AMERICAN MUSEUM

Novttates 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

KARI F. KOOPMAN Bats of the Virgin Islands in Relation to Those of the Greater and Lesser Antilles

AMERICAN MUSEUM Novitates 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 fauna is a depauperate derivative of that of Vieques and Culebra) are discussed in relation to Puerto Rico. Of the eight passages of bat species distribution, feeding, and roosting habits. The here discussed between the Greater and Lesser bat fauna of these islands is compared with those Antilles, four probably went from west to east, of the northern Lesser Antilles and Puerto Rico three from east to west, and one is indetermi- and it is concluded that the Virgin Island bat nate.

INTRODUCTION The present paper represents, to some extent, My field work in the Virgin Islands was aided an extension of my earlier paper (Koopman, by many people and I am indebted to them. Of 1968), in which the bats of the Lesser Antilles the personnel of the Virgin Islands National Park were considered in relation to those of the South I am especially indebted to Messrs. Stephen American mainland and those of the Greater Edwards, Hugh Muller, Edward Rothfuss, and Antilles. In this-paper I examine in greater detail Dr. Alan Robinson. I have also been given much the nature of the relationship between the assistance by Dr. G. A. Seaman of the Fish and Greater Antillean (particularly Puerto Rican) bat Wildlife Service in St. Croix, Virgin Islands. fauna and that of the northern Lesser Antilles. Thanks are due Drs. Arthur E. Dammann and 0. The Virgin Islands, lying between these two Marcus Buchanan, both formerly of the Virgin areas, are important in understanding this Islands Ecological Research Station. Finally I relationship. As the number of bat species in the express appreciation to my sister, Miss Elinor Virgin Islands is only six, I would like to sum- Koopman, for assistance in the frequently frus- marize this limited fauna, based on collections I trating business of setting up mist nets for made over a 10-year period and other specimens catching bats. Dr. J. Knox Jones, Jr., gave me largely unreported. This includes the Puerto access to the specimens and field notes of Mr. Rican islands of Vieques and Culebra and the James Bee at the Museum of Natural History, Virgin Islands proper. University of Kansas, for which I am grateful.

'Associate Curator, Department of Mammalogy, the American Museum of Natural History.

Copyright i The American Museum of Natural History 1975 ISSN 0003-0082 / Price 60 cents 2 AMERICAN MUSEUM NOVITATES NO. 2581

THE VIRGIN ISLAND BAT FAUNA by James Bee on Lovango Cay, a small island just north of the western end of St. John. This spe- Noctilio leporinus: This large widespread fish- cies is very labile in its roosting requirements. I and insect-eating species is known from St. have collected it from an abandoned house on St. Thomas (Dobson, 1878; Goodwin, 1928); St. John and from a small shed on St. Croix. Bee John (Hall and Bee, 1960; also two specimens in obtained a number of specimens from small the American Museum of Natural History col- caves, both on St. John and Lovango, often in lected by me); and St. Croix (Vahl, 1797, the cliffs at the edge of the sea. Elsewhere in the original description of mastivus, now considered West Indies it is well known from both large and a subspecies of N. leporinus; also four specimens small caves. Artibeus jamaicensis is probably the in the American Museum of Natural History species in the Virgin Islands most frequently collected by Harry Beatty). Goodwin (1928, p. netted, as Bee and I have found on St. John and 106) gave a sight record from Vieques. I have St. Croix, and La Bastille on Anegada. little information about roosting sites for this Stenoderma rufum: This rare fruit-eating bat species. Beatty collected his specimens from an is known only from St. Thomas (Hall and unused chimney. Elsewhere in the West Indies, Tamsitt, 1968) and St. John (Hall and Bee, 1960; specimens have been obtained from hollow trees Hall and Tamsitt, 1968). I have had no field and from a small sea cave. The fish-catching experience with this bat and my only informa- abilities of N. leporinus are well known and I tion on roosting sites is from 0. Marcus have witnessed this at Caneel Bay on St. John. Buchanan who obtained three from a hollow silk- The stomachs of Goodwin's specimens, however, cotton tree at Mandahl Gut west of Lameshur, contained only insects. I have caught two speci- St. John (oral commun.). These bats were mens: one in a net set across a small salt pond released after being banded. Bee obtained his near Cruz Bay, St. John, where the individual specimens from nets. might have come to fish; the other also near Cruz Brachyphylla cavernarum: This Puerto Rican Bay, but in a net set across the bed of a small and Lesser Antillean fruit and nectar-feeding bat stream (locally called a "gut") which contained is known from St. Thomas (specimens in the water only after heavy rains. Bee's specimen was American Museum of Natural History collected also netted. by G. G. Goodwin); St. John (Hall and Bee, Artibeus jamaicensis: This widespread fruit- 1960; also specimens-in the American Museum of eating bat is known from Vieques (specimens in Natural History collected by G. G. Goodwin and the American Museum of Natural History col- by me); and St. Croix (Bond and Seaman, 1958; lected by G. G. Goodwin); Culebra (Heatwole et also specimens in the American Museum of al., 1963; also specimens in the American Natural History collected by G. G. Goodwin and Museum of Natural History collected by G. G. me). The American Museum of Natural History Goodwin); St. Thomas (specimens in the Ameri- also has specimens from three much smaller can Museum of Natural History collected by G. islands. These include a series I collected on G. Goodwin); St. John (Hall and Bee, 1960; also Norman Island, just to the east of St. John. specimens in the American Museum of Natural Single specimens have also been collected by History collected by G. G. Goodwin, 0. Marcus William Rainey from two small islands to the Buchanan, and me); Tortola (specimens in the north of the eastern end of St. Thomas (Thatch American Museum of Natural History collected Cay and Grass Cay). Bee and I each collected by Harry Beatty); Virgin Gorda (J. A. Allen, specimens from caves (at Lameshur on St. John 1890; G. M. Allen, 191 1); Anegada (J. A. Allen, and on Norman Island) in cliffs opening directly 1890; La Bastille and Richmond, 1973); St. on the sea. Neither cave was large, but both were Croix (specimens in the American Museum of deep enough so that the bats were roosting in Natural History collected by G. G. Goodwin and complete darkness. A similar situation obtained by me, also specimens in the United States in an underground unused sugar house at Sion National Museum in Washington). There are 10 Hill, St. Croix. As far as I am aware, Brachy- specimens at the University of Kansas collected phylla cavernarum has never been found roosting 1975 KOOPMAN: VIRGIN ISLAND BATS 3 in either trees or wooden structures and it is Anegada only Artibeus jamaicensis; and St. Croix probable that a combination of complete or has Noctilio leporinus, Artibeus jamaicensis, nearly complete darkness and a rough stony Brachyphylla cavernarum, and Molossus place to hang is necessary for this species. Al- molossus. It is possible that the low dry islands though not netted as frequently as Artibeus, I of Vieques, Culebra, and Anegada do indeed have caught Brachyphylla a number of times in support only Artibeus and Molossus, although the above-mentioned net across the "gut" at Goodwin, 1928, p. 106, gave a sight record of Cruz Bay, St. John. Noctilio for Vieques. Tortola and Virgin Gorda, Tadarida brasiliensis: To my knowledge, the however, are, like St. Thomas and St. John, only record of this widespread insectivorous spe- higher (and therefore wetter) islands. It therefore cies from the Virgin Islands is Bee's collection seems very likely that Noctilio, Brachyphylla, (Hall and Bee, 1960) from Lameshur on St. and even Stenoderma occur on, but are not John. Bee netted his specimens, and as far as I am known from, Tortola and Virgin Gorda because aware there are no data on roosting sites in the they have been less adequately collected than St. Virgin Islands. Elsewhere in the West Indies, this Thomas or St. John. species is found in both caves and houses. Molossus molossus: This widespread insectiv- TAXONOMY OF THE VIRGIN ISLAND orous bat is known from Culebra (Heatwole et BATS al., 1963; also specimens in the American Museum of Natural History collected by G. G. Noctilio leporinus: Davis (1973) has shown Goodwin); St. Thomas (Dobson, 1878, p. 413; that mastivus is not only the subspecies found also specimens in the American Museum of throughout the West Indian range but also has a Natural History collected by Harry Beatty); St. circum-Caribbean distribution. John (Hall and Bee, 1960; also specimens in the Artibeus jamaicensis: The nominate sub- American Museum of Natural History collected species occurs over most of the West Indies by G. G. Goodwin, 0. Marcus Buchanan, and including the entire area under consideration. A me); Tortola (specimens in the American number of synonyms are listed by Hall and Museum collected by Harry Beatty); Virgin Kelson (1959, p. 137). La Bastille and Richmond Gorda (J. A. Allen, 1890); St. Croix (specimens (1973, p. 96) seemed to imply that J. A. Allen's in the American Museum of Natural History col- (1890, pp. 170-173) Anegada specimens were lected by G. G. Goodwin and by G. A. Seaman). different in color from the specimens they col- I have collected this species only from roosts, lected in 1970. In part this resulted from their usually corrugated iron roofs, but also once from misinterpretation of Allen's "the pelage below a crevice above a doorway. These roosting sites the surface nearly pure white" to mean that the would certainly be usual throughout the West pelage on the ventral side was white, whereas Indies. Bee picked a bat of this species off a wall examination of one of Allen's specimens (AMNH at Annaberg on St. John. I have never caught any 3073/2382) shows that Allen was referring to the in nets but Bee obtained most of his specimens in bases of the hairs being white. It is true that this this fashion. I have often seen what is probably specimen has "facial stripes two, grayish, very this species flying at dusk on St. John and have inconspicuous." Most Virgin Island Artibeus lack caught a few (definitely this species) that blun- these stripes altogether, but a few such as AMNH dered into enclosed spaces and could not escape. 214229 from St. John do show them as clearly Summing up the distributional pattern, we as does the above-mentioned Anegada specimen. now know that Vieques has Artibeus jamaicensis, Stenoderma rufum: Two subspecies have been and perhaps Noctilio leporinus; Culebra has described recently, S. r. anthonyi (Choate and Artibeus jamaicensis and Molossus molossus; St. Birney, 1968) based on a Pleistocene population Thomas has all the Virgin Island species except from west-central Puerto Rico and S. r. darioi Tadarida brasiliensis; St. John has all six species; (Hall and Tamsitt, 1968) for living populations Tortola has Artibeus jamaicensis and Molossus in the northeastern part of the island. Hall and molossus; Virgin Gorda the same two species; Tamsitt restricted the type locality of S. r. rufum 4 AMERICAN MUSEUM NOVITATES NO. 2581 to the Virgin Islands. Although there is little Rico (Schwartz and Jones, 1967) and must have doubt that S. r. anthonyi can be distinguished occured, if it does not now occur, somewhere in from the other two subspecies, I believe that the the Virgin Islands. Ardops is closely related to distinction of these two from each other is much and may be said to be represented by Steno- less clear. This is based entirely on the shade of derma of Puerto Rico and the Virgin Islands. In color, which in my experience is a most unreli- fact Varona (1974) considers them to be con- able character in bats, certainly without ex- generic. Natalus mzajor of Hispaniola, Jamaica, tensive series of skins of both forms taken at and Cuba is closely related to and possibly con- various times of the year. specific with Natalus stramineus. Thus only Brachyphylla cavernarum: The nominate sub- Myotis can be said to be unrepresented in or species occurs throughout the area under con- beyond the Virgin Islands. sideration. See Koopman (1968) for reasons for Sixteen species are now known to occur or to considering minor of Barbados a subspecies ofB. have occured in Puerto Rico (Starrett, 1962; cavemarum Starrett and Rolle, 1963; Choate and Birney, Tadarida brasiliensis: T. b. antillularum is the 1968). The species are as follows: Noctilio lepo- subspecies throughout the area discussed here rinus, Pteronotus fuliginosus, P. pamellii, (Shamel, 1931;Schwartz, 1955). Mormoops blainvillii, Macrotus waterhousii Molossus molossus: In recent years, two spe- (extinct), Monophyllus plethodon (extinct), M. cies of this genus, M. fortis of Puerto Rico and redmani, Artibeus jamaicensis, Stenoderma M. debilis of the northern Lesser Antilles have rufum, Brachyphylla cavernarum, Erophylla been recognized. I have previously shown (Koop- bombifrons, Phyllonycteris major (extinct), man, 1968) that debilis is at most a subspecies of Eptesicus fuscus, Lasiurus borealis, Tadarida M. molossus. I am now inclined to follow Varona brasiliensis, Molossus molossus. Five (Noctilio, (1974) in regarding fortis as another subspecies Artibeus, Brachyphylla, Tadarida, Molossus) of M. molossus. Specimens from Culebra, St. occur, as we have seen, both in the Virgin Islands Thomas, St. John, Tortola, and Virgin Gorda are and in the northern Lesser Antilles. Stenoderma all referable to the Puerto Rican M. m. fortis. occurs in the Virgin Islands and has a repre- Those from St. Croix, however, are better allo- sentative (Ardops) in the Lesser Antilles. Only cated to the Lesser Antillean M. m. debilis, al- Monophyllus plethodon occurred on Puerto Rico though showing intergradation with fortis. and still occurs in the northem Lesser Antilles, but is not known to occur on the Virgin Islands. ZOOGEOGRAPHICAL RELATIONSHIPS Of course, since no Pleistocene fossil remains of BETWEEN LESSER AND GREATER bats are known from the Virgin Islands, there is ANTILLEAN BAT FAUNAS no reason to assume that the species did not occur there. It may since have become extinct Both Jones and Phillips (1970) and I (Koop- there as it has on Puerto Rico. The remaining man, 1968) have discussed the bats of the nine species are not known to occur east of northern Lesser Antilles (north of Guadeloupe Puerto Rico in the West Indies. but not including the Virgin Islands). Excluding The subgenusNatalus poses a special problem. the dubious record of Carollia perspicillata from Natalus stramineus is known from several Lesser Redonda, there are nine species: Noctilio lepo- Antillean islands. The closely related N. major rinus, Monophyllus plethodon, Artibeus (Varona, 1974, considered them to be con- jamaicensis, Ardops nichollsi, Brachyphylla ca- specific) is known from Hispaniola, Jamaica, and vernarum, Natalus stramineus, Myotis nigricans Cuba, the Hispaniolan N. m. major being the (although judged from La Val, 1973, this may smallest and therefore most like N. stramineus. not be the correct name for the northern Lesser There is a considerable gap between the ranges of Antillean populations), Tadarida brasiliensis, and N. stramineus and N. major (nothing between Molossus molossus. Five (all except Mono- Anguilla and Saba on the east and Hispaniola on phyllus, Ardops, Natalus, and Myotis) have been the west). It is indeed odd that no Natalus is recorded from the Virgin Islands. Monophyllus known on Puerto Rico. There is, of course, plethodon is known (as a fossil) from Puerto always the possibility that it does or did occur. 1 975 KOOPMAN: VIRGIN ISLAND BATS 5

These bats are never abundant and are usually gone in the opposite direction. Particularly in the found in small numbers far back in large caves. light of Davis's revision (1973), I think it is The chance of fossilization is also poor as the impossible to say in which direction Noctilio Natalus skull is unusually delicately built and went and it is perfectly possible that some popu- therefore quite fragile. lation movement has gone in both directions. In short, there are nine Greater Antillean spe- This leaves for further discussion the three cases cies that reach (or reached) Puerto Rico, but are involving endemic West Indian genera. The find- not known farther eastward. Of these only ing by Silva and Pine (1969) that Brachyphylla is Phyllonycteris major was endemic to Puerto not a stenodermine, but a member of the other- Rico. Five species extend from Puerto Rico wise Greater Antillean Phyllonycterinae means through the Virgin Islands to the Lesser Antilles that all the species of this subfamily occur in the and of these, only Brachyphylla cavernarum does Greater Antilles and all but one are endemic to not occur on islands west of Puerto Rico. Even it. To me this means that movement of Brachy- here, there is a close relative, B. nana (including phylla cavernarum from the Greater to the Lesser B. pumila, both considered to be subspecies ofB. Antilles is far more probable than the reverse. cavernarum by Varona, 1974). One species, Schwartz and Jones (1967) revised the genus Monophyllus plethodon, occurred both on Monophyllus, reducing it to two species, allo- Puerto Rico (but not on islands west of Puerto patric except for Pleistocene sympatry on Puerto Rico) and in the Lesser Antilles, but is not Rico. It may still be considered to be more known from the Virgin Islands. Stenoderma diversified on the Greater than on the Lesser rufum is endemic to Puerto Rico and the Virgin Antilles, but the difference is slight. I still think Islands, but has a representative in the Lesser Monophyllus was probably derived from Glosso- Antilles (Ardops) and others on islands west of phaga longirostris (Varona, 1974, considered Puerto Rico (Phyllops, Anteus). One species them to be congeneric) and the origins ofMono- group (probably at most a superspecies), Natalus phyllus in the Lesser Antilles and its later spread stramineus-major occurs in the Lesser Antilles to the Greater Antilles seems most probable to and the islands west of Puerto Rico, but not me. Jones and Schwartz (1967) also revised from the islands between. Finally, a member of Ardops and recognized a single Lesser Antillean the Myotis nigricans group occurs in the northern species. However, here the Greater Antillean Lesser Antilles, but has no representative in the representation of the group to which it belongs is Virgin Islands, Puerto Rico, or any islands farther much richer. There are at least four good species west. Thus, whereas a majority of Puerto Rican (haitensis may be only a subspecies of Phyllops bats are Greater Antillean forms that do not falcatus), some of them quite distinct. It seems reach the Lesser Antilles, the northem Lesser most reasonable, therefore, to regard the Steno- Antilles are dominated by species that also occur derma complex as having originated in the in Puerto Rico, the Virgin Islands, or the more Greater Antilles and the Lesser Antillean Ardops western Greater Antilles. This might imply that as being derived from it, albeit earlier than other the Greater Antilles rather than the Lesser invasions into the Lesser Antilles. My count of Antilles was the main source for species going the eight passages stands, therefore, at four from "around the comer" of the Antilles. west to east (Artibeus, Ardops, Brachyphylla, Jones and Phillips (1970) and I (Koopman, Tadarida), three from east to west (Monophyllus, 1968) have discussed this general problem, but I Natalus, Molossus), and one unknown (Noctilio). think it should be re-examined in a broader con- text, using any additional new data that may be ZOOGEOGRAPHICAL RELATIONSHIPS OF available. Of the eight passages of bats here VIRGIN ISLAND BATS enumerated between the Greater and Lesser Antilles, I have no reason to change my earlier The Virgin Islands actually form two quite opinion that two (Artibeus jamaicensis, Tadarida distinct geographical units. Vieques, Culebra, St. brasiliensis) have clearly gone from the Greater Thomas, St. John, Tortola, Virgin Gorda, to the Lesser Antilles, whereas two others Anegada, and most of the smaller islands are on (Natalus stramineus, Molossus molossus) have the Puerto Rican bank, which formed a land 6 AMERICAN MUSEUM NOVITATES NO. 2581 extension of Puerto Rico up until about 10,000 Noctilio leporinus. Jour. Mammal., vol. years ago (Heatwole and MacKenzie, 1967). St. 54, pp. 862-874. Croix (together with a few small associated Dobson, G. E. islands), on the other hand is on its own small 1878. Catalogue of the Chiroptera of the Brit- bank, separated from the Puerto Rican bank by a ish Museum. London, British Museum (Natural History), 567 pp. deep channel, the Anegada Passage. St. Croix Goodwin, G. G. might be expected to lack some Puerto Rican 1928. Observations on Noctilio. Jour. elements present on the other Virgin Islands but Mammal.,vol. 9,pp. 104-113. absent in the Lesser Antilles. This is true only of Hall, E. R., and J. W. Bee Stenoderma rufum. All four bat species known 1960. The red fig-eating bat Stenoderma from St. Croix are present on both Puerto Rico rufum Demarest found alive in the West and the Lesser Antilles. The only indication of Indies. Mammalia, vol. 24, pp. 67-75. any special Lesser Antillean affinity on St. Croix Hall, E. R., and K. R. Kelson is the fact that the St. Croix Molossus molossus is 1959. The mammals of North America. New more like M. m. debilis of the Lesser Antilles York, Ronald Press, 2 vols., 1083 pp. Hall, E. R., and J. R. Tamsitt than like M. m fortis of other Virgin Islands and 1968. A new subspecies of the red fig-eating Puerto Rico. All the other Virgin Islands have, as bat from Puerto Rico. Royal Ontario might be expected, a depauperate Puerto Rican Mus., Life Sci. Occas. Papers, no. 11, bat fauna (particularly depauperate on the pp. 1-5. smaller, drier islands). As is usual in border zones Heatwole, H., L. Kelts, R. Levins, and between distinctive faunas, there is not so much F. Torres a mixing of distinctive elements as a disap- 1963. Faunal notes on Culebra Island, Puerto pearance of distinctive elements of both faunas, Rico. Caribbean Jour. Sci., vol. 3, pp. leaving only the widespread varicants. (See 29-30. Hershkovitz, 1958, p. 589, for use of this term.) Heatwole, H., and F. MacKenzie This is true of the Virgin Islands and it is also 1967. Herpetogeography of Puerto Rico. IV. Paleogeography, Faunal Similarity and true of the northern Lesser Antilles. The Endemism. Evolution, vol. 21, pp. "coiner" of the Antilles is indeed an area that 429-438. lacks most of the distinctive elements of both Hershkovitz, P. Greater and Lesser Antilles. 1958. A geographic classification of Neo- tropical mammals. Fieldiana: Zool., LITERATURE CITED vol. 36, pp. 579-620. Jones, J. K., and C. J. Phillips Allen, G. M. 1970. Comments on systematics and zoo- 191 1. Mammals of the West Indies. Bull. Mus. geography of bats in the Lesser Comp. Zool., vol. 54, pp. 175-263. Antilles. Studies on the Fauna of Allen, J. A. Curacao and other Caribbean islands, 1890. Notes on a small collection of West vol. 32, pp. 131-145. Indian bats, with description of an Jones, J. K., and A. Schwartz apparently new species. Bull. Amer. 1967. Bredin-Archbold-Smithsonian biological Mus. Nat. Hist., vol. 3, pp. 169-173. survey of Dominica. 6. Synopsis of Bond, R. M., and G. A. Seaman bats of the Antillean genus Ardops. 1958. Notes on a colony of Brachyphylla Proc. U. S. Natl. Mus., vol. 124, no. cavernarum. Jour. Mammal., vol. 39, 3634, pp. 1-13. pp. 150-151. Koopman, K. F. Choate, J. R., and E. C. Birney 1968. Taxonomic and distributional notes on 1968. Sub-recent Insectivora and Chiroptera Lesser Antillean bats. Amer. Mus. Novi- from Puerto Rico, with the description tates, no. 2333, pp. 1-13. of a new bat of the genus Stenoderma. La Bastille, A., and M. Richmond Jour. Mammal., vol. 49, pp. 400-412. 1973. Birds and mammals of Anegada Island, Davis, W. B. British Virgin Islands. Caribbean Jour. 1973. Geographic variation in the fishing bat, Sci., vol. 13, pp. 91-1 09. 1975 KOOPMAN: VIRGIN ISLAND BATS 7

La Val, R. for the relationships between the bat 1973. A revision of the Neotropical bats of genus Brachyphylla and the Phyllonyc- the genus Myotis. Nat. Hist. Mus. Los terinae. Biotropica, vol. 1, pp. 10-19. Angeles Co., Sci. Bull. 15, pp. 1-54. Starrett, A. Schwartz, A. 1962. The bats of Puerto Rico and the Virgin 1955. The status of the species of the brasil- Islands, with a check-list and keys for iensis group of the genus Tadarida. identification. Caribbean Jour. Sci., vol. Jour. Mammal., vol. 36, pp. 106-109. 2, pp. 1-7. Schwartz, A., and J. K. Jones. Starrett, A., and F. J. Roile 1967. Bredin-Archbold-Smithsonian biological 1963. A record of the genus Lasiurus in survey of Dominica. 7. Review of bats Puerto Rico. Jour. Mammal., vol. 44, p. of the endemic Antillean genus Mono- 264. phyllus. Proc. U.S. Natl. Mus., vol. 124, Vahl, M. no. 3635, pp. 1-20. 1797. Beskivelse paa tre nye Arter Flager- Shamel, H. H. muse. Skr. Naturhist., Selsk. Kjoben- 1931. Notes on the American bats of the havn, vol. 4, pp. 121-138. genus Tadarida. Proc. U. S. Natl. Mus., Varona, L. S. vol. 78, pp. 1-27. 1974. Catilogo de los Mamiferos Vivientes y Silva Taboada, G., and R. H. Pine Extinguidos de las Antillas. Acad. Sci. 1969. Morphological and behavioral evidences Cuba, 139 pp.