BULLETIN

BATS OF THE BAHAMAS: NATURAL HISTORY AND CONSERVATION

Kelly A. Speer, J. Angel Soto-Centeno, Nancy A. Albury, Zachary Quicksall, Melina G. Marte, and David L. Reed

Vol. 53, No. 3, pp. 45–95 August 12, 2015 ISSN 2373-9991 UNIVERSITY OF FLORIDA GAINESVILLE BATS OF THE BAHAMAS: NATURAL HISTORY AND CONSERVATION

Kelly A. Speer1,4, J. Angel Soto-Centeno2, Nancy A. Albury3, Zachary Quicksall1, Melina G. Marte1, and David L. Reed1

ABSTRACT The Bahamas are well known for their karst landscape, with vast cave systems and inundated caves called blue holes that formed during the Pleistocene. Despite this, the biota of these islands has not been well documented when compared to other West Indian islands. It is important that Bahamians have the knowledge and tools to preserve the flora and fauna of their islands in the face of increasing development. This review provides an accessible resource for Bahamians and researchers by consolidating the available knowledge on bat biology from literature and natural history collections. Bats represent the most diverse group of mammals in The Bahamas and no study has thoroughly characterized their biology on these islands. There are currently 10 species of bat in The Bahamas, one of which is endemic and considered “near threatened” by the IUCN. While 9 of the 10 are considered species of “least concern,” bats in The Bahamas may be at higher risk of local extinction than in other parts of their ranges. As previous local conservation efforts have shown, it is imperative to gain support for bat research and conservation through outreach and education. We hope this review enables outreach efforts and research projects key to the conservation of bat populations in The Bahamas.

Key words: Chiroptera, Bahamas, species accounts, natural history collections, outreach.

TABLE OF CONTENTS Introduction...... 46 Materials and Methods...... 49 Species Accounts...... 49 Conclusions...... 69 Acknowledgements...... 72 Literature Cited...... 72 Table 1. Bahamian bats localities...... 77

1Division of Mammals, Florida Museum of Natural History, University of Florida, Gainesville, Florida, 32611-7800 2Department of Mammalogy, American Museum of Natural History, New York, New York, 10024 3National Museum of The Bahamas/Antiquities, Monuments and Museums Corporation, Marsh Harbour, Great Abaco, The Bahamas 4Corresponding Author: [email protected] Speer, K. A., J. A. Soto-Centeno, N. A. Albury, Z. Quicksall, M. G. Marte, and D. L. Reed. 2015. Bats of the Bahamas: natural history and conservation. Bulletin of the Florida Museum of Natural History 53(3):45–95. https://www.flmnh.ufl.edu/bulletin/publications/ 46 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

INTRODUCTION largely unknown, they likely play an important role in controlling insect populations on these islands. Conservation of roosting and foraging habitat is Frugivorous (fruit-eating) and nectarivorous crucial to the maintenance of bat biodiversity. This (nectar-eating) bats provide a direct benefit to is of special importance on islands, where bats are ecosystems through seed dispersal and pollination often the most abundant native terrestrial mammals (Kunz et al., 2011). Commercially important bat- (Gannon et al., 2005; Fleming and Racey, 2010). pollinated plants (e.g., agave, bananas, mangos, Excluding livestock, there are 15 species of etc.) are artificially pollinated, but bats help native and introduced terrestrial mammals in The maintain genetic diversity in wild populations of Bahamas, 10 of which are bats (Table 1; Buden, these plants through pollination (Kunz et al., 2011). 1986). As these islands become increasingly Frugivorous bats are able to transport seeds great developed, human disturbance of bat roosting and distances and are more likely to disperse them foraging habitat poses a threat to terrestrial mammal in open, disturbed areas, which have favorable diversity. However, it can be difficult to gain public conditions for early successional plants responsible support for conservation of bats because they are for regrowth in disturbed areas (Muscarella and culturally associated with fear, danger, ugliness, Fleming, 2007; Kunz et al., 2011). Through insect and disease (Hutson et al., 2001; Knight, 2008; population control, seed dispersal, and pollination, Kunz et al., 2011). Based on limited records, there bats provide economic benefits for humans and are have been no documented transmissions of rabies instrumental in maintaining healthy ecosystems, from wildlife to humans in The Bahamas (Belotto making preservation of bat populations an et al., 2005). More information is needed on the role important conservation issue. bats play in disease maintenance and transmission, but negative perception of bats in The Bahamas Effects of Climate and Human Colonization may be caused by a lack of knowledge and Bats represent a significant proportion of the misrepresentations of bats in folklore and media, extant mammal fauna in The Bahamas, but little rather than fear associated with disease potential effort has been made to protect their populations. (Hutson et al., 2001). The lack of information about Compared to other West Indian islands (e.g., Cuba, the natural history of bats poses the greatest threat Hispaniola, Jamaica, and Puerto Rico), vertebrate to their conservation (Hutson et al., 2001). species diversity is low in The Bahamas. This is partly due to island area and the unique geologic Bats Support Healthy Ecosystems and climatic history of these islands (Hedges, 1996; Bats are economically beneficial to humans Dávalos and Russell, 2012). All exposed land in and make important contributions toward The Bahamas was deposited relatively recently, less maintaining healthy ecosystems (Hutson et al., than 500,000 years ago, and underwent significant 2001; Kunz et al., 2011). Insectivorous bats naturally changes in response to past sea level changes control insect populations, often consuming over (Carew and Mylroie, 1995; Mylroie, 2008). As The 25% of their body weight in insects in a single night Bahamas have maintained their geographic position (Kunz et al., 2011). In southwest Texas (United at least through the Quaternary, colonization of States), the presence of bats increased the amount The Bahamas by terrestrial animals likely occurred of harvestable cotton and decreased the number by over-water dispersal, with non-volant animals of pesticide sprays for an estimated monetary aided by floating vegetation, flotsam, or by human benefit of $688,000 per year (McCracken et al., introduction. 2012). Six of the 10 bat species in The Bahamas Sea level changes during glacial-interglacial consume members of the insect Order Diptera, cycles of the Pleistocene (ca. 2.5 Ma–11 ka) affected which includes mosquitoes. Although the amount the occurrence and distribution of animals in The and type of insects that Bahamian bats consume is Bahamas, and other islands of the West Indies, by SPEER ET AL.: Bats of the Bahamas: natural history and conservation 47

Figure 1. Map of The Bahamas indicating bathymetry. Black areas represent land, dark gray represents ocean < 200 m in depth, medium gray represents 201–500 m in depth, light gray represents 501–2500 m in depth and white represents depths > 2500 m. In the case of The Bahamas, most of the dark gray area is < 30 m in depth and is only slightly larger in area than the land area that would have been exposed during Pleistocene glacial periods 48 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) altering island size and distance between landmasses (Pregill and Olson, 1981). Human colonization of (Pregill and Olson, 1981). These ancient, global The Bahamas after 1.1 ka may have also changed climate fluctuations caused periods of extreme habitat composition (Kjellmark, 1996). Evidence cooling (glacials) followed by periods of warming from a sediment core in Andros indicates that (interglacials). During periods of glaciation, ice human-induced fires associated with agriculture sheets would form in temperate latitudes and freeze may have caused recent, rapid changes in plant seawater to such an extent that global sea levels communities (Kjellmark, 1996), which resulted in decreased. During interglacials, ice sheets would habitat loss for terrestrial animals. Some evidence melt, releasing water and causing global increases suggests that Pleistocene climatic fluctuations and in sea level. The Bahamas are low-lying islands on more recent habitat changes may not have had as top of shallowly submerged platforms and even great an impact on animal extinctions, which instead small changes in sea level influenced the amount seem to correspond better to human colonization of available habitat and the distance to other events (Steadman et al., 2014; Soto-Centeno and landmasses (Pregill and Olson, 1981). During Steadman, 2015). Fossils from Abaco indicate that the last interglacial (130–117 ka), sea level was 41% of terrestrial animals became locally extinct between 6 –10 m higher than it is today (Hearty following human colonization (Steadman et al., and Kaufman, 2000). The amount of exposed land 2014). Bats did not experience the level of local in The Bahamas would have been less than half extinctions seen in other terrestrial mammals, but of its current area (Morgan, 1989). Inundation of underwent the greatest loss in diversity in The large portions of land caused loss of habitat and Bahamas compared to other West Indian islands an increase in the distance between landmasses, (Morgan, 2001). potentially leading to isolated populations and Aims local extinctions (Dávalos and Russell, 2012). During the last glacial period (80–10 ka) sea levels Conservation of bats in The Bahamas remained at least 15 m below present until glaciers is dependent on increasing our knowledge of began to melt, approximately 17 ka (Richards et island populations and changing negative public al., 1994). At the lowest estimated sea levels during perception of bats to viewing bats as important this period (120 m below present), the islands of providers of ecosystem services. This review of the Great Bahama Bank (Andros; Bimini; the Berry literature and historical collections is meant to Islands; Cat Island; Eleuthera; the Exuma Islands, provide information to the people of The Bahamas collectively known as the Exumas; Long Island; allowing a better understanding of bat species that New Providence; and Ragged Island) and those inhabit their islands. We reviewed studies conducted of the Little Bahama Bank (the Abaco Islands, on island bats from the literature and collected collectively referred to as the Abacos, and Grand data from natural history collections. We note Bahama Island) would have formed two large that very few studies have been conducted in The islands, the extent of which follows the contour of Bahamas and the most recent review of Bahamian the carbonate platforms that create the shallow sea bats was almost 30 years ago (Buden, 1986). To surrounding the Bahamas (Fig. 1; Morgan, 1989). enable future research on ecology and natural Decreases in water depth and increases in island history of bats in The Bahamas, we have updated size may have facilitated colonization and dispersal the locality data where bats have been vouchered by animals. and preserved in natural history collections. By Dramatic habitat change following providing foundational information on the presence colonization by humans may have also caused local and distribution of bat populations, we can enable animal extinctions. Prevalence of arid-adapted effective conservation efforts directly involving the species in the fossil record suggests that The public and encourage future research focusing on Bahamas were much drier during the Pleistocene the natural history of Bahamian bat populations. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 49

MATERIALS AND METHODS accurate geographic coordinates when possible. Species Accounts We also identified potentially redundant collecting Buden (1986) counts 15 species of bat in The sites using GPS coordinates and specific locality descriptions associated with museum records. We Bahamas, excluding Brachyphylla nana, which is consolidated sites that were geographically close found in the Turks and Caicos. However, two of these and/or had matching locality descriptions using (Pteronotus parnellii and Mormoops blainvillei) are only known from fossils. An additional three species local and professional knowledge. We pair specific descriptions taken from museum databases with (Lonchorhina aurita, Glossophaga soricina, and our updated information to clarify and add detail Lasionycteris noctivagans) are documented from to locality data. Our database of updated and single specimens, which are likely vagrants that do consolidated information on every site where bats not represent resident populations in The Bahamas. have been collected in The Bahamas is presented After excluding these, we are left with 10 species of in Table 1. We do not necessarily recommend bats in The Bahamas (Table 1; Tadarida brasiliensis, that locality records curated at natural history Chilonatalus tumidifrons, Nyctiellus lepidus, collections be updated based on the data in Table Noctilio leporinus, Artibeus jamaicensis, Erophylla 1. Curators and collection managers must adhere to sezekorni, Macrotus waterhousii, Monophyllus the locality descriptions provided by collectors; in redmani, Eptesicus fuscus, and Lasiurus minor). contrast, we have the luxury of extrapolating from Most of these solely use cave roosts, but one roosts these descriptions based on our local expertise. Due in trees under foliage (Lasiurus minor), and four to the sensitivity of cave ecosystems to tourism and have been collected or observed in both caves and human disturbance, and their cultural importance buildings (Eptesicus fuscus, Erophylla sezekorni, as burial sites for Lucayan Taino Indians, we only Macrotus waterhousii, and Tadarida brasiliensis). provide coordinate data to two decimal places. Bat populations on other islands of the West Indies Maps were constructed in ArcGIS™ v.10.2 (ESRI, (e.g., Cuba and Puerto Rico) are better studied and Redlands, California) and indicate presence or provide some insight into the ecology of island bats. absence of a species on a given island. Presence of We review information on the 10 extant bat species in The Bahamas, focusing on studies conducted on a species on an island is defined as any island where specimens have been collected and preserved in a island populations when possible. Conservation natural history collection. Field observations of a status is taken from the International Union for species, without voucher specimens or tissues are Conservation of Nature (IUCN). Species collected not included in these maps. from the same cave are listed in most accounts, because previous research suggests that bats form SPECIES ACCOUNTS non-random species assemblages possibly related Molossidae (Free-Tailed Bats) to the roosting climate (Rodríguez-Durán, 1998). Tadarida brasiliensis (Saint-Hilaire, 1824) These lists are preliminary and caves have dynamic microclimates that may also influence the species Common Name.—Brazilian free-tailed bat composition of a roost. Type Locality.—“Curityba, Paraná, Brazil” (Shamel, 1931) Locality Information Conservation Status.—“least concern” We downloaded locality information for (IUCN, 2014) all bats collected in The Bahamas from VertNet Tadarida brasiliensis is a small (10–12 g), (vertnet.org). In many cases historical localities insectivorous bat with distinctive vertical wrinkles were georeferenced at the center of the island or grooves on the upper lip. Pelage is typically from which specimens were collected, because short and dense, and hairs are uniformly brown many bat specimens were collected prior to the with ventral pelage slightly lighter than dorsal availability of handheld GPS. We provide more pelage (Wilkins, 1989). Like other members of the 50 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Figure 2. Presence/absence map of Tadarida brasiliensis in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which T. brasiliensis is found are indicated ZLWKEODFN¿OO3KRWRJUDSKRIT. brasiliensis by J. A. Soto-Centeno. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 51 family Molossidae, approximately half of the tail normally occurs above the canopy or in open areas extends freely beyond the tail membrane, giving (Norberg and Rayner, 1987). To compensate for the name “free-tailed bat”. The characteristically lower maneuverability, it has been suggested that wrinkled upper lip makes T. brasiliensis easily the wrinkled upper lip of T. brasiliensis allows it distinguishable from other bats in The Bahamas to open its mouth wider to more effectively catch (Fig. 2). insects in air (Wilkins, 1989). Wing and flight Tadarida brasiliensis is widely distributed adaptations allow T. brasiliensis populations across the southern United States, Mexico and in temperate regions to fly long distances to Central America, portions of South America, foraging sites and undertake annual long-distance and the Greater and Lesser Antilles, making it migrations (Fleming and Eby, 2003). In Cuba and one of the most widely distributed species of bat Puerto Rico, populations do not seasonally migrate in the Americas. In The Bahamas, T. brasiliensis and it is likely that T. brasiliensis in The Bahamas was previously known only from Great Abaco, show a similar behavior, because food is available Eleuthera, Long Island, and the Exumas (Koopman throughout the year (Silva-Taboada, 1979; Gannon et al., 1957). Current records available from mus- et al., 2005). eum databases extend this distribution to include As in many insectivorous bats, foraging is Grand Bahama Island, Crooked and Acklins focused at dusk and dawn during peaks of insect Islands, and Long Cay (Table 1; Fig. 2). Fossil activity (Silva-Taboada, 1979). Large colonies of evidence suggests T. brasiliensis has become T. brasiliensis normally leave the roost together locally extinct in New Providence (Morgan, 2001). at dusk, flying in a serpentine pattern across the Approximately 270 T. brasiliensis individuals sky before dispersing (Wilkins, 1989). Generally, have been collected from The Bahamas and are insular colonies are smaller than those on the preserved in natural history collections, attesting to mainland and emergence may be more diffuse its abundance in The Bahamas. (Gannon et al., 2005; Rodríguez-Durán and Island colonies of T. brasiliensis roost in Christenson, 2012). In Cuba and Puerto Rico, T. buildings or caves with good ventilation and brasiliensis feeds mainly on flies (Diptera), wasps cool air (Koopman et al., 1957; Silva-Taboada, and ants (Hymenoptera), moths (Lepidoptera), 1979; Gannon et al., 2005; Rodríguez-Durán and true bugs (Homoptera; Silva-Taboada, 1979; and Christenson, 2012). Solitary individuals or Whittaker and Rodríguez-Durán, 1999), while small groups are found in fissures or on walls and populations in New Mexico and Texas feed mainly ceilings of caves and manmade structures, whereas on moths and beetles (Coleoptera; McWilliams, larger groups of tens of bats occupy bell holes, 2005). In more temperate regions, seasonal climate and groups of hundreds or thousands roost in large fluctuations strongly impact abundance and flat places on the ceiling of caves (Silva-Taboada, diversity of available food sources (McWilliams, 1979). Normally, individuals are tightly clustered 2005). Dietary variation may also be caused by forming a mass, but this may vary depending composition of bat communities in the West Indies on the temperature of the roost (Koopman et al., or on the mainland (Gannon et al., 2005). On some 1957; Wilkins, 1989; Gannon et al., 2005). In The islands, mustached bats (family Mormoopidae) Bahamas, T. brasiliensis has been documented feed mainly on moths and beetles and may exclude in the same roost as Eptesicus fuscus, Erophylla T. brasiliensis from this niche (Gannon et al., sezekorni and Macrotus waterhousii (Koopman et 2005). Similarly, bats of the genera Myotis and al., 1957; Wilkins, 1989; Speer, pers. obs.). Pipistrellus (family Vespertilionidae) may compete Tadarida brasiliensis is highly adapted for with T. brasiliensis on the mainland, but are absent rapid, sustained flight due to its narrow, pointed from the Greater Antilles, and may account for wings (Norberg and Rayner, 1987). This flight variation of fly consumption between mainland pattern allows for fast aerial capture of prey and insular populations (Whittaker and Rodríguez- insects, but decreases maneuverability so foraging Durán, 1999). 52 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

As in other bats, T. brasiliensis gives birth endemic to The Bahamas, and one of only two to one pup a year in the late spring and early endemic terrestrial mammals along with the summer. Little is known about the reproduction of Bahamian hutia (Geocapromys ingrahami, Order T. brasiliensis in the Greater Antilles, but Silva- Rodentia). C. tumidifrons is a small bat (ca. 4–7 Taboada (1979) documented gestation from April g) with wide ears that are shaped like a funnel. to June and lactation in July and August in Cuban The ears are so expanded that they cover the eyes populations. Studies conducted on mainland when observed from the side (Fig. 3A). Pelage is populations and our field observations in The dense and long, ranging from light brown to orange Bahamas suggest that there is little parental care of brown. Hair is bi-colored with light color at the pups, with females depositing their young in large base and dark at the tips (Allen, 1905). The ears groups, returning to nurse their young until weaned can appear light yellowish brown and darkening (Gustin and McCracken, 1987). to grayish brown toward the edges (Miller, 1903). Ecology of insular populations of T. C. tumidifrons has a characteristic and prominent brasiliensis is known mainly from Cuba and Puerto “moustache” of hair that is ventrally curved along Rico, and future research should focus on better the upper lip (Fig. 3A). Males have a large and characterizing annual movement, dietary variation, externally prominent natalid organ, about half the social and community structure, reproduction, and length of the skull, located between the ears (Miller, roost ecology of Bahamian colonies. T. brasiliensis 1903; Dalquest, 1950; Tejedor, 2011). The natalid is currently listed by the IUCN (2014) as “least organ is only present in the family Natalidae and its concern,” but population declines in other regions function is not well known. Some authors suggest have occurred due to destruction or disturbance it may aid in communication among individuals, of large colony roost sites and indirect poisoning or have sensory or glandular functions (Dalquest, by pesticides (Gannon et al., 2005). T. brasiliensis 1950; Tejedor, 2011). C. tumidifrons is similar to is known to occupy buildings in The Bahamas, other members of the family Natalidae, having a and extermination of these colonies may threaten short body, long tail, and thin legs that support a large portions of the bat communities on certain large tail membrane. These characteristics may Bahamian islands. For example, T. brasiliensis on cause C. tumidifrons to be confused with Nyctiellus the Abacos has only been collected from buildings lepidus (Natalidae) in The Bahamas. Nonetheless, and bats at these sites likely represent a large C. tumidifrons can be differentiated by the presence portion of the populations on these islands (Table of hairs along the free edge of the tail membrane 1). We advocate non-lethal exclusion of bats from and by a natalid organ that is more prominent than buildings (information is available through Bat in N. lepidus (Dalquest, 1950). Conservation International, http://www.batcon.org/ Chilonatalus tumidifrons is known from resources/for-specific-issues/bats-in-buildings) and Andros, the Abacos, and San Salvador (Fig. 3; preservation of cave roosts to decrease interaction Allen, 1911; Dávalos, 2005; Tejedor, 2011). Fossils between humans and bats and to maintain healthy have been reported from Andros, Cat Island, the populations of T. brasiliensis. Exumas, and New Providence (Morgan, 2001). Natalidae (Funnel-Eared Bats) C. tumidifrons is an uncommon species and only Chilonatalus tumidifrons (Miller, 1903) 120 specimens, collected over the course of more Common Name.—Bahamian funnel-eared bat than 100 years, are preserved in natural history Type Locality.—“Watling Island [= San Sal- collections (Table 1). vador],” Bahamas Chilonatalus tumidifrons exclusively roosts Conservation Status.—“near threatened” in caves in The Bahamas and prefers chambers with (IUCN, 2014) constant warm temperatures and high humidity Chilonatalus tumidifrons is the only bat (Table 1). Individuals form small colonies ranging from 10 to 500 individuals, roosting away from one SPEER ET AL.: Bats of the Bahamas: natural history and conservation 53

Figure 3. Presence/absence map of Chilonatalus tumidifrons and Nyctiellus lepidus in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which C. tumidifrons is found are indicated with black hatching and those on which N. lepidusDUHIRXQGDUH¿OOHGLQEODFN Photographs of C. tumidifrons (A) and N. lepidus (B) by J. A. Soto-Centeno. 54 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) another, often hanging by one foot (Allen, 1905; information for this species is speculative, primarily Soto-Centeno, pers. obs.). C. tumidifrons has been based on the life history accounts of other members found to share cave roosts with Eptesicus fuscus, of the family Natalidae. Fossil evidence suggests Erophylla sezekorni, and Macrotus waterhousii, that this species was once widespread on the Great but are never observed roosting in the immediate Bahama Bank, but currently only inhabits three vicinity of any of these species (Allen, 1905; Soto- islands (Great Abaco, Andros, and San Salvador). Centeno, pers. obs.). Some roosting groups have As C. tumidifrons has previously undergone local been observed to move between roosts (ca. 1 km extinction from much of its historic range, we apart) on a daily basis (Tejedor, 2011). agree with the IUCN classification of this species Chilonatalus tumidifrons begins to forage at as “near threatened.” Until the status of this species dusk and is insectivorous, like other members of the is properly assessed via research and population family Natalidae, but no study has comprehensively surveys, we encourage conservation of cave habitat analyzed the diet of this bat. The small body and and coppice and pine woodland. short broad wings of C. tumidifrons allow it to have Nyctiellus lepidus (Gervais, 1837) a slow and very maneuverable flight pattern that can be compared to that of the large “bat moth” Common Name.—Gervais’ funnel-eared bat (also known as money moth or black witch moth; Type Locality.—“Cuba” (specific locality Ascalapha odorata). This flight pattern allows and collector unknown) C. tumidifrons to use habitats with dense, woody Conservation Status.—“least concern” vegetation like coppice (dry broadleaf forests) and (IUCN, 2014) shrubs (Andersen, 1994). The maneuverable of Nyctiellus lepidus is the smallest of all flight of C. tumidifrons allows it to easily detect Bahamian bats (ca. 3–5 g) and one of the smallest and avoid mist-nets, even when netting in areas of bats in the world (Tejedor, 2011). It has short high activity near roosts (Soto-Centeno, pers. obs.). and narrow ears that are somewhat shaped like a Nothing is known about the reproduction funnel (Fig. 3B). Like other members of the family of C. tumidifrons. Allen (1905) observed a colony Natalidae, N. lepidus is characterized by its small of about 300 individuals on Great Abaco that was size, short body, long tail, and thin legs that support composed primarily of males. This observation a large tail membrane (Tejedor et al., 2005; Tejedor suggests that, as in other species of bats, males and 2011). As in Chilonatalus tumidifrons, males can females may segregate after mating. be distinguished from females by the presence Efforts towards the conservation of C. of the natalid organ located between the ears tumidifrons are perhaps more imperative than (Tejedor et al., 2005). Pelage is dense and long, any other bat species in The Bahamas. The IUCN ranging from grayish brown to reddish brown, (2014) considers C. tumidifrons near threatened and is lighter ventrally (Silva-Taboada, 1979). The due to its limited distribution and the potential head of N. lepidus is characterized by a low brain for loss of habitat and roosts. Preservation of case and prominent “moustache” of hair along the coppice communities in the midst of increasing upper lip (Fig. 3B; Tejedor, 2011). It can be easily deforestation (Larkin et al., 2012) will protect differentiated from C. tumidifrons by the absence valuable foraging grounds for C. tumidifrons. of hairs along the free edge of the tail membrane Maintenance of pristine cave ecosystems is also and among males, a less prominent natalid organ important for the conservation of C. tumidifrons, (Dalquest, 1950; Tejedor et al., 2005; Tejedor, which is especially susceptible to disturbance due 2011). to its strict roosting conditions and preference for In the Greater Antilles, N. lepidus is widely small roosting colonies (Dávalos, 2005). distributed on Cuba and Isla de Pinos (Allen and C. tumidifrons is the only endemic bat of Sanborn, 1937; Dalquest, 1950; Silva-Taboada, The Bahamas, however, the majority of available 1979; Tejedor et al., 2005). The Bahamian SPEER ET AL.: Bats of the Bahamas: natural history and conservation 55 populations of this species occur on Cat Island, 1937). Bats are known to have some of the largest Eleuthera, the Exumas, and Long Island (Fig. offspring known in mammals (Kurta and Kunz, 2B; Allen and Sanborn, 1937; Dalquest, 1950; 1987; Hayssen and Kunz, 1996; Soto-Centeno and Koopman et al., 1957; Tejedor, 2011). Fossils of Kurta, 2003) and newborn N. lepidus often average N. lepidus have been reported from Andros and the more than 35% of the mother’s weight (Silva- Exuma Islands (Morgan, 2001). Currently, there Taboada, 1979). Nonetheless, females commonly are over 600 specimens of N. lepidus preserved carry their pups during flight while foraging until in natural history collections, which attests to the pups begin to fly in late spring (Silva-Taboada, abundance of this species throughout its range 1979). (Table 1). Nyctiellus lepidus is considered a species Nyctiellus lepidus is exclusively a cave of “least concern” by the IUCN (2014). Tejedor dwelling species in The Bahamas. In Cuba, it has (2011) suggests N. lepidus may be resilient to been found primarily in caves with high humidity disturbance as long as roost habitats are preserved, (Silva-Taboada, 1979). Individuals in large colonies citing its preference for forming large colonies and can be observed widely spread out from one another its ability to use a variety of feeding habitats. N. (Soto-Centeno, pers. obs.). On Long Island, N. lepidus is a cave specialist that prefers sheltered lepidus has been observed roosting in deep, narrow areas of caves that are humid and maintain a bell holes of well-ventilated chambers, forming constant temperature (Silva-Taboada, 1979). The small groups of 10–15 individuals (Soto-Centeno, availability of climatically stable cave roosts may pers. obs.). The shape of these narrow bell holes limit the possibility of dispersal of this species may serve as a small thermal chamber that helps into new localities if current roosts are disturbed. maintain stable temperature and humidity. N. Alteration of the microclimate in caves has been lepidus has been observed in the same cave as implicated in the local extinction of other members Eptesicus fuscus, Erophylla sezekorni, Macrotus of the family Natalidae from some Bahamian waterhousii, and Tadarida brasiliensis in The islands (Dávalos, 2005) and colonies of N. lepidus Bahamas (Soto-Centeno, pers. obs.). may be at risk if human disturbance changes the The small size and highly maneuverable conditions of roosting sites. Information about flight of N. lepidus allows it to feed in cluttered sustainable guano mining can be found through environments within the forest understory and in Emerging Wildlife Conservation Leaders (http:// grasslands (Tejedor, 2011). Although the diet of wildlifeleaders.org/projects/bats/). N. lepidus on The Bahamas has not been studied, Noctilionidae ( ishing ats) populations on Cuba feed exclusively on insects F B Noctilio leporinus (Linnaeus , 1758) like true bugs (Homoptera), flies (Diptera), small bees (Hymenoptera), moths (Lepidoptera), and Common Name.—Greater Bulldog Bat or termites (Isoptera; Silva-Taboada, 1979). Foraging Fishing Bat bouts occur around dusk and dawn when most Type Locality.—“America” (Linnaeus, small insects are swarming (Tejedor, 2011). N. 1758), limited to “Suriname” by Thomas (1911) lepidus normally forages for short periods of time, Conservation Status.—“least concern” likely using small home ranges and traveling only (IUCN, 2014) short a distance from the roost (Tejedor 2011). Noctilio leporinus is an insectivorous and Reproduction in N. lepidus takes place during piscivorous (fish-eating) bat, living in tropical winter (December–February) and pregnant females lowland areas of the Neotropics (Hood and can be observed between March and July (Tejedor, Jones, 1984). It is the largest bat (ca. 50–65 g) 2011). Female-only maternity colonies form after inhabiting The Bahamas, being slightly heavier mating and males can be seen forming groups in than Artibeus jamaicensis (Hood and Jones, 1984; the external areas of the caves (Allen and Sanborn, Ortega and Castro-Arellano, 2001). N. leporinus 56 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) has a characteristically pronounced snout and during foraging, a trait that is rare in bats (Brooke, lips and the chin has well-defined ridges, giving 1997). In each harem, resident males are replaced it a bulldog-like appearance (Fig. 4B). It also has approximately every two years, but females within cheek pouches, unique to the family Noctilionidae, the group are stable and roosting groups tend to used for food storage during foraging (Hood and occupy the same physical place in a roost over Jones, 1984). Pelage is short and ranges in color several years (Brooke, 1997; Gannon et al., 2005). from orange to grayish brown. N. leporinus can be Noctilio leporinus forages in short bouts identified by its enlarged hind feet and claws (Hood totaling 1.3–2.5 hr./night (Brooke, 1997). It and Jones, 1984), and is not likely to be confused consumes a diversity of small fish (less than 15 with any other bat species in The Bahamas. cm in length) and insects from bays and coastlines, Noctilio leporinus is distributed from estuaries, streams, and ponds (Hood and Jones, northeastern Mexico to northern Argentina, and 1984; Brooke, 1994; Bordignon, 2006). When is widespread across the West Indies (Hood and catching fish, these bats use echolocation to detect Jones, 1984). Populations are focused along disturbances at the water surface and use their large coastlines, estuaries and lowlands, and are rare in hind feet to rake across the surface, spearing their arid or montane regions across its range (Gannon prey with their claws (Altenbach, 1989; Schnitzler et al., 2005). Schwartz and Thomas vouchered et al., 1994). N. leporinus does not use its tail the only specimens collected in The Bahamas in membrane during capture of fish, but may use it 1967 on Great Inagua from a single cave locality during capture of insects from the air (Altenbach, (Table 1). N. leporinus has not been documented in 1989; Schnitzler et al., 1994). N. leporinus can The Bahamas since this initial collection, although catch insects off the surface of water, in the air, or we do not know of an expedition revisiting this from the ground (Schnitzler et al., 1994). In the wet locality. Artibeus jamaicensis was collected in season, insects represent the majority of the diet the same roost, but was not likely roosting in of N. leporinus, where fish are the major dietary the immediate vicinity of N. leporinus (Table 1). component in the dry season (Brooke, 1994). The Fossils are known from Cuba, Isla de Pinos, Puerto diet of N. leporinus may also vary between the Rico, and Antigua, but there are no fossil records sexes. In Brazil, females feed solely on insects of N. leporinus in The Bahamas (Morgan, 2001), during the spring (October–December), potentially suggesting that this species recently colonized The corresponding with lactation (Bordignon, 2006). Bahamas. Due to a paucity of information, we do During this time, males feed on fish, insects, and not know the current status of N. leporinus in The crustaceans (Bordignon, 2006). Bahamas. Mating occurs in November and December While N. leporinus has only been found in and females give birth from April to June (Hood and a cave in The Bahamas, in other parts of its range Jones, 1984). Females give birth to a single young it also roosts in abandoned buildings, hollow during each pregnancy and only leave their pup to trees and rock crevices that have a fairly stable forage at night. Pups are able to fly once they reach temperature and high humidity (Brooke, 1997; nearly adult size at 1 to 2 months of age (Silva- Silva-Taboada, 1979). Normally, they roost in Taboada, 1979; Brooke, 1997). Births may be timed close contact with each other in small groups of with an increase in insect abundance during the wet less than 30 individuals. The groups can be harems, season, allowing lactation to occur during months with one or two males and the rest female, or in of highest resource availability (Brooke, 1997). In bachelor colonies where all individuals are young some cases, females may become pregnant for a males (Brooke, 1997). There can be multiple second time in a single year, following the birth small groups in a single roost (Brooke, 1997). of their first young, but this is not true for every Females in the same roosting group form tightly population (Hood and Jones, 1984; Gannon et al., linked social groups that are sometimes maintained 2005). SPEER ET AL.: Bats of the Bahamas: natural history and conservation 57

Figure 4. Presence/absence map of Noctilio leporinus, Artibeus jamaicensis, and Monophyllus redmani in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which A. jamaicensis is found are indicated with black hatching. Islands on which N. leporinus and A. jamaicensis are found are indicated by cross-hatching. Presence of M. redmaniLVLQGLFDWHGE\EODFN¿OO Photographs of M. redmani (A), N. leporinus (B), and A. jamaicensis (C) by J. A. Soto-Centeno. 58 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Noctilio leporinus is classified as “least 1). Noctilio leporinus was collected from the same concern” according to the IUCN (2014) due to its locality on Inagua, but neither bat is known to form widespread distribution and population abundance. mixed species colonies (Silva-Taboada, 1979). In However, our knowledge of N. leporinus in The the West Indies, fossils have been documented in Bahamas is based on individuals collected almost Cuba, Isla de Pinos, Jamaica, Hispaniola, Ile de la 50 years ago from a single locality, and the status of Gonâve (Haiti), and Puerto Rico (Morgan, 2001). current populations is unknown. Given the fidelity Molecular analyses and fossil data suggest that of roosting groups at a single site across multiple A. jamaicensis colonized the West Indies through years, we recommend preservation of cave and Central America during the Pleistocene, and forest habitats to minimize disturbance of colonies. began inhabiting the southern Bahamas relatively Without further fieldwork to document populations recently (Morgan, 2001, Larsen et al., 2007, Ruiz of N. leporinus in The Bahamas, we are unable to et al., 2013). assess threats to this species. Artibeus jamaicensis occurs throughout Phyllostomidae (Leaf-Nosed Bats) many habitat types including evergreen forests, Artibeus jamaicensis (Leach, 1821) cloud forests, and arid habitats (Ortega and Castro- Arellano, 2001; Gannon et al., 2005). On the Common Name.—Jamaican Fruit Bat mainland and Cuba, A. jamaicensis can be found in Type locality.—“Jamaica” (Leach, 1821:75) Conservation Status.—“least concern” trunks and foliage of trees, palm leaf tents, caves and (IUCN, 2014) manmade structures (Silva-Taboada, 1979; Ortega Artibeus jamaicensis is a medium to large and Castro-Arellano, 2001). However, in Puerto bat (ca. 36–48 g) and one of only two frugivorous Rico, A. jamaicensis has only been documented bats found on The Bahamas along with Erophylla in manmade structures and caves (Gannon et al., sezekorni. The face is characterized by a prominent 2005). In The Bahamas, A. jamaicensis has only nose-leaf and white stripes above and below the been documented from caves. eyes, which are more defined in populations with In caves, juveniles roost solitarily or in dark pelage (Fig. 4C; Ortega and Castro-Arellano, small, unstructured groups in exposed areas or 2001; Gannon et al., 2005). The ears are broad and near solution cavities (Kunz et al., 1983; Ortega triangular separated at the base, and the tragus is and Arita, 1999). Adults form harems consisting of short with 4–5 small serrations on the outer border several females and their offspring are defended by (Ortega and Castro-Arellano, 2001). The wings one or two males (Kunz et al., 1983; Ortega and are broad and dark gray, and the tail membrane Arita, 1999). Dominant males sire the majority is thin and U-shaped when stretched out (Ortega of offspring in harem groups and maintain their and Castro-Arellano, 2001; Gannon et al., 2005). dominance status for two reproductive seasons Pelage ranges from ashy gray to brown to black and (Ortega et al., 2003). Harems are relatively stable, is white at the base (Ortega and Castro-Arellano, but females often change harems between periods 2001; Gannon et al., 2005). Unlike other bats on of reproduction (Ortega and Arita, 1999). Females The Bahamas, A. jamaicensis has no external tail give birth to one pup twice per year from March to (Ortega and Castro-Arellano, 2001). It is unlikely July, which coincides with the rainy season when that A. jamaicensis would be confused with any fruits are more abundant (Gannon et al., 2005). other bat in The Bahamas. Artibeus jamaicensis is a generalist frugivore, Artibeus jamaicensis is distributed from but also consumes other plant materials, including Mexico and Central America to northwest South pollen, flowers, leaves, or nectar (Ortega and Castro- America, and is found throughout the Greater Arellano, 2001; Gannon et al., 2005). Figs are often and Lesser Antilles (Larsen et al., 2007). In The the primary fruit consumed by A. jamaicensis, Bahamas, A. jamaicensis has been collected from but dietary composition varies geographically. In cave localities on Inagua and Mayaguana (Table Puerto Rico, trumpet tree, turkey berry, jag, almond, SPEER ET AL.: Bats of the Bahamas: natural history and conservation 59 and elder were the most commonly consumed fruits obs.). The tail membrane is narrow and U-shaped (Gannon et al., 2005). In Panamá, A. jamaicensis with the tail extending only slightly beyond the begins foraging after sunset and returns before edge of the membrane. Wing morphology of E. dawn, but may return to the day roost on especially sezekorni is indicative of maneuverable but slow moonlit nights as a predator avoidance technique flight (Jennings et al., 2004). Pelage is white at the (Morrison, 1978). However, this behavior is not base and tipped with brown (Baker et al., 1978). exhibited by individuals on Puerto Rico, which is Hair on the face and head is uniform in color and not inhabited by many of the nocturnal predators lighter than hair on the body (Baker et al., 1978). E. that prey on bats (Gannon et al., 2005). While sezekorni is easily distinguished from other bats in foraging, A. jamaicensis carries fruit or plant The Bahamas by its small nose-leaf and blonde or material to a proximal feeding roost or back to light brown coloration. the day roost to eat, subsequently dispersing seeds The genus Erophylla is endemic to the West from the parent plant (Silva-Taboada, 1979; Ortega Indies and consists of two species: E. sezekorni and Castro-Arellano, 2001; Gannon et al., 2005). distributed in Cuba, The Bahamas, Jamaica, In some regions, A. jamaicensis is known to visit Turks and Caicos, and the Cayman Islands; and the same fruiting plant on consecutive nights, but E. bombifrons, which occurs on Puerto Rico in Puerto Rico and potentially The Bahamas, which and Hispaniola (Baker et al.,1978; Timm and have low fruiting plant density, A. jamaicensis does Genoways, 2003). Until recently, there was not revisit foraging sites on consecutive nights considerable debate on the recognition of these (Gannon et al., 2005). two species and Baker et al. (1978) considered E. According to IUCN (2014), A. jamaicensis bombifrons a subspecies of E. sezekorni. Because is of “least concern” across their range. However, E. bombifrons and E. sezekorni are closely rela- this species only occurs on Inagua and Mayaguana ted and have similar ecological habits, we use in The Bahamas and the status of these populations information collected from E. bombifrons to infer is not known. 1977 was the last time A. jamaicensis aspects of the natural history of E. sezekorni where was documented on Inagua and also the last data is sparse. In The Bahamas, E. sezekorni is time any bats were collected from this island. A. the most widespread bat, found on the Abacos, jamaicensis transports seeds of the fruit it eats into Andros, Cat Island, Crooked and Acklins Islands, disturbed areas, preserving populations of this bat Eleuthera, Great Exuma, Grand Bahama, Great can increase the health of the ecosystems on Inagua Inagua, Long Island, Mayaguana, New Providence, and Mayaguana. Future surveys in the Bahamas are and San Salvador (Table 1; Fig. 5; Morgan, 1989). especially needed in the southern islands, whose It is known from fossil deposits on Great Abaco, bat populations have not been examined in over 30 Andros, the Exumas, and New Providence, as well years. as Cuba, and Jamaica (Morgan, 2001). Notably, Erophylla sezekorni (Gundlach, 1861) E. sezekorni populations persist on every island Common Name.—Buffy Flower Bat where they are documented in the fossil record, Type Locality.—“Cuba” (Gundlach, 1861) meaning this species has not undergone a single Conservation Status.—“least concern” local extinction in The Bahamas (Morgan, 1989). (IUCN, 2014) Little is known about the habitat preference Erophylla sezekorni is a medium-sized bat of E. sezekorni. It has mainly been captured in (ca. 14–18g) with a flat forehead and elongated, lowland areas of variable vegetation (Baker et al., conical snout (Fig. 5; Baker et al. 1978). Its nose- 1978). E. sezekorni has been vouchered mainly leaf is often notched at the tip and is generally from caves. In Grand Bahama, it has been collected less prominent than that of other phyllostomids from one abandoned building (Table 1) and its (Baker et al., 1978). The bottom lip is split and feces have been observed in an abandoned hotel the tongue extends through this slit (Speer pers. (Soto-Centeno and Speer, pers. obs.), suggesting 60 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Figure 5. Presence/absence map of Erophylla sezekorni in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which E. sezekorni is found are indicated with black fill. Photograph ofE. sezekorni by J. A. Soto-Centeno. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 61 these bats might opportunistically roost in man- Erophylla sezekorni is listed as a species made structures. E. sezekorni is sensitive to roost of “least concern” by IUCN (2014) due to its temperature and humidity changes and prefers large population and widespread distribution. cooler portions of hot caves or cavities of cooler E. sezekorni is one of three bats species in The caves, but roosts in variable levels of light (Silva- Bahamas that is currently found on all islands Taboada, 1979; Murray and Fleming, 2008). In a where it has been recorded in the fossil record. This study conducted on E. bombifrons (referred to as suggests that populations have withstood climate E. sezekorni, op. cit.), individuals preferred stable and habitat changes. More information specific to ambient cave temperatures 25–28° C and only Bahamian populations is necessary to assess the roosted in tight clusters in cooler temperatures risks that temperature sensitivity, habitat loss, and (Rodriguez-Durán and Soto-Centeno, 2003). This roost disturbance potentially pose. suggests that members of the genus Erophylla Macrotus waterhousii (Gray, 1843) are likely sensitive to overheating (Rodriguez- Common Name.—Big-eared bat or Water- Durán and Soto-Centeno, 2003). In The Bahamas house’s leaf-nosed bat and Caymans, colonies range in size from a few Type Locality.—“Hayti” (Gray, 1843) individuals to a few hundred (Hall et al., 1998; Conservation Status.—“least concern” Murray and Fleming, 2008). (IUCN, 2014) Originally thought to be predominantly Macrotus waterhousii is one of the most nectarivorous, the diet of E. sezekorni consists widespread bats of the Greater Antilles and The of insects, fruits, and nectar (Silva-Taboada, Bahamas along with Erophylla sezekorni (Koopman 1979; Soto-Centeno and Kurta, 2006). Dietary et al., 1957). M. waterhousii is a medium-to-large studies based on analysis of feces suggest that sized bat with broad wings and a large tail membrane E. bombifrons in Puerto Rico has an omnivorous (Anderson, 1969). Pelage is typically uniformly diet (Soto-Centeno and Kurta, 2006). Similarly, brown-gray across the body with Bahamian Silva-Taboada (1979) found E. sezekorni in Cuba populations having lighter pelage than other West consumed insects in addition to fruit and nectar. Indian populations (Anderson, 1969; Buden, Recent evidence shows that although insects are 1975a). M. waterhousii can be easily distinguished consumed frequently, fruit and nectar play a greater from other Bahamian bats by its distinct nose leaf role in the diet of this species (Soto-Centeno et al., and large ears (Fig. 6; Buden, 1975a). 2014). Macrotus waterhousii occurs on the Females give birth to one pup per year, with mainland from the southwestern United States, gestation occurring from February to May and through western Mexico to Guatemala, and in the lactation occurring from June through September West Indies in Cuba, Hispaniola, Jamaica, Turks (Silva-Taboada, 1979). On the Exuma Islands, E. and Caicos, The Bahamas, and the Cayman Islands sezekorni formed mixed-gender groups that were (Anderson and Nelson, 1965). There are two maintained during mating and the maternity season subspecies in the West Indies, with M. w. minor (Murray and Fleming, 2008). Males use wing and occurring on Cuba, the northern islands of the Great vocal displays, but non-displaying males were Bahama Bank (as far south as Long Island), and the shown to father just as many young as displaying Cayman Islands and M. w. waterhousii occurring males (Murray and Fleming, 2008). This suggests on the southern islands of the Great Bahama Bank, that other strategies to increase mating success Jamaica, and Hispaniola (Buden, 1975a). Genetic are likely employed by males to attract females analyses suggest that there has been historic gene (Murray and Fleming, 2008). E. sezekorni employs flow between populations on Cuba and those on a promiscuous mating strategy, with males and Hispaniola, Jamaica, and islands of The Bahamas females in a small group mating multiple times (Muscarella et al., 2011). It is hypothesized that (Murray and Fleming, 2008). there is little recent gene flow between populations 62 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Figure 6. Presence/absence map of Macrotus waterhousii in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which M. waterhousii is found are indicated ZLWKEODFN¿OO3KRWRJUDSKRIM. waterhousii by J. A. Soto-Centeno. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 63 on different islands (Muscarella et al., 2011). As visits a nocturnal roost to eat rather than consuming populations on each island are distinct from those its prey directly at the capture site (Silva-Taboada, on neighboring islands, M. waterhousii may be 1979). Nocturnal roosts are visited frequently and more susceptible to local extinction events than can be caves or the external architecture of buildings bats that disperse more frequently. West Indian (Silva-Taboada, 1979). The wings of insect prey fossil records of M. waterhousii are known from are often found beneath nocturnal roosting sites Cuba, Jamaica, Hispaniola, Puerto Rico, Grand used by M. waterhousii (Silva-Taboada, 1979). In Caicos, Grand Cayman, Cayman Brac, Anguilla, The Bahamas, wings of cockroaches and moths Barbuda, and from The Bahamas in Great Abaco, have been found in hypothesized nocturnal roosts Andros, the Exumas, and New Providence (Morgan, (Albury, pers. obs.) It has also been suggested 2001). Based on current distributions (Fig. 6B), M. that M. waterhousii consumes fruit in some waterhousii has become locally extinct in Puerto portions of its range, but this is not evidenced in Rico, Anguilla, and Barbuda, but is currently Cuban populations and is unlikely in Bahamian found on all islands in The Bahamas where it has populations (Silva-Taboada, 1979). been documented in the fossil record (Morgan, Little is known about the mating system or 2001). The genetic isolation of populations and social structure of M. waterhousii, especially in the local extinctions of M. waterhousii suggest West Indian populations. Females give birth to that colonization events are rare for this species one pup a year and have a uniquely slow gestation (Muscarella et al., 2011). in which embryos undergo delayed development Macrotus waterhousii is found primarily (Anderson, 1969; Silva-Taboada, 1979). Copulation in dry areas and rarely in evergreen lowland occurs in the fall, but females do not show any forests (Reid, 1997). This bat roosts in caves and signs of pregnancy until February (Silva-Taboada, occasionally buildings, forming colonies of up to 1979). Embryos can be detected from February to 500 individuals (Reid, 1997). In The Bahamas, April and lactation occurs from March to August solitary individuals have been observed in (Silva-Taboada, 1979). Females leave pups that are abandoned buildings and caves (Albury pers. obs.). unable to fly in the cave while they hunt at night, M. waterhousii generally prefers humid, sheltered but carry pups with them when roosting during the caves, but can be found in more exposed areas day (Silva-Taboada, 1979). (Silva-Taboada, 1979). When roosting in caves, M. Macrotus waterhousii is listed as “least waterhousii tends to be found in very dark areas, concern” according to the IUCN (2014) due to but when roosting in buildings, it tolerates more large population, widespread distribution, and light (Silva-Taboada, 1979). occurrence in protected habitats. However, genetic Macrotus waterhousii is a gleaning analyses suggest that M. waterhousii populations insectivore, meaning it captures insect prey from are vulnerable to local extinction events, because a surface rather than catching insects directly from movement between populations and colonization the air. M. waterhousii typically hunts in densely events are infrequent (Muscarella et al., 2011). This foliated habitats (Emrich et al., 2013). In Cuba, its bat is already known to have suffered extirpations diet is composed primarily of moths and butterflies on several West Indian islands (Morgan, 2001). (Lepidoptera) and grasshoppers and crickets Because populations in The Bahamas may represent (Orthoptera; Silva-Taboada, 1979). However, a unique subset of M. waterhousii (Buden 1975a), in Jamaica its diet is more heavily composed of disturbance of roosting and foraging habitat may beetles (Coleoptera) and flies (Diptera; Emrich et have unforeseen negative consequences for this al., 2013). This variation in diet is likely due to bat. We recommend future research efforts to competition and resource partitioning, which can clarify the natural history of M. waterhousii in The vary across small geographic areas (Emrich et al., Bahamas and conservation efforts should seek to 2013). After M. waterhousii catches its prey, it maintain caves and dense shrub and forest habitat. 64 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Monophyllus redmani (Leach, 1821) and diet, soft-bodied insects like flies (Diptera), Common Name.—Greater Antillean long- wasps (Hymenoptera), and butterflies and moths tongued bat or Leach’s single leaf bat (Lepidoptera) also represent a significant portion Type Locality.—“Jamaica” (Leach, 1821) of the diet (Gannon et al., 2005; Soto-Centeno Conservation Status.—“least concern” and Kurta, 2006). This bat is well adapted for (IUCN, 2014) hovering, potentially enabling it to access nectar Monophyllus redmani is the only primarily more efficiently than other bats that cannot hover nectarivorous bat in The Bahamas. It is a small bat, (Norberg and Rayner, 1987; Soto-Centeno and Kurta, 2006). Its dietary focus on nectar may allow weighing ca. 8–14 g (Silva-Taboada, 1979), and is M. redmani to better recover after hurricanes, found only on islands of the Greater Antilles and The while fruit-eating bats often take longer to attain Bahamas (Homan and Jones, 1975). Its elongated pre-hurricane population levels (Soto-Centeno snout is characteristic of nectar-feeding bats, and and Kurta, 2006). In Cuba, bats with specialized distinguishes this species from other bats found on morphology for consuming nectar also tended to The Bahamas (Fig. 4A). Erophylla sezekorni also have high incidence of insect remains in their feces has an elongated snout, but it is lighter in color (Clairmont et al., 2014). Even among other nectar- than M. redmani and has a less-prominent nose specialized omnivores, dietary reliance on insects leaf, with a small ridge just behind it on the snout in M. redmani is greater than expected given its (Gannon et al., 2005). M. redmani ranges in color classification as a nectarivorous bat (Soto-Centeno from brown to gray and is normally darker dorsally et al., 2014). (Buden, 1975b; Silva-Taboada, 1979). Unlike other bats, M. redmani undergoes two Monophyllus redmani is only known from reproductive peaks per year, but females give birth Cuba, Hispaniola, Jamaica, Puerto Rico, and The to only one pup per year (Gannon et al., 2005). In Bahamas. In The Bahamas, M. redmani has only Cuba, pregnant females have been recorded mainly been documented from Crooked and Acklins from March to June and again in October (Silva- islands (Fig. 4; Buden, 1975b). Fossil evidence Taboada, 1979). Pregnant females form maternity suggests this bat previously occurred on Great colonies, roosting separately from males (Gannon Abaco, Andros, and New Providence but has et al., 2005). Females give less parental care to become locally extinct on these islands (Morgan, young than other bats and newborns are often 2001). found roosting alone (Gannon et al., 2005). Monophyllus redmani primarily roosts in hot, According to IUCN (2014), M. redmani humid caves and can be found in large colonies of is characterized as a species of “least concern” many thousands of individuals (Silva-Taboada, due to its abundance within its restricted range 1979; Gannon et al., 2005). In The Bahamas, M. and occurrence in protected habitats. While M. redmani has been seen and collected from dry redmani remains prevalent on larger islands across caves that are normally well ventilated (Buden, its range, this is not true for populations in The 1975b). These bats were found roosting in bell Bahamas. M. redmani has not been collected from holes in groups of 15–20 individuals (Buden, any of the islands (Great Abaco, Andros, and New 1975b). In Cuba and Jamaica, M. redmani has been Providence) where it has been documented in the documented in mixed species roosting groups with fossil record. The current distribution of this species Pteronotus parnellii (Silva-Taboada, 1979). in The Bahamas is limited to Crooked and Acklins, Monophyllus redmani is an omnivorous and the presence of these populations has not been bat, mainly feeding on nectar and insects, only confirmed since 1973, the year of the most recent sometimes consuming fruit (Soto-Centeno and collecting on these islands. Updated occurrence Kurta, 2006). While nectar seems to be the primary data will allow accurate conservation assessment food item according to studies of dental morphology of this species in The Bahamas. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 65

Vespertilionidae (Evening Bats) needed to fully elucidate the taxonomy of E. fuscus Eptesicus fuscus (Beauvois, 1796) in The Bahamas (Timm and Genoways, 2003). Common Name.—Big Brown Bat Eptesicus fuscus is common in urban areas in Type Locality.—Philadelphia, Pennsylvania the North American part of its range, and has been (Beauvois, 1796 ) documented in forested highlands and suburban Conservation Status.—“least concern” areas in Puerto Rico (Gannon et al., 2005; Soto- (IUCN, 2014) Centeno pers. obs.). This bat is commonly found Eptesicus fuscus is a small bat (ca. 11–23 g) roosting in buildings and caves in Cuba, but has that is widely distributed (Kurta and Baker, 1990). only occasionally been documented from buildings Pelage is dark brown on the dorsum, but slightly in The Bahamas (Table 1; Silva-Taboada, 1979; paler ventrally, and the face and ears are also Albury, pers. obs.). In caves, they roost in a similar darkly colored. Pelage is long and soft and has an habit as T. brasiliensis, with many individuals oily appearance (Fig. 7; Kurta and Baker, 1990). being tightly packed together (Gannon et al., The ears are thick, short, and rounded. The tail 2005). E. fuscus roosts in large colonies in well membrane is large and has hair at the base (Kurta ventilated, open areas of caves, or in smaller groups and Baker, 1990). E. fuscus may be confused with in solution cavities (Silva-Taboada, 1979). In Tadarida brasiliensis in The Bahamas, because Cuba, T. brasiliensis and E. fuscus are occasionally they are both small and darkly colored. However, found roosting together (Silva-Taboada, 1979). In E. fuscus is distinguishable because it does not have Hillside Grocery Cave, Long Island, 2 individuals the characteristic wrinkled lips of T. brasiliensis. In of E. fuscus were observed roosting on the outside addition, E. fuscus has a prominent, smooth snout, edge of a colony of T. brasiliensis (Soto-Centeno, and its tail only slightly extends beyond the edge of pers. obs.). the tail membrane (Kurta and Baker, 1990). Eptesicus fuscus is an aerial insectivore The range of E. fuscus is nearly as broad and is known for the incredible precision of its as that of T. brasiliensis, reaching from southern echolocation (Kurta and Baker, 1990). E. fuscus Canada to very northern edge of South America, has long, narrow wings and is less maneuverable and includes islands of the Greater Antilles, The than other species, typically flying above the Bahamas, Grand Cayman, Barbuda, and Dominica canopy or in open areas (Kurta and Baker, 1990). (Kurta and Baker, 1990; Gannon et al., 2005). It mainly consumes beetles (Coleoptera), but also In The Bahamas, E. fuscus is known from fossil preys on cockroaches (Dictyoptera), flies (Diptera), deposits on Great Abaco, Andros, the Exumas, wasps (Hymenoptera), butterflies (Lepidoptera), and New Providence (Morgan, 2001). E. fuscus is and lacewings (Neuroptera; Silva-Taboada, 1979). still present on these islands, in addition to Long There have been no dietary studies on these bats Island, San Salvador, and Crooked and Acklins in The Bahamas. Foraging occurs throughout the (Table 1; Fig. 7). Based on morphological data, night, beginning soon after sunset and ending just E. fuscus in The Bahamas has been split into two before sunrise (Silva-Taboada, 1979). In mainland subspecies (Buden, 1985, 1986). E. f. bahamensis portions of their range, bats fly 1–2 km from occurs on New Providence and San Salvador and roost sites to foraging areas with total flight times E. f. dutertreus is found on Andros, the Exumas, averaging 100 min. per night (Kurta and Baker, Crooked and Acklins, and Long Island, and 1990), but in Cuba they stay close to the roost intermediate specimens are found on San Salvador (Silva-Taboada, 1979). (Timm and Genoways, 2003). However, more In cooler portions of its range, E. fuscus recent morphological analyses of specimens in hibernates in response to food shortage that occurs the Albert Schwartz Collection suggest that there in the winter when it is too cold for insects to survive is little difference between these subspecies, and (Kurta and Baker, 1990; Gannon et al., 2005). In further genetic and morphometric analyses are the tropics, there is no evidence of hibernation, but 66 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Figure 7. Presence/absence map of Eptesicus fuscus in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which E. fuscus is found are indicated with black fill. Photograph of E. fuscus by J. A. Soto-Centeno. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 67 these bats may become torpid if temperatures drop North America and the Greater Antilles, including below 20o C rather than leaving the roost to hunt The Bahamas and Turks and Caicos (Koopman, (Silva-Taboada, 1979). 1993). However, bats found on Hispaniola and Females give birth once a year to twins and Puerto Rico are smaller and probably belong to occasionally single pups (Gannon et al., 2005). a different species (i.e., L. minor; Miller, 1931). Gestation occurs from March to May in Cuba and Allen (1905) examined a single specimen from this seems to be true for populations in The Bahamas Nassau, New Providence, which he considered based on limited sampling (Silva-Taboada, 1979; “indistinguishable in color and proportions” of Timm and Genoways, 2003). While giving birth, the body and skull from L. borealis in Florida, females roost with their heads up and use their tail USA. Allen and Sanborn (1937) examined a single membrane to catch pups as they are born (Gannon specimen from Cat Island, which they assigned to et al., 2005). Lactation occurs in June (Silva- L. minor. This specimen was not an adult, yet they Taboada, 1979) and juveniles begin to fly at 18– still considered it different than L. borealis. Given 35 days old (Kurta and Baker, 1990). The female the paucity of the specimens examined, Koopman does not carry her offspring while hunting at night et al. (1957) suggested the use of L. borealis minor (Silva-Taboada, 1979). More research is necessary for the populations on The Bahamas, Hispaniola, on populations in the West Indies to understand and Puerto Rico. Until further evidence is social structure. uncovered and to avoid confusion with its mainland Eptesicus fuscus is indicated as of “least congenerics, we adopt the name L. minor to refer to concern” by IUCN (2014), because of its wide the small species of insular red bat that occurs on distribution, high population size, occurrence in The Bahamas, Hispaniola, Puerto Rico, and Turks protected habitats, and resilience against some and Caicos (Miller, 1931; Simmons, 2005). degree of habitat modification. E. fuscus has not Lasiurus minor is a small bat (ca. 6–11 g) suffered any local extinctions in The Bahamas, with short rounded ears and a small rostrum (Fig. suggesting that populations have persisted through 8; Miller, 1931; Gannon et al., 2005; Rodríguez- past climate and habitat change. As this bat has the Durán and Christenson, 2012). This bat has dense propensity to roost in buildings and is commonly fur that extends from the body onto to the large found in urban areas, it may be at risk from improper wing membrane, which appears visibly furred. removal or extermination when found in man- Perhaps the most striking characteristic of L. minor made structures. Instructions for excluding bats is its distinctive reddish or yellow-orange pelage, from buildings can be found at Bat Conservation which in some individuals show small patches of International (http://www.batcon.org/resources/ white hair giving it a frosted appearance (Allen for-specific-issues/bats-in-buildings). We recom- and Sanborn, 1937; Gannon et al., 2005). The mend future work to investigate the genetics and dark brown wings of L. minor show a marked ecology of the two proposed subspecies of E. contrast with the reddish coloration of the fur. The fuscus occurring in The Bahamas. coloration and densely furred tail membrane make Lasiurus minor (Miller, 1931) this bat easy to distinguish among other Bahamian Common Name.—Minor Red Bat bats. However, this species is infrequently seen or Type Locality.—“Trouin Voute L’Eglise,” captured probably because of its solitary nature and Haiti (Miller ,1931) preference for foliage and tree roosts. Conservation Status.—“vulnerable” (IUCN, Information on L. minor in The Bahamas is 2014) limited and is based on about 15 specimens. These The validity for the use of the species name records are from Cat Island, Great Inagua, Long Lasiurus minor for red bats in The Bahamas is Island, and New Providence (Table 1; Fig. 8; Allen, uncertain. Some authors consider red bats a single 1905, 1911; Allen and Sanborn, 1937; Koopman et species (Lasiurus borealis) that expanded across al., 1957). Two road kill specimens collected from 68 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Figure 8. Presence/absence map of Lasiurus minor in The Bahamas. The Bahamas are indicated in gray. See Figure 1 for geographic reference. The islands on which L. minor is found are indicated with EODFN¿OO3KRWRJUDSKRIL. borealis, which looks very similar to L. minor, by Merlin D. Tuttle, Bat Conservation International, www.batcon.org. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 69

Great Abaco, have been tentatively identified as L. Rodríguez-Durán (1999) documented the capture minor at the Florida Museum of Natural History. of a single lactating female carrying three pups on Fossils of L. minor have been identified from Great Puerto Rico during the month of June. Based on Abaco (Soto-Centeno and Steadman, 2015) Cuba, the appearance of the pups, he speculated that they and Hispaniola (Morgan, 2001). Because L. minor were likely born around late May. Silva-Taboada is considered rare across its distribution and in some (1979) examined embryos in February, April, and instances could be identified as L. borealis, the May and identified the earliest date of birth as late number of specimens preserved in natural history May. museum collections is difficult to determine. Lasiurus minor is considered “vulnerable” Almost nothing is known of the life history by the IUCN (2014) due to population decline and of L. minor in The Bahamas, because of the lack restricted geographic range. The solitary nature and of field observations and paucity of specimens. high reliance on densely foliaged trees for roosting L. borealis in the southeastern United States are makes L. minor susceptible to habitat changes, such solitary mammals that roost primarily in the foliage as deforestation. Population surveys using multiple of trees, making them rare to document. They often sampling techniques are required to formally assess hang by one foot and their coloration gives them the status of L. minor in The Bahamas and would the appearance of dead leaves (Harvey et al., 2011). greatly increase the available knowledge on the No roost trees of L. minor have been observed on natural history of this species. the islands, but on the mainland, red bats roost in trees with dense foliage and broad leaves (Gannon CONCLUSIONS et al., 2005). While most species of bat in The Bahamas are Lasiurus minor, like its congenerics, is classified as “least concern” by the IUCN (2014), insectivorous and probably feeds on beetles this may not be a good indicator of the local health (Coleoptera), flies (Diptera), leafhoppers of bat populations. The IUCN aims to prevent the (Homoptera), termites (Isoptera), and moths global extinction of species and does not specifically (Lepidoptera) as evidenced from data of one assess local extinction risks (www.iucnredlist.org). individual from Puerto Rico (Rodríguez-Durán, Global extinction risk may not accurately reflect 1999). Wings of L. minor are elongated and the conservation needs of regional populations. narrow, which give this species a swift but not For example, T. brasiliensis is widespread in the very maneuverable flight. Therefore, L. minor can Americas and is classified as a species of “least typically be found foraging in open areas above concern” (IUCN, 2014), but this classification may the tree canopy or along forest edges. Hickey and not reflect risk to populations in The Bahamas. Fenton (1990) reported L. borealis foraging around IUCN classifications are more reflective of local streetlights where concentration of insects was conservation needs in the case of endemic species, higher. A red bat (likely L. minor) was observed like C. tumidifrons, which is limited to The Bahamas foraging around a streetlight in Conch Bar, Turks and is classified as “near threatened” (IUCN, and Caicos (Soto-Centeno, pers. obs.). Therefore, it 2014). Bat populations in The Bahamas may be is possible that island populations of L. minor also at a greater risk than throughout the rest of their takes advantage of streetlights as foraging habitat. range, especially given the widespread distribution Reproduction of L. minor on islands is not and abundance of some species on larger islands well known. Female L. borealis give birth to three and the mainland. Bats represent two-thirds of the pups on average and carry their pups with them mammals in The Bahamas (excluding domesticated while flying (Harvey et al., 2011). L. minor from species; Buden, 1986). Local extinction of just one Puerto Rico seem also to be able to give birth or two species will result in a significant loss of to multiple pups, where most species of bats in mammalian biodiversity on these islands. the West Indies give birth to one pup per year. Islands of the Great Bahama Bank support 70 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) more bat species (7 of 10 species) than islands of surprising given all bat fossils collected in The the Little Bahama Bank (6 of 10), Crooked and Bahamas to date have come from caves and blue Acklins (5 of 10), Inagua (5 of 10), Mayaguana holes, which are inundated caves. Four species of bat (3 of 10), and San Salvador (3 of 10; Table 1). (Chilonatalus tumidifrons, Monophyllus redmani, Species richness does not seem to correlate Nyctiellus lepidus, and Tadarida brasiliensis) were with island area, but may reflect the effects of once more widespread in The Bahamas than they Pleistocene glacial-interglacial cycles on island are today (Morgan, 2001:table 1). M. redmani is area and habitat (Morgan, 1989). For example, the not found on any of the islands where fossils of this Abacos and Long Island are each inhabited by 6 species have been documented and C. tumidifrons of 10 bat species, while larger islands by area, like is found on one of the four islands it previously Andros and Inagua, are inhabited by 3 and 5 bat inhabited. There is no fossil evidence of Artibeus species, respectively. In the case of Inagua, it may jamaicensis and Noctilio leporinus, which are be that there has not yet been enough sampling currently only found on Inagua and Mayaguana. conducted on the island to categorize all of the Fossil deposits have not been documented from bat diversity there. Habitat may also influence these islands and it is possible that A. jamaicensis bat species diversity (Pregill & Olson, 1981). The and N. leporinus have occupied these islands Abacos and Grand Bahama in the north receive historically without colonizing other Bahamian the highest rainfall, with the lowest mean annual islands. However, A. jamaicensis is documented rainfall recorded in southern islands of Crooked from fossil and extant collections on Cuba, Jamaica, and Acklins, Mayaguana, and Inagua (Buchan, Hispaniola, and Puerto Rico (Morgan, 2001:table 2000). The habitat of southern islands is dominated 1) and it is more likely that it recently colonized by coppice and scrub vegetation, but lacks the pine Inagua and Mayaguana. N. leporinus is known woodlands and palmetto understory common in from fossil deposits on Cuba and Puerto Rico and the northern Bahamas (Pregill and Olson, 1981). is extant on these islands, as well as Jamaica and However, Grand Bahama has considerably fewer Hispaniola. As it has only been collected from a bat species (2 of 10) than Great Abaco (6 of 10), single locality in The Bahamas, but is widespread despite having similar habitat and rainfall. Perhaps across the Greater Antilles, N. leporinus is likely distribution and availability of roosting habitat also a recent colonizer. corresponds more closely with bat species diversity. Comparing the fossil record to current Comprehensive information on locality of caves presence/absence data, the Bahamas lost more and use by bats is not known, but N. A. Albury in bat species than other islands of the West Indies conjunction with other Bahamian collaborators is (Morgan, 2001). Pleistocene glacial-interglacial working to accumulate these data. cycles significantly altered the available habitat Sixteen species of fossil bats have been for terrestrial organisms in The Bahamas and these identified from cave deposits in Great Abaco, changes may have led to bat extinctions (Dávalos Andros, Cat Island, the Exumas, and New and Russell, 2012). However, recent evidence Providence (Morgan, 2001:table 1). Of these, indicates human colonization of the West Indies only four (Eptesicus fuscus, Erophylla sezekorni, played a driving role in bat extinctions (Soto- Lasiurus minor and Macrotus waterhousii) inhabit Centeno and Steadman, 2015). In the past, humans all of the Bahamian islands where they have been have been linked to other major extinctions in the documented in the fossil record (Morgan, 2001; West Indies (Steadman et al., 2005), and currently Soto-Centeno and Steadman, 2015). However, the effects of land development and habitat loss in fossils of L. minor have only been documented from The Bahamas are threatening the local extinction of one locality on Great Abaco, so its historic range several others. The Bahamian hutia (Geocapromys is unknown. L. minor is a tree-roosting species, ingrahami, Order Rodentia) was once widespread so the paucity of data from the fossil record is not and abundant across The Bahamas, but hunting by SPEER ET AL.: Bats of the Bahamas: natural history and conservation 71 native peoples likely caused their local extinction focused on educating school children, providing (Morgan, 1989). Extant populations are now information for employees of museums and nature limited to East Plana Cay (near Acklins Island) centers, and updating community members on and introduced populations in the Exuma Land our research. J. A. Soto-Centeno has also created and Sea Park on Little Wax Cay and Warderick an informational brochure on the “Common Bats Wells Cay (Morgan, 1989). Increasing human of the Bahamas,” which can be found online at populations and lack of habitat protection have also the Friends of the Environment website (http:// influenced the once widespread Bahamas Parrot www.friendsoftheenvironment.org/wp-content/ (Order Psittaciformes: Amazona leucocephala uploads/2012/08/Bat-brochure.pdf) and has been bahamensis; Williams and Steadman, 2001; distributed to nature centers and museums on Russello et al., 2010). Where The Bahamas Parrot several islands. These efforts are motivated by our has been documented through fossils on Crooked desire to change negative public perception of bats and Acklins, New Providence, Long Island, Fortune in The Bahamas through education and outreach Cay, and Grand Turk, it is currently limited to designed to encourage public participation in populations on Inagua and Great Abaco (Williams conservation efforts. and Steadman 2001). As preservation of Bahamian Beyond the importance of public support, terrestrial mammals, specifically bats, is dependent conservation of caves and coppice and evergreen on minimizing human effects, engaging the public forest habitat will protect roosting and foraging is key to successful conservation efforts. grounds used by all 10 species of Bahamian Education and outreach have been used bats. National parks have been established for effectively in the past in The Bahamas to increase the purpose of maintaining these habitats in The public awareness of and concern for pertinent Bahamas, but most parks are focused on Grand conservation issues. One locally famous example Bahama, the Abacos, New Providence and the is the “Size Matters” campaign implemented by Exuma Islands (excluding wetland parks). As each Friends of the Environment and Rare Planet on island is unique in terms of species composition, Great Abaco (http://www.friendsoftheenvironment. not all of the bat species in The Bahamas are org/page-1/marine-resources/size-matters). This protected by these national parks. The majority of program aimed to prevent further decline of the the bat species diversity occurs south of the Little local spiny lobster (Panulirus argus) population Bahama Bank (Grand Bahama and the Abacos; by educating fishermen, restaurant owners, and Table 1) and additional protected areas on the the public on the negative effects of harvesting Great Bahama Bank, Crooked and Acklins, San undersized lobsters. The insinuating logo, “Size Salvador, Inagua, and Mayaguana are necessary Matters,” was printed on rulers that were handed for preservation of the complete diversity of bats out to fishermen, an educational song by a local in this archipelago. Decreasing disturbance of cave musician (titled “Size Matters”) was played on ecosystems is also important to preservation of bat the radio, and workshops were held to educate populations. It is common to find trash and graffiti community members. Data on the health of the in caves, indicating destructive human use of these spiny lobster population is insufficient, but follow- ecosystems. In the past, most accessible caves up polling of community members indicates were mined for guano as a commercial export and increased awareness of the problem. Outreach to agricultural fertilizer. Unfortunately, guano was increase public awareness has also been employed not sifted and much of the fossil record of The to aid conservation of mangroves (My Island, My Bahamas was lost due to removal of sediments Mangrove) and conch (Community Conch, www. (Mylroie and Mylroie and, 2013). While guano communityconch.org). Presentations that we mining is not currently a commercial venture in have conducted on Great Abaco, Grand Bahama, The Bahamas, guano continues to be harvested Eleuthera, Long Island, and New Providence have from caves on a small scale and can force bats 72 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) to abandon cave roosts. However, guano mining Kakuk, David Knowles, d’Shan Maycock, Kris can be performed sustainably by minimizing the Newman and Jean Pierce, Judd and Dale Rosen, disturbance of bat roosting colonies (information David Steadman (DWS), and Ellsworth Weir for available through IUCN, http://www.batcon.org/ invaluable guidance and local knowledge for which pdfs/GuanoGuidelinesVersion1.pdf). Human im- the authors are incredibly grateful. The Theodore pact on bat habitat may be minimal on islands with Roosevelt Memorial Grant funded fieldwork by fewer inhabitants, but human effects are likely to KAS and work by JAS-C was funded by Theodore increase over time. By preserving roosting and Roosevelt and Gerstner Scholar postdoctoral foraging habitat used by bats, we can maintain the fellowships at AMNH. Fieldwork was also partially health of these populations in The Bahamas. funded by U.S. National Science Foundation grant Few of the studies reviewed for this (BCS 1118369) awarded to DWS. The project was publication were conducted specifically on bat supported in part by a UF Faculty Enhancement populations in The Bahamas. Given the unique Award to DLR. Printed copies of this manuscript climate, environmental pressures, and geologic were paid for by a UF Women’s Club Scholarship history of these islands in comparison to other awarded to KAS. Thanks also to Dr. Bret Boyd for islands in the West Indies, it is necessary to conduct comments on early drafts of this manuscript. studies focused on bats in The Bahamas. Our current knowledge of Bahamian bat populations LITERATURE CITED is based on observational information, and there Allen, G. M. 1905. Notes on Bahama bats. have not been thorough examinations of the Proceedings of the Biological Society of natural history, ecology, and genetic diversity of Washington 18:65–71. bats on these islands. The information reviewed Allen, G.M. 1911. Mammals of the West Indies. in this publication can serve as a foundation from Bulletin of the Museum of Comparitive which future research can build. Minimal available Zoology 54:175–263. data prevents accurate assessment of the health Allen, G. M., and C. C. Sanborn. 1937. Notes on bats and trends of bat populations, making it difficult from The Bahamas. Journal of Mammalogy to develop specific conservation solutions. By 18:225–228. consolidating, updating, and verifying all known Altenbach, J. S. 1989. Prey capture by the fishing locality and natural history information on bats bats Noctilio leporinus and Myotis vivesi. in The Bahamas, areas of missing information Journal of Mammalogy 70(2):421–424. are identified and can be targeted in future Andersen, K. W. 1994. On the status of bats on research. Through this research we will better our Andros and San Salvador island, Bahamas. understanding of Bahamian fauna and increase Proceedings of the Fifth Symposium on the public engagement in a shared initiative to preserve Natural History of The Bahamas 5:1–4. the natural biodiversity of these islands. Anderson, S. 1969. Macrotus waterhousii. Mammalian Species 1:1–4. ACKNOWLEDGEMENTS Anderson, S., and C. E. Nelson. 1965. A systematic The authors thank The National Museum of The revision of Macrotus (Chiroptera). American Bahamas/Antiquities, Monuments and Museums Museum Novitates no. 2212. Corporation, Abaco-based NGO, Friends of Baker, R. J., P. V. August, and A. A. Stueter. 1978. the Environment on Abaco, the Rand Nature Erophylla sezekorni. Mammalian Species Center, the Long Island Museum, and the Exuma 115:1–5. Foundation for providing opportunities to conduct Belotto, A., L. F. Leanes, M. C. Schneider, H. fieldwork and outreach. We also thank Michael Tamayo, and E. Correa. 2005. Overview of Albury, Allison Ball, Cecilia Bodie, Catherine rabies in the Americas. Virus Research 111: Booker, Patsy Cartwright, Jane Earlie, Brian 5–12. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 73

Bordignon, M. O. 2006. Diet of the fishing bat explains bat extinction in the Caribbean. Noctilio leporinus (Linnaeus) (Mammalia, Ecology and Evolution 2(12):3045–3051. Chiroptera) in a mangrove area of southern Emrich, M. A., E. L. Clare, W. O. C. Symondson, S. Brazil. Revista Brasileira de Zoologia 23(1): E. Koenig, and M. B. Fenton. 2013. Resource 256–260. partitioning by insectivorous bats in Jamaica. Brooke, A. P. 1994. Diet of the fishing bat,Noctilio Molecular Ecology 23(15):3648–3656. leporinus (Chiroptera: Noctilionidae). Journal Fleming, T. H., and P. A. Racey, eds. 2010. Pp. of Mammalogy 75(1): 212–218. 1-17 in Island Bats: Evolution, Ecology, and Brooke, A. P. 1997. Social organization and Conservation. University of Chicago Press, foraging behaviour of the fishing bat, Noctilio Chicago, Illinois. leporinus (Chiroptera: Noctilionidae). Ethol- Fleming, T. H., and P. Eby. 2003. Ecology of bat ogy 103(5):421–436. migration. Pp. 156-208 in T. H. Kunz and M. Buchan, K. C. 2000. The Bahamas. Marine B. Fenton, eds. Bat Ecology. University of Pollution Bulletin 41(1):94–111. Chicago Press, Chicago, Illinois. Buden, D. W. 1975a. A taxonomic and Gannon, M. R., A. Kurta, A. Rodríguez-Durán, and zoogeographic appraisal of the big-eared bat M. R. Willig. 2005. Bats of Puerto Rico. Texas (Macrotus waterhousii Gray) in the West Tech University Press, Lubbock, Texas, 224 p. Indies. Journal of Mammalogy 56(4):758–769. Gustin, M. K., and G. F. McCracken. 1987. Scent Buden, D. W. 1975b. Monophyllus redmani Leach recognition between females and pups in the (Chiroptera) from The Bahamas, with notes on bat Tadarida brasiliensis mexicana. Animal variation in the species. Journal of Mammalogy Behaviour 35(1):13–19. 56(2):369–377. Hall, J. S., C. W. Stihler, and P. L. Dougherty. Buden, D. W. 1985. Additional records of bats 1998. Bat populations on San Salvador and from The Bahama islands. Caribbean Journal New Providence islands. Bahamas Journal of of Science 21(2):19–25. Science 6:22–27. Buden, D. W. 1986. Distribution of mammals Harvey, M. J., J. S. Altenbach, and T. L. Best. 2011. of The Bahamas. Florida Field Naturalist Bats of the United States and Canada. The 14(3):53–63. John Hopkins University Press, Baltimore, Carew, J. L., and J. E. Mylroie. 1995. Quaternary Maryland, 199p. tectonic stability of the Bahamian archipelago: Hayssen, V. and T. H. Kunz. 1996. Allometry evidence from fossil coral reefs and flank of litter mass in bats: maternal size, wing margin caves. Quaternary Science Reviews morphology, and phylogeny. Journal of 14(2):145–153. Mammalogy 77(2):476–490. Clairmont, L., E. C. Mora, and M. B. Fenton. Hearty, P. J., and D. S. Kaufman. 2000. Whole-rock 2014. Morphology, diet and flower‐visiting aminostratigraphy and Quaternary sea-level by phyllostomid bats in Cuba. Biotropica history of The Bahamas. Quaternary Research 46(4):433–440. 54(2):163–173. Dalquest, W. W. 1950. The genera of the chiropteran Hedges, S. B. 1996. Historical biogeography of family Natalidae. Journal of Mammalogy West Indian vertebrates. Annual Review of 31(4):436–443. Ecology and Systematics 27:163–196. Dávalos, L. M. 2005. Molecular phylogeny of Hickey, M. B., and M. B. Fenton. 1990. Foraging by funnel-eared bats (Chiroptera: Natalidae), with red bats (Lasiurus borealis): Do intraspecific notes on the biogeography and conservation. chases mean territoriality? Canadian Journal Molecular Phylogenetics and Evolution of Zoology 68:2477–2482. 37(1):91–103. Hood, C. S., and J. K. Jones. 1984. Noctilio lepo- Dávalos, L. M., and A. L. Russell. 2012. Deglaciation rinus. Mammalian Species 216:1–7. 74 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Homan, J. A., and J. K. Jones. 1975. Monophyllus Kurta, A., and R. H. Baker. 1990. Eptesicus fuscus. redmani. Mammalian Species 57:1–3. Mammalian Species 356:1–10. Hutson, A. M., S. P. Mickleburgh, and P. A. Racey, Larkin, C. C., C. Kwit, J. M. Wunderle, E. H. eds. 2001. Microchiropteran bats: global status Helmer, M. H. H. Stevens, M. T. K. Roberts, survey and conservation action plan. IUCN, and D. N. Ewert. 2012. Disturbance type and Gland, Switzerland and Cambridge, United plant successional communities in Bahamian Kingdom, 258 p. dry forests. Biotropica 44(1):10–18. The IUCN Red List of Threatened Species. Version Larsen, P. A., S. R. Hoofer, M. C. Bozeman, S. C. 2014.3. www.iucnredlist.org on 03 February Pedersen, H. H. Genoways, C. J. Phillips, D. 2015. E. Pumo, and R. J. Baker. 2007. Phylogenetics Jennings, N. V., S. Parsons, K. E. Barlow, and M. and phylogeography of the Artibeus R. Gannon. 2004. Echolocation calls and wing jamaicensis complex based on cytochrome-b morphology of bats from the West Indies. Acta DNA sequences. Journal of Mammalogy Chiropterologica 6(1):75–90. 88(3):712–727. Kjellmark, E. 1996. Late Holocene climate McCracken G. F., J. K. Westbrook, V. A. Brown, change and human disturbance on Andros M. Eldridge, P. Federico, and T. H. Kunz. Island, Bahamas. Journal of Paleolimnology 2012. Bats track and exploit changes in insect 15(2):133–145. pest populations. PLoS ONE 7(8): e43839. Knight, A. J. 2008. “Bats, snakes and spiders, oh doi:10.1371/journal.pone.0043839. my!” How aesthetic and negativistic attitudes, McWilliams, L. A. 2005. Variation in diet of the and other concepts predict support for Mexican free-tailed bat (Tadarida brasiliensis mexicana). Journal of Mammalogy 86(3):599- species protection. Journal of Environmental 605. Psychology 28(1):94–103. Miller, G. S. 1903. A new nataline bat from The Koopman, K. F. 1993. Order Chiroptera. Pp. 137– Bahamas. Proceedings of The Biological 241 in D. E. Wilson and D. M. Reeder, eds. Society of Washington 16:119-20. Mammal Species of the World: A taxonomic Miller, G. S. 1905. Mammals of the Bahama and geographic reference (2nd edition). Islands. Pp. 371-386 in G. B. Shattuck, ed. The Smithsonian Institution Press, Washington, Bahama Islands. The Johns Hopkins Press, DC. New York, New York. Koopman, K. F., M. K. Hecht, and E. Ledecky- Miller, G. S. 1931. The red bats of the Greater Janecek. 1957. Notes on the mammals of The Antilles. Journal of Mammalogy 12:409–410. Bahamas with special reference to the bats. Morgan, G. S. 1989. Fossil Chiroptera and Journal of Mammalogy 38:164–174. Rodentia from the Bahamas, and the historical Kunz, T. H., P. V. August, and C. D. Burnett. biogeography of the Bahamian mammal fauna. 1983. Harem social organization in cave Pp. 685–740 in C. A. Woods, ed. Biogeography roosting Artibeus jamaicensis (Chiroptera: of the West Indies: Past, Present, and Future. Phyllostomidae). Biotropica 15(2):133–138. Sandhill Crane Press, Gainesville, Florida. Kunz, T. H., E. Braun de Torrez, D. Bauer, T. Morgan, G. S. 2001. Patterns of extinction in West Lobova, and T. H. Fleming. 2011. Ecosystem Indian bats. Pp. 369-408 in C. A. Woods and services provided by bats. Annals of the New F. E. Sergile, eds. Biogeography of the West York Academy of Sciences, 1223:1–38. Indies: Patterns and Perspectives. CRC Press, Kurta, A. and T. H. Kunz. 1987. Size of bats at birth Boca Raton, Florida. and maternal investment during pregnancy. Morrison, D. W. 1978. Foraging ecology and Symposia of the Zoological Society of London energetics of the frugivorous bat Artibeus 57:79–107. jamaicensis. Ecology 59(4):716–723. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 75

Muscarella, R. A., K. L. Murray, D. Ortt, A. L. Ortega, J., J. E. Maldonado, G. S. Wilkinson, Russell, and T. H. Fleming. 2011. Exploring H. T. Arita, and R. C. Fleischer. 2003. Male demographic, physical, and historical dominance, paternity, and relatedness in the explanations for the genetic structure of Jamaican fruit‐eating bat (Artibeus jamai- two lineages of Greater Antillean bats. PloS censis). Molecular Ecology 12(9):2409–2415. ONE 6(3): e17704. doi:10.1371/journal. Pregill, G. K., and S. L. Olson. 1981. Zoogeography pone.0017704. of West Indian vertebrates in relation to Muscarella, R., and T. H. Fleming. 2007. The role of Pleistocene climatic cycles. Annual Review of frugivorous bats in tropical forest succession. Ecology and Systematics 12:75–98. Biological Reviews 82:573–590. Reid, F. 1997. A field guide to the mammals of Murray, K. L., and T. H. Fleming. 2008. Social Central America and Southeast Mexico. structure and mating system of the buffy Oxford University Press, New York, 300 p. flower bat, Erophylla sezekorni (Chiroptera, Richards, D. A., P. L. Smart, and R. L. Edwards. Phyllostomidae). Journal of Mammalogy 1994. Maximum sea levels for the last glacial 89(6):1391–1400. period from U-series ages of submerged Mylroie, J. E. 2008. Late Quaternary sea-level speleothems. Nature 367:357–360. Rodríguez-Durán, A. 1998. Nonrandom aggre- position: Evidence from Bahamian carbonate gations and distribution of cave-dwelling deposition and dissolution cycles. Quaternary bats in Puerto Rico. Journal of Mammalogy International 183(1):61–75. 79(1):141−146. Mylroie, J. E., and J. R. Mylroie. 2013. Caves and Rodríguez-Durán, A. 1999. First record of repro- karst of the Bahama islands. Pp. 147–176 in M. ductive Lasiurus borealis minor (Miller) from J. Lace and J. E. Mylroie, eds. Coastal Karst Puerto Rico (Chiroptera). Caribbean Journal Landforms. Springer, Dordrecht, Netherlands. of Science 35:143–144. Norberg, U. M., and J. M. V. Rayner. 1987. Rodríguez-Durán, A., and K. Christenson. 2012. Ecological morphology and flight in bats Brevario sobre los murciélagos de Puerto Rico, (Mammalia; Chiroptera): wing adaptations, La Española, e Islas Vírgenes. Publicaciones flight performance, foraging strategy and Puertorriqueñas, Inc., Bayamón, Puerto Rico, echolocation. Philosophical Transactions of 98 p. the Royal Society B: Biological Sciences Rodríguez-Durán, A., and J. A. Soto-Centeno. 316(1179): 335–427. 2003. Temperature selection by tropical bats Northrop, A. R. 1910. Bahaman trip, general notes. roosting in caves. Journal of Thermal Biology Pp 1-26 in H. F. Osborn, ed. A Naturalist in 28(6):465–468. The Bahamas. The Columbia University Press, Ruiz, E. A., B. Vargas-Miranda, and G. Zúñiga. New York, New York. 2013. Late-Pleistocene phylogeography and Olson, S. L., G. K. Pregill, and W. B. Hilgartner. demographic history of two evolutionary 1990. Studies on Fossil and Extant Vertebrates lineages of Artibeus jamaicensis (Chiroptera: from San Salvador (Watling’s) Island, Phyllostomidae) in Mexico. Acta Chirop- Bahamas. Smithsonian Institution Press, terologica 15(1):19–33. Washington, D.C, 15 p. Russello, M. A., C. Stahala, D. Lalonde, K. L. Ortega, J., and H. T. Arita. 1999. Structure and Schmidt, and G. Amato. 2010. Cryptic diversity social dynamics of harem groups in Artibeus and conservation units in the Bahama parrot. jamaicensis (Chiroptera: Phyllostomidae). Conservation Genetics 11(5):1809–1821. Journal of Mammalogy 80(4):1173–1185. Schnitzler, H. U., E. K. V. Kalko, I. Kaipf, and A. Ortega, J., and I. Castro-Arellano. 2001. Artibeus D. Grinnell. 1994. Fishing and echolocation jamaicensis. Mammalian Species 662:1–9. behavior of the greater bulldog bat, Noctilio 76 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

leporinus, in the field. Behavioral Ecology and A. J. T. Jull, H. G. McDonald, C. A. Woods, M. Sociobiology 35(5):327–345. Iturralde-Vinent, and G. W. L. Hodgins. 2005. Silva-Taboada, G. 1979. Los murciélagos de Cuba. Asynchronous extinction of late Quaternary Editorial Academia, La Habana, 423 p. sloths on continents and islands. Proceedings Simmons, N. B. 2005. Order Chiroptera. Pp. 312– of the National Academy of Sciences 529 in D. E. Wilson and D. M. Reeder, eds. 102:11763–11768. Mammal Species of the World (3rd edition). Tejedor, A. 2011. Systematics of funnel-eared The John Hopkins University Press, Baltimore, bats (Chiroptera: Natalidae). Bulletin of the Maryland. American Museum of Natural History 353:1– Soto-Centeno, J. A. and A. Kurta. 2003. Description 140. of fetal and newborn brown flower bats, Tejedor, A., V. D. C. Tavares, and G. Silva-Taboada. Erophylla sezekorni (Phyllostomidae). Carib- 2005. A revision of extant Greater Antillean bean Journal of Science 39:233–234. bats of the genus Natalus. American Museum Soto-Centeno, J. A., and A. Kurta. 2006. Diet of Novitates 3493:1–23. two nectarivorous bats, Erophylla sezekorni Timm, R. M., and H. H. Genoways. 2003. and Monophyllus redmani (Phyllostomidae), West Indian mammals from the Albert on Puerto Rico. Journal of Mammalogy Schwartz collection: biological and historical 87(1):19–26. information. Mammalian Papers: University Soto-Centeno, J. A., D. L. Phillips, A. Kurta, and K. of Nebraska State Museum, no. 107. A. Hobson. 2014. Food resource partitioning Whittaker, J. O., and A. Rodríguez-Durán. 1999. in syntopic nectarivorous bats on Puerto Rico. Seasonal variation in the diet of Mexican free- Journal of Tropical Ecology 30:359–369. tailed bats, Tadarida brasiliensis antillarum Soto-Centeno, J. A., and D. W. Steadman. 2015. (Miller) from a colony in Puerto Rico. Fossils reject climate change as the cause Caribbean Journal of Science 35:23–28. of extinction of Caribbean bats. Scientific Wilkins, K. T. Tadarida brasiliensis. 1989. Mam- Reports 5(7971). malian species 331:1–10. Steadman, D. W., N. A. Albury, P. Maillis, J. I. Mead, Williams, M. L., and D. W. Steadman. 2001. The J. Slapcinsky, K. L. Krysko, H. M. Singleton, historic and prehistoric distribution of parrots and J. Franklin. 2014. Late-Holocene faunal (Psittacidae) in the West 1ndies. Pp. 175– and landscape change in The Bahamas. The 190 in C. A. Woods and F. E. Sergile, eds. Holocene 24(2):220–230. Biogeography of the West Indies: Patterns and Steadman, D. W., P. S. Martin, R. D. E. MacPhee, Perspectives. CRC Press, Boca Raton, Florida. SPEER ET AL.: Bats of the Bahamas: natural history and conservation 77 Notes Likely Israel's Point Cave Consolidated locality, abandoned building, coordinates are approximate Likely Sonny's Cave, but may also be Prophet's Cave, coordinates given for Sonny's Cave Sometimes confused Wall with Hole in the Cave Likely Eight mile Cave and the collec- tors are probably I. Lothain and D. Bethel Either Little Harbour Cave or Israel's Point Cave (Allen 1905) Collector G. M. Allen (1904) G. S. Morgan (1989) Gelder Van R. G. (1966) G. S. Morgan (1989), J. A. Soto-Centeno et al. (2009, 2011) I. Bethel (1985) G. M. Allen (1904) Museum FMNH FLMNH, USNM AMNH FLMNH, USNM FLMNH KU, MCZ Datum WGS84 WGS84 WGS84

-77.09 -77.56 -77.20 Longitude

Abaco Islands 26.29 26.32 25.98 Latitude Species Chilonatalus tumidi- frons brasiliensis Tadarida sezekorni Erophylla Chilonatalus tumidi- Eptesicus frons, fuscus, Erophylla sezekorni, Macrotus waterhousii Chilonatalus tumidi- frons sezekorni Erophylla Tadarida brasiliensis and Tadarida waterhousii, Macrotus sezekorni, Lasiurus minor, Eptesicus fuscus, Erophylla Chilonatalus tumidifrons, Additional Descriptions near Marsh Harbour of Flint W 0.5 km 13 km N of Bay, Wall; Hole in the Abaco National Park Species present (6): Locality Bahi Table 1. All Bahamian localities where bats have been collected, including museum where specimens are curated, collector, date collected, island, and latitude collected, date collector, are curated, museum where specimens including where bats have been collected, localities All Bahamian 1. Table and longitude to two decimal places. Localities were taken from and VertNet consolidated where the island is known. only show localities where We when available. information and provides additional localities notes. "Notes" all consolidated column indicates possible based on expert local knowledge and collector field Museum of Natural American are: Museum abbreviations locations. sensitive to preserve culturally places to two decimal are only provided and longitude Latitude History (FLMNH); University Museum of Natural History (FMNH); Florida (CUMV); Field Museum of Natural Vertebrates Museum of History (AMNH); Cornell Zoology (MCZ); Michigan State of Kansas Natural History Museum (KU); Louisiana State University Zoology (LSUMZ); Comparative University Museum (MSU); Royal Ontario (ROM); and United States National Museum, Smithsonian Institution (USNM). Casuarina Cave 0.5 mi N of Hard Bargain, Mores Island Eight Mile Cave Golden Cave Abaco Great 78 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Likely refers to Nor- man's Castle, an aban- doned settlement. Cay, Green Turtle Cay, now Treasure is the closest active settlement. Coordin- ates are provided for Norman's Castle. Sometimes confused with Eight Mile Cave, these are commonly used names for the same cave Consolidated locality Consolidated locality, see Allen (1905) Collected from a building on Little Abaco, settlement unknown (Allen 1905) Consolidated locality Collector Gelder Van R. G. (1966) Gel- Van R. G. der (1966), D. L. Reed et al. (2009), J. A. Soto-Centeno et al. (2009) G. S. Morgan (1989), D. L. Reed et al. A. (2009), J. Soto-Centeno et al. (2009, 2011), Speer and Soto- Centeno (2013) Allen et al. G. M. (1904) Allen et al. G. M. (1904) G. M. Allen (1904), Speer et al. (2013) Museum AMNH AMNH, FLMNH FLMNH, USNM AMNH, MCZ MCZ FLMNH, MCZ Datum WGS84 WGS84 WGS84 WGS84 WGS84

-77.45 -77.19 -77.19 -77.15 -77.00 Longitude

26.72 25.86 26.14 26.58 26.32 Latitude Abaco Islands (continued) Species sezekorni Erophylla Chilonatalus tumidi- Eptesicus frons, fuscus, Erophylla sezekorni, Macrotus waterhousii Eptesicus fuscus, sezekorni, Erophylla waterhousii Macrotus Chilonatalus tumidi- Erophylla frons, sezekorni brasiliensis Tadarida sezekorni Erophylla Additional Descriptions of imperial W 3 mi Lighthouse; Imperial Lighthouse Cave Long Bay Cave, 2 km N of Crossing Rocks, Island Homes Development Cave 7 mi from Marsh Harbour, Abaco; Central Great Abaco Cave at Hurricane Hole; Sitting Duck Cave; Abaco Central Table 1. Continued Table Locality Cay, Green Turtle Normans Castle Wall Hole in the Cave Island Homes Cave Israel's Point Abaco Little Little Harbour Cave SPEER ET AL.: Bats of the Bahamas: natural history and conservation 79 Notes Collected from buildings in Marsh Harbour (Allen, 1905), coordinates provided for the settlement Likely refers to Israel's Point Cave (Allen 1905) Consolidated locality Notes Consolidated locality Collector N. A. Albury (2009) G. M. Allen (1904) Allen et al. G. M. (1904) G. S. Morgan (1989), J. A. Soto-Centeno et al. (2011), Speer and Soto- Centeno (2013) B. Sweeting and N. A. Albury (2011) Unknown Collector Buden D. W. (1972) Buden D. W. (1973) Museum FLMNH MCZ MCZ FLMNH, USNM FLMNH FMNH Museum LSUMZ LSUMZ Datum WGS84 WGS84 WGS84 WGS84 Datum WGS84 WGS84

-77.00 -77.06 -77.09 -77.04 -73.89 -73.99 Longitude Longitude Acklins Island

26.49 26.54 26.34 26.55 22.72 22.41 Latitude Latitude Abaco Islands (continued) Species Lasiurus minor brasiliensis Tadarida sezekorni Erophylla Eptesicus fuscus, brasiliensis Tadarida Lasiurus minor Chilonatalus tumidifrons Species brasiliensis Tadarida sezekorni, Erophylla Monophyllus redmani, brasiliensis Tadarida Tadarida brasiliensis and Tadarida Monophyllus redmani, waterhousii, sezekorni, Macrotus Eptesicus fuscus,Erophylla Additional Descriptions Owens-Illinois Sugar Cane Processing Abandoned Plant, Building Additional Descriptions Cave Atwood near Harbour Delectable Bay SW Species present (5): Table 1. Continued Table Locality Lubber's Quarters Marsh Harbour near Marsh Central Harbour, Abaco Old Sugar Mill RBOR, the Ferry Parking Lot Locality Atwood Harbour Cave Duncan Pond Cave 80 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Likely refers to a series of 3 caves in the region. Salina Point Cave or Nibbles is the central and of these three largest caves. Consolidated locality Specimens collected from a cave (Timm and Genoways, 2003) Consolidated locality, J. D. Hall collection may have been made in 1994 Notes Collector G. Clough (1973) Bartsch (1930) P. Buden D. W. (1973) A. Schwartz et al. (1968) M. Langworthy (1981) King W. F. (1964), J. D. Hall et al. (1974), R. Franz (1981) Collector Museum LSUMZ USNM LSUMZ KU FLMNH FLMNH, MSU Museum Macrotus waterhousii and Macrotus Datum WGS84 WGS84 Datum

-73.89 -78.01 Longitude Longitude Andros

22.49 25.15 Latitude Latitude Acklins Island (continued) Eptesicus fuscus, Erophylla sezekorni, Eptesicus fuscus, Erophylla Species waterhousii Macrotus waterhousii Macrotus Eptesicus fuscus, waterhousii, Macrotus Monophyllus redmani, brasiliensis Tadarida sezekorni Erophylla sezekorni Erophylla Eptesicus fuscus, sezekorni, Erophylla waterhousii Macrotus Species Species present (3): Additional Descriptions Cave between Snug Corner and Spring Point Cave, Bluff Morgan's Pleasant Harbour, Pleasant Island; Bat cave, .4 mi SSE of “New” Morgan’s Road on “Old” Bluff Road Bluff Morgan’s Additional Descriptions Table 1. Continued Table Locality East Plana Cay, in cave Salt Jamaica Bay, Point Jumbie Hole Ashton Cave, near surface King's Cave #1 Locality 2.5 mi S of Airstrip, Man- grove Cay SPEER ET AL.: Bats of the Bahamas: natural history and conservation 81 Consolidated locality Consolidated locality, Consolidated locality, coordinates are approximate Consolidated locality, Consolidated locality, and see Timm Genoways (2003) for cave description Sometimes confused with Sheep Hill Cave, coordinates are approximate. Notes Likely collected from King's Cave #1 or see Northrop nearby, (1910) Likely Stepwell Cave, 4 mi NE of Cutlass Bay May be different from Man O'War Rock Caves Notes A. Schwartz and D. R. Paulson (1963) W. J. Clench et al. W. (1935) A. Schwartz and J. R. Thomas (1966) J. Clench et al. W. (1935) J. Clench et al. W. (1935) Collector J. I. Northrop (1890) Buden (1986) W. D. A. Schwartz and J. R. Thomas (1966) J. Clench et al. W. (1935) Collector and Lasiurus minor KU MCZ KU MCZ MCZ Museum AMNH MCZ KU MCZ Museum WGS84 WGS84 WGS84 WGS84 WGS84 Datum Datum

-75.55 -75.64 -75.38 -75.71 -75.76 Longitude Longitude Cat Island

24.46 24.61 24.15 24.67 24.67 Andros (continued) Andros Latitude Latitude Erophylla sezekorni, Macrotus waterhousii, Nyctiellus lepidus, waterhousii, sezekorni, Macrotus Erophylla Macrotus waterhousii Macrotus Erophylla sezekorni Erophylla Species waterhousii Macrotus sezekorni Erophylla waterhousii Macrotus sezekorni, Erophylla waterhousii Macrotus sezekorni, Erophylla Nyctiellus lepidus sezekorni Erophylla Lasiurus minor, waterhousii Macrotus Species Species present (4): 4.1 mi N of Tea Tea 4.1 mi N of Cave Bay, Crown Cave Additional Descriptions E end Town, Bain's of Cutlass Bay Additional Descriptions James Cystern Cave Industrious Hill Cave Dumfries Rock Man O'War Caves, Orange Creek Table 1. Continued Table Locality 3 mi NE of Cut- Cave lass Bay, Brackenwell Cave Orange Creek Locality 3 km SE of Smith Bay 82 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3)

Notes Consolidated locality, sometimes confused with James Cystern Cave Coordinates are approximate, there is a string of caves be- tween McCay's bluff and Major's Cay May be Gordon Hill Cave Long Cay was for- mally Fortune Island, coordinates are pro- vided for the church that was in the center of the settlement Consolidated locality Notes Collector Buden D. W. (1986) J. Clench et W. al. (1935) L. M. Dávalos (2004) Buden D. W. (1972) Barbour and T. J. C. Greenway (1932) Buden D. W. (1972) Buden D. W. (1972) Collector Museum MCZ MCZ AMNH LSUMZ MCZ LSUMZ LSUMZ Museum Datum WGS84 NAD27BAH WGS84 WGS84 Datum

-75.71 -74.17 -74.34 -74.25 Longitude Longitude

Crooked Island Crooked 24.67 22.75 22.60 22.78 Latitude Latitude Cat Island (continued) Species sezekorni, Erophylla waterhousii Macrotus sezekorni, Erophylla Nyctiellus lepidus Nyctiellus lepidus sezekorni Erophylla sezekorni Erophylla brasiliensis Tadarida Eptesicus fuscus, sezekorni, Erophylla waterhousii, Macrotus Monophyllus redmani, brasiliensis Tadarida Species Tadarida brasiliensis and Tadarida Monophyllus redmani, waterhousii, sezekorni, Macrotus Eptesicus fuscus, Erophylla Additional Descriptions Griffin Cave Cave Turtle between Sound and Cripple Hill Additional Descriptions Species present (5): Table 1. Continued Table Locality seaside cave at Port Howe Sheep Hill Cave, N side of Griffinland cave on north shore Fortune Island, in Albert Town, church Jingle Hill Cave Locality Cave between McCay's Bluff and Major's Cay SPEER ET AL.: Bats of the Bahamas: natural history and conservation 83 Notes Consolidated locality Coordinates provided for set- tlement, which is near many caves Notes Collector Buden (1972) W. D. D. W. Buden (1972) W. D. D. L. Reed et al. (2011) J. Riley (1903) A. Schwartz and Klinikowski R. F. A. Soto- (1961), J. Centeno et al. (2010), D. L. Reed et al. (2011) A. Speer and N. K. A. Albury (2013) J. A. Soto-Centeno et al. (2010) Collector D. L. Reed et al. (2011) Tadarida brasiliensis and Tadarida Museum LSUMZ LSUMZ FLMNH USNM FLMNH, KU FLMNH FLMNH Museum FLMNH Datum NAD27BAH NAD27BAH WGS84 NAD27BAH WGS84 WGS84 Datum NAD27BAH

-74.17 -76.16 -76.56 -76.52 -76.60 -75.16 -76.17 Longitude Longitude Eleuthera

22.75 24.62 25.39 25.37 25.55 24.85 24.64 Latitude Latitude Crooked Island (continued) Crooked Erophylla sezekorni, Macrotus waterhousii, Nyctiellus lepidus, waterhousii, sezekorni, Macrotus Erophylla Tadarida brasiliensis Tadarida Species brasiliensis Tadarida waterhousii Macrotus sezekorni, Erophylla waterhousii Macrotus sezekorni, Erophylla Nyctiellus lepidus sezekorni Erophylla sezekorni, Erophylla waterhousii Macrotus Species sezekorni Erophylla Species present (4): Cave at McKay's Bluff Additional Descriptions near Bannerman Town Hatchet Bay Plan- tation; 7.25 mi S of Glass Window Bridge on Queen's HWY 0.6 mi S of Ocean Hole on Queens across from HWY, Rock Sound AME Church Additional Descriptions 2.1 mi SSE of Town Bannerman McKays Bluff McKays Bluff Cave Table 1. Continued Table Locality Crigley Hill Cave Gregorytown Hatchet Bay Cave Hector's Cave Rock Sound Cave Locality Butter Hole Cave 84 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Likely Thompson's Cave, coordinates unknown Likely Nursery Caves Koopman et al. (1957) indicates this cave is near Williamstown Likely Nursery Caves Coordinates are approximate Notes Collector J. C. Greenway (1934) A. Soto-Centeno J. (2010), D. L. Reed A. K. et al. (2011), A. Speer and N. Albury (2013) A. Speer and N. K. A. Albury (2013) S. H. Derickson (1903) A. Schwartz and J. R. Thomas (1966) Koopman and K. F. M. K. Hecht (1953) A. Schwartz and J. R. Thomas (1966) G. B. Rabb and E. Hayden (1953) A. Speer et al. K. (2014) Collector Tadarida brasiliensis and Tadarida Museum MCZ FLMNH FLMNH USNM KU AMNH KU AMNH FLMNH Museum Datum WGS84 WGS84 WGS84 WGS84 Datum

-76.15 -76.71 -76.19 -75.72 Longitude Longitude

Exuma Islands 25.13 25.55 23.82 23.47 Latitude Latitude Eleuthera (continued) Species Nyctiellus lepidus sezekorni, Erophylla waterhousii, Macrotus brasiliensis Tadarida sezekorni Erophylla waterhousii Macrotus Eptesicus fuscus Eptesicus fuscus, brasiliensis Tadarida waterhousii Macrotus waterhousii Macrotus brasiliensis Tadarida Species Eptesicus fuscus, Erophylla sezekorni, Macrotus waterhousii, Nyctiellus lepidus, waterhousii, sezekorni, Macrotus Eptesicus fuscus, Erophylla Additional Descriptions View 3.3 mi S of Sea Drive on Queens HWY Additional Descriptions Species present (5): S Eleuthera, Bight, Wemyss John Miller's Town Table 1. Continued Table Locality Bay Cave Ten Cave Valentine of W 3 mi Town Williams Cave 1 mi of Forest SSW Settlement Darby Island, dry side, cave on SW Exuma Cays Drydock Cave Locality of 1 mi SW Forest SPEER ET AL.: Bats of the Bahamas: natural history and conservation 85 Notes Consolidated locality Consolidated locality, see Koopman et al. (1957) Consolidated locality May be Pasture Cave, Boatyard Cave, or Drydock Cave, which are all very close to the old ferry Consolidated locality, see Koopman et al. (1957) Consolidated locality Consolidated locality Collector Koopman K. F. and M. K. Hecht (1953) K. F. Koopman K. F. and M. K. Hecht (1953) Koopman K. F. and M. K. Hecht (1953) Koopman K. F. and M. K. Hecht (1953) A. Schwartz and J. R. Thomas (1966) Koopman K. F. and M. K. Hecht (1953), Speer et al. (2014) Koopman K. F. and M. K. Hecht (1953), K. A. Speer et al. (2014) Koopman K. F. and M. K. Hecht (1953), A. Schwartz and J. R. Thomas (1966) Museum AMNH AMNH AMNH AMNH KU AMNH, FLMNH AMNH, FLMNH AMNH, KU Datum NAD27BAH WGS84 WGS84 NAD27BAH

-75.91 -75.95 -75.64 -75.88 Longitude

23.57 23.65 23.46 23.55 Latitude Exuma Islands (continued) Eptesicus fuscus Species Eptesicus fuscus, waterhousii Macrotus Eptesicus fuscus, brasiliensis Tadarida Eptesicus fuscus, waterhousii Macrotus brasiliensis Tadarida Eptesicus fuscus, sezekorni, Erophylla Nyctiellus lepidus Eptesicus fuscus, sezekorni, Erophylla waterhousii, Macrotus brasiliensis Tadarida Eptesicus fuscus Nursery Caves Additional Descriptions 2 mi E of the ferry Isaac's Bay Cave 3 mi E of the ferry Bay Turnaround Cave Little Exuma, SE of Forbes Hill; Hill Cave Forest Settlement Georgetown Table 1. Continued Table Locality Hog Pen Cave Isaac's Cay Little Exuma, Christian Farm Cave; 1 mi S of Ferry Pasture Cave Pigeon Cay Robeson Cave 86 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Koopman et al. (1957) indicates this locality is a cave Collected from an old school house (Koopman et al., 1957), coordinates provided for Settlement of Rolleville Connected by passages to Burial Mound Cave Connected by passages to Ben's Cave Abandoned building Notes Collector K. F. Koopman K. F. and M. K. Hecht (1953) Koopman K. F. and M. K. Hecht (1953) A. Speer and J. K. A. Soto Centeno (2013) D. L. Reed et al. (2011) D. L. Reed et A. al. (2010), K. Speer et al. (2014) Collector D. L. Reed et A. K. al. (2011), Speer et al. (2014) Museum AMNH AMNH FLMNH FLMNH FLMNH Museum FLMNH Datum WGS84 NAD27BAH WGS84 WGS84 Datum WGS84 Tadarida brasiliensis and Tadarida

-75.98 -78.40 -78.40 -78.57 -78.56 Longitude Longitude

23.67 26.61 26.61 26.55 26.56 Erophylla sezekorni Erophylla Latitude Latitude Grand Bahama Island Exuma Islands (continued) Species present (2): Species Eptesicus fuscus, brasiliensis Tadarida brasiliensis Tadarida sezekorni Erophylla sezekorni Erophylla sezekorni Erophylla Species brasiliensis Tadarida Additional Descriptions Garden of the Groves Additional Descriptions 7 mi E of Freeport, Abandoned Hotel, Royal Oasis Hotel Rolletown Table 1. Continued Table Locality Rolleville Ben's Cave, Lu- cayan National Park Burial Mound Cave, Lucayan National Park Shannon Golf Course Locality Arawak Hotel SPEER ET AL.: Bats of the Bahamas: natural history and conservation 87 Notes Specimens collected and from a cave (Timm Genoways, 2003) Likely either Salt Pond Hill Cave or Maroon Hill Cave Specimens collected from abandoned kiln and building (Timm Genoways, 2003) Consolidated locality, coordinates are approximate Matthew Town Collector A. Schwartz and J. R. Thomas (1967) A. Schwartz and D. C. Leber (1960) A. Schwartz and J. R. Thomas (1967) Bartsch (1930) P. G. C. Klingel (1938) R. McLean and B. Shreve (1938) R. McLean and B. Shreve (1938) Buden D. W. (1975, 1977) (1962) J. Wing G. B. Rabb (1953) R. McLean and B. Shreve (1938) Museum KU KU KU USNM USNM MCZ MCZ LSUMZ CUMV, FLMNH AMNH MCZ Datum WGS84 WGS84 WGS84

-73.65 -73.68 -73.58 Longitude Inagua

21.05 20.95 20.97 Latitude Species Artibeus jamaicensis waterhousii Macrotus waterhousii Macrotus Artibeus jamaicensis Artibeus jamaicensis sezekorni Erophylla sezekorni Erophylla Artibeus jamaicensis waterhousii Macrotus waterhousii Macrotus waterhousii, Macrotus Lasiurus minor Macrotus waterhousii, and Noctilio leporinus waterhousii, Macrotus sezekorni, Lasiurus minor, Artibeus jamaicensis, Erophylla Species present (5): Additional Descriptions Undercut solution hole inland from NW Point Abandoned building located in Matthew Town Locality 2 mi N of Matthew Town Table 1. Continued Table 7 mi NE of Matthew Town 8 mi N of Matthew Town, Lake Windsor Magnasite Barn Maroon Hill Cave Great Inagua Great Inagua Great Inagua In cave at Jackline, 12 miles east of Matthew Town Little Inagua, sink hole, ca. 2 mi inland from point NW Matthew Town 88 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Consolidated locality Notes

Tadarida brasiliensis and Tadarida Collector G. C. Klingel Coleman and W. (1931), R. McLean and B. Shreve (1938), G. B. Rabb and E. B. Hayden (1953), A. Schwartz and J. R. Thomas (1967), D. Buden (1972) W. Collector T. Barbour and T. J. C. Greenway (1933) I. Dunbar (1963) C. B. Cory (1901) J. Pearson (1937) J. Pearson (1937) J. Pearson (1936) J. Clench et al. W. (1935) L. A. Hodsdon (1935) Museum AMNH, KU, LSUMZ, MCZ Museum MCZ USNM FMNH FMNH FMNH FMNH MCZ FMNH Datum WGS84 Datum

-73.58 Longitude Longitude Long Island

Inagua (continued) 20.95 Latitude Latitude Artibeus jamaicensis, waterhousii, Macrotus Noctilio leporinus Species Species Artibeus jamaicensis Artibeus jamaicensis Lasiurus minor waterhousii Macrotus brasiliensis Tadarida waterhousii, Macrotus brasiliensis Tadarida brasiliensis Tadarida brasiliensis Tadarida Eptesicus fuscus, Erophylla sezekorni, Lasiurus minor, Macrotus waterhousii, Nyctiellus lepidus, waterhousii, Macrotus sezekorni, Lasiurus minor, Eptesicus fuscus, Erophylla Cave 5.5 mi E of near Mattew Town, Salt Pond Hill; 89 km NNE of Cape Maisi (E Cuba); Matthew Town collected in cave Additional Descriptions Additional Descriptions Species present (6): Salt Pond Hill Cave Locality Adderley's Table 1. Continued Table Locality Burnt Ground, N End Carlton's Hill Cave Cave 2.25 mi SW of Clarencetown Cave at Tatnails SPEER ET AL.: Bats of the Bahamas: natural history and conservation 89 Notes May be Salt Pond Cave Consolidated locality; see Miller (1905) for clarification of J. Riley specimens Consolidated locality Collector W. J. Clench et al. W. (1936) J. Pearson (1936) Barbour and T. J. C. Greenway (1934), J. Pearson (1936), D. L. Reed et al. (2012) J. Riley (1903), L. A. Hodson J. (1935), W. Clench et al. (1936), J. Pearson (1936), D. L. Reed et al. (2010), A. Speer et al. K. (2014) D. L. Reed et A. al. (2010), K. Speer et al. (2014) J. Pearson (1936) J. Pearson (1936) J. Pearson (1937) Museum MCZ FMNH, KU FLMNH, FMNH, KU, MCZ FLMNH, FMNH, KU, MCZ, USNM FLMNH FMNH, KU FMNH, KU FMNH Datum WGS84 NAD27BAH NAD27BAH WGS84

-74.86 -75.05 -75.13 -75.21 Longitude

22.92 23.12 23.35 23.40 Latitude Long Island (continued) Species Eptesicus fuscus waterhousii Macrotus Eptesicus fuscus, sezekorni, Erophylla waterhousii, Macrotus Nyctiellus lepidus, brasiliensis Tadarida Eptesicus fuscus, sezekorni, Erophylla waterhousii, Macrotus Nyctiellus lepidus, brasiliensis Tadarida sezekorni, Erophylla waterhousii, Macrotus brasiliensis Tadarida Eptesicus fuscus, brasiliensis Tadarida Nyctiellus lepidus Lasiurus minor Additional Descriptions Mortimer's, S end, 15 mi SE of Clarencetown; near Mortimer's; S Point Cartwright's Cave, of Clar- 10 mi NW 14 mi Town; ence of Clarence NW Town of ca. 28 mi NW Clarencetown; Whalehead Clarencetown Table 1. Continued Table Locality Fox's Cave, ca. of 26 mi NW Clarencetown Cave Lakes Great Hamilton's Cave Hillside Grocery Cave McKinnon's Cave Miller's Cave, of ca. 30 mi NW Clarencetown Millerton 90 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Consolidated locality Consolidated locality Specimens collected from a cave (Timm and Genoways, 2003) There are two salt pond caves in close vicinity Collected from one of the two Salt Pond Caves in close vicinity Coordinates provided for Simms Settlement, St. Peter's Anglican Church Consolidated locality Collector J. Pearson (1936) Malone, J. V. J. Pearson (1936) A. Schwartz and J. Malone (1950) V. A. Speer et al. K. (2013) J. Pearson (1936) J. Pearson (1937) D. L. Reed et al. (2010) A. Schwartz and J. Thomas (1966), R. J. S. Davis (1980), D. L. Reed et al. (2010) Malone (1938) V. J. Malone (1938) V. J. Museum FMNH, KU FMNH, KU FLMNH, KU, UMMZ FLMNH FMNH, KU FMNH FLMNH FLMNH MCZ MCZ Datum NAD27BAH WGS84 WGS84 WGS84

-75.12 -75.24 -75.27 -74.92 Longitude

23.34 23.48 23.59 23.06 Latitude Long Island (continued) Macrotus waterhousii Macrotus Species waterhousii, Macrotus Nyctiellus lepidus, brasiliensis Tadarida waterhousii, Macrotus Nyctiellus lepidus brasiliensis Tadarida brasiliensis Tadarida Eptesicus fuscus, Lasiurus minor sezekorni, Erophylla waterhousii Macrotus Eptesicus fuscus, waterhousii Macrotus sezekorni Erophylla waterhousii Macrotus ca. 32 mi NW of ca. 32 mi NW Clarencetown; Banana Hole Additional Descriptions Grotto Bay Mimms Cave, near Dunmore; 4.9 mi SE of Clarencetown; Miley Cave Morris' Cave Table 1. Continued Table Locality Pinder's Hill, ca. of 24 mi NW Clarencetown Pinder's Settlement, ca. of 24 mi NW Clarencetown Salt Pond Cave Salt Pond Cave, of ca. 26 mi NW Clarencetown Simm's Church, Episcopal Cave Maris Stella Cave Valet SPEER ET AL.: Bats of the Bahamas: natural history and conservation 91 Notes Miller (1905) says this of back "a cave is locality Clarence Harbor" Later in the same publication, he says Clarence Harbor is a town, so perhaps he means Clarencetown Miller (1905) says "Mr. Riley procured sixty- one skins near Clarence Long Island… harbor, This species was found in a rather light cave about a mile and half back of Clarence Harbor, Long Island, and in the Hamilton cave, about five mi from the same town." Collector is likely J. Riley Consolidated locality Collector J. Riley (1903) Malone J. V. (1938) Barbour (1934) T. J. Riley (1903) Unknown (1903) Buden D. W. (1972) Museum USNM MCZ MCZ USNM USNM LSUMZ Datum WGS84

-73.07 Longitude

22.35 Latitude Long Island (continued) Species waterhousii Macrotus Nyctiellus lepidus brasiliensis Tadarida brasiliensis Tadarida brasiliensis Tadarida Lasiurus minor Additional Descriptions Near site of former military base (in cave) Table 1. Continued Table Locality Christmas Hill Cave 92 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) Notes Consolidated locality Likely Hunt's Cave Consolidated locality, see Miller (1905) Consolidated locality; Allen (1905, 1937) see and Miller (1905) Notes Collector Bartsch (1930), N. P. J. Clench et al. (1936), G. B. Rabb and E. W. Hayden (1953), D. Buden (1972) J. C. Barlow (1971) (1949) Tyne J. Van C. J. Maynard (1884), S. H. Hamilton (1902), A. Schwartz and J. R. Thomas (1966) C. Maynard (1894), S. H. Hamilton (1902), J. Riley (1903), G. M. A. Allen (1904), L. Hodsdon (1935), J. C. Dickinson (1958), A. Schwartz and R. Porter (1950) Collector Museum AMNH, FMNH, LSUMZ, MCZ, USNM ROM UMMZ AMNH, KU AMNH, FLMNH, FMNH, KU, MCZ, USNM Museum and Lasiurus minor Datum WGS84 WGS84 WGS84 Datum

-72.95 -77.45 -77.36 Longitude Longitude Mayaguana New Providence

22.39 25.07 25.08 Latitude Latitude Artibeus jamaicensis, Erophylla sezekorni, Artibeus jamaicensis, Erophylla Macrotus waterhousii sezekorni, Lasiurus minor , and Macrotus Eptesicus fuscus, Erophylla Artibeus jamaicensis, sezekorni Erophylla sezekorni Erophylla sezekorni Erophylla Eptesicus fuscus, sezekorni Erophylla Eptesicus fuscus, sezekorni, Erophylla waterhousii Macrotus Species Species Species present (3): ) ); Species present (4): Abraham's Hill Cave; Abraham's cave E of Bay; cave on top of Abraham's Hill of W Sea caves, Nassau; Cave Junction, Nassau Nassau; in dungeon of Fort Charlotte ( Eptesicus fuscus in out-building sezekorni ( Erophylla Additional Descriptions Additional Descriptions Low Point Hill Cave Table 1. Continued Table Adelaide Caves Point Cave Fort Charlotte Locality Locality of Nassau W 4 mi SPEER ET AL.: Bats of the Bahamas: natural history and conservation 93 Notes Consolidated locality, this cave was leveled for quarry rock. or Fort Charlotte Likely Caves Point Cave Likely collected by J. Benedict Likely collected by J. Benedict Miller (1905) says "cave practically unex- plored and not very long known, several miles back from Nas- from a sau" different "much better known cave somewhere back of Nassau" that likely refers to Cave Point Cave Likely collected by J. Benedict Collector H. Kennedy (1900), J. Clench et al. W. (1935), G. B. Rabb and E. B. Hayden (1953), J. C. Dickinson and M. Dickinson (1958) C. B. Cory (1879) R. Porter (1950) Unknown (1886) Unknown (1886) H. Kennedy (1900) J. Benedict (1886) Unknown (1886) Museum AMNH, FMNH, FLMNH, MCZ, USNM FMNH FLMNH, UMMZ USNM USNM AMNH, USNM FMNH, USNM USNM Datum WGS84

-77.38 Longitude

25.03 Latitude New Providence (continued) New Providence Eptesicus fuscus, sezekorni Erophylla Species Eptesicus fuscus Eptesicus fuscus Eptesicus fuscus Eptesicus fuscus sezekorni Erophylla sezekorni Erophylla sezekorni Erophylla 0.5 mi S of Harold Pond; near Fort Charlotte Additional Descriptions Hunt's Cave Nassau Table 1. Continued Table Locality Nassau Nassau Nassau Nassau Nassau Nassau 94 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 53(3) L. Notes This bat was caught after it flew inside a building (Allen 1905), coordinates given for Nassau; likely minor , not L. seminolus accessioned at USNM in 1882 and Genoways Timm (2003) says specimens were "netted in a for- mal garden." Notes Specimens were mist-netted between Graham's Harbour and Reckley Hill Set- tlement; coordinates are approximate Collector O. F. Bryant (1904) O. F. C. B. Cory (1879) C. K. Brace J. Clench (1935) W. (1971) West R. M. S. H. Hamilton (1902) A. Schwartz and J. R. Thomas (1966) Collector S. L. Olson et al. (1981) Macrotus waterhousii and Macrotus Museum MCZ FMNH USNM FMNH AMNH AMNH KU Museum USNM Datum WGS84 Datum

-77.26 Longitude Longitude San Salvador

25.04 Latitude Latitude New Providence (continued) New Providence Chilonatalus tumidifrons, Eptesicus fuscus, Erophylla sezekorni, Eptesicus fuscus, Erophylla Chilonatalus tumidifrons, Species Lasiurus seminolus waterhousii Macrotus waterhousii Macrotus sezekorni Erophylla Eptesicus fuscus waterhousii Macrotus sezekorni Erophylla Species sezekorni Erophylla Species present (4): Additional Descriptions Additional Descriptions Nassau Table 1. Continued Table Locality Nassau Nassau near Nassau end West Winton Locality 0.5 km S of Graham's Harbour 4.2 mi N of Cockburn Town SPEER ET AL.: Bats of the Bahamas: natural history and conservation 95 Notes Timm and Genoways Timm (2003) says specimens were "netted in a formal garden." Coordinates provided for abandoned and Timm church, see Genoways (2003) Consolidated locality, see Miller (1905) Barbour took E. T. fuscus from an old church in February 1934 (Olson et al. 1990), coordinates are provided for the oldest church in Cockburntown Unable to identify coordinates locality, provided for Victoria Hill settlement Likely collected from Alter Cave Collector A. Schwartz and J. R. Thomas (1966) A. Schwartz and J. R. Thomas (1966) J. Riley (1903), S. Olson et al. (1981), D. L. Reed et al. (2012) Barbour (1934) T. D. L. Reed et al. (2012) D. L. Reed et al. (2012) Bartsch (1923) P. S. Derickson (1903) Unknown (1903) Museum KU KU FLMNH, USNM MCZ FLMNH FLMNH USNM USNM USNM Datum WGS84 WGS84 WGS84 WGS84 WGS84 WGS84

-74.52 -74.56 -74.53 -74.45 -74.52 -74.46 Longitude

24.11 23.95 23.95 24.05 24.10 24.07 Latitude San Salvador (continued) San Salvador Species sezekorni Erophylla Eptesicus fuscus Chilonatalus tumidi- Erophylla frons, sezekorni Eptesicus fuscus Chilonatalus tumidi- frons sezekorni Erophylla Chilonatalus tumidi- frons Chilonatalus tumidi- frons waterhousii Macrotus Additional Descriptions Sandy Point, Island Watling's Garden Cave Likely collected by J. Riley and S. Derickson 6.9 mi NE of Cockburn Town Table 1. Continued Table Locality 9 mi S of Cockburn Town Alter Cave Cockburntown Lighthouse Cave Major's Cave at Hoy Cay North Victoria Hill The FLORIDA MUSEUM OF NATURAL HISTORY is Florida’s state museum of natural history, dedicated to understanding, preserving, and interpreting biological diversity and cultural heritage.

The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is an on-line, open-ac- cess, peer-reviewed journal that publishes results of original research in zoology, botany, paleontology, archaeology, and museum science. New issues of the Bulletin are published at irregular intervals, and volumes are not necessarily completed in any one year. Volumes contain between 150 and 300 pages, sometimes more. The number of papers contained in each volume varies, depending upon the number of pages in each paper, but four numbers is the current standard. Multi-author issues of related papers have been published together, and inquiries about putting together such isues are welcomed. Address all inqui- ries to the Editor of the Bulletin.

Richard C. Hulbert Jr., Editor

Bulletin Committee Ann S. Cordell Richard C. Hulbert Jr. Jacqueline Miller Larry M. Page Roger W. Portell, Treasurer Irvy R. Quitmyer David L. Reed, Ex officio Member

ISSN: 2373-9991

Copyright © 2015 by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, images and other media are for nonprofit, educational, and personal use of students, scholars, and the public. Any commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum.

Publication Date: August 12, 2015 This and other recent issues of the Bulletin can be freely downloaded at: http://www.flmnh.ufl.edu/bulletin/publications/

Send communications about this publication to:

Editor of the Bulletin Florida Museum of Natural History University of Florida P.O. Box 117800 Gainesville, FL 32611-7800 USA

FAX: 352-846-0287 Email: [email protected] URL: https://www.flmnh.ufl.edu/bulletin/home/