^

Occasional Papers

Museum of Texas Tech University ^

Number 217

31 October 2002

Caroffia brevicauda

A New Central American Species

FROM THE CaROLLIA BREVICAUDA COMPLEX

Editor’s Comment: This paper describes a previously unrecognized species
in the genus Carollia. Although it is probable that morphological differences
distinguish this new species from the remainder of the genus, it is clear that
this species would not have been recognized based on morphology alone.
Methods developed since 1986 allow easy access to DNA sequence data from
native species permit scientists to employ data other than morphological to
document biological uniqueness and species boundaries. The use of the
mitochondrial cytochrome-i» gene to indicate biological species in mammals
was recently reviewed by Robert Bradley and Robert Baker (Journal of
Mammalogy, vol. 82:960-973). If their conclusions are correct, there will be
numero us currently unrecognized species that will be detected through studies
of DNA sequence and application of the genetic species concept. The total
may be 25% more than the current list in Wilson and Reeder, 1992 (Mammals
Species of the World, Smithsonian Institution Press, Washington,
DC). Clearly, there is a substantial need for detailed studies of the
implications of genetic variation to species boundaries in natural populations
of mammals. One important implication to the hypothesis that many
biological species remain unrecognized is that museum collections, especially
those that archive material for DNA analysis, are not sufficient to resolve
many of these systematic and biodiversity issues. Additional museum
collections that save a maximum amount of material for genetic studies will
be needed. Collections of mammal specimens for this purpose should be
archived (voucher specimens and tissues) in accredited collections that are
funded for perpetuity. Storage of samples in individual laboratories greatly
increases the likelihood that these specimens will be unavailable for research
by others to better understand the diversity of life on Earth.

RJB

Front cover: Color representation of the cytochrome-^ branding pattern of the Carollia brevicauda complex.
Specimens are: Carollia brevicauda, 7TU 85130; Carollia perspicillata, FTU 63655; Carollia sp,
nov,, TTU 82495; Carollia subrufa, TTU 63681; and Carollia castanea, TTU 84903.

1

A New Central American Species from the Carollia brevicauda Complex

Robert 1 Baker, Sergio Solarj and Federico G Hoffmann

In a study of the sequence divergence in the mi-
tochondnal cytoclii’ome-Z) gene, Wright et al. (1999)
suggested that the species Carollia brevicauda as rec¬
ognized currently may consist of two biological spe¬
cies primarily because C. brevicauda was paraphyletic
relative to C. perspicillata. Because of the sample size
of the proposed new species (N=l) as compared to C.
brevicauda (sensu stricto) (N=3) and the geographic
areas represented in the Wright et al. (1999) study, it
was impossible to estimate the limits of geographic
distribution as well as the range of genetic variation
within and among populations and ciades. From a
taxonomic standpoint, it was necessary to establish
the geographic limits of each clade and relate them to
available species-level names. Here we further exam¬
ine the biogeography of the two ciades that Wright et
al. (1999) suggested might represent biological spe¬
cies.

To do this we sequenced 1140 base pairs of the
mitochondrial cytochrome-Z> gene from 21 additional
individuals of the C brevicauda complex representing
as many geographic localities as were available to us
for DNA samples. Additionally, we sequenced the
cytochrome-^j gene from 30 specimens of the other
recognized species of Carollia. The new sequences
were combined with the 10 reported by Wright et al.
(1999) to generate a phylogenetic tree, which provides
additional information concerning the extent of geo¬
graphic variation within members of the genus
Carollia. The new data aie consistent with the pro¬
posal that a major subdivision exists in what currently
IS recognized as Carollia brevicauda (Fig. 1). Three
methods (neighbor-joining, parsimony and likelihood,
as implemented by PAUP*, Swofford, 1999) all pro¬
duce a tree with the same topology as shown in figure
2. The results and implications of these additional data
beyond the resolution of the possibility of two biologi¬
cal species within the Carollia brevicauda complex
will be published elsewhere. Based on the cytochrome-

b data, the South American clade of C. brevicauda
shares a common ancestor with Cawllia perspicillata
after diverging from the Central American representa¬
tives oT Carollia brevicauda. One clade of C brevicauda
{sensu lato) is restricted to Central America, north of
Panama, and includes Western Panama. The other clade
of C. brevicauda is distributed from Eastern Panama
to Bolivia (Fig. 1).

The potential application of the sequence data
from the cytoclirome-6 gene to predicting situations
where unrecognized biological species might exist was
reviewed by Bradley and Baker (2001) and the con¬
clusion that C. brevicauda comprises two biological
species is supported by both distance values, and the
observation that samples from C. brevicauda {sensu
lato) do not fonn a monophyletic clade (genetic spe¬
cies concept, Dobzhansky, 1950; phylogenetic spe¬
cies concept, Cracraft, 1983). In fact, C brevicauda
{sensu stricto) is sister to C. perspicillata^ whereas the
Central American portion of C, brevicauda {sensu lato)
is sister to the common ancestor of C perspicillata
and C. brevicauda {sensu stricto).

This resolution of the geographic distribution of
the two ciades permits assignment of the species level
names that have been recognized as available for
Carollia brevicauda {sensu lato). The type locality of
brevicauda is Rio do Espinto Santo, Brazil. The four
synonyms that are available (Koopman, 1993) are bi¬
color (Wagner, 1840), type locality Brazil; grayi
(Waterhouse, 1838), type locality Pernambuco, Bra¬
zil; lanceolatum (Gray, 1843), type locality South
America; and minor {Gray, 1866), type locality Bahia,
Brazil. However, according to Pine (1972) the name
lanceolatum is a nomen nudum. There is no name avail¬
able for representatives of the clade of C brevicauda
{sensu lato) from Central America. Below we describe
a new taxon to fill this need.

Baker et al,, 2002. A New Central American Species from the Carolua brevicauda Complex
OCCASION.AL Papers, Museum of Texas Tech University 217:1-12

2

Occasional Papers, Museum of Texas Tech University

Figure 1, Geographic distribution of Carollia brevicauda sensu Koopman (1993). Triangles and dots correspond
to collecting localities of individuals for which we have cytochrome-Z? sequence that relate to the two clades
defined by Wright et al, (1999).

Baker et al— A New Species of Carollia

3

Outgroups

—

B

C. castanea

“k

— C

—r C. subrufa

z C. sp. n.

I

I— Lie C. perspicillata
\ I—E

I-^

_

_—^ C. brevicauda

Figure 2. Parsimony tree depicting phylogenetic relationships among the spe¬
cies of Carollia.

4

Occasional Papers, Museum of Texas Tech University

Carollia sowellU New Species

Holotype- Adult male, skin, skull and skeleton,
Museum of Texas Tech, TTU 82495 from Honduras,
Comayagua, Cueva de Taulabe,(14“4U42”N,
87=57>0rw), (UTMzone 16:397511 E 1624803 N),
collected on 11 July, 2001, by a Texas Tech field party
on the Sowell Expedition 2001 led by Robert D, Brad¬
ley. Original number, Ronald A. Van Den Bussche 1869,
TK number 101341 identifies tissue samples and
karyotype preparations that are deposited in the Natu¬
ral Science Research Laboratory, Texas Tech Univer¬
sity

Distribution - From Western Panama, north
through Middle America to the states of Veracruz and
San Luis Potosi in Mexico on the Atlantic versant and
to the state of Oaxaca on the Pacific versant (Fig. 3).

Diagnosis - The initial diagnosis is based on
nucleotide order in the mitochondrial cytochrome-^i
gene. These sequences are deposited in GenBank
(pending) along with the sequences of C hrevicauda
{sensu siricto). The differences between these two

involve nucleotide characters at positions in the cyto-
chrome-i gene (Table 1), A summary of pairwise com¬
parisons within and between C. sowelli, C, brevicauda
and C. perspicillata can be found in Table 2.

Morphologically C sowelli can be distinguished
from C. perspicillata, C, subrufa and C. castanea by
the same morphological suite of characters used to
distinguish C. brevicauda. These include a pelage that
is long and thick with the size of the shafts of the
individual hairs being fine; forearm hairy, hair on nape
of neck with broad, dark band contrasting strongly
with and demarcated from a broad whitish band distal
to it (Hall, 1981).

This is a large species of Carollia, slightly larger
than brevicauda but smaller than perspicillata, with
long and lax dorsal fur. Fur on back tricolor, almost
like brevicauda, but sowelli lacks the dark brown tips
that would make conspicuous its medial light band.
Consequently, the appearance is fighter than
Forearm haired but not furred. Forearm, tibia, and
feet almost the same length as those of brevicauda,
but smaller than those oiperspicillata.

Figure 3, Geographic distribution of Carollia sowelli. Dots indicate sample localities for which cytochrome-^)
sequence data are available. Type locality is identified by an asterisk.

Baker et al—A New Species of Carollia

Table L List of fixed changes in the mitochondrial
cytochrome-h gene among Carollia sowelli,
C. brevicauda and C. perspicillata,
A = adenine, C = cytosine, G = Guanine, and

T= Thymine.

Position

C- brevicauda

C. perspiciliala

C sowelli

39

T

T

C

72

C

T

C

102

C/T

T

c

183

T

C

c

207

C

T/C

T

276

c

T

C

309

T

C

C

348

C

C/T

T

363

c

T

T

390

A

A

C

483

C

A

T

573

T

C

GT

585

G

A/G

C

588

C/T

T

C

603

C

C

T

630

c

C

T

675

C/T

T

C

708

c

C

T

71 1

T

T/G

C

759

T

T/C

C

777

G/T

C

T

816

G

G

A

885

T

T

C

888

G

G

A

912

A

A

G

925

G/A

G

A

999

C

C

T

Skull larger than brevicauda especially in its total
length; but also with a longer palate, longer maxillary
and mandibular toothrow, mandibular length, and coro-
noid height (Table 3). There is a gap between the
upper premolars, but the space is highly variable. It is
because the posterior edge of the anterior premolar
(PM3), which is almost straight, and the anterior pro¬
jection of the cingulum of the posterior premolar
(PM4), wliich is not well developed but, always present
(Fig. 4). Bas 1 sphenoidal pits are elongated. Supraor¬
bital processes are well-developed, and the interorbital
(postorbital) breadth is conspicuously narrower. The
posterior palate is long and wide and not as slender as
in perspicillata. Lateral projection of the mastoid pro¬
cess IS evident on the external side of the skull, form¬
ing a low crest that connects with the lambdoidal crest.

5

The anterior projection of this process is low, and fused
to the side of the skull Just behind the middle ear.

Selected measurements. - External measurements
(in millimeters) recorded in the field by Ronald A. Van
Den Bussche are: total length - 66; tail length - 6;
hind foot - 10; ear - 20; tragus - 10. Weight was 16.5
grams. Length of forearm on the dried specimen is
41.8. Cranial measurements (in millimeters) ofthe
holotype (Fig. 4) are as follows: Greatest length of
skull - 23.1; condylobasal length - 21.2; breadth of
brain case - 9.1; depth of brain case - 9.0; mastoid
breadth - 11.5; length of mandibular toothrow - 8.5;
length of maxillary toothrow - 7.4; post-orbital breadth-
5.6; width across upper canines - 5.6; length of man¬
dible - 14.6 (see also Table 3).

Karyological Data.- The karyotype of the holo¬
type is indistinguishable from that reported by Patton
and Gardner (1971) for C. brevicauda. The diploid
number is 21 and the fundamental number is 36. The
X is a subtelocentric and tlie 2 Ys are acrocentnc.

Lower jaw and the lower toothrows are bowed,
but not to the same extent as seen in brevicauda. Base
and cingulum of lower canines conceal in part the ex¬
ternal lower incisors. The middle lower incisors are
enlarged, and bigger than the external ones, but not as
much as those of perspicillata. Little or no space
between the premolars although some variation is
present. As seen in other species of Carollia, there is
some evidence of sexual dimorphism, with develop¬
ment of processes and crests more conspicuous in
the males.

A full comparison of Carollia sowelli, witli its
congeners is beyond the scope of this description.
There is a previously recognized wide range of mor¬
phological (Pine, 1972; Owen et al., 1984; lum and
Engstrom, 1998) and morphometric (McLeilan, 1984;
Owen et al., 1984) variation between and within spe¬
cies of Carollia. McLeilan (1984) provides a refer¬
ence for the scale of the morphometric variation in
terms of the geographic range of the species, even
recognizing the existence of a large difference between
populations of the northern range (our new species,
see above) and the southern range (C. brevicauda as
here restricted) of her concept of C brevicauda.

6

Occasional Papers, Museum of Texas Tech University

Baker et al-—ANew Species of Carolua

1

Table 2. Average genetic distance (uncorrected) between and within samples
f?/Carollia sowelli, C. brevicauda and C* perspicillata.

C. brevicauda

C. perspicdlata

C sowelli

C, brevicauda

1,98 ±0.11%

C. perspicillata

3.67 ±0.02%

1.46 ±0.05%

C. sowelli

4.95 ±0.03%

4.79 ±0.02%

1.58 ±0,17%

Table 3. Measurements ^?/Carollia sowelli new species (n=I5) and C. brevicauda
{n=I0), For each measurement, the average plus/minus one standard deviation is
shown. Differences are significant (P < 0.05) in the variables marked with *

brevicauda

sowelli

Braincase breadth (BRJ3)

9.570 ±0.190

9.595 ±0.177

Coronoid height (CH)*

4.821 ±0.175

5.005 ± 0.240

Greater skull length (GSL)*

21.711 ±0,465

22,095 ±0.360

Least interorbital breadth (LIB)

5.492 ±0.131

5.465 ±0.150

Breadth of upper M2 (M2M2)

7,548 ±0.248

7.482 ±0.238

Mandibular length (MDL)*

13.973 ±0.503

14.498 ±0.470

Mastoid breadth (MST)

10.797 ±0.295

10.980 ±0.300

Maxi liar toothrow (MTR)’*'

6.934 ±0.195

7.131 ±0.113

Mandibular toothrow (MDTR)*

7.508 ± 0.265

7.788 ±0.165

Palate length (PL)*

9.036 ±0.326

9.495 ± 0.220

Rostra! breadth (RB)

4.817±0.211

4,7S7±0,169

Supraorbital breadth (SOB)

6.340 ±0.225

6.362 ±0,231

Intraspecific Genetic Variation in the
Cytochrome-^ gene within CarolUa sowelli

We have sequenced 14 individuals from Middle
America that are referable to C. sowelli^ and the cyto-
chrome-i» variation is partitioned into 2 lineages that
are quite distinct. The average distance values that sepa¬
rate those clades are 3.6%. Perhaps the most striking
aspect of these data is the nature of the changes rela¬
tive to the codon positions when compared to the dif¬
ferences that distinguish C sowelli from C. brevicauda.
Of the 19 fixed changes between C sowelli and C
brevicauda (Table 1) all are in 3""* codon positions, in¬
volving 17 transitions and 2 transversions and no fixed
amino acid replacements. Within C. sowelli, (Table 4)
the 2 clades are distinguished by 23 fixed differences,
which involve 8 fixed changes in P' codon positions,
1 in the 2”^ codon position, and 14 in 3"'' codon posi¬

tions. These fixed differences include 22 transitions
and 1 transversion, and involve 4 amino acid replace¬
ments. In each of these amino acid changes one clade
of C. sowelli is unique from C. brevicauda, whereas
the other shares the amino acid condition in C.
brevicauda. hi all 4 examples the unique condition is
present in the specimens from western Panama and
Costa Rica.

The application of genetic data to taxonomic rec¬
ognition is a relatively unexplored field, but the general
ideas have been presented philosophically (Avise and
Walker, 1999; Bradley and Baker, 2001; Cracraft 1983,
Templeton, 1989, 2001). Clearly the magnitude of dif¬
ferences that separate the C. sowelli clades typically is
present in other species of mammals that have been
recognized on a morphological basis (Bradley and Baker,
2001). Indeed, C. sowelli may be a composite species

8

Occasional Papers, Museum of Texas Tech University

Table 4. Fixed changes in the cytochrome-h between the
2 lineages of C, so we 111. A - adenine, C = cytosine,
G - Guanine, and T = Thymine.

Position

Northwest

Southeast

27

C

A/G

84

T

C

117

C

A

180

T

C

264

C

T

291

C

T

304

T

C

444

C

T

459

T

C

489

A

G

573

C

T

609

T

C

672

C

T

685

A

G

688

T

C

721

A

G

795

C

T

846

C

T

886

c

T

1066

A

G

1069

C

T

1078

T

C

1106

T

C

or alternatively, may be comprised of two subspecies.
While all these are intriguing possibilities, the best so-
lution to these questions must await resolution from
other forms of data (morphology, nuclear genes, etc.).

Etymology - It is our pleasure to name this spe¬
cies in honor of Mr. James E. Sowell, who has been a
major benefactor to Texas Tech University Mr. Sowell
has funded the Sowell Expeditions, which have tre¬
mendously benefited the Natural Science Research
Laboratory’s research collections and provided an op¬
portunity for many Tech students to experience the
natural history and ecology of the tropics.

AcKNOW'LEDGMENTS

We thank Robert Bradley for leading the expedi¬
tion that collected the holotype as well as other speci¬
mens valuable to this study. We thank Ron A. Van Den
Bussche for preparing the holotype and Meredith
Elamilton for karyotypic preparation. We thank Miguel
Quintana for logistical and organizational support for
the trip. Other members of the field party were Brian
Amman, Dann Carroll, Nevin Durish, Carl Dick, Steve

Hoofer, Francisca Mendez-Harclerode, Lisa Mitchell,
Serena Reeder, John Suchecki, with the collaboration
of Reyna Teresa Velasquez and Catalina Sherman of
the Honduras Secretaria de Salud. For discussions on
the description of species using genetic data we thank
Robert Bradley, David Hafner, Ennque Less a and Bruce
Patterson.

Baker et al—A New Species ok Carolua

9

List of Specimens Examined

Specimens examined and their geographic locali¬
ties are given below: TK numbers correspond to
samples from the frozen tissue collection at the Natu¬
ral Science Research Laboratory from Texas Tech Uni¬
versity, Lubbock, Texas; MVZ numbers correspond
to samples from the Museum of Vertebrate Zoology,
Berkeley, California; NK to the Museum of Southwest¬
ern Biology (Albuquerque), FMNH and MDE num¬
bers correspond to samples from the Field Museum
of Natural History, Chicago, Illinois. Voucher number
are given in parentheses.

Carollia hrevicauda- BOLIVIA: Santa Cruz,
BuenRetiroNK 12171 (MSB 55142); Santa CruzNK
15417 (MSB 59775); ECUADOR: Esmeraldas, San
Lorenzo TK 104530 (TTU 85302); Napo, Parque
NacionalYasuni FMNH37060;GUYANA: Northwest
District, Baramita TK 86502; PANAMA: Panama,
Parque Nacional Altos de Campana FMNH 38117;
PERU: Loreto, Aguas Negras TK 46009, TK 46010;
Cuzco, La Convencion TK 70412; SURINAME;
Saramacca, Raleigh Falls TK 10218 (CMNH 63727);
VENEZUELA: Barinas, Bariiiitas TK 19316 (CMNH
78409); Bolivar, El Palmar TK 19273 (CMNH 78400).

Carollia castanea- BOLIVIA: Beni, Yucumo NK
25385 (MSB 68356); Cochabama, Villa Tunaria NK
30033 (MSB 70298); Sajta NK 30150 (MSB 70297);
COSTARICA: Limon, Estacioii Biologica Cano Palma
FMNH 44029; Ibrtuga Lodge FMNH 44016; ECUA¬
DOR: Esmeraldas, San Lorenzo TK 104506 (TTU
85278); TK 104508 (TW 85280), TK 104681 (TTU
85453); Napo, Parque Nacional Yasuni FMNH 37061,
FMNH 37065; HONDUIUVS: Comayagua, Cueva de
TaulabeTK 101378 (TTU 84037); Atlantida, Lancetilla
TK 101462 (TTU 84121); PANAMA: Chinqui,Ojo de
Agua FMNH 38156; Darien, Parque Nacional Darien
FMNH 38195-

Carolliaperspicillata- BRAZIL: Minas Gerais,
Municipio de Caratinga MVZ 185533; Pernambuco,
Miinicipio Tamandare MVZ 185518; Rio Grande do
Norte, Municipio Baia Formosa MVZ 185806; Sergipe,
Municipio Santo Amaro Das Brotas MVZ 185813; EC¬
UADOR: Esmeraldas, San Lorenzo TK 104613 (TTU
85385), TK 104631 (TTU 85403); Napo, Parque
Nacional Yasuni FMNH 37084, FMNH 37107; GUA¬
TEMALA: El Peten, Poptun FMNH 31809; GUYANA:
Berbice District, Dubulay Ranch TK 86671, TK 86691;
North West District, Baramita TK 86503; MEXICO:
Campeche, Escarcega FMNH 33206; Chiapas, Agua
Azul NK 8644 (MSB 55645), NK 8645 (MSB 55643);
Quintana Roo, Laguna Noh-Bee FMNH 30973; Tulum
MDE 6004; PERU: Cuzco, 1^ Convencion TK 70435;
SURINAME: Nickene, Kabalebo TK 17466 (CMNH
68804); VENEZUELA: Bannas, Barinitas TK 19315
(CMNH 78397).

Carollia sowelli- COSTA RICA: Limon, Estacion
Biologica Cano Palma FMNH 44027; Tortuga Lodge
FMNH 44017; GUATEMALA: £1 Peten, Poptun FMNH
31769; Poptun FMNH 31805, FMNH 31824; HON¬
DURAS: Francisco Morazan, Parque Nacional La Tigra
TK 101005 (TTU 83668), TK 101010 (TTU 82496),
TK 101013 (TTU 82497); Comayagua, Cueva de
Taulabe TK 101341 (TTU 82495), TK 101377 (TTU
82498); MEXICO: Chiapas, Agua Azul NK 8641 (MSB
55644); Quintana Roo, Laguna Noh-Bec FMNH 30976;
Tabasco, Jonuta FMNH 30002; PANAMA: Chiriqui,
Oj 0 de Agua FMNH 38140.

Carollia subrufa - EL SALVADOR: Auachapan,
El Refugio TK 15818; MEXICO: Jalisco, Chamela TK
19550 (TTU 37719), TK 19551 (TTU 37720).

Loan of Tissues

Tissue loans were generously provided by C.
Cicero and J. Patton from the Museum of Vertebrate
Zoology (Berkeley), C, Parmenter, J. Salazar and T.
Yates from the Museum of Southwestern Biology (Al¬

buquerque), M. Engstrom and B. Lim from the Royal
Ontario Museum (Ontario), and R, Monk from the
Natural Science Research Laboratory at Texas Tech
University.

10

Occasional Papers, Museum of Texas Tech University

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Wright, A, J., R. A. Van Den Bussche, B. K. Lim, M. D. Engstrom,
and R. J. Baker, 1999. Systcmatics of the genera Carollia
and Rhinophylla based on the Cytochrome-i Gene. Jour¬
nal of Manunalogy 80(4): 1202-1213.

Baker et al—A New Species of Carollia

11

Addresses of authors:

Robert J. Baker Federico G, Hoffmann

Department of Biological Sciences and Museum Department of Biological Sciences

Texas Tech University Texas Tech University

Lubbock, TX 79409-3 BI Lubbock, TX 79409-3131

e-mail: rjbaker@ttu.edu e-mail: fhoffmann@ttacs.ttu.edu

Sercio Solarj

Museum of Texas Tech University
Box 43191

Lubbock, TX 79409-3191
e-mail: ssolari@ttacs. itu.edu

Publications of the Museum of Texas Tech University

It was through the efforts of Horn Professor J Knox Jones, as director of Academic
Publications, that Texas Tech University initiated several publications series including the
Occasional Papers of the Museum. This and future editions in the series are a memorial to his
dedication to excellence in academic publications. Professor Jones enjoyed editing scientific
publications and served the scientific community as an editor for the Journal of Mammalogy,
Evolution, The Texas Journal of Science, Occasional Papers of the Museum, and Special
Publications of the Museum. It is with special fondness that we remember Dr. J Knox Jones.

Institutional subscriptions are available through the Museum of Texas Tech University,
attn: NSRL Publications Secretary, B ox 43191, Lubbock, TX 79409-3191. Individuals may also
purchase separate numbers of the Occasional Papers directly from the Museum of Texas Tech
University.

Layout and Design; Jacqueline B, Chavez

Cover Design: R. Richard Monk

Copyright 2002, Museum of Texas Tech University

All rights reserved. No portion of this book may be reproduced in any form or by any means,
including electronic storage and retrieval systems, except by explicit, prior written permission
of the publisher.

This book was set in Times New Roman and printed on acid-free paper that meets the guidelines
for pennanence and durability of the Committee on Production Guidelines for Book Longevity of
the Council on Library Resources,

Printed: 31 October, 2002

Library of Congress Cataloging-in-Publication Data

Occasional Papers, Number 217
Series Editor: Robert J. Baker

A NEW CENTRAL AMERICAN SPECIES FROM THE CAROLLIA BREVICAIJDA COMPLEX
By: Robert J, Baker, Sergio Solan and Federico G. Hoffmann
ISSN 0149-175X

Museum of Texas Tech University
Lubbock, TX 79409-3191 USA
(806)742-2442