NEW SPECIES OF STURNIRA (CHIROPTERA: PHYLLOSTOMIDAE) FROM NORTHERN SOUTH AMERICA

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1 Journal of Mammalogy, 86(5): , 2005 NEW SPECIES OF STURNIRA (CHIROPTERA: PHYLLOSTOMIDAE) FROM NORTHERN SOUTH AMERICA CORNELIO SÁNCHEZ-HERNÁNDEZ,* M. LOURDES ROMERO-ALMARAZ, AND GARY D. SCHNELL Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México, A.P , Coyoacán, México, Distrito Federal, 04510, Mexico (CS-H) Escuinapa No. 92 bis. Col. Pedregal de Santo Domingo, C.P , México, Distrito Federal, Mexico (MLR-A) Sam Noble Oklahoma Museum of Natural History and Department of Zoology, University of Oklahoma, 2401 Chautauqua Avenue, Norman, OK 73072, USA (GDS) Yellow-shouldered bats (genus Sturnira Gray, 1842), found throughout Latin America, are not well understood taxonomically. Four specimens of Sturnira from Bolivia and Venezuela possess dental characteristics differing from those of currently known species, leading us to regard them as representing a new species. We describe external, dental, and cranial characteristics and conduct a multivariate comparison using 29 morphometric characters for 44 male specimens of Sturnira erythromos, S. ludovici, S. hondurensis, and the new species. In a principal components analysis, most of the variation was summarized on component I (78.5% of total character variance), a general size vector. A multiple-comparisons test of component I projections showed all differences among species pairs to be highly significant, except that between S. ludovici and S. hondurensis. For the 29 individual characters, the new species was statistically different from S. erythromos in 21, S. ludovici in 13, and S. hondurensis in 14. The new species has a unique combination of dental characteristics, including bicuspid upper incisors, trilobed lower incisors, and smooth 1st and 2nd lower molars. Key words: Bolivia, new species, Sturnira, Venezuela Latin American yellow-shouldered bats belong to the genus Sturnira. Although they have been relatively well studied (de la Torre 1961; Iudica 2000; Pacheco and Patterson 1991; Villalobos and Valerio 2002), their taxonomy has undergone numerous changes. Recently, Sturnira mistratensis was described from Colombia (Contreras-Vega and Cadena 2001). Koopman (1993) recognized 12 species in the genus, considering S. oporaphilum and S. hondurensis as synonyms of S. bogotensis and S. ludovici, respectively. S. hondurensis initially was described by Goodwin (1940), who noted that it approached S. ludovici from Ecuador, but judged that they were specifically distinct; Goodwin (1942, 1946) reiterated this position, extending the range of S. hondurensis in Costa Rica. Hershkovitz (1949), ignoring Goodwin s (1940) opinion, lumped S. hondurensis into S. ludovici, a treatment followed by a number of subsequent authors (e.g., de la Torre 1961; Jones and Carter 1976; Koopman 1978, 1993). Timm et al. (1989) and Timm and LaVal (1998) considered S. ludovici to represent a composite species and that Central American bats * Correspondent: cornelio@servidor.unam.mx Ó 2005 American Society of Mammalogists under this name might prove to be specifically distinct, for which the name S. hondurensis was available. Iudica (2000) concluded that S. hondurensis was a valid species represented by Central American specimens and a sister taxon to a S. oporaphilum S. ludovici clade. One well-supported clade within S. hondurensis consisted of individuals from Panama and Costa Rica, which is the southernmost limit of the species as described by Goodwin (1946). We examined specimens from Venezuela that shared morphological characteristics with those cited by Goodwin (1940) for S. hondurensis. In Venezuela this genus is represented by 8 of the 13 currently recognized species (S. aratathomasi, S. bidens, S. bogotensis, S. erythromos, S. hondurensis, S. lilium, S. ludovici, and S. tildae). However, in Bolivia only 4 species have been recorded (S. bogotensis, S. erythromos, S. lilium, and S. magna Koopman 1993). Although it is relatively easy to distinguish among some species of Sturnira, there is considerable intraspecific variation in color, size, and skull shape, which can make it difficult to identify some specimens to species. However, most authors (Davis 1980; de la Torre 1966; Iudica 2000) agree that the dentition provides some of the most consistent characters, and the presence and number of dental cusps on molars and other teeth have been used as the primary bases for separation of species (Albuja 1999; Davis 1980; Giannini and Barquez 2003; 866

2 October 2005 SÁNCHEZ-HERNÁNDEZ ET AL. NEW SPECIES OF STURNIRA 867 TABLE 1. Loadings of 29 characters on principal component I based on 44 male specimens of Sturnira erythromos (n ¼ 17), S. sp. novo (n ¼ 3), S. ludovici (n ¼ 16), and S. hondurensis (n ¼ 8). Character Loading 1 Forearm length (FL) Metacarpal length of 3rd digit (ML) Length of 1st phalanx of 3rd digit (F1L) Length of 2nd phalanx of 3rd digit (F2L) Tibia length (TL) Greatest skull length (GSL) Condyloincisive length (CIL) Condylobasal length (CBL) Condylocanine length (CCL) Braincase breadth (BCB) Mastoid breadth (MST) Zygomatic breadth (ZB) Skull height (SKH) Skull depth (SKD) Interorbital breadth (INB) Interorbital constriction (INC) Rostral breadth (RB) Maxillary toothrow length (MTR) Width across C C (CC) Width across M1 M1 (M1M1) Width across M2 M2 (M2M2) M1 length (M1L) M1 width (M1W) M2 length (M2L) M2 width (M2W) Mandible length (MDL) Mandibular toothrow length (MDTR) Mandibular height at base of inner incisors (MANH) Coronoid process height (CPH) 0.92 Hall 1981). The presence of 1 or 2 pairs of lower incisors is used to separate the genus into 2 subgenera (Corvira with 1 pair, although occasionally 3 or 4 lower incisors can be found, and Sturnira with 2 pairs Davis 1980). The presence or absence of lingual cusps (entoconid and metaconid) on the lower molars (serrate or smooth) led to the separation of the genus into 2 groups (Iudica 2000). In an attempt to better understand the genus and to devise more detailed criteria for identification of species of Sturnira, we visited the Colección de Vertebrados de la Universidad de Los Andes (CVULA) in Mérida, Venezuela, in 1997 and the collection of mammals of the American Museum of Natural History (AMNH) in New York in At these museums, we examined 4 specimens of Sturnira that exhibit notable dental differences from previously described species, which led us to consider these specimens as representing an undescribed species. MATERIALS AND METHODS External, cranial, and dental measurements were recorded for 44 male specimens of 4 Sturnira species: S. erythromos (n ¼ 17), S. sp. novo (n ¼ 3), S. ludovici (n ¼ 16), and S. hondurensis (n ¼ 8). We confined the statistical analysis to adult males because 3 of the specimens in question are adult males and because sexual dimorphism is known in several species of Sturnira. Specimens included in our comparisons (Appendix I) were from the CVULA, the Estación Biológica de Rancho Grande (EBRG), the AMNH, and the National Museum of Natural History (USNM). All measurements follow Hall (1981) and Sánchez- Hernández and Romero-Almaraz (1995). Measurements obtained from dried specimens were described as dry. Measurements of external, cranial, and dental characters were taken with digital calipers to the nearest 0.05 mm. Characters and abbreviations are listed in Table 1. Statistical analysis was based on adult specimens that were aged based on ossification of the skull and epiphyses of the phalanges of the wings. A multivariate comparison was made with the 29 morphometric characters (Table 1). Each character was standardized (i.e., X ¼ 0 and SD ¼ 1), a character product-moment correlation matrix was obtained, and principal components were computed by using version 2.1T of the computer program NTSYSpc (Rohlf 2000). Specimens were projected onto the resulting principal components based on standardized data. The Tukey Kramer procedure (Sokal and Rohlf 1995) was used to conduct multiple comparisons among means of projections on principal component I for the 4 species, as well as of the individual measurements. This procedure is appropriate for unequal sample sizes and employs the studentized range and average sample sizes in the standard errors. The method, although conservative, provides evaluations closer to the intended significance level than some other multiple-comparison methods, which tend to be too conservative (Sokal and Rohlf 1995). RESULTS Statistical comparisons. In the principal components analysis based on 29 characters and 44 male specimens of S. erythromos, S. sp. novo, S. ludovici, and S. hondurensis, most of the variation was summarized on component I (eigenvalue 22.77, 78.5% of total character variance). Other components summarized relatively little of the total character variance and none reflected interspecific differences. All but 1 of the characters (M2 length) had high loadings on principal component I (Table 1), indicating that the component represents a general size vector (Marcus 1990). Projections of specimens of the 4 species on principal component I are shown in Fig. 1. The Tukey Kramer procedure for multiple comparisons among means showed all differences among species in component I to be highly significant (P, 0.01) except for the difference between S. ludovici and S. hondurensis (P. 0.05). Thus, in overall size the males of the new species are notably larger than those of S. erythromos, but smaller than males of either S. hondurensis or S. ludovici (Fig. 1). In addition to the general size difference, clear differences in individual morphometric characters also were evident (Table 2). For 21 of the 29 characters, males of the new species were significantly different from those of S. erythromos. Furthermore, the new species was significantly different in 13 characters from S. ludovici and in 14 characters from S. hondurensis. Separation of the new species from these 3 other species was notably greater than that between S. ludovici and S. hondurensis, which were significantly different in only 1 of the 29 characters (i.e., character 14, SKD; Table 2). DISCUSSION Sturnira sorianoi, new species Fig. 2; Table 2 Holotype. Number I-2498 from the CVULA. The specimen is an adult male (inguinal testes) prepared as skin and

3 868 JOURNAL OF MAMMALOGY Vol. 86, No. 5 skull, in good condition, collected on 15 January 1983, with field number 1461 by Pascual Soriano. No external measurements are found on the original label. Type locality. Venezuela, Asentamiento Monterrey, 8 km NNE from Mérida, ME. El Valle, 2,300 m, 88379N, W. Distribution. Known only in Venezuela and Bolivia. Etymology. The name sorianoi is in honor of our friend Dr. Pascual J. Soriano, who collected some of the specimens and who has contributed in significant ways to the understanding of Venezuelan bats. Diagnosis. Medium-sized bat with pale gray brown to gray blackish dorsal coloration; ventral coloration pale brown to gray brown. Coloration of epaulettes is blended whitish to cinnamon reddish. Nose leaf and ears are small. Maxillary toothrow has a horseshoe shape. Upper incisors are bicuspid, lower incisors trilobed, and 1st and 2nd lower molars smooth (Fig. 2). Description. Medium size, dorsal coloration pale gray and venter pale brown; face blackish; without distinguishable epaulettes on shoulder and neck, but these areas are paler than rest of body. Dorsal hairs are 8 mm in length, abundant and silky, tetracolored with whitish base, blackish subbasal band, brown and whitish subterminal band, and pale gray or brown tip. Ventral coloration is pale brown, with tricolor or bicolored hairs, with white or black base, blackish intermediate band, and pale brown tip. Wing membrane, nose leaf, and ears are blackish. Nose leaf is triangular, 7 mm high and 4 mm wide at base. Ears measure 14.5 mm in the dry specimen. Internal edges of legs have hairs 4 5 mm long. External membrane border has abundant hair 2 mm long. Skull is long and robust, with square and short face. Rostrum is flat with a slope of about 458 and a light concave depression at center. Braincase is bowed outward. Orbital border is oval. Supraorbital border, and sagittal and occipital crests are well developed. Zygomatic arches are thin and bowed outward, with middle part almost straight. Palate is dome-shaped, short and wide in its posterior border, ending in short V-shaped cavity. Maxillary toothrow is horseshoe-shaped. Upper incisors are 1.4 mm long and bicuspid, with internal lobe larger than external lobe. Canines are 3.1 mm in length. Mandibular coronoid process has triangular shape, and slope of posterior border is abrupt. Height of mandible from base of lower incisors is 2.4 mm and from coronoid process is 5.4 mm. Lower incisors are trilobed, and cingulum of canine covers both crown and posterior part of external incisors. First and 2nd lower molars are smooth (Fig. 2). Measurements ( a, dry; b, broken). Foot length a, 14 mm; ear length a, 16 mm; tibia length, 15.9 mm; forearm length b, 41.4 mm; thumb length, 13.0 mm; metacarpal II length, 34.7 mm; metacarpal IV length, 38.0 mm; phalanx I length, 14.8 mm; phalanx II length, 14.1 mm; metacarpal V length, 42.6 mm; phalanx I length, 10.3 mm; phalanx II length, 12.6 mm; nose-leaf length a, 7.1 mm; nose-leaf width a, 4.3 mm; and weight, 18.5 g. Additional measurements are given in Table 2. Paratypes. Three specimens, 1 female and 2 males. CVULA number I-1398 is a young female collected on 31 October 1980, in Monte Zerma, 6 km N of Mérida, Venezuela, FIG. 1. Bar chart summarizing projections of 44 male specimens of Sturnira erythromos, S. sp. novo, S. ludovici, and S. hondurensis onto principal component I resulting from analysis of 29 morphometric characters. For the new species, holotype (CVULA I-2498) is leftmost specimen in panel, AMNH is in middle, and AMNH is on right. 2,000 m, 88379N, W, by Pascual Soriano and Jesus Molinari, with field number 1374 by P. Soriano. Two specimens from the AMNH originally were identified as S. oporaphilum. Number is a male collected by C. G. Schmitt, number 2948, on 11 January 1984, in the Department of Santa Cruz, Valle Grande Province, 13 km by road NW of Masicurs, Bolivia, 1,000 m (188469S, W). Number is a male collected by B. R. Riddle, with field number 2215, on 19 August 1985, in Santa Cruz, 4.5 km N and 1.5 km E of Cerro Amborro, Río P. Tasama, Bolivia, 620 m (178459S, W). Characteristics of paratypes. Paratypes share most characteristics of the holotype. Number I-1398 is smaller than the type. Dorsum is pale brown mixed with dark brown. Dorsal hairs are tetracolored, with a whitish basal band followed by a narrow black subbasal band, a brown subterminal band, and a dark brown tip. Ventral coloration is darker, and tricolored

4 October 2005 SÁNCHEZ-HERNÁNDEZ ET AL. NEW SPECIES OF STURNIRA 869 TABLE 2. Measurements of 29 characters of male Sturnira erythromos, S. sp. novo, S. ludovici, and S. hondurensis shown as mean 6 SD (n, range), as well as measurements for type specimen of S. sp. novo. Superscripts denote lack of statistical significance of comparisons based on Tukey-Kramer procedure (Sokal and Rohlf 1995). If for a given character superscript a is shared by S. erythromos and a species to its right in the table, the two are not significantly different statistically (P. 0.05); species not sharing superscript a with S. erythromos are statistically different (P, 0.05) from it. If superscript b is shared by S. sp. novo and a species to its right, the two are not different; those to the right not having superscript b are different from S. sp. novo. Likewise, when superscript c is shared by S. ludovici and S. hondurensis they are not different; if they do not share superscript c they are different. In general, means for species change from low to high from left to right, but there are exceptions (e.g., character 19, CC). All measurements in millimeters. Value for FL in type obtained from broken element. Character Sturnira erythromos sp. novo ludovici hondurensis Type specimen 1 FL (17, ) (2, ) (16, ) c (8, ) c ML (17, ) a (2, ) a (16, ) c (8, ) c F1L (17, ) a (2, ) ab (16, ) bc (8, ) bc F2L (17, ) (2, ) b (16, ) bc (8, ) c TL (17, ) (2, ) b (16, ) bc (7, ) bc GSL (17, ) (3, ) (16, ) c (8, ) c CIL (17, ) (3, ) (16, ) c (8, ) c CBL (17, ) (3, ) (16, ) c (8, ) c CCL (17, ) (3, ) (16, ) c (8, ) c BCB (17, ) (3, ) (16, ) c (8, ) c MST (17, ) (3, ) (16, ) c (8, ) c ZB (16, ) (3, ) b (16, ) bc (8, ) bc SKH (11, ) (2, ) b (15, ) bc (6, ) c SKD (11, ) (2, ) b (16, ) b (6, ) b INB (17, ) (3, ) b (16, ) bc (8, ) bc INC (17, ) a (3, ) ab (16, ) bc (8, ) bc RB (17, ) (3, ) b (16, ) bc (8, ) bc MTR (16, ) (3, ) (16, ) c (8, ) c CC (13, ) a (2, ) ab (16, ) c (8, ) bc M1M (17, ) (3, ) b (16, ) bc (8, ) bc M2M (16, ) (3, ) b (16, ) bc (8, ) bc M1L (17, ) a (3, ) ab (16, ) abc (8, ) bc M1W (17, ) a (3, ) ab (16, ) bc (8, ) bc M2L (16, ) a (3, ) ab (16, ) abc (8, ) abc M2W (16, ) (3, ) b (16, ) bc (8, ) bc MDL (17, ) (3, ) (16, ) c (8, ) c MDTR (16, ) (3, ) (16, ) c (7, ) c MANH (13, ) a (2, ) a (16, ) c (8, ) c CPH (13, ) (2, ) b (16, ) bc (8, ) bc 5.42 hairs have a white basal band, a gray intermediate band, and pale brown tips. Posterior border of uropatagium and internal border of the legs have no hair because of the poor preparation of the specimen. Skull is smaller than the type, and the rostrum is somewhat straight. Skin is somewhat damaged, with hair lost in the internal border of legs. The teeth of the maxilla are incomplete, but a single bicuspid internal incisor is in place on the left side. It also has the canines and premolars, and on the right side there are only 2 molars. All teeth are present in the mandible, with trilobed incisors and smooth 1st and 2nd molars. Specimens AMNH and have abundant long and silky hair; the hair is tricolored on the back and belly. The posterior border of the uropatagium is hairy (4 6 mm), and the hair on the external border of the wing membrane at the tibia also is abundant, measuring 2.6 mm. Measurements of paratypes ( a, dry). Specimens I-1398 and , respectively: total length, 57, 69 mm; foot length, 13, 14 mm; ear length, 15, 18 mm; tibia length, 12.7, 18.5 mm; forearm length, 42.8, 43.7 mm; thumb length, 12.1, mm; metacarpal II length, 34.1, mm; metacarpal IV length, 39.8, mm; phalanx I length, 15.2, mm; phalanx II length, 13.1, mm; metacarpal V length, 42.5, mm; phalanx I length, 10.0, mm; phalanx II length, 10.5, mm; nose-leaf length a, 6.0, mm; nose-leaf width a, 4.0, mm; weight, 17.0, 21.8 g. Additional measurements are given in Table 2. Comparisons. Sturnira sorianoi is similar to S. oporaphilum, S. ludovici, S. hondurensis, S. bogotensis, S. bidens, and S. mordax in having smooth molars, but different from these species because it has a tricuspid, rather than bicuspid, lower incisor. In addition, S. sorianoi differs from these species because of its different skull shape and size. In general, the lengths of forearm and skull in S. sorianoi are longer (FL, mm; GSL, mm) compared to S. erythromos (FL, mm, GSL, mm), but shorter than in other similar species S. hondurensis (FL, mm; GLS, mm), S. ludovici (FL, mm; GLS, mm), and S. oporaphilum (FL. 44 mm Barquez et al. 1993). Although S. magna has smooth molars and trilobed lower incisors as in S. sorianoi, the former has a longer forearm ( mm compared to

5 870 JOURNAL OF MAMMALOGY Vol. 86, No. 5 FIG. 2. Lateral, dorsal, and ventral views of cranium and lateral view of mandible of type specimen of Sturnira sorianoi (CVULA I-2498), as well as frontal view of upper and lower incisors and lateral view of lower molars. mm in S. sorianoi), as well as some differences in the skull shape, and unicuspid instead of bicuspid upper inner incisors as in S. sorianoi (Davis 1980; de la Torre 1966; Tamsitt and Hauser 1985; Tamsitt and Valdivieso 1986). Because of its smooth lower molars, S. sorianoi is different from S. aratathomasi (Peterson and Tamsitt 1968; Soriano and Molinari 1987), S. l. lilium (Carter and Dolan 1978; Gannon et al. 1989; Geoffroy Saint Hilaire 1810; Sánchez-Hernández and Romero-Almaraz 2003), S. l. parvidens (Goldman 1917; Sánchez-Hernández and Romero-Almaraz 2003), S. luisi (Davis 1980), S. thomasi (de la Torre and Schwartz 1966), S. tildae (de la Torre 1959), and S. mistratensis (Contreras- Vega and Cadena 2001), which have well-defined lingual cusps of m1 and m2, with entoconid and metaconid separated by a vertical notch, so that each molar is serrated. In addition, other size, skull-shape, and dental characteristics differentiate S. sorianoi from a number of these species. S. sorianoi also differs from S. nana (Gardner and O Neill 1971) and S. bidens in that these have 2 pairs of lower incisors instead of a single pair. Sturnira sorianoi is a medium-sized bat with pale gray brown to gray blackish dorsal coloration; ventral coloration is pale brown to gray brown. This species has a unique combination of dental characters (bicuspid upper incisors, trilobed lower incisors, and smooth 1st and 2nd lower molars). The species with the most similar dentition is S. magna, which has smooth molars and trilobed lower incisors. However, S. magna has a longer forearm ( mm compared to mm for S. sorianoi), as well as some differences in the skull shape, and unicuspid instead of bicuspid upper inner incisors as found in S. sorianoi (Davis 1980; de la Torre 1966; Tamsitt and Hauser 1985; Tamsitt and Valdivieso 1986). Principal components analysis, based on 29 characters of male S. sorianoi and 3 of the more similar species, S. erythromos, S. ludovici, and S. hondurensis, provides clear evidence of a separation of S. sorianoi from the others based on size. In addition, the specimens of S. sorianoi statistically are separable from those of the other 3 species on the basis of numerous individual measurements. In overall size males of S. sorianoi are larger than those of S. erythromos, but smaller than males of either S. hondurensis or S. ludovici. Acquisition of additional specimens of S. sorianoi and a complete review of the genus Sturnira are needed for further taxonomic clarification. Albuja (1999) and Pacheco and Patterson (1991) have mentioned the existence of Sturnira sp. A, and Albuja (1999) the existence of Sturnira sp. B; however, these 2 species remain undescribed. Sturnira sp. A, a specimen from Ecuador, Chimborazo, Pallatanga, EPN E- 6722, has the following main characteristics: upper incisors, procumbent bicuspid, 4 triangular-shaped and trilobed cuspids, lower incisors, smooth lower molars, and a 3rd lower molar. This specimen shares all of these characteristics with S. sorianoi; however, it does not have a zygomatic arch, and has spaces between the lower and upper premolars and molars, apparently produced by the reduction of the paraconed(id). In addition, the lengths of forearm and skull in S. sorianoi are shorter (FL, mm; GSL, mm versus FL, 51 mm; GSL, 26 mm) than in Sturnira sp. A. Thus, Sturnira sp. A is probably a different species from S. sorianoi.

6 October 2005 SÁNCHEZ-HERNÁNDEZ ET AL. NEW SPECIES OF STURNIRA 871 Sturnira sp. B (Albuja 1999) is similar in size to S. sorianoi (FL, 42.9 mm; GLS, 22 mm in Sturnira sp. B and FL, mm; GSL, mm in S. sorianoi). However, Sturnira sp. B has serrated molars, which suggests that the 2 species are different. Natural history. The localities of Monterrey and Monte Zerma, where specimens of S. sorianoi were obtained, belong to the Cordillera de Mérida in the Venezuelan Andes. The vegetation is cloud forests with at least 3 strata. The 1st stratum consists of trees between 25 and 35 m in height, the intermediate stratum has trees that range from 5 to 10 m, and the 3rd stratum is herbaceous plants of m. In addition, this forest is characterized by an irregular canopy and a richness of tree species that varies from 40 to 60 species per hectare, mainly evergreen, which support a great variety of epiphytes (Bromeliaceae, Orchidaceae, Araceae, Piperaceae, and pteridophites). The most characteristic feature of the shrubby and herbaceous strata is the presence of arborecent ferns from the family Cyatheaceae. Under favorable climatic conditions these forests occupy the altitudinal belt between 1,700 and 3,000 m. Annual rainfall is 1,000 2,600 mm, distributed in a bimodal pattern with peaks in May and September. The driest season is from December to February. Annual mean temperatures oscillate between 10 and 188C, with day night variations from 6 to 88C. Fog is frequent in the afternoon, which reduces both solar radiation and the temperature, conferring high levels of relative humidity to the environment (Soriano et al. 1999). RESUMEN Los murciélagos de charreteras (género Sturnira) se encuentran en Latinoamérica y han sido relativamente poco estudiados desde el punto de vista taxonómico. Cuatro especímenes de Sturnira de Bolivia y Venezuela, poseen características morfológicas, dentarias y craneales diferentes de las de especies actualmente reconocidas. Se describen las características externas, dentarias y craneales de la especie nueva y se presenta una comparación mutivariada realizada con 29 caracteres morfométricos de 44 especímenes machos de Sturnira erythromos, S. ludovici, S. hondurensis y la especie nueva. En el análisis de componentes principales, la mayoría de la variación fue sumarizada por el componente I (78.5% del total de la varianza de los caracteres), que representa un vector de tamaño. Una prueba de comparación múltiple entre las medias, mostró que todas las especies son significativamente diferentes (P, 0.01) en el componente I, excepto S. ludovici y S. hondurensis Para los 29 caracteres individuales la especie nueva fue estadísticamente diferente de S. erythromos en 21, S. ludovici en 13 y S. hondurensis en 14. La especie nueva tiene una combinación única de características dentarias, como son los incisivos superiores bicúspides, los incisivos inferiores trilobados y los molares inferiores 1 y 2, lisos. ACKNOWLEDGMENTS We express our gratitude to A. Gardner, K. Rodríguez, A. Guerrero- Enríquez, V. H. Reynoso, and R. D. Owen for constructive comments on an earlier draft of the manuscript. The curators of the museums mentioned in this work allowed us to review the type specimens and other specimens of Sturnira: P. J. Soriano of the CVULA, Javier Sánchez of the EBRG, S. Anderson and G. Musser of the AMNH, and D. E. Wilson of the USNM. Finally, we would like to thank the authorities of the Dirección General del Personal Académico and the Instituto de Biología, Universidad Nacional Autónoma de México, for their support of this project. LITERATURE CITED ALBUJA, L Murciélagos del Ecuador. 2nd ed. Cicetrónic Compañía Limitada Offset, Quito, Ecuador. BARQUEZ, R. M., N. P. GIANNINI, AND M. A. MARES Guide to the bats of Argentina (Guía de los murciélagos de Argentina). Oklahoma Museum of Natural History, Norman. CARTER, D. C., AND P. G. DOLAN Catalogue of type specimens of neotropical bats in selected European museums. Special Publication, The Museum, Texas Tech University 15: CONTRERAS-VEGA, M., AND A. CADENA Una nueva especie del género Sturnira (Chiroptera: Phyllostomidae) de los Andes Colombianos. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 26: DAVIS, W. B New Sturnira (Chiroptera: Phyllostomidae) from Central and South America, with key to currently recognized species. Occasional Papers, The Museum, Texas Tech University 70:1 5. DE LA TORRE, L A new species of bat of the genus Sturnira (Phyllostomidae) from the island of Trinidad, West Indies. Chicago Academy of Science, Natural History Miscellanea 166:1 6. DE LA TORRE, L The evolution, variation, and systematics of the neotropical bats of the genus Sturnira. Ph.D. dissertation, University of Illinois, Champaign. DE LA TORRE, L New bats of the genus Sturnira (Phyllostomidae) from the Amazonian lowlands of Peru and the Windward Islands, West Indies. Proceedings of the Biological Society of Washington 79: DE LA TORRE, L., AND A. SCHWARTZ New species of Sturnira (Chiroptera: Phyllostomidae) from the islands of Guadeloupe and Saint Vincent, Lesser Antilles. Proceedings of the Biological Society of Washington 79: GANNON, M. R., M. R. WILLING, AND J. K. JONES, JR Sturnira lilium. Mammalian Species 333:1 4. GARDNER, A. L., AND J. P. O NEILL A new species of Sturnira (Chiroptera: Phyllostomidae) from Peru. Occasional Papers of the Museum of Zoology, Louisiana State University, Baton Rouge 42:1 7. GEOFFROY SAINT HILAIRE, É Sur les phyllostomes et les megadermes, deux genres de la famille des chauve-souris. Annales du Muséum d Histoire Naturelle, Paris 15: GIANNINI, N. P., AND R. M. BARQUEZ Sturnira erythromos. Mammalian Species 729:1 5. GOLDMAN, E. A New mammals from North and Middle America. Proceedings of the Biological Society of Washington 30: GOODWIN, G. G Three new bats from Honduras and the first record of Enchisthenes harti (Thomas) for North America. American Museum Novitates 1075:1 3. GOODWIN, G. G Mammals of Honduras. Bulletin of the American Museum of Natural History 79: GOODWIN, G. G Mammals of Costa Rica. Bulletin of the American Museum of Natural History 87:

7 872 JOURNAL OF MAMMALOGY Vol. 86, No. 5 GRAY, J. E Descriptions of some new genera and fifty unrecorded species of Mammalia. Annals and Magazine of Natural History 10: HALL, E. R The mammals of North America. 2nd ed. Vol. 1. John Wiley and Sons, New York. HERSHKOVITZ, P Mammals of northern Colombia, preliminary report no. 5: bats (Chiroptera). Proceedings of the United States National Museum 99: IUDICA, C. A Systematic revision of the neotropical fruit bats of the genus Sturnira: a molecular and morphological approach. Ph.D. dissertation, University of Florida, Miami. JONES, J. K., JR., AND D. C. CARTER Annotated checklist, with keys to subfamilies and genera. Part I. Pp in Biology of bats of the New World family Phyllostomatidae (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Special Publications, The Museum, Texas Tech University, Lubbock 10: KOOPMAN, K. F Zoogeography of Peruvian bats with special emphasis on the role of the Andes. American Museum Novitates 2651:1 33. KOOPMAN, K. F Order Chiroptera. Pp in Mammal species of the world: a taxonomic and geographic reference (D. E. Wilson and D. M. Reeder, eds.). Smithsonian Institution Press, Washington, D.C. MARCUS, L. F Traditional morphometrics. Pp in Proceedings of the Michigan morphometrics workshop (F. J. Rohlf and F. L. Bookstein, eds.). Special Publication, University of Michigan Museum of Zoology 2: PACHECO, V., AND B. D. PATTERSON Phylogenetic relationships of the New World bat genus Sturnira (Chiroptera: Phyllostomidae). Pp 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: PETERSON, R. L., AND J. R. TAMSITT A new species of bat of the genus Sturnira (family Phyllostomatidae) from northwestern South America. Royal Ontario Museum, Life Sciences Occasional Papers 12:1 8. ROHLF, F. J NTSYSpc: user guide. Version 2.1. Exeter Software, Setauket, New York. SÁNCHEZ-HERNÁNDEZ, C., AND M. L. ROMERO-ALMARAZ Los Murciélagos de Tabasco y Campeche. Una propuesta para su conservación. Cuadernos del Instituto de Biología, Universidad Nacional Autónoma de México, México 24: SÁNCHEZ-HERNÁNDEZ, C., AND M. L. ROMERO-ALMARAZ Redescription of Sturnira lilium lilium and Sturnira lilium parvidens (Mammalia: Chiroptera: Phyllostomidae). Southwestern Naturalist 48: SOKAL, R. R., AND F. J. ROHLF Biometry. 3rd ed. W. H. Freeman and Company, New York. SORIANO, P., AND J. MOLINARI Sturnira aratathomasi. Mammalian Species 284:1 3. SORIANO, P. J., A. D. PASCUAL, J.OCHOA, AND M. AGUILERA Biogeographic análisis of the mammal communities in the Venezuelan Andes. Interciencia 24: TAMSITT, J. R., AND C. HAUSER Sturnira magna. Mammalian Species 240:1 4. TAMSITT, J. R., AND D. VALDIVIESO Variación morfométrica en el murciélago Sturnira magna (Chiroptera: Phyllostomidae). Caldasia 15: TIMM, R. M., AND R. K. LAVAL A field key to the bats of Costa Rica. Occasional Publication Series, Center of Latin American Studies, University of Kansas, Lawrence 22:1 30. TIMM, R. M., D. E. WILSON, B.L.CLAUSON, R.K.LAVAL, AND C. S. VAUGHAN Mammals of the La Selva Braulio Carrillo Complex, Costa Rica. North American Fauna 75: VILLALOBOS, F., AND A. VALERIO The phylogenetic relationships of the bat genus Sturnira Gray, 1842 (Chiroptera: Phyllostomidae). Mammalian Biology 67: Submitted 28 April Accepted 7 April Associate Editor was Robert D. Bradley. APPENDIX I Specimens examined. The 44 specimens included in the morphometric analysis are listed below. Collection acronyms are as follows: AMNH, American Museum of Natural History; CVULA, Colección de Vertebrados, Universidad de Los Andes, Venezuela; EBRG, Estación Biológica de Rancho Grande; USNM, National Museum of Natural History. Sturnira erythromos (n ¼ 17): Venezuela, CVULA 1328, 1431, 1443, 1447, 2097, 2098, 2148, 2483, 3008, 3141, 5892; EBRG 10056, 10058; USNM ; Bolivia, AMNH , , S. sp. novo (n ¼ 3): Venezuela, CVULA 2498; Bolivia, AMNH , S. ludovici (n ¼ 16): Venezuela, CVULA 2099, 2136, 3109; EBRG 8076, 14435, 14440, 14441, 14454, 14467, 14468, 14470, 14474, 14475, 14482, 14487; Ecuador, AMNH S. hondurensis (n ¼ 8): Venezuela, CVULA 2759, 2993, 3108, 3530; EBRG 57, 1777, 14431,