Systematics and Taxonomy of Great Striped-Faced Bats of the Genus Vampyrodes Thomas, 1900 (Chiroptera: Phyllostomidae) Author Velazco, PaúL M. Author Simmons, Nancy B. text American Museum Novitates 2011 2011-04-20 2011 3710 1 36 http://www.bioone.org/doi/abs/10.1206/3710.2 journal article 10.1206/3710.2 0003-0082 5359357 Vampyrodes caraccioli ( Thomas, 1889 ) Caracciolo’s Stripe-faced Bat Figures 1 , 2 , 8 , 9 , and 10 Vampyrops Caraccioloe Thomas, 1889: 167 ; type locality “ Trinidad .” Vampyrops Caraccioli Thomas, 1893a: 186 ; corrected original spelling. V [ ampyrops . ( Vampyrodes )] Caraccioli : Thomas, 1900: 270 ; name combination. Vampyrodes caraccioloe : Miller, 1907: 156 ; first use of current name combination and incorrect sub- sequent spelling of Vampyrops caraccioloi Thomas. Vampyrodes ornatus Thomas, 1924: 532 ; type locality “ San Lorenzo , Rio Marañon , nearly opposite mouth of Huallaga. Alt. 500′,” Loreto , Peru . V [ ampyrodes ]. caraccioloi Pittier and Tate, 1932: 273 ; incorrect subsequent spelling of Vampyrodes caraccioli Thomas. Vampyrodes caraccioloi Goodwin and Greenhall, 1961: 257 ; unjustified emendation of Vampyrops caraccioli Thomas. Vampyrodes ornata Goodwin and Greenhall, 1961: 257 ; incorrect subsequent spelling of Vampyrodes ornatus Thomas. TYPE MATERIAL: The holotype of V. caraccioli is BMNH 89.6.10.2, a subadult of undetermined sex prepared as a skin and skull. The skin is in good condition, but the skull is damaged. The braincase was removed, a distal section of the parietals are missing, and there is a hole on the right parietal. The occipital bone is mostly missing, although the proximal section of the basioccipital is present; the left squamosal and zygomatic arch are missing. The mandibles are intact but were separated at some point and glued together afterward. V. caraccioli was collected by Henry Caracciolo in Trinidad at an unspecified locality. DISTRIBUTION: Vampyrodes caraccioli is known from eastern Colombia , eastern Ecuador , Peru , northern Bolivia , Venezuela , Trinidad and Tobago , French Guiana, Guyana , Suriname , and Brazil (fig. 6). EMENDED DIAGNOSIS: Dorsal fur pale brown to dark brown, 7–9 mm long; two genal vibrissae present; uropatagium with inverted U-shaped posterior margin fringed with short (< 2 mm ) dense hair along its free edge; metacarpal III longer than metacarpal V; rostrum slender; well-developed anterior notch present in nasals; parietal foramina well separated from nuchal crest; weakly developed groove present between occipital condyle and paracondylar process; paraoccipital processes well developed; perikymata present on all upper and lower teeth; I1 broad and bilobed but appears single lobed in older individuals with worn teeth; M1 postentoconule absent or poorly developed; M2 parastyle absent or poorly developed; M2 postentoconule absent or weakly developed; lower incisors robust and bilobed; lingual accessory cuspule present on p4; cuspule on m1 and m2 paracristid absent. DESCRIPTION AND COMPARISONS: A medium-sized Vampyrodes (FA 47.28–55.98 mm ; GLS 25.14–27.97 mm ; CCL 21.98–24.98 mm ; table 4). All linear measurements of V. caraccioli show a slight overlap with those of V. major , with V. caraccioli being the smaller of the two species (tables 4–5). Dorsal pelage in V. caraccioli is long ( 7–9 mm ) and brown, with individual hairs bicolored with darker tips. Compared with V. major , the pelage tends to be slightly lighter and the hairs shorter ( 9–10 mm in V. major ). The ventral pelage is similar but slightly darker, with individual hairs tricolored, with a basal pale brownish band that makes up some 70% to 80% of the total length of each hair. Each hair also has a short dark brown (~ 10% of the total length of each hair) subterminal band, and a tiny pale brownish terminal band. The uropatagium in V. caraccioli has an inverted U-shaped posterior margin with dense and short hair (< 2 mm ) along the trailing edge (V-shaped posterior margin in V. major with dense and long hair (> 2 mm ) along the trailing edge). The width of uropatagium in V. caraccioli is 5–9 mm at midline ( 6–10 mm in V. major ). The proximal half of forearm is covered with dense, short hair. Metacarpal III is longer than metacarpal V in V. caraccioli (the metacarpal III is shorter than metacarpal V in V. major ). The plagiopatagium inserts onto the tarsal bones. Two genal vibrissae are present in V. caraccioli (three genal vibrissae are present in V. major ). V. caraccioli has six vibrissae surrounding the margin of the noseleaf in a single array; two vibrissae on each side of upper lip below the vibrissae surrounding the noseleaf; four submental vibrissae on each side of chin; and two interramal vibrissae. The noseleaf is longer than it is wide and the inferior border of the nasal horseshoe is completely free of upper lip. The skull of V. caraccioli has a slender rostrum (broad and robust in V. major ) and a welldeveloped anterior notch in the nasals (absent or weakly developed in V. major ). Two infraorbital foramina usually present (three infraorbital foramina are present on one specimen examined: USNM 405129). The parietal foramina are well separated from the nuchal crest in V. caraccioli (USNM 405129; figs. 8A, 10A) whereas in V. major (FMNH 127114; figs. 8C, 10B) these foramina are closer to the nuchal crest. The groove between the occipital condyle and paracondylar process is weakly developed (USNM 405129; FMNH 139776; fig. 10A). Compared with V. major , the groove between the occipital condyle and paracondylar process is well developed (FMNH 58263; fig. 10B). The paraoccipital processes are well developed (moderately developed in V. major ). FIGURE 7. Labial view of the left P3–M1 illustrating presence and absence of perikymata. Top , Vampyrodes caraccioli (AMNH 230653) with distinct perikymata (arrow). Bottom , Platyrrhinus lineatus (AMNH 23771) without distinct perikyma. Perikymata are present on all upper and lower teeth (fig. 7, top). The upper inner incisors are broad; both the outer and inner incisors are bilobate (USNM 361711), but may appear single lobed in older individuals with worn teeth (USNM 528341). By comparison, in V. major the upper inner incisors are slender. P3 is more than half the size of P 4 in V. caraccioli , and two stylar cuspules are present on posterior cristid of P4. The M1 lacks a parastyle but both a mesostyle and metastyle are present, and a stylar cuspule is present on the labial cingulum of the metacone. A sulcus is present on the posterior cristid of the M1 paracone, the protocone is well developed, and a postentoconule is absent or poorly developed on M1. On M2 the parastyle is absent or poorly developed (FMNH 139776) (well developed in V. major ), the labial cingulum on the paracone is absent or poorly developed, and the postentoconule is absent or poorly developed (well developed in V. major ). The lower incisors are robust and bilobed (small in V. major ). A p4 lingual accessory cuspule is present. A cuspule on the m1 paracristid is absent in V. caraccioli (present in V. major ). A cuspule on the m2 paracristid is always absent in V. caraccioli . In V. major , this cuspule is sometimes present (AMNH 186381) and sometimes absent (USNM 314717). NATURAL HISTORY: Vampyrodes caraccioli is a frugivorous bat that has been reported to feed on at least six plant species representing three genera in two families: Spondias mombin (Anacardiaceae) and Ficus insipida , F. obtusifolia , F. yoponensis , F . sp, and Poulsenia armata (Moraceae) ( Foster et al., 1986 ; Kalko and Handley, 2001 ; Lobova et al., 2009 ). Lobova et al. (2009) reported an epizoochorous dispersal by V. caraccioli of Cyathula prostrata (Amaranthaceae) , a terrestrial herb with diaspores that adhere to the fur of its dispersal agents. Very few reports of roosts of Vampyrodes caraccioli have been published. Day roosts include foliage, branches, and palm fronds where groups of two to four have been recorded ( Husson, 1954 ; Goodwin and Greenhall, 1961 ). Two species of ectoparasite ( Periglischrus iheringi : Spinturnicidae ; Speleochir brasiliensis : Ereynetidae ) have been reported from V. caraccioli from a Brazilian specimen ( Confalonieri, 1976 ; Fain and Aitken, 1969 ) and one from a Venezuelan specimen ( Paratrichobius sp. , salvini complex: Streblidae ) ( Wenzel, 1976 ). Nogueira et al. (2004) reported that 66% of V. caraccioli captured in western Amazonia of Brazil were infested with trematode Hasstilesia tricolor in their small intestines. Like other stenodermatines, Vampyrodes caraccioli has a litter size of one ( Tuttle, 1970 ; Graham, 1987 ). Reproductive data suggest possible seasonal polyestry; pregnant females have a The sample mean plus or minus one standard deviation, the observed range (in parentheses), and the sample size are provided for each sex. TABLE 5. Measurements (mm) and sample statisticsa of Vampyrodes major

MCZ 6756b	Adult females	Adult males
FA	55.35	55.11 ± 1.23 (52.06–58.64) 103	54.19 ± 1.21 (51.36–56.93) 109
GLS	27.74	27.88 ± 0.49 (26.65–28.96) 102	27.66 ± 0.48 (26.26–29.06) 109
CIL	25.61	25.25 ± 0.44 (23.88–26.14) 101	25.07 ± 0.49 (23.71–26.52) 109
CCL	24.80	24.62 ± 0.43 (23.39–25.53) 102	24.41 ± 0.48 (23.09–25.71) 109
BB	12.00	11.90 ± 0.28 (11.24–12.70) 102	11.89 ± 0.29 (11.09–12.65) 109
ZB	18.36	17.56 ± 0.33 (16.87–18.48) 103	17.40 ± 0.37 (16.33–18.47) 109
PB	7.14	6.70 ± 0.21 (6.20–7.35) 103	6.72 ± 0.19 (6.33–7.28) 109
C–C	7.00	7.02 ± 0.21 (6.58–7.67) 103	6.95 ± 0.23 (6.35–7.47) 109
MB	13.23	12.89 ± 0.23 (12.33–13.53) 103	12.83 ± 0.25 (12.20–13.63) 109
PL	14.25	14.50 ± 0.36 (13.68–15.51) 103	14.39 ± 0.36 (13.54–15.44) 109
MTRL	10.43	10.31 ± 0.23 (9.78–10.91) 103	10.23 ± 0.25 (9.68–10.78) 109
MLTRL	8.50	8.57 ± 0.20 (8.14–9.07) 103	8.49 ± 0.22 (7.98–9.04) 109
M1–M1	12.63	12.19 ± 0.30 (11.60–12.94) 103	12.03 ± 0.33 (10.97–12.79) 109
M2–M2	12.58	12.44 ± 0.29 (11.93–13.32) 103	12.30 ± 0.34 (10.95–13.14) 109
MXBR	6.86	6.51 ± 0.21 (6.06–7.05) 103	6.42 ± 0.25 (5.46–6.97) 109
M1W	2.90	2.99 ± 0.16 (2.58–3.37) 103	2.98 ± 0.15 (2.59–3.30) 109
M2W	3.20	3.35 ± 0.12 (3.00–3.74) 103	3.34 ± 0.12 (3.05–3.60) 109
DENL	19.38	19.08 ± 0.36 (18.28–19.81) 102	18.88 ± 0.41 (17.49–19.89) 109
MANDL	11.44	11.56 ± 0.23 (11.07–12.14) 102	11.46 ± 0.26 (10.86–12.06) 109
COH	7.42	7.87 ± 0.28 (7.27–8.55) 102	7.71 ± 0.29 (6.90–8.41) 109
m1W	2.07	2.06 ± 0.08 (1.90–2.28) 102	2.04 ± 0.08 (1.79–2.30) 109

b Holotype of V. major , an adult female. FIGURE 8. Dorsal ( A ) and ventral ( B ) views of the skull of Vampyrodes caraccioli (USNM 405129; male) from Amazonas, Venezuela; the stylohyals were reconstructed from USNM 582872, a female from Cuzco, Peru. Dorsal ( C ) and ventral ( D ) views of the skull of V. major (FMNH 127114; male) from Veracruz, Mexico. been captured in July, September, October, November, December, and January ( Tuttle, 1970 ; Davis and Dixon, 1976 ; Graham, 1987 ; Moya and Arteaga, 2007 ). Lactating females have been captured in October ( Moya and Arteaga, 2007 ). KARYOLOGY: Vampyrodes caraccioli has a diploid chromosome number (2 n ) of 30 and a fundamental number (FN) of 56. The X chromosome is subtelocentric and the Y chromosome is submetacentric ( Baker and Hsu, 1970 ; Baker, 1973 ). REMARKS: The holotype of Vampyrodes ornatus BMNH 24.3.1.63, is an adult female with a hole on the right parietal, missing the right tympanic bula and both m3. Measurements of this holotype are shown in table 4. Forman and Genoways (1979) reported that the head morphology of the sperm of V. caraccioli is unique in being long and having an unusually narrow apex and base.