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Urban Roosts: Use of Buildings by Florida Bonneted Bats

Elysia N. Webb1, Holly K. Ober1,*, Elizabeth C. Braun de Torrez1,2,
Jeffery A. Gore3, and Ricardo Zambrano4

1Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL 32611 USA. 2Florida Fish and Wildlife Conservation Commission, Gainesville, FL 32601 USA. 3Florida Fish and Wildlife Conservation Commission, Panama City, FL 32409 USA. 4Florida Fish and Wildlife Conservation Commission, West Palm Beach, FL 33412 USA. *Corresponding author.

Urban Naturalist, No. 42 (2021)

Abstract
Florida Bonneted Bats, Eumops floridanus, were first documented in Miami, Florida, USA, in the 1930s. We summarized reports of these bats in the greater Miami area throughout the past 80 years and documented new roosts by radio-tracking bats captured in semi-natural areas. Florida Bonneted Bats in Miami consistently roosted in buildings, in contrast to other portions of the species’ range where they use trees and bat houses. Throughout the past 60 years, reports of building use have been confined to a small (40 km2) portion of the city. Bats regularly selected buildings with architectural similarities (Mediterranean Revival style; characterized by stucco exteriors, open chimneys with integrated arch covers, and clay tile roofs). To ensure adequate conservation measures are taken to minimize harm to this federally endangered species in urban areas, we outline four topics in need of additional research and suggest four topics that should be covered through targeted educational campaigns.

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No. 42 Urban Naturalist 2021 Urban Roosts: Use of Buildings by Florida Bonneted Bats Elysia N. Webb, Holly K. Ober, Elizabeth C. Braun de Torrez, Jeffery A. Gore, and Ricardo Zambrano Urban Naturalist The Urban Naturalist (ISSN # 2328-8965) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680- 0009. Phone 207-546-2821 Ext. 4, FAX 207-546-3042. E-mail: office@eaglehill.us. Webpage: http://www.eaglehill.us/urna. Copyright © 2021, all rights reserved. Published on an article by article basis. Special issue proposals are welcome. The Urban Naturalist is an open access journal. Authors: Submission guidelines are available at http://www.eaglehill.us/urna. Co-published journals: The Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, and Eastern Paleontologist, each with a separate Board of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Federal ID # 010379899). Board of Editors Myla Aronson, Rutgers University, New Brunswick, NJ, USA Joscha Beninde, University of California at Los Angeles, CA, USA ... Co-Editor Sabina Caula, Universidad de Carabobo, Naguanagua, Venezuela Sylvio Codella, Kean University, Union New Jersey, USA Julie Craves, University of Michigan-Dearborn, Dearborn, MI, USA Ana Faggi, Universidad de Flores/CONICET, Buenos Aires, Argentina Leonie Fischer, Technical University of Berlin, Berlin, Germany Chad Johnson, Arizona State University, Glendale, AZ, USA Kirsten Jung, University of Ulm, Ulm, Germany Erik Kiviat, Hudsonia, Bard College, Annandale-on- Hudson, NY, USA Sonja Knapp, Helmholtz Centre for Environmental Research–UFZ, Halle (Saale), Germany David Krauss, City University of New York, New York, NY, USA Mark Laska, Great Ecology, consulting, La Jolla, CA, USA Zdenka Lososova, Masaryk University, Brno, Czechia Joerg-Henner Lotze, Eagle Hill Institute, Steuben, ME ... Publisher Kristi MacDonald, Hudsonia, Bard College, Annandale-on- Hudson, NY, USA Ian MacGregor-Fors, Insituto de Ecología Mexico, Veracruz, Mexico ... Co-Editor Tibor Magura, University of Debrecen, Debrecen, Hungary Brooke Maslo, Rutgers University, New Brunswick, NJ, USA Mark McDonnell, Royal Botanic Gardens Victoria and University of Melbourne, Melbourne, Australia Mike McKinney, University of Tennessee, Knoxville, TN, USA Desirée Narango, City University of New York, New York, NY, USA Mitchell Pavao-Zuckerman, University of Arizona, Tucson, Arizona, USA Joseph Rachlin, Lehman College, City University of New York, New York, NY, USA Travis Ryan, Center for Urban Ecology, Butler University, Indianapolis, IN, USA Michael Strohbach, Technische Universität Braunschweig, Institute of Geoecology, Braunschweig, Germany Katalin Szlavecz, Johns Hopkins University, Baltimore, MD, USA Paige Warren, University of Massachusetts, Amherst, MA, USA Alan Yeakley, Portland State University, Portland, OR, USA Iriana Zuria, Universidad Autónoma del Estado de Hidalgo, Hidalgo, Mexico ♦ The Urban Naturalist is a peer-reviewed and edited interdisciplinary natural history journal with a global focus on urban areas (ISSN 2328- 8965 [online]). ♦ The journal features research articles, notes, and research summaries on terrestrial, freshwater, and marine organisms and their habitats. ♦ It offers article-by-article online publication for prompt distribution to a global audience. ♦ It offers authors the option of publishing large files such as data tables, and audio and video clips as online supplemental files. ♦ Special issues - The Urban Naturalist welcomes proposals for special issues that are based on conference proceedings or on a series of invitational articles. Special issue editors can rely on the publisher’s years of experiences in efficiently handling most details relating to the publication of special issues. ♦ Indexing - The Urban Naturalist is a young journal whose indexing at this time is by way of author entries in Google Scholar and Researchgate. Its indexing coverage is expected to become comparable to that of the Institute's first 3 journals (Northeastern Naturalist, Southeastern Naturalist, and Journal of the North Atlantic). These 3 journals are included in full-text in BioOne.org and JSTOR.org and are indexed in Web of Science (clarivate.com) and EBSCO.com. ♦ The journal's staff is pleased to discuss ideas for manuscripts and to assist during all stages of manuscript preparation. The journal has a page charge to help defray a portion of the costs of publishing manuscripts. Instructions for Authors are available online on the journal’s website (http://www.eaglehill.us/urna). ♦ It is co-published with the Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, Eastern Paleontologist, Eastern Biologist, and Journal of the North Atlantic. ♦ It is available online in full-text version on the journal's website (http://www.eaglehill.us/urna). Arrangements for inclusion in other databases are being pursued. Cover Photograph: A Florida Bonneted Bat (Eumops floridanus) captured during this study. Dade County, Florida. Photograph © Elysia Webb. Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 1 2021 Urban Naturalist 42:1–11 Urban Roosts: Use of Buildings by Florida Bonneted Bats Elysia N. Webb1, Holly K. Ober1,*, Elizabeth C. Braun de Torrez1,2, Jeffery A. Gore3, and Ricardo Zambrano4 Abstract - Florida Bonneted Bats, Eumops floridanus, were first documented in Miami, Florida, USA, in the 1930s. We summarized reports of these bats in the greater Miami area throughout the past 80 years and documented new roosts by radio-tracking bats captured in semi-natural areas. Florida Bonneted Bats in Miami consistently roosted in buildings, in contrast to other portions of the species’ range where they use trees and bat houses. Throughout the past 60 years, reports of building use have been confined to a small (40 km2) portion of the city. Bats regularly selected buildings with architectural similarities (Mediterranean Revival style; characterized by stucco exteriors, open chimneys with integrated arch covers, and clay tile roofs). To ensure adequate conservation measures are taken to minimize harm to this federally endangered species in urban areas, we outline four topics in need of additional research and suggest four topics that should be covered through targeted educational campaigns. Introduction Eumops floridanus (Allen 1932) (Florida Bonneted Bat) is a federally endangered species endemic to southern and central Florida, with a geographic distribution believed to be among the smallest of all bat species in the New World (FWC 2011, Timm and Genoways 2004). The US Fish and Wildlife Service (USFWS) listed Florida Bonneted Bats as federally endangered in 2013, citing concerns about habitat loss, degradation, and modification from human population growth as well as the species’ small population size, slow reproduction, low fecundity, and relative isolation (USFWS 2013). The ongoing loss and modification of natural areas is of great concern to the future wellbeing of these bats, given the small size of their geographic range and the likelihood of climate change causing coastal portions of the current range to become unsuitable (USFWS 2013). Location and characterization of structures used as roosts for the Florida Bonneted Bats has been identified as a key research priority of both the USFWS (USFWS 2013) and the Florida Fish and Wildlife Conservation Commission (FWC) (FWC 2013). Most recent efforts to identify roosts have occurred in natural areas, where the species has been documented using Pinus palustris Mill and Pinus elliottii Engelm, Roystonea spp. (Royal Palm Trees), and Taxodium distichum (L.) Rich, as well as artificial structures, including bat houses and a utility pole (Angell and Thompson 2015; Belwood 1992; Braun de Torrez et al. 2016, 2020 unpubl. data; Timm and Genoways 2004; Webb 2017 unpubl. data). Although decades-old records indicate the species resided in buildings in the greater Miami area (Barbour and Davis 1969, Gore et al. 2015, Owre 1978, Robson et al. 1989, Timm and Genoways 2004), little effort has been made to characterize the types of structures used by bats or changes in use over time as urbanization has intensified. Guidelines recently established by the USFWS to minimize potential negative im- 1Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL 32611 USA. 2Florida Fish and Wildlife Conservation Commission, Gainesville, FL 32601 USA. 3Florida Fish and Wildlife Conservation Commission, Panama City, FL 32409 USA. 4Florida Fish and Wildlife Conservation Commission, West Palm Beach, FL 33412 USA. *Corresponding author: holly.ober@ufl.edu. Manuscript Editor: Brooke Maslo Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 2 pacts to existing habitat for these bats (Florida Bonneted Bat Consultation Guidelines, available at https://www.fws.gov/verobeach/ListedSpeciesMammals.html) delineate not only the region where the species is presumed to occur (the “Consultation Area”) but also the urban development boundary (the “South Florida Urban Bat Area”), where the species is suspected to rely largely on artificial structures as roosts (Fig. 1A). Given that a substantial portion of their geographic range is undergoing urbanization, there is an urgent need to better understand use of urban environments by this endangered species. Our objective was to increase understanding of Florida Bonneted Bat roost use in the most intensely urbanized region within their geographic range. Specifically, we identified locations where these bats roosted in the greater Miami area and evaluated how the location and type of roost structures has changed over time. Because the current Florida Bonneted Bat Consultation Guidelines do not include recommendations for urban environments, this synthesis of data from the greater Miami area can provide insight that directly informs conservation and management recommendations for these bats in this region as well as other urbanizing areas across the species range. Figure 1. Maps showing (A) the location of Florida within the USA, the region designated by the US Fish and Wildlife Service (USFWS) as the currently accepted geographic range for Florida Bonneted Bats, Eumops floridanus (“Consultation Area”), and the location of our study region, designated by USFWS as the “South Florida Urban Bat Area”; (B) the location of 2017 Florida Bonneted Bat (FBB) capture attempts within the “South Florida Urban Bat Area” and historical and recent records of buildings occupied by Florida Bonneted Bats and; and (C) the area of most intense use over time by Florida Bonneted Bats in Coral Gables (Source: Esri, DigitalGlobe, GeoEye, Earthstar Geographics, CNES/Airbus DS, USDA, USGS, AeroGRID, IGN, and the GIS User Community). Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 3 Materials and Methods Historical Roosts We used two approaches to compile a comprehensive list of all known roosts used by Florida Bonneted Bats in the greater Miami area prior to the species’ listing as federally endangered in 2013, which we refer to as the historical period. First, we completed a thorough literature review of all reports using the Web of Science (Thomson Reuters) database. Because the species name has changed over time from Eumops glaucinus to Eumops glaucinus floridanus to Eumops floridanus, we used each of these three scientific names as keywords in our search as well as "Florida Bonneted Bat”, “Florida Mastiff Bat”, “Wagner’s Mastiff Bat”, OR “Florida” AND “bat”. The literature search included publications between 1900 and 15 August 2020. We then searched the reference lists of these publications for additional relevant sources. Second, we contacted the four museums that hold E. floridanus or E. glaucinus floridanus specimens (American Museum of Natural History in New York, NY; Field Museum of Natural History in Chicago, IL; Florida Museum of Natural History in Gainesville, FL; and Kansas University Natural History Museum in Lawrence, KS) and requested details on the location where each specimen was obtained. From each published report and each museum specimen, we recorded information on roost location, and for those reported in buildings, we used publicly available information from commercial realty websites to determine the year built and architecture type. Recent Roosts We also used two approaches to document structures used as roosts by Florida Bonneted Bats in Miami recently, which we define as the period starting when the species was listed as endangered in 2013 until 2020. First, we compiled a list of all reports to FWC, USFWS, and University of Florida (UF) during this period. Reports included written, oral, and electronic communication. Second, we captured bats and radio-tracked them to roost sites. We selected four semi-natural areas evenly distributed across the greater Miami area where Florida Bonneted Bats had been detected acoustically (Marks and Marks 2012; F. Ridgley, Zoo Miami, Miami, FL, 2014 pers. comm.) to serve as potential bat capture locations. The southernmost area was Zoo Miami, located in Miami; the northernmost and easternmost locations were the Granada Golf Course and Fairchild Tropical Botanic Garden in Coral Gables; and the remaining site was Kendall Indian Hammocks Park in Kendall (Fig. 1B). We attempted to capture bats in each area during June and July 2017 by setting up triple-high stacked mist nets on 7.3-m poles (Bat Conservation and Management, Carlisle, PA) at two locations within Zoo Miami, three locations within the Granada Golf Course, two locations within Fairchild Tropical Botanic Garden, and two locations within Kendall Indian Hammocks Park. Mist nets were opened shortly before sunset, closed at 0200 h, and checked every 10 minutes. Short-duration rain events were common many evenings; we paused capture efforts whenever precipitation began and resumed once it ended. Adjacent to each net, we situated an acoustic lure (BatLure™, Apodemus Field Equipment, Mheer, Netherlands) to attract bats to the net. The lure broadcast Florida Bonneted Bat social calls pre-recorded from a roost in Florida Panther National Wildlife Refuge, which is a tactic demonstrated to markedly increase the capture rate of this species in mist nets (Braun de Torrez et al. 2017). We netted for 26 nights: 18 nights at Granada Golf Course, two nights at Fairchild Tropical Botanic Garden, three nights at Zoo Miami, and three nights at Kendall Indian Hammocks Park. Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 4 We recorded age, sex, reproductive status, mass, and forearm length for each captured individual. Age was categorized as adult or juvenile, determined by the degree of fusion of epiphyseal cartilages in the fourth metacarpal-phalangeal joint (Kunz and Anthony 1982). Reproductive status for females was categorized as nonreproductive, pregnant, lactating, or postlactating through palpation of the abdomen, presence of swollen nipples or milk. Reproductive status for males was categorized as nonreproductive or reproductive through the presence of abdominal or descended testes. We calculated body condition index (mass/ forearm length) of each individual to serve as a relative measure of bat condition (Jonasson and Willis 2011, Reynolds et al. 2009) that could be compared with reports for Florida Bonneted Bats roosting in a natural area (reported in Ober et al. 2017b). We attached VHF radio-transmitters (model LB-2, Holohil Systems Ltd., Carp, Ontario, Canada) to all Florida Bonneted Bats captured, with the exception of juveniles, pregnant or lactating female adults, or any bat for which the mass of the transmitter would exceed 5% of the bat’s mass (Aldridge and Brigham 1988), in accordance with our permit (TE23583B-2) and protocol approved by University of Florida IACUC (#201609497). Each radio-transmitter was sewed onto a break-away collar closed with absorbable suture, and each individual was released at the site of capture immediately after collar attachment. Each bat to which a transmitter was attached was tracked by vehicle and on foot during subsequent days until a structure used as a roost was identified. Each roost structure identified was verified with emergence observations after sunset that included colony size counts. Because Florida Bonneted Bats typically emerge from their roosts long after sunset, and because of the challenges associated with observing the emergence of bats from buildings in residential areas with high housing densities, it was generally not possible to discern what portion of each building bats were emerging from (e.g., beneath roof tiles, from chimney). Lastly, we recorded information on the architecture of each building used as a roost and the year it was built. Results Historical Roosts Publications and museum records confirmed that Eumops specimens were collected regularly in buildings in Miami during the 3-decade period extending from the mid-1930s to the mid-1960s (Table 1; Figs. 1B, 1C). The first record of Eumops in Miami indicated a bat was found by a student from “Edison High School” (Barbour 1936), likely Miami Edison Senior High School located in Miami, but no indication of the location where the bat was originally found was provided, so it is not included further in our analyses. Schwartz (1952) subsequently described 5 male specimens recovered from the gymnasium and grounds of “Miami High School” during the preceding 10 years (i.e., 1942–1952). As the oldest high school in the county, Miami High School (i.e., Miami Senior High School) has been in its current location since 1928 and is noted for its Mediterranean Revival architecture. Jennings (1958) reported that 20 Florida Bonneted Bats had been collected from Coral Gables, Coconut Grove, and Miami since the first report by Barbour in 1936, but did not specify precise locations from which they originated. Robson (1989) reported that Florida Bonneted Bat specimens were collected regularly in Miami during the midcentury. In a table summarizing all bat specimens available in museums in 1989, he documented 18 individuals collected in the greater Miami area. These included three bats from Miami High School, three from North Miami High School, one male from University of Miami in Coral Gables in 1955, two males from chimneys in homes in Coral Gables in 1961 and Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 5 Table 1. Description of buildings used by Florida Bonneted Bats, Eumops floridanus, in Miami, FL, indicating the year bats were observed, type of building, type of architecture, year built, location within Miami, description of bat use, and method used to determine use. Numbered superscripts denote museum reports, and lettered superscripts denote published reports. Year used by bats Type of structure Architectural type Year built Location in greater Miami area Description of bat use Location method 1949, 1951, 1953 Miami High School (i.e., Miami Senior High School) Mediterranean 1928 Miami 7 individuals found singly Museum1,2, indirect reporta,b 1952 North Miami High School Other 1951 North Miami 4 individuals Museum2, indirect reportb 1955 University of Miami Unknown Unknown Coral Gables 1 male Indirect reportb 1961 House Unknown Unknown Coral Gables 1 male in house chimney Indirect reportb 1962 House Unknown Unknown Coral Gables 1 male in house chimney Indirect reportb 1988 Office Building Art deco 1986 Coral Gables 1 female on exterior balcony Citizen reportc 1995 House Mediterranean 1926 Coral Gables 1 male entered through chimney Homeowner reportd 1997, 2014, 2017 House Mediterranean 1929 Coral Gables 1 male entered through chimney, colony roosted, 1 male roosted Homeowner reportd, citizen scientistse, radio- telemetry 2015 Condominium Other 1981 Kendall Colony (n = 10) roosted Building renovator report 2017 House Other 1986 West Kendall 1 male roosted Radio-telemetry 2017 House Mediterranean 1925 Coral Gables 1 male roosted Radio-telemetry 2017 House Mediterranean 1928 Coral Gables Colony (n = 10) roosted Radio-telemetry 2018 House Mediterranean 1925 Coral Gables 1 male entered through chimney Citizen report 2020 Bell tower Mediterranean Unknown Pinecrest Colony (n = 15) roosted Citizen report 1 Field Museum of Natural History, Chicago, IL 2 Florida Museum of Natural History, Gainesville, FL a Schwartz 1952 b Robson 1989 c Robson et al. 1989 d Gore et al. 2015 Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 6 1962 (Robson 1989), and one female from an office building balcony in Coral Gables in 1988 (Robson et al. 1989). Collection locations for eight other records simply indicate Coral Gables (n = 7) or Miami (n = 1). The only published records of Florida Bonneted Bats collected in the greater Miami area anywhere other than in buildings included three bats reportedly taken on the ground (two of which were associated with construction projects), all undated but presumably midcentury (Owre 1978); one bat under rocks in Coral Gables in 1955 (Robson 1989); bats taken in low shrubbery, undated but presumably midcentury (Jennings 1958); and bats in shafts of royal palm leaves, undated (Belwood 1992). Our review of records obtained directly from the four museums now housing E. floridanus/E. glaucinus floridanus specimens documented six additional records, leading to a total of 20 individuals collected in the greater Miami area between 1949 and 1965. Only 10 of these records had precise descriptions of locations where specimens were collected; all 10 were from either Miami High School or North Miami High School. No specimens were reported during the 21-year period extending from 1966 to 1987, leading to speculation that the species might have been extirpated from the region (Belwood 1992). Subsequently, a single pregnant female Florida Bonneted Bat was found roosting on the 7th-floor balcony of an office building in Coral Gables in 1988 (Robson et al. 1989). A single male Florida Bonneted Bat entered a home in Coral Gables through a chimney in 1995, and another male Florida Bonneted Bat entered a home with Mediterranean Revival architecture in Coral Gables through a chimney in 1997 (Gore et al. 2015). Following these records, there was again a period with no confirmed specimens in Miami for well over a decade (1998–2013), although acoustic surveys suggested the species persisted (Marks and Marks 2012). Recent Roosts Four incidents of Florida Bonneted Bats in buildings have been reported to FWC, USFWS, and/or UF since the species was federally listed (i.e., 2013–2020; Fig. 1B). The first was a colony of bats tracked through auditory means to a home in Coral Gables in 2014 by a Miami-based group of citizen scientists, the Bat Squad (Staletovich 2014). This colony of bats emerged from beneath the clay tile roof of the same home with Mediterranean Revival architecture that a male Florida Bonneted Bat had entered through a chimney in 1997 (Gore et al. 2015). The second was a colony auditorily identified as Florida Bonneted Bats roosting in two condominiums located in a complex in Kendall in 2016. The third was a single male bat found within the living space of a home with Mediterranean Revival architecture in Coral Gables in 2018; the homeowner believed the bat had entered through the chimney. The fourth was a colony of bats located in a church bell tower in Pinecrest when the clay tile roof was removed in 2020. The species identities of individuals in the third and fourth incidents were confirmed by a local veterinarian (F. Ridgley, 2020, pers. comm). During our radio-telemetry effort to locate roosts in 2017, we captured 11 Florida Bonneted Bats in greenspaces within the greater Miami area. Eight of these individuals were captured at Granada Golf Course, two at Zoo Miami, and one at Kendall Indian Hammocks Park; none were captured at Fairchild Tropical Botanic Garden. We captured four adult males, three subadult males, three adult females, and one subadult female. One adult female was pregnant and another was lactating, confirming that bats in the greater Miami area were reproductively active. We deployed transmitters on five bats (one adult female and four adult males), which led us to discover four roost structures, all located in residences (Figs. 1B, 1C). One home, located 5.8 km west of the capture site, was used by a lone male captured at Kendall Indian Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 7 Hammocks Park. The other three homes were in Coral Gables, had Mediterranean style architecture, and were within one km of each bat’s capture site, the Granada Golf Course. Two of the homes used in Coral Gables were vacant and undergoing renovation. One of these vacant homes was used as a roost by a colony of at least 10 Florida Bonneted Bats, and the other which was the same home where Florida Bonneted Bats had been documented in 1997 and 2014, was used by a lone male. The third home used in Coral Gables was also used by a lone male. The body condition indices (BCI) of bats of all sex and reproductive condition categories captured during this investigation were lower than those reported for Florida Bonneted Bats in a natural area (Ober et al. 2017b) (Table 2). Small sample sizes precluded our ability to determine if differences were statistically significant. Discussion There is a long history of building use by Florida Bonneted Bats in the greater Miami area, the most urbanized region in Florida, despite urban intensification. This species was first documented in the region in the 1930s, when Miami-Dade County had <200,000 people. Intermittent reports confirm the bats’ continued presence as the human population ballooned to 2.8 million over the subsequent eight decades. The past 80 years have been punctuated by periods with many reports of Florida Bonneted Bats in buildings (1936–1965, 1988–1997, and 2014–2020) interspersed with periods having no reports (Table 1). We cannot conclude definitively whether these alternating intervals reflect fluctuations in bat abundance, changes in bat roosting patterns, or simply incidental trends in reporting. Regardless, the species has clearly continued to reside in the same portion of the city for over a half century, with perhaps a slight southward and westward shift away from areas that experienced the greatest growth in human population density during this time (Nijman and Clery 2015; Fig. 1B). Many of the buildings used by Florida Bonneted Bats in the greater Miami area were built during 1925–1930, and many of these have Mediterranean Revival architecture. The link between these bats and buildings with this architectural style has been reported repeatedly (Barbour and Davis 1969, Belwood 1992, Gore et al. 2015, Owre 1978). Several records mention bats entering homes through, or roosting within, chimneys in Coral Gables (Gore et al. 2015, Owre 1978, Robson 1989, homeowner report received in 2018). In most cases where the address could be verified, these buildings shared a Mediterranean architectural style with chimneys exhibiting an integrated arch cover over the flue opening. Bats may find these small spaces appealing (Gore et al. 2015; Fig. 2). Recent citizen reports and our recent radio-tracking efforts confirmed that bats continue to use buildings Table 2. BCI (Body condition indices) (mean ± SD) of Florida Bonneted Bats, Eumops floridanus, in Miami, FL (this study), and in a natural area (Fred C. Babcock/Cecil M. Webb Wildlife Management Area [BWWMA], FL, reported in Ober et al. 2017b). Numbers in parentheses indicate sample sizes. Sex Status BCI Miami bats BCI BWWMA bats Male Reproductively active adult 0.647 ± 0.078 (3) 0.686 ± 0.062 (31) Non-reproductive adult 0.607 (1) 0.642 ± 0.088 (16) Non-reproductive subadult 0.486 ± 0.031 (3) 0.608 ± 0.053 (35) Female Pregnant adult 0.654 (1) 0.750 ± 0.063 (56) Non-reproductive adult 0.439 (1) 0.645 ± 0.053 (46) Non-reproductive subadult 0.431 (1) 0.609 ± 0.059 (35) Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 8 with this architecture, as two of the three roosts reported by citizens and three of the four roosts identified through radio-tracking were homes with similar architectural features. Interestingly, bats used one of the same homes in 2017 that they had used in 1997 and 2014, despite the choice of millions of artificial structures across the 150 km2 area comprising the greater Miami area. This residence was located within a neighborhood in Coral Gables used consistently by Florida Bonneted Bats over the past three decades (Fig. 1C). Future research should investigate what specific aspects of the buildings selected by Florida Bonneted Bats in urban areas attract the bats. We found it difficult to distinguish exactly where bats were exiting from structures during emergence observations and therefore recommend the use of infrared or thermal-imaging video cameras during future emergence observations to build a deeper understanding of which portion of the buildings bats are using as roosts. Characteristics of homes in Coral Gables that may be appealing to roosting bats are the clay tile roofs that provide ample crevices, the stucco substrate that minimizes temperature fluctuations, the unusual chimney features typical of Mediterranean Revival homes, or the large yards with lush vegetation that are more prevalent in this region than elsewhere in the greater Miami area (Nijman and Clery 2015). The recent documentation of Florida Bonneted Bats roosting in Kendall and Pinecrest, two regions that have not experienced the densification of buildings typical of much of the rest of the greater Miami area, suggests Florida Bonneted Bats may prefer areas with lower building densities and greenery (Nijman and Clery 2015). Alternatively, it may be the clay tile roof of the belltower in Pinecrest, which is similar to Figure 2. Many buildings established in Miami during the 1920s and 1930s featured Mediterranean Revival architecture, characterized by elements depicted here, which may attract Florida Bonneted Bats, Eumops floridanus. Illustrative credit: Elizabeth Braun de Torrez. Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 9 that of many homes used as roosts in Coral Gables, that attracted bats to roost there. Clearly, additional investigation is warranted to determine the relative importance to the bats of factors such as roofing material, building style, chimney architect ure, and vegetation. In contrast to natural areas, where Florida Bonneted Bats regularly roosts in trees (Angell and Thompson 2015; Braun de Torrez et al. 2016, 2020 unpubl. data), use of vegetation by the species in Miami has not been documented in recent years. The last time Florida Bonneted Bats were reported using natural vegetation in the greater Miami area was at least 3 decades ago, and it has possibly been much longer (bats taken from low shrubbery reported in Jennings 1958; bats found in shafts of Roystonea regia Kunth reported in Belwood 1992; neither reported precise locational information or the year of these observations). It is unclear whether bats in this urban region have been using buildings exclusively during recent times because these structures provide roost conditions superior to those provided by natural vegetation, whether vegetation with requisite characteristics has become limited in availability over time, or whether bats have continued to use both natural and artificial structures, but the former have not been investigated with adequate intensity to detect use. To date, the greater Miami region is the only area within the species’ geographic range where Florida Bonneted Bats have been reported to roost in buildings. Additional research in other urban areas is recommended to determine if this pattern reflects a true lack of use of buildings by the species in any other urban area. We recommend comparisons of demography and fitness of Florida Bonneted Bats roosting in urban areas relative to bats roosting in natural areas to determine if urban conditions impart negative long-term population level impacts. Larger sample sizes are needed to confirm or refute the trends suggested by the data reported here. The comparatively lower body condition indices of bats of all sex and reproductive condition categories captured during this investigation relative to those reported for Florida Bonneted Bats in a natural area suggest that individuals in urban areas may be less fit. These differences may be due to increased stress experienced by bats in Miami compared to natural areas, perhaps due to artificial light pollution, elevated noise levels, or inferior d iets (Jung and Threlfall 2016). We also recommend additional research to clarify the timing of reproduction of urban Florida Bonneted Bats. Given that purposeful exclusions of bats from buildings and roof renovations are two primary pathways through which bat mortality or injury could occur in urban areas, a better understanding of the timing of reproductive peaks among urban populations of Florida Bonneted Bats could enable refinement of guidelines pertaining to such activities. Currently, bats of all species cannot legally be excluded from man-made structures in Florida when non-volant pups are present, which is designated as the period extending from 15 April to 15 August (Florida Administrative Code Chapter 68A-4.001 and 68A-9.010). Research on Florida Bonneted Bats in a natural area in southwest Florida suggests that the species may be aseasonally polyestrous with a peak pregnancy period in spring (Ober et al. 2017a). A clearer understanding of when non-volant pups are least likely to be present in buildings in urban areas would enable refinement of guidelines so that dis - turbance to these bats is minimized during periods when they are most vulnerable. Current guidelines specify that, if bats are discovered in a building undergoing renovation at any time of year within the geographic range of Florida Bonneted Bats, consultation with USFWS should occur prior to proceeding if the bats are suspected to be Florida Bonneted Bats (Florida Bonneted Bat Consultation Guidelines, available at https://www.fws. gov/verobeach/ListedSpeciesMammals.html). This suggests the need for an educational campaign to increase awareness that an endangered species of bat sometimes resides in buildings in Miami. Such a campaign should include homeowners, who might encounter Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 10 bats but consider them to be a nuisance or a human health concern and be unaware that the animals are protected by conservation regulations or be unable to identify the species. Efforts should prioritize those areas—such as Coral Gables—where older homes (built in the 1920s and 1930s) are most common, particularly those with Mediterranean architecture. Also, a training program should target individuals in professions likely to encounter roosting bats, such as roofers and nuisance wildlife control operators, to ensure they can identify Florida Bonneted Bats, know who to contact if they suspect they have found this species, and are aware of the time of year of bat eviction restrictions. Our study provides the initial foundation for understanding roost use by Florida Bonneted Bats in urban areas and highlights aspects of roost use about which our knowledge is limited. As urbanization continues throughout south and central Florida, additional research and targeted education efforts can play a critical role in the development and adoption of appropriate conservation actions to ensure the continued existence of Florida Bonneted Bats in Miami and other urban areas across the species’ range. Acknowledgments We thank Shalana Gray for participating in all aspects of field work. We also thank the following individuals for facilitating access to properties where captures were attempted and assisting with radiotracking and/or roost emergence observations: Troy Hall, Amy Padolf, Frank Ridgely, Eduardo Salcedo, and Sonya Thompson. We also thank Adam Ferguson, Verity Mathis, and Sandra Sneckenberger for providing information. Lastly, we thank FWC, UF, and Bat Conservation International for funding. Literature Cited Aldridge, H.D.J.N., and R.M. Brigham. 1988. Load carrying and maneuverability in an insectivorous bat: A test of the 5% “rule” of radio-telemetry. Journal of Mammalogy 69:379–382. Allen, G.M. 1932. A Pleistocene bat from Florida. Journal of Mammalogy 13:256–259. Angell, E.N., and G. Thompson. 2015. Second record of a natural Florida Bonneted Bat (Eumops floridanus) roost in a Longleaf Pine (Pinus palustris). Florida Field Naturalist 43:185–188. Barbour, R.W., and W.H. Davis. 1969. Bats of America. University Press of Kentucky, Lexington, KY, USA. 269 pp. Barbour, T. 1936. Eumops in Florida. Journal of Mammalogy 17:414. Belwood, J.J. 1992. Florida Mastiff Bat Eumops glaucinus floridanus. Pp. 216–223, In S.R. Humphrey (Ed.). Rare and Endangered Biota of Florida. Vol. I. Mammals. University Press of Florida, Gainesville, FL, USA. 392 pp. Braun de Torrez, E.C., H.K. Ober, and R.A. McCleery. 2016. Use of a multi-tactic approach to locate an endangered Florida Bonneted Bat roost. Southeastern Naturalist 15:235–242. Braun de Torrez, E.C., S.T. Samoray, K.A. Silas, M.A. Wallrichs, M.W. Gumbert, H.K. Ober, and R.A. McCleery. 2017. Acoustic lure allows for capture of a high-flying, endangered bat. Wildlife Society Bulletin 41:322–328. Florida Fish and Wildlife Conservation Commission (FWC). 2011. Biological status review report for the Florida Bonneted Bat Eumops floridanus. Tallahassee, FL, USA. 11 pp. FWC. 2013. A species action plan for the Florida Bonneted Bat Eumops floridanus. Tallahassee, FL, USA. 35 pp. Gore, J.A., M.S. Robson, R. Zambrano, and N.J. Douglass. 2015. Roosting sites of a Florida Bonneted Bat (Eumops floridanus). Florida Field Naturalist 43:179–184. Jennings, W.L. 1958. The ecological distribution of bats in Florida. Ph.D. Dissertation. University of Florida, Gainesville, FL, USA. 126 pp. Jonasson, K.A., and C.K.R. Willis. 2011. Changes in body condition of hibernating bats support the thrifty female hypothesis and predict consequences for populations with white-nose syndrome. PLoS ONE 6:e21061. Urban Naturalist E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 2021 No. 42 11 Jung K., and C.G. Threlfall. 2016. Urbanisation and its effects on bats—A global meta-analysis. Pp. 13-33, In C. Voigt and T. Kingston (Eds.). Bats in the Anthropocene: Conservation of Bats in a Changing World. Springer, Cham. 606 pp. https://doi.org/10.1007/978-3-319-25220-9_2. Kunz, T.H., and E.L.P. Anthony. 1982. Age estimation and post-natal growth in the bat Myotis lucifugus. Journal of Mammalogy 63:23–32. Marks, G.E., and C.S. Marks. 2012. Status of the Florida Bonneted Bat (Eumops floridanus). Report submitted by the Florida Bat Conservancy for the US Fish and Wildlife Service under grant agreement number 40181AG121. Florida Bat Conservancy, Bay Pines, FL, USA. 22 pp. Nijman, J., and T. Clery. 2015. Rethinking suburbia: A case study of metropolitan Miami. Environmental Planning A: Economy and Space 47:69–88. Ober, H.K., E.C. Braun de Torrez, J.A. Gore, A.M. Bailey, J.K. Myers, K.N. Smith, and R.A. Mc- Cleery. 2017a. Social organization of an endangered subtropical species, Eumops floridanus, the Florida Bonneted Bat. Mammalia 81:375–383. Ober, H.K., E.C. Braun de Torrez, R.A. McCleery, A.M. Bailey, and J.A. Gore. 2017b. Sexual dimorphism in the endangered Florida Bonneted Bat, Eumops floridanus (Chiroptera: Molossidae). Florida Scientist 80:38–48. Owre, O.T. 1978. The Florida Mastiff Bat, Eumops glaucinus floridanus. Pp. 43–44, In J.N. Layne (Ed.). Rare and Endangered Biota of Florida. Vol. 1. Mammals. University Press of Florida, Gainesville, FL, USA. 52 pp. Reynolds, D.S., J.C. Sullivan, and T.H. Kunz. 2009. Evaluation of total body electrical conductivity to estimate body composition of a small mammal. Journal of Wildlife Management 73:1197–1206. Robson, M. 1989. Status survey of the Florida Mastiff Bat. Final performance report. Florida Game and Freshwater Fish Commission, Tallahassee, FL, USA. 18 pp. Robson, M.S., F.J. Mazzotti, and T. Parrott. 1989. Recent evidence of the Mastiff Bat in southern Florida. Florida Field Naturalist 17:81–82. Schwartz, A. 1952. The land mammals of southern Florida and the upper Florida Keys. Ph.D. Dissertation. University of Michigan, Ann Arbor, MI, USA. 189 pp. Staletovich, J. 2014. “Citizen scientist” finds rare bat roost near Gables Golf Course. Miami Herald. 2 October 2014. Available online at https://www.miamiherald.com/news/local/community/miamidade/ coral-gables/article2489882.html. Accessed 21 August 2020. Timm, R.M., and H.H. Genoways. 2004. The Florida Bonneted Bat, Eumops floridanus (Chiroptera: Molossidae): Distribution, morphometrics, systematics, and ecology. Journal of Mammalogy 85:852–865. US Fish and Wildlife Service (USFWS). 2013. Endangered and threatened wildlife and plants: Endangered species status for the Florida Bonneted Bat. Federal Register 78:61003–61043.