Quaternary bat diversity in the Dominican Republic

AMERICAN MUSEUM NOVITATES

Number 3779, 20 pp.

June 21, 2013

Quaternary Bat Diversity in the Dominican Republic

PAUL M. VELAZCO, 1 HANNAH O’NEILL, 2 GREGG F. GUNNELL, 3
SIOBHAN B. COOKE, 4 RENATO RIMOLI, 5 ALFRED L. ROSENBERGER, 1 - 6

AND NANCY B. SIMMONS 1

ABSTRACT

The fossil record of bats is extensive in the Caribbean, but few fossils have previously been
reported from the Dominican Republic. In this paper, we describe new collections of fossil bats from
two flooded caves in the Dominican Republic, and summarize previous finds from the Island of
Hispaniola. The new collections were evaluated in the context of extant and fossil faunas of the
Greater Antilles to provide information on the evolution of the bat community of Hispaniola. Eleven
species were identified within the new collections, including five mormoopids ( Mormoops blainvillei,
fMormoops magna, Pteronotus macleayii, P. parnellii, and P. quadridens), five phyllostomids ( Brachy-
phylla nana, Monophyllus redmani, Phyllonycteris poeyi, Erophylla bombifrons, and Phyllopsfalcatus),
and one natalid ( Chilonatalus micropus). All of these species today inhabitant Hispaniola with the
exception of fMormoops magna, an extinct species previously known only from the Quaternary of
Cuba, and Pteronotus macleayii, which is currently known only from extant populations in Cuba
and Jamaica, although Quaternary fossils have also been recovered in the Bahamas. Differences
between the fossil faunas and those known from the island today suggest that dispersal and extirpa¬
tion events, perhaps linked to climate change or stochastic events such as hurricanes, may have
played roles in structuring the modern fauna of Hispaniola.

1 Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History.

2 Ecology and Evolutionary Biology, Brown University, Providence, Rhode Island.

3 Division of Fossil Primates, Duke University Lemur Center, Durham, North Carolina.

4 Department of Anthropology, Northeastern Illinois University, Chicago, Illinois.

5 Department of Biology, Universidad Autonoma de Santo Domingo (UASD), Ciudad Universitaria, Santo
Domingo, Dominican Republic; and Museo del Hombre Dominicano, Santo Domingo, Dominican Republic.

6 Department of Anthropology and Archaeology, Brooklyn College, and the Graduate Center, the City University
of New York, New York; and New York Consortium in Evolutionary Primatology (NYCEP), New York.

Copyright © American Museum of Natural History 2013 ISSN 0003-0082

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AMERICAN MUSEUM NOVITATES

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FIGURE 1. Flooded floor of Olegs Bat Cave in eastern Dominican Republic, where numerous bat cranial and
postcranial remains can be observed. Photograph courtesy of the Dominican Republic Speleological Society
and Phillip Lehman.

INTRODUCTION

The West Indies harbors a diverse fauna and flora with high levels of endemism (Myers et
ah, 2000; Hedges, 2001; Willig et ah, 2009; Acevedo-Rodriguez and Strong, 2012; Davalos and
Turvey, 2012). Of the 53 extant bat species currently known from the West Indies, nearly half
are endemic to the region (Davalos and Turvey, 2012). There are 18 species of bats today living
on the island of Hispaniola (Davalos and Turvey, 2012; Nunez Novas and Leon, 2011; Tejedor
et ah, 2005) and the sparse fossil record provides little evidence as to how this assemblage of
species evolved (Griffiths and Klingener, 1988; McFarlane et ah, 2000). Hispaniola has lost
much of its mammalian diversity in the last 100,000 years, but the chronology of these extinc¬
tions is poorly documented (McFarlane et ah, 2000). Understanding the timing of these extinc¬
tions is essential to understanding the evolution of the contemporary fauna of Hispaniola.
Fossil bats have been recorded from superhcial deposits and fossilized owl pellets from the
Dominican Republic (Miller, 1929b, 1930; Morgan, 2001) and Haiti (Koopman, 1955; Miller,
1918, 1929a, 1930; Silva Taboada, 1952). New fossil faunas recovered from two sinkhole caves
in the Dominican Republic represent the biggest chiropteran fossil collection recorded from

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VELAZCO ET AL.: QUATERNARY BAT DIVERSITY

3

74°

70®

68 ®

100 KM

FIGURE 2. Map showing the localities where fossil bats have been recorded in Hispaniola. Circles indicate
localities reported by other studies. Squares indicate localities presented by this study. Haiti: (1) Diquini, (2)
Gonave Island, (3) Port-de-Paix, (4) Saint-Michel-de-FAtalaye, and (5) La Selle. Dominican Republic: (6)
Cueva de Lily, (7) Olegs Bat Cave, (8) Cerro de San Francisco, (9) Constanza, and (10) San Gabriel.

eastern Hispaniola (figs. 1-2). Here we describe these new collections in an effort to evaluate
the taxonomic diversity of the fossil bat fauna of the island, and to provide context for under¬
standing the bat diversity found in the Caribbean region today.

MATERIALS AND METHODS

The fossil collections described in this paper were recovered from two sinkhole caves on
the eastern coastline of Hispaniola: Cueva de Lily (19°33'51.19" N, 69°54'27.32" W) in the
Maria Trinidad Sanchez province, and Oleg’s Bat Cave (10 km West of Bavaro, precise locality
information can be provided upon request) in the La Altagracia province (fig. 2). Specimens
in both caves were collected as part of a joint project involving Brooklyn College and the
Museo del Hombre Dominicano, focusing on the recovery of primate and other vertebrate
remains from underwater caves. They were retrieved from the cave floors by a team of scuba
divers. The Dominican Republic Speleological Society worked on behalf of the Museo del
Hombre Dominicano.

Cueva de Lily is approximately a 900 m long system of fully freshwater-flooded passages
and caverns, with a maximum depth of 21 m. The bat fossils were collected from two areas
within the cave: one approximately 100 m from a secondary cave entrance, and the other fur¬
ther in at 180 m. The depth of the cave in both cases was between 3-8 m. The size of the tunnel
at both collection sites was approximately 5 m wide by 5-6 m high, large enough to support a

Table 1. Diversity of extant, fossil, and subfossil remains of bats from Hispaniola reported by this study, a Morgan (2001), b Miller (1930), c Miller
(1929b), d Miller (1929a), e Silva Taboada (1952)/ Koopman (1955),s and Miller (1918). h The records presented here include extant records (e) and
remains from three different sources: fossils (x), fossilized owl pellets (xx), and superficial deposits (yy).

4

AMERICAN MUSEUM NOVITATES

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FIGURE 3. Mormoopidae skulls: dorsal, lateral, and ventral views. A. Mormoops blainvillei , B. Pteronotus
macleayii, C. Pteronotus parnellii, D. Pteronotus quadridens. Scale bar = 5 mm.

FIGURE 4. Phyllostomidae skulls: dorsal, lateral, and ventral views. A. Brachyphylla nana, B. Erophylla bom-
bifrons, C. Monophyllus redmani, D. Phyllonycteris poeyi. Scale bar = 5 mm.

bat colony if dry. In the dry part at the second entrance of Cueva de Lily, there is currently a
small bat colony.

Like Cueva de Lily, Oleg’s Bat Cave is an approximately 900 m long system of fully fresh¬
water-flooded passages and caverns, with a maximum depth of 11 m. The bat fossils were col¬
lected from the surface of a rocky area (fig. 1), located approximately 15 m away from the

2013

VELAZCO ET AL.: QUATERNARY BAT DIVERSITY

7

FIGURE 5. Chiropteran mandibles. A. Mormoops blainvillei, B. Pteronotus macleayii , C. Pteronotus parnellii,
D. Pteronotus quadridens, E. Brachyphylla nana, F. Erophylla bombifrons, G. Monophyllus redmani, H. Phyll-
onycteris poeyi. Scale bar = 5 mm.

nearest entrance/exit. Although there was some silt present, the collecting process did not
involve any excavation, only picking specimens off the substrate surface. The rocky plateau was
at a depth of 8 m (fig. 1).

All specimens were hand collected and removed from the caves in water-filled plastic con¬
tainers. Specimens were placed on screens to dry and many were sprayed lightly with White Rain®
hairspray as a means of hardening them. After collection and drying, all specimens were subse¬
quently processed and identified at Duke University and the American Museum of Natural His¬
tory. Specimens were identified based on comparisons with skeletal material of the 18 extant
chiropteran species of Hispaniola as well as closely related species from elsewhere in the Carib¬
bean region, Central America, and South America. Extant specimens utilized in the comparative

AMERICAN MUSEUM NOVITATES

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analysis (see appendix) were taken from the collections of the American Museum of Natural
History, New York (AMNH), the Royal Ontario Museum, Toronto, Canada (ROM), and the
National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM). The
fossil specimens described in this paper are housed in the AMNH Mammalogy collections and
the Museo del Hombre Dominicano, Santo Domingo, Dominican Republic (MHD).

RESULTS

A total of 497 skeletal elements were identified, 165 from Oleg’s bat cave and 332 from
Cueva de Lily Eleven species from three families were identified from these collections (figs.
3-6). Many of these species inhabit Hispaniola today, including all five phyllostomids, the
single natalid, and three of the five identified mormoopid species.

Species Accounts

FAMILY MORMOOPIDAE SAUSSURE, 1860
Mormoops blainvillei Leach, 1821
Figures 3, 5

Material examined: Cueva de Lily: 2 complete skulls, 1 skull fragment, 1 femur, 2 radii.
Oleg’s Bat Cave: 6 complete skulls, 1 skull fragment, 4 dentaries, 1 scapula.

Extant distribution: Cuba, Jamaica, Hispaniola, and Puerto Rico (Davalos and Turvey,

2012 ).

Fossil record: In Hispaniola Mormoops blainvillei has been recovered from superficial
deposits (Saint-Michel-de-l’Atalaye) and fossilized owl pellets (Gonave Island) in Haiti, and
from cave fossils (Cerro de San Francisco) from the Dominican Republic (fig. 2; table 1). Addi¬
tionally, M. blainvillei has been found in Pleistocene or Holocene cave deposits in Anguilla,
Antigua and Barbuda, the Bahamas, Cuba, Jamaica, and Puerto Rico (Gundlach, 1878; Anthony,
1918; Koopman, 1951; Koopman and Williams, 1951; Koopman et al., 1957; Choate and Birney,
1968; Silva Taboada, 1974; Olson and Pregill, 1982; Steadman et al., 1984; Morgan and Woods,
1986; Morgan, 2001).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material of modern bats examined
(appendix).

f Mormoops magna Silva Taboada, 1974
Figure 6

Material examined: Oleg’s Bat Cave: 2 humeri.

Distribution: Cuba and Hispaniola (Silva Taboada, 1974; Jimenez Vazquez et al., 2005;
this report).

Remarks: f Mormoops magna is a large-bodied Mormoops known only from humeral frag¬
ments and diagnosed only on the basis of size (Silva Taboada, 1974; Jimenez Vazquez et al.,
2005). The two humeri recovered in this study correspond well with the measurements of
f Mormoops magna provided by Silva Taboada (1974) and Jimenez Vazquez et al. (2005) (table

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AMERICAN MUSEUM NOVITATES

NO. 3779

2). Previous to this study, f M. magna was known only from two cave deposits in Cuba (Silva
Taboada, 1974; Jimenez Vazquez et al., 2005). Our record represents a range extension of over
1200 km from the localities in Cuba.

Pteronotus macleayii (Gray, 1839)

Figures 3, 5, 6

Material examined: Cueva de Lily: 4 radii. Oleg’s Bat Cave: 6 complete skulls, 2 skull
fragments, 2 dentaries, 1 humerus, 3 femora.

Extant distribution: Cuba and Jamaica (Davalos and Turvey, 2012).

Fossil record: Fossil and subfossil remains are known from Pleistocene and Holocene
deposits in Cuba and the Bahamas (Silva Taboada, 1974; Morgan, 1989).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material (appendix). This is the first record of
P. macleayii for Hispaniola. Absence of this species from the modern fauna despite years of
extensive collecting suggests that it has been extirpated from the island.

Pteronotus parnellii (Gray, 1843)

Figures 3, 5

Material examined: Cueva de Lily: 4 complete skulls, 1 skull fragment. Oleg’s Bat Cave:
19 complete skulls, 6 dentaries.

Extant distribution: Cuba, Jamaica, Puerto Rico, Hispaniola, Saint Vincent, and pos¬
sibly Trinidad and Tobago (Davalos and Turvey, 2012; Clare et al., 2013).

Fossil record: In Hispaniola Pteronotus parnellii has been recovered from fossilized
owl pellets (Diquini and Gonave Island) in Haiti and from a Quaternary deposit (Cerro de
San Francisco) in the Dominican Republic (fig. 2; table 1). Additionally, P. parnellii sensu
stricto (see below) has been found in Pleistocene or Holocene cave deposits in Antigua, the
Bahamas (New Providence), Cuba, Grand Cayman, Jamaica, Puerto Rico, and Tobago (Mar¬
tin, 1972; Morgan, 1989, 2001).

Remarks: No consistent differences in cranial morphology or size were found between
our sample and comparative material from the modern fauna of Hispaniola (appendix). The
taxonomy and biogeography of bats of the Pteronotus parnellii complex is currently in a state
of flux. Although traditionally recognized as a single species ranging through the Greater
Antilles and from Mexico south to Peru and Brazil (e.g., Simmons, 2005), recent authors
have found multiple diagnosable species within what was once called Pteronotus parnellii.
Morphological and molecular studies have demonstrated that this complex includes at least
five species and perhaps more, only some of which seem to correspond to previously delim¬
ited subspecies (Gutierrez and Molinari, 2008; Clare et al., 2013). Because the holotype P.
parnellii is from Jamaica and Antillean populations are typically much smaller than mainland
forms, it seems likely that the name P. parnellii properly applies to all these bats including
those from Hispaniola.

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VELAZCO ET AL.: QUATERNARY BAT DIVERSITY

11

Pteronotus quadridens (Gundlach, 1840)

Figures 3, 5

Material examined: Cueva de Lily: 1 complete skull, 2 skull fragments, 1 dentary, 2
radii. Oleg’s Bat Cave: 1 complete skull, 1 dentary.

Extant distribution: Cuba, Hispaniola, Jamaica, and Puerto Rico (Davalos and Turvey,
2012; Simmons, 2005).

Fossil record: Pteronotus quadridens has previously been recovered from a Quaternary
cave deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table 1). Additionally,
P. quadridens has been found in Pleistocene or Holocene cave deposits in Cuba (Silva Taboada,
1974, 1979; Woloszyn and Silva Taboada, 1977) and the Bahamas (Andros, Great Abaco, and
New Providence; Morgan, 2001).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material (appendix).

FAMILY PHYLLOSTOMIDAE GRAY, 1825

SUBFAMILY GLOSSOPHAGINAE BONAPARTE, 1845
Brachyphylla nana Miller, 1902
Figures 4-6

Material examined: Oleg’s Bat Cave: 25 complete skulls, 8 mandibles, 5 dentaries, 7
scapula, 11 pelvises, 16 humeri, 4 femora.

Extant distribution: Cayman Islands (Grand Cayman), Cuba, Hispaniola, and Turks
and Caicos Islands (Middle Caicos) (Simmons, 2005).

Fossil record: Brachyphylla nana has been recovered from fossilized owl pellets (Port-de-
Paix) and superficial deposit material (Saint-Michel-de-lAtalaye) in Haiti and from a Quater¬
nary cave deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table 1).
Additionally, B. nana has been found in Pleistocene or Holocene cave deposits in the Bahamas
(Andros and New Providence), Cayman Islands (Cayman Brae), Cuba, and Jamaica (Peterson,
1917; Anthony, 1919; Miller, 1929a; Koopman and Williams, 1951; Williams, 1952; Koopman
and Ruibal, 1955; Arredondo, 1970; Mayo, 1970; Silva Taboada, 1974; Woloszyn and Silva
Taboada, 1977; Swanepoel and Genoways, 1978; Morgan, 2001).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material (appendix).

Erophylla bombifrons (Miller, 1899)

Figures 4-6

Material examined: Cueva de Lily: 1 complete skull, 2 skull fragments, 3 dentaries, 16
humeri, 2 femora, 19 radii. Oleg’s Bat Cave: 12 complete skulls, 6 mandibles, 2 dentaries, 1
scapula, 2 humeri, 2 femora.

Extant distribution: Hispaniola and Puerto Rico (Davalos and Turvey, 2012; Simmons,
2005).

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AMERICAN MUSEUM NOVITATES

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Fossil record: The only previous fossil record for this species is from Hispaniola, consist¬
ing of specimens recovered from fossilized owl pellets (Port-de-Paix) and superficial deposit
material (Saint-Michel-de-lAtalaye) from Haiti, and from a Quaternary cave deposit (Cerro de
San Francisco) from the Dominican Republic (fig. 2; table 1).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material (appendix).

Monophyllus redmani Leach, 1821
Figures 4-6

Material examined: Cueva de Lily: 14 complete skulls, 14 skull fragments, 2 dentaries,
7 humeri, 9 radii. Oleg’s Bat Cave: 20 complete skulls, 9 dentaries, 8 humeri, 2 femora.

Extant distribution: The Bahamas (Acklins, Crooked Island), Cuba, Hispaniola, Jamaica,
Puerto Rico, Turks and Caicos Islands (Middle Caicos, North Caicos, and Providenciales)
(Davalos and Turvey, 2012).

Fossil record: Monophyllus redmani has previously been recovered from fossilized
owl pellets (Diquini and Gonave Island) from Haiti as well as from Quaternary cave
deposits (Cerro de San Francisco) from the Dominican Republic (fig. 2; table 1). Addi¬
tionally, M. redmani has been found in Pleistocene or Holocene cave deposits in the
Bahamas, Cayman Islands, Cuba, Jamaica, Puerto Rico, and Middle Caicos in the Turks
and Caicos Islands (Anthony, 1925; Koopman and Willliams, 1951; Williams, 1952; Koop-
man, 1955; Koopman and Ruibal, 1955; Choate and Birney, 1968; Silva Taboada 1974;
Morgan, 2001).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material (appendix).

Phyllonycteris poeyi Gundlach, 1861
Figures 4-6

Material examined: Cueva de Lily: 151 complete skulls, 84 skull fragments, 4 mandibles,
17 dentaries, 2 scapulae, 3 femora, 32 humeri, 29 radii. Oleg’s Bat Cave: 6 complete skulls, 2
mandibles, 2 scapulae, 1 femur.

Extant distribution: Cuba and Hispaniola (Davalos and Turvey, 2012; Simmons, 2005).

Fossil record: Phyllonycteris poeyi has previously been recovered from fossilized owl
pellets (Diquini) and superficial deposit material (Saint-Michel-de-lAtalaye) in Haiti and from
a Quaternary cave deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table
1). Additionally, P. poeyi has been found in late Quaternary cave deposits in the Bahamas
(Abaco and New Providence), Cuba, and Cayman Islands (Cayman Brae) (Anthony, 1919;
Koopman and Ruibal, 1955; Silva Taboada 1974; Morgan, 2001).

Remarks: No consistent differences in cranial or postcranial morphology or size were
found between our sample and the comparative material (appendix).

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VELAZCO ET AL.: QUATERNARY BAT DIVERSITY

13

SUBFAMILY STENODERMATINAE GERVAIS, 1856
Phyllops falcatus (Gray, 1839)

Material examined: Oleg’s Bat Cave: 2 left scapulae.

Extant distribution: Cayman Islands (Grand Cayman and Cayman Brae), Cuba, and
Hispaniola (Morgan, 2001; Tavares and Mancina, 2008; Davalos and Turvey, 2012).

Fossil record: Phyllops falcatus has previously been recovered from fossilized owl pellets
(Diquini and Port-de-Paix) and superficial deposit material (Saint-Michel-de-lAtalaye) in Haiti
and from fossilized owl pellets (Constanza) and Quaternary cave deposits (Cerro de San Fran¬
cisco) in the Dominican Republic (fig. 2; table 1). Additionally, P. falcatus has been found in
late Quaternary cave deposits in Cuba (Anthony, 1919; Koopman and Ruibal, 1955; Arredondo,
1970; Torres and Rivero de la Calle, 1970; Silva Taboada and Woloszyn, 1975; Suarez and Dfaz-
Franco, 2003).

Remarks: No consistent differences in scapula shape or size were found between the Olegs
Bat Cave samples and the comparative material (appendix).

FAMILY NATALIDAE GRAY, 1866
Chilonatalus micropus (Dobson, 1880)

Material examined: Cueva de Lily: 1 radius.

Distribution: Colombia (San Andres and Providencia islands), Hispaniola, and Jamaica
(Tejedor, 2011).

Fossil record: Chilonatalus micropus has previously been recovered from a late Quater¬
nary deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table 1; Morgan, 1994,
2001). The fossil records of C. micropus from Cuba and Grand Cayman Island reported by
Morgan (2001) correspond to C. macer (Tejedor, 2011).

Remarks: No differences in radius shape or size were found between the Cueva de Lily
sample and the comparative material (appendix).

DISCUSSION

The flooded nature of both cave localities makes it impossible to determine the age of the
fossils or the time span in which they were deposited. However, the presence of extinct (f Mor-
moops magna ) and extirpated ( Pteronotus macleayii) species along with other vertebrate taxa
recovered from these caves that are currently under study (final identifications not yet avail¬
able) suggests that these fossils may be from the Late Pleistocene. Extinct sloth remains have
been recovered from Cueva de Lily. Among the more interesting remains from Oleg’s Bat Cave,
which is more complex geologically and may be more heterogeneous taphonomically, are
sloths, extinct rodents, a solenodon, an extinct bird with Cuban affinities, and the extirpated
Cuban crocodile, Crocodylus rhombifer.

Historically, 23 species of bats have been recorded from Hispaniola of which only 18 spe¬
cies are currently extant on the island (table 1). Of the five species that no longer occur on
Hispaniola, two are extinct mormoopid species (f Mormoops magna and f Pteronotus sp.) and

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AMERICAN MUSEUM NOVITATES

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FIGURE 6. Chiropteran humeri. A. Mormoops blainvillei (extant, ROM 89973), B. Mormoops megalophylla
(extant, AMNH 25589), C. f Mormoops magna, D. f Mormoops magna (mirror image), E. Pteronotus macleayii
(mirror image), F. Brachyphylla nana (mirror image), G. Erophylla bombifrons (mirror image), H. Monophyllus
redmani (mirror image), I. Phyllonycteris poeyi. Scale bar = 10 mm.

three are extant species that occur elsewhere in the Caribbean ( Mormoops megalophylla, Pter¬
onotus macleayii, and Lasiurus insularis). f Mormoops magna, apparently endemic to the islands
of Cuba and Hispaniola, is known only from humeri remains from three localites. Similarly,
f Pteronotus sp. is known only from a single mandible collected at Cerro de San Francisco
(Morgan, 2001). The abundance of fossil bat remains in Olegs Bat Cave and Cueva de Lily
opens the possibility that future collecting expeditions may be able to collect additional mate¬
rial for these two species. Little is known about these extinct taxa other than that they occurred
sympatrically with congeners, suggesting that diversity of sympatric mormoopid communities
may have been even greater in the Pleistocene than it is today A recently discovered correlation
between loss of species and loss of island area due to rising sea levels since the last glacial
maximum (LGM) suggests that climate change may have been one of the major drivers of
extinction of Caribbean bats since the Pleistocene (Davalos and Russell, 2012).

Mormoops megalophylla, Pteronotus macleayii, and Lasiurus insularis are species that have
been recorded in Hispaniola only as fossils. Extant populations of Pteronotus macleayii are cur¬
rently found on the adjacent islands of Cuba and Jamaica, with Pleistocene records from Cuba,
Hispaniola, and the Bahamas (New Providence). P. macleayii is considered an obligate cave¬
dwelling species (Silva Taboada, 1979; McFarlane, 1986; Rodriguez-Duran and Kunz, 2001;
Genoways et al., 2005). The slightly larger Pleistocene range of this taxon suggests that it was
extirpated relatively recently from the more northern and eastern parts of its range, perhaps as a
result of flooding of roost caves due to rising sea levels and climate change (Morgan, 2001).

Extant populations of Mormoops megalophylla have a wide range in mainland Central and
South America but a very restricted distribution in the Caribbean, apparently limited to Aruba,

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15

Curasao, and Bonaire (Netherlands Antilles), Trinidad, and Margarita Island (Simmons, 2005).
In the West Indies, Pleistocene remains of M. megalophylla had been found in the Bahamas
(Andros, Great Abaco), Cuba, Hispaniola, Jamaica, and Tobago, indicating that this taxon was
once widespread in the Caribbean (Morgan, 2001; Rojas Martin, 2006). In the case of Lasiurus
insularis , extant populations occur today only in Cuba, but fossils of this species are known
from both Cuba and Hispaniola (Morales and Bickham, 1995; Morgan, 2001; Simmons, 2005;
Davalos and Turvey, 2012; Nunez Novas and Leon, 2011). Reasons for the local or regional
extinctions of these taxa could have included a variety of factors including competition with
other bat species (Koopman and Williams, 1951; Williams, 1952), natural habitat changes (e.g.,
increased xerification; Pregill and Olson, 1981), deforestation (Gannon et al., 2005), flooding
of roost caves due to sea-level changes (Morgan, 2001; Davalos and Turvey, 2012), or more
complex ecological factors associated with reduced island areas after the LGM (Davalos and
Russell, 2012). In the case of Lasiurus insularis, a tree-roosting species (Silva Taboada, 1979),
anthropogenic deforestation and stochastic events such as hurricanes might have played a
significant role; in the case of Mormoops and Pteronotus species, which rely on caves for roosts,
rising sea levels and cave flooding seem more likely.

Only two of the extant species currently distributed in Hispaniola are thus far completely
absent from the fossil record of the Island —Noctilio leporinus and Molossus molossus. Both
species share a widespread distribution that extends from Mexico southward to Argentina and
the West Indies. In the Caribbean region their fossil record is sparse, with the former species
reported as fossils only from Barbuda, Cuba, and Puerto Rico, while the latter is known from
fossils only from Antigua and possibly Jamaica 7 (Morgan, 2001; Olson and Nieves-Rivera,
2010). This may at least in part reflect the roosting habits of these taxa, both of which in natural
situations prefer roosts in hollow trees to those in caves (Hood and Jones, 1984; Morgan, 2001;
Genoways et al., 2005). Caves, which offer many opportunities for fossilization of vertebrates
trapped or deposited within them, are by far the greatest source of fossil bats in the Caribbean
region (Morgan, 1989, 1994, 2001). It is perhaps ironic that the flooding of these caves as a
result of postglacial climate change may have significantly contributed to the extirpation and
extinction of multiple populations of bats on Hispaniola and other Caribbean islands (Morgan,
2001; Davalos and Turvey, 2012; Davalos and Russell, 2012). However, in the case of Oleg’s Bat
Cave and Cueva de Lily, flooding has helped to preserve ancient records of bat diversity that
provide new insights into the fauna of Hispaniola.

ACKNOWLEDGMENTS

We especially thank Arq. Christian Martinez, Director of the Museo del Hombre Dominicano,
for his support, divers from the Dominican Republic Speleological Society (Cristian Pittaro, Vika
Alexandrova, and Phillip Lehman) who assisted by collecting the specimens described in this pub¬
lication, and Gary Morgan for allowing us to use his data from Cerro de San Francisco. The follow-

7 According to Morgan (2001) it is unclear whether this record based on Koopman and Williams (1951) cor¬
responds to fossil deposits or recent owl pellet deposits.

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ing curators and collection staff graciously provided access to specimens under their care: Eileen
Westwig (AMNH), Burton Lim (ROM), and Alfred Gardner and Suzanne Peurach (USGS Patuxent
Wildlife Research Center/United States National Museum). Gary Morgan and an anonymous
reviewer read early drafts of this report and made helpful suggestions for its improvement, for which
we are grateful. Patricia J. Wynne drew figure 2. This research was supported by the National Science
Foundation Research Experience for Undergraduates (REU) program at the AMNH, and NSF grant
DEB 0949859 to N.B.S. Support was also provided by grants from the Leakey Foundation and Waitt
Foundation/National Geographic Society to A.L.R. and the Explorers Fund Grant to S.B.C.

REFERENCES

Acevedo-Rodriguez, P., and M.T. Strong. 2012. Catalogue of seed plants of the West Indies. Smithsonian
Contributions to Botany 98: 1-1192.

Anthony, H.E. 1918. The indigenous land mammals of Porto Rico, living and extinct. Memoirs of the
American Museum of Natural History (new series) 2 (2): 333-435.

Anthony, H.E. 1919. Mammals collected in eastern Cuba in 1917. With descriptions of two new species.

Bulletin of the American Museum of Natural History 41 (20): 625-643.

Anthony, H.E. 1925. Mammals of Porto Rico, living and extinct. In Scientific survey of Porto Rico and
the Virgin Islands 9: 1-241. New York: New York Academy of Sciences.

Arredondo, O. 1970. Dos nuevas especies subfosfles de mamfferos (Insectivora: Nesophontidae) del
Holoceno Precolombino de Cuba. Memorias de la Sociedad de Ciencias Naturales La Salle 86 (30):
122-152.

Choate, J.R., and E.C. Birney. 1968. Sub-recent Insectivora and Chiroptera from Puerto Rico, with the
description of a new bat of the Genus Stenoderma. Journal of Mammalogy 49: 400-412.

Clare, E.L., et al. 2013. Diversification and reproductive isolation: cryptic species in the only New World
high-duty cycle bat, Pteronotus parnellii. BMC Evolutionary Biology 13: 26.

Davalos, L.M., and A.L. Russell. 2012. Deglaciation explains bat extinction in the Caribbean. Ecology
and Evolution 2: 3045-3051.

Davalos, L.M., and S. Turvey. 2012. West Indian mammals: the old, the new, and the recent. In B.D.
Patterson and L.P. Costa (editors), Bones, clones, and biomes. The history of recent Neotropical
mammals: 157-202. Chicago: University of Chicago Press.

Gannon, M.R., A. Kurta, A. Rodrfguez-Duran, and M.R. Willig. 2005. Bats of Puerto Rico: an island
focus and a Caribbean perspective. Lubbock: Texas Tech University Press.

Genoways, H.H., R.J. Baker, J.W. Bickham, and C.J. Phillips. 2005. Bats of Jamaica. Special Publication
of the Museum, Texas Tech University 48: 1-155.

Griffiths, T., and D. Klingener. 1988. On the distribution of Greater Antillean bats. Biotropica 20: 240-
251.

Gundlach, J. 1878. Apuntes para la fauna Puerto-Riquena. Mamiferos. Anales de la Sociedad Espanola
de Historia Natural 7: 139-141.

Gutierrez, E.E., and J. Molinari. 2008. Morphometries and taxonomy of bats of the genus Pteronotus
(subgenus Phyllodia) in Venezuela. Journal of Mammalogy 89: 292-305.

Hedges, S.B. 2001. Biogeography of the West Indies: an overview. In C.A. Woods and EE. Sergile (edi¬
tors), Biogeography of the West Indies: patterns and perspectives: 15-33. Boca Raton, Florida:
CRC Press.

2013

VELAZCO ET AL.: QUATERNARY BAT DIVERSITY

17

Hood, C.S., and J.K. Jones, Jr. 1984. Noctilio leporinus. Mammalian Species 216: 1-7.

Jimenez Vazquez, O., M.M. Condis, and E. Garcia Cancio. 2005. Vertebrados post-glaciales en un resid-
uario fosil de Tyto alba scopoli (Aves: Tytonidae) en el occidente de Cuba. Revista Mexicana de
Mastozoologia 9: 85-112.

Koopman, K.F. 1951. Fossil bats from the Bahamas. Journal of Mammalogy 32: 229.

Koopman, K.F. 1955. A new subspecies of Chilonycteris from the West Indies and a discussion of the
mammals of La Gonave. Journal of Mammalogy 36: 109-113.

Koopman, K.F., and E.E. Williams. 1951. Fossil Chiroptera collected by H.E. Anthony in Jamaica, 1919—
1920. American Museum Novitates 1519: 1-29.

Koopman, K.F., and R. Ruibal. 1955. Cave-fossil vertebrates from Camaguey, Cuba. Breviora 46: 1-8.

Koopman, K.F., M.K. Hecht, and E. Ledecky-Janecek. 1957. Notes on the mammals of the Bahamas with
special reference to bats. Journal of Mammalogy 38: 164-174.

Martin, R.A. 1972. Synopsis of late Pliocene and Pleistocene bats of North America and the Antilles.
American Midland Naturalist 87: 326-335.

Mayo, N.A. 1970. La fauna vertebrada de Punta Judas. In A. Grana Gonzalez and J. Izquierdo Bordon
(editors), Sistema subterraneo de Punta Judas. Academia de Ciencias de Cuba. Serie Espeleologica
y Carsologica 30: 38-45.

McFarlane, D.A. 1986. Cave bats in Jamaica. Oryx 20: 27-30.

McFarlane, D.A., et al. 2000. New specimens of late Quaternary extinct mammals from caves in Sanchez
Ramirez Province, Dominican Republic. Caribbean Journal of Science 36: 163-166.

Miller, G.S. 1918. Three new bats from Haiti and Santo Domingo. Proceedings of the Biological Society
of Washington 31: 39-40.

Miller, G.S. 1929a. A second collection of mammals from caves near St. Michel, Haiti. Smithsonian
Miscellaneous Collections 81 (9): 1-30.

Miller, G.S. 1929b. Mammals eaten by Indians, owls and Spaniards in the coast region of the Dominican
Republic. Smithsonian Miscellaneous Collections 82 (5): 1-16.

Miller, G.S. 1930. Three small collections of mammals from Hispaniola. Smithsonian Miscellaneous
Collections 82 (15): 1-16.

Morales, J.C., and J.W. Bickham. 1995. Molecular systematics of the genus Lasiurus (Chiroptera: Vesper-
tilionidae) based on restriction-site maps of mitochondrial ribosomal genes. Journal of Mammalogy
76: 730-749.

Morgan, G.S. 1989. Fossil Chiroptera and Rodentia from the Bahamas, and the historical biogeography
of the Bahamian mammal fauna. In C.A. Wood (editor), Biogeography of the West Indies: past,
present and future: 685-740. Gainesville: Sandhill Crane Press.

Morgan, G.S. 1994. Late Quaternary fossil vertebrates from the Cayman Islands. In M.A. Brunt and J.E.
Davies (editors), The Cayman Islands: natural history and biogeography: 465-580. Dordrecht: Klu-
wer Academic.

Morgan, G.S. 2001. Patterns of extinction in West Indian bats. In C.A. Woods and F.E. Sergile (editors),
Biogeography of the West Indies: patterns and perspectives: 369-406. Boca Raton: CRC Press.

Morgan, G.S., and C.A. Woods. 1986. Extinction and zoogeography of West Indian land mammals.
Biological Journal of the Linnean Society 28: 167-203.

Myers, N., R.A. Mittermier, C.G. Mittermeier, G.A.B. da Fonseca, and J. Kent. 2000. Biodiversity hotspots
for conservation priorities. Nature 403: 853-858.

Nunez Novas, M.S., and Y.M. Leon. 2011. Analisis de la coleccion de murcielagos (Mammalia: Chirop¬
tera) del Museo Nacional de Historia Natural de Santo Domingo. Novitates Caribaea 4: 109-119.

18

AMERICAN MUSEUM NOVITATES

NO. 3779

Olson, S.L., and G.K. Pregill. 1982. Introduction to the paleontology of Bahaman vertebrates. In S.L. Olson
(editor), Fossil vertebrates from the Bahamas. Smithsonian Contributions to Paleobiology 48: 1-7.

Olson, S.L., and A.M. Nieves-Rivera. 2010. Fossil evidence and probable extinction of the greater fishing
bat Noctilio leporinus (Chiroptera: Noctilionidae) on Isla de Mona, Puerto Rico. Mastozoologia
Neotropical 17: 167-170.

Peterson, O.A. 1917. Report upon the fossil material collected in 1913 by the Messrs. Link in a cave in
the Isle of Pines. Annals of the Carnegie Museum 11: 359-361.

Pregill, G.K. and S.L. Olson. 1981. Zoogeography of West Indian vertebrates in relation to Pleistocene
climatic cycles. Annual Review of Ecology and Systematics 12: 75-98.

Rodriguez-Duran, A., and T.H. Kunz. 2001. Biogeography of West Indian bats: an ecological perspective.
In C.A. Woods and F.E. Sergile (editors), Biogeography of the West Indies: patterns and perspectives:
333-368. Boca Raton: CRC Press.

Rojas Martin, D. 2006. Notes on biogeography of Cuban bats. Chiroptera Neotropical 12: 268-273.

Silva Taboada, G. 1952. Notas sobre los murcielagos colectados en Jamaica y Haiti durante la expedicion
cientifica Cubana. Revista de Arqueologia y Etnologia 15-16: 203-214.

Silva Taboada, G. 1974. Fossil Chiroptera from cave deposits in central Cuba, with description of two
new species (genera Pteronotus and Mormoops) and the first West Indian record of Mormoops
megalophylla. Acta Zoologica Cracoviensia 19: 33-73.

Silva Taboada, G. 1979. Los murcielagos de Cuba. La Habana: Editorial Academia, Academia de Ciencias.

Silva Taboada, G., and B.W. Woloszyn. 1975. Phyllops vetus (Mammalia: Chiroptera) en Isla de Pinos.
Miscelania Zoologica (Cuba) 1: 3.

Simmons, N.B. 2005. Order Chiroptera. In D.E. Wilson and D.M. Reeder (editors), Mammal species of
the world: a taxonomic and geographic reference. 3rd ed., 1: 312-529. Baltimore, MD: Johns Hop¬
kins University Press.

Steadman, D.W., G.K. Pregill, and S.L. Olson. 1984. Fossil vertebrates from Antigua, Lesser Antilles:
evidence for late Holocene human-caused extinctions in the West Indies. Proceedings of the
National Academy of Sciences 81: 4448-4451.

Suarez, W., and S. Diaz-Franco. 2003. A new fossil bat (Chiroptera: Phyllostomidae) from a Quaternary
cave deposit in Cuba. Caribbean Journal of Science 39: 371-377.

Swanepoel, P., and H.H. Genoways. 1978. Revision of the Antillean bats of the genus Brachyphylla (Mam¬
malia: Phyllostomatidae). Bulletin of the Carnegie Museum of Natural History 12:1-53.

Tavares, V.C., and C.A. Mancina. 2008. Phyllopsfalcatus (Chiroptera: Phyllostomidae). Mammalian Spe¬
cies 811: 1-7.

Tejedor, A. 2011. Systematics of funnel-eared bats (Chiroptera: Natalidae). Bulletin of the American
Museum of Natural History 353: 1-140.

Tejedor, A., V.C. Tavares, and D. Rodriguez-Hernandez. 2005. New records of hot-cave bats from Cuba
and the Dominican Republic. Boletin de la Sociedad Venezolana de Espeleologia 39: 10-15.

Torres, P.V., and M. Rivero de la Calle. 1970. La Cueva de la Santa. Academia de Ciencias de Cuba, Serie
Espeleologia 13: 1-42.

Williams, E.E. 1952. Additional notes of fossil and subfossil bats from Jamaica. Journal of Mammalogy
33: 171-179.

Willig, M.R., S.J. Presley, C.P. Bloch, and H.H. Genoways. 2009. Macroecology of Caribbean bats:
effects of area, elevation, latitude, and hurricane-induced disturbance. In T.H. Fleming and P.A.
Racey (editors), Island bats: evolution, ecology and conservation: 216-264. Chicago: University
of Chicago Press.

2013

VELAZCO ET AL.: QUATERNARY BAT DIVERSITY

19

Woloszyn, B.W., and G. Silva Taboada. 1977. Nueva especie fosil de Artibeus (Mammalia: Chiroptera)
de Cuba, y tipificacion preliminar de los depositos fosiliferos cubanos contentivos de mamiferos
terrestres. Poeyana 161: 1-17.

APPENDIX
Specimens Examined

The following list includes all the specimens used in the comparative analysis of this study.
Specimens examined belong to the collections of the American Museum of Natural History,
New York (AMNH), the Royal Ontario Museum, Toronto (ROM), or the United Stated National
Museum, Smithsonian Institution, Washington, D.C. (USNM).

FAMILY MORMOOPIDAE

Mormoops blainvillei: Dominican Republic: San Rafael, Rancho La Guardia (AMNH
213897); Barahano, Pedernales, Cabo Rio (AMNH 238144). Jamaica: Saint Catherine, Saint
Clair Cave (ROM 89973).

Mormoops megalophylla: Mexico: Nayarit, Amatlan de Canas, Rancho Palo Amarillo
(AMNH 25589, 25602).

Pteronotus macleayii: Jamaica: Saint James, Montego Bay (AMNH 45256, 45258, 45260,
45261, 45266, 60917); Saint Elizabeth, Balaclava, Oxford Cave (AMNH 45268).

Pteronotus parnellii: Dominican Republic: Santiago Rodriguez, Moncion, Cueva Duran
(AMNH 212996). French Guiana: Paracou, near Sinnamary (AMNH 267284). Jamaica: Saint
James Parish, Montego Bay, Sewell Cave (AMNH 271546, 271547); Saint Mary Parish, Lucky
Hill, Mount Plenty Cave (AMNH 271542, 271543); Manchester Parish, Auchtembeddie, Oxford
Cave (AMNH 271544).

Pteronotus quadridens: Dominican Republic: Santiago Rodriguez, Moncion, Cueva Duran
(AMNH 212995). Haiti: Sud, Sapoti (AMNH 236654). Jamaica: Saint Catharine Parish,
Ewarton, Saint Clair Cave (AMNH 271545, 271554, 271555); Saint Elizabeth, Balaclava, Oxford
Cave (AMNH 45248). Puerto Rico: San Juan, Pueblo Viejo, Cueva de Fari (AMNH 39359);
Trujillo Alto (AMNH 39397, 39405, 39417).

FAMILY PHYLLOSTOMIDAE

Artibeus jamaicensis: French Guiana: Paracou, near Sinnamary (AMNH 266345).

Haiti: Sud, Paillant (AMNH 236678, 236679).

Brachyphylla cavernarum: U.S. Virgin Islands: Saint John, Lameshur (AMNH 188237);
Saint John, Cruz Bay (AMNH 208181).

Brachyphylla nana: Dominican Republic: San Cristobal, Los Haitises (AMNH 244909,
244910, 244912, 244914); Barahona, Los Patos (AMNH 97597). Cayman Islands: Grand Cay¬
man, Prospect (USNM 538177).

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Erophylla bombifrons: Dominican Republic: La Vega, Bonao (USNM 538347). Puerto
Rico: Pueblo Viejo (AMNH 39339, 39341).

Macrotus waterhousii: Cuba: Guantanamo Bay, Kittery Beach Road (USNM 598957).
Dominican Republic: Samana, San Juan River (AMNH 91343).

Monophyllus redmani: Haiti: Sud, Paillant (AMNH 236663,236669). Jamaica: Trelawny, Quick
Step (USNM 546355). Puerto Rico: Trujillo Alto, La Cueva de Mollfulleda (USNM 178028).

Phyllonycteris poeyi: Cuba: Habana, Aguacate, Cueva de la Numancia (AMNH 176027);
Habana, Guanajay (AMNH 23758; USNM 103445).

Phyllops falcatus: Dominican Republic: Elias Pina, Rio Limpio (USNM 542273). Haiti:
Sud, Paillant (AMNH 236696).

FAMILY NATALIDAE

Chilonatalus macer : Cuba: Isla de la Juventud, Cueva de Punta Brava (AMNH 186978).
Chilonatalus micropus: Dominican Republic: Samana, Samana, Vicenti cove (AMNH
216128).

Natalus jamaicensis: Jamaica: Saint Catharine Parish, Ewarton, Saint Clair Cave (AMNH
246127).

Natalus major: Dominican Republic: Barahona, Maniel Viejo (AMNH 97589).

FAMILY NOCTILIONIDAE

Noctilio Leporinus: Bolivia: Beni, Mamore, Mamore River (AMNH 210667); Beni, Yacuma,
Apere River (AMNH 210666). Dominican Republic: Pedernales, Oviedo, La Poza (AMNH
244903, 244904).

FAMILY MOLOSSIDAE

Molossus molossus: Dominican Republic: Distrito Nacional, Santo Domingo, La Bracita
(AMNH 62469); Santiago Rodriguez, Moncion, Cueva Duran (AMNH 213000). French Gui¬
ana: Paracou, near Sinnamary (AMNH 267250). British Virgin Islands: Guana Island (AMNH
256412). U.S. Virgin Islands: Saint John, Cruz Bay (AMNH 206704).

Nyctinomops macrotis: Dominican Republic: Distrito Nacional, Domingo, Santo Domingo
(AMNH 244932, 244936). Jamaica: (USNM 210546).

Tadarida brasiliensis: Bolivia: Cochabamba, Tablas Monte (AMNH 268655). Haiti: Sud,
Sapoti (AMNH 236705).

FAMILY VESPERTILIONIDAE

Eptesicusfuscus: Dominican Republic: San Cristobal, Cueva Santa Maria (AMNH 244927).
United States of America: Arizona, Cochise, south fork of Cave Creek (AMNH 207699).

Lasiurus borealis: United States of America: New York, New York City, American Museum
of Natural History Building (AMNH 238155); New York, New York City, 138 Convent Avenue,
City College (AMNH 203072).