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A new species of Eumops (Chiroptera:

Molossidae) from southwestern Peru

ARTICLE in ZOOTAXA OCTOBER 2014

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 Zootaxa 3878 (1): 019036 ISSN 1175-5326 (print edition)
www.mapress.com /zootaxa /
Copyright 2014 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3878.1.2
http://zoobank.org/urn:lsid:zoobank.org:pub:FDE7F7A4-7DCC-4155-8D96-A0539229DBFE

A new species of Eumops (Chiroptera: Molossidae) from southwestern Peru

CSAR E. MEDINA1, RENATO GREGORIN2, HORACIO ZEBALLOS1,3,

HUGO T. ZAMORA1 & LIGIANE M. MORAS4
1
Museo de Historia Natural de la Universidad Nacional San Agustn (MUSA). Av. Alcides Carrin s/n. Arequipa, Per.
E-mail: cmedinap1234@yahoo.com
2
Departamento de Biologia, Universidade Federal de Lavras, Lavras, Minas Gerais, CEP 37200-000, Brazil.
E-mail: rgregorin@dbi.ufla.br
3
Instituto de Ciencias de la Naturaleza, Territorio y Energas Renovables, Pontificia Universidad Catlica del Per, Av. Universitaria
1801, San Miguel, Lima 32, Per. E-mail: horaciozeballos@gmail.com
4
Departamento de Zoologia, Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil.
E-mail: ligimoras@yahoo.com.br

Abstract

The genus Eumops is the most diverse genera of molossid bats in the Neotropics. In Peru this genus is widely distributed
and represented by nine species: E. auripendulus, E. delticus, E. hansae, E. maurus, E. nanus, E. patagonicus, E. perotis,
E. trumbulli, and E. wilsoni. After several years of mammalian diversity surveys in the coastal desert and western slopes
of southwestern Peru, a specimen of Eumops was collected whose unique set of traits allows us to assert that deserves to
be described as a new species. Based on molecular and morphological evidence, the new species is related to medium-
large sized species (i.e. E. glaucinus, E. auripendulus, and E. perotis). Cytochrome b genetic divergence between the new
species and the other species of the genus was high (> 12%) and it is consistent with morphological divergence presented
for this new species. This new species, endemic to Peru, increases the diversity of Eumops to 16 species.

Key words: Bonneted bat, coastal desert, endemism, Molossidae, phylogeny, Peru

Introduction

The Neotropical genus Eumops Miller, 1906, belongs to the nearly cosmopolitan bat family Molossidae (Eger
2008). The combined geographic ranges of the species of the genus extend from southern United States to
Patagonia in Argentina (Eger 1977). Eumops occurs in a great variety of habitats from sea level up to 3000 m (Eger
1977; Eisenberg & Redford 1999; Eger 2008). Currently Eumops includes 15 species: E. auripendulus (Shaw,
1800); E. bonariensis (Peters, 1874); E. dabbenei Thomas, 1914; E. delticus Thomas, 1923; E. ferox (Gundlach,
1861); E. floridanus Allen, 1932; E. glaucinus (Wagner, 1843); E. hansae Sanborn, 1932; E. maurus (Thomas,
1901a); E. nanus (Miller, 1900); E. patagonicus Thomas, 1924; E. perotis (Schinz, 1821); E. trumbulli (Thomas,
1901b); E. underwoodi Goodwin, 1940; and E. wilsoni Baker et al., 2009.
Since the description of Eumops in the early 1900s (Miller 1906), two major revisions were focused on its
taxonomy (Sanborn 1932; Eger 1977) and several studies analyzed relationships within the genus based on
phenetic analyses of similarity (Eger 1977), biochemical approaches (Dolan & Honeycutt 1978), and cladistic
analyses based on morphological and molecular data (Gregorin 2009; Bartlett et al. 2013).
Nine species of Eumops occur in Peru (E. auripendulus, E. delticus, E. hansae, E. maurus, E. nanus, E.
patagonicus, E. perotis, E. trumbulli, and E. wilsoni). They are widely distributed throughout the country, including
northwestern dry forest, coastal desert, mountain forest, Amazonian forest and savannah portions (Eger 1977; Eger
2008; Baker et al. 2009; Pacheco et al. 2009; Daz 2011; Medina et al. 2012). In 2010, after four years of assessing
the mammalian diversity of the coastal desert and western slopes of southwestern Peru, a specimen of Eumops was
collected in the Department of Moquegua whose unique set of traits warrants it recognition as a new species.

Accepted by P. Velazco: 7 Oct. 2014; published: 22 Oct. 2014 19

Herein we describe this specimen as a new species. We also analyze genetic, morphological, and morphometric
data in order to assess the phylogenetic relationships of this new species with the other species of the genus.

Material and methods

Skull morphology nomenclature follows Freeman (1981), Hand (1990), Giannini et al. (2006), and Giannini &
Simmons (2007). External measurements were recorded from the original field tags. Wing measurements were
taken following the definitions and illustration of Freeman (1981), whereas cranial measurements followed
Bernardi et al. (2009). Measurements were taken with digital calipers to the nearest 0.1 mm as follows: fourth
metacarpal length (LIV), first phalanx of fourth digit length (fal1-IV), greatest length of skull (GLS), braincase
breadth (BB), condyle-incisive length (CIL), zygomatic breadth (ZB), postorbital breadth (PB), palate length (PL),
maxillary toothrow length (MTRL), breadth across upper molars (M-M), width across upper canines (C-C),
mandible length (DENL), and mandibular toothrow length (MANDL).
We examined a total of 357 specimens from the following institutional collections: American Museum of
Natural History, New York, USA (AMNH); Louisiana State University, Baton Rouge, USA (LSU); Museum of
Comparative Zoology, Harvard University, Cambridge, USA (MCZ); Museum of Vertebrate Zoology, Berkeley,
USA (MVZ); Museu de Zoologia da Universidade de So Paulo, So Paulo, Brazil (MZUSP); Museu de Histria
Natural, Universidade Estadual de Campinas, Campinas, Brazil (ZUEC); Museo de Historia Natural, Universidad
Mayor de San Marcos, Lima, Peru (MUSM); Museo de Historia Natural, Universidad Nacional de San Agustn,
Arequipa, Peru (MUSA); National Museum of Natural History, Washington, USA (USNM); Royal Ontario
Museum, Toronto, Canada (ROM); Texas Cooperative Wildlife Research Collection, Texas A&M University,
College Station, USA (TCWC); The Field Museum, Chicago, USA (FMNH); Universidade do Estado de So
Paulo, So Jos do Rio Preto, Brazil (DZSJRP); Universidade Federal do Lavras, Lavras, Brazil (UFLA);
Universidade Federal Rural do Rio de Janeiro, Seropdica, Brazil (ALP); and, University of Michigan, Museum of
Zoology, Ann Harbor, USA (UMMZ). The list of examined specimens, their geographic locations, museum
numbers, and acronyms of the institutions where they are housed are presented in Appendix I.
Molecular analyses. We obtained the entire mitochondrial Cytochrome-b (Cytb) gene of five bats of the genus
Eumops, including the new species and two individuals of E. bonariensis which has not been included in previous
Eumops phylogenies, and two of Molossus molossus (Pallas, 1766) that was used as part of the outgroup.
Additionally, we gathered 28 sequences from GenBank, 27 sequences of Eumops and one of Tadarida brasiliensis
(I. Geoffroy St.-Hilaire, 1824) (outgroup), including 13 of the 15 recognized species of Eumops (Appendix II). As
the sequences downloaded from GenBank were partial segments (~700 bp), we used the same partial segment (688
bp) for all sequences. The sequences produced in this study were deposited on GenBank under accession numbers:
KJ152625KJ152631 (Appendix II).
DNA extractions were performed with DNeasy extraction kits (Qiagen). PCR amplifications were carried
out in 15 L reactions containing 4060 ng of DNA, 0.3U of Platinum Taq (Invitrogen), 1x Platinum Taq PCR
buffer, 2.5 M MgCl2, 200 M dNTPs set (Invitrogen) and 0.3 M of each primer. The primers used for
amplifications were L14121 and H15318 (Redondo et al. 2008). We used the following cycling scheme for PCR: 5
min at 94C followed by 35 cycles of 30 s at 94C, 40 s at 50 C for primer annealing and 85 s at 72C for
extension, and a final 10 min extension at 72C after the last cycle. PCR products were purified using a solution of
20% Polyethylene-glycol 8000 in 2.5 M of NaCl using a protocol described by Sambrook et al. (2001).
Sequencing of both strands were carried out in a ABI 3130 (Applied Biosystems) automated sequencer using
Big Dye Terminator Cycle Sequencing methodology (Applied Biosystems) and we employed the same primers
used in PCR amplifications, besides two additional internal primers: MVZ4 (Kocher et al. 1989) and L14881
(Redondo et al. 2008). Sequences were assembled and checked for quality using DNA Baser Sequence Assembler
v3.x 2011.
Sequences were aligned using the Muscle algorithm (Edgar 2004) implemented in MEGA 5.05 (Tamura et al.
2011). MEGA 5.05 was also used to test if the new species sequence was not a pseudogene and to calculate the
genetic distance as intraspecific and interspecific sequence divergence with a Kimura 2-parameter (K2P) model
(Kimura 1980).

20 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

 MrModeltest 2.2 (Nylander 2004) was used to select the best fit model of sequence evolution using the Akaike
Information Criterion (AIC; Akaike 1974).
A Bayesian Inference analysis was conducted in MrBayes 3.2 (Huelsenbeck & Ronquist 2001). The Akaike
Information Criterion indicated that the optimal model was the General Time Reversible model incorporating
invariant sites and a gamma distribution (GTR+I+G). Four (one cold and three heated) simultaneous Markov
chains were run for 20 million generations, trees were sampled every 200 generations, with the rst 25% of the
generations were discarded as burn-in.
A Maximum Likelihood (ML) analysis was performed using RAxML 7.2.8 in the CIPRES Gateway (Miller et
al. 2010; Stamatakis 2006). This was implemented using the GTR+I+G model (Stamatakis 2006). Bootstrap
resampling was used to assess support for tree nodes and was set to stop automatically in RaxML. (RaxML
options: -s infile -n result -p 12345 -m GTRGAMMAI -x 12345 -N 1000 -k -f a o <identified outgroup>).
Morphological cladistic analysis. We used 38 of the 39 morphological characters used by Gregorin (2009).
We reanalyzed Gregorins (2009) data and made some modifications due to the inclusion of the new taxon and
more specimens of other taxa in order to describe the variation and better define the characters and their states
(Appendix III). Thus, we excluded character 23 (medial sagittal crest), because we were unable to accurately define
the states with the inclusion of more specimens. We also modified character 4 (size of the ears) considering three
states rather than two, and we morphometrically defined them based on the ratio between total length of ears and
mean length of forearm, therefore the ranges for the three states were: small (ratio between 0.26 to 0.38), medium-
sized (0.42 to 0.46), and large (> 0.49). Similarly, we delimited three states in character 10 based on the mean of the
length of forearm for each taxon: small (3448 mm), medium (5262 mm), and large (> 65 mm). We also modified
characters 12 and 31 considering them to include two states rather than three. The matrix of characters is presented
in Table 1.

TABLE 1. Morphological matrix used in the phylogenetic analysis. Question marks ? denote missing data and dashes
denote characters that are not applicable to that particular taxon.
Taxon\character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Molossus molossus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 0
Tadarida brasiliensis 1 0 1 0 0 1 0 0 2 0 0 2 0 1 0 0 1 1 0 1
Eumops. a. 1 1 1 0 0 1 1 2 0 0 1 0 1 2 1 1 1 1 3 0
auripendulus
E. a. major 1 1 1 0 0 1 1 2 0 0 1 0 1 2 1 1 1 1 3 0
E. bonariensis 1 1 1 1 0 1 1 1 1 1 1 2 1 2 1 1 1 1 0 1
E. dabbenei 1 1 1 0 0 0 1 2 0 2 1 2 1 2 1 1 1 1 2 0
E. delticus 1 1 1 0 0 1 1 1 1 1 1 2 1 2 1 1 1 1 0 1
E. glaucinus 1 1 1 1 0 0 1 0 0 0 1 2 1 2 1 1 1 1 2 0
E. hansae 2 1 1 1 2 2 1 1 1 1 1 2 1 2 1 1 1 1 0 1
E. maurus 1 1 1 1 0 0 1 2 0 0 1 1 ? 2 1 1 1 1 3 0
E. nanus 1 1 1 1 0 1 1 1 1 1 1 2 1 2 1 1 1 1 0 1
E. patagonicus 1 1 1 1 0 1 1 1 1 1 1 2 1 2 1 1 1 1 0 1
E. p. californicus 1 1 1 2 0 0 1 0 0 2 1 2 1 2 1 1 1 1 2 0
E. p. perotis 1 1 1 2 0 0 1 0 0 2 1 2 1 2 1 1 1 1 2 0
E. sp. nov. 1 1 1 1 0 0 1 0 0 1 1 2 1 2 1 1 1 1 1 0
E. trumbulli 1 1 1 2 0 0 1 0 0 2 1 2 1 2 1 1 1 1 1 0
E. u. sonoriensis 1 1 1 1 0 0 1 2 0 2 1 2 1 2 1 1 1 1 2 0
E. u. underwoodi 1 1 1 1 0 0 1 2 0 2 1 2 1 2 1 1 1 1 2 0

......continued on the next page

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 21
TABLE 1. (Continued)
Taxon\character 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
Molossus molossus 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 ? 0 0
Tadarida brasiliensis 1 1 0 0 1 1 0 0 0 0 1 1 1 0 ? 0 0
Eumops. a. 1 0 0 0 0 1 0 0 1 2 1 1 1 1 1 1 1 0
auripendulus
E. a. major 1 0 0 0 0 1 0 0 1 2 1 1 1 1 ? ? ? 0
E. bonariensis 1 0 0 0 1 1 1 0 0 0 1 1 1 1 1 0 1 0
E. dabbenei 1 1 1 0 0 1 0 1 1 1 1 1 1 0 ? ? ? ?
E. delticus 1 0 0 0 1 1 1 0 0 0 1 1 1 1 ? ? ? ?
E. glaucinus 1 1 0 0 0 1 0 1 1 1 1 1 1 0 1 1 1 0
E. hansae 1 0 0 0 1 2 1 0 0 0 1 1 1 1 ? ? ? ?
E. maurus 1 0 0 0 0 0 0 1 0 2 1 1 1 0 ? ? ? ?
E. nanus 1 0 0 0 1 1 1 0 0 0 1 1 1 1 ? ? ? ?
E. patagonicus 1 0 0 0 1 1 1 0 0 0 1 1 1 1 ? ? ? 0
E. p. californicus 2 1 0 1 1 2 1 0 1 2 1 1 1 1 ? ? ? ?
E. p. perotis 2 1 0 1 1 2 1 0 1 2 1 1 1 1 1 1 1 1
E. sp. nov. 2 1 0 1 1 1 1 0 1 1 1 1 1 1 ? ? ? ?
E. trumbulli 2 1 0 1 1 2 1 0 1 2 1 1 1 1 1 1 1 1
E. u. sonoriensis 1 1 1 0 0 1 0 1 1 1 1 1 1 0 ? ? ? ?
E. u. underwoodi 1 1 1 0 0 1 0 1 1 1 1 1 1 0 ? ? ? 0

Cladistic analysis methods, such as character analysis, homology criterion, and outgroup methodology follows
that of Gregorin (2009). We performed the analysis considering all multistate characters as unordered.
Phylogenetic analysis (maximum parsimony) was performed using TNT version 1.1 (Goloboff et al. 2008) for
Windows. Heurist searches to find the most parsimonious tree(s) were performed using tree bisection-reconnection
(TBR) branch-swapping method with 100 replicates. A strict consensus was calculated because more than one
most parsimonious tree were obtained. A bootstrap analysis was performed, with 1,000 replications, to evaluate the
support of the resulting clades.
Morphometric analysis. Principal component analysis (PCA) was carried out using a variance-covariance
matrix for 14 external and cranial measurements of the new species and female specimens of five medium to large
sized species of Eumops that are closely related to the new species based on the phylogenetic analyses. All the
measurements were log-transformed to achieve normalization for the analysis. The first three components were
retained and the loadings of all variables were evaluated as a measure of importance in this preliminary separation.
Statistical differences between groups were assessed using a discriminant function analysis (DFA) (Brown &
Wicker 2000) and cross validation was performed using the option leave out in PAST. The analyses were carried
out using the software PAST 3.0 for Windows (Hammer 2013).

Results

Molecular analyses. Bayesian inference and maximum-likelihood analyses recovered similar topologies, with
some strongly supported clades (Fig. 1). The new species was recovered sister to E. auripendulus and E. maurus,
but this clade was weakly supported (Fig. 1). The average Cytb pairwise distance among the species of Eumops
included in this study ranged from 1.9% (E. feroxE. floridanus) to 23.1% (E. hansaeE. underwoodi). The
pairwise distance between the new species and the other species of Eumops ranged from 12.3% (E. perotis) to

22 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

19.2% (E. hansae) (Table 2). The new species is distinguished by 64 fixed nucleotides differences when compared
with E. auripendulus and E. maurus (Table 3).

FIGURE 1. Phylogeny of Eumops. Left, Bayesian phylogeny with Molossus molossus and Tadarida brasiliensis as outgroups
(number above branches are Bayesian posterior probabilities and those below are maximum-likelihood bootstrap values).
Right, strict consensus of three most parsimonious trees calculated from a heuristic search of 18 taxa and 38 discrete
morphological characters (number above branches indicate bootstrap values).

FIGURE 2. Plots summarizing the PCA (left) and DFA (right) of adult females of Eumops (n=201) extracted from 14
measurements. Eumops sp. nov. (star), E. wilsoni (empty circles), E. glaucinus (filled circles), E. auripendulus (cross), E.
trumbulli (triangles) and E. perotis (squares).

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 23
TABLE 2. Average Kimura 2-parameter distances (in percentages) between species of Eumops based on 688 base pairs
of the cytb. Intraspecific divergence is on the diagonal. Outgroup taxa was not included.
n 1 2 3 4 5 6 7 8 9 10 11 12 13 14
1 Eumops 5 0.9
auripendulus
2 E. bonariensis 2 16.8 0.5
3 E. dabbenei 1 14.0 18.0
4 E. ferox 4 14.6 15.6 11.1 0.7
5 E. floridanus 2 16.7 17.6 12.9 1.9 0.9
6 E. glaucinus 2 15.2 17.5 11.7 4.6 5.7 1.9
7 E. hansae 3 20.3 19.3 22.2 20.9 22.1 20.6 0.9
8 E. maurus 1 2.7 16.9 14.8 14.5 16.7 15.7 19.6
9 E. nanus 2 18.3 6.0 18.2 17.3 19.0 17.2 20.9 19.1 1.1
10 E. patagonicus 2 18.0 3.9 17.9 7.0 15.9 17.7 19.6 17.9 7.0 0.6
11 E. perotis 2 13.9 17.0 12.8 12.8 14.0 14.6 18.6 13.5 17.6 17.3 0.2
12 E. sp. nov. 1 13.6 18.6 14.6 14.6 16.7 16.1 19.2 13.2 18.1 18.6 12.3
13 E. underwoodi 2 16.5 18.8 6.4 11.2 13.2 13.0 23.1 15.3 20.0 18.8 12.4 16.3 1.5
14 E. wilsoni 2 14.3 14.3 14.4 7.3 8.3 9.0 19.3 14.3 16.6 15.8 10.5 13.5 12.3 0.2

TABLE 3. Fixed nucleotide changes in cytb among Eumops sp. nov., E. auripendulus (N = 5) and E. maurus (N = 1). A
= adenine, C = cytosine, G = guanine, and T = thymine. Position refers to the nucleotide position in the cytb gene from
the complete mitochondrial genome of Artibeus jamaicensis (Pumo et al. 1998).
Position Eumops sp. nov. E. auripendulus/ E. maurus
14257 C A
14258 T C
14266 C T
14275 C T
14311 C T
14323 T C
14353 C T
14365 T C
14377 T A
14393 T C
14404 T C
14413 C T
14416 G A
14422 C T
14429 C T
14441 A G
14461 C T
14464 A C
14467 C A
14476 C T
14501 C A
14503 A T
......continued on the next page

24 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

TABLE 3. (Continued)
Position Eumops sp. nov. E. auripendulus/ E. maurus
14524 T C
14542 T C
14545 A C
14548 G A
14569 T C
14608 T C
14614 T C
14620 T C
14623 C T
14624 G A
14641 C T
14662 T C
14674 A C
14683 T C
14699 C T
14707 C A
14710 T C
14711 A G
14719 T C
14728 T C
14749 A T
14752 T C
14767 C T
14770 T C
14773 C T
14782 C T
14785 G A
14821 T C
14837 T C
14839 A T
14845 G C/ T
14855 C A
14859 C G
14861 A G
14862 T C
14877 C T
14890 G T
14908 C A
14911 C T
14914 T C
14917 T C
14920 C T

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 25
 Morphological cladistic analysis. Our morphological analysis resulted in three most parsimonious trees of 69
steps (Fig. 1). Eumops monophyly was weakly supported (67% of bootstrap). Although poorly resolved, the
consensus tree is fully resolved in deeper portions of the tree.
The new species is sister to the clade including E. perotis + E. trumbulli supported with a high bootstrap value
(97%) (Fig. 1). This clade presents two synapomorphies: characters 21, state 2 (long rostrum) and 24, state 1
(narrow post-orbital constriction); these characters contribute to the odd shape of the skull shared by the three
species. The clade comprised by E. perotis + E. trumbulli is supported by two synapormophies that distinguished
them from the new species: character 4, state 2 (very large ears) and character 10, state 2 (large body sized).
Morphometric analysis. The first three principal components accounted for 97.2% of the total variation
(Table 4). PC1 accounted for the highest percentage (93.3%), and this vector exhibit only positive loadings,
suggesting that it is size factor (with high loadings for C-C, MTRL, and CIL). Correspondingly, the small-bodies
species (Eumops auripendulus, E. glaucinus, E. wilsoni, and the new species) have lower scores along this axis,
medium-sized E. trumbulli have intermediate values, and large-bodied E. perotis presents high scores. Along PC2
the new species overlaps with E. auripendulus, E. trumbulli, and E. perotis, reflecting shape similarities among
these species (Fig. 2; Table 4).

TABLE 4. Factor loadings of characters for PCA and DFA extracted from the character variance-covariance matrix of
medium-large sized Eumops (Eumops sp. nov., E. wilsoni, E. glaucinus, E. auripendulus, E. trumbulli, and E. perotis).
Principal component Discriminant function
Characters PC1 PC2 PC3 Axis 1 Axis 2 Axis 3
GLS 0.275 -0.118 -0.050 0.0063 -0.0024 -0.0001
CIL 0.296 0.086 0.080 0.0068 0.0001 -0.0006
PL 0.293 -0.190 -0.180 0.0066 -0.0036 -0.0015
ZB 0.250 0.142 -0.014 0.0057 0.0002 -0.0034
PB 0.130 0.637 0.346 0.0030 0.0047 -0.0058
BB 0.201 0.229 0.158 0.0047 0.0014 0.0015
C-C 0.313 -0.056 -0.539 0.0070 -0.0034 -0.0076
MTRL 0.302 0.148 -0.074 0.0069 0.0003 -0.0037
M-M 0.238 0.159 -0.272 0.0054 -0.0004 -0.0075
DENL 0.277 -0.003 -0.106 0.0063 -0.0015 -0.0041
MANDL 0.284 0.120 -0.169 0.0065 -0.0001 -0.0044
FA 0.291 -0.068 0.368 0.0067 -0.0007 0.0032
LIV 0.271 -0.056 0.394 0.0063 -0.0003 0.0016
fal1-IV 0.264 -0.626 0.337 0.0059 -0.0055 0.0012
Eigenvalue 0.0259 0.0007 0.0003 49.389 7.129 0.687
Variance (in %) 93.31 2.64 1.30 85.53 12.35 1.19

The DFA analysis showed that 85.5% and 12.3% of the total variation is expressed by the first and second
canonical axes, respectively (Table 4). The bivariate plot of the first and second discriminant function axes shows a
good separation among all the species, except Eumops glaucinus and E. wilsoni that exhibit an extensive overlap
(Fig. 2). The discriminant factor matrix indicates that C-C, MTRL, and CIL have the highest positive correlation on
the first discriminant function axis, whereas PB and BB have the highest positive loadings on the second
discriminant function axis (Table 4). There was some extensive overlap between Eumops glaucinus and E. wilsoni,
but no overlap among the other species, as illustrated by an 87.8% (E. glaucinus), 88.8% (E. wilsoni), 98.6% (E.
auripendulus), and 100% (E. perotis, E. trumbulli, and new species) post hoc classification of the original cases
indicated (Fig. 2).

26 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

Systematics

Family Molossidae Gervais, 1856

Genus Eumops Miller, 1906

Eumops chiribaya, sp. nov.

Chiribaya's Bonneted Bat
Figure 4

Holotype. Dried skin, skull, and partial skeleton of an adult female (MUSA 8493), deposited at the Museo de
Historia Natural de la Universidad Nacional de San Agustn (MUSA). The specimen was captured on 1 June 2010
by a local person and was taken to the MUSA, where it was received and prepared by Hugo T. Zamora (original
number HZM 670). Tissues are deposited at MUSA. The specimen is in good condition, except by the broken
lower left canine.
Type locality. El Algarrobal, 111 m; 8.4 km NE of Ilo; Ilo Province; Moquegua Department; Peru, 1737'21.6"
S, 7116'06.6" W (Fig. 3).

FIGURE 3. Map of southwestern Peru illustrating the type locality of Eumops chiribaya (star).

Etymology. The specific epithet chiribaya is given in honor of the Chiribaya culture (9001,350 A.D.).
Chiribayans were settled in the basin of the Ilo River, and expanded north to the Tambo valley (Arequipa) and the
south to the Azapa valley (Chile), including the high altitude regions, up to nearly 3,000 m of elevation.
Distribution. Eumops chiribaya is known only from the type locality (Fig. 3).

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 27
 Diagnosis. Eumops chiribaya is distinguished from all other species of the genus by the following combination
of characters: medium body size; relatively large ears (25.7 mm); dorsal coloration light olive-brown, with slightly
lighter belly; dorsal hairs bicolored, with white base that conspicuously contrast with the tip; square-shaped tragus;
cranium with elongated rostrum; narrow post-orbital constriction; tiny incisive foramen; basisphenoid pits
rectangular, deep and separated by a narrow septum; opisthocranion curved dorsally and anteriorly; mastoid
foramen present; premetacrista equal to half the length of postparacrista on M3; mandible gracile with a well-
developed keel (Fig. 4).

FIGURE 4. Dorsal, ventral, and lateral views of the skulls and lateral view of the mandibles of five species of genus Eumops
from Peru, from left to right: E. chiribaya (MUSA 8493), E. auripendulus (MUSA 5995), E. wilsoni (holotype, TTU 103281),
E. trumbulli (MUSM 243), and E. perotis (MUSM 6049). Scale bar = 10 mm.

Description. Eumops chiribaya is a medium size species (Table 5). Dorsal coloration is light olive-brown,
with a lighter belly. Pelage is soft, long (6 mm taken at the level of the shoulder region) with bicolored dorsal hairs
(basal third of each hair white in color). Face naked; upper lip smooth to weakly wrinkled on distal edge of snout.
Nose blackish; upper border of narial region surrounded by small and pointed warts, internarial ribs covered by
small, spoon-like and blackish hairs. Ears large (25.7 mm), round, blackish and joined on the forehead; upper
border smooth, but the inner face with brown hairs distributed from the dermal keel to the top of the ear. Tragus
square-shaped. Antitragus semicircular, wider than higher. Dorsal surface of propatagium and plagiopatagium near
to forearm and dactilopatagium at basal middle of fifth metacarpal covered with short light olive-brown hairs. Feet
blackish; external edges and ventral surface of the first and fifth toes of the feet covered by short hairs, silver in
color, with claws covered by longer hairs of the same color. Calcar longer than feet. Uropatagium dull brown with
free portion of the tail blackish.
Skull with elongated and narrow rostrum in dorsal view, and straight in profile (Fig. 4). Braincase rounded,
tending to be angular anteriorly; opisthocranion curved dorsally and anteriorly. Nasal process of the premaxilla
well-developed. Postorbital constriction narrow. Medial sagittal crest poorly developed. Palate long and slightly
wide. Medial process of posterior margin of palate bridge absent. Incisive foramina tiny, with two concentric cusps
longitudinally. Mesopterygoid fossa broad and rectangular. Foramen ovale well-developed. Sphenorbital fissure
narrow. Basisphenoid pits large, deep, rectangle-shaped, and separated from each other by a narrow septum.
Mastoid foramen present. Premetacrista equal to half the length of postparacrista on M3. Mandible gracile with
well-developed keel.
Measurements of the holotype. The external measurements (mm) and weight (g) are: total length 134; tail
length 52; hindfoot length 13.5; ear length 25.7; forearm length 61.1; and weight 20.3. Measures used in the
morphometric analysis are indicated in table 5.
Comparisons. Eumops chiribaya is a medium-sized member of Eumops (Table 5) phylogenetically related to
medium-large sized species and morphologically more similar to E. perotis and E. trumbulli (Fig. 1). The smaller

28 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

size distinguishes E. chiribaya from the large species E. perotis and E. trumbulli, also E. dabbenei and E.
underwoodi (FA greater than 68 mm; Eger 1977).
Eumops bonariensis, E. patagonicus, E. nanus, and E. hansae (here, E. bonariensis clade) are comparatively
small (FA less than 49 mm; Table 1, Eger 1977) and differentiated in morphology and molecular data (Fig. 1). E.
chiribaya is also distinguished from E. bonariensis and its relatives by smooth upper lips (finely wrinkled in E.
bonariensis clade), blackish pelage (light or grayish brown in E. bonariensis clade), long rostrum (shorter in E.
bonariensis clade), occipital region in lateral view is mildly curved (straight in E. bonariensis clade complex and
E. wilsoniFig. 4), and reduced premetacrista on M3 (long, N-shapped in E. bonariensis clade).

TABLE 5. Comparative measurements (mm) of the holotype of Eumops chiribaya sp nov., and females of selected
species of genus Eumops. Numbers include mean SD and ranges in brackets.
Eumops E. wilsoni E. glaucinus E. auripendulus E. trumbulli E. perotis
chiribaya (N = 18) (N = 66) (N = 74) (N = 11) (N = 31)
GLS 23.36 23.76 0.27 24.40 0.42 24.68 0.53 28.62 0.44 32.02 0.65
(23.2824.27) (22.8825.54) (23.5125.94) (27.7729.18) (30.4433.01)
CIL 22.51 22.69 0.27 23.37 0.40 22.81 0.48 27.29 0.41 30.78 0.59
(22.2623.30) (22.2724.27) (21.7323.91) (26.5427.84) (29.5231.93)
PL 9.38 10.49 0.22 10.71 0.26 10.98 0.26 12.94 0.31 14.36 0.40
(10.1510.97) (10.0111.56) (10.4511.67) (12.5813.38) (13.7915.55)
ZB 13.23 14.02 0.21 14.52 0.31 14.13 0.36 16.29 0.28 18.22 0.46
(13.5714.27) (13.8115.21) (13.3414.89) (15.8216.64) (17.2518.90)
PB 4.55 4.87 0.13 4.94 0.17 4.57 0.18 4.99 0.14 5.41 0.20
(4.635.19) (4.585.49) (4.255.08) (4.785.24) (5.025.96)
BB 11.59 10.96 0.20 11.05 0.34 10.71 0.29 12.25 0.28 13.27 0.31
(10.6011.30) (10.2911.8) (10.0411.40) (11.7312.67) (12.5913.73)
C-C 4.93 5.82 0.24 5.92 0.17 6.04 0.21 6.76 0.24 8.14 0.33
(5.376.31) (5.566.38) (5.626.54) (6.297.04) (7.378.70)
MTRL 8.55 9.39 0.22 9.60 0.20 9.33 0.19 11.13 0.22 12.72 0.23
(8.909.75) (9.2010.04) (8.939.79) (10.8211.40) (12.3013.06)
M-M 8.40 9.82 0.24 10.03 0.29 9.85 0.25 10.98 0.19 12.54 0.30
(9.3910.20) (9.4610.73) (9.3010.60) (10.7111.28) (11.8613.03)
DENL 15.92 17.71 0.33 18.38 0.37 18.26 0.45 21.21 0.43 23.92 0.48
(17.1018.44) (17.2219.01) (16.8419.19) (20.6421.79) (23.1424.78)
MANDL 9.41 10.23 0.25 10.49 0.25 10.25 0.28 11.88 0.28 13.69 0.29
(9.7410.66) (9.9811.09) (9.7310.92) (11.4012.30) (13.0614.21)
FA 61.1 57.40 1.12 58.60 1.60 58.04 1.51 69.85 0.77 77.50 2.04
(55.3059.48) (50.4061.32) (54.9162.34) (68.3870.92) (72.0281.09)
LIV 58.02 56.90 1.14 58.58 1.47 57.60 1.65 68.56 0.68 75.51 1.97
(54.9459.20) (54.1662.14) (52.8162.59) (67.2869.46) (71.6579.84)
fal1-IV 20.66 19.22 0.75 20.53 0.75 21.50 0.89 24.08 0.63 26.96 1.02
(18.0320.59) (18.1522.24) (19.6023.30) (23.0425.34) (25.3029.03)

From the species that occur in Peru, Eumops chiribaya has similar external and cranial dimensions to E.
wilsoni and E. auripendulus (Table 5). Eumops auripendulus (and also E. maurus) is easily distinguished from E.
chiribaya because it has a blackish pelage (cocoa brown in E. maurus), pointed tragus, different shape of skull, in
particular, extended posterior portion of basicranium, and reduced premetacrista on M3 (V-shaped) (Fig. 4).
Eumops wilsoni (and also E. glaucinus) is similar to E. chiribaya (i.e. light brown dorsal pelage, square-shaped
tragus, and premetacrista on M3 slightly longer), but the shape of skull (shorter rostrum and wider post-orbital
constriction) and ears relatively smaller (21.8 mm in E. wilsoni, 17-23 mm in E. glaucinus) distinguish both species
from the E. chiribaya (Fig. 5). Very deep and wide basisphenoid pits also distinguish E. chiribaya from E.
auripendulus, E. glaucinus, and E. wilsoni (Fig. 4).

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 29
 Eumops chiribaya differs from E. perotis and E. trumbulli based on the following combination of
characteristics: smaller size (E. chiribaya has a medium body size and forearm shorter than in E. trumbulli and E.
perotis - FA = 61, 65, and 79 mm, respectively); smaller ears, with ratio of length of ears/length of forearm
reaching 0.42 in comparison with E. trumbulli and E. perotis, which have very developed ears (> to 0.49);
basisphenoid pits are proportionally deeper in E. chiribaya than E. trumbulli and E. perotis; and its opisthocranion
is curved dorsally and anteriorly whereas it is extended posteriorly in E. trumbulli and E. perotis (Fig. 4).
Natural history. Eumops chiribaya inhabits in Coastal desert of southern Peru. The holotype was captured
alive by a police officer during his patrolling in the locality El Algarrobal, Moquegua, Peru. He mentioned that the
bat struck the top of a gate (approximately 5 meters high) and fell on the road, allowing him to catch the specimen
with his own hands.
The holotype had no ectoparasites nor showed signs of reproductive activity. The stomach content was not
analyzed.

Discussion

Phylogenetic relationships. The molecular and morphological data supported similar results: 1) Eumops splits in
two main clades (one clade composed by E. bonariensis and its relatives, small species, and other including the
medium and large-sized species) (Fig. 1), 2) E. auripendulus and E. maurus grouping together and both are sister to
the remaining medium-large species, and 3) E. glaucinus is sister to the clade composed by E. dabbenei and E.
underwoodi, and this clade in turn is sister to E. perotis + E. trumbulli. The affinities of Eumops chiribaya within
its clade appear poorly supported in the Cytb tree (Fig. 1). Based on the Cytb genetic distance values, the most
similar species to E. chiribaya is E. perotis with 12% of Cytb divergence (Table 2), a greater value than those
typically found within mammal species (Siles et al. 2013); however this result is very interesting because both
species belong to different clades: E. chiribaya + (E. maurus + E. auripendulus) and E. perotis + (E. glaucinus
species complex + (E. dabbenei + E. underwoodi)). Thus, the combination of molecular, morphological, and
morphometric data support the recognition of E. chiribaya as a new species.
The molecular phylogenetic relationships found among the species of Eumops is similar to those recovered by
Bartlett et al. (2013), where E. perotis is related with medium-large sized species rather than smaller ones (E.
hansae, E. bonariensis, and relatives) as proposed by Gregorin (2009). Although, E. chiribaya is a full species, its
phylogenetic position is controversial within the medium-large sized species. However, there are notable
similarities to E. perotis in the relative size and morphology of the ears, the color pattern, cranial morphology (long
rostrum, narrow post-orbital constriction, and mildly curved occipital region). Also it shares similarities on the M3
morphology with E. trumbulli a relative to E. perotis.
Our morphological phylogeny results are quite distinct from those of Gregorins (2009) and we considered
them more robust because after a reassessment of some morphological characters and addition of other species in
the resulting topology is overall congruent with our cytb tree (Fig. 1) and with the recent molecular phylogeny of
Bartlett et al. (2013) for Eumops that was based on two mitochondrial and one nuclear genes. Two main differences
between our analysis and that of Gregorin (2009) are: 1) the clade including E. perotis + E. trumbulli is closer to
the medium-large sized species groupings E. glaucinus + (E. underwoodi + E. dabbenei), instead Gregorin (2009)
recovered that clade closed to the small sized species (E. bonariensis species complex + E. hansae); and 2) the E.
bonariensis species complex + E. hansae comprise an isolated clade and sister-group of all remaining species of
Eumops. The composition of these basal clades was recovered in others studies (Ammerman et al. 2012; Barlett et
al. 2013) with E. hansae placed as a basal species of the genus (Barlett et al. 2013), in agreement with previously
propositions based on biochemical (Dolan & Honeycutt 1978) and phenetical approaches (Eger 1977).
Conservation. Eumops is a New World molosid genus whose majorities of species were described between
1800s to 1940s (Sanborn 1932; Eger 1977). After seven decades, two new species were described, one from the
lowlands of western Ecuador and Peru (Baker et al. 2009) and the one described in this article. Eumops now
includes 16 species, 12 of them occurring in South America, becoming the most specious genus of Molossidae in
the Neotropic. Moreover, E. chiribaya represents the first bat described from the coastal desert and western slopes
of southwestern Peru.
The diversity of bats in the coastal desert of southwestern Peru is high, and includes insectivorous (i.e.

30 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

Molossus molossus, Promops davisoni, Tadarida brasiliensis, Histiotus montanus, among others), nectarivorous
(Platalina genovensium), and frugivorous species (Sturnira bogotensis), some of which are endemic and threatened
(Tomopeas ravus, Amorphochilus schnablii, and P. genovensium) (Zeballos et al. 2001; Pacheco et al. 2009).
Since November 2010, new expeditions have been carried out to El Algarrobal attempting to capture more
individuals of E. chiribaya, without success. However, five individuals of Kalinowski's free tail bat (Mormopterus
kalinowskii) were captured in the mouth of Osmore River (7.7 km E type locality of E. chiribaya). Despite nearly
2,000 net-hours with 100 individuals caught in almost a decade of fieldwork in southwestern Peru (Tacna,
Moquegua, and Arequipa), not a single individual of E. chiribaya was captured. This rarity may be explained
because mostly Neotropical molossids forage far above net range and, therefore, are underestimated in inventories
that are based solely on mist-net capture (Jung et al. 2014) or by its natural rarity, reduced populations, and
restricted distribution (Arita 1993).
El Algarrobal is located in a narrow valley surrounded by steep hills (slope > 60) and has notable human
activity resulting from changes in land use, such as intensive agriculture of Olive trees (Olea europaea), urban
expansion, mining activities and other anthropogenic factors. The sparse native vegetation is limited to river banks
and the annual average temperature and precipitation are 18.3 C and 74.4 mm, respectively (INRENA 1994).
Furthermore the nearest Protected Area is 80 km NW of El Agarrobal (Lagunas de Mejia National Sanctuary).
The continuous loss of habitat and restricted distribution of E. chiribaya warrants its placement under some
category of protection, however with only one specimen it is impossible to assess population stability and so it
must be considered Data Deficient until more long-term data is obtained (International Union for Conservation of
Nature and Natural Resources 2011). Because of the threats facing this species we consider necessary the
implementation of conservation actions (i.e., identified and protection of roosts, educate local people through
workshops on insectivorous bats, and short courses on the importance of bats in the ecosystem).

Acknowledgements

We thank the curators of the museums we visited or who loaned us material, and for the use of their facilities to
study specimens: Judith Chupasko (MCZ), Mario de Vivo (MZUSP), Judith Eger (ROM), Mark S. Hafner (LSU),
Paulina Jenkins (BMNH), Emygdio Leite (ZUEC), Jessica E. Light (TCWC), Evaristo Lpez (MUSA), Philip
Myers (UMMZ), Vctor Pacheco (MUSM), Bruce D. Patterson (FMNH), James Patton (MVZ), Adriano L.
Peracchi (ALP), Nancy B. Simmons (AMNH), Valdir Taddei (DZSJRP), and Don Wilson (USNM). We are
especially grateful to Pal M. Velazco who carefully read an earlier version of this manuscript and gave insightful
comments for its improvement. Thanks to Don Wilson for the pertinent suggestions and improvements to the
English. This study was partially supported by Fundao de Amparo Pesquisa do Estado de Minas Gerais
(FAPEMIG) and Conselho Nacional de Desenvolvimento Cientfico e Tecnolgico (CNPq).

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Thomas, O. (1901b) On a collection of mammals from the Kanuku Mountains, British Guiana. Annals Magazine of Natural
History, 8, 139154. [series 7]
http://dx.doi.org/10.1080/03745480109442900
Thomas, O. (1914) New Callicebus and Eumops from S. America. Annals Magazine of Natural History, 13, 480481. [series 8]
http://dx.doi.org/10.1080/00222931408693514
Thomas, O. (1923) Two new mammals from Maraj Island. Annals Magazine of Natural History, 12, 341342. [series 9]
http://dx.doi.org/10.1080/00222932308632954
Thomas, O. (1924) New South American small mammals. Annals Magazine of Natural History, 13, 234237. [series 9]
http://dx.doi.org/10.1080/00222932408633035
Wagner, J.A. (1843) Diagnosen neuer Arten brasilischer Handflgler. Archiv fur Naturgeschichte, 9, 365368.
Zeballos, H., Pacheco, V. & Baraybar, E. (2001) Diversidad y conservacin de los mamferos de Arequipa, Per. Revista
Peruana de Biologa, 8, 94104.

APPENDIX I

The following list includes all specimens examined in this study, with their respective localities. Specimens used in the
morphometric analysis are indicated with asterisk (*).

Eumops auripendulus (92)BOLIVIA: Beni, Rurrenabaque: AMNH 248209*, 24821113*, 248215*, 248220*, 248221,
248222*, 24822324, 248225*, 248226, 248227*, 248228, 248229*, 248230. BRAZIL: Acre, Plcido de Castro: MZUSP
728187, 729093*, 7297*, 7302*; MACN 16574*; Tarauac: MZUSP 8245*; Faro, Rio Amazonas: AMNH 93852*,
9385556*; Amazonas, Humaita: DZSJRP 13357*, 13358*, 14194*; Iroanga, Tapajs: FMNH 140794*. COLOMBIA:
Meta, Villavicencio: AMNH 149256*, 183321*, ROM 45530*; Guaicarama: AMNH 74359*, MCZ 27862*; Noros:
USNM 281251*, 281252*. ECUADOR: Manabi, Rio Mongoya: FMNH 5351113. GUYANA: Barima-Waini, Arakaka:
AMNH 207327*, 20732930*, 207332*, 207334*, 20733637*, 207340*. FRENCH GUIANA: Cayenne, Sinnamary,
Paracou: AMNH 267537*. PANAMA: Boca del Toro, Changuinola: USNM 31578788*; Bohio, Canal Zone: USNM
17820709*; Boim: MZUSP 565960*; Fort Kobbe, Canal Zone: USNM 317628*, 312120*; Gamboa, Canal Zone: MVZ

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 33
 118820*; Summit, Canal Zone: USNM 317631*; Tapia, Panama: MCZ 33612*. PERU: Junn, Colonia del Peren: USNM
15554849*, 155552*; San Martn, Guaybamba: AMNH 1012527; Hunuco, Hacienta San Antonio, Rio Chincha:
FMNH 2493435, Vista Alegre: FMNH 24936, 24937*, Pachitea - Agua Caliente: MUSA 5995. SURIMANE: Sipaliwini,
Alalapadoo, near Rio Sipaliwini: ROM 39497*. VENEZUELA: Carabobo, 10 km NW Urama: USNM 37416567*;
Aragua, El Dorado, Ro Cuyuni: AMNH 3064041*, 30643*, 3064648*, 30650*; Amazonas, San Carlos de Rio Negro:
USNM 56064244*.
Eumops bonariensis (21).BRAZIL: Rio Grande do Sul, Quinta: AMNH 235405-18. URUGUAY: Ro Negro, Arroyo Negro:
AMNH 20565357, 20566061.
Eumops chiribaya new species (1).PERU: Moquegua, Ilo: MUSA 8493*.
Eumops dabbenei (4).PARAGUAY: Presidente Hayes, Villa Hayes: UMMZ 133763-64. VENEZUELA: FMNH 108310;
Carabobo, Rio Uacauy: USNM 374169.
Eumops delticus (2).BRAZIL: Amazonas, Humait: DZSJRP 12770, Itacoatiara: MZUSP 15221.
Eumops glaucinus (69).BRAZIL: Amazonas, Itaboca: FMNH 140795*; Par, Faro: AMNH 9385860, Pinhy: MCZ
30634*, Tamary: MCZ 30653*; Paran, Maring: DZSJRP 1046869*, 1047273*, 10509*, 10512*, 10515*; So Paulo,
So Jos do Rio Preto: DZSJRP 2714*, 15077*, 15672, 15674, 15691, 16313, 16529, 16536, Bilac: DZSJRP 10910*,
10911*, 10912*, Fazenda Cajueiro: DZSJRP 1104347*, Regente Feij: DZSJRP 10908*, 10909*. COLOMBIA: Bolivar,
Sincelejo: TCWC 12445*; Boyaca, El Yopal: ROM 45287*; Honda: AMNH 69185*; Meta, Puerto Lopez: ROM 62544*,
62547*. GUYANA: Essequibo, rio Achimere: ROM 4282930*, 42832*, 42834*, Rio Aroquoi: ROM 5782325*,
5782728*. PANAMA: Chiriqui, Progreso: USNM 363142*. PARAGUAY: Presidente Hayes, 275 km NW Villa Hayes:
MVZ 145134*. VENEZUELA: Yaracuy, Anoa, rio Yaracuy: USNM 44186263*; Carabobo, Mont Alban: USNM
441868*; Lara, Rio Tocuyu: AMNH 130700*, 130702*; Amazonas, San Juan: USNM 40958485*, 409587*, 409591*,
40959495*, 409605*, 409607*, 409614*, 40961617*, 40961922*, 409627*, 409630*.
Eumops hansae (6).FRENCH GUIANA: Cayenne, Paracou, Sinnamary: AMNH 267538. GUYANA: Alto Demerara-
Berbice, Ituni: ROM 73014; Upper Takutu-Upper Essequibo, Maruranau, Ishi Wau: ROM 35517; Alto Takutu-Alto
Esequibo, Rupununi, Machawira: ROM 35636. PANAMA: Darien, Tarcacuna: USNM 310278. VENEZUELA: Bolvar,
29 km El Dorado: USNM 387799.
Eumops maurus (2)ECUADOR: Napo, Pompeya Sur: ROM 106326. GUYANA: Kanaku: BMNH 1.6.4.34.
Eumops nanus (13).COLOMBIA: Guajira, 32 km W. Maracaibo: USNM 44184855. PANAMA: Chiriqu, Boquern:
USNM 104892, 33197172. VENEZUELA: Falcn, Capatarida: USNM 441856, 495016.
Eumops perotis (75).ARGENTINA: Jujuy, Yuto: AMNH 185208, Ledesma: ROM 53959*; Tucuman, Concepcin: FMNH
23637*, 73738; Chaco, General Vedia: ROM 5020102, 53960; Salta, Campo Santo (General Guemes): ROM 50203,
5026769*, 50198, 50199*, 53958*, Capital: ROM 50200; Santiago del Estero, Nueva Esperanza: ROM 5395354*.
BOLIVIA: Beni, San Joaqun: FMNH 116690, 116699*, 11670203*, 116704, 116705*, 116707, 116709, 11671112,
11671618, 116719*, 116720, 116724*. BRAZIL: Rio de Janeiro, Seropdica: ALP 32, 127, 45758, 1267, 1763, 1764*,
2482, 3084*, ZUEC 276; Braslia, Universidade de Braslia (UnB): DZSJRP 10018*; Minas Gerais, Lagoa Santa: FMNH
20976, 20984; Lavras: UFLA 1048, 4039*, 4057*, 4080, 4100, Rio das Velhas: MCZ 4163; Maranho, Barra do Corda:
FMNH 23953*, 23954, 23955*, 23956; So Paulo, So Jos do Rio Preto: DZSJRP 11673, 13588, 1663133*, Cedral:
DZSJRP 16465*, Ribeiro Preto: DZSJRP 14027, Montemor: MZUSP 15645, Campinas: ZUEC 1566*, 1567, 1568*,
Atibaia, Parque Florestal Itapetinga: ZUEC 1919, 1920. PARAGUAY: Chaco, 170km W. de Bahia Negra: AMNH
248391*; Cordillera, Juan de Mena: USNM 552739*. PERU: Lima, Iglesia Inmaculada: MUSM 245, 246, 6049.
VENEZUELA: Anzotegui, Cantaura: USNM 296600*.
Eumops trumbulli (42).BOLIVIA: Beni, Guayaramerin: AMNH 209900-02. BRAZIL: Par, Rio Tocantins, Mocajuba:
AMNH 97016, 97018, 97020, 9702122*, Camet: MCZ 30608*, Rio Xingu, Villaringo do Monte: AMNH 96022,
97328; Amazonas, Rio Juru, Eirunep: FMNH 140796. COLOMBIA: Meta, Puerto Lopez: ROM 45291*, 4529395*,
6072, 62491, 62512, 62546; Amazonas, Leticia: ROM 62492*, 6249394, 62515*, 62516. GUYANA: Alto Takatu - Alto
Essequibo, ros Takatu-Achimerewau: ROM 35712, ros Takatu-Rewa: ROM 38456, 16 km N. of Dadanawa: ROM
40595, Dadanawa: ROM 48787, 61900. PERU: Ucayali, Ro Ucayali, Yarinacocha: FMNH 62112, 62114*, 62115;
MUSM 24344, 24748; Loreto, Ro Amazonas, Boca de ro Peruate: FMNH 8901920*, 8902122, 89165.
Eumops underwoodi (8).MEXICO: Sonora, Rancho Costa Rica: MVZ 8215153, 10 mi NW Noche Buena: MVZ 8531114.
UNITED STATES: Arizona, Sasabe, Pima Co.: USNM 31469697.
Eumops wilsoni (22).ECUADOR: Guayaquil, 19 km N. of Bridge over Rio Catatumbo: USNM 154492. PERU:
Lambayeque, Olmos: LSU 2131112*, 21313, 21314*, 2121921*, 2721315*, 27217*, 27218*, 27221, 27225*, 20km.
Olmos: MVZ 139991*, Cerro la Vieja: LSU 27224*; Piura, Suyo: TCWC 12456*, 12460*, 1246263*, 12464.

34 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.

APPENDIX II. Locality, catalog/tissue numbers, and GenBank accession number for the sequences used in this study.
Taxon Locality Catalog no./tissue no. GenBank accession no.
(Cyt-b)
Tadarida brasiliensis - - JF489126
Molossus molossus Brazil, Para, UFMG3191/VCT386 KJ152630
Molossus molossus Brazil, Para, UFMG3331/VCT956 KJ152631
Eumops auripendulus Brazil, Para, VCT3279/VCT3279 KJ152626
Eumops auripendulus Costa Rica, Alajuela TTU29815/TK12554 JQ731822
Eumops auripendulus Jamaica, St. Ann Parish CM46610/TK925931 JQ731824
Eumops auripendulus Ecuador, Napo ROM105526/F37759 JQ731823
Eumops auripendulus Brazil, Para, VCT3219/VCT3219 KJ152625
Eumops bonariensis Brazil, Sao Paulo, R641/R641 KJ152627
Eumops bonariensis Brazil, Sao Paulo, R6447/R6447 KJ152628
Eumops chiribaya Peru, Moquegua MUSA8493/MUSA8493 KJ152629
Eumops dabbenei Venezuela, Guarico TTU33406/TK15063 JQ731806
Eumops ferox Mexico, Yucatan TTU47519/TK13585 EU350036
Eumops ferox Mexico, Yucatan TTU47521/TK13589 EU350040
Eumops ferox Cuba, Guantanamo TTU52613/TK32033 EU350013
Eumops ferox Cuba, Guantanamo TTU52617/TK32052 EU350015
Eumops floridanus United States, Florida KU163656/RMT4610 EU350024
Eumops floridanus United States, Florida KU163657/RMT4611 EU350025
Eumops glaucinus Paraguay, Concepcin TTU80255/TK64163 EU350005
Eumops glaucinus Paraguay, Presidente Hayes TTU80542 /TK64926 EU35004
Eumops hansae Ecuador, Napo ROM105642/F37887 JQ731815
Eumops hansae Guyana, Potaro-siparuni ROM108361/F43341 JQ731814
Eumops hansae Venezuela, Miranda TTU33409/TK15296 JQ731813
Eumops maurus Ecuador, Napo ROM106326/TK40481 JQ731821
Eumops nanus Mexico, Yucatan TTU47516/TK13557 JQ731829
Eumops nanus Mexico, Yucatan TTU47517/TK13558 JQ731826
Eumops patagonicus Paraguay, Presidente Hayes TTU62499/TK34937 JQ731833
Eumops patagonicus Paraguay, Neembucu TTU80582/TK64367 JQ731830
Eumops perotis United States, Texas ASNHC12238/ASK5379 EU349990
Eumops perotis United States, Texas ASNHC13295/ASK6271 EU349991
Eumops underwoodi Mexico, Chiapas TTU29322/TK12144 JQ731808
Eumops underwoodi Nicaragua, Boaco TTU29311/TK12366 EU349989
Eumops wilsoni Ecuador, Guayas TTU103278/TK134816 JQ731804
Eumops wilsoni Ecuador, Guayas TTU103466/TK134989 JQ731827

NEW EUMOPS FROM PERU Zootaxa 3878 (1) 2014 Magnolia Press 35
APPENDIX III. Morphological characters and states used in the analysis.

C1. Upper border of narial region: without any structure (smooth) (0); surrounded by small and pointed warts (1); covered by
small hairs (2).
C2. Internarial ribs: smooth (0); covered by small hairs (1).
C3. Shape of ears: triangular (0); rounded (1).
C4. Size of ears: small (0); large (1); very large (2).
C5. Level of insertion of the ears at forehead: slightly separated (0); joined in a common point (1); united by a membrane (2).
C6. Upper border of ears: smooth (0), with pointed and fleshy warts (1), with pointed and rounded warts (2).
C7. Long and semicircular antitragus: absent (0), present (1).
C8. Shape of the tragus: squarish (0); slightly squared with the upper edge directed in a posterior position (1); pointed (2).
C9. Upper lips: smooth (0); slightly wrinkled (1); deeply wrinkled (2).
C10. Size of the body: small (0), medium (1), large (2).
C11. Ratio between second phalanx of fourth finger and total length of the body: large (> 0.1; 0.130.2) (0), small (< 0.1;
0.050.08) (1).
C12. Dorsal pelage coloration: dark brown to blackish (0); lighter brown to grayish (1).
C13.Frequency of fungiform papillae on the dorsal and lateral regions of the tongue: abundant (0), reduced (1).
C14. Relative position of upper incisors: convergent (0), parallel (1), divergent (2).
C15. Shape of incisors: flattened (0), conical (1).
C16. Protofossa on the upper incisors: absent (0), present (1).
C17. Number of upper premolars: one (0), two (1).
C18. Metaconule on M1 and M2: low (0), high (1).
C19. M3 premetacrista length: very long (0), medially developed (1), reduced (2), vestigial or absent (3).
C20. Trigonid on M3: reduced (0), developed (1).
C21. Length of rostrum: short (0), medium (1), long (2).
C22. Outline of mid-dorsal region of skull: curved (0), flattened (1).
C23. Width of rostrum: narrow (0); wide (1).
C24. Width of post-orbital region: enlarged (0); narrowed (1).
C25. Shape of basisphenoid pits: slightly oval (0); squarish (1).
C26. Depth of basisphenoid pits: shallow (0); deep (1); very deep (2).
C27. Rib between basisphenoid pits: wide (0); narrowed (1).
C28. Lateral wall of braincase in dorsal view: curved (0); enlarged posteriorward (1).
C29. Shape of occipital in posterior view: domed (0); straight (1).
C30. Opisthocranion in lateral view: straight (0); curved dorsally and anteriorly (1), extended posteriorly (2).
C31. Bar crossing the optic canal: absent (0); present (1).
C32. Basicochlear fissure: narrow (0); wide (1).
C33. Number of incisive foramina: two (0); one (1).
C34. Development of mandible: massive (0); gracile (1).
C35. Glans penis: covered with spines (0); spineless (1).
C36. Shape of glans penis: oval with three basal lobes (0); oval but without three-lobed structure (1).
C37. Length of glans penis: long (0); short (1).
C38. Os penis: present (0); absent (1).

36 Zootaxa 3878 (1) 2014 Magnolia Press MEDINA ET AL.