New records of the Western Rounded Ear Bat , Lophostoma occidentalis ( Davis & Carter , 1978 ) ( Chiroptera : Phyllostomidae ) , from Colombia

We report new records of the western rounded ear bat Lophostoma occidentalis for the Colombian territory, represented by ten specimens collected on the western portion of the country, in the departments of Cauca, Chocó, and Valle del Cauca, all in the Colombian Pacific Region. The new records of L. occidentalis represent a substantial geographic and ecological extension in the known distribution of this species and corroborate its occurrence in Colombia. In addition, we discuss on the morphological variation of representatives of large Lophostoma species in Colombia.


INTRODUCTION
In Colombia, phyllostomid bats of the genus Lophostoma are currently represented by two small-sized species-L.brasiliense Peters, 1867 andL. carrikeri (Allen, 1910) -and two large species -L.occidentalis (Davis & Carter, 1978) and L. silvicolum d'Orbigny, 1836.Lophostoma occidentalis, however, has only recently been recognized as a valid species (Velazco and Cadenillas 2011), and Colombian records from the western versant of the Andes were all assigned as L. s. occidentalis (Alberico 1994).
The taxonomic history of L. occidentalis began with Davis and Carter (1978), who described this taxon as a geographic variant of L. silvicolum from western Peru (L.s. occidentalis).On their part Baker et al. (2004) found consistent morphological and molecular differences between typical L. silvicolum and specimens of the genus from several localities in western Ecuador.Baker et al. (2004) concluded that divergent specimens from western Ecuador should be included in an undescribed species they named L. aequatorialis and suggested that Colombian material previously identified as L. s. occidentalis may represent L. aequatorialis.Baker et al. (2004) advice that at the time of their work no karyotypes or tissues were available for L. s. occidentalis, and despite the authors were allowed to extract DNA from the holotype of this taxon (Texas Cooperative Wildlife Collection -TCWC 11704), it was not possible to obtain DNA sequences, preventing molecular analyses.Williams and Genoways (2008) restricted the distribution of L. s. occidentalis to Peru and Ecuador influenced by Baker et al. (2004) suggestion that Colombian material of L. s. occidentalis may represent L. aequatorialis.Velazco and Cadenillas (2011) analyzed new museum vouchers of L. s. occidentalis, with tissues available, and failed in identifying the evolutionary independence of typical L. s. occidentalis from L. aequatorialis, suggesting the elevation of L. s. occidentalis to the specific level.These findings open new questions about the identity of large Lophostoma specimens from western Colombia, and our aim here is to review the identification of those specimens, including the material assigned as L. s. occidentalis by Alberico (1994).We also examined specimens previously identified as L. silvicolum from other Colombian natural regions.

MATERIALS AND METHODS
Forty seven Colombian specimens of Lophostoma assigned as L. silvicolum or L. s. occidentalis in previous studies were contrasted against L. occidentalis morphological diagnostic characters.These include specimens from 12 departments, and from the five Colombian regions (Amazon, Andes, Caribbean, Orinoco, and Pacific).Three appendicular (Ear, E; forearm, FA; and third metacarpal, Mt3) and 16 craniodental measurements (greatest skull length, GSL; condylobasal length, CBL; zygomatic width, ZW; mastoid width, MW; braincase width, BW; braincase height, BH; palatal length, PL; postpalatal length, PPL; postorbital constriction width, PO; interorbital width, IW; distance across upper molars, M-M; distances across canines, including cingula, C-C; maxilar tooth-row length, TR; mandibular tooth-row length, MTR; coronoid process length, CR; and mandibular length, ML) were taken of each specimen reviewed.A total of 30 specimens with complete measurements (marked with an asterisk in Appendix 1), were analysed through a principal component analysis (PCA), in order to explore whether there is a morphological gap or intraspecific geographic structure separating L. occidentalis

NOTES ON GEOGRAPHIC DISTRIBUTION
The average of the measurements of external and craniodental characters tend to be larger in L. silvicolum than in L. occidentalis (Table 1).However, all variables overlap in their range.Morphometric variation of Colombian L. silvicolum and L. occidentalis specimens analyzed in our PCA based on skull measurements show no interspecific segregation, or intraspecific geographic structure in L. silvicolum.In the PCA, the first two components accounted for most of the observed variation (PC1=55.5;PC2=9.3), with higher loadings on PC1 associated with GSL, CBL, MW, and ML, whereas GSL, MW, and BH had the highest loadings on PC2 (Figure 4).

DISCUSSION
The herein presented records of L. occidentalis for Colombia represent a significant geographic and ecological extension in the distribution of this taxon.Additionally, we confirmed that the three specimens assigned as L. s. occidentalis from Department of Valle del Cauca by Alberico (1994), all deposited at UV collections (UV 5673 ♂, UV 5674 ♂, UV 5675 ♀), conform with the diagnosis provided by Velazco and Cadenillas (2011) for L. occidentalis.and L. silvicolum from Colombia.This analysis was performed using the software PAST (Hammer et al. 2001) and included only skull measurements (log-transformed) of adult specimens (identified by closed epiphyses).Males and females were analysed together because both species lack sexual dimorphism (Velazco and Cadenillas 2011).Examined material is deposited at the following institutions in Colombia: Instituto Alexander von Humboldt (IAvH); Instituto de Ciencias Naturales, Universidad Nacional de Colombia (ICN); Museo de Historia Natural Universidad de la Amazonia (UAM); Museo de Historia Natural Universidad de La Salle (MLS); Museo de Historia Natural Universidad del Cauca (MHNUC); and Universidad del Valle (UV).

RESULTS
As product of our taxonomic assessment, ten specimens were identified as L. occidentalis, all obtained in localities west of the Andean system (Figure 1 and 2, Appendix 1), in the Chocó-Magdalena biogeographic province, as defined by Hernández-Camacho et al. (1992).These specimens match the following characters in L. occidentalis re-description (Velazco and Cadenillas 2011): mastoid process developed; clinoid processes laterally curved; cingula absent on upper canines; distal accessory cusp on P3 and P4 present; labial cingula on P4 absent; mesostylar crest on M1 and M2 present, and lingual cingula on M1 and M2 present (Figure 3).Prior Velazco and Cadenillas (2011), it was difficult to define the northern limits of L. s. occidentalis and this is probably the reason for which the taxon was not reported for the western portion of the country in recent checklists (Alberico et al. 2000;Mantilla-Meluk and Jiménez-Ortega 2006;Mantilla-Meluk et al. 2009).As mentioned above, Colombian L. occidentalis match all cranial characters in Velazco and Cadenillas (2011) re-description of the species.However, Colombian specimens share a paler dorsal coat color, grayish brown rather than chocolate brown, as in typical L. silvicolum.These Colombian specimens also proved to be variable for the following external diagnostic characters: whitish venter, as described by Davis and Carter (1978); paler throat and naked forearm, as described by Velazco and Cadenillas (2011); and white distinctive postauricular patches, as presented in both descriptions.These variations were found in specimens from the Department of Chocó.The adult male IAvH 4490 has less distinctive postauricular patches, paler throat, and whitish venter, while the adult female ICN 5928 presents distinctive postauricular patches and a frosted throat of same color as the venter.Interestingly, all herein presented Colombian specimens of L. occidentalis have a dorsal frosted coat color pattern, contrasting the non-frosted L. silvicolum dorsal pelage of Colombian specimens also analyzed in this work.In addition, although Davis and Carter (1978) used the size of the middle lower premolar as a diagnostic character of L. s. occidentalis -not discussed by Velazco and Cadenillas (2011) -we found size and placement of this dental piece variable among Colombian analyzed samples.
Finally, despite specimens from the five Colombian natural regions were included in our PCA, there was no evidence of a morphological gap among L. occidentalis and L. silvicolum.These results are in agreement with findings of Velazco and Cadenillas (2011).

Table 1 .
Average, standard deviation, and range of measurements (mm) of external and craniodental characters of Colombian Lophostoma silvicolum (n=25) and L. occidentalis (n=4) (see Material and Methods for abbreviations).