Occasional Papers Museum of Texas Tech University NUMBER 176 1 MAY 1998 TWO NEW SUBSPECIES OF BATS OF THE GENUS STURNIRA FROM THE LESSER ANTILLES, WEST INDIES Hugh H. Genoways The last systematic review of the yellow-shoul¬ dered bats of the Neotropical genus Sturnira in the Lesser Antilles was in 1976 (Jones and Phillips, 1976). At that point in time, two species —Sturnira lilium and Sturnira thomasi —were known from these islands. Sturnira lilium was represented by five subspecies, beginning with Trinidad and moving northward, these subspecies were lilium on Trinidad (Goodwin and Greenhall, 1961), paulsoni on St Vincent (de la Torre and Schwartz, 1966), luciae on St. Lucia (Jones and Phillips, 1976), zygomaticus on Martinique (Jones and Phillips, 1976), and angeli on Dominica (de la Torre, 1966). By way of contrast to Sturnira lilium , S. thomasi is endemic to the Antilles, with a single population de¬ scribed from the island of Guadeloupe. Interestingly, S. thomasi must either be rare or difficult to capture in mist nets (or both) because the original description of this species was based upon a single specimen (de la Torre and Schwartz, 1966). Koopman (1968) was “in¬ clined to regard paulsoni , angelic and even thomasi as successive modifications of lilium out of contact with other species of Sturnira ” In the case of S. thomasi , this recommendation has not been supported by subse¬ quent authors (Genoways and Jones, 1975; Jones and Phillips, 1976; Baker and Genoways, 1978; Baker et ai., 1978; Owen, 1987; Koopman, 1989, 1993; Jones, 1989; Pacheco and Patterson, 1991; Pedersen et ai,, 1996). Genoways and Jones (1975) and Owen (1987) have reported an additional six specimens of S. thomasi from Guadeloupe. The genus Sturnira thus was known from five is¬ lands in the Lesser Antilles with Guadeloupe being the furthest north. Of the Lesser Antillean islands to the south of Guadeloupe, according to Jones and Phillips (1976), representatives of the genus Sturnira were not known from Marie Galante, Barbados, The Grenadines, and Grenada. Subsequent to the paper by Jones and Phillips (1976), members of the genus have been cap¬ tured on Grenada (reported herein) and Montserrat, which lies about 55 km to the northwest of Guadeloupe (Pedersen et al., 1996). Study of these specimens has convinced me that these populations represent two undescribed subspecies, which are characterized and named below. In the two accounts below, all measurements are given in millimeters. Statistical analyses were per¬ formed using the St at Gew® software package (Sager, 1992). The paired t-test gave statistical significance of differences in group means. I am grateful to the follow¬ ing people for allowing me access to specimens housed in their research collection: the lafe Karl F, Koopman, American Museum of Natural History (AMNH); Rob- 2 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY ert M. Timm, University of Kansas (KU); Robert J. Baker, Texas Tech University (TTU); Suzanne B. McLaren, Carnegie Museum ofNatural History (CM); Patricia W. Freeman, University of Nebraska State Museum (UNSM). 1 also wish to thank my field com¬ panions, who were involved in collecting these bats: Robert J. Baker, Carleton J. Phillips, Dorothy Pumo, and Scott Pedersen. Figure 1 was prepared by Angie Fox, Staff Artist, University ofNebraska State Museum. DESCRIPTIONS Sturnira lilium serotinus new subspecies Holotype. — Adult male, skin and skull, UNSM 16493, from Birch Grove, St. Andrew Parish, Grenada; obtained by Hugh H. Genoways on 26 May 1987, origi¬ nal no. HHG 5609. Etymology .— serotinus Latin, meaning happen¬ ing late; referring to the fact that the description of this subspecies is happening late in comparison to other Antillean populations o f Sturnira lilium. Distribution .— Known only from Grenada (Fig. 1 ). Diagnosis. — Forearm short for Antillean repre¬ sentatives of the species; cranium narrow (Table 1); the first upper molar with a small anterior labial shelf and a prominent metacone; second upper molar with a poorly-developed anterior labial shelf; third upper mo¬ lar of average size; in the first lower molar the paraconid is located near the lingual edge of the tooth so that the paraconid, metaconid, and entoconid nearly forma lin¬ ear lingual sequence; entoconid well developed into an elongate cone; third lower molar small with only two cuspids with the lingual the larger of the two; the pel- age is a reddish to yellowish brown with the “ydlow shoulders” present. Comparisons. — Morphometrically, the speci¬ mens from Grenada have a length of forearm that is significantly smaller than those of the other Antillean populations (Table 1), but is matched by S. I lilium from Trinidad. In length measurements of the cranium, the Grenada population matches those from the Antillean islands and Trinidad for greatest length of skull, and in length of maxillary toothrow, the Grenada population does not significantly differ from populations of S. lilium on Trinidad, Martinique, and Dominica, but is signifi¬ cantly longer than the toothrow of specimens from St. Lucia and St. Vincent. In three measurements of the width of the cranium (postorbital constriction, mastoid breath, and breadth across upper molars), the popula¬ tion on Grenada has a significantly narrower cranium than all populations except in breadth of mastoid of specimens from Martinique where there is no signifi¬ cant difference. In the fourth cranial breadth measure¬ ment, zygomatic breadth, the Grenada population was significantly smaller than S. I. lilium from Trinidad and S. i zygomaticus from Martinique. In characteristics of the first upper molar, the specimens from Grenada most closely resemble those from Trinidad in the reduction of the anterior labial shelf and next most closely resemble the Dominica and the Martinique populations, with the populations from St. Lucia and St. Vincent having the most highly devel¬ oped anterior labial shelf. The third upper molars is smallest in the Dominican population and largest in the St. Lucian population, with populations from St. Vincent, Martinique, Grenada, and Trinidad on the averaging an intermediate size between these extremes (Jones and Phillips, 1976). The morphology of the lower first molar groups the Grenada population with populations on St. Lucia and St. Vincent where the paraconid is located on the lingual edge of the tooth so that the paraconid, meta¬ conid, and entoconid form a linear lingual sequence and the entoconid is enlarged as opposed to the condition found in populations from Trinidad, Martinique, and Dominica where the paraconid is located nearer to the center line of the tooth than the lingual margin so that the paraconid does not form a linear lingual sequence with the metaconid and entoconid and the entoconid is a low ridge. The third lower molar of the Grenada popu¬ lation was the smallest among the populations exam¬ ined with only two cuspids present. s a red- Trinid >aler be- variati ic genus m from i OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY Table 1 .— Forearm and cranial measurements of six subspecies o/ Sturnira lilium from the southern Lesser Antilles and Trinidad. Testing for significant differences between the means of S. 1. serotinus and the other subspecies was performed using paired t-tests. Measurements and statistics serotinus lilium paulsoni iuciae zygomaticus angeli Length of forearm N 6 10 6 8 8 10 Mean 42.0 42.3 43.3* 43.9** 43.6** 43.6* Range 40.142.6 40.8-44.3 42.6-44.2 42.7-44.8 42.8-45,0 41.7-45.2 ±SE ±0.39 ±0.35 ±0.23 ±0.25 ± 0.29 ± 0.37 Greatest length of skull N 6 10 6 8 8 10 Mean 22.7 23.0 23.1 22.6 22.7 23.1 Range 21,8-23.3 22.2-23.6 22.7-23.5 22.1-23.0 22.4-23.0 22,7-23.7 ± SE ±0.22 ±0.15 ±0. 17 ±0.15 ± 0.07 ±0.11 Zygomatic breadth N 6 10 6 8 8 10 Mean 13.0 13.5* 13.2 13.2 13.5* 13.0 Range 12.8-13.3 12.844.2 12.8-13.5 12.8-13.7 13.0-13.9 12.7-13.2 ± SE ±0.08 ±0.13 ±0.08 ±0.10 ± 0.10 ±0.08 Postorbital constriction N 6 10 6 8 8 10 Mean 5.7 6.1*** 5.9** 9* * 6.0** 5.9* Range 5.5-5.8 5.7-6.3 5.7-6.0 58-6.2 5.6-6.3 5.7-6.4 ± SE ±0.04 ±0.07 ± 0.05 ±0.06 ±0.07 ±0.07 Mastoid breadth N 6 10 6 8 8 10 Mean 11.5 12.0** 11,8* 11,8* 117 11.8* Range 11.141.7 11.6-12.7 11.6-12.1 11.2-12.1 11.3-11.9 11.542.1 dtSE ± 0.10 ± 0. H ± 0.09 ± 0.10 ±0.08 ± 0.08 Length of maxillary toothrow N 6 10 6 6 8 10 Mean 6,7 6.8 6.4*** 6.4** 6.7 6.8 Range 6,6-6.8 6.4-7.1 6.3-6.5 6.2-66 6.5-6.8 6.5-7.0 ± SE ± 0,03 ±0.07 ±0,03 ± 0.07 ±0,05 ±0.08 Breadth across upper molars N 6 10 6 7 8 10 Meap 7.8 8.1*** 8.0* 8.0** 8.3*** g 1*** Range 7.7-8.0 7.9-8.5 7.8-8.3 7.8-8.1 8.1-8.5 7.9-8.3 ± SE ± 0.05 ± 0.06 ± 0.08 ± 0.04 ± 0.05 ± 0.04 km B Vendome, St. George Parish, 12°05’ N, 61°42.5' W, 4 (CM). Comparative specimens examined .— Sturnira lilium angeli : Clarke Hall Estate, 100 ft., St. Joseph Parish, Dominica, 5 (K.U); l mi from mouth of Layou River, St. Joseph Parish, Dominica, 2 (ITU); Mt. Joy Estate, St. Paul Parish, Dominica, 3 (TTU); Springfield, St. Paul Parish, Dominica, 2 (TTU). Sturnira lilium GENOWAYS— NEW SUBSPECIES OF STURNIRA 5 lilium: Caura Valley, Trinidad, i (KU); Guayaguayare, Trinidad, 1 (KU); Las Cuevas, Trinidad, 2 (TTU); St. George, San Rafael, Trinidad, 8 (ITU). Stumira lilium luciae: 1/2 mi, SE Bogius, 100 ft., Dauphin Parish, St. Lucia, 7 (KU); Union Agricultural Station, 100 ft., Gros- Islet Parish, St. Lucia, 1 (KU); 1 1/2 km E Marigot Bay, St. Lucia, 1 (UNSM). Stumira lilium paulsoni : Clifton Hill, 400 ft., St. George Parish, St. Vincent, 6 (KU), Stumira lilium zygomaticus: Balata, Martinique, 10(AMNH). Comments. — The population of Stumira lilium on Grenada can be distinguished from populations to the north in the Antilles by its shorter length of forearm and from populations both to the north and south in hav¬ ing a much narrower cranium, which is especially evi¬ dent in postorbital constriction, mastoid breadth, and breadth across upper molars. It probably is more dis¬ tinctive in these morphological characters than are other Antillean populations from each other. In characteris¬ tics of the maxillary and mandibular dentition, S . /. serotinus displays a unique combination of character¬ istics, but with the possible exception of the m3, these characteristics are within the range of characters dis¬ played by the other Antillean populations. The m3 of S. L serotinus appears to be smaller than in other popu¬ lations with only two distinct cuspids; however, given the variability ofthis tooth, more individuals need to be examined before the validity of this characteristic can be determined. The specimens from Grenada came from higher elevation (Vendome at 300 m) and wet interior valleys of the island (Birch Grove). No specimen was taken in the dry coastal areas of the island despite considerable collecting effort in this area. Three specimens were captured in nets over a branch of the Balthazar River and an adjacent area behind the Police Station in Birch Grove. The vegetation in the area was a mixture of fruit and native trees. There was no canopy over the fairly broad river, which contains huge boulders with water braiding among them. At Vendome, nets were set over a small stream as it emerged from Grand Etang Forest Reserve. The stream was one of the small upper tributaries of the Beausejour River, which enters the Caribbean Sea approximately 5 km to the west-north¬ west. The habitat at this site was characterized by wet, dense montane forest, which became fairly open under the canopy away from disturbed areas. More informa¬ tion on the natural history of the new taxon will appear in Genoways et al. (1998). Stumira thomasi vulcanensis new subspecies Holotype .— Adult female, skin, skull, and par¬ tial skeleton, UNSM 20062. from Paradise Estate, 1 kilometer S Harris [1000 m], St. George’s Parish, Montserrat; obtained by Scott C. Pedersen on 14 May 1994, original no. SCP 261. Etymology .— Vulcan was the Roman God of Fire or Volcanoes; -ensis Latin, meaning place, country, or land; thus, the name is intended to mean the place of fire or volcanoes referring to the Soufriere that has se¬ riously damaged the natural habitats and the lives of the citizens of Montserrat with its recent eruptions. Distribution. — Known only from Montserrat (Fig. 1). Diagnosis. — The holotype was compared with two specimens of Stumira thomasi thomasi and two specimens of Antillean Stumira lilium to establish its specific identity. The holotype is morphologically simi¬ lar to the S. thomasi in the characters used to differen¬ tiated. thomasi and S. lilium , including a relatively large skull with a long, narrow cranium (Genoways and Jones, 1975; Jones and Phillips. 1976). The holotype of 5. t. vulcanensis differs from the specimens of S. t. thomasi from Guadeloupe in that its cranium is proportionally and absolutely narrower al¬ though of the same length (see measurements below) and all crests and ridges are less pronounced, especially noticeable in the sagittal crest. The pelage of the holo- type is uniformly grayish brown dorsally and ventrally and lacks the “yellow shoulder” characteristic of the genus, whereas the pelage of specimens from Guadeloupe is reddish brow n and reddish-yellow shoul¬ der is clearly evident. Forearm and cranial measurements of the holo¬ type followed by measurements of two specimens from Guadeloupe are as follows: length of forearm, 44.7, 46.1, 47.7; greatest length of skull, 24.9, 24.9, 25.1; condylobasal length, 23.3, 22.9, 23.6; zygomatic 6 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY breadth, 11.6, 12.2, 12.5; mastoid breadth, 10.9, 11.7, 11.8; breadth ofbraincase, 9.3, 9.8, 9.6; breadth of in¬ terorbital constriction, 5.8, 5.9, 6.0; breadth of the pos¬ torbital constriction, 5.8, 5.5, 5.9; length of maxillary toothrow, 7.3, 6.9, 6.9; width across upper molars, 7.6, 8.0,8.0; length of mandibular toothrow (i-m2), 8.4,7.7, 7.8. The holotype lacks the m3 on both sides of the mandible as do both the specimens from Guadeloupe. Specimen examined (1).— Montserrat: Paradise Estate, 1 km S Harris [1000 m], St. George’s Parish, 1 (UNSM). Comparative specimens examined .— Sturnira thomasi thomasi: 1 km W Vernou. Basse-Terre, Guadeloupe, I (TTU); 1 km S, 4 km W Vernou, Basse- Terre, Guadeloupe, 1 (TTU). Sturnira lilium serotinus: Birch Grove, St. Andrew Parish, Grenada, 2 (UNSM). Comments .— This unique specimen was first re¬ ported by Pedersen et al. (1996). It was thought at that time that it would be best to await additional specimens before formally describing this new taxon. However, with the natural disaster currently occurring on Montserrat because of volcanic eruptions, it is believed that it is best to get this taxon recorded and recognized in the scientific literature. Because this population was probably confined to the more mesic and older vegeta¬ tion on the slopes of the Soufriere, there seems to be a reasonable chance that this new taxon is now extinct. The holotype was pregnant when captured so a breed¬ ing population did occur on the island at that time. Sturnira possibly is relatively uncommon on many of the Antillean islands where it occurs and unlike some fruit bats possibly needs native humid forest instead of tropical agriculture to flourish. The morphological data all point to the fact that these bats apparently do not migrate very well and therefore gene flow is frequently interrupted. Low population levels on the various is¬ lands could contribute to the pattern in two ways: 1) no intraland pressure to disperse long distances and 2) greater possibility of local genetic drift and consequent subspecific geographic differences. LITERATURE CITED Baker, R. J., and H. H. Genoways, 1978. Zoogeogra¬ phy of Antillean bats. Pp. 53-97, in Zoogeog¬ raphy in the Caribbean: The 1975 Leidy Medal Symposium (F. B. Gill, ed,), Special Publica¬ tion, Academy of Natural Sciences of Phila¬ delphia, 13 : HI + 1-128. Baker, R. J., H. H. Genoways, and J. C. Patton. 1978. Bats of Guadeloupe. Occasional Papers of the Museum, Texas Tech University, 50: 1-16. de la Torre, L. 1966. New bats of the genus Sturnira (Phyllostomidae) from the Amazonian low¬ lands of Peru and the Windward Islands, West Indies. Proceedings of Biological Society Washington, 79: 267-272. de la Torre, L., and A. Schwartz. 1966. New species of Sturnira (Chiroptera: Phyllostomidae) from the island of Guadeloupe and Saint Vincent, Lesser Antilles. Proceedings of Biological Society Washington, 79: 297-303. Genoways, H, H., and J. K. Jones, Jr. 1975. Addi¬ tional records of the stenodermine bat, Sturnira thomasi, from the Lesser Antillean island of Guadeloupe. Journal ofMammalogy, 56:924- 925. Genoways, H. H., C. J. Phillips, and R. J. Baker. 1998. Bats of the Antillean island of Grenada: A new zoogeographic perspective. Occasional Papers ofthe Museum, Texas Tech University, in press. Goodwin, G. G., and A. M. Greenhall. 1961. A re¬ view of the bats of Trinidad and Tobago. Bul¬ letin ofthe American Museum of Natural His¬ tory, 122: 187-302. Jones, J. K., Jr. 1989. Distribution and systematics of bats in the Lesser Antilles. Pp. 645-660, in Biogeography ofthe West Indies (C. A. Woods, ed.). Sandhill Crane Press, Inc., Gainesville, FL, xvii + 878 pp. GENOWAYS— NEW SUBSPECIES OF STURNIRA 7 Jones, J. K., Jr., and C. J. Phillips. 1976. Bats of the genus Stumira in the Lesser Antilles. Occa¬ sional Papers of the Museum, Texas Tech Uni¬ versity, 40: 1-16. Koopman, K. F. 1968. Taxonomic and distributional notes on Lesser Antillean bats. American Museum Novitates, 2333: 1-13. _. 1989. A review and analysis of the bats of the West Indies. Pp. 635-644, in Biogeography of the West Indies (C. A, Woods, ed), Sandhill Crane Press, Inc., Gainesville, FL, xvii 4 878 pp. _. 1993. Order Chiroptera. Pp. 137-241, in Mammal Species of the World: A Taxonomic and Geographic Refemence(D. E. Wilson and D. M. Reeder, eds.), Smithsonian Institution Press, Washington, DC, second ed„ xviii + 1207 pp. Owen, R. D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatidae (Mammalia: Chiroptera). Special Publication of the Mu¬ seum, Texas Tech University, 26: 1-65. Pacheco, V., and B. D. Patterson. 1991. Phylogenetic relationships of the New World bat genus Stumira (Chiroptera: Phyllostomidae). Pp. 101-121, in Contributions to mammalogy in honor of Karl F. Koopman (T. A. Griffiths and D. Klingener, eds.), Bulletin of the American Museum of Natural History, 206: 1-432. Pedersen, S. C., H. H. Genoways. and P. W. Freeman. 1996. Notes on bats from Montserrat (Lesser Anti lles) with comments concerning the effects ofHurricane Hugo. Caribbean Journal ofSci- ence, 32: 206-213. Sager, S. 1992. StatVieW*. Abacus Concepts, Inc., Berkeley, CA, x + 466 pp. Address of Author: HUGH H. GENOWAYS University of Nebraska State Museum, and School of Natural Resource Sciences W436 Nebraska Hall University of Nebraska-Lincoln Lincoln , NE 68588-0514 e-mail . hge noway (if uni info. uni. edu PUBLICATIONS OF THE MUSEUM OF TEXAS TECH UNIVERSITY It was through the efforts of Horn Professor J Knox Jones, as director of Academic Publications, that Texas Tech University initiated several publications series including the Occasional Papers of the 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 of Mammalogy, Evolution, The Texas Journal of Science, Occasional Papers of the Museum, and Special Publica¬ tions of the Museum. It is with special fondness that we remember Dr. J Knox Jones, Institutional subscriptions are available through the Museum of Texas Tech University, attn: NSRL Publica¬ tions Secretary, Box 43191, Lubbock, TX 79409-3191. Individuals may also purchase separate numbers of the Occasional Papers directly from the Museum of Texas Tech University. ISSN 0149-175X Museum of Texas Tech University, Lubbock, TX 79409-3191