Northeastern Naturalist 32 L.E. Burns, J.L. Segers, and H.G. Broders 22001155 NORTHEASTERN NATURALIST V2o2l.( 12)2:,3 N2–o4. 01 Bat Activity and Community Composition in the Northern Boreal Forest of South-central Labrador, Canada Lynne E. Burns1,2, Jordi L. Segers2, and Hugh G. Broders2,* Abstract - Little is known about composition of bat communities and their acoustic activity in the northern boreal forests. Acoustic and capture surveys were conducted in the Mecatina River Ecoregion (MRE) in south central Labrador. We acoustically surveyed forest edges, streams, and ponds and netted and trapped bats near streams and along forest edges. The acoustic survey showed greater bat activity at streams and ponds. The capture survey confirmed the presence of Myotis lucifugus (Little Brown Bat) and M. septentrionalis (Northern Long-eared Bat) in the MRE, the latter showing signs of reproduction. Together these results support the idea that riparian areas in the boreal forest are important landscape features for bats in the genus Myotis. Introduction Knowledge of the distribution and biology of bats in the boreal regions of North America is quite limited and derived primarily from work in the northern boreal forest of Alaska and Western Canada (Kalcounis et al. 1999, Olson and Barclay 2013, Parker et al. 1997, Randall et al. 2011). In the northeastern boreal forest of mainland Canada, information on roost requirements and foraging ecology is lacking, and predicted distributions have been extrapolated from sparse location records (Mills et al. 2013, Naughton 2012, van Zyll de Jong 1985). Only limited surveys for bat species diversity and abundance have been conducted on the boreal regions of the Quebec–Labrador Peninsula (Broders et al. 2013). The only species confirmed to occur there are Myotis lucifugus Le Conte (Little Brown Bat) and M. septentrionalis Trouessart (Northern Long-eared Bat) (Broders et al. 2013, Naughton 2012, van Zyll de Jong 1985). However, previous reports have been based on opportunistic records or were focused on known bat maternity roosts rather than characterizing bat habitat or quantifying the magnitude of activity within the forests of Labrador. The Little Brown Bat is an aerial-hawking bat with a wide distribution extending through forest and prairie landscapes from central Alaska, south through most of the United States, into Mexico, and east into central Labrador and Newfoundland. The range of the Northern Long-eared Bat, a forest-dwelling gleaning bat, extends from northeastern British Colombia to central and eastern United States, southern Quebec, Nova Scotia, and the island of Newfoundland (Mills et al. 2013, Naughton 2012, van Zyll de Jong 1985). The documented range of Lasiurus cinereus Palisot 1Department of Biology, Dalhousie University, 6299 South Street, Halifax, NS, B3H 3J5 Canada. 2Department of Biology, Saint Mary’s University, 923 Robie Street, Halifax, NS, B3H 3C3 Canada. *Corresponding author - hugh.broders@smu.ca. Manuscript Editor: Jacques Veilleux Northeastern Naturalist Vol. 22, No. 1 L.E. Burns, J.L. Segers, and H.G. Broders 2015 33 de Beauvois (Hoary Bat) extends into the boreal forest of northern Ontario and Quebec to approximately the Saguenay River (Naughton 2012, van Zyll de Jong 1985). However, records are known from the islands of Newfoundland, Prince Edward Island, Southampton Island in Nunavut, and Iceland, which suggests they could occur in Labrador (Broders et al. 2003, Hayman 1959, Henderson et al. 2009, Hitchcock 1943, Maunder 1988). At this time, studies are needed to provide baseline data on bat distribution and resource use. Then, inferences on bat biology in the boreal ecosystem may be made, and the baseline data can be used as a metric of ecosystem change. Climate change is considered one of the most important threats to biodiversity, and one major effect is shifts in the distribution of species (Parmesan et al. 1999, Thomas et al. 2006). The greatest impact of climate change may occur at the most northern extent of a species’ range (Virkkala et al. 2008). Bioenergetic models for hibernating mammals have led some researchers to predict that climate change may shift the current range distribution of some bats, including Little Brown Bat and Northern Long-eared Bat, further north (Humphries et al. 2004, Naughton 2012). Another cause of environmental change is anthropogenic fragmentation of natural areas. Animals using such areas respond differently depending on their habitat requirements and ability to adapt (Loehle and Li 1996, Marvier et al. 2004, Ricklefs 2001, Webala et al. 2011). Bats are no exception to this general principle, and those with more specialized life histories (i.e., forest specialization) are more vulnerable to forest fragmentation (Lane et al. 2006, Russ and Montgomery 2002). In North America, the Little Brown Bat and Northern Long-eared Bat occur in sympatry but occupy different niches (Broders and Forbes 2004, Owen et al. 2002, Ratcliffe and Dawson 2002) and therefore may be impacted differently by fragmentation of landscapes (Segers and Broders 2014, Webala et al. 2011). The Little Brown Bat is more of a generalist, using a wide variety of landscape elements relative to the more forest-dependent Northern Long-eared Bat (Henderson and Broders 2008, Jung et al. 2004, Sasse and Pekins 1996). Increasing extraction of resources (e.g., mining) and the construction of large hydro-electric power projects in Labrador (Newfoundland Department of Natural Resources 2012) have the potential to alter large tracts of forest, which may impact bats. In addition to these threats, bats in eastern Canada are in serious decline due to the psychrophilic fungus Pseudogymnoascus destructans (formerly known as Geomyces destructans) (Blehert & Gargas) Minnis & D.L. Lindner, which causes white-nose syndrome and major mortality amongst hibernating bats (Blehert et al. 2009, Minnis and Lindner 2013). The aim of this study was to investigate the composition of the bat community and their acoustic activity in the northern boreal forest of south-central Labrador. Specifically, our objectives were to quantify the magnitude and distribution of bat activity in the boreal forest of the Labrador Peninsula by (1) quantifying differences in foraging activity between habitat types through analyses of feeding buzzes and (2) capturing bats to identify community composition and make inferences about their reproductive biology. Northeastern Naturalist 34 L.E. Burns, J.L. Segers, and H.G. Broders 2015 Vol. 22, No. 1 Methods The study was conducted from 20 June to 02 July 2013, in the Mecatina River Ecoregion (MRE) of the Minipi Ecodistrict (52°17'51.99"N, 60°59'16.03"W) of southern Labrador (Notzl et al. 2013). This area is characterized by cool summers and cold, snowy winters. The dominant vegetation there is a closed-canopy forest of Picea mariana (Mill.) Britton, Sterns & Poggenb. (Black Spruce), Picea glauca (Moench) Voss (White Spruce), and Abies balsamea (L.) Mill. (Balsam Fir). Wildfires are common in the region and, as a result, many areas are in some stage of transition from lichen-covered ground to closed-canopy forest resulting in sprucedominated forest over much of the landscape (Roberts et al. 2006, Wiken 1986). We captured bats using mist nets and a harp trap set during the first 3 h after sunset. There were 8 capture locations (Fig. 1), which included forest edges, streams, trails, and natural canopy gaps in the forest interior. Distance between capture sites averaged 2227 m (range = 313–4410 m). Nets and traps were checked every 10 minutes. We identified to species all captured bats and recorded their sex, weight, forearm length, and wing-index scores (Reichard and Kunz 2009). For females, we determined reproductive condition (pregnant, lactating, non-reproductive) by palpating the abdomen or by noting signs of lactation (the expression of milk; Racey and Swift 1985). All capture and handling equipment had either only been used in Labrador or was new to minimize the risk of exposing captured bats to white-nose Figure 1. Study site within the Mecatina River Ecoregion (MRE), south-central Labrador, to assess bat species composition and activity. In-map ratios (M. septentrionalis : M. lucifugus) represent the number of bats captured in 2013 for the three sites where at least one bat was captured. (Service layer credits: © 2013 ESRI, DeLorme, NAVTEQ). Northeastern Naturalist Vol. 22, No. 1 L.E. Burns, J.L. Segers, and H.G. Broders 2015 35 syndrome. We released all bats at the site of capture following processing. Methods for the capture and handling of bats were approved by the Saint Mary’s Animal Care Committee and under permit from the Wildlife Division, Department of Environment of Newfoundland and Labrador. We conducted acoustic surveys for bat activity using 4 automated bat detectors (model Song Meter SM2Bat+, Wildlife Acoustics, Maynard, MA) at 16 sites within the MRE (Fig. 1). Detectors were deployed at 8 forest edges, along 5 streams, and at the edge of 3 ponds and were left to passively sample for bats over 2–3 nights. We oriented microphones towards a water feature or perpendicular to forest edges and placed them at least 1 m off the ground and slightly less than parallel to the ground to shield them from rain. All recorded bat call sequences were converted to zero-crossing file formats using KaleidoscopeTM software (Wildlife Acoustics) and were imported into AnalookW software (Titley Scientific, version 3.8) for identification and analysis. We identified calls based on the minimum, maximum, and characteristic frequencies and the slope of calls (O’Farrell et al. 1999). Calls from bats in the genus Myotis were not identified to species since they have highly overlapping call structure that makes reliable species identification difficult (Broders et al. 2004). We defined a bat call sequence as containing at least two discrete echolocation pulses, and a detector night as the passive recording of bat calls from 20:00 to 07:00. We also quantified the number of call sequences containing feeding buzzes (Griffin et al. 1960) as a relative measure of feeding activity in each habitat. We tested for differences in the number of bat call sequences recorded among site types (streams, ponds, forest edges) using a Kruskal-Wallis test in R (R Development Core Team 2013) because the data were not normally distributed. Results and Discussion Over 11 nights of sampling, 7 bats were captured at 3 of 8 capture locations in 143.5 mist-net hours (6 m of net set for 1 hour = 1 mist-net hour) and 22.4 harptrap hours. Captures included 1 male and 1 female Little Brown Bat and 2 male and 3 female Northern Long-eared Bats. Both Little Brown Bats were captured at the same site on 2 separate nights and the Northern Long-eared Bats were captured at 3 sites on separate nights. The 3 female Northern Long-eared Bats were captured at the same site within 5 minutes of each other and approximately 90 minutes after sunset. The spatio-temporal clustering of these captured females may represent individuals commuting from an unidentified maternity roost. One of these females was obviously pregnant, and the others were either not pregnant or in such an early stage of pregnancy that we were unable to confirm. Together, these captures represent the first record of females and the first record of reproduction for Northern Long-eared Bat in Labrador (Broders et al. 2013). In comparison to our study in Labrador, Broders et al. (2003) caught 46 bats during 52.2 mist-net hours and 75.8 harp-trap hours in southwest Nova Scotia. In New Brunswick in the Greater Fundy Ecosystem, Broders et al. (2006) caught 277 bats during 106 mist-net hours and 363 harp-trap hours over 3 years of sampling. Factors to explain the low capture success in this current study may include low Northeastern Naturalist 36 L.E. Burns, J.L. Segers, and H.G. Broders 2015 Vol. 22, No. 1 nightly temperatures approaching 0 °C in south-central Labrador through our trapping period in June, the open-forest characteristics of the boreal forest with fewer forest edges and corridors to capture bats along, or lower numbers in the area and fewer bats in the area. At all sites where bats were captured, stands of mature spruce-dominated forest with dead and dying trees were present. We speculate that these mature trees and those in higher-decay classes could be used as roost sites for bats in the area. The Northern Long-eared Bat is a forest-associated bat which typically roosts in cavities of tall, large-diameter trees in forests that have an open canopy and high density of snags (Barclay and Kurta 2007, Kalcounis-Rüppell et al. 2005). Although Northern Long-eared Bats often roost in deciduous trees (Broders and Forbes 2004, Henderson and Broders 2008, Sasse and Pekins 1996), they also roost in conifers (Garroway and Broders 2008, Jung et al. 2004, Park and Broders 2012), which were the dominant tree type present within the MRE. Little Brown Bats tend to be more flexible in their roosting habits and use both buildings and trees (coniferous and deciduous; Fenton and Barclay 1980, Olson and Barclay 2013). Selection of roost sites by bats reflects local availability of trees and their associated microclimates (Boyles 2007, Broders and Forbes 2004). Thus, bats in Labrador, likely use locally abundant mature stands of coniferous trees, or small patches of deciduous stands that comprise the northern boreal forest. We recorded 1023 identifiable bat call sequences at 15 of 16 bat detector locations over 44 detector nights, 16.5–31% of which were feeding buzzes (Table 1). The mean number of call sequences per detector night was 24 (range = 0–194), and all sequences were attributed to Myotis species, which suggests the bat community in summer is composed of Myotis species (Little Brown Bat and Northern Longeared Bat). The high proportion of sites with bat call sequences recorded (94%) suggests that Myotis occur throughout the MRE, although they may be patchily associated with specific landscape features. The number of call sequences recorded differed significantly across the 3 habitat types, with greater acoustic activity recorded at sites associated with water (stream or pond edges) relative to forest edges (H = 10.82, df = 2, P < 0.01; Table 1). The site where no acoustic activity was recorded was along a forest edge within 100 m of water. In comparison, Segers and Broders (2014) also recorded on average more Myotis call sequences per night at water-associated sites in Nova Scotia (pond edges: 112, streams: 115, forest edges: 90) although between-habitat differences were not statistically tested. Call sequences recorded on forest edges in Labrador may primarily represent commuting calls as bats move from day-roosts to foraging locations, since fewer feeding Table 1. Number of Myotis call sequences and percentage of call sequences containing feeding buzzes recorded by site type and sampling effort in the MRE in Labrador, 20 June to 2 July 2013. Number of Number of call sequences % of sequences Site type Number of sites detector nights per night: mean (range) with feeding buzzes Pond edge 3 8 54 (1–180) 31.0 Stream edge 5 14 39 (0–194) 29.3 Forest edge 8 22 4 (0–24) 16.5 Northeastern Naturalist Vol. 22, No. 1 L.E. Burns, J.L. Segers, and H.G. Broders 2015 37 buzzes were recorded at forest edges (16.5%) compared to pond edges (31%) and stream edges (29.3%) (Table 1). Some bats, including Northern Long-eared Bats, may opportunistically forage along forest edges or in the interior of forests, although forest edges associated with water (i.e., stream or pond edges) tend to show higher bat activity (Grindal et al. 1999, Henderson and Broders 2008, Krusic and Neefus 1996). Riparian areas are important for many species of bats because they typically have greater concentrations of prey, provide drinking areas, and can act as unobstructed commuting corridors (Downs and Racey 2006, Racey and Swift 1985). The recording of feeding buzzes at several sites in the MRE demonstrates foraging activity by bats, including along forest edges. Understanding bat species composition and patterns of activity in the boreal forest of Labrador may be important for management for these species. The results from both the acoustic and capture survey in this study indicate the importance of riparian areas (e.g., streams and ponds) for bats in south central Labrador. In addition, this study shows the presence of at least 2 species of bat (Northern Long-eared Bat and Little Brown Bat) in the MRE. Additional surveys could provide a higher resolution of habitat-specific bat activity and species composition and may show variation between seasons and years. Bats have many life-history traits such as small size, high mobility, and long life spans with low reproductive potential that facilitate stable populations over time (Fenton 1997, Findley 1993). However, these same traits may make them sensitive to environmental changes whereby they may be used as bioindicators of ecosystem change (Jones et al. 2009). 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