studies, ob- servers noted 586 instances of adult passerines provi- sioning food to individual fledglings, of which only 26 (4.4%) instances involved hosts feeding cowbird fledglings. These 26 observations occurred between 4 June and 7 July and involved eight host species: Clay- colored Sparrow (5 cases), Grasshopper Sparrow (1 case), Le Conte’s Sparrow (1 case), Savannah Sparrow (6 cases), Song Sparrow (4 cases), Brewer’s Blackbird (2 cases), Red-winged Blackbird (5 cases), and Orchard Oriole (2 cases). These eight species accounted for 83.6% of the total cowbird parasitism in these two stud- ies (Table 2). Although we cannot establish conclu- sively the true foster-parentage of these fledgling cow- birds, we did not observe any non-parasitized species (i.€., species whose nests were not parasitized during the two studies) feeding cowbird fledglings during the two studies. Discussion Brown-headed Cowbirds parasitize nests of a wide variety of host species (Friedmann 1963; Lowther 1993), including some inappropriate hosts. As in other Distribution of cowbird eggs in parasitized nests* Parasitism intensity (mean + SD) Frequency of multiple parasitism (%) 21.4 1.2+0.4 random 39.8 ilove IL non-random 46.3 1.8+ 1.1 random 25.0 1.3+0.5 random 42.9 1.5+0.7 random 58.7 Dp), = M5) random 45.2 1.7+0.9 random 59.2 AAdes IIail random 50.0 Allee i} random studies (Ortega 1998), we found almost no cowbird parasitism of nonpasserine nests. Most nonpasserines are considered unsuitable hosts, and their nests usu- ally are avoided by Brown-headed Cowbirds (Roth- stein and Robinson 1998). Although the only parasi- tized nonpasserine, an Upland Sandpiper, was clearly an unsuitable host (1.e., a large-bodied host with pre- cocial, nidifugous young), several other studies have documented cowbird eggs in Upland Sandpiper nests (Higgins and Kirsch 1975; Faanes and Lingle 1995*; Davis and Duncan 1999). Our overall frequency of cowbird parasitism (25%) for passerine species was within the range that Ortega (1998) considered moderate (20-40%). This overall rate seems low given that cowbird densities are highest in the northern Great Plains (Sauer et al. 2005*) and that cowbirds are more abundant than many of their common hosts in this region (Ig] and Johnson 1997; Ig] et al. 1999). However, our combined parasitism rate is based on nests from two studies, an extensive region, 17 years, and numerous passerine species, including several species that ostensibly have developed behav- ioral defenses against cowbird parasitism (see below). After we excluded nests of ejector species, our over- all parasitism rate remained moderate (28.2%). Some studies in this region have found higher combined rates of cowbird parasitism than those reported here- in. For example, Koford et al. (2000) found an over- all parasitism rate of 40.3% for passerines nesting in seeded and native grasslands and cropland in North Dakota and Minnesota, and Davis and Sealy (2000) found an overall rate of 32.6% for species nesting in grasslands in southwestern Manitoba. In both of these studies, some individual species experienced moder- ate-to-heavy levels of cowbird parasitism (e.g., >40% parasitism for Western Meadowlarks in both studies). Robinson and Smith (2000) suggested that some of these commonly cited reports of high parasitism rates from the northern Great Plains might not be represen- tative of this region. Indeed, some recent reviews of geographic patterns of cowbird parasitism (e.g., Peer et al. 2000; Chace et al. 2005) focused on the extreme 2007 > oO y = 4.2169x + 3.1958, R? = 0.6437 ho ta abe Ch Cee it eo Parasitism rate (%) DR inidise oii ode Bion iarn8 o 9 Cowbird Densities (99 / 100 ha) FIGURE 3. Relationship between female cowbird densities and the percentage of the total number of passerine nests that were parasitized in Conservation Reserve Program grasslands in North Dakota, South Dakota, Minnesota, and Montana, 1990-2006. The data from the North Dakota (N) statewide study were included for compar- ison (1992 and 1993 were averaged). cases of parasitism in the northern Great Plains (e.g., Linz and Bolin 1982; Davis and Sealy 2000; Koford et al. 2000). There are many less-cited studies of cowbird parasitism (see Shaffer et al. 2003) with low-to-mod- erate rates of cowbird parasitism in this region (e.g., Hill and Sealy 1994; Granfors et al. 2001; Davis 2003: Woolfenden et al. 2004). For example, in Manitoba, Davis (2003) found an overall parasitism rate of 19.9% for six grassland species in southern Saskatchewan over a 5-year period. Although our overall rate of cowbird parasitism was 25%, we did find higher rates of parasitism in some years (e.g., 35.7% in 2001), weeks (e.g., 37.3% in the second week of June), counties (e.g., 51.5% in McPherson County, South Dakota), and individual species (e.g., 43.1% for the Red-winged Blackbird). The results from our studies can serve as a reminder that cowbird parasitism rates are not constant across space or time (i.e., among years or within a breeding season) and that cowbirds differentially parasitize hosts within the same habitats. Differences in parasitism lev- els among studies in the northern Great Plains may simply reflect variation among species, regions, habi- tats, or years. For example, Davis and Sealy’s (2000) overall parasitism rate of 32.6% was based on data col- lected at three study sites over two years; parasitism rates were low-to-moderate at two of the study sites (18 and 20%) and heavy at the third study site (67%). We echo Hahn and Hatfield’s (1995) caveat that patterns of cowbird parasitism are complex and variable, and that field studies of cowbird parasitism would be more effective if they were long-term, community-wide stud- ies set in strategically chosen communities in different regions. Our nest data were slightly unorthodox compared to nest data from many other studies. We located all of IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS 249 en v— Oo Oo ot Ba z ® (=) a] With Grant: y = 0.0616x - 5.0673, R? = 0.715 ri Without Grant: y = 0.0707x - 6.0706, R? = 0.908 = °o ° 100 120 140 160 180 200 22 Breeding Bird Densities (pairs / 100 ha) = 10 f= oOo aug = 6 a= y = 0.2357x - 4.6603, R? = 0.687 oO Cowbird Densities (4 ho ine) or 30 35 40 45 50 55 Species Richness (al! species) FiGuRE 4. Relationship between (a) mean female cowbird densities and average densities of all other breeding bird species, and (b) mean female cowbird densities and mean species richness within the breeding bird community (excluding cowbirds) in Conservation Reserve Program (CRP) grasslands in North Dakota, South Dakota, Minnesota, and Montana, 1990-2006. The circle in Figure 4a indicates a potential outlier, Grant County in Minnesota, which is discussed in the text. our nests fortuitously and observed each nest only once. Any cowbird eggs ejected before our surveys, any nests abandoned before our surveys, any cowbird eggs buried completely in the nest lining, and any cowbird eggs added or removed after our brief observations were not recorded in our studies, and thus our frequencies of cowbird parasitism should be considered minimal esti- mates. These limitations, however, are not unique to our data but rather are common features of all studies of cowbird parasitism that do not involve cameras, because most nests are not monitored continuously throughout the nesting cycle but rather are visited at relatively fixed intervals (e.g., every 24 days) for rela- tively brief periods of time (e.g., less than a few min- utes). Even systematic nest searches are neither inces- sant nor comprehensive, but rather occur at fixed intervals (e.g., Koford et al. 2000; Davis 2003; Winter 250 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 7. Five levels of avian abundance (density) and richness (number of species) that female Brown-headed Cowbirds could use as proximate cues for settlement in Conservation Reserve Program grassland fields in nine counties in the northern Great Plains, 1990-2006. Linear regression analyses were performed to examine the relationship between female cowbird densities and each different subset of avian abundance and richness at the five levels of the avian community (also see Figure 4). Level of avian community* Breeding bird density Number of breeding species R? F P R? F IP All breeding birds 0.72 17.53 0.004 0.69 15.56 0.006 All passerines 0.63 12.02 0.011 0.40 4.65 0.068 All passerines that have raised cowbirds 0.66 13.64 0.008 0.32 325) 0.115 All passerines that are non-ejectors 0.62 11.39 0.012 0.38 4.32 0.076 Preferred host (Red-winged Blackbird) 0.54 8.35 0.023 - = = 4 Excluding cowbirds. et al. 2004). In addition, many studies that use sys- tematic nest searches also include a fortuitous com- ponent to their nest searching methodology (e.g., Koford et al. 2000; Davis 2003). Thus, nearly all esti- mates of cowbird parasitism in the literature could be considered minimal estimates. In our two studies, female cowbirds did not avoid laying eggs in previously parasitized nests. Nearly one- half of the parasitized nests in our two studies con- tained more than one cowbird egg, with the average parasitized nest containing about two cowbird eggs. The assumption that female cowbirds laid eggs ran- domly in parasitized nests could only be rejected in two of eight weekly periods, one of 16 years, and one of nine counties in the CRP grassland study. Deviations from a random Poisson distribution were likely relat- ed to a greater number of nests with multiple cowbird eggs or nestlings than would be expected by chance. Multiple parasitism has been widely documented in nests parasitized by the Brown-headed Cowbird (Mc- Laren et al. 2003). Although multiple parasitism can vary by region or host species, it typically occurs in about one-third of all reported cases of cowbird para- sitism (Friedmann 1963; Lowther 1993; Johnsgard 1997; Ortega 1998) and seems to be more common in nests of larger hosts (Lorenzana and Sealy 1999; Trine 2000). Little is known about the factors that might influence multiple parasitism, but some mechanisms have been proposed or investigated. For example, mul- tiple parasitism might occur if (1) a female cowbird exhibits host preference (Smith and Myers-Smith 1998), (2) there is a high ratio of female cowbirds to host nests (McGeen 1972), (3) there is a shortage of alternative host nests (Smith and Myers-Smith 1998), or (4) an inexperienced and nonselective female makes a poor host choice (McLaren et al. 2003). Cowbird hosts in the Great Plains seem to be particularly vul- nerable to multiple parasitism (e.g., Elliott 1978; Davis and Sealy 2000; this study), which may reflect higher densities of cowbirds (Elliott 1977; Ortega 1998) or lower densities of available or suitable hosts (Robin- son and Smith 2000). We likely underestimated the frequency of cowbird parasitism of those species that eject or bury cowbird eggs or that abandon parasitized nests (Ortega 1998). Less than 5% of the nests of ejector species in the two studies contained cowbird eggs, which was much lower than the overall parasitism rate (28.2%) for acceptor species. Ejection of cowbird eggs by a host is an effec- tive anti-parasite strategy among some North America passerines (Ortega 1998; Peer and Sealy 2004), espe- cially among edge-nesting passerines that are common in the northern Great Plains (Ig] and Johnson 1997; Table 1). Even within acceptor species, however, some individuals are prone to abandon parasitized nests and then re-nest (Lorenzana and Sealy 1999). Hosoi and Rothstein (2000) showed that the frequency of deser- tion of parasitized nests was higher in non-forest than forest species and suggested that increased nest deser- tion was an evolved response to cowbird parasitism. Among small-sized hosts, nest desertion seems to be a common response to parasitism or defense against parasitism (Ortega 1998). Many species that breed in the northern Great Plains are known to abandon their nests occasionally as a result of parasitism or cowbird removal of host eggs (e.g., Ortega 1991; Hosoi and Rothstein 2000), including some grassland and shrub- grassland species, although the frequency of abandon- ment may be quite low (Sealy 1999; Davis and Sealy 2000). The Red-winged Blackbird was the third most abundant species observed in both of our studies. Despite its aggressive nest-defense behaviors against cowbirds and its colonial-nesting tendencies (Free- man et al. 1990), the Red-winged Blackbird was the most commonly parasitized species in the two studies (43.1% parasitism, 49.6% multiple parasitism, 71.2% of all cases of parasitism) and appears to be a pre- ferred cowbird host in the northern Great Plains. Our data support Hanka’s (1979) contention that cowbirds may show a phylogenetic preference for Red-winged Blackbirds and other Icterine species (e.g., Brewer’s Blackbird, Orchard Oriole; Table 2) in the cowbird’s range. Linz and Bolin (1982) and Koford et al. (2000) reported similar parasitism rates (42 and 43%, respectively) for Red-winged Blackbirds in_ this region. In the eastern portion of the cowbird’s range, Red-winged Blackbirds are rarely parasitized (e.g., 2007 Hahn and Hatfield 1995). Although we categorized the Red-winged Blackbird as a wetland species (Table 2; Igl and Johnson 1997), many of the red- wing nests that we found during the CRP grassland study were scattered in upland habitats (grassland, shrub-grassland), where group defense is generally lower (Robertson and Norman 1977; Ortega 1991) and parasitism rates are generally higher (Robertson and Norman 1976; Krapu 1978). Several host species are known to avoid or reduce parasitism by acting aggressively toward cowbirds (Robertson and Nor- man 1976, 1977), but aggression is not always an effective host defense against cowbird parasitism, espe- cially in habitats in which host densities are low be- cause cowbirds may use host aggression (Robertson and Norman 1977) or vocalizations (Clotfelter 1998) to locate nests. Nonetheless, the Red-winged Blackbird is one of a few species reported to reduce parasitism by mobbing (Freeman et al. 1990; Chace et al. 2005). The presence of Red-winged Blackbirds may be a double-edged sword for other passerines nesting in the same habitats, such as CRP grasslands. Red-winged Blackbirds are conspicuous breeding birds in CRP grasslands and thus might increase parasitism in nests of other species. Barber and Martin (1997) found that a conspicuous, co-occurring species in a breeding bird community can increase the risk of cowbird parasitism on alternative host species. Alternatively, other species may benefit from nesting near Red-winged Blackbirds if (1) the defensive behavior of blackbirds toward cow- birds reduces cowbird activity in the area, or (2) higher cowbird parasitism of blackbird nests reduces para- sitism pressure on other species. Fretwell (1972) noted higher rates of cowbird parasitism in Dickcissel nests that were built near Red-winged Blackbird nests, whereas Clark and Robertson (1979) found the oppo- site for Yellow Warblers. Woolfenden et al. (2004) speculated that Red-winged Blackbirds might have in- creased the risk of parasitism on Yellow Warblers in their Manitoba study sites. The influence of Red-winged Blackbird presence and abundance on cowbird para- sitism rates of other species requires further study. Mayfield (1965b) suggested that grassland birds may have evolved defenses against brood parasitism because of their long evolutionary history of co- occurrence with the Brown-headed Cowbird in the Great Plains. To date, however, the species that have shown the strongest anti-parasite behaviors (i.e., egg ejection) have not been the grassland species but rather have been the species associated with woody vegetation (Peer and Sealy 2004, Table 2). Our overall rate of cow- bird parasitism for grassland bird nests (16.4%) was much lower than those reported in some studies (e.g. Elliott 1978 [57.7%]; Davis and Sealy 2000 [32.6%]; Jensen and Finck 2004 [42.9%]) but comparable to or higher than those reported in other studies (e.g., Straus- berger and Ashley 1997 [0%]; Kershner and Bollinger 1998 [1.7%]; Robinson et al. 2000 [7.8%]; Granfors et al. 2001 [14%]; Davis 2003 [19.9%]; Winter et al. IGL and JOHNSON: COWBIRDS IN THE NORTHERN GREAT PLAINS 251 2004 [6.7%]). Nonetheless, we found a higher fre- quency of parasitism in nests of grassland species than in nests of species that nest in edge and woodland habi- tats (9.5%). In contrast to our results, in Illinois, Rob- inson et al. (1999, 2000) and Strausberger and Ashley (1997) found a higher frequency of parasitism in spe- cies nesting in edge and woodland habitats than in nests of grassland species. Although the difference between the two regions for grassland birds is consistent with the pattern that cowbird parasitism declines with dis- tance from the center of the cowbird’s range in the northern Great Plains, the difference for species asso- ciated with woody vegetation is contrary to this gen- eralization. Again, these patterns emphasize the com- plexity and variability of cowbird parasitism in North America (Hahn and Hatfield 1995). Robinson and Smith (2000) and others (Peer et al. 2000; Jensen and Finck 2004) have suggested that the higher levels of parasitism in grasslands in the Great Plains than those in the Midwest might reflect the near absence of large forested areas, where more tolerant hosts can absorb cowbirds and cowbird parasitism from grasslands. Seemingly, our data support this contention. In our most-forested county in the CRP grassland study (Grant County in west-central Minnesota; Ig] and Johnson, unpublished data), the density of female cowbirds (and cowbird parasitism) was much lower than would be expected given the density of breeding birds in the avian community (Figure 4a). Moreover, Minnesota typical- ly is lumped with midwestern states rather than with Great Plains states. In the other eight counties, there are few, if any, large forested areas that could absorb cowbird parasitism from the open grasslands. Trees and shrubs in these eight counties occur largely in small, often linear, patches, where cowbird egg ejectors (e.g., kingbirds, thrashers, catbirds, robins) dominate the breeding bird community. The low parasitism rates of edge and woodland species in our study may reflect the predominance of ejector species in woody habitats in the northern Great Plains (Igl and Johnson 1997) and the avoidance by cowbirds of a habitat that may con- tain many less-tolerant hosts (Sealy and Bazin 1995). In Illinois, Peer et al. (2000) recently found that Western Meadowlarks rejected (i.e., ejected or dam- aged) 78% of artificial and real cowbird eggs experi- mentally added to nests. In contrast, in the northern Great Plains, some Western Meadowlark populations are moderately to heavily parasitized (23-67%: Davis and Sealy 2000; 19-47%: Koford et al. 2000), and the species often experiences extreme levels of multiple parasitism in this region (Davis and Sealy 2000; Davis 2003; this study). It is unclear why some meadowlark individuals or populations experience heavy or intense parasitism (e.g., average of 3.4 and 3.1 cowbird eggs per parasitized nest in this study and Davis and Sealy 2000, respectively), whereas others apparently expe- rience little or none. Davis and Sealy (2000) suggest- ed that the apparent geographic variation in para- sitism levels in Western Meadowlark populations in 252 the northern Great Plains and elsewhere might reflect differences in the size of suitable grassland patches and the availability of other host species in the com- munity. Peer et al. (2000) suggested that geographic differences in parasitism levels in the northern Great Plains and the Midwest might reflect geographic vari- ation in egg rejection by Western Meadowlarks. Egg rejection behavior in the Western Meadowlark should be tested in more geographic locations or landscapes to resolve these issues. Moreover, the Western Mead- owlark and its congener, the Eastern Meadowlark (Sturnella magna), use a similar tactic to destroy eggs or clutches of other birds (Creighton and Porter 1974; Schaef and Picman 1988; Picman 1992). The function of this egg-destroying behavior has been speculative (e.g., predation, interference competition for limited resources), but in view of the similarities (e.g., egg removal, puncture) between this egg-destroying behav- ior at nests of other birds and cowbird egg ejection at their own nests, experiments are needed to evaluate the relationship between general egg destruction and cowbird egg rejection by meadowlarks. In our CRP grassland study, female cowbird den- sities varied greatly among the nine counties in the northern Great Plains, and rates of cowbird parasitism were higher in the counties in which densities of fe- male cowbirds were higher. Our results support the interpretation of Miles and Buehler (2000) and Chace et al. (2005) that female cowbird densities are a rea- sonable predictor of the frequencies of cowbird para- sitism. Our results also support the contention that cow- bird parasitism declines with distance from the center of the cowbird’s range, although on a smaller scale (Hoover and Brittingham 1993; Smith and Myers- Smith 1998; Thompson et al. 2000). Few studies have evaluated the relationship be- tween cowbird abundance and parasitism levels using cowbird and nest data collected concurrently at the same study sites over a large region (e.g., Robinson et al. 2000; Jensen and Cully 2005a, 2005b). Large- scale evaluations of this relationship typically have relied on data from unconnected sources, such as abun- dance data from the North American Breeding Bird Survey (BBS) and nest data from the Cornell Labora- tory of Ornithology or from multiple studies (e.g., Hoover and Brittingham 1993; Basili 1997; Smith and Myers-Smith 1998; Herkert et al. 2003). Most evaluations of this relationship have focused on single species (Hoover and Brittingham 1993; Basili 1997; Smith and Myers-Smith 1998; Herkert et al. 2003). Nonetheless, many studies have found similar positive relationships between cowbird densities and cowbird parasitism levels. For example, Basili (1997) and Herkert et al. (2003) showed that cowbird densities on BBS routes were positively related to frequencies of cowbird parasitism of Dickcissel nests from multiple studies across several states. Jensen and Cully (2005a,b) found a similar, positive relationship between female cowbird densities and parasitism levels of Dickcissel THE CANADIAN FIELD-NATURALIST Vol. 121 nests at eight study sites in Kansas and Oklahoma. In a meta-analysis of nest and cowbird data from six stud- ies from four midwestern states, Thompson et al. (2000) similarly found that cowbird abundance and para- sitism levels of multiple species were positively cor- related across study areas. In contrast, Woolfenden et al. (2004) found that female cowbird abundance was not related to parasitism frequency among three species in an avian community in Manitoba, and Robinson et al. (2000) found that the percentage of nests para- sitized was not related to cowbird abundance in mul- tiple forest sites across Illinois. In CRP grasslands, we also found a positive rela- tionship between cowbird abundance and all five lev- els of the avian community, including all avian species, all passerine species, all passerines that are non-ejec- tors, all passerines known to raise cowbird young, and a single, preferred host (i1.e., Red-winged Blackbird). The abundance and richness of all avian species were the strongest predictors of cowbird abundance, which suggests that cowbirds might use the overall breeding bird community as a settlement cue for optimal areas to breed rather than more specific passerine or host categories (e.g., all passerines that are known to have raised cowbird young). The number of individuals and the number of species represent different components of a cowbird’s breeding resource. The presence of more individuals and more species with different breeding chronologies ensures a larger number of potential hosts within the breeding bird community throughout the cowbird’s breeding season (Farmer 1999). Although we evaluated these relationships using only data from the CRP grassland study, cowbirds also might use other cues (e.g., vegetation or landscape factors) as an indi- cator of the richness or abundance of the avian com- munity (Chace et al. 2005). Chace et al. (2005) tabulated 16 published studies and one unpublished study that evaluated the relation- ship between different measures of the avian commu- nity and cowbird abundance or occurrence. As with our results, many studies found positive relationships between cowbirds and different measures of avian den- sities and richness. To our knowledge, Farmer (1999) is the only other published study to have evaluated mul- tiple host and avian community measures using both abundance and richness data. Farmer (1999) found positive relationships for nearly every level of the avian community, including abundance and richness cate- gories. Because host species are necessary for cow- birds to be reproductively successful, most studies have evaluated these relationships at the host level. Results of such evaluations have been inconsistent. Moreover, comparisons among studies using host categories are difficult because a variety of definitions of host com- munities were used (Chace et al. 2005). For example, Thompson et al. (2000) found that cowbird abundance was significantly correlated with host abundance (i.e., hosts included species that bred on the study area and accepted cowbird eggs), whereas Robinson et al. (2000) 2007 found no relationship between cowbird abundance and host abundance (i.e., hosts included species that accept cowbird eggs regularly but excluding cavity nesters, cowbird egg ejectors, and large species). Evans and Gates (1997) found that cowbird abundance was positively related to the abundance of all avian species combined, but the relationship was non-significant for richness of all avian species and richness and abun- dance of host species (1.e., known cowbird hosts). In summary, our data show that, even within the northern Great Plains, there is considerable variation in parasitism rates among species and across space and time. Our results demonstrate that parasitism rates over- all are moderate in the northern Great Plains, although some individual species, counties, years, and weeks ex- perience heavier rates of parasitism or multiple para- sitism than others. Our study is one of the first to show a strong, positive relationship between regional cow- bird abundance and nest parasitism rates using both cowbird abundance and nest data collected at the same study sites. We also found a clear, positive relation- ship between female cowbird densities and the over- all breeding bird communities in CRP grasslands in the northern Great Plains. More studies are needed to determine if these relationships hold elsewhere in the northern Great Plains with different host communi- ties or within other habitats. Acknowledgments We thank K. L. Andersson, I. Balodis, K. A. Dal- ton, J. E. Doster, D. A. Hobbick, C. J. Johnson, K. F. Kuehnl, J. M. Legge, R. L. Manson, J. W. Marlow, M. C. Marlow, L. A. Murphy, K. L. Richardson, T. R. Runia, M. D. Schwartz, C. M. Shoemaker, M. L. Sondreal, J. M. Steiner, and K. A. Ward for their assis- tance in the field. We are grateful to the numerous land owners and operators who allowed us access to their property. J. E. 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Gibbs, C. M. McLaren, and S. G. Sealy. 2004. Community-level patterns of parasitism: use of three common hosts by a brood parasitic bird, the Brown-headed Cowbird. Ecoscience 11: 238-248. Received 27 August 2007 Accepted 12 March 2008 Wolf, Canis lupus, Behavior in Areas of Frequent Human Activity ELLEN HEILHECKER!, RICHARD P. THIEL’, and WAYNE HALL, JR.? University of Wisconsin-Stevens Point, College of Natural Resources, Stevens Point, Wisconsin 54481 USA *Wisconsin Department of Natural Resources, Sandhill Wildlife Area, Box 156, Babcock, Wisconsin 54413 USA Heilhecker, Ellen, Richard P. Thiel, and Wayne Hall, Jr. Wolf, Canis lupus, behavior in areas of frequent human activity. Canadian Field-Naturalist 121(3): 256-260. We report incidental observations of Wolves (Canis lupus) tolerating human activity in central Wisconsin. Three monitored packs raised pups in close proximity to varying levels of human activity. Wolf pups were raised <350m from rearing pens of the endangered Whooping Crane (Grus americana), which saw daily human activity. One pack used cornfields as ren- dezvous sites within 175 m of a maintenance shed visited regularly by workers. Another pack centered their activities along a well-traveled state highway using both the verge and the road center for activity. Aerial locations of 10 yearling and adult dispersing Wolves were plotted to evalute human densities in natal territories relative to dispersal and post-dispersal territo- ries. Township densities (¥ = 9.02 humans/km?, SE = 4.015) and residential densities (¥ = 5.59 housing units/km?, SE =2.12 ) in natal pack territories were significantly greater (P <.01) for dispersal and post-dispersal township densities (¥ = 43.98 humans/km?, SE =7.37) and residence densities (¥ = 23.12 housing units/km*, SE =3.49). Furthermore, a pup negotiated the densely populated region of northern Illinois and dispersed from central Wisconsin to east-central Indiana, a distance of at least 690 km. As Wolves live 1n closer proximity to humans, living in areas of higher township and residential densities, they can be expected to be more habituated to people, increasing the probability of human/Wolf conflicts. Key Words: Wolf, Canis lupus, pups, humans, habituation, behavior, Wisconsin. Historically, Gray Wolves (Canis lupus) were one of the most widely distributed land mammals in North America inhabiting all regions except arid deserts (Young 1944; Mech 1970; Mech and Boitani 2003). However, human settlement and persecution restrict- ed Wolves to northern forested regions and eventually the only substantial population within the lower 48 states occurred in northern Minnesota (Mech 1970). Under the protection of the federal Endangered Species Act of 1973, Wolves have returned to some areas within their historical range (Mech 1995). They began recolonizing Wisconsin in the mid-1970s (Mech and Nowak 1981; Thiel and Welch 1981; Wydeven et al. 1995). Wolves have since repopulated much of north- ern Wisconsin and an isolated area in the central part of the state referred to as the Central Forest Region (CFR). In 1999, the Wisconsin Department of Natural Resources (WDNR) downlisted the Wolf from an endangered to a threatened species (WDNR 1999*). Wolves continued to surpass target recovery levels. In April 2003, the U.S. Fish and Wildlife Service pro- posed to delist the Eastern Distinct Population Seg- ment of Gray Wolf, which includes Wisconsin (Fed- eral Register 2003*). Wolves initially recolonized areas in Wisconsin with significantly lower road densities (y = 0.23 km/km/?) and human densities (y = 1.52 people/ km*) (Mlade- noff et al. 1995). As recovery progresses, Wolves are demonstrating an increasing tolerance to human activ- ity throughout the upper Great Lakes region, including Wisconsin (Thiel et al. 1998). In recent years, Wolves have denned near active sphagnum moss drying sites, occupying homesites inside artillery impact zones on military bases, crossed four lane highways and large expanses of non-forested areas, and bypassed major U.S. cities (Licht and Fritts 1994; Mech et al. 1995; Thiel et al. 1998). While such behavior is not unique to | the species, it is unique to Wolves living in the conti- nental United States (McNay 2002; Mech and Boitani 2003). Observations of Wolves displaying increased tolerance towards humans are of value insofar as it af- fects management of Wolf recovery. This paper reports on the success of Wolves attempting to colonize areas of higher human densities than previously reported in the continental United States. Study Area and Methods Wisconsin’s Central Forest Region (CFR) encom- passes 7767 km? of deciduous forest (Acer saccharum, Tilia americana, Quercus alba, Q. borealis, Q. veluti- na), coniferous swamps (Picea mariana, Larix larici- na), aspen stands (Populus tremuloides, P. grandiden- tata), pine barrens (Pinus banksiana, P. resinosa, P. strobus), sphagnum bogs, and marshes (Curtis 1959; Finley 1976*). A 12 141-hectare swamp, the largest in Wisconsin, lies within the CFR (Martin 1965). The») CER includes portions of Eau Claire, Chippewa, Clark, || Jackson, Wood, Monroe, Juneau, Adams, and Mar- quette counties. The majority of the CFR consists of | private industrial, county, state and federal forestland. Major economic industries include logging and cran- berry agriculture as well as dairy operations (WDNR 2003*). Road density within the CFR is 1.23 km/km? (U. S. Census Bureau 2000*; WDNR 2003%*). Wolves were captured on or near rendezvous sites, | located by howling and track surveys, using offset, 256 2007 modified #14 Newhouse traps during summers 2002 and 2003, following guidelines established by Ani- mal Care and Use committees within the Wisconsin Department of Natural Resources and University of Wisconsin-Stevens Point. With the exception of setting traps, researchers minimized time spent in pack ren- dezvous sites to avoid Wolf-human interactions with study animals. Wolves were sedated with a 5:1 mL mixture of ketamine hydrochloride and xylazine and were weighed, sexed, and examined to assess general condition (Wydeven et al. 1995). Radio collars were placed on adults. Pups received ear tag transmitters developed for a Wolf pup mortal- ity study (Advanced Telemetry Systems, Isanti, Min- nesota) programmed to transmit from 0700 — 1900. Pups and adults were located from the ground using a receiver with either a 5-element hand held Yagi anten- na or a 9-element vehicle mounted antenna once daily from time of capture to 15 January each study sea- son. All Wolves were located from fixed-wing air- craft 1-2 times/week. Human activity observed near pack homesites was documented. Additionally, we interviewed people and collected reports from people who encountered study pack Wolves to assess the relative level of tolerant be- havior of study packs towards humans. Aerial locations of 10 dispersing Wolves were plot- ted in Universal Transverse Mercator (UTM) coordi- nates utilizing ArcView 3.X (ESRI, Inc., Redlands, California). Township (human) and residential (struc- tural) densities for each county in which Wolf loca- tions were plotted were queried (U. S. Census Bureau 2005*). Measurements obtained included township densities and residential densities in the county for (1) natal territories from which Wolves dispersed, (2) locations while Wolves were dispersing, and (3) after dispersers settled and either became territorial or be- came breeders. Results We monitored 17 Wolves from three study packs in 2002 and nine Wolves in five study packs in 2003. Seven Wolves and three packs were new in 2003. In three of the six study packs we were able to detail repeated interactions with humans. Suk Cerney pack: Three male pups, one female pup, a yearling female, and an adult male Wolf were radioed near their den site between 3-6 July 2002 on Necedah National Wildlife Refuge (NNWR) in Juneau County (44°04.957'N, 90°10.399'W). The den site was locat- ed approximately 350 m from an observation bunker and several rearing pens used as part of a Whooping Crane (Grus americana) reintroduction program locat- ed on the NNWR. Between July and October employ- ees and volunteers of the crane project, NNWR staff. along with visiting news media and guests, visited the cranes at least twice per day. Crane staff and Wolf re- searchers routinely observed pups before and during HEILHECKER, THIEL, and HALL: WOLF BEHAVIOR tN A ~~ . LY. from inet Of f Wisconsin YA Ai \ Uy . fee J \ ( Madison Q j f y |Miwauked ey ) |} Michigan ya 1, Chicago / ( jf - oe SS _/ Gary i ew ) | | , Minis | * Jet Ne } Indiana | “be ’ ~ 2 é t 4 ¥ i; ) ma ARAN ( s we =) \ a x ae ra bela of ~ W E FIGURE |. Capture site of Wolf 409 in central Wisconsin on 10 August 2002 and death site in eastcentral Indiana. June 2003. our study period. One pup watched Wolf researchers from the den, located in oak savannah dominated by Hill’s oak (Quercus ellipsoidalis) for approximately 10 minutes as traps were set on 3 July 2002, approxi- mately 10.5 m away. Adult Wolves were rarely seen. However, radio signals confirmed they were often in the immediate vicinity. The 4 radioed pups stayed in the area despite daily human activity, until their deaths between July 15 and 5 September 2002 (Heilhecker 2003*). Bear Bluff pack: Six Wolves were captured in and adjacent to cornfields on private land in Jackson Coun- ty between 20 July and 3 August 2002 (44°15.355'N, 90°20.582'W). Two male pups and two female pups, a yearling female, and a yearling male were radio-tagged. A partially planted, sandy soil cornfield with stunted corn stalks < 0.9 m tall, used as a daily rendezvous site from July to early November in 2002 and 2003, was approximately 175 m from the maintenance building used by employees of the cranberry farm at least twice a week. Distance from the rendezvous site to the main- tenance building and cranberry beds were approximate- ly 450 m and 1300 m, respectively. White-tailed Deer (Odocoileus virginianus), the Wolves’ primary prey in the region, were abundant and cause $35000/year dam- age to local commercial cranberry beds (K. Rice, Flying Dollar Cranberry Company, personal commu- nication). Deer flies (Chrysops sp.) and mosquitoes (Culicidae) were generally less abundant in the corn- 258 fields compared to the surrounding forested land. In both years, and for years prior to our study, employees of the cranberry company regularly observed Wolves. Once during our study they witnessed a Wolf in their parking lot sniffing vehicle tires. The property man- ager also reported his dog interacting with Wolves in what looked like play behavior. Noch Hanai pack: Wolf 429, a yearling female, was radio-collared on 3 June 2002. In summer 2002, she was observed multiple times acting indifferently to vehicles along State Highway 54 on the northern border of her natal pack territory (44°20.15'N, 90°35.51'W). In the autumn she dispersed and created a new pack that occupied a 102-km/? territory centered along the same state highway. The pack’s den site was discov- ered approximately 230 m from State Highway 54 ina Red Pine (Pinus resinosa) plantation with no under- story. Vehicles passing on the state highway were visi- ble from the den. The wolves’ rendezvous sites, as determined by matted grass, trails, human garbage with teeth marks, and deer legs, were on embankments along the shoulder of this moderately traveled state highway, adjacent side roads, and recently logged forests span- ning a distance of 7.7 km by 60 m along the state high- way. In summer 2003, over 60 citizen phone calls and e- mails reported a radio-collared Wolf with pups walk- ing, playing, and lying in the middle of the state high- way. Initially four pups and a radio-collared adult were observed. On 19 July 2003, a dead male pup was col- lected from the shoulder of the highway. The necropsy revealed injuries consistent with a vehicle collision. None of the Wolves appeared to be afraid of vehi- cles. On 20 July 2003, a citizen reported watching three pups and a collared female in the middle of the state highway. They moved off to the shoulder of the road to let vehicles pass and then returned to the middle of the road. E. Heilhecker and W. Hall, Jr., observed Wolf 429 looking both directions before stepping onto the highway. Pups were observed sitting on the side of the road chewing on deer legs as vehicles passed. Pups once walked next to Heilhecker and Hall’s vehicle parked on the shoulder of the road. In all instances the pups appeared oblivious to the presence of vehicles. It was later learned that the Wolves had been fed by loggers working in this area during the late spring/early summer. On 13 January 2004, WDNR received a report of an uncollared Wolf on State Highway 54 jumping into the bed of a truck containing deer hides and carcass- es after its occupant had stopped to take pictures of the Wolf (M. Windsor and T. Babros, WDNR, person- al communication). On 29 March 2004, an uncollared Wolf was observed walking back and forth across State Highway 54 stopping traffic. At one point the Wolf picked up a yellow plastic oil container, crossed the road, and lay down within 3 m of a vehicle. Several vehicles slowed down to watch the Wolf. While no one exited their vehicles, people rolled down their win- THE CANADIAN FIELD-NATURALIST Vol. 121 dows and continued to talk within hearing range of the Wolf. The Wolf did not appear to be affected by the commotion. On 27 April 2004, the radio-collared female, Wolf 429, was humanely dispatched along Hwy 54 because she was unable to ambulate on her hind legs. A preliminary necropsy was inconclusive, but her paralysis was not caused by a vehicle collision. This Wolf did have a healed injury to the pelvic girdle that was caused by blunt trauma suggestive of an ear- lier collision with a vehicle (A. Dassow, University of Wisconsin, Madison, personal communication). Dispersal: We compared aerial locations for 10 Wolves, as yearlings and adults, dispersing between 1998 and 2004. Seven Wolves (W002, W269, W309, W338, W341, W426, W429) established new territo- ries; five of which (W309, W338, W341, W426, W429) were known to have successfully bred. Three Wolves (W337, W427, W480) continued to disperse until their deaths and never set up known territories. Town- ship densities (x = 9.02 humans/km?, SE = 4.015) and residential densities (y = 5.59 housing units/km?, SE =2.12 ) in natal pack territories were significantly lower (P < 0.01) than township densities (z = 43.98 humans/km?, SE = 7.37) and residential densities (x = 23.12 housing units/km?, SE = 3.49) in dispersal and post-dispersal territories (U. S. Census Bureau 2005*). A male pup, W409, dispersed to eastern Indiana in less than 156 days. The pup was last monitored on 15 January 2003 and found dead on 20 June 2003. If the pup dispersed in a straight line, it would have passed through relatively densely settled areas of northern Illinois and the greater metropolitan Chicago area to reach eastern Indiana (40°06.4'N, 85°04.3'W) (Figure 1). The human density of Jackson County, its birth- place, was 7.8 people/km?. The least populated coun- ties along the Wisconsin/Illinois state border had 22.9 people/km? (Green County, Wisconsin) and 14.5 peo- ple/km? (Jo Daviess County, Illinois). The human den- sity in Randolph County, Indiana, where the Wolf was found shot was 22.84 people/km? (U. S. Census Bureau 2005*). This was the first documented Wolf in Indiana since their extirpation in 1908 (Mumford and Whitaker 1982), and the southern-most movement of a radio-collared Wisconsin Wolf to date (A. Wyde- ven, WDNR, personal communication). Discussion Wolves’ reactions to humans are a reflection of their experiences with people and the relative tolerance of humans to Wolves (McNay 2002; Fritts et al. 2003). By the mid-1900s Wolves were largely restricted to wilderness areas in North America due to intense human persecution (Young 1944; Mech 1995). With legal protection from the Endangered Species Act of 1973, a gradual increase in acceptance of Wolves (Fritts et al. 2003), and an expanding population, Wolves have inevitably come in closer contact with humans (Mech 1995; Thiel et al. 1998; Mech and 2007 Boitani 2003). By the 1990s an increasing number of Wolves began dispersing into and colonizing more settled regions and agricultural areas in the northern United States (Licht and Fritts 1994; Mech 1995; Mech and Boitani 2003). Thiel et al. (1998) and Merrill and Mech (2000) reported tolerance of humans close to den and rendezvous sites in Minnesota and Wisconsin. As Wolves colonize more fragmented habitat dom- inated by humans they will come into more frequent contact with humans, as our findings demonstrate. Dis- persing Wolves from the numerous packs presently inhabiting the forest-farmland zones in Wisconsin and Minnesota are capable of penetrating great distances into landscapes highly dominated by humans (Licht and Fritts 1994; Merrill and Mech 2000; Fritts et al. 2003; Mech and Boitani 2003). The male Wolf pup, 409, provides an example of this progression of tolerance to humans, a behavior that can be expected to be exhibited by Wolves in succeeding generations. Although movements of = 670 km from their homesites have been well docu- mented in Wolves (Van Camp and Gluckie 1979; Bal- lard et al. 1983; Fritts 1983; Ream et al. 1991; Licht and Fritts 1994; Merrill and Mech 2000; Mech and Boitani 2003), this is the first documentation of a wild North American Wolf moving through areas of such high human densities. While Wolf 409 was not known to have any contact with nor was known to have been conditioned to humans as a pup, it was subjected to increasing levels of human activity as it dispersed south and east of Lake Michigan. Interestingly, Wolf 429, the founder of what became the human-habituated Noch Hanai pack, came from the very same pack in which Wolf pup 409 was born. We observed two forms of habituation of Wolves in our study: (1) pups that became accustomed to the pres- ence of humans through repeated, benign association, and (2) association of humans with a food reward. In the two cases where litters were raised in close proxim- ity to human activity and were frequently seen, the pups fled at the close presence or approach of a human(s). These Wolves created no known threats to public safe- ty or caused any damage to personal property. Wolves displaying avoidance or fear behavior during direct encounters with humans are considered to be showing “normal” behavior (Smith and Stahler 2003*). In the remaining case in our study, pups received food awards from humans. Wolves are not known to attack people during their first human encounter but require repeated exposure to humans before attack- ing (McNay 2002; Smith and Stahler 2003*). A pre- requisite for aggressive Wolf behavior towards humans is habituation, with food being the most influential factor (McNay 2002; Smith and Stahler 2003*). Our human-habituated Wolves’ whose bold behavior rap- idly evolved into a public nuisance issue. We expect more Wolf-human encounters and human- habituated Wolves due to the recovering Wolf popula- HEILHECKER, THIEL, and HALL: WOLF BEHAVIOR 259 tion in the Great Lakes area. Wisconsin's Wolf popu- lation has increased from 83 Wolves in 1995 to 425 Wolves in 2005 (Wydeven and Wiedenhoeft 2005*) The growing Wolf population and increased number of Wolf sightings have raised safety concerns from the general public. A continuous educational program that focuses on how human actions affect Wolf behav- ior may minimize the establishing of human-habituated Wolves. Periodic news releases and public education programs at the federal and state forests, posters at road side kiosks emphasizing the reasons why feeding of Wolves is improper, writing stipulation in public con- tracts on public land prohibiting feeding of Wolves especially in areas where Wolf habituation is occur- ring, or fining individuals who feed Wolves may be useful approaches. When habituation occurs, attempts should be made to deter Wolves displaying bold behav- ior with aversion techniques such as rubber bullets and cracker shells before Wolf aggression develops. If aversion conditioning is unsuccessful, for public safety reasons, aggressively bold Wolves should be humanely euthanized. Acknowledgments This study was funded by the Zoological Society of Milwaukee County, National Fish and Wildlife Foun- dation’s Budweiser Conservation Scholarship, Univer- sity of Wisconsin-Stevens Point, and the WDNR. We thank Angela Dassow (UW-Madison), Adrian Wyde- ven (WDNR), Paul Anderson (WDNR), Peggy Calla- han and the Wildlife Science Center, Kirby Rice and the Flying Dollar Cranberry Company, Advanced Telemetry Systems, Inc., Neal Paisley and the employ- ees of Sandhill Wildlife Area (WDNR), Michele Windsor (WDNR), Tim Babros (WDNR), Tim Beyer (WDNR), the staff at the NNWR, the Bloomington Field Office of the Indiana Department of Natural Resources (IDNR), Lori Pruitt, Scott Pruitt, Larry Har- ris (USFWS), Kent Hanauer (IDNR), and Jayne Bel- sky (NNWR volunteer). Documents Cited (marked * in text) Federal Register. 2003. USFWS. Endangered and threat- ened wildlife and plants; removing the eastern Gray Wolf from the list of endangered and threatened wildlife. | April 2003, Docket Number 03-7020. Finley, R. W. 1976. Original vegetation cover of Wisconsin (map). http://www.dnr.state.wi.us/org/aV/et/geo/map_gal/ landcov/orgveg. Heilhecker, E. 2003. 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Merrill, S., and L. D. Mech. 2000. Details of extensive move- ments by Minnesota Wolves (Canis lupus). American Mid- land Naturalist 144: 428-433. Mladenoff, D. J., T. A. Sickley, R. G. Haight, and A. P. Wydeven. 1995. A regional landscape analysis and pre- diction of favorable gray wolf habitat in the northern Great Lakes region. Conservation Biology 9: 279-294. Mumford, R. E., and J. O. Whitaker, Jr. 1982. Mammals of Indiana. Bloomington Indiana University Press, Bloom- ington, Indiana. 537 pages. Ream, R. R., M. W. Fairchild, D. K. Boyd, and D. H. Pletscher. 1991. Population dynamics and home range changes in a colonizing wolf population. Pages 349-366 in The greater Yellowstone ecosystem: redefining America’s wilderness heritage. Edited by R. B. Keiter and M. S. Boyce. Yale University Press, New Haven, Connecticut. Thiel, R. P., and R. J. Welch. 1981. Evidence of recent breeding activity in Wisconsin wolves. American Mid- land Naturalist 106: 401-402. Thiel, R. P., S. Merrill, and L. D. Mech. 1998. Tolerance by denning wolves, Canis lupus, to human disturbance. Canadian Field-Naturalist 112: 340-342. Van Camp, J., and R. Gluckie. 1979. A record long dis- tance move by a wolf (Canis lupus). Journal of Mammal- ogy 60: 236. Wydeven A., R. N. Schultz, and R. P. Thiel. 1995. Moni- toring of a recovering gray wolf population in Wisconsin, 1979-1999. Pages 147-156 in Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S. H. Fritts, and D. R. Seip. Canadian Circumpolar Insti- tute, Canada. Young, S. P. 1944. The wolves of North America. American Wildlife Institute, Washington, D.C. 385 pages. Received 11 April 2006 Accepted 5 June 2008 Influence of Gender and Den Type on Home Range Shape for Striped Skunks, Mephitis mephitis, in Saskatchewan SERGE LARIVIERE!, DAVID HOWERTER?, and FRANCOIS MESSIER® 'Delta Waterfowl Foundation, R.R. 1, Box 1, Site 1, Portage La Prairie, Manitoba RIN 3A1 Canada. Present address: Cree Hunters and Trappers Income Security Board, 2700 boulevard Laurier, Champlain #1110, Québec (Québec) G1 V 4K5 Canada “Institute for Wetland and Waterfowl Research, Ducks Unlimited Canada, P.O. Box 1160, Stonewall, Manitoba ROC 270 Canada 3 Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada Lariviere, Serge, David Howerter, and Francois Messier. 2007. Influence of gender and den type on home range shape for Striped Skunks, Mephitis mephitis, in Saskatchewan. Canadian Field-Naturalist 121(3): 261-264. We investigated gender differences in shape of home ranges for Striped Skunks, Mephitis mephitis, in southcentral Saskatchewan, Canada, during 1993-1994. Data collected on free-ranging Striped Skunks indicated that shape of female home ranges was unaffected by den type (building versus burrow) and was similar to shape of male home ranges. Moreover, type of maternity den did not influence the position of the den within the home range (inside or outside of core area). We conclude that the dis- tribution of Striped Skunks in summer, and the distribution of their foraging activity, are not affected by the availability of anthropogenic den structures. Key Words: Striped Skunk, Mephitis mephitis, home range, denning site, spatial distribution, Canadian prairies. Fragmentation of habitats affects not only how ani- mals are distributed in the landscape, but also their use of space. In fragmented landscapes, suitable habitats for foraging may not occur in proximity to other criti- cal habitats for denning, parturition, or rearing of young. Therefore, mammalian females that live in frag- mented landscapes face two problems: establishing a home range that includes suitable foraging habitat and finding a site for parturition and rearing of young. For males that do not provide parental care, obtain- ing food is a primary drive affecting behavior during summer. In contrast, females give birth and raise a lit- ter during the same period, and habitat preferences for raising young may not coincide with good forag- ing habitats. In fragmented landscapes, the compro- mise between optimal foraging and optimal rearing habitats may affect the establishment of home ranges, and consequently their shape. Thus, the location of maternity dens may have several implications for the distribution of foraging activity of females. First, den location may affect the shape of the home range or the position of the den site within the home range. For example, species that rely on a burrow and central place for refuge and storage of food may benefit from a centrally located den and circular home range | (Bowers 1995). However, in fragmented landscapes, females may have to prioritize either establishment of home range (find a good foraging area first, then find a suitable place to rear young within the home range), or _ the establishment of maternity den (find a good den _ first, then find a good place to forage). We investigated the influence of gender and den _ types on home range shape of Striped Skunks (Mephi- _ tis mephitis) in the parklands of Saskatchewan, Cana- ae! da. Striped Skunks are distributed across most of Cana- da and the United States (Rosatte and Lariviére 2003). Females give birth in mid-May, and because males pro- vide no parental care, the progeny are dependent on the mother until mid-July, when the young disperse (Lariviére and Messier 1997). In Saskatchewan, female Striped Skunks use two main types of shelter for mater- nity dens, either anthropogenic structures (e.g., build- ings, ca. 60% of maternity dens) or underground burrows (ca. 40% of maternity dens) (Lariviere and Messier 1998c). The distribution and availability of both den types differ, and thus, we hypothesized that den type could influence home range shape. For example, build- ings may provide good denning structure, but their position may not be in proximity to good foraging areas, a scenario that would yield an elongated home range. In contrast, a burrow may be dug presumably closer to good foraging areas, and thus home range of females using burrows could be more circular in shape. Thus, Striped Skunks provide a good model species to test predictions of den site positioning and home range shape by animals occupying fragmented landscapes. Study Area and Methods We conducted this study in the Prairie Pothole Region of southcentral Saskatchewan (52°45'N, 107°08'W). Small grain (1.e., wheat, barley, oats) and oil crops (mostly canola but also flax) occupy 60% of the land- scape. Numerous wetlands and stands of Trembling Aspen (Populus tremuloides) occur throughout the area. Topography is gently rolling, and an extensive network of roads divides the land. Areas managed for nesting waterfowl are common throughout the area, and consist mostly of dense nesting cover, a mixture of native and tame grasses seeded specifically to attract upland nesting waterfowl. General characteristics of the Prairie Pothole Region are detailed elsewhere (Greenwood et al. 1995). 261 262 From April to August, 1993-1994, Striped Skunks were captured in opaque-sided livetraps (Larivieére and Messier 1999) and anesthetized using halothane and Telazol® (Lariviére and Messier 1996a). All individuals were equipped with a 5-sec delay motion-sensitive radio-collar (150-152 Mhz, Telonics Inc., Mesa, Ari- zona, USA) to monitor activity during tracking (Lari- viere and Messier 1997). Skunks were released at the site of capture. Radio-collared animals were located by an observer using hand-held receiving equipment from 18:00 to 06:00, during the period of greatest activity (Lariviere and Messier 1997). Individuals were located every 15 min by direct observation, auditive location, or short- range (<50 m) triangulation (Lariviere and Messier 1998b). Light amplifying, night-vision goggles (AN- PVS 5, Bill’s Electronics Ltd., Mildmay, Ontario) enabled visual observations at night. Tracking effort was systematically distributed among all skunks, and no animals were tracked during two consecutive nights. Striped Skunks exhibit aposematic behavior, and defensive postures are obvious (Lariviére and Messier 1996b). Locations immediately following an observer- induced defensive posture were discarded from analy- ses due to possible observer disturbance (Lariviére and Messier 1998b). Universal Tranverse Mercator coordinates were ob- tained from the exact location of the skunk following its departure from the area using a portable Global Positioning System (Ensign GPS, Trimble Naviga- tion, Cansel Survey Equipment, Burnaby, British Columbia). To increase the precision of the coordinates, we only considered locations for which =3 satellites were available (D’ Eon 1995). Our radio-tracking schedule was intensive (1.e., loca- tions every 15 min), and designed to maximize direct behavioral information instead of statistical independ- ence within an animal’s home range. Using Shoener’s ratio, we estimated that statistical independence be- tween locations occurred at intervals of 270 min. How- ever, Striped Skunks are highly mobile, and are capable of moving >1 km in 15 min, even in dense vegetation (Lariviére and Messier 1998b). Thus, we assumed that short movements reflected behavioral decisions by the skunk, not physical constraints. We defined maternity dens as any sites for which at least two of the following criteria were satisfied: repeated use for >3 consecutive days during the par- turition/rearing period (15 May - 30 June), collection of grass for preparation of nest chamber, or presence of young during the rearing period (Lariviére and Messier 1998a). Because some females may relocate their prog- eny during the rearing period, and because we could not assess why females changed maternity dens (e.g., to reduce parasite loads, disturbance, relocation to a better foraging area, etc.), we only used data on females with a single maternity den for spatial analysis of den locations within home ranges. THE CANADIAN FIELD-NATURALIST Vol. 121 Our entire study area was digitized into a Geograph- ic Information System (SPANS GIS, PCI Geomatics, Richmond Hill, Ontario, Canada). Locations of the maternity dens were plotted for all female skunks, as well as all telemetry locations to determine home range size (Lariviere and Messier 1998c) and shape. Home range shape was quantified using a shape index (McGarigal and Marks 1994) that was calculated by the following formula: shape index = perimeter / (2* Vv (pi * area)). Thus, a smaller shape index indicates a more cir- cular range. We compared average shape indices for males and females with a Student t-test for unpaired samples. Spatial analyses of activity centres were performed using software package TRACKER and RANGES IV, and core areas (50% minimum convex polygon; Lariviere and Messier 1998c) were consid- ered activity centers (Samuel et al. 1985). Results In 1993 and 1994, we radiotracked Striped Skunks during 1858 hours, and obtained home range informa- tion for 20 females and 5 males. Of the 20 females, 11 used only | maternity den during the entire period. First, we predicted that because of the fixed position of buildings in the landscapes, females using buildings as maternity dens would have home ranges less circular (larger shape index) than those using burrows. This pre- diction was not supported by our data: the shape index of females using buildings (mean = 1.53, SE = 0.08, n=6) did not differ (one-tailed t= 0.45, df =9, P=0.33) from the shape index of females using burrows as maternity dens (mean = 1.48, SE = 0.07, n =5). Second, we hypothesized that if females try to min- imize the travel distance to any point within their res- pective home ranges due to lactation constraints, then home range of females should be more circular (small- er shape index) than home ranges of males. Our pre- diction that home ranges of females are more circular than those of males was not supported: the average shape index for females (mean = 1.46, SE = 0.03, n = 20) did not differ (two-tailed t = 0.59, df = 23, P =0.56) from the average shape index of male home ranges (mean = 1.50, SE = 0.06, n = 5). Third, we hypothesized that because buildings are preferred denning sites (Lariviére and Messier 1998a), some buildings may be used as maternity dens but not be located within the core area of the home range. We predicted that underground burrows should occur in the core areas more often than dens in buildings. Our data also did not support this prediction: there were no associations between den type and whether or not the den was located inside the core area (n = 12 fe- males; Fisher exact test, P = 0.52). Overall, three underground burrows were inside versus two outside the core area, whereas six buildings were inside versus one outside the core area. In total, 75% of dens were inside core areas. 2007 In this study, we did not detect any effect of den type (anthropogenic or not) on shape of female home ranges or position of the den. Further, the shape of Striped Skunk home ranges was similar among males and fe- males. These findings suggest that female skunks estab- lish home ranges irrespective of den type and with rules similar to males. Examination of the natural history of Striped Skunks may help explain the patterns observed. First, Striped Skunks in northern environments undergo long periods of inactivity during winter, and to survive such period requires sufficient body reserves (Gunson and Bjorge 1979; Hwang et al. 2007; see also Rogers 1987). Thus, establishing a home range that includes good foraging areas may be more critical than choice of maternity den, but whether choice of maternity den influences juvenile survival remains unknown. In our study area, many buildings that were deemed suitable for occupa- tion by skunks were not utilized (Lariviere et al. 1999). As well, numerous females that used underground bur- rows to rear young had suitable unoccupied buildings within the boundaries of their home ranges. Thus, al- though buildings may be convenient denning sites, they may not be necessary or provide reproductive advan- tages to Striped Skunks. If juvenile mortality in skunks mostly occurs after emergence, then anthropogenic den types such as buildings may be used for other reasons such as thermoregulation, protection from the elements, or simply convenience. During the parturition/rearing period of Striped Skunks, females make nocturnal foraging trips but return to the maternity den by morning to nurse the young (Lariviére and Messier 1997). Foraging move- ments are highly variable (0-20 km per night; Lariv- iére et al. 1998b) and highly opportunistic, although mostly focused on insects and small mammals (Green- wood et al. 1999) and targeted to specific habitats (Bix- ler and Gittleman 2000; Lariviere and Messier 2000). Moreover, habitat availability around used and unused buildings in our study area was similar, suggesting that distribution of resources may not affect location of maternity dens, and conversely that choice of maternity den does not affect access to resources. Suitable den sites likely were not limiting for Striped Skunks (La- riviére et al. 1999). The preference of Striped Skunks for buildings may be most obvious where buildings are abundant, but absence of buildings as maternity dens probably does not affect distribution of skunks in the landscape during summer. Hence, our study reinforces the previous suggestion that food, instead of denning structures, probably affects distribution of Striped Skunks and their foraging activity in the land- scape. Acknowledgments This study was financed by Ducks Unlimited Cana- da (Institute for Wetland and Waterfowl Research), the Canadian Wildlife Service, and logistic support from Delta Waterfowl Foundation. SL is grateful to LARIVIERE, HOWERTER, and MESSIER: HOME RANGE SHAPE FOR SKUNKS 263 Fonds pour la Formation de Chercheurs et I’ Aide a la Recherche (FCAR), Québec, for postgraduate and postdoctoral support, as well as to the National Sci- ence and Engineering Research Council for an oper- ating research grant. Technical assistance was provid- ed by B. Dinter, E. Howard, G. Poon, L. R. Walton, B. Fry, and D. Anderson. W. A. Gorsuch reviewed an earlier draft of this manuscript. Methods used in this study were approved by the University of Saskatche- wan Animal Care Committee (Approval #920091). Literature Cited Bixler, A., and J. L. Gittleman. 2000. Variation in home range and use of habitat in the striped skunk (Mephitis mephitis). Journal of Zoology (London) 251: 525-533. Bowers, M. A. 1995. Use of space and habitats by the eastern chipmunk, Jamias striatus. Journal of Mammalogy 76: |2- Pile D’Eon, S. P. 1995. Accuracy and signal reception of a hand- held Global Positioning System (GPS) receiver. Forestry Chronicle 71: 192-196. Greenwood, R. J., A. B. Sargeant, D. H. Johnson, L. M. Cowardin, and T. L. Shaffer. 1995. Factors associated with duck nest success in the prairie pothole region of Canada. Wildlife Monographs 128: 1-57. Greenwood, R. J., A. B. Sargeant, J. L. Piehl, D. A. Buhl, and B. A. Hanson. 1999. Foods and foraging of prairie striped skunks during the avian nesting season. Wildlife Society Bulletin 27: 823-832. Gunson, J. R., and R. R. Bjorge. 1979. Winter denning of the striped skunk in Alberta. Canadian Field-Naturalist 93: 252-258. Hwang, Y. T., S. 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Forest Science Department, Ore- gon State University, Corvallis, Oregon, USA. THE CANADIAN FIELD-NATURALIST Vol. 121 Rogers, L. L. 1987. Effects of food supply and kinship on social behaviour, movements, and population growth of black bears in northeastern Minnesota. Wildlife Mono- graphs 97: 1-72. Rosatte, R., and S. Lariviere. 2003. Skunks. Pages 692- 707 in Wild mammals of North America: biology, man- agement, and economics. Edited by G. A. Feldhamer, B. Thompson, and J. A. Chapman. Johns Hopkins University Press, Baltimore. Samuel, M. D., D. J. Pierce, and E. O. Garton. 1985. Iden- tifying areas of concentrated use within the home range. Journal of Animal Ecology 54: 711-719. Received 18 April 2006 Accepted 7 July 2008 Road-kill of Mammals in Nova Scotia DAvID FupGE!*, BILL FREEDMAN!, MICHAEL CROWELL”, TONY NetrTe?, and VINCE POWER ' Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada ? Jacques-Whitford Environment Limited, 3 Spectacle Lake Road, Dartmouth, Nova Scotia B3B 1Q9 Canada 4 Nova Scotia Department of Natural Resources, Wildlife Branch, Kentville, Nova Scotia B4N 4E5 Canada Fudge, David, Bill Freedman, Michael Crowell, Tony Nette, and Vince Power. 2007. Road-kill of mammals in Nova Scotia Canadian Field-Naturalist 121(3): 265-273. We examined road mortality of wild mammals in Nova Scotia using data from (a) five years of province-wide data on wildlife collisions involving larger mammals, and (b) 20 months of observations of smaller mammals along a 160-km route near Halifax. An average of 2079 White-tailed Deer was reported killed annually on highways during 1999 through 2003, along with 14 Moose and 33 Black Bear. Female White-tailed Deer were more likely to be road-killed than males (by a fac- tor of 1.8), yearlings more so than older animals, and there were peaks of mortality in the late spring and late autumn. The road-kill of smaller mammals was highest in the summer and least in the winter, and the most frequent mortalities were of Raccoon (28% of observations), Porcupine (27%), Skunk (17%), Snowshoe Hare (6%), Coyote (4%), and Groundhog (3%). The standardized kill-rate (number/100 km) was highest on a 100-series highway compared to trunk highways or urban streets, but the rate standardized to vehicle density was highest on trunk highways. Key Words: White-tailed Deer, Odocoileus virginianus, Moose, Alces alces, Black Bear, Ursus americanus, Raccoon, Procyon lotor, Porcupine, Erethizon dorsatum, Skunk, Mephitus mephitis, Snowshoe Hare, Lepus americanus, Coyote, Canis latrans, Groundhog, Marmota monax, road-kill, mortality, wild mammals, Nova Scotia. Roads are a defining feature of modern transportation systems. Networks of public roads connect population centers, allowing the movement of people and goods in increasingly larger numbers of fast-moving vehicles. Because much highway infrastructure traverses wildlife habitat, an unfortunate consequence is accidental col- _ lisions that threaten the lives of both people and ani- mals, while also causing damage to property (Romin and Bissonette 1996). In Nova Scotia, for example, 503 collisions between vehicles and large wild mam- mals were reported in 2003, with 68 of the incidents resulting in personal injury requiring emergency serv- ices (NSDTPW 2006*). Data relating to the economic _ costs of such accidents have been quantified in Maine, where they result in US$32 million/year in property damage and medical costs (MEDOT 2001*). Overall in the United States, vehicle accidents involving only White-tailed Deer (Odocoileus virginianus) annually -tesult in about 200 human fatalities, 30 thousand injuries, and more than US$1 billion in property dam- age (Conover 1997). In Europe, collisions with ungu- | lates result in 300 human fatalities and US$1 billion in property damage annually (Bruinderink and Haze- broek 1996). Clearly, a large socio-economic cost is associated with collisions with large animals on roads. Roads also have important ecological impacts, in- cluding the following: (a) damage to populations of some species through mortality; (b) acting as barriers to movement or disrupting behaviour in other ways; (Cc) destroying and fragmenting natural habitat; (d) caus- » ing pollution by de-icing salt, metals, and pesticides; _and (e) enhancing access for hunters (Bennet 1991; Evink 1999). The direct mortality of larger animals due to vehicle collisions is relatively apparent and well- quantified (Clarke et al. 1998; Cain et al. 2003; Cle- venger et al. 2003; Saeki and MacDonald 2004). Road- kill may affect the conservation status of species — about 65% of the mortality of the endangered Florida Panther (Puma concolor coryi) is caused by vehicular collisions (Harris and Gallagher 1989). Other at-risk populations threatened by road-kill mortality include Ocelot (Leopardus pardalis) in the U.S. southwest (Tewes and Blanton 1998), the European Badger (Meles meles) in Great Britain (Clarke et al. 1998), and the endangered mainland Moose of Nova Scotia (Alces alces americana) (NSDNR 2008*). Many jurisdictions have implemented mitigations to reduce the frequency of collisions between vehicles and wildlife. Often, this is limited to warning signs in- stalled along highway sections known to have high collision rates (AMEC 2004*). In some other places, highways may have perimeter fencing or even wildlife under- and over-passes to keep animals off the roads, particularly in national parks and wilderness areas (Clevenger 1997; AMEC 2004*; NBDOT 2008*). Such mitigations help prevent animals from entering the right-of-way and are successful in reducing mortality, particularly of mammals (Foster and Humphrey 1995). Interestingly, the effectiveness of animal-crossing signs in reducing collision rates has not yet been quantified (MEDOT 2001*). Despite the fact that collisions with vehicles are an important source of mortality of wildlife, there has been little research into the phenomenon. Putman (1997) argued that such work is necessary, because understanding the biological and ecological influences 265 266 on road-kill rates is integral to the design and imple- mentation of effective mitigations to deal with this problem. Within this context, the great majority of research into road-kill has dealt with larger mammals. This focus is due to several socio-economic factors, including the danger to people associated with colli- sions with large mammals, their economic importance in hunting, and their high public profile (Cain et al. 2003; Clevenger et al. 2003). A few studies have also examined factors influencing road-kills of smaller mammals (Garland and Bradley 1984; Clevenger et al. 2003; Saeki and Macdonald 2004). In the present study the road-kill mortality of larger- and medium-sized mammals is examined for a variety of roads in Nova Scotia. Two data sets were used: (a) province-wide incident data for larger mammals over a five-year period, and (b) data for medium-sized mam- mals over a 20-month period along a 160-km work-day commute. We examine biological and environmental factors possibly influencing the road-kill rates, includ- ing gender, age, and proximity to distinctive habitats. Methods This study was done in Nova Scotia, an eastern province with a population of about 934 thousand peo- ple and 23 thousand km of public roads (NSDTPW 2008*). Data for larger mammals were obtained from incident reports of the Nova Scotia Department of Natural Resources (NSDNR), which records road-kill of White-tailed Deer, Moose (Alces alces), and Black Bear (Ursus americanus), including the location, date, species, sex, and age of the dead animals. Although the reporting of kills is not mandatory, NSDNR esti- mates that 60-90% of bear and deer collisions and all involving Moose are reported to them (see Conover et al. 1997). The original NSDNR data were taken from a FoxPro database and converted to Excel spread- sheets. The original location coordinates were con- verted to UTM using ArcView software to allow for analysis by a geographical information system (GIS). The number of road-kills for each species was summed monthly and annually from 1999 through 2003. The data for White-tailed Deer and Moose were also analyzed by gender and by yearling and multi- year animals. A GIS analysis was used to plot the distribution of road-kills on digital maps, and spatial relationships with road type and traffic volume were examined. In cases where the carcass was not found immediately beside a road, because the wounded ani- mal had moved before it died, the nearest road was inferred as the place of impact. The traffic volume of highway segments was identified for reaches between successive exits and interchanges, using archived data from NSDTPW (2008*). The 100-series highways of Nova Scotia have relatively high traffic volumes and are often four lanes with controlled access, while the trunk highways are relatively low-volume, two-laned, and have free access. The annual road-kill data were standardized on a per-km and per-10* vehicles basis, THE CANADIAN FIELD-NATURALIST Vol. 121 and were compared across highway types and on the basis of adjacent habitat (categories were: mature soft- wood forest (>60% conifer trees), mature hardwood (>60% angiosperm trees), mature mixedwood, imma- ture forest, agricultural, and urban land-use; a sepa- rate analysis involved surface water within 100 m. The relationship of road-kill density was also com- pared with indicators of local species abundance for White-tailed Deer and Moose. For deer, density was indicated by county-level Pellet Group Inventory (PGI) data (obtained by NSDNR in the spring, after snowmelt but before green-up, along 460 random | km x 2 m transects throughout the province). For Moose, the abundance polygons of Snaith (2001) were used. Data for smaller mammals were collected over a 20-month period during 1993 through 1994 along a 160-km weekday commuting route (by M. Crowell) from the rural village of Urbania to the urban Burn- side Industrial Park in Dartmouth. The route included 106 km of 100-series highway (part of Highway 102), 38 km of trunk highway (Highway 215), and 16 km of urban road. Animals observed dead beside the road were identified to species, as juvenile or adult, and the location was mapped. The annual road-kill data were standardized per km of road, per km? of road-bed, and per 10? vehicles of traffic density (using archived data from NSDTPW (2008*)). The total-kill data and that for Raccoon (Procyon lotor), Porcupine (Erethizon dorsatum), and Striped Skunk (Mephitis mephitis) were analyzed with respect to adjacent habitat along the surveyed reaches of 100-series highway, including: (a) roadbed being lower than, raised, or level with the adjacent terrain; (b) adjacent habitat being mature softwood forest, mature hardwood, mature mixed- wood, immature forest, agricultural, or urban land-use; and (c) surface water within 100 m. A coefficient of similarity was used to make pair-wise comparisons of the relative dominance of species among road-types: S, = 2C/(A+B), where C is the number of species killed on both road-types, while A and B are the num- bers on particular road-types. Observations and Discussion White-tailed Deer A total of 10 397 White-tailed Deer were reported killed in vehicle collisions from 1999 through 2003, or an annual average of 2 079. This mortality is equiv- alent to about 6% of the estimated deer population, and is a 28% increment on top of the reported hunt (on average, hunters take 20% of the Nova Scotia herd during the fall hunt; NSDNR 2007*).. The road-kill rate was similar during 1999 through 2001 (average + SD: 2 137 + 102), but greater than during 2002 and 2003 (1 449 + 44: P = 0.003 for difference between the groups of years). The smaller road-kill of 2002- 2003 is likely related to a reduced deer herd during that period (NSDNR 2007*). There was a bimodal pattern of peak mortality of multi-year individuals (of both sexes) during June- 2007 FUDGE, FREEDMAN, CROWELL, NETTE, AND POWER: ROAD-KILL OF MAMMALS IN NOVA SCOTIA 267 July and October-November (Figure 1a). Other authors have reported a fall peak of deer road-kill, and have attributed this to breeding activities (Puglisi et al. 1974; Case 1978). During the rut, bucks move about in search of receptive females, and does are also relatively vagile during that period (Banfield 1974). The fall hunting season may also increase road-kill rates (Puglisi et al. 1974; Allen and McCullough 1976), although Kilgo et al. (1998) found that deer in Florida generally avoid roads and other open areas when under hunting pres- sure. The increase of deer road-kill in June-July could be related to such factors as increased rural traffic asso- ciated with tourists and cottagers, the availability of high-quality forage and residual de-icing salt along verges, harassment by biting flies and partial refuge from them in open areas, and increased mobility of does with fawns during this time. Puglisi et al. (1974) commented on the attractiveness of early-greening road verges to deer in the springtime. We observed a par- ticularly large peak of road-kill in the spring of 2001 following an extremely harsh winter, during which about half of the provincial herd starved to death because of deep snow cover (NSDNR 2007*). Adult females were 1.8-times more likely to be killed in vehicle collisions than adult males. This is likely due to the Nova Scotian deer herd being disproportionate- ly female because of the selective removal of males by hunters (Feldhamer et al. 1986; Bruinderink and Haze- broek 1996). Yearling mortality was 11% of the total road-kill mortality, and the kill rate was similar to that of adults (when standardized by estimated population size), although there was little seasonality of the year- ling mortality. The road-kill rate was higher on two-lane secondary highways (0.38 kills/km.yr) than on four-lane 100-series highways (0.13/km.yr). Bashore et al. (1985) made a similar observation in Pennsylvania, where there were more deer road-kills along two-lane roads than along a larger interstate highway. Putman (1997) found that deer would routinely cross secondary roads within their territory, but would avoid larger interstate highways, likely because of their greater width and faster and more plentiful traffic. The road-kill rate per 10* vehicles was higher on secondary (0.092/10+ vehicles) than on 100-series high- ways (0.012/10*). Overall, however, we did not find a clear relationship between road-kill rates and traffic volume at most of the tested spatial scales, suggesting that more cars do not directly relate to more acci- dents. Carbaugh et al. (1975) also did not find a linear relationship between traffic density and deer kills, and Armstrong (2005*) reported a slight inverse relation- ship. Although we did not examine the phenomenon, it is likely that high traffic rates during crepuscular hours would be particularly lethal to deer trying to cross roads (Allen and McCullough 1976). We observed seven clusters of deer road-kills, each located near a human population centre with a high- density road network. A linear regression using average county data found a positive correlation between the estimated density of deer and that of road-kill (R* = 0.71; P<0.01). In some areas, road-kill may exert a controlling in- fluence on local deer populations and slow their recov- ery from high winter mortality. There are economic implications of such effects because White-tailed Deer are the most valuable wildlife resource in North Amer- ica in terms of hunting (Conover 1997; DuWors et al. 1999*). Romin and Bissonette (1996) estimated that the inclusive value of deer in North America is about $2 200 per animal (Canadian dollars adjusted for infla- tion to 2005). Moose Seventy Moose were reported killed in vehicle colli- sions from 1999 through 2003 (average = 14/yr). About two-thirds of the mortality involved the relatively abun- dant population on the highlands of Cape Breton, which is derived from 18 released individuals (Alces alces andersoni) imported from Alberta in the late 1940s (Pulsifier and Nette 1997). The annual road-kill mor- tality is equivalent to about 0.23% of the Cape Breton population, and 3.7% of the reported harvest by hunters (in 2003; this does not include the Mi’ kmaq harvest, which is estimated to be similar to or exceeding the licenced hunter harvest). However, the native mainland population of Moose (Alces alces americana) is much less abundant, likely fewer than a thousand individuals, and since 2003 it has been listed as endangered under the Nova Scotia Endangered Species Act. The annual road-kill mortality of mainland Moose is equivalent to about 0.5% of the population (there is no legal hunt- ing of this population). Moose road-kills occurred in all months except April, with a peak of male mortality in October, and of fe- males during June-July (Figure 1b). The autumn peak of male mortality occurs during the rut, when they are competing and moving about in search of receptive females (Banfield 1974). Males also become more aggressive during the rut and may be less averse to roads and traffic (Joyce and Mahoney 2001). The female peak of road-kill in early summer is related to the high volume of tourism-related traffic that occurs then on Cape Breton highways, particularly on the Cabot Trail. Similar numbers of males and females were killed on the mainland (10 vs. 12), but 1.6-times more fe- males were killed on Cape Breton roads. The latter observation is related to the female-skewed sex dis- tribution of Moose on Cape Breton (the gender ratio of kills by hunters is about three males per two fe- males). Yearling mortality was 11% of the total, but their kill rate was similar to that of adults (when stan- dardized by their estimated population size). There was no obvious seasonality of the yearling mortality. Of the road-kills on Cape Breton, 85% occurred along eastern sections of the Cabot Trail, where traf- fic volumes are relatively high. The highest kill rate (0.4/10 km. yr) occurred along a stretch from French 268 River Bridge to Wreck Cove along the Cabot Trail, within Cape Breton Highlands National Park. On mainland Nova Scotia, the highest road-kill rate was on Highway 104 on the upland Cobequid Pass Toll highway, where 38% of the mainland kills occurred (0.2/10 km.yr). These high-kill areas coincide with regions of Nova Scotia that have the highest density of Moose (Snaith 2001). Black Bear One hundred and sixty-four Black Bears were report- ed killed in vehicle collisions from 1999 through 2003 (average = 33/yr). The annual mortality is equivalent to about 0.5% of the estimated population of Black Bears in Nova Scotia, and 7.2% of the reported har- vest by hunters (in 2003). The lowest kill-rate was in 1999 (10 individuals) and the highest in 2003 (57). An unusually large number of 27 bears were road-killed in June 2003, compared to the 1999-2002 June average of 5. This may have been due to an increasing bear population, or the springtime of 2003 being unusually cool and wet, so that animals emerging from hiberna- tion had trouble finding adequate natural food and were attracted to areas near human settlement. There was a bimodal pattern of road-kill mortality, with peaks in May-June and in October-November (Figure Ic). The first peak is likely due to activity as- sociated with the breeding season, when males wander in search of females (Klenney 1987). The early sum- mer is also when sub-adult animals, at about 15 months of age, leave their mothers to find their own territory (Matson 1967). The fall peak of mortality is likely relat- ed to activity associated with pre-hibernation feeding and movement to denning sites (Matson 1967; Davis and Browne 1996). No road-kills were recorded from mid-November to early April, when bears are in hiber- nation. It was not possible to factor the influence of age or sex on bear road-kill, as these data were not con- sistently recorded. Bear road-kills occurred throughout the province, except in southeastern Cape Breton, where the species does not have an established population (Macmichael 2007). The 100-series highways had a higher linear mortality rate (1.58 kills/100 km.yr) than secondary highways (0.032/100 km.yr). According to traffic den- sity, however, the 100-series and secondary highways had similar mortality rates (both 0.02 kills/10° vehi- cles.year). Relative to their total estimated population size in Nova Scotia, Black Bear road-kill rates are lower than those of White-tailed Deer. This may be due to bears being relatively wary of humans and roads, except for habituated animals (Herrero 1985). In addition, deer tend to be relatively abundant in areas of Nova Scotia with high populations of peo- ple, especially in agricultural and coastal regions. Other Mammals Three hundred and ninety-two smaller mammals of sixteen species were recorded as road-kill on a 160-km THE CANADIAN FIELD-NATURALIST Vol. 121 study route during a 20-month survey (Figure Id). The most frequent mortalities were of Raccoon (28% of kills), Porcupine (27%), and Striped Skunk (18%). These are relatively abundant in the study area, often occurring near roads and other habitat edges, and are at inherent risk because they are relatively deliberate in their movements and have a body too large to pass beneath most moving vehicles without impact (Gar- land and Bradley 1984; Rolley and Lehmen 1992; Cle- venger et al. 2003). All species recorded as road-kill are indigenous to the study area, except for Coyote, Canis latrans, domestic cat, Felis domesticus, and domestic dog, Canis familarus (Banfield 1974; Davis and Browne 1996). Additionally, a Red-necked Wallaby (Macropus rufogriseus) that had been a 6-month escapee from a wildlife park was found by one of us as road-kill in the Annapolis Valley of Nova Scotia. Discoveries of small-mammal road-kills (rodents and shrews) were uncommon, but this is likely due to the difficulty of detecting these animals when scan- ning from a moving vehicle, plus the likelihood that they are quickly scavenged. In addition, small mam- mals are reluctant to cross multi-lane highways (Cle- venger et al. 2003). An innate reluctance to cross open spaces may also explain why Snowshoe Hare, Lepus americanus (the most abundant medium-sized mam- mal in the study area), was killed less frequently than Raccoon, Porcupine, or Striped Skunk, and why their kill rate was higher on the two-lane Highway 215 than on the four-lane Highway 102. Clevenger et al. (2003) also found that Snowshoe Hare was more likely to be killed on narrow roads than on wider ones in Banff National Park, Alberta. The road-kill rate, when standardized by distance surveyed, was highest on the four-lane Highway 102 (6.0 kills/km) than on the two-lane Highway 215 (4.0/km) or city streets (1.3/km). However, when stan- dardized to traffic density, Highway 215 had a greater rate of mortality (6.3 kills/10* vehicles) than Highway 102 (1.9/10* vehicles). The kill rate per 10* vehicles approached zero on urban streets because of the large traffic volumes, lower vehicle speeds, and likely a rel- atively small abundance of wild mammals. The lower rate on Highway 102 is likely due to its much greater width and larger volume of faster-moving vehicles, which likely discouraged animals from crossing. Cle- venger et al. (2003) also found that relatively wide and high-traffic roads discouraged small- and medium-sized mammals from crossing in Alberta, as did Clarke et al. (1998) for badgers in England. Similar numbers of species were observed killed on the four-lane Highway 102 (n=12) and on the second- ary Highway 215 (n=11), and both exceeded urban streets (only Red Squirrel, Zamiasciurus hudsonicus, were observed road-killed in the urban area studied). The most frequent mortalities on the secondary High- way 215 were Raccoon (19% of the total), Striped Skunk (18%), Red Squirrel (16%), Snowshoe Hare 2007 FUDGE, FREEDMAN, CROWELL, NETTE, AND POWER: ROAD-KILL OF MAMMALS IN Nova SCOTIA 269 (a) White-tailed Deer 14 — 7 —— = 12 —_ — 10 Ble i re M A S&S 6 Tot: 4 2 oO Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec (b) Moose 30 — 25 | 20 a 3 oF epee te) M 6 Tota S 10 a mE g _ Dai. ig we eo 1 zZ 3 t 5 6 P 8 9 10 11 12 (c) Black Bear 35 ; eee es £ 20 ee ey as EX =ans SS I CE ee a ae ae | ee a SP EN ee ee Ses eee ie) Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec (d) Smaller Mammals S = qy is) SS Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Ficure |. Average monthly road-kill mortality of large mammals from 1999 through 2003: (a) White-tailed Deer (m = 10 397): (b) Moose (n = 70); (c) Black Bear (n = 164); (d) smaller mammals (7m = 392). Sum of genders may not add to totals, because the sex of some animals was not identified. 270 (14%), and Porcupine (11%). The most frequent on the four-lane Highway 102 were Porcupine (31%), Raccoon (30%), and Striped Skunk (18%). The species spec- trum of mortality of Highways 102 and 215 had a similarity coefficient of 0.78, but their similarity with urban streets was very low — only 0.00 and 0.08, respec- tively. The greatest rate of road-kill occurred in the summer months of June (35 mortalities), July (34) and August (22) (Figure 1d). Smaller numbers were recorded in the winter months of November through February, when animals are hibernating or generally less active. The road-kill of Raccoons was highest in May and June, when adult males search widely for females with which to mate (Banfield 1974). In addition, 43% of the Rac- coon mortality in May and June was yearlings, which disperse during that time. The mortality of Striped Skunks was highest in September, when juveniles dis- perse from their natal home range. The mortality of Porcupines was relatively even throughout the spring, summer, and fall. Partial-deviance tests indicate that roadside topogra- phy and habitat were significant predictors of Raccoon road-kill. The likelihood of a Raccoon road-kill was much greater at level roadside topography (x? = 11.3, df = 3, P <0.001) and along segments of highway bisecting mature forest (y? = 9.4, df = 3, P <0.05). Proximity to water did not have a significant effect (x° = 0.50, df = 1, P >0.05). The likelihood of Rac- coon road-kill at different roadside topographies was assessed using log-likelihood ratios, and was much greater at level crossings (T-value = 3.1; P <0.05) and along segments bordering mature forest (T-value = 5.9; P <0.0025). For Porcupines, topography (x2 = 29.4, df = 4, P <0.005) and proximity to water (7 = 5.9, df = 1, P <0.025) were significant predictors of road- kill. Log-likelihood ratios indicate that road-kill was most frequent at level roadside topography (T-value 4.1; P <0.025) and adjacent to mature softwood forest (T-value 4.0; P <0.025). For Striped Skunks, topogra- phy (x? = 27.4, df = 2, P <0.005) was the only signifi- cant predictor of road-kill, and the log-likelihood of mortality was much higher at level roadside topogra- phy (T-value = 4.1; P <0.025). Overall, no road-kills of any species were observed at sites where the roadbed was raised (higher than the adjacent terrain), and the likelihood of road-kill was higher on roads bisecting mature forest (mixed, soft- wood, or hardwood) than adjacent to immature for- est, agricultural fields, or urban habitat (T-value 3.8; P <0.05). Proximity to water did not have an overall significant predictive effect. The results suggest that far fewer road-kills occurred on raised than on level or buried roadbeds. Raised roadbeds could serve as a visual barrier to habitat on the other side, and the banked incline may act as a physical deterrent to crossing. Clevenger et al. (2003) made similar observations for smaller-animal road- THE CANADIAN FIELD-NATURALIST Vol. 121 kills in Banff National Park, where raised roadbeds had a lower risk of mortality than other topographies. Another factor potentially affecting road-kill rates is the presence of vegetated center medians on divid- ed highways, such as the grassy ones occurring on Highway 102 in the present study. Bellis and Graves (1971) suggest that these habitats function as temporary refuge areas for animals attempting to cross major highways. Our study does not allow for an assessment of the impact of road-kill mortality on the local abundances of smaller mammals, as population estimates are not available. If our data were extrapolated to the total length of 100-series (3 x 10° km) and secondary (4 x 10° km) highways in Nova Scotia, the estimated annual mortality within this size range would be 20 x 10°? mammals. It can be concluded that the num- bers of animals killed on Nova Scotian highways is large, and that this is a significant anthropogenic stres- sor. Our field data for smaller mammals were collect- ed in 1993-1994, and since then traffic volumes on Nova Scotian roads have increased by about 24% since then (NSDTPW 2008*), so it is likely that road-kill rates have also increased. Moreover, some new roads are being built and the 100-series highways are being expanded, which may also be increasing the rates of road-kill. Mitigation Clearly, large numbers of mammals are being killed on Nova Scotian roads, and the further creation and upgrading of highways and increasing traffic may wors- en this situation. Moreover, there are important socio- economic consequences of collisions with wildlife. These include injuries to people involved in the colli- sions, the associated medical costs and time off work, damages to property, and decreases in the popula- tions of animals available to be hunted or trapped. For example, during our study period of 1999 through 2003, White-tailed Deer were involved in 204 vehicle collisions in which people suffered injuries requiring hospitalization, and seven incidents involved a human fatality (NSDTPW 2006%*). In fact, White-tailed Deer cause more collision-related prop- erty damage and human mortality than any other wild animal in North America (Conover 1997). According to Putman (1996), an average collision with a deer costs about $3 400 in vehicle damage (Canadi- an dollars, adjusted for inflation to 2005). Using this cost estimate, deer collisions in Nova Scotia during our study period resulted in about $6.8 million in annual property damage, not including costs associ- ated with health care and emergency services. Fortunately, there are ways to reduce road-kill along highways (Woods 1990*; Clevenger et al. 2001; USDOT 2002*; AMEC 2004*). Some mitigations are intended to change the behavior of animals in ways that make them less likely to enter the right-of-way. For example, fencing is used to prevent animals from 2007 FUDGE, FREEDMAN, CROWELL, NETTE, AND POWER: ROAD-KILL OF MAMMALS IN Nova ScoTIA 27] TABLE |. Road-kill of smaller mammals over a 20-month study period. Highway 101 i Species n Raccoon (Procyon lotor) 97 Porcupine (Erethizon dorsatum) 10] Striped Skunk (Mephitis mephitis) 60 Snowshoe Hare (Lepus americanus) 14 Domestic Cat (Felis domesticus) 17 Coyote (Canis latrans) 13 Groundhog (Marmota monax) 10 Red Squirrel (Jamiasciurus hudsonicus) 0 Domestic Dog (Canis familiaris) 4 Red Fox (Vulpes vulpes) 4 Eastern Chipmunk (7amias striatus) 0 Beaver (Castor canadensis) | Mink (Mustela vison) ] Muskrat (Ondatra zibethicus) | Red-backed Vole (Clethrionomys gapperi) 0 Woodland Jumping Mouse (Zapus hudsonicus) 0 Unknown mammal | Total Species = 16 324 crossing major roadways in at least eleven U.S. states (MEDOT 2001*), New Brunswick, and in the Cana- dian Rockies. In Banff National Park, for example, fencing resulted in a 96% reduction in large-mammal road-kill (Clevenger 1997). It is generally recommend- ed that fencing be at least 2.6 m in height, including above winter snow levels (USDOT 2002*). However, fencing also has drawbacks, because it can fragment populations and may trap animals within a right-of- way that they may enter via a break or at a fence-end (Phillips 1999*). Phillips (1999*) estimated that fenc- ing along a major highway in New Brunswick typi- cally costs $40 000 to $80 000 (Canadian) per kilome- ter. At present, the only areas in Nova Scotia with wildlife fencing along road verges are on the Cabot Trail on Cape Breton Island to prevent devastating col- lisions with Moose in segments where there are steep drop-offs along the highway, and an area near Maga- zine Hill in Dartmouth that had experienced a high number of accidents with deer. Although extremely expensive, structural crossings can be installed to allow for the movement of animals across a right-of-way. This mitigation can help pre- vent local and even regional populations of certain animals from becoming fragmented, with potentially deleterious consequences, while also avoiding risks to people and property from collisions. Crossing structures include underpasses, overpasses, and modified water- course structures (a kind of underpass). However, not all animals will use structural crossings. Reed (1981) observed that Mule Deer (Odocoileus hemionus) were reluctant to use enclosed underpasses, and they did not acclimatize to them over a study period of 10 years. He suggested that deer were far more willing to use open “bridge” style overpasses than underpasses. A review by USDOT (2002*) also concluded that ungu- Highway 215 Urban lotal c ‘ n c n n a 29.9 1 19.3 0 108 27.5 Ships 6 10.5 0 107 27.3 18.5 10 17.5 0 70 17.9 4.3 8 14.0 0 22 35 5.2 4 7.0 0) 21 5.4 4.0 9) 0.0 0 13 3.3 2 BiG 0 12 3.) 0.0 9 15.8 3 9 2.3 1.2 2 35 5) 6 ics 1.2 0 0.0 6) 4 1.0 0.0 3 5.3 (0) 3 0.8 0.3 0 0.0 0 ] 0.3 0.3 0 0.0 0 ] 0.3 0.3 0 0.0 0 l 0.3 0.0 | 1.8 0 l 0.3 0.0 | 1.8 0 | 0.3 0.3 0 0.0 0 l 0.3 57 3 392 lates were most likely to use overpass structures. The installation of a single overpass crossing structure can cost Canadian $1.4-2.6 million, while underpass- es and modified watercourse structures are generally $92 000 to $420 000 (MEDOT 2001*). Another useful mitigation is to make drivers more aware of potential dangers and to reduce vehicular speeds (AMEC 2004*). These driver-orientated ap- proaches include signage and educational programs. Passive warning signs (sometimes with flashing lights) of White-tailed Deer and Moose are used in many places in Nova Scotia and elsewhere to alert motorists to increased risks of collision with wildlife, generally in places where repeated collisions have occurred (AMEC 2004*). Although there are no data supporting the efficacy of signs, they likely have limited effective- ness because they rely on driver attentiveness, yet peo- ple quickly become desensitized to them (USDOT, 2002*). Passive signs are, however, relatively inexpen- sive to install and maintain. Driver education related to road-kill may make motorists more aware of the potential for wildlife en- counters, and therefore better prepared to avoid them. For example, a program called “Watch out for Wild- life” is delivered in the public-school system and by driver-education messaging in Florida (Evink 1999). The effectiveness of education programs in reducing road-kill is difficult to quantify, although people skilled at spotting animals and predicting their behaviour tend to be involved in fewer collisions (Joyce and Mahoney 2001). In Nova Scotia, enhanced public awareness of wild- life-collision risks could be promoted by an increased educational program. In addition, fencing verges in stretches of all major roads known to have high kill rates of larger mammals would reduce hazards, partic- 272 ularly for conservation of the at-risk mainland Moose population along the Cobequid Pass of Highway 104 (which also has high rates of bear and deer road-kills). However, roadside fencing could also detrimentally fragment the habitat of the already-small populations of mainland Moose in these areas. Of course, mitigations have significant costs, but they provide large benefits by reducing property damage and risks of injury to people and wildlife. Acknowledgments This work was supported by a Discovery Grant to BF from the Natural Sciences and Engineering Research Council of Canada. Lester Hanley of the Nova Scotia Department of Transportation provided traffic statis- tics for roads, Jennifer Milne and James Boxall of Dalhousie University assisted with the GIS analysis, and Cathy Wambolt with statistical analysis. Documents Cited (marked * in text) AMEC. 2004. Mainland Moose: Status, Potential Impacts, and Mitigation Considerations of Proposed Highway 113. Unpublished Research Report by AMEC Industrial to Nova Scotia Department of Transportation and Public Works. Armstrong, K. E. 2005. Deer-Related Automobile Acci- dents: Using GIS and GPS to Analyze Trends and Find Solutions. GIS Café, GIS Technical Papers. http://www. giscafe.com/technical_papers/Papers/paper005/. DuWors, E., M. Villeneuve, F. L. Filion, R. Reid, P. Bouchard, D. Legg, P. Boxall, T. Williamson, A. Bath, and S. Meiss. 1999. 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Pages 559-564 in Proceedings of the International Conference on Ecology and Transportation, Keystone, Colorado. Received 13 September 2006 Accepted 15 April 2008 Third Census of Seabird Populations of the Gaspé Peninsula, Québec, 2002 RICHARD COTTER and JEAN-FRANCOIS RAIL Canadian Wildlife Service, P. O. Box 10100, Ste-Foy, Québec G1V 4H5 Canada Cotter, Richard, and Jean-Francois Rail. 2007. Third census of seabird populations of the Gaspé Peninsula, Québec, 2002. Canadian Field-Naturalist 121(3): 274-286. In the tradition of the quinquennial census of seabirds in the Migratory Bird Sanctuaries along Québec’s North Shore, which began in 1925, this paper presents the results of the first three seabird censuses of the Gaspé Peninsula conducted in 1979, 1989, and 2002, with an emphasis on the third census and on changes in seabird populations that occurred between the sec- ond and third censuses. In 1979 the population of seabirds was estimated at 134 163 birds, and this increased 72% to 231 186 birds in 1989. The upward trend in population continued in the 1990s but less sharply. The 2002 census recorded 279 276 birds, a 21% increase over the second census. The seabird community of the Gaspé Peninsula comprises 14 species. In each of the three censuses, three species accounted for over 75% of the total seabird population: Northern Gannet, Black-legged Kittiwake, and Common Murre. Between 1989 and 2002, Common Eider, Northern Gannet, Great Cormorant, Ring-billed Gull, Razorbill, and Black Guillemot increased in numbers, whereas Herring Gull, Black-legged Kittiwake, and Common Tern all suffered declines. Populations for Double-crested Cormorant, Great Black-backed Gull, and Common Murre all remained stable. Very small populations (<25 birds) of Leach’s Storm-Petrel and Atlantic Puffin have been recorded during the censuses at only one location, Bonaventure Island. In the 2002 census there were no petrels observed, neither of adults nor of active burrows. A Vinstar des inventaires quinquennaux des oiseaux marins dans les refuges d’ oiseaux migrateurs de la Céte-Nord, entrepris en 1925, cet article présente les résultats des trois premiers inventaires complets des oiseaux de mer nicheurs en Gaspésie, menés: en 1979, 1989 et 2002. Une attention particuliére est mise sur le statut des espéces d’aprés le troisiéme inventaire, et sur les tendances des populations entre les deux derniers inventaires. En 1979, le nombre d’oiseaux marins nicheur était évalué a 134 163 oiseaux, puis ce total a augmenté de 72% pour atteindre 231 186 oiseaux en 1989. Cette tendance a la hausse a con- tinué dans les années 90, mais moins rapidement. En 2002 l’estimation était de 279 276 oiseaux, soit une hausse de 21 % par rapport au deuxieme inventaire. La communauté d’ oiseaux de mer de la Gaspésie est formée de 14 especes nicheuses. Dans chacun des trois inventaires, trois espéces représentaient plus de 75 % de la population totale des oiseaux marins: le Fou de Bassan, la Mouette tridactyle, et le Guillemot marmette. Entre 1989 et 2002, I’ Eider a duvet, le Fou de Bassan, le Grand Cor- moran, le Goéland a bec cerclé, le Petit Pingouin et le Guillemot a miroir ont vu leurs nombres croitre, tandis que le Goéland. argenté, la Mouette tridactyle et la Sterne pierregarin subissaient des déclins. Les populations de Cormoran a aigrettes, Goéland marin et Guillemot marmette sont demeurées stables. Les petites populations (<25 oiseaux) d’Océanite cul-blanc et de Macareux moine n’ont été vues durant ces inventaires qu’a l’ile Bonaventure. Cependant, aucun nid actif d’océanite ne fut localisé a cet endroit en 2002. Key Words: Seabirds, populations, larids, alcids, Gaspé Peninsula, Bonaventure Island, Forillon National Park, Québec. Seabird populations of the Gaspé Peninsula have long been well known due to two very popular sum- mer tourist destinations: Forillon National Park of Canada and the town of Percé. At Percé boat tours are offered around Rocher Percé and Bonaventure Island, which together form a provincial park as well as a fed- eral Migratory Bird Sanctuary. The most striking fea- tures of these sites are the high vertical cliffs holding large Black-legged Kittiwake colonies and the spectac- ular Northern Gannet colony on Bonaventure Island. (Scientific names are in Table 1.) With eight breeding seabird species numbering more than 100 000 pairs in 2002, Bonaventure Island is undoubtedly the most important seabird colony in the area; nevertheless, the smaller mainland cliffs, together with a few islands and sand bars distributed around the peninsula, are also home to numerous colonies of cormorants, gulls, guillemots and Razorbills. Chapdelaine and Brousseau (1992) published the first account the distribution and abundance of all 274 seabird species breeding along the entire Gaspé Penin- sula in eastern Québec. They presented the results of the first systematic censuses of seabirds for the region—in 1979 and 10 years later in 1989—as well as changes in’ populations and distribution for each species. After ani interval of 13 years, in 2002 the Canadian Wildlife Service conducted a third seabird census. The objec- tive of this paper is to present the results of all three censuses, but with an emphasis on describing thei changes in seabird populations and distribution that occurred between 1989 and 2002. This paper thereforet complements and updates Chapdelaine and Brousseau (1992) who discussed the changes that occurred: between 1979 and 1989. Methods On the Gaspé Peninsula, seabird species (Table 1) were censused at 59 breeding sites (Table 2, Figure 1) during June and early July in 1979, 1989, and 2002 by the Canadian Wildlife Service-Québec Region. There, 2007 COTTER AND RAIL: SEABIRD POPULATIONS OF GASPE PENINSULA 275 TABLE 1. Species names and summary of census methods used for each species in the 1979, 1989, and 2002 seabird census- es of the Gaspé Peninsula. English name Scientific name Common Eider Somateria mollissima Oceanodroma leucorhoa Morus bassanus Phalacrocorax auritus Leach’s Storm-Petrel Northern Gannet Double-crested Cormorant Phalacrocorax carbo Larus delawarensis Larus argentatus Great Cormorant Ring-billed Gull Herring Gull Great Black-backed Gull Larus marinus Black-legged Kittiwake Rissa tridactyla Sterna hirundo Uria aalge Common Tern Common Murre Razorbill Black Guillemot Alca torda Cepphus grylle Atlantic Puffin Fratercula arctica were three exceptions with respect to the year of the census, and these were: Black Guillemots at Clori- dorme to Marsoui were counted in 1980 instead of in 1979, Common Eiders at Pointe de Sandy Beach were counted in 2003 instead of in 2002, and Northern Gannets at Bonaventure Island were counted in 2004 instead of in 2002. The census techniques employed varied among species and colonies but were consis- tent for each colony among the three censuses (i.e., 1979, 1989, and 2002). A detailed explanation of the techniques used in 1979 and 1989 can be found in Chapdelaine and Brousseau (1992); with few excep- tions the same methods were used in 2002. A brief description of the techniques used in 2002 for each species is presented in Table 1. Certain species, such as Black Guillemots and Herring and Great Black- backed gulls, nest in colonies as well as solitary pairs or in very small groups. As such, for these species the distribution and colony size data may pertain to a sec- tion of shoreline rather than a discrete colony. At colonies where a species was believed to have been present but there are no population estimate available, a “+” is indicated in the corresponding table. To exam- ine changes in population sizes between the three cen- suses, that is from 1979 to 1989 and from 1989 to 2002, the compound annual growth rate (CAGR) was calculated for each species using the formula: CAGR = (((N(t)-N(0))/N(0)+1)!)-l where M(t) = population size at time t, N(0) = population size at time 0 and t = number of years between two censuses. Methodology systematic counts of all nests, except for llot Mahy nord where flushed females were counted systematic counts of active burrows population analysis from aerial photography nests counted from a boat, except for the Upper Plateau of Rocher Percé where nests were counted from a look out point using a spotting scope nests counted from a boat systematic counts of all nests systematic counts of nests; ratio calculated where nested with Great Black-backed Gulls; counts of incubating adults on cliffs made from a boat systematic counts of nests (sometimes from boat); ratio calculated where nested with Herring Gull Bonaventure Island, Cap d’Espoir, Forillon Peninsula: counts derived from photographs (taken from boat); elsewhere nests were counted from a boat systematic counts of all nests counts of birds at colonies on cliffs; used a conversion factor of 0.76 pair/bird observed counts of birds at colonies on cliffs counts of birds at colonies on cliffs and on the water; used Cairns (1979) conversion factor to estimate the number of breeding pairs counts of adult birds Results During the surveys in 1979, 1989, and 2002, a total of 14 species of seabirds was recorded at 59 breeding sites (colonies and/or shoreline sectors) around the Gaspé Peninsula (Tables 1, 2). Twelve species were observed in all three censuses, with the remaining two absent in one of the census years (Tables 3-8). In 2002, 13 species were observed, with only the Leach’s Storm- Petrel not recorded. In terms of total abundance (num- ber of individuals) and breeding sites, the censuses recorded 134 163 birds at 44 sites in 1979, 231 186 birds at 58 sites in 1989, and 279 276 birds at 57 sites in 2002 (Table 8). In each of the three survey years, three species accounted for over 75% of the total sea- bird population—Northern Gannet, Black-legged Kit- tiwake, and Common Murre. Individual species ac- counts are as follows: Common Eider: In 2001, the total population of breeding eiders (subspecies dresseri) in the St. Lawrence Estuary and Gulf was estimated at 42 000 pairs (The Joint Working Group on the Management of the Common Eider 2004). The 2002 seabird cen- sus of the Gaspé Peninsula recorded only 235 pairs of this eider, which represents less than 1% of the total population in Québec. Nevertheless, the popula- tion is steadily increasing, from 41 pairs in 1979 to 128 pairs in 1989 to 235 pairs in 2002 (Table 8). In 2002 this seabird bred at [lot Mahy nord, Ile Plate, and Pointe de Sandy Beach (Table 6). A few pairs are also believed to nest each year at the foot of the cliff on the Forillon Peninsula. 276 THE CANADIAN FIELD-NATURALIST Vol. 121 TABLE 2. Number, name, and centroid coordinates of sites on the Gaspé Peninsula surveyed for nesting seabirds in 1979, 1989, and 2002. (Colony numbers correspond to locations in Figure 1.) Miguasha to [lot Mahy Sud sector Colony Coordinates! 1. Miguasha to Saint-Omer 48°04'N, 66°18'W 2. Saint-Omer 48°07'N, 66°15'W 3. Banc de Carleton 48°06'N, 66°08'W 4. Ile au Pique-Nique 48°06'N, 66°07'W 5. Ile Taylor 48°09'N, 65°51'W 6. New Richmond to Bonaventure 48°09'N, 65°50'W 7. Pointe Howatson 48°08'N, 65°49'W 8. Caps Noirs 48°07'N, 65°48'W 9. Caplan 48°06'N, 65°39'W 48°0S'N, 65°37'W 48°04'N, 65°33'W 48°02'N, 65°29"'W 48°03'N, 65°28'W 10. Ruisseau Leblanc 11. Marais de Saint-Siméon 12. Bonaventure to Paspébiac 13. [lots de la riviére Bonaventure 14. Paspébiac Ouest 48°02'N, 65°17'W 15. Marais de Paspébiac 48°02'N, 65°15'W 16. Paspébiac to Shigawake 48°O1'N, 65°15'W 48°02'N, 65°13'W 48°03'N, 65°11"'W 48°03'N, 65°09'W 48°0S'N, 65°06'W 48°06'N, 65°04'W 48°06'N, 65°03'W 48°08'N, 64°59'W 48°11'N, 64°56'W 48°11'N, 64°53'W 48°11'N, 64°52'W 48°12'N, 64°49'W 48°15'N, 64°45'W 17. Pointe aux Corbeaux 18. Pointe Huntington 19. Pointe a Ritchie 20. Saint-Godefroi 21. Shigawake to Chandler 22. Shigawake 23. Colline de Port-Daniel 24. La Vieille (Pointe Pillar) 25. Gascons-Ouest 26. Pointe Reddish 27. Anse a Pierre Loiselle 28. Tlot Mahy sud llot Mahy Nord to Forillon sector Coordinates! 48°15'N, 64°45'W 48°20'N, 64°42'W 48°21'N, 64°41'W 48°21'N, 64°40'W 48°21'N, 64°39'W 48°23'N, 64°35'W 48°24'N, 64°29'W 48°25'N, 64°26'W 48°25'N, 64°25'W 48°25'N, 64°19"'W 48°26'N, 64°19'W 48°27'N, 64°19'W 48°29'N, 64°17'W 48°30'N, 64°13'W 48°30'N, 64°10'W 48°32'N, 64°12'W 48°32'N, 64°12'W 48°32'N, 64°13'W 48°34'N, 64°18'W 48°36'N, 64°18'W 48°38'N, 64°09'W 48°37'N, 64°10'W Colony 29. lot Mahy nord 30. Banc de la baie du Grand Pabos 31. Barachois de Chandler 32. Ile Dupuis 33. Chandler to Grande-Riviére 34. Barachois de Pabos 35. Grande-Riviere to Cap d’ Espoir 36. Grande-Riviere 37. Sainte-Thérese-de-Gaspé 38. Cap d’Espoir to Percé 39. Cap d’Espoir 40. Cap Malin 41. Weygand 42. Cap Blanc 43. Bonaventure Island 44. Rocher Percé 45. Percé to Cannes de Roches 46. Les Trois Soeurs 47. Coin-du-Banc to Pte St-Pierre 48. Barachois de Malbaie 49. Ile Plate 50. Pointe St-Pierre to Douglastown 51. La Grande Anse 48°41'N, 64°14"'W 52. Bois-Brilé 48°46'N, 64°22'W 53. Douglastown to Sandy Beach 48°46'N, 64°22'W 54. Cap Haldimand 48°48'N, 64°23'W 55. Pointe de Sandy Beach 48°50'N, 64°24'W 56. Pointe Jacques Cartier (Gaspé) 48°51'N, 64°29'W 57. Forillon Peninsula 48°47'N, 64°12'W 58. Cloridorme to Marsoui 49°11'N, 64°50'W 59. Marsoui to Baie des Capucins 49°13'N, 66°04'W ' For shoreline sections, the coordinates of the starting locations are given. Leach’s Storm-Petrel: On the Gaspé Peninsula, this petrel has only ever been known to nest at two locations, Forillon Peninsula and Bonaventure Island. By the 1930s, the Forillon colony no longer existed (Ball 1938), leaving only the Bonaventure Island colony. This colony was probably never large—in the 1979 and 1989 censuses only 10 and 21 pairs were counted, respectively. Unfortunately, no petrels (nor active burrows) were observed in the 2002 census (Table 7). In fact, this species’ breeding status in Québec is currently uncertain. Historically, it nested at several locations along the North Shore of the Gulf of St. Lawrence and on the Magdalen Islands. Howev- er, the only active colony found in the 1998-1999 census of North Shore Migratory Bird Sanctuaries (Corossol Island with 718 active burrows, Rail and Chapdelaine 2004) was deserted in the 2005 census (Rail and Cotter 2007), and no active burrow has been recorded on the Magdalen Islands in the last 20 years, although observations in 2007 of adults in the [le Brion Ecological Reserve during a survey for this species sug- gest a small colony still exists there (CWS, unpublished data). In some cases, including Bonaventure Island, the presence of Red Foxes (Vulpes vulpes) might be responsible of the abandonment of the colony. Northern Gannet: On the Gaspé Peninsula, this species nests only on Bonaventure Island. During the last census the Northern Gannet was the most abundant of all seabirds with an estimated 53 635 pairs (Table 8). Since 1979 this gannet colony has grown at a fairly consistent rate; the compound annual growth rate (CAGR) was 2.8% between 1979 and 1989 and 5.5% between 1989 and 2004 (Table 8). This colony was also censused in 1999 and was estimated at 36 936 pairs; the CAGR for 1999 to 2004 was 7.7%. In 1999 and 2004 the colony showed no sign of density-depen- dant effects, and because breeding success estimates obtained in 1999 and 2004 were high and stable (CWS, unpublished data), we think that the colony should con- tinue to increase in the next few years. Double-crested Cormorant: After a doubling in population size between 1979 and 1989, the size of this cormorant’s population on the Gaspé Peninsula did not change substantially between 1989 and 2002 (CAGR of only 0.4%), reaching 4 889 pairs in 2002 (Table 8). A stabilization in populations between the mid-1980s 2007 GULF OF ST. LAWRENCE GASPE PENINSULA > 47° 45'N 67° 00' W COTTER AND RAIL: SEABIRD POPULATIONS OF GASPI 77 Py NINSULA 49° 25'N _- 28-29 30- A 24.26 ~27 FIGURE 1. Location of colonies and shoreline sectors on the Gaspé Peninsula surveyed for breeding seabirds in 2002. See Table 2 for location listings and coordinates. and the early 2000s has also been recorded in Nova Scotia (Boyne and Beukens 2004) and New Brunswick (Boyne and Hudson 2002). In Québec, the population on the Magdalen Islands also remained fairly stable between 1990 and 2002 (CWS, unpublished data), as did the overall population in breeding colonies within the Migratory Bird Sanctuaries along the St. Lawrence North Shore between 1993 and 2005 (Rail and Cotter 2007). Interestingly, the increase in the size of the Gaspé population between 1979 and 1989 was accom- panied with an additional nine new colonies in 1989, bringing to a total of 29 active colonies that year. In 2002, however, there was a contraction in the number of sites where it nested, as it was recorded at only 25 sites (Tables 4, 6, 7). Of the nine new colonies in 1989, three were not active in 2002. Additionally, six colonies that were active in both 1979 and 1989 har- boured no cormorants in 2002. The loss of these nine colonies was offset by five new colonies, including one comprising 393 pairs in the New Richmond- Bonaventure region and, interestingly, a small colony on Bonaventure Island. As Chapdelaine and Brousseau (1992) stated, the majority of Double-crested Cor- morant colonies on the Gaspé Peninsula are found on sea-cliffs that in some areas are unstable and subject to erosion. Thus it is not surprising that between cen- suses with an interval of =10 years there are colonies that become inactive and new ones are created. Great Cormorant: World-wide this species has the largest breeding distribution of all cormorant species, but only a small fraction of its population nests in North America (Hatch et al. 2000). In Québec there are approximately 20 small colonies (total population <1500 pairs) (CWS, unpublished data), with histori- cally only one colony on the Gaspé Peninsula at Rocher Percé. In 1979 this colony had 16 nesting pairs, and in the 1989 census this number had doubled to 35 pairs. This upward trend continued in the 1990s as the total Gaspé population reached 96 pairs in 2002 (Table 7). In that census, the Rocher Percé colony had changed little in size since 1989, but a new and fairly large colony of 50 pairs was found at Cap d’Espoir (Table 7). Small numbers (1-3 pairs) were also observed at four other sites in 2002 (Tables 4, 6), which along with the new colony at Cap d’Espoir may indicate the population is expanding not only in size but also in its distribution. Rail and Chapdelaine (2004) also report- ed on the recent appearance of small Great Cormorant colonies in North Shore Migratory Bird Sanctuaries, although the most recent survey of these sanctuaries, in 2005, recorded only 48 Great Cormorants, a sharp decrease from the previous survey in 1998-1999 when 342 adults were counted (Rail and Cotter 2007). Ring-billed Gull: This gull, whose first breeding record in the Maritime Provinces was in 1965 (Lock 1988), is also a newcomer to the Gaspé Peninsula. It 278 was absent during the 1979 census, and in 1989 only 21 pairs were counted at two sites, Barachois de Chandler and Pointe de Sandy Beach (Table 5). In the 2002 census, however, there was only one colony and it was not at either of those two sites. Instead it was at Saint-Omer, and the colony was surprisingly quite large with 1663 pairs (Table 3). There were more Ring-bills at this site than the traditional species that nest there, that is Herring and Great Black-backed gulls and Common Terns. With numbers of Ring-billed Gulls still rising in the Maritimes (Boyne and Hudson 2002; Boyne et al. 2006) and apparently also on the North Shore (Roberge 2004), it is not surprising to observe the same trend around the Gaspé Peninsula. Herring Gull: Among all seabirds on the Gaspé Peninsula, the Herring Gull is the only species whose population declined in size between 1979 and 1989 and again between 1989 and 2002 (Table 8). From 1979 to 1989 its CAGR was -1.8%, and this rate of decline doubled to -3.5% between 1989 and 2002. Between 1979 and 2002, the total population declined from 9796 to 5134 pairs. This decline is attributable to colonies decreasing in size as opposed to a reduction in the number of breeding sites (24 in 1979 vs 29 in 2002; Table 8). In 2002, 22 of the 29 colonies had fewer gulls than in 1989. There were, however, four colonies/sectors that increased in size by more than 50% between 1989 and 2002: Saint-Omer, Ile Taylor, Marais de Paspébiac, and the New Richmond to Bona- venture sector (Tables 3, 5). These four areas along with [lot Mahy sud and Pointe de Sandy Beach com- prised 64% of the total Herring Gull population on the Gaspé Peninsula in 2002. The decline in colony size is probably not a result of competition, such as for nest or foraging areas, with other larids. Among the 22 sites where the population declined between 1989 and 2002, there were only two-Ilot Mahy sud and Ile Plate-that had a sizable increase in the popu- lation of another seabird, and at both sites it was of Great Black-backed Gulls (three other sites had small- er increases in their Great Black-backed Gull popula- tions; Tables 3, 5). Along coastal New England, how- ever, the more aggressive Great Black-backed Gulls have displaced Herring Gulls from certain good breed- ing sites, and Rome and Ellis (2004) found evidence there that Great Black-backed Gulls may suppress foraging by Herring Gulls. Elsewhere in Canada, pop- ulations of Herring Gulls have also declined in New- foundland (Robertson et al. 2004), New Brunswick (Boyne and Hudson 2002), Nova Scotia (Boyne and Beukens 2004), and Prince Edward Island (Boyne and McKnight 2005). Chapdelaine and Rail (1997) showed that there was a positive relationship between Herring Gull populations in the North Shore sanctu- aries and cod fishery activities in the Gulf of St. Lawrence, the latter providing substantial amounts of fish offal and discards to breeding gulls. After a thriv- ing period that lasted up to the end of the 1980s, the cod fishery collapsed, a situation which probably trig- THE CANADIAN FIELD-NATURALIST Vol. 121 gered the decline of the Herring Gull populations on the North Shore (-70% between 1988 and 1993; Chap- delaine 1995) and around the Gaspé Peninsula as well (-37% between 1989 and 2002). We think that the disappearance of this abundant anthropogenic food source brought Herring Gull populations back to a more “natural” level, which will stabilize in the future, as has been observed in North Shore sanctuaries since the early 1990s (Rail and Cotter 2007). Great Black-backed Gull: This large gull often nests in mixed colonies with its more abundant rela- tive, the Herring Gull. Unlike that species, however, Great Black-backs increased dramatically in num- bers on the Gaspé Peninsula between 1979 and 1989, with a CAGR of 23.7%. After reaching a population of 1 337 pairs in 1989, the population in 2002 was rel- atively unchanged with 1 322 pairs. The large increase from 1979 to 1989 was due in part to colonies increas- ing in size but also to a doubling in the number of colonies (Table 8). Comparing 2002 with 1989, there were a few changes in colony locations. In 2002 this gull did not nest at Barachois de Chandler but colo- nized the Banc de la baie du Grand Pabos in the adja- cent bay, less than 1 km away. It also deserted the cliffs from Paspébiac to Shigawake, but became established between New Richmond and Bonaventure. Elsewhere, in each of the neighbouring Maritime Provinces Great Black-backed Gull numbers declined between the late 1980s and early 2000s (Boyne and Beukens 2004; Boyne and Hudson 2002; Boyne and McKnight 2005). In the Migratory Bird Sanctuaries (excluding Gros Mécatina and Saint-Augustin) along the St. Lawrence North Shore, however, the population has been rela- tively stable, with | 883 individuals counted in 1988 and | 835 individuals counted in 2005 (Rail and Chapdelaine 2004; Rail and Cotter 2007). Black-legged Kittiwake: This small larid was the most abundant seabird species in both 1979 and 1989 (Table 8). The size of the kittiwake population on the Gaspé Peninsula doubled in size in the 1980s, from 21 827 pairs in 1979 to 43 423 pairs in 1989. This species’ distribution and abundance between 1974 and 1985 have been described in detail for the entire Gulf of St. Lawrence by Chapdelaine and Brousseau (1989). In the 1990s, the population stopped grow- ing, and in 2002 there were 13% fewer pairs (37 649) than there were in 1989. Recent downward trends (1.e., negative annual growth rates) have been observed in other regions of eastern Canada: Magdalen Islands (Québec), -1.0%/year, 1989-2000; Anticosti Island (Québec), -2.3%/year, 1985-2004; North Shore Migra- tory Bird Sanctuaries (Québec), -8.0%, 1988 to 1998- 1999 but followed by a stabilization with a rate of 0.64%, 1998-1999 to 2005; Gull Island (Newfound- land), -2.2%, 1971-2001; Great Island (Newfound- land), -9.9%, 1994-2003; and Baccalieu Island (New- foundland), -3.6%, 1984-2003 (CWS, unpublished data; Rail and Cotter 2007; Robertson et al. 2004; G. Robertson, personal communication). 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CV Ire LOI u- IL8 ly czIIS 8C 6S L L8e SOV = 8C7 9 Oc Iv LVS Gy 18 LOI 611 clc 696 GE tol 6L L8 EG LC TOOT 6861 6261 COOT JOUWSTIND Yul| ‘086] Ul posnsuan - ‘(UOTROTUNUIUIOD [eUOSJad ‘INo=poIg 9519S) ENOT UL pesnsuaD | 8 0 t 0 0 BLOT eecl 6cr SEC 8cl 9¢ [RIOL, suronded sap avg 0} Inosie] °6S INOSILIA, 0} DULIOPLIO[D “8G cle = 8S 1 (adse) soniej-sonbovr a1Ul0g “9¢ 1Sl € yorog Apurg op ajulod *¢¢ OV esl —- 861 pueunpreH dep “ps yorag Apurs 0} uMo}se[snog “€¢ ICI OIC Cl g1O1G-Slog “CS SL eC asuy opursH eT 1S I Iv UMOJSE[SNOG O} ILAIG-S ULOg “(IS Lec — CI Se OSI COI ce ld Al ‘6r aleq[ey op sloyseieg ‘Sp ILIIIg-1S Aq 0} UBG-NP-UIOD “Lp 9 SINIOG SIOLL, SAT “OP soyooy op souuRD 0} 919g “Sfp G Cl ouvir de ‘Tr bl Ol pursAaa “|p Z ure ded ‘Or 2019 0} Ilodsy ,p ded “ge (6 89 Syl gdsey-op-asaigy [-oules “L¢ OIQIATY-OPUuBIY) “OF todsq ,p ded 0} o1gIAry-apurly “Ce soged op sloyovieg ‘pe JIQIALY-OpuvIH O} JB]|purYD EE cIS 09 19] sindnq of] ‘Ze Jo[pueyO ap sloyoeieg ‘[¢ soqeg puvin np aieq ve] ap surg ‘O¢ 9EL EET OL x6 pou Aye IO[] “67 6861 6L61 COOT 6861 6L6I TOOT 6861 = 6L6l TOOT §=66861 ~—s OL [[Iquozey JURIOWIOD IIIH JURIOUIIOD po}so19-9]qnoq Joprq UOWIWOD AuojoD (‘Spiig [RNprAIPUT SI YOIYA [[Iglozey Joy }dooxa sated Ur UAAIS ore S9ZIs uone[ndodg) ‘ZOOT PUP “6861 “6L6I Ul P[NsUIUed adsey oy) uo sutondes sap aig pur piou Aye] IO[] Uaamjoq SUIPII1g SPlO[e PUL *SJULIOUTIOS *STOPlo JO 9Z1S uonejndod ut sasuvyo pur ‘uontsoduios sarads ‘uonnqinsiq °9 Atay, 2007 COTTER AND RAIL: SEABIRD POPULATIONS OF GASPE PENINSULA 283 three Gaspé Peninsula censuses, five colonies com- prised 295% of the total kittiwake population: Cap al onocoeonn > enn a S € peed a é 0 o d’ Espoir, Bonaventure Island, Les Trois Soeurs, Rocher E =| ve aA Percé, and Forillon Peninsula (Tables 5, 7). Compar- 3 2s) Rete ale a nak * ing colony sizes in 2002 with 1979, the former three o e 2 is rm i colonies had changed by less than +35%, whereas & aline ms too aa S +r the latter two had increased substantially in size, in par- Re Bel de Ean Gh mS ticular Rocher Percé, which increased from 16 pairs ro “ a a = in 1979 to | 094 pairs in 2002 (Table 7). In each of 2 the three censuses, the Bonaventure Island colony 2 harboured the most kittiwakes, with an estimated 18 cE” ||| ee aa 2 alent Sale whe Serr ee ee 550 pairs nesting in 2002. After the 1989 census, at AE a + = - 3 “= which time there were 23 650 pairs at this colony, o — ~ ay Chapdelaine and Brousseau (1992) postulated that bb ia alle a, ME A the Bonaventure Island colony may have reached its =| ZR oF as zl TR peak due to a lack of available nesting habitat. é S “ = Common Tern: Of all species censused in 1979, ‘@ ee es CcuMccve®) as a 1989, and 2002, the Common Tern had the greatest s ies a o z aS change in CAGR over the 1989 to 2002 period as 2 .. e compared with 1979 to 1989 (Table 8). In that first 5 interval, the population increased at an annual rate of S a Terr ws =ee 18.0%, from 459 pairs in 1979 to 2 407 in 1989. From s |3s aS 2 z SSS 1989 to 2002, however, the tern population declined S 2 7. at a CAGR of -13.1%. The 2002 population was esti- a | So A) Sere wey mated at only 388 pairs, which is fewer birds than Ei BG SC AS a Sue} there were in 1979 (Table 8); only one other species, ie) ae im the Leach’s Storm-Petrel, had a smaller population in 2 + D Beas ont 2002 than in 1979. Terns have never nested at a large = 3 Se number of sites on the Gaspé Peninsula. In 1979 the ef Common Tern nested only at Bane de Carleton and % Bs Pointe de Sandy Beach, and both of these colonies had z BQ wena & ; ON 0 Fe tn increased considerably in size by 1989 and were the ra aa Ro) | Sane largest of all tern colonies, with each containing over e : ve eas 800 pairs (Tables 3, 5). In addition to these two = 5 % Ss ci om SS g iB m S ; colonies, there were four new colonies in 1989, and 6 Sai a as among these six colonies four had 200 or more nest- 2 a EN) Sree sty SA aI) | NOES == ing pairs. In 2002, however, there were only four 3S ee e g ae 3 colonies, of which three consisted of 20 or fewer pairs. & oh Pr yee = During this census there were no terns at Banc de =