nvItate MUSEUM 
Novitates 
PUBLISHED BY 
THE AMERICAN MUSEUM 
OF NATURAL HISTORY 
CENTRAL PARK WEST AT 79TH STREET 
NEW YORK, N.Y. 10024 U.S.A. 
NUMBER 2581 
JUNE 20, 1975 
KARL F. KOOPMAN 
Bats of the Virgin Islands in Relation to 
Those of the Greater and Lesser Antilles 
AMERICAN MUSEUM 
Novttates 
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY 
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 
Number 2581, pp. 1-7 
June 20, 1975 
Bats of the Virgin Islands in Relation to 
Those of the Greater and Lesser Antilles 
/ 
KARL F. KOOPMAN! 
ABSTRACT 
The 
bats 
of the Virgin 
Islands (including 
Vieques and Culebra) are discussed in relation to 
distribution, 
feeding, and roosting habits. The 
bat fauna of these islands is compared with those 
of the northern Lesser Antilles and Puerto Rico 
and it is concluded that the Virgin Island bat 
fauna 
is a depauperate 
derivative 
of that of 
Puerto Rico. Of the eight passages of bat species 
here discussed between the Greater and Lesser 
Antilles, four probably went from west to east, 
three from east to west, and one is indetermi- 
nate. 
INTRODUCTION 
The present paper represents, to some extent, 
an extension of my earlier paper (Koopman, 
1968), in which the bats of the Lesser Antilles 
were considered in relation to those of the South 
American mainland and those of the Greater 
Antilles. In this paper I examine in greater detail 
the nature 
of the relationship between the . 
Greater Antillean (particularly Puerto Rican) bat 
fauna and that of the northern Lesser Antilles. 
The Virgin Islands, lying between these two 
areas, 
are 
important 
in understanding 
this 
relationship. As the number of bat species in the 
Virgin Islands is only six, I would like to sum- 
marize this limited fauna, based on collections I 
made over a 10-year period and other specimens 
largely 
unreported. 
This includes the Puerto 
Rican islands of Vieques and Culebra and the 
Virgin Islands proper. 
My field work in the Virgin Islands was aided 
by many people and I am indebted to them. Of 
the personnel of the Virgin Islands National Park 
I am especially indebted 
to Messrs. Stephen 
Edwards, Hugh Muller, Edward Rothfuss, and 
Dr. Alan Robinson. I have also been given much 
assistance by Dr. G. A. Seaman of the Fish and 
Wildlife 
Service 
in St. Croix, Virgin Islands. 
Thanks are due Drs. Arthur E. Dammann and O. 
Marcus Buchanan, both formerly of the Virgin 
Islands Ecological Research Station. Finally I 
express appreciation to my sister, Miss Elinor - 
Koopman, for assistance in the frequently frus- 
trating business of setting up mist nets for 
catching bats. Dr. J. Knox Jones, Jr., gave me 
access to the specimens and field notes of Mr. 
James Bee at the Museum of Natural History, 
University of Kansas, for which I am grateful. 
‘Associate Curator, Department of Mammalogy, the American Museum of Natural History. 
Copyright © The American Museum of Natural History 1975 
ISSN 0003-0082 / Price 60 cents 
2 
AMERICAN MUSEUM NOVITATES 
THE VIRGIN ISLAND BAT FAUNA 
Noctilio leporinus: This large widespread fish- 
and insect-eating 
species is known from St. 
Thomas (Dobson, 1878; Goodwin, 1928); St. 
John (Hall and Bee, 1960; also two specimens in 
the American Museum of Natural History col- 
lected by me); and St. Croix (Vahl, 1797, the 
original description of mastivus, now considered 
a subspecies of NV. leporinus; also four specimens 
in the American Museum of Natural History 
collected by Harry Beatty). Goodwin (1928, p. 
106) gave a sight record from Vieques. I have 
little information about roosting sites for this 
species. Beatty collected his specimens from an 
unused chimney. Elsewhere in the West Indies, 
specimens have been obtained from hollow trees 
and from a small sea cave. The fish-catching 
abilities of N. leporinus are well known and I 
have witnessed this at Caneel Bay on St. John. 
The stomachs of Goodwin’s specimens, however, 
contained only insects. I have caught two speci- 
mens: one in a net set across a small salt pond 
near Cruz Bay, St. John, where the individual 
might have come to fish; the other also near Cruz 
Bay, but in a net set across the bed of a small 
stream (locally called a “gut”) which contained 
water only after heavy rains. Bee’s specimen was 
also netted. 
Artibeus jamaicensis: This widespread fruit- 
eating bat is known from Vieques (specimens in 
the American Museum of Natural History col- 
lected by G. G. Goodwin); Culebra (Heatwole et 
al., 
1963; 
also specimens 
in the American 
Museum of Natural History collected by G. G. 
Goodwin); St. Thomas (specimens in the Ameri- 
can Museum of Natural History collected by G. 
G. Goodwin); St. John (Hall and Bee, 1960; also 
specimens in the American Museum of Natural 
History collected by G. G. Goodwin, O. Marcus 
Buchanan, and me); Tortola (specimens in the 
American Museum of Natural History collected 
by Harry Beatty); Virgin Gorda (J. A. Allen, 
1890; G. M. Allen, 1911); Anegada (J. A. Allen, 
1890; La Bastille and Richmond, 
1973); St. 
Croix (specimens in the American Museum of 
Natural History collected by G. G. Goodwin and 
by me, also specimens in the United States 
National Museum in Washington). There are 10 
specimens at the University of Kansas collected 
NO. 2581 
by James Bee on Lovango Cay, a small island just 
north of the western end of St. John. This spe- 
cies is very labile in its roosting requirements. I 
have collected it from an abandoned house on St. 
John and from a small shed on St. Croix. Bee 
obtained 
a number. of specimens from small 
caves, both on St. John and Lovango, often in 
cliffs at the edge of the sea. Elsewhere in the 
West Indies it is well known from both large and 
small caves. Artibeus jamaicensis is probably the 
species in the Virgin Islands most frequently 
netted, as Bee and I have found on St. John and 
St. Croix, and La Bastille on Anegada. 
Stenoderma rufum: This rare fruit-eating bat 
is known 
only from St. Thomas (Hall and 
Tamsitt, 1968) and St. John (Hall and Bee, 1960; 
Hall and Tamsitt, 1968). I have had no field 
experience with this bat and my only informa- 
tion 
on 
roosting 
sites 
is from O. Marcus 
Buchanan who obtained three from a hollow silk- 
cotton tree at Mandahl Gut west of Lameshur, 
St. John 
(oral commun.). 
These 
bats were 
released after being banded. Bee obtained his 
specimens from nets. 
Brachyphylla cavernarum: This Puerto Rican 
and Lesser Antillean fruit and nectar-feeding bat 
is known from St. Thomas (specimens in the 
American Museum of Natural History collected 
by G. G. Goodwin); St. John (Hall and Bee, 
1960; also specimens in the American Museum of 
Natural History collected by G. G. Goodwin and 
by me); and St. Croix (Bond and Seaman, 1958; 
also specimens in the American Museum of 
Natural History collected by G. G. Goodwin and 
me). The American Museum of Natural History 
also has specimens from three much smaller 
islands. These include 
a series I collected on 
Norman Island, just to the east of St. John. 
Single specimens have also been collected by 
William Rainey from two small islands to the 
north of the eastern end of St. Thomas (Thatch 
Cay and Grass Cay). Bee and I each collected 
specimens from caves (at Lameshur on St. John 
and on Norman Island) in cliffs opening directly 
on the sea. Neither cave was large, but both were 
deep enough so that the bats were roosting in 
complete darkness. A similar situation obtained 
in an underground unused sugar house at Sion 
Hill, St. Croix. As far as I am aware, Brachy- 
phylla cavernarum has never been found roosting 
1975 
in either trees or wooden structures and it is 
probable 
that 
a combination of complete or 
nearly complete 
darkness and a rough stony 
place to hang is necessary for this species. Al- 
though not netted as frequently as Artibeus, I 
have caught Brachyphylla a number of times in 
the above-mentioned net across the “gut’’ at 
Cruz Bay, St. John. 
Tadarida brasiliensis: To my knowledge, the 
only record of this widespread insectivorous spe- 
cies from the Virgin Islands is Bee’s collection 
(Hall and Bee, 1960) from Lameshur on St. 
John. Bee netted his specimens, and as far as 
[am 
aware there are no data on roosting sites in the 
Virgin Islands. Elsewhere in the West Indies, this 
species is found in both caves and houses. 
Molossus molossus: This widespread insectiv- 
orous bat is known from Culebra (Heatwole et 
al, 
1963; 
also specimens 
in the American 
Museum of Natural History collected by G. G. 
Goodwin); St. Thomas (Dobson, 1878, p. 413; 
also specimens in the American Museum of 
Natural History collected by Harry Beatty); St. 
John (Hall and Bee, 1960; also specimens in the 
American Museum of Natural History collected 
by G. G. Goodwin, O. Marcus Buchanan, and 
me); 
Tortola 
(specimens 
in 
the 
American 
Museum 
collected 
by Harry 
Beatty); Virgin 
Gorda (J. A. Allen, 1890); St. Croix (specimens 
in the American Museum of Natural History col- 
lected by G. G. Goodwin and by G. A. Seaman). 
I have collected this species only from roosts, 
usually corrugated iron roofs, but also once from 
a crevice above a doorway. These roosting sites 
would certainly be usual throughout the West 
Indies. Bee picked a bat of this species off a wall 
at Annaberg on St. John. I have never caught any 
in nets but Bee obtained most of his specimens in 
this fashion. I have often seen what is probably 
this species flying at dusk on St. John and have 
caught a few (definitely this species) that blun- 
dered into enclosed spaces and could not escape. 
Summing up the distributional pattern, we 
now know that Vieques has Artibeus jamaicensis, 
and perhaps Noctilio 
leporinus; Culebra has 
Artibeus jamaicensis and Molossus molossus, St. 
Thomas has all the Virgin Island species except 
Tadarida brasiliensis, St. John has all six species; 
Tortola has Artibeus jamaicensis and Molossus 
molossus; Virgin Gorda the same two species; 
KOOPMAN: VIRGIN ISLAND BATS 
3 
Anegada only Artibeus jamaicensis; and St. Croix 
has Noctilio 
leporinus, Artibeus jamaicensis, 
Brachyphylla_ 
cavernarum, 
and 
 Molossus 
molossus. It is possible that the low dry islands 
of Vieques, Culebra, and Anegada do indeed 
support only Artibeus and Molossus, although 
Goodwin, 1928, p. 106, gave a sight record of 
Noctilio for Vieques. Tortola and Virgin Gorda, 
however, are, like St. Thomas and St. John, 
higher (and therefore wetter) islands. It therefore 
seems very likely that Noctilio, Brachyphylla, 
and even Stenoderma occur on, but are not 
known from, Tortola and Virgin Gorda because 
they have been less adequately collected than St. 
Thomas or St. John. 
TAXONOMY OF THE VIRGIN ISLAND 
BATS 
Noctilio leporinus: Davis (1973) has shown 
that mastivus is not only the subspecies found 
throughout the West Indian range but also has a 
circum-Caribbean distribution. 
Artibeus 
jamaicensis: 
The 
nominate 
sub- 
species occurs 
over most of the West Indies 
including the entire area under consideration. A 
number of synonyms are listed by Hall and 
Kelson (1959, p. 137). La Bastille and Richmond 
(1973, p. 96) seemed to imply that J. A. Allen’s 
(1890, pp. 170-173) Anegada specimens were 
different in color from the specimens they col- 
lected in 1970. In part this resulted from their 
misinterpretation of Allen’s “the pelage below 
the surface nearly pure white” to mean that the 
pelage on the ventral side was white, whereas 
examination of one of Allen’s specimens (AMNH 
3073/2382) shows that Allen was referring to the 
bases of the hairs being white. It is true that this 
specimen has “‘facial stripes two, grayish, very 
inconspicuous.” Most Virgin Island Artibeus lack 
these stripes altogether, but a few such as AMNH 
214229 from St. John do show them as clearly 
as does the above-mentioned Anegada specimen. 
Stenoderma rufum: Two subspecies have been 
described recently, S. r. anthonyi (Choate and 
Birney, 1968) based on a Pleistocene population 
from west-central Puerto Rico and S. r. darioi 
(Hall and Tamsitt, 1968) for living populations 
in the northeastern part of the island. Hall and 
Tamsitt restricted the type locality of S. r. rufum 
4 
AMERICAN MUSEUM NOVITATES 
to the Virgin Islands. Although there is little 
doubt that S. r. anthonyi can be distinguished 
from the other two subspecies, I believe that the 
distinction of these two from each other is much 
less clear. This is based entirely on the shade of 
color, which in my experience is a most unreli- 
able character 
in bats, certainly without ex- 
tensive series of skins of both forms taken at 
various times of the year. 
Brachyphylla cavernarum: The nominate sub- 
species occurs throughout the area under con- 
sideration. See Koopman (1968) for reasons for 
considering minor of Barbados a subspecies of B. 
cavernarum. 
Tadarida brasiliensis: T. b. antillularum is the 
subspecies throughout the area discussed here 
(Shamel, 1931; Schwartz, 1955). 
Molossus molossus: In recent years, two spe- 
cies of this genus, M. fortis of Puerto Rico and 
M. debilis of the northern Lesser Antilles have 
been recognized. I have previously shown (Koop- 
man, 1968) that debilis is at most a subspecies of 
M. molossus. 1 am now inclined to follow Varona 
(1974) in regarding fortis as another subspecies 
of M. molossus. Specimens from Culebra, St. 
Thomas, St. John, Tortola, and Virgin Gorda are 
all referable to the Puerto Rican M. m. fortis. 
Those from St. Croix, however, are better allo- 
cated to the Lesser Antillean M. m. debilis, al- 
though showing intergradation with fortis. 
ZOOGEOGRAPHICAL RELATIONSHIPS 
BETWEEN LESSER AND GREATER 
ANTILLEAN BAT FAUNAS 
Both Jones and Phillips (1970) and I (Koop- 
man, 
1968) have discussed 
the bats of the 
northern Lesser Antilles (north of Guadeloupe 
but not including the Virgin Islands). Excluding 
the dubious record of Carollia perspicillata from 
Redonda, there are nine species: Noctilio lepo- 
rinus, 
Monophyllus 
 plethodon, 
 Artibeus 
jamaicensis, Ardops nichollsi, Brachyphylla ca- 
vernarum, Natalus stramineus, Myotis nigricans 
(although judged from La Val, 1973, this may 
not be the correct name for the northern Lesser 
Antillean populations), Tadarida brasiliensis, and 
Molossus 
molossus. 
Five 
(all except Mono- 
phyllus, Ardops, Natalus, and Myotis) have been 
recorded from the Virgin Islands. Monophyllus 
plethodon 
is known (as a fossil) from Puerto 
NO. 2581 
Rico (Schwartz and Jones, 1967) and must have 
occured, if it does not now occur, somewhere in 
the Virgin Islands. Ardops is closely related to 
and may be said to be represented by Steno- 
derma of Puerto Rico and the Virgin Islands. In 
fact Varona (1974) considers them to be con- 
generic. Natalus major of Hispaniola, Jamaica, 
and Cuba is closely related to and possibly con- 
specific 
with Natalus 
stramineus. Thus only 
Myotis can be said to be unrepresented in or 
beyond the Virgin Islands. 
Sixteen species are now known to occur or to 
have occured in Puerto Rico (Starrett, 1962; 
Starrett and Rolle, 1963; Choate and Birney, 
1968). The species are as follows: Noctilio lepo- 
rinus, 
Pteronotus 
fuliginosus, 
P. 
parnellii, 
Mormoops _ blainvillii, 
Macrotus 
 waterhousii 
(extinct), Monophyllus plethodon (extinct), M. 
redmani, 
Artibeus 
jamaicensis, 
Stenoderma 
rufum, 
Brachyphylla 
cavernarum, 
Erophylla 
bombifrons, 
Phyllonycteris 
major 
(extinct), 
Eptesicus fuscus, Lasiurus 
borealis, 
Tadarida 
brasiliensis, Molossus molossus. Five (Noctilio, 
Artibeus, 
Brachyphylla, 
Tadarida, 
Molossus) 
occur, as we have seen, both in the Virgin Islands 
and in the northern Lesser Antilles. Stenoderma 
occurs in the Virgin Islands and has a repre- 
sentative (Ardops) in the Lesser Antilles. Only 
Monophyllus plethodon occurred on Puerto Rico 
and still occurs in the northern Lesser Antilles, 
but is not known to occur on the Virgin Islands. 
Of course, since no Pleistocene fossil remains of 
bats are known from the Virgin Islands, there is 
no reason to assume that the species did not 
occur there. It may since have become extinct 
there as it has on Puerto Rico. The remaining 
nine species are not known to occur east of 
Puerto Rico in the West Indies. 
The subgenus Natalus poses a special problem. 
Natalus stramineus is known from several Lesser 
Antillean islands. The closely related NV. major 
(Varona, 
1974, considered them to be con- 
specific) is known from Hispaniola, Jamaica, and 
Cuba, the Hispaniolan N. m. major being the 
smallest and therefore most like N. stramineus. 
There is a considerable gap between the ranges of 
N. stramineus and N. major (nothing between 
Anguilla and Saba on the east and Hispaniola on 
the west). It is indeed odd that no Natalus is 
known 
on Puerto Rico. There 
is, of course, 
always the possibility that it does or did occur. 
1975 
These bats are never abundant and are usually 
found in small numbers far back in large caves. 
The chance of fossilization is also poor as the 
Natalus skull is unusually delicately built and 
therefore quite fragile. 
In short, there are nine Greater Antillean spe- 
cies that reach (or reached) Puerto Rico, but are 
not known 
farther eastward. Of these only 
Phyllonycteris major was endemic 
to Puerto 
Rico. Five species extend from Puerto Rico 
through the Virgin Islands to the Lesser Antilles 
and of these, only Brachyphylla cavernarum does 
not occur on islands west of Puerto Rico. Even 
here, there is a close relative, B. nana (including 
B. pumila, both considered to be subspecies of B. 
cavernarum 
by Varona, 
1974). One species, 
Monophyllus_ 
plethodon, 
occurred 
both 
on 
Puerto Rico (but not on islands west of Puerto 
Rico) and in the Lesser Antilles, but is not 
known 
from the Virgin Islands. Stenoderma 
rufum is endemic to Puerto Rico and the Virgin 
Islands, but has a representative in the Lesser 
Antilles (Ardops) and others on islands west of 
Puerto Rico (Phyllops, Ariteus). One species 
group (probably at most a superspecies), Natalus 
Stramineus—major occurs in the Lesser Antilles 
and the islands west of Puerto Rico, but not 
from the islands between. Finally, a member of 
the Myotis nigricans group occurs in the northern 
Lesser Antilles, but has no representative in the 
Virgin Islands, Puerto Rico, or any islands farther 
west. Thus, whereas a majority of Puerto Rican 
bats are Greater Antillean forms that do not 
reach the Lesser Antilles, the northern Lesser 
Antilles are dominated by species that also occur 
in Puerto Rico, the Virgin Islands, or the more 
western Greater Antilles. This might imply that 
the Greater 
Antilles 
rather 
than the Lesser 
Antilles was the main source for species going 
“around the comer” of the Antilles. 
Jones and Phillips (1970) and I (Koopman, 
1968) have discussed this general problem, but I 
think it should be re-examined in a broader con- 
text, using any additional new data that may be 
available. Of the eight passages of bats here 
enumerated 
between the Greater and Lesser 
Antilles, I have no reason to change my earlier 
opinion that two (Artibeus jamaicensis , 
Tadarida 
brasiliensis) have clearly gone from the Greater 
to the 
Lesser 
Antilles, 
whereas 
two 
others 
(Natalus stramineus, Molossus molossus) have 
KOOPMAN: VIRGIN ISLAND BATS 
5 
gone in the opposite direction. Particularly in the 
light of Davis’s revision (1973), I think it is 
impossible to say in which direction Noctilio 
went and it is perfectly possible that some popu- 
lation movement has gone in both directions. 
This leaves for further discussion the three cases 
involving endemic West Indian genera. The find- 
ing by Silva and Pine (1969) that Brachyphylla is 
not a stenodermine, but a member of the other- 
wise Greater Antillean Phyllonycterinae means 
that all the species of this subfamily occur in the 
Greater Antilles and all but one are endemic to 
it. To me this means that movement of Brachy- 
phylla cavernarum from the Greater to the Lesser 
Antilles is far more probable than the reverse. 
Schwartz and Jones (4967) revised the genus 
Monophyllus, reducing it to two species, allo- 
patric except for Pleistocene sympatry on Puerto 
Rico. 
It may still be considered to be more 
diversified on the Greater than on the Lesser 
Antilles, but the difference is slight. I still think 
Monophyllus was probably derived from Glosso- 
phaga 
longirostris 
(Varona, 
1974, considered 
them to be congeneric) and the origins of Mono- 
phyllus in the Lesser Antilles and its later spread 
to the Greater Antilles seems most probable to 
me. Jones and Schwartz (1967) also revised 
Ardops and recognized a single Lesser Antillean 
species. However, here the Greater Antillean 
representation of the group to which it belongs is 
much richer. There are at least four good species 
(haitensis may be only a subspecies of Phyllops 
falcatus), some of them quite distinct. It seems 
most reasonable, therefore, to regard the Steno- 
derma 
complex 
as having originated in the 
Greater Antilles and the Lesser Antillean Ardops 
as being derived from it, albeit earlier than other 
invasions into the Lesser Antilles. My count of 
the eight passages stands, therefore, at four from 
west to east (Artibeus, Ardops, Brachyphylla, 
Tadarida), three from east to west (Monophyllus, 
Natalus, Molossus), and one unknown (Noctilio). 
ZOOGEOGRAPHICAL RELATIONSHIPS OF 
VIRGIN ISLAND BATS 
The Virgin Islands actually form two quite 
distinct geographical units. Vieques, Culebra, St. 
Thomas, 
St. 
John, 
Tortola, 
Virgin 
Gorda, 
Anegada, and most of the smaller islands are on 
the Puerto Rican bank, which formed a land 
6 
AMERICAN MUSEUM NOVITATES 
extension of Puerto Rico up until about 10,000 
years ago (Heatwole and MacKenzie, 1967). St. 
Croix (together with a few small associated 
islands), on the other hand is on its own small 
bank, separated from the Puerto Rican bank by a 
deep channel, the Anegada Passage. St. Croix 
might be expected to lack some Puerto Rican 
elements present on the other Virgin Islands but 
absent in the Lesser Antilles. This is true only of 
Stenoderma rufum. All four bat species known 
from St. Croix are present on both Puerto Rico 
and the Lesser Antilles. The only indication of 
any special Lesser Antillean affinity on St. Croix 
is the fact that the St. Croix Molossus molossus is 
more like M. m. debilis 
of the Lesser Antilles 
than like M. m. fortis of other Virgin Islands and 
Puerto Rico. All the other Virgin Islands have, as 
might be expected, a depauperate Puerto Rican 
bat 
fauna 
(particularly 
depauperate 
on 
the 
smaller, drier islands). As is usual in border zones 
between distinctive faunas, there is not so much 
a mixing of distinctive elements 
as a disap- 
pearance of distinctive elements of both faunas, 
leaving 
only 
the widespread 
varicants. 
(See 
Hershkovitz, 1958, p. 589, for use of this term.) 
This is true of the Virgin Islands and it is also 
true 
of the northern 
Lesser 
Antilles. 
The 
“comer” of the Antilles is indeed an area that 
lacks most of the distinctive elements of both 
Greater and Lesser Antilles. 
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1911. 
Mammals of the West Indies. Bull. Mus. 
Comp. Zool., vol. 54, pp. 175-263. 
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1890. 
Notes on a small collection of West 
Indian 
bats, with 
description 
of an 
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Mus. Nat. Hist., vol. 3, pp. 169-173. 
Bond, R. M., and G. A. Seaman 
1958. 
Notes 
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a colony of Brachyphylla 
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Choate, J. R., and E. C. Birney 
1968. 
Sub-recent Insectivora and Chiroptera 
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1973. 
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Jour. 
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H., 
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Levins, 
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1967. 
Herpetogeography of Puerto Rico. IV. 
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21, 
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429-438. 
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1958. 
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tropical 
mammals. 
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Zool., 
vol. 36, pp. 579-620. 
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1970. 
Comments 
on 
systematics 
and 
zoo- 
geography 
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the 
Lesser 
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on 
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Curacao and other Caribbean islands, 
vol. 32, pp. 131-145. 
Jones, J. K., and A. Schwartz 
1967. 
Bredin-Archbold-Smithsonian biological 
survey 
of Dominica. 
6. Synopsis of 
bats of the Antillean genus Ardops. 
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3634, pp. 1-13. 
Koopman, K. F. 
1968. 
Taxonomic and distributional notes on 
Lesser Antillean bats. Amer. Mus. Novi- 
tates, no. 2333, pp. 1-13. 
La Bastille, A.. and M. Richmond 
1973. 
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British Virgin Islands. Caribbean Jour. 
Sci., vol. 13, pp. 91-109. 
(1975 
La Val, R. 
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