2009 NORTHEASTERN NATURALIST 16(1):131–140 Summer Distribution and Status of the Bats of Prince Edward Island, Canada Lynne E. Henderson1, Lesley J. Farrow1, and Hugh G. Broders1,* Abstract - The distribution and status of the bats of Prince Edward Island (PEI) is largely unknown. We addressed this information gap by compiling records from museums and published documents, and surveying for bats with traps and ultrasonic detectors during the summers of 2004 and 2005. Myotis lucifugus (Little Brown Bat) and M. septentrionalis (Northern Long-eared Bat) were the most abundant and widespread species of bats on the island. These species form maternity colonies, but the potential for there to be large natural hibernacula on the island is considered low. Lasiurus cinereus (Hoary Bat) is the only other species of bat that has been recorded on PEI, but its occurrence is rare. Diversity and abundance of bats on PEI may be limited by its northern insular geography, patchy distribution of forested areas, and lack of hibernacula. These limitations make protecting such resources on PEI and in neighboring provinces important for conserving the region’s bat fauna. Introduction Conservation problems facing bat populations have attracted considerable attention (Mickleburgh et al. 2002). Despite widespread efforts to document regional distribution and abundance patterns of bats, the distribution and status of the bats of Prince Edward Island, Canada (hereafter termed PEI) have not been studied. Seven bat species are known to occur in Atlantic Canada, including the long-distance migratory species Lasiurus cinereus Palisot de Beauvois (Hoary Bat), Lasiurus borealis Müller (Red Bat), and Lasionycteris noctivagans Le Conte (Silver-haired Bat) and presumed breeding populations of Myotis lucifugus Le Conte (Little Brown Bat), Myotis septentrionalis Trouessart (Northern Long-eared Bat), Eptesicus fuscus Palisot de Beauvois (Big Brown Bat), and Perimyotis subflavus F. Cuvier (Eastern Pipistrelle) (Broders et al. 2001, 2003; McAlpine 1976; van Zyll de Jong 1985). Of these, only the Little Brown and the Northern Long-eared Bat are widespread in the region, with breeding records available for Little Brown Bats, Northern Long-eared Bats, Red Bats, and Eastern Pipistrelles (Broders et al. 2003, 2006). Records exist for each long-distance migrant species from off-shore of Maritime Canada and the New England region of the United States, mostly during the fall migratory period (Broders et al. 2003, Maunder 1988, McAlpine et al. 2002a, Morris 1948, Tremblay 1992). Published records of bats on PEI exist for the Little Brown Bat, the Northern Long-eared Bat, and the Hoary Bat. Cameron (1952) reported three Little Brown Bats at Mount Herbert (Queens County), while Jones and Thomas (1983) reported five specimens from Ellerslie (Prince County), 22 from Cavendish (Queens County), and 11 from Cherry Valley (Queens County). Brown et 1Saint Mary’s University, Department of Biology, 923 Robie Street, Halifax, NS, Canada B3H 3C3. *Corresponding author - hugh.broders@smu.ca. 132 Northeastern Naturalist Vol. 16, No. 1 al. (2007) report 645 Little Brown Bats and 136 Northern Long-eared Bats from a hibernaculum located in Culloden (Queens County). They report a further two Northern Long-eared Bats from Breadalbane (Queens County). McAlpine et al. (2002b) reported the first Hoary Bat record for the province, collected from Charlottetown (Queens County) on 17 August 1999. Cameron (1958) reported seeing a large bat in flight near Fortune Bridge (Kings County) in June 1954, which he suggested was a Red Bat, but there is no way to verify this report. These records do not provide enough information to assess the distribution and conservation needs of any bat species on PEI. The purpose of this study was to provide a current assessment of the distribution and status of the bats occurring on PEI. We gathered museum records and conducted a province-wide survey during the summers of 2004 and 2005. These combined data represent the first assessment of the status of the bats of PEI. Field-site Description Prince Edward Island has an approximate area of 5660 km2, with two major urban centers at Charlottetown (Queens County) and Summerside (Kings County). Much of the original Acadian mixedwood forests on the island, characterized by both broadleaf deciduous and boreal coniferous tree species, have been removed or modified by agricultural activities (Loo and Ives 2003). The remaining forest is a mosaic of disturbed and remnant patches interspersed among agricultural areas (Sobey and Glen 2004). The geology of the island is part of the late Palaeozoic Maritimes Basin, consisting of flat to gently dipping conglomerates, sandstones, and siltstones that rise from sea level to a maximum height of 140 m inland (van de Poll 1995). PEI is in the Atlantic Maritime Ecozone and has warm summers (daily average of 19 ºC in July), and snowy, mild winters (daily average of -8.0 ºC in January). Mean annual daily temperature and precipitation are 5.5 °C and 110 cm, respectively (Environment Canada 2004). Methods We contacted curatorial staff or accessed online databases for the New Brunswick Museum (NBM), Nova Scotia Museum (NSM), Acadia University Wildlife Museum (AUWM), Canadian Museum of Nature (CMN), American Museum of Natural History (AMNH), Carnegie Museum of Natural History (CMNH), US National Museum of Natural History (NMNH), Museum of Comparative Zoology at Harvard (MCZH), Royal Ontario Museum (ROM; accessed through a MaNIS data portal http://manisnet.org on 20 February 2008), and Northeastern University Vertebrate Collection (NEU) to identify specimen records from PEI. Field surveys were conducted from 27 July to 25 August 2004, and from 9 June to 17 August 2005. In 2004, we sampled sites within the Greater Prince Edward Island National Park Ecosystem (263 km2), but expanded the study in 2005 to include additional sites across PEI. Each site was sampled twice to control for temporal variation in bat activity, while allowing us to survey as many sites as possible (1–3 sites per night). Bats were captured using free-standing 4.2-m2 two-bank harp 2009 L.E. Henderson, L.J. Farrow, and H.G. Broders 133 traps (Austbat Research Equipment, Lower Plenty, Victoria, Australia). We placed traps across forested trails or single-track old roads at locations where surrounding vegetation closed off potential bat flyways around the trap. This method of trapping was chosen to target the Northern Long-eared Bat, a forest-interior species (Broders et al. 2006; Caceres and Barclay 2000), as surveys were conducted as part of a separate study exploring the effects of forest fragmentation on the species (Henderson et al. 2008). In our trapping survey, 26 sites were sampled over 52 trap nights in 2004 and 62 sites over 124 trap nights in 2005 for totals of 88 sites and 176 trap nights. Herein, we define a trap night as deployment of one trap from sunset until five to eight hours after sunset. Traps were checked for captures once at the end of each trap night. However, on several occasions, we found only bat feces in the trap, indicating that some bats had been captured but escaped. Therefore, in retrospect, we do not recommend leaving traps unattended for extended periods of time due to missed captures. Captured bats were assessed for species, sex, reproductive status, and relative age (adult or juvenile; Anthony 1988) and released at the site of capture at least 30–60 min before sunrise. Methods of capturing and handling bats were approved by the Saint Mary’s Animal Care Committee and permitted by the Prince Edward Island Department of Energy, Environment and Forestry, and Parks Canada. Efforts to catch bats in 2004 were accompanied by passive acoustic surveys of bats using three Anabat II detectors (Titley Electronics Pty. Ltd., Ballina, NSW, Australia) interfaced to laptop computers via a zero-crossing analysis interface module (ZCAIM; Titley Electronics Pty. Ltd.), and calls were recorded using Anabat6 software (Version 4.9J, Titley Electronics Pty. Ltd.). During sampling, bat detectors were placed on the ground along forest edges or along the edges of lakes because species such as Hoary Bat and Little Brown Bat are more likely to be recorded in these areas than in the forest interior (Grindal and Brigham 1999; Krusic and Neefus 1996). Detector sites were ≤1000 m from harp-trap sites. In total, 33 sites were monitored over 60 detector nights. A detector night is defined as the passive recording of bat calls from dusk until dawn. Echolocation calls were qualitatively identified (O’Farrell et al. 1999) using Analook software (version 4.7J; Titley Electronic Pty. Ltd.) and published information regarding the calls of bats in eastern North America (Barclay 1986, Barclay et al. 1999, Betts 1998, Broders et al. 2001, Fenton and Bell 1981, Fenton et al. 1983, MacDonald et al. 1994). Call sequences were identified based on the minimum, maximum, and characteristic frequencies, as well as the slope of the calls (O’Farrell et al. 1999). Recordings that were identified as not containing any clearly identifiable bat calls were discarded. Echolocation calls from bats in the genus Myotis were not identified to species because identification is difficult due to the similarity of their echolocation call structure under certain conditions (Broders et al. 2004). Results Museum records for the Little Brown Bat, the Northern Long-eared Bat, and the Hoary Bat were found for PEI (Table 1). The majority of museum 134 Northeastern Naturalist Vol. 16, No. 1 records (66%) were for the Little Brown Bat, with records from all three counties. Thirty-one percent of records were for the Northern Long-eared Bat, with records from both Queens and Kings counties. Lastly, a single museum record for the Hoary Bat was available from Queens County. Forty-four Little Brown Bats and 12 Northern Long-eared Bats were captured in 2004 at seven sites each. In 2005, 60 Little Brown Bats and 45 Northern Long-eared Bats were captured at nine and 21 sites, respectively (Table 2). These data represent the addition of 37 new distributional records of bats on PEI, including 16 for the Little Brown Bat and 28 for the Northern Long-eared Bat (Fig. 1). Records of both species were widespread but patchily distributed on the island. Sex ratios of both species were not significantly different from 1:1 (binomial test: Little Brown Bats, P = 0.922; Northern Long-eared Bats, P = 0.570) during both years. No pregnant females were observed for either species. Twenty-five and 31 percent of adult female Little Brown Bats and Northern Long-eared Bats, respectively, displayed evidence of lactation (active or post-lactation), and all were caught between 04 July and 24 August. Overall capture success rate was 0.9 captures per trap night, with no bats captured on 133 of the 176 trap nights. Sixty-eight percent of Little Brown Bats captured in 2004 (13 adult females, 13 adult males, 4 juveniles) and 59% of Little Brown Bats captured in 2005 (8 adult females, 7 adult males, 22 juveniles) were caught at single sites over two nights of sampling (i.e., two trap nights at each site). Both of these sites were located in mature, hardwood-dominated Table 1. Records of bats on Prince Edward Island, Canada, as determined by published accounts and specimens occurring in museum collections. Species/date Individuals County MuseumA RemarksB Little Brown Bat 10 August 1922 2/3 Queens NMNH NMNH lists 2; Cameron (1952) lists 3 15 June 1938 5 Prince ROM Jones and Thomas (1983) 19 June 1958 22 Queens AUWM Jones and Thomas (1983) 12 August 1980 11 Queens NEU 7 M, 4 F; Jones and Thomas (1983) 05 July 1984 5 Queens CMN 4 M, 1 F 19 July 1984 1 Queens CMN F 31 July 1984 1 Prince CMN F 01 August 1984 3 Queens CMN 3 F 10 January 1989 1 Queens NBM M; Brown et al. (2007) Northern Long-eared Bat 24 September 1988 2 Queens CMN 1 M, 1 F; Brown et al. (2007) 10 January 1989 22 Kings NBM 21 F, 1 unlisted Hoary Bat 17 August 1999 1 Queens NBM McAlpine et al. (2002) ASee text for full names of museums. BF = female, M = male. Table 2. Age and gender of bats captured on Prince Edward Island, Canada, 2004 and 2005. Male Female Species Adult Juvenile Unknown Adult Juvenile Total Little Brown Bat 34 16 1 31 22 104 Northern Long-eared Bat 22 8 1 16 10 57 2009 L.E. Henderson, L.J. Farrow, and H.G. Broders 135 Figure 1. Distribution of Myotis lucifugus (Little Brown Bat), M. septentrionalis (Northern Long-eared Bat), and Lasiurus cinereus (Hoary Bat) on Prince Edward Island, as determined by museum records and survey efforts during 2004 and 2005. 136 Northeastern Naturalist Vol. 16, No. 1 stands with nearby large wetlands and farms containing old wooden barns. We did not attempt to search for maternity colonies in these barns. The largest sample of Northern Long-eared Bats occurred at one site in 2005, where four adult males and three adult females were captured. This site was located in a 385-ha provincially owned forest parcel dominated by hardwoods. A total of 6044 identifiable bat echolocation call sequences was recorded during acoustic sampling (mean of 101 call sequences recorded per detector night). Except for one call sequence, all remaining calls were attributed to Myotis species. The single non-Myotis call sequence was recorded on 18 August 2004. This sequence was recorded along the forested edge of a field in Prince Edward Island National Park (492840E, 5140435N, North American Datum 1983; Fig.1). The sequence included four echolocation calls with an average (±SD) maximum frequency of 19.9 ± 0.2 kHz, average minimum frequency of 18.5 ± 0.3 kHz, and average duration of 6.6 ± 1.6 ms. These call characteristics were consistent with those of a Hoary Bat (Barclay 1986, Barclay et al. 1999). Discussion Combined with museum records, data from this study suggest Little Brown Bats and Northern Long-eared Bats are widespread on PEI. In this respect, the bat fauna of PEI is similar to other areas of Atlantic Canada, where both species are widespread (Broders et al. 2003, 2006). However, prior to our survey, published records for Little Brown Bats were restricted to four locations for PEI. The current study adds an additional 16 capture localities. Little Brown Bats are widely distributed in the US (Fenton and Barclay 1980) and in Canada ranging from the Yukon through to Newfoundland (van Zyll de Jong 1985), being common in adjacent southern New Brunswick and Nova Scotia (Broders et al. 2003, 2006). The Little Brown Bat is a generalist species that roosts in buildings and trees and forages over water and along forests edges (Crampton and Barclay 1998, van Zyll de Jong 1985). Little Brown Bats are likely distributed across PEI where these resources are available. Their flexible foraging and roosting behaviour may be well suited for the agriculturally dominated landscape of PEI. Large numbers of juveniles and adult females captured at two sites sampled suggest maternity colonies are present on PEI. Two critical components for the persistence of bat populations are the availability of roosting and foraging resources, although the relative of importance of each may be uncertain for most species (Fenton 2003). Regardless of their status as limiting factors, maintaining suitable foraging and roosting resources is essential to the conservation of maternity colonies in order to maintain local summer populations. Regionally, Northern Long-eared Bats are widespread and common in New Brunswick and Nova Scotia, where they breed and hibernate (Broders et al. 2003, 2006). Previous published records for the species on PEI were from two locations where bats were found during the fall or winter. Our survey provided 28 new summer locality records across PEI and suggests that the Northern Long-eared Bat is widespread but patchily distributed on the island. Unlike Little Brown Bats, Northern Long-eared Bats are dependent 2009 L.E. Henderson, L.J. Farrow, and H.G. Broders 137 on forests for roosting and foraging (Broders et al. 2006; Caceres and Barclay 2000; Henderson and Broders 2008) and are likely restricted to forested areas on PEI. Forested areas are distributed across the island as remnant patches interspersed among agricultural areas (Sobey and Glen 2004). In the northern portions of the species’ range, adult females form maternity colonies in tree hollows or under exfoliating bark (Broders and Forbes 2004, Foster and Kurta 1999), where they care for young following a mid-July parturition period (Broders et al. 2006, Caceres and Barclay 2000). In 2005, lactating and postlactating females, as well as juveniles, were captured in late July and August suggesting similar reproductive phenology occurred on PEI. Similar to those of Little Brown Bats, maternity colonies of Northern Long-eared Bats are likely distributed on the island where suitable roosting and foraging resources are found together. However, their forest-dependent nature likely means that Northern Long-eared Bats are more restricted in their distribution, following the pattern of forest cover on the island (Henderson et al. 2008). Results of our acoustic survey suggested the presence of Hoary Bats on PEI. Although Hoary Bats are the most widespread of North American bats and are widely distributed in Canada, they are encountered infrequently in eastern Canada (van Zyll de Jong 1985), and only one specimen has been collected from PEI (McAlpine et al. 2002b). Broders et al. (2003) suggested that there are no resident populations or significant migratory movements of Hoary Bats through Nova Scotia. Our results suggest the same is true on PEI, where the species also appears to be a rare migrant. We did not record or capture any of the other species that have been occasionally documented in the region including Big Brown, Eastern Pipistrelle, Silver-haired, and Red Bats. This result was not surprising based on the apparent rarity of these species in the region (Maunder 1988, McAlpine et al. 2002a). However, our capture efforts likely were biased toward Myotis bats, and our acoustic sampling occurred mainly on the northern shore in the National Park rather than across the entire island. Big Brown Bats have been found in Nova Scotia and New Brunswick (McAlpine et al. 2002a, Taylor 1997), and because they have been associated with human dominated environments for roosting, foraging, and hibernating (Brigham 1991, Furlonger et al. 1987, Hamilton and Barclay 1994, Whitaker and Gummer 2000), it is possible that further work in more human-populated areas could reveal their presence on the island. Although it is possible that we missed capturing or recording the remaining larger species on the island, it is unlikely that our inferences regarding the status of these species on the island would change significantly based on our understanding of the distribution and status of these species in the adjacent mainland provinces. Our work indicates that Little Brown Bats and Northern Long-eared Bats constitute most of the bat community on PEI. Captures of adults and juveniles of these species confirmed the presence of maternity colonies on PEI, and a recent study confirms the presence of Northern Long-eared Bat maternity colonies on the island in the summer (Henderson and Broders 2008). However, only a single hibernaculum (in a basement) has been recorded on PEI, and the population has been exterminated (Brown et al. 2007). The geology of the 138 Northeastern Naturalist Vol. 16, No. 1 island does not include karst features (van de Poll 1995); therefore, suitable natural hibernacula may be rare on PEI. Bats can move significant distances to hibernacula (Davis and Hitchcock 1965, Fenton 1969); thus, bats spending the summer on PEI may use hibernacula in other provinces, such as those in adjacent New Brunswick and Nova Scotia (McAlpine 1983, Moseley 2007). If so, protecting such resources in neighboring provinces will be important for conserving the bats of PEI. Managing summer colonies on PEI also may require conservation of existing roosting and foraging sites. For Little Brown Bats, it will be important to identify and protect buildings that contain colonies, particularly since an increasing number of old barns are being destroyed, as well as wetland and forested foraging areas. To manage for Northern Long-eared Bats, maintaining mature forest stands that provide roosting and foraging opportunities will be essential for ensuring viable populations of the species in the agriculturally dominated landscape of PEI. Acknowledgments We thank the land owners on PEI who allowed us to conduct surveys on their property and Parks Canada and the PEI Department of Tourism for providing accommodations. 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