Southeastern Naturalist V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 140 2017 SOUTHEASTERN NATURALIST 16(2):140–148 Baseline Capture Rates and Roosting Habits of Myotis septentrionalis (Northern Long-eared Bat) Prior to White-nose Syndrome Detection in the Southern Appalachians Vanessa G. Rojas1,*, Joy M. O’Keefe1, and Susan C. Loeb2 Abstract - Myotis septentrionalis (Northern Long-eared Bat) is a federally threatened insectivorous bat facing devastating population declines due to white-nose syndrome (WNS). Our study provides pre-WNS (2009) capture rates and roosting-behavior data for Northern Long-eared Bats in the southern Appalachians. We conducted mist-net surveys at 37 sites and radio-tracked female Northern Long-eared Bats to their day roosts in eastern Tennessee and western North Carolina. We compared tree and plot characteristics for roosts and corresponding random trees using Wilcoxon rank-sum tests. Our 43 survey nights yielded 302 bats of 11 species; Northern Long-eared Bats were the most commonly captured species (n = 97). We located 14 unique roosts for 7 radio-tracked bats; Pinus strobus (White Pine) snags (n = 8) were the most common roost sites. We observed a colony of 72 bats using a White Pine snag as a maternity roost. Roost trees were significantly larger in diameter and had more solar exposure above the roost and within the plot than random trees. Our data show the high abundance of Northern Long-eared Bats pre-WNS, highlight the use of White Pine roosts in an area impacted by a Dendroctonus frontalis (Southern Pine Beetle) outbreak, and support previous determinations of roost-selection flexibility by Northern Long-eared Bats across their range. Introduction Myotis septentrionalis (Northern Long-eared Bat) is a small (6–9 g) insectivorous bat found in eastern North America, ranging from southern Canada into the Northeast, Midwest, and most of the southeastern US (Caceres and Barclay 2000). Northern Long-eared Bats hibernate in cold caves and mines during winter (Caceres and Barclay 2000), but roost in forests and forage along forested hillsides and ridges during summer (Foster and Kurta 1999). Despite their wide distribution, Northern Long-eared Bat populations are decreasing rapidly across a large portion of their range due to white-nose syndrome (WNS), a devastating fungal disease that has reduced overwintering populations by >90% in many infected winter hibernacula (USFWS 2016). Due to significant population declines, Northern Long-eared Bats were listed as a federally threatened species (USFWS 2015). Roosts are crucial to bats for rearing young, protection from weather and predators, and hibernation (Kunz and Lumsden 2003). During spring and summer, 1Department of Biology, Indiana State University, Terre Haute, IN 47809. 2USDA Forest Service, Southern Research Station, Clemson, SC 29634. *Corresponding author - vanessaxrojas@gmail.com. Manuscript Editor: Roger Perry Southeastern Naturalist 141 V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 Northern Long-eared Bats primarily roost in cracks, crevices, and exfoliating bark of trees in upland forests (Lacki et al. 2009, Silvis et al. 2016) and wetlands (Foster and Kurta 1999). However, roost characteristics and tree species used as roosts vary across the species’ range. For example, Jung et al. (2004) found that Northern Long-eared Bats use Pinus strobus L. (White Pine) snags more than expected in Ontario, Canada. In Michigan, Northern Long-eared Bats mainly use Acer saccharinum L. (Silver Maple) and Fraxinus pennsylvanica Marsh. (Green Ash), and roost in live trees and snags in about equal proportions (Foster and Kurta 1999). In a pine-dominated landscape in Arkansas, 71% of roosts used by male and female Northern Long-eared Bats were in Pinus echinata Mill. (Shortleaf Pine; Perry and Thill 2007). It is important to know more about the roosting ecology of the Northern Longeared Bat across its range in order to protect summer populations and habitats, especially during the critical maternity stage, and to protect survivors in areas impacted by WNS. White-nose syndrome was first documented in the southern Appalachian Mountains during the winter of 2009–2010 (USFWS 2010), which followed the period of this study; hence, our work provides pre-WNS data on capture rates and roosting habits of Northern Long-eared Bats in this region. Our data can serve as a reference for future recovery efforts. Field-site Description We worked in a 281,788-ha area in the Great Smoky Mountains National Park, Cherokee National Forest, and Nantahala National Forest, in eastern Tennessee and western North Carolina. Elevation within this area varied from 250 m to 2025 m above sea level (asl), but we focused survey efforts in mixed pine–hardwood forests at <1000 m asl. Quercus (oak; 68% of forest cover) and Pinus spp. (yellow pine; 15%) (SAMAB 1996) were the most common forest types in our survey areas. The primary vegetation communities used by bats were low-elevation pine, Tsuga canadensis (L.) Carriére (Eastern Hemlock), Eastern Hemlock–White Pine, acidic cove, and oak–hickory forests (Schafale 2012). Most of the study area was forested (>90%), mainly with mid-successional forest, but it also contained some young and old-growth forest. Management in this region included prescribed fire and timber harvests (national forests only), however bats were not tracked to recently burned or harvested stands during our study. Natural disturbances included Dendroctonus frontalis Zimmermann (Southern Pine Beetle; Nowak et al. 2008) and Adelges tsugae Annand (Hemlock Woolly Adelgid; Nuckolls et al. 2009) outbreaks, which caused widespread formations of pine and Eastern Hemlock snags, respectively. Mean minimum and maximum daily temperatures were 15.7 °C and 28.6 °C in June, and 15.7 °C and 27.2 °C in July. Total precipitation was 6.4 cm both in June and July. The State Climate Office of North Carolina Raleigh, NC, provided weather data obtained from a station near the center of our study area (Robbinsville, NC, Station NCHE, elevation 640 m asl). Southeastern Naturalist V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 142 Methods We conducted our study from 1 June to 29 July 2009. We used mist nets (Avinet, Inc., Dryden, NY) to survey for bats at 37 sites located on roads and trails beside perennial streams 10–20 m in width. Our surveys occurred between 2045 and 0200 EDT, with nets checked at 8-minute intervals. For each survey, we placed 2–7 netsets (single or double-high nets) across roads, trails, and streams with edge and canopy cover that created forested corridors for potential flyways. Capture rates were defined as: number of bats captured x 1000, (total net area) x (total hours) where total net area is expressed as m2 and capture rate is presented as captures per 1000 m2h. We identified captured bats to species and marked them with a uniquely numbered, lipped aluminum forearm band (USFS-SRS or USFS-NC; Lambournes, Ltd., Birmingham, UK) of the appropriate size (2.9 mm or 4.2 mm). We recorded sex, age, reproductive condition, weight (g), and forearm length (mm) of each bat. Based on radio-transmitter availability and time constraints, we selected 8 Northern Long-eared Bats that weighed ≥7 g for radio telemetry; for each bat, we trimmed fur and attached a 0.42-g radio transmitter (Holohil Systems, Ltd., ON, Canada) between the scapulae using surgical glue (Torbot Group, Inc., Cranston, RI). We released all bats at the point of capture. Animal capture and handling methods were approved by the Clemson University Animal Research Committee (Animal Use Protocol 2009-016) and conducted under the American Society of Mammalogists’ guidelines (Animal Care and Use Committee 1998). Field work was conducted under permits held by J.M. O'Keefe: USFWS federal recovery permit TE206872, North Carolina permit ES261, Tennessee permit 3148, and National Park Service Permits GRSM-2009-SCI-0075 and GRSM-2012-SCI-0085. Using a Telonics TR-5 receiver (Telonics, Inc., Mesa, AZ) and a 3-element Yagi antenna, we radio-tracked bats to day roosts and recorded GPS coordinates for each tree location. For 6 roosts, we were able to visually locate the specific roost location on the tree and confirm roost sites with observations at dusk, counting bats that emerged from the tree. For roost trees containing ≥2 bats (all snag s), we identified a random snag with visible roost potential (i.e., bark peeling from the tree trunk or a crevice). We used a method described by O’Keefe and Loeb (2017) to locate a random snag to pair with a known roost, which facilitated stand-level comparisons between roosts and random trees. We recorded tree and plot characteristics for 8 of the 14 roost trees and 6 corresponding random trees. Time and personnel constraints made it impossible for us to complete random plots for every roost. Thus, we prioritized measuring random-tree characteristics for roosts with ≥2 bats. Two roost trees were within the same 0.1-ha plot, which was centered between the 2 roosts and matched with 1 random plot; thus, we measured only 7 plots for 8 roost trees. At each tree (roost or random; hereafter, focal tree), we recorded species, diameter at breast height (DBH, cm), and tree height (m). We documented total Southeastern Naturalist 143 V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 number of live trees and snags ≥10 cm DBH in a 0.1-ha plot around each focal tree. We tallied all saplings ≤8.9 m from the focal tree. We estimated percent canopy closure to the nearest 25% for the entire plot and directly above the focal tree. For each focal tree, we measured distance to the nearest road (m) and stream (m) in a GIS (ArcMap v10, Esri, Redlands, CA). The National Park Service and USDA Forest Service provided spatial data for major roads and minor roads/trails; we acquired stream data from the National Hydrography Dataset (USGS 2013). We used Wilcoxon rank-sum tests to compare quantitative traits of roost and random plots, and present means ± 1 standard error for these variables and for elevation, aspect, and slope of roost locations. Results We conducted 43 nights of netting, for a total effort of 11,202 m2h (net area × hours), and captured 302 bats of 11 species (Table 1). We averaged 7 captures per survey night, with a variation of 0–29 bats; only 2 nights did not yield captures. Northern Long-eared Bats and Lasiurus borealis (Eastern Red Bat) were captured most often; 97 and 58 captures, respectively. We captured 38 adult female, 31 adult male, and 26 juvenile Northern Long-eared Bats, plus 2 individuals (1 male) with incomplete data. The capture rate across the entire season for Northern Long-eared Bats was 8.66 bats per 1000 m2h (Table 1). A typical survey night (81.5 m2 total net area × 3.15 hours) yielded 2.22 Northern Long-eared Bats with a maximum of 10 bats captured during 1 net night; we did not capture this species on 10 survey nights. We radio-tracked 8 reproductive female Northern Long-eared Bats (2 pregnant, 4 lactating, 2 post-lactating) and documented 14 unique roosts for 7 bats. Roost trees were used 1−4 days each over tracking periods of 2–5 days/bat (mean = 4.3 days/bat). Of the 14 roost trees we located, 11 were snags: 8 White Pine, 2 Pinus virginiana Mill. (Virginia Pine), and 1 Quercus rubra L. (Northern Red Oak). The Table 1. Capture results from 43 summer mistnetting surveys of Northern Long-eared Bats in the Cherokee National Forest, Nantahala National Forest, and Great Smoky Mountains National Park in June and July 2009. Capture results include total number of individuals captured during 43 surveys (# bats), and capture rate ([# bats captured / (total net area × total hours)] x 1000) or captures per 1000 m2h. Species Authority Common name # bats Rate Corynorhinus rafinesquii Lesson Rafinesque’s Big-eared Bat 3 0.27 Eptesicus fuscus Palisot de Beauvois Big Brown Bat 35 3.12 Lasiurus borealis Müller Eastern Red Bat 58 5.18 Lasiurus cinereus Palisot de Beauvois Hoary Bat 2 0.18 Lasionycteris noctivigans La Conte Silver-haired Bat 9 0.80 Myotis leibii Audubon and Bachman Eastern Small-footed Bat 1 0.09 Myotis lucifugus Le Conte Little Brown Bat 29 2.59 Myotis septentrionalis Trouessart Northern Long-eared Bat 97 8.66 Myotis sodalis Miller and Allen Indiana Bat 46 4.11 Nycticeius humeralis Rafinesque Evening Bat 3 0.27 Perimyotis subflavus F. Cuvier Tri-colored Bat 19 1.70 Southeastern Naturalist V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 144 remainder of the roosts were in live trees: 1 Acer rubrum L. (Red Maple), 1 Quercus alba L. (White Oak), and 1 damaged hardwood sapling (unidentified species). We recorded and analyzed tree and plot characteristics for 8 focal roost-trees, all snags: 6 White Pine, 1 Virginia Pine, and 1 Northern Red Oak. White Pine roosts were at all slope positions (lower, mid, and upland), whereas the Virginia Pine and the Northern Red Oak were upland roosts. Roost trees were at a mean elevation of 473 ± 78 m, a mean slope of 32.3 ± 22.2%, and were usually south-facing (mean aspect = 187 ± 75°). Roost trees had an average diameter of 58.2 ± 8.1 cm and were significantly larger than random trees (P = 0.008; Table 2). Canopy closure above roost trees (22 ± 9%) was less than half the closure above random trees (P = 0.03). Within the 0.1-ha plot, canopy closure was significantly less than closure in random plots (P = 0.002). Roosts were closer to streams than roads, with a mean distance of 98 ± 24 m to a stream and 1140 ± 367 m to a road. We conducted 1 emergence count at 6 focal trees. The largest colony sizes detected were 72 bats roosting under exfoliating bark of a White Pine snag (27.0 m tall, 94.2 cm DBH) and 19 individuals under exfoliating bark of another White Pine snag (22.0 m tall, 89.5 cm DBH), both located in Great Smoky Mountains National Park. We noted 4 other roosts under exfoliating bark of snags (11.9–39.0 m tall, 42–62 cm DBH) that held 1–4 bats each. We conducted all emergence counts in early to mid-June (pre-volant period), except 1, which was conducted in early July and yielded only 1 bat. We counted only bats that emerged from roost trees. Discussion This study presents capture rates and the first description of roost-tree characteristics for Northern Long-eared Bats in the southern Appalachian Mountains of North Carolina and Tennessee. During the study period, Northern Long-eared Bats were relatively common; we captured more of them per typical net-night than all Table 2. Mean (± SE) characteristics of Northern Long-eared Bat tree roosts, random trees, and habitat within 0.1-ha plots centered on focal trees in the Cherokee National Forest, Nantahala National Forest, and Great Smoky Mountains National Park in June and July 2009. W = Wilcoxon rank-sum test statistic and P = significance measured at α < 0.05. Random Roost plots (n = 8) plots (n = 6) Characteristic Mean Variation Mean W P Height (m) 20.5 ± 4.0 4.6–39.0 12.4 ± 1.5 15.0 0.282 Diameter (cm) 58.2 ± 8.1 28.8–94.2 22.8 ± 6.4 4.0 0.008 % bark remaining 59 ± 7 30–80 63 ± 17 29.0 0.558 % canopy closure above roost 22 ± 9 0–50 67 ± 15 40.5 0.032 % plot canopy closure 50 ± 6 25–69 83 ± 3 48.0 0.002 # snags in plot 11 ± 2 4–19 10 ± 5 11.5 0.120 # live trees in plot 40 ± 7 17–79 49 ± 4 38.5 0.069 # saplings 80 ± 18 34–168 59 ± 15 20.5 0.698 Distance to road (m) 1140 ± 367 367–3313 1227 ± 427 26.0 0.852 Distance to water (m) 98 ± 24 14–228 452 ± 234 34.5 0.196 Southeastern Naturalist 145 V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 other bat species. Our documentation of a large pre-volant colony (72 bats emerged) in a White Pine snag that was likely killed by Southern Pine Beetles (Nowak et al. 2008), is similar to findings in other regions. For example, pre-volant colony sizes for Northern Long-eared Bats have been as high as 60 bats using a cavity roost in a live Silver Maple in Michigan (Foster and Kurta 1999), 65 in a hardwood in West Virginia (Menzel et al. 2002), and 75 in a Northern Red Oak snag in North Carolina (O’Keefe 2009). This study was done in tandem with a higher priority Myotis sodalis (Indiana Bat) project. Our limited data for Northern Long-eared Bats relate to time and personnel constraints, and we recognize these limitations (e.g., small sample-size collected and data for only 1 year; Silvis et al. 2015); however, given the decline of Northern Long-eared Bat populations (USFWS 2016) and our limited ability to obtain further data on this species, the data presented here are highly valuable for developing conservation and recovery strategies for this species in the Southern Appalachians. Northern Long-eared Bat roost-tree selection appears to vary by region, suggesting they are a flexible species. Our study was not the first to document the use of pine by Northern Long-eared Bats. Male Northern Long-eared Bats in central Ontario use White Pine roosts most often and more than would be expected if they utilized that tree species in proportion to availability (Jung et al. 2004). In northeastern Kentucky, Northern Long-eared Bat roosting is mainly solitary and in Shortleaf Pine and Oxydendrum arboreum (L.) DC. (Sourwood) (Lacki and Schwierjohann 2001). Northern Long-eared Bats in Arkansas prefer Shortleaf Pine snag roosts over hardwood snags (Perry and Thill 2007). In South Dakota, female Northern Longeared Bats roost in Pinus ponderosa L. (Ponderosa Pine), most of which are snags (81%; Cryan et al. 2001). Although Northern Long-eared Bats use primarily oak species along with other hardwoods in northwestern North Carolina, they have also been documented using White Pine and 2 unknown Pinus species (O’Keefe 2009). However, selection for hardwood roosts predominates for this species throughout much of its distribution. Examples of hardwoods used by lactating females include Robinia pseudoacacia L. (Black Locust) (WV; Johnson et al. 2009, Menzel et al. 2002), Sassafras albidum (Nutt.) Nees (Sassafras) (KY; Silvis et al. 2012), Silver Maple (MI; Foster and Kurta 1999), Fagus grandifolia Ehrh. (American Beech) (NH; Sasse and Pekins 1996), and Quercus palustris Münchh. (Pin Oak) (IL; Carter and Feldhamer 2005). All of the roosts we observed were under bark, yet it is also common for Northern Long-eared Bats to use crevices and cavities. We might expect roost type (bark or crevice) to be predicted by tree species, but there is variation. Our results were similar to findings of Jung et al. (2004) who found that Northern Long-eared Bats roost primarily under exfoliating bark (81.8% of roosts), mainly in White Pine snags. However, current data suggest Northern Long-eared Bats are flexible with regard to where in the tree they roost, using both cavities and crevices (Johnson et al. 2009, Lacki et al. 2009, Silvis et al. 2012). For example, in Arkansas, female Northern Long-eared Bats roost under bark and in crevices at similar rates (43%), and cavities minimally (14%), whereas males use bark roosts (61%) more Southeastern Naturalist V.G. Rojas, J.M. O’Keefe, and S.C. Loeb 2017 Vol. 16, No. 2 146 often than cavities (25%) and crevices (15%); most roosts were in Shortleaf Pine snags (67%; Perry and Thill 2007). Nearly equal bark- and cavity-use proportions have been documented in hardwood roosts used by Northern Long-eared Bats in Michigan (Foster and Kurta 1999). Many researchers do not present roost- and available-tree characteristics such as decay stage, percentage of bark available, or presence of cavities. With limited information available on specific characteristics of Northern Long-eared Bat roosts across the species’ range, we cannot conclude that the use of bark, crevice, or cavity is dependent on tree species. Large trees with moderate to abundant solar exposure are important maternity roosts for Northern Long-eared Bats. Although roost and random plots were similar in snag, live tree, and sapling counts, they differed in canopy closure, both above the roost and within the plot. Canopy closure at roosts varies from 22% to 83% across various Northern Long-eared Bat habitats in different portions of its range. Bats in our study used trees with greater mean solar exposure (22 ± 9% canopy closure) than populations in the Midwest (39–44%; Carter and Feldhamer 2005, Foster and Kurta 1999, Jung et al. 2004), Northeast (83%; Sasse and Pekins 1996), and Central Appalachians (66–92%; Lacki and Schwierjohann 2001, Menzel et al. 2002). Regional variation might be explained by differences in ambient conditions and characteristics of available roosts (Patriquin et al. 2016). We have presented data on capture rates and the roosting characteristics of Northern Long-eared Bats in the southern Appalachians to aid in defining suitable summer habitat for the species across its range. Our capture-rate data provide pre- WNS records in a region now heavily impacted by this disease. Our data support the notion that roosting flexibility helps explain why Northern Long-eared Bats were once a common forest bat with a wide distribution (Lacki et al. 2009). Recognizing that populations are being drastically impacted by WNS, we recommend future surveys to locate roosts and determine relative abundance of Northern Longeared Bats. Acknowledgments We thank Dylan Horvath, Caroline Byrne, Dottie Brown, Mary Miller, and Eric Winters for field assistance. 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