New species of Anoura (Chiroptera: Phyllostomidae) from Colombia, with systematic remarks and notes on the distribution of the A. geoffroyi complex

TEXAS TECH UNIVERSITY

Natural Science Research Laboratory

Occasional Papers

Museum of Texas Tech University

Number 292 19 May 2010

New Species of Anoura (Chiroptera: Phyllostomidae)
from Colombia, with Systematic Remarks and Notes on
the Distribution of the A. geoffroyi Complex

Front cover: Map representing the collecting localities of new species Anoura carishina in Colombia. Upper right
inset: Anoura geoffroyi peruana specimen FMNH 49178, from Cundinamarca, Colombia. Lower right inset: Skull
comparisons between carishina paratype $ ICN 14531 (left, within box), from Taminango, Narino, Colombia, and
other members of the A. geoffroyi complex. Illustration by Hugo Mantilla-Meluk.

New Species of Anoura (Chiroptera: Phyllostomidae) from Colombia, with
Systematic Remarks and Notes on the Distribution of the A. geoffroyi

Complex

Hugo Mantilla-Meluk and Robert J. Baker

Abstract

A new species of nectar-feeding tailless bat in the genus Anoura (Chiroptera: Phyllosto¬
midae) is described from the highlands of the Colombian Andes and the independent mountain
system of the Sierra Nevada de Santa Marta. Complete zygomatic arches, a relatively wide
uropatagium, and wide first upper molars with poorly developed paracones are proposed as
synapomorphies of the new species and A. geoffroyi geoffroyi. However, the two taxa are allo-
patrically distributed, and the new species is morphologically distinguished from A g. geoffroyi
by a smaller skull and body size, more massive and squared molars with wider hypoconal basins,
smaller P4 that are not laterally compressed, reduced anterobasal cusps, and medial internal cusps
that are enlarged. Based on morphological analysis the distributional ranges of A. geoffroyi sub¬
species were interpreted as follows: A. g. lasiopyga is restricted to Central America from Costa
Rica north to Mexico; A. g. peruana is restricted to the mid to high elevations of the Andean
system from Bolivia to Colombia; and A. g. geoffroyi is restricted to the mid and low elevations
of eastern versant of the Andes from Brazil to northern South America, including the island of
Trinidad. Further, the morphological affinities between A g. apolinari and members of the A.
geoffroyi complex support its current recognition as a junior synonym of A. g. peruana. Based
on the morphological distinction observed between A. g. peruana and A. g. geoffroyi , including
the absence of complete zygomatic arches, a more delicate rostrum, less massive molars, and
overall darker coat coloration, as well as the ecological differentiation of the areas inhabited by
these two taxa, we recommend the elevation of A. peruana to specific level.

Key words: Anoura , bats, Colombia, new species

Resumen

Se describe una nueva especie de murcielago nectarivoro sin cola del genero Anoura
(Chiroptera: Phyllostomidae) proveniente de los Andes y el sistema montanoso independiente
de la Sierra Nevada de Santa Marta en Colombia. La presencia de arcos zigomaticos comple¬
tes, un uropatagio relativamente amplio y molares superiores amplios, son propuestos como
sinapomorfias para la nueva especie y A. geoffroyi geoffroyi. Sin embargo, estos dos taxa se
encuentran alopatricamente distribuidos y la nueva especie se distingue morfologicamente de
A. g. geoffroyi por tener un menor tamano de craneo y menor tamano corporal, molares mas
masivos y cuadrados, con las fosas de los hipoconos mas amplias, los P4 de un tamano menor,
no lateralmente comprimidos y cuspides anterobasales de mayor tamano. Basados en analisis
morfologicos interpretamos la distribucion de las subespecies de A. geoffroyi como sigue: A. g.
lasiopyga es restringida a las elevaciones medias y altas de Centro America desde Costa Rica
hasta Mexico; A. g. peruana es restringida a las elevaciones medias y altas del sistema Andino
desde Bolivia hasta Colombia; y A. g. geoffroyi es restringida a las tierras medias y bajas de la

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Occasional Papers, Museum of Texas Tech University

vertiente oriental de los Andes desde Brasil hasta el norte de Sur America, incluyendo la isla
de Trinidad. Adicionalmente, las afinidades morfologicas entre A. g. apolinari y los miembros
del complejo A. geoffroyi apoyan su actual reconocimiento como sinonimo menor de A. g. pe¬
ruana. Basados en las diferencias morfologicas observadas entre A. g. peruanay A. g. geoffroyi,
incluyendo la ausencia de arcos zigomaticos completos, rostro mas delicado, molares menos
masivos y una coloracion del pelaje mas oscura, asi como tambien la diferenciacion ecologica
entre las areas habitadas por estos dos taxa recomendamos la elevacion de A. peruana a estado
especifico.

Palabras clave: Anoura , Colombia, murcielagos, nueva especie

Introduction

Nectarivorous bats of the genus Anoura Gray
1838 (Glossophaginae: Choeronycterini) (Baker et
al. 2003) currently are recognized as eight species:
A. caudifer (Geoffroy St.-Hilaire 1818); A. geoffroyi
Gray 1838; A. aequatoris (Lonnberg 1921); A. cultrata
Handley 1960; A. latidens Handley 1984; A. luismanu-
eli Molinari 1994; A. fistulata Muchhala, Mena, and
Albuja 2005; and A. cadenai Mantilla-Meluk and Baker
2006. Anoura geoffroyi is the most widely distributed
and morphometrically variable species in the genus and
includes three recognized subspecies: A. g. geoffroyi
Gray 1838, from Rio de Janeiro, Brazil (type locality
accompanied by a question mark in the description), re¬
stricted to Rio de Janeiro by C. O. C. Vieira (1942:324);
A. g. peruana (Tschudi 1844), from Junin, Peru; and
A. g. lasiopyga Peters 1868, type locality “Mexico”
restricted to the state of Veracruz, Mexico, by Arroyo-
Cabrales and Gardner (2003:740). The uncertainty
of the geographic origin of some of the holotypes has
made it difficult to define the geographic boundaries of
the subspecies, or to validate potential new taxa within
the A. geoffroyi complex. Two junior synonyms exist
for Anoura g. peruana : A. g. apolinari (Allen 1916),
described from Boqueron de San Francisco, near Bo¬
gota, Cundinamarca, Colombia; and A. g. antricola

Anthony 1921, described from Loja, Ecuador. Al¬
though A. geoffroyi has been documented in Colombia
(Mantilla-Meluk et al. 2009), the hypothesized presence
of A. geoffroyi geographic variants in northern South
America often has lacked a detailed systematic analysis,
and their geographic limits remain poorly defined. In a
recent examination of Anoura specimens deposited in
the Colombian collections of the Instituto de Ciencias
Naturales of the Universidad Nacional de Colombia,
we discovered a distinctive and undescribed species of
Anoura previously identified as A. geoffroyi. Herein,
we describe this new species of Anoura and use mul¬
tivariate methods to evaluate the phenetic similarities
among recognized taxa within the A. geoffroyi complex
(A. g. geoffroyi , A. g. lasiopyga , and A. g. peruana).
Due to the unique opportunity to conduct systematic
analyses of a relatively large number of A. geoffroyi
voucher specimens from Cundinamarca, Colombia, we
also re-examine the taxonomic status of A g. apolinari.
Further, we evaluate the presence of A. g. geoffroyi and
A. g. lasiopyga in Colombia and take advantage of the
wide geographic extent of the available specimens to
further refine the distributional limits of these two taxa
in Central America.

Materials and Methods

Data collection. —We examined 268 adult A.
geoffroyi specimens (149 males and 119 females),
from Bolivia (2), Brazil (3), Colombia (155), Costa
Rica (2), Ecuador (6), Guatemala (8), Mexico (27),
Peru (33), and Trinidad (31), deposited in the follow¬

ing institutions: American Museum of Natural History
(AMNH); Instituto de Ciencias Naturales of the Univer¬
sidad Nacional de Colombia (ICN); Field Museum of
Natural History (FMNH); Musee D’Histoire Naturelle
de Geneve (MHNG); Museo de Historia Natural de la

Mantilla-Meluk and Baker—New Species of Anoura from Colombia

3

Universidad del Cauca (MHNUC); Museo de la Uni-
versidad Distrital Francisco Jose de Caldas (MUD);
Southwestern Biology Collection of the University of
New Mexico (MSB); and the Museum of Texas Tech
University (TTU) (Appendix). Relative age of each
specimen was determined based on the degree of os¬
sification of phalangeal epiphyses and the completeness
of ossification of the basisphenoid suture. For each
specimen, we recorded 12 craniodental measurements
with digital calipers to the nearest 0.01 mm as follows:
greatest length of the skull (GLS), greatest distance
from the anteriormost projection of the nasal bones to
the posteriormost portion of the occipital bone; con-
dylobasal length (CB), distance from the anteriormost
projection of the premaxillae to the posteriormost
projection of the exoccipital condyles; palatal length
(PAL), distance from the anteriormost point of the pre¬
maxilla (excluding incisors) to the posterior margin of
the horizontal process of the palatine, just in the midline
of the horizontal process of the palatine; postorbital
constriction (PO), least width across the interorbital
constriction at a right angle to the longitudinal axis of
the cranium; mastoid breadth (MB), greatest breadth
across the lateral margins of the parietal at the posterior
region to the suture coronalis (measured at a right angle
to the longitudinal axis of the cranium); anterior brain-
case breadth (BCW); braincase height (BCH), distance
from the juncture of the midline and frontal-parietal
sutures to the inferiormost point of the glenoid fossae;
maxillary toothrow length (LTR); M3 breadth (M-M),
greatest breadth across the lateral margins of M3 at a
right angle to the longitudinal axis of the tooth; greatest
breadth across the canines (C-C), distance between the
externalmost point of the canine alveoli; mandibular
condylocanine length (ML), greatest distance from the
anteriormost point of the alveolus of the lower canines
to the posteriormost point of the mandibular condyles;

and mandibular toothrow length (MTR), greatest dis¬
tance from the anteriormost surface of il to the poste¬
riormost surface of m3. Although for this work only
dry skins were analyzed, we recorded the relative size
of the uropatagium (UL), relative size of the vibrissae
(VL), and the relative size of the thumb (TL).

Sexual dimorphism. —To identify those measure¬
ments that contribute the most to skull variation among
large Anoura , we performed a Principal Components
Analysis (PCA) among 234 Anoura specimens (125
males and 109 females) with complete data in this
work, including all recognized A. geoffroyi subspecies
and the newly described taxon; then, a MANOVA was
performed on six craniodental variables that accounted
for most of the variation (GLS, CB, PAL, M-M, C-C,
ML) between male and female A. geoffroyi to test
for sexual dimorphism in the R statistical package,
script available at: http://faculfybiol.ttu.edu/strauss/
Multivar/R/SampleDFAManova.R.txt.

Multivariate analysis. —To investigate and de¬
scribe the morphometric variation and phenetic similar¬
ity between the new species and other representatives
of A. geoffroyi across its geographic range, we applied
Principal Component Analyses to the twelve cranioden¬
tal variables listed above of260^4. geoffroyi specimens
(143 males and 117 females) in the statistical package
PAST, http ://folk.uio .no/ohammer/past/download.html.
Due to the existence of missing data within the sample
(0.99%), the “missem” MATLAB function, available at
www.ttu.edu/strauss/Matlab/Matlab.htm, was applied
to find the most likely values for those missing data.
Female and male measurements were analyzed sepa¬
rately because of significant differences observed in
skull size in our MANOVA.

Results

Sexual dimorphism. —Sexual dimorphism was
confirmed among 234 A. geoffroyi specimens in this
study (125 males and 109 females). A MANOVA
performed on the six craniodental variables that con¬
tributed the most to skull variation in this analysis
(GLS, CB, PAL, M-M, C-C, ML), show significant
differences (F = 9.33; p < 0.001) between^, geoffroyi
males and females.

Principal Component Analyses. —Principal
Component Analyses revealed high levels of intra¬
subspecific morphological variation for both male and
female A. geoffroyi. All subspecies of A. geoffroyi
extensively overlapped in morphospace. Anourag. pe¬
ruana had the widest range of morphometric variation
with A. g. geoffroyi and A. g. lasiopyga present within
the morphospace of A. g. peruana (Fig. 1). The greater

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Occasional Papers, Museum of Texas Tech University

PC 1 (65.46 %)

PC 1 (61.8 %)

Figure 1. Scatter plot showing the results of Principal Component Analyses applied to 12 craniodental
measurements of adult males (top) and adult females (bottom) representing: new species Anoura
carishina (black stars), holotype A. g. apolinari AMNH 37376 (star in grey circle), A. g. peruana , A.
g. lasiopyga , and A. g. geoffroyi.

Mantilla-Meluk and Baker—New Species of A no lira from Colombia

5

Table 1. Principal component scores of the percentage of variation explained by the
first two components and loadings of the analyzed variables for both A. geoffroyi
(sensu lato) males andfemales. GLS: Greatest length of skull; CB: Condylobasal
length; PAL: Palatal length; PO: Post orbital constriction width; MB: Mastoid
breadth; BCW: Braincase width; BCH: Braincase height; LTR: Tooth-row length;
M-M: Distance across upper molars; C-C: Distance across upper canines; ML:
Mandible length; MTR: Longitude of the mandibular tooth-row.

s

(

$

PCI

PC2

PCI

PC2

% Variation

61.8

9.7

65.466

11.35363

GLS

-0.4848

0.237

-0.2478

0.4858

CB

-0.5252

0.1312

-0.3028

0.5304

PAL

-0.4763

-0.6418

0.8558

0.4518

PO

-0.0071

0.132

-0.057

0.03148

MB

-0.0485

0.3904

0.05467

0.0784

BCW

-0.0242

0.298

0.08931

0.07177

BCH

0.01131

0.4585

0.1636

0.03369

LTR

-0.1987

0.05901

-0.1708

0.186

M-M

-0.0442

0.02493

-0.0548

0.05301

C-C

-0.0318

0.09404

-0.0214

0.09798

ML

-0.4099

0.1659

-0.1096

0.4082

MTR

-0.2207

0.07332

-0.1703

0.2267

variation found among//, g. peruana can be also related
to its larger sample size in this analysis.

Palatal length accounted for most of the variation
among males and females, followed by condylobasal
length and greatest length of the skull, with all having
negative loadings in females and positive loadings in
males (Table 1).

Anourag. peruana from the departments of Aya-
cucho, Huanuco, and Ollantaytambo in Peru, Pichincha
in Ecuador, and Caldas and Quindio in Colombia were
characterized by a larger average size (GLS >25.5 mm)
in comparison with specimens of A. g. peruana from

other localities, including specimens from Cundina-
marca, Colombia. Differences in size between A. g.
peruana from Peru and A. g. peruana from Cundina-
marca, Colombia, were more evident among females
than in males (Fig. 1).

Some specimens from the mid- to high elevations
in the departments of Magdalena, Narino, and Valle del
Cauca were among the smallest representatives of the
A. geoffroyi complex (GLS <24.5 mm). The above
mentioned specimens shared the complete zygomatic
arches (or evidence of this condition) and relatively
large uropatagia of A. g. geoffroyi from the lowlands.

Discussion

Taxonomic remarks. —The systematics of the ge¬
nus Anoura has been a source of debates on the validity
of morphology to identify independent evolutionary
units (Sanborn 1933; Cabrera 1957; Mantilla-Meluk

and Baker 2006; Jarrin-V and Kunz 2008). The ob¬
served morphological variation among large Anoura
in the A. geoffroyi complex has been interpreted as
geographic variants, resulting in the designation of

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Occasional Papers, Museum of Texas Tech University

three subspecies: A. g. geoffroyi , A. g. lasiopyga, and
A. g. peruana. Up to date, it has been difficult to
establish the geographic boundaries of A. geoffroyi
geographic variants. Geographic distributions of these
three taxa have been established based primarily on the
indiscriminate usage of available Linnean epithets. In
addition, two currently recognized subspecies, A. g.
peruana and A. g. lasiopyga, were described before
1868 in different genera than Anoura ( Choeronyct-
eris peruana and Glossonycteris lasiopyga ), making
difficult the identification of diagnostic characters to
separate them from other members of the genus. We
used morphological criteria derived from the study of
the holotypes of Glossophaga apolinari Allen 1916
(A. g. apolinari ) and Glossonycteris lasiopyga Peters
1868 (A. g. lasiopyga ) to reassess the taxonomic status
of A. peruana and these two taxa and to define their
geographic ranges.

Taxonomic status of Anoura g. apolinari.—
Anoura g. apolinari is currently considered a junior
synonym of A. g. peruana (Simmons 2005). This
taxon, with type locality in Boqueron de San Fran¬
cisco, near Bogota, Cundinamarca, Colombia, was
described based on six specimens (sex not defined by
the author Allen 1916). However, the holotype of G.
apolinari (AMNH 37376, GLS = 26.0 mm) has the
long and delicate rostrum, the smaller and less later¬
ally divergent canines, and less pronounced zygomatic
processes typical of Peruvian female specimens of A.
g. peruana (Fig. 2). Sexual dimorphism was observed
among A. g. peruana specimens included in this work,
with females having an overall larger size than males
in all analyzed measurements (Table 2). Based on
the morphology observed, as well as morphometric
ranges documented, we concluded that specimen
AMNH 37376 used by Allen (1916) as holotype of
G. apolinari is within the measurements of female A.
g. peruana. In our PC A, the holotype of G. apolinari
was within the morphospace of A. g. peruana. Based
on the observed similarities in discrete characters, as
well as the overlapping morphometric ranges, it is dif¬
ficult to distinguish G. apolinari from A. g. peruana.
Therefore, we recommend maintaining G. apolinari as
junior synonym of A. g. peruana and the application of
the epithet A. g. peruana for A. geoffroyi Colombian
specimens characterized by a large skull (GLS > 25.5),
incomplete zygomatic arches, reduced uropatagium,
and dark coat color, ranging from Mummy-Brown to
Black (Ridgeway 1912) as well as long hair lengths.

Geographic distributions. —The most recent
works including distributional maps for A. geoffroyi
subspecies in northern South America differ in their
interpretation of the geographic limits of A. g. peruana
and A. g. lasiopyga in Colombia (Gardner 2008 [2007];
Ortega and Alarcon-D 2008). Gardner (2008 [2007])
reported A. g. peruana for the Colombian territory
and mentioned that A. lasiopyga , only reported for
Ecuador in South America (Albuja 1983), is potentially
present on the western side of Colombia. In contrast,
although Ortega and Alarcon-D (2008) accepted G.
apolinari (holotype from Cundinamarca, Colombia) as
a synonym of A. g. peruana , the taxon was excluded
from Colombia in their distribution map. In addition,
Ortega and Alarcon-D (2008) extended the distribu¬
tion of A. g. lasiopyga to the entire Andean region in
Colombia. We did not find support for the hypothesis
of Ortega and Alarcon-D (2008) in the morphology
observed in our specimens examined. In our samples,
the Colombian specimens of A. geoffroyi with incom¬
plete zygomatic arches matched the description of A.
g. peruana. Compared with the G. lasiopyga holotype
(MHNG 515.88), Colombian samples presented a more
acute palatal angle, a darker coat color, and relatively
longer hair (Figs. 2 and 3). The same characteristics
were observed in specimens from Ecuador analyzed
in this work (N = 6; GLS mean = 25.58 mm, StDv
0.51) (Appendix).

Until 1957, A. g. lasiopyga was known only from
41 specimens, all from localities north of Guatemala
(Goodwin 1934). Anderson (1957) extended the dis¬
tribution of A. g. lasiopyga into Costa Rica based on
one specimen (KU 39249) from San Rafael, Canton
de Tarrazu (misspelled on tag as Terrazi), 30 km S
from San Jose (1300 m). Anderson (1957) compared
specimen KU 39249 with a series of 13 specimens (KU
60635-46, KU 60978-61076) from Teopisca, Chiapas,
and mentioned that despite differences in external
color and cranial measurements, specimen KU 39249
was within the ranges proposed for A g. lasiopyga. In
addition, specimen KU 39249 was paler than typical
A. g. peruana from South America and both the Costa
Rican and the Mexican specimens averaged smaller
than those of A. g. peruana in all analyzed measure¬
ments (Anderson 1957) (Table 2). Among A. geoffroyi
from Middle America examined in this work, two color
phases were distinguished: a) pale phase (Sanborn
1933) (Light-Gray to Light-Drab, Ridgeway 1912), also
characterized by short hair, represented in our samples

Mantilla-Meluk and Baker—New Species of Anoura from Colombia

7

Figure 2. Comparison between A. g. peruana (FMNH S 169842, Pucataitambo,
Peru; FMNH $ 75160, Andahuaylas, Peru) and the holotype of A g. apolinari (Allen
1916) (AMNH $ 37376, San Francisco, Bogota, Colombia) (upper row). Difference
in palatal angle between holotype of Glossonycteris lasiopyga (MHNG 515.88 <$)
and A. g. peruana from Colombia (ICN 15157 S) (lower row).

Peru

Colombia

FMNH 169842 FMNH 75160 AMNH 37376
Male Female Female

G. lasiopyga MHNG 515.88 (Male)

A. g. peruana ICN 15157 (Male)

Table 2. Average values of the 12 analyzed craniodental measurements (mm) of large Anoura specimens. Abbreviations: GLS: Greatest

8

Occasional Papers, Museum of Texas Tech University

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Mantilla-Meluk and Baker—New Species of A no lira from Colombia

9

A. g. lasiopyga

Hidalgo, Mexico

icn 14530 (Male) icn 14531 (Female)

Figure 3. Differences in coat color and hair length among ,4. g. lasiopyga from Mexico (TTU 15474
S, TTU 15479 S, and TTU 38035 <J) (top). Skin of A. carishina holotype (ICN 14530 <$) and a
female (ICN 14531 9) of the type series (bottom).

10

Occasional Papers, Museum of Texas Tech University

by specimens from the Mexican state of Hidalgo; and b)
dark phase (dusky-brown to blackish-brown, Ridgeway
1912), also characterized by longer hair, represented
in our sample by specimens from the Mexican states
of Hidalgo, Tamaulipas, Tlaxcala, and Veracruz, as
well as samples from Olancho, Honduras, and Punta
Arenas, Costa Rica that to some extent resemble the
typical coat color pattern observed in A. g. geoffroyi
(Fig. 3) (Appendix). Further analyses using datasets
other than morphology will be necessary to elucidate
the taxonomic affinities between Middle American
A. geoffroyi populations and A. g. geoffroyi from the
lowlands to mid-elevations on the eastern versant of
the Andes and the island of Trinidad.

Although A. g. peruana had the largest range of
variation in skull size among samples in this work,
specimens from Bolivia, Colombia, Ecuador, and Peru
were clearly distinguishable from typical A. g. geof¬
froyi from the lowlands along eastern South America,
including specimens from Brazil, by the absence of
zygomatic arches, more delicate rostrum, less massive
molars and premolars, and a darker general coat color
from Mummy-Brown to Black (Ridgeway 1912), with
no intermediate forms. Based on the morphological
distinction between A. g. peruana and A. g. geoffroyi
we support the elevation of A. peruana (Tschudi 1844)
to specific status.

Size variation in Anoura peruana.—Samples
of Anoura peruana in this work included the widest
ranges of size variation for all the measurements ana¬
lyzed (Table 2). Greatest length of the skull (GLS),
representing the measurement that accounted for most
of the variation among all taxa analyzed, ranged from
24.05 to 26.51 mm (mean = 25.04; StDev = 0.61) in
A. peruana. Among our samples, large and small A.
peruana were found sympatrically in some localities on
the Eastern Cordillera of the Colombia Andes as well
as the northernmost range of the Central Cordillera; in
contrast, A. peruana from Peru and the southern portion
of the Central and Western Cordilleras of the Colombia
Andes only included specimens characterized by skulls
larger than 25.05 mm. Anoura peruana had the widest
range of geographic distribution including contrasting
ecosystems across the Andean range. We suggest that
size variation in A. peruana may be interpreted as an
adaptive response of this taxon to the heterogeneity of

the environments included in its distributional range.
Nonetheless, a more detailed analysis including inde¬
pendent data sets is necessary to clarify the taxonomic
affinities of A. peruana specimens with small skull
size (GLS < 25.5) from the Eastern Cordillera and the
northern range of the Colombian Andes.

Undescribed morphological variation in our
samples. —Among the smaller A. geoffroyi specimens
(GLS <24.5 mm) examined in our morphological as¬
sessment we found an unrecognized taxon character¬
ized by complete zygomatic arches and relatively wide
uropatagium from the mid- and high elevations of the
Colombian Andes and from the geographically isolated
range of the Sierra Nevada de Santa Marta. A formal
description of this taxon is presented below.

Family Phyllostomidae Gray 1825
Genus Anoura Gray 1838
Anoura carishina sp. nov.

Holotype. —Adult male ICN 14530 from Tami-
nango, corregimiento de Remolino, department of Nari-
no, collected by A. Albesiano, C. Ariza, J. Alvarez, and
H. Mantilla-Meluk (in the field notes the holotype of
A. carishina was identified with group field collection
number 21). The holotype, preserved as a skin-and-
skull voucher specimen in excellent condition (Figs. 3
and 4) (Table 3), was collected on 5 December 1996.

Paratypes. —The type series consists of four indi¬
viduals prepared as skin-and-skull voucher specimens
as follows: an adult female (ICN 14531), collected by
A. Albesiano, C. Ariza, J. Alvarez, and H. Mantilla-
Meluk (collectors number 36) on 9 December 1996 at
the type locality; an adult male (ICN 5224) collected at
San Pedro de La Sierra, tinea Tierra Grata, department
of Magdalena, Colombia (10°54'06"N, 74°02'00"W)
at 1320 m, on 26 May 1976 by A. Cadena Garcia
(collector number ACG 1147); an adult female (ICN
5225) from the same locality collected at 1320 m by
A. Cadena Garcia, M. Romer, and B. Moreno (without
an assigned field number and date); and an adult male
(ICN 5938) from Pance, department of Valle del Cauca,
Colombia (3° 19'42"N, 76°38T9"W), collected at 1200
m, on 5 February 1947 by M. Thomas (collector number
3299) (Fig. 5, Table 3).

Mantilla-Meluk and Baker—New Species of Anoura from Colombia

11

Figure 4. Ventral (upper photos) and lateral (lower photo) views of Anoura carishina
holotype skull and mandibles.

Table 3. Average values of the 12 craniodental measurements analyzedfor Anoura carishina holotype (*) and the type
series. Abbreviations: GLS: Greatest length of skull; CB: Condylobasal length; PAL: Palatal length; PO: Postorbital
constriction width; MB: Mastoid breadth; BCW: Braincase width; BCH: Braincase height; TR: Tooth-row length;
M-M: Distance across upper molars; C-C: Distance across upper canines; ML: Mandible length; MTR: Longitude of
the mandibular tooth-row.

ICN

Sex

GLS

CB

PAL

PO

MB

BCW

BCH

TR

M-M

C-C

ML

MTR

14530*

dP

24.52

23.5

12.05

4.88

9.03

9.77

7.65

9.3

6.06

4.38

16.58

9.65

5224

4

24.13

23.3

12.3

5.33

9.48

9.96

7.49

9.12

6.31

4.48

16.83

9.41

5938

24.27

23.3

12.52

5.16

9.29

9.77

7.41

9.18

5.97

4.5

16.61

9.52

14531

•$

24.04

23.28

11.77

4.99

9.07

9.8

7.48

8.93

6.09

4.15

16.66

9.49

5225

?

24.04

23.34

12.18

4.72

8.69

9.3

6.69

9.03

5.88

4.07

16.49

9.45

12

Occasional Papers, Museum of Texas Tech University

Figure 5. Type series of A. carishina'. Holotype ICN 14530 S from Taminango, Narino; ICN 5224 S from
San Pedro de la Sierra, Magdalena; ICN 5938 S Pance, valle del Cauca; ICN 5225 $ San Pedro de la Sierra,
Magdalena; ICN 14531 $ from Taminango, Narino; and A g. peruana ICN 12368 S from Quindio.

Diagnosis. —The new form can be distinguished
from other species in the genus by the following char¬
acteristics: small skull (GLS < 24.5 mm); presence of
complete fragile zygomatic arches (sometimes broken);
small canines; P4 massive with a triangular base; para-
cone poorly developed on the first upper molar; large,
wide, and squared upper molars, with a well-developed
hypoconal basin; and rostrum wide in its base at the
area between M3 and the P4, which is particularly
noticeable from a dorsal view (Fig. 6). Externally,
A. carishina can be distinguished from other species
in the A. geoffroyi complex by relatively short lateral
vibrissae (VL < 7.0 mm); reduced thumb (TL < 5.24
mm); and absence of tail.

Description.—Anoura carishina is among the
smallest representative of the A. geoffroyi complex,
best indicated by cranial measurements, including GLS,
CBL, and PAL (Table 2) and its shorter thumb size (TL
< 5.24 mm). General coloration is cinnamon-brown
(Ridgway 1912) resembling that of A. g. geoffroyi.

The dorsal and ventral pelage typically is long and
bicolored. Individual hairs are basally white for two
thirds of the length of the hair and cinnamon-brown
(Ridgway 1912) on their distal edge. Ears and mem¬
branes average paler than in A. g. geoffroyi. Although
all specimens included in this work correspond to dry
skins likely to have modifications in some of the ex¬
ternal characters, mystacial vibrissae in A. carishina
appeared to be shorter than in other representatives
of the A. geoffroyi complex, and the uroptagium was
clearly more developed in the holotype and specimens
in the type series of A. carishina when compared with
A. g. peruana specimens (UW >3.5). Uropatagium of
A. carishina is hairy on both ventral and dorsal surfaces,
with a well-developed whorl of hairs on the edge.

Skull, mandible, and teeth are similar to those of
A. geoffroyi , but the rostrum is shorter and the braincase
is more globoid, particularly in males; the zygomata is
complete, but thin and sometimes broken; the canines
are small, not projected laterally, and are almost parallel

Mantilla-Meluk and Baker—New Species of Anoura from Colombia

13

Figure 6. Differences in size and development of the cusps of the second upper premolar (P4) between
A. carishina and A. geoffroyi. Praconid (PaC); Protoconid (PrC); Hypoconid (HC); Entoconid (EC).
Lingual side on the left of the drawing.

to the tooth row; the posterolateral edges of the palate
are pointed; the P3 and P4 are relatively wide in com¬
parison to those in A. geoffroyi , A. lasiopyga, and A.
g. peruana ; the new species has a reduced, thick, and
shorter anterobasal cusp on the P4, while the medial
internal cusp of the P4 is enlarged, rendering the basal
outline of the tooth to be approximately triangular; the
molars are thick and squared, with a wide hypoconal
basin; the posterior portion of the rostrum is wide; and
the maxillae are slightly projected laterally, making
them visible from a dorsal view.

Type locality. —The holotype was collected in
Taminango, corregimiento de Remolino, department

of Narino. This area of Taminango is part of the xe-
rophitic enclave of the Patia River, where local arid
conditions are the results of a rain-shadow effect cre¬
ated by the transversal position of the Patia depression.
The Patia region is located in southwestern Colombia
between the Western and Central Cordilleras (Fig. 7),
in the upper Patia River basin, Department of Narino,
Colombia (1°34'23"N, 77°16'59"W). The average
annual temperature is 25.2°C, and the average relative
humidity is 79%. The annual average precipitation
is 809.3 mm distributed in a bimodal-tetraseasonal
pattern. Rainfall is greatest between March and May,
and between October and December. In this zone,
the shrubs Lippia origanoides, Lantana canescens,

14

Occasional Papers, Museum of Texas Tech University

COLOMBIA

VENEZUELA

COLOMBIA

ECUADOR

77H6’W

Taminago, Narino

1 P 34'N

^ ‘ij

ECUADOR

Figure 7. Type locality of Anoura carishina in Taminago, Narino, Colombia (1°34'23" N, 77T6'59" W)
at 1,465 m. Inset; geographic location of type series (site 1 designates holotype locality).

and Senna pallida , and the cacti Stenocereus griseus,
Pilosocereus sp., and Opuntia dillenii are abundant.
Pollen of S. griseus, Pilosocereus sp., and O. dillenii
was found on the fur of two nectarivorous bats captured
in the area (A. carishina and Choeroniscus godmani ).
Other bat species found locally included Artibeus
jamaicensis, Carollia perspicillata, Phyllostomus dis¬
color, P. hastatus, Micronycteris megalotis, Desmodus
rotundus, Sturnira erythromos, S. lilium, Enchistenes
hartii , and Chiroderma sahini.

Distribution.—Anoura carishina is widely dis¬
tributed in Colombia from the department of Narino
on the border with Ecuador to the piedmonts of the
Sierra Nevada de Santa Marta in the department of
Magdalena in the Colombian Caribbean. There is no
evidence of A. carishina on the eastern versant of the
Eastern Cordillera in the Colombian Andes. Based on
the wide variety of ecosystems included in the distribu¬
tion of A. carishina , it is probable that this taxon may
extend its distribution along the Andes southward into
Ecuador (Fig. 7).

Mantilla-Meluk and Baker—New Species of A no dr a from Colombia

15

Anoura caudifer Anoura geoffroyi

II

1 mm

Figure 8. Electron microphotograph of first upper molars (Ml) (lingual view) of A. caudifer complex (A.
aequatoris, A. cadenai, A. caudifer , and A.fistulata) (left) and A. geoffroyi sensu lato (A. g. geoffroyi, A. g.
lasiopyga , and A. g. peruana ) (right). We followed the nomenclature system for dental homologies on Ml
proposed by Hershkovitz (1977): eocone or paracone (1); protocone (2); metacone (4); parastyle (a);metastyle
(b); epiconule or protoconule (c); precentrocrista (I); paracrista (II); preprotocrista (III); postcentrocrista
(IV); postprotocrista (entocrista) (V); hypoconal basin (Hb). Images obtained by B. S track, FMNH.

Comparisons.—Anoura carishina is sympatric
at sampling localities with three other recognized taxa
in Anoura (A. cadenai, A. cultrata, and A. g. peruana).
The new species is easily distinguished from A. cade¬
nai by the triangular shape of P4 (in contrast with the
laterally compressed P4 of A. cadenai ) and the absence
of a well-developed paracone (eocone) on Ml, which
is always well-developed in all species within the A.
caudifer complex (A. aequatoris, A. cadenai, A. caudi¬
fer, A. fistulata, and A. luismanueli ) (Fig. 8). Anoura
carishina lacks the robust and knife-shaped canines
as well as the blade-shaped lower first premolar of A.
cultrata. Anoura carishina also is found near localities
where A. g. peruana is present in the elevations of the
Sierra Nevada de Santa Marta, although the two taxa
occur at different altitudes. Anoura carishina is eas¬
ily distinguished from A. g. peruana by its paler color
(Ridgway 1912), smaller size (Table 2), shallow notch
on the posterior edge of the palatal bones (sometimes
erroneously called fossa mesopterigoydea), smaller
thumb, shorter lateral vibrissae, and relatively large

uropatagium; these characteristics along with the com¬
plete zygomata and the longer hair, also differentiate
A. carishina from A. lasiopyga. Traits separating A.
carishina from A. lasiopyga also apply to distinguish
it from A. latidens.

Etymology.—Anoura carishina takes its name
from the Quichua word “carishina” which means
“woman who resembles a man” (Cordero 2003). Car¬
ishina in the Inca culture is a male character who wears
female clothes as a costume in festivities. We consider
the Quichua word carishina appropriate to highlight
the following significant aspects of the new species: 1)
its geographic origin - the A. carishina holotype was
collected in the Andes of southern Colombia originally
inhabited by ethnic groups that speak Quichua; and
2) its smaller size with respect to other species within
the A. geoffroyi complex and lower degree of sexual
dimorphism. We chose this name as a tribute to the
senior author’s father who has dedicated his life to the
study of the Inca culture, and to our colleagues who

16

Occasional Papers, Museum of Texas Tech University

have been dedicating their efforts to the understanding
of the Andean mammalian chiropterofauna, particularly
Alfred L. Gardner, Bruce D. Patterson, Victor Pacheco,
Rene M. Fonseca, Luis Albuja, Luis Fernando Aguirre,
Paul Velazco, and Sergio Solari.

This work would not be possible without the
many efforts of collectors who generously have en¬

riched scientific collections through the years, pro¬
viding in this way the evidence that feeds the always
healthy scientific discussion. Through our work we
would like to encourage the community of scientists
interested in this fascinating group of organisms to
validate the museological work by contributing with
specimens that would be a valuable asset to present and
coming generations of mammalogists.

Acknowledgments

We thank Y. Munoz-Saba, and the staff of the
ICN; A. Rodriguez-Bolanos and the staff of the MUD;
M. Rodriguez-Posada, M. Ruedi and the MHNG for
access to the images of G. lasiopyga holotype; the
AMNH for making available images of the holotype
of G. apolinari ,; H. E. Ramirez-Chaves for valuable
comments on the taxonomy of the group; and H. Gamer
and the staff of the NSRL Museum of TTU museum.
This work would not have been possible without the
generous cooperation of B. D. Patterson, L. R. Heaney,

W. B. Stanley, B. Starck, and the staff of the Field
Museum of Natural History. We thank all of them for
their support and assistance. Financial support was
provided by the Thomas J. Dee and Karl P. Schmidt
Funds of the Field Museum’s Scholarship Committee,
the Texas Tech University Horn Professors Scholarship
Award, and the Department of Biological Sciences. We
thank A. Daugherty, R. Marchan, L. Siles, M. Pinto,
C. Gomes de Sotero Caio, and L. Bradley for valuable
editorial assistance.

Literature Cited

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35:83-87.

Anderson, S. 1957. New records of the bat, Anoura geoffroyi
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Addresses of authors:

Hugo Mantilla-Meluk

Department of Biological Sciences
Texas Tech University
Lubbock, TX, 79409
hugo. mantilla@ttu. edu

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Robert J. Baker

Department of Biological Sciences
Texas Tech University
Lubbock, TX, 79409
rjbaker@ttu.edu

18

Occasional Papers, Museum of Texas Tech University

Appendix

List of specimens examined. Acronyms are described in the text.

Anoura carishina (n = 5): COLOMBIA: Magdalena,Pedro de La Sierra, Finca Tierra Grata, ICN 5224 S
and ICN 5225 ?; Narino, Taminango, ICN 14530 S and 14531 ?; and Valle del Cauca, Pance ICN 5938 S-

Anourageoffroyi apolinari (n = 1): COLOMBIA: Cundinamarca: Boqueron de San Francisco, near Bogota
AMNH 37376 ?.

Anoura geoffroyi geoffroyi (n = 52): BRAZIL, Ceara FMNH 116852-54 ?; COLOMBIA: Caqueta: Rio
Mesay, Puerto Abeja, (Vorizal - Tepuy), southeastern, Serrania de Chiribiquete, ICN 14538 ?, ICN 14539-40 S',
Parque Natural Nacional Chiribiquete, Rio Cunare, Raudal El Tubo, southeastern, Serrania de Chiribiquete, ICN
14711 c?; Parque Natural Nacional Chiribiquete western side Serrania Norte, ICN 12769 S', Meta: Acacias, Vereda
San Jose, ICN 9700-01 S', Restrepo, Caney Alto, ICN 8060 S, ICN 8061 ?, ICN 8062 S, ICN 8063 ?, ICN 10110
S, ICN 10111-13 ?; Restrepo, Surroundings CREAD, Cano Seco, ICN 10932 S', La Macarena, souther portion
of the Serrania de La Macarena, Cano Cristales, ICN 7727-28 S', Restrepo, Caney Alto, Villavicencio, Pozo Azul
Cave, ICN 9750 S', TRINIDAD: Blanchisseuer, TTU 5463 ?, TTU 5370 S, TTU 5403 S', Maracas Valley, TTU
5363 S', San Rafael, TTU 5384 S, TTU 5823 S, TTU 5826 S, 37665-66 S', Saint Andrew, TTU 5464 $, TTU
8977 ?, TTU 8979 S, TTU 26463-65 ?, TTU 26466 S, TTU 26467-68 ?, TTU 26469-70 S, TTU 26471-72 ?,
TTU 26473-76 S, TTU 26788 ?; Saint George, TTU 26780 ?, TTU 26787 S, TTU 37664 ?.

Anoura geoffroyi lasiopyga. (n = 36): Color phase a) MEXICO: Hidalgo, TTU 15465-68 S, TTU 15470
S, TTU 15471 ?, TTU 15472 cJ,TTU 15473 ?, TTU 15474 S, TTU 15476 ?, TTU 15477-81 S', Veracruz,
TTU 10029 ?, FMNH 34162 ?; MHNG 515.88 S', Color phase b) COSTA RICA: Punta Arenas, Finca Las
Cruces County, MSB 26789-90 ?; GUATEMALA: Santa Elena, FMNH 41654 S, FMNH 41832-33 ?, FMNH
41834-35 $, FMNH 41837-38 S', Jalapa, FMNH 73362 ?; MEXICO: Hidalgo, TTU 36704 $, TTU 38035 ?;
Tamulipas, TTU 7345-46 S, TTU 7347-48 ?; San Luis Potosi TTU 9931 ?; Tehuetlan, TTU 5469 S', Tlaxcala,
TTU 25342 ?.

Anoura peruana (n = 172): BOLIVIA: La Paz, 0.5 km Rio Zongo MSB 70293 S', San Matias, FMNH
39294 S', COLOMBIA: Antioquia: ICN, 8753 S', Jardin, Vereda La Linda, Quebrada La Linda, ICN 16454 S',
San Luis Corregimiento El Prodigio, Vereda Las Confusas, Quebrada Corinto, Las Confusas Cave, ICN 13316
?, ICN 13317 S', ICN 16502 ?; Urrao, Paramo de Frontino, FMNH 72191 S', FMNH 72192-96 ?; Venecia,
Santa Catalina Caves; Boyaca: ICN 1704 ?, ICN 5314 ?; ICN 5315 S', ICN 5420 S', Caldas: MHNUC, 450-
51 S, MHNUC 460 S, MHNUC 462; Manizalez, Alto del Laurel, La Navarra, ICN 16730 ?, MNHUC 452 ?;
Cauca: Inza, Vereda Tierras Blancas, school surroundings, ICN 8439 ?, ICN 8440 S', El Retiro, Reserva Rio
Blanco, ICN 15157 S', Cundinamarca: ICN 8048 S, ICN 12971 S', MUD, 162-65 S, MUD 170-72 S, MUD
177 S, MUD 184 S, MUD 189 S', ICN 5274 ?, ICN, 17694 ?; Cabrera, Granja Experimental Agropecuaria,
ICN 9552 S', Cabrera, Vereda Penas Blancas, El Infiernito Cave, ICN12629 S, ICN 12630-31 9. ICN 12632 $,
ICN 12633 ?, ICN 12634 S', Choachi, Vereda Ferralara, Pena Fonte, ICN 8854 9; La Calera, Los Patios, ICN
8850-51 9; Tena, Laguna de Pedro Palo, ICN 5494 9, ICN 8367 S', Villapinzon, headwaters Bogota River, ICN
17675 ?* ICN 17677-78 S', Ubala, Exit to Bogota, ICN 15100 ?; Zipaquira, Paramo de Guerrero, FMNH 72204
S, FMNH 72206-07 S, FMNH 72208-13 9; Huila: Las Cuevas Parque, Indian Cave, 160m from exit, FMNH
58678 ?, FMNH 58680-81 ?; FMNH 58679 S', Magdalena: Serrania San Lorenzo, Inderena Station, ICN 5353
?, Serrania San Lorenzo, Inderena Station ICN 5354-55 S', Serrania San Lorenzo, Hacienda La Victoria ICN
5356 S', Serrania San Lorenzo, Hacienda La Victoria ICN 5357-60 ?; Narino: La Victoria FMNH 113482 S',
Llorente, FMNH 113449 ?, FMNH 113489 ?, FMNH 113490 S, FMNH 113491 ?; FMNH 223492 ?; FMNH
114029 S', Quindio: Salento; Natural Reserve Canon Quindio, frente de reforestacion La Montana, ICN 12368

Mantilla-Meluk and Baker—New Species of Anoura from Colombia

19

Appendix (cont.)

c?, 12369-70 9 , ICN 12371-73 g, ICN 12374 9 , ICN 12375-78 g\ Natural Reserve Canon Quindio, Rente de
reforestacion La Romelia, ICN 12380-83 $, ICN 12385-87 g; Natural Reserve Canon Quindio, frente de refo-
restacion La Picota, ICN 12388-90 5; Risaralda: ICN 12538 g; Santuario, Verada, Los planes, ICN 11791 5;
Santuario, Verada, El Campamento, ICN 11833-34 5; Santander: MUD, 641-43 g; Charala, Vereda El Salitre,
finca El Mirador, Aida Cave, ICN 17498 5, ICN 17499 g; Encino, Vereda Rio Negro, sitio Las Tapias, finca El
Aserradero, ICN 17515 9, ICN 17517 g, ICN 17518 9, ICN 17519 g; Ocamonte, Finca Macanal, La Virgen
Cave, ICN 17632 $; Ocamonte, Vereda Miraflorez, finca Macanal, La Virgen Cave, ICN 17633 g, ICN 17634
9 ; Tolima: Km. 140 carretera Libano-Murillo, Hacienda Casas Viejas, ICN 8380 g; Herveo, Vereda Albania,
ICN 3470 9 ; Icononzo, El Tigre Caves, ICN 8812 9 , ICN 8681 g, ICN 8682-83 $; ECUADOR: Chimborazo,
Pallatngo, USNM 513439 g, USNM 513440 9: Guayas, Balao, Guertas Negras, USNM 498832 g, USNM
534301 g; Pichincha, Zapadores, USNM 513437-38 g; PERU: Amazonas: Bongara, Road Utcubamba, between
Churuja y Pedro Ruiz, FMNH 128643 g, FMNH 128645 g; Chachapoyas, Balsas, 19 km by road E, FMNH
128639 g; Apurimac: Abancay, Bosque Ampay, FMNH 110922 9, FMNH 110923 g; Andahuaylas, FMNH
75178 9 , FMNH 75179-80 g, FMNH 75159-62 9 , FMNH 75163-64 g. Hacienda La Laguna, FMNH 75157-58
g, FMNH 75177 g; Ayacucho: La Mar, Ocros, Hacienda Pajonal, FMNH 75165 9 , FMNH 75166-69 g, FMNH
75170-71 9, FMNH 75181 9; Huanuco: Tingo Maria Las Lechuzas Cave, MSB 49924-28 g; Junin, Cueva de
Guarapo, ca. 35 km Tarma, MSB 53085 9 , San Ramon, USNM 507176 g, USNM 507177 9 ; Ollantaytambo,
USNM 195127 g; UNDETERMINED LOCALITY: ICN 18052 9 , ICN 18053 g, ICN 18054

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Library of Congress Cataloging-in-Publication Data

Occasional Papers of the Museum of Texas Tech University, Number 292
Series Editor: Robert J. Baker

New species of Anoura (Chiroptera: Phyllostomidae) from Colombia, with systematic remarks and
notes on the distribution of the A. geoffroyi complex

Hugo Mantilla-Meluk and Robert J. Baker

ISSN 0149-175X

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