OCCASIONAL PAPERS ...... __ .~_1IEl! __....._ Museum of Texas Tech University NUMBER 177 31 AUGUST 1998
BATS OF THE ANTILLEAN ISLAND OF GRENADA: A NEW ZOOGEOGRAPHIC PERSPECTIVE
HUGH H GENOWAYS, CARLETON J PHILliPS, AND ROBERT J BAKER
The island ofGrenada is the southernmost ofthe We have studied and collectedbats onGrenada on Lesser Antilles, lying 130 kmnorth ofTrinidad and 135 5 occasions-lO-12 August 1967 (Phillips); 22-26 May km north ofthe Venezuelan mainland. It measures 34 1980 (Baker and Genoways); 28 May to 2 June 1986 km north to southand 19 km eastto west andhas an area (Genoways andPhillips); 26 to 29 May 1987 (Genoways of311 square km. Grenada and the Grenadines north and Phillips), and 18-19 July 1989 (Phillips). Vegeta ward to Bequia stand on the large submarine Grenada tion and sites that we visited are discussed in the species Bank. At 183 m depth, the bank is 179 km long. The accounts below. Generally, there is great variety in the Grenadines cover the bank to its northern end, but the island's plant communities. Mostofthe original native bank extends for 39 km south of Grenada with no is forest was cleared during the plantation era so what is lands. During the last Ice Age, Grenada and the Grena seen today is primarily secondary growth. The Grand dines musthave beenone large island that extended from Etang Reserve in the southern mountains covers 1520 within 2 km ofSt. Vincent in the north to within 80 km ha andprotects the forest and the island's watershed. Be ofSouth America in the south. cause the prevailing wind is from the east, the western side ofthe island is relatively arid in comparison to the Grenada is a volcanic island with many extinct cra eastern coastal zone and montane slopes. In the south ters, including the one occupied by Grand Etang Lake, west and along the rocky west coast, vegetation is domi located at 530 m and covering 14 ha. A central ridge of nated by arid tropical and scrubland plants, including mountains runs like a north to south "spine" with Mount agave, prickly pearand other cacti, acacia, and calabash St. Catherine at 840 m the highest point. The island has trees. Other areas, especially the east coast, have man an average annual temperature of29°C. Rainfall varies grove swamps, low wet areas used for tropical agricul from 75 cm annually in the Point Salines area on the ture, and coconuts. The higher elevations have rainforest southwestern coast to 150 cm in other coastal areas to with trees, including mahogany, teak, saman, and blue 380 cm in the mountainous interior. The rainy season mahoe, reaching up to 30 m to form a canopy. These lasts from June to December, withNovemberusually the areas include a variety ofothernative plant species such wettest month. Grenada lies south ofthe primary hurri as bamboo, helicona, balsa, mountain cabbage palm, and cane belt and was last hit by a major hurricane in 1955, birdofparadise flower. Grenada is known as the Island when Hurricane Janet hit the island with winds ofup to ofSpice and has numerous exotic cultivated spices and 210kph. Neotropical fruit trees, including bananas, bay, breadfruit, 2 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY cocoa, guava, mango, nutmeg, papaya, passionfruit, pi from the island. In fact, it was the first species to be geonpeas, pimento, sugarapple, soursop, turmeric, and reported from Grenada (Dobson, 1878),butwe find no vanilla. evidence ofa contemporary population ofthis bat on Grenada. We, thus, considerthe one existing specimen Based on our studies, the chiropteran fauna of to be an erroneous record or an accidental occurrence. Grenada consists of12 species includingrepresentatives G. M. Allenwas the first scientistto make a systematic of the families Emballonuridae (1 species), collection ofbats from Grenada whenhe visitedthe is Mormoopidae (1), Noctilionidae (1), Phyllostomidae land in August and September of 1910. His report of (7), Vespertilionidae (1), and Molossidae (1). A thir this collection (Allen, 1911) recorded the occurrence teenth species, Carolliaperspicillata, hasbeenreported ofsix species on the island.
MEmODS Species accounts givenbeloware arrangedinsys meters (m), area in square kilometers (sq kIn) and hect tematic orderandpresentdata onsystematics, habitats, ares (ha), and rainfall in centimeters (cm). Statistical distribution, and reproduction for eachspecies. Alllin analyses were performedusing the StatViewe software earmeasurements are giveninmillimeters andwere taken package (Sager, 1992). The paired t-test gave standard with electronic calipers. Distances are recorded in ki statistics for each sample and statistical significance of lometers (km) or miles (mi), depending upon the units differences ingroup means. from the original collector. Elevations and depths are in
ACKNOWLEDGMENTS
We would like to thank the museums serving as ofNebraska State Museum (UNSM). Funding for our repositories for specimens used in this study, including workcame from a variety ofsources andrelatedprojects, AmericanMuseum ofNatural History (AMNH); British inparticularNIH (R01 to CJP and DEP), Hofstra Uni Museum (Natural History) (BMNH); Section ofMam versity HCLAS grants (CJP), and Research Council, mals, Carnegie Museum ofNatural History (CM); Mu University ofNebraska-Lincoln(HHG). We would like seum ofNatural History, University ofKansas (KU); to express gratitude to our field companions in doing MuseumofComparative Zoology, Harvard University this study, especially Dorothy E. Pumo (DEP) and Scott (MCZ); National Museum of Natural History, C. Pedersen. Figure 1was preparedbyAngie Fox, Staff SmithsonianInstitution (NMNH); The MuseumofTexas Artist, University ofNebraska State Museum. TechUniversity (TTU); Division ofZoology, University
SPECIES ACCOUNTS
Peropteryx macrotisphaea documents that the species has been a member ofthe G. M. Allen, 1911 chiropteran fauna ofthe island. Because the most re LesserDoglike Bat cent date ofcapture for this species is 29 August 1910, it is tempting to think that the species no longer inhabits Specimens examined (50).-ST. ANDREW: Mt. the island. The authors have spent extensive time sur Pleasant Estate [nearAdelphi], 24 (NMNH). ST. DAVID: veying areas where the species was taken previously, but Westerhall,2 (NMNH). ST. GEORGE: Point Salines, our work was almost exclusively done using mist nets. 23 (MCZ). PARISH UNKNOWN: no specific locality, G. M. Allen (1911) made the following statement con 1 (BMNH). cerning the collecting ofPeropteryx at Point Salines: " We found these bats in butone spot, a rather open cave Remarks.-This species was not takenduring our on the seacliffs at Point Saline [= Point Salines], the ex various surveys on Grenada; however, it was taken on treme southern end ofthe island. They clung by both several occasions by earlier investigators, which firmly hind feet to the rough surfaces ofthe rocks, usually in GENOWAYSETAL.-BATSOFGRENADA 3 well-shaded, overlumgingplaces; but, onbeing disturbed, (1976) stated no reasons for this decision in Venezuela, would flit farther into the darker recesses ofthe cave. but Brosset and Charles-Dominique (1990) working in Others, however, flew about under a tree near the mouth French Guiana distinguished the two taxa based on the ofthe cave, but eventually took shelter in adjoining fis differing shape ofthe ears and tragus and the shorter skull sures." A note on the specimens in the large collection (greatest length ofskull: 14.0-15.2 versus 13.2, 13.2) ofbats from Mt. Pleasant Estate stated "taken in rocks" with smaller teeth (length ofmaxillary toothrow: 5.3 and a note on the two specimens from Westerha11 Estate 6.0 versus 4.7, 4.8) ofP. trinitatus. Jones and Hood stated "taken in hollow rock." We have not searched rock (1993) in a synopsis ofSouth American emballonurids crevices and small caves on the island and this will be included both trinitatus and phaea as subspecies ofP. required before P. macrotis canbe said to no longer oc macrotis, although this was done without comments on cur on Grenada. their reasons. Koopman (1994) uses the name P. m. phaea for the population on Grenada. Although it is difficult to evaluate the gender ratio ofbats in research collections (it is impossible to know Comparing the measurements ofour specimens the selective actions, ifany, ofthe collector), it is inter from Grenada (Table I) with those from other regions esting to notice that the ratio offemales to males in our where similar data are available, it is clear that the two largest samples were 12: 12 at Mt. Pleasant Estate Grenadan material most closely resembles the sample and 8:15 at Point Salines. G. M. Allen (1911) consid from Trinidad (Dalquest, 1951; Goodwin and Greenhall, ered seven ofthe bats he collected at Point Salines on 1961; Carter et al., 1981). Variation in the sample from 29 August 1910 to not be adults. No other reproductive Grenada encompasses the variation in the Trinidad sample data were recorded for any ofthe specimens examined. for length offorearm, greatest length ofskull, zygomatic breadth, and length ofthe maxillary toothrow, whereas G. M. Allen (1911) described P. m. phaea based variation in the Trinidad samples encompasses the varia upon the material from Point Saline [= Point Salines]. tion in the Grenada sample for breadth ofpostorbital He distinguished the Grenada populations from those on constriction, which is the only other measurement avail the SouthAmerican mainland basedprimarily upon color, able. The sample from Tobago is the most morphometri which was nearly uniform Prout's brown above and be cally distinct of those compared with the Grenadan low as opposed to having a reddish cast. From the taxon sample. There is no overlap in the measurements ofthe trinitatus from Trinidad, phaea was distinguished by its these two samples in length offorearm, greatest length longer length offorearm, being 42.5 (41-44.5) as op ofskull, and length ofthe maxillarytoothrow. The sample posed to 40.0 (39.0-41.0). Sanborn (1937) believed that from Grenada fills the gap in the range measurements the population on Grenada averaged smaller than main between the two samples from French Guiana that are land populations, but he was not able to characterize supposed to represent the species Peropteryx macrotis trinitatus with the material that he had at hand. Goodwin and P. trinitatus. Infour measurements the sample from and Greenhall (1961) assigned specimens from Tobago Grenada overlaps the lower end ofthe range ofvariation to the mainland subspeciesP. m. macrotis, while restrict in the sample ofP. macrotis, including length offorearm ing the taxon P. m. trinitatus to Trinidad. They believed (overlap from 43.0 to 43.2), greatest length ofskull (14.0 that the larger external and cranial size ofthe Tobago to 14.1), zygomatic breadth (7.9 to 8.2), and length of material supported this decision, with mean length of maxillary toothrow (5.3 to 5.5). The two specimens of forearm ofTobago specimens (three females) being 46.5 P. trinitatus from French Guiana fall at the lower end of (46.0-47.2) as opposed to 42.1 (41.0-43.0) for speci the range ofvariation ofthe Grenada sample in length of mens from Trinidad (one male and three females), great forearm, greatest length ofskull, and zygomatic breadth, est length ofskull 14.8 (14.7-15.0) versus 13.6 (13.2 whereas for breadth ofthe postorbital constriction the 13.9), and length ofmaxillary toothrow 5.9 (5.8-6.0) specimens from French Guiana are at the upper end of versus 5.4 (5.3-5.5). Handley (1976) and Brosset and the range ofvariation and in length ofmaxillary toothrow Charles-Dominique (1990) more recently have treated at 4.7 and 4.8, these are the two smallest specimens that P. macrotis and P. trinitatus as distinct species. Handley have been considered in this study. 4 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY
It is difficult to draw any meaningful taxonomic north ofGrenada. This was the fIrst record ofPamell's conclusions from these data. Morphologically, the mustachedbatfrom the LesserAntilles and indicates that sample from Grenada is clearly most similar to the ma this larger relative ofDavy's naked-backed bat can be terial from Trinidadreportedby Goodwinand Greenhall expectedonGrenada. These species canbedistinguished (1961), but this leaves the perplexing specimens from bythe much larger size ofP. parnellii and that the wing Tobago. Italso is clearthat to gobeyondthis point will membranes ofP. davyi fuse along the mid-dorsal line, require a comprehensive analysis of the Peropteryx whereas the wingmembranes ofP.pamelliiattach lower macrotis complex in northeastern South America to de on the sides so that the mid-dorsal area is covered by a termine iftwo species are present and, ifthere are two band ofhair. P. d. davyi was described based upon a species, to thoroughly document their morphological specimenfrom Trinidad. The measurements ofoursingle differences. We would not be surprised to ultimately specimen (fable 1) fit withinthe range ofmeasurements fmd that material from Grenada is best assigned to the ofsamples from Trinidad and Dominica (Smith, 1972). same species and subspecies ofPeropteryx as is found on Trinidad. The taxonphaea was poorly distinguished The specimen from Grenada was anadultmalepre byG. M. Allen (1911), whobased its distinctiveness pri served in fluid. The skull has been removed from the marily onpelage color, which is known to be polytypic. individual allowingmeasurementsto betaken. The speci menwas obtained in 1894 from an unspecified location Itis worthy ofnote thatPeropteryx is the only rep on GrenadabyN. H. Smith (Jones, 1951). resentative ofthe family Emballonuridae known to have reached the Antilles (Baker and Genoways, 1978). Al though the family Emballonuridae is believed to be one Noctilio leporinus mastivus ofthe oldest ofextant chiropteran families, Peropteryx (Vahl,1797) probably is a recent invader based upon its weakly de GreaterFishing Bat fined morphological distinctiveness, reaching the island from the south from eitherTrinidad orthe South Ameri Specimens examined (11).- ST. GEORGE: can mainland. Webelieve thatthis invasionhas occurred Chemin River, 1/2 km E Confer 12°01' N, 61 °43' W, 6 as over water dispersal. It is not understood why (5 CM, 1 UNSM); 1/2 km E Vendome, 12°05' N, emballonurids are not more extensively distributed in 61°42.5'W, 1(UNSM). PARISH UNKNOWN: E of the Antilles. Bats of similar size and habits, such as Crawford, sea level (BMNH); no specific locality, 3 (2 Natalus andMyotis, are widely distributed inthe islands. NMNH, 1MCZ).
Remarks.-Large fish-eating bats ofthe species Pteronotus davyi davyi Noctilio leporinus are common inhabitants ofthe islands Gray, 1838 ofthe Caribbean, and Grenada is no exception. These Davy's Naked-backed Bat bats often are seen fishing over quiet bays and lagoons along the coast, and overpools and slow-moving areas Specimen examined(1 ).-PARlSH UNKNOWN: offreshwater rivers. The species commonly roosts in no specific locality, 1 (BMNH). small caves andhollow trees nearthese areas. Six ofour specimens were taken in mist nets set over the Chemin Remarks.-The single specimenofDavy'snaked River at a pointthat was no more than a kilometer from backed bat from Grenadawas assigned to the nominate where it entered the sea. All were taken as they were subspecies by Smith (1972). He also reported this spe hunting for small fish and invertebrates in the river. Most cies from the Antillean islands ofMartinique, Dominica, of the area adjacent to the river was open sugarcane and Marie Galante. Masson etal. (1990) have reaffirmed fields, but several large trees on both sides ofthe river that P. davyi does occur as far north as Marie Galante; formed a canopy in some areas. The specimen from however, more recently Vaughan(1995) andVaughan and Vendome was taken in another habitat type at an eleva Hill (1996) have reported the related species Pteronotus tion ofabout 300 m. Nets were set over a small stream parnellii from St. Vincent, the first major island to the as it emerged from Grand Etang Forest Reserve. The Table i.-Length offorearm and cranial measurements ofi2 species ofbats from Grenada, West Indies. All measurements are ~ in millimeters. Z 0
Sex/Cat. no. Length of Greatest Condylobasal Zygomatic Postorbital Mastoid Length of Breadth ~ and Statistics foreann length length breadth breadth breadth maxillary across 00 ~ ofskull toothrow upper molars ~ ~ PEROPTERYXMACROTIS PHAEA 0, Males = N 6 6 6 6 6 6 6 6 ~> Mean 42.6 13.7 12.3 8.0 2.6 7.1 5.3 6.0 00 SE ±0.32 ±0.15 ±0.15 ±0.05 ±0.03 ±0.04 ±0.05 ±0.09 0 ~ Minimum 41.3 13.2 11.8 7.8 2.5 6.9 5.1 5.6 ~ Maximum 43.2 14.1 12.7 8.2 2.7 7.2 5.4 6.2 ~ Females Z N 7 7 7 6 7 7 7 7 ~ Mean 41.6 13.8 12.3 7.9 2.5 7.1 5.3 5.9 > SE ±0.61 ±0.09 ±0.13 ±0.12 ±0.04 ±0.05 ±0.06 ±0.11 Minimum 39.6 13.4 11.8 7.5 2.4 6.9 5.0 5.4 Maximum 43.1 14.1 12.7 8.2 2.6 7.2 5.5 6.3
PTERONOTUS DAVYIDAVYI
Male 94.9.7.1 BMNH 44.8 16.2 14.9 8.8 3.7 8.8 6.7 6.0
NOCTILIO LEPORINUS MASTIVUS
Females N 4 5 5 5 5 5 5 5 Mean 84.9 26.4 23.6 18.4 6.7 16.6 10.1 12.1 SE ±1.30 ±0.21 ±0.26 ±0.30 ±0.11 ±0.17 ±0.04 ±0.28 Minimum 82.0 25.8 22.8 17.4 6.5 16.2 10.0 11.1 Maximum 88.1 26.9 24.2 19.2 7.1 17.1 10.2 12.6
Ul Table 1.- (continued.) ~
Sex/Cat. no. Length of Greatest Condylobasal Zygomatic Postorbital Mastoid Length of Breadth and Statistics foreann length length breadth breadth breadth maxillary across ofskull toothrow upper molars
GLOSSOPHAGA LONGIROSTRIS ROSTRATA
Males 0 N 6 6 6 6 6 6 5 6 ~ Mean 37.6 22.8 21.4 10.0 4.6 9.4 8.0 5.7 ~ SE ±0.13 ±O.II ±0.12 ±O.IO ±0.06 ±0.08 ±0.06 ±0.07 >rn Minimum 37.2 22.4 21.1 9.6 4.5 9.2 7.8 5.5 .... Maximum 38.1 23.2 21.8 10.2 4.8 9.7 8.1 5.9 ~ ~ Females lo'l:l N 6 6 6 6 6 6 5 5 i!; Mean 37.9 23.1 21.5 9.8 4.7 9.4 8.0 5.9 ~ SE ±0.43 ±0.15 ±0.16 ±0.08 ±0.06 ±0.05 ±0.14 ±0.07 ~ Minimum 36.0 22.7 20.9 9.5 4.6 9.3 7.7 5.7 .. Maximum 39.0 23.7 22.0 10.1 4.9 9.6 8.3 6.1 arn ~ ANOURA GEOFFROYI GEOFFROYI ~ Males ~ N 10 10 10 10 10 10 10 Io"l Mean 42.0 25.0 11.0 5.0 10.5 9.4 6.2 ~ SE ±0.22 ±0.15 ±0.07 ±0.03 ±0.05 ±0.05 ±0.06 ~ Minimum 40.6 24.4 10.5 4.8 10.2 9.1 5.9 rn Maximum 43.2 25.6 11.2 5.1 10.8 9.5 6.5 Io"l ~ ~ Females N 10 10 10 10 10 8 10 = Mean 42.9 25.2 10.9 5.0 10.4 9.5 6.2 SE ±0.28 ±0.06 ±0.07 ±0.03 ±0.08 ±0.04 ±0.05 ~ Minimum 41.2 24.9 10.6 4.9 10.1 9.3 5.9 ~ Maximum 44.3 25.5 11.3 5.2 10.9 9.6 6.4 .... -
Female 14599MCZ 39.8 21.0 18.8 11.7 5.0 10.9 6.7 8.7
ARTIBEUSJAMAICENSIS GRENADENSIS
Males N 10 10 9 10 10 10 10 10 Mean 58.0 28.2 25.3 17.5 7.1 15.0 10.1 12.8 SE ±0.62 ±0.19 ±0.17 ±0.14 ±0.07 ±0.11 ±0.08 ±0.11 Minimum 55.7 27.5 24.9 16.6 6.6 14.4 9.8 12.2 Maximum 61.8 29.4 26.0 18.0 7.3 15.5 10.6 13.3
Females N 32 32 32 32 32 32 32 32 Mean 58.1 28.2 25.0 17.2 6.9 15.0 10.0 12.6 SE ±0.27 ±0.10 ±0.09 ±0.08 ±0.03 ±0.08 ±0.05 ±0.06 Minimum 55.7 26.8 23.8 16.5 6.7 13.7 9.4 11.8 Maximum 61.3 29.2 25.9 18.1 7.4 15.9 10.3 13.2 ~ Table J.- (continued.) 00
Sex/Cat. no. Length of Greatest Condylobasal Zygomatic Postorbital Mastoid Length of Breadth and Statistics forearm length length breadth breadth breadth maxillary across ofskull toothrow upper molars
ARTIBEUSLITURATUSPALMARUM
Males 0 N 8 8 8 8 8 8 8 8 n Mean 70.1 30.9 27.1 18.7 6.5 16.6 10.7 13.6 n SE ±0.72 ±0.23 ±0.16 ±0.17 ±0.08 ±0.13 ±0.09 ±0.12 >00 Io-ol Minimum 67.7 29.8 26.5 17.6 6.2 16.1 10.4 13.0 0 Maximum 72.4 31.5 27.8 19.0 6.7 17.2 11.2 14.1 Z ~ Females "'d N 7 7 7 7 7 7 7 7 Mean 72.0 30.8 27.0 18.5 6.5 16.4 10.7 13.4 ~ t'!:l SE ±0.81 ±0.32 ±0.28 ±0.38 ±0.09 ±0.21 ±0.12 ±0.24 Minimum 69.0 29.5 25.9 16.9 6.2 15.3 10.2 12.7 ~ Maximum 75.7 31.6 27.9 19.6 6.7 16.9 11.1 14.2
a00 t'!:l STURNlRA LILIUMSEROTINUS a:~ Males 0 ~ 63412 CM 41.9 23.0 20.4 13.2 5.5 11.6 6.8 7.7 l-3 63413 CM 42.4 23.3 20.7 13.3 5.8 11.7 6.8 8.0 t'!:l 16493 UNSM 42.5 23.1 20.7 13.0 5.7 11.6 6.8 7.7 ~ 00 Females l-3 63414 CM 42.6 22.5 20.0 13.0 5.7 11.5 6.7 7.7 t'!:l 83426CM 42.5 22.6 19.9 12.9 5.6 11.2 6.6 7.7 n 16494 UNSM 40.1 21.8 19.2 12.8 5.6 11.1 6.7 7.8 =~
Sex/Cat. no. Length of Greatest Condylobasal Zygomatic Postorbital Mastoid Length of Breadth and Statistics forearm length length breadth breadth breadth maxillary across ~ ofskull toothrow upper molars tI.l ~ l-3 MYOTIS NIGRICANS NIGRICANS ~ 0, Males 83427 CM 33.4 13.5 12.4 8.3 3.4 6.8 4.9 5.2 252600NMNH 32.8 12.8 12.1 8.2 3.5 6.7 4.7 5.1 ~ tI.l 0 ~ MOLOSSUS MOLOSSUS MOLOSSUS Males ~ 63415 CM 39.5 17.5 15.3 10.8 3.6 10.0 6.1 7.8 63423 CM 38.7 16.7 14.6 10.5 3.7 10.0 5.8 7.6 ~ 63424 CM 39.0 16.8 14.9 10.6 3.6 10.0 5.8 7.4 > Females 63421 CM 37.4 16.0 14.0 10.0 3.4 9.5 5.5 7.0 63422 CM 37.5 16.1 14.2 10.0 3.6 9.5 5.7 7.3 63428 CM 38.2 16.2 14.2 10.0 3.5 9.3 5.7 7.2 \C 10 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY stream was one of the small upper tributaries of the that our specimen could represent a Grenadan popula Beausejour River, which enters the Caribbean Sea ap tion Vaughan (1995) and Vaughan and Hill(1996) found proximately 5kIn to the west-northwest. Habitat at this M mega/otis roosting in coastal caves and feeding un site was characterized by wet, dense montane forest, der rain forest canopy and along side ofbanana planta which became fairly open underthe canopy away from tions near the rain forest. We netted many nights on disturbed areas. To the south ofthe edge ofthe Grand Grenada in areas similar to where this species was found Etang Forest Reserve, much ofthe native vegetation was feeding on St. Vincent without taking any specimens of cleared for cocoa, bananas, mangos, and small garden this species, so if M mega/otis presently occurs on plots, which were situated behind several homes that Grenada, itmust be in relatively low numbers. faced the nearby road. The stream was about 4 m wide and 30 cm deep. It ran fairly straight with ahigh canopy Many recent authors (Handley, 1976; Brosset and for nearly 50 m before dropping about 8 m over a con Charles-Dominique, 1990; Simmons, 1996) have treated crete spillway. M mega/otis andM microtis as separate species in con trast to the arrangement ofSanborn (1949) and Koopman Five females that were captured along the Chemin (1993, 1994) where microtis was placed as a subspecies River were lactating on 23 May 1980 (4) and 28 May ofmega/otis. We are following the arrangement ofrec 1987. A sixth female from this location evinced no re ognizing two species with the limited material at hand, productive activity on 23 May 1980. No reproductive which results in M mega/otis being considered a mono data were recorded for three male specimens that we typic species. examined (BMNH, MCZ, NMNH). The specimen of M. mega/otis in the National We have followed Davis (1973) in assigning our Museum ofNatural History is an adult female with a specimens to the subspecies N. /. mastivus, with a type length offorearm of 34.2. The specimen is stored in locality ofSt. Croix, U. S. Virgin Islands. Specimens alcohol and the skull has not been extracted for study. from throughout the Caribbean islands, Mexico, Central The specimen has large ears characteristic ofthis spe America, and northern South America were placed by cies, but the membranes and pelage are so faded that no Davis (1973) into this subspecies. The five females in color determination is possible. The specimen does dis our sample (Table 1) average slightly smaller in the five play the band between ears characteristic ofthe subge measurements reported by Davis (1973) than a sample nus Micronycteris and, as Sanborn (1949) described it, from Trinidad and a pooled sample from throughout the the band is relatively low at the base ofthe ears rising to West Indies. These average differences never exceed a triangle at the center. The relatively small notch in the 0.6 mm and we believe are not oftaxonomic significance. band is located at the apex ofthe triangle. Micronycteris megalotis Glossophaga longirostris rostrata (Gray, 1842) Miller, 1913 Brazilian Large-eared Bat Miller's Long-tongued Bat Specimen examined (1).-PARISH UNKNOWN: Specimens examined (231).- ST. ANDREW: no specific locality, 1 (NMNH). Birch Grove, 19 (UNSM); Grand Etang, 4 (MCZ); Grenville,S (NMNH); Grenville Vale [near Granton], 6 Remarks.-Because the Brazilian large-eared bat (5 AMNH, 1NMNH); 0.4 kIn S, 0.4 kIn W S1. Margaret, is represented from Grenada by a single, long-preserved 1600ft, 12°06'N,61°41.5'W,3 (CM). ST. DAVID: 1 museum specimen, we have questioned, as did Koopman 1/4 kIn S, 3/4 kIn W Beaton, 12°01.5' N, 61 °42' W, 2 (1958), whether it represented an existing population or (CM); WesterhallEstate 13 (NMNH). ST. GEORGE: a single accidental record. However, with report ofa Chemin River, 1/2 kIn E Confer, 12°01'N, 61°43' W,30 population ofthis species on St. Vincent by Vaughan (22 CM, 8 UNSM); Fort Frederick, 2 (KU); French (1995) and Vaughan and Hill (1997), it seems possible Ammunition Tunnels, 6(KU); Point Salines, 3(AMNH); GENOWAYSETAL.--BATSOFGRENADA 11 3 mi N, 1/2 mi E St. George's, 4 (KU); 1/2 mi NE St. amined); 24 May (3); 30 May (3); 16 August (2); 17 George's,2 (1 KU, 1TIU); St. George's, 9 (MCZ); St. August (1). A female netted on24 May at St. Paul's was Paul's, 12°04' N, 61 °44' W, 15 (11 CM, 4 TIU); Salt lactating and carried a non-volantjuvenile male (length Pond, 1(KU); True Blue 4 (3 AMNH, 1NMNH); 1/2 Ian offorearm, 26.7), whereas females taken on 26 May and E Vendome, 12° 05' N, 61°42.5' W, 53 (32 CM, 15 17 August were lactating, but did not carry young. Vol UNSM, 6 TIU); 1/2 mi E Vendome, 1000 ft, 9 (KU). ant subadult (as indicated by unfused wing phalangeal ST. JOHN: 3/4 km S, 1/2 Ian W Concord, 12°05.5'N, epiphyses) individuals were taken onthe following dates: 61 °44.5' W, 13 (11 CM, 2TTU). PARISH UNKNOWN: 24 May (2 males and 1 female, length offorearm, 37.2, no specific locality, 28 (NMNH). 38.4, 38.0); 25 May (female, 35.6); 26 May (1 female and 2 males, 38.8, 36.7, 36.9); 28 May (female, 38.8). Remarks.-Miller's long-tongued bat was one of Inasample of27 individuals preservedbyA. M. Greenhall the two most common and widespread species taken dur between 11 and 13 August 1955, 13 individuals were ing oursurveys on Grenada. It is known from the driest adults while 14 were subadults. The following testes areas ofthe island around Point Salines and True Blue to lengths were recorded from adult males: 25 May, 5, 7; the wettest areas that we visited at Vendome and Birch 26 May, 2, 4, 4, 5, 5, 5, 5; 27 May, 2,3.5,3.5,4,4,4.5; Grove. The rain forest at the site we visited nearVendome 28 May, 3,4,4; 29 May, 2; 16August,2.5,3,5; 17 Au was relatively undistributed, but at Beaton and Chemin gust, 4, 4. Although this species appears to breed during River the original vegetation is highly disturbedby agri the rainy season, there are still not enough data available cultural activities. The sample ofMiller's long-tongued to describe the annual reproductive cycle (Wilson, bats from near Concord were netted in a banana planta 1979). Our reproductive data do agree with those pub tion with a few ofthe old tall forest trees providing the lished previously for this species from Trinidad overstory. Specimens from near Concord, as well as many (Goodwin, 1958; Goodwin and Greenhall, 1961) and others that we captured, had pollen covering their heads Margarita Island, Venezuela (Smith andGenoways, 1974). and shoulders. The yellow, powdery pollen was clearly visible in our lights at night. Table 1gives the length offorearm and eight cra nial measurements for samples ofsix males and six fe Allen (1911) commented on the capture of the males from Grenada. Using a t-test, we found no evi specimens from St. George's and Grand Etang that are dence ofsignificant secondary sexual variation between housed in the Museum ofComparative Zoology. Speci these samples. mens from St. George's were obtained "from holes, in the old fort on Richmond Hill." At Grand Etang (1800 ft The taxon rostrata originally was described by =545 m) specimens were observed "hanging to the ridge Miller (1913a) as a distinct species based on material pole in a room ofa disused stable, whence they obtained from Westerhall Estate on Grenada, but quickly was egress by means ofa partially open window." The speci moved by Miller (1913c) to a subspecies oflongirostris. mens from Westerhall Estate were "taken in hollow tree" This taxonomic arrangement was maintained by the most in 1900 by PeterGellineau (Miller, 1913a; Handley and recent reviewers ofthe species (Webster and Handley, Webster, 1987). 1986). Glossophaga I. rostrata was distinguished by Webster and Handley (1986) from other subspecies by Females containing embryos, with crown-rump its narrow rostrum and reduced postorbital swellings. The length ofembryos indicated, were taken on the follow subspecies rostrata currently is know to occur on ing dates (number offemales from date with data in pa Grenada, Trinidad, Tobago, Grenadines, and St. Vincent. rentheses): 26 May, 5, 5, 7 (total of 6 adult females Two specimens previously reported by Miller (1913a, preserved); 27 May, 6 (2); 28 May, 5, 5 (2); 29 May, 5, 5 1913c) from Dominica were shown by Handley and (4); 2 June, 9 (1); 18 July, no c-r (1); 19 July, no c-r for Webster (1987) to be mislabeled individuals from the 2 (2). Females evincing no reproductive activity were original type series from Grenada. taken on the following additional dates: 23 May (6 ex- 12 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY Anourageo//roy;geo//roy; All ofour specimens were adults. None ofthe Gray, 1838 females taken onthe following dates evinced any repro Geoffroy's Tailless Bat ductive activity: 24 May (7 females); 25 May (3); 26 May (9); 27 May (1); 30 May (4); 1 June (2); 19 July Specimens examined (120).- ST. ANDREW: (10); 16 August (2); 17 August (8). These observations Birch Grove, 27 (UNSM); 0.4 Ian S, 0.4 IanW St. Mar would be in line with those made by Goodwin and garet, 1600 ft, 12°06' N, 61°41.5'W, 8 (CM). ST. Greenhall (1961) on Trinidad where they found gravid GEORGE: 3 mi N, 1/2 mi E St. George's, 2 (KU); 1/2 females with fetuses inadvanced stages ofdevelopment kmNE St. George's, 3 (KU); St. Paul's, 12°04'N, 61 °44' on20 November, butfetuses were not found in females W, 14 (CM); 1/2 IanEVendome, 12°05'N,61°42.5'W, taken on 12 June or 17 October. Wilson (1979) com 39 (34 CM, 5 UNSM); 1/2 mi E Vendome, 1000 ft, 27 mented on the rather unusual discrete late rainy season (KU). breedingperiodofAnourageoffroyi. The following tes tes length were recorded for this species on Grenada: Remarks.- Geoffroy's tailless bat first was re 25 May, 6, 6, 7; 26 May, 2, 5, 8; 30 May 6, 7; 1 June, 7; ported from GrenadabyJones and Phillips (1970) based 16 August, 6; 17 August, 6, 6, 6, 7, 7, 7, 7, 7, 7, 7, 8. on the specimens listed above from the University of Kansas. Our work has shown that this species is more Selected samples of males and females from widespread onthe island than the original records indi Grenada were tested for secondary sexual variation us catedwith Geoffroy's tailless batoccurring inthe more ing t-tests (see Table 1). No secondary sexual variation mesic areas on the island at locations that are above 250 was found in six cranial measurements, but the length of m. This elevationallimitation was shown in our work forearm offemaleswas significantly (P< .05) longerthan when Anoura was captured at the location 3 mi north that ofthe males. and 1/2 mi east ofSt. George's at an elevation that is about 250 to 300 m, whereas less than 3 km to the north Jones and Phillips (1970) made the following state ofthis point at our locationjustsouth ofConcord at about ment concerning the systematic relationships of this 60 m no specimens ofthis species were taken, but the newly discoveredpopulation to the knownpopulation on related species, Glossophaga longirostris, was ex Trinidad: "Preliminary study indicates that Grenadan tremelyabundant. We worked several sites along the Anoura average slightly larger than specimens from southern and eastern coast and feel confident that Trinidad and have a somewhat more inflatedbraincase, Geoffroy's tailless bat does not occur in these low, dry but they do not differ otherwise and probably represent areas dominated by thorn scrub, cacti, and agricultural the same subspecies (A. g. geofJroyi Gray)." Jones fields. Goodwin and Greenhall (1961) believed that this (1989) later confirmed this preliminary subspecific de species strictly roosted in caves on Trinidad. We can termination (see also Koopman, 1994). We tested a not confirm this observation on Grenada because all of sample (8 males, 2 females in collections ofMuseum our specimens were netted away from caves as they fed, ofTexas TechUniversity) from Trinidad against ourcom particularly nearplanted orchards and bananaplantations. bined sample from Grenada (Table 1) using t-tests to We took the series of14 individuals at St. Paul's in just determine ifany significant geographic variation was such a situation on the evening of24 May 1980. We present. None ofthe seven measurement tested revealed placed our nets in the open grassy areas that had been any significant geographic variation between these two cleared ofnative plants for fruit trees, which included island populations. We, therefore, follow the earlier bananas, mangos, papaya, guava, andnutmeg. There was authors in assigning the Grenadan population to same little native vegetation left within several hundred meters subspecies as on Trinidad, which is currently considered ofthe area being netted, but a number ofhomes were to be Anoura geoffroyi geoffroyi. scattered among the orchard and along the nearby road. Only Anoura andArtibeusjamaicensis andA.lituratus were taken during the evening. GENOWAYS ETAL.-BATS OF GRENADA 13 Carolliaperspicillataperspicillata Artibeus glaucus bogotensis (Linnaeus, 1758) Andersen, 1906 Seba's Short-tailedBat LesserGray Fruit-eatingBat Specimen examined(1).-PARlSHUNKNOWN: Specimens examined(5).-ST. ANDREW: Grand no specific locality, 1 (BMNH). Etang, 1 (MCZ). ST. GEORGE: 8 mi N St. George's, 1 (KU); 1/2 km E Vendome, 12°05' N, 61°42.5 W, 3 (2 Remarks.-Carolliaperspicillata is represented CM,1 UNSM). from Grenada by a single specimen ofan adult female deposited in the British Museum (Natural History). Remarks.- The lesser gray fruit-eating bat has Based upon the catalogue number of the specimen been reported previously from Grenada based upon a (67.5.4.4), it would appear to have been registered into single specimen from Grand Etang (Koopman, 1958). the collection in 1867. lbis specimen first was reported lbis female bat was captured in November 1912. The byDobson (1878) underthe name Carollia brevicauda. species was not taken on the island again until 2 May He indicatedthatthe specimenhadbeenacquired through 1980 when we netted two males (length oftestes, 2, 5) purchase. Subsequent authors (Allen, 1911; Pine, 1972) at the edge ofthe Grand Etang Forest Reserve. Subse have assigned this specimen to the species C. quently, we captured another male (length oftestes, 4) perspicillata and we agree with this assignment. Exam on 19 July 1989 at the exactplace where the specimens ining the characters used by Pine (1972) to distinguish were captured in 1980. Therefore, all captures ofthis species within the genus, this specimen possesses the exceedingly rare species on Grenada have come from following characteristics in agreement with C. undisturbed montane forests at elevations above 1000 perspicillata: size relatively large (see Table 1); the lower feet [=300 m]. We hypothesize that the continued exist outer incisors, when viewed from above, obscured by ence ofthis species on Grenada is dependent upon the canines; maxillary toothrow straight, not bowed orwith continued existence and health ofthese montane forests. a step; short haircovering the forearm; hair on mid-back about6mm. Koopman (1958, 1994) and subsequent authors have reported this species under the name Artibeus Koopman (1989) omitted Carollia from his analy cinereus. Owen (1987) resurrected the generic name sis ofthe bats from the Antilles and Jones (1989) con Dermanura for most ofthe smaller representatives of sidered the status ofCarollia on Grenada as question the genus Artibeus and treated this species as Dermanura able. We here considerthat the specimen from Grenada cinerea. We have not followed his arrangement prefer represents either an erroneous record or an accidental ring instead to follow Koopman (1993, 1994), Lim occurrence on Grenada. With a very old record based (1993), Lim and Wilson (1993), Van Den Bussche et al. upon a purchased specimen, we feel that it is very likely (1993, 1998), and Pumo et al. (1996) in retaining the that the locality data for this specimen are incorrect, but generic name Artibeus for these bats. Handley (1987) because we can not absolutely rule out this being an ac in describing a new species from Venezuela rearranged cidental record from Grenada, we have retained the spe the small Artibeus into 10 species and six species groups. cies in this compilation ofbats from the island. It is He restricted Artibeus cinereus to the Guiana region, actually of some considerable interest that Carollia coastal Brazil, and lower Amazon Basin and applied the perspicillata is not found in the Antilles. The species is name Artibeus glaucus to the northern South American abundant andwidespread onTrinidad, Tobago, and Little form. The name Artibeus glaucus bogotensis appears Tobago (Goodwin and Greenhall, 1961). Seba's short to be the most appropriate trinomial to apply to popula tailed batwould appearto be successful in habitats dis tions on Trinidad and Tobago formerly designated as turbedby human orweatheractivities; therefore, the ap Artibeus cinereus cinereus (Handley, 1987; Koopman, propriate habitats would appear to be available on 1993), although Koopman (1994) has usedA. cinereus Grenada. bogotensis. 14 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY Because the mostobvious route ofinvasion ofthis 46 (MCZ); St. Paul's, 12°04'N, 61°44'W, 14 (12 CM, 2 species into the Antilles was from the south (Koopman, TTU); 1/2 mi E Vendome, 1000 ft, 8 (KU); 1/2 km E 1968; Jones and Phillips, 1970; Jones, 1989), we have Vendome, 12°05'N, 61 °44.5' W, 76 (38 CM, 36UNSM, compared our specimens with a sample from Trinidad 2 TTU). ST. JOHN: 3/4 km S, 1/2 km W Concord, (five males and five females from Las Cuevas in The 12°05.5'N, 61°44.5' W, 11 (9 CM, 2 TTU). PARISH Museum ofTexas Tech University) to determine their UNKNOWN: no specific locality, 51 (48 NMNH, 2 subspecific status. Measurements ofthe four specimens MCZ, 1 BMNH). from Grenada are presented in Table 1. Because the sample from Grenada was too small to test for second Additionalrecord.-Douglaston (Jones, 1951). ary sexual variation, the Trinidad sample was tested to determine ifthe sexes couldbe combined for an analy Remarks.- The Jamaican fruit bat, along with sis ofgeographic variation. None ofthe eightmeasure Miller's long-tonguedbat, are the two mostcommon and ments tested revealed any significant (usingt-test) sec widespread species ofbat on Grenada. It occurs in all ondary sexual variation so the sexes were combined in habitats and atall elevations with the possible exception the next analysis. We compared the measurements of ofthe driest areas in the southwestern corner ofthe is the four specimens from Grenadawith the sample of10 land around Point Salines. Allen (1911) reported a breed specimens from Trinidad using a t-test. The only sig ing colony ofJamaican fruit bats found in the recesses nificant (P< 0.5) difference was for zygomatic breadth ofthe old fort on Richmond Hill in St. George's. Ofthe the sample from Grenada averaged 11.7, whereas the 30 adults in his sample, 26 were females. AtDouglaston Trinidad sample averaged 12.3. Means for the two a large colony was found inthe roofofa distillery (Jones, samples (Grenada followed byTrinidad) for the remain 1951). ingsevenmeasurements were as follows: length offore arm, 39.4, 40.6; greatest length of skull, 20.9, 20.8; Atleast since the workofHershkovitz(1949), the condylobasallength, 18.8, 18.7; postorbital constriction, subspecies A. j. grenadensis Andersen has been consid 5.0,5.0; mastoid breadth, 10.7, 10.7; length ofmaxillary ered by most authors to be a junior synonym ofA. j. toothrow, 6.7, 6.7; breadth across uppermolars, 8.7, 8.8. trinitatis witha type locality onTrinidad. However, we Clearly, the specimens from Grenada are morphologi believe that we have developed bothmorphological and cally close to those from Trinidad and shouldbe assigned genetic data that support the recognition ofpopulation to the same subspecies, which currently is believed to on Grenada as a distinct subspecies to which the name A. be bogotensis. All three specimens that we captured j. grenadensis would apply. Table 2 presents the results during our work also exhibited the dental character of of t-tests comparing samples of A. j. trinitatis from only two rather than three lower molars that Handley Trinidad andA.j. schwartzi from St. Vincent with the (1987) believed was typical ofA. g. bogotensis. population on Grenada. The sample ofgrenadensis is significantly smaller than schwartzi in length of fore arm and the seven cranial measurements at the P <0.001 Artibeusjamaicensis grenadens;s level. Infact the range ofmeasurements ofthe two taxa Andersen, 1906 only overlap in three measurements-breadthofpostor Jamaican Fruit-eating Bat bital constriction, mastoid breadth, length ofmaxillary toothrow. The sample ofgrenadensis averaged signifi Specimens examined (330).- ST. ANDREW: cantly larger at the P <0.001 level than the sample of Birch Grove, 20 (UNSM); 0.4 km S, 0.4 km W St. Mar trinitatis for length offorearm, greatest length ofskull, garet, 1600 ft, 12°06' N, 61°41.5' W, 2 (CM). ST. condylobasallength, and zygomatic breadth and at the P DAVID: 1 1/4 km S, 3/4 km W Beaton, 12°01.5' N, <0.05 level for breadth ofpostorbital constriction and 61 °42' W, 11 (CM); Westerhall Estate, 18 (NMNH). ST. breadth across upper molars. Means ofthe two samples GEORGE: Chemin River, 1/2 km E Confer, 12°01'N, were notsignificantly different for mastoid breadth and 61°43'W, 45 (25 CM, 20 UNSM); Fort Frederick, 8 length of maxillary toothrow. Therefore, A. j. (KU); Grenville Vale [near Grafton], 1 (NMNH); 8 mi grenadensis is morphologically distinct from the geo NE St. George's, 1 (KU); 3 mi N, 1/2 mi E St. George's, graphically nearest taxa, although it would appear to be 9 (KU); 1/2 mi NE St. George's, 9 (KU); St. George's, slightly closer to trinitatis morphologically. GENOWAYS ETAL.-BATS OF GRENADA 15 Table 2.-Morphometric comparisons ofone external and seven cranial measure- ments ofpopulations ofArtibeus jamaicensisfrom Grenada, St. Vincent, and Trinidad. ns = not significant,' *= P Measurements and Trinidld Grenada StVincent Statistics Significance Significance Length offoreann Number 16 42 8 Mean 55.8 ••• 58.1 ••• 63.6 Range 53.6-57.9 55.6-61.8 62.1-64.6 SE ±O.30 ±O.25 ±O.29 Greatest length ofskull Number 16 42 8 Mean 27.6 ••• 28.2 ••• 30.9 Range 26.8-28.7 26.8-29.4 30.1-31.8 SE ±O.l4 ±O.09 ±O.21 Condylobasallength Number 16 41 8 Mean 24,5 ••• 25.1 ••• 27,2 Range 23.7-25.1 23.8-26.0 26.5-28.0 SE ±O.11 ±O.09 ±O.24 Zygomatic breadth Number 15 42 8 Mean 16.8 ••• 17.3 ••• 19.3 Range 16.3-17.3 15.8-18.0 18.6-20.1 SE ±O,08 ±O.07 ±O.22 Breadth ofpostorbital constriction Number 16 42 8 Mean 6.9 • 7.0 ••• 7.4 Range 6.7-7.1 6.6-7.4 6.9-7.6 SE ±O.05 ±O.03 ±O.08 Mastoid breadth Number 16 42 8 Mean 15.1 ns 15.0 ••• 16.3 Range 14.5-16.4 13.7-15,9 15.7-16,7 SE ±O,I2 ±O.07 ±O.13 Length ofmaxillary toothrow Number 16 42 8 Mean 10,0 ns 10.0 ••• 10.8 Range 9.3-10.6 9.5-10.4 10.4-11.4 SE ±O.06 ±O.04 ±O.12 Breadth across upper molars Number 16 42 8 Mean 12.4 • 12.7 ••• 14.1 Range 11.6-13.3 11.8-13.2 13.5-14.7 ±O,I6 SE ±O.09 ±O.05 16 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY Artibeus}amaicensis is the only Grenadanbatspe Comparing the means ofmeasurements ofmale cies for which we have genetic data. These data are in and femaleA.}amaicensis from Grenada (Table 1) using the fonn ofmitochondrial DNA (mtDNA) haplotypes t-tests, we found that there were no secondary sexual based on restriction endonuclease analyses (phillips et differences in the eight measurements studied. The Ja aI., 1989). Two aspects ofthe mtDNA data are highly maican fruit bat canpotentially be confused with the other relevant to our understanding ofsubspeciation as dis large species of Artibeus occurring on Grenada, A. cussed in the previous paragraph, and to our overall un lituratus, when attempting to identify the species under derstanding ofthe zoogeographic position ofGrenada field conditions. However, the two species canbe sepa and its bat populations. First, in terms ofmtDNA, A. rated based solely using the measurement ofthe length }amaicensis grenadensis previously was reported to be offorearm. We have measured the length offorearm genetically diversified in comparison to other island for 200 A.}amaicensis from the island, giving a mean of populations examined (phillips etal., 1989). We recently 58.0. The smallest individual, an adult male from along reanalyzed the restriction site data bymeans ofArlequin the Chemin River, had a forearm that measured 52.5, genetic analysis software (ver. 1.0, Schneideret al.,1996) whereas the largest individual, an adult female from near and expanded the survey to include bats from additional Vendome, measured 62.2. This can be compared with islands and Yucatan. We obtained the following estimates the range oflength offorearm for 31 A. lituratus, which ofnucleotide diversity in mtDNA in Antillean island was 66.9 to 75.7. populations of A. }amaicensis: Grenada, 0.0331; St. Lucia, 0.01226; St. Vincent, 0.00984; Barbados, Table 3presents the reproductive data available for 0.00795; Puerto Rico, 0.00326; Jamaica, 0.00174; Artibeus}amaicensis from Grenada. Although these data Anguilla, 0; Bequia, O. Nucleotide diversity obviously is are incomplete, only covering the period of March independent ofisland size, and in general there is a de through August, they fit well with the data presented by crease from south to north. The high nucleotide diver Wilson et ai. (1991) for populations ofthis species on sity ofmtDNA in A.}. grenadensis in part reflects the Barro Colorado Island, Panarrui. They concludedthat the fact that all ofthe haplotype lineages (labeled G, SV, and reproductive pattern ofthe Jamaican fruit bat is bimodal J) discovered in the Caribbean are represented on the polyestry with females palpably pregnant inJanuary and island (phillips et aI., 1989, 1991). Another component giving birth in late February and early March and again ofthe diversity is a haplotype lineage (G) that was repre beingpalpablypregnant in April, May, orJune with births sented in 61 % ofthe 28 bats examined from Grenada. in July and August. Although ourdata set is limited for This haplotype lineage is not unique to Grenada, butmust the period ofJanuary through April, we have data for six be extremely rare elsewhere in the Antilles. Indeed, it females taken in early March (Table 3). One ofthese was recovered from only one of134 (0.7%) bats from females was carrying ajuvenile male with a total length Bequia and St. Vincentnorth through Jamaica This single of58, two were noted as having near term fetuses, and individual with aG haplotype was obtained on S1. Vincent three were noted as having small fetuses. Although not (1/20 bats examined). A third component ofthe genetic conclusive, these data would fit the pattern reported by diversity on Grenada comes from the SV haplotype lin Wilson et ai. (1991). Our data are more complete for eage, which was obtained from 10 individuals (36%). the period covering the secondpeak ofreproductive ac This mtDNA is typical ofA.}. schwartzi on St. Vincent tivity and they do fit very nicely with the second peak of (pumo et aI., 1988) and also has been isolated from pregnancy in May-June with births in July-August as seen Artibeusplanirostris collected in French Guiana (Pumo in Panama (Wilson et aI., 1991). In addition to the data et aI., 1996). Finally, it is noteworthy that the J haplo in Table 3, we took three flying subadults in a sample of type lineage, which predominates in the Greater Antilles, 57 individual in late May. Lengths offorearms of58.2, is rare inA.}. grenadensis (Phillips et aI., 1991), being 58.7, and 59.0 indicated that these individuals had reached found so far in a single individual (3.6%). The rarity of adult size even though some ofthe phalangeal epiphyses the J haplotype is consistent with the hypothesis that had not closed. In a sample 39 individuals taken on 11 Artibeus}amaicensis on Grenada reached the island from August, 16 were considered to bejuveniles or subadults the south rather than the north. with the length offorearm ranging from 39.1 to 59.1. GENOWAYS ETAL.--BATS OF GRENADA 17 Table 3.-Reproductive data recordedfor Artibeus jamicensisfrom Grenada. Date ofCapture Females Males Total no. No. pregnant Crown-rump No. No. Length length lactating oftestes 2 March 6 5 1 0 23 May 3 3 15.7 (13-20) 0 2 5,5 24 May 6 3 24.0 (10-33) 0 1 1 25 May 5 5.2(2-7) 26 May 15 10 14.5 (6-22) 2 4 4.5 (4-6) 27 May 11 6 14.0(6-20) 2 2 7,7 28 May 7 4 17.3(6-25) 1 5 5.9 (4-8) 29 May 5 3 11.0 (10-20) 0 0 30 May 29 25 16.2(5-40) 1 1 7 31 May 1 0 0 0 1June 1 1 23 0 0 18 July 1 1 0 3 7.8(7-9) 19 July 8 4 0 2 5,7.5 16-17 August 15 0 2 2 5,9 These data again are directly comparable to those reported stagnant, but formed several relatively large and deep by Wilson etal. (1991). pools. The report from Douglaston (Jones, 1951) was based on a single individual taken along with specimens ofA.jamaicensis from the roofofa distillery. Artibeus lituratuspalmarum J. A. Allen and Chapman, 1897 We follow previous authors in assigning our speci GreaterFruit-eating Bat mens to the subspecies palmarum with a type locality at Port ofSpain, Trinidad. Comparing the measurements Specimens examined(46).-ST.ANDREW: Birch ofourspecimens in Table 1with those given by Dalquest Grove, 2 (UNSM); 0.4 km S, 0.4 km W St. Margaret, (1951), Goodwin and Greenhall (1961), and Davis 1600ft, 12°06'N,61°41.5'W, 1(CM). ST. DAVID: 11/ (1984) for specimens from Trinidad revealed mean val 4kmS, 3/4kmWBeaton, 12°01.5 N,61°42'W,4 (CM). ues that were similar and had abroadoverlap in the range ST. GEORGE: Chemin River, 1/2 km E Confer, 12°01' ofthe measurements. T-tests ofthe eight measurements N, 61 °43' W, 29 (23 CM, 6 UNSM); St. Paul's, 12°04' presented in Table 1revealed no significant secondary N, 61°44' W, 6 (CM); 1/2 km E Vendome, 12°05' N, sexual variation in our sample. As pointed out previ 61 °42.5' W, 4 (2 CM, 2 TTU). ously, it is not always easy to separate the greater fruit bat from the Jamaican fruit bat under field conditions. Additional record.-Douglaston (Jones, 1951). On Grenada, however, there is no overlap in the length of forearm ofthese two species. Thirty-one greater fruit Remarks.-The greater fruit-eating bat is wide bats had an average length offorearm of70.4 with arange spread on Grenada, but we obtained large numbers of of66.9 to 75.7, as compared with 52.5 to 62.2 for the specimens only along the Chemin River. This species Jamaican fruit bat. The smallest ofthe greater fruit bats was taken at several sites along with it congener A. was an adult female from near Vendome and the largest jamaicensis, which always was the more abundant ofthe was an adult female from St. Paul's. two. The site along the Chemin River was near open sug arcane fields, but several large trees on both sides ofthe Reproductive data for the greater fruit bat on river formed a canopy in some areas. Away from the Grenada are limited and does not allow discerning the river vegetation was semitropical shrub forest and brush. pattern ofreproduction. The reproductive cycle ofthis At the time that we visited this area the river was nearly species is variable on the mainland (Wilson, 1979), with 18 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY a single young produced in northern Middle America and Females taken on 25 and 26 May were lactating, Mexico, in Costa Rica and Panama it has abimodal cycle, whereas a female taken on 30 May evinced no gross re and in Colombia the large fruit bats is a continuous productive activity. Males taken on 25 and 26 May had breederwith bimodal activity peaks. Onlyone pregnant testes that measured 6 and 3, respectively, in length. female was recorded in oursample from Grenada, which was taken on 29 May with an embryowitha crown-romp length of7. Six non-pregnant[ema1es were noted on Myotis n;gr;cans n;gr;cans 23-24 May. Lactating females were takenon 23 and 29 (Schinz, 1821) May. The mean length oftestes of11 adult males cap BlackMyotis tured between 23 May and 2 June was 5.6 (3-7). Specimens examined (3).-ST. DAVID: 11/4 km S, 3/4 km W Beaton, 12°01.5' N, 61°42' W, 1 (CM). Sturn;ra [ilium serotinus PARISH UNKNOWN: no specific locality, 2 (NMNH). Genoways, 1998 Yellow-shouldered Bat Remarks.- Koopman (1968, 1994) assigned a specimen from Grenada to M n. nigricans and we have Specimens examined (7).-ST.ANDREW: Birch maintained this arrangement here. LaVal (1973) in his Grove, 3 (UNSM). ST. GEORGE: 1/2 kmEVendome, revision ofthe Neotropical bats ofthe genus Myotis de 12°05' N, 61 °42.5' W, 4 (CM). scribed a new species from Martinique and Barbados in the central Antilles and recognized a distinct species Remarks.-These specimens constitute the first occurring on Dominica Ithas subsequentlybeen reported records for yellow-shouldered bats from Grenada. This from Guadeloupe (Masson and Breuil (1992), but he did species is known as far north in the Lesser Antilles as not examine or assign specimens from Grenada to a spe Dominica (Jones and Phillips, 1976) and a closely re cies. We have compared ourspecimens with the descrip lated species, Sturnira thomasi, is known from tions givenbyLaVal (1973) and believe that Koopman's Guadeloupe (Genoways and Jones, 1975) and Montserrat initial identification was correct. Our specimen from (pedersen et aI., 1996). This material recently has been near Beaton has a longer tibia (tibia to forearm ratio is described as a new subspecies (Genoways, 1998) based 0.43) characteristic of M. nigricans rather than M. upon its cranium being proportionally narrower than the dominicensis (tibia/forearm 0.35). The specimens from population ofSturnira lilium lilium from Trinidad and Grenada are smallerthanM martiniquensis and closely overall smaller size (Table 1) than other Antillean forms match measurements given by LaVal (1973) for M ofthe species (angeli, luciae,paulsoni, andzygomati nigricans from Venezuela. They have dorsal pelage that cus; Jones and Phillips, 1976). Males were significantly is darker at the base than the tip, which is characteristic (t-test) larger than females in greatest length ofskull, ofM nigricans, and lack a sagittal crest on the cranium, condylobasallength, length ofmaxillary toothrow, and which usually is present in M martiniquensis. mastoid breadth (Table 1). The specimen from Beaton was an adult male with We collected three ofour specimens in nets over testes length of3 on 2 June 1986. Other specimens are a branch ofthe Balthazar River and an adjacent area be males taken in March 1928 and on 25 August 1929, with hind the Police Station in Birch Grove. Vegetation in the former taken by Dr. Paul Bartsch and the latter by the area was a mixture offruit and native trees. When we Rev. Sebastian Gates. visited the site on 26 May 1987 the fruits on the mango, guava, and nutmeg trees were ripening and there was con Our specimen from near Beaton was netted over a siderable activity by bats during the night. There was no small stream in an area that was just being slashed and canopy over the fairly broad river, which contains huge burned to open it for agricultural purposes. A poorly boulders with water braiding among them. The site at developed gallery forest formed a canopy at intermit Vendome where the other four specimens were captured tent points along the stream. Away from the stream veg is described in the account for Noctilio. etation was semi-deciduous, thorny bushes and small GENOWAYSET AL.-BATS OF GRENADA 19 trees. The area was so disturbed that several hundred and roof. A large colony inhabited the roofofa cottage active snakes were observed during the evening ofnet at St. George's, and here a small series was easily ob ting. tained by placing a dipnet over one ofthe openings. The owner ofthe house told us that the bats were active all night, constantly coming and going." This species also Molossus molossus molossus is captured easily in mist nets set over large calm pools (pallas, 1766) ofwaterwith little orno canopy. These areas allow these Pallas' MastiffBat bats to drink, while only slightly slowingtheir rapid flight. Specimens examined (50).- ST. GEORGE: The systematic relationships ofthe small mastiff Chemin River, 1/2 km E Confer, 12°01'N,61°43'W,30 bats occurring in the Antilles is still a matter ofconsid (28 CM, 2 UNSM); St. George's, 17 (MCZ); 1/2 km erable debate (Miller, 1913b; Koopman, 1968; Dolan, Vendome, 12°05'N,61°42.5'W,2(CM). PARISHUN 1989; Jones, 1989). Dolan (1989) made the most thor KNOWN: no specific locality, 1(NMNH). ough analysis ofthe complex nomenclatural history and relationships ofthese small bats and we follow her ar Remarks.-This insectivorous species is far more rangement and that ofKoopman (1994) in assigningpopu abundant on Grenada than our collections would indi lations on Grenada to the nominate subspecies. Our cate. Iteasily is identified by its high rapid flight in the specimens are clearly larger (Table 1) than specimens earliest evening, while swifts and other birds are still from southern Mexico, Guatemala, and EI Salvador re active. Pallas' mastiffbats are undoubtedly far more ported byDolan (1989), but a detailed analysis ofvaria abundant currently on Grenada than priorto human habi tion throughout the range ofspecies will be necessary tation because man-made structures are now their pri before a final decision will be possible on allocating all mary roosting site. As pointed out by Allen (1911) this specimens to the appropriate taxon. species "lives in colonies underneath the roofs of houses. Buildings covered with galvanized iron sheeting Of10 females examined for reproductive activity are especially favored by it since the small holes left on 23 May 1980, nine were pregnant. Embryos ranged open where the convexities of the sheeting meet the in crown-rump length from 12 to 16.5, with a mean of rafters afford ready ingress to spaces between ceiling 13.2. DISCUSSION Grenada, being situated at the southern end ofthe The origin ofthe chiropteran fauna ofGrenada Lesser Antilles, occupies an interesting zoogeographic stands in contrast to the other group of volant verte position. Itis positioned to receive bat species invading brates-birds. The avifauna ofTobago is composed of the Antilles from the south via Trinidad or Tobago and 70 species ofwhich 50 are ofprobable South American the South American mainland and for Antillean endemic origin and 20 are widespread species, whereas the avi species to arrive from the north via St. Vincent and the fauna ofGrenada consists of35 species ofwhich 9 are Grenadines. We do believe that all ofthe species ofbats ofAntillean origin, 6 ofSouth American origin, and 20 currently on islands in the Caribbean region have arrived are widespread species (Lack 1976). Lack (1976) was via over water migration (Baker and Genoways, 1978; so struck by the faunal differences between these two Pregill, 1981; Morgan and Woods, 1986; Pumo et aI., islands that he proposed "Bond's Line" as the faunal 1988, 1996; Phillips et aI., 1989, 1991) rather than be boundary between the West Indian fauna ofGrenada and ing the result ofvicariance (Rosen, 1975). All species the South American fauna ofTobago. There has been a ofbats on Grenada can be classified as either recent in plethora ofrecent investigations ofmammalian fauna of vaders from the south or as widespread species for which the West Indies, especially ofbats, to the point that we no dispersal route is discernible (Koopman, 1968, 1989; believe that the zoogeographic boundary ofthe West In Jones and Phillips, 1970; Baker and Genoways, 1978; dian subregion can now be defined with relative certainty. Jones, 1989). We here propose to call this boundary demarking the 20 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY West Indian Faunal subregion as "Koopman's Line" in Currently, the relationship ofsouth Florida and the honor ofour late colleague Dr. Karl F. Koopman, who Florida Keys to the West Indian Faunal subregion is open published many ofthese studies, including the first de to question because the bat fauna is poorly known and limitingthe boundaries ofthe West Indian subregion (Fig. bats seem to be relatively rare in the area; however, there 1). is a growing body ofdata that would seem to relate this area to the subregion. Recently, Frank (1997) reported Our West Indian Faunal subregion ofthe Neotro Molossus molossus tropidorhynchus, the Cuban sub pica! region differs from the one presented by species ofthis bat, from four ofthe Keys, including Key Hershkovitz (1958) inseveral details. We are excluding West, Vaca Key, Boca ChicaKey, and Stock Island. He Grenada and the Grenadines from the region. We have found three established colonies, but could not be cer questioned the placement ofthe Grenadines because the tain whether their origin involved over water dispersal bat fauna ofthis series ofislands is poorly known. The from Cuba (Silva Taboada, 1979) orhuman transporta four species reported from this complex of islands tion. Fossil and Recent specimens ofEumops glaucinus (Jones, 1989) are all widespread in distribution and give floridanus document that this Neotropical species has no indication oftheir faunistic relationships. We antici been in Florida since the Late Pleistocene (Koopman, pate that one or two (Monophyllus plethodon and/or 1971; Morgan, 1991). Although the nearest population Brachyphylla cavernarum) ofthe Antillean endemics ofthis species is on Cuba (Silva Taboada, 1979), Mor known from St. Vincentprobably also occur at least on gan (1991) could not rule out that this species had dis Bequia, which is justover 2 Ian south ofSt. Vincent and persed into Florida via the GulfCoast from mainland can be seen from there. This means that Koopman's Line populations ofthe species in Mexico. There are at least may eventually be found to bisectthe Grenadines. two reports ofArtibeus jamaicensis from Key West (Maynard, 1872; Lazell and Koopman, 1985), but be Along the south side ofthe subregion we are ex cause the specimen was not preserved in either case, cludingTrinidad, Tobago, Margarita, Aruba, Bonaire, and these reports can not be verified. Further doubt was cast Curas:ao (Koopman, 1958, 1959; Husson, 1961; byHumphrey and Brown (1986) who disputed the iden Goodwin and Greenhall, 1961; Smith and Genoways, tification ofthe photographedbatinLazell and Koopman 1974; Genoways and Williams, 1979; Carteretal., 1981; (1985). Morgan (1991) acknowledged that A. Eshelman and Morgan, 1985; Morgan andWoods, 1986). jamaicensis could occur on Key West, but concluded In the west we exclude Isla Cozumel, Isla Mujeres, Half that they were only accidental occurrences and that no moon Cay, Roatan, Bonacca, and SanAndreas (Koopman, population had been established. Fossils oftwo Neotro 1959; Jones et aI., 1973). We differ from Koopman pical bats lend some additional evidence to the West In (1959) in excluding San Andreas, which he included be dian relationships ofsouth Florida and the Florida Keys. cause ofit proximity to Providencia; however, the only The modem species Mormoops megalophylla only is species currently known from San Andreas is Artibeus known as a fossil from Florida, Bahamas, Dominican jamaicensis, but this species is widely distributed so its Republic, Cuba, and Jamaica. Morgan (1991) concluded mere presence does not provide much insight into zoo that the Florida and West Indian specimens were "virtu geographic patterns. We do agree with Koopman (1959) ally identical in size" and, furthermore, these specimens in including Providencia in the West Indian subregion could not be distinguished from modem mainland popu because ofthe presence ofAntillean endemic Natalus lations ofthe species from Mexico and South America. brevimanus in addition toA.jamaicensis. We also in Morgan (1991) also reported a fossil bat from Florida clude the Swan Islands in the West Indian subregion be as Pteronotus cf. P. pristinus, which is a fossil species cause of the occurrence of the endemic hutia described by Silva Taboada (1974) from Cuba. Although (Geocapromys thoracatus), which is ofAntillean affin the evidence is not compelling enough to include south ity (Koopman, 1959; Morgan, 1985). In the north we Florida and the Florida Keys in the West Indian Faunal include all ofthe Bahamas (Koopman et al., 1957; Baker subregion without question, the evidence does seem to and Genoways, 1976; Buden, 1986; Morgan and Woods, be accumulating and we should certainly be aware that 1986; Jones, 1989). this relationship may exist. GENOWAYS ETAL.--BATS OF GRENADA 21 II 7. 72 II •• •• 'DO. 2DO ... ..- ~ ..lDO 2 11 - c=::7 .-" 11 - ~ - .... 0 • Koopmanls Line 12 I' '. ".-.> .. II 71 II Figure 1. A map of the Caribbean region showing the limits of Koopman's Line, which defmes the boundary of the West Indian Faunal subregion based upon mammalian distribution patterns. Like the avifauna data, our data reflect a disparity (Wunderle, 1985). Previous authors (Eshelman and in the size ofthe chiropteran fauna between Tobago and Morgan, 1985; Wunderle, 1985) have not found an ex Grenada (Lack, 1976;Wunderle, 1985). Twelve species planation for this phenomenon. Itcan not be attributed ofbats are known from Grenada, 20 species are known to the size of the islands, which Morgan and Woods from Tobago (Eshelman and Morgan, 1985), and 64 spe (1986) found to account for 69% ofthe mammalian spe cies are known from Trinidad (Carter et al., 1981). In cies diversity on Antillean islands. They believed that the avifauna, 35 species have been recorded from Grenada the other 31% could be related to "variables such as habi and 70 from Tobago. The chiropteran fauna does not tat diversity and distance from source areas." Habitat quite reach a doubling ofthe number ofspecies on To diversity can not explain differences because in the bago compared to Grenada that the avifauna does, but it Antilles habitat diversity is directly affected by eleva is 1.66 times larger. The difference in the size ofthe tion, which in this case would favor higher diversity on fauna between Trinidad and Grenada is consistent with Grenada. We believe that the most important factor af island biogeographic theory, which predicts that island fecting the mammalian species diversity ofTobago and size and distance to the source area (mainland) are im Grenada is distance from the source area, which in the portant factors. Tobago on the other hand is quite simi case ofTobago is primarily Trinidad, whereas the source lar to Grenada in size (300 sq km versus 311 sq km, re for the fauna on Grenada is either Trinidad or the South spectively), elevation (580 m versus 840 m), distance American mainland, or both. This makes the source for from South American mainland (both about 130 km, al Grenada at least three times further away than for To though Tobago is only about 35 to 40 km from Trinidad), bago. climate, cultivated crops, and area under cultivation 22 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY McFarlane (1989) examined patterns ofco-occur son we also included 12 specimens from the Yucatan rence ofspecies inthe Antilleanbat fauna and concluded Peninsula. We scored each animal based on our previ that..."The structure ofbat assemblages on the Greater ous work and usedpreviously unavailable analytical soft Antilles is dominated by the co-occurrence ofsingle is ware (Arlequin, ver 1.0, Schneideret al., 1996; Holsinger land endemics, whereas a similar structure in the Lesser and Mason-Gamer, 1996) to estimate nucleotide diver Antilles isdominated bythe co-occurrence ofundiffer sity per island and genetic distances among islands in entiated South American taxa." We believe that the situ pairwise comparisons. The complete analysis goes be ation in the southern Lesser Antilles is more complex yond the present paper and will be presented elsewhere than this conclusion would indicate. If the co-occur (phillips et al., in preparation), but the data strongly sup rence of bat species southward from Montserrat to port our idea that Grenada falls outside the West Indian Grenada is considered, the species Noctilio leporinus, faunal subregion. Indeed, genetic distance (FST) values Artibeusjamaicensis, and Molossus molossus are found between Grenada and the otherseven islands were 0.435 to occur on all nine ofthese islands or island groups. (St. Vincent), 0.446 (Bequia), 0.450 (St. Lucia), 0.92 This would support the conclusions ofMcFarlane (1989) (Barbados), 0.509 (Anguilla), 0.578 (Jamaica), and 0.609 because these are widespreadspecies. Onthe otherhand, (Puerto Rico), and all comparisons were highly signifi ifGrenada and the Grenadines are eliminated from this cant (P< 0.01). By way ofcomparison, the FST value group ofislands as being outside ofthe West Indian sub between Puerto Rico and Anguilla and Anguilla and Bar region, Monophyllus plethodon and Brachyphylla bados was 0.009 and was nonsignificant. The striking cavernarum will be added to the list ofco-occurring nature ofthe data is further illustrated by the fact that species ofbats on these islands. Additionally, Ardops although the FST value (0.362) between Grenada and the nichollsi occurs on all except Barbados. These are three Yucatan Peninsula is significant (P LITERATURE CITED Allen, G. M. 1911. Mammals ofthe West Indies. Bulle Frank, P. A. 1997. First record ofMolossus molossus tin ofthe Museum ofComparative Zoology, tropidorhynchus Gray (1839) from the United 54: 175-263. States. Journal ofMammalogy, 78: 103-105. Baker, R 1. andH. H. Genoways. 1978. 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O. Smith, J. D. and H. H. Genoways. 1974. Bats of Handley, Jr., D. E. Wilson, and A L. Gardner, Margarita Island, Venezuela, withzoogeographic eds.), Smithsonian Contributions to Zoology, comments. Bulletinofthe Southern California 511: iii + 1-173. Academy ofScience, 73: 64-79. Wunderle, J. M., Jr. 1985. An ecological comparison VanDenBussche, R. A, R. 1. Baker, H. A Wichman, and ofthe avifauna ofGrenada and Tobago, West M. J. Hamilton. 1993. Molecular phylo Indies. Wilson Bulletin, 97: 356-365. genetics ofstenodermatine bat genera: Congru ence of data from nuclear and mitochondrial DNA. Molecular Biology and Evolution, 10: 944-959. 28 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY Addresses ofauthors: HUGHRGENOWAYS ROBERT J. BAKER University ofNebraska State Museum, and Department ofBiological Sciences, and School ofNatural Resource Sciences Museum ofTexas Tech University W436 Nebraska Hall Lubbock, TX 79409-3131 University ofNebraska-Lincoln e-mail: [email protected] Lincoln, NE 68588-0514 e-mail: [email protected] CARLETON J. PIDLLIPS Department ofBiological Sciences lllinois State University Normal, IL 61761 e-mail: [email protected] PUBLICATIONS OF THE MUSEUM OF TEXAS TECH UNIVERSITY Itwas through the efforts ofHorn Professor J Knox Jones, as director ofAcademic Publications, that Texas TechUniversity initiated several publications series including the Occasional Papers ofthe Museum. This and future editions in the series are a memorial to his dedication to excellence in academic publications. Professor Jones enjoyed editing scientific publications and served the scientific community as an editor for the Journal ofMammal ogy, Evolution, The Texas Journal ofScience, Occasional Papers ofthe Museum, and Special Publications ofthe Museum. Itis with special fondness that we remember Dr. J Knox Jones. Institutional subscriptions are available through the Museum ofTexas Tech University, attn: NSRL Publications Secretary, Box 43191, Lubbock, TX 79409-3191. Individuals may also purchase separate numbers ofthe Occa sional Papers directly from the Museum ofTexas Tech University. ISSN0149-175X Museum ofTexas Tech University, Lubbock, TX 79409-3191