University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Mammalogy Papers: University of Nebraska State Museum, University of Nebraska State Museum

2003 of Nevis, Northern Lesser Antilles Scott .C Pedersen South Dakota State University, [email protected]

Hugh H. Genoways University of Nebraska - Lincoln, [email protected]

Matthew .N Morton Durrell Wildlife Conservation Trust, [email protected]

James W. Johnson Whitehall, Nevis

Siân E. Courts Durrell Wildlife Conservation Trust

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Pedersen, Scott .;C Genoways, Hugh H.; Morton, Matthew N.; Johnson, James W.; and Courts, Siân E., "Bats of Nevis, Northern Lesser Antilles" (2003). Mammalogy Papers: University of Nebraska State Museum. 263. http://digitalcommons.unl.edu/museummammalogy/263

This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Mammalogy Papers: University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Pedersen, Genoways, Morton, Johnson & Courts in Acta Chiropterologica (2003) 5(2): 251-267. Copyright 2003, Polish Academy of Sciences. Used by permission.

Acta Chiropterologica, 5(2): 251-267, 2003 PL ISSN 1508-1109 © Museum and Institute ofZoology PAS

Bats of Nevis, northern Lesser Antilles

3 SCOTT C. PEDERSEN', HUGH H. GENOWAYS2, MATHEW N. MORTON , JAMES W. JOHNSON4, and SLAN E. COURTS 5

'South Dakota State University, Brookings, South Dakota 57007, USA; E-mail: [email protected] 2University ofNebraska State Museum, University ofNebraska-Lincoln, Lincoln, Nebraska 68588, USA 3c1o 22 Kensal Avenue, Bedminster, Bristol, BS3 4QY, United Kingdom 4Whitehall, Parish ofSt. James, Nevis, Eastern Caribbean 5Durrell Wildlife Conservation Trust, Trinity, Jersey, JE3 5BP, United Kingdom

Only one species of , Molossus molossus, previously has been documented as occurring on the northern Lesser Antillean island of Nevis. Field research and reviews of existing museum collections have provided documentation based on voucher specimens for an additional seven species occurring on the island ~ Noctilio leporinus, cavernarum, plethodon, Ardops nichollsi, jamaicensis, Natalus stramineus, and Tadarida brasiliensis. The biological diversity of the chiropteran fauna on Nevis is similar to that found on other islands in the northern Lesser Antilles. Ecologically, this is a simple chiropteran fauna, including one piscivore (N. leporinusi, one omnivore (B. cavernarumi, one pollenivore/nectivore (M. plethodoni, two frugivores (A. nichollsi, A. jamaicensisi, and three insectivorous species (N. stramineus, T. brasiliensis, M. molossus). Species-area and species-elevation analyses for the chiropteran fauna of the Greater and Lesser Antillean islands gave ?-values of 0.74 and 0.33, respectively. Tn the species-area analysis the bat fauna ofNevis falls above the regression line and in the species-elevation analysis it falls almost on the line. The chiropteran fauna of Nevis lies outside the Lesser Antillean Faunal Core and would be best characterized as a generalized Lesser Antillean fauna that appears to be characteristic of the northern Lesser Antilles.

Key words: Chiroptera, biogeography, ecology, systematics, island, Nevis, West Indies

INTRODUCTION currently is the accepted scientific name for this taxon (Dolan, 1989). Bats are the only native Islands ofsimilar size and topography to known to occur on Nevis. The first and only Nevis in the northern Lesser Antilles typi­ published report of a bat from the island of cally support six to 10 species ofbats. With Nevis was by Miller (1913) when he report­ only one species of bat documented in the ed Molossus molossus. This record was not­ scientific literature for Nevis, it appeared at ed in the description of Molossus debilis, the beginning ofour study that either the is­ with a type locality of St. Kitts and other land possesses a depauperate fauna or this is records from Antigua and Montserrat. The a simple case of under-sampling. To exam­ name Molossus molossus molossus with ine this question, two field surveys of the the type locality restricted to Martinique bats of Nevis were performed and visits 252 S. C. Pedersen, 1-1 . H. Genoways, M. N. Morton, J. W.Johnson, and S. E. Courts were made to the major research collections climate with high temperatures ranging to 30.5°C in in order to examine voucher specimens col­ August and Septemb er and low temperatures ranging lected from throughout the region for com­ to 22.2°C in February. The majority of Nevis is relatively low volcanic parative purposes. Although some mist-net­ country, which is under cultivation or is covered with ting was performed during the first survey secondary scrub and thorn-bush, known in the West by M.M. , S.C. , and J.J. in 1999, that effort Indies as ruinate vegetat ion (Beard, 1949). The dom­ focused primarily on roost surveys, aiming inant vegetation in these low-lying areas include I to provide a resource for future work con­ birch gum iBursera simarubai, loblolly (Pisonia sub­ cordata and P. fragra nsi, dogwood (Lonchocarp us cerning sites that would benefit from local latifoliusy; and white cedar (Tabebuia pallida). The conservation efforts and indicated that fur­ slopes of the mountain are so steep and exposed in ther study would be necessary. The survey most places that the low secondary forest adjacent to conducted by S.P. and J.J . in July 2001 pro­ the cultivated fields is quickly replaced by palm vides the first results from mist netting of bats in a wide variety of foraging habitats on Nevis. The sur vey of museum holdings re­ vealed that some unreported earlier work on Nevis had produced voucher specimens. In 1937 Chester Roys visited Nevis and l rlO ·OO~ collected 18 individuals of M. molossus I -/------+------+-- in Charleston on June 12. These speci­ mens were subsequently deposited in the Field Museum ofNatural History in Chic a­ go. David W. Nellis also collected bats when he visited Nevis between April 4 to 7, 1980. He collected at three sites and obtained individuals representing four species that were subsequently depo sited in the in American Museum of Natural History in New York. We have combin­ 5km ed the results of our field surveys with the data from these earlier efforts to provide FIG. I. Mist-netting localities and roost sites of note a description of the chiropteran fauna of on Nevis. Numbers refer to locations shown in the figure: I) Combermere School: roost within semi­ Nevis. abandon ed school building; 2) Mt, Pleasant: tarrish pits/caves situated among ruin s; 3) Butlers M ATERIALS ANDM ETHODS Government storage build ings; 4) Butlers Source: bog situated in deep ravine with moderate canopy Study Area cover; 5)New River Source: pools along river bed under dense canopy; 6) New River Estate buildings. Nevis is a small island located at i7 °10'N, 7) Golden Rock Estate: along trail above estate 62°35'W in the northern Lesser Antilles. Nevis is house; 8) Zetland s; 9) Herbert Heights; 10) Old joined politically with the island ofSt. Kitts in the na­ Manor Estate buildin gs; II ) Rosehill: along Handley tion of SI. Kitts and Nevis. Even by the standards of road ; 12) Indian Castle Estate buildings; 13) the Lesser Antilles, Nevis is quite small with an area Montpell ier Estate buildings; 14) M'Lady Well. 15) of 93 km2 (Fig. I). Nevis is nearly round with an av­ Prospect Estate; 16) Hamilton's Estate: along gravel erage diamete r of 12 km and is dominated by a steep acce ss roads ; 17) Charlestown: collected from volcanic cone reaching 985 m. The adjacent island building s; 18) Barnes Ghaut: flyway along gravel ofSt. Kitts lies across a shallow 3.5 km wide channel road above Jessup; 19) Nelson Spring. 20) Cades to the northwest of Nevis. Nevis has a mild tropical Estate ·'·Bats of Nevis 253

brake, which continues to the summit. The vegetation Roost Surveys in this area is primarily (50--60%) mountain.cabbage (Euterpe globosa sand 15% tree ferns (Cyathea arbo­ An additional 141 bats (Brachyphylla caverna­ rea), with such associated trees as blue tongue (Ma­ rum, Tadarida brasiliensis, Molossus molossus) were rila racemosa ), pigeonberry (Hirella triandra), and captured by hand from a dozen bat roosts, caves, and blue box (flex sideroxyloides). Good rain forest is buildings during the two survey periods February 2 only present on the northwestern face of Nevis Peak through March 13, 1999, and July I to 8, 2001. above Jessop's. Two small areas of other vegetation I types are dry evergreen forest along the top of a high ridge that runs to the east from the primary peak and Visual Records montane thicket that is intermediate between the palm brake and the rain forest above Jessop's. Natalu s stramineus was observed on numerous Historically, several hurricanes have hit the re­ occasions in a cave on the Mount Pleasant Estate. One gion, especially Hurricane Georges, which caused ofus (1.1.) has observed greater fishing bats on at least considerable damage on St. Kitts and Nevis when it two occasions on the west coast of Nevis. made landfall southeast of Basseterre, St. Kitts, on September 21, 1998 (see also Rodriquez-Duran and Measurements Vazquez, 200 I). In subsequent years, Hurricanes Jose and Lenny in September and November 1999, respec­ Voucher specimens from the 200 I survey were tively, have had major impacts on the economy and deposited in the research collections at the University forest ecology on Nevis. of Nebraska State Museum. All measurements are in millimeters and body masses are given in grams. Forearms and crania were measured with digital Mist-n etting calipers (Table 2). External measurements other than forearm are those recorded on specimen labels by col" Mist-netting for bats was conducted in a variety lectors. of habitats, including sites associated with pools of water in rivers or bogs, flyways adjacent to fruiting SYSTEMATIC ACCOUNTS mango trees, fig trees, and piper plants (Piper di­ latumi, and access roads during the two census peri­ ods. Twenty net-nights were conducte d between Noctilio leporinus mastivus February 6 and March 10, 1999 (6 captures), and 34 (Vahl, 1797) net-nights between July 2 to 8, 2001 (56 captures). Netting efforts during the latter survey involved erect­ Specimen examined(l). - St. Thomas ing five or six mist-nets at each site and monitoring Lowland Parish: Nelson Spring [17° I I'N, them for four to six hours depending on activity and weather. Together, the two mist-netting efforts pro­ 62°37'W], I (AMNH). vided 59 captures of six of the eight species of bat Davis (1973) reviewed geographic vari­ found on Nevis (Table I). ation in Noctilio leporinus. He assigned all

TABLE I. Bat captures (including the year) on northern Lesser Antillean island of Nevis ------,------.- - - - ,, -- - - ,,- _ _ _Roost Captures_ _ Habitat Captures Species Total 1937 1999 2001 1980 1999 2001 Noctilio leporinus visual I I Monop hyllus plethodon 1 1 Artibeus jamaicensis 4 21 27 53 Ardops nicholsi 19 19 Brachyphylla cavernarum 5 9 12 2 29 Natalu s stramin eus 1* 1 Tadarida brasiliensis 10 9 6 25 Molossus molos sus 18 98 6 7 130 259 ------,, ------_.- * killed by ceiling fan, May 5, 2003 T ARLE tv 2. Length of forearm and cranial measurements for eight species of bats occurring on the northern Lesser Antillean island ofNevis Vl ~ - _._----_._....~-- --_..--_.----_•...•._-_. ------~--- . - ..__ ._.._--_._ .~- ._------Length of Catalog numbers, Length of Greatest length Condylobasal Zygomatic Postorbital Mastoid Breadth across maxillary statistics, and sex forearm of skull length breadth breadth breadth upper molars toothrow -_._-----,.__.._-- _.__. _----_.~.•- --_.. ------,-- Noctilio leporinus mastivus ~ AMNH 246993, s> 83.8 26.6 23.3 18.5 7.0 17.1 9.9 12.0 (') "tl (l> Brachyphylla cavernarum cavernarum 0.. (l>.., UNSM 27649, 0 64.4 31.8 28.1 17.6 6.5 15.1 10.8 11.8 ", UNSM 27663, 0 65.0 31.8 28.5 17.2 6.7 14.7 II.I 11.4 .?'" UNSM 27665, 0 64.4 32.4 28.8 17.3 6.6 15.3 11.2 11.6 ::r: UNSM 27667, 0 64.7 32.2 28.6 17.8 6.6 15.2 10.9 11.6 ::I: 0 Monophyllus plethodon luciae :s 0'" UNSM 27670, 0 39.6 23.1 21.4 10.2 4.6 9.8 7.9 5.6 ~ co -e Ardops nichollsi montserra tensis .'" UNSM 27653, 0 50.1 22.5 19.6 14.9 5.8 12.2 7.5 9.8 ~ UNSM 27654, s> 50.0 24.3 21.4 15.8 7.1 12.9 8.0 10.6 Z ~ Artibeus jamaicensis jamaicensis 0 :4 00 n 6 6 6 6 6 6 6 6 0 x ± SE 59.4 ± 0.56 28.2 ± 0.25 25.1 ± 0.21 16.9 ± 0.10 7.3 ± 0.08 14.8 ± 0.09 9.9 ± 0.14 12.6 ± 0.19 .? ~ (Range) (57.9- 61.4) (27.5-29.2) (24.6-25.9) (16.5-17.2) (7.1-7.6) (14.4-15.0) (9.6-10.4) (12.0-13.2) ~ S> S> n 4 4 4 4 4 4 4 4 '- 0 x ± SE 60.5 ± 0.4 027.9 ± 0.27 24.9 ± 0.25 16.8 ± 0.04 7.1 ± 0.06 14.6 ± 0.05 9.8 ± 0.08 12.3 ± 0.14 5 ", (Range) (59.8-61.6) (27.2-28.4) (24.2-25.3) (16.7-1 6.9) (6.9- 7.2) (14.4-14.6) (9.6-9.9) (11.9-1 2.5) 0 .? co Natalus stramineus stramineus :s UNSM 28159, 0 39.5 17.0 15.6 8.7 3.3 7.6 7.4 5.7 0.. ~ Tadarida brasiliensis antillularum rt'l UNSM 27675, 0 37.7 15.9 14.8 9.5 3.7 9.1 5.6 6.5 o e0 AMNH 247029, 0 37.1 15.7 14.7 9.4 3.6 8.9 5.5 6.7 :4 AMNH 247030, 0 37.9 15.8 14.8 9.7 3.7 9.0 5.5 6.7 ", AMNH 247031, 0 37.4 15.8 14.6 9.2 3.7 8.7 5.4 6.5 UNSM 27674, s> 38.9 15.2 14.7 9.5 3.7 9.0 5.5 6.2 AMNH 247032, s> 37.9 15.6 14.3 9.0 3.6 8.5 5.5 6.5 0_0' ___ ._-- _ . - _ ._--..,-- - - ".__._- - -_.-_...._-- -_._- - _ .__ ...._-_..,_.,--_._ ._._----'•..~. _ --- - ,------_. . _.. Bats of Nevis 255

specimens from the Antillean islands and circum-Caribbean mainland to the subspe­ cies N. I. mastivus, which he recognized based on it large-size for the species (Table 2).The specimen of fishing bat that we ex­ amined was taken on the night of April 7, 1980, by D. W. Nellis at Nelson Spring in northwestern Nevis. It isa non-pregnant adult female. Nelson Spring is a small brackish water lagoon located on the lee­ ward (western) side ofNevis. This lagoon is fed by a small spring, surrounded by marshy ground and dense sedges, and occa­ sionally opens into the sea. The depth is about 0.5 to I m with a clay bottom. Small fish, shrimp, and crabs come into the area and stay. Fish species observed in the spring included juvenile mangrove snapper, small barracuda, young gars (needlefish), and ti­ lapia (introduced). One of us (1.J.) has observed greater fishing bat tNoctilio leporinus) on at least two occasions on the island. A deep recess located at Cades Point, St. Thomas Low": land Parish, contained three N leporinus "i when visited on one occasion in the early J;>..c:'" '"0 ­ ~ !>l) 1990's. Unfortunately the building of a re­ >. S:: "0 0 s::~ sort (Cliffdwellers) on the overlying bluffin o U 1994, disturbed the bats sufficiently that they vacated this shelter soon thereafter. When this Cades Point site was visited in '

I dock atthe Four Seasons Resort [2.0 km N I I Charleston, St. Thomas Lowland Parish] in I the late evening (lO:45 and 11 :00 P.M.). "0"00+ 0+ Several of the bats were observed catching -:'NO-:' r-. t> l.r) l£) small fish of an unknown species, although 11 \0 \0 \0 \0 t-- r-- t-- r­ N N N N large numbers of needlefish (14-16 mm ~~~~ ~ I Vl Vl Vl Vl in length) were observed to be schooling ~ ! 5555 , nearby.

------_._---_. .

256 S. C. Pedersen, H. H. Genoways, M. N. Morton, J. W. Johnson, and S. E. Courts

There is some question in the museum Vincent. This subspecies was distinguished records about the location ofNelson Spring, by its large size. Our specimens from Nevis which in part can be attributed to confusion support this decision because they match about the name of country - St. Kitts and closely the mean and range ofsamples from Nevis - and the name of the island - Ne­ Montserrat and St. Martin (Swanepoel and vis. According to the U. S. Board on Geo­ Genoways, 1978). graphic Names gazetteer number 7 (British Reproductive data are sparse for B. ca­ West Indies and Bermuda) there is only one vernarum on Nevis. Four of eight males Nelson Spring in the former British posses­ taken on April 6, 1980, had testes lengths sions in the Lesser Antilles. This place is of6.5, 7, 9, and 10. Two ofthe four females located at 17° 11 'N, 62°3TW, which places taken on this date carried embryos, meas­ it in the northwestern part of the island of uring 7.5 and 22 in crown-rump length. Nevis near the village of Cotton Ground. During February of 1999, one male and Nelson Spring does appear on the 1984 three females were non-reproductive, and United Kingdom topographic map of the a single female was clearly a juvenile with island. unfused phalangeal epiphyses. In 200 I, 10 males were found to be non-reproductive Brachyphylla cavernarum cavernarum and the single female captured at Hamil­ Gray, 1834 ton's Estate was lactating (July 5, 2001). The male from near Zion taken on July Specimens examined (22). - St. James 4 had a testis length of 6.5 and those of Windward Parish: New River Source, 0.6 three individuals from Mt. Pleasant Upper km N, 0.8 km W Zion, 180 m, 1 (UNSM); Cave taken on July 7 were 6, 7, and 8. The Mt. Pleasant Upper Cave, 0.8 km S, 0.8 km male from near Zion weighed 53.1 g when E Mt. Lily Estate, 9 (UNSM). St. John taken on July 4, whereas the nine males Figtree Parish: Prospect Estate [l7°08'N, from Mt. Pleasant Upper Cave collected on 62°36'W], 12 (AMNH). July 7 had a mean body mass of 50.1 g Specimens captured/released (7).- St. (47.9-51.4). John Figtree Parish: Hamilton 's Estate, 3.5 Brachyphylla has only been found in km E Charlestown, I; M'Lady Well, 0.5 km cave-like roosts on Nevis, however, in re­ S Prospect Estate, 3. St. George Gingerland cent years, such places have become in­ Parish: Herbert Heights, 0.5 km N Old creasingly scarce owing to the practice of Manor Estate, 1. St. James Windward Par­ filling in these large tunnels and pits for ish: Mt. Pleasant Cave, 0.8 km S, 0.8 km E public safety reasons. We have heard sever­ Mt. Lily Estate, 2. al reports that some of these chambers had Swanepoel and Genoways (1978) re­ housed large numbers of bats; one notable viewed the systematic relationships of example being a tunnel running several members of the genus Brachyphylla. They hundred meters east from Gingerland Meth­ recognized a single species - B. caver­ odist Church. narum - in the Lesser Antilles, ranging M'lady Well is a large vertical shaft from Puerto Rico in the Greater Antilles in thought to have been a mine for either sul­ the north to Barbados in the south. Their fur or mercury. It is located on the southeast analyses of the geographic variation in this corner of the island and serves as a large species lead to the assignment ofspecimens permanent roost for Brachyphylla. The 22 from St. Croix to St. Vincent to the nomi­ m deep shaft is reinforced with cut stone nate subspecies, with its type locality on St. and is and approximately 7 m in diameter Bats ofNevis 257

and is; surrounded by lowland desert scrub km southwest), or to other unknown roosts. and acacia. The sheer, vertical walls of this These movements may be related to the for­ defunct mine were covered with active mation of breeding aggregations, the avail­ B. cavernarum during July of 2001. M.M. ability offood, roost disturbance by humans and S.C. estimated that this colony included or livestock , or a complex balance of the 1,000 individuals in 1999 and no attempt at these factors. A conservative estimate ofthe a census was made in 2001. Due to the ease number of B. cavernarum at the Mt. Pleas­ of access to the area surrounding the en­ ant roost complex on July 8, 2001, was trance to this well by humans and livestock 6,000 located in the middle cave. - a potential safety hazard - this roost is Of note, bat guano is periodically collected in great need ofprotection by the local con­ by local agriculturists from the upper cave servation authorities. at Mount Pleasant. Each of the three 'caves' at the Mount During. periods of drought, hurricane, Pleasant site is a tarrish pit - a crude, pit or heavy ash fall, Brachyphylla may 'resort mine for fine volcanic gravel used in the to foraging on alternate forage such as manufacture of concrete. The upper cave legumes (pigeon peas ; Pedersen et al., is the largest (10m in diamet er, 7 m in dome 1996) or possibly the flowers, leaves, and height), sitting 75 m uphill from the ruins, young pods of false tamarind (Leuca­ while the middle and lower caves skirt ena leucocephala). The use of alternate the lower portions of the ruins. The middle forage by Brachyphylla is not unprecedent­ and lower caves were smaller, cooler, and ed because these bats have been observed more humid than the upper cave. The mid­ by one of us (J.J.) crawling out from sour dle cave is composed of two chambers, the orange fruits (Citrus aurantiumi during I - larger of the two was more or less spherical times of hard drought on Nevis. Bats are but did not exhibit a complete roof. The suspected of eating cocoa pods on Nevis smaller chamber (4-5 m in height) was in the Green Ghaut above Fountain Village. more enclosed and was occupied by nu­ As these soft pods ripen, bats can gain merous Brachyphylla, which were located access to the jellied inner mass. Monkeys along the walls of the cave within easy tear off the pods and eat them whole, rats reach from the cave floor. The lower cave chew the stem where they attach to the at the Mt. Pleasant site has a more tubular trees, but it appears that bats are perforat­ enclosed shape and was the most humid ing and eating them from the soft distal of the three caves. Brachyphylla were noted end while they are still attached to the tree. in the upper and middle caves, whereas only The liquid guano that accumulates under Natalus was noted in the lower cave. The the pods does not exhibit white uric acid area surrounding the ruins at Mt. Pleasant that characterizes the guano of fruit-fed is strewn with rocky outcrops, and is pre­ birds. dominantly covered by highland dry forest There is some question in the museum intermingled with larger trees in the deeper records about the location of Prospect Es­ ghauts. tate. According to the U. S. Board on Geo­ The B. cavernarum cave roosts at Mount graphic Names gazetteer number 7 (British Pleasant are not continuously occupied, West Indies and Bermuda) there is only one suggesting a frequent movement of bats Prospect Estate in the former British pos­ between the caves at Mount Pleasant, to sessions in the Lesser Antilles . This place is the large B. cavernarum roost at M'Lady located at 17°08'N, 62°36'W, which places Well on the opposite side of the island (6.5 it in the southwestern part of the island of ------

258 S. C. Pedersen, H. H. Genoways, M. N. Morton, 1. W. Johnson , and S. E. Courts

Nevis and the -1984 United Kingdom topo­ Artibeus and Ardops) foraging in the graphic map of the island shows 'Prospect canopy. Estate' at this location. Ardops nichollsi montserratensis Monophyllus plethodon luciae (Thomas, 1894) Miller, 1902 Specimens examined (3). - St. James Specimen examined ( I). - St. James Windward Parish: New River Source, 0.6 Windward Parish: Butler's Source, 0.75 km km N, 0.8 km W Zion , 180 m, 2 (UNSM); S, 2.3 km W Butlers, 360 m, 1 (UNSM). Butlers Source, 0.75 km S, 2.3 km W But­ Schwartz and Jones (1967) reviewed the lers, 360 m, 1 (UNSM). systematic relationships of members of the Specimens captured/released (16 ). ­ genus Monophyllus. They recognized a sin­ St. George Gingerland Parish: Golden Rock gle species in the Lesser Antilles, being rep­ Estate, 1. St. James Windward Parish: New resented on the islands of Anguilla, Barbu­ River Source , 0.6 km N, 0.8 km W Zion, da, Antigua, Dominica, and St. Lucia by 180 m, 4; Butlers Source, 0.75 km S, 2.3 km the subspecies M. plethodon lucia e. Subse­ W Butlers, 360 m, 11. quently, Baker et al. (1978) and Genoways Jones and Schwartz (1967) reviewed the et al. (200 I) have publi shed measurements genus Ardops, recognizing a single species ofseries ofthis taxon from Guadeloupe and - A. nichollsi - which is confined to the Dominica, respectively. The measurements Lesser Antilles. Determining the subspecif­ of our single male specimen fall at the low­ ic status of our specimens from Nevis is er end ofthe range ofmeasurements ofboth problematic, however. Plotting the Nevis of these samples in cranial measurements. specimens on to figure 1 of Jones and The length of forearm for the specimen Schwartz (1967) , places the male from Ne­ from Nevis is the shortest of all of these vis close to A. n. annectens with a type lo­ bats. Clearly, additional material will be cality of Guadeloupe, whereas the female needed to fully assess the meaning of these from Nevis falls with specimens of A. n. differences. External measurements of our montserratensis. We tentati vely assign our single male were as follows : total length , specimens to the latter taxon on geographic 75; length of tail, 12; length of hind foot, grounds because Nevis lies between 12; length of ear, 13 (for cranial measure­ Montserrat and St. Eustatius where A. n. ments see Table 2). This individual taken on montserratensis was reported by Jones and July 7 weighed 15.9 g and had testes that _Schwartz (1967). However, we believe the measured 3 mm in length. subspecific variation of this species should Althou gh 10 to 15 individuals ofM. ple­ be re-investigated because far more materi­ thodon were observed feeding on piper fruit al is now available than that studied by (Piper dilatum) at Butler's Source , capture Jones and Schwartz (1967). External meas­ success was severely hampered by difficul­ urements of a female from near Zion were ties in placing mist-nets along the steep hill­ as follows: total length, 82; length of hind sides at this location. This deep ravine was foot, 15; length of ear, 15. A single male filled with large fruiting trees and a small from this same locality weighed 23.7 g and stream. Indeed , the ground was covered the pregnant female taken on July 4 weigh­ with the fruits of pitch apples (Clau sia ma­ ed 34.3 g. (Table 2). j or) and hogplums iSpondia mombin) that The reproductive status of the 19 A. n. had been knocked down by bats (probably montserratensis captured during July 2001 Bats ofNevis 259 can be summarized as follows: four scrotal et al. (1989) and Pumo et al. (1996) have and nine non-scrotal males; three pregnant, presented mtDNA analyses of the relation­ two lactating, and one non-reproductive fe­ ships among Antillean populations of the male. The male captured on July 4 near . Specimens from Cuba Zion had a testis length of 5. The female and to which the subspecific from there taken on the same date was preg­ name parvipes is applicable, are smaller, nant with a near-term embryo, measuring both externally and cranially, than are those 32 in crown-rump length. The male cap­ from Jamaica south to Dominica and be­ tured on July 7 from near Butlers was a yond. A comparison of specimens in exter- subadult judging by its unfused phalangeal . nal or cranial dimensions from Jamaica epiphyses. The length offorearm for the lat­ southward to Dominica revealed no signifi­ ter specimen was 49.4, which approached cant differences among the populations. the size of the adult male, but its weight at Thus, the specimens from Nevis, which fit 18.5 was slightly less than 80% of that of within this, pattern. of variation, are appro­ the adult male. All Ardops captures were priately referred to the nominate subspecies. along rivers or in very boggy areas adjacent The six males and four females did not to trees in fruit and thickets of piper plants reveal significant secondary sexual varia­ (P dilatum). Individuals of Ardops were tion in any of the eight measurements (Ta­ noted feeding upon a variety of both 'small ble 2). Males did average larger than fe­ and large figs, pitch apples (c. major), and males in all measurements except length of hogplums (S' mombin). forearm. In the latter measurement, females averaged a millimeter longer than the Artibeus jamaicensis jamaicensis males. Two males taken on July 4 weighed Leach, 1821 41.7 and 42.0 g, whereas a pregnant female taken on that date weighed 42.3 g. Specimens examined (26). - St. James We hand-netted one of the clusters of Windward Parish: New River sugar mill A. jamaicensis in a disused house at Rose­ ruins [New River Estate, l7°09'N, hill, and found one adult male and three 62°33 'W], 6 (AMNH); New River Source, adult females. This is typical of the small 0.6 km N, 0.8 km W Zion, 180 m, 5 harems formed by males of this species (UNSM). St. John Figtree Parish: Prospect (Handley et al., 1991) - this type ofcluster Estate [17°08'N, 62°36'W], 1 (AMNH). St. was commonly noted at other sites during Thomas Lowland Parish: Nelson Spring roost survey work in 1999, All roosts of [17°11 'N, 62°37'W], 14 (AMNH). A.jamaicensis located during these surveys Specimens captured/released (27). ~ were assumed to be temporary due to the St. Thomas Lowland Parish: 1.5 km W Jes­ lack ofaccumulated guano at each site. sup, 2. St. George Gingerland Parish: Gold­ Two males taken on April 4 had testes en Rock Estate, 2. St. James Windward Par­ measurements of6 and 10 and those offour ish: New River Source, 0.6 km N, 0.8 km W males taken on April 7 were 7.5, 8, 9, and Zion, 7; Zetlands, 1; Rosehill, 4. St. John 10. Nine female Jamaican fruit-eating bats Figtree Parish: Hamilton's Estate, 3.5 km E from early April were examined for repro­ Charlestown, 6. St. James Windward Par­ ductive activity with the following results: ish: Butlers Source, 0.75 km S, 2.3 km W April 4, two pregnant, with crown-rump Butlers, 360 m, 5. length of embryos being 6 and 7; April 6, Genoways et al. (1998, 2001) have pre­ one non-reproductive; April 7, two preg­ sented morphometric analyses, and Phillips nant, with embryo crown-rump lengths of 260 S. C. Pedersen, H. H. Gcnoways, M. N. Morton, J. W. Johnson, and S. E. Courts

11 and 12, three lactating, and one non-re­ rather than with the nearby village of New productive, Together, the reproductive sta­ River, St. George Gingerland Pari sh. tus ofthe 27 Asjamaicensis captured during Jul y 2001 can .be summarized as follows: Natalus stramineus stramineus one juvenile, nine scrotal, and three non­ Gray, 1838 scrotal males; one juvenile, nine pregnant, one lactating, and three non-reproductive Specimen examined (1). - St. Thomas femal es; one animal was released before Lowland Parish: Cades Estate, 15 m, 1 I reproductive data could be regi stered. Of (UNSM). the three males and two femal es taken as The appropriate scientific name for spe­ vouchers on July 4, 2001 , testes lengths of cimens of this species coming from the is­ the two scrotal males were 5.5 and 8, the land of Nevis is N. stramineus stramineus third male was non-reproductive; one fe­ based on the work of Goodwin (1959) and male was non-reproductive while the sec­ Arroyo-Cabrales et al. (1997). Goodwin ond female was lactating. (1959) provided evidence that the type lo­ Three females collected at Nelson cality of N. stramineus was not Lagoa San­ Spring in 1980 , show varying degrees of ta, Minas Gerais, Brazil, and fixed the type alopecia. One individual (AMNH 247020) locality as Antigua in the Lesser Antilles. was naked on its cheeks and showed thin­ He assigned specimens from Anguilla and ning of the hair on its chin and upper che st. Dominica in addition to those from the type Another individual (AMNH 247026) was locality from Antigua to the nominate form . naked on its chin and upper chest, while the Handley and Gardner (1990) made a more third (AM NH 247025 ) was nak ed on the detailed investig ation of this situation, but crown of its head, chin, che st, and upper ab­ ultimately they agreed with Goodwin's domen. None of these indi viduals was not­ ( 1959) decision. Arroyo-Cabrales et al. ed as lactating. ( 1997) used gen etic data to study the rela­ Of the 10 specimens examined, all tionship among island and mainland pop­ lacked both M3 and all possessed both m3. ulations ofNatalus. They concluded that the Two other specimens did exhibit dental allozymic data supported the presence of a anomalies. A female (AMNH 247012) from single species in the Antillean islands to Pro spect Estate had mal-aligned lower teeth which the name N. stramineus would apply. with both ofthe m3 displaced lingually. The Forearm and cranial measurements of our other individual was a male from Nelson specimen from Nevis (Table 2) fall within Spring (AMNH 247014) that exhibited the range of a sample of nine males from heavily worn teeth. This specimen had part . Dominica (Genoways et al., 2001), but are of the crown of m2 arid all of the crown of noticeably larger than the measurements of m3 missin g on both sides of the lower jaws. a female reported from Guadeloupe (Baker Right i1 and i2 and left i1 were missing. et al., 1978). ' Left 11 and 12 were missin g, but the root of Our specimen was obtained on May 5; 12 was visible. All of the dental changes in 2003, by a 6-year old boy, Shaun Brear. Th e this male appeared to have occ urred during bat wa s found after it had collided with its life. The exact location of 'New River a ceiling fan and was brought to one of the sugar mill ruin s' (AMNH specimens) is not authors (J.J .) for identification. Testes of kno wn for certain, but in all likelihood the adult male measured 1.5 in length . In refers to the mill ruin s associated wit h New addition, three N. stramineus were observed River Estate, St. James Windward Pari sh, on Jul y 8, 2001, in a cave on the Mount Bats ofNevis 261

Pleasant Estate [1.25 km northeast from based on its small size. Finally, Schwartz the village ofFountain]. Although we were (1955) reduced this monotypic species to not able to capture any of these bats with subspecific rank within the widespread mist-nets or by hand, their proximity and mainland species Tadarida brasiliensis, characteristic external features left no doubt which is the arrangement that we have fol­ as to their identification. Their occupation lowed. of the smallest, most protected cave at External and cranial measurements of Mount Pleasant Estate (lower cave has a four males and two females (Table 2) agree distinctly tubular enclosed shape and was closely with those presented for bats from the coolest and most humid of the three Guadeloupe (Baker et al., 1978) and Do­ caves at the Mt. Pleasant site; 3 m wide, 6 m minica (Genoways et al., 2001). The male dome height) is not surprising given their from Scarborough weighed 9.0 and the predilection for humid microenviron­ pregnant female from there taken on July ments. One of us (J.J.) has observed these 8 weighed 11.7. In the March 1999 sur­ bats on several occasions in this lower cave vey, six of the 10 captured animals were during daylight hours throughout 2001 and non-reproductive males, two were post­ 2002, but their occupation of this roost has lactating and two were non-reproductive been inconsistent, often with the animals females. Two males taken on April 5, 1999, being r,resent or absent for several weeks at had testes lengths of 1.5 and 2 and the fe­ a time. male from this date evinced no reproduc­ tive activity. In July 2002, four of the nine Tadarida brasiliensis antillularum captured bats were non-reproductive ma­ (Miller, 1902) les, three were pregnant, and two were non-reproductive females. The male taken Specimens examined (9). - St. James on July 8, 2002, had testes measuring 3.5 Windward Parish: Combermere School in length. Both females taken on this [abandoned], Scarborough, 30 m, 3; New date were pregnant each with single em­ River sugar mill ruins [New River Estate, bryos measuring 11.5 and 13 crown-rump 17°09'N, 62°33'WJ, 6 (AMNH). length. Specimens captured/released (16). ­ Tadarida and Molossus roosted in small St. James Windward Parish: Comber­ clusters (5 to 100 animals) throughout the mere School [abandoned], Scarborough, 30 unused Combermere Primary School cen­ m,16. tered within the small community of Scar­ Miller (1902) described Nyctinomus an­ borough. The building is surrounded by tillularum from Roseau, Dominica, and a large, gravel, playing field. The nearest assigned specimens from Montserrat, tree was located 50 meters from the build­ St. Kitts, St. Lucia, and Tobago to the new ing. The numerous species-specific clus­ taxon. Miller (1902) distinguished the ters noted within the building were typical­ new species based on its size being small­ ly separated from each other by a distance er than populations in the Greater Antil­ of at least one meter and were most often les. Shamel (1931) in his revision of the associated with rafters or wedged into I ' genus Tadarida treated this species under crevices in the suspended ceiling. Despite the name T. antillularum. Shamel (1931) windows or doors that were ajar, the bats added material from Guadeloupe, Antigua, left through narrow cracks in the masonry and Puerto Rico to the monotypic species located at three of the four corners of the and continued to distinguish the taxon building. 262 S. C. Pedersen, H. H. Genoways, M. N. Morton, J. W. Johnson, and S. E. Courts

Molossus molossus molossus captured in February and March of 1999, (Pallas, 1766) each of the 45 males was non-reproductive. There were five pregnant females, eight Specimens examined (23). - St. James were post-lactation, 32 were non-reproduc­ Windward Parish: New River Source, 0.6 tive, and nine were juveniles. Of the 13 M. km N, 0.8 km W Zion, 180 m, 3; Com­ molossus captured in July 2001, four were bermere School [abandon], Scarborough, non-reproductive males, one was a non-re­ 30 m, 2. St. Paul Charlestown Parish: productive female, and the remaining eight Charlestown, 18 (FMNH). were pregnant. The two males from Scar­ Specimens captured/released (107). ­ borough in 2001 had testes lengths of 3 and St. George Gingerland Parish: Hotel, Mont­ 4. The three females captured in 2001 near pellier Estate, 1; Indian Castle Estate, 33. Zion were pregnant each containing a single St. James Windward Parish: Combermere embryo measuring 16, 16, and 18 in crown­ School [abandoned], Scarborough, 30 m, rump length. 17; Government storage buildings, Butlers, 52; Butlers Source, 0.75 km S, 2.3 km W Butlers, 360 m, 1. St. Thomas Lowland Par­ DISCUSSION ish: Barnes Ghaut, 3. Data presented here provide new and The previous record of Molossus from previously unpublished records of seven Nevis was noted in the description of Mo­ lossus debilis, with a type locality of St. species of bats for the island of Nevis: N. Kitts and other records from Antigua and leporinus, B. cavernarum, M. plethodon, A. Montserrat (Miller, 1913). This name and a nichollsi, A. jamaicensis, N. stramineus, number of others have been applied to this and T. brasiliensis. This clearly answers our species in the Antillean islands. It now is initial question about the number of species believed that the most appropriate name to ofbats occurring on the island and indicates apply to these bats is Molossus molossus, that the initial low number was a sampling which is a widespread Neotropical species. artifact. The biological diversity of the chi­ Husson (1962) restricted the type locality of ropteran fauna on Nevis is similar to that M. molossus to the island of Martinique, found on other island in the Lesser Antilles which lead Dolan (1989) to apply the name to the north of Guadeloupe (Baker and M. m. molossus to this species throughout Genoways, 1978; Jones, 1989). Ecologi­ the Lesser Antilles. Two males were col­ cally, this is a simple chiropteran fauna lected in the Combermere Primary School of eight species including one piscivore in Scarborough and two females were col­ (N. leporinus), one omnivore (B. caverna­ lected near Zion. The males taken on July 8 rum), one pollenivore/nectarivore (M. ple­ weighed 15.3 and 12.2 and the females, thodon), two frugivores (A. nichollsi, A. ja­ both pregnant, taken on July 4 weighed 13.6 maicensisi, and three insectivorous species and 14.3 (Table 2). (N. stramineus, T. brasiliensis, M. mo­ The 18 individuals obtained by Chester lossus), representing four families - No­ Roys in Charleston on June 12, 1937, in­ ctilionidae, Phyllostomidae, Natalidae, and cluded five males and 13 females. The Molossidae. males had testes length of4, 4.5, 4.5. 5, and In theory, the number of species found 5. All five of the females examined for re­ on an island is correlated with the size productive activity were pregnant with em­ (area) of the island, but this effect is influ­ bryos having crown-rump lengths of 10, 10, enced by other factors, including for ex­ 10.5, 10.5, and 12.5. Of the 99 M. molossus ample distance from a source area such ~: I

Bats of Nevis 263

as a continental mainland. The number 1989; Masson et af., 1990; McCarthy and of species occurring on an island also is Henderson, 1992; Genoways et af., 2001 ; dependent on the diversity ofhabitats avail­ Timm and Genoways, 2003) and data pre­ able, which in most cases is directly affect­ sented here. Under this general model, the ed by elevation of the island. Increased ele­ island of Nevis falls above the regression vation usually results in increased rainfall line relative to other islands in the region. and a more diverse vegetation (MacArthur, With the inclusion of new species records 1972). Species-area curve models have reported herein, the position of the island L been used in biogeographic studies of insu­ relati ve to the empirically derived curve has lar amphibians and reptiles (Preston, 1962a, been altered (former position indicated by 1962b), birds (Hamilton et al., 1964), and arrow) in a predictable manner, but it is now mammals (Morgan and Woods, 1986; Grif­ understood that the former position on the fiths and Klingener, 1988). For example, graph was simply because of under-sam­ Morgan and Woods (1986) studying the pling. The ?-value for the analysis using whole West Indian mammalian fauna found the cunent Nevis data is 0.74 , indicating L that 69% of the variance in species diversi­ that for the islands included in Fig. 2, size of ty could be explained by area of the islands the islands explains 74% of the species di­ alone. They concluded: "The remaining versity. The new position above the regre s­ 31% of the variance must be dependent sion line for the bat fauna ofNevis indicates upon other variables such as habitat diversi­ that there are more species ofbats on the is­ ty and distance from source areas. " (Morgan land than its size alone would predict. ~, and Woods, 1986). Because the remaining variation in We provide a species-area curve for the species diversity in part may be attributed to chiropteran fauna of the Greater and Lesser the elevation of the island and its resulting Antilles (Fig. 2), which includes data culled floral diversity, we compared the number of from the literature (Pinchon, 1967; Jones , species on the same islands to the elevation

1.4 Jamaica o

Dominica o Puerto Rico Grenada I SI. Vincent 0 Guad eloupe Monserrat 0 Marie • Galan te Martinique NeVIS 0 0 0 SI. Lucia Antigua 51.John I 51.Maa rtcn

y = 0.206x + 0.368, / = 0.74 0.2 -'------.------'-r------r------r------,.------' 1.0 2.0 3.0 4.0 5.0 Log (island area - km') FrG. 2. Species/area curve based on the published data (August, 200 I) for the chiropteran fauna in the Antilles. The vertical line indicates the shift in location of Nevis on this curve due to our report of an additional seven species ofbat from this island, from the long-standing record of a single species (M. molossusi on Nevis 264 S. C. Pedersen, H. H. Genoways, M. N. Morton, J. W. Johnson, and S. E. Courts of the island (Fig. 3). The r2-value is only the majority of the original chiropteran 0.33, much lower than found for species­ fauna of the island as compared to islands area analysis. Nevertheless, this analysis with lower elevations or uncomplicated does indicate that 33% of the bat species terrain. diversity on these islands could be ex­ Nevis is situated just to the north of the plained by elevation. The fauna from Ne­ islands that exhibit a fauna Genoways et at. vis falls almost on the regression line (2001) designated as the Lesser Antillean (Fig. 3). This indicates to us that the eleva­ Faunal Core. The eight species in the chiro­ tion of Nevis certainly is important in pteran fauna ofNevis can be divided in two general biogeographic units. The first is explaining the diversity of bats on the is­ composed of five species that have wide­ land and helps account in part for why it spread Neotropical distributions that extend falls above the regression line in the well beyond the Antillean to the mainland species-area analysis. Another important of South and Central America and Mexico fact to consider is that the interiors ofmoun­ - N. /eporinus, A. jamaicensis, N. stra­ tainous islands in the West Indies, such as mineus, M. motossus, and T. brasiliensis. Nevis, were harder to access by colonists These species are not informative about bio­ and consequently have suffered (and are geographic relations within the Lesser An­ suffering) less anthropogenic deforesta­ tilles because their distribution extend so tion. Returning to the issue of 'under-sam­ far beyond the West Indies. The second pIing', the inaccessibility of these habitats biogeographic unit is composed of three also creates logistical difficulties for biol­ species that are Lesser Antillean regional ogists attempting to adequately survey an endemics. These species are widely distrib­ island. However, it is exactly these sites uted in the Lesser Antilles and either do not that must to be sampled, as they may extend out of the region or only extend as be functioning as refugia, maintaining far as Puerto Rico - A. nichollsi occurs

1.4 o Cuba o Jamaica Hispanola 0 Puerto Rico 51.Vincent Guadeloupe Grenada 0 000 Dominica Monserrat 0 51.Lucia Marie Galante 0

o Barbuda 51.John / SI. Maarten o 0 St. Eustatius 51.Thomas o 51.Croix 0 51.Kilts

Virgin Gorda 0 0 0 Saba Tortola

y = 0.327x - 0.049, / = 0.33 0.2 L..-----r------,------..------,,.------r------' 1.5 2.0 2.5 3.0 3.5 Log (island altitude - m)

F IG. 3. Species/altitude curve based on the published data (August, 2001) for the ehiropteran fauna in the Antilles Bats ofNevis 265

from St. Eustatius to St. Vincent; B. caver­ provided logistical support for the 1999 Survey. narum occurs from Puerto Rico to Barba­ Funding was provided by the Totten Trust, United Na­ tions Development Programme (St. Kitts & Nevis dos; M. plethodon occurs from Anguilla to Biodi versity Project), and Mary Maxwell of the UK Barbados, with a fossil representative on Foreign & Commonwealth Office. We respe ctfully Puerto Rico. acknowledge Elvin Bailey ofthe Ministry ofAgricul­ One of the more interesting biogeo­ ture - Nevis, Raymond Solomon of the Department graphie characteristics of the bat fauna on ofthe Environm ent - St. Kitts, and Dr. Patricia Bart­ lette-Powell, Veterinary Officer - Nevis, for their Nevis relates to the species that are appar­ assistance and issuance of collecting and export per­ ently missing from the fauna, including mits. Curatorial support was provided by the Division improvisum, Sturn ira thomasi of Zoology, University of Nebraska State Museum or S. lilium, Myotis dominicensis, M. mar­ (UNSM), with special thanks to Patricia W. Freeman tiniqu ensis, or M. nigricans (Husson, 1960), and Thom as Labedz. For assistance and access to their collections we wish to thank Nancy B. Simmons and Eptesicus guadeloupensis or E. fuscus. and S. Jean Spank, American Museum of Natural The Lesser Antillean endemic species C. History (AMNH), and Bruce D. Patterson and Law­ improvisum and S. thomasi are known from rence R. Heaney, Field Museum of Natural History Montserrat (Jones and Baker, 1979; Peder­ (FMNH). sen et al., 1996; Genoways, 1998) and Gua­ deloupe (de la Torre and Schwartz, 1966; LITERATURE CITED Baker et al., 1978), M. dominicensis from Dominica (Genoways et al., 200 I) and Gua­ ARROYO-CABRA LES, J., R. A. VAN DEN BUSSCH E, K. H. SIGLER . R. K. CHESSER, and R. J. BAKER. 1997. deloupe (Masson and" Breuil, 1992), and E. Genic variation of mainland and island popul a­ guadeloupensis from Guadeloupe (Baker et tions of Natalus stramineus (Chiroptera: Nata­ al., 1978). These species along with S. lili­ lidae). Occasional Papers of the Museum , Texas um, which occurs on islands from Domini­ Tech University, 171: 1-9. ca southward, are important species that de­ BAKER, R. 1., and H. H. GENOWAYS . 1978. Zooge­ ography of Antillean bats . Acad emy Natur al fine the Lesser Antillean Faunal Core. The Science of Philadelphia Special Publication, 13: absence of any of these species on Nevis 563-597. confirms the conclusion of Genoways et al. BAKER, R. J., H. H. GENOWAYS, and J. C. PATTON. (2001) that the fauna of Nevis lies outside 1978. Bats of Guadeloupe. Occa sional Pape rs of of the Lesser Antillean Faunal Core. In the Museum , Texas Tech University, 50: 1- 16. BEARD, 1. S. 1949. The natural veget ation of the summary, the chiropteran fauna of Nevis Windward and Leeward Island. Oxford Univer­ would be best characterized as a generalized sity Press, London, England, 192 pp. Lesser Antillean fauna that appears to be DAVIS, W. B. 1973. Geographic variation in the fish­ characteristic ofmany islands located in the ing bat, Noctilio leporinus. Journal of Mam­ northern Lesser Antilles. malogy, 54: 862-874. DE LA TORRE, L., and A. SCHWARTZ. 1966. New species of (Chiroptera: Phyllostomidae) from the islands of Guadeloupe and Saint Vin­ A CKNOWLEDGMENTS cent, Lesser Antilles. Proceedings of the Biolog­ ical Society ofWashington, 79: 297- 303. We wish to thank the various Estate owners on DOLAN, P. G. 1989. Systematics of Middl e American Nevis for their good humor and access to their prop­ mastiff bats ofthe genus Molossus. Special Publi ­ ertie s. We wish to gratefully acknowledge the assis­ cation ofthe Museum, Texas Tech University, 29: I _ tance and support of Nikki Johnson, David and Joan 1-71. Robin son, the staff at the Nevi s Historical and Con­ GENOWAYS, H. H. 1998. Two new species of bats of servation Society, Edward and Michael Herbert, Mi­ the genu s Sturn ira from the Lesser Antille s, West randa Liburd, Jesse James, Jame s Eaton , Nancy Hall, Indies . Occa sional Papers of the Museum, Texas Hugh Price , Herb and Flo Taylor, Ben and Mari­ Tech University, 176: 1-7. lyn Carr. Mark Day of Fauna & Flora International GENOWAYS, H. H., C. J. PHILLIPS, and R. 1. BAK ER. 266 S. C. Pedersen, I-I. H. Genoways, M. N. Mort on, J. W. Johnson, and S. E. Co urts

1998. Bats of the Antillean island of Grenada : a MASSON, D., M. BREUI L, and A. BREUI L. 1990. new zoogeographic perspective. Occasional Pap­ Premier inventaire des chauves-souris de l'ile de ers of the Mu seum, Texas Tech Uni versity, 177: Marie-Galante (An tilles francaises), Mammalia, 1-28. 54: 656- 658. GENOWAYS, H. H., R. M. TIMM, R. J. BAKER, C. J. MCCARTH Y, T. J., and R. W. HENDERSON. 1992. Con­ PHILLI PS, and D. A. SCHLITTER . 200 I. Bats of the firmation of Ardops nichollsi on Marie-Galante, West Indian island of Dom inica: natu ral history, Lesser Antilles, and comments on other bats. areography, and trophic struc ture . Special Publi­ Caribbea n Journal of Science, 28 : 106-107. cations of the Mu seum. Texas Tech University. MILL ER , G. S., JR. 1902. Twenty new Ame rica n bats. 43 : 1-43. Proceedings of the Academy of Na tural Sciences GOODWI N, G. G. 1959. Bats of the subgenus Natalus. of Philadelphia, 54: 389-412. American Mu seum Novitates, 1977 : 1- 22. MILLER, G. S., JR. 19 13. Notes on the bats of the ge­ GRIFFITH S, T. A., and D. KLINGENER. 1988. On the nus Moloss us. Proceedin gs of the United State distribut ion ofGreater An tillea n bat s. Biotropica, Na tional Museum, 46: 85-92. 20: 240 -25 1. MORGAN, G. S., and C. A. WOODS. 1986. Extinction HAMILTON, T. 1-1. , R. H., BART H, and I. RUBI NOFF. and the zoogeography of West Indi an land mam­ 1964 . The environmental co ntrol ofinsular var ia­ mals. Biological Journal of the Linnean Society, tion in bird species abundance. Proceedings of 28 : 167- 203. the Nat ional Aca de my Scie nce , 52 : 132-140. PEDERSEN, S. C. 2001. The imp act of volcanic erup­ HANDLEY. C. 0. , JR., and A. L. GARDNER. 1990. The tions on the bat popul ations of Montse rrat, BWI. holotype of Na talus stramineus Gray (Mamma­ American Zoologist, 40: 1167A. lia: Chirop tera: Natalidae). Proceedings of the PEDERSEN, S. c, H. H. GENOWAYS, and P. W. FREE­ Biological Soc iety of Washington, 103: 966-972. MA N. 1996. No tes on the bats from Mon tserrat HANDLEY, C. 0 ., JR., D. E. WILSON, and A. L. G ARD­ (Lesser Antilles) , with co mments co ncerni ng the NER. 1991. Demo graph y and natural history of effects of Hurricane Hugo. Caribbean Journa l of the common fruit bat , Artibeus jamaicensis, on Scie nce, 32: 206-213. Barro Co lorado Island, Pana ma. Smith soni an PHILLI PS, C. J., D. E. PUMO, H. H. GENO WAYS, and Contributions to Zoology, 5 11: 1-173. P. E. RAY. 1989 . Caribbean island zoogeography: HUSSON, A. M. 1960 . De zoogdieren van de Neder­ a new approach using mitochondrial DNA to land se An tillcn. Uitgaven van de Natuurwetens­ study Neotropical bats. Pp. 66 1-684, in Biogeog­ chappe lijke Werkgroep Nederlandse Antille n, rap hy of the West Indi es (c. A. WOODS, ed. ). Curacao, 12: 1- 83. Sandhill Crane Press, Ga inesville, xv ii + 878 pp . HUSSON,A. M. 1962. Th e bats of Suriname. Z06 10­ PINCHON,R. 1967.Quelques aspects de la nature aux gische Mo nographicen, Rijksmuseum van Nat u­ Antilles . M. M. Ozanne and Cie., Caen, Fort-de­ urlij ke Histor ic, Leid en, 2: xxx iv + 1- 569. France , Mart inique, 254 pp. JONES, J. K., JR. 1989. Distribution and sys tema tics of PRESTON, F. W. 1962a. The canonical distribution of bat s in the Less er Antilles . Pp. 645-660, in Bio­ commonness and rarity: part I. Eco logy, 43: geography of the West Indies: past, pre sent, and 185- 215 . future (c. A. WOODS, ed.). Sandhill Crane Press PRESTON , F. W. 1962b. The canonical distribution of Inc ., Gai nesville, xvii + 878 pp com mo nness and rari ty: part n. Ecology, 43 : JONES, J. K. JR., and R. J. BAKER. 1979. No tes on 410-432. a co llec tion of bats from Mon tserrat, Lesser An­ PUMO, D. E., I. KIM, J. REMSEN, C. J. PHI LLIPS, and tilles. Occasional Papers of the Museum., Texas H. H. GENOWAYS. 1996. Mo lecu lar sys tema tics of Tech University, 60 : 1-6. the fruit bat, Artibeus jamaicensis: origin of an JONES, J. K., JR., and A. SCHWARTZ . 1967. Bredin­ unusual island pop ulation. Journal of ­ Archbo ld-Sm ithso nian Bio logical Survey of Do­ ogy, 77: 49 1-503 . min ica. 6. Synopsis of bats ofthe Antillea n genus RODRiGUEZ-D URAN, A., and R. VAZQUEZ, 200 I. The Ardops. Proceedings ofthe United States Natio n­ bat Artibeusjamaicensis in Puert o Rico (West In­ al Mu seum , 124 (363 4) : 1-13. dies): seasonality of diet, activity, and effect of a MACARTHUR, R. H. 1972. Geographical ecology -­ hurricane. Acta Chiropterologica, 3: 53-6 1. patterns in the distribution ofspecies. Harper and SCHWARTZ, A. 1955 . The status of the spec ies of the Ro w, New York, 270 pp . brasiliensis group of the genus Tadarida. Journa l MASSON, D., and M . BREUI L. 1992. Un Myotis (C hi­ of Mammalogy, 36 : 106-1 09. rop tera, Vespertilionidae) en Guadeloupe (Petites SCHWARTZ, A., and J. K. JONES, JR. 1967. Bredin­ Antille-s). Mammalia. 56: 473-475. A rchbold-Smi thsonia n Biologi cal ' Survey of Bats ofNevis 267

Dominica. 7. Review of bats of the endemic An­ of the Antillean bats of the genus Brachyphylla tillean genus Monophyllus. Proceedings of the (Mammalia: Phyllostomatidae). Bulletin of the United States National Museum, 124 (3635): Carnegie Museum ofNatural History, 12: 1-53. 1-20. TIMM, R. M., and H. H. GENOWAYS. 2003. West SHAMEL, H. H.. 1931. Notes on the American bats of Indian mammals from the Albert Schwartz Col­ the genus Tadarida. Proceedings of the United lection: biological and historical information. States National Museum, 78 (19): 1-27. Scientific Papers of the University of Kansas SWANEPOEL, P., and H. H. GENOWAYS. 1978. Revision Natural History Museum, 29: 1--47.

Received 11 February 2003, accepted 04 July 2003

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