Regular issues
Special Issues

Urban Naturalist
    URNA Home
    Range and Scope
    Board of Editors
    Editorial Workflow
    Publication Charges

Other Eagle Hill Journals
    Northeastern Naturalist
    Southeastern Naturalist
    Caribbean Naturalist
    Neotropical Naturalist
    Eastern Paleontologist
    Journal of the North Atlantic
    Eastern Biologist

Eagle Hill Institute Home

Bats in the Bronx: Acoustic Monitoring of Bats in New York City
Kaitlyn L. Parkins, Michelle Mathios, Colleen McCann, and J. Alan Clark

Urban Naturalist, No. 10 (2016): 1–16

Full-text pdf complete with cover.


Site by Bennett Web & Design Co.
Urban Naturalist 1 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 22001166 URBAN NATURALIST No. 10N:1o–. 1160 Bats in the Bronx: Acoustic Monitoring of Bats in New York City Kaitlyn L. Parkins1,*, Michelle Mathios1, Colleen McCann2, and J. Alan Clark1 Abstract - Studies on urban bat ecology, mainly focusing on large-scale landscape features that affect biodiversity, have flourished in the last decade. In the present study, we investigated the presence of bats on a local scale in a highly urbanized area, Bronx Borough, in New York City (NYC), 1 of 2 North American megacities. We used passive and active monitoring of bat echolocation calls at 4 sites in the Bronx to determine which species were present and to examine seasonal variation in activity levels. We recorded 5 species of bats at our study sites: Eptesicus fuscus (Big Brown Bat), Lasiurus borealis (Eastern Red Bat), L. cinereus (Hoary Bat), Lasionycteris noctivagans (Silver-haired Bat), and Perimyotis subflavus (Tri-colored Bat). The majority of recorded activity was by Eastern Red Bat at all sites. Activity peaked in August and November during the migratory periods for Eastern Red Bat and Silver-haired Bat. During the winter months, we recorded activity by Eastern Red Bat, Silver-haired Bat, and Hoary Bat, and found that we detected greater bat activity on nights with higher maximum daily temperatures. Our study provides preliminary documentation of tree-bat migration through NYC and evidence of winter bat activity in NYC. Further acoustic and mist-net surveys will help us better understand the diversity and seasonal activity of bats in NYC. Introduction In recent years, the body of research on North American bats has grown substantially, and this interest in bat conservation is owed, in large part, to 2 major and relatively new causes of bat mortality: white-nose syndrome (WNS) and wind turbines. WNS has triggered mass mortality in some species of hibernating bats and continues to spread across North America (Cryan et al. 2013, Turner et al. 2011). Recent surveys of bat mortality due to collisions with turbines have raised concerns about wind development and its impact on migratory bat populations (Arnett and Baerwald 2013, Cryan and Barclay 2009). These increasing threats, often acting in concert with the pressures of habitat loss (Agosta 2002), roost disturbance (Agosta 2002), climate change (Sherwin et al. 2012), and urbanization (Weller 2009), have motivated efforts to understand their effect on bat populations. Here, we explore the basic ecology of urban bat populations. Landscapes are becoming increasingly urbanized (United Nations 2012) and for most wildlife, urbanization results in negative overall effects on species richness and diversity (Czech et al. 2000; McKinney 2002, 2008). For insectivorous bat assemblages, urbanization tends to result in both a reduction of species diversity 1Fordham University, 441 East Fordham Road, Bronx, NY 10458. 2Wildlife Conservation Society, 2300 Southern Boulevard, Bronx, NY 10460 *Corresponding author - Manuscript Editor: Mark Laska Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 2 and dominance of a few, open-adapted, generalist species (Avila-Flores and Fenton 2005, Coleman and Barclay 2012, Duchamp and Swihart 2008, Loeb et al. 2008, Luck et al. 2013, Threlfall et al. 2011, Ulrey et al. 2005). In some cases, however, urbanization can provide a heterogeneous environment that is beneficial to particular species (Gehrt and Chelsvig 2003, 2004). Hence, the effects of urbanization on bats are largely species and context specific. Because of the growing number of “megacities”—urban agglomerations with a population over 10 million—worldwide (United Nations 2014), a better understanding of the effects of various levels of urbanization on bat communities is needed. Most previous studies on this topic are broad-scale examinations of bats in an urbanized landscape. Here, we focus on local-scale recording of bats within New York City (NYC), the largest city in the US and second-largest in North America (US Census Bureau 2014). However, we are not aware of published studies of bats within NYC. Baseline data on bat activity and species occupancy is important, including data from a variety of locations, habitats, and seasons (Weller et al. 2009). Baseline surveys and monitoring of bats can also provide a basis for future research and are necessary for quantifying the impact of new threats as they arise. Before designing studies on the behavior and ecology of bats, species-occurrence data are needed. One of the most common methods of determining bat species’ distributions and activity patterns is acoustic monitoring. Insectivorous bats produce echolocation calls to navigate and locate prey. The ability to record and visualize those calls provides a non-intrusive method of monitoring bats. Bat species can generally be distinguished by their echolocation calls (O’Farrell 1997), and recordings of calls can be used to compare relative levels of bat activity (Hayes 2000). For this study, we used acoustic monitoring to gather baseline data on bat species presence and seasonal levels of activity at 4 relatively close sites in the Bronx, 1 of NYC’s 5 boroughs. We utilized range maps (Arnett and Baerwald 2013, Cryan and Barclay 2009, Harvey et al. 2011) and New York State Department of Environmental Conservation reports (Carl Herzog, New York State Department of Environmental Conservation, Albany, NY, pers. comm.) to predict presence of bat species in the Bronx. Based on these data, we determined that bat species possibly present in the Bronx included 9 species: Eptesicus fuscus (Beavois) (Big Brown Bat), Lasiurus borealis (Müller) (Eastern Red Bat), L. cinereus (Palisot de Beauvois) (Hoary Bat), Lasionycteris noctivagans (Le Conte) (Silver-haired Bat), Perimyotis subflavus F. Couvier (Tricolored Bat), Myotis lucifugus (Le Conte) (Little Brown Bat), M. septentrionalis (Trouessart) (Northern Long-eared Bat), M. lebeii (Audubon and Bachman) (Eastern Small-footed Bat), and M. sodalis Miller & Allen (Indiana Bat). Eastern Red Bat, Hoary Bat, and Silver-haired Bat are migratory, foliage-roosting species (tree bats). The 6 other species hibernate in caves during the winter and use tree cavities, buildings, or other structures as summer roosts (Harvey et al. 2011). We expected that Big Brown Bat would be the most commonly detected species, based on anecdotal reports and because, in many surveys, this species was the most common bat in North American urban centers (Agosta 2002, Damm et al. 2015, Duchamp and Swihart 2008, Everette et al. 2001, Loeb et al. 2008, Ulrey et al. 2005). Urban Naturalist 3 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 Most bat surveys are conducted during the summer; however, little is known about winter bat behavior even though this information could prove valuable in determining migration routes and overwintering strategies (Boyles et al. 2006). For this reason, we also explored bat activity in the Bronx during winter months, with the expectation of possibly detecting Eastern Red Bat. To date, Eastern Red Bat has not been detected in NYC in the winter months, but NYC is within the possible wintering range for this species (Cryan 2003, Sherwin et al. 2012), which is known to feed and switch roosts during winter and has been reported flying in locations with winter temperatures as low as -2 °C (Boyles et al. 2006, J ones et al. 2009). Methods Study site We conducted this study within the Bronx borough of New York, NY. NYC has a population of ~8.4 million people within 487 km2 (US Census Bureau 2014). We monitored 4 sites for bat activity: the Rose Hill Campus of Fordham University (Rose Hill), the New York Botanical Garden (NYBG), the Bronx Zoo, and Hughes Avenue, in the Belmont neighborhood of the Bronx (Fig. 1). The Fordham, NYBG, Figure 1. Map of the 5 boroughs of New York City and bat-survey sites in the Bronx, NY. Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 4 and Bronx Zoo sites were located within urban parks. These 3 parks are divided by 2 major thoroughfares, and NYBG and the Bronx Zoo are part of a larger contiguous green space called Bronx Park. The Hughes Avenue site is located near the parks but is in a commercial and residential area of dense buildings, high impervious surface, and little vegetation. Passive acoustic monitoring We recorded bat activity from 1 May 2012 to 15 February 2013 using stationary SongMeter SM2BAT+ (Wildlife Acoustics, Concord, MA) full-spectrum ultrasonic, acoustic-recording units (detectors) placed on the rooftop of buildings at each site. We placed detectors on rooftops to maximize their height and reduce the risk of vandalism. We used external SMX-US omni-directional microphones attached to the top of 2.7-m poles to minimize echolocation-bounce off hard surfaces and maximize the number and quality of calls recorded. We set detectors to automatically record calls continuously throughout the night (from civil twilight to civil twilight), set at a 192-kHz sample rate, 12-kHz digital high-pass filter, 18-dB trigger level, 36-dB gain, and the microphone bias off. These settings were those recommended for the SongMeter SM2BAT+ with minor alterations for use in an urban environment (J. Tyburec, Janet Tyburec Consulting, Tucson, AZ; pers. comm). We used a 2.0-sec trigger-window minimum and 8.0-sec maximum file length so that calls would be an appropriate length for the call-analysis software. We recorded calls in .wav format onto SD data cards and copied them to hard drives for later analysis. We changed data cards and batteries approximately every 2 weeks. Active acoustic monitoring We also conducted bi-weekly active surveys at each site beginning the week of 25 June 2012 and ending the week of 27 August 2012. Active surveys consisted of walking a set transect using an Echometer 3 (EM3) ultrasonic recording unit (Wildlife Acoustics, Concord, MA). Each active survey began at sunset and lasted one hour. We recorded bat passes from the active surveys onto SD data cards and then copied to hard drives for later analysis. Bat call analysis We used the software’s default settings and passed recordings from all sites through the SonoBat Batch Scrubber Utility 5.2 (DND Design, Arcata, CA) to remove the majority of files that did not contain bat passes. We defined a bat pass (or call) as a file with 2 or more pulses with each pass separated by 1 or more seconds (Kalcounis et al. 1999, Weller et al. 2009, White and Ghert 2001). Due to the large amount of high- and low-frequency ambient noise in our urban setting, we then visually inspected each file on a time-frequency spectrogram for the presence of bat echolocation calls in order to manually eliminate files that contained only non-bat noise. We examined bat passes for quality before species analysis, and we used only regular, non-fragmented search-phase calls from a single echolocating bat for Urban Naturalist 5 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 species analysis because low-quality, overlapping, and fragmented calls are likely to result in misidentification (Szewczak 2002, Weller et al. 2009). We used default settings in the SonoBat NNE 3.2.0 automated classifier to analyze passes. We set a required call quality of 0.80 and a decision threshold of 0.90, the default settings for Sonobat 3.2.0 (see Jameson and Willis 2014, Kalcounis-Ruppell et al. 2013). We considered a species confirmed when the Sonobat maximum-likelihood estimate was ≥0.90. A trained individual checked identifications using Sonobat NNE reference calls to confirm species presence at each site and to investigate any unusual identifications made by the automated classifier. The use of acoustic recordings to make interspecific comparisons is limited due to differences in the ability to record and correctly identify passes from each species (Hayes 2000, O’Farrell and Gannon 1999). To account for these differences and to compare relative activity between species, we used the Miller activity index (Miller 2001), the number of 1-minute time blocks in which a species is present at each site. We used all bat passes recorded, including those that we could not positively identify to the species level, to calculate an index of activity for each site and for each month. We quantified the activity index as the number of passes per recording night (Avila-Flores and Fenton 2005, Coleman and Barclay 2012, Duchamp and Swihart 2008, Gehrt and Chelsvig 2003, Loeb et al. 2008, Luck et al. 2013, Threlfall et al. 2011, Ulrey et al. 2005). The activity index does not provide an estimate of abundance of bats in an area (Gehrt and Chelsvig 2003, 2004; Hayes 2000) but is, instead, an estimate of the relative use of a site and can be used to make relative comparisons between sites (Hayes 2000, Parsons and Szewczak 2009). To compare activity levels on a temporal scale, we standardized the number of passes to recording hour, which controlled for the changes in nighttime length throughout the year. We obtained data on the duration of darkness in NYC on each recording night from the US Naval Observatory Astronomical Application Department online database ( We used only the data from active surveys for the confirmation of species’ presence at each site. Because of time and personnel limitations, we could be present at only 1 site each night and, hence, we did not use the active-recording data to compare activity levels between sites. We examined activity levels using calendar months as divisions. This approach allowed us to examine seasonality of bat activity in the same manner as Cryan (2003). We obtained daily minimum and maximum temperatures for the portion of the winter months during which we conducted surveys (1 December–15 February) from the National Oceanic and Atmospheric Administration Climate Data Online database ( We obtained temperature data recorded at Laguardia Airport, which is ~9.5 km from our survey sites. We expected that higher temperatures would be positively correlated with higher winter bat activity and that lower temperatures would be negatively correlated with lower winter bat activity; thus, we compared temperatures on days during which bat activity was and was not recorded using a one-tailed t-test. Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 6 Results Species presence We conducted passive acoustic surveys on 470 nights (5410 hours) between 1 May 2012 and 15 February 2013. The nights were not evenly distributed between the sites due to detector failures. We recorded for 59 nights at Hughes Avenue, 88 nights at the Bronx Zoo, 146 nights at NYBG, and 177 nights at Rose Hill. We conducted active acoustic surveys on 20 nights, 5 at each site, between 26 June and 1 September 2012. We recorded >6000 bat passes among the 4 sites (5708 passive, 348 active). Of the recorded passes, we classified 57% (56% passive; 59% active) into the Sonobat- designated high-frequency call clade, which for the northeastern US includes: Eastern Red Bat, Tri-colored Bat, and Myotis spp. We classified the remaining 43% (44% passive; 41% active) into the low-frequency call clade comprised of Big Brown Bat, Silver-haired Bat, and Hoary Bat. Species-level identification was possible for 49% of passes (49% of passive; 48% active). Overall, we confirmed 5 species: Eastern Red Bat, Silver-haired Bat, Big Brown Bat, Tri-colored Bat, and Hoary Bat. All species were present at all sites (Table 1). Sonobat analysis identified 2 passes as being from Little Brown Bat Table 1. Bat species confirmed as present in the Bronx, NY, from 1 May 2012–15 February 2013, using passive and active acoustic monitoring techniques. Site Survey method Species present Bronx Zoo Active Lasiurus borealis Passive Perimyotis subflavus L. borealis Eptesicus fuscus Lasionycteris noctivagans L. cinereus New York Botanical Garden Active L. borealis E. fuscus Passive P. subflavus L. borealis E. fuscus L. noctivagans L. cinereus Fordham University, Rose Hill Active L. borealis Passive P. subflavus L. borealis E. fuscus L. noctivagans L. cinereus Hughes Avenue Active None Passive P. subflavus L. borealis E. fuscus L. noctivagans L. cinereus Urban Naturalist 7 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 (1 each at the Bronx Zoo and NYBG); however, the Sonobat likelihood analysis for those recordings was 0.42, and we could not confirm using an acoustic key that either pass was from Little Brown Bat. In general, active surveys recorded fewer species than passive surveys (Table 1). Only Eastern Red Bat and Big Brown Bat were recorded during active surveys. In no case did active surveys result in the confirmation of a species that was not also detected on a passive recorder. Eastern Red Bat made up 62% of identified passes from active surveys, and we classified the remaining 38% as Big Brown Bat. We classified 47% of passive recordings across all sites as Eastern Red Bat, 24% as Big Brown Bat, 21% as Sliver-haired Bat, 6% as Tri-colored Bat, and 2% as Hoary Bat. Miller indices showed the same trends—Eastern Red Bat had the highest overall value, followed by Big Brown Bat (Table 2). Eastern Red Bat had the highest Miller index value at every site (Fig. 2). The highest relative Miller index value for Big Brown Bat occurred at NYBG. At the Bronx Zoo, Silver-haired Bat had nearly as high a Miller index value as Eastern Red Bat. Hoary Bat and Tri-colored Bat made up the smallest proportion of active minutes at all sites and were relatively evenly distributed acr oss the four sites. Activity levels The sites averaged 12 passes per night (n = 4, SD = 8.8). The Bronx Zoo had the highest activity level overall with a mean of 22.9 passes per night, and Hughes Avenue had the lowest with a mean of 4.7 passes per night. Rose Hill and NYBG had means of 15.3 and 5.0 passes per night, respectively . Bat activity peaked at all sites in August and October and was lowest in the winter months (Fig. 3); however, we recorded bat activity in all 9 months of this study. Species activity was not distributed evenly over the course of the sampling period (Fig. 4). From June through September, we recorded more activity by Eastern Red Bat than any other species, and activity for this species peaked in August. Table 2. Count of passes, estimated likelihood of presence, and Miller activity index values (the number of 1-minute periods during which a species is present) for each bat species recorded in the Bronx, NY, 1 May 2012–15 February 2013. MYLU = Myotis lucifugus, PESU = Perimyotis subflavus, EPFU = Eptesicus fuscus, LANO = Lasionycteris noctivagans, LABO = Lasiurus borealis, and LACI = Lasiurus cinereus. Number of passes = total (and percent) of all passes identified to species level Number of Survey type MYLU PESU EPFU LANO LABO LACI passes Passive Pass count 2 178 665 578 1318 62 2803 (49%) Estimated likelihood 0.42 1.00 1.00 1.00 1.00 0.99 Percent of total passes 0 6 24 21 47 2 Miller Index 0 175 571 529 1261 60 Active Pass count 0 5 102 5 166 0 278 (48%) Estimated likelihood 0.00 0.44 1.00 0.18 1.00 0.00 Percent of total passes 0 0 38 0 62 0 Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 8 Figure 2. Proportion of total activity using the Miller activity index (the number of 1-minute periods in which a bat is present) recorded 1 May 2012–15 February 2013 for each bat species at 4 sites in the Bronx, NY. Figure 3. Average bat activity (passes/hour) from passive acoustic-recording devices at 4 sites in the Bronx, NY from 1 May 2012–15 February 2013. NYBG = New York Botanical Garden. Urban Naturalist 9 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 Big Brown Bat activity also peaked in August. In October, there was a substantial increase in activity by Silver-haired Bat, and this species had higher activity levels than any other species in October and November. Hoary Bat and Tri-colored Bat activity levels were low overall. Winter activity We followed Cryan (2003) and considered December, January, and February as the winter months for bats in the US and Canada. In December, we recorded 11 passes (1 at NYBG, 10 at the Bronx Zoo), 3 in January (all at NYBG), and 2 in February (both at NYBG). All identifiable winter bat activity was from tree bats; we recorded passes by Eastern Red Bat and Silver-haired Bat in December, and Figure 4. (A) Bat activity (passes/hour) recorded on passive acoustic-recording devices at 4 sites in the Bronx, NY from 1 May 2012–15 February 2013. (B) Subset of data: 1 December 2012–15 February 2013. Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 10 Hoary Bat in February. The December recordings fell into both the low- and highfrequency clades (5 and 6 passes, respectively), while all January and February recordings were classified in the low-frequency clade. Daily high temperatures during the winter sampling dates ranged from -6.7 ºC to 16.1 ºC. The mean high temperature when bat activity was recorded (8.9 ºC) was higher than when bat activity was not recorded (5.6 ºC) (1-tailed t-test, P = 0.03). We found no relationship between bat activity and daily low temperature (1-tailed t-test, P = 0.19). Although the mean temperature was higher when bat activity was detected, we recorded bats on both high- and low-temperature days throughout the winter (Fig. 5). Discussion Species presence Of the 9 species that are expected to occur in New York State, we detected the following 5 species in the Bronx during our sampling period: Eastern Red Bat, Silver-haired Bat, Big Brown Bat, Tri-colored Bat, and Hoary Bat. Our prediction that the majority of bat recordings would be from Big Brown Bat was not supported. The majority of the calls we recorded during both active and passive surveys were from Eastern Red Bat, and this species had the highest Miller Index at every site. In fact, tree bats—Eastern Red Bat, Silver-haired Bat, and Hoary Bat—made up 70% of all passively recorded calls in this study. This result may be partially explained by the natural history of the species detected and the fact that 3 of the 4 sites were in parks. However, passes from Eastern Red Bat made up the majority (71%) of recorded sequences even at the least vegetated site (Hughes Avenue), while the largest proportion of Big Brown Bats was at a highly vegetated site Figure 5. Daily high temperatures for the location of bat recording sites in the Bronx, NY, during winter months (1 Dec 2012–15 Feb 2013). Days on which bat activity was recorded are indicated by “*”. The mean high temperature was higher when bat activity was recorded (8.9 ºC) than when bat activity was not recorded (5.6 ºC) (1-tailed t-test, P = 0.03). There was no relationship between bat activity and daily low temperature (1-tailed t-test: P = 0.19). Urban Naturalist 11 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 (NYBG). Our results are similar to those from surveys in locations with high densities of buildings in Manhattan, Queens, and the Bronx, where Eastern Red Bat comprised 66% of all passes recorded and tree bats compromised 95% of all passes recorded (Parkins and Clark 2015). Although other studies have documented the dominance of Big Brown Bat in urban areas (e.g., Damm et al. 2015, Loeb et al. 2008, Ulrey et al. 2005), the landscape of NYC may be particularly well suited for tree bats. Eastern Red Bat is a generalist insectivore (Clare et al. 2009) and adapted to flight in open areas, 2 traits that make it more likely to thrive in cities (Duchamp and Swihart 2008). Other studies found a relationship between tree/vegetation cover and Eastern Red Bat activity in urban environments (Dixon 2011, Li and Wilkins 2014). Since 2007, hundreds of thousands of trees have been planted in NYC as part of the PlaNYC campaign. Twenty-one percent of NYC was covered with tree canopy as of 2010, with the goal of reaching 30% by 2030 (McPhearson 2011, McPhearson et al. 2013). The increasing tree cover in NYC may be conducive to the presence of tree bats. Additionally, the relatively low proportion of Big Brown Bat recordings at Hughes Avenue, compared to higher proportions at the 3 park sites, may be indicative of Big Brown Bat roosting in buildings and flying to parks to forage; this species is known to have a large home range and to fly long distances to feed in higher-quality habitat (Everette et al. 2001). Little Brown Bat, which previously may have been common in NYC, was not documented in this study. White-nose syndrome (WNS) has greatly reduced northeastern Little Brown Bat populations (Turner et al. 2011); hence, it is not surprising that we did not record this species, particularly because NYC is close to the epicenter of the disease outbreak. Unfortunately, there are no published surveys of Little Brown Bat in NYC prior to the WNS outbreak to which we could compare activity patterns and species presence. Additionally, our lack of confirmation of Little Brown Bat at these 4 sites does not necessarily mean that the species was not present. Little Brown Bat is known to rely on water for foraging on aquatic insects, and only 1 of the 4 passive-recorder locations was near a source of non-moving water. In the future, recording near potential Little Brown Bat foraging habitat using both passive and active surveys to further explore this species’ status in NYC would be beneficial. Temporal activity Each bat species exhibited distinctive changes in activity level over the course of the sampling period. Big Brown Bat is a common resident in the NYC region, and summer colonies often occur in close proximity to humans due to this species’ habit of using buildings for roosting (Agosta 2002, Harvey et al. 2011). Our data also showed steady summer activity by this species. The peak in Big Brown Bat activity in August may indicate either increased foraging activity or movement prior to regional migration to winter hibernacula; this species is known to roost in buildings during the summer and migrate to less-developed areas for the winter (Neubaum et al. 2006). Tri-colored Bat activity was steady, but low, from June through October, suggesting that this species may be a low-density summer Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 12 resident in NYC. Activity by Hoary Bat, a long distance migrant, was consistently very low, which suggests it may also be a low-density summer resident or migrant passing through NYC. Hoary Bat was the only species recorded in February. Occurrence records and stable isotope analyses show a southern and coastal migration of this species during the autumn from inland summer ranges (Cryan 2003, Cryan et al. 2014). Most winter Lasiurus spp. (hairy-tailed bats) activity occurs below 40° N latitude in regions where winter temperatures do not often dip below freezing (Cryan and Veilleux 2007), and NYC is located on the northern edge of this region. Furthermore, Hoary Bats may be capable of roosting under leaves and surviving periods of freezing temperatures in the winter in a manner similar to Eastern Red Bat (Cryan et al. 2014); hence, we believe individuals of this species are winter ing in NYC. The peaks of bat activity in the late summer and fall, which are driven by increased activity by Eastern Red Bats and Silver-haired Bats, are particularly interesting. Eastern Red Bats disperse to wintering grounds south and east during the autumn, where they roost in trees and leaf litter (Cryan 2003, Cryan and Veilleux 2007) and remain somewhat active (see Boyles 2006). Summer activity by Eastern Red Bats is indicative of a potential summer population, but the increase in activity in July, with a peak in August and sharp decline in September, also suggests migratory movement through NYC. These data are consistent with similar patterns in Eastern Red Bat migratory activity found in other acoustic surveys conducted in the Midwest and East Coast (Johnson et al. 2010, Walters et al. 2006) and documented by Eastern Red Bat mortalities at wind farms (Arnett et al. 2008). A relatively large jump in Silver-haired Bat activity occurred later, in October, which is consistent with the timing of coastal migratory movement of these bats detected by Johnson et al. (2010) and, again, is suggestive of a pulse of bats migrating through NYC. The winter records of both of these species indicate that some individuals are likely also wintering in NYC. As expected, the average temperature on days we detected bats was higher than days when we did not; however, individual records showed more variation. Two of the days on which we recorded bats were the coldest of the sampling period, at -3.9 ºC and -6.7 ºC. These occurrences were at NYBG, which is a public garden, so it is possible reason that this low-temperature activity resulted from human disturbance at their winter roosts. Conclusions This study is the first to document the migratory movement of Eastern Red Bats and Silver-haired Bats through NYC. These species are likely much more prevalent in NYC than previously thought, and Eastern Red Bat is possibly more common than Big Brown Bat, which is contrary to the generally accepted status of bats in NYC. Little is known about tree-bat migration ecology, and our study contributes to the understanding of migration timing and winter ranges for these species. Migration is a critical life stage, during which bats are susceptible to several threats (Cryan and Veilleux 2007). Results of further research on migratory movement of Urban Naturalist 13 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 tree bats through urban landscapes should help inform management decisions and conservation planning for these species. Although our survey added ecological information to our knowledge of bats in NYC, additional surveys are needed. Continued year-round recording should be combined with mist-net surveys, harp trapping, and active recording in a variety of landscapes to gain a fuller picture of the bat species assemblage in NYC. Also, surveys should be expanded to other boroughs and take place in areas of both high and low impervious-surface and building density, as well as in parks of various sizes in order to more thoroughly understand the habitat selection by bats in a North American megacity. Acknowledgments We thank those who allowed access to our study sites: Jessica Arcate-Schuler and the New York Botanical Garden, LAL properties, Inc., and Fordham College at Rose Hill. Adele Heib assisted with call analyses. M. Mathios and J.A. Clark were supported by a Fordham College at Rose Hill Faculty Undergraduate Research Grant. Two anonymous reviewers provided comments that improved this manuscript. Literature Cited Agosta, S.J. 2002. Habitat use, diet, and roost selection by the Big Brown Bat (Eptesicus fuscus) in North America: A case for conserving an abundant species. Mammal Review 32:179–198. Arnett, E.B., and E.F. Baerwald. 2013. Impacts of wind-energy development on bats: Implications for conservation. Pp. 435–456, In R.S. Adams and S.C. Pedersen (Eds.). Bat Evolution, Ecology and Conservation. Spring Science and Business Media, New York, NY. 640 pp. Arnett, E.B., W.K. Brown, W.P. Erickson, J.K. Fiedler, B.L. Hamilton, T.H. Henry, A. Jain, G.D. Johnson, J. Kerns, R.R. Koford, C.P. Nicholson, T.J. O’Connell, M.D. Piorkowski, and R.D. Tankersley Jr. 2008. Patterns of bat fatalities at wind-energy facilities in North America. Journal of Wildlife Management 72:61–78. Avila-Flores, R., and M.B. Fenton. 2005. Use of spatial features by foraging insectivorous bats in a large urban landscape. Journal of Mammalogy 86:1193–1204. Boyles, J.G., M.B. Dunbar, and J.O. Whitaker. 2006. Activity following arousal in winter in North American vespertilionid bats. Mammal Review 36:267–280. Clare, E.L., E.E. Fraser, H.E. Braid, M.B. Fenton, and P.D.N. Hebert. 2009. Species on the menu of a generalist predator, the Eastern Red Bat (Lasiurus borealis): Using a molecular approach to detect arthropod prey. Molecular Ecology 18:2532–2542. Coleman, J.L., and R.M.R. Barclay. 2012. Urbanization and the abundance and diversity of prairie bats. Urban Ecosystems 15:87–102. Cryan, P.M. 2003. Seasonal distribution of migratory tree bats (Lasiurus and Lasionycteris) in North America. Journal of Mammalogy 84:579–593. Cryan, P.M., and R.M.R. Barclay. 2009. Causes of bat fatalities at wind turbines: Hypotheses and predictions. Journal of Mammalogy 90:1330–1340. Cryan, P.M., and J.P. Veilleux. 2007. Migration and the use of autumn, winter, and spring roosts by tree bats. Pp. 153–175, In M.J. Lacki, J.P. Hayes, and A. Kurta (Eds.). Bats in Forests. The Johns Hopkins University Press, Baltimore, MD. 368 pp. Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 14 Cryan, P.M., C.U. Meteyer, J.G. Boyles, and D.S. Blehert. 2013. White-nose syndrome in bats: Illuminating the darkness. BMC Biology 11:1–4. Cryan, P.M., C.A. Stricker, and M.B. Wunder. 2014. Continental-scale, seasonal movements of a heterothermic migratory tree bat. Ecological Applications 24:602–616. Czech, B., P. Krausman, and P. Devers. 2000. Economic associations among causes of species endangerment in the United States. BioScience 50:593–601. Damm, J.P., D.W. Sparks, and J.O. Whitaker Jr. 2015. Bat-species diversity at an urban– rural interface: Dominance by one species in an urban area. Urb an Naturalist 8:1–13. Dixon, M.D. 2011. Relationship between land cover and insectivorous bat activity in an urban landscape. Urban Ecosystems 15:683–695. Duchamp, J.E., and R.K. Swihart. 2008. Shifts in bat-community structure related to evolved traits and features of human-altered landscapes. Landscape Ecol ogy 23:849–860. Everette, A.L., T.J. O’Shea, L.E. Ellison, L.A. Stone, and J.L. McCance. 2001. Bat use of a high-plains urban wildlife refuge. Wildlife Society Bulletin 29:967–973. Gehrt, S.D., and J.E. Chelsvig. 2003. Bat activity in an urban landscape: Patterns at the landscape and microhabitat scale. Ecological Applications 13:939–950. Gehrt, S.D., and J.E. Chelsvig. 2004. Species-specific patterns of bat activity in an urban landscape. Ecological Applications 14:1–12. Harvey, M.J., J.S. Altenbach, and T.L. Best. 2011. Bats of the United States and Canada. The Johns Hopkins University Press, Baltimore, MD. 202 pp. Hayes, J.P. 2000. Assumptions and practical considerations in the design and interpretation of echolocation-monitoring studies. Acta Chiropterologica 2:225–236. Jameson, J.W., and C.K.R. Willis. 2014. Activity of tree bats at anthropogenic tall structures: Implications for mortality of bats at wind turbines. Animal Behaviour 97:145–152. Johnson, J.B., J.E. Gates, and N.P. Zegre. 2010. Monitoring seasonal bat activity on a coastal barrier island in Maryland, USA. Environmental Monitoring and Assessment 173:685–699. Jones, G., D.S. Jacobs, T.H. Kunz, M.R. Willig, and P.A. Racey. 2009. Carpe noctem: The importance of bats as bioindicators. Endangered Species Researc h 8:93–115. Kalcounis, M.C., K.A. Hobson, R.M. Brigham, and K.R. Hecker. 1999. Bat activity in the boreal forest: Importance of stand type and vertical strata. Journal of Mammalogy 80:673–682. Kalcounis-Ruppell, M.C., K.M. Briones, J.A. Homyack, R. Petric, M.M. Marshall, and D.A. Miller. 2013. Hard forest-edges act as conduits, not filters, for bats. Wildlife Society Bulletin 37:571–576. Li, H., and K.T. Wilkins. 2014. Patch or mosaic: Bat activity responds to fine-scale urban heterogeneity in a medium-sized city in the United States. Urban Ecosystems 17:1013–1031. Loeb, S.C., C.J. Post, and S.T. Hall. 2008. Relationship between urbanization and batcommunity structure in national parks of the southeastern US. Urban Ecosystems 12:197–214. Luck, G.W., L. Smallbone, C. Threlfall, and B. Law. 2013. Patterns in bat functional guilds across multiple urban centres in south-eastern Australia. Landscape Ecology 28:455–469. McKinney, M.L. 2002. Urbanization, biodiversity, and conservation. BioScience 52:883–890. McKinney, M.L. 2008. Effects of urbanization on species richness: A review of plants and animals. Urban Ecosystems 11:161–176. Urban Naturalist 15 K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 McPhearson, P.T. 2011. Toward a sustainable New York City: Greening through urban forest restoration. Pp. 181–203, In Sustainability in America’s Cities. Island Press/Center for Resource Economics, Washington, DC. 304 pp. McPhearson, T., D. Maddox, B. Gunther, and D. Bragdon. 2013. Local assessment of New York City: Biodiversity, green space, and ecosystem services. Pp. 355–383, In T. Elmqvist, M. Fragkias, J. Goodness, B. Güneralp, P.J. Marcotullio, R.I. McDonald, S. Parnell, M. Schewenius, M. Sendstad, K.C. Seto, and C. Wilkinson (Eds.). Urbanization, Biodiversity and Ecosystem Services: Challenges and Opportunities: A Global Assessment. Springer Netherlands, Dordrecht, Netherlands. 755 p p. Miller, B.W. 2001. A new method for determining relative activity of free-flying bats using a new activity index for acoustic monitoring. Acta Chirpoterologica 3:93–105. Neubaum, D.J., T.J. O’Shea, and K.R. Wilson. 2006. Autumn migration and selection of rock crevices as hibernacula by Big Brown Bats in Colorado. Journal of Mammalogy 87:470–479. O’Farrell, M.J. 1997. Use of echolocation calls for the identification of free-flying bats. Transactions of the Western Section of the Wildlife Society 33:1–8. O’Farrell, M.J., and W.L. Gannon. 1999. A comparison of acoustic versus capture techniques for the inventory of bats. Journal of Mammalogy 80:24–30 . Parkins, K.L., and J.A. Clark. Green roofs provide habitat for urban bats. 2015. Global Conservation and Ecology 4:349–357. Parsons, S., and J.M. Szewczak. 2009. Detecting, recording, and analyzing the vocalizations of bats. Pp. 91–111, In T.H. Kunz and S. Parsons (Eds.). Ecological and Behavioral Methods for the Study of Bats. The Johns Hopkins University Press, Baltimore, MD. 901 pp. Sherwin, H.A., W.I. Montgomery, and M.G. Lundy. 2012. The impact and implications of climate change for bats. Mammal Review 43:171–182. Szewczak, J.M. 2002. Advanced analysis techniques for identifying bat species. Pp. 121–126, In R.M. Brigham, E.K.V. Kalko, G. Jones, S. Parsons, and H.J.G.A. Limpens (Eds.). Bat Echolocation Research: Tools, Techniques, Analysis. Bat Conservation International, Austin, TX. 174 pp. Threlfall, C., B. Law, T. Penman, and P.B. Banks. 2011. Ecological processes in urban landscapes: Mechanisms influencing the distribution and activity of insectivorous bats. Ecography 34:814–826. Turner, G.G., D.M. Reeder, and J.T.H. Coleman. 2011. A five-year assessment of mortality and geographic spread of white-nose syndrome in North American bats and a look to the future. Bat Research News 52:13–27. Ulrey, W.A., D.W. Sparks, and C.M. Ritzi. 2005. Bat communities in highly impacted areas: Comparing Camp Atterbury to the Indianapolis airport. Proceedings of the Indiana Academy of Science 114:73–76. United Nations. 2012. World Urbanization Prospects: The 2011 revision. United Nations, Department of Economic and Social Affairs Population Division, New York, NY. 318 pp. United Nations. 2014. World urbanization prospects: The 2014 revision, highlights. United Nations, Department of Economic and Social Affairs Population Division, New York, NY. 32 pp. US Census Bureau. 2014. Annual estimates of the resident population: 1 April 2010 to 1 July 2014—Metropolitan Statistical Area. Available online at http://factfinder.census. gov. Accessed 15 May 2015. Urban Naturalist K.L. Parkins, M. Mathios, C. McCann, and J.A. Clark 2016 No. 10 16 Walters, B.L., D.W. Sparks, J.O. Whitaker, and C.M. Ritzi. 2006. Timing of migration by Eastern Red Bats (Lasiurus borealis) through Central Indiana. Acta Chiropterologica 8:259–263. Weller, T.J., P.M. Cryan, and T.J. O’Shea. 2009. Broadening the focus of bat conservation and research in the USA for the 21st century. Endangered Species Research 8:129–145. White, E.P., and S.D. Ghert. 2001. Effects of recording media on echolocation data from broadband bat detectors. Wildlife Society Bulletin 29:974–978.