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B RE V I O R A 



us ISSN 0006-9698 



Cambridge, Mass. 10 April 2002 Number 51 1 

THE BATS OF FLORES, INDONESIA, WITH REMARKS 
ON ASIAN TADARIDA 

Kristofer M. Helgen'^ and Don E. Wilson"* 



Abstract. The Museum of Comparative Zoology at Harvard University holds 
an unreported collection of bats from Flores in the Lesser Sundas. These speci- 
mens, including a remarkable record of the Palearctic molossid Tadarida teniotis 
from the island, are reported here. The Flores specimen of T. teniotis is compared 
with other Tadarida from the region, including three specimens of T. latoiichei 
from Laos, reported here for the first time. A list of bats recorded to date from 
Flores is presented. Additional field collecting must ensue before a coherent un- 
derstanding of mammalian patterns of zoogeography and species richness in the 
Lesser Sundas, and especially Flores, will emerge. 



INTRODUCTION 

Flores is the third largest of the Lesser Sundas (Nusa Teng- 
gara), the archipelago of Indonesian islands extending east from 
Java (Fig. 1), comprising many medium-sized islands, including 
Bali, Lombok, Sumba, Sumbawa, Komodo, and Timor. The Less- 
er Sundas are especially interesting to biogeographers; bounded 
to the west by Java, to the south by Australia, and to the north 
and east by the Moluccas and New Guinea, these islands form 



' Mammal Department, Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts 02138. 

- Current address: South Australian Museum, North Terrace, Adelaide, South Aus- 
tralia 5000, Australia; e-mail: helgen@post.harvard.edu. 

^ Division of Mammals, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560. 

MCZ 
JBRARY 



SEP - 5 200Z 

HARVARD 
UNIVER31TY 



B RE V I O R A 

M. TUL s e ui nm of v^onriparafive z£/oology 



us ISSN 0006-9698 



Cambridge, Mass. 10 April 2002 Number 511 

THE BATS OF FLORES, INDONESIA, WITH REMARKS 
ON ASIAN TADARIDA 

Kristofer M. Helgen' - and Don E. Wilson^^ 



Abstract. The Museum of Comparative Zoology at Harvard University holds 
an unreported collection of bats from Flores in the Lesser Sundas. These speci- 
mens, including a remarkable record of the Palearctic molossid Tadarida teniotis 
from the island, are reported here. The Flores specimen of T. teniotis is compared 
with other Tadarida from the region, including three specimens of T. latouchei 
from Laos, reported here for the first time. A list of bats recorded to date from 
Flores is presented. Additional field collecting must ensue before a coherent un- 
derstanding of mammalian patterns of zoogeography and species richness in the 
Lesser Sundas, and especially Flores, will emerge. 



INTRODUCTION 

Flores is the third largest of the Lesser Sundas (Nusa Teng- 
gara), the archipelago of Indonesian islands extending east from 
Java (Fig. 1), comprising many medium-sized islands, including 
Bali, Lombok, Sumba, Sumbawa, Komodo, and Timor. The Less- 
er Sundas are especially interesting to biogeographers; bounded 
to the west by Java, to the south by Australia, and to the north 
and east by the Moluccas and New Guinea, these islands form 



' Mammal Department, Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts 02138. 

- Cun-ent address: South Australian Museum, North Terrace, Adelaide, South Aus- 
tralia 5000, Australia; e-mail: helgen@post.harvard.edu. 

^ Division of Mammals, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560. 

MCZ ^ 
• JBRARY 



SEP - 5 2002 

HARVARD 
UNIVERSITY 



BREVIORA 



No. 511 




Figure 1. Map of the Malay Archipelago spanning the distance between 
Southeast Asia and Australia, showing the central position of Flores in the Lesser 
Sundas. 



the southern and eastern Hmit in the region for insular faunas that 
are characteristically Asian, rather than Australian, in origin. 

The Museum of Comparative Zoology (MCZ) holds zoological 
collections made by the Reverend J. A. J. Verheijen on Flores in 
the late 1950s and early 1960s. The birds and native murids that 
Verheijen collected have been discussed (Musser, 1981; Paynter, 
1963); however, a small collection of bats that he assembled re- 
mains unreported. 

The past decade has seen numerous systematic revisions of the 
bats of the Lesser Sundas, which we draw from in identifying 
bats in Verheijen's collection (see species accounts below). De- 
spite the increased attention paid in recent years to mammal fau- 
nas in the Lesser Sundas, many of these islands have not been 
surveyed adequately. The Lesser Sundas have revealed a much 
larger degree of mammalian endemism than was previously sus- 
pected (Kitchener and Suyanto, 1996:10), and this rate of alpha- 
level systematic discovery is unlikely to subside any time soon 
if survey work is continued. Despite its relatively large size, 
knowledge of the mammalian fauna of Flores remains obscure, 
and new discoveries likely await future expeditions and continued 



2002 THE BATS OF FLORES 3 

investigation of museum specimens from the island. Thus, al- 
though a coherent understanding of Lesser Sundaic mammalian 
biogeography is emerging, many questions regarding distribution 
and species richness among the islands in this region remain to 
be answered. 

Verheijen's collection of bats from Flores comprises nine spe- 
cies, described below. Common names follow Wilson and Cole 
(2000). 

SPECIES ACCOUNTS 

Pteropus lombocensis heudei Matschie, 1899 
Lombok Flying Fox 

Specimen. MCZ 56952, male, skin only, collected at Endeh on 
the southern coast of Flores, East Nusa Tenggara, Indonesia, on 
30 November 1959. 

The taxonomy and zoogeography of P. lombocensis were re- 
viewed by Kitchener and Maryanto (1995) and Kitchener et al. 
(1995b). 

Measurements. Length of forearm, 106.7 mm. 

Aceroclon mackloti floresii (Gray, 1 87 1 ) 
Sunda Fruit Bat 

Specimen. MCZ 56960, unsexed, immature, skull, collected on 
Flores (locality unspecified). East Nusa Tenggara, Indonesia, in 
"early 1960s.'' 

Flores is the type locality of this bat, as suggested by its tri- 
nomial epithet. Koopman (1994:27) retained this form as a valid 
subspecies restricted to Sumbawa and Flores, but a thorough ex- 
amination of specimens from throughout the Lesser Sundas will 
probably bring marked changes to the classification of A. mackloti 
at the subspecific level. Specimen MCZ 56960 is an immature 
animal, with its dentition not fully erupted and its braincase and 
nasal sutures poorly ossified. 

Measurements. Greatest length of skull, 51.6 mm; condylobas- 
al length, 48.8 mm; interorbital breadth, 7.0 mm; maxillary tooth 
row, broken, unavailable. 



4 BREMOR.A No. 511 

Dobsonia peronii peronii (E. Geoffroy. 1810) 
Western Naked-backed Fruit Bat 

Specimen. MCZ 51111. female, skin only, collected at Mano 
(10 km east of Rutengi. western Flores. East Nusa Tenggara. 
Indonesia, in December 1958. 

Geographic variation in this Lesser Sundaic endemic was con- 
clusively discussed by Kitchener et al. (1997a) and Bergmans 
(1978). This specimen is a skin in good condition, unlike many 
of Verheijen's smdy skins, which were damp and putrid when his 
shipment of mammals was received by the MCZ. 

Measuremenis. Length of forearm. 119.3 mm. 

Cynopterus nusatenggara Jiusatenggara Kitchener and 

Maharadamnkamsi. 1991 

Nusa Tenggara Shon-nosed Fruit Bat 

Specimen. MCZ 56953. unsexed. immature, skin and skull, col- 
lected at Potjong. Lamba Leda. Flores. East Nusa Tenggara. In- 
donesia, on 5 January 1960. 

This recently-described bat has been \ ariably assigned to the 
svTionymy of Cynopterus brachyoris (Hill. 1992:70) or treated as 
a vahd species (K<3opman. 1993:139). This specimen is a juvenile 
individual, represented by a skull and a skin in poor condition. 
Cynopterus nusatenggara is the only Cynopterus species known 
from Flores. to which this specimen is tentatively attributed. 

Measurements. Length of forearm. 52.1 mm: greatest length of 
skuU. 26.9 mm: condylobasal length. 25.1 mm: interorbital 
breadth. 5.3 mm: maxillary tooth row. 8.7 mm. 

RJiinoIophus ajfinis princeps .-Vndersen. 1905 
Intermediate Horseshoe Bat 

Specimens. MCZ 51118. male. skin, collected at Ruteng. west- 
em Flores. East Nusa Tenggara. Indonesia, on 16 June 1959: 
MCZ 56962. unsexed. skull, collected on Flores (localir> un- 
specified). East Nusa Tenggara, Indonesia. recei\ed by MCZ in 
"early 1960s." 

This subspecies has been recorded previously from Flores by 
Bergmans and van Bree (1986:335). Hill and Rozendaal (1989: 
100). and Pumomo and Banss (1995:32). It is also found on 



2002 THE BATS OF FLOREi 5 

Lombok. Sumbawa. and Sumba fKoopman. 1994:54). The MCZ 
holds two additional female specimens of this subspecies col- 
lected in 192" at Wawo on Sumbawa. 

MeasureTiieiirs. MCZ 51 US: Length of forearm. 53.2 m. MCZ 
56962: Greatest length of skull. 23.1 mm: condylobasal length. 
20.6 mm: mterorbital breadth. 2.2 mm: maxiUar\- tooth row. 9.3 
mm. 

Hipposideros diadefna diadema lE. Geoffroy. 1813 • 
Diadem Roundleaf Bat 

Specimens. MCZ 51109. female, skin, collected 2 May 1959. 
and MCZ 51110. male, skin and skniU. collected 25 June 195S. 
both taken at Mano i 10 km east of Rutengi: MCZ 56961. un- 
sexed. skull, collected on Flores tlocaht}' unspecified i. East Nusa 
Tenggara. Indonesia, received by MCZ in "early 1960s."" 

Kitchener er al. <1992i reviewed geographic variation in H. 
diadema from the Lesser Sundas. \erheijen"s notes state that 
MCZ 51109 was collected at an altimde of 9O0 m. 

Measuremenrs. MCZ 51109: Length of forearm. S5.9 mm. 
MCZ 51110: Length or forearm. S3.4 mm: greatest length of 
skull, broken, unavailable: condylobasal length, broken, unavail- 
able: mterorbital breadth. 3^ rom: niaxillar> tooth row. 12.8 mm. 
MCZ 56961: Greatest length of skull. 30.'' mm: condylobasal 
length. 2S.4 mm: interorbital breadth. 3.5 mm: maxillary- toodi 
row. 12.6 mm. 

Myotis muricola muric .a Gray. 1^46 1 
\Miiskered Myotis 

Specimen. MCZ 51114. unsexed. skull, collected on Rores ' lo- 
cality- unspecified". East Nusa Tenggara. Indonesia, received by 
MCZ m October 1961. 

MxoTis muhcola is a wide-ranging bat of southern and south- 
east Asia. Hill ( 19S3 » discussed the taxonomy of this species, and 
considered representatives oi M. muhcola fromlh^ Lesser Sundas 
to represent the nominate subspecies. Interestingly, no names in 
the svnonym> of M. fnuricola originate from either the Greater 
or Lesser Sundas. with the exception of the distinctive M. '':. 
niasensis of Pulau Nias off western S.:::-..-:.- 'Hill. 1*^S3: Lyon. 



6 BREVIORA No. 511 

1916). A careful review of geographic variation in this species 
across its wide distribution is needed. 

Measurements. Greatest length of skull, 13.2 mm; condylobas- 
al length, 12.5 mm; interorbital breadth, 2.9 mm; maxillary tooth 
row, 5.1 mm. 

Scotophiliis colliniis Sody, 1936 
Sody's Yellow Bat 

Specimen. MCZ 51115, unsexed, skull and postcranial skele- 
ton, collected at Ruteng, western Flores, East Nusa Tenggara, 
Indonesia, on 15 June 1959. 

Kitchener et al. (1997b) reviewed the taxonomy of Scotophilus 
in the Lesser Sundas; previously, Scotophilus specimens from 
Flores were considered to represent S. kuhlii temmincki (see 
Koopman, 1994:128). 

Measurements. Greatest length of skull, 19.9 mm; condylobas- 
al length, 18.6 mm; interorbital breadth, 4.8 mm; maxillary tooth 
row, 6.6 mm. 

Tadarida teniotis (Rafinesque, 1814) 
European Free-tailed Bat 

Specimen. MCZ 56950, unsexed, skin and skull, collected at 
Endeh on the southern coast of Flores, East Nusa Tenggara, In- 
donesia in July 1960. 

Specimen MCZ 56950 is the most interesting bat collected on 
Flores by Verheijen. It represents a remarkable range extension 
for T. teniotis, a bat widely distributed throughout Europe, North 
Africa, and Asia. Previously, southern China was the most south- 
eastern geographic record for T. teniotis; this Flores specimen 
extends the range of this bat almost 2,500 miles (= 4,000 km). 

We have compared MCZ 56950 with Tadarida from other 
Asian localities (Table 1). Three specimens from Laos, examined 
in the Natural History Museum, London, are considerably smaller 
than other Asian, even south Chinese, T. teniotis. These speci- 
mens provide further support for Kock's (1999) recognition of 
Tadarida latouchei as a southeast Asian species distinct from T. 
teniotis, and firmly establish the previously disputed claim that 
this bat is present in Laos (Salter, 1993; see Kock, 1999:239). 



2002 



THE BATS OF FLORES 



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8 BREVIORA No. 511 

Verheijen's specimen of Tadarida from Flores is larger than 
the southeast Asian T. latouchei, and agrees closely in cranial and 
dental measurements with specimens of T. teniotis from China 
and Central Asia (Table 1). and with measurements for European 
T. teniotis reported in the literature (Kock and Nader, 1984). The 
presence of outer lower incisors and an unreduced anterior upper 
premolar are dental traits that demonstrate that MCZ 56950 is 
allied with T. teniotis and T. latoiichei of Asia, and not with 
Australasian (New Guinean T. kubohensis or Australian T. aus- 
tralis) or Indian {T. aegyptiaca of the Indian subcontinent) bats 
of the genus Tadarida (sensu Koopman, 1993). The other bats of 
the island also bear close affinity to Asian rather than Australian 
bats, despite the proximity of Flores to Australia and the Moluc- 
cas. (The zoogeographic relationships of the murid rodents from 
Flores are presently less clear; see Musser, 1981:163.) 

Remarkable range extensions of bats reported in the literature 
can often be the result of mistaken locality information, or of rare 
vagrants. The first seems to be unlikely in this case, because 
Verheijen sent one specimen of T. teniotis in the midst of a ship- 
ment of other mammals assuredly from Flores. and it is accom- 
panied by good original tags and unambiguous locality infor- 
mation. We believe this specimen does not represent an unusual 
vagrant, for several reasons. First, its collection locality, on the 
southeastern coast of Flores, is about the most distant site on the 
island for an individual traveling from mainland Asia. However, 
because long-distance seasonal migrations have been reported in 
North American Tadarida brasiliensis (Glass, 1982; McCracken 
et aL, 1994), the possibility that this was an individual on a cur- 
rently undocumented migration route should not be disregarded. 
In addition, it should be noted that T. teniotis is rarely collected 
even over wide areas within its known range (see Kock, 1992; 
St. Pandurska, 1992), and that the microchiropteran fauna of the 
Lesser Sundas is assuredly far from being fully characterized. 
Tadarida teniotis is the first molossid species reported for 
Flores; further mammal collecting on Flores and possibly other 
Sundaic islands may reveal additional individuals or populations 
of T teniotis in the region. If this is the case, representatives 
of this species from the Lesser Sundas may prove upon exami- 



2002 THE BATS OF FLORES 9 

Table 2. The bat fauna of Flores; compiled from Laurie and Hill (1954), 
Hill and Rozendaal (1989), Kitchener and Maharadatunkamsi (1991), Kitch- 
ener et al. (1992), Kitchener and Maryanto (1993), Kitchener et. al (1995a), 
Kitchener et. al. (1995c), Maharadatunkamsi and Kitchener (1997), Kitch- 
ener ET AL. (1997b), and this paper. 

Pteropodidae 
Pteropiis lombocensis heudei Matschie, 1899 
Acerodon mackloti floresii (Gray, 1871) 
Rousettus amplexicaudatus infumatus (Gray, 1871) 
Dobsonia peroni peroni (E. Geoffroy, 1810) 

Cynopterus nusatenggara misatenggara Kitchener and Maharadatunkamsi, 1991 
Eonycteris spelaea glandifera Lawrence, 1939 

Emballonuridae 
Taphozous longimanus leiicopleurus Dobson, 1875 

Rhinolophidae 
Rhinolophiis ajfinis princeps Andersen, 1905 
Rhinolophus simplex simplex Andersen, 1905 
Hipposideros bicolor bicolor (Temminck, 1834) 
Hipposideros diadema diadema (E. Geoffroy, 1813) 
Hipposideros sumbae sumbawae Kitchener and Maryanto, 1993 

Vespertilionidae 
Myotis miiricola muricola (Gray, 1846) 
Myotis adversus adversus (Horsfield, 1824) 
Scotophilus collimis Sody, 1936 
Pipistrellus javanicus (Gray, 1838) 
Murina florium jiorium Thomas, 1 908 
Kerivoula flora Thomas, 1914 

Molossidae 
Tadarida teniotis (Rafinesque, 1814) 



nation of additional specimens to be subspecifically distinct from 
the Asian T. t. ins ignis. 

The presence of T. teniotis on Flores brings the recorded bat 
fauna of the island to 19 species (Table 2), encompassing 15 
genera and five families. This number will no doubt increase with 
future fieldwork, and we predict that the Lesser Sundas will con- 
tinue to yield both new records and new species of mammals, 
providing over time a more complete understanding of patterns 
of zoogeography and species richness in the region. 



10 BREVIORA No. 5 1 1 

ACKNOWLEDGMENTS 

We are grateful to the late Karl Koopman of the American 
Museum of Natural History, who pointed out to us the need to 
make this report, and to Paula Jenkins for access to specimens in 
the mammal collections at the Natural History Museum in Lon- 
don. We also thank Linda Gordon of the National Museum of 
Natural History and Terri McFadden, Maria Rutzmoser, and An- 
drew Biewener of the MCZ for their support and assistance. 

NOTE ADDED IN PROOF: 

In their catalogue of the Heude Collection of mammals at the 
Institute of Zoology, Chinese Academy of Sciences, Beijing, 
Braun et al. (2001) listed a bat collected on Flores in 1895, at- 
tributable to the megachiropteran genus Nyctimene (a juvenile 
female, skin and skull). We know of only one other Nyctimene 
specimen collected in the Lesser Sundas, an adult male (skin and 
skull) from Timor (BMNH 9.1.4.8; see Andersen 1912). The Flo- 
res specimen should provisionally be referred to Nyctimene ce- 
phalotes (otherwise known from Sulawesi and the Central Mo- 
luccas), as has been done in the past for the Timor specimen 
(Andersen, 1912; Goodwin, 1979; Kitchener ^T «/., 1995). 

This additional record brings the total number of bats recorded 
from Flores to 20 species and 16 genera. 

Andersen, K. 1912. Catalogue of the Chiroptera in the Collection of the British 

Museum. Second edition. London: British Museum (Natural History), 854 pp. 
Braun, A., C. P. Groves, P. Grubb, Yang Q.. and Xia L. 2001. Catalog of the 

Musee Heude collection of mammal skulls. Acta Zootaxonomica Sinica, 26: 

608-660. 
Goodwin, R. E. 1979. The bats of Timor: systematics and ecology. Bulletin of 

the American Museum of Natural History, 163: 73-122. 
Kitchener, D. J., W. C. Packer, and A. Suyanto. 1995. Systematic review of 

Nyctimene cepholotes and N. albiventer (Chiroptera: Pteropodidae) in the 

Maluku and Sulawesi regions, Indonesia. Records of the Western AustraHan 

Museum, 17: 125-142. 

LITERATURE CITED 

Bergmans, W. 1978. On Dobsonia Palmer 1898 from the Lesser Sunda Islands 
(Mammalia, Megachiroptera). Senckenbergiana Biologica, 59: 1-18. 

Bergmans, W., and P. J. H. van Bree. 1986. On a collection of bats and rats 
from the Kangean Islands, Indonesia (Mammalia: Chiroptera and Rodentia). 
Zeitschrift fur Saugetierkunde, 51: 329-344. 



2002 THE BATS OF FLORES 11 

Glass, B. P. 1982. Seasonal movements of Mexican freetail bats Tadarida bras- 
iliensis rnexicana banded in the Great Plains. Southwestern Naturalist, 27: 
127-133. 

Hill, J. E. 1983. Bats (Mammalia: Chiroptera) from Indo- Australia. Bulletin of 
the British Museum (Natural History), 45: 103-208. 

. 1992. Order Chiroptera, pp. 54-156. In G. B. Corbet and J. E. Hill (eds.). 

The Mammals of the Indomalayan Region. Oxford, United Kingdom, Oxford 
University Press. 488 pp. 

Hill, J. E., and E G. Rozendaal. 1989. Records of bats (Microchiroptera) from 
Wallacea. Zoologische Mededelingen, Leiden 63: 97-122. 

Kitchener, D. J., N. Cooper, and I. Maryanto. 1995a. The Myotis adversus 
(Chiroptera: Vespertilionidae) species complex in eastern Indonesia, Austra- 
lia, Papua New Guinea, and the Solomon Islands. Records of the Western 
Australian Museum. 17: 191-212. 

Kitchener, D. J., S. Hisheh, L. H. Schmitt, and Maharadatunkamsi. 1997a. 
Morphological and genetic variation among island populations of Dobsonia 
peronii (Chiroptera: Pteropodidae) from the Lesser Sunda Islands, Indonesia. 
Tropical Biodiversity, 4: 35-51. 

Kitchener, D. J., R. A. How, N. K. Cooper, and A. Suyanto. 1992. Hipposideros 
diadema (Chiroptera Hipposideridae) in the Lesser Sunda Islands, Indonesia: 
Taxonomy and geographic morphological variation. Records of the Western 
Australian Museum, 16: 1-60. 

Kitchener, D. J., and Maharadatunkamsi. 1991. Description of a new species 
of Cynopterus (Chiroptera: Pteropodidae) from Nusa Tenggara, Indonesia. 
Records of the Western Australian Museum, 15: 307-363. 

Kitchener, D. J., and I. Maryanto. 1993. Taxonomic reappraisal of the Hippos- 
ideros larvatus species complex (Chiroptera: Hipposideridae) in the Greater 
and Lesser Sunda Islands. Records of the Western Australian Museum, 16: 
119-173. 

. 1995. Small Pteropus (Chiroptera: Pteropodidae) from Timor and sur- 
rounding islands, Indonesia. Records of the Western Australian Museum, 17: 
147-152. 

Kitchener, D. J., W. C. Packer, and Maharadatunkamsi. 1995b. Morphological 
variation in Pteropus lombocensis (Chiroptera: Pteropodidae) in Nusa Teng- 
gara, Indonesia. Records of the Western Australian Museum, 17: 61-67. 

Kitchener, D. J., W. C. Packer, and I. Maryanto. 1997b. Morphological vari- 
ation among populations of Scotophilus kuhlii (sensu lato) Leach, 1821 (Chi- 
roptera, Vespertilionidae) from the Greater and Lesser Sunda Islands, Indo- 
nesia. Tropical Biodiversity, 4: 53-81. 

Kitchener, D. J., L. H. Schmitt, P Strang, A. Wheeler, and A. Suyanto. 
1995c. Taxonomy of Rhinolophus simplex Andersen, 1905 (Chiroptera, Rhin- 
olophidae) in Nusa Tenggara and Maluku, Indonesia. Records of the Western 
Australian Museum, 17: 1-28. 

Kitchener, D. J., and A. Suyanto. 1996. Intraspecific morphological variation 
among island populations of small mammals in southern Indonesia, pp. 7- 



12 BREVIORA No. 511 

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J ■ • ;' KOCK, D. 1992. Tadarida teniotis (Rafinesque, 1814): Zweiter nachweis fur Ma- 
H^^/ rokko, w-palaarktische Arealgrenzen und taxonomische Anmerkung (Chirop- 

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. 1999. Tadarida (Tadarida) latouchei, a separate species recorded from 

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KocK. D., AND I. A. Nader. 1984. Tadarida teniotis (Rafinesque. 1814) in the W. 
Palaearctic and a lectotype for Dysopes rupelii Temminck, 1826 (Chiroptera: 
Molossidae). Zeitschrift fiir Saugetierkunde, 49: 129-135. 

KooPMAN, K. F 1993. Order Chiroptera. pp. 135-239. In D. E. Wilson and D. M. 
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