Journal of Mammalogy, 93(2):605–614, 2012
A new species of Lophostoma d’Orbigny, 1836 (Chiroptera: Phyllostomidae) from Panama PAU´L M. VELAZCO*
AND
ALFRED L. GARDNER
Department of Mammalogy, American Museum of Natural History, New York, NY 10024, USA (PMV) United States Geological Survey Patuxent Wildlife Research Center, National Museum of Natural History MRC-111, Smithsonian Institution, Washington, DC 20013-7012, USA (ALG) * Correspondent:
[email protected] We report the discovery of a new species of Lophostoma from Panama, which we name L. kalkoae. This new species resembles L. carrikeri and L. yasuni in possessing a white venter, but is distinguishable from both by external and cranial characteristics. The new species is similar in size to L. carrikeri and L. schulzi. Lophostoma sp. nov. can be most easily recognized by its combination of white venter, postauricular patches connected by a thin line of pale hair to the white fur on the chest, elongated clitoris and swollen labia, less strongly developed lateral projection of mastoid processes, well-marked indentation on the lingual cingulum of the upper canine, well-developed P3, well-developed posterior lingual cusp on the cingulum of P4, and parastyle absent on M1 and M2. We present a dichotomous key for the genus Lophostoma and a map showing all the localities where white-bellied Lophostoma have been recorded. Key words:
dichotomous key, Lophostoma, masculinization, Panama, Phyllostomidae, Phyllostominae, taxonomy
E 2012 American Society of Mammalogists DOI: 10.1644/11-MAMM-A-217.1
The Neotropical bat genus Lophostoma d’Orbigny, 1836 (type species Lophostoma silvicolum d’Orbigny, 1836, by monotypy), is distributed from southern Mexico to central Paraguay (Simmons 2005; Williams and Genoways 2008). Currently, Lophostoma includes 7 recognized species of small to medium-sized bats (forearm 30–60 mm, greatest length of skull 18–31 mm—Velazco and Cadenillas 2011): L. silvicolum d’Orbigny, 1836; L. brasiliense Peters, 1867; L. carrikeri (Allen, 1910); L. evotis (Davis and Carter, 1978); L. occidentalis (Davis and Carter, 1978); L. schulzi (Genoways and Williams, 1980); and L. yasuni Fonseca and Pinto, 2004. Treated as a synonym of Tonatia Gray, 1827, for most of the 20th century, Lophostoma was resurrected by Lee et al. (2002). With few exceptions (e.g., ‘‘Vampyrini’’—Wetterer et al. 2000), Lophostoma has been allied within the Phyllostomini along with the genera Lonchorhina, Macrophyllum, Mimon, and Phyllostomus, and either as a synonym of Tonatia (Baker et al. 1989, 2000) or as a separate genus in association with Phylloderma, Phyllostomus, Tonatia, and Trachops (Datzmann et al. 2010). Two species of Lophostoma are characterized by having white fur on the chest and abdomen: L. yasuni, known only from its type locality in eastern Ecuador (Fonseca and Pinto 2004), and L. carrikeri, described by Allen (1910) from specimens from central Venezuela, and now known from
Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Peru, and Bolivia (Genoways and Williams 1980, 1984; Gribel and Taddei 1989; Lim 2009; Lim et al. 1999, 2005; McCarthy et al. 1983; McCarthy and Handley 1988; Vizotto et al. 1980; Williams and Genoways 2008; Fig. 1). Herein, we describe a new species of white-ventered Lophostoma from Panama. We analyzed morphometric and morphological data to distinguished this new taxon from other species of the genus, and provide a map (Fig. 1) of all the localities where white-ventered Lophostoma have been collected.
MATERIALS AND METHODS External and osteological characters examined were based on, but not restricted to, those defined by Baker et al. (2004), Fonseca and Pinto (2004), and Velazco and Cadenillas (2011). The dental homology nomenclature for the premolars follows that of Miller (1907): 1st upper premolar (P3), 2nd upper premolar (P4), 1st lower premolar (p2), 2nd lower premolar (p3), and 3rd lower premolar (p4).
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FIG. 1.—Map of southern Central America and northern South America, showing the type localities of Lophostoma kalkoae sp. nov. (star) and L. yasuni (square), and all reported localities for L. carrikeri (triangles).
For the morphometric analyses, we examined 288 specimens of adult Lophostoma (152 males and 136 females) representing all known species of Lophostoma (see Appendix I). We used a digital caliper to take 8 external and 16 craniodental measurements to the nearest 0.01 mm on each specimen (Fig. 2). Descriptive statistics (mean and range) were calculated for all samples for selected measurements. Definitions for craniodental, mandibular, and external measurements are listed in Table 1. All craniodental and 2 external (FA and MET–III) measurements were log-transformed to achieve normalization for statistical analyses. We evaluated differences among morphological groups by principal component analysis using a covariance matrix. Principal component (PC) scores were plotted to show relationships between species groups in morphospace. Principal component analysis was performed using IBM SPSS Statistics for Windows, version 19 (IBM, Armonk, New York). We constructed a dichotomous identification key for all recognized species of Lophostoma. Photographs of the holotype and paratype of Lophostoma sp. nov. (32 images: M54224–M54255) and of the female genitalia of L. schulzi (AMNH 257128 [M54312 and M54313]; AMNH 267420 [M54314 and M54315]; and AMNH 267421 [M54316]) are available at Morphobank (http://morphobank.org/permalink/?P421). We reference some of these images throughout our description (the image reference numbers begin with the letter M). We examined specimens from the following institutional collections: American Museum of Natural History, New York (AMNH); Carnegie Museum of Natural History, Pittsburgh (CM); Field Museum of Natural History, Chicago (FMNH); Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogota´ (ICN); Instituto Nacional de Pesquisas da Amazoˆnia, Manaus (INPA); Museum of Natural Science, Louisiana State University, Baton Rouge (LSUMZ); Museu
FIG. 2.—Dorsal and ventral views of the cranium and lateral view of the cranium and mandible illustrating most of the measurements used in the morphometric analyses. For definitions of abbreviations see Table 1.
Paraense Emı´lio Goeldi, Bele´m (MPEG); Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima (MUSM); Museo de Vertebrados de la Universidad de Panama´, Panama´ (MVUP); Museo de Zoologı´a, Pontificia Universidad Cato´lica del Ecuador, Quito (QCAZ); Royal Ontario Museum, Toronto (ROM); Museum of Texas Tech University, Lubbock (TTU); National Museum of Natural History, Smithsonian Institution, Washington (USNM).
RESULTS Family Phyllostomidae Gray, 1825 Subfamily Phyllostominae Gray, 1825 Genus Lophostoma d’Orbigny, 1836 Lophostoma kalkoae, new species Kalko’s Round-eared Bat Fig. 3, M54224–M54255
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TABLE 1.—Definition of craniodental, mandibular, and external measurements used in the present study. Measurement Forearm length (FA) Metacarpal III length (MET–III) Metacarpal IV length (MET–IV) Metacarpal V length (MET–V) Tail length (TL) Hind-foot length (HF) Tibia length (TiL) Calcar length (CL) Greatest length of skull (GLS) Condyloincisive length (CIL) Condylocanine length (CCL) Braincase breadth (BB) Zygomatic breadth (ZB) Postorbital breadth (PB) Palatal width at canines (C–C) Mastoid width (MSTW) Mastoid process width (MPW) Palatal length (PL) Maxillary toothrow length (MTRL) Molariform toothrow length (MLTRL) Width at M2 (M2–M2) Dentary length (DENL) Mandibular toothrow length (MANDL) Coronoid height (COH)
Measurement definition Distance from the elbow (tip of the olecranon process) to the wrist (including the carpals). This measurement is made with the wing at least partially folded. Distance from the joint of the wrist (carpal bones) with the 3rd metacarpal to the metacarpophalangeal joint of 3rd digit. Distance from the joint of the wrist (carpal bones) with the 4th metacarpal to the metacarpophalangeal joint of 4th digit. Distance from the joint of the wrist (carpal bones) with the 5th metacarpal to the metacarpophalangeal joint of 5th digit. Distance from dorsal flexure at base of the tail to the tip of the last caudal vertebra. From the proximal edge of the base of the calcar to the tip of the claw of the longest toe. From the proximal end of the tibia to the distal base of the calcar. From the joint with the ankle to the calcar tip. Greatest distance from the occiput to the anteriormost point on the premaxilla (including the incisors). Distance between a line connecting the posteriormost margins of the occipital condyles and the anteriormost point on the upper incisors. Distance between a line connecting the posteriormost margins of the occipital condyles and a line connecting the anteriormost surfaces of the upper canines. Greatest breadth of the globular part of the braincase, excluding mastoid and paraoccipital processes. Greatest breadth across the zygomatic arches. Least breadth at the postorbital constriction. Least width across palate between lingual margins of the alveoli of upper canines. Least breadth across skull immediately behind jugal base of zygomatic arches. Greatest breadth across skull, including mastoid processes. Distance from the posterior palatal notch to the anteriormost border of the incisive alveoli. Distance from the anteriormost surface of the upper canine to the posteriormost surface of the crown of M3. Distance from the anteriormost surface of P3 to the posteriormost surface of the crown of M3. Greatest width of palate across labial margins of the alveoli of M2s. Distance from midpoint of condyle to the anteriormost point of the dentary. Distance from the anteriormost surface of the lower canine to the posteriormost surface of m3. Perpendicular height from the ventral margin of mandible to the tip of coronoid process.
Holotype.—An adult male (USNM 582249; Fig. 3, M54237–54246) caught on Pipeline Road, near the former Limbo Hunt Club (9u99500N, 79u459100W), Soberania National Park, Colo´n Province, Panama´, by Elisabeth K. V. Kalko (field number EKVK 118) on 11 October 1998. The holotype is preserved in alcohol with the skull removed and cleaned. The upper and lower dentition lost some of the enamel during initial preservation in unbuffered formalin. Paratype.—An adult pregnant female (EKVK 119 to be deposited in the MVUP, M54224–M54236, M54247– M54255) caught at the type locality by Elisabeth K. V. Kalko (original field number EKVK 119) on 11 October 1998. The paratype is preserved in alcohol with the skull removed and cleaned. As with the holotype, the dentition has lost some enamel due to decalcification in unbuffered formalin. Distribution.—The new species is currently known only from the type locality (Fig. 1). Etymology.—The name kalkoae is in honor of our late friend Dr. Elisabeth K. V. Kalko, a remarkable scientist who collected both specimens and who has contributed in significant ways to the understanding of bat behavior and ecology worldwide. Measurements.—External and craniodental measurements are presented in Table 2. Diagnosis.—Postauricular patches connected by a thin line of pale hairs to the white fur on the chest; ventrally the
proximal one-third of the forearm covered with long white hair; the clitoris elongated and labia swollen (M54253– M54255); hairs on rim of pinna are whitish; lateral projection of mastoid region of the skull less developed that in other Lophostoma; lingual cingulum of the upper canine strongly indented; P3 well developed; posterior lingual cusp on the cingulum of P4 well developed; M1 parastyle absent; M1 hypocone absent or weakly developed; M1 lingual cingulum present; M2 parastyle absent; M2 hypocone absent or weakly developed; M2 lingual cingulum present. Description and comparisons.—A medium-sized Lophostoma (FA 44.6–45.8 mm; GLS 23.2–23.8 mm; CCL 19.2– 19.6 mm; Table 2). All linear measurements of L. kalkoae show overlap with those of L. carrikeri and L. schulzi; L. kalkoae is larger than L. brasiliense, and smaller than L. evotis, L. occidentalis, L. silvicolum, and L. yasuni (Table 3). Dorsal pelage in all species of Lophostoma is dark brown and long; individual hairs tricolored with a short, white base (approximately 15% of the length of each hair), a long, dark brown subterminal band (approximately 80% of each hair), and a very short, pale to whitish terminal band. Gular fur is dark brown in L. kalkoae, L. brasiliense, L. evotis, L. schulzi, and L. silvicolum, but pale to whitish in L. carrikeri, L. occidentalis, and L. yasuni. L. kalkoae, L. occidentalis, and L. evotis have white to pale gray postauricular patches (absent in L. brasiliense, L. carrikeri, L. schulzi, L. silvicolum, and L. yasuni) that are connected by a
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FIG. 3.—Dorsal and ventral views of the cranium and lateral view of the cranium and mandible of Lophostoma kalkoae sp. nov. (USNM 582249, holotype). See Table 2 for measurements.
thin line of pale hairs to the white fur on the chest in L. kalkoae (pale line also present in L. evotis, which has a dark venter). The ventral fur is white across the chest, but restricted laterally over the abdomen by the pale brown fur of the sides of the body in L. kalkoae, L. carrikeri, and L. yasuni (venter pale brown in L. brasiliense, L. occidentalis, and L. schulzi; pale to dark brown chest in L. evotis, and L. silvicolum). Abdominal fur is white in L. kalkoae, L. carrikeri, and L. yasuni (pale brown in L. occidentalis, L. brasiliense, and L. schulzi; dark brown in L. evotis and L. silvicolum). Pinnae are sparsely haired and have a whitish rim; folds in the pinna are well marked; a band of skin across the head connects the internal bases of the pinnae. Uropatagium is essentially naked. The dorsal surface of the forearm appears naked in L. kalkoae, L. occidentalis, L. schulzi, and L. silvicolum (the proximal one-third is conspicuously covered with sparse, short hair in L. brasiliense, L. carrikeri, L. evotis, and L. yasuni); ventrally the proximal one-third of the forearm is covered with long, white hair in L. kalkoae (long, pale brown hair in L. brasiliense, L. carrikeri, L. evotis, and L. occidentalis; short, pale brown hair in L. schulzi and L. silvicolum). The dorsal surface of the forearm, digits, legs, and sagittal midline of nose leaf lack wartlike granulations in
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L. kalkoae, L. brasiliense, L. carrikeri, L. occidentalis, L. evotis, and L. silvicolum (present in L. schulzi). Dorsal surfaces of the feet are covered with short hair. Females have an elongated clitoris and swollen labia (dark red in the live specimen) in L. kalkoae and L. carrikeri (the clitoris is remarkably elongated and resembles the penis of males in L. schulzi; clitoris elongated but labia not swollen in L. brasiliense, L. occidentalis, and L. silvicolum). As is characteristic of other Lophostoma, metacarpal III is shorter than metacarpal V; there are 2 genal vibrissae, along with approximately 10 submental vibrissae on each side of the chin, and 2 interramal vibrissae implanted in a basal bulb. The skull of L. kalkoae has a slender rostrum with an accentuated postorbital constriction resembling that of L. brasiliense, L. carrikeri, and L. occidentalis (rostrum is robust in L. evotis, L. schulzi, L. silvicolum, and L. yasuni). Sagittal crest well developed in the female (EKVK 119) and weakly developed in the male (USNM 582249). Lateral development of the mastoid region is less in L. kalkoae and L. brasiliense; moderate in L. carrikeri; strongly developed and projecting well beyond sides of braincase in L. evotis, L. occidentalis, L. schulzi, L. silvicolum, and L. yasuni. The basioccipital is narrow; basisphenoid pits shallow and the midline septum comparatively wide; whereas in L. carrikeri basisphenoid pits are deeper and the septum conspicuously narrower. Upper central incisors (I1) are well developed and orthodont; outer upper incisors (I2) well developed and convergent in L. kalkoae (smaller in L. carrikeri). I2 is not displaced from occlusion toothrow. A deep indentation is present on the lingual cingulum of the upper canine in L. kalkoae and L. silvicolum (not as well marked in L. brasiliense, L. carrikeri, L. evotis, L. occidentalis, and L. schulzi). P3 is tall and well developed in L. kalkoae (shorter in L. carrikeri); labial cingulum of P3 weakly developed in L. kalkoae, L. carrikeri, L. evotis, L. occidentalis, L. schulzi, and L. silvicolum (absent in L. brasiliense). P4 is longer than wide in occlusal view in L. kalkoae, L. brasiliense, L. carrikeri, L. evotis, L. occidentalis, and L. silvicolum (length and width subequal in L. schulzi); posterior lingual cusp on the cingulum of P4 well developed in L. kalkoae (weakly developed in L. carrikeri). Metacone and paracone of M1 are subequal in height; postparacrista contacts premetacrista on labial aspect of M1, hence the trigon is closed labially in L. kalkoae, L. brasiliense, L. carrikeri, and L. schulzi (postparacrista does not contact premetacrista; therefore, trigon open labially in L. evotis, L. occidentalis, and L. silvicolum). M1 parastyle is absent in L. kalkoae and L. carrikeri (present in L. brasiliense, L. evotis, L. occidentalis, L. schulzi, and L. silvicolum); M1 hypocone either absent or weakly developed in L. kalkoae, L. carrikeri, and L. evotis (moderately to well developed in L. brasiliense, L. occidentalis, L. schulzi, and L. silvicolum); M1 lingual cingulum present in L. kalkoae and L. occidentalis (absent in L. brasiliense, L. carrikeri, L. evotis, L. schulzi, and L. silvicolum). Metacone and paracone of M2 are subequal in height; postparacrista contacts premetacrista on labial aspect of M2 closing the trigon labially in L. kalkoae,
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TABLE 2.—Measurements of type series of Lophostoma kalkoae sp. nov. (in mm). Character
Holotype, USNM 582249 =
Paratype, EKVK 119 R
Greatest length of skull Condyloincisive length Condylocanine length Braincase breadth Zygomatic breadth Postorbital breadth Palatal width at canines Mastoid width Mastoid process width Palatal length Maxillary toothrow length Molariform toothrow length Width at M2 Dentary length Mandibular toothrow length Coronoid height Forearm length Metacarpal III length Metacarpal IV length Metacarpal V length Tail length Hind-foot length Tibia length Calcar length
23.8 20.4 19.6 9.5 11.2 3.9 4.7 9.4 11.3 10.1 8.0 6.7 7.8 14.7 9.1 5.5 45.8 36.1 38.2 40.8 8.9 12.8 23.4 16.1
23.2 19.8 19.2 9.5 11.4 3.6 4.3 9.5 11.3 9.9 7.9 6.4 7.5 14.3 9.0 5.1 44.6 35.7 38.0 40.2 8.4 12.5 22.8 13.7
L. brasiliense, L. carrikeri, and L. schulzi (postparacrista does not contact premetacrista, hence trigon open labially in L. evotis, L. occidentalis, and L. silvicolum); M2 without parastyle in L. kalkoae and L. carrikeri (present in L. brasiliense, L. evotis, L. occidentalis, L. schulzi, and L. silvicolum); M2 hypocone either absent or weakly developed in L. kalkoae and L. carrikeri (moderately or well developed in L. brasiliense, L. evotis, L. occidentalis, L. schulzi, and L. silvicolum); M2 lingual cingulum present in L. kalkoae and L. occidentalis (absent in L. brasiliense, L. carrikeri, L. evotis, L. schulzi, and L. silvicolum). M3 is compressed labiolingually. Length and width of the crown of the lower inner incisors are subequal in L. kalkoae, L. brasiliense, L. carrikeri, and L. schulzi (crown longer than wide in L. evotis; width greater than length in L. occidentalis and L. silvicolum). The p3 is well developed in L. kalkoae, L. carrikeri, L. evotis, L. occidentalis, L. schulzi, and L. silvicolum (p3 minute in L. brasiliense); p3 aligned in toothrow (occlusal view) in L. kalkoae, L. carrikeri, L. occidentalis, L. schulzi, and L. silvicolum (p3 labially displaced, not in line with other teeth in L. brasiliense and L. evotis). Labial cingulid of p4 is undulate in L. kalkoae, L. brasiliense, L. carrikeri, and L. evotis (straight in L. occidentalis, L. silvicolum, and L. schulzi). Multivariate analysis.—We compared the 2 specimens of Lophostoma kalkoae with 70 specimens of L. brasiliense, 13 of L. carrikeri, 3 of L. evotis, 32 of L. occidentalis, 4 of L. schulzi, 163 of L. silvicolum, and 1 of L. yasuni (Appendix I). The first 3 PCs accounted for 91% of the overall variation (Table 4). A plot of factor scores on the first 2 axes (Fig. 4) shows that L. kalkoae clustered with L. carrikeri and L. schulzi
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on PC1, which represents overall size. We did not plot factor scores for L. brasiliense in Fig. 4 because we wanted a better depiction of the spatial distribution of the other species with L. kalkoae. L. brasiliense is the smallest species in the genus, so its specimens would plot lower on PC1 than all other species of the genus. L. evotis, L. occidentalis, L. silvicolum, and L. yasuni plot higher than L. carrikeri, L. kalkoae, and L. schulzi on PC1, corresponding to their larger sizes (Fig. 4). Natural history.—Both specimens of L. kalkoae were collected in their roost, a hollowed-out abandoned Azteca ant (Azteca sp.—Hymenoptera: Formicidae) nest, hanging approximately 12 m above the ground on a tree (unknown species) of about 30–40 cm diameter at breast height. During their capture, 10 other bats in the roost, presumably of the same species, managed to escape. Other species of the genus have been reported as roosting in hollowed-out termite nests, including L. brasiliense (York et al. 2008), L. carrikeri (M. A. Carriker in Allen 1911; McCarthy et al. 1983), and L. silvicolum (Dechmann et al. 2004, 2005; Handley 1976; Kalko et al. 1999, 2006; McCarthy et al. 1983; Sanborn 1951; Tuttle 1970). Both bats were placed in a flight cage to record their echolocation calls and be photographed. During the photography session the bats where fed katydids, which they readily consumed (M54224–M54236). Both bats emitted echolocation calls that were typical for phyllostomid bats (Kalko 2004): steep frequency-modulated, multiharmonic calls with the main energy in the higher harmonics (mostly in the 2nd and 3rd or in the 3rd to the 4th, rather than the 1st).
DISCUSSION In several species of mammals (e.g., Crocuta crocuta and Lemur catta), the appearance of female genitalia resembles those of the male. Such ‘‘masculinized’’ females may show elongation of the clitoris, increased overall size, and aggressively mediated social dominance over males (Drea 2009). This pattern is expressed to greater or lesser extent in some species of Lophostoma. Based on a specimen of L. schulzi (AMNH 257128 [M54312 and M54313]), McCarthy et al. (1989) reported that the clitoris is pendulous, resembling a penis. Simmons and Voss (1998) confirmed the observations of McCarthy et al. (1989), based on 3 specimens collected in French Guiana (AMNH 267105, 267420 [M54314 and M54315], and 267421 [M54316]). The clitoris of L. kalkoae (M54253–M54255) is similar to that of L. carrikeri in being elongated, although not to the same degree as in L. schulzi (M54312–M54316), and the labia are swollen (dark red in live L. kalkoae). Our observations indicate that the clitorides of L. brasiliense, L. occidentalis, and L. silvicolum are elongated like those of L. kalkoae and L. carrikeri, but the labia are not swollen. Further studies are needed in order to understand the significance of and the physiological mechanisms behind this pattern in some species of Lophostoma. Although L. brasiliense and L. silvicolum are each currently regarded as species, we found considerable morphological and
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TABLE 3.—Selected measurements (in mm) of Lophostoma species. Summary statistics (mean and standard deviation [1st line], observed range and sample size [2nd line]) of measurements are given for each species (see Appendix I for a list of specimens measured and Table 1 for definitions of measurement abbreviations). All measurements are in millimeters. Character
L. brasiliense
L. carrikeri
L. evotis
L. occidentalis
L. schulzi
L. silvicolum
L. yasuni
GLS ¯ X 6 SD Range (n)
20.0 6 0.9 18.6–21.8 (70)
24.3 6 0.8 23.0–25.3 (13)
25.3 6 0.3 24.9–25.5 (3)
26.6 6 0.7 25.5–28.7 (32)
23.2 6 0.3 22.8–23.6 (4)
27.6 6 1.1 25.4–30.5 (163)
26.6
CIL ¯ X 6 SD Range (n)
17.6 6 0.8 16.4–19.3 (70)
21.0 6 0.5 20.2–21.9 (13)
21.9 6 0.2 21.6–22.0 (3)
23.1 6 0.6 22.1–24.4 (32)
20.1 6 0.4 19.8–20.6 (4)
24.1 6 0.9 22.4–26.2 (163)
23.5
CCL ¯ X 6 SD Range (n)
17.1 6 0.8 15.9–18.8 (70)
20.4 6 0.5 19.5–21.3 (13)
21.1 6 0.2 20.9–21.3 (3)
22.3 6 0.5 21.5–23.7 (32)
19.5 6 0.4 19.0–20.0 (4)
23.3 6 0.8 21.6–25.5 (163)
22.6
BB ¯ X 6 SD Range (n)
8.1 6 0.2 7.5–8.6 (70)
9.6 6 0.3 9.1–10.3 (13)
10.0 6 0.3 9.7–10.2 (3)
10.3 6 0.3 9.8–11.2 (32)
9.5 6 0.2 9.3–9.8 (4)
10.6 6 0.3 10.0–11.7 (163)
10
ZB ¯ X 6 SD Range (n)
9.5 6 0.5 8.5–10.6 (70)
11.2 6 0.4 10.2–12.1 (13)
12.0 6 0.3 11.6–12.3 (3)
12.7 6 0.3 12.1–13.4 (32)
11.4 6 0.3 11.0–11.8 (4)
13.4 6 0.5 12.4–14.7 (163)
12.7
PB ¯ X 6 SD Range (n)
3.2 6 0.1 2.8–3.4 (70)
3.8 6 0.2 3.3–4.1 (13)
4.0 6 0.2 3.8–4.2 (3)
4.1 6 0.1 3.9–4.4 (32)
3.8 6 0.2 3.7–4.1 (4)
4.1 6 0.2 3.7–4.5 (163)
4
MSTW ¯ X 6 SD Range (n)
8.4 6 0.4 7.5–9.5 (70)
9.7 6 0.3 9.0–10.1 (13)
10.3 6 0.2 10.1–10.5 (3)
10.5 6 0.3 10.1–11.3 (32)
9.7 6 0.2 9.5–10.0 (4)
11.0 6 0.4 10.0–12.2 (163)
10.3
MPW ¯ X 6 SD Range (n)
9.3 6 0.4 8.6–10.2 (70)
11.6 6 0.6 10.8–12.8 (13)
12.3 6 0.0 12.3–12.3 (3)
12.8 6 0.5 11.9–13.8 (32)
11.9 6 0.2 11.6–12.1 (4)
13.5 6 0.5 12.4–15.0 (163)
12.7
PL ¯ X 6 SD Range (n)
8.9 6 0.4 8.2–10.0 (70)
10.7 6 0.4 10.0–11.1 (13)
11.4 6 0.2 11.3–11.6 (3)
12.2 6 0.3 11.6–13.0 (32)
10.1 6 0.4 9.7–10.6 (4)
13.0 6 0.6 11.8–14.6 (163)
11.8
MTRL ¯ X 6 SD Range (n)
7.0 6 0.3 6.4–7.7 (70)
8.2 6 0.2 7.9–8.6 (13)
8.7 6 0.1 8.6–8.8 (3)
9.5 6 0.2 9.2–10.0 (32)
7.9 6 0.2 7.7–8.2 (4)
9.8 6 0.4 9.0–10.6 (163)
9.4
MLTRL ¯ X 6 SD Range (n)
5.8 6 0.3 5.0–6.5 (70)
7.0 6 0.2 6.6–7.3 (13)
7.2 6 0.1 7.1–7.2 (3)
7.6 6 0.3 7.0–8.1 (32)
6.4 6 0.2 6.0–6.6 (4)
7.9 6 0.4 7.0–9.9 (163)
7.9
M2–M2 ¯ X 6 SD Range (n)
6.3 6 0.3 5.6–7.0 (70)
7.5 6 0.2 7.1–7.9 (13)
7.9 6 0.1 7.7–8.0 (3)
8.5 6 0.2 8.2–9.0 (32)
7.4 6 0.2 7.2–7.6 (4)
8.8 6 0.4 8.0–9.7 (163)
8.8
DENL ¯ X 6 SD Range (n)
12.6 6 0.6 11.7–14.1 (70)
15.0 6 0.5 13.8–15.9 (13)
15.8 6 0.3 15.6–16.1 (3)
17.0 6 0.4 16.1–18.4 (32)
14.6 6 0.2 14.2–14.8 (4)
17.9 6 0.7 16.4–19.9 (163)
16.9
MANDL ¯ X 6 SD Range (n)
7.7 6 0.4 7.0–8.6 (70)
9.3 6 0.3 8.8–9.7 (13)
9.9 6 0.1 9.7–10.0 (3)
10.6 6 0.3 10.1–11.2 (32)
9.1 6 0.3 8.8–9.3 (4)
11.0 6 0.4 10.1–12.1 (163)
10.7
FA ¯ X 6 SD Range (n)
35.3 6 1.9 30.1–39.9 (70)
46.1 6 1.1 44.1–47.7 (13)
49.7 6 1.5 48.7–51.4 (3)
53.9 6 1.4 51.2–56.8 (32)
43.0 6 1.1 41.6–44.4 (4)
53.8 6 2.7 45.4–59.5 (163)
43.9
morphometric evidence suggesting that both taxa (Table 3) are composite and deserving of revisory studies (see Velazco and Cadenillas 2011). The morphological patterns found in these 2 species are similar to those of species with distributions in both Central and South America and that now, after analyses based on large sample sizes and a combination of molecular and
morphological approaches, are found to be composites of 2 or more species (e.g., Artibeus jamaicensis [Larsen et al. 2010; Marcha´n-Rivadeneira et al. 2010], Platyrrhinus helleri [Velazco et al. 2010], and Vampyrodes caraccioli [Velazco and Simmons 2011]). L. kalkoae is the only species of white-bellied Lophostoma present in Central America (Fig. 1) and its
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TABLE 4.—Factor loadings for the first 3 factors extracted from a principal component (PC) analysis of 18 variables for Lophostoma brasiliense, L. carrikeri, L. evotis, L. kalkoae sp. nov., L. occidentalis, L. schulzi, L. silvicolum, and L. yasuni. See Table 1 for definitions of abbreviations of variables.
2. 29.
3.
Correlations Variable
PC1
PC2
PC3
GLS CIL CCL BB ZB PB C–C MSTW MPW PL MTRL MLTRL M2–M2 DENL MANDL COH FA MET–III Proportion of variation (%)
0.956 0.980 0.981 0.871 0.945 0.650 0.940 0.909 0.903 0.955 0.967 0.867 0.922 0.980 0.968 0.929 0.802 0.865 84.8
0.031 0.034 0.046 20.099 20.151 20.074 20.163 20.124 20.175 0.085 0.091 0.169 20.006 20.010 0.008 20.192 0.503 0.317 3.4
0.012 20.021 20.024 0.111 0.002 0.673 0.092 20.011 20.052 20.058 0.054 20.008 0.067 0.004 0.069 20.228 20.023 20.051 2.8
recognition increases the number of known species in the genus to 8.
KEY TO THE SPECIES OF LOPHOSTOMA 1.
Fur of underparts entirely white, except on chin and sides of abdomen . . . . . . . . . . . . . . . . . . . . . . . . 2 19. Fur of underparts brown to gray, a pale patch on the chest can be present . . . . . . . . . . . . . . . . . . . . . . 4
39.
4. 49. 5.
59.
6. 69. 7.
79.
611
Forearm longer than 44 mm; greatest length of skull 25.5 mm or less . . . . . . . . . . . . . . . . . . . . . . . . . 3 Forearm shorter than 44 mm; greatest length of skull 25.5 mm or more; known only from Ecuador. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophostoma yasuni Conspicuous pale to white postauricular patches; strong indentation on lingual cingulum of upper canine; known only from Panama . . . . . Lophostoma kalkoae Pale to white postauricular patches lacking; weak indentation on lingual cingulum of the upper canine; known only from South America . . . . . . . Lophostoma carrikeri Forearm shorter than 45 mm; greatest length of skull 24 mm or less . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Forearm longer than 45 mm; greatest length of skull 24 mm or more . . . . . . . . . . . . . . . . . . . . . . . . . 6 Forearm shorter than 40 mm; lacking small wartlike granulations on head, wings, and legs; greatest length of skull less than 22 mm . . . Lophostoma brasiliense Forearm longer than 40 mm; small wartlike granulations on dorsal surfaces of forearms, digits, legs, and nose leaf; greatest length of skull more than 22 mm . . . . . . . . . . . . . . . . . . . Lophostoma schulzi M1 hypocone moderately to well developed; p3 aligned in toothrow when seen in occlusal view . . . 7 M1 hypocone absent; p3 displaced labially from toothrow, not in line with other teeth . . . Lophostoma evotis Pale to white postauricular patches; indentation on the lingual cingulum of the upper canine either absent or weakly developed . . . . . . . Lophostoma occidentalis Postauricular patches lacking; strong indentation present on the lingual cingulum of the upper canine . . . . . . . . . . . . . . . . Lophostoma silvicolum
RESUMEN Reportamos el descubrimiento de una nueva especie del ge´nero Lophostoma de Panama´, que nombramos L. kalkoae. Esta nueva especie se asemeja a L. carrikeri y L. yasuni por su vientre blanco, pero es diferenciable de ambas por caracter´ısticas externas y craniales. Morfometricamente esta especie nueva es similar en taman˜o a L. carrikeri and L. schulzi. Esta nueva especie se puede reconocer por su vientre blanco; parches pos-auriculares conectados por una linea de pelo claro con el pelo blanco en el pecho; clitoris alargado con los labios hinchados; ausencia de la proyeccio´n lateral del proceso mastoideo; fuerte indentacio´n presente en el cı´ngulo labial del canino superior; P3 bien desarrollado; cuspide bien desarrollada en el cı´ngulo lingual de P4; parastilo ausente en el M1 y M2, entre otros cara´cteres. Se presenta una clave dicoto´mica para todas las especies del ge´nero Lophostoma y un mapa incluyendo todas las localidades donde han sido colectados especies de Lophostoma de vientre blanco.
FIG. 4.—Plot of scores on 1st and 2nd axes from a principal component analysis of 18 variables from 13 Lophostoma carrikeri, 3 L. evotis, 2 L. kalkoae sp. nov., 32 L. occidentalis, 4 L. schulzi, 163 L. silvicolum, and 1 L. yasuni. PC 5 principal component.
ACKNOWLEDGMENTS The following curators and collection staff graciously provided access to specimens under their care: N. B. Simmons and E. Westwig,
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American Museum of Natural History, New York; B. D. Patterson and J. Phelps, Field Museum of Natural History, Chicago; M. S. Hafner, Museum of Natural Science, Louisiana State University, Baton Rouge; V. Pacheco, Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima; S. Burneo, Museo de Zoologı´a, Pontificia Universidad Cato´lica del Ecuador, Quito; R. J. Baker, Museum of Texas Tech University, Lubbock; and S. C. Peurach, Patuxent Wildlife Research Center, United States Geological Survey. We thank P. J. Wynne for the illustration of Fig. 2, J. P. Carrera and C. M. Pinto for their measurements of the holotypes of Lophostoma aequatorialis and L. yasuni, and D. von Staden for the live bat photographs. For critical comments on an early draft of this manuscript, we thank T. Chesser, K. Kline, R. Pine, N. Woodman, and 1 anonymous reviewer. Funding for this project was provided to PMV by the Gerstner and Roosevelt postdoctoral fellowships at the American Museum of Natural History.
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Submitted 8 June 2011. Accepted 9 October 2011. Associate Editor was Elizabeth R. Dumont.
APPENDIX I The following list includes all specimens examined in this study, with their respective localities. See ‘‘Materials and Methods’’ for museum abbreviations. Individuals or series in bold were used only in the morphological comparisons. In Lophostoma carrikeri, individuals or series with an asterisk were used in the morphometric and morphological analyses; the remaining localities were used only in the construction of Fig. 1. Lophostoma brasiliense (70).—BOLIVIA: Cochabamba: Chapare, Sajta (AMNH 264927). BRAZIL: Para´: Bele´m (USNM 336990, 460096); Igarape Mirim, Tapajo´s River (AMNH 95497, 95498). COLOMBIA: Antioquia: Zaragoza, 25 km S, 22 km W, at La Tirana (USNM 499293, 499294). COSTA RICA: Guanacaste: Palo Verde Wildlife Refuge, OTS (USNM 566438). Heredia: Parque Nacional Braulio Carrillo, San Miguel, 1 km S, 11.5 km E (USNM 562760). ECUADOR: Pichincha: Santo Domingo, 47 km S (by road), Rı´o Palenque Science Center (USNM 528484, 528485). FRENCH GUIANA: Cayenne: Sinnamary, Paracou (AMNH 267101, 267103, 267916, 267917). GUYANA: Barima–Waini: North West District, Baramita (USNM 582273). PANAMA: Bocas del Toro: Changuinola, 7 km SSW (USNM 315219); Sibube (USNM 335106, 335107); Isla San Cristobal, Bocatorito (USNM 449604). Colo´n: Fort Sherman (USNM 314221); Buena Vista Peninsula, 1.75 km NNW Frijoles (USNM 503446). Chiriquı´: San Vincente (USNM 331113); Progreso, 1 mile SW (USNM 362456); Progreso, 1.5 miles SW (USNM 362457); Progreso, 8 miles S (USNM 362458); San Vincente, 2 miles W (USNM 331114). Darie´n: Jaque, Junction Rı´os Jaque and Imamado (USNM 362453); Jaque, Rı´o Jaque, 20 min below junction with Rı´o Imamado (USNM 362454); Jaque, junction Rı´os Jaque and Imamado, 2 h downstream (USNM 362455). Los Santos: Cerro Hoya (USNM 323067). Panama´: Barro Colorado Island, Lutz Ravine (USNM 582080). San Blas: Armila, Quebrada Venado (USNM 335108, 335109); Mandinga (USNM 305185). PERU: Loreto: Curaray River (AMNH 71620, 71623, 71625, 71626). TRINIDAD AND TOBAGO: Trinidad: Nariva Parish, Rı´o Claro (AMNH 205370); Saint George Parish, Port of Spain (AMNH 207062). VENEZUELA: Amazonas: San Juan, 163 km ESE Puerto Ayacucho, Rı´o Manapiare (USNM 407278–407282); Puerto Ayacucho, 65 km SSW Puerto Ayacucho, near Morganito (USNM 407285). Apure: Nulita, 29 km SSW Santo Domingo, Selvas de San Camilo (USNM 418965–418968). Barinas: Altamira (USNM 418960, 418961). Bolı´var: El Manaco, 59 km SE El Dorado, km 74 (USNM 385453). Carabobo: Palmichal, 23 km N Bejuma (USNM 562891); San Esteban (USNM 142567). Falco´n: Urama, 19 km NW Urama, km 40 (USNM 371448, 371454, 371456–371462). Monagas: Hato Mata de Bejuco, 55 km SSE Maturı´n, near Rı´o Tigre (USNM 385454). Trujillo: Valera, 19 km N Valera, near Agua Viva (USNM 371463). Vargas: Macuto (USNM 102919). Yaracuy: Urama, 11 km NW Urama, El Central (USNM 373484, 373487). Lophostoma carrikeri (13 measured).—BOLIVIA: Beni: Mamore, opposite Costa Marques, Brazil, Itenez River (AMNH
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JOURNAL OF MAMMALOGY
209322*). BRAZIL: Para´ : Bele´ m, Mocambo, Igapo (USNM 393005*); Bele´m, Varzea (USNM 460095*); Belterra, 30 km SW Santare´m (MPEG 9221, 9222); Rı´o Iriri, 85 km SW of Altamira. Piauı´: Teresina. Roraima: Ilha de Maraca´ (INPA 1394). COLOMBIA: Meta: Hacienda Los Guaduales near San Martı´n (ICN 5144); San Juan de Arama (FMNH 87942–87945, 88081–88091). FRENCH GUIANA: Cayenne: Paracou (AMNH 267918). GUYANA: Potaro– Siparuni: Clearwater Camp (ROM 107391, 107392); Iwokrama Field Station (ROM 107461); Three Mile Camp (ROM 107190). PERU: Loreto: Estacio´n Biolo´gica Isla Muyuy (MUSM 21175); Requena, Centro de Investigaciones Jenaro Herrera (MUSM 6977). Ucayali: Coronel Portillo, Cerro Tahuayo, approximately 65 km ENE of Pucallpa (MUSM 3158); Rı´o Curanja, Balta (LSUMZ 14076*, 14077*). SURINAME: Brokopondo: Brownsberg Nature Park, Jeep Trail (ROM 114041). Para: Zanderij; Sipaliwini: Sipaliwini Airstrip (CM 77172). Sipaliwini: Bakhuis, Area 22 Recon Fly Camp (ROM 117005); Voltzberg (CM 63668). VENEZUELA: Amazonas: Cerro Neblina Base Camp, Rı´o Mawarinuma (USNM 560556*); San Juan, 163 km ESE Puerto Ayacucho, Rı´o Manapiare (USNM 407274*). Bolı´var: Mocho River (AMNH 30177–30180*, 30182*, 30183*). Lophostoma evotis (3).—BELIZE: Toledo: Columbia Forest, Crique Negro, Collin’s Trail (USNM 506465); Columbia Forest Station (USNM 506466). GUATEMALA: Izabal: El Estor, El Estor airstrip (AMNH 267635). Lophostoma kalkoae sp. nov. (2).—PANAMA: Colo´n: Soberania National Park, Pipeline road (EKVK 119; USNM 582249 [holotype]). Lophostoma occidentalis (32).—ECUADOR: Esmeraldas: Comuna San Francisco de Bogota´ (TTU 103215); Estacio´n Experimental La Chiquita, near San Lorenzo town (QCAZ 6500 [holotype of Lophostoma aequatorialis]). Guayas: Reserva Ecolo´gica Manglares Churute, Cerro Pancho Diablo (TTU 103573, 103583, 103584). Los Rı´os: El Papayo, 7 km SW Puebloviejo, near San Juan (USNM 522064). Pichincha: Santo Domingo, 47 km S (by road), Rı´o Palenque Science Center (USNM 528483). PERU: Piura: Ayabaca, Paymas (FMNH 81126); Morropon, Hacienda Bigote (FMNH 81113–81125). Tumbes: Tumbes, Pampas de Hospital, Quebrada Angostura (MUSM 19332–19336, 22164–22168); Zarumilla, Matapalo, Rı´o Zarumilla, Caban˜a INRENA (MUSM 22169). Lophostoma schulzi (7).—FRENCH GUIANA: Cayenne: Sinnamary, Paracou (AMNH 267105, 267106, 267420, 267421, 267920). Saint Laurent du Maroni: Sau¨l (AMNH 257128). GUYANA: Barima–Waini: North West District, Baramita (USNM 582274). Lophostoma silvicolum (163).—BOLIVIA: Pando: Federico Roman, left bank of Beni River (AMNH 262425). Santa Cruz: Velasco Province, Parque Nacional Kempff Mercado, El Refugio (USNM 584477). BRAZIL: Amazonas: Sa˜o Gabriel do Cachoeira, Taua, Uaupes River (AMNH 78637, 78639, 78642, 78643). Para´: Tapajo´s River, Limoal (AMNH 95443, 95444); Tapajo´s River, Limontuba (AMNH 95431, 95438); Baiao, Tocantins River (AMNH 97011, 97012); Cameta´, Tocantins River, Ilha do Taiuna (AMNH 96978, 96980–96984, 96990, 96992–96994, 96996, 97000–97002, 97005, 97006); Igarape Mirim, Tapajo´s River (AMNH 95442); Mocajuba, Tocantins River (AMNH 97007). ECUADOR: Napo: Suno River (AMNH 64026); Loreto, San Jose Nuevo (AMNH 64031, 66798, 67938, 67940, 67942, 67943, 67946–67949, 67952). Pastaza: Curaray River mouth (AMNH 71464, 71469, 71471). FRENCH
Vol. 93, No. 2
GUIANA: Cayenne: Sinnamary, Paracou (AMNH 267107, 267108, 267923). GUYANA: Cuyuni–Mazaruni: Kartabo Point, Cuyuni River (AMNH 142906, 142907). Upper Demerara–Berbice: Kwakwani, Aroaima Mining Company (USNM 588492); Tacama Trail, 5.5 miles from Ituni Junction (USNM 582275). Upper Takutu–Upper Essequibo: Dadanawa, Rupununi Savanna, Imprenza (AMNH 182718). PANAMA: Bocas del Toro: Isla San Cristobal, Bocatorito (USNM ˜ urı´ (USNM 575457, 575459–575463); Tierra Oscura, 449605); N 3.5 km S Tiger Key (USNM 449606). Colo´n: Fort Sherman (USNM 396400, 396401); Frijoles, Bohio Peninsula, 4 km NNW (USNM 461096, 461097); Frijoles, Bohio Peninsula, 4.5 km NW (USNM 461095); Frijoles, Bohio Peninsula, 6 km NWW (USNM 461099); Frijoles, Buena Vista Peninsula, 1.75 km NNW (USNM 461100, 461101); Gamboa, 6 miles N (USNM 520547–520549). Darie´n: Cerro Mali (USNM 337974, 337975); Jaque, 30 min downstream from rı´os Jaque and Imamado (USNM 362459, 362460); Rı´o Paya, mouth (USNM 306549); Tacarcuna Village Camp (USNM 309357). Los Santos: Guanico Arriba, Jobero (USNM 323080, 323082); Las Palmitas, Jobero (USNM 323068, 323069, 323071–323077). Panama´: Barro Colorado Island (USNM 304876, 332045, 519698); Cerro Azul (USNM 306548). San Blas: Armila, Quebrada Venado (USNM 335114–335116). PERU: Cuzco: La Convencio´n, La Convencio´n, 2 km SW of C.N. Tangoshiari (MUSM 13400); La Convencio´n, Camisea, Armihuari (USNM 582791, 582793); Paucartambo, Consuelo, 15.9 km SW Pilcopata (MUSM 19706). Junı´n: Chanchamayo, Chanchamayo (MUSM 1191–1197). Loreto: Alto Amazonas, Nuevo San Juan, Galvez River (AMNH 272800, 272801, 272829, 272833, 273074, 273087; MUSM 13272–13276, 15284); Maynas, Alto Nanay, Quebrada Agua Blanca (MUSM 24421); Maynas, Puerto Indiana, Amazon River (AMNH 73526, 73527); Maynas, Rı´o Lagartococha, Campamento Catalino (MUSM 21183, 21184); Maynas, Orosa, Amazon River (AMNH 73721, 74101, 74102). Madre de Dios: Manu, Est. Biol. Cocha Cashu (MUSM 5098); Manu, Rı´o Manu, 40 km above mouth (MUSM 187); Manu, Manu, Pakitza (MUSM 6816); Tambopata, Rı´o Tambopata, 30 km above mouth (MUSM 186); Tambopata, Reserva Cusco Amazo´nico, 15 km NE of Puerto Maldonado (MUSM 6256). Pasco: Oxapampa, Pozuzo, Yanahuanca (MUSM 10991); Oxapampa, San Juan (USNM 364278, 364282, 364283, 364295–364300). San Martı´n: Moyobamba, Area de Conservacio´n Municipal Mishquiyacu RumiyacuAlmendra, Orquidiario Waqanki (FMNH 203542). Ucayali: Coronel Portillo, Pucallpa, Rı´o Ucayali (MUSM 1343); Coronel Portillo, Yarinacocha (MUSM 1201). VENEZUELA: Amazonas: Belen, 56 km NNW Esmeralda, Rı´o Cunucunuma (USNM 388748, 388752); Cerro Neblina, Base Camp (USNM 560800, 560801); Capibara, 106 km SW Esmeralda, Brazo Casiquiare (USNM 407296–407299); Puerto Ayacucho, 25 km S Puerto Ayacucho, Paria (USNM 407302). Bolı´var: Los Patos, 28 km SE El Manteco (USNM 385455). Falco´n: Urama, 19 km NW Urama, km 40 (USNM 373489, 373490). Trujillo: Valera, 25 km NW Valera, near Agua Santa (USNM 370084, 371468–371471). Zulia: El Rosario, 60 km WNW Encontrados, Boca del Rı´o de Oro (USNM 418970). Lophostoma yasuni (1).—ECUADOR: Orellana: Yasunı´ National Park and Biosphere Reserve, Yasunı´ Research Station (QCAZ 4935 [holotype]).
