Issues and Perspectives Status of an Anomalous Population of Northern Long- Eared in Coastal North Carolina

Gary W. Jordan*

U.S. Fish and Wildlife Service, P.O. Box 33726, Raleigh, North Carolina 27636

Abstract

The northern long-eared Myotis septentrionalis was discovered in coastal North Carolina in 2007. Work began in Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 2015 to document the species’ distribution and behavior in eastern North Carolina, and the known range of the species has expanded from 4 coastal counties to 19. Captures occurred in all months of the year and mostly occurred in or adjacent to wetland forest. Captures occurred exclusively in the Middle Atlantic Coastal Plain ecoregion of the state, and the species has not been documented in the Southeastern Plains or Piedmont ecoregions. The lack of captures in the middle of the state suggests spatially disjunct populations in North Carolina. The bats were observed to be active throughout most of the winter and roosted in trees. During late fall–winter 2015–2018, 43 bats were tracked to 165 winter roost trees located mostly in wetland forest. The species’ winter activity in coastal North Carolina represents a novel survival strategy as opposed to the hibernation behavior it is assumed to use in the rest of its range. This portion of the state is nearly devoid of caves or mines suitable for hibernacula, but has milder winters with insect activity. During spring 2019, 21 reproductive females were captured in the northern coastal plain and tracked to 64 maternity roost trees located mostly in wetland forest. Pregnant females began to be captured on April 25 and juveniles on June 16, indicating that pups are likely born in late May. Swab samples collected during late fall–winter from species susceptible to white-nose syndrome provided no evidence of Pseudogymnoascus destructans. Since northern long-eared bats in coastal North Carolina are active most of the winter and not dependent upon caves or mines for hibernation, they are likely not susceptible to white-nose syndrome. With the species in sharp decline elsewhere due to white-nose syndrome, this coastal population may serve as a refugium.

Keywords: Myotis septentrionalis; northern long-eared bat Received: March 27, 2020; Accepted: June 9, 2020; Published Online Early: August 2020; Published: December 2020 Citation: Jordan GW. 2020. Status of an anomalous population of northern long-eared bats in coastal North Carolina. Journal of Fish and Wildlife Management 11(2):665–678; e1944-687X. https://doi.org/10.3996/JFWM-20-018 Copyright: All material appearing in the Journal of Fish and Wildlife Management is in the public domain and may be reproduced or copied without permission unless specifically noted with the copyright symbol &. Citation of the source, as given above, is requested. The findings and conclusions in this article are those of the author(s) and do not necessarily represent the views of the U.S. Fish and Wildlife Service. * Corresponding author: [email protected]

Introduction primary hibernacula (Caceres and Barclay 2000; Federal Register 2016). Despite formerly being one of the most The northern long-eared bat Myotis septentrionalis is a abundant bat species in much of its range, their small (~5–8 g) insectivorous bat that occurs in much of populations are declining rapidly across a large portion the eastern and north-central United States and southern of their range because of white-nose syndrome (WNS), a Canada (Caceres and Barclay 2000; Federal Register disease caused by the fungus Pseudogymnoascus de- 2015). During spring and summer months, they roost structans (USFWS 2020). primarily in cavities, cracks, crevices, and exfoliating bark Although the presence of the northern long-eared bat of trees of a variety of species and sizes (Foster and Kurta was well documented within the Blue Ridge ecoregion of 1999; Caceres and Barclay 2000; Lacki et al. 2009; Silvis et western North Carolina before WNS (Rojas et al. 2017; al. 2016). Until recently, all northern long-eared bats were North Carolina Wildlife Resources Commission, unpub- thought to hibernate, with caves and mines being the lished data), there was little information regarding the

Journal of Fish and Wildlife Management | www.fwspubs.org December 2020 | Volume 11 | Issue 2 | 665 Northern Long-Eared Bats in Coastal North Carolina G.W. Jordan presence of the species in eastern North Carolina. It had intensively managed loblolly pine Pinus taeda, is also only recently been discovered in coastal North Carolina common in the study area. when it was listed as a federally threatened species in 2015 under the U.S. Endangered Species Act (ESA 1973, Methods as amended; Federal Register 2015). The species was first mist-netted and identification confirmed through deox- Review of previous museum records yribonucleic acid analysis in Washington County in 2007 All known information on previous northern long- (Morris et al. 2009) and subsequently caught in Camden eared bat records from eastern North Carolina was County in 2012 (Grider et al. 2016) and Currituck County reviewed. Inquiries were made to the University of North in 2014 (Tetra Tech 2015). In addition to mist-netting Carolina Wilmington Natural History Collection and records, a specimen of unknown origin presumably North Carolina Museum of Natural Sciences regarding collected from New Hanover County in 1996 is stored in specimens of questionable identification. The the University of North Carolina Wilmington Natural curators at each institution subsequently re-examined all History Collection (UNCW 4925). These county records Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 occur within the Coastal Plain where no known specimens suspected of misidentification or mislabeling. hibernacula occur, presumably because of the near absence of caves or mines suitable for hibernation. Since Presence–absence mist-net surveys they are not considered long-distance migrators (Federal To map out the distribution of northern long-eared Register 2015), Morris et al. (2009) suggested that bats in eastern North Carolina, environmental consulting northern long-eared bats in coastal North Carolina either companies were contracted to conduct mist-net pres- 1) travel much farther to hibernacula than previously ence–absence surveys in multiple counties across the thought, 2) utilize alternative hibernacula such as tree study area over three spring–summer seasons. Consul- cavities, or 3) do not hibernate at all in the milder winters tants netted a combined 210 nights during 2016–2018. of the Coastal Plain. Work began in 2015 to explore these Additionally, the author conducted 110 nights of netting possibilities and to determine the range of the species in throughout all seasons March 2016–March 2020, and eastern North Carolina. An ad hoc and adaptive North Carolina Wildlife Resources Commission biologists approach was utilized for these investigations, and most conducted 51 nights of netting across the spring, of the fieldwork was conducted by environmental summer, and fall seasons of 2016–2019. Single-, dou- consultants through multiple contracts. The following ble-, or triple-high mist nets (Avinet Research Supplies, information and analysis are a synthesis of multiple Portland, Maine) 2.6–18 m wide were used to capture reports submitted to the U.S. Fish and Wildlife Service by bats along trails, forest roads, or other flyways. Netting environmental consultants and supplemented by data collected by the author, North Carolina Wildlife Resourc- generally occurred from sunset to 5 h after sunset, es Commission biologists, and other wildlife profession- although occasional deviations occurred. The number of als. My objectives are to provide a current and nets utilized per night varied. Data recorded for each bat comprehensive status of the northern long-eared bat in captured included date, time, location, forest type, eastern North Carolina, to report novel behavior for the species, sex, age, reproductive status, forearm length, species, and to provide natural history data for this weight, wing damage index score (Reichard and Kunz unique population. 2009), and band number. Each northern long-eared bat was fitted with a 2.9-mm numbered band (Porzana Ltd., Study area East Sussex, UK). In addition to the efforts of the The study area included numerous sites within the aforementioned consultants and government biologists, easternmost 59 counties of North Carolina (Figure 1). additional entities (e.g., universities and other consul- Three Environmental Protection Agency Level III ecor- tants working on unrelated projects) performed mist-net egions occur within the study area: from west to east, surveys within the study area during the spring, summer, Piedmont, Southeastern Plains, and Middle Atlantic and fall seasons of 2015–2019. Northern long-eared bat Coastal Plain (Griffith et al. 2002a, 2002b). Although records from all known surveys were pooled for this some small caves are present in the Piedmont, caves review. potentially suitable as bat hibernacula are extremely rare in the Southeastern Plains and Middle Atlantic Coastal Late fall–winter mist netting and radiotelemetry Plain of North Carolina (Culver et al. 1999; Grotto Center To determine whether northern long-eared bats were 2020). Underground mines are common in the Piedmont present and active during the winter in coastal North but are absent from the Southeastern Plains and Middle Carolina, mist netting and radiotelemetry were con- Atlantic Coastal Plain (Carpenter 1976; North Carolina ductedduringaseasonwhensuchactivitiesare Department of Environmental Quality 2020). Although normally not conducted. Environmental consulting many natural forest types are present in the study area, companies were contracted to capture bats and track they were lumped into either wetland forest or upland them to roost sites over three late fall–winter seasons forest on the basis of descriptions in Schafale and 2015–2018. Study locations occurred in the best Weakley (1990) and Schafale (2012). Pine plantation, an available habitat across multiple counties in the Middle unnatural forest type usually consisting of planted and Atlantic Coastal Plain ecoregion (Figure 2). Initially,

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Figure 1. Map depicting study area in 2015–2020 with superimposed Environmental Protection Agency Level III ecoregions. Study area corresponds to the boundaries of the U.S. Fish and Wildlife Service’s Raleigh Ecological Services Field Office work area. netting occurred when temperatures were 7.28Cbut etry Systems, Isanti, Minnesota), or Lotek PicoPip (Lotek was later revised to 4.48C because of activity Wireless, Newmarket, Ontario, Canada) radio transmit- documented at lower temperatures. Data recorded for ter. Tracking was conducted primarily by terrestrial each bat captured included date, time, location, forest (pedestrian or vehicular) means; however, on a few type,species,sex,age,reproductivestatus,forearm occasions an airplane was used. Various models of length, weight, wing damage index score, band receivers and antennae were utilized. An attempt was number, and temperature at time of capture. Each made to track each bat for a minimum of 21 d. Data northern long-eared bat was fitted with a 2.9-mm recorded for each roost tree included location, date, numbered band. During late fall–winter 2016–2017, all forest type, type of roost, tree species, diameter at captured WNS-susceptible species were swabbed for breast height (dbh), and tree condition (live or dead). Pseudogymnoascus destructans and samples were sub- Ten of the radio transmitters deployed in 2017–2018 mitted to the Southeastern Cooperative Wildlife Disease were temperature-sensitive models, and model Study (University of Georgia, Athens). During late fall– winter 2017–2018, all captured WNS-susceptible species R4500SD receiver data loggers (Advanced Telemetry were swabbed and samples were submitted initially to Systems) and DS1923 iButtont temperature–humidity the BioInnovation Laboratory at Kennesaw State Uni- data loggers (Maxim Integrated, San Jose, California) versity (Kennesaw, Georgia), with some retests later were placed near roost trees to collect information on being sent to the National Wildlife Health Center bat body temperature and activity level (normothermic (Madison, Wisconsin). or torpid). An attempt was made to conduct at least Northern long-eared bats 6.0 g were fitted with one emergence survey at each roost tree and was either a Holohil model LB-2X (Holohil Systems Ltd, Carp, generally conducted from 1 half hour before dusk to 1 h Ontario, Canada), ATS model #A2414 (Advanced Telem- after dusk.

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Figure 2. Map depicting general locations of northern long-eared bat Myotis septentrionalis tracking and roost studies in Coastal Plain of North Carolina during late fall–winter seasons of 2015–2018 and spring maternity season of 2019.

Maternity studies condition (live or dead). One emergence survey was To determine the maternity season for northern long- conducted at each roost tree and was generally eared bats in coastal North Carolina, environmental conducted from 1 half hour before dusk to 1 h after consulting companies were contracted to capture bats at dusk. In addition to the consultants’ efforts, researchers North River Game Land (Camden and Currituck counties), from the Conservation Management Institute at Virginia Merchants Millpond State Park (Gates County), and Tech also captured and tracked reproductive female Alligator River National Wildlife Refuge (Dare County) northern long-eared bats during June 2019 at North from late April 2019 to mid-July 2019 and use River Game Land. These data were pooled with the radiotelemetry to track pregnant and lactating females consultants’ data for this review. to roost sites (Figure 2). Data recorded for each bat captured included date, time, location, forest type, Results species, sex, age, reproductive status, forearm length, weight, wing damage index score, and band number. Review of previous museum records Most adult northern long-eared bats and some juveniles Three specimens within the University of North were fitted with a 2.9-mm numbered band. Pregnant or Carolina Wilmington Natural History Collection repre- lactating females were fitted with a Holohil model LB-2X senting purported county records for the northern long- radio transmitter. Tracking was conducted by terrestrial eared bat within the Piedmont ecoregion in Lee (UNCW means (pedestrian or vehicular), and various models of 9143), Davidson (UNCW 9148), and Mecklenburg (UNCW receivers and antennae were utilized. An attempt was 22483) counties were re-examined by the mammal made to track each bat for a minimum of 12 d. Data curator in 2017. A range map provided in White et al. recorded for each roost tree included location, date, (2018) was partially based on these museum records. forest type, type of roost, tree species, dbh, and tree After re-examination, two of these specimens were

Journal of Fish and Wildlife Management | www.fwspubs.org December 2020 | Volume 11 | Issue 2 | 668 Northern Long-Eared Bats in Coastal North Carolina G.W. Jordan determined to be evening bats humeralis and Engineering 2018). Winter roost trees occurred in one specimen was determined to be a little brown bat wetland forest (94.6%), upland forest (1.2%), pine Myotis lucifugus (W.D. Webster, University of North plantation (1.2%), and unknown forest (3.0%) and were Carolina Wilmington, personal communication). The represented by at least 22 species (Table 2). The mean mammal curator re-examined the two specimens within dbh of winter roost trees was 30.9 cm (27.6–34.3 cm, the North Carolina Museum of Natural Sciences (NCSM 95% confidence interval [CI]; Figure 4). Most trees were 45 and NCSM 3582) representing purported county live (84.2%), but dead trees (9.1%) and those of unknown records in Wake County (Piedmont ecoregion) in 2020. status (6.7%) were also used. Roost types included Range maps provided in Morris et al. (2009) and White et cavities (55.8%), cracks or crevices (11.5%), exfoliating al. (2018) were partially based on these museum records. bark (4.8%), and unknown (27.9%). Emergence surveys After re-examination, these specimens were determined were conducted at most roost trees, with some being to be little brown bats (L.J. Gatens, North Carolina repeated one or more times for a total of 195 surveys. Museum of Natural Sciences, personal communication). The mean number of bats observed emerging per survey Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 was 0.5 (0.4–0.6, 95% CI). Presence–absence mist-net surveys Utilizing temperature-sensitive transmitters, Copper- From 2007 when the species was first identified in head Environmental Consulting (2018) documented 10 eastern North Carolina to March 2020, 181 unique northern long-eared bats in sustained torpor during late northern long-eared bats were captured by mist netting, fall–winter 2017–2018. The longest sustained torpor with 163 captured since 2015 (Table S1, Supplemental bout for each radio-tagged bat averaged 6.8 d (4.1–9.5 Material). Captures included 146 adults, 32 juveniles, and d, 95% CI), and the coldest temperature for a normo- 3 of unknown age and occurred in all months of the year, thermic bat was 3.08C. However, Ecological Engineering with 35 captures in winter (December–February). Cap- (2018) documented a northern long-eared bat on tures occurred in wetland forest (73.5%), upland forest December 31, 2017 that moved 4.9 km from one roost (24.9%), and pine plantations (1.7%). However, upland tree to another sometime after sunset when the air forest and pine plantation capture sites were generally in temperature was 2.08C (Figure 5). proximity (, 0.5 km) to wetland forest. Captures None of the northern long-eared bats captured in occurred exclusively within the Middle Atlantic Coastal coastal North Carolina showed clinical symptoms for Plain ecoregion. All but one capture occurred , 50 km WNS. Swab samples collected from 67 bats of three from the Atlantic Ocean or one of its major sounds, with species (northern long-eared bat, southeastern myotis the lone outlier captured ~87 km from the Atlantic Myotis austroriparius, and big brown bat fuscus) Ocean. Despite significant survey efforts (Table S2, during late fall–winter 2016–2017 were submitted to the Supplemental Material), northern long-eared bats have Southeastern Cooperative Wildlife Disease Study. Pseu- never been captured within the Piedmont or Southeast- dogymnoascus destructans was not detected on any ern Plains ecoregions. Nineteen coastal counties in North sample (Southeastern Cooperative Wildlife Disease Study Carolina currently have records for the species (Figure 3). 2017). Swab samples collected from 131 bats of five These data were synthesized from multiple sources species (northern long-eared bat, southeastern myotis, (Morris et al. 2009; Tetra Tech 2015, 2016; Ecological big brown bat, , and little brown bat) Engineering 2016, 2018; Grider et al. 2016; Three Oaks during late fall–winter 2017–2018 were submitted to the Engineering 2016, 2017, 2018; Calyx Engineers þ BioInnovation Laboratory at Kennesaw State University. Consultants 2017a, 2017b, 2018a, 2018b, 2019; Copper- Four samples initially tested positive for P. destructans (C. head Environmental Consulting 2017, 2018, 2019; VHB Cornelison, Kennesaw State University, unpublished Engineering NC 2019; De La Cruz et al. 2020; North data). These four samples were subsequently submitted Carolina Wildlife Resources Commission, unpublished to the National Wildlife Health Center, and more refined data; U.S. Fish and Wildlife Service, unpublished data). testing yielded a negative result for three samples and an equivocal result for one sample (National Wildlife Health Late fall–winter mist netting and radiotelemetry Center 2019). The northern long-eared bat was documented to be active during most of the winter at multiple locations Maternity studies throughout the Middle Atlantic Coastal Plain ecoregion During spring–summer 2019, 14 pregnant females of North Carolina. The lowest temperature at which the were captured April 25–June 1, 12 lactating females were species was captured was 7.88C, but it is important to captured June 3–June 27, 6 postlactating females were note that a tricolored bat Perimyotis subflavus was captured June 16–July 15, and 24 juveniles were captured at 3.38C. Cumulatively, 43 northern long-eared captured June 16–July 15. Twenty-one pregnant or bats were tracked to 165 winter roost trees (Table 1; lactating northern long-eared bats were fitted with Figure 2; Table S3, Supplemental Material; Ecological transmitters and tracked for 1–15 d to 64 maternity Engineering 2016; Copperhead Environmental Consult- roost trees (Figure 2; Table S4, Supplemental Material; ing 2017; Calyx Engineers þ Consultants 2018a; Copper- Calyx Engineers þ Consultants 2019; Copperhead Envi- head Environmental Consulting 2018; Ecological ronmental Consulting 2019; VHB Engineering NC 2019;

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Figure 3. Map depicting the known range of the northern long-eared bat Myotis septentrionalis as of March 2020 in North Carolina by county. Note that some western county boundaries overlap both the Blue Ridge and Piedmont ecoregions, and some eastern county boundaries overlap both the Middle Atlantic Coastal Plain and Southeastern Plains ecoregions. However, the record collection locations in the western counties occur entirely within the Blue Ridge ecoregion, and the record collection locations in the eastern counties occur entirely within the Middle Atlantic Coastal Plain ecoregion.

De La Cruz et al. 2020). Maternity roost trees were found mean number of bats observed emerging per survey was in wetland forest (92.2%) and upland forest (7.8%) and 4.7 (1.8–7.6, 95% CI); however, the median number was were represented by at least 14 species (Table 3). The one. mean dbh of maternity roost trees was 20.6 cm (17.5– 23.6 cm, 95% CI; Figure 4). Both live trees (59.4%) and Discussion dead trees (40.6%) were used. Roost types included cavities (59.4%), exfoliating bark (29.7%), cracks or At the time of listing as a threatened species under the crevices (4.7%), and unknown (6.2%). Emergence surveys ESA (1973) in 2015 and accounting for recent museum were conducted at all 64 maternity roost trees, and the record identification corrections, only four eastern North

Table 1. Number of northern long-eared bats Myotis septentrionalis tracked, number of days each bat tracked, mean days tracked, and number of roost trees documented in the Coastal Plain of North Carolina during late fall–winter seasons of 2015–2018.

Tracking dates November 17–December 18, 2015 November 17, 2016–March 20, 2017 November 16, 2017–March 21, 2018 Number tracked 7 10 26 Days each tracked 6–29 6–23 5–30 Mean days tracked 16.4 18.8 19.7 Number of roost trees 25 41 99

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Table 2. Percentage of total for each tree species used for 165 roost sites by northern long-eared bats Myotis septentrionalis in the Coastal Plain of North Carolina during late fall–winter seasons of 2015–2018.

Common name Scientific name % Swamp tupelo Nyssa biflora 20.6 Water tupelo Nyssa aquatica 20.6 Bald cypress Taxodium distichum 10.3 Red maple Acer rubrum 10.3 Unknown Unknown 6.7 Sweetgum Liquidambar styraciflua 4.8 Tupelo spp. Nyssa spp. 3.0 Swamp bay Persea palustris 3.0

Swamp cottonwood Populus heterophylla 2.4 Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 Loblolly pine Pinus taeda 2.4 Redbay Persea borbonia 1.8 Blackgum Nyssa sylvatica 1.8 Atlantic white cedar Chamaecyparis thyoides 1.2 Wax myrtle Morella cerifera 1.2 Sweetbay Magnolia virginiana 1.2 Oak spp. Quercus spp. 1.2 Pond cypress Taxodium ascendens 1.2 Bay spp. Persea spp. 0.6 American holly Ilex opaca 0.6 Longleaf pine Pinus palustris 0.6 Ash spp. Fraxinus spp. 0.6 Loblolly bay Gordonia lasianthus 0.6 Swamp titi Cyrilla racemiflora 0.6 Sourwood Oxydendrum arboreum 0.6 Pine spp. Pinus spp. 0.6 Carolina ash Fraxinus caroliniana 0.6 Pond pine Pinus serotina 0.6 Figure 5. Photograph of a bald cypress tree Taxodium distichum cavity occupied by a northern long-eared bat Myotis Carolina counties had records for northern long-eared septentrionalis on January 5, 2018 in the Croatan National Forest, Craven County, North Carolina. The snow depicts a rare bats. The increase to 19 counties (Figure 3) reflects weather event for the southern Coastal Plain of North Carolina. substantial improvement in the knowledge of the Photograph credit: Dottie Brown. species’ range in eastern North Carolina. This expansion is almost certainly the result of increased survey effort and not a true range expansion. Despite the encouraging number of recent captures, the numbers caught in mist nets are relatively low when compared with other parts

Table 3. Percentage of total for each tree species used for 64 maternity roost sites by northern long-eared bats Myotis septentrionalis in the northern Coastal Plain of North Carolina during spring 2019.

Common name Scientific name % Red maple Acer rubrum 21.9 Water tupelo Nyssa aquatica 18.8 Carolina ash Fraxinus caroliniana 14.1 Loblolly pine Pinus taeda 12.5 Bald cypress Taxodium distichum 7.8 Swamp bay Persea palustris 6.3 Sweetgum Liquidambar styraciflua 3.1 Slippery elm Ulmus rubra 3.1 Figure 4. Distribution of sizes (diameter at breast height) of Longleaf pine Pinus palustris 3.1 trees used as winter roosts (n ¼ 165) during late fall–winter Sweetbay Magnolia virginiana 1.6 seasons of 2015–2018 and as maternity roosts during spring Elm spp. Ulmus spp. 1.6 2019 (n ¼ 64) by northern long-eared bats Myotis septentrionalis American holly Ilex opaca 1.6 in the coastal plain of North Carolina. Of 165 total winter roost Loblolly bay Gordonia lasianthus 1.6 trees, 12 trees of unknown size were not factored into the White oak Quercus alba 1.6 analysis. Swamp tupelo Nyssa biflora 1.6

Journal of Fish and Wildlife Management | www.fwspubs.org December 2020 | Volume 11 | Issue 2 | 671 Northern Long-Eared Bats in Coastal North Carolina G.W. Jordan of its range pre-WNS (e.g., Rojas et al. 2017), but this low in the rest of its range (Federal Register 2015). Although capture rate could be a result of coastal North Carolina low-level winter activity has previously been document- being on the periphery of the species’ overall range. ed for northern long-eared bats, those observations were Captures occurring in every month of the year confirm near cave or mine openings at known hibernacula the presence of a year-round population of northern (Whitaker and Rissler 1992; Boyles et al. 2006; Bernard long-eared bats in the Middle Atlantic Coastal Plain and McCracken 2017). Despite the near absence of caves ecoregion within North Carolina. However, it is unclear if and mines as suitable hibernacula, Grider et al. (2016) the bats are an entirely resident population or if some documented year-round activity of multiple species of individuals make seasonal movements. The lack of bats while using acoustic methods in the Coastal Plain of captures from recent and previous efforts within the North Carolina. In contrast to the Piedmont ecoregion, Piedmont (including the portion outside the study area) the Middle Atlantic Coastal Plain ecoregion maintains and Southeastern Plains ecoregions (Table S2, Supple- insect activity throughout most of the winter, though at mental Material; Lambiase et al. 2000, 2005; Cameron et a lower level than the rest of the year (personal al. 2004; Kalcounis-Rueppell and Li 2016) strongly observation). With higher nightly winter temperatures Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 suggests spatially disjunct populations of northern (Grider et al. 2016), the Coastal Plain appears to provide long-eared bats in North Carolina. A formerly robust more opportunities for winter foraging than does the population in the Blue Ridge ecoregion of western North Piedmont. For example, Whitaker et al. (1997) docu- Carolina appears separated from the population in the mented feeding in eastern red bats borealis Middle Atlantic Coastal Plain ecoregion by a large during winter in the Great Dismal Swamp of southeast- unpopulated area in the middle of the state (Figure 3), ern Virginia and northeastern North Carolina by exam- despite the populations being genetically nearly identi- ining the stomach contents of sacrificed bats. Moths cal (Morris et al. 2009). This separation is similar to the (Lepidoptera) and flies (Diptera) are commonly observed findings of the North American Bat Monitoring Program during winter in coastal North Carolina (Padgett and (Li and Kalcounis-Rueppell 2019). This bimodal distribu- Rose 1991). tion in North Carolina is also comparable with the Although southeastern myotis and Rafinesque’s big- species’ known range in South Carolina where a Blue eared bats rafinesquii in the southeastern Ridge Mountains population appears separated from a United States are known to roost in tree cavities during recently discovered population along South Carolina’s winter (Clement and Castleberry 2013; Fleming et al. Atlantic Coast (South Carolina Department of Natural 2013), this work provides the first documentation of Resources 2019). northern long-eared bats utilizing trees as winter roost All captures occurred 87 km from either the Atlantic sites. Most (84.2%) of the tracked bats selected live trees Ocean or its major sounds. This distance roughly over dead trees, but the species and size varied greatly. corresponds to the western boundary of the Middle Although the mean winter roost tree dbh was 30.9 cm, Atlantic Coastal Plain ecoregion within North Carolina. bats utilized trees as small as 4.1 cm. This may be Nearly all captures, winter roost sites, and maternity indicative of the relatively warmer winter temperatures roost sites occurred either within or very close to in coastal areas and therefore less need for thermal wetland forest. Northern long-eared bats were often protection. The winter roost emergence surveys revealed captured along with southeastern myotis, a species that the northern long-eared bats roosted in very small associated with swamps (Clement and Castleberry 2013; numbers, with the majority roosting alone. Fleming et al. 2013). The Middle Atlantic Coastal Plain Though northern long-eared bats were observed to be has a preponderance of swamps (Griffith et al. 2002b), active most of the winter, the use of temperature- and these wetland forests are the most intact and least sensitive transmitters during late fall–winter 2017–2018 disturbed habitat on the landscape. Thus, it is unclear if did document bats in torpor. Torpor is used to minimize northern long-eared bats are selecting for specific the thermoregulatory costs during periods of extreme qualities of wetland forests, or if they are simply utilizing conditions such as low ambient temperature or de- the largest intact forest tracts available regardless of creased food availability (Dunbar and Tomasi 2006). The hydrology. Nearly all northern long-eared bat captures in longest sustained torpor bout for each bat averaged 6.8 coastal North Carolina have occurred on large public d, and the lowest temperature at which a bat was lands, which indicates the importance of these conser- recorded to be normothermic was 3.08C. Since torpor vation lands. Moving westward from the Middle Atlantic duration is inversely related to ambient temperature Coastal Plain ecoregion into the Southeastern Plains (Dunbar and Tomasi 2006), these relatively short ecoregion, the dominant land use is agriculture, and the sustained torpor bouts are indicative of nighttime winter forest tracts are generally smaller, more fragmented, and temperatures commonly being greater than 3.08Cin more disturbed. Farther west, urbanization and agricul- coastal North Carolina. ture similarly affect forest tracts in the Piedmont Captures of juvenile northern long-eared bats begin- ecoregion. These land uses may contribute to the ning on June 16 and juvenile volancy occurring by 21 d seeming absence of northern long-eared bats in the (Krochmal and Sparks 2007) indicates that birth of pups Southeastern Plains and Piedmont ecoregions. begins no later than late May. However, weather and The documentation of extensive winter activity staff scheduling issues may have prevented earlier confirms a novel survival strategy as opposed to the documentation of juveniles. Additionally, it must be hibernation strategy that the species is assumed to use noted that these data were obtained from the northern

Journal of Fish and Wildlife Management | www.fwspubs.org December 2020 | Volume 11 | Issue 2 | 672 Northern Long-Eared Bats in Coastal North Carolina G.W. Jordan portion of the coastal plain in North Carolina. Kindel long-eared bat Myotis septentrionalis captures in eastern (2019) captured heavily pregnant northern long-eared North Carolina through March 2020 with location, sex, bats in late April and juveniles in early June in coastal age, forest type, and source. South Carolina, indicating birth occurring in early to mid- Found at DOI: https://doi.org/10.3996/JFWM-20-018.S1 May. Birth time in the Coastal Plain of North Carolina (27 KB XLSX). likely occurs along a continuum from north to south. Given this birth chronology, and due to the fact that no Table S2. Bat mist-netting level of effort within pregnant females were captured after June 1, I suggest a eastern North Carolina counties from all sources April conservative pup season for eastern North Carolina as 2015 to March 2020. The number of nights indicated are May 1–June 30. This is in contrast to the June 1–July 31 calendar nights, not net nights. A calendar night was pup season published in the Federal Register (2015). generally one netting crew at one site, but because of The species and size of maternity roost trees varied inconsistency in reporting across multiple sources, in greatly, but the mean maternity roost tree dbh (20.6 cm) some instances a calendar night may have represented was much smaller than the mean winter roost tree dbh multiple netting crews at different sites on the same date Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 (30.9 cm). Despite larger trees being available, some in the same county. Thus the number of calendar nights females selected trees as small as 6.1 cm. This may be should be considered minimum numbers. Netting effort indicative of the relatively warmer spring temperatures per calendar night (i.e., number of nets and number of in coastal areas and therefore less need for thermal hours that nets were open) varied greatly. The number of protection. The maternity roost tree emergence surveys calendar nights of netting in some counties was heavily indicate variability in roosting strategy. Although the biased because of certain productive sites being selected mean number of bats observed emerging at roost sites as focus areas for winter behavior or maternity studies. was 4.7, the median number was 1. However, notable Therefore, this data should not be used to determine outliers of 61 and 63 occurred. Since northern long-eared catch per unit effort. bats were often captured with the similar southeastern Found at DOI: https://doi.org/10.3996/JFWM-20-018.S2 myotis, this raises the question of possible coroosting (21 KB XLSX). with this species. This bears further study. Although WNS has decimated the western population Table S3. List of all northern long-eared bat Myotis of northern long-eared bats in North Carolina (Caldwell septentrionalis winter roost trees documented in eastern 2016), there is currently no evidence of WNS or P. North Carolina 2015–2018 with species, size (diameter at destructans in the Middle Atlantic Coastal Plain or breast height), status (live or dead), roost type, forest Southeastern Plains ecoregions of North Carolina, likely type, and emergence survey data. The numbers in the due to the rarity of caves and subsurface mines and emergence column indicate the number of bats ob- milder winters permitting bats to be active during served per emergence survey, with some winter roost winter. The discovery of a population of northern long- trees having multiple surveys. A blank cell in the eared bats that forgoes long-term hibernation is emergence column indicates that no emergence survey particularly important from a conservation standpoint. was conducted at that roost tree. Since this coastal population is not dependent upon Found at DOI: https://doi.org/10.3996/JFWM-20-018.S3 hibernation in caves or mines and exhibits only relatively (32 KB XLSX). short torpor durations, it is likely not susceptible to WNS- induced mortality. If the coastal population is truly Table S4. List of all northern long-eared bat Myotis spatially separated from the population in western North septentrionalis maternity roost trees documented in Carolina, the lack of contact between the two popula- eastern North Carolina in spring 2019 with species, size tions would further reduce the likelihood of the spread (diameter at breast height), status (live or dead), roost of P. destructans into coastal areas. Northern long-eared type, forest type, and emergence survey data. The bats were recently discovered in coastal South Carolina numbers in the emergence column indicate the number (White et al. 2018), an area with similar conditions to of bats observed per emergence survey. One emergence coastal North Carolina. With the species in sharp decline survey was conducted for each maternity roost tree. elsewhere from WNS, coastal North Carolina and South Found at DOI: https://doi.org/10.3996/JFWM-20-018.S4 Carolina could ultimately serve as a refugium for the (25 KB XLSX). species. However, the viability and population trends of this unique population of bats are currently unknown Reference S1. Caldwell K. 2016. White-nose syndrome and thus further study is warranted. surveillance and bat monitoring in North Carolina, October 1, 2015–September 30, 2016. Final performance Supplemental Material report under white-nose syndrome grant to states, grant number E-17. North Carolina Wildlife Resources Com- Please note: The Journal of Fish and Wildlife Manage- mission, Raleigh, North Carolina. ment is not responsible for the content or functionality of Found at DOI: https://doi.org/10.3996/JFWM-20-018.S5 any supplemental material. Queries should be directed (403 KB PDF). to the corresponding author for the article. Reference S2. Calyx Engineers þ Consultants. 2017a. Table S1. Chronological list of all known northern Federally protected bat survey report; eastern North

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Carolina; northern long-eared bat (Myotis septentrionalis) Department of Transportation and U.S. Fish and Wildlife research study; Beaufort, Bladen, Craven, New Hanover, Service, Raleigh, North Carolina. Pamlico, Pender, and Wayne Counties, North Carolina; Found at DOI: https://doi.org/10.3996/JFWM-20-018. survey phase 2: summer 2016. Report to North Carolina S12 (64.2 MB PDF). Department of Transportation, Raleigh, North Carolina. Found at DOI: https://doi.org/10.3996/JFWM-20-018.S6 Reference S9. Copperhead Environmental Consulting, (25.26 MB PDF). Inc. 2019. Final report: 2019 eastern North Carolina northern long-eared bat research project, phase VII, Dare Reference S3. Calyx Engineers þ Consultants. 2017b. County, North Carolina. Report to North Carolina Federally protected bat survey report; eastern North Department of Transportation and U.S. Fish and Wildlife Carolina; northern long-eared bat (Myotis septentrionalis) Service, Raleigh, North Carolina. research study; Brunswick, Columbus, Onslow, Pender, Found at DOI: https://doi.org/10.3996/JFWM-20-018. Robeson, Rockingham, and Sampson Counties, North S13 (25.15 MB PDF).

Carolina; survey phase 2: summer 2017. Report to North Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 Carolina Department of Transportation, Raleigh, North Reference S10. De La Cruz JL, True M, Taylor H, Brown Carolina. D, Ford WM. 2020. Occupancy and roost ecology of the Found at DOI: https://doi.org/10.3996/JFWM-20-018.S7 northern long-eared and Indiana bat on the coastal plain (87.63 MB PDF). of North Carolina. U.S. Geological Survey Science Support Program Project G17AC00288 to Virginia Tech, Blacks- Reference S4. Calyx Engineers þ Consultants. 2018a. burg. Report to U.S. Fish and Wildlife Service, Raleigh, Federally protected bat survey report; eastern North North Carolina. Carolina; northern long-eared bat (Myotis septentrionalis) Found at DOI: https://doi.org/10.3996/JFWM-20-018. research study; Camden, Currituck, and Gates Counties, S14 (15.01 MB PDF). North Carolina; survey phase v winter 2017/2018. Report to North Carolina Department of Transportation, Raleigh, Reference S11. Ecological Engineering. 2016. Protect- North Carolina. ed bat species survey report, northern long-eared bat Found at DOI: https://doi.org/10.3996/JFWM-20-018.S8 research project, eastern North Carolina, survey phase 1, (33.81 MB PDF). sessions 1 & 2: fall 2015 and winter 2016. Report to North Carolina Department of Transportation, Raleigh, North Reference S5. Calyx Engineers Consultants. 2018b. þ Carolina. Federally protected bat survey report; eastern North Found at DOI: https://doi.org/10.3996/JFWM-20-018. Carolina; northern long-eared bat (Myotis septentrionalis) S15 (274.97 MB PDF). research study; Edgecombe, Halifax, Hertford, and North- ampton Counties, North Carolina; spring/summer 2018 phase vi survey. Report to North Carolina Department of Reference S12. Ecological Engineering. 2018. Protect- Transportation, Raleigh, North Carolina. ed bat species survey report, northern long-eared bat Found at DOI: https://doi.org/10.3996/JFWM-20-018.S9 research project, eastern North Carolina, fall 2017 and (16.26 MB PDF). winter 2018, phase v. Report to North Carolina Depart- ment of Transportation, Raleigh, North Carolina. Reference S6. Calyx Engineers þ Consultants. 2019. Found at DOI: https://doi.org/10.3996/JFWM-20-018. Federally protected bat survey report; eastern North S16 (47.73 MB PDF). Carolina; northern long-eared bat (Myotis septentrionalis) research study; Gates County, North Carolina; survey Reference S13. Kalcounis-Rueppell MC, Li H. 2016. phase vii spring/summer 2019. Report to North Carolina Bats of the Uwharrie National Forest: mist-netting (2014 Department of Transportation, Raleigh, North Carolina. and 2015) and acoustic sampling (2013–2015). University Found at DOI: https://doi.org/10.3996/JFWM-20-018. of North Carolina at Greensboro. Report to North S10 (26.21 MB PDF). Carolina Wildlife Resources Commission, Raleigh, North Carolina. Reference S7. Copperhead Environmental Consulting, Found at DOI: https://doi.org/10.3996/JFWM-20-018. Inc. 2017. Final report: fall 2016/winter 2017 NCDOT S17 (48.03 MB PDF). northern long-eared bat research project, Bertie, Bladen, Hyde, Martin, and Tyrrell Counties, North Carolina. Report Reference S14. Lambiase SJ, Clark MK, Gatens LJ. to North Carolina Department of Transportation and U.S. 2000. Bat (chiroptera) inventory of North Carolina state Fish and Wildlife Service, Raleigh, North Carolina. parks 1999–2000. North Carolina Division of Parks and Found at DOI: https://doi.org/10.3996/JFWM-20-018. Recreation, Raleigh, North Carolina. Unpublished report. S11 (121.54 MB PDF). Found at DOI: https://doi.org/10.3996/JFWM-20-018. S18 (658 KB PDF). Reference S8. Copperhead Environmental Consulting, Inc. 2018. Final report: fall 2017/winter 2018 NCDOT Reference S15. Lambiase SJ, Clark MK, Gatens LJ. northern long-eared bat research project, Dare and Hyde 2005. Bat (chiroptera) inventory of North Carolina state Counties, North Carolina. Report to North Carolina parks, 2005 addendum. North Carolina Division of Parks

Journal of Fish and Wildlife Management | www.fwspubs.org December 2020 | Volume 11 | Issue 2 | 674 Northern Long-Eared Bats in Coastal North Carolina G.W. Jordan and Recreation, Raleigh, North Carolina. Unpublished Found at DOI: https://doi.org/10.3996/JFWM-20-018. report. S27 (12.76 MB PDF). Found at DOI: https://doi.org/10.3996/JFWM-20-018. Reference S24. VHB Engineering NC. 2019. Protected S19 (204 KB PDF). bat species survey report, northern long-eared bat research project, eastern North Carolina, spring and Reference S16. Li H, Kalcounis-Rueppell M. 2019. Final summer 2019, phase vii. Report to North Carolina report for WM-0321: acoustic monitoring of bats in North Department of Transportation, Raleigh, North Carolina. Carolina May 2017–October 2019. University of North Found at DOI: https://doi.org/10.3996/JFWM-20-018. Carolina at Greensboro. Report to North Carolina Wildlife S28 (212.9 MB PDF). Resources Commission, Raleigh, North Carolina. Found at DOI: https://doi.org/10.3996/JFWM-20-018. Acknowledgments S20 (1.49 MB PDF). The North Carolina Department of Transportation Reference S17. National Wildlife Health Center. 2019. provided funding for this work, and I thank the following Downloaded from http://meridian.allenpress.com/jfwm/article-pdf/11/2/665/2884157/i1944-687x-11-2-665.pdf by guest on 24 September 2021 Diagnostic services case report: case 29023. Madison, former and current staff for all their assistance with study Wisconsin. designs, contracts, and fieldwork: Heather Wallace, Mary Found at DOI: https://doi.org/10.3996/JFWM-20-018. Frazer, Chris Manley, Neil Medlin, and Cheryl Knepp. I S21 (26 KB PDF). thank all the staff from the following environmental consulting companies that did most of the fieldwork: Reference S18. Southeastern Cooperative Wildlife Ecological Engineering, Ecological Solutions, Three Oaks Disease Study. 2017. White-nose syndrome surveillance Engineering, Calyx Engineers þ Consultants, Copperhead diagnostic results. Case number CC17-204 A-NNN. Environmental Consulting, and VHB Engineering NC. I University of Georgia, Athens. Found at DOI: https://doi.org/10.3996/JFWM-20-018. thank Katherine Etchison (North Carolina Wildlife Re- S22 (146 MB PDF). sources Commission) and Jesse De La Cruz (Conservation Management Institute at Virginia Tech) for sharing their Reference S19. Tetra Tech, Inc. 2015. Northern long- data. I also thank all the volunteers who assisted me with eared bat survey report, NSAHR Northwest Annex, my mist-netting efforts. Valuable comments on my first Virginia and North Carolina. Report to Naval Facilities draft of this article were provided by Piper Roby, Theresa Engineering Command Mid-Atlantic, Norfolk, Virginia. Wetzel, Mary Frazer, Heather Wallace, and Kathryn Found at DOI: https://doi.org/10.3996/JFWM-20-018. Cunningham. The comments of the Associate Editor S23 (21.91 MB PDF). and two anonymous reviewers also helped me focus and substantially improve this article. Reference S20. Tetra Tech, Inc. 2016. Northern long- Any use of trade, product, website, or firm names in eared bat survey report, NSAHR Northwest Annex, this publication is for descriptive purposes only and does Virginia and North Carolina. Report to Naval Facilities not imply endorsement by the U.S. Government. Engineering Command Mid-Atlantic, Norfolk, Virginia. Found at DOI: https://doi.org/10.3996/JFWM-20-018. References S24 (26.72 MB PDF). Bernard RF, McCracken GF. 2017. Winter behavior of bats Reference S21. Three Oaks Engineering. 2016. North- and the progression of white-nose syndrome in the ern long-eared bat research project, eastern North southeastern United States. Ecology and Evolution Carolina, survey phase 2: summer 2016, northeast section 7:1487–1496. report. Report to North Carolina Department of Trans- Boyles JG, Dunbar MB, Whitaker JO. 2006. Activity portation, Raleigh, North Carolina. following arousal in winter in North American Found at DOI: https://doi.org/10.3996/JFWM-20-018. vespertilionid bats. Mammal Review 36:267–280. S25 (23.51 MB PDF). Caceres MC, Barclay RMR. 2000. Myotis septentrionalis. Reference S22. Three Oaks Engineering. 2017. North- Mammalian Species 634:1–4. ern long-eared bat research project, eastern North Caldwell K. 2016. White-nose syndrome surveillance and Carolina, summer 2017 survey phase iv, northeast section bat monitoring in North Carolina, October 1, 2015– report. Report to North Carolina Department of Trans- September 30, 2016. Final performance report under portation, Raleigh, North Carolina. white-nose syndrome grant to states, grant number E- Found at DOI: https://doi.org/10.3996/JFWM-20-018. 17. North Carolina Wildlife Resources Commission, S26 (30.46 MB PDF). Raleigh, North Carolina (see Supplemental Material, Reference S1). Reference S23. Three Oaks Engineering. 2018. North- Calyx Engineers þ Consultants. 2017a. Federally protect- ern long-eared bat research project, eastern North ed bat survey report; eastern North Carolina; northern Carolina, spring/summer 2018 survey phase vi, southeast long-eared bat (Myotis septentrionalis) research study; section report. Report to North Carolina Department of Beaufort, Bladen, Craven, New Hanover, Pamlico, Transportation, Raleigh, North Carolina. Pender, and Wayne Counties, North Carolina; survey

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phase 2: summer 2016. Report to North Carolina to North Carolina Department of Transportation and Department of Transportation, Raleigh, North Carolina U.S. Fish and Wildlife Service, Raleigh, North Carolina (see Supplemental Material, Reference S2). (see Supplemental Material, Reference S7). Calyx Engineers þ Consultants. 2017b. Federally protect- Copperhead Environmental Consulting, Inc. 2018. Final ed bat survey report; eastern North Carolina; northern report: fall 2017/winter 2018 NCDOT northern long- long-eared bat (Myotis septentrionalis) research study; eared bat research project, Dare and Hyde Counties, Brunswick, Columbus, Onslow, Pender, Robeson, North Carolina. Report to North Carolina Department Rockingham, and Sampson Counties, North Carolina; of Transportation and U.S. Fish and Wildlife Service, survey phase 2: summer 2017. Report to North Raleigh, North Carolina (see Supplemental Material, Carolina Department of Transportation, Raleigh, North Reference S8). Carolina (see Supplemental Material, Reference S3). Copperhead Environmental Consulting, Inc. 2019. Final Calyx Engineers þ Consultants. 2018a. Federally protect- report: 2019 eastern North Carolina northern long-

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