A new species of Platyrrhinus (Chiroptera: Phyllostomidae) from western Colombia and Ecuador, with emended diagnoses of P. aquilus, P. dorsalis, and P. umbratus

Abstract

The Neotropical bat genus Platyrrhinus (Chiroptera: Phyllostomidae: Stenodermatinae) currently comprises 15 species. Our morphological and morphometric analysis of large and medium-sized Platyrrhinus revealed a distinctive Undescribed species from western South America. We also recognize P. aquilus (Handley & Ferris 1972) and P. umbratus (Lyon 1902) as valid species. We describe P. nitelinea sp. nov. from western Colombia and Ecuador and provide emended diagnoses along with descriptions of P. aquilus, P.. dorsalis, and P. umbratus. Phylogenetic analysis of Platyrrhinus based on morphological characters indicates that P. aquilus is closely related to P. aurarius and P. nigellus, P. umbratus to P. chocoensis, and P. nitelinea to P. vittatus.

Full text

A new species of Platyrrhinus (Chiroptera: Phyllostomidae) from western
Colombia and Ecuador, with emended diagnoses of P. aquilus,P. dorsalis,
and P. umbratus
Pau´l M. Velazco* and Alfred L. Gardner
(PMV) Department of Zoology, Field Museum of Natural History, 1400 S. Lake Shore Dr.,
Chicago, Illinois 60605-2496, U.S.A., and Department of Biological Sciences, University of
Illinois at Chicago, 845 W. Taylor St, Chicago, Illinois 60607, U.S.A.,
e-mail: pvelazco@fieldmuseum.org;
(ALG) USGS Patuxent Wildlife Research Center, National Museum of Natural History MRC-
111, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A., e-mail: gardnera@si.edu
Abstract.—The Neotropical bat genus Platyrrhinus (Chiroptera: Phyllos-
tomidae: Stenodermatinae) currently comprises 15 species. Our morpho-
logical and morphometric analysis of large and medium-sized Platyrrhinus
revealed a distinctive undescribed species from western South America. We
also recognize P. aquilus (Handley & Ferris 1972) and P. umbratus (Lyon
1902) as valid species. We describe P. nitelinea sp. nov. from western
Colombia and Ecuador and provide emended diagnoses along with
descriptions of P. aquilus,P. dorsalis,andP. umbratus. Phylogenetic
analysis of Platyrrhinus based on morphological characters indicates that P.
aquilus is closely related to P. aurarius and P. nigellus,P. umbratus to P.
chocoensis,andP. nitelinea to P. vittatus.
The genus Platyrrhinus (Phyllostomi-
dae: Stenodermatinae) includes at least 15
species of frugivorous bats endemic to the
Neotropics. The combined geographic
ranges of these species extends from
southern Mexico into Paraguay and
northern Argentina (Velazco 2005, Gard-
ner 2008, Velazco & Patterson 2008). The
greatest diversity of Platyrrhinus occurs
along the eastern slopes of the Andes
where as many as four species can be
found sympatrically (e.g., Solari et al.
2006). Platyrrhinus occurs primarily in
tropical lowland and montane forest from
sea level to at least 2550 m (Velazco 2005,
Gardner 2008).
During recent years, the use of
morphometric, morphological, and mo-
lecular techniques to study Platyrrhinus
have enormously improved our knowl-
edge of its taxonomy and phylogenetic
relationships (Velazco & Solari 2003,
Velazco 2005, Velazco & Patterson
2008). The erstwhile species P. dorsalis,
P. helleri,andP. vittatus all proved to
constitute species complexes comprising
three, three, and two species, respective-
ly. In addition, several junior synonyms
have been resurrected such as P. incarum
for the South American population of P.
helleri (excluding the recently described
P. matapalensis), and P. nigellus, which
previously was treated as a synonym of
P. lineatus (see Velazco & Solari 2003,
Velazco 2005). Recently, Velazco &
Patterson (2008) evaluated the species
relationships in Platyrrhinus (with the
exception of P. chocoensis), using four
molecular markers, and found phyloge-
netic support for all species included in
Gardner (2008) and Velazco (2005) and
identified the Andes as one of the most
* Corresponding author.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
122(3):249–281. 2009.

important regional centers for Platyrrhi-
nus diversification.
While reviewing the collections of
several museums and examining the
holotypes of all described species of
Platyrrhinus, we found that P. aquilus
(Handley & Ferris, 1972) and P. umbratus
(Lyon, 1902), both generally regarded as
synonyms of P. dorsalis (Thomas, 1900),
deserve specific status. We also found
specimens of a large distinctive Platyr-
rhinus from the Pacific lowlands of
southwestern Colombia and southwestern
Ecuador that appears to represent an
undescribed species. Herein, we provide
evidence supporting the recognition of P.
aquilus and P. umbratus as valid species
and describe a new species from western
Colombia and Ecuador. Also resulting
from our investigations is an improved
understanding of the distributions of
medium-sized and larger species of Pla-
tyrrhinus, particularly of P. aurarius
(including the first confirmed records
from Brazil) and P. nigellus as is reflected
in the list of additional specimens exam-
ined (Appendix I).
Material and Methods
Lists of specimens examined are in their
respective species accounts with a list of
additional specimens provided in Appen-
dix I. The following institutional names
and abbreviations are used in the text,
figures, tables, and the appendix: Natu-
ral History Museum, London, U.K.
(BMNH); Carnegie Museum of Natural
History, Pittsburgh, Pennsylvania, U.S.A.
(CM); Colec¸a˜o de Mamı
´feros da Uni-
versidade Federal de Lavras, Lavras,
Minas Gerais, Brazil (CMUFLA); The
Field Museum, Chicago, Illinois, U.S.A.
(FMNH); Instituto de Investigacio´n de
Recursos Biolo´ gicos Alexander von
Humboldt, Villa de Leyva, Boyaca´, Co-
lombia (IAvH-M); Instituto de Ciencias
Naturales, Universidad Nacional de Co-
lombia, Bogota´ , Colombia (ICN); Uni-
versidad de los Andes, Bogota´ , Colombia
(ULA); Museum of Natural Science,
Louisiana State University, Baton Rouge,
Louisiana, U.S.A. (LSUMZ); Museum of
Comparative Zoology, Harvard Univer-
sity, Cambridge, Massachusetts, U.S.A.
(MCZ); Muse´um National d’Histoire
Naturelle, Paris, France (MNHN); Mu-
seo de Ciencias Naturales de la Universi-
dad de Antioquia, Medellı
´n, Colombia
(MUA); Museo de Historia Natural de la
Universidad Nacional Mayor de San
Marcos, Lima, Peru (MUSM); Museum
of Vertebrate Zoology, University of
California, Berkeley, California, U.S.A.
(MVZ); Royal Ontario Museum, Tor-
onto, Ontario, Canada (ROM); Field
numbers of Rafael Zerbini Coutinho
(RZ)—specimens deposited in the Cole-
c¸a˜ o de Mamı
´feros do Departamento de
Zoologia da Universidade Federal de
Minas Gerais, Minas Gerais, Brazil
(DZ-UFMG); Staatliches Museum fu¨r
Naturkunde Stuttgart, Baden-Wu¨rttem-
berg, Germany (SMNS); Texas Coopera-
tive Wildlife Collection, Texas A&M
University, College Station, Texas,
U.S.A. (TCWC); Museo de Historia
Natural de la Universidad de la Amazo-
nia, Florencia, Caqueta´, Colombia
(UAM); Museum of Zoology, University
of Michigan, Ann Arbor, Michigan,
U.S.A. (UMMZ); National Museum of
Natural History, Smithsonian Institution,
Washington, D.C., U.S.A. (USNM); Sec-
cio´n de Zoologı
´a, Departamento de
Biologı
´a, Universidad del Valle, Cali,
Colombia (UV); Field numbers of Vale´ria
da Cunha Tavares (VCT); Zoologisches
Forschungsmuseum Alexander Koenig,
Bonn, North Rhine-Westphalia, Ger-
many (ZFMK); Museum fu¨ r Naturkunde
der Humboldt-Universita¨t zu Berlin, Ber-
lin, Germany (ZMB).
All observations reported here are
based on adults as defined by closed
wing-bone epiphyses and fused cranial
sutures. All linear measurements are in
millimeters (mm); weights, in grams.
250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Standard external measurements (TL,
total length; HF, hind foot; E, ear) are
those recorded on the specimen labels.
Definitions for measurements not defined
here can be found in Velazco (2005).
Certain measurements (with their abbre-
viations) are illustrated in Figure 1. Fore-
arm length and 21 cranial and dental
dimensions were measured with a digital
caliper and recorded to the nearest
0.01 mm as follows: GLS, greatest length
of skull; CIL, condyloincisive length;
CCL, condylocanine length; BB, breadth
of braincase; ZB, zygomatic breadth; PB,
postorbital breadth; C–C, palatal width
at canines (width across palate between
cingula of upper canines); MB, mastoid
breadth; PL, palatal length (distance
between posterior palatal notch to ante-
rior border of the incisive alveolus);
MTRL, maxillary toothrow length;
MLTRL, molariform toothrow length
(posterior border of M3 alveolus to
anterior border of P3); M1–M1, width
across first upper molars; M2–M2, width
across second upper molars; MXBR,
breadth across maxillae (least breadth
across maxillae between the lingual mar-
gins of M2s); M1W, greatest width of
crown of M1; M2W, greatest width of
crown of M2; DENL, length of dentary
(distance from midpoint of mandibular
condyle to anterior-most margin of den-
tary); MANDL, length of mandibular
toothrow (distance from anterior-most
surface of the lower canine to the
posterior-most surface of m3); COH,
coronoid height (perpendicular height
from ventral surface of the mandible to
tip of coronoid process); WMC, width at
mandibular condyles (greatest width be-
tween inner margins of mandibular con-
dyles; m1W, crown width of m1; FA,
forearm length.
In our metric comparisons of Platyr-
rhinus aquilus,P. umbratus,andP. sp.
nov. with other species of Platyrrhinus of
similar size, we conducted Principal Com-
ponent Analyses (PCA) using a correla-
tion matrix of 22 measurements (GLS,
CIL, CCL, BB, ZB, PB, C–C, MB, PL,
MTRL, MLTRL, M1–M1, M2–M2,
MXBR, M1W, M2W, DENL, MANDL,
COH, WMC, m1W, FA) from 265
individuals (see specimens examined in
species accounts and Appendix I).
To assess the phylogenetic position of
Platyrrhinus aquilus,P. umbratus,andP.
sp. nov. based on morphology, we ana-
lyzed variation in 60 external, cranial, and
dental characters identified by Velazco
(2005) as useful in assessing relationships
within Platyrrhinus. These characters and
their states are discussed and illustrated in
Velazco (2005) along with descriptions of
species not included here in the Species
Accounts. Character states for the new
species and for P. aquilus and P. umbra-
tus, species not included in Velazco’s
(2005) analysis, are presented in Table 1.
We used PAUP* version 4.0b10 (Swof-
ford 2002) to analyze the resulting matrix
of 21 taxa and 60 character states, using
unordered states and the heuristic search
specifying the tree bisection-reconnection
option. The topology of shortest-length
trees was then subjected to bootstrap
analysis (1000 replicates). The dichoto-
mous identification key, modified from
Gardner (2008), follows the species ac-
counts and includes all species we recog-
nize in Platyrrhinus.
Phylogenetic Analyses
Our phylogenetic analysis of the matrix
of 60 characters from Velazco (2005)
identified a single tree of 186 steps
(Fig. 2; CI 50.478; CI excluding unin-
formative characters 50.458; RI 5
0.559). The single most parsimonious tree
was completely resolved showing unam-
biguous relationships among all species,
both in the ingroup and outgroup taxa
(Fig. 2), although all groupings were not
supported by the bootstrap analysis.
Platyrrhinus monophyly was weakly sup-
ported (60%of bootstraps); however,
VOLUME 122, NUMBER 3 251

Fig. 1. Lateral and dorsal views of the cranium and mandible, and ventral view of the cranium,
illustrating most of the measurements used in this study. For abbreviations see Material and Methods.
252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

other morphological and molecular stud-
ies show that Platyrrhinus is monophylet-
ic (Velazco 2005, Velazco & Patterson
2008). Noteworthy are the cladistic posi-
tions depicted for P. aquilus and P.
umbratus (Fig. 2), two forms that had
been synonymized with P. dorsalis in
recent studies (Velazco 2005, Gardner
2008). Their distant relationship from P.
dorsalis, together with the morphological
and morphometric comparisons present-
ed below, indicate that P. aquilus and P.
umbratus should be recognized as sepa-
rate species. These two species and P. sp.
nov. were recovered in different group-
ings inside Platyrrhinus but with low
support (,50%bootstrap value): P.
vittatus and P. sp. nov. are portrayed as
sister taxa, as are P. chocoensis and P.
umbratus;P. aquilus falls between P.
aurarius and P. nigellus.
Species Accounts
Family Phyllostomidae Gray, 1825
Subfamily Stenodermatinae Gervais,
1856
Genus Platyrrhinus Saussure, 1860
Platyrrhinus aquilus (Handley & Ferris,
1972)
Darien Broad-nosed Bat
Fig. 3
Vampyrops aquilus Handley & Ferris,
1972:521–522; type locality ‘‘on the
head of the Rı
´oPucro,4100ft
[1250 m], Cerro Malı
´,Darie´n, Pana-
ma´.’’
V[ampyrops]dorsalis: Carter & Rouk,
1973:976; not Vampyrops dorsalis Tho-
mas, 1900.
[Platyrrhinus]umbratus: Koopman, 1993:
191; not Platyrrhinus umbratus (Lyon,
1902).
Platyrrhinus dorsalis: Velazco, 2005:24;
not Vampyrops dorsalis Thomas, 1900.
Type material.—The holotype USNM
338025, a lactating female when collected,
and two paratypes: USNM 338026, an
adult male from the type locality, and
USNM 338027, an adult male, all from
Cerro Malı
´(Table 2).
Distribution.—Platyrrhinus aquilus is
known only from Cerro Malı
´and Cerro
Pirre in provincia Darie´n, Panama, at
elevations from 1250–1433 m (Fig. 4).
Table 1.—Morphological character states for Platyrrhinus aquilus,P. umbratus,andP. sp. nov. Character
state defined by Velazco (2005).
1234567891011 121314 1516 17 18 1920
Platyrrhinus aquilus 111011302 2 3 0 1 2 2 1 2 0/1 1 2
Platyrrhinus umbratus 101021312 2 2 0 1 1 2 1 1 1 1 1
Platyrrhinus sp.nov. 111131321 2 3 0 0 1 1 1 1 1 2 2
21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40
Platyrrhinus aquilus 12100111111101120111
Platyrrhinus
umbratus 11/210011111100/12120/1101
Platyrrhinus sp.nov.10100111111002120111
41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60
Platyrrhinus aquilus 10101010200121101221
Platyrrhinus
umbratus 11101010200121200221
Platyrrhinus sp.nov.11111010200121201221
VOLUME 122, NUMBER 3 253

Emended diagnosis.—Platyrrhinus aqui-
lus is a medium-sized bat (FA 45.2–
46.8 mm, GLS 26.5–27.4 mm, CCL
23.6–25.1 mm; Tables 2–4), easily distin-
guished from P. aurarius and P. ismaeli
by its shorter skull, and from P. lineatus,
P. nigellus,P. recifinus,andP. umbratus
by its longer skull (Tables 3, 4). Cranial
measurements overlap those of P. cho-
coensis,P. dorsalis,P. masu,andP.
umbratus. Dorsal fur is blackish, ventral
fur pale brown and gray; facial stripes
well marked but dark; mid-dorsal stripe
whitish and well defined; fold lines in the
pinnae well marked; hair along trailing
edge of the uropatagium dark, long, and
Fig. 2. Most parsimonious tree of the phylogenetic analysis with the characters unordered. Bootstrap
values higher than 50%are presented above each branch.
254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Dorsal and ventral views of the cranium and lateral view of the cranium and mandible of
Platyrrhinus aquilus (USNM 338026, paratype). See Table 2 for measurements.
VOLUME 122, NUMBER 3 255

dense; frontals steeply sloping; postorbital
processes only moderately developed; and
a parastyle present on M1.
Description.—A medium-sized species
of Platyrrhinus with dense, blackish dor-
sal fur; dense, pale brown and gray
ventral fur; dorsal and ventral hair have
three color bands. Dorsomedial and
ventrolateral facial stripes are the same
color (whitish hairs with dusky tips).
Dorsal hairs are 8–10 mm long on the
back; dorsal stripe well defined and
brighter than the facial stripes; fold lines
in the pinnae well marked; noseleaf longer
than wide; lower border of nasal horse-
shoe completely free of upper lip; long,
dense hair on upper surface of feet;
inverted ‘‘U-shaped’’ porterior margin of
uropatagium bordered with dense fringe
of long (4–4.5 mm), dark hair; uropata-
gium (dry) extends 2–3.5 mm beyond
body at midline; proximal 0.67 of forearm
densely covered with long hair; plagiopa-
tagium attached to leg on metatarsal I;
metacarpal III shorter than metacarpal V;
posterior border of hard palate an invert-
ed ‘‘U’’-shape in USNM 338025 and
338026 but closer to ‘‘V-shaped’’ in
USNM 308027; postorbital processes
moderately developed; paraoccipital pro-
cesses well developed; fossa on the squa-
mosal portion of the zygomatic arch
shallow; upper inner incisors convergent
and in contact; upper outer incisors
monolobate; two stylar cuspules on pos-
terior crista of P4; deep fossa on hypoco-
nal basin of P4; parastyle present on M1,
but mesostyle absent; labial and lingual
cingula present on M1 metacone; sulcus
on posterior crista of paracone continuous
with groove bordered by lingual cingulum
of M1 metacone; M1 metastyle present;
M1 protocone moderately developed;
labial cingulum present on M2 paracone
and metacone; stylar cuspule on lingual
face of the M2 paracone and metacone
absent; M2 metastyle present; stylar
cuspule on the lingual face of the M2
metacone absent; lingual cingulum of M2
metacone not joined with paracone; M2
hypoconal basin developed; labial and
lingual cingulids of p4 present; stylar
cuspulids on anterior and posterior cris-
tids of p4 lacking; m1 paraconid absent;
m1 metaconid well developed; m2 hypo-
conid present; stylid cuspulid present
between the metaconid and protoconid
on m2; labial and lingual cingulids present
on m2.
Comparisons.—Platyrrhinus aquilus oc-
curs sympatrically with P. chocoensis,P.
dorsalis,P. helleri,andP. vittatus.Platyr-
rhinus aquilus is easily distinguished from
P. vittatus and P. helleri on the basis of
size; P. helleri (FA 35–40 mm, GLS 19.0–
22.5 mm; Velazco 2005) is much smaller,
and P. vittatus (FA 56.7–61.9 mm, GLS
30.6–32.8 mm; Tables 3, 4) is much larg-
er. P. aquilus has been confused with P.
dorsalis and P. umbratus,anditsmea-
surements overlap those of P. chocoensis
(Tables 3, 4). Therefore, the following
comparison focuses on differentiating
these four species.
Externally, Platyrrhinus aquilus can be
distinguished from these species by its
darker, almost black dorsal color (P.
chocoensis,P. dorsalis,andP. umbratus
are pale brown to brown dorsally); the
Table 2.—Measurements (mm) of type material of
Platyrrhinus aquilus.
Character
Holotype
USNM
338025 R
Paratype
USNM
338026 =
Paratype
USNM
338027 =
GLS 27.3 27.3 26.9
CIL 25.6 25.5 25.0
CCL 24.9 25.1 24.5
BB 11.5 11.2 11.0
ZB 15.4 15.6 15.0
PB 6.1 6.3 6.2
MB 12.3 12.0 11.7
MTRL 10.4 10.7 10.6
M2–M2 11.5 11.8 11.4
DENL 19.2 19.2 19.0
MANDL 11.4 11.8 11.6
FA 45.2 45.4 46.8
TL 78.0 74.0 70.0
HF 12.0 12.3 10.8
E 20.0 20.0 19.0
256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

hairs of the fringe on the trailing edge of
the uropatagium in P. aquilus are long (4–
4.5 mm) and abundant (shorter [less than
4 mm, usually less than 3 mm] and sparse
in P. chocoensis and P. dorsalis); facial and
dorsal stripes are brighter and well defined
in P. aquilus and P. dorsalis (thin and not
contrasting as strongly with the surround-
ing fur in P. chocoensis and P. umbratus);
fold lines in the pinnae are well developed
in P. aquilus,P. chocoensis,andP.
umbratus (poorly marked in P. dorsalis);
ventral fur has three color bands in P.
aquilus and P. dorsalis (two in P. chocoen-
sis and P. umbratus); hair on upper surface
of feet is long and dense in P. aquilus and
P. umbratus (comparatively short and
sparse in P. chocoensis and P. dorsalis).
Cranially, the postorbital processes are
moderately developed in Platyrrhinus
aquilus and P. umbratus (poorly devel-
oped in P. chocoensis and P. dorsalis). In
addition, P. aquilus has well-developed
paraoccipital processes (only moderately
Fig. 4. Map of northwestern South America, illustrating the distribution of specimens of Platyrrhinus
aquilus (dots), P. umbratus (stars), and P. nitelinea (triangles). Elevations: no shading, 0–1000 m; pale gray,
1000–2000 m; dark gray, .2000 m.
VOLUME 122, NUMBER 3 257

Table 3.—Selected measurements
a
(mm) of Platyrrhinus species.
Character P. albericoi P. aquilus P. aurarius P. chocoensis P. dorsalis P. infuscus P. ismaeli
GLS 32.0 60.68 27.1 60.31 28.2 60.46 27.6 60.64 27.2 60.51 30.8 60.63 29.0 60.59
31.7–33.9 (15) 26.5–27.4 (7) 27.4–29.3 (22) 26.5–29.0 (21) 25.9–28.2 (16) 29.8–31.8 (14) 28.0–30.4 (22)
CCL 30.3 60.59 24.4 60.53 25.6 60.46 24.8 60.66 24.9 60.46 27.8 60.55 26.5 60.51
29.4–31.2 (15) 23.6–25.1 (7) 24.9–26.9 (22) 23.6–25.9 (21) 23.8–25.7 (16) 27.1–28.7 (14) 25.6–27.6 (22)
CIL 31.1 60.62 24.9 60.59 26.3 60.48 25.2 60.67 25.4 60.56 28.5 60.52 27.1 60.49
30.0–31. 9 (15) 24.1–25.6 (7) 25.6–27.4 (22) 24.3–26.6 (21) 24.1–26.3 (16) 27.8–29.3 (14) 26.4–28.2 (22)
BB 13.6 60.29 11.3 60.22 11.9 60.25 11.8 60.19 11.5 60.23 12.9 60.32 12.0 60.28
13.0–14.1 (15) 11.0–11.6 (7) 11.4–12.4 (22) 11.3–12.2 (21) 10.9–11.8 (16) 12.1–13.4 (14) 11.6–12.5 (22)
ZB 20.2 60.41 15.2 60.32 16.8 60.44 16.5 60.43 15.8 60.54 18.4 60.59 17.2 60.56
19.4–20.8 (15) 14.7–15.6 (7) 15.9–17.7 (22) 15.6–17.2 (21) 14.9–16.5 (16) 17.6–19.3 (14) 16.2–18.2 (22)
PB 7.3 60.16 6.3 60.10 6.7 60.20 6.4 60.21 6.3 60.20 6.9 60.22 6.5 60.21
7.0–7.5 (15) 6.1–6.4 (10) 6.4–7.3 (22) 6.0–6.8 (21) 5.9–6.6 (16) 6.6–7.3 (14) 6.2–6.9 (22)
MB 15.0 60.29 12.0 60.21 13.0 60.30 12.6 60.19 12.3 60.32 13.8 60.30 13.0 60.28
14.3–15.4 (15) 11.7–12.3 (7) 12.4–13.6 (22) 12.2–12.9 (21) 11.8–12.9 (16) 13.4–14.3 (14) 12.4–13.6 (22)
PL 18.3 60.33 14.0 60.48 14.9 60.36 14.1 60.45 14.5 60.60 16.4 60.49 15.70 60.44
17.6–18.8 (15) 13.4–14.8 (7 14.0–15.6 (22) 13.2–15.0 (21) 12.9–15.2 (16) 15.5–17.0 (14) 14.8–16.4 (22)
MTRL 13.6 60.29 10.4 60.25 11.1 60.25 10.7 60.32 10.9 60.43 12.2 60.28 11.8 60.38
13.2–14.1 (15) 10.0–10.9 (10) 10.5–11.5 (22) 10.1–11.3 (21) 10.0–11.4 (16) 11.8–12.8 (14) 11.2–12.5 (22)
M1–M1 14.6 60.37 10.8 60.35 11.9 60.40 11.0 60.36 11.4 60.48 13.4 60.45 12.5 60.49
13.7–15.2 (15) 10.3–11.4 (10) 11.1–12.6 (22) 10.2–11.6 (21) 10.3–12.1 (16) 12.6–14.2 (14) 11.6–13.3 (22)
M2–M2 15.3 60.29 11.4 60.33 12.6 60.38 11.5 60.32 11.9 60.47 13.6 60.41 13.1 60.51
14.7–15.6 (15) 10.8–11.8 (10) 11.9–13.4 (22) 10.7–12.1 (21) 10.9–12.5 (16) 12.9–14.2 (14) 13.3–14.0 (22)
DENL 24.2 60.51 19.0 60.32 20.0 60.35 19.6 60.54 19.5 60.41 22.17 60.51 20.9 60.39
23.3–25.3 (15) 18.5–19.4 (10) 19.3–20.8 (22) 18.8–20.5 (21) 18.6–20.1 (15) 21.5–22.8 (14) 20.3–21.8 (22)
MANDL 14.5 60.29 11.5 60.29 11.9 60.26 11.6 60.31 11.8 60.37 13.3 60.28 12.7 60.42
14.1–14.9 (15) 11.0–11.8 (10) 11.4–12.6 (22) 11.0–12.1 (21) 11.1–12.3 (15) 12.9–13.9 (14) 11.9–13.4 (22)
FA 59.9 61.52 45.8 60.85 52.2 61.03 48.8 61.20 47.6 60.89 57.3 61.42 52.4 61.68
57.4–62.6(15) 45.2–46.8 (3) 50.5–54.3 (22) 46.9–50.7 (19) 46.6–49.5 (16) 55.1–60.6 (14) 49.2–56.2 (22)
a
Summary statistics (mean and standard deviation followed by range and sample size) of measurements for each species. See lists of specimens examinedin
species accounts and Appendix I for catalog numbers of the specimens measured.
258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.—Selected measurements
a
(mm) of Platyrrhinus species.
Character P. lineatus P. masu P. nitelinea P. nigellus P. recifinus P. umbratus P. vittatus
GLS 24.6 60.44 26.9 60.47 30.6 60.61 25.3 60.67 23.8 60.72 25.9 60.35 31.7 60.62
23.8–25.5 (26) 25.9–27.4 (15) 29.3–31.9 (48) 23.8–26.8 (57) 23.1–24.5 (3) 25.1–26.7 (36) 30.6–32.8 (33)
CCL 22.1 60.42 24.3 60.39 28.2 60.64 23.0 60.70 21.6 60.48 23.7 60.35 29.1 60.50
21.5–23.0 (26) 23.4–24.9 (15) 26.9–29.6 (48) 21.5–24.5 (57) 21.0–21.9 (3) 23.0–24.4 (36) 28.4–30.0 (33)
CIL 22.7 60.46 24.8 60.41 28.8 60.65 23.5 60.72 22.1 60.40 24.2 60.40 29.8 60.51
21.9–23.5 (26) 23.9–25.4 (15) 27.6–30.5 (48) 21.9–25.2 (57) 21.7–22.5 (3) 23.4–25.1 (34) 29.0–30.7 (33)
BB 10.9 60.26 11.7 60.21 12.7 60.20 10.8 60.20 10.3 60.00 10.9 60.27 13.2 60.27
10.3–11.5 (28) 11.3–12.0 (15) 12.3–13.2 (48) 10.3–11.5 (57) 10.3 (3) 10.4–11.6 (36) 12.6–13.7 (33)
ZB 14.5 60.43 16.0 60.19 18.4 60.44 14.4 60.43 14.1 60.17 14.9 60.33 19.1 60.49
13.9–15.3 (27) 15.6–16.3 (15) 17.3–19.6 (47) 13.4–15.5 (57) 14.0–14.3 (3) 14.4–15.5 (34) 18.3–20.2 (33)
PB 6.3 60.18 6.3 60.18 7.2 60.16 6.1 60.18 5.7 60.11 6.2 60.21 7.6 60.22
5.9–6.6 (28) 6.0–6.6 (15) 6.7–7.5 (48) 5.6–6.5 (57) 5.6–5.8 (3) 5.6–6.6 (36) 7.1–8.2 (33)
MB 11.5 60.26 12.3 60.19 13.8 60.30 11.5 60.26 11.2 60.16 11.8 60.20 14.5 60.35
11.0–12.0 (28) 12.1–12.7 (15) 13.1–14.5 (48) 10.8–12.2 (57) 11.0–11.3 (3) 11.4–12.2 (36) 13.7–15.1 (33)
PL 12.0 60.35 13.9 60.34 16.2 60.51 12.8 60.46 11.9 60.40 13.1 60.38 16.6 60.40
11.2–12.5 (28) 13.2–14.4 (15) 15.0–17.2 (48) 11.8–13.5 (57) 11.5–12.3 (3) 12.2–13.9 (34) 15.8–17.6 (33)
MTRL 8.9 60.21 10.5 60.18 12.5 60.35 9.7 60.40 8.9 60.03 10.1 60.18 12.9 60.33
8.5–9.4 (28) 10.1–10.7 (15) 11.8–13.4 (48) 8.8–10.5 (57) 8.9–9.0 (3) 9.7–10.5 (36) 12.3–13.7 (33)
M1–M1 9.8 60.28 11.3 60.20 13.1 60.44 10.3 60.46 10.0 60.20 10.7 60.29 13.7 60.40
9.1–10.4 (28) 10.9–11.7 (15) 12.4–14.5 (48) 9.4–11.3 (57) 9.8–10.2 (3) 10.0–11.4 (35) 13.0–14.6 (33)
M2–M2 10.2 60.26 12.0 60.20 13.9 60.41 10.6 60.28 10.3 60.22 11.0 60.28 14.2 60.36
9.5–10.8 (28) 11.5–12.4 (15) 13.1–15.1 (48) 10.0–11.1 (57) 10.1–10.5 (3) 10.5–11.8 (36) 13.5–15.0 (33)
DENL 16.7 60.46 19.1 60.29 22.6 60.56 17.5 60.59 16.5 60.42 18.0 60.36 23.0 60.41
16.2–18.0 (28) 18.5–19.4 (15) 21.4–23.7 (48) 16.3–19.0 (57) 16.0–16.8 (3) 17.3–18.7 (36) 22.1–23.9 (33)
MANDL 9.8 60.23 11.3 60.20 13.5 60.35 10.5 60.40 9.7 60.13 10.9 60.23 13.8 60.31
9.4–10.3 (28) 11.0–11.7 (15) 13.0–14.6 (48) 9.7–11.4 (57) 9.6–9.8 (3) 10.4–11.4 (36) 13.2–14.5 (33)
FA 47.1 60.81 48.5 61.42 55.5 61.38 44.0 61.79 43.0 60.80 45.4 61.33 59.1 61.25
46.0–48.6 (26) 46.5–51.0 (15) 52.9–58.4 (46) 40.6–48.0 (54) 42.4–43.5 (2) 42.0–47.8 (36) 56.7–61.9 (33)
a
Summary statistics (mean and standard deviation followed by range and sample size) of measurements for each species. See lists of specimens examinedin
species accounts and Appendix I for catalog numbers of the specimens measured.
VOLUME 122, NUMBER 3 259

developed in P. chocoensis,P. dorsalis,
and P. umbratus).
Dentally, Platyrrhinus aquilus,P. cho-
coensis,P. dorsalis,andP. umbratus differ
as follows: the M1 in P. aquilus has a
parastyle (absent in P. chocoensis,P.
dorsalis,andP. umbratus); P. aquilus
lacks a stylar cuspule on the labial
cingulum of the M1 metacone (present
in P. chocoensis,P. dorsalis,andP.
umbratus); P. aquilus has a moderately
developed M1 protocone (small and blunt
in P. chocoensis and P. dorsalis but well
developed in P. umbratus); P. aquilus
lacks a stylar cuspule on the lingual face
of the M2 paracone (present in P.
umbratus; variably developed in P. cho-
coensis and P. dorsalis); P. aquilus,P.
chocoensis,andP. umbratus lack a stylar
cuspule on the lingual face of the M2
metacone (present in P. dorsalis); the
lingual cingulum of the M2 metacone is
restricted to the metacone in P. aquilus
and P. umbratus (continuous to paracone
in P. chocoensis and P. dorsalis); both
labial and lingual cingulids are present on
p4 in P. aquilus,P. dorsalis,andP.
umbratus (only labial cingulid present in
P. chocoensis); m1 metaconid is weakly
developed in P. aquilus (lacking in P.
chocoensis,P. dorsalis,andP. umbratus);
m1 hypoconid present in P. aquilus and P.
umbratus (absent in P. chocoensis and P.
dorsalis); a stylid cuspulid is present
between the metaconid and protoconid
of m2 in P. aquilus and P. dorsalis (absent
in P. chocoensis and P. umbratus).
Multivariate analysis.—We compared
the type material of Platyrrhinus aquilus
with 21 P. chocoensis (Colombia and
Panama), 13 P. dorsalis (Ecuador and
Colombia), and 36 P. umbratus (Colombia
and Venezuela). The first three PCs ac-
counted for 82.4%of the overall variation.
A plot of factor scores on the first two axes
(Fig. 5) shows that the type material of P.
aquilus and P. chocoensis overlaps com-
pletely on PC1, which represent overall size
(Table 5). In contrast, P. aquilus falls
between P. dorsalis and P. umbratus,with
P. dorsalis being larger and P. umbratus
smaller than P. aquilus. Along the PC2
axis, P. aquilus overlaps nearly completely
with P. dorsalis and P. umbratus but is
entirely separated from P. chocoensis,
reflecting its lower coronoid, narrower
braincase, narrower palate at canines,
and shorter forearm (Fig. 5, Table 5).
Remarks.—A year after its description,
Carter & Rouk (1973) treated Platyrrhi-
nus aquilus as a junior synonym of P.
dorsalis stating ‘‘V. aquilus Handley &
Ferris is unlike either of the species
named and described as new to science
by Rouk & Carter (1972) or Gardner &
Carter (1972a), but it does not differ
substantially from Peruvian specimens of
V. dorsalis reported by Gardner & Carter
(1972b).’’ Subsequently, P. aquilus has
been treated either as a junior synonym of
P. dorsalis (see Alberico 1990, Velazco
2005, Gardner 2008) or as a synonym of
P. umbratus (see Koopman 1993, Sim-
mons 2005). After examining the three
specimens constituting the type material
of P. aquilus and comparing them with all
specimens of Platyrrhinus available to us
in the FMNH and USNM, we find P.
aquilus to be distinctive morphologically
and morphometrically; therefore, we rec-
ognized it as a valid species.
Specimens examined (10).—Three spec-
imens (marked with an asterisk) were
used in the elaboration of Tables 2–5 and
in the morphometric analyses. PANA-
MA: Darie´n: Cerro Mali, Head of Rı
´o
Pucro (USNM 338025* [Holotype of
Vampyrops aquilus], 338026*); Cerro Mali
(USNM 338027*); 9 km NW of Cana on
the slopes of Cerro Pirre (LSUMZ 25123,
25126, 25131, 25132, 25139–25141).
Platyrrhinus dorsalis (Thomas, 1900)
Thomas’s Broad-nosed Bat
Vampyrops dorsalis Thomas, 1900:269–
270; type locality ‘‘Paramba, [Imba-
bura,] N. Ecuador. Alt. 1100 m.’’
260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Platyrrhinus dorsalis: Alberico & Velasco,
1991:237; part; first use of current name
combination.
Type material.—The holotype of Pla-
tyrrhinus dorsalis, BMNH 99.12.5.1, is a
subadult male missing the left M3.
Distribution.—Platyrrhinus dorsalis is
found at elevations from 150 to above
2000 m in Colombia and south into
Ecuador along both slopes of the Andes.
Emended diagnosis.—Platyrrhinus dor-
salis is a medium-sized bat (FA 46.6–
49.5 mm, GLS 25.9–28.2, CCL 23.8–
25.7 mm; Tables 3, 4) that has dark
brown dorsal fur, brownish ventral fur;
pale brownish upper and lower facial
stripes; narrow dorsal stripe; lacks obvi-
ous fold lines in the pinnae; a small, blunt
protocone on M1; only the labial cingu-
lum present on P4; stylar cuspule present
on lingual face of metacone on M2; both
labial and lingual cingulids present on p4.
Remarks.—Gardner (2008) and Ve-
lazco (2005) included the names Vampyr-
ops aquilus,V. oratus,andV. umbratus as
synonyms of P. dorsalis. Here we treat P.
aquilus and P. umbratus as valid species
and relegate V. oratus to the synonymy of
P. umbratus. Specimens that Gardner
(2008) and Velazco (2005) reported as P.
dorsalis from Venezuela represent P.
umbratus. Specimens that Gardner
(2008) reported from Panama (Darie´n)
as P. dorsalis represent P. aquilus.
Fig. 5. Plot of scores on first and second axes from PCA of 22 variables from 3 Platyrrhinus aquilus,21
P. chocoensis,13P. dorsalis,and36P. umbratus (see Table 5).
VOLUME 122, NUMBER 3 261

Specimens examined (144).—Individu-
als or series marked with an asterisk were
used in the elaboration of Tables 3–6 and
in the morphometric analyses. COLOM-
BIA: Antioquia: Cocorna, Vereda La
Granja, quebrada La Granja (ICN
9875); Urrao, Calles (MUA 10776). Boy-
aca´: Almeida, Corregimiento Chivor, ver-
eda Camoyo, cerca al tunel Chamizo
(ICN 8868); Villa de Leyva, al S de la
poblacio´n Cabeceras de la Quebrada del
Jacal (camino a Chiquisa) (ICN 656).
Cauca: Alto Micay, Betania (FMNH
133372, 113380, 113385, 113394–113396);
Buenos Aires, El Ceral, Finca El Dia-
mante (UV 2165); Charguayaco (FMNH
113538, 113539*); Parque Natural Na-
cional ‘‘Munchique,’’ Caban˜ a La Rome-
lia (IAvH-M 3313); Popaya´ n (FMNH
90327*). Choco´:Bahı
´a Solano (Mutis),
5 km by road SW Alto de Gala´ pagos
(UV 4571); San Jose´ del Palmar, 4 km N
La Italia (UV 4575, 10034–10035); San
Jose´ del Palmar, Alto del Oso, 10 km W
La Italia (UV 4559–4561, 10837); San
Jose´ del Palmar, Quebrada La Guagua,
8 km W La Italia (UV 7448). Cundina-
marca: San Francisco, Vereda San Miguel
(ICN 8742). Huila: Las Cuevas Parque,
Upper Caban˜a (FMNH 58740). Meta:
Restrepo (UV 3851). Narin˜o:ElCarmen
(FMNH 113397, 113890, 113893); Barba-
coas, Junı
´n, Planada de Maindes (UV
3050–3055); Ricaurte, Hacienda La Pla-
nada (Buenos Aires) (UV 2942–2957).
Quindı
´o: Municipio Ge´nova, Vereda El
Dorado, Bosque San Isidro (IAvH-M
7040); Municipio Pereira, Can˜on del Rı
´o
Barbas (IAvH-M 7038). Risaralda: Pueb-
lo Rico, Camino a La Bocatoma (ICN
11518); Pueblo Rico, Santa Cecilia, 8 km
approx. Plosan (ICN 12268, 12269);
Pueblo Rico, Vereda San Jose´, Quebrada
San Jose´ (ICN 11517, 11932, 11933);
Table 5.—Factor loadings for first three axes
from PCA of 22 variables from Platyrrhinus aquilus,
P. chocoensis,P. dorsalis, and P. umbratus.
Variable
Correlations
PC 1 PC 2 PC 3
GLS 0.936 0.112 20.071
CIL 0.921 20.170 20.125
CCL 0.926 20.125 20.108
BB 0.842 0.365 20.197
ZB 0.899 0.304 20.042
PB 0.548 0.104 20.510
C–C 0.877 0.336 0.007
MB 0.892 0.281 20.123
PL 0.894 20.263 0.024
MTRL 0.938 20.187 0.096
MLTRL 0.802 20.487 0.095
M1–M1 0.813 20.425 20.111
M2–M2 0.835 20.394 0.041
MXBR 0.669 20.475 20.383
M1W 0.593 0.213 0.450
M2W 0.762 0.144 0.519
DENL 0.964 0.063 0.020
MANDL 0.931 20.152 0.110
COH 0.776 0.502 0.080
WMC 0.671 0.375 20.094
m1W 0.624 20.398 0.376
FA 0.760 0.301 20.033
Proportion of
variation 67.4%9.7%5.3%
Table 6.—Factor loadings for first three axes
from PCA of 22 variables from Platyrrhinus
chocoensis,P. dorsalis,P. nigellus, and P. umbratus.
Variable
Correlations
PC 1 PC 2 PC 3
GLS 0.958 0.030 20.063
CIL 0.958 20.178 20.038
CCL 0.958 20.156 20.038
BB 0.862 0.272 20.233
ZB 0.932 0.184 20.145
PB 0.640 20.025 20.459
C–C 0.919 0.262 20.044
MB 0.936 0.147 20.166
PL 0.929 20.165 0.078
MTRL 0.957 20.128 0.103
MLTRL 0.881 20.328 0.130
M1–M1 0.901 20.164 0.154
M2–M2 0.883 20.346 0.007
MXBR 0.806 20.460 20.170
M1W 0.725 0.235 0.284
M2W 0.694 0.436 0.469
DENL 0.969 0.020 20.016
MANDL 0.956 20.096 0.115
COH 0.846 0.401 20.067
WMC 0.759 0.260 20.225
m1W 0.686 20.085 0.552
FA 0.863 0.038 20.130
Proportion of
variation 75.7%5.7%5.0%
262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Santa Rosa de Cabal, Laguna del Otun
(UV 2519). Santander: Charala, Vereda
El Salitre, Finca El Mirador, Cueva Aı
´da
(ICN 17502, 17503); Encino, Vereda Rı
´o
Negro, Sitio Cachalu´ , Finca La Desdi-
chada (ICN 17583); Mogotes, El Hoyo
(ICN 5536). Valle del Cauca: Buenaven-
tura, Quebrada La Delfina (UV 11728);
Buenaventura, San Antonio de Anchi-
caya, Bajo Anchicaya (UV 2631–2634,
2636); Buga, La Magdalena, El Janeiro,
Hacienda Santelina (UV 11701, 11829,
11952); Buga, San Jose´, Nogales, Cuenca
Rı
´oTulu´a (UV 11223, 11224); Cali,
Campamento Corea (UV 2557); Cali,
Finca Coro, Km 18, carretera Cali-
Buenaventura (UV 7175, 7177, 7178,
7180); Cali, Kilometro 18, San Antonio
(UV 3529); Cali, Pen˜ as Blancas-Carretera
a Cristo Rey (UV 2310–2313); Calima,
Finca La Guayacana, 2 km W del muro
(UV 3414–3423); Calima, Rı
´oBravo,
campamento CVC (ICN 9369); Calima,
Vereda Rı
´o Bravo, proyecto Calima III
CVC, Estacio´n Rı
´o Bravo (ICN 8945,
8946, 8948); Dagua, El Queremal, Esta-
cio´n Tokio (UV 3425–3427); Dagua, La
Cascada, Vereda La Cascada, en cerca-
nı
´as Quebrada La Estrella (ICN 13083–
13085); Dapa, 12 mi NW Cali (USNM
483573*); El Cairo, Estacio´ n Cerro de
Ingle´s (UV 12239); El Cairo, Quebrada
Charco Azul, via a San Jorge del Palmar
(UV 12305, 12308); El Silencio (UMMZ
169038); Florida, Hacienda Los Alpes
(UV 3521–3524, 3527, 3528, 7529, 7530,
10578–10581, 10833–10835); La Cumbre,
Finca La Cataisa, 3 km S, 2 km E Bitaco
(UV 7328); Pance, approx. 20 km SW
Cali (USNM 483580–483585*, 483593*,
483598*; UV 806, 1243); Pichinde´, 10 km
SW Cali (USNM 483577*, 483578); Toro,
Vereda Ventaquemada, Finca Monte
Bonito (UV 12110). ECUADOR: Imba-
bura, Paramba (BMNH 99.12.5.1* [Ho-
lotype of Vampyrops dorsalis]). El Oro:
Minas Miranda, 3 km N Zaruma (USNM
534249*); Pastaza: Mera (USNM 548216,
548225).
Platyrrhinus umbratus (Lyon 1902)
Shadowy Broad-nosed Bat
Fig. 6
Vampyrops lineatus: Bangs, 1900:100; not
Phyllostoma lineatum Geoffroy St.-Hi-
laire, 1810.
Vampyrops umbratus Lyon, 1902:151–
152; type locality ‘‘San Miguel,’’ La
Guajira, Colombia.
Vampyrops oratus Thomas, 1914:411–
412; type locality ‘‘Galifari, Sierra del
Avila, [Distrito Federal,] N. Venezuela.
Alt. 65009.’’
Platyrrhinus umbratus: Koopman, 1993:
191; first use of current name combi-
nation.
Type material.—The holotype, MCZ
B8180, is an adult female. There are two
paratypes: MCZ B8300 is an adult male
from ‘‘San Antonio,’’ La Guajira, Co-
lombia; MCZ B8301 is an adult male
from ‘‘Palamina’’ (5Palomino), La Gua-
jira, Colombia. The holotype is the only
specimen from the hypodigm that still has
an associated skull, the paratypes are
skins only. Proximal portions of the
forearms from all three specimens were
removed during preparation.
Distribution.—Platyrrhinus umbratus is
found in Colombia and northern Vene-
zuela (Fig. 4) where the species has been
found at elevations from approximately
250toover2000m.
Measurements of the holotype (mm).—
MCZ B8180: GLS 26.3; CIL 24.6; CCL
24.0; BB 11.2; ZB 15.4; PB 6.4; C–C 6.8;
MB 12.2; PL 13.4; MTRL 10.3; MLTRL
8.6; M1–M1 11.0; M2–M2 11.0; MXBR
6.9; M1W 2.1; M2W 2.5; DENL 18.4;
MANDL 11.0; COH 6.0; WMC 7.5;
m1W 1.9; FA [44.1; incomplete].
Emended diagnosis.—Platyrrhinus um-
bratus is a medium-sized bat (FA 42.0–
47.8 mm, GLS 25.1–26.7 mm, CCL 23.0–
24.4 mm; Tables 3, 4) most easily distin-
guished from P. aquilus,P. aurarius,and
P. ismaeli by its shorter skull, and from P.
recifinus by its longer skull (Tables 3, 4).
VOLUME 122, NUMBER 3 263

Fig. 6. Dorsal and ventral views of the cranium and lateral view of the cranium and mandible of
Platyrrhinus umbratus (USNM 370410).
264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Most measurements of P. umbratus over-
lap those of P. chocoensis,P. dorsalis,P.
lineatus,P. masu,andP. nigellus (Ta-
bles 3, 4). Dorsal and ventral fur is pale to
dark brown; ventral fur has three color
bands; facial stripes well marked but
dusky; dorsal stripe narrow, but conspic-
uous; seven vibrissae surround margin of
noseleaf; two vibrissae on each side of the
upper lip below vibrissae surrounding
noseleaf; fold lines in pinnae well marked;
long (4+mm), dense hair on the upper
surface of feet; long, dense fringe of hair
on trailing edge of uropatagium paler
than fur of dorsum; postorbital processes
moderately developed; M1 protocone
large, well developed.
Description.—A medium-sized Platyr-
rhinus (FA 42.0–47.8 mm, GLS 25.1–
26.7 mm, CCL 23.0–24.4 mm; Tables 3,
4) that has dense, brown dorsal and
ventral fur with individual hairs having
three color bands; dorsomedial and ven-
trolateral facial stripes the same color
(dusky); fur of dorsum 8–10 mm long;
dorsal stripe thin and brighter than the
facial stripes; well-marked fold lines in
pinnae; genal vibrissae lack a basal
protuberance; a single array of seven
vibrissae surround margins of noseleaf;
one vibrissa on each side of upper lip
ventral to vibrissae surrounding noseleaf;
four submental vibrissae on each side of
chin; one interramal vibrissa present;
noseleaf longer than wide; inferior border
of nasal horseshoe completely free of
upper lip; dense hair covers upper surfac-
es of the feet; inverted ‘‘U’’-shaped
posterior margin of uropatagium fringed
with long, pale hair; uropatagium (dry)
extends 2–3 mm beyond body at midline;
proximal half of forearm covered with
long hair; insertion of posterior margin of
plagiopatagium on middle of metatarsal
I; metacarpal III shorter than metacarpal
V; postorbital and paraoccipital processes
moderately developed; fossa on squamo-
sal root of zygomatic arch variable in
development, shallow to deep; upper
inner incisors convergent and in contact;
upper outer incisors monolobate; two
stylar cuspules on posterior cristid of P4;
deep fossa on the hypoconal basin of P4;
M1 parastyle absent; a stylar cuspule
present on both labial and lingual cingula
of M1 metacone; metastyle present on
M1; protocone well developed on M1;
labial cingulum present on M2 paracone;
stylar cuspule present on the lingual face
of M2 paracone; metastyle present on
M2; lacks a stylar cuspule on lingual face
of M2 metacone; lingual cingulum of the
M2 metacone not continuous with para-
cone; M2 hypoconal basin developed;
labial and lingual cingulids present on
p4; anterior and posterior cristids of p4
lack stylid cuspulids; lacks both paraco-
nid and metaconid on m1; hypoconid
present on m2; stylid cuspulid between
the metaconid and protoconid absent on
m2; labial and lingual cingulids present
on m2.
Comparisons.—Platyrrhinus umbratus
is sympatric with P. albericoi,P. chocoen-
sis,P. dorsalis,P. infuscus,P. ismaeli,P.
nigellus,andP. vittatus. Easily distin-
guished from P. albericoi,P. infuscus,P.
ismaeli,andP. vittatus by its shorter
forearm and greatest length of skull
(Tables 3, 4), P. umbratus has been
confused with P. dorsalis and its mea-
surements overlap those of P. aquilus,P.
chocoensis and P. nigellus (Tables 3, 4).
Therefore, the following comparisons
focus on differentiating these five species.
Externally, Platyrrhinus umbratus and
P. chocoensis can be distinguished from P.
dorsalis and P. nigellus by paler dorsal
color (P. dorsalis is dark brown and P.
aquilus and P. nigellus are blackish
dorsally); seven vibrissae surround the
noseleaf in a single array in P. chocoensis
and P. umbratus (P. aquilus has eight, P.
dorsalis and P. nigellus have six); P.
umbratus and P. aquilus have two vibris-
sae on each side of upper lip below
vibrissae surrounding noseleaf (P. cho-
coensis and P. dorsalis have one on each
VOLUME 122, NUMBER 3 265

side; P. nigellus shows variation in this
character, having either one or two
vibrissae on each side); fold lines in the
pinnae are well marked in P. aquilus,P.
chocoensis,P. nigellus,andP. umbratus
(fold lines are poorly marked or lacking in
P. dorsalis); dorsal hair is long (8+mm) in
P. aquilus,P. dorsalis,P. nigellus,andP.
umbratus (shorter than 8 mm in P.
chocoensis); ventral fur has two color
bands in P. chocoensis and P. umbratus
(three bands in P. aquilus,P dorsalis and
P. nigellus); P. aquilus,P. nigellus,andP.
umbratus have long, dense hair on upper
surface of the feet (fur is short in P.
chocoensis and less dense in P. dorsalis);
the uropatagial fringe of hair is abundant
and long in P. aquilus,P. nigellus,andP.
umbratus (shorter and sparse in P. cho-
coensis and P. dorsalis).
Cranially, Platyrrhinus aquilus,P. cho-
coensis,P. dorsalis,P. nigellus,andP.
umbratus differ as follows: The skull is
slender in P. umbratus, but longer in P.
aquilus and more robust in P. chocoensis,
P. dorsalis, and P. nigellus; the postorbital
processes are moderately developed in P.
aquilus and P. umbratus, but poorly
developed in P. chocoensis,P. dorsalis,
and P. nigellus.
Dentally, Platyrrhinus aquilus,P. cho-
coensis,P. dorsalis,P. nigellus,andP.
umbratus differ as follows: P. aquilus and
P. umbratus have a deep fossa on the
hypoconal basin of P4 (the fossa is
shallow in P. chocoensis;P. dorsalis and
P. nigellus show variation in this charac-
ter); P. umbratus has a well-developed
protocone on M1 (moderately developed
in P. aquilus and P. nigellus; small and
blunt in P. dorsalis and P. chocoensis); P.
aquilus and P. umbratus have a labial
cingulum on M2 paracone (lacking in P.
nigellus; variably developed in P. cho-
coensis and P. dorsalis); P. aquilus,P.
umbratus,P. chocoensis,andP. nigellus
lack a stylar cuspule on lingual face of M2
paracone (present in P. dorsalis); the
lingual cingulum of M2 metacone does
not extend beyond the metacone in P.
aquilus and P. umbratus (lingual cingulum
of M2 metacone continuous to paracone
in P. chocoensis and P. dorsalis); both
labial and lingual cingulids of p4 present
in P. aquilus,P. umbratus,P. dorsalis,and
P. nigellus (only labial cingulid present in
P. chocoensis); hypoconid present on m2
in P. aquilus and P. umbratus (absent in
P. chocoensis,P. dorsalis,andP. nigellus);
a stylid cuspulid between metaconid and
protoconid of m2 is lacking in P. umbra-
tus and P. chocoensis (present in P.
aquilus,P. dorsalis,andP. nigellus).
Multivariate analysis.—In addition to
the analysis graphed in Figure 5, we
compared the holotype and 35 specimens
of Platyrrhinus umbratus from Colombia
and Venezuela with 21 P. chocoensis
(Colombia and Panama), 13 P. dorsalis
(Ecuador and Colombia), and 57 P.
nigellus (Colombia, Ecuador, and Peru).
The first three PCs accounted for 86.4%
of the variation. A plot of factor scores on
the first two axes (Fig. 7) shows that P.
umbratus overlaps completely with P.
nigellus on PC1, which represents overall
size, but is completely separated from P.
chocoensis and P. dorsalis, a reflection of
the larger size of P. chocoensis and P.
dorsalis. Along PC2, P. umbratus overlaps
completely with P. nigellus and P. dorsalis
and partially with P. chocoensis, reflecting
shape similarities among these species
(Fig. 7, Table 6).
Remarks.—Probably because Sanborn
(1955), in his revision of the genus (as
Vampyrops), regarded Platyrrhinus um-
bratus as a junior synonym of P. dorsalis,
many authors did not question his
decision (Gardner & Carter 1972b, Alber-
ico 1990, Velazco 2005, Gardner 2008).
Other authors recognized P. umbratus as
a valid species, but treated aquilus and
oratus as junior synonyms (e.g., Koop-
man 1994, Simmons 2005). After exam-
ining the type material of P. umbratus and
comparing it with holotypes and speci-
mens of other species of Platyrrhinus,we
266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

found P. umbratus to be distinctive
morphologically and morphometrically,
and easily distinguishable from conge-
ners. Therefore, we recognize P. umbratus
as a valid species and treat oratus as a
junior synonym.
The location of the type locality of
Platyrrhinus umbratus has been matter of
confusion due to vague information on
the provenance of the specimens (Bangs
1990, Lyon 1902). Simmons (2005) and
Gardner (2008) placed the type locality in
the Colombian department of Magda-
lena; the MCZ placed it in the department
of Santander. Here we follow Helgen &
MacFadden (2001) in identifying San
Miguel, department of La Guajira, as
the correct locality for the holotype and
paratypes of P. umbratus (see Paynter
1997).
The holotype of Vampyrops oratus,
BMNH 14.7.27.1, is an adult male
missing the right M3 and left upper
incisor. The mandible does not match
the cranium and obviously belongs to a
smaller Platyrrhinus. Based on the size of
the skin, we believe that the cranium and
the skin are from the same specimen.
Therefore, we restrict the holotype of V.
oratus to the skin and cranium.
Specimens examined (177).—Specimens
marked with an asterisk were used in the
elaboration of Tables 3–6 and in the
morphometric analyses. COLOMBIA:
Boyaca´: Pajarito, Corinto, Finca El Des-
canso, Quebrada Conguta (ICN 8351);
Pajarito, Hacienda Camijoque (ICN
8039, 17111*). El Cesar: Valledupar,
Villanueva, Sierra Negra (USNM 281924,
281925, 281303, 281305). Choco´: Bahia
Fig. 7. Plot of scores on first and second axes from PCA of 22 variables from 35 Platyrrhinus umbratus,
21 P. chocoensis,13P. dorsalis,and57P. nigellus (see Table 6).
VOLUME 122, NUMBER 3 267

Solano (Mutis) (UV 4149, 4150, 4152).
Cundinamarca: Tena, Laguna de Pedro
Palo (ICN 5292, 5294*, 5537*, 5538,
5540). Huila/Cauca: Moscopas, Rı
´oLa
Plata (USNM 483574*, 483576*). La
Guajira: San Antonio (MCZ B8300);
San Miguel (MCZ B8180* [Holotype of
Vampyrops umbratus]); Palomino (MCZ
B8301). Magdalena: Santa Marta, Serra-
nı
´a San Lorenzo, estacio´ n Inderena (ICN
5388–5391*). Meta: Restrepo (UV 3850).
Risaralda: Santa Rosa de Cabal, Laguna
de Otun (UV 2517, 2518, 2520). Santan-
der:Charala,Virolı
´n, Margen izquierda
Rı
´o Oibita, carretera Can˜averal-Olival
(ICN 6695–6697*). Valle del Cauca: Cali,
Corea, Los Farallones (UV 7574); Cali,
Pance (USNM 483586–483592*, 483594–
483596*, 483597, 483599–483601*; UV
769, 1234); Dagua, El Queramal, Tokio,
Antena Tele (UV 2027); Florida, Hacien-
da Los Alpes (UV 7575, 10836). VENE-
ZUELA: Aragua: Rancho Grande Bio-
logical Station, 13 km NW Maracay
(USNM 370514–370516, 517465); El Por-
tachuelo, 14 km NW Maracay (USNM
517466). Carabobo: Montalban (USNM
440651, 440653); Montalban, 4 km NW
Montalban, La Copa (USNM 440652,
440654–440656, 443363). Distrito Federal:
Alto N
˜oLeo´ n, 33 km WSW Caracas
(USNM 408559–408564); Boca Tigre
Valley, 5 km NW Caracas, near Claveli-
tos (USNM 370470, 370472, 370473,
370477, 370478); 0.5 km W junction
Puerto Cruz and Colonia Tovar highways
(USNM 562985–562987); Los Venados
(USNM 370407–370416*, 370418, 370429,
370431–370447); Pico A
´vila, Estacio´n de
Bombeo (UV 11467, 11468); Pico A
´vila,
Near Hotel Humboldt, 5 km NNE Car-
acas (USNM 370452–370456, 370462,
370480–370490, 370509–370511, 372128);
Pico A
´vila, near Boca Tigre, 6 km NNW
Caracas (USNM 370491–370494, 370500–
370508); Sierra del A
´vila, Galifari (BMNH
14.7.27.1 [Holotype of Vampyrops ora-
tus]). Me´rida: La Carbonera, 12 km SE La
Azulita (USNM 387116–387118); Tabay,
4 km E Tabay, near Mucuy (USNM
373837–373839); Tabay, 6 km ESE Ta-
bay, Middle Refugio (USNM 387110–
387115). Miranda: Curupao, 5 km NNW
Guarenas (USNM 387126–387141). Mon-
agas: San Agustı
´n, 3 km NW Caripe
(USNM 408566); San Agustı
´n, 5 km
NW Caripe (USNM 408567, 408568).
Trujillo: Hacienda Misisi, 13 km E Tru-
jillo (USNM 373834–373836). Yaracuy:
Minas de Aroa, 20 km NW San Felipe
(USNM 440647).
Platyrrhinus nitelinea, new species
Western Broad-nosed Bat
Fig. 8
Vampyrops sp. A: Alberico, 1990:348.
Holotype.—Dried skin and skull of an
adult female, National Museum of Nat-
ural History (USNM 513461), obtained 9
Aug 1976 by Alfred L. Gardner (original
field number ALG 13227). The skin and
skull are in good condition.
Type locality.—1 km SW of Puente de
Moromoro, 3480 ft (1060 m); El Oro
Province; Ecuador (03u449S, 79u449W).
Paratypes.—The skin and skull of two
adult males and two adult females
(USNM 513460, 513462, 513463, 513464),
all from the type locality, and one skin and
skull of an adult female (USNM 522431)
that was lactating when caught on 5
Sep 1974 at Huerta Negra (02u599S,
79u389W), 10 km ESE Balao, E of Tenguel,
Guayas Province, Ecuador. Measurements
ofthetypematerialarepresentedin
Table 7.
Etymology.—From Latin niteo mean-
ing to shine, look bright, glisten in
combination with linea meaning a thread,
line, or mark, and referring to the
distinctive bright dorsal stripe in this
species; gender feminine and treated as a
noun in apposition to Platyrrhinus.
Distribution.—Platyrrhinus nitelinea is
known from western Colombia in the
departments of Choco´ , Narin˜ o, and Valle
del Cauca and from western Ecuador in
268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Dorsal and ventral views of the cranium and lateral view of the cranium and mandible of
Platyrrhinus nitelinea (USNM 513461, the holotype). See Table 7 for measurements.
VOLUME 122, NUMBER 3 269

the provinces of El Oro and Guayas
(Fig. 4).
Diagnosis.—Platyrrhinus nitelinea is a
moderately large Platyrrhinus (FA 52.9–
58.4 mm, GLS 29.3–31.9 mm, CCL 26.9–
29.6 mm; Tables 3, 4), easily distin-
guished from P. aquilus,P. aurarius,P.
chocoensis,P. dorsalis,P. lineatus,P.
masu,P. nigellus,P. recifinus, and P.
umbratus by its larger size and longer
skull (Tables 3, 4). This species is too
large to be confused with the diminuitive
P. brachycephalus,P. helleri,andP.
matapalensis. Measurements of P. niteli-
nea overlap with those of P. albericoi,P.
infuscus,P. ismaeli,andP. vittatus.
Dorsal and ventral fur is blackish; facial
stripes well marked; basal protuberance
present at base of genal vibrissae; two
vibrissae present on each side of upper lip
below the series of vibrissae surrounding
margin of noseleaf; inferior border of the
horshoe partially joined to upper lip;
pinnae have well-marked fold lines; dor-
sal stripe brilliant white; dorsal fur has
two color bands (bicolored); hair on
upper surface of feet short and moder-
ately dense; fringe of hair on trailing edge
of uropatagium is short (#3 mm) and
conspicuously pale; postorbital processes
well developed; paraoccipital processes
well developed; metaconid lacking on
m1; hypoconid present on m2.
Description.—Platyrrhinus nitelinea is
one of the four largest species in the
genus (Tables 3, 4). Dorsal pelage is 8–
11 mm long, bicolored with darker tips;
ventral pelage pale brown to dark brown,
monocolored on throat and upper chest,
bicolored (hairs have gray tips) on re-
mainder of venter; dorsal stripe wide and
brighter than the facial stripes; folds in
pinnae well marked; hair on the upper
surface of feet and along trailing edge of
uropatagium short and moderately dense;
porterior margin of uropatagium has
shape of an inverted ‘‘U’’; width of
uropatagium 2–3 mm (dry) at midline;
proximal third of forearm covered with
dense, long hair; metacarpal III is shorter
than metacarpal V; insertion of plagiopa-
tagium on metatarsal I. Genal vibrissae
arise from a basal protuberance; six
vibrissae surround margin of noseleaf in
a single array; two vibrissae present on
each side of upper lip below vibrissae
surrounding noseleaf; four submental
vibrissae present on each side of chin;
two interramal vibrissae present; noseleaf
longer than wide; inferior border of nasal
horseshoe partially fused to t upper lip.
The cranium has well-developed postor-
Table 7.—Measurements (mm) of the type material of Platyrrhinus nitelinea.
Character
Holotype USNM
513461 RParatype USNM
513460 RParatype USNM
513462 =Paratype USNM
513463 RParatype USNM
513464 =Paratype USNM
522431 R
GLS 31.6 31.3 31.5 31.9 31.6 31.2
CIL 29.8 29.7 29.6 30.5 29.9 29.8
CCL 29.2 28.9 28. 9 29.6 29.2 28.9
BB 12.7 12.8 12.7 13.0 13.1 12.7
ZB 19.2 18.7 19.4 19.6 18.8 18.5
PB 7.1 7.0 7.3 7.2 7.4 7.0
MB 13.9 13.8 14.0 14.5 14.1 14.1
MTRL 13.3 12.7 13.4 13.2 13.3 12.9
M2–M2 14.4 14.1 14.8 14.3 14.3 14.5
DENL 23.5 23.2 23.5 23.7 23.5 23.5
MANDL 14.6 13.9 14.5 14.4 14.2 14.1
FA 57.6 58.2 56.9 56.8 57.1 58.4
TL 85.0 90.0 90.0 87.0 89.0 84.0
HF 14.0 16.0 16.0 14.0 17.0 17.0
E 22.0 22.0 22.0 23.0 23.0 23.0
270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bital processes, an inverted ‘‘V’’-shaped
posterior border of the hard palate; well-
developed paraoccipital processes; fossa
on the squamosal root of the zygomatic
arch lacking or only barely perceptible.
Upper inner incisors convergent and in
contact; upper outer incisors monolobate;
posterior cristid of P4 has two stylar
cuspules; hypoconal basin of P4 a deep
fossa; M1 lacks both a parastyle and
mesostyle; stylar cuspule present on labial
cingulum of M1 metacone; stylar cuspule
present on lingual cingulum of M1 meta-
cone; sulcus on posterior cristid of para-
cone continous with cingulum on lingual
face of metacone on M1; metastyle
present on M1; M1 protocone moderately
developed; parastyle present on M2; labial
cingulum present on M2 paracone; stylar
cuspule present on lingual face of M2
paracone; metastyle present on M2; stylar
cuspule present on lingual face of M2
metacone; lingual cingulum of the M2
metacone not extending to the paracone;
hypoconal basin well developed on M2;
labial and lingual cingulids present on p4;
stylid cuspulids lacking on anterior and
posterior cristids of p4; paraconid lacking
on m1; labial and lingual cingulids present
on m1; stylid cuspulid present on anterior
cristid of m1 protoconid; metaconid
absent on m1, hypoconid present on m2;
stylid cuspulid between the metaconid and
protoconid present on m2; labial and
lingual cingulids present on m2.
Comparisons.—Platyrrhinus nitelinea is
most easily distinguished from P. aquilus,
P. aurarius,P. brachycephalus,P. cho-
coensis,P. dorsalis,P. helleri,P. lineatus,
P. masu,P. matapalensis,P. nigellus,P.
recifinus,andP. umbratus by its larger
size (GLS .29.3). Although its its longer,
blackish dorsal pelage distinguishes P.
nitelinea from other large Platyrrhinus (P.
albericoi,P. infuscus,P. ismaeli,andP.
vittatus), their external and cranial mea-
surements overlap (Tables 3, 4). There-
fore, the following comparisons focus on
differentiating these five species.
Externally, Platyrrhinus nitelinea can be
distinguished from P. albericoi,P. infus-
cus,P. ismaeli,andP. vittatus by its
darker, almost black color (P. albericoi,
P. infuscus,P. ismaeli,andP. vittatus
range in color from pale brown to dark
brown); dorsal hair in P. nitelinea,P.
albericoi,P. ismaeli,andP. vittatus is
longer than 8 mm (shorter than 6.3 mm
in P. infuscus); dorsal pelage bicolored in
P. nitelinea (three color bands in P.
albericoi,P. infuscus,P. ismaeli,andP.
vittatus); ventral pelage bicolored in P.
nitelinea,P. albericoi,andP. vittatus
(three color bands in P. infuscus and P.
ismaeli); dorsal stripe wide and brilliant
white in P. nitelinea and P. vittatus
(conspicuous, but narrow in P. albericoi
and P. ismaeli; thin and inconspicuous in
P. infuscus). Fold lines in the pinnae are
well marked in P. nitelinea and P. vittatus
(poorly marked but distinguishable in P.
albericoi and P. ismaeli); hair on upper
surface of feet is short and intermediate in
density in P. nitelinea (dense and long in
P. albericoi,P. ismaeli,andP. vittatus;
short and sparse in P. infuscus); uropata-
gial fringe is sparse in P. nitelinea,P.
infuscus,andP. vittatus (dense in P.
albericoi and P. ismaeli); metacarpal III
shorter than metacarpal V in P. nitelinea,
P. ismaeli,andP. vittatus (metacarpals III
and V subequal in P. albericoi; metacar-
pals subequal or metacarpal III longer in
P. infuscus). Basal protuberance present
at base of genal vibrissae in P. nitelinea
(protuberance lacking in P. albericoi,P.
infuscus,P. ismaeli, and P. vittatus); two
vibrissae present on each side of upper lip
below vibrissae surrounding margin of
noseleaf in P. nitelinea (one vibrissa on
each side in P. albericoi,P. infuscus,P.
ismaeli,andP. vittatus); two interramal
vibrissae present in P. nitelinea,P. in-
fuscus,andP. vittatus (one interramal
vibrissa in P. ismaeli; one or two in P.
albericoi); inferior border of the horseshoe
partially joined to the upper lip in P.
nitelinea (inferior border of horseshoe
VOLUME 122, NUMBER 3 271

completely free in P. albericoi,P. ismaeli,
and P. vittatus; variably free or partially
joined in P. infuscus).
Cranially, Platyrrhinus nitelinea,P. al-
bericoi,P. infuscus,P. ismaeli,andP.
vittatus differ as follows: P. nitelinea has
well-developed postorbital processes (mod-
erately developed in P. albericoi,P. infus-
cus,andP. vittatus); P. nitelinea,P.
albericoi,andP. infuscus have well-devel-
oped paraoccipital processes (moderately
developed in P. ismaeli and P. vittatus);
fossa on the squamosal end of the zygo-
matic arch absent or barely perceptible in
P. nitelinea,P. albericoi,andP. vittatus
(shallow in P. infuscus and P. ismaeli).
Dentally, Platyrrhinus nitelinea,P. al-
bericoi,P. infuscus,P. ismaeli,andP.
vittatus differ as follows: P. nitelinea,P.
albericoi,P. ismaeli,andP. vittatus lack a
parastyle on M1 (present in P. infuscus);
P. nitelinea and P. vittatus lack a mesos-
tyle on M1 (present in P. albericoi;
present or absent in P. infuscus and P.
ismaeli); the sulcus on the posterior cristid
of paracone is continuous to the cingulum
of lingual face of metacone on M1 in P.
nitelinea,P. albericoi,P. infuscus, and P.
vittatus (not continuous to cingulum of
metacone in P. ismaeli); P. nitelinea,P.
albericoi,andP. infuscus have a moder-
ately developed protocone on M1 (proto-
cone small and blunt in P. ismaeli;
variably developed in P. vittatus); P.
nitelinea,P. infuscus,P. ismaeli, and P.
vittatus have a metastyle on M2 (absent in
P. albericoi); P. nitelinea,P. albericoi,P.
infuscus,andP. ismaeli lack a stylid
cuspulid on the anterior cristid of p4
(present in P. vittatus); P. nitelinea lacks a
metaconid on m1 (well developed in P.
infuscus; poorly developed in P. albericoi,
P. ismaeli, and P. vittatus); P. nitelinea
has a hypoconid on m2 (absent in P.
albericoi,P. infuscus,P. ismaeli,andP.
vittatus); P. nitelinea,P. albericoi,P.
ismaeli,andP. vittatus have a stylid
cuspulid between the metaconid and
protoconid on m2 (lacking in P. infuscus).
Multivariate analysis.—We compared
the 48 specimens of Platyrrhinus niteli-
nea, incuding the type material from
Ecuador, with 15 P. albericoi (Colombia,
Peru, and Venezuela), 14 P. infuscus
(Peru), 22 P. ismaeli (Colombia, Ecua-
dor, and Peru), and 33 P. vittatus
(Colombia and Panama). Two PCs were
extracted that account for 85.6%of the
variation. A plot of factor scores on the
first two axes (Fig. 9) shows that P.
nitelinea overlaps completely with P.
infuscus and P. vittatus on PC1, which
represent overall size, and is completely
separated from P. albericoi and P.
ismaeli, with P. albericoi being larger
and P. ismaeli smaller than P. nitelinea.
On PC2, P. nitelinea overlaps with P.
albericoi,P. infuscus,P. ismaeli,andP.
vittatus, reflecting shape similarities
among these species (Fig. 9, Table 8).
Specimens examined (52).—Individuals
or series marked with an asterisk were
used in the elaboration of Tables 3, 4, 7,
and 8 and in the morphometric analyses.
COLOMBIA: Choco´: Quebrada La Gua-
gua, 8 km W La Italia, subiendo por la
Quebrada Toro Viejo (UV 7456–7460*);
Municipio San Jose´ del Palmar, 14 km N
La Italia (UV 10042, 10043*); Municipio
San Jose´ del Palmar, Vereda La Italia,
Alto del Oso (UV 4562–4569*, 10121–
10124*). Narin˜o: Municipio Junı
´n, La
Planada de Maindez, ,97 km W, 13 km
N of Pasto (UV 3045–3049*, 3056–3058).
Valle del Cauca: Municipio Buenaven-
tura, Llano Bajo (UV 10541*); Municipio
Darie´n, Campamento Rı
´o Azul, Calima
Medio (UV 11355*); Municipio Darie´n,
Rı
´o Azul (UV 3395–3397*, 3398, 3399–
3406*); Anchicaga´ (bajo), carretera vieja
Cali-Buenaventura, ,Km 90 (UV 2635*);
La Loma del A
´guila, subiendo Rı
´o
Cajambre, frente a Cerro Cajas (UV
3704, 3705*); Planta Ele´ ctrica, Alto An-
chicaya (UV 3181, 3182*). ECUADOR:
El Oro: Puente de Moromoro, 1 km SW
(USNM 513460*, 513461* [Holotype of
Platyrrhinus nitelinea], 513462–513464*).
272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Guayas: Huerta Negra, 10 km ESE Ba-
lao, east of Tenguel (USNM 522431*).
Key to species of the genus Platyrrhinus
1. Forearm longer than 53 mm; condy-
lobasal length more than 26 mm . . . 2
19. Forearm equal to or shorter than
56 mm, usually shorter than 54 mm;
condylobasal length usually less than
26mm ........................ 5
2. Pelage dark, usually blackish brown;
four facial stripes and a conspicuous
dorsal stripe; p4 with long posterior
‘‘heel’’ (posterolabial cuspulid) . . . . 3
29. Dorsal pelage buffy to dark brown,
not blackish; facial and dorsal stripes
inconspicuous; p4 with a short pos-
terior ‘‘heel’’ (posterolabial cuspu-
lid) ........... Platyrrhinus infuscus
3. Forearm 55 mm or longer; condylo-
basal length 28 mm or longer . . . . . 4
39. Forearm 55 mm or shorter; condylo-
basal length less than 28 mm . . . . . 5
4. Forearm shorter than 62 mm; acces-
sory cuspulid present or absent on
Fig. 9. Plot of scores on first and second axes from PCA of 22 variables from 48 Platyrrhinus nitelinea,
15 P. albericoi,14P. infuscus,22P. ismaeli,and33P. vittatus (see Table 8).
VOLUME 122, NUMBER 3 273

anterolingual cristid of p4; M1
mesostyle absent; M2 metastyle pre-
sent ........................... 6
49. Forearm longer than 61 mm; ante-
rolingual cristid of p4 smooth, ac-
cessory cuspulids lacking on p4; M1
mesostyle present; M2 metastyle
absent ......... Platyrrhinus albericoi
5. Pelage pale brown to dark brown;
facial stripes always white and usu-
ally conspicuous, forearm 32–
48 mm ........................ 7
59. Pelage brown to blackish brown;
facial stripes buff, sometimes incon-
spicuous,forearm 40–56mm .... 11
6. Anterolingual cristid of p4 smooth,
accessory cuspulids lacking on p4
................ Platyrrhinus nitelinea
69. Accessory cuspulid present on ante-
rolingual cristid of p4 ............
................ Platyrrhinus vittatus
7. Size smaller, forearm 42 mm or
shorter . ....................... 8
79. Size larger, forearm 42 mm or longer
................................ 10
8. Anterolingual cristid of last lower
premolar (p4) bears one or two
tubercle-like cuspulids ............ 9
89. Anterolingual cristid of last lower
premolar (p4) essentially smooth,
lacking accessory cuspulids ........
............ Platyrrhinus matapalensis
9. Anterolingual cristid of p4 bears two
well-developed accessory cuspulids;
outline of lateral margin of ptery-
goid process narrowly concave (small
c-shaped) in posterior view because
of well-developed thin ridge of bone
that extends posterolaterally from
near mid-height of pterygoid process
to base of braincase .............
.......... Platyrrhinus brachycephalus
99. Anterolingual cristid of p4 has one
or two small accessory cuspulids;
posterior cristid of p4 bears one or
more erect cuspulids anterior to and
in addition to a short ‘‘heel’’; outline
of lateral margin of pterygoid pro-
cess broadly concave (large C-
shaped) in posterior view, lateral
bony ridge not well developed or
elevated ......... Platyrrhinus helleri
10. Size larger, forearm more than
45 mm; crown of p2 relatively nar-
row in labial outline and triangular
in cross-sectional outline (posterolin-
gual cristid well developed), labial
surface conspicuously convex, and
‘‘heel’’ 20%or more of anterior-
posterior length of tooth in lateral
view .......... Platyrrhinus lineatus
109.Size smaller, forearm 42–46 mm,
usually less than 45 mm; crown of
p2 ‘‘sail-shaped’’ in lateral outline
and blade-like in cross-sectional out-
line (posteroligual cristid weakly
developed), labial surface nearly flat,
and ‘‘heel’’ less than 20%of anteri-
or-posterior length of tooth in lateral
view .......... Platyrrhinus recifinus
11. Size larger, forearm 49–55 mm; up-
per inner incisors slender and long,
with tips parallel for more than half
their length (crown height)
................ Platyrrhinus aurarius
Table 8.—Factor loadings for first three axes
from PCA of 22 variables from Platyrrhinus
albericoi,P. infuscus,P. ismaeli,P. nitelinea,and
P. vittatus.
Variable
Correlations
PC 1 PC 2
GLS 0.969 20.065
CIL 0.971 20.105
CCL 0.972 20.103
BB 0.880 20.149
ZB 0.963 20.066
PB 0.755 20.510
C–C 0.890 0.047
MB 0.927 20.177
PL 0.901 0.222
MTRL 0.959 20.002
MLTRL 0.942 0.065
M1–M1 0.947 0.085
M2–M2 0.948 0.117
MXBR 0.821 20.329
M1W 0.597 0.657
M2W 0.750 0.560
DENL 0.944 20.116
MANDL 0.960 0.012
COH 0.855 0.228
WMC 0.817 20.291
m1W 0.769 0.346
FA 0.827 20.185
Proportion of
variation 78.4%7.2%
274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

119.Size small to large, forearm 40–
56 mm; upper inner incisors broad
and short with convergent tips . . . . 12
12. Forearm 40–56 mm; stylid cuspulid
present between protoconid and
paraconid of m2; ventral pelage has
three color bands; dorsal coloration
dark .......................... 13
129.Forearm 46–51 mm; stylid cuspulid
absent between protoconid and
metaconid of m2; ventral pelage with
two color bands, dorsal coloration
comparatively paler . . . . . . . . . . . . . . 17
13. Size larger, forearm 49–56 mm; lin-
gual cingulum not continuous from
paracone to metacone (sulcus not
continuous between paracone and
metacone) of M1; pinnae lack lateral
folds ........... Platyrrhinus ismaeli
139.Size smaller, forearm 40–51 mm;
lingual cingulum continuous from
paracone to metacone (sulcus con-
tinuous between paracone and meta-
cone) of M1; pinnae have lateral
folds .......................... 14
14. Size smaller, forearm 40–47 mm;
stylar cuspule absent on lingual
margin of paracone on M2 . . . . . . . 15
149.Size larger, forearm 46–51 mm; sty-
lar cuspule present on lingual margin
of paracone on M2 . . . . . . . . . . . . . 16
15. GLS more than 26.8 mm; eight
vibrissae surrounding the margins
of the noseleaf in a single array;
interramal vibrissae absent; paraocci-
pital processes well developed; hypo-
conid present on m2 . . . . . . . . . . . . .
................. Platyrrhinus aquilus
159.GLS less than 26.8 mm; six vibrissae
surrounding the margins of the
noseleaf in a single array; one
interramal vibrissae present; paraoc-
cipital processes moderately devel-
oped; hypoconid present on m2
................ Platyrrhinus nigellus
16. Fold lines in pinnae relatively week;
dorsal stripe relatively narrow, fur
on dorsum long (more than 8 mm);
fossa in squamosal branch of zygo-
matic arch deep; upper outer incisors
unilobate; lingual cingulum of M2
metacone continuous to paraco-
ne ............ Platyrrhinus dorsalis
169.Fold lines in pinnae well marked;
dorsal stripe conspicuous and wide;
fur on dorsum shorter (less than
8 mm); fossa in squamosal branch of
zygomatic arch shallow to almost
imperceptible; upper outer incisors
bilobate; lingual cingulum of M2
metacone not continuous to para-
cone ............ Platyrrhinus masu
17. Size larger, forearm 47–51 mm, GLS
26.5–29.0 mm; one vibrissa on each
upper lip below the series of vibris-
sae surrounding the noseleaf; post-
orbital processes almost impercepti-
ble; fossa on the hypoconal basin of
P4 shallow; M1 protocone small and
blunt; lingual cingulum of M2 meta-
cone continuous to paracone; hypo-
conid absent on m2 .............
.............. Platyrrhinus chocoensis
179.Size smaller, forearm 42–48 mm,
GLS 25.1–26.6 mm; two vibrissae
on each upper lip below the series
of vibrissae surrounding the noseleaf;
postorbital processes moderately de-
veloped; fossa on the hypoconal
basin of P4 deep; M1 protocone
well developed; lingual cingulum of
M2 metacone does not continue to
paracone; hypoconid present on
m2 ........... Platyrrhinus umbratus
Acknowledgments
This study was part of PMV’s disserta-
tion submitted to the University of Illinois
at Chicago as partial fulfillment of his
doctoral degree. PMV thanks committee
members, M. Ashley, H. F. Howe, R.
Mason-Gamer, B. D. Patterson, and R. H.
Ree. Special recognition is due to B. D.
Patterson for his constant support and
encouragement to PMV. The following
curators and collection staff graciously
provided access to specimens under their
care: Daphne M. Hills and Paula Jenkins
(BMNH), Suzanne B. McLaren (CM),
Renato Gregorin (CMUFLA), Bruce D.
Patterson and John Phelps (FMNH), J.
Enrique Castillo (IAvH-M), Yaneth Mu-
n˜ oz-Saba (ICN), Gustavo A. B. da Fon-
VOLUME 122, NUMBER 3 275

seca, Raquel Texeira de Moura, Valeria C.
Tavares, Yuri Leite, and Leonora Pires
Costa (DZ-UFMG); Santiago Madrin˜a´n
(ULA), Mark S. Hafner (LSUMZ), Judith
Chupasko (MCZ), Ce´cile Callou
(MNHN), Carlos Delgado, Javier Mun˜oz,
and Danny Zurc (MUA); Vı
´ctor Pacheco
(MUSM), James L. Patton (MVZ), Judith
L. Eger and Burton K. Lim (ROM), Philip
Myers (UMMZ), Linda K. Gordon,
James G. Mead (USNM), and Suzanne
C. Peurach (USGS-PWRC); Oscar Mur-
illo Garcia (UV), Hendrik Turni and
Robert Asher (ZMB). Many of the spec-
imens were collected with NSF support
(DEB 9870191 to Bruce D. Patterson and
colleagues; OISE 0630149 to BDP and
PMV). The research was also supported
by a Grant-in-Aid of Research from the
American Society of Mammalogists, and
the Ellen Thorne Smith Fund (FMNH),
Barbara E. Brown Fund for Mammal
Research (FMNH), the Lester Armour
Graduate Fellowship (FMNH), the
Smithsonian Institution Pre-doctoral Fel-
lowship, Ernest Mayr Travel Grant in
Animal Systematics from the Museum of
Comparative Zoology at Harvard Univer-
sity, The Albert R. and Alma Shadle
Fellowship in Mammalogy from the
American Society of Mammalogists, and
the Provost’s Award for Graduate Re-
search from the University of Illinois at
Chicago. Lastly, we appreciate the con-
structive comments that were provided by
M. Monica Diaz, Michael D. Carleton,
and one anonymous reviewer.
Literature Cited
Alberico, M. 1990. Systematics and distribution of
the genus Vampyrops (Chiroptera: Phyllosto-
midae) in northwestern South America.
Pp. 103–111 in G. Peters and R. Hutterer,
eds., Vertebrates in the tropics: Proceedings
of the International Symposium on Verte-
brate Biogeography and Systematics in the
Tropics, Bonn, June 5–8, 1989. Alexander
Koenig Zoological Research Institute and
Zoological Museum, Bonn, Germany.
Alberico, M. S., & E. Velasco. 1991. Description of
a new broad-nosed bat from Colombia.—
Bonner Zoologische Beitra¨ ge 42:237–239.
Bangs, O. 1900. List of the mammals collected in the
Santa Marta region of Colombia by W. W.
Brown, Jr.—Proceedings of the New England
Zoo¨logical Club 1:87–102.
Carter, D. C., & C. S. Rouk. 1973. Status of recently
described species of Vampyrops (Chiroptera:
Phyllostomatidae).—Journal of Mammalogy
54:975–977.
Gardner, A. L. 2008. Genus Platyrrhinus Saussure,
1860. Pp. 329–342 in A. L. Gardner, ed.,
Mammals of South America. Vol. 1: Marsu-
pials, xenarthrans, shrews, and bats. Univer-
sity of Chicago Press, Chicago, xviii +669 pp.
[Copyright 2007; published March 2008.]
———, & D. C. Carter. 1972a. A new Stenoderma-
tinae bat (Phyllostomidae) from Peru.—Oc-
casional Papers, The Museum Texas Tech
University 2:1–4.
———, & ———. 1972b. A review of the Peruvian
species of Vampyrops (Chiroptera: Phyllosto-
matidae).—Journal of Mammalogy 53:72–82.
Geoffroy St.-Hilaire, E
´. 1810. Sur les Phyllostomes
et les Me´gadermes, deux genres de la famille
des Chauve-souris.—Annales du Muse´um
d’Histoire Naturelle, Paris 15:157–198.
Gervais, P. 1855. Deuxie`me me´moire. Documents
zoologiques pour servir a la monographie des
Che´iropte`res Sud-Ame´ricains. Pp. 25–88 in P.
Gervais, ed., Mammife`res. In Animaux nou-
veaux ou rares recueillis pendant l’expe´dition
dans les parties centrales de l’Ame´rique du
Sud, de Rio de Janeiro a Lima, et de Lima au
Para; exe´cute´e par ordre du gouvernement
franc¸ais pendent les anne´es 1843 a 1847, sous
la direction du comte Francis de Castelnau. 1.
Anatomie. ed. F. de Castelnau. P. Bertrand,
Paris, 1855, 1: 1–116, 20 pls.
Gray, J. E. 1825. An attempt at a division of the
family Vespertilionidae into groups.—Zoo-
logical Journal 2:242–243.
Handley, C. O., Jr., & K. C. Ferris. 1972. Description
of new bats of the genus Vampyrops.—
Proceedings of the Biological Society of
Washington 84:519–524.
Helgen, K. M., & T. L. McFadden. 2001. Type
specimens of recent mammals in the Museum
of Comparative Zoology.—Bulletin of the
Museum of Comparative Zoology 157(2):
93–181.
Koopman, K. F. 1993. Order Chiroptera.
Pp. 137–242 in D. E. Wilson and D. M.
Reeder, eds., Mammal Species of the
World. A Taxonomic and Geographic Refer-
ence. 2nd edition. Smithsonian Institution
Press, Washington, D.C.
276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

———. 1994. Chiroptera: Systematics. In J.
Niethammer, H. Schliemann and D. Starck,
eds., Handbuch der Zoologie. Vol. 8, Mam-
malia, part 60. Walter de Gruyter, Berlin, 6 +
217 pp.
Lyon, M. W., Jr. 1902. Description of a new
bat from Colombia.—Proceedings of the
Biological Society of Washington 15:151–
152.
Paynter, R. A., Jr. 1997. Ornithological gazetteer
of Colombia, 2nd edition. Bird Department,
Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts, 537
pp.
Peters, W. 1859. Neue Beitra¨ge zur Kenntniss der
Chiropteren.—Monatsberichte der Ko¨ nigli-
chen Preussischen Akademie der Wis-
senschaften zu Berlin 1860:222–225.
Rouk, C. S., & D. C. Carter. 1972. A new species of
Vampyrops (Chiroptera: Phyllostomatidae)
from South America.—Occasional Papers,
Museum Texas Tech University 1:1–7.
Sanborn, C. C. 1955. Remarks on the bats of
the genus Vampyrops.—Fieldiana: Zoology
37(14):403–413.
Saussure, H. de. 1860. Note sur quelques mammi-
fe` res du Mexique.—Revue et Magasin de
Zoologie, serie 2, 12:429.
Simmons, N. B. 2005. Order Chiroptera.
Pp. 312–529 in D. E. Wilson and D. M.
Reeder, eds., Mammal species of the
world: a taxonomic and geographic refe-
rence. 3rd edition. The Johns Hopkins
University Press, Baltimore, Maryland,
1:xxxix +743 pp.
Solari, S., V. Pacheco, L. Luna, P. M. Velazco, & B.
D. Patterson. 2006. Mammals of the Manu
Biosphere Reserve. Pp. 13–22 in B. D.
Patterson, D. F. Stotz and S. Solari, eds.,
Mammals and Birds of the Manu Biosphere
Reserve, Peru.—Fieldiana: Zoology, new
series 110:1–49.
Swofford, D. L. 2002. PAUP*: phylogenetic analysis
using parsimony (* and other methods).
Version 4. Sinauer Associates, Sunderland,
Massachusetts,
Thomas, O. 1900. Descriptions of new Neotropical
Mammals.—Annals and Magazine of Natu-
ral History, series 7, 5:269–274.
———. 1901. On a collection of bats from Para.—
Annals and Magazine of Natural History,
series 7, 8:189–193.
———. 1914. Four new small mammals from
Venezuela.—Annals and Magazine of Natu-
ral History, series 8, 14:410–414.
———. 1924. New South American small mam-
mals.—Annals and Magazine of Natural
History, series 9, 13:234–237.
Velazco, P. M. 2005. Morphological phylogeny of
the bat genus Platyrrhinus Saussure, 1860
(Chiroptera: Phyllostomidae) with the de-
scription of four new species.—Fieldiana:
Zoology, new series 105:1–53.
———, & B. D. Patterson. 2008. Phylogenetics and
biogeography of the broad-nosed bats, genus
Platyrrhinus (Chiroptera: Phyllostomidae).—
Molecular Phylogenetics and Evolution
49:749–759.
———, & S. Solari. 2003. Taxonomy of Platyr-
rhinus dorsalis and Platyrrhinus lineatus (Chi-
roptera: Phyllostomidae) in Peru.—Masto-
zoologı
´a Neotropical 10(2):303–319.
Associate Editor: Michael D. Carleton.
Appendix I
The following list includes all other specimens
examined in this study, with their respective
localities. See Materials and Methods for abbrevi-
ations. Individuals or series marked with an asterisk
were used in the elaboration of Tables 3–8 and in
the morphometric analyses. Specimens examined of
the species P. aquilus,P. dorsalis,P. umbratus,and
P. nitelinea are listed in their respective species
accounts.
Platyrrhinus albericoi (99).—COLOMBIA: Anti-
oquia: Sonson, La Soledad (MUA 11141). Boyaca´:
Pajarito, Hacienda Camijoque (ICN 5449); Santa
Maria, Vereda La Calichana, sitio La Almenara
(ICN 16329); Togui, 2 km SW de Togui, Finca
Versalles (ICN 5344, 5345). Cundinamarca: El Ocaso
(IAvH-M 2578); Tena, Laguna de Pedro Palo (ICN
5296, 5541). Magdalena: Santa Marta, Cerro de San
Lorenzo (ICN 3468); Santa Marta, Serranı
´aSan
Lorenzo, Estacio´ n Inderena (ICN 5395–5397, 5399);
Santa Marta, Serranı
´a San Lorenzo, Hacienda La
Victoria (ICN 5392). Norte de Santander: Municipio
Toledo, Rı
´o Negro, Finca ‘‘San Isidro,’’ Parque
Natural Nacional Tama (IAvH-M 6699); Municipio
Toledo, Vereda el Diamante, Cerro San Agustı
´n,
Parque Natural Nacional Tama (IAvH-M 6740).
Quindı
´o: Municipio Armenia, Vereda San Juan de
Carolina, Finca La Irlanda, Bosque San Pedro
(IAvH-M 7046); Municipio Quimbaya, Vereda El
Laurel, Finca Balmaral (IAvH-M 7047). Risaralda:
Mistrato, Puerto de Oro (ICN 11788); Pueblo Rico,
Vereda San Jose´, Quebrada San Jose´ (ICN 11941,
11942); Santa Rosa, Finca La Selva, Vereda El
Cedralito (UV 13044). Santander: Charala, Inspec-
cio´n de policı
´a Virolin, margen derecha Rı
´o
Guillermo (ICN 8151); Encino, Vereda Los Pericos,
Finca Vegaleo´ n (ICN 17590, 17591). Valle del
Cauca: Cali, Finca Coro, Km 18, carretera Cali-
VOLUME 122, NUMBER 3 277

Buenaventura (UV 7174, 7179); Darie´n, Finca La
Guayacana, 2 km W del muro (UV 3413); Darie´n,
Rı
´o Bravo, campamento CVC (ICN 9395–9397); El
A
´guila, Corregimiento La Maria, Vereda El Cor-
azo´n (UV 12146); El Cairo, Estacio´n Cerro de Ingle´s
(UV 12631–12633, 13092); El Tambor, 15 km NW
Vijes (USNM 483579*, 483646–483650*, 483651–
483653*); Pance, 20 km SW Cali (USNM 483654,
483656*); Vijes, Vereda Villa Marı
´a, Finca La Selva
(UV 12113); Yumbo, Corregimiento Dapa, Vereda
La Paz, Finca La Rivera (UV 12725); Yumbo,
Dapa, Finca Humberto Arango (ICN 5015, 5016).
PERU: Ayacucho: La Mar, Santa Rosa, Yuraccyacu
(MUSM 993). Cusco: La Convencio´ n, Echarate,
Campamento Wayrapata (MUSM 14598, 14599);
La Convencio´ n, Echarate, a 2 km SW C.N.
Tangoshiari (MUSM 13395); La Convencio´n,
Quimbiri, Campamento Llactahuaman (FMNH
177477*, MUSM 14580–14597); Paucartambo, Cos-
n˜ ipata, Pillahuata (FMNH 172108, MUSM 19445);
Paucartambo, San Pedro (MUSM 19149 [Holotype
of Platyrrhinus albericoi Velazco, 2005]); Paucar-
tambo, Suecia, Km 138.5, Carretera Shintuya
(FMNH 170145*). Hua´nuco: Leoncio Prado,
Rupa-Rupa, ca. Tingo Maria (MUSM 3204–3209).
Junı
´n: Satipo, Rı
´o Tambo, Cordillera del Vilca-
bamba (MUSM 12996); Chanchamayo, Vitoc, ex-
campamento minero SIMSA (MUSM 20774).
Pasco: Oxapampa, Pozuzo, Palmira (MUSM
10973, 10974); Oxapampa, Pozuzo, Yanahuanca
(MUSM 10975). Ucayali: Coronel Portillo, Calleria,
WbankRı
´o Shesha, ca. 65 km ENE Pucallpa
(MUSM 3210). VENEZUELA: Aragua:ElPorta-
chuelo, 14 km NW Maracay, (USNM 517467).
Barinas: Altamira, 2 km SW Altamira (USNM
440658*). Distrito Federal: Los Venados, 4 km
NNW Caracas (USNM 370519, 370520*, 370521–
370523); Pico A
´vila, 6 km NNW Caracas, near Boca
Tigre (USNM 370524).
Platyrrhinus aurarius (78).—BRAZIL: Amazonas:
Barcelos, Alto da Serra do Araca´ , 1113 m. (RZ 100,
104–106, 109, 110, 113); Barcelos, Missa˜o Marari,
margem esquerdo do rio Katana, 340 m (RZ 020).
GUYANA: Cuyuni-Mazaruni: Namai Creek, 5 km
W of Parmina, 800 m (ROM 108220); N Slope Mt.
Roraima (USNM 588470). Potaro-Siparuni: Mount
Ayanganna, Toe Slope Camp, 700 m (ROM
114634, 114660, 114679); Mount Ayanganna, First
Plateau Camp, elevation 1100 m (ROM 114702).
SURINAME: Saramacca: Tafelberg, Arrowhead
Basin (CM 76804, 76805*); Tafelberg, Geyskes
Creek (CM 76806*). VENEZUELA: Amazonas:
Cerro de la Neblina, base of Pico Maguire (FMNH
137294–137298*, 137299–137307*, 137308–137311);
Cerro de la Neblina, left bank of Rı
´o Baria (FMNH
137312, 137313*); Cerro Neblina, Camp II (USNM
560803); Cerro Neblina, Camp V (USNM 560596–
560601, UV 11471, 11472); Cerro Neblina, Camp
VII (USNM 560682, 560683, 560804); Cerro Duida,
Cano Culebra, 50 km NNW Esmeralda (USNM
405044, 405046). Bolı
´var:Km125,85kmSSEEl
Dorado (USNM 387143–387145*, 387147, 387148*,
387150*, 387151, 387155, 387156*, 387157–387161,
387163* [Holotype of Vampyrops aurarius Handley
& Ferris, 1972], 387166, 387170–387172, 387174–
387178, 387180–387182, 387184).
Platyrrhinus chocoensis (158).—COLOMBIA:
Cauca: Alto Micay, Betania (FMNH 113745*,
113821*, 113822–113824*, 113825–113829*, 113830,
113831*, 113832–113835*). Choco´ : Alto Baudo (Pie
de Pepe´), Quebrada Platinero, Km 12, vı
´a a Itsmina
(UV 3817 [Holotype of Platyrrhinus chocoensis
Alberico and Velasco, 1991], 3818–3822, 10100–
10103); Alto Baudo (Pie de Pepe´), Rı
´o Pato, Km 51,
vı
´a Panamericana (UV 3823); Bahia Solano (Mutis),
Ensenada de Utria (UV 3645–3648); Bajo Baudo
(Pizarro), 2 km NE (UV 11289); Bajo Baudo
(Pizarro), Casa de la Maquina, confluencia quebrada
Angostura y Rı
´o Cubarrado (UV 11332); Bajo
Baudo (Pizarro), Quebrada Sen˜or Sen˜orcito (UV
11302, 11310); Municipio Riosucio, Reserva Forest-
al ‘‘Las Teresitas’’ Rı
´o Iruando (IAvH-M 3316);
Parque Natural Nacional Utrı
´a, Ensenada de Utrı
´a
(IAvH-M 6943); San Jose´ del Palmar, La Italia,
Quebrada La Guagua (UV 7446, 7447, 7449).
Narin˜o: La Guayacana (USNM 309018*, 309065,
309066). Valle del Cauca: Buenaventura, Bajo
Calima, Campamento Pulpare (UV 5748–5750);
Buenaventura, Bajo Calima, Cuartel B-V-83
(FMNH 140696, 140697*; MNHN C.G. 1989-1,
UV 5751, 5753–5755); Buenaventura, Bajo Calima,
Fuente de Material (UV 5566–5575); Buenaventura,
Bajo Calima, Granja Agroforesta (UV 2810–2812,
3183–3185, 10167); Buenaventura, Bajo Calima,
Quebrada San Joaquı
´n, 3 km E Bajo Calima (UV
2153); Buenaventura, El Mirador, Subiendo Agua-
sucia, (Rı
´o Cajambre), margen derecha, frente a
cerro caja (UV 3707–3709); Buenaventura, Llano-
bajo (UV 2162–2164, 2166, 2167, 10539, 10540);
Quebrada Caimancito, Rı
´o Calambre (UV 3706);
Buenaventura, Rı
´o Escalerete (UV 3186); Buena-
ventura, Rı
´o Raposo (USNM 339395, 339396*; UV
4875); Rı
´o Zabaletas, 29 km SE Buenaventura
(FMNH 85838, UMMZ 169039, 169040, 169048;
USNM 483533*, 483534–483539*, 483540–483572;
UV 281, 972, 2287–2291, 2294, 4257–4259). PANA-
MA: Darie´n: Parque Nacional Darie´n; Rancho Frı
´o
(FMNH 128141*); Tacarcuna Village Camp
(USNM 309601–309616).
Platyrrhinus ismaeli (126).—COLOMBIA: Anti-
oquia: Andes, Santa Rita, Vereda La Soledad, Finca
La Reina (ICN 16503); Carmen del Viboral, Los
Rı
´os (MUA 10291); Jardı
´n, Vereda La Linda,
quebrada La Linda (ICN 16474). Boyaca´ : Otanche,
Vereda La Y, Escuela La Y (ICN 16270); Pajarito,
Hacienda Camijoque (ICN 5442–5445, 8040, 8041);
Santa Marı
´a, Margen izquierda Rı
´o Bata, sendero
ecolo´gico (ICN 15067); Santa Marı
´a, Vereda Can˜o
278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Negro, margen derecha Rı
´o Bata (ICN 16326).
Caqueta: Florencia, Corregimiento EL Caran˜o,
Vereda Las Brisas, Finca Los Lirios (UAM 193,
194). Cundinamarca: Medina, Vereda Choapal,
alrededores Mesa del Cura (ICN 9585); Tena,
Laguna de Pedro Palo (ICN 5539). Huila: Parque
Natural Nacional ‘‘Cueva de los Huacharos’’
(FMNH 58732–58738*, IAvH-M 1930, 1932, 1934,
1990, 1992, 1994, 1996); Suaza, Vereda Alto Campo
Hermoso, Finca Marllins (UAM 159, 160, 227).
Huila/Cauca: Moscotas (USNM 483575). Meta:
Parque Natural Nacional ‘‘La Macarena’’ (IAvH-
M 1998). Norte de Santander: Municipio Toledo,
Rı
´o Negro, Parque Natural Nacional Tama, Finca
‘‘San Isidro’’ (IAvH-M 6681, 6682). Putumayo:
Municipio Mocoa, El Mirador (IAvH-M 6818,
6823). Quindı
´o: Filandia, Vereda El Roble, Reserva
Forestal Bremen La Popa (ICN 12476); Municipio
Salento, Vereda Boquia, Reserva La Pata Sola,
Finca La Betulia (UV 13229, 13230). Risaralda:
Quimbaya, La Florida, Vereda La Suiza, Santuario
de Flora y Fauna Otun (UV 12694); Santa Rosa,
Vereda El Cedralito, Finca La Selva (UV 13042,
13043). Valle del Cauca: Alto Gala´pagos, Limite
Valle-Choco (UV 12310); Buga, Corregimiento de la
Habana, Vereda El Janeiro, Finca Santelina (UV
12402, 12403, 12405, 12407, 13022, 13023); Buga,
San Jose´, Nogales, Cuenca Rı
´o Tulu´ a (UV 11225,
11226); Cali, Finca Coro (UV 7176); El A
´guila, La
Maria, El Corazo´ n (UV 12144); El Cairo, Estacio´n
Cerro de Ingle´s (UV 12615). ECUADOR: Napo:
Cascada San Rafael (FMNH 124988*). Pastaza:
Mera (USNM 548214, 548215, 548217–548224*).
PERU: Amazonas: Chachapoyas, 19 km by rd E
Balsas (FMNH 129132–129134*, 129135–129137*,
129138; MUSM 4944–4946 [Holotype of Platyr-
rhinus ismaeli Velazco, 2005]). Cajamarca: Celendin,
Hacienda Limo´ n (FMNH 129139*, 129140–
129143*, 129144–129146*; MUSM 4947, 4948);
San Miguel, 12 km SSW by road (FMNH 129147,
MUSM 4949); San Ignacio, Namballe, El Sauce
(MUSM 18171, 18224, 18225); San Ignacio, Chir-
inos, Nuevo Chalaquito. ‘‘El Chaupe’’ (MUSM
12884–12889); San Ignacio, Tabaconas (MUSM
10650). La Libertad:Sa´ nchez Carrio´ n, Sanagora´n
(MUSM 17268, 17269). Lambayeque:Ferren˜ afe,
Bosque Chin˜ama (MUSM 925, 926). Piura: Aya-
baca, ca. 44 km ESE by road Ayabaca, ladera Cerro
Mayordomo (MUSM 996–998); Huancabamba,
Carmen de la Frontera, Alto Samaniego (MUSM
18172). San Martı
´n: Huallaga, between ‘‘La Mor-
ada’’ and ‘‘La Rivera’’ (MUSM 16167–16169);
Mariscal Caceres, Parque Nacional Rı
´o Abiseo, La
Playa (MUSM 7280–7284); Mariscal Caceres, Par-
que Nacional Rı
´o Abiseo, Las Palmas (MUSM
7285–7293); Mariscal Caceres, Parque Nacional Rı
´o
Abiseo, Las Papayas (MUSM 7294).
Platyrrhinus lineatus (215).—BOLIVIA: El Beni:
Yacuma, Santiago (FMNH 105902). Santa Cruz:
Cercado, Andres Iban˜ ez (FMNH 50990*); Chiqui-
tos, Santiago (FMNH 105844*, 105845–105847*,
105848, 105849*, 105850–105853*, 105854–105856*,
105857–105862*, 105899–105901, 105903–105909*,
105910–105913). BRAZIL: Ceara´: Floresta Nacio-
nal Araripe, Crato (USNM 555706); Sitio Luanda,
Itaitera, Crato (USNM 555707). Ceara´ /Piauı
´: Serra
da Ibiapaba (FMNH 19516). Espı
´rito Santo: Rio
Doce (BMNH 23.12.12.9* [Holotype of Vampyrops
lineatus sacrillus Thomas, 1924]). Mato Grosso:
Serra do Roncador, 264 km N (by road) Xavantina
(USNM 393703). Mato Grosso do Sul: Rio Vacaria,
Fazenda Capa˜ o Bonita (FMNH 47962–47999,
48008; UV 10203); Urucu´ m (FMNH 26775–
26777*, 26778–26781*, 26782–26784, 30032–30048,
136867–136872; UV 10202). Minas Gerais:APA
Coqueiral, Southern MG (CMUFLA 112*, 115*);
Pains (USNM 391088); 3 mi ESE Sete Lagoas
(USNM 391087); Vicosa (USNM 391085, 391086,
391089, 541481–541484). Parana´: Salto Grande
(USNM 141392). Sa˜o Paulo: an Hembe [5municı
´-
pio Anhembe?] (USNM 484021); Iguape (USNM
542613, 542614); Piracicaba (ZMB 39963); Valpar-
aı
´so (FMNH 41645*). PARAGUAY: Alto Para-
guay: Fuerte Olimpo (FMNH 145257, 145258*).
Amambay: Parque Nacional Cerro Cora (USNM
552728, 554536). Central: Asuncio´n (MNHN 953*);
Asuncio´ n Recoleta (UMMZ 124321–124330,
124332, 125422–125443, 125876, 125877, 125879–
125901, 133734). La Cordillera: 1.6 km by road S
Tobati (UMMZ 125878). Misiones: 2.7 km (by
road) N San Antonio (UMMZ 124331). Paraguarı
´:
Parque Nacional Ybycuı
´(UMMZ 133732, USNM
531177, 531178); Saltos de Pirareta (UMMZ
133733); Sapucaı
´(BMNH 1.11.1.33*, 1.11.1.36*;
FMNH 48791*, USNM 115033, 115035–115043).
SURINAME: Nickerie: Sipaliwini Airstrip (CM
77663*).
Platyrrhinus masu (106).—BOLIVIA: Cocha-
bamba: 50 km NW Villa Tunari (UMMZ 126759,
126760). La Paz: Serranı
´a Ballavista (UMMZ
158068). PERU: Cusco: La Convencio´ n, Echarate,
Campamento Wayrapata (MUSM 14567–14572);
Huayopata (MUSM 989–992); La Convencio´n,
Quimbiri, Campamento Llactahuaman (MUSM
14559–14566); La Convencio´ n, Vilcabamba, Sum-
bayo, Pacaypata (MUSM 17034, 19150); Paucar-
tambo, Cosn˜ ipata, Challabamba, P. V. Acjanaco
(FMNH 170112, MUSM 8851); Paucartambo,
Cosn˜ipata. Consuelo (FMNH 123917* [Holotype
of Platyrrhinus masu Velazco, 2005], 123918,
123919; MUSM 9854); Paucartambo, Cosn˜ ipata,
La Esperanza (FMNH 174757–174760, MUSM
19732, 19733); Paucartambo, Cosn˜ipata, Pillahuata
(FMNH 171825, 172102; MUSM 11793–11795);
Paucartambo, Cosn˜ ipata, Parque Nacional Manu,
San Pedro (FMNH 170111, 170113, 172098–172101,
172103; MUSM 8853, 8854, 11792, 19446–19448;
UMMZ 160627, 160633); Paucartambo, Cosn˜ ipata,
VOLUME 122, NUMBER 3 279

Quitacalzon (MUSM 8852); Paucartambo, Cosn˜i-
pata, Suecia (MUSM 16706); Quispicanchi, Ca-
mante (FMNH 68455); Quispicanchi, Collpa de San
Lorenzo (FMNH 93594*); Quispicanchi, Hacienda
Cadena (FMNH 93588*); Urubamba, Machupic-
chu, Santuario Historico Machupijchu, Win˜ ay-
wayn˜a (MUSM 5442). Hua´ nuco:Hua´nuco,
Chinchao, E. slope Cordillera Carpish (MUSM
1011). Madre de Dios: Manu, Alto Rı
´o Madre de
Dios, Hacienda Amazonia (FMNH 139590,
139591*, 139592, 139593*, 139594, 139595, 139692,
139764; 9847, 9848); Manu, Cerro de Pantiacolla
(FMNH 122136*, 139596–139599*, 139600–
139603*, 139604–139607*, 139608; MUSM 9849–
9853). Pasco: Oxapampa, San Alberto (MUSM
10272, 10273, 14943); Oxapampa, Pozuzo, Delfin
(MUSM 12101–12105); Oxapampa, Pozuzo, Pal-
mira (MUSM 10945); Pasco, Paucartambo, Auqui-
marca, Anexo Santa Isabel (MUSM 15879–15884).
Platyrrhinus nigellus (303).—BOLIVIA: La Paz:
20 km NNE Caranavi (UMMZ 127174). COLOM-
BIA: Boyaca´: Municipio Villa de Leyva, Detras del
Instituto Humboldt, Sitio Hosteria (IAvH-M 6872);
Pajarito, Hacienda Camijoque (ICN 5441); Santa
Maria, Can˜ o Negro (ICN 17165*); Santa Maria,
margen izquierda Rı
´o Bata, sendero ecolo´gico (ICN
15066*); Santa Maria, Sitio Represa Chivor (ICN
16352*). Caqueta´ : Florencia, Corregimiento El
Caran˜o, Vereda Las Brisas, Finca Los Lirios
(UAM 190, 192). Cauca: Inza, Vereda Tierras
Blancas, Km 78, carretera Popayan – Inza (ICN
8467, 8468); Parque Natural Nacional ‘‘Munchi-
que,’’ Caban˜ a La Romelia (IAvH-M 3314, 3315).
Cesar: San Sebastia´ n (FMNH 69484*); Valledupar,
Villanueva, Sierra Negra (USNM 281304). Cundi-
namarca: Tena, alrededores laguna de Pedro Pablo
(ICN 5293*, 5295). Huila: Parque Natural Nacional
‘‘Cueva de los Huacharos’’ (IAvH-M 3311); Suaza,
Vereda Alto Campo Hermoso, Finca Marllins
(UAM 228). Magdalena: Santa Marta, Alto de
Mira, 3 km W Rı
´o Buritaca, Sierra Nevada de Santa
Marta (ICN 13087). Meta: Cubarral, Vereda Aguas
Claras, Escuela Santa Clara, Finca La Reforma
(ICN 14800*); Restrepo, Caney Alto, Botacoma
(ICN 10147*); Restrepo, Salinas de Upin (ICN
14372*). Narin˜o: El Carmen (FMNH 113713,
113721, 113730–113732, 113734, 113891, 113892,
113894–113897). Norte de Santander: Municipio
Herra´n, Parque Natural Nacional Tama, Sector
Orocue´ (IAvH-M 6631–6637, 6672); Municipio
Toledo, Rı
´o Negro, Finca ‘‘San Isidro’’ de Pablo
Contreras, Parque Natural Nacional Tama (IAvH-
M 6678, 6685, 6689, 6701–6704, 6710, 6715, 6782);
Municipio Toledo, Vereda el Diamante, Cerro San
Agustin, Parque Natural Nacional Tama (IAvH-M
6719, 6721, 6722, 6734, 6739). Putumayo: Municipio
Mocoa, carretera entre Sibundoy y Mocoa, Locali-
dad el Mirador (IAvH-M 6817, 6819, 6825).
Quindı
´o: Salento, Reserva Natural Can˜on Quindı
´o
(ICN 12441–12448*). Risaralda: Pueblo Rico (ICN
11934*, 11937*). Santander: Charala, Inspeccio´n de
policı
´a Virolin (ICN 8150, 8972*, 8973); Encino,
Vereda La Cabuya, Finca San Benito (ICN 17588);
Encino, Vereda Los Pericos, Finca Vegaleo´ n (ICN
17587); Encino, Vereda Rı
´o Negro, sitio Las Tapias,
Finca El Aserradero (ICN 17585, 17586); Tona,
Sitio El Mortin˜ o (ICN 17187, 17188*); Tona,
Vereda Guarumales, Finca El Pajal (ICN 16660,
16661). Valle del Cauca: Bolivar, La Tulia, Hacienda
La Argelia (UV 12112); Buga, La Magdalena,
Janeiro, Hacienda Santelina (UV 13024); El A
´guila,
Quebrada Charco Azul (UV 12522, 12559); El
Cairo, Estacio´ n Cerro de Ingle´s (UV 12242,
12243); El Cairo, Quebrada Charco Azul (UV
12301–12304, 12306, 12307); Florida, Hacienda
Los Alpes (UV 3525, 3526). ECUADOR: Azuay:
Yunguilla Valley (FMNH 53504). El Oro: 1 km SW
Puente de Moromoro (USNM 513465*). Napo:
Cascada San Rafael (FMNH 124987*). Pastaza:
Mera (USNM 548189–548195*, 548196–548210,
548399). PERU: Amazonas: Cordillera del Condor,
Valle Rı
´o Comaina; Puesto Vigilancia 3, Alfonso
Ugarte (USNM 581960*). Ayacucho: La Mar, Ayna,
Ccentabamba (MUSM 21405). Cajamarca: Jae´n,
Chontali, Las Ashitas (MUSM 10652); San Ignacio,
Namballe, El Sauce (MUSM 18174, 18226–18229);
San Ignacio, Chirinos, Nuevo Chalaquito, ‘‘El
Chaupe’’ (MUSM 12890–12892); San Ignacio,
Tabaconas, 4 km W El Chaupe (MUSM 10651).
Cusco: La Convencio´ n, Echarate, Camisea, Cashir-
iari (MUSM 13863, 13864); La Convencio´n, Echa-
rate, Campamento Wayrapata (MUSM 14579); La
Convencio´ n, Echarate, 2 km SW C. N. Tangoshiari
(MUSM 13394); La Convencio´ n, Pichari, Catarata
(MUSM 21406); La Convencio´ n, Quimbiri, Cam-
pamento Llactahuaman (MUSM 14573–14578); La
Convencio´ n, Ridge Campamento (USNM 588031*);
Paucartambo, Challabamba, Bosque de las Nubes
(MUSM 8857); Paucartambo, Cosn˜ ipata, Consuelo
(FMNH 123934–123947*, 123948, 123949, 174773–
174783; MUSM 9970–9975, 19734–19738); Paucar-
tambo, Cosn˜ ipata, Parque Nacional Manu, San
Pedro (FMNH 172104–172106, MUSM 8861,
11796, 15409, 19449); Paucartambo, Cosn˜ipata,
Tono (FMNH 139589, 139759, 140019; MUSM
9977); Paucartambo, Cosn˜ ipata, Quitacalzon
(MUSM 8858–8860); Quispicanchi, Collpa de San
Lorenzo (FMNH 93589–93593*, 93595–93597*,
93607*); Quispicanchi, Hacienda Cadena (FMNH
93598–93601*, 93602–93606*). Junı
´n: Satipo, Rı
´o
Tambo, Cordillera del Vilcabamba (MUSM 12995).
Madre de Dios: Manu, Alto Rı
´o Madre de Dios,
Hacienda Amazonia (FMNH 126031, 126032*,
139573, 139761–139763, 139765–139775, 139778–
139784; MUSM 9955–9962); Manu, Cerro de
Pantiacolla (FMNH 139576–139578*, 139579,
139580*, 139786–139803; MUSM 9963–9969,
9976); Manu, Rı
´o Palotoa (FMNH 139804). Pasco:
280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Oxapampa, Pozuzo, Palmira (MUSM 10943, 10944,
10971, 10972). San Martı
´n: Huallaga, entre ‘‘La
Morada’’ y ‘‘La Rivera’’ (MUSM 16170–16179);
Mariscal Caceres, Parque Nacional Rı
´o Abiseo, Las
Palmas (MUSM 7295, 7296). VENEZUELA: Me´r-
ida: Pedregosa, Pedregosa Norte, 3 km N Puente la
Pedregosa (UV 11832).
Platyrrhinus recifinus (29).—BRAZIL: Espı
´rito
Santo: Municı
´pio Linhares, Fazenda Santa Tere-
zinha (MVZ 185898); Municı
´pio Santa Teresa,
Museu Mello Leitao (MVZ 185897). Minas Gerais:
APA Coqueiral (CMUFLA 113*, 114*); Marlie´ria,
PERD-Campo de Pouso (VCT 06); Marlie´ria,
PERD-Campolina (VCT 08, 09, 11). Pernambuco:
Pernambuco [Recife] (BMNH 81.2.16.4* [Holotype
of Vampyrops recifinus Thomas, 1901]). Sa˜o Paulo:
(USNM 545001, 545002); Guaratuba (USNM
542611, 542612); Ilha de Sa˜ o Sebastia˜ o, Parque
Estadual de Ilhabela (MVZ 185604–185606, 185610,
185895, 185896); Itanhae´m (USNM 542615); Mu-
nicı
´pio de Cajuru, Fazenda Santa Carlotta (MVZ
185899, 185900); Municı
´pio de Saleso´ polis, Estac¸a˜o
Biolo´gica de Boraceia, Museu Zoologia da USP
(MVZ 185901). GUYANA: Potaro-Siparuni:Iwok-
rawa Reserve, 35 km SW of Kurupukari (ROM
108487). Upper Takutu-Upper Essequibo: Pobawau
Creek Mouth (ROM 113465). SURINAME: Bro-
kopondo: Brownsberg Nature Park, Headquarters
(ROM 113991, 114070); Brownsberg Nature Park,
Km 2.4, Wittie Kreek Trail (ROM 114195).
Platyrrhinus vittatus (136).—COLOMBIA: Anti-
oquia: 25 km S and 22 km W of Zaragoza, La
Tirana (IAvH-M 918, 928; USNM 499456); 24 km S
and 22 km W of Zaragoza, Buenos Aires (USNM
499455*); 26 km S and 22 km W of Zaragoza,
Aljibes (USNM 499457*); Carmen del Viboral, Los
Rı
´os (MUA 10140); Urrao, Calles (MUA 10768,
10881); Vereda Santiago, Anori (MUA 10943).
Magdalena: San Pedro de la Sierra, Finca Tierra
Grata (ICN 5258); Santa Marta, Sierra Nevada de
Santa Marta, Alto de Mira, 3 km W Rı
´o Buritaca
(ICN 13025); Santa Marta, Serranı
´a San Lorenzo,
estacio´n de Inderena (ICN 5394, 5398); Santa
Marta, Serranı
´a San Lorenzo, Hacienda La Victoria
(ICN 5393). Narin˜o: Ricaurte, Hacienda La Pla-
nada, Buenos Aires (UV 2940, 2941, 3006). Valle del
Cauca: Buga, Estacio´ n Biolo´ gica El Vincu´lo (UV
8144, 8145); Dagua, El Queramal, Tokio. Antena de
Telecori (UV 2028); El Cairo, Estacio´ n Cerro de
Ingle´s (UV 12447); Yotoco, Reserva Forestal de
Rocoto (UV 2029). COSTA RICA: Puntarenas:
Monteverde (ROM 97305, UMMZ 116681,
116682). PANAMA: Bocas del Toro: Upper Rı
´o
Changena, Rancho Caballero, 20 mi SSW Changui-
nola (USNM 319278, 319279*); Upper Rı
´o Chan-
gena, Rancho Mojica, 20 mi SSW Changuinola
(USNM 319277*, 319280, 319281); Rı
´o Changena
Camp (USNM 319390–319393*, 319394–319399*,
319400–319403, 319494, 319495*); 25 km NNE San
Felix (USNM 541098). Chiriquı
´: Cerro Punta
(USNM 314244*); 1 mi E Cuesta de Piedra (USNM
331674, 331675*); Finca Santa Clara, 14.5 km NW
El Volca´n (USNM 518012, 537597); Osta Clara,
15.5 km NW El Volca´ n (USNM 539879–539881);
4 mi S El Volca´ n (USNM331676*). Cocle´: La Mesa,
3.7 km NE El Valle (USNM 457950*). Darie´n: ca.
6 km NW Cana, E Slope Cerro Pirre (LSUMZ M-
515); Cerro Mali (USNM 338029, 338030*, 338031–
338040); Tacarcuna Mali Camp (USNM 309755*);
Tacarcuna Village Camp (USNM 309691–309733*,
309734–309737*, 309738–309741*, 309742, 309743*,
309744–309754*). VENEZUELA: Carabobo, Puerto
Cabello (ZMB 568 [Holotype of Artibeus vittatus
Peters, 1859]).
VOLUME 122, NUMBER 3 281