Caribbean Journal of Science, Vol. 38, No. 3-4, 281?284 Copyright 2002 College of Arts and Sciences University of Puerto Rico, Mayagu?ez A New Southern Distributional Limit for the Central American Rodent Peromyscus stirtoni NEAL WOODMAN1, ELISA SCHNEIDER2, PAUL GRANT3, DERRICK SAME4, KAREN E. SCHMALL5, AND JOHN T. CURTIS6 1USGS/ Patuxent Wildlife Research Center, Smithsonian Institution, PO Box 37012 National Mu- seum of Natural History, NHB 378, MRC-111 Wash- ington, DC 20013-7012, USA, 2c/o Peace Corps, P.O. Box 208, Lilongwe, Malawi 34901 Cherry Tree Bend, Victoria, BC V8Y 1S1, Canada, 41116 North Point Drive, Stevens Point, WI 54481, USA, 53144 Cowley Way #3, San Diego, CA 92117, USA, 66010 N. Calle de la Culebra, Tucson, AZ 85718, USA Stirton?s deer mouse, Peromyscus stirtoni Dickey, 1928, is a Central American endemic species with a discontinuous distribution in dry and seasonally dry lowland habitats ranging from southeastern Guate- mala to western Nicaragua (Jones, 1990; Reid, 1997). In Nicaragua, P. stirtoni was known from seven speci- mens at three localities in the west-central portion of the country (Jones and Yates, 1983; Jones, 1990). Its southernmost record was from Los Cocos, Boaco De- partment. Our field work during 1998 and 2000 on Isla de Ometepe in Lake Nicaragua (= Lago Colcibolca), at least 80 km south of Los Cocos, yielded new, vouchered records of P. stirtoni that extend its south- ern distribution. Collecting in 2000 and 2001 by a sepa- rate group from the Royal Ontario Museum (ROM) yielded additional specimens from the same locality. Specimens were identified by comparison with iden- tified museum specimens and by using characters and measurements outlined by Dickey (1928), Huckaby (1980), Jones and Yates (1983), and Jones (1990). Mice of the genus Peromyscus inhabit a number of nearshore oceanic and lacustrine islands (e.g., Hall, 1981; Lomolino, 1993). Island populations typically are subsets of mainland species with broad geographic and habitat distributions. In contrast, the distribution of Stirton?s deer mouse on the mainland is spotty, probably resulting from an ability for long-distance dispersal among scarce patches of favorable habitat, the fragmentation of a much broader prehistoric dis- NOTES 281 tribution, or a combination of vicariance and dis- persal. As in other areas, the distribution of P. stirtoni on Isla de Ometepe is restricted. All captures were made in Parque Natural ?Pe?a Inculta La Cabuya,? near Santo Domingo, Rivas Department. La Cabuya is a ca. 1190-hectare reserve that extends from 50?150 m el- evation. It lies on a flattened ridge of sharp-edged, volcanic cobbles and boulders with little soil or leaf litter. The rocky terrain probably kept the area from being cleared for farming, as has happened in most lowland areas on Isla de Ometepe. Old-growth, high- canopied, evergreen dry tropical forest grows out of this volcanic rubble, the trees supported by roots in- tertwined amongst the rocks, reaching down to the shallow water table. The forest in the study area is dominated numerically by Hura crepitans (jabillo) and Sterculia apetala (camajondura), with fewer Terminalia oblonga (guayab?n), Ceiba pentandra (ceiba), Tabebuia rosea (roble de sabana), Bursera simaruba (gumbo- limbo), and Stemmadenia donnell-smithii (huevos de ca- ballo). The understory contains many Urera baccifera (chichicaste) and Luehea candida (gu?cimo). Despite se- lective cutting of some trees (e.g., Cedrela odorata [ce- dro], for canoes), the forest appears mostly undis- turbed. This impression was reinforced by the relative abundance of emballonurid and phyllostomid bats in La Cabuya (29 individuals of at least 7 species in a single net during 10 hours in 1998) and the lack of the common vampire bat, Desmodus rotundus, a species abundant (17% of captures in 1998) elsewhere on Isla de Ometepe before eradication efforts in early 2000. We trapped for small mammals at five other sites on Isla de Ometepe within the ca. 30?975 m elevational range of P. stirtoni. Previous expeditions there in 1964 and 1968 sampled at least seven other localities (J. R. Choate, H. H. Genoways, T. E. Lawlor, J. D. Smith? unpublished field notes and catalogs, University of Kansas Natural History Museum [KU]). Although Lio- mys salvini (Salvin?s spiny pocket mouse) and Sigmo- don hispidus (hispid cotton rat) were present in dis- turbed rocky areas elsewhere on the island, the relatively undisturbed forest at La Cabuya was the only site where P. stirtoni was taken. Other workers have reported capturing P. stirtoni in disturbed habi- tats elsewhere in Central America (e.g., Peppers et al., 1999). Peromyscus stirtoni appeared to be locally abundant TABLE 1. Descriptive statistics for adult Peromyscus stirtoni from Isla de Ometepe and mainland Nicaragua. Statistics presented are mean ? SD and range. All measurements in millimeters, weight in grams. Cranial measurements are defined and illustrated by Musser et al. (1998). Isla de Ometepe Mainland external measurements (n = 7) (n = 6) length of head and body (HB) 102 ? 7 103 ? 5 (94?111) (97?109) length of tail (LT) 87 ? 7 96 ? 7 (75?95) (86?107) relative tail length (LT/HB ? 100) 85 ? 4% 93 ? 10% (79?90) (79?107) length of hind foot (LF) 23 ? 0.5 23 ? 1 (23?24) (22?24) weight (n = 7) (n = 4) 29 ? 4 31 ? 3 (23?35) (28?35) cranial measurements (n = 7) (n = 7) occipitonasal length (ONL) 30.0 ? 1.5 29.5 ? 0.6 (28.0?31.8) (28.6?30.2) greatest zygomatic breadth (ZB) 14.1 ? 0.6 13.8 ? 0.4 (13.2?14.6) (13.0?14.1) interorbital breadth (IB) 5.0 ? 0.1 4.9 ? 0.1 (4.8?5.2) (4.7?5.1) breadth of zygomatic plate (BZP) 2.3 ? 0.2 2.4 ? 0.2 (2.0?2.5) (2.2?2.6) crown length of maxillary toothrow (CLM1?3) 4.2 ? 0.1 3.9 ? 0.1 (4.0?4.4) (3.7?4.1) length of incisive foramina (LIF) 5.5 ? 0.4 5.8 ? 0.3 (5.3?6.4) (5.4?6.2) NOTES282 in 1998, but less abundant in subsequent years. We captured 10 P. stirtoni and 5 Liomys salvini using 30 Sherman traps at La Cabuya on the nights of 9 June and 11 June 1998 (60 total trapnights). We kept two adult males, two adult females, and one subadult fe- male P. stirtoni as vouchers. On the night of 15 June 2000, we captured one adult male P. stirtoni in 30 Sher- man traps set at La Cabuya. Specimens from La Cabuya in the ROM include two adult females, one juvenile female, and one juvenile male from 17 Febru- ary 2000, and an adult female from 23 February 2001. Reproductive data from the individuals captured on Isla de Ometepe are informative but represent only two short portions of the year. None of the females was gravid but three captured from 17?23 February were lactating, and one of the two females captured on 12 June had enlarged teats suggestive of recent lacta- tion. This last individual had an extensive nematode infestation of the mouth, pharynx, and esophagus. The adult males from June had abdominal testes, measur- ing 5 ? 3, 5 ? 3, and 12 ? 6 mm. Individuals in paler gray juvenile pelage were captured on 17 February and 10 June. Initial comparisons of selected cranial variables from specimens we collected in 1998 suggested that individuals from La Cabuya might be larger than those on the mainland, possibly corresponding to the ?island rule? of biogeography (Lomolino, 1985). To test this possibility, we used Minitab Release 8 to carry out principal components analysis (PCA) of a correla- tion matrix of six log-transformed cranial variables (Table 1; Musser et al., 1998) from seven adult P. stir- toni from Isla de Ometepe and seven adults from the mainland. These samples represent the total number of adults (as determined by complete adult pelage and FIG. 1. Plot of scores on PC I and PC II from PCA of adult Peromyscus stirtoni from Isla de Ometepe and mainland Nicaragua. TABLE 2. Factor loadings for PC I and PC II from PCA of six cranial variables from samples of P. stirtoni from Isla de Ometepe and mainland Nicaragua. Vari- ables are listed in descending order by their loadings on PC I. Abbreviations of measurements are explained in Table 1. Variable Loadings PC I PC II ONL ?0.568 0.089 ZB ?0.504 0.248 IB ?0.061 0.667 BZP ?0.466 ?0.301 CLM1?3 ?0.204 0.464 LIF ?0.401 ?0.424 eigenvalue: 2.7113 1.7699 proportion of variation explained: 45.2% 29.5% NOTES 283 fully erupted, worn dentition) currently available from Nicaragua. The resulting plot of specimens on PC I and PC II (Fig. 1) indicates modest separation between the two samples along PC II. This distribu- tion in part reflects the tendency of the island sample to have a broader zygomatic plate, longer maxillary toothrow, and shorter incisive foramen (Table 2). Al- though these results provide a tantalizing suggestion of morphological divergence, the low sample sizes preclude drawing firm conclusions about the distinc- tiveness of the two groups. There is little difference in overall size as measured by PCA (PC I in Fig. 1), head- and-body length (Table 1), weight, or cranial variables (e.g., for ONL, Mann-Whitney U = 33; n1 = n2 = 7; P > 0.10). In fact, the overall size variation exhibited by the sample from La Cabuya is greater than that exhibited by the mainland sample, which includes mice from three departments of Nicaragua (Fig. 1). The longer relative tail length (Table 2) of the mainland mice is not statistically significant (Mann-Whitney U = 32.5; n1 = 7, n2 = 6; P > 0.05). Subfossil remains of Stirton?s deer mouse reported from cave sediments further south in Guanacaste Province, Costa Rica (Woodman, 1988), were dated recently at 2855 ? 50 yrs B.P. (NSF-Arizona AMS Laboratory sample number AA35317). These remains were recovered from a region that still lacks a com- plete survey of its modern mammalian fauna, and it is possible that P. stirtoni still occurs there. The above records emphasize the incompleteness of our knowl- edge of the distribution and status of many small mammals in Nicaragua (and throughout much of Cen- tral America) and the continued need for basic bio- logical surveys and for samples adequate to character- ize populations of poorly known species. Acknowledgments.? Permits for field work on Isla de Ometepe were kindly granted by MARENA. Logisti- cal support was provided by the Isla de Ometepe Bio- logical Field Station. Teresa Snyder-Leiby identified the plants. Robert P. Anderson, Maria Eifler, Thor Hol- mes, and Burton K. Lim provided valuable assistance with field notes at their respective institutions. We thank the curators and collection managers at KU and ROM for loans and/or permission to study specimens under their care. Robert P. Anderson, Robert M. Timm, and an anonymous reviewer provided substantive comments that greatly improved on previous drafts of this paper. LITERATURE CITED Dickey, D. R. 1928. Five new mammals of the genus Peromyscus from El Salvador. Proc. Biol. Soc. Wash. 41:1-6. Hall, E. R. 1981. Mammals of North America. 2nd ed. John Wiley & Sons, New York, 2: 601?1181. Huckaby, D. G. 1980. Species limits in the Peromyscus mexicanus group (Mammalia: Rodentia: Muroidea). Contrib. Sci. Nat. Hist. Mus. Los Angeles County, 326:1?24. Jones, J. K., Jr. 1990. Peromyscus stirtoni. Mammalian Species 361:1?2. Jones, J. K., Jr., and T. A. Yates. 1983. Review of the white-footed mice, genus Peromyscus, of Nicara- gua. Occas. Pap. Mus., Texas Tech Univ. 82:1? 15. Lomolino, M. V. 1985. Body size of mammals on is- lands: the island rule reexamined. Amer. Nat. 125: 310?316. Lomolino, M. V. 1993. Winter filtering, immigrant se- lection and species composition of insular mam- mals of Lake Huron. Ecography 16:24?30. Musser, G. G., M. D. Carleton, E. M. Brothers, and A. L. Gardner. 1998. Systematic studies of Oryzomy- ine rodents (Muridae, Sigmondontinae): diagnoses and distributions of species formerly assigned to Oryzomys ?capito.? Bull. Amer. Mus. Nat. Hist. 236: 1?376. Peppers, J. A., J. G. Owen, and R. D. Bradley. 1999. The karyotype of Peromyscus stirtoni (Rodentia: Muri- dae). Southwest. Nat. 44:109?112. Reid, F. A. 1997. A field guide to the mammals of Central America and Southeast Mexico. Oxford University Press, New York, 334 pp. Woodman, N. 1988. Subfossil remains of Peromyscus stirtoni (Mammalia: Rodentia) from Costa Rica. Rev. Biol. Trop. 36:247?253. APPENDIX 1: SPECIMENS EXAMINED Peromyscus stirtoni (18).?NICARAGUA: BOACO: Los Cocos, 220 m, 14 km S Boaco (KU 115570?115572). MANAGUA: 8 km (by road) N Las Maderas, 380 m (KU 106797, 106798). MATAGALPA: 11 mi SE Dario (KU 71541, 71542). RIVAS: Parque Natural ?Pe?a Inculta de La Cabuya,? 50 m, 1 km NNW Santo Domingo, Isla de Ometepe (KU 157735?157739,158892; ROM 112210? 112213, 112731). NOTES284