Urban Roosts: Use of Buildings by Florida Bonneted Bats - Elysia N. Webb, Holly K. Ober, Elizabeth C. Braun de Torrez, Jeffery A. Gore, and ...
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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 No. 42 Urban Naturalist 2021
Urban Naturalist Board of Editors ♦ The Urban Naturalist is a peer-reviewed and Myla Aronson, Rutgers University, New Brunswick, NJ, edited interdisciplinary natural history journal USA with a global focus on urban areas (ISSN 2328- Joscha Beninde, University of California at Los Angeles, 8965 [online]). CA, USA ... Co-Editor Sabina Caula, Universidad de Carabobo, Naguanagua, ♦ The journal features research articles, notes, Venezuela and research summaries on terrestrial, fresh- Sylvio Codella, Kean University, Union New Jersey, USA water, and marine organisms and their habitats. Julie Craves, University of Michigan-Dearborn, Dearborn, ♦ It offers article-by-article online publication MI, USA for prompt distribution to a global audience. Ana Faggi, Universidad de Flores/CONICET, Buenos Aires, Argentina ♦ It offers authors the option of publishing large Leonie Fischer, Technical University of Berlin, Berlin, files such as data tables, and audio and video Germany clips as online supplemental files. Chad Johnson, Arizona State University, Glendale, AZ, ♦ Special issues - The Urban Naturalist wel- USA comes proposals for special issues that are based Kirsten Jung, University of Ulm, Ulm, Germany Erik Kiviat, Hudsonia, Bard College, Annandale-on- on conference proceedings or on a series of in- Hudson, NY, USA vitational articles. Special issue editors can rely Sonja Knapp, Helmholtz Centre for Environmental on the publisher’s years of experiences in ef- Research–UFZ, Halle (Saale), Germany ficiently handling most details relating to the David Krauss, City University of New York, New York, publication of special issues. NY, USA Mark Laska, Great Ecology, consulting, La Jolla, CA, USA ♦ Indexing - The Urban Naturalist is a young Zdenka Lososova, Masaryk University, Brno, Czechia journal whose indexing at this time is by way of Joerg-Henner Lotze, Eagle Hill Institute, Steuben, ME ... author entries in Google Scholar and Research- Publisher gate. Its indexing coverage is expected to become Kristi MacDonald, Hudsonia, Bard College, Annandale-on- comparable to that of the Institute's first 3 journals Hudson, NY, USA Ian MacGregor-Fors, Insituto de Ecología Mexico, (Northeastern Naturalist, Southeastern Natural- Veracruz, Mexico ... Co-Editor ist, and Journal of the North Atlantic). These 3 Tibor Magura, University of Debrecen, Debrecen, Hungary journals are included in full-text in BioOne.org Brooke Maslo, Rutgers University, New Brunswick, NJ, and JSTOR.org and are indexed in Web of Science USA (clarivate.com) and EBSCO.com. Mark McDonnell, Royal Botanic Gardens Victoria and University of Melbourne, Melbourne, Australia ♦ The journal's staff is pleased to discuss ideas Mike McKinney, University of Tennessee, Knoxville, TN, for manuscripts and to assist during all stages of USA manuscript preparation. The journal has a page Desirée Narango, City University of New York, New York, charge to help defray a portion of the costs of NY, USA publishing manuscripts. Instructions for Authors Mitchell Pavao-Zuckerman, University of Arizona, Tucson, Arizona, USA are available online on the journal’s website Joseph Rachlin, Lehman College, City University of New (http://www.eaglehill.us/urna). York, New York, NY, USA ♦ It is co-published with the Northeastern Natu- Travis Ryan, Center for Urban Ecology, Butler University, ralist, Southeastern Naturalist, Caribbean Natu- Indianapolis, IN, USA ralist, Eastern Paleontologist, Eastern Biolo- Michael Strohbach, Technische Universität Braunschweig, Institute of Geoecology, Braunschweig, Germany gist, and Journal of the North Atlantic. Katalin Szlavecz, Johns Hopkins University, Baltimore, ♦ It is available online in full-text version on the MD, USA journal's website (http://www.eaglehill.us/urna). Paige Warren, University of Massachusetts, Amherst, MA, Arrangements for inclusion in other databases USA are being pursued. Alan Yeakley, Portland State University, Portland, OR, USA Iriana Zuria, Universidad Autónoma del Estado de Hidalgo, Hidalgo, Mexico Cover Photograph: A Florida Bonneted Bat (Eumops floridanus) captured during this study. Dade County, Florida. Photograph © Elysia Webb. 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).
2021 Urban Naturalist No. 42 2021 Urban Naturalist 42:1–11 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 con- servation 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 spe- cies 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 feder- ally endangered in 2013, citing concerns about habitat loss, degradation, and modification from human population growth as well as the species’ small population size, slow reproduc- tion, low fecundity, and relative isolation (USFWS 2013). The ongoing loss and modifica- tion 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 intensi- fied. Guidelines recently established by the USFWS to minimize potential negative im- 1 Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL 32611 USA. 2 Florida Fish and Wildlife Conservation Commission, Gainesville, FL 32601 USA. 3Florida Fish and Wildlife Conservation Commission, Panama City, FL 32409 USA. 4Florida Fish and Wildlife Conserva- tion Commission, West Palm Beach, FL 33412 USA. *Corresponding author: holly.ober@ufl.edu. Manuscript Editor: Brooke Maslo 1
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 con- servation 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 (“Consulta- tion 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). 2
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 en- dangered 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. Be- cause 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 addi- tional 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 web- sites 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 loca- tions. The southernmost area was Zoo Miami, located in Miami; the northernmost and east- ernmost 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. 3
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 (Jonas- son 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, On- tario, 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 identi- fied 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 ar- chitecture. 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 4
Table 1. Description of buildings used by Florida Bonneted Bats, Eumops floridanus, in Miami, FL, indicating the year bats were observed, type of building, 2021 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 Type of structure Architectural Year built Location in Description of bat use Location method by bats type greater Miami area 1949, 1951, Miami High School Mediterranean 1928 Miami 7 individuals found singly Museum1,2, indirect reporta,b 1953 (i.e., Miami Senior High School) 1952 North Miami High Other 1951 North Miami 4 individuals Museum2, indirect reportb School 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 5 1995 House Mediterranean 1926 Coral Gables 1 male entered through chimney Homeowner reportd 1997, 2014, House Mediterranean 1929 Coral Gables 1 male entered through chimney, Homeowner reportd, citizen Urban Naturalist 2017 colony roosted, 1 male roosted 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 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 No. 42 Gore et al. 2015
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 Cor- al 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, US- FWS, 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 roost- ing 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 Bon- neted 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 6
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 catego- ries 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
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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. In- terestingly, 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. Character- istics 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. 8
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 fac- tors such as roofing material, building style, chimney architecture, and vegetation. In contrast to natural areas, where Florida Bonneted Bats regularly roosts in trees (An- gell 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 re- ported 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 roost- ing 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 diets (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 renova- tion 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 Bon- neted 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 9
2021 Urban Naturalist No. 42 E.N. Webb, H.K. Ober, E.C. Braun de Torrez, J.A. Gore, R. Zambrano 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 architec- ture. 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 Bon- neted 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 radio- tracking and/or roost emergence observations: Troy Hall, Amy Padolf, Frank Ridgely, Eduardo Sal- cedo, 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. Hum- phrey (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. 10
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